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For negative controls, both NaRC and HRC differentiations were largely undisturbed in either foxi3a MOmis (Figure 9B) or foxi3b MOmis (data not shown).

However, when we forced foxi3a expression in bmp7 morphants, it was sufficient to restore both NaRC (100%, n = 62) and HRC (100%, n = 62) differentiation with high penetrance (Figure 8G).

However, when we forced foxi3a expression in Notch-ON embryos, it was sufficient to restore both NaRC (100%, n = 74) and HRC (100%, n = 74) differentiation with high penetrance (Figure 8D).

In 24-hpf bmp7 morphants, although we still could detect P63 expression, both NaRC (0%, n = 88) and HRC (0%, n = 88) differentiations were completely undetectable (Figure 8F) due to the loss of epidermal IC identity (detected by a foxi3a in situ study, data not shown).

This observation suggests that both NaRCs (atp1b1b-positive cells) and HRCs (ca2a-positive cells) might come from common progenitors and then subsequently differentiate.

In foxi3b morphants, the apical openings of both NaRCs and HRCs were reduced (Figure 9O), while they were no longer visible in foxi3a morphants (Figure 9N).

By counting the number of epidermal ICs (including both NaRCs and HRCs) and epidermal SCs (by P63 antibody staining) at 24 hpf, we detected significantly higher epidermal ICs (NaRCs, 41±5 vs. 28±5, n = 7, p<0.05; HRCs, 88±8 vs. 21±3, n = 7, p<0.05) and lower epidermal SCs (135±8 vs. 212±11, n = 7, p<0.05) in mibta52b mutants.

However, when foxi3a mRNA was misexpressed, we found that the elevated expression level of foxi3a was sufficient to promote precocious differentiation of both NaRCs (73%, n = 171) and HRCs (4%, n = 171) not only within the epidermal IC domain but also in the ectopic sites of the cephalic epidermis at the 5-s stage (Figure 7B).

This result clearly demonstrates that both foxi3a and foxi3b are sufficient to promote the primary setting of NaRC differentiation, while the secondary setting of HRC differentiation strongly relies on a higher concentration of foxi3a.

Although the loss-of-function assay suggested that only foxi3a but not foxi3b is required for NaRC and HRC differentiation, the gain-of-function assay showed that both exogenous foxi3a and foxi3b are sufficient to promote ectopic NaRC and HRC differentiation in a wild-type background.

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