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Bownes (1992) used their model to compare the sex-determination system in
the house fly ( Figure 10.4 ). According to the Nothiger and Steinmann-Zwicky
(1985) model, the male-determining factor ( M ) in M. domestica would corre-
spond to the repressor (R) of Sxl + . The genes tra + and Ag + may be equivalent to
da + in Drosophila . The F gene of M. domestica could be equivalent to Sxl + . Sxl +
is involved in dosage compensation in Drosophila , but dosage compensation is
not needed in species such as M. domestica with heterochromatic sex chromo-
somes (which usually contain few coding regions) or no sex chromosomes. As a
result, insects with heterochromatic sex chromosomes or no sex chromosomes
can survive mutations of tra + , Ag + , and F ; such mutations can alter sex determi-
nation, but are not lethal to one sex.
Different genera of mosquitoes have several different sex-determination
systems, but these systems still may conform to the Nothiger and Steinmann-
Zwicky model ( Bownes 1992 ). Anopheles gambiae and Anopheles culicifacies
have XY males and XX females. Sex in Aedes is determined by a dominant male-
determining factor. Intersex flies with phenotypes similar to the ix , dsx , and tra
mutants of Drosophila have been found in Aedes aegypti and Culex pipiens .
A single gene located on an autosome determines sex in Culex ; Culex gynan-
dromorphs have been found, suggesting that sex determination is cell autono-
mous, as it is in Drosophila . The sex of some northern strains of Aedes depends
upon the temperature at which they are reared, with males transformed into
intersexes at higher temperatures. This suggests that an allele equivalent to ix +
is temperature sensitive in these populations. In Cx. pipiens , a sex-linked gene
cercus ( c ) changes females into intersexes; these intersexes are sterile and fail
to take blood meals. Bownes (1992) speculated that it is possible that cercus + is
similar to tra + , ix + , or dsx + of Drosophila .
Sexual differentiation among different organisms (flies, nematodes, and
mammals) has superficially similar patterns of hierarchical control (for reviews,
see Marin and Baker 1998, McAllister and McVean 2000 ). Comparative genetic
analysis suggests that the functions of tra + , tra-2 + , and dsx + may be conserved
throughout higher eukaryotes ( Verhulst et  al. 2010a ). Sex-determining mecha-
nisms are, however, variable and the function of Sxl + is not conserved among
all arthropods. Verhulst et al. (2010a) evaluated what is known about insect sex
determination and conclude
“The primary signal that starts sex determination is processed by a cascade
of genes ending with the conserved switch doublesex that controls sexual
differentiation. Transformer is the doublesex splicing regulator and has
been found in all examined insects, indicating its ancestral function as a sex-
determining gene.”
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