A study of haplotypes of the Y-chromosome, published in 2000, addressed the paternal origins of Ashkenazi Jews. Hammer et al. found that the Y-chromosome of Ashkenazi and Sephardic Jews contained mutations that are also common among Middle Eastern peoples, but uncommon in the general European population. This suggested that the male ancestors of the Ashkenazi Jews could be traced mostly to the Middle East. The proportion of male genetic admixture in Ashkenazi Jews amounts to less than 0.5% per generation over an estimated 80 generations, with "relatively minor contribution of European Y chromosomes to the Ashkenazim," and a total admixture estimate "very similar to Motulsky's average estimate of 12.5%." This supported the finding that "Diaspora Jews from Europe, Northwest Africa, and the Near East resemble each other more closely than they resemble their non-Jewish neighbors." "Past research found that 50–80 percent of DNA from the Ashkenazi Y chromosome, which is used to trace the male lineage, originated in the Near East," Richards said.