贝氏高原鳅消化系统的结构与发育
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摘要
2005年12月至2006年12月，应用解剖学、组织学、组织化学等方法，以及透射电镜技术对贝氏高原鳅消化系统的解剖结构、显微结构、亚显微结构和消化系统发育的显微结构做了详细的研究，结果表明：
口咽腔宽阔，顶壁和底壁均由粘膜层、粘膜下层和肌肉层三层结构组成，区别在于顶壁粘膜层杯状细胞与味蕾丰富，而底壁的棒状细胞含量较多。
食道的复层上皮主要由鳞状上皮细胞组成。鳞状细胞顶端有短的微绒毛，胞内除线粒体外，其他细胞器较少。鳞状细胞中间夹杂大量的粘液细胞。有发达的微管泡系统。基膜内有少量的网状纤维和纤维细胞及大量成束的胶原纤维。未见食道腺。仅一层环肌，为横纹肌。
胃为“V”形，其上皮为典型的单层柱状上皮。上皮细胞内含大量的分泌颗粒，PAS反应呈强阳性，线粒体发达，多位于细胞基部。粘膜层无杯状细胞。仅贲门和胃体部有发达的单管状腺体，胃腺细胞内含有大量的酶原颗粒、线粒体、糙面内质网及发达的微管泡系统。幽门部无任何腺体。在贲门与胃体处的肌层均为内纵肌外环肌，幽门部则为内环肌外纵肌。
肠绕胃呈“(?)”形，绒毛发达，其柱状上皮游离面有很多微绒毛。在上皮细胞之间分散有大量的粘液细胞，其内充满均质的分泌颗粒；在整个肠段，粘液细胞自前向后数量逐渐减少。粘膜层细胞间有发达的微管泡系统。
肛门为半开放结构，复层扁平上皮。粘膜下层为一层致密的结缔组织。由一层厚厚的横纹肌层支持。
肝脏外形不规则，外覆被膜，肝小叶不明显。肝细胞形态多样，偶见枯否氏细胞分布，血管极其发达，汇管区不明显。肝细胞胞质内有丰富的细胞器及内含物，以糙面内质网和线粒体最为常见。肝血窦由内皮细胞和枯否氏细胞组成。狄斯隙及毛细胆管内均可见肝细胞微绒毛伸入。
胰腺为独立结构，附于肝侧和肠侧，为很薄的条带。有外分泌部和内分泌部两部分，血管组织发达。
贝氏高原鳅仔稚鱼生长呈线性生长趋势，内源性营养阶段所需营养大部分来自勺形卵黄的头端。
口凹在出膜时就已经出现，3dph，与外界相通；9dph，口咽腔基本发育完成。8dph，食道发育完成。初孵仔鱼，消化道雏形已现，但胃肠未见明显分化；64dph，在胃小凹处有胃腺的雏形出现，胃开始具有全功能。肠细胞在出膜时就已经分化，27dph，肠基本发育完成。肝细胞2dph开始分化；7dph，肝脏中出现明显的中央静脉和肝细胞索，肝脏组织结构与成体差异不大。27dph开始肝细胞内含脂质泡明显累积。5dph，在食道和胃侧出现胰腺组织，胰腺腔间有大量嗜曙红酶原颗粒物质。9dph，胰岛出现。胰腺组织发育完成。消化系统组织学结构发育在64dph完成。
64dph，仔鱼消化道仅前端膨大，无任何弯曲。85dph，胃与食道呈直角弯曲后下行，但胃肠无明显分界。120dph，胃弯曲为“Z”形后笔直下行，胃肠仍无明显分界；肝脏为一整体，未见分叶。1龄幼鱼，消化系统解剖结构与成鱼相似，仅肝脏缺少右叶，而肠缺少胃底处的圆环形弯曲。
Anatomy,histological,histochemical and TEM methods were used to describe the structure and differentiation of the digestive system in Triplophysa bleekeri (Sauvage et Dabry) during December 2005 to December 2006. The results showed:
The digestive system of Triplophysa bleekeri (Sauvage et Dabry) was composed of digestive tract and digestive gland. The digestive tract was divided into five parts : oropharyngeal cavity, esophagus, stomach, intestine and anus. Digestive gland include liver,pancreas,and so on. The oropharyngeal cavity of Triplophysa bleekeri was wide, the top wall and the end wall had mucosa, submucosa and muscle layer three-layer structure. The difference was that mucous cells and taste buds were more abundance in the top wall and the end wall had more baculiform cells.
