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In 2014 Animal Genetics sequenced the TYRP1 gene of several brown French bulldogs but was not able to find any functional variants that corresponded to brown coat color, eliminating the TYRP1 gene as the primary cause of French Bulldog chocolate. Recently, research conducted in University of Bern in Switzerland revealed a nonsense variant in HPS3, c.2420G>A or p.(Trp807*). In this study, the brown or chocolate dog was homozygous for the mutant allele.