Thorogood P
J Embryol Exp Morphol. 1979 Dec;54:185-207.
In the avian embryo ectomesenchyme cells, derived from the mesencephalic level of the cranial neural crest, migrate into the presumptive maxillary region and subsequently differentiate into the membrane bones and associated secondary cartilage of the upper jaw skeleton. The cartilage arises secondarily within the periosteum at points of articulation between membrane bones and provides an embryonic articulating surface. The stimulus for the differentiation of secondary cartilage is believed to be intermittent pressure and shear created at the developing embryonic movement. The development of one such system--the quadratojugal, has been analysed using organ and explant culture techniques and studied with particular reference to the differentiation of periosteal cells into secondary cartilage. A number of conclusions were reached. (1) Normally only cells at discrete loci express a chondrogenic potential in vivo: the periosteal cells at these sites of future articulation become committed to chondrogenesis during stage 35, more than 24 h before cartilage is identifiable in vivo. (2) However, cells with a 'latent' chondrogenic potential are widespread in membrane bone periosteum and occur over most, if not all, of the surface area of the bone. This potential is expressed in the 'permissive' environment created by submersion of the tissue in explant culture or in submerged organ culture. (3) This chondrogenic potential exists long before the time at which commitment of cartilage-forming cells occurs and even presumptive maxillary ectomesenchyme at stage 29 has a limited ability to form cartilage in vitro. It is suggested that spatial position is a principal factor controlling the differentiation of secondary cartilage. Ectomesenchyme cells with the potential to form secondary cartilage are widespread but it is only those cells whose migration from the neural crest positions them and their progeny at the site of a presumptive joint which subsequently express this potential. This epigenetic interpretation is discussed in the general context of development mechanisms underlying the spatial and temporal patterns in which neural crest-derived cells differentiate to produce bone and cartilage during the formation of the head skeleton.
在鸟类胚胎外胚间充质细胞中,这些细胞源自颅神经嵴的中脑水平,迁移至假定的上颌区域,随后分化为上颌骨骼的膜性骨和相关的次级软骨。软骨在膜性骨之间的关节点处的骨膜内继发形成,并提供一个胚胎关节表面。次级软骨分化的刺激因素被认为是发育中的胚胎运动产生的间歇性压力和剪切力。利用器官和外植体培养技术分析了一个这样的系统——方轭骨,并特别参考骨膜细胞向次级软骨的分化进行了研究。得出了一些结论。(1)正常情况下,体内只有离散位点的细胞表现出软骨形成潜能:在第35阶段,这些未来关节部位的骨膜细胞开始致力于软骨形成,这比体内可识别软骨的时间早24小时以上。(2)然而,具有“潜在”软骨形成潜能的细胞广泛存在于膜性骨骨膜中,并且在骨的大部分(如果不是全部)表面积上都有。这种潜能在组织浸没于外植体培养或浸没器官培养所创造的“允许”环境中得以表达。(3)这种软骨形成潜能在软骨形成细胞开始定向分化之前就已存在很长时间,甚至第29阶段的假定上颌外胚间充质在体外形成软骨的能力也有限。有人提出空间位置是控制次级软骨分化的主要因素。具有形成次级软骨潜能的外胚间充质细胞广泛存在,但只有那些从神经嵴迁移而来并将它们及其后代定位在假定关节部位的细胞,随后才会表达这种潜能。在神经嵴衍生细胞在头部骨骼形成过程中分化产生骨和软骨的时空模式所依据的发育机制的总体背景下,讨论了这种表观遗传学解释。