Higher order structure in metaphase chromosomes. I. The 250 A fiber

Metaphase chromosomes released from cells in the presence of Joklik's suspension media by vortex-mixing with 0.5 mm glass beads have been analyzed by electron microscopy. In these preparations the chromosomes are composed of series of loops (200-300 A in diameter) which are, in turn, composed of closely-apposed arrays of nucleosomes. Negative-staining of these preparations has allowed the identification of several distinct patterns within the loop which appear to arise from variations in nucleosome packing. Analogous patterns are also observed in chromatin fragments generated by brief micrococcal nuclease digestion. From these data we have deduced certain features of nucleosome-nucleosome interactions in higher-ordered chromatin fibers.