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Summary 1.Trophic position is a fundamental feature of food-web structure, knowledge of which is being improved by stable isotope approaches which assume a constant enrichment in heavier isotopes in consumers relative to their diet. 2. We argue that the typical enrichment reflects a dynamic equilibrium between fractionation vectors associated with assimilation and excretion. 3. We develop a linear model to characterize the relationship between the equilibrium isotopic signature and the feeding rate influenced by isotopic discrimination during assimilation and excretion. 4. We present new data for both diet switching and starvation experiments using a marine worm Nereis virens and use this, and previously published data for a fish, bird and mammal to calculate controlling parameters from observations of the isotopic signature following diet-switching and the onset of starvation. 5. We show that the observed variance in isotopic signatures at each trophic step carries substantially more information than has been used hitherto and is influenced by feeding rate in addition to the isotopic signature of the food source. 6. Using the linear model as a tool we predict that parasitic organisms may be depleted relative to the tissues of their host.

Retinoid X receptor (RXR) and Ultraspiracle (USP) play a central role as ubiquitous heterodimerization partners of many nuclear receptors. While it has long been accepted that a wide range of ligands can activate vertebrate/ mollusc RXRs, the existence and necessity of specific endogenous ligands activating RXR-USP in vivo is still matter of intense debate. Here we report the existence of a novel type of RXR-USP with a ligand-independent functional conformation. Our studies involved Tribolium USP (TcUSP) as representative of most arthropod RXR-USPs, with high sequence homology to vertebrate/mollusc RXRs. The crystal structure of the ligand-binding domain of TcUSP was solved in the context of the functional heterodimer with the ecdysone receptor (EcR). While EcR exhibits a canonical ligand-bound conformation, USP adopts an original apo structure. Our functional data demonstrate that TcUSP is a constitutively silent partner of EcR, and that none of the RXR ligands can bind and activate TcUSP. These findings together with a phylogenetic analysis suggest that RXR-USPs have undergone remarkable functional shifts during evolution and give insight into receptor-ligand binding evolution and dynamics.

We have characterised a P-element-induced prepupal mutant of Drosophila melanogaster which after an apparently normal embryonic and larval development fails to complete head eversion, an essential step in metamorphosis. The P-element insertion disrupts an ecdysone-regulated transcript which, although expressed during embryonic and larval stages, appears critical for preparing the late prepupal response to ecdysone. By a combination of molecular and genetic studies, in which we recovered new alleles, we show that the locus is complex, containing at least two distinct promoters. Its transcripts contain a short region described previously by R. Schüh et al. (1986, Cell 47, 1025-1032), who screened for homologues of the Krüppel gene. Our studies on the corresponding gene, named Krüppel-homolog (Kr-h), add to a growing body of evidence that specific isoforms of a number of key genes are implicated in both embryogenesis and metamorphosis.