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The EMBO Journal (2003) 22, 2741–2751, doi:10.1093/emboj/cdg271 Figure 3 The magnitude of hedgehog signaling activity defines skin tumor phenotype Vladimir Grachtchouk, Marina Grachtchouk, Lori Lowe, Tim Johnson, Lebing Wei, Aiqin Wang, Fred de Sauvage and Andrzej A. Dlugosz

Figure 3 Histology, marker expression and increased proliferation rate of basaloid follicular hamartomas in K5-M2SMO mice. H&E-stained skin sections from control (A, C and E) and K5-M2SMO (B, D and F) mice at 14 weeks (A–D) and 23 days (E and F) of age. Basaloid follicular hamartomas were detected throughout the integument, including dorsal (A and B) and tail (C–F) skin. In some areas, a cleft separated abnormal epithelium from the surrounding mesenchyme (inset in D), mimicking changes seen in human BCCs. Hair follicles in age-matched control skin (A and C) are in the resting phase of the hair cycle, telogen, with compact clusters of dermal papilla cells (dp) at the tips of the follicles. epi, epidermis; derm, dermis; hf, hair follicle; sq, subcutaneous fat; dp, dermal papilla; seb, sebaceous gland. Analysis of tail skin from young mice (23 days) revealed multiple epithelial downgrowths originating directly from K5-M2SMO epidermis (arrowheads in F, compared with control in E exhibiting a single hair follicle). Note the similar morphology of mouse lesions and human basaloid follicular hamartoma in (G). Marker analysis and BrdU labeling in control (H, J, L and N) and K5-M2SMO transgenic (I, K, M and O) tail skin. Note the widespread expression of K17 in hamartomas (I); expansion of K5 expression (K), which is more evident in immunofluorescence studies (not shown); and focal expression of K10 (M). Scattered pigmented cells frequently are present in the dermis of K5-M2SMO mice (inset in M). Immunostaining for BrdU revealed focally increased proliferation in transgenic mouse skin (O) compared with control (N). (P and Q) Normal growing (anagen) hair follicles. H&E-stained section (P) reveals dermal papilla cells with abundant extracellular material enveloped by hair matrix cells. K17 is expressed in the outer root sheath (Q), but not matrix cells or inner root sheath. ors, outer root sheath; irs, inner root sheath.

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