中华鳖肠道、脾脏和脑垂体的主要功能细胞学研究
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摘要
中华鳖是我国重要的水产养殖动物,具有很高的营养价值和药用价值。在生物学方面,中华鳖属于爬行纲,具有冬眠现象和变温特点,其消化、免疫和内分泌等的生理活动和代谢过程有别于其它动物。随着我国人工饲养的规模不断扩大,中华鳖一些传染病的流行和扩散也越来越严重。以往,关于中华鳖的试验研究较多都集中在与疾病相关领域,对于其特殊结构的基础性研究,资料比较缺乏。本文以具有我国特色的中华鳖为模式动物,详细研究了其代表性的器官组织：消化系统的肠道,免疫系统的脾脏,内分泌系统的脑垂体等的细胞学特征,并分析了其功能意义。论文主要内容包括鳖肠道黏膜机械屏障和免疫屏障细胞组成和结构特征；Cajal间质细胞在肠道的分布差异和超微结构特征；脾脏椭球高内皮毛细血管的发现及淋巴细胞迁移的细胞学基础；腺垂体内分泌细胞超微结构及其季节性变化。从以上三个方面,阐述了中华鳖的细胞特征及其意义,为进一步研究其生理功能、免疫机能、冬眠期和非冬眠期的生理周期变化奠定了理论的基础,同时为人工饲养、疾病预防和治疗方面提供科学依据。
试验Ⅰ中华鳖肠道上皮内黏膜屏障的结构与细胞特征通过光镜和电镜观察正常情况下,中华鳖肠道上皮内黏膜屏障的结构特征,统计分析了中华鳖不同月份肠免疫屏障的主要成分—上皮内淋巴细胞(IEL)在小肠和大肠的分布规律。结果显示：小肠黏膜上皮为单层柱状或假复层柱状上皮。黏膜上皮顶部的上皮细胞之间排列紧密,有发达的紧密连接,中间连接和桥粒等细胞连接复合体。黏膜上皮中部和基底部的上皮细胞之间也存在桥粒和嵌合连接。小肠黏膜上皮大多数上皮细胞胞质内含有少量线粒体,少量上皮细胞胞质含有丰富线粒体和脂滴。大肠黏膜上皮为假复层柱状上皮,所有的上皮细胞胞质内未见脂滴分布。黏膜上皮顶部的上皮细胞排列紧密,而中部和基底部的上皮细胞排列疏松,细胞之间的间隙宽大而明显。许多黏液细胞散在分布于整个肠道的黏膜上皮细胞之间,有的细胞胞质内黏液颗粒电子密度低,而有的电子密度高。黏液细胞分泌的黏液覆盖于肠黏膜表面。肠黏膜上皮内缺乏微皱褶细胞。肠道黏膜上皮细胞之间存在直径为1-7gm的通道,有时可见通道内有淋巴细胞分布。IEL(直径4-9μm)位于肠道黏膜上皮细胞之间,并在它们之间伸出伪足,显示出IEL具有迁移趋势。肠腔内可见完整的游离淋巴细胞。在上皮细胞之间发现了浆细胞的分布,一般位于肠道黏膜上皮的中部和基底部,浆细胞有两种形态：一种粗面内质网为扩张状态,另一种是扁平囊状。IEL细胞数量从5月、9月到12月逐渐减少。以上试验结果提示,鳖肠道黏膜上皮具有黏膜屏障功能。上皮细胞及其紧密连接复合体、黏液细胞分泌的黏液构成机械屏障；浆细胞、IEL和通道构成免疫屏障。鳖肠道黏膜免疫细胞分布也受季节影响。
试验Ⅱ中华鳖肠黏膜免疫屏障中CD3+T、CD22+B淋巴细胞的分布采用免疫细胞化学SABC法,观察中华鳖肠道免疫屏障内CD3+T、CD22+B淋巴细胞的分布特征,统计并分析了中华鳖各肠段CD3+T淋巴细胞、CD22+B淋巴细胞的分布规律。结果发现：CD3+T淋巴细胞主要分布在固有膜内,少量见于黏膜上皮内。而大部分CD22+B淋巴细胞分布于黏膜上皮,固有膜内却少见。在上皮内,CD3+T淋巴细胞在小肠从前到后依次递增,小肠后段数量最多；而在大肠CD3+T淋巴细胞依次递减。在固有膜内,CD3+T淋巴细胞在小肠中段密度最高,而在大肠依次递减。上皮内,固有膜内CD3+T淋巴细胞各肠段相比,大部分肠段之间差异显著。固有膜CD3+T淋巴细胞数量是上皮内的3.11倍,小肠CD3-T淋巴细胞数量是大肠的2倍。对于CD22+B淋巴细胞而言,在上皮内,CD22+B淋巴细胞在小肠后段数量最多,而在大肠从前到后逐渐减少；在固有膜,CD22+B淋巴细胞从小肠前到小肠后段依次减少,在大肠依次降低。上皮内CD22+B淋巴细胞各肠段相比,大部分肠段之间差异不显著,固有膜内CD22+B淋巴细胞在各肠段之间没有显著差异。上皮内CD22+B淋巴细胞数量是固有膜的7.99倍,小肠CD22+B淋巴细胞数量是大肠的1.75倍。以上试验结果说明鳖肠道各段免疫屏障中T、B淋巴细胞的分布存在差异,提示它们在肠道黏膜免疫应答中发挥着不同的生物学作用。
试验Ⅲ中华鳖肠道S-100蛋白免疫细胞化学反应研究应用免疫组织化学技术,研究了S-100蛋白阳性细胞在中华鳖肠道的分布规律。结果显示：S-100蛋白阳性细胞可以分为两种类型：第一种类型S-100阳性细胞形态不规则,细胞突起粗细长短不一,主要分布于小肠固有膜和黏膜下层,大肠内几乎没有阳性细胞分布。第二种类型S-100蛋白阳性细胞分为双极和星形细胞。双极S-100蛋白阳性细胞突起有的长,有的短。它们分布于小肠和大肠的黏膜下层和环形肌之间,环形肌内,环形肌和纵形肌之间,大肠的黏膜下层。星形S-100蛋白阳性细胞仅分布于大肠前段固有膜,这些细胞含有多个突起,突起之间相互交织在一起形成网络状结构。S-100蛋白是树突状细胞的特异性抗体,但在Cajal司质细胞、神经胶质细胞也有表达。根据形态和分布位置,本实验第一种类型S-100蛋白阳性细胞为树突状细胞,第二种类型S-100蛋白阳性细胞可能是Cajal间质细胞或神经胶质细胞。目前研究认为,树突状细胞具有抗原呈递作用,Cajal间质细胞是肠道蠕动的起搏点,神经胶质细胞可以维护肠黏膜上皮完整性。因此,推测树突状细胞参与的免疫屏障功能主要在鳖小肠,Cajal间质细胞和神经胶质细胞分别具有调节肠道运动和肠黏膜屏障功能。
试验ⅣCajal间质细胞在中华鳖肠道的分布差异和超微特征本文应用c-Kit免疫细胞化学技术和透射电镜研究了Cajal间质细胞(ICC)在中华鳖肠道中的分布和超微结构特征。c-Kit免疫细胞化学反应显示,Cajal间质细胞分为双极和星形细胞。双极c-Kit+ICC细胞突起有的长,有的短或不明显。这些细胞分布于大肠黏膜下层,小肠和大肠的黏膜下层与环肌层之间、环形肌内、环形肌和纵形肌之间、浆膜。在黏膜下层,双极c-Kit+ICC围绕血管分布。在黏膜下层与环形肌之间,环形肌和纵形肌之间,呈线形或呈簇分布,有时围绕在血管周围。在环形肌内,双极c-Kit+ICC围绕在血管周围或与平滑肌细胞平行分布。在浆膜内,双极c-Kit+ICC散在分布。星形c-Kit+ICC细胞只分布在大肠固有膜内,这些细胞具有多个突起,突起之间相互交织形成网状结构,主要围绕在血管周围分布。电镜下,双极和星形ICC细胞胞质内含有线粒体,膜内陷小泡,粗面内质网和多个不规则的胞浆突起。ICC细胞与ICC细胞之间、ICC细胞与神经、ICC细胞与平滑肌细胞之间具有缝隙连接。以上试验结果提示中华鳖肠道ICC细胞具有独特的分布特点和超微结构,其形态特点与其在维护肠道蠕动中的重要作用相一致。
