All insects possess homologous segments, but segment specification differs radically among insect orders. In Drosophila, maternal morphogens control the patterned activation of gap genes, which encode transcriptional regulators that shape the patterned expression of pair-rule genes. This patterning cascade takes place before cellularization. Pair-rule gene products subsequently 'imprint' segment polarity genes with reiterated patterns, thus defining the primordial segments. This mechanism must be greatly modified in insect groups in which many segments emerge only after cellularization. In beetles and parasitic wasps, for instance, pair-rule homologues are expressed in patterns consistent with roles during segmentation, but these patterns emerge within cellular fields. In contrast, although in locusts pair-rule homologues may not control segmentation, some segment polarity genes and their interactions are conserved. Perhaps segmentation is modular, with each module autonomously expressing a characteristic intrinsic behaviour in response to transient stimuli. If so, evolution could rearrange inputs to modules without changing their intrinsic behaviours. Here we suggest, using computer simulations, that the Drosophila segment polarity genes constitute such a module, and that this module is resistant to variations in the kinetic constants that govern its behaviour.

Loss of biodiversity is one of the world's overriding environmental challenges. Reducing those losses by creating reserve networks is a cornerstone of global conservation and resource management. Historically, assembly of reserve networks has been ad hoc, but recently the focus has shifted to identifying optimal reserve networks. We show that while comprehensive reserve network design is best when the entire network can be implemented immediately, when conservation investments must be staged over years, such solutions actually may be sub-optimal in the context of biodiversity loss and uncertainty. Simple decision rules, such as protecting the available site with the highest irreplaceability or with the highest species richness, may be more effective when implementation occurs over many years.