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Furthermore, the reported relationships between expression and intron features are also observed after removing ribosomal proteins from the analyses in Ostreococcus.
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This finding reflected a peculiar relationship between gene expression and intron chromatin structure, described in more detail below.
Analyses done on animal models however have reported a more diverse pattern when investigating relationships between expression levels and intron size and density.
Genes that are highly expressed indeed tend to have shorter introns but genes expressed at low to medium levels show a positive correlation between intron length and expression; hence a roughly bell-shaped dependency between expression level and intron length [ 167].
The Spearman's rank correlation coefficient analysis between expression levels and intron size was performed using the MATLAB program.
In accord with the previous findings (Carmel, Rogozin, et al. 2007), there was a significant positive correlation between expression level and intron density in three of the four analyzed organisms (table 2).
This is confirmed by Spearman's rank correlation coefficient analysis, which shows a statistically significant negative correlation between expression levels and intron size in the four groups, with correlation of the TATA-less group being 2 fold lower than that of the TATA group (Table 4).
This finding suggests that introns might contribute to optimal gene expression [ 155] although this effect is confounded by the stronger connection between expression and evolution rate.
The negative correlation between gene expression and intronic burden observed in this study supports the hypothesis that introns have a biological cost.
Also significant, although of a lesser magnitude, was the positive correlation between gene expression level and intron gain rate and the converse negative of expression with intron loss rate.
We find a positive correlation between expression levels and both intron number (R = +0.0893, p = <0.0005) and intron density (number of introns/kb of CDS; R = +0.0753, p = <0.005).
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