试验V中华鳖脾脏椭球高内皮毛细血管的发现及淋巴细胞迁移的细胞学基础本文应用光镜和电镜等技术,对中华鳖脾脏白髓组织结构进行了研究,发现中华鳖脾脏椭球血管不同于其它动物,其血管属于高内皮毛细血管,内皮细胞呈高柱状或立方形,相当于哺乳动物高内皮微静脉。脾脏内存在血-脾屏结构,一方面可以阻止外来异物抗原(碳粒)进入椭球周围淋巴鞘(PELS),另一方面可以形成特殊临时性通道,允许淋巴细胞在其中迁移,到达PELS.通道结构是由网状细胞、网状纤维、迁移细胞(淋巴细胞、巨噬细胞、红细胞)、支持细胞、椭球相关细胞、通道共同构成的复合结构。它们形成了一个有利于淋巴细胞运动的微环境。本试验发现淋巴细胞迁移过程如下：椭球高内皮毛细血管内皮细胞之间形成通道,淋巴细胞沿着通道,穿过椭球高内皮毛细血管基膜,到达椭球；淋巴细胞在支持细胞及其突起作用下,沿着通道,从椭球到达PELS外；淋巴细胞与椭球相关细胞及其突起相联系,从PELS外进入PELS。以上试验结果为深入研究爬行动物淋巴细胞归巢提供了细胞学基础。
试验Ⅵ中华鳖腺垂体内分泌细胞超微结构及其季节性变化研究采用透射电镜技术对冬眠期和非冬眠期的中华鳖腺垂体中各种内分泌细胞的超微结构进行了观测分析。结果显示,腺垂体主要有5种内分泌细胞：生长激素(GH)细胞、催乳激素(PRL)细胞、促甲状腺激素(TSH)细胞、促肾上腺皮质激素(ACTH)细胞和促性腺激素(GTH)细胞。它们各有其超微结构特征：GH细胞呈圆形或椭圆形,直径为10-12μm；分泌颗粒分布稠密,呈圆形。PRL细胞椭圆形或不规则形,长径为7-9μm,细胞核形状不规则,胞质电子密度高。TSH细胞呈卵圆形或不规则的多角形,直径为8-12μm,细胞核椭圆形,分泌颗粒数量不多,主要分布在细胞周边。ACTH细胞不规则,直径8-10μm,细胞核为椭圆形,分泌颗粒较少。GTH细胞圆形或卵圆形,直径12-15μm,细胞核为圆形或椭圆形,胞质内有大、小两种分泌颗粒。同时,鳖腺垂体中各种内分泌细胞的结构特征和分泌颗粒密度等存在一定的季节性变化。冬眠期,GH细胞胞质内细胞器比非冬眠期明显减少；TSH细胞在非冬眠期数量高于冬眠期；冬眠期内GTH细胞内的分泌颗粒出现融合变大的现象,颗粒直径明显大于非冬眠期,最大的可以达到2μm。PRL细胞和ACTH细胞超微结构没有明显的季节变化。
Soft-shelled turtle, Pelodiseus Sinensis distributed widely in China and this species was famous for its important value in economy and in pharmacy. In the biology aspect, Pelodiseus Sinensis were belonged to Reptilia, they were ectothermic animal and had hibernation. The physiological action and metabolic process of the digest, immunity and endocrine secretion were different from other kinds of animals. With the scale of the turtle hand-feeding has rapid development, the communicable diseases became more and more serious. Most of the scholars were focused on the researches of turtle disease, there was almost no studies on the special structural feature of turtle. The present study was used Pelodiseus Sinensis as research target. We selected three typical tissues in the turtle, they were intestine of the digestive system, the spleen of the immune system and the pituitary gland of the endocrine system. The text included three parts, in the first part, we examined the structural feature and cell component of mechanical barrier and immunologic barrier, the distribution and ultrastructure of interstitial cells of Cajal in the turtle gut. In the second part, the ellipsoid high endothelial capillary and lymphocytes migration cytologic basement in the spleen were studied. In the third part, the ultrastructure and seasonal change of endocrine cell were investigated in the adenohypophysis. All the aboved three parts, the turtle cytological feature and their significance were expounded in detail. The studies provided theoretical foundation for the further research on the physiologic and immunologic function, physiological aspection during the hibernation and non-hibernation of the turtle. At the same time, they also provided sciencific basis for the hand-feeding, disease prevention and cure in the Pelodiseus Sinensis.
