“If species have a common origin where did they pick up the ingredients
which produce this sexual incompatibility? Almost certainly it is a
variation in which something has been added. We have come to see that
variations can very commonly—I do not say always—be distinguished as
positive and negative.... Now we have no difficulty in finding evidence
of variation by loss, but variations by addition are rarities, even
if there are any such which must be so accounted. The variations to
which interspecific sterility is due are obviously variations in which
something is apparently added to the stock of ingredients. It is one of
the common experiences of the breeder that when a hybrid is partially
sterile, and from it any fertile offspring can be obtained, the
sterility, once lost, disappears. This has been the history of many,
perhaps most, of our cultivated plants of hybrid origin.
“The production of an indubitably sterile hybrid from completely
fertile parents which has arisen under critical observation is the
event for which we wait. Until this event is witnessed, our knowledge
of evolution is incomplete in a vital respect. From time to time such
an observation is published, but none has yet survived criticism.”
(_Science_, Jan. 20, 1922, pp. 58, 59.)
But what of the chromosomal mutant? For our knowledge of this type
of mutation we are largely indebted to Blakeslee’s researches and
experiments on the Jimson weed (_Datura stramonium_). According to
Blakeslee, chromosomal mutants result from duplication, or from
reduction, of the chromosomes, and they are classified as _balanced_ or
_unbalanced_ types according as all, or only some, of the chromosomal
linkage-groups are similarly doubled or reduced. If only one of the
homologous chromosomes of a synaptic pair is doubled, the mutant
is termed a _triploid_ form. It is balanced when one homologous
chromosome is doubled in every synaptic pair, but if one or more
chromosomes be added to, or subtracted from, this balanced triploid
complex, the mutant is termed an unbalanced triploid. When all the
chromosomes of the normal diploid complex are uniformly doubled, we
have a balanced _tetraploid_ race. The subtraction or addition of
one or more chromosomes in the case of a balanced tetraploid complex
renders it an unbalanced tetraploid mutant. The retention in somatic
cells of the haploid number of chromosomes characteristic of gametes
and gametophytes gives a balanced _haploid_ mutant, from which hitherto
no unbalanced haploids have been obtained. The normal diploid type and
the balanced tetraploid type are said to constitute an _even_ balance,
while balanced triploids and haploids constitute an _odd_ balance. The
odd balances and all the unbalanced mutants are largely sterile. Thus,
for example, more than 80% of the pollen of the haploid mutant is bad.
“The normal Jimson Weed,” says Blakeslee, “is diploid (2n) with a total
of 24 chromosomes in somatic cells. In previous papers the finding of
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