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In early development, genes are expressed in spatial patterns which later define cellular identities and tissue locations. The mechanisms of such pattern formation have been studied extensively in early  Drosophila  (fruit fly) embryos. The gap gene  hunchback  ( hb ) is one of the earliest genes to be expressed in anterior-posterior (AP) body segmentation. As a transcriptional regulator for a number of downstream genes, the spatial precision of  hb  expression can have significant effects in the development of the body plan. To investigate the factors contributing to  hb  precision, we used fine spatial and temporal resolution data to develop a quantitative model for the regulation of  hb  expression in the mid-embryo. In particular, modelling  hb  pattern refinement in mid nuclear cleavage cycle 14 (NC14) reveals some of the regulatory contributions of simultaneously-expressed gap genes. Matching the model to recent data from wild-type (WT) embryos and mutants of the gap gene  Krüppel  ( Kr ) indicates that a mid-embryo Hb concentration peak important in thoracic development (at parasegment 4, PS4) is regulated in a dual manner by Kr, with low Kr concentration activating  hb  and high Kr concentration repressing  hb . The processes of gene expression (transcription, translation, transport) are intrinsically random. We used stochastic simulations to characterize the noise generated in  hb  expression. We find that Kr regulation can limit the positional variability of the Hb mid-embryo border. This has been recently corroborated in experimental comparisons of WT and  Kr-  mutant embryos. Further, Kr regulation can decrease uncertainty in mid-embryo  hb  expression (i.e. contribute to a smooth Hb boundary) and decrease between-copy transcriptional variability within nuclei. Since many tissue boundaries are first established by interactions between neighbouring gene expression domains, these properties of Hb-Kr dynamics to diminish the effects of intrinsic expression noise may represent a general mechanism contributing to robustness in early development.

Mid-Embryo Patterning and Precision in Drosophila Segmentation: Krüppel Dual Regulation of hunchback