Lynch J et al. (2011), Analysis of the expression of retinoic acid met...

XB-ART-42247

Gene Expr Patterns 2011 Jan 01;111-2:112-7. doi: 10.1016/j.gep.2010.10.003.

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Analysis of the expression of retinoic acid metabolising genes during Xenopus laevis organogenesis.

Lynch J , McEwan J , Beck CW .

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Retinoic acid (RA) is a known teratogen that is also required endogenously for normal development of the embryo. RA can act as a morphogen, through direct binding to receptors and RA response elements in the genome, and classical studies of limb development and regeneration in amphibians have shown that it is likely to provide positional information. Availability of RA depends on both metabolic synthesis and catabolic degradation, and specific binding proteins act to further modulate the binding of RA to response elements. Here, we describe the expression of seven genes involved in metabolism (Raldh1-3), catabolism (Cyp26a and b) and binding of RA (Crabp1 and 2) during organogenesis in the clawed frog Xenopus laevis. Taken together, this data indicates regions of the embryo that could be affected by RA mediated patterning, and identifies some differences with other vertebrates.

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Species referenced: Xenopus laevis
Genes referenced: aldh1a1 aldh1a2 aldh1a3 crabp1 crabp2 cyp26a1 cyp26b1

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	Fig. 1. Expression of Raldh1-3 during organogenesis. In situ hybridisation of Raldh1-3, dark blue stain indicates expression. Panels A–C, H–J and O–Q are whole mount lateral views with head facing left and dorsal uppermost. Panels C’, J’ and Q’ show detail of head area. Panels D–G, K–N and R–U are paraffin sections of previously stained embryos. The blue/purple stain indicates expression and sections are counterstained with eosin (pink). (A) Raldh1 expression at stage 28. (B) Raldh1 expression at stage 30. (C) Raldh1 expression in stage 34, red arrows indicate relative positions of transverse sections in panels D to G. (C’) Detail of head of embryo shown in panel C. (D–G) Transverse sections of stage 34 embryo showing expression of Raldh1 in blue/purple. (H) Raldh2 expression at stage 28. (I) Raldh2 expression at stage 30. (J) Raldh2 expression at stage 32, red arrows indicate relative positions of transverse sections in panels K–M. (J’) Detail of head of embryo shown in panel J. (K–M) Transverse sections through stage 34 embryo showing expression of Raldh2 in dark blue. (N) Sagittal section through eye of a Raldh2 stained embryo at stage 34. (O) Raldh3 expression in stage 26 embryo. (P) Raldh3 expression in stage 30 embryo. (Q) Raldh3 expression in stage 34 embryo, red arrows indicate relative positions of transverse sections in panels R–T. (Q’) Detail of head of embryo shown in panel Q. (R–T) Transverse sections of stage 34 embryo showing expression of Raldh3 in dark blue. (U) Sagittal section of Raldh3 st. 34 embryo oriented as for whole mounts. Abbreviations: di, diencephalon; cf, choroid fissure; dr, dorsal retina; e, endoderm; ey, eye; hb, hindbrain; hy, hypophysis; le, lens; ll, lateral line precursors; lpm, lateral plate mesoderm; m/h, midbrain hindbrain boundary; no, notochord; nt, neural tube; op, olfactory placode; ov, otic vesicle; pk, pronephritic kidney; pd, pronephritic duct; re, retina; rp, roof plate; s, somite; te, telencephalon. Scale bars are 100 μM.
