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Our goal is to understand how chromosomes are faithfully transmitted during cell division and how chromosomes are structured and organized in the nucleus. Many cellular processes act on the chromosome to specialize different chromosomal domains for unique functions in the biology of cells. One of the best examples of a specialized chromatin domain is the eukaryotic centromere and kinetochore that forms at a single site on each human chromosome and ensures the proper segregation of the genome during each cell division cycle. The laboratory studies the genetic and epigenetic mechanisms that control the formation of human centromeres and kinetochores. Cells use a variety of different mechanisms to change the function of chromosomes. We have also focused on understanding how RNAs that associate with chromosomes regulate the reorganization of chromatin into silent heterochromatic domains. Our recent efforts have been directed at understanding how long range interactions between chromosomes are used to organize the genome within the nucleus and to control gene expression and chromosome dynamics. We use a combination of digital microscopy to extract quantitative information about the dynamics of chromosomes in living cells, biochemical reconstitution to assemble chromatin and chromosomes in vitro, and genetics to manipulate the chromosome segregation process in order to study how chromosome-distribution systems function in eukaryotes.