Translin and its intimate binding partner TRAX are highly conserved proteins. Translin has been implicated in numerous biological functions ranging from cancer-associated chromosomal rearrangements in some types of leukaemia and other cancers to the control of mRNA transport in the brain. Translin is essential for TRAX stability whilst its own stability is unaffected by loss of TRAX. Two recent finding now give a more detailed insight into Translin and TRAX functions. First, these two proteins combine to form the C3PO complex, which functions in the removal of the passenger strand in RNAi mediated gene silencing. Second, Translin and TRAX are implicated in tRNA processing. However, in S. pombe,deletion of the two genes encoding these proteins causes no measurable phenotype change in mitotically proliferating cells.In this study, our aim was to investigate the biological role of TraX, in particular, in the regulation of the RNAi machinery used by cells to maintain the heterochromatin state of key functional genomic regions. We used different approaches to examine whether TraX (and Translin) play redundant roles with other RNAi regulators. Some intriguing observations led us to propose a novel Dicer-independent, Translin-dependent pathway for maintaining chromosomal stability. Additionally, whilst TraX does not seem to be essential for this novel pathway, its loss suppresses the genome instability phenotype of Argonaute-deficient cells and this suppression is Translin-dependent. This finding suggests that TraX functions to prevent Translin mediation of chromosome stability in the absence of Argonaute. Remarkably, microarray analysis to test this proposal indicates that TraX might link Ago1 (Argonaute) function to chromosome stability through the regulation of Rif1 levels or by the control of some replicative or recombinogenic process in which the Tlh2 RecQ-like helicase can serve as a substitute. This could indicate a major new role for Ago1, TraX and Translin in genome maintenance. Moreover, the functional relationship between Translin and TraX is cast into doubt as clear evidence emerges from this study for distinct roles for each of thesefission yeast genes.