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Random Segregation Versus Coupling in Mendelian Inheritance

Author(s): T. H. Morgan
Source: Science, New Series, Vol. 34, No. 873 (Sep. 22, 1911), p. 384
Published by: American Association for the Advancement of Science
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384 SCIENCE [N. S. VOL.XXXIV. No. 873

SPECIAL ARTICLES a repeller of at least three other factors, .. .

RANDOM SEGREGATION VERSUS COUPLING IN some of them may be found able to take
MENDELIAN INHERITANCE precedence of the others in such a way as to
MENDEL'S law of inheritance rests on the annul the present repulsion with subsequent
assumption of random segregation of the fac- coupling as a consequence."
tors for unit characters. The typical propor- In place of attractions, repulsions and or-
tions for two or more characters, such as ders of precedence, and the elaborate systems
9:3:3:1, etc., that characterize Mendelian of coupling, I venture to suggest a compara-
inheritance, depend on an assumption of this tively simple explanation based on results of
kind. In recent years a number of cases inheritance of eye color, body color, wing
have come to light in which when two or mutations and the sex factor for femaleness in
more characters are involved the propor- Drosophila. If the materials that represent
tions do not accord with Mendel's assumption these factors are contained in the chromo-
of random segregation. The most notable somes, and if those factors that "couple" be
cases of this sort are found in sex-limited in- near together in a linear series, then when
heritance in Abraxas and Drosophila, and in the parental pairs (in the heterozygote) con-
several breeds of poultry, in which a coupling jugate like regions will stand opposed. There
between the factors for femaleness and one is good evidence to support the view that
other factor must be assumed to take place, during the strepsinema stage homologous
and in the case of peas where color and shape chromosomes twist around each other, but
of pollen are involved. In addition to these when the chromosomes separate (split) the
cases Bateson and his collaborators (Punnett, split is in a single plane, as maintained by
DeVilmorin and Gregory) have recently pub- Janssens. In consequence, the original ma-
lished1 a number of new ones. terials will, for short distances, be more likely
In order to account for the results Bateson to fall on the same side of the split, while
assumes not only coupling, but also repulsions remoter regions will be as likely to fall on
in the germ cells. The facts appear to be the same side as the last, as on the opposite
exactly comparable to those that I have dis- side. In consequence, we find coupling in
covered in Drosophila, and since these results certain characters, and little or no evidence
have led me to a very simple interpretation, at all of coupling in other characters; the
I venture to contrast Bateson's hypothesis difference depending on the linear distance
with the one that I have to offer. apart of the chromosomal materials that repre-
The facts on which Bateson bases his inter- sent the factors. Such an explanation will
pretation may be briefly stated in his own account for all of the many phenomena that
words, namely: " that if A, a and B, b are two I have observed and will explain equally, I
allelomorphic pairs subject to coupling and think, the other cases so far described. The
repulsion, the factors A and B will repel each results are a simple mechanical result of the
other in the gametogenesis of the double location of the materials in the chromosomes,
heterozygote resulting from the union Ab X and of the method of union of homologous
aB, but will be coupled in the gametogenesis chromosomes, and the proportions that result
of the double heterozygote resulting from the are not so much the expression of a numerical
union AB X ab," and further, "We have as system as of the relative location of the
yet no probable surmise to offer as to the factors in the chromosomes. Instead of ran-
essential nature of this distinction, and all dom segregation in Mendel's sense we find
that can yet be said is that in these special "associations of factors" that are located
cases the distribution of the characters in the near together in the chromosomes. Cytology
heterozygote is affected by the distribution in furnishes the mechanism that the experi-
the original pure parents." Bateson further mental evidence demands.
points out that since " sex in the fowls acts as T. H. MORGAN
'Proc. Royal Soc., Vol. 84, 1911 September 10, 1911.