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otx2xenopus thalamus 

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Experiment details for otx2

Juraver-Geslin HA et al. (2014) Assay

The conserved barH-like homeobox-2 gene barhl2 acts downstream of orthodentricle-2 and together with iroquois-3 in establishment of the caudal forebrain signaling center induced by Sonic Hedgehog.

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Gene Clone Species Stages Anatomy
otx2.S laevis NF stage 29 and 30 thalamus

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  Fig. 1. The ZLI develops inside the rostral alar p2 territory that expresses barhl2, otx2, irx3 and does not express irx1 and irx2. ISH or Double ISH on wt embryos, shown as dissected neural tubes from a side view, dorsal up, anterior left. The markers and stages are indicated. (D)–(P) Enlargement views centered on p2 as indicated by the red square on A. The pineal gland (red star) located on top of p2 is used as a morphological landmark. The black dashed lines are indicative of the putative anterior and posterior borders of p2. The scale bar stands for 0.5 mm. p: prosomere; MDF: Mid Diencephalic Furrow. (A–C) Time course analysis of shh expression in the Xenopus forebrain: (A) at st. 29 and st. 30 shh is only expressed in the forebrain floor and basal plates; (B) at st. 32 shh is detected in the forming ZLI; (C) at st. 37 development of the ZLI is mostly achieved. (D–L) At st. 30 and st. 31 the p2 alar plate contains two subdomains. (D) The ZLI forms in a domain that expresses barhl2. (E) The p2 anterior limit of barhl2 abuts that of fezf2, which marks the p3/p2 boundary; in the p2 alar plate barhl2 is co-expressed with (F) otx2 and (G) irx3. (H) pax6 marked p1 and p3 but is excluded from p2, except for the most dorsal part, which gives rise to the epithalamus. barhl2 is expressed in the part of the p2 domain devoid of pax6 expression, i.e. the mid-diencephalic furrow. A comparative analysis of barhl2 and (I–L) iroquois expression revealed two subdomains inside the alar p2: a rostral p2 domain that expresses barhl2 (I, K), otx2 (F), irx3 (G, J, L) and a caudal p2 domain that expresses barhl2 (I, K), otx2 (F), irx3 (G, J, L), irx1 (I, J) and irx2 (K, L). (M–P) The ZLI develops inside the rostral p2 domain. Double ISH on st. 32 and st. 36 dissected neural tubes with shh and iroquois genes that mark either the rostral p2 territory (M, N) irx3, or the caudal p2 territory (O, P) irx1 as probes. The alar progression of shh expression occurs in a domain that expresses irx3 (M, N) but not irx1 (O, P). (Q) Schematic of diencephalic markers at st. 31. Prosomere p2 is indicated in blue; Areas of expression are shown for fezf2 (green) a marker of the p3–p2 border; nkx2.1 (purple) that marks basal p2; shh (red) marks the p2 basal plate and the ZLI; p1, p3 and the epithalamus are revealed using pax6 (yellow). (R) Schematic of dynamics of p2 alar plate markers expression: The basal plate expresses shh (light blue). Within the p2 territory devoid of pax6 but expressing barhl2 and otx2: At stage 30 the rostral domain expresses irx3 and not irx1/2 (orange) the caudal domain expresses irx1/2/3 (green). At stage 37 the rostral domain expressed the same markers and shh (light blue) and the caudal domain expresses irx1/2 (green).

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Gene Clone Species Stages Anatomy
otx2.S laevis NF stage 32 thalamus