Experiment I The fine structure and cell character of barrier in the intestine mucosal epithelium of Chinese soft-shelled turtle, Pelodiseus Sinensis. Light and transmission electron microscopy were used to study the structural feature of barrier and the distribution pattern of intestinal intraepithelial lymphocyte (IELs) in different month in the normal turtles intestine mucosal epithelium. IELs was one of the most important part of the immunologic barrier. The results showed that in the small intestine, the mucosal epithelium was single layered or pseudostratified columnar epithelium. In the apical parts of the epithelium, enterocytes were arranged tightly. There have cell junctional complex of tight junction, intermediate junction and desmosomes. In the middle and basal parts of mucosal epithelium, enterocytes were connected by desmosomes and interdigitations. In the small intestinal epithelium, most of enterocytes contained several mitochondria in the cytoplasm. A few enterocytes were loaded with numerous mitochondria and lipid droplets. In the large intestine, the epithelium was pseudostratified. All the enterocytes did not contain lipid droplets. Enterocytes were arraged tight in the apical parts of the epithelium. They were arranged loosely and formed the obvious intercellular spaces in the middle and basal parts of mucosal epithelium. Numerous mucous cells were scattered between the enterocytes of the entire mucosal epithelium. Some of them contained mucus granules of low electron density in the cytoplasm, others had the high electron density. The mucus was secreted by mucous cells and covered the surface of intestinal mucosa. No microfold cells (M cells) were observed throughout intestinal epithelium. In the whole intestine mucosal epithelium, channels with a diameter of 1-7μm were found between the enterocytes. Sometimes, a lymphocyte resided in the channel. IELs (4-9μm in diameter) located in the whole intestinal epithelium, some of them had protruded pseudopodium between the enterocytes, showing the migration tendency, and others were found in the lumen of the intestine. In the nuclear layer and basal layer of the small and large intestinal epithelium, there were two types of plasma cells, one had the dilated rough endoplasmic reticulum and the other the parallel. The amounts of IELs decreased gradually from May to September, up to December. It was therefore suggested that the intestine mucosal epithelium have the mucosal barrier function in the normal turtle. Enterocytes and their tight junction complex, the mucus of mucous cell were composed of the mechanical barrier. Immunologic barrier contained plasma cell, IEL and channel. The distribution of mucosal immune cell in the turtle intestine was affected by the season.
Experiment II Distribution of CD3+T、CD22+B lymphocytes in the intestine mucosal immunologic barrier of Chinese soft-shelled turtle, Pelodiseus Sinensis. Using immunocytochemistry SABC, we studied the distribution feature of CD3+T lymphocytes、CD22+B lymphocytes in the intestine mucosal immunologic barrier of Pelodiscus sinensis. At the same time, the distribution regularity of the CD3+T、CD22+B lymphocytes have been statisticed and analyzed in the different segment of turtle intestine. The results showed that CD3+T lymphocytes predominanted in the lamina propria and fewer in the epithelium. By contrast, great numbers of CD22+B lymphocytes were present in the epithelium and small numbers in the lamina propria. In the epithelium, the density of CD3+T lymphocytes in the distal small intestine was much greater than in any region of the small intestine. They increased from proximal to distal small intestine, decreased from proximal to distal large intestine. In the lamina propria, the highest density of CD3+T lymphocytes occurred in the middle small intestine. They reduced from proximal to distal large intestine. In the epithelium and lamina propria, CD3+T lymphocytes showed significant difference among the most segment of small and large intestine. Throughout the intestine, the total quantity of CD3+T lymphocytes in the lamina propria were 3.11 times of these cells in the epithelium. The number of CD3+T lymphocytes in the small intestine were 2 times of these positive cells in the large intestine. With regard to CD22+B lymphocytes, in the epithelium, the highest numbers of these cells were found in the distal small intestine, and they decreased from proximal to distal large intestine. In the lamina propria, CD22+B lymphocytes decreased in turn from proximal to distal small intestine, reduced from proximal to distal large intestine. In the epithelium, the amount of CD22+B lymphocytes showed no obvious difference during the most segment of small and large intestine. In the lamina propria, there was no remarkable difference among the each segment of small and large intestine. The numbers of CD22+B lymphocytes in the small intestine were 1.75 times of in the large intestine, and the total numbers of these cells in the epithelium were 7.99 times of these positive cells in the lamina propria of the whole intestine. The distributional difference of T and B lymphocytes in the turtle intestinal immunologic barrier suggested that they may play the different biologic function in the immune response of the intestinal mucosa.
