Transplantation and ablation experiments have led to the generalization that in insects the mesoderm is naive, and that pattern is imposed upon it by the ectoderm. This has been demonstrated directly by mosaic analysis for the case of one muscle in Drosophila. The unique character of this muscle depends on the activity of sex-determining and homeotic genes, not in the muscle itself, but in the nerve that innervates it. Indirect evidence suggests, however, that homeotic genes specify some aspects of mesoderm patterning autonomously. Homeotic genes are expressed in the mesoderm, and are regulated in a segment-specific pattern analogous to, but different from, that seen in the ectoderm. Moreover, the effects of homeotic mutations on the muscles do not always mirror transformations seen in the epidermis. Here we examine this problem directly, by expressing homeotic genes ectopically in the mesoderm without altering their expression in the overlying ectoderm. We find that the pattern of adult muscle precursor cells characteristic of the thorax can be converted to that seen in the abdomen by expressing the homeotic gene abdominal-A specifically in the mesoderm.

Extradenticle (Exd) and Homothorax (Hth) function as positive transcriptional cofactors of Hox proteins, helping them to bind specifically their direct targets. The posterior Hox protein Abdominal-B (Abd-B) does not require Exd/Hth to bind DNA; and, during embryogenesis, Abd-B represses hth and exd transcription. Here we show that this repression is necessary for Abd-B function, as maintained Exd/Hth expression results in transformations similar to those observed in loss-of-function Abd-B mutants. We characterize the cis regulatory module directly regulated by Abd-B in the empty spiracles gene and show that the Exd/Hth complex interferes with Abd-B binding to this enhancer. Our results suggest that this novel Exd/Hth function does not require the complex to bind DNA and may be mediated by direct Exd/Hth binding to the Abd-B homeodomain. Thus, in some instances, the main positive cofactor complex for anterior Hox proteins can act as a negative factor for the posterior Hox protein Abd-B. This antagonistic interaction uncovers an alternative way in which MEIS and PBC cofactors can modulate Abd-B like posterior Hox genes during development.

The specific requirement for abd-A and not Ubx in gonad development does not reflect differences in the properties of the proteins that these genes encode, but presumably reflects differences in their regulation. In normal development, the restriction of gonad formation to the posterior abdomen does not depend on the overlap of abd-A and Abd-B expression, but must depend on the regulation of abd-A and Ubx in the sub-population of the mesoderm that forms the gonad. Factors other than homeotic gene expression provide some cues that direct gonadal mesoderm to condense in the correct location.

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SUMMARYDetergent extracts of Trypanosoma cruzi epimastigotes catalysed the hydrolysis of a range of amino-acyl and peptidyl />-nitro-anilides and aminomethylcoumarins. At least three enzymes were detected that cleave Z-Phe-Arg-MCA. Two of these were optimally active at alkaline pH, the other at pH 4-0. Of the two enzymes with alkaline pH optima, one was a cysteine peptidase and was unable to cleave Bz-Arg-MCA readily, whilst the other cleaved Bz-Arg-MCA and was inhibited by diisopropyl fluorophosphate. The acidic enzyme was similar to cathespin L of other eukayrotes with respect to its pH profile, substrate-specificity and inhibitor-sensitivity. Evidence was presented that epimastigotes contain a cysteine-type dipeptidyl aminopeptidase, one or more aminopeptidases, and a serine peptidase that cleaves Boc-AlaAla-pNA. Digitonin solubilization of the activities from cells supports the hypothesis that the cathespin L-like enzyme and the dipeptidyl aminopeptidase are lysosomal, whilst the Bz-Arg-MCA hydrolase, the aminopeptidases and the Boc-Ala-Ala-pNA serine peptidase are cytosolic.