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  Fig. 2. Barhl2-depleted embryos exhibit defects in p2 alar plate patterning. Gene expression profiles of forebrain markers in Barhl2 morphants (MObarhl2) are shown at st. 27, st. 32 and st. 37 as indicated. Both the control (CT) and the injected (INJ) sides of a representative neural tube are shown, anterior left, dorsal up, allowing direct comparison of expression territories. n: number of embryos analyzed. The scale bar stands for 0.4 mm. In Barhl2-depleted embryos (A) the p3–p2 limit, marked by the caudal limit of the dorsal prethalamic marker fezf2 (n=24), or (B) arx2 (n=24) is established properly. The p2-like cells in MObarhl2-injected embryos are present and, at least in part, specified as shown by (C) the expression of the p2 basal plate marker nkx2.1 (n=16), (D) that of otx2 (n=24) and (E) tcf4 (n=20). However, the p2 alar plate is misspecified: (F) the domain in which pax6 is expressed spreads ventrally inside the mid-diencephalic furrow (n=20); In the rostral p2 (G) irx3 expression remains unchanged compared to the CT side (n=24); Conversely (G) irx3 expression is expanded in the caudal p2 of barhl2-depleted embryos (n=24), and expression levels of both caudal p2 markers (H) irx1 (n=24) and (I) irx2 (n=36) were down-regulated upon depletion of Barhl2 activity. (J) At st. 37 the progression of shh inside the p2 alar plate is inhibited (n=16).

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Gene Clone Species Stages Anatomy
otx2.S laevis NF stage 32 thalamus

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  Fig. 3. Within alar p2, depletion of Irx1 and Irx2 activities, or an increase in Irx3 activity, promotes ZLI specification. Double ISH or ISH on embryos (A–D) either depleted for both Irx1 and Irx2 activities or (E–H) overexpressing irx3, or (I) overexpressing irx3-GR using barhl2 (A, E), otx2 (B, F), irx3 (C), irx1 (G), shh (D, H, I) probes as indicated. The scale bar stands for 0.5 mm. RNA encoding for irx3, or irx3-GR were co-injected with a MO control used as tracer, coupled to fluorescein detected by immunohistochemistry (red). (A–D) Co-depletion of Irx1/2 shifts the ZLI caudal border. In st. 32 embryos depleted for irx1/2 (A) barhl2 (n=24) expression is not significantly affected, and (B) otx2 (n=20) expression is slightly decreased. We observed (C) a weak increase of irx3 in the thalamic domain (n=36). (D) The ZLI territory, characterized by shh expression, is enlarged (n=36); DISH using barhl2 together with shh probes shows that the barhl2 anterior border and the ZLI anterior border coincided, indicating that the ZLI posterior boundary is caudally shifted. (E–I)irx3 or irx3-GR overexpression promotes ZLI specification. At st. 32 in irx3 overexpressing embryos (E) barhl2 (n=40) expression is modified compared with the CT sides whereas (F) otx2 (n=40) and (G) irx1 (n=40) expression are weakly decreased in the caudal p2 domain. (H) In irx3 overexpressing embryos the surface of the ZLI territory marked by shh is enlarged (n=22). DISH using barhl2 together with shh as probes demonstrates that the ZLI anterior border is not affected. Note that in embryos overexpressing Irx3 the expansion of the ZLI territory is associated to a decrease in shh staining intensity. (I) In embryos overexpressing a hormone-inducible form of irx3 (irx3-GR) and exposed to dexamethasone at st. 20, the surface of the ZLI territory marked by shh is enlarged (n=10). (J) The average width of p2 is not modified in embryos depleted of Irx1/2 or overexpressing Irx3. Using barhl2 as a p2 marker, the width of p2 was measured (Image J) on both the non-injected and the injected sides of embryos injected with GFP (CT, n=11), depleted for Irx1/2 (MOirx1/2, n=12) or overexpressing Irx3 (Irx3, n=14). The ratio of the p2 width of the injected side relative to the control side is shown. The error bars indicate the standard deviation. (K and L) The ZLI surface is significantly increased in Irx3 overexpressing embryos. (K) We delimited the alar/basal plate boundary of the diencephalon by drawing a line at the base of the ZLI area. We measure the ZLI area as the alar p2 area expressing Xshh. (L) Using Image J the surface of the ZLI territory was measured on both the non-injected and the injected sides of embryos injected with GFP (CT, n=9), overexpressing Irx3 (Irx3, n=9), or overexpressing a hormone-inducible form of Irx3 (Irx3-GR, n=9). The average ratio of the ZLI surface of the injected size relative to the control size is shown. The error bars indicate the standard deviation. We observed a significant increase of the ZLI surface in both Irx3 overexpressing embryos (R=1.5±0.19, t-test p≤0.004) and Irx3-GR overexpressing embryos (R=1.2±0.17, t-test p≤0.03).
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