But crossing produces, in the second filial generation (F₂), pure
(homozygous) as well as hybrid (heterozygous) forms.⅖ In some cases
these pure forms are new, the phenotype being different from that
of either pure grandparent. Such a result is produced by _random
assortment_ of the chromosomes in gamete and zygote formation, and
occurs when the genes for two or more pairs of contrasted characters
are located in different chromosome pairs. The phenomenon is formulated
in Mendel’s Second Law, the law of independent assortment. The novelty,
however, of the true-breeding forms thus produced is not absolute,
but relative. There is no origination of new hereditary factors. It
is simply a recombination of the old genes of different stocks, the
genes themselves undergoing no intrinsic alteration. The combination
is new, but not the elements combined. In addition to chromosomal
recombination, we have factorial recombination by means of crossovers.
This, too, can produce new and true-breeding forms of a fixed nature,
but here, likewise, it is the combination, and not the elements
combined, which is new. The “new” forms thus produced are called, as we
have seen, pseudomutants. When pseudomutations, that is, crossovers,
occur in conjunction with the condition of balanced lethals, they
closely simulate genuine factorial mutations. This is exemplified in
the case of De Vries’ _Œnothera Lamarckiana_, which is the product of
a crossover supervening upon a situation of balanced lethals. In cases
of this kind, the crossover releases hitherto suppressed recessive
characters, giving the appearance of real mutation. “The workers with
Drosophila,” says Davis, “seem inclined to believe that much of the
phenomena simulating mutation in their material is in reality the
appearance of characters set free by the breaking of lethal adjustments
which held the characters latent. Well-known workers have arrived at
similar conclusions for _Œnothera_ material and are not content to
accept as evidence of mutations the behavior of _Lamarckiana_ and some
other forms when they throw their marked variants.” (_Science_, Feb. 3,
1922, p. 111.)
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