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Fig. 4. Genetic interaction of Pbx1 and Hox11 during spleen organogenesis. (A-D) Immunostaining of sagittal sections of E11.5 (A,B) and E 12.5 (C,D) Hox11^lacZ/+ embryonic upper abdominal organs. Arrowheads in A-D indicate cells expressing Pbx1 but not Hox11. Pbx1 is visualized in the splenic anlage with DAB (brown staining), and Hox11 by ß-galactosidase staining (blue). (B,D) Enlargements of the black rectangle depicted on the splenic anlage in A,C. Arrows in B,D indicate cells in which Pbx1 and Hox11 colocalize. (E,F) The spectrum of malformations of double heterozygous spleens (F) is compared with the normal gross morphology of Pbx1^+/- spleens (E). The Pbx1^+/-;Hox11^+/- double heterozygous spleens are hypoplastic (F), and display sickle shapes, indentations, tubercles and nodules, as well as fusions of two spleens (arrowheads). All spleens were isolated from 6- to 8-week-old mice. (G,H) Hematoxylin and Eosin-stained spleen sections reveal no abnormalities in splenic structure of Pbx1^+/-;Hox11^+/- double heterozygous mice. Distribution of white (arrows) and red (arrowheads) pulp is normal within the spleen parenchyma of double heterozygous mice (H) compared with Pbx1^+/- (not shown), Hox11^+/- (not shown) and wild-type (G) littermates. (I,J) PNA-stained spleen sections of mice immunized with sheep red blood cells (SRBC). Formation of germinal centers (GC; arrows), as indicated by PNA staining (brown), appears normal in Pbx1^+/-;Hox11^+/- double heterozygous mice (J) compared with Pbx1^+/- (not shown), Hox11^+/- (not shown) and wild-type (I) littermate controls. L, liver; Sp, spleen; St, stomach.

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