ExperimentⅢThe immunocytochemistry reaction of S-100 protein in the intestine of Chinese soft-shelled turtle, Pelodiseus Sinensis. By using immunohistochemistry, this paper studied the distribution of S-100 protein positive cells in the Pelodiscus sinensis intestinal tract. S-100 protein positive cells were classified into two groups. S-100 protein positive cells in the first group had irregular shape and processes with different thickness, length. Most of these cells were located in the lamina propria and submucosa of small intestine. Fewer cells were found in the large intestine. Cells in the second group included bipolar and stellate S-100 protein positive cells. Bipolar S-100 protein positive cells had long and short processes. They were observed in the interspace between the submucosa and circular muscle layer, within the circular muscle layer, as well between the longitudinal and circular muscle layers of whole intesinte and submucosa of large intestine. Stellate S-100 protein positive cells were only found in the large intestine lamina propria, their irregular processes were connected each other forming a network. S-100 protein was regarded as specific antibody for dendritic cell, it also expressed in the interstitial cells of Cajal and glial cells. According to the morphology and distribution of S-100 protein positive cells in the intestine. Cells in the first group were regard as dendritic cells. Cells in the second group were either interstitial cells of Cajal or glial cells. Previous researches suggested that dendritic cells could paly the function of antigen presentation. Interstitial cells of Cajal were regarded as pacemaker and glial cells were maintained the intestinal mucosal epithelium integrity. Base on the findings, we presumed that the mucosal immunologic barrier was correlated with dendritic cells in the turtle small intestine. Interstitial cells of Cajal and glial cells could regulate intestinal movement and barrier function, respectively.
Experiment IV The distributional difference and ultrastructure of interstitial cells of Cajal in the intestinal tract of Chinese soft-shelled turtle, Pelodiseus Sinensis. By using c-Kit immuocytochemistry and transmission electron microscopy, this paper investigated the distribution and ultrastructure of interstitial cells of Cajal (ICCs) in the intestinal tract of Pelodiscus sinensis. The c-Kit immuocytochemistry results showed that ICCs were bipolar and stellate cells. The processes of bipolar c-Kit+ICCs some were long, some were short or no obvious. Bipolar c-Kit+ICCs were present in the large intestine submucosa, in the interspace between the submucosa and circular muscle layer, the circular muscle layer, between circular and longitudinal muscle layer, serosa of small and large intestine. In the submucosa, bipolar c-Kit+ICCs were found arrounding the blood vessel. In the interspace between the submucosa and circular muscle layer, between circular and longitudinal muscle layer, bipolar c-Kit+ICCs showed as shape of line or cluster. Sometimes, these cells were surrounded the blood vessel. In the circular muscle layer, bipolar c-Kit+ICCs were appeared surrounding the blood vessel or parallel to the adjacent smooth muscle cell. Bipolar c-Kit+ICCs scattered in the serosa. Stellate c-Kit+ICCs had numerous long and short processes that formed a network in the villus. These cells were found only in the large intestine lamina propria, they were surrounded the blood vessel. Bipolar and stellate ICCs were identified in the turtle intestine by transmission electron microscope which contain many characteristics; viz., numerous mitochondria, caveolae, thin cytoplasmic extensions or processes, rough endoplasmic reticulum. Gap junctions were formed between ICCs and ICCs, ICCs and nerve, ICCs and smooth muscle cells. All the finding suggested that ICCs have unique cell networks and ultrastructure characterwhich were suitable with its functions.
Experiment V The discovering of ellipsoid high endothelial capillary and lymphocytes migration cytologic base in the spleen of Chinese soft-shelled turtle, Pelodiseus Sinensis. By using microscopy and electron microscopy, the architecture feature of spleen white plup of the Chinese soft-shelled turtle, Pelodiseus Sinensis were examined in detail. The results showed that the ellipsoid capillary was different from capillary in the other kinds of animals, it belonged to high endothelial capillary. The endothelial cells of the ellipsoid capillary were cube or cylindroid. So, it was equaled to the mammalian high endothelial venule. There had blood-spleen barrier (BSB) in the turtle spleen. BSB could prevent the foreign antigen (carbon particles) enter the periellipsoidal lymphocyte sheath (PELS). At the same time, they could form special and temporary channel structure, allow lymphocyte movement and arrive at the PELS. The channel structure was composed of reticular cells and fibers, migrating cells (lymphocytes, macrophages, blood cells), supporting cells, ellipsoid associated cells (EACs), channels. All the components formed a microenvironment specific for lymphocytes migration. The processes of lymphocyte migration was showed following. First, channels were formed between the endothelial cells of ellipsoid high endothelial capillary. Lymphocytes were along the channel and passed the basement membrane of ellipsoid high endothelial capillary, moved in the ellipsoid. Then, by the supporting cells and their processes, lymphocytes along the channel, arrived outside of PELS. At last, lymphocytes were moved from external to the inner PELS by connecting with the EACs and their processes. This study was provided cytologic groundwork for further researching on the reptile lymphocyte homing.
Experiment VI The ultrastructure and seasonal change of endocrine cell in the Pelodiscus sinensis's adenohypophysis. The ultrastructure of endocrine cells in the Pelodiscus sinensis's adenohypophysis were studied by the transmission electron microscope, during the hibernation and non-hibernation period. The results showed that it contained 5 types of endocrine cells in the adenohypophysis.They were growth hormone (GH) cells, prolactin (PRL) cells, thyrotropin (TSH) cells, adrenocorticotrophic hormone (ACTH) cells, gonadotrophin (GTH) cells. Their ultrastructural features appeared characteristically. GH cells were rounded or elliptic in shape, which about 10-12μm in diameter. Numerous round secretory granules were found in the cytoplasm. PRL cells were ovoid or irregular in shape, contained irregular nucleus. The cytoplasm had high electron density. Their diameter was about 7-9μm. TSH cells were ovoid or irregular polygon in shape, contained elliptic nucleus. The diameter was about 8-12μm. A few secretory granules which were occurred in cytoplasmic periphery. ACTH cells had irregular shape and elliptic nucleus. The diameter was reached 8-10μm. Small numbers of secretory granules were observed in the cytoplasm. GTH cells had rounded or elliptic shape, contained round or elliptic nucleus. The diameter was about 12-15μm. They contained two types of secretory granules, the large secretory granules and small ones. In the different seasons, the structural feature and secretory granules of endocrine cell in the turtle adenohypophysis were changed. The organelle of GH cells in the hibernation period was decreased than in the non-hibernation period. TSH cells had the higher numbers in the non-hibernation period. The several secretory granules of GTH cells were fused and became large in the hibernation period, their diameter were larger than in the non-hibernation period, the largest one was reached at 2μm. The ultrastructure of PRL cells and ACTH cells had no obvious differences during the different season.