The histological structure in esophagus, stomach and intestinal were similar composed of mucosa, submucosa, muscle layer and adventitia.
The stratified epithelium of esophageal mainly from squamous cells.There was short microvilli on the top of squamous cell. There were not many organelles except for mitochondria. Squamous cell inclusions among a large number of mucous cells.There were developed tublovesicular system in esophagus. A small reticular fiber and fibrocyte and large bundles of collagen fibers in basement membrane.No esophageal glands.Only one layer striated muscle around esophagus.
The "V"-shaped stomach could be divided into three parts : cardiac, stomach and pylorus,those had typical simple columnar epithelium.The columnar epithelial cells contained a large number of secretory granules, that were strongly positive by the PAS reaction. Mitochondrial located in the lower part of cells, also advanced. Ther were no goblet cells in mucous layer. Advanced single tubular glands located in cardiac and body of stomach. The gastric glandular cells contained a large number of zymogen granule, mitochondria,rough surfaced endoplasmic reticulum and tublovesicular system. There was no gastric gland in pylorus. The inside muscular layer in gastric cardia and body of stomach was longitudinal muscle layer,and outer was circular muscle layer. The muscular layer of pylorus was in opposition to gastric cardia and body of stomach's.
The intestine circled around the stomach like "φ",was provided with developed villus. the columnar epithelium had lots of microvilli.A large number of mucous cells which scattered among the epithelial cells ,were filled with homogeneous secretory granules. The amount of mucous cells had been declining from the former to backward volume in the entire colon. Mucosa cells had developed tublovesicular system.
The structure of anus was semi-opened,which had stratified squamous epithelium.The dense connective tissue of submucosa layer was supported by thick striated muscle layer.
Liver was irregular in shape,which had capsule outside. The lobules of liver could not distinguish clearly.The hepatocyte varied in shape. Kupffer cells could be seen occasionally. Vascular was very prosperous. Portal area was not obvious. Liver cells cytoplasm was enriched with organelle and cell contents, in which rough surfaced endoplasmic reticulum and mitochondria most common. The liver sinusoid was composed of endothelial cells and Kupffer cells. The hepatocytes microvilli could be seen in Disse's space and capillary bile duct.
Pancreas was an independent thin strip, attached to the side of the liver and intestinal. Pancreas had exocrine and endocrine parts, where vascular tissue developed.
The growth of Triplophysa bleekeri larvae is linear growth trends.Nutrition mostly came from the first end of the spoon-shaped egg yolk in endogenous nutrition phase.
The mouth had already can be seen at the larvar hatching and linked to the outside world at 3dph. Oropharyngeal cavity completed basic development at 9dph. Esophagus completed development at 8dph. Newly hatched larvar, the digestive tract was a prototype, but no obvious differentiation gastrointestinal; At 64dph,the rudiment of gastric gland appeared in gastric pit,which showed the stomach started with the whole function.
Intestine completed the basic development at 27dph.Liver cells begin to differentiate at 2dph;At 7dph, central vein and hepatic cord appeared. There was no obviously distinguish between larvar liver and adult's. The liver cells started to accumulate lipid vesicle at 27dph. Pancreas appeared at the side of the esophagus and stomach at 5dph. Pancreatic cavity with a large number of eosinophilic granular material. At 9dph, the appeared of pancreatic islet completed the development of pancreatic tissue. Digestive histological structure development completed in 64dph.
Threr was no any bending only front enlargement in larval digestive tract at 64dph. At 85dph, stomach and esophagus was right angle after bending down, but no significant gastrointestinal boundaries. At 120dph, stomach bending to "Z" shape after straight down, no noticeable gastrointestinal boundaries; Liver of a whole, not divided. At one-year age, digestive anatomy was similar to adult's, except that liver lacking of the dexter liver and intestine lacking of a round ring bending.
