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Tenomodulin (Tnmd) is a type II transmembrane glycoprotein predominantly expressed in tendons and ligaments. We found that scleraxis (Scx), a member of the Twist-family of basic helix-loop-helix transcription factors, is a transcriptional activator of  Tnmd  expression in tenocytes. During embryonic development,  Scx  expression preceded that of  Tnmd .  Tnmd  expression was nearly absent in tendons and ligaments of  Scx -deficient mice generated by transcription activator-like effector nucleases-mediated gene disruption.  Tnmd  mRNA levels were dramatically decreased during serial passages of rat tenocytes.  Scx  silencing by small interfering RNA significantly suppressed endogenous  Tnmd  mRNA levels in tenocytes. Mouse  Tnmd  contains five E-box sites in the ~1-kb 5′-flanking region. A 174-base pair genomic fragment containing a TATA box drives transcription in tenocytes. Enhancer activity was increased in the upstream region (−1030 to −295) of  Tnmd  in tenocytes, but not in NIH3T3 and C3H10T1/2 cells. Preferential binding of both Scx and Twist1 as a heterodimer with E12 or E47 to CAGATG or CATCTG and transactivation of the 5′-flanking region were confirmed by electrophoresis mobility shift and dual luciferase assays, respectively. Scx directly transactivates  Tnmd  via these E-boxes to positively regulate tenocyte differentiation and maturation.

Scleraxis is a transcriptional activator that regulates the expression of Tenomodulin, a marker of mature tenocytes and ligamentocytes