双峰驼下丘脑—垂体—肾上腺轴(HPAA)的功能形态
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摘要
为了给研究双峰驼(Camelus bactrianus)应激反应能力提供理论基础,本文对其下丘脑-垂体-肾上腺轴(HPAA)各器官进行了形态学测量、CT图像分析、组织结构和超微结构的观察,并与其它偶蹄动物的同类器官进行了比较,结果表明:
     1.双峰驼的下丘脑重约2.4 g,体积约为2.0 cm~3,与大脑的重量之比约为0.42%,体积比约为0.45%。下丘脑的大部分神经元兼具有腺上皮细胞的结构特征,具有分泌功能,电镜下观察可见其含有神经分泌颗粒。双峰驼的EQ值为1.11,具有中等智力水平,下丘脑发达程度也属中等,这说明双峰驼在接到外界信号刺激后,能较快速的把神经信号转变成激素信号,并通过下丘脑-垂体系影响垂体。
     2.双峰驼的垂体重约1.54 g,体积约为5.0 cm~3,体重比约为0.00446 g/kg,和同体形的偶蹄动物很接近。双峰驼垂体由发达的远侧部、中间部和不发达的神经部组成。垂体远侧部的细胞主要有7种,分别是生长激素细胞、催乳素细胞、促甲状腺激素细胞、促肾上腺皮质激素细胞、促性腺激素细胞、嫌色细胞和星形细胞,还有一些间质细胞,这与大多数动物垂体远侧部的细胞类型及特点相同。其中比较有争议的是星形细胞,从形态结构上,完全可以把星形细胞从嫌色细胞里面区分出来。发达的中间部可能与双峰驼具有很强的耐渴性有关;神经部不很发达,可能与双峰驼的生产力较低有关。双峰驼神经部中间区域的神经纤维束间有大量的赫令体,而在其周边位置,赫令体的数量则比较稀少,这与大多数动物的赫令体分布不同。
     3.双峰驼单个肾上腺重量约为12.449±2.111 g,占体重比约为0.0341±0.0039 g/kg,体积约为10.78±1.42 cm~3,在偶蹄动物中属于中等。双峰驼肾上腺的髓质体积在肾上腺总体积中所占的比例比大多数的动物大;双峰驼肾上腺有发达的被膜和较发达的球状带,在肾上腺和肾脏之间还存在一个由神经纤维束、血管和结缔组织组成的混合束,其功能可能是将球状带分泌的盐皮质激素不用经过体循环直接输送入肾脏,以调节肾脏的泌尿功能,这对生活在干旱、半干旱生态区的双峰驼有着重要的意义;雌性双峰驼肾上腺有明显的中间带,而雄性双峰驼却不明显;双峰驼的肾上腺皮质和髓质内经常可见在小动脉由被膜延伸向髓质时,周围围绕着圆形或椭圆形的皮质环,最后一般消失在髓质;双峰驼肾上腺皮质细胞内的脂滴形状不规则,一般呈边缘不整的圆形,部分是含有很低电子密度的脂滴,其中以束状带细胞内的脂滴分布最为丰富,球状带和网状带细胞内只有少量的脂滴;线粒体一般呈圆形或是椭圆形,嵴呈管泡状;肾上腺细胞胞质内一般有丰富的滑面内质网,粗面内质网分布很少;肾上腺皮质内还可见到少量的微体分布。肾上腺髓质细胞最明显的特征是含有大量膜包的电子致密颗粒,据此可区分去甲肾上腺素细胞和肾上腺素细胞。
     双峰驼的下丘脑、垂体和肾上腺的基本形态结构与其它偶蹄动物的相似,但也有一些特殊或独特的结构。双峰驼的下丘脑、垂体和肾上腺的发达程度中等,组织结构发育系统完善。这说明双峰驼能较快地应对各种刺激,并迅速做出反应,这对生活在特殊环境下的双峰驼有着重要的意义。
The ability of Bactrian camel to respond to the stress of environment was studied by exploring the properties of hypothalamus-pituitary-adrenal axis(HPAA) with morphological measuring,computed tomographical analyzing,microscope and ultrastructure observation,and comparative anatomy with other artiodactyls.The results show as follows:
     1.The hypothalamus of Bactrian camel is about 2.4 g in weight,0.42%of the brain and 2 cm~3 in volume,0.45%of the brain.The hypothalamus consist of nerve and neuroglia cells.Most of neurons own the structural properties of epithelioglandular cells.The neurosecretory granules were found in electron microscope,so these neurons owns secretive abilities.The emotional quotient of Bactrian camel is 1.11.It owns moderate intelligence and hypothalamus development status.We concluded that the nerve signal will translate to hormonal signal and release to the hypophysis through the hypothalamus-pituitary linkage when the Bactrian camel received the stimulation from outside.
     2.The pituitary gland of the Bactrian camel is about 1.54 g in weight,5 cm~3 in volume and the weight ratio is 0.00446 g/kg to body,which is quite close to adjacent body size of artiodactyls.It comprises with the developed pars distalis,pars intermedia which may relate to the resistance of the water depletion,and the undeveloped pars nervosa,by which the low productivity may relevant.Pars distalis of pituitary gland mainly consists of seven types of cells,growth hormone,prolactin, thyroid-stimulating hormone,ACTH,gonadotropic,chromophobe and stellate cells. These cells are the same as the pars distali of most animals' pituitary gland in morphology.The stellate cells are controversial,which can be separated from chromophobe cells by focusing on the morphological structure.Moreover,the Herring's bodies are observed in the nerve fiber bundles of intermedial pars nervosa. But in the peripheral zone,there are few Herring bodys,which is different from most of other animals.
     3.A single adrenal gland of Bactrian camel is 12.449±2.111 g in weight,10.78±1.42 cm~3 in volumn and the weight ratio is 0.0341±0.0039 g/kg to body,while the ratio is moderate in artiodactyls.The volume ratio of medulla to adrenal gland is higher than most of other animals.The adrenal gland of Bactrian camel owns not only developed capsule and comparatively developed zona glomerulosa,but a combined beam which composed by nerve fiber bundles,blood vessels and some connective tissues between the adrenal gland and kidney.So,the mineralocorticoid secreted by zona glomerulosa may enter into kidney directly without through systemic circulation to regulate the urinary function,which is significant to the adaptability of Bactrian camel to its arid and semi-arid condition.The intermediate zone can be seen evidently in female adrenal gland,but which is not the case in male.The round or elliptical cortical cuff which surrounded the small artery usually disappeared ultimately in medulla during the process of small artery extending from capsule to medulla.The lipid droplet shapes within adrenocortical cell is irregular and presents round shape with incomplete edge and low electron density.The fascicular zone has rich lipid droplets.Few lipid droplets present in glomerulosa zone and zona reticularis. Mitochondria are round or oval-shaped,in which the cristae shapes is bubbly pipe like. The adrenal gland cytoplasm presents rich smooth endoplasmic reticulum and the rough endoplasmic reticulum is hardly to be seen.Few microbodies were found within the adrenal cortex.The most obvious property of adrenal medulla cells are that they contain a huge number of electron-dense granules enveloped by the membrane, and can be further divided into noradrenaline cells and adrenaline cells.
     The morphologies of hypothalamus,pituitary and adrenal in Bactrian camel are similar with other artiodactyls,but it owns special or unique structures.The development status of the hypothalamus,pituitary and adrenal in Bactrian camel is moderate with consummate and systematic histological structure.It shows that the Bactrian camels are able to quickly respond to any stimulation,which has significant meaning for Bactrian camel to adapt to arid and semi-arid circumstances.
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