highly significantP < 0.0001
As expected, predicted expression varied in a manner consistent with the codon usage bias measures: genes near O2 and O3 are predicted to be expressed most and those distant from origins significantly less ( supplementary table S2 , Supplementary Material online), and the genome-wide regression of MELP on distance to the nearest origin was also highly significant ( Fig 4 , F = 96.2, P < 0.0001), albeit weakly predictive ( r 2 = 0.05).