Alibardi Lorenzo
Comparative Histolab Padova, Italy; Department of Biology, University of Bologna, Bologna, Italy.
Dev Biol. 2023 Sep;501:60-73. doi: 10.1016/j.ydbio.2023.05.007. Epub 2023 May 25.
General cellular aspects of skin development in vertebrates are presented with emphasis on the epidermis of sauropsids. Anamniote skin develops into a multilayered mucogenic and soft keratinized epidermis made of Intermediate Filament Keratins (IFKs) that is reinforced in most fish and few anurans by dermal bony and fibrous scales. In amniotes, the developing epidermis in contact with the amniotic fluid initially transits through a mucogenic phase recalling that of their anamniotes progenitors. A new gene cluster termed EDC (Epidermal Differentiation Complex) evolved in amniotes contributing to the origin of the stratum corneum. The EDC contains numerous genes coding for over 100 types of corneous proteins (CPs). In sauropsids 2-8 layers of embryonic epidermis accumulate soft keratins (IFKs) but do not form a compact corneous layer. The embryonic epidermis of reptiles and birds produces small amount of other, poorly known proteins in addition to IFKs and mucins. In the following development, a resistant corneous layer is formed underneath the embryonic epidermis that is shed before hatching. The definitive corneous epidermis of sauropsids is mainly composed of CBPs (Corneous beta proteins, formerly indicated as beta-keratins) derived from the EDC. CBPs belong to a gene sub-family of CPs unique for sauropsids, contain an inner amino acid region formed by beta-sheets, are rich in cysteine and glycine, and make most of the protein composition of scales, claws, beaks and feathers. In mammalian epidermis CPs missing the beta-sheet region are instead produced, and include loricrin, involucrin, filaggrin and various cornulins. Small amount of CPs accumulate in the 2-3 layers of mammalian embryonic epidermis and their appendages, that is replaced with the definitive corneous layers before birth. Differently from sauropsids, mammals utilize KAPs (keratin associated proteins) rich in cysteine and glycine for making the hard corneous material of hairs, claws, hooves, horns, and occasionally also scales.
本文介绍了脊椎动物皮肤发育的一般细胞特征,重点是蜥形纲动物的表皮。无羊膜动物的皮肤发育成多层产黏液且轻度角质化的表皮,由中间丝角蛋白(IFK)构成,大多数鱼类和少数无尾两栖类动物的皮肤通过真皮骨鳞和纤维鳞得到强化。在羊膜动物中,与羊水接触的发育中的表皮最初会经历一个产黏液阶段,这让人想起它们无羊膜动物祖先的阶段。一个名为表皮分化复合体(EDC)的新基因簇在羊膜动物中进化出来,促成了角质层的起源。EDC包含众多编码100多种角质蛋白(CP)的基因。在蜥形纲动物中,胚胎表皮的2 - 8层积累了软角蛋白(IFK),但没有形成致密的角质层。爬行动物和鸟类的胚胎表皮除了IFK和黏蛋白外,还会产生少量其他鲜为人知的蛋白质。在随后的发育过程中,在胚胎表皮下方形成一层抗性角质层,在孵化前脱落。蜥形纲动物的定型角质表皮主要由源自EDC的角质β蛋白(CBP,以前称为β - 角蛋白)组成。CBP属于蜥形纲动物特有的CP基因亚家族,包含一个由β折叠形成的内部氨基酸区域,富含半胱氨酸和甘氨酸,构成了鳞片、爪子、喙和羽毛的大部分蛋白质组成。而在哺乳动物表皮中,产生的是缺少β折叠区域的CP,包括兜甲蛋白、内披蛋白、丝聚蛋白和各种兜甲蛋白相关蛋白。少量CP在哺乳动物胚胎表皮及其附属器的2 - 3层中积累,在出生前被定型角质层取代。与蜥形纲动物不同,哺乳动物利用富含半胱氨酸和甘氨酸的角蛋白相关蛋白(KAP)来制造毛发、爪子、蹄、角,偶尔也用于制造鳞片的硬角质材料。