Disorders of sex development (DSDs) are congenital conditions resulting
in atypical chromosomal, gonadal or anatomic sex. Around 80% of 46,XY DSD cases
still remain unexplained genetically. Phenotypes include gonadal dysgenesis,
male-to-female sex reversal and hypospadias, where the urethra is abnormally
positioned along the penis. Mutations in SOX9,
a gene essential for testicular development, have been implicated in DSDs, but
its tissue-specific regulation is not fully understood. SOX9 tissue-specific
enhancer mutations can cause isolated clinical phenotypes. Through genome wide
mapping, mutations around SOX9 have
been identified in isolated DSD patients. All patients had gonadal defects and
hypospadias. We hypothesised that a 78 kb region encompassing these mutations
contains a novel gonadal enhancer that we aimed to identify. Comparative
genomic analysis identified seven highly conserved sub-regions within TES2.
Unexpectedly, transgenic mouse analysis revealed that the most highly conserved
sub-region (SR4) was expressed in the genital tubercle (GT; penis primordium)
instead of the gonad. Immunofluorescence analysis on mouse GT sections
confirmed a novel domain of SOX9 expression within the urethral plate
epithelium. These data suggest that loss of SOX9 expression due to SR4
mutations may have contributed to the
hypospadias phenotype. To investigate whether SOX9 plays a direct role in
hypospadias, we conditionally ablated Sox9
in the GT using mice transgenic for Shh-Cre. Immunofluorescence analysis confirmed loss of SOX9 expression
in the urethra of Shh-Cre/+;Sox9flox/flox mice. Abnormal tail and limb development were observed, consistent
with the role of Sox9 in bone
development. However, preliminary analysis at stages before masculinisation
revealed no abnormalities in the GT. GT development in these mice is currently
being analysed at later stages. This study may reveal a novel cause of
hypospadias. In conclusion, we have identified putative SOX9 enhancers that may be required for human testicular and GT
development, the disruption of which cause DSDs.