口咽腔宽阔，顶壁和底壁均由粘膜层、粘膜下层和肌肉层三层结构组成，区别在于顶壁粘膜层杯状细胞与味蕾丰富，而底壁的棒状细胞含量较多。
食道的复层上皮主要由鳞状上皮细胞组成。鳞状细胞顶端有短的微绒毛，胞内除线粒体外，其他细胞器较少。鳞状细胞中间夹杂大量的粘液细胞。有发达的微管泡系统。基膜内有少量的网状纤维和纤维细胞及大量成束的胶原纤维。未见食道腺。仅一层环肌，为横纹肌。
胃为“V”形，其上皮为典型的单层柱状上皮。上皮细胞内含大量的分泌颗粒，PAS反应呈强阳性，线粒体发达，多位于细胞基部。粘膜层无杯状细胞。仅贲门和胃体部有发达的单管状腺体，胃腺细胞内含有大量的酶原颗粒、线粒体、糙面内质网及发达的微管泡系统。幽门部无任何腺体。在贲门与胃体处的肌层均为内纵肌外环肌，幽门部则为内环肌外纵肌。
肠绕胃呈“(?)”形，绒毛发达，其柱状上皮游离面有很多微绒毛。在上皮细胞之间分散有大量的粘液细胞，其内充满均质的分泌颗粒；在整个肠段，粘液细胞自前向后数量逐渐减少。粘膜层细胞间有发达的微管泡系统。
肛门为半开放结构，复层扁平上皮。粘膜下层为一层致密的结缔组织。由一层厚厚的横纹肌层支持。
肝脏外形不规则，外覆被膜，肝小叶不明显。肝细胞形态多样，偶见枯否氏细胞分布，血管极其发达，汇管区不明显。肝细胞胞质内有丰富的细胞器及内含物，以糙面内质网和线粒体最为常见。肝血窦由内皮细胞和枯否氏细胞组成。狄斯隙及毛细胆管内均可见肝细胞微绒毛伸入。
胰腺为独立结构，附于肝侧和肠侧，为很薄的条带。有外分泌部和内分泌部两部分，血管组织发达。
贝氏高原鳅仔稚鱼生长呈线性生长趋势，内源性营养阶段所需营养大部分来自勺形卵黄的头端。
口凹在出膜时就已经出现，3dph，与外界相通；9dph，口咽腔基本发育完成。8dph，食道发育完成。初孵仔鱼，消化道雏形已现，但胃肠未见明显分化；64dph，在胃小凹处有胃腺的雏形出现，胃开始具有全功能。肠细胞在出膜时就已经分化，27dph，肠基本发育完成。肝细胞2dph开始分化；7dph，肝脏中出现明显的中央静脉和肝细胞索，肝脏组织结构与成体差异不大。27dph开始肝细胞内含脂质泡明显累积。5dph，在食道和胃侧出现胰腺组织，胰腺腔间有大量嗜曙红酶原颗粒物质。9dph，胰岛出现。胰腺组织发育完成。消化系统组织学结构发育在64dph完成。
64dph，仔鱼消化道仅前端膨大，无任何弯曲。85dph，胃与食道呈直角弯曲后下行，但胃肠无明显分界。120dph，胃弯曲为“Z”形后笔直下行，胃肠仍无明显分界；肝脏为一整体，未见分叶。1龄幼鱼，消化系统解剖结构与成鱼相似，仅肝脏缺少右叶，而肠缺少胃底处的圆环形弯曲。
Anatomy,histological,histochemical and TEM methods were used to describe the structure and differentiation of the digestive system in Triplophysa bleekeri (Sauvage et Dabry) during December 2005 to December 2006. The results showed:
The digestive system of Triplophysa bleekeri (Sauvage et Dabry) was composed of digestive tract and digestive gland. The digestive tract was divided into five parts : oropharyngeal cavity, esophagus, stomach, intestine and anus. Digestive gland include liver,pancreas,and so on. The oropharyngeal cavity of Triplophysa bleekeri was wide, the top wall and the end wall had mucosa, submucosa and muscle layer three-layer structure. The difference was that mucous cells and taste buds were more abundance in the top wall and the end wall had more baculiform cells.
