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Figure 1.

Histological Analysis of vasculature in embryonic thymus of Foxn1Δ/Δ Mice.

Haematoxylin & Eosin staining on paraffin section of fetal thymus (E13.5–E14.5). (A) E13.5 Foxn1+/Δ embryonic thymus with RBC detected throughout rudiment (B) Absence of RBC in E13.5 Foxn1Δ/Δ embryonic thymus (C) RBC present in E14.5 Foxn1+/Δ and (D) Foxn1Δ/Δ embryonic thymus. Scale bar = 50 µm; n = 3.

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Figure 2.

Initial embryonic thymic vascularization is defective in Foxn1Δ/Δ Mice.

Immunostaining on frozen transverse sections of fetal thymus (E12.5–E14.5). Endothelial and stromal cell markers used are listed above each column in the corresponding color: CD31+/CD144+ for endothelial cells (green); PDGFR-β+ for neural crest mesenchyme (red); Cytokeratin (blue) or Keratin 5 (red, K5) for epithelial cells. Embryonic stages in the first column and genotypes to the left apply to the entire row unless otherwise labeled. (A–e) CD31+/CD144+ endothelial cells and PDGFR-β+ neural crest cells are present in the thymic capsule region in Foxn1+/Δ (A–E) and Foxn1Δ/Δ mice (a–e) at E12.5. (F–o) CD31+/CD144+ cells followed by PDGFR-β+ cells initially immigrate into the thymus at E13.5 in heterozygotes (F–J) and at E14.5 in homozygotes (k–o). Scale bar, 50 µm; n = 3.

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Figure 3.

Thymic vascularization is sensitive to Foxn1 levels.

Quantification of CD31+/Thymus Area from immunostained frozen sections of embryonic thymi (A) E12.5 Foxn1+/Δ (n = 8) and Foxn1Δ/Δ (n = 6); p>0.05 and E13.5 Foxn1+/Δ (n = 5) and Foxn1Δ/Δ (n = 7); p<0.001. (B) E14.5 Foxn1+/+ (n = 7), Foxn1+/Δ (n = 8); p>0.05, Foxn1Δ/Δ (n = 10); p>0.05, Foxn1Δ/nu (n = 9); p = <0.0001, Foxn1nu/nu (n = 6); p<0.0001. (C–G) CD31+ endothelial cells (green) and PDGFR-β+ neural crest mesenchyme (red) can be detected in the thymic capsule and inside the keratin-positive thymus (blue) in E14.5 (C) Foxn1+/+ (D) Foxn1+/Δ (E) Foxn1Δ/Δ (F) but in the capsule only in Foxn1Δ/nu (G) and Foxn1nu/nu mice. Scale bar, 100 µm; n = 3.

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Figure 4.

Initial LPC thymic immigration is normal in Foxn1Δ mice.

(A) CD45+ LPCs (green) colonize the Foxn1+/Δ and (B) Foxn1Δ/Δ thymus at E11.5. (C) At E11.5, the frequency of CD45+ cells/section was similar between Foxn1+/Δ (n = 10) and Foxn1Δ/Δ (n = 9) thymi; (p>0.05). (D) Immunostaining for CCL21 (green) expression is similar in Foxn1+/Δ and (E) Foxn1Δ/Δ mouse thymus. (F–I) Reduced expression of CCL25 (white) in (G and I) Foxn1Δ/Δ compared to (F and H) Foxn1+/Δ thymus at E11.5. Cytokeratin (red). (J) CCL25 expression was significantly reduced in E13.5 Foxn1Δ/Δ (n = 4), E15.5 Foxn1Δ/Δ (n = 3), and E15.5 Foxn1Δ/nu (n = 6), compared to Foxn1+/Δ control thymi. CD45+ cells (green) were noticeably reduced in (K–L) E12.5, (M–N) E13.5, and (O–P) E14.5 Foxn1Δ/Δ thymi compared to control littermates. Scale bar, 100 µm. qRT experiments represent relative RNA expression of pooled thymi. Controls were set to 1. Asterisks denote statistical significance.

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Figure 5.

Peripheral circulation is connected to the thymus at E14.5.

FITC-dextran (green) facial vein injections and immunostaining for CD31 (red) and cytokeratin (blue) on frozen sagittal sections of fetal mouse thymus. (A–B) FITC-dextran is detected in E14.5 Foxn1+/Δ thymi, tightly associated with CD31+ blood vessels. (C–D) In Foxn1Δ/Δ embryos, FITC-dextran is also present, but the signal is more diffusely associated with vessels. (E–F) FITC-dextran is present throughout E18.5 Foxn1+/Δ thymi tightly associated with branched blood vessels. (G–H) FITC-dextran is more diffusely present in the thymus of E18.5 Foxn1Δ/Δ mice. Scale bar, 100 µm; n = 3.

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Figure 6.

Thymus vascular patterning altered in Foxn1Δ mice.

Immunofluorescence analysis on frozen sagittal sections of Foxn1+/Δ and Foxn1Δ/Δ newborn thymus for CD31+ (red) and PDGFR-β+ (green) cells in (A–C) Foxn1+/Δ and (D–F) Foxn1Δ/Δ mice. (G) Average mean fluorescence intensity for CD31 in Foxn1+/Δ (n = 8) Foxn1Δ/Δ (n = 6) thymus sections. Asterisks denote statistical significance (P<0.05). Scale bar, 50 µm; n = 3.

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Figure 7.

Electron Microscopic Analysis of E18.5 thymus vascular defects.

Electron microscopy analysis of (A–B) Foxn1+/Δ thymus show compact arrangement of cells including endothelial cells and pericytes, while the (C–D) Foxn1Δ/Δ thymus display loose arrangement of cells, vacuolated endothelium, and indistinct vessel walls. Endothelial Cell (EC), Pericyte (P), Lumen of Blood Vessel (L), and Red Blood Cell (RBC) n = 3.

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Figure 8.

VEGF-A and PDGF-B expression reduced in Foxn1Δ/Δ thymus.

(A) VEGF-A expression was significantly reduced in E13.5 Foxn1Δ/Δ (n = 4), E15.5 Foxn1Δ/Δ (n = 3), and E15.5 Foxn1Δ/nu (n = 6), compared to Foxn1+/Δ control thymi. PDGF-B expression was also reduced in E13.5 Foxn1Δ/Δ (n = 4), E15.5 Foxn1Δ/Δ (n = 3), and E15.5 Foxn1Δ/nu (n = 6), compared to Foxn1+/Δ control thymi. Experiments represent relative RNA expression of pooled thymi. Controls were set to 1. Asterisks denote statistical significance (P<0.05). (B–E) Immunofluorescence analysis of VEGF-A expression performed on frozen transverse sections of embryonic thymus for CD31+ (blue), VEGF-A (green) and Cytokeratin (red). VEGF-A expression was detected in thymic endothelium, perivascular cells, and TECs in Foxn1+/Δ and Foxn1Δ/Δ mice (B–E). VEGF-A expression was reduced in (D–E) E13.5 Foxn1Δ/Δ thymus compared to (B–C) E13.5 Foxn1+/Δ controls. Scale bar, 100 µm; n = 3 or more.

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