Młocicki Daniel
Instytucie Parazytologii im. W. Stefańiskiego PAN w Warszawie.
Wiad Parazytol. 2007;53(2):153-5.
In this study the cleavage divisions and the ultrastructural analysis of early embryos as well as cellular organisation of infective oncosphere of the anoplocephalid cestode Mosgovoyia ctenoides are described. The early cleavage is unequal and results in the formation of three types of blastomeres: 2 large macromeres containing large electron dense granules, 3 medium-size mesomeres and several small micromeres. In the early stage of oncospheral morphogenesis, formation of three following primary embryonic envelopes takes place: (1) the capsule replaced by thick, rigid outer coat originated form the uterine material secretion, (2) the outer envelope and (3) the inner envelope. The capsule is formed from the vitellocyte material. Two macromeres contribute to the formation of the outer envelope and three mesomeres take part in the formation of the inner envelope. The inner envelope undergoes differentiation into three sublayers: (1) a thick extraembryophoral cytoplasmic layer, (2) an electron-dense embryophore, as a stiff pyriform apparatus, and (3) a thin intraembryophoral cytoplasmic layer containing mesomere nuclei. The oncosphere is located in the extended cupule-like part of the pyriform apparatus. Four egg envelopes surround the mature infective oncosphere of M. ctenoides: (1) a thick outer coat, (2) the outer envelope, (3) the inner envelope with a characteristic pyriform apparatus and (4) the oncospheral membrane. Hook morphogenesis takes place inside six symmetrically arranged oncoblasts, each of which shows a characteristic large nucleus of semi-lunar shape. At the beginning the "hook-forming center" appears in the cytoplasmic part of each oncoblast. It consists of numerous free ribosomes, polyribosomes, mitochondria and Golgi complexes. The hook-forming center is involved in synthesis of a hook primordium, which undergoes differentiation and elongation into the fully developed hook. Mature hook consists of three parts: (1) blade, (2) shank, (3) base, and at the site of its protrusion from the oncosphere, is surrounded by a circular septate junction. Wide bands of hook muscles are attached to the basal and collar parts of the hook. The hook blades project outside the oncospheral body into a large cavity that is delimited by the hook region membrane. In the fully developed oncosphere of M. ctenoides three pairs of oncospheral hooks together with specialized hook muscles form a complex of "hook muscle system", responsible for coordinated hook action. The surface of the infective oncosphere is covered by a thin cytoplasmic layer of oncospheral tegument connected with the so-called "binucleate subtegumental cell", situated deeper in the oncospheral body. Below the cytoplasmic layer are situated wide bands of the somatic musculature responsible for oncospheral body movements. Five major types of oncospheral cells have been distinguished in the infective oncosphere: (1) a binucleate subtegumental cell, (2) a binucleate penetration gland, (3) two nerve cells, (4) numerous somatic cells, and (5) six germinative cells. During development of the oncosphere, changes in the concentration of glycogen and number of lipid droplets were observed. In the early embryos glycogen particles were most abundant in the macromere cytoplasm, whereas in micromeres concentration of glycogen was observed to be lower. In the course of the differentiation of the oncospheral envelopes glycogen was progressively distributed to other parts of the developing embryo. Simultaneously, a great increase in the number of lipid droplets was detected. However, during the preoncospheral phase of development a progressive reduction of lipid droplets was observed. This may indicate that lipids play a role of the energy source for developing oncosphere.
本研究描述了多头绦虫莫氏绦虫早期胚胎的卵裂分裂、超微结构分析以及感染性六钩蚴的细胞组织。早期卵裂不均等,导致形成三种类型的卵裂球:2个含有大电子致密颗粒的大型大卵裂球、3个中等大小的中卵裂球和几个小型小卵裂球。在六钩蚴形态发生的早期阶段,形成了以下三种初级胚胎包膜:(1)由子宫物质分泌产生的厚而硬的外层取代的囊膜,(2)外膜和(3)内膜。囊膜由卵黄细胞物质形成。两个大卵裂球参与外膜的形成,三个中卵裂球参与内膜的形成。内膜分化为三个亚层:(1)厚的胚外细胞质层,(2)作为坚硬梨形结构的电子致密胚体,(3)含有中卵裂球细胞核的薄的胚内细胞质层。六钩蚴位于梨形结构延伸的杯状部分。四个卵膜围绕着莫氏绦虫成熟的感染性六钩蚴:(1)厚的外层,(2)外膜,(3)带有特征性梨形结构的内膜和(4)六钩蚴膜。钩的形态发生发生在六个对称排列的成钩细胞内,每个成钩细胞都有一个特征性的半月形大细胞核。起初,“钩形成中心”出现在每个成钩细胞的细胞质部分。它由许多游离核糖体、多核糖体、线粒体和高尔基体组成。钩形成中心参与钩原基的合成,钩原基经过分化和伸长形成完全发育的钩。成熟的钩由三部分组成:(1)钩尖,(2)钩柄,(3)基部,在其从六钩蚴突出的部位,被一个环形分隔连接所包围。宽的钩肌带附着在钩的基部和颈部。钩尖突出到六钩蚴体外,进入由钩区膜界定的大腔中。在莫氏绦虫完全发育的六钩蚴中,三对六钩蚴钩与专门的钩肌形成一个“钩肌系统”复合体,负责协调钩的作用。感染性六钩蚴的表面覆盖着一层薄的六钩蚴皮层细胞质层,与位于六钩蚴体更深部位的所谓“双核皮层下细胞”相连。在细胞质层下方是负责六钩蚴体运动的宽的体壁肌肉带。在感染性六钩蚴中已区分出五种主要类型的六钩蚴细胞:(1)双核皮层下细胞,(2)双核穿刺腺,(3)两个神经细胞,(4)许多体细胞,和(5)六个生殖细胞。在六钩蚴发育过程中,观察到糖原浓度和脂滴数量的变化。在早期胚胎中,糖原颗粒在大卵裂球细胞质中最丰富,而在小卵裂球中糖原浓度较低。在六钩蚴包膜分化过程中,糖原逐渐分布到发育中胚胎的其他部位。同时,检测到脂滴数量大幅增加。然而,在六钩蚴发育前阶段,观察到脂滴逐渐减少。这可能表明脂质在发育中的六钩蚴中起能量来源的作用。