The evolution of multivariate maternal effects (panel A) when the evolutionary optimum <i>θ</i>₂(<i>t</i>) lags <i>φ</i> timesteps behind the evolutionary optimum <i>θ</i>₁(<i>t</i>).

<p>Grey area: fluctuations in optimum precede those of optimum . White area: precedes . Insets on the top of the panel depict the fluctuating selective optima (blue line) and (red line) over time. Panel B: selective cross-correlations that result from varying the time-lag between both selective optima. Note that each point in panel A depicts the maternal effect (at generation ) averaged over ten replicate generations, while error bars depict corresponding standard deviations. Error bars are large when time lags are small, as evolution then leads to alternatively stable states observed in <a href="http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1003550#pcbi-1003550-g003" target="_blank">Figures 3A,B</a>. The results in this figure also extend to a periodic environment in which the time delay <i>φ</i> is stochastically distributed according to a normal distribution with mean and variance varying from 0.1 to 4 (results not shown). Parameters: .