Avian sex chromosomes in the lampbrush form: the ZW lampbrush bivalents from six species of bird.

The ZW bivalent has been identified and characterized in detail in its lampbrush form in oocytes of chicken, quail, turkey, pigeon, chaffinch and sparrow. The sex bivalent in all six species looks like a single highly asymmetrical chromosome. Most of it has the typical lampbrush organization. The terminal one-fifth is relatively thick and condensed and bears only a few pairs of lateral loops: this condensed terminal region is the W chromosome; the part with normal lampbrush morphology is the Z. The two are connected by a single near terminal chiasma. The fine scale morphology and arrangement of loops and markers on Z and W chromosomes are described for each species and lampbrush maps have been constructed. The identification of the lampbrush sex bivalent is based on the following criteria. The asymmetrical chromosome has two centromere regions. In the interstitial region of the asymmetrical chromosome where the junction between Z and W chromosomes is supposed to be, there are telomere-specific loops and telomeric DNA sequences and there is good morphological evidence for the presence of a chiasma. There are W chromosome specific DNA sequences in the region of the asymmetrical lampbrush chromosome that is thought to represent the W. Breed-specific variations in the morphology of the chicken W chromosome with respect to the sizes, numbers and arrangements of axial chromomeres and distributions of specified repeated DNA sequence families have been identified, offering one of the first examples of definitive correlation between a repeat family and a single chromomere. The lampbrush chromosomes of all the birds examined, except quail, terminate in distinctive free hanging loops. These are a novel feature in the sense that at the end of each chromatid there is a large transcription unit terminating in a cluster of telomeric DNA sequences.