试验Ⅰ中华鳖肠道上皮内黏膜屏障的结构与细胞特征通过光镜和电镜观察正常情况下,中华鳖肠道上皮内黏膜屏障的结构特征,统计分析了中华鳖不同月份肠免疫屏障的主要成分—上皮内淋巴细胞(IEL)在小肠和大肠的分布规律。结果显示：小肠黏膜上皮为单层柱状或假复层柱状上皮。黏膜上皮顶部的上皮细胞之间排列紧密,有发达的紧密连接,中间连接和桥粒等细胞连接复合体。黏膜上皮中部和基底部的上皮细胞之间也存在桥粒和嵌合连接。小肠黏膜上皮大多数上皮细胞胞质内含有少量线粒体,少量上皮细胞胞质含有丰富线粒体和脂滴。大肠黏膜上皮为假复层柱状上皮,所有的上皮细胞胞质内未见脂滴分布。黏膜上皮顶部的上皮细胞排列紧密,而中部和基底部的上皮细胞排列疏松,细胞之间的间隙宽大而明显。许多黏液细胞散在分布于整个肠道的黏膜上皮细胞之间,有的细胞胞质内黏液颗粒电子密度低,而有的电子密度高。黏液细胞分泌的黏液覆盖于肠黏膜表面。肠黏膜上皮内缺乏微皱褶细胞。肠道黏膜上皮细胞之间存在直径为1-7gm的通道,有时可见通道内有淋巴细胞分布。IEL(直径4-9μm)位于肠道黏膜上皮细胞之间,并在它们之间伸出伪足,显示出IEL具有迁移趋势。肠腔内可见完整的游离淋巴细胞。在上皮细胞之间发现了浆细胞的分布,一般位于肠道黏膜上皮的中部和基底部,浆细胞有两种形态：一种粗面内质网为扩张状态,另一种是扁平囊状。IEL细胞数量从5月、9月到12月逐渐减少。以上试验结果提示,鳖肠道黏膜上皮具有黏膜屏障功能。上皮细胞及其紧密连接复合体、黏液细胞分泌的黏液构成机械屏障；浆细胞、IEL和通道构成免疫屏障。鳖肠道黏膜免疫细胞分布也受季节影响。
试验Ⅱ中华鳖肠黏膜免疫屏障中CD3+T、CD22+B淋巴细胞的分布采用免疫细胞化学SABC法,观察中华鳖肠道免疫屏障内CD3+T、CD22+B淋巴细胞的分布特征,统计并分析了中华鳖各肠段CD3+T淋巴细胞、CD22+B淋巴细胞的分布规律。结果发现：CD3+T淋巴细胞主要分布在固有膜内,少量见于黏膜上皮内。而大部分CD22+B淋巴细胞分布于黏膜上皮,固有膜内却少见。在上皮内,CD3+T淋巴细胞在小肠从前到后依次递增,小肠后段数量最多；而在大肠CD3+T淋巴细胞依次递减。在固有膜内,CD3+T淋巴细胞在小肠中段密度最高,而在大肠依次递减。上皮内,固有膜内CD3+T淋巴细胞各肠段相比,大部分肠段之间差异显著。固有膜CD3+T淋巴细胞数量是上皮内的3.11倍,小肠CD3-T淋巴细胞数量是大肠的2倍。对于CD22+B淋巴细胞而言,在上皮内,CD22+B淋巴细胞在小肠后段数量最多,而在大肠从前到后逐渐减少；在固有膜,CD22+B淋巴细胞从小肠前到小肠后段依次减少,在大肠依次降低。上皮内CD22+B淋巴细胞各肠段相比,大部分肠段之间差异不显著,固有膜内CD22+B淋巴细胞在各肠段之间没有显著差异。上皮内CD22+B淋巴细胞数量是固有膜的7.99倍,小肠CD22+B淋巴细胞数量是大肠的1.75倍。以上试验结果说明鳖肠道各段免疫屏障中T、B淋巴细胞的分布存在差异,提示它们在肠道黏膜免疫应答中发挥着不同的生物学作用。
试验Ⅲ中华鳖肠道S-100蛋白免疫细胞化学反应研究应用免疫组织化学技术,研究了S-100蛋白阳性细胞在中华鳖肠道的分布规律。结果显示：S-100蛋白阳性细胞可以分为两种类型：第一种类型S-100阳性细胞形态不规则,细胞突起粗细长短不一,主要分布于小肠固有膜和黏膜下层,大肠内几乎没有阳性细胞分布。第二种类型S-100蛋白阳性细胞分为双极和星形细胞。双极S-100蛋白阳性细胞突起有的长,有的短。它们分布于小肠和大肠的黏膜下层和环形肌之间,环形肌内,环形肌和纵形肌之间,大肠的黏膜下层。星形S-100蛋白阳性细胞仅分布于大肠前段固有膜,这些细胞含有多个突起,突起之间相互交织在一起形成网络状结构。S-100蛋白是树突状细胞的特异性抗体,但在Cajal司质细胞、神经胶质细胞也有表达。根据形态和分布位置,本实验第一种类型S-100蛋白阳性细胞为树突状细胞,第二种类型S-100蛋白阳性细胞可能是Cajal间质细胞或神经胶质细胞。目前研究认为,树突状细胞具有抗原呈递作用,Cajal间质细胞是肠道蠕动的起搏点,神经胶质细胞可以维护肠黏膜上皮完整性。因此,推测树突状细胞参与的免疫屏障功能主要在鳖小肠,Cajal间质细胞和神经胶质细胞分别具有调节肠道运动和肠黏膜屏障功能。
试验ⅣCajal间质细胞在中华鳖肠道的分布差异和超微特征本文应用c-Kit免疫细胞化学技术和透射电镜研究了Cajal间质细胞(ICC)在中华鳖肠道中的分布和超微结构特征。c-Kit免疫细胞化学反应显示,Cajal间质细胞分为双极和星形细胞。双极c-Kit+ICC细胞突起有的长,有的短或不明显。这些细胞分布于大肠黏膜下层,小肠和大肠的黏膜下层与环肌层之间、环形肌内、环形肌和纵形肌之间、浆膜。在黏膜下层,双极c-Kit+ICC围绕血管分布。在黏膜下层与环形肌之间,环形肌和纵形肌之间,呈线形或呈簇分布,有时围绕在血管周围。在环形肌内,双极c-Kit+ICC围绕在血管周围或与平滑肌细胞平行分布。在浆膜内,双极c-Kit+ICC散在分布。星形c-Kit+ICC细胞只分布在大肠固有膜内,这些细胞具有多个突起,突起之间相互交织形成网状结构,主要围绕在血管周围分布。电镜下,双极和星形ICC细胞胞质内含有线粒体,膜内陷小泡,粗面内质网和多个不规则的胞浆突起。ICC细胞与ICC细胞之间、ICC细胞与神经、ICC细胞与平滑肌细胞之间具有缝隙连接。以上试验结果提示中华鳖肠道ICC细胞具有独特的分布特点和超微结构,其形态特点与其在维护肠道蠕动中的重要作用相一致。
试验V中华鳖脾脏椭球高内皮毛细血管的发现及淋巴细胞迁移的细胞学基础本文应用光镜和电镜等技术,对中华鳖脾脏白髓组织结构进行了研究,发现中华鳖脾脏椭球血管不同于其它动物,其血管属于高内皮毛细血管,内皮细胞呈高柱状或立方形,相当于哺乳动物高内皮微静脉。脾脏内存在血-脾屏结构,一方面可以阻止外来异物抗原(碳粒)进入椭球周围淋巴鞘(PELS),另一方面可以形成特殊临时性通道,允许淋巴细胞在其中迁移,到达PELS.通道结构是由网状细胞、网状纤维、迁移细胞(淋巴细胞、巨噬细胞、红细胞)、支持细胞、椭球相关细胞、通道共同构成的复合结构。它们形成了一个有利于淋巴细胞运动的微环境。本试验发现淋巴细胞迁移过程如下：椭球高内皮毛细血管内皮细胞之间形成通道,淋巴细胞沿着通道,穿过椭球高内皮毛细血管基膜,到达椭球；淋巴细胞在支持细胞及其突起作用下,沿着通道,从椭球到达PELS外；淋巴细胞与椭球相关细胞及其突起相联系,从PELS外进入PELS。以上试验结果为深入研究爬行动物淋巴细胞归巢提供了细胞学基础。
