Sex-linked Inheritance in DrosophilaMorgan, Thomas Hunt
Science
Sex-linked Inheritance in Drosophila
Morgan, Thomas Hunt
Drosophila; Heredity
Ordinarily all the sons and none of the daughters show the recessive
sex-linked characters of the mother when the father carries the dominant
allelomorph. The peculiarity of non-disjunction is that sometimes a female
produces a daughter like herself or a son like the father, although the
rest of the offspring are perfectly regular. For example, a vermilion
female mated to a wild male produces vermilion sons and wild-type
daughters, but rarely also a vermilion daughter or a wild-type son. The
production of these exceptions (primary exceptions) by a normal XX female
must be due to an aberrant reduction division at which the two X
chromosomes fail to disjoin from each other. In consequence both remain in
the egg or both pass into the polar body. In the latter case an egg without
an X chromosome is produced. Such an egg fertilized by an X sperm produces
a male with the constitution XO. These males received their single X from
their father and therefore show the father's characters. While these XO
males are exceptions to sex-linked inheritance, the characters that they do
show are perfectly normal, that is, the miniature or the bar or other
sex-linked characters that the XO male has are like those of an XY male,
showing that the Y normally has no effect upon the development of these
characters. But that the Y does play some positive role is proved by the
fact that all the XO males have been found to be absolutely sterile.
While the presence of the Y is necessary for the fertility of the male, it
has no effect upon sex itself. This is shown even more strikingly by the
phenomenon known as secondary non-disjunction. If the two X chromosomes
that fail to disjoin remain in the egg, and this egg is fertilized by a Y
sperm, an XXY individual results. This is a female which is like her mother
in all sex-linked characters (a matroclinous exception), since she received
both her X chromosomes from her mother and none from her father. As far as
sex is concerned this is a perfectly normal female. The extra Y has no
effect upon the appearance of the characters, even in the case of eosin,
where the female is much darker than the male. The only effect which the
extra Y has is as an extra wheel in the machinery of synapsis and
reduction; for, on account of the presence of the Y, both X's of the XXY
female are sometimes left within the ripe egg, a process called secondary
non-disjunction. In consequence, an XXY female regularly produces
exceptions (to the extent of about 4 per cent). A small percentage of
reductions are of this XX-Y type; the majority are X-XY. The XY eggs,
produced by the X-XY reductions, when fertilized by Y sperm, give XYY
males, which show no influence of the extra Y except at synapsis and
reduction. By mating an XXY female to an XYY male, XXYY females have been
produced and these are perfectly normal in appearance. We may conclude from
the fact that visibly indistinguishable males have been produced with the
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