Fujimi TJ and Aruga J (2008), Upstream stimulatory factors, USF1 and USF2 are...

XB-ART-38037

Gene Expr Patterns 2008 Jul 01;86:376-381. doi: 10.1016/j.gep.2008.05.003.

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Upstream stimulatory factors, USF1 and USF2 are differentially expressed during Xenopus embryonic development.

Fujimi TJ , Aruga J .

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Upstream stimulatory factors (USF) 1 and 2 are members of the basic helix-loop-helix leucine zipper transcription factor family. They are considered to play critical roles in cell-cycle regulation and chromatin remodeling. Their gene expression patterns are considered ubiquitous but have not been fully investigated in terms of embryogenesis. We examined the expression of the genes encoding USF1 and USF2 in Xenopus laevis during embryonic development. Expression of both genes was first detected as maternal transcripts and was observed continuously throughout development. However, in situ hybridization analysis revealed that the two genes were expressed differentially. In the late blastula, both genes were expressed in the blastocoel roof and marginal zone. At the gastrula stage, USF2 was strongly expressed in the sensorial layer of the ectoderm and in the mesoderm, whereas USF1 expression was hardly detectable. From the neurula stage onward, expression of both genes was markedly enhanced in the neural tissues, neural crest, eye and otic vesicle. However, spatial expression of the genes within the neural tube differed in that the strongest USF1 signals were observed in the lateral region of the basal plate and the strongest USF2 ones in the dorsal region of the neural tube. Expression of the two genes occurred in different mesoderm derivatives at the tailbud stage (USF1, somite; USF2, pronephros and lateral plate mesoderm of the tail region). USF1 was expressed in the notochord of the early neurula, but was lost at the stage.

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Species referenced: Xenopus laevis
Genes referenced: foxa2 lhx1 lhx3 msx1 pax2 tbx2 uqcc6 usf1 usf2

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	Fig. 3. Expression profiles of Xenopus laevis USF genes during embryonic development. WMISH analysis with digoxigenin-labeled probes was performed to determine the spatial expression patterns of USF genes during development. Stained embryos are shown at six stages as labeled in the figure. In the images of late blastula- (stages 9–9.5) and gastrula-stage (stages 10.5–11) embryos, arrowheads indicate the dorsal lip positions (blastula, initial dorsal lip pigment; gastrula, fine dorsal lip). A higher-magnification view of the gray rectangular region in panel O is shown in panel O′ [black dashed line: mesoderm (meso), white dashed line: border of epithelial layer (e.l.) and sensorial layer (s.l.); bc: blastocoel]. White dashed lines in panels E and P indicate the positions of the cuts made to give the cross-sections in panels F and Q. Panels F and Q are WMISH images of cut embryos at stages 14–15 (nc, notochord; e.l., epithelial layer of neuroectoderm; s.l., sensorial layer of neuroectoderm; s.m., somitic mesoderm; en, endoderm). The borders of the three germ layers are indicated at the left side of the embryo as dashed lines. Different views of stage 19 and 25 embryos are shown (H, J, S and U dorsal; G, I, R and T, lateral). Panels K and V are lateral views of late tailbud-stage embryos (stage 33). The dashed line with arrowheads indicates the somite stained with the USF1 probe (panel K). Arrowhead (white) indicates the lateral plate mesoderm stained with the USF2 probe (panel V). Small lines (a–d) in panels K and V point to where the transverse sections of the stage 33 WMISH sample (Fig. 5) were taken. Panels K′ and V′ are higher-magnification views of the gray boxes in panels K and V (br, brain; ov, otic vesicle; ey, eye; ba, branchial arches). Inset in V is a highly contrasted view of the area indicated by dashed box in panel V (pronephros).
	Fig. 5. Expression of Xenopus laevis USF genes in the CNS at the tailbud stage. Sections of embryos stained by WMISH are shown. A and B are images from embryos stained by USF probes (A, USF1; B, USF2). The positions of sections a–d in the embryos are indicated in Fig. 3K and V. Higher-magnification views of the neural tube in panels a–d are shown in panels e–h. C–G are the corresponding sections [from the embryos stained by WMISH with CNS specification markers from dorsal to ventral: C, Msx1 as marker of roof plate ([Liem et al., 1995] and [Schlosser and Ahrens, 2004]); D, Lim1 as alar plate (Taira et al., 1994); E, Pax2 as lateral part of CNS ([Mansouri et al., 1996] and [Heller and Brandli, 1999]); F, Lim3 as basal plate (Taira et al., 1993); G, FoxA2 as floor plate (Lee et al., 1997)]. Arrowheads indicate the pronephros. br, brain; nc, notochord; ov, otic vesicle; ey, eye; ba, branchial arches.
	usf1 (upstream transcription factor 1 [provisional]) gene expression in Xenopus laevis embryo via in situ hybridization, NF stage 9, lateral view, dorsal up.
	usf1 (upstream transcription factor 1 [provisional]) gene expression in Xenopus laevis embryo via in situ hybridization, NF stage 15, dorsal view, anterior up.
	usf1 (upstream transcription factor 1 [provisional]) gene expression in Xenopus laevis embryo via in situ hybridization, NF stage 19, dorsal view, anterior up.
	usf1 (upstream transcription factor 1 [provisional]) gene expression in Xenopus laevis embryo via in situ hybridization, NF stage 25, lateral view, anterior left, dorsal up.
	usf1 (upstream transcription factor 1 [provisional]) gene expression in Xenopus laevis embryo via in situ hybridization, NF stage 33, lateral view, anterior left, dorsal up.
	usf2 (upstream transcription factor 2 [provisional]) gene expression in Xenopus laevis embryo via in situ hybridization, NF stage 9, vegetal view, dorsal up.
	usf2 (upstream transcription factor 2 [provisional]) gene expression in Xenopus laevis embryo via in situ hybridization, NF stage10.5, lateral view, dorsal up, anterior left.
	usf2 (upstream transcription factor 2 [provisional]) gene expression in Xenopus laevis sectioned gastrula embryo, via in situ hybridization, NF stage 11, mid-saggital section dorsal up.
	usf2 (upstream transcription factor 2 [provisional]) gene expression in Xenopus laevis embryo via in situ hybridization, NF stage 15, dorsal view, anterior up.
	usf2 (upstream transcription factor 2 [provisional]) gene expression in Xenopus laevis embryo via in situ hybridization, NF stage 25, lateral view, dorsal up, anterior left.
	usf2 (upstream transcription factor 2 [provisional]) gene expression in Xenopus laevis embryo via in situ hybridization, NF stage 33,head region, lateral view, dorsal up, anterior left.