试验Ⅵ中华鳖腺垂体内分泌细胞超微结构及其季节性变化研究采用透射电镜技术对冬眠期和非冬眠期的中华鳖腺垂体中各种内分泌细胞的超微结构进行了观测分析。结果显示,腺垂体主要有5种内分泌细胞：生长激素(GH)细胞、催乳激素(PRL)细胞、促甲状腺激素(TSH)细胞、促肾上腺皮质激素(ACTH)细胞和促性腺激素(GTH)细胞。它们各有其超微结构特征：GH细胞呈圆形或椭圆形,直径为10-12μm；分泌颗粒分布稠密,呈圆形。PRL细胞椭圆形或不规则形,长径为7-9μm,细胞核形状不规则,胞质电子密度高。TSH细胞呈卵圆形或不规则的多角形,直径为8-12μm,细胞核椭圆形,分泌颗粒数量不多,主要分布在细胞周边。ACTH细胞不规则,直径8-10μm,细胞核为椭圆形,分泌颗粒较少。GTH细胞圆形或卵圆形,直径12-15μm,细胞核为圆形或椭圆形,胞质内有大、小两种分泌颗粒。同时,鳖腺垂体中各种内分泌细胞的结构特征和分泌颗粒密度等存在一定的季节性变化。冬眠期,GH细胞胞质内细胞器比非冬眠期明显减少；TSH细胞在非冬眠期数量高于冬眠期；冬眠期内GTH细胞内的分泌颗粒出现融合变大的现象,颗粒直径明显大于非冬眠期,最大的可以达到2μm。PRL细胞和ACTH细胞超微结构没有明显的季节变化。
Soft-shelled turtle, Pelodiseus Sinensis distributed widely in China and this species was famous for its important value in economy and in pharmacy. In the biology aspect, Pelodiseus Sinensis were belonged to Reptilia, they were ectothermic animal and had hibernation. The physiological action and metabolic process of the digest, immunity and endocrine secretion were different from other kinds of animals. With the scale of the turtle hand-feeding has rapid development, the communicable diseases became more and more serious. Most of the scholars were focused on the researches of turtle disease, there was almost no studies on the special structural feature of turtle. The present study was used Pelodiseus Sinensis as research target. We selected three typical tissues in the turtle, they were intestine of the digestive system, the spleen of the immune system and the pituitary gland of the endocrine system. The text included three parts, in the first part, we examined the structural feature and cell component of mechanical barrier and immunologic barrier, the distribution and ultrastructure of interstitial cells of Cajal in the turtle gut. In the second part, the ellipsoid high endothelial capillary and lymphocytes migration cytologic basement in the spleen were studied. In the third part, the ultrastructure and seasonal change of endocrine cell were investigated in the adenohypophysis. All the aboved three parts, the turtle cytological feature and their significance were expounded in detail. The studies provided theoretical foundation for the further research on the physiologic and immunologic function, physiological aspection during the hibernation and non-hibernation of the turtle. At the same time, they also provided sciencific basis for the hand-feeding, disease prevention and cure in the Pelodiseus Sinensis.