	Fig. 2. Expression of Cyp26a and b during organogenesis. In situ hybridisation of Cyp26a and b, dark blue stain indicates expression. Panels A–F are Cyp26a and G–L are Cyp26b. Panels A–C and G–I are whole mount whole mount lateral views with head facing left and dorsal uppermost. (A) Expression of Cyp26a at stage 28. (B) Expression of Cyp26a at stage 30. (C) Expression of Cyp26a at stage 34. (B’) Close-up view of intersomitic staining in embryo shown in panel B. (D–F) 12 μm sagittal sections through previously stained stage 34 embryo showing Cyp26a expression in dark blue and with eosin (pink) used as a counterstain. Heads are to the left and dorsal uppermost. (D) Section through eye. (E) Deeper section (more towards midline) through head area. (F) Section through trunk somites. (G) Cyp26b expression in stage 28 embryo. (H) Cyp26b expression in stage 30 embryo. (I) Cyp26b expression in stage 34 embryo. Red arrows indicate the level of sections used in panels K and L. (H’) Closer detail of the rhombomere staining in panel H. (J) Sagittal section through stage 34 embryo showing expression in numbered rhombomeres of the hindbrain. Anterior is to the left and dorsal uppermost (J). (K) Transverse section at the level of the otic vesicles showing hindbrain staining localised towards the lumenal side. (L) Transverse section through ventral embryo at the level of the heart showing staining in ventral aortae. In K and L, dorsal is uppermost. All scale bars are100 μm. Abbreviations: ba, branchial arches; cf, choroid fissure; e, endoderm; he, heart; isv, intersomitic vessels; le, lens; lpm, lateral plate mesoderm; ov, otic vesicle; r, rhombomere; re, retina; s, somite; tb, tailbud; va, ventral aortae.
	Fig. 3. Expression of Crabp2 during organogenesis. In situ hybridisation of Crabp2, dark blue stain indicates expression. (A) whole mount embryos with anterior to the left and dorsal uppermost. (A) Expression at stage 28. (B) Expression at stage 30. (C) Expression at stage 34. Red arrows indicate positions of transverse sections in panels D. (C Detail of head region of embryo pictured in panel C. (D) Transverse sections through stage 34 embryo. Scale bars are 100 lm. (D) Staining in head mesenchyme ventral to olfactory placodes, at the level of the telencephalon. (E) Staining in branchial arches and dorsal retina at eye level. (F) Staining in the hindbrain is strongest at the lumenal surfaces. (G) Sagittal section though stage 34 embryo showing staining in branchial arches, head mesenchyme, midbrain/hindbrain boundary region and posterior hindbrain. Scale bar is 100 lm. Abbreviations: ba, branchial arches; dr, dorsal retina; hb, hindbrain; hm, head mesenchyme; le, lens; m/h, midbrain hindbrain boundary; no, notochord; op, olfactory placode; ov, otic vesicle; pd, pronephritic duct; s, somite; tb, tailbud; te, telencephalon.
	aldh1a1 (aldehyde dehydrogenase 1 family member A1 ) gene expression in Xenopus laevis embryos, NF stage 34, as assayed by in situ hybridization, lateral view, anterior left, dorsal up. (key: op= olfactory placode; cf= choroid fissure; pk= pronephric kidney; pd= pronephric duct; no= posterior notochord)
	crabp2 (cellular retinoic acid binding protein 2) gene expression in Xenopus laevis embryos, NF stage 30, as assayed by in situ hybridization, lateral view: anterior left, dorsal up.
	cyp26a1 (cytochrome P450, family 26, subfamily A, polypeptide 1 ) gene expression in Xenopus laevis embryos, NF stage 34, as assayed by in situ hybridization, lateral view, anterior left, dorsal up.
	cyp26b1 ( cytochrome P450, family 26, subfamily B, polypeptide 1 ) gene expression in Xenopus laevis embryos, NF stage 34, as assayed by in situ hybridization, lateral view, anterior left, dorsal up.
	crabp2 () gene expression in X. during organogenesis. In situ hybridisation of crabp2, dark blue stain indicates expression. Stage 30.
