Experimental studies of the origin and expression of metameric pattern in the chick embryo

Abstract

On either side of Hensen's node of the fully extended primitive streak of the chick embryo (stage 4) the mesoderm is already organized into circular domains called somitomeres. As Hensen's node regresses, paraxial somitomeres are added in tandem and are early morphological representatives of metameric pattern in the mesoderm. These organized circular domains of mesenchyme cells are best visualized with stereo pair scanning electron microscopy. Experiments suggested that a prepattern of segmentation exists in and around the fully extended primitive streak. Streaks divested of Hensen's node can generate some paraxial somites, but only if surgically split down the midline. We assessed metameric pattern formation in nodeless streaks, both severed and unsevered down the midline. Operated blastoderms were cultured 15 hours, fixed, dissected, processed for scanning electron microscopy, and photographed in stereo. Split nodeless streaks produced a cranial to caudal sequence of somitomeric development. This sequence is similar to the sequential maturational events seen in the segmental plate of older embryos. The least mature somitomeres, toward the posterior end of the severed edge, appear as circular domains of radially oriented cells, looking much like the first somitomeres to emerge near Hensen's node of the stage 4 streak. More cranially along the severed edge, somitomeres are morphologically more mature, being more condensed, with cells oriented about a central myocoele. At the most cranial end of the severed piece, somitomeres are the most mature, having contracted about their centers to create intersomitomeric gaps that permit their identification with light microscopy as individual “somites.” Embryos from which the node was removed, but the streak left intact, generated only the most primitive somitomeric pattern repetitively along either side of the primitive groove. We conclude that regression of Hensen's node provides for the timely initiation of morphogenesis of somitomeres from a prepattern of segmentation that already exists.