Experiment I The fine structure and cell character of barrier in the intestine mucosal epithelium of Chinese soft-shelled turtle, Pelodiseus Sinensis. Light and transmission electron microscopy were used to study the structural feature of barrier and the distribution pattern of intestinal intraepithelial lymphocyte (IELs) in different month in the normal turtles intestine mucosal epithelium. IELs was one of the most important part of the immunologic barrier. The results showed that in the small intestine, the mucosal epithelium was single layered or pseudostratified columnar epithelium. In the apical parts of the epithelium, enterocytes were arranged tightly. There have cell junctional complex of tight junction, intermediate junction and desmosomes. In the middle and basal parts of mucosal epithelium, enterocytes were connected by desmosomes and interdigitations. In the small intestinal epithelium, most of enterocytes contained several mitochondria in the cytoplasm. A few enterocytes were loaded with numerous mitochondria and lipid droplets. In the large intestine, the epithelium was pseudostratified. All the enterocytes did not contain lipid droplets. Enterocytes were arraged tight in the apical parts of the epithelium. They were arranged loosely and formed the obvious intercellular spaces in the middle and basal parts of mucosal epithelium. Numerous mucous cells were scattered between the enterocytes of the entire mucosal epithelium. Some of them contained mucus granules of low electron density in the cytoplasm, others had the high electron density. The mucus was secreted by mucous cells and covered the surface of intestinal mucosa. No microfold cells (M cells) were observed throughout intestinal epithelium. In the whole intestine mucosal epithelium, channels with a diameter of 1-7μm were found between the enterocytes. Sometimes, a lymphocyte resided in the channel. IELs (4-9μm in diameter) located in the whole intestinal epithelium, some of them had protruded pseudopodium between the enterocytes, showing the migration tendency, and others were found in the lumen of the intestine. In the nuclear layer and basal layer of the small and large intestinal epithelium, there were two types of plasma cells, one had the dilated rough endoplasmic reticulum and the other the parallel. The amounts of IELs decreased gradually from May to September, up to December. It was therefore suggested that the intestine mucosal epithelium have the mucosal barrier function in the normal turtle. Enterocytes and their tight junction complex, the mucus of mucous cell were composed of the mechanical barrier. Immunologic barrier contained plasma cell, IEL and channel. The distribution of mucosal immune cell in the turtle intestine was affected by the season.
Experiment II Distribution of CD3+T、CD22+B lymphocytes in the intestine mucosal immunologic barrier of Chinese soft-shelled turtle, Pelodiseus Sinensis. Using immunocytochemistry SABC, we studied the distribution feature of CD3+T lymphocytes、CD22+B lymphocytes in the intestine mucosal immunologic barrier of Pelodiscus sinensis. At the same time, the distribution regularity of the CD3+T、CD22+B lymphocytes have been statisticed and analyzed in the different segment of turtle intestine. The results showed that CD3+T lymphocytes predominanted in the lamina propria and fewer in the epithelium. By contrast, great numbers of CD22+B lymphocytes were present in the epithelium and small numbers in the lamina propria. In the epithelium, the density of CD3+T lymphocytes in the distal small intestine was much greater than in any region of the small intestine. They increased from proximal to distal small intestine, decreased from proximal to distal large intestine. In the lamina propria, the highest density of CD3+T lymphocytes occurred in the middle small intestine. They reduced from proximal to distal large intestine. In the epithelium and lamina propria, CD3+T lymphocytes showed significant difference among the most segment of small and large intestine. Throughout the intestine, the total quantity of CD3+T lymphocytes in the lamina propria were 3.11 times of these cells in the epithelium. The number of CD3+T lymphocytes in the small intestine were 2 times of these positive cells in the large intestine. With regard to CD22+B lymphocytes, in the epithelium, the highest numbers of these cells were found in the distal small intestine, and they decreased from proximal to distal large intestine. In the lamina propria, CD22+B lymphocytes decreased in turn from proximal to distal small intestine, reduced from proximal to distal large intestine. In the epithelium, the amount of CD22+B lymphocytes showed no obvious difference during the most segment of small and large intestine. In the lamina propria, there was no remarkable difference among the each segment of small and large intestine. The numbers of CD22+B lymphocytes in the small intestine were 1.75 times of in the large intestine, and the total numbers of these cells in the epithelium were 7.99 times of these positive cells in the lamina propria of the whole intestine. The distributional difference of T and B lymphocytes in the turtle intestinal immunologic barrier suggested that they may play the different biologic function in the immune response of the intestinal mucosa.
ExperimentⅢThe immunocytochemistry reaction of S-100 protein in the intestine of Chinese soft-shelled turtle, Pelodiseus Sinensis. By using immunohistochemistry, this paper studied the distribution of S-100 protein positive cells in the Pelodiscus sinensis intestinal tract. S-100 protein positive cells were classified into two groups. S-100 protein positive cells in the first group had irregular shape and processes with different thickness, length. Most of these cells were located in the lamina propria and submucosa of small intestine. Fewer cells were found in the large intestine. Cells in the second group included bipolar and stellate S-100 protein positive cells. Bipolar S-100 protein positive cells had long and short processes. They were observed in the interspace between the submucosa and circular muscle layer, within the circular muscle layer, as well between the longitudinal and circular muscle layers of whole intesinte and submucosa of large intestine. Stellate S-100 protein positive cells were only found in the large intestine lamina propria, their irregular processes were connected each other forming a network. S-100 protein was regarded as specific antibody for dendritic cell, it also expressed in the interstitial cells of Cajal and glial cells. According to the morphology and distribution of S-100 protein positive cells in the intestine. Cells in the first group were regard as dendritic cells. Cells in the second group were either interstitial cells of Cajal or glial cells. Previous researches suggested that dendritic cells could paly the function of antigen presentation. Interstitial cells of Cajal were regarded as pacemaker and glial cells were maintained the intestinal mucosal epithelium integrity. Base on the findings, we presumed that the mucosal immunologic barrier was correlated with dendritic cells in the turtle small intestine. Interstitial cells of Cajal and glial cells could regulate intestinal movement and barrier function, respectively.
