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cluster is found on the X chromosome, and one gene cluster is on the third chro-
mosome. Chorion proteins could not be synthesized quickly and in sufficiently
large quantities unless gene amplification occurred because each chorion-gene
cluster is represented only once in the haploid genome. A 20-fold amplifica-
tion of the chorion genes on the X chromosome and an 80-fold amplification of
the genes on chromosome 3 is found in follicle cells. Amplification is achieved
by replicating the DNA segments at multiple replication origins ( Heck and
Spradling 1990 ). DNA amplification extends bidirectionally for a distance of up
to 40-50 kb to produce a multiforked “onion-skin” structure that contains mul-
tiple copies of DNA containing the chorion genes ( Figure 4.1 ).
Gene amplification also occurs in the chorion genes of the Mediterranean
fruit fly Ceratitis capitata . The overall organization of the cluster is similar to
that of Drosophila , with the same four genes maintained in tandem, in the same
order, and with similar spacing ( Konsolaki et al. 1990 ). Despite the divergence of
Drosophila and Ceratitis family lineages, 120 million years ago, there is high con-
servation in coding sequences and regulatory properties of their chorion genes.
Silk moth chorion proteins are produced over a longer time and involve larger
numbers of genes that have probably arisen by gene duplication . More than 100
Figure 4.1 Amplification of the Drosophila chorion genes in follicle cells. The first three rounds of
DNA replication at the 66D locus on chromosome 3 are shown. The three small arrows represent
three well-characterized chorion genes in this cluster. The polarity of a fourth chorion gene and the
precise location of the origin are unknown. The boundaries of the amplified DNA are much larger
than the chorion protein transcription units within it.
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