
Seattle Central Community College
Science & Math Department
1701 Broadway
Seattle, WA USA
"The ecdysone receptor and the
parallel evolution of paedogenesis (larval reproduction) in two species
of gall midges (Diptera: Cecidomyiidae)" (abstract number 1515)
Although most insects reproduce in the adult stage, facultative larval or pupal reproduction (paedogenesis) has evolved at least six times indepently in insects, twice in gall midges of the family Cecidomyiidae (Diptera). Paedogenesis in cecids involves the precocious growth and differentiation of the ovary in an otherwise larval form. We have previously shown (Hodin & Riddiford, 1998) that the timing of expression of the Ecdysone Receptor (EcR) and Ultraspiracle (USP), the two proteins that constitute the functional receptor for the steroid hormone 20-hydroxyecdysone, regulates the timing and progression of ovarian differentiation in Drosophila melanogaster (Diptera: Drosophilidae). Here I test the hypothesis that precocious activation of EcR and USP in the ovaries of paedogenetic gall midges allows for precocious ovarian differentiation. Using monoclonal antibodies directed against insect EcR and USP proteins, I first show that when these gall midges are reared under conditions that promote typical, metamorphic development, up-regulation of EcR and USP occurs in the final larval stage. By contrast, in the paedogenetic life cycle, EcR and USP are up-regulated early in the first larval stage. A similar pattern is seen for two independently-evolved paedogenetic gall midges, Heteropeza pygmaea and Mycophila speyeri. Thus, I conclude that the mechanisms underlying precocious growth of the ovary evolved in parallel in these two lineages.
Of the six independent instances of paedogenesis
(defined broadly to include pupal reproduction, as seen in some chironomid
midges), four of them are among the Diptera. I argue that the paucity
of instances of paedogenesis outside of the "lower" Diptera represents
a developmental constraint on the evolution of paedogenesis. I will
propose that certain ontogenetic features unique to the lower Diptera predispose
them to evolve a paedogenetic life cycle.
PERSONAL STATEMENT
I'm interested in evolutionary patterns, and how
those patterns may be generated by constraints or other biases in ontogenetic
mechanisms. I look for developmentally-tractable organisms that: 1) exhibit
variability in some developmentally interesting character or characters;
2) come from a group with some ideas as to their phylogenetic relationships;
and 3) show some evolutionary pattern (such as evolutionary novelty or
homoplasy) for that (those) character(s). From such a starting point, I
try to determine the developmental underpinnings of the evolutionary pattern.
In this context, I have studied the developmental basis of plasticity (environmentally-based
variability within a genotype) and inter-species variation (genetically-fixed
variability) for ovariole number (a reproductive character) in the fruit
fly Drosophila melanogaster and its close relatives. I have undertaken
a similar study of worker reproduction in the Cape honeybee (Apis mellifera
capensis). At this symposium, I will discuss some work on parallel
evolution of larval reproduction (paedogenesis) in the Cecidomyiidae (the
"gall midge" family of flies). I feel that by uncovering the ontogenetic
bases for differences among closely related species, we can begin to build
a theoretical frame-work for the nascent field of Evo-Devo.
RECENT PUBLICATIONS
Hodin, J. (2000) Plasticity and constraints in
development and evolution.
Journal
of Experimental Zoology (Molecular Development and Evolution) 288: 1-20.
Hodin, J. and Riddiford, L.M. (2000) Different
mechanisms underlie phenotypic plasticity and interspecific variation for
a reproductive character in drosophilids (Insecta: Diptera).
Evolution
(in press).
Hodin, J. and Riddiford, L.M. (2000) Parallel
alterations in the timing of ovarian Ecdysone Receptor and Ultraspiracle
expression characterize the independent evolution of larval reproduction
in two species of gall midges (Diptera: Cecidomyiidae).
Development,
Genes and Evolution 210: 358-372.
Hodin, J. and Riddiford, L.M. (1998) The Ecdysone
Receptor and Ultraspiracle regulate the timing and progression of ovarian
differentiation during Drosophila metamorphosis.
Development,
Genes and Evolution 208: 304-317.
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for further information, contact hodin@alumni.washington.edu