several years; dozens of experiments take place simulta-
neously, often involving hundreds of fish. Kallman never
became overwhelmed by this orderly mass of extremely
complicated, interrelated information. His complete famil-
iarity stood him in good stead when, in April 1959, Gordon
suddenly died. Less than a year later, Kallman was put in
charge of the laboratory, which continued to function with
little noticeable change. Unquestionably, Kallman knew
more about what was going on in the New York Zoological
Societys fish genetics laboratory than did anyone else. He
had become the man of the moment, and he continued to
lead this premier laboratory for the next 33 years.
While overseeing all the laboratorys daily operations,
including shipping live fish to investigators in other insti-
tutions, Kallman carried on his own vigorous research pro-
gram. In all, he published at least 70 substantive papers,
amounting to 1164 pages. A dozen of his articles concerned
the unique sex chromosomes of the platyfish (Xiphophorus
maculatus), which has 3 kinds instead of the usual 2: XY and
YY males, and XX, WX, and WY females. The WW female
has not been found in the wild, but was produced in the
laboratory. In a most impressive review, Kallman (1984)
marshals the evidence for a theory of how this system of sex
differentiation evolved and argues convincingly against the
idea that it simply results from the primitiveness of piscine
chromosomes. However, the existence of atypical sex deter-
mination complicates the matter because individuals that
exhibit normal sexual development and function can pos-
sess the sex chromosomes of the opposite sex: XY and YY
females and XX, WX, WY, and even WW males. Kallman
showed that the interactions of the sex chromosomes with
specific autosomal factors played important roles in such
often highly predictable events. He also convincingly laid to
rest the phenomenon of sex reversal in swordtails from
functional female to functional male.
In 1973, Kallman, Martin P. Schreibman, and Valerie
Borkoski announced the discovery of a sex-linked gene in
Xiphophorus maculatus, the alleles of which determined
when the pituitary gonadotrops were activated and, thus,
the onset of sexual maturation of males and females (Kall-
man et al., 1973). Inasmuch as the release of androgens
terminates growth, early-maturing males were significantly
smaller than late-maturing ones. So far, 9 different alleles
have been identified in this species (Kallman, 1989). The
polymorphism of this gene differed in each of the 10 species
of Xiphophorus that Kallman studied. Moreover, it was as-
sociated