The histological structure in esophagus, stomach and intestinal were similar composed of mucosa, submucosa, muscle layer and adventitia.
The stratified epithelium of esophageal mainly from squamous cells.There was short microvilli on the top of squamous cell. There were not many organelles except for mitochondria. Squamous cell inclusions among a large number of mucous cells.There were developed tublovesicular system in esophagus. A small reticular fiber and fibrocyte and large bundles of collagen fibers in basement membrane.No esophageal glands.Only one layer striated muscle around esophagus.
The "V"-shaped stomach could be divided into three parts : cardiac, stomach and pylorus,those had typical simple columnar epithelium.The columnar epithelial cells contained a large number of secretory granules, that were strongly positive by the PAS reaction. Mitochondrial located in the lower part of cells, also advanced. Ther were no goblet cells in mucous layer. Advanced single tubular glands located in cardiac and body of stomach. The gastric glandular cells contained a large number of zymogen granule, mitochondria,rough surfaced endoplasmic reticulum and tublovesicular system. There was no gastric gland in pylorus. The inside muscular layer in gastric cardia and body of stomach was longitudinal muscle layer,and outer was circular muscle layer. The muscular layer of pylorus was in opposition to gastric cardia and body of stomach's.
The intestine circled around the stomach like "φ",was provided with developed villus. the columnar epithelium had lots of microvilli.A large number of mucous cells which scattered among the epithelial cells ,were filled with homogeneous secretory granules. The amount of mucous cells had been declining from the former to backward volume in the entire colon. Mucosa cells had developed tublovesicular system.
The structure of anus was semi-opened,which had stratified squamous epithelium.The dense connective tissue of submucosa layer was supported by thick striated muscle layer.
Liver was irregular in shape,which had capsule outside. The lobules of liver could not distinguish clearly.The hepatocyte varied in shape. Kupffer cells could be seen occasionally. Vascular was very prosperous. Portal area was not obvious. Liver cells cytoplasm was enriched with organelle and cell contents, in which rough surfaced endoplasmic reticulum and mitochondria most common. The liver sinusoid was composed of endothelial cells and Kupffer cells. The hepatocytes microvilli could be seen in Disse's space and capillary bile duct.
Pancreas was an independent thin strip, attached to the side of the liver and intestinal. Pancreas had exocrine and endocrine parts, where vascular tissue developed.
The growth of Triplophysa bleekeri larvae is linear growth trends.Nutrition mostly came from the first end of the spoon-shaped egg yolk in endogenous nutrition phase.
The mouth had already can be seen at the larvar hatching and linked to the outside world at 3dph. Oropharyngeal cavity completed basic development at 9dph. Esophagus completed development at 8dph. Newly hatched larvar, the digestive tract was a prototype, but no obvious differentiation gastrointestinal; At 64dph,the rudiment of gastric gland appeared in gastric pit,which showed the stomach started with the whole function.
Intestine completed the basic development at 27dph.Liver cells begin to differentiate at 2dph;At 7dph, central vein and hepatic cord appeared. There was no obviously distinguish between larvar liver and adult's. The liver cells started to accumulate lipid vesicle at 27dph. Pancreas appeared at the side of the esophagus and stomach at 5dph. Pancreatic cavity with a large number of eosinophilic granular material. At 9dph, the appeared of pancreatic islet completed the development of pancreatic tissue. Digestive histological structure development completed in 64dph.
Threr was no any bending only front enlargement in larval digestive tract at 64dph. At 85dph, stomach and esophagus was right angle after bending down, but no significant gastrointestinal boundaries. At 120dph, stomach bending to "Z" shape after straight down, no noticeable gastrointestinal boundaries; Liver of a whole, not divided. At one-year age, digestive anatomy was similar to adult's, except that liver lacking of the dexter liver and intestine lacking of a round ring bending.
引文
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