Experiment IV The distributional difference and ultrastructure of interstitial cells of Cajal in the intestinal tract of Chinese soft-shelled turtle, Pelodiseus Sinensis. By using c-Kit immuocytochemistry and transmission electron microscopy, this paper investigated the distribution and ultrastructure of interstitial cells of Cajal (ICCs) in the intestinal tract of Pelodiscus sinensis. The c-Kit immuocytochemistry results showed that ICCs were bipolar and stellate cells. The processes of bipolar c-Kit+ICCs some were long, some were short or no obvious. Bipolar c-Kit+ICCs were present in the large intestine submucosa, in the interspace between the submucosa and circular muscle layer, the circular muscle layer, between circular and longitudinal muscle layer, serosa of small and large intestine. In the submucosa, bipolar c-Kit+ICCs were found arrounding the blood vessel. In the interspace between the submucosa and circular muscle layer, between circular and longitudinal muscle layer, bipolar c-Kit+ICCs showed as shape of line or cluster. Sometimes, these cells were surrounded the blood vessel. In the circular muscle layer, bipolar c-Kit+ICCs were appeared surrounding the blood vessel or parallel to the adjacent smooth muscle cell. Bipolar c-Kit+ICCs scattered in the serosa. Stellate c-Kit+ICCs had numerous long and short processes that formed a network in the villus. These cells were found only in the large intestine lamina propria, they were surrounded the blood vessel. Bipolar and stellate ICCs were identified in the turtle intestine by transmission electron microscope which contain many characteristics; viz., numerous mitochondria, caveolae, thin cytoplasmic extensions or processes, rough endoplasmic reticulum. Gap junctions were formed between ICCs and ICCs, ICCs and nerve, ICCs and smooth muscle cells. All the finding suggested that ICCs have unique cell networks and ultrastructure characterwhich were suitable with its functions.
Experiment V The discovering of ellipsoid high endothelial capillary and lymphocytes migration cytologic base in the spleen of Chinese soft-shelled turtle, Pelodiseus Sinensis. By using microscopy and electron microscopy, the architecture feature of spleen white plup of the Chinese soft-shelled turtle, Pelodiseus Sinensis were examined in detail. The results showed that the ellipsoid capillary was different from capillary in the other kinds of animals, it belonged to high endothelial capillary. The endothelial cells of the ellipsoid capillary were cube or cylindroid. So, it was equaled to the mammalian high endothelial venule. There had blood-spleen barrier (BSB) in the turtle spleen. BSB could prevent the foreign antigen (carbon particles) enter the periellipsoidal lymphocyte sheath (PELS). At the same time, they could form special and temporary channel structure, allow lymphocyte movement and arrive at the PELS. The channel structure was composed of reticular cells and fibers, migrating cells (lymphocytes, macrophages, blood cells), supporting cells, ellipsoid associated cells (EACs), channels. All the components formed a microenvironment specific for lymphocytes migration. The processes of lymphocyte migration was showed following. First, channels were formed between the endothelial cells of ellipsoid high endothelial capillary. Lymphocytes were along the channel and passed the basement membrane of ellipsoid high endothelial capillary, moved in the ellipsoid. Then, by the supporting cells and their processes, lymphocytes along the channel, arrived outside of PELS. At last, lymphocytes were moved from external to the inner PELS by connecting with the EACs and their processes. This study was provided cytologic groundwork for further researching on the reptile lymphocyte homing.
Experiment VI The ultrastructure and seasonal change of endocrine cell in the Pelodiscus sinensis's adenohypophysis. The ultrastructure of endocrine cells in the Pelodiscus sinensis's adenohypophysis were studied by the transmission electron microscope, during the hibernation and non-hibernation period. The results showed that it contained 5 types of endocrine cells in the adenohypophysis.They were growth hormone (GH) cells, prolactin (PRL) cells, thyrotropin (TSH) cells, adrenocorticotrophic hormone (ACTH) cells, gonadotrophin (GTH) cells. Their ultrastructural features appeared characteristically. GH cells were rounded or elliptic in shape, which about 10-12μm in diameter. Numerous round secretory granules were found in the cytoplasm. PRL cells were ovoid or irregular in shape, contained irregular nucleus. The cytoplasm had high electron density. Their diameter was about 7-9μm. TSH cells were ovoid or irregular polygon in shape, contained elliptic nucleus. The diameter was about 8-12μm. A few secretory granules which were occurred in cytoplasmic periphery. ACTH cells had irregular shape and elliptic nucleus. The diameter was reached 8-10μm. Small numbers of secretory granules were observed in the cytoplasm. GTH cells had rounded or elliptic shape, contained round or elliptic nucleus. The diameter was about 12-15μm. They contained two types of secretory granules, the large secretory granules and small ones. In the different seasons, the structural feature and secretory granules of endocrine cell in the turtle adenohypophysis were changed. The organelle of GH cells in the hibernation period was decreased than in the non-hibernation period. TSH cells had the higher numbers in the non-hibernation period. The several secretory granules of GTH cells were fused and became large in the hibernation period, their diameter were larger than in the non-hibernation period, the largest one was reached at 2μm. The ultrastructure of PRL cells and ACTH cells had no obvious differences during the different season.
引文
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