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Changing role of even-skipped during the evolution of insect pattern
formation Nipam H. Patel*†, Eldon E. Ball*‡ & Corey S. Goodman*
*Howard Hughes Medical Institute, Department of Molecular and Cell Biology, 519 LSA,
University of California, Berkeley, California 94720,
USA
†Carnegie Institution, 115 West University Parkway,
Baltimore, Maryland 21210, USA
‡ Molecular Neurobiology
Group, Research School of Biological Sciences, PO Box 475, Canberra, Australian
Capital Territory 2601, Australia
THE development of Drosophila is typical of the so-called long germband mode
of insect development, in which the pattern of segments is established by the
end of the blastoderm stage1,2. Short germband insects, such as
the grasshopper Schistocerca americana, by contrast, generate all or most of
their metameric pattern after the blastoderm stage by the sequential addition
of segments during caudal elongation3. This difference is
discernible at the molecular level in the pattern of initiation of the segment
polarity gene engrailed
4, and the homeotic gene abdominal-A
(ref. 5). For example, in both types of insects, engrailed is expressed by the
highly conserved germband stage4,6 in a pattern of regularly
spaced stripes, one stripe per segment7–9. In Drosophila, the
complete pattern is visible by the end of the blastoderm stage, although
engrailed appears initially in alternate segments in a pair-rule
pattern9–10 that reflects its known control by pair-rule genes
such as even-skipped
11*15. In contrast, in
the grasshopper, the engrailed stripes appear one at a time after the
blastoderm stage as the embryo elongates4. To address the
molecular basis for this difference, we have cloned the grasshopper homologue
of the Drosophila pair-rule gene even-skipped and show that it does not serve a
pair-rule function in early development, although it does have a similar
function in both insects during neurogenesis later in development.
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