Precision and accuracy in the timing of transcription. (A) Examples of differences in the timing of transcriptional activation of a gene between wild-type (wt) and perturbed/mutant (mut) early Drosophila embryos. (B) Transcriptional synchrony in a population of cells can be determined by measuring the time when transcription is activated in a given fraction of cells, e.g. twt and tA are the times when 50% of wt and mut-A cells are active, respectively. Accuracy in the timing of transcription can refer to changes in the average value of this time, as demonstrated between mut-A and wt embryos. (C) Precision in timing can be measured in absolute time, or relative to a developmental event, or relative to other cells within an organism. Synchrony of the population of cells can be determined by measuring the time between initial activation (when the signal surpasses a threshold, thd) and t50 (dti). Here, activation in mut-B is less synchronized than in the wt (dtB is larger than dtwt) and therefore the timing of activation is less precise. Since wt and mut-B embryos have the same t50 value, there is no change in the accuracy of the timing of activation. It should be noted that temporal dynamics can affect transcript levels in cells/nuclei. For example, earlier transcription activation can result in the accumulation of more transcripts over time.