Some trends in the evolution of very large chromosomes

Abstract

The general features of the arrangement and cytological distribution of repeated sequences in animal chromosomes are reviewed. These features include internal repetitiveness, conservation of clearly functional sequences, rapid divergence of certain classes of repeated sequence with subsequent fixation of families of diverged sequences, and well defined cytological localization of large blocks of sequences in specific parts of the chromosome set. Moderately or ‘middle’ repetitive (m.r.) sequences constitute a large part of the genomes of higher organisms, they seem to accumulate in a balanced manner within chromosome sets, they are mainly responsible for genome growth, and they are interspersed with sequences of other kinds. Little is known about their cytological distribution. Four experiments are described, each of which aimed to locate middle repetitive sequences in the chromosomes of a salamander and a newt. Tritiated m.r. DNA from Plethodon cinereus binds in a non-random fashion to the meiotic diplotene and mitotic chromosomes of the same species, suggesting a non-random distribution of m.r. sequences on these chromosomes. The same DNA, hybridized in situ to the RNA transcripts on the loops of lampbrush chromosomes, produces light and widespread labelling of many loops, but intense labelling of six pairs of loops, each of which lies near to a centromere. Similar experiments in which newt m.r. DNA was hybridized in situ to newt lampbrush chromosomes showed heavy labelling of about 30 loop pairs on each of the long heteromorphic arms of chromosome I, but very little labelling elsewhere. Autoradiographs of newt mitotic chromosomes hybridized with newt m.r. DNA showed rather even labelling of all chromosomes including chromosome I. The significance of the heavy labelling of lampbrush loops near centromeres and on the heteromorphic arms of the newt chromosome I is discussed in relation to possible cytological and molecular mechanisms for generating and preserving families of tandemly linked and cytologically localized m.r. sequences.