Optimal encoding design for one-dimensional positioning. (A) Optimal encoding for two morphogens (M = 2). For positively correlated noises, the oppositely directed gradients are better than identically directed gradients, and vice versa for negatively correlated noises. (Left) (Blue and red curves) Variability of morphogen gradients over 150 embryos. (Bold curves) Average profiles are u1(x) = exp(−5x), u2(x) = exp(4(x − 1)) (upper left), and u2 (x) = exp(−4x) u2 (x) = exp(−4x) (lower leftt). As a noise source, we assumed the variability of the morphogen source levels that obeys two-variable Gaussian distribution, where the SD was set as σ1 = σ2 = 0.2. (Upper right) When the variability of the morphogen source levels are correlated with each other, the concentrations of the two morphogens (u1′, u2′) observed at any position x0 are also correlated with each other, where the correlation coefficient is denoted by ρ12. The upper right panels show the distributions of the relative deviations of concentrations from their averages at x0, δui(x0) ≡ u′i/ui (x0) −1. ρ12 = 0.7 (left) and ρ12 = −0.7 (right). (Lower right) The histograms (∝) of the estimated error relative to the organ size. It becomes sharper when the appropriate choice of relative directions of gradients is made in the presence of correlation (ρ ≠ 0). (B) Optimal encoding for three morphogens when inequality |ζ12| > |ζ23| > |ζ31| holds (see the text for the definition of ζij). The direction of a morphogen gradient is fixed as u1 (x) = exp(−5x). The directions for the rest two are u2 (x) = exp(−4x) or u2 (x) = exp(4(x − 1)) for the second morphogen, and u3 (x) = exp(−3x) or u3 (x) = exp(3(x − 1)) for the third morphogen. Parameters: (|η1|, |η2|, |η3|) = (50, 40, 30) for PP values and (σ1, σ2, σ3, ρ12, ρ23, ρ31) = (0.1, 0.1, 0.1, ±0.5, ±0.5, 0) for the variability of morphogen source levels.