Abstract. Interphase chromosome organization in four different Drosophila melanogaster tissues, covering three to four levels of polyteny, has been analyzed. The results are based primarily on three-dimensional reconstructions from unfixed tissues using a computerbased data collection and modeling system. A characteristic organization of chromosomes in each cell type is observed, independent of polyteny, with some packing motifs common to several or all tissues and others tissue-specific. All chromosomes display a righthanded coiling chirality, despite large differences in size and degree of coiling. Conversely, in each cell type, the heterochromatic centromeric regions have a unique structure, tendency to associate, and intranuclear location. The organization of condensed nucleolar chromatin is also tissue-specific. The tightly coiled prothoracic gland chromosomes are arrayed in a similar fashion to the much larger salivary gland chromosomes described previously, having polarized orientations, nonintertwined spatial domains, and close packing of the arms of each autosome, whereas hindgut and especially the unusually straight midgut chromosomes display striking departures from these regularities. Surprisingly, gut chromosomes often appear to be broken in the centric heterochromatin. Severe deformations of midgut nuclei observed during gut contractions in living larvae may account for their unusual properties. Finally, morphometric measurements of chromosome and nuclear dimensions provide insights into chromosome growth and substructure and also suggest an unexpected parallel with diploid chromatin organization.
IN a polytene interphase nucleus, chromosomes undergo repeated rounds of DNA replication without nuclear division. The homologous chromatids remain in close lateral alignment, rendering chromosomes with distinctive banding patterns visible in the light microscope. These nuclei provide a useful model system for directly analyzing interphase chromosome organization. Previously, we determined the spatial organization of chromosomes in a set of nuclei from Drosophila melanogaster salivary glands, which have the highest level of polyteny in the larva (22, 34). Characteristic packing motifs were found, including a polarized (Rabl) orientation of chromosomes across the nucleus, separation of chromosomes into noninterwound spatial domains, nearest-neighbor packing of the metacentric autosome arms, high-frequency contacts between a specific set of loci and the nuclear surface, invariant apposition of the aggregated centromeric regions (the chromocenter) to the nuclear surface, and a striking right-handed coiling chirality that does not require homologue pairing. Despite these nonrandom features, precise chromosome configurations appear not to be specified. Thus, while the folding of chromosomes is not identical in every salivary gland cell, several other facets of their organization are quite well defined.Dr. Hochstrasser's present address is