	Expression of Raldh1-3 during organogenesis. In situ hybridization of Raldh1-3, dark blue stain indicates expression. Stage 34. di, diencephalon; cf, choroid fissure; dr, dorsal retina; e, endoderm; ey, eye; hb, hindbrain; hy, hypophysis; le, lens; ll, lateral line precursors; lpm, lateral plate mesoderm; m/h, midbrain hindbrain boundary; no, notochord; nt, neural tube; op, olfactory placode; ov, otic vesicle; pk, pronephritic kidney; pd, pronephritic duct; re, retina; rp, roof plate; s, somite; te, telencephalon. Scale bars are 100 Î¼M.
	Expression of Raldh3 during organogenesis. In situ hybridization of Raldh3, dark blue stain indicates expression. Stage 34. di, diencephalon; cf, choroid fissure; dr, dorsal retina; e, endoderm; ey, eye; hb, hindbrain; hy, hypophysis; le, lens; ll, lateral line precursors; lpm, lateral plate mesoderm; m/h, midbrain hindbrain boundary; no, notochord; nt, neural tube; op, olfactory placode; ov, otic vesicle; pk, pronephritic kidney; pd, pronephritic duct; re, retina; rp, roof plate; s, somite; te, telencephalon. Scale bars are 100 Î¼M.
	Expression of Raldh3 during organogenesis. In situ hybridization of Raldh3, dark blue stain indicates expression. Stage 34. di, diencephalon; cf, choroid fissure; dr, dorsal retina; e, endoderm; ey, eye; hb, hindbrain; hy, hypophysis; le, lens; ll, lateral line precursors; lpm, lateral plate mesoderm; m/h, midbrain hindbrain boundary; no, notochord; nt, neural tube; op, olfactory placode; ov, otic vesicle; pk, pronephritic kidney; pd, pronephritic duct; re, retina; rp, roof plate; s, somite; te, telencephalon. Scale bars are 100 Î¼M.
	Expression of Raldh1-3 during organogenesis. In situ hybridization of Raldh1-3, dark blue stain indicates expression. Stage 34. di, diencephalon; cf, choroid fissure; dr, dorsal retina; e, endoderm; ey, eye; hb, hindbrain; hy, hypophysis; le, lens; ll, lateral line precursors; lpm, lateral plate mesoderm; m/h, midbrain hindbrain boundary; no, notochord; nt, neural tube; op, olfactory placode; ov, otic vesicle; pk, pronephritic kidney; pd, pronephritic duct; re, retina; rp, roof plate; s, somite; te, telencephalon. Scale bars are 100 Î¼M.
	Expression of Raldh3 during organogenesis. In situ hybridization of Raldh3, dark blue stain indicates expression. Stage 34. di, diencephalon; cf, choroid fissure; dr, dorsal retina; e, endoderm; ey, eye; hb, hindbrain; hy, hypophysis; le, lens; ll, lateral line precursors; lpm, lateral plate mesoderm; m/h, midbrain hindbrain boundary; no, notochord; nt, neural tube; op, olfactory placode; ov, otic vesicle; pk, pronephritic kidney; pd, pronephritic duct; re, retina; rp, roof plate; s, somite; te, telencephalon. Scale bars are 100 Î¼M.
	Expression of Raldh3 during organogenesis. In situ hybridization of Raldh3, dark blue stain indicates expression. Stage 34. di, diencephalon; cf, choroid fissure; dr, dorsal retina; e, endoderm; ey, eye; hb, hindbrain; hy, hypophysis; le, lens; ll, lateral line precursors; lpm, lateral plate mesoderm; m/h, midbrain hindbrain boundary; no, notochord; nt, neural tube; op, olfactory placode; ov, otic vesicle; pk, pronephritic kidney; pd, pronephritic duct; re, retina; rp, roof plate; s, somite; te, telencephalon. Scale bars are 100 Î¼M.
	Raldh3 expression in stage 34 embryo, red arrows indicate relative positions of transverse sections in panels RâT. (Qâ) Detail of head of embryo shown in panel Q. Abbreviations: di, diencephalon; cf, choroid fissure; dr, dorsal retina; e, endoderm; ey, eye; hb, hindbrain; hy, hypophysis; le, lens; ll, lateral line precursors; lpm, lateral plate mesoderm; m/h, midbrain hindbrain boundary; no, notochord; nt, neural tube; op, olfactory placode; ov, otic vesicle; pk, pronephritic kidney; pd, pronephritic duct; re, retina; rp, roof plate; s, somite; te, telencephalon. Scale bars are 100 Î¼M.