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The Chromosomal Theory

Society of Natural Science in 1865, and published it the


following year. While his findings were often cited and discussed,
their significance went unappreciated for about 35
years. Many explanations have been proposed for this delay.
First, Mendels adherence to mathematical analysis of
probability events was quite unusual for biological studies
in those days. Perhaps it seemed foreign to his contemporaries.
More important, his conclusions did not fit well
with existing hypotheses concerning the cause of variation
among organisms. The topic of natural variation intrigued
students of evolutionary theory. This group, stimulated
by the proposal developed by Charles Darwin and Alfred
Russel Wallace, subscribed to the theory of continuous
variation, which held that offspring were a blend of their
parents phenotypes. As we mentioned earlier, Mendel theorized
that variation was due to a dominancerecessive
relationship between discrete or particulate units, resulting
in discontinuous variation. For example, note that the F2
flowers in Figure 31 are either white or violet, never something
intermediate. Mendel proposed that the F2 offspring
of a dihybrid cross are expressing traits produced by new
combinations of previously existing unit factors. As a result,
Mendels hypotheses did not fit well with the evolutionists
preconceptions about causes of variation.
It is also likely that Mendels contemporaries failed to
realize that Mendels postulates explained how variation was
transmitted to offspring. Instead, they may have attempted
to interpret his work in a way that addressed the issue of why
certain phenotypes survive preferentially. It was this latter
question that had been addressed in the theory of natural
selection, but it was not addressed by Mendel. The collective
vision of Mendels scientific colleagues may have been
obscured by the impact of Darwins extraordinary theory of
organic evolution.
The Chromosomal Theory
of Inheritance
In the latter part of the nineteenth century, a remarkable observation
set the scene for the recognition of Mendels work:
Walter Flemmings discovery of chromosomes in the nuclei
of salamander cells. In 1879, Flemming described the behavior
of these thread-like structures during cell division. As a
result of his findings and the work of many other cytologists,
the presence of discrete units within the nucleus soon
became an integral part of scientists ideas about inheritance.
In the early twentieth century, hybridization experiments
similar to Mendels were performed independently

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