The avian yolk sac is a multifunctional extraembryonic organ that serves not only as a site of nutrient (yolk) absorption, but also for early hemopoiesis, and formation of blood vessels. Although the yolk sac membrane being specialized to function as an extraembryonic absorptive organ, it is neither morphologically nor functionally part of the embryonic gut. Yolk absorption is by the phagocytic activity of the extraembryonic endoderm. I used cryohistology and resin embedding histology of complete developmental series of Japanese quail to document the development of the avian yolk sac and changes of the microscopic anatomy throughout development. This material is complemented by complete series of MRT-scans of live ostrich embryos from beginning of incubation through hatching. Considerable changes of size and shape of the yolk mass are documented and discussed as resulting from water flux from albumen to yolk associated with the biochemical activation of yolk sac proteins.During embryogenesis, the yolk sac endoderm forms villi that increase the absorptive surface and reach into the yolk ball. The histology of the absorptive epithelium is specialized for phagocytic absorption of yolk. During early developmental stages, the extraembryonic endoderm is single layered, but it eventually becomes several layers thick during later stages. The extraembryonic mesoderm forms an extensive layer of hematopoietic tissue; deep in this tissue lie the yolk sac vessels. During late stages of development, the erythropoietic tissue disappears, blood vessels are obliterated, and the yolk sac epithelium becomes apoptotic. Results are discussed in the light of the evolutionary history and phylogeny of the amniote egg.
| INTRODUCTIONThe avian egg is an almost paradigmatic model of amniote eggs. However, diverging evolutionary pathways have led to different functional, developmental, and morphological features of the eggs in the diverse groups of amniotes (e.g. Packard & Packard, 1980;Elinson, Stewart, Bonneau, & Blackburn, 2015; Starck, Stewart & Blackburn, this issue of JMOR). However, knowledge of the amniote egg is scattered through the published literature, incomplete for morphology and clades providing the necessary phylogenetic coverage, and rarely comprehensive and integrative in an evolutionary context. While the eggshell and the chorioallantois have been covered to some degree by morphological studies, the analysis of the morphology of the yolk mass and the cellular yolk sac has been neglected. Therefore, this paper focuses on the morphology and histology of the avian yolk sac so that comparisons with other amniote eggs may be made and placed in a phylogenetic and evolutionary framework.The avian egg is a cleidoic egg; that is, a protective layer of eggshell membranes, and a calcified, but semi-conductive eggshell (only water vapor and respiratory gases diffuse across shell membranes and shell) enclose proportionally large volumes of yolk and albumen. Yolk, albumen, and eggshell are the main constituents of the egg. The ovary