双峰驼皱胃淋巴集结区的结构和功能及其与年龄之间的相关性研究
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摘要
双峰驼皱胃淋巴集结区是王雯慧于2003年在我国阿拉善双峰驼的胃中发现并报道的特殊结构,为骆驼所特有(在其它动物和人未见有类似报道)。为了进一步探讨该结构的免疫形态学特征和功能及其与年龄之间的关系,本文首先以正常双峰驼为研究对象,运用解剖学、组织学、组织化学和电子显微镜技术等研究方法,将不同年龄的阿拉善双峰驼分为驼羔(0.5～2岁)、青年驼(3～5岁)、壮龄驼(6～16岁)和老龄驼(17～20岁)4个组,分别对其皱胃淋巴集结区的解剖学位置、黏膜的解剖学特点、微观结构特点、超微结构特点,及这些结构特点与年龄之间的关系进行了详细的观察、测定与研究。在此基础之上,本文又以枯草芽孢杆菌作为口服抗原示踪物,建立了双峰驼动物模型,通过各项相关指标的检测,对双峰驼皱胃淋巴集结区的功能和免疫机制进行了探讨。此外,本文还从发育形态学的角度研究了双峰驼淋巴集结区在胚胎发育过程中的形态结构变化及其发育时间。结果显示:(1)双峰驼皱胃淋巴集结区不仅具有特殊的解剖学位置和黏膜结构,而且其发达程度与年龄之间存在密切的关系。该结构呈长三角带状,位于胃颈之后,沿胃小弯壁分布;其黏膜褶较其周围区域宽厚,与周围非淋巴集结区形成明显界限;根据其黏膜褶表现形式的不同,可分为网格黏膜褶区和纵行黏膜褶区;对各年龄组皱胃淋巴集结区的面积、重量,及黏膜皱褶的高度和数量的测量和统计结果显示,皱胃淋巴集结区在双峰驼青春期之前是逐渐增大的,到青春期达到其一生中最发达时期,之后逐渐减小、萎缩,但20岁时仍然存在,没有完全消失。(2)双峰驼皱胃淋巴集结区既具有与其他肠道相关淋巴组织相似之处,又具有其自身的一些特点,并随年龄而有所改变。驼羔和青年驼皱胃淋巴集结区的黏膜褶高而宽厚,淋巴小结几乎都沿黏膜褶轴心两侧均匀分布,多为圆形、椭圆形或楔形,呈单层连续排列于黏膜下层,黏膜褶厚的地方淋巴小结可呈2～3层排列;黏膜褶内滤泡相关上皮丰富;生发中心明显:淋巴小结的数量及黏膜褶中网状纤维和浆细胞的数量都在随年龄逐渐增多,到青春期达到其一生中最发达时期。而壮龄驼和老年驼皱胃淋巴集结区的黏膜褶变低变薄,淋巴小结仍分布于黏膜褶轴心两侧,但不均匀,常常是一侧多,而另一侧稀少,甚至缺如:淋巴小结多为椭圆形或不规则形,呈不连续排列,且老龄驼的淋巴组织多以弥散形式存在;淋巴小结生发中心仍然明显;其数量及黏膜褶中网状纤维和浆细胞的数量都在逐渐减少。这些结果与大体解剖学研究结果一致,它们从微观水平进一步证实并解释了双峰驼皱胃淋巴集结区的结构特征随年龄变化的规律及免疫学意义。(3)超微结构研究显示,与皱胃非淋巴集结区相比,淋巴集结区具有其特殊的超微结构特征,不同年龄组之间也存在差异。青年驼皱胃淋巴集结区贲门腺的开口较小,其中的腺体细胞较多,黏膜表面的抗原摄取部位-滤泡相关的上皮数量较多,其表面的黏液也较多。而老龄驼皱胃淋巴集结区贲门腺的开口较大,其中的腺体细胞很少,黏膜表面滤泡相关上皮的数量也较少。因此,随着年龄的增长,双峰驼皱胃淋巴集结区的机械屏障和黏膜免疫屏障均发生了不同程度的萎缩和退化。(4)动物模型的观察结果显示,实验组皱胃淋巴集结区的黏膜表面,尤其是滤泡相关上皮表面有很多枯草芽孢杆菌黏附,并有少量细菌穿过黏膜表面进入黏膜固有层,而没有滤泡相关上皮的黏膜表面细菌数量相对较少,但比非淋巴集结区多,对照组中没有发现枯草芽孢杆菌。此外,实验组皱胃淋巴集结区的上皮内淋巴细胞、浆细胞、肥大细胞和亲银细胞等免疫相关细胞的数量均发生了显著增殖;超微研究显示,皱胃淋巴集结区黏膜表面的黏液量增加,上皮细胞发生增殖,M细胞的数量增加,黏膜的表面结构从原来的“蜂窝”状变成了表面有许多小孔的多孔状结构。(5)双峰驼皱胃淋巴集结区的发育形态学研究结果显示,双峰驼皱胃淋巴集结区的初级淋巴小结在妊娠中期开始发育,并在妊娠最后1～2个月内发育成熟;此外,皱胃淋巴集结区内淋巴小结的最初发育部位在黏膜固有层,之后部分淋巴小结穿过黏膜肌层到达黏膜下层。
综上所述,双峰驼皱胃淋巴集结区不仅具有特殊的消化结构,还具有特殊的免疫形态学结构,其随年龄的增长而逐渐萎缩的现象属于一种生理性萎缩现象,是机体免疫衰老和黏膜免疫衰老的一个有力证据。当有外源性抗原侵入时,双峰驼皱胃淋巴集结区会产生一系列与防御相关的免疫细胞学和黏膜屏障结构的变化;其抗原摄取部位主要位于滤泡相关上皮处,其黏膜免疫机制和肠道派伊尔结相似。另外,双峰驼皱胃淋巴集结区在胚胎时期就已经发育成熟,并在青春期之后逐渐萎缩、退化,具有与牛、羊等动物回肠派伊尔结类似的发育特点,可能属于一种初级免疫器官。
The aggregated lymphoid nodule area(ALNA) in the third compartment of the stomach of Chinese Alashan Bactrian camels is a recently(2003) described species-specific anatomical structure by Wang Wen-hui.To further investigate the immunological morphological characteristics and function of it,and establish the relationship between this structure and animal age,normal Bactrian camels were firstly studied using the technologies of anatomy, histology,histochemistry and electron microscopy while the age groups were defined as follows:young camels(0.5-2 years old),pubertal camels(3-5 years old),middle-aged camels (6-16 years old) and old camels(17-20 years old).The exact anatomical location,and the anatomical,microstuctural and ultrastructural characteristics of the mucosa of the ALNA were described in detail as was the relationship between this structure andanimal age.On the above basis,Bactrian camel models were imitated using Bacillus subtilis as an oral tracer antigen,then the function and immune mechanisms of the ALNA were discussed after detecting relevant indicators.In addition,the changes of morphological characteristics of the ALNA and its development time during the ontogenic courses were also studied.The results showed that:(1) The ALNA of the Bactrian camels not only has a particular anatomical location and distinct mucosal structure,but also changes with age.The ALNA extended along the ventral wall of the isthmus,from the origin of the proximal enlargement and along the curvatura ventriculi minor.The mucosal folds of the ALNA were much thicker than and were clearly demarcated from those of adjacent non-ALNA.Based on the morphology of the mucosal folds,the ALNA could be divided into reticular and longitudinal regions.On statistical analysis,the ALNA was found to enlarge prior to puberty,peak in size at puberty, and then regress.A small remnant ALNA was found in 20-year-old camels,although the germinal centres of the contained lymphoid nodules were not as obvious at this time.(2) In addition to some similarities to gut-associated lymphoid tissues,the ALNA also has its special histological characteristics which may change with ageing.The mucosal folds on the ALAN of the young and pubertal camels are higher and wider than those of older animals,and most lymphoid nodules are round,oval,and cuneiform shaped aggregating in the submucosa continuously although some nodules are arranged in two or more rows,especially at the top of wide mucosal folds.All the lymphoid nodules are with distinct germinal centers,and are distributed evenly on both sides of the axis of the mucosal folds.The number of the follicular associated epithelium(FAE) and lymphoid nodules in the mucosal folds are very abundant as well as the number of reticular fibers and plasma cells.On the contrary,the mucosal folds on the ALNA of the middle-aged and old camels become lower and thinner than those of the young animals.Most lymphoid tissues of this age are diffuse lymphoid tissues while lymphoid nodules are oval and irregular shapes,and are arranged interruptedly on both sides of the mucosal fold axis unevenly,which are generally more on one side,fewer on the other side,or even absent.Furthermore,the germinal centers are still obvious while the number of the lymphoid nodules in the mucosal folds as well as the number of reticular fibers and plasma cells decrease gradually.These results are consistent with the anatomical findings,and can further confirm and explain the regularity and immunological significance of structural characteristics changes with ageing of the ALNA.(3) The ultrastructural studies suggested that the ALAN has its unique ultrastructural characteristics compared to the non-ALNA of stomach,and the ultrastructural characteristics are different between age groups.The cardiac glands in the ALNA of young Bactrian camels have more gland cells with small opening diameter,and the numbers of antigen uptake sites -FAE on the mucosal surface are very abundant as is the mucus.Whereas the cardiac glands in the ALNA of the old camels have wider opening and fewer gland ceils while the quantity of FAE and mucus on the mucosal surface also becomes fewer.Therefore,both the mechanical barrier and mucosal immune barrier of the ALNA regress with increasing age in Bactrian camels.(4) Observations on animal models showed that:while many Bacillus subtilis adhere to the mucosal surface of the FAE,there are a fewer on the non-FAE and non-ALNA in the experimental group,and there is none anywhere of the control group.In addition,there are a few Bacillus subtilisis penetrating the epithelium into the laminae propria of the FAE in the experimental group. Moreover,the numbers of immune-related cells,such as intraepithelial lymphocytes,plasma cells,mast cells and argyrophilic cells have all significant proliferated in the experimental group.Ultrastructural studies indicated that after feeding bacteria,the mucus and M cells of the ALNA increase whlie the epithelial cells proliferate prominently,which leads to the surface structure of the ALNA changed from "honeycomb" into porous structure.(5) The results of the ontogenic morphology of the ALNA showed that the first primordial lymphoid nodules of the ALNA was encountered in the midterm of gestation whereas the lymphoid nodules development was observed to have been completed in the last one or two months of pregnancy.In addition,the initial development location of lymphoid nodules location was in the lamina propria,and partial lymphoid nodules penetrated the lamina muscularis into the submucosa.
In conclusion,the ALNA of the Bactrian camels not only has special digestive structure, but also has special immune morphological structure.The changes of ALNA with ageing were related to the physiological atrophy,and is a useful evidence of immunesenescence and mucosal immunesenescence.A series of the immune celluar reactions and mucosal barrier changes of ALNA take place to defend the invasion of exogenous antigen,and the mucosal immune mechanism of the ALNA is similar to PP in the intestine while the FAE is the main antigen uptake sites.In addition,similar to the development characteristics of the ileal Peyer's patches that reported in bovine and sheep,the development of the ALNA has been completed in the gestation,and it regress after puberty,so it may be a primary immune organ.
综上所述,双峰驼皱胃淋巴集结区不仅具有特殊的消化结构,还具有特殊的免疫形态学结构,其随年龄的增长而逐渐萎缩的现象属于一种生理性萎缩现象,是机体免疫衰老和黏膜免疫衰老的一个有力证据。当有外源性抗原侵入时,双峰驼皱胃淋巴集结区会产生一系列与防御相关的免疫细胞学和黏膜屏障结构的变化;其抗原摄取部位主要位于滤泡相关上皮处,其黏膜免疫机制和肠道派伊尔结相似。另外,双峰驼皱胃淋巴集结区在胚胎时期就已经发育成熟,并在青春期之后逐渐萎缩、退化,具有与牛、羊等动物回肠派伊尔结类似的发育特点,可能属于一种初级免疫器官。
The aggregated lymphoid nodule area(ALNA) in the third compartment of the stomach of Chinese Alashan Bactrian camels is a recently(2003) described species-specific anatomical structure by Wang Wen-hui.To further investigate the immunological morphological characteristics and function of it,and establish the relationship between this structure and animal age,normal Bactrian camels were firstly studied using the technologies of anatomy, histology,histochemistry and electron microscopy while the age groups were defined as follows:young camels(0.5-2 years old),pubertal camels(3-5 years old),middle-aged camels (6-16 years old) and old camels(17-20 years old).The exact anatomical location,and the anatomical,microstuctural and ultrastructural characteristics of the mucosa of the ALNA were described in detail as was the relationship between this structure andanimal age.On the above basis,Bactrian camel models were imitated using Bacillus subtilis as an oral tracer antigen,then the function and immune mechanisms of the ALNA were discussed after detecting relevant indicators.In addition,the changes of morphological characteristics of the ALNA and its development time during the ontogenic courses were also studied.The results showed that:(1) The ALNA of the Bactrian camels not only has a particular anatomical location and distinct mucosal structure,but also changes with age.The ALNA extended along the ventral wall of the isthmus,from the origin of the proximal enlargement and along the curvatura ventriculi minor.The mucosal folds of the ALNA were much thicker than and were clearly demarcated from those of adjacent non-ALNA.Based on the morphology of the mucosal folds,the ALNA could be divided into reticular and longitudinal regions.On statistical analysis,the ALNA was found to enlarge prior to puberty,peak in size at puberty, and then regress.A small remnant ALNA was found in 20-year-old camels,although the germinal centres of the contained lymphoid nodules were not as obvious at this time.(2) In addition to some similarities to gut-associated lymphoid tissues,the ALNA also has its special histological characteristics which may change with ageing.The mucosal folds on the ALAN of the young and pubertal camels are higher and wider than those of older animals,and most lymphoid nodules are round,oval,and cuneiform shaped aggregating in the submucosa continuously although some nodules are arranged in two or more rows,especially at the top of wide mucosal folds.All the lymphoid nodules are with distinct germinal centers,and are distributed evenly on both sides of the axis of the mucosal folds.The number of the follicular associated epithelium(FAE) and lymphoid nodules in the mucosal folds are very abundant as well as the number of reticular fibers and plasma cells.On the contrary,the mucosal folds on the ALNA of the middle-aged and old camels become lower and thinner than those of the young animals.Most lymphoid tissues of this age are diffuse lymphoid tissues while lymphoid nodules are oval and irregular shapes,and are arranged interruptedly on both sides of the mucosal fold axis unevenly,which are generally more on one side,fewer on the other side,or even absent.Furthermore,the germinal centers are still obvious while the number of the lymphoid nodules in the mucosal folds as well as the number of reticular fibers and plasma cells decrease gradually.These results are consistent with the anatomical findings,and can further confirm and explain the regularity and immunological significance of structural characteristics changes with ageing of the ALNA.(3) The ultrastructural studies suggested that the ALAN has its unique ultrastructural characteristics compared to the non-ALNA of stomach,and the ultrastructural characteristics are different between age groups.The cardiac glands in the ALNA of young Bactrian camels have more gland cells with small opening diameter,and the numbers of antigen uptake sites -FAE on the mucosal surface are very abundant as is the mucus.Whereas the cardiac glands in the ALNA of the old camels have wider opening and fewer gland ceils while the quantity of FAE and mucus on the mucosal surface also becomes fewer.Therefore,both the mechanical barrier and mucosal immune barrier of the ALNA regress with increasing age in Bactrian camels.(4) Observations on animal models showed that:while many Bacillus subtilis adhere to the mucosal surface of the FAE,there are a fewer on the non-FAE and non-ALNA in the experimental group,and there is none anywhere of the control group.In addition,there are a few Bacillus subtilisis penetrating the epithelium into the laminae propria of the FAE in the experimental group. Moreover,the numbers of immune-related cells,such as intraepithelial lymphocytes,plasma cells,mast cells and argyrophilic cells have all significant proliferated in the experimental group.Ultrastructural studies indicated that after feeding bacteria,the mucus and M cells of the ALNA increase whlie the epithelial cells proliferate prominently,which leads to the surface structure of the ALNA changed from "honeycomb" into porous structure.(5) The results of the ontogenic morphology of the ALNA showed that the first primordial lymphoid nodules of the ALNA was encountered in the midterm of gestation whereas the lymphoid nodules development was observed to have been completed in the last one or two months of pregnancy.In addition,the initial development location of lymphoid nodules location was in the lamina propria,and partial lymphoid nodules penetrated the lamina muscularis into the submucosa.
In conclusion,the ALNA of the Bactrian camels not only has special digestive structure, but also has special immune morphological structure.The changes of ALNA with ageing were related to the physiological atrophy,and is a useful evidence of immunesenescence and mucosal immunesenescence.A series of the immune celluar reactions and mucosal barrier changes of ALNA take place to defend the invasion of exogenous antigen,and the mucosal immune mechanism of the ALNA is similar to PP in the intestine while the FAE is the main antigen uptake sites.In addition,similar to the development characteristics of the ileal Peyer's patches that reported in bovine and sheep,the development of the ALNA has been completed in the gestation,and it regress after puberty,so it may be a primary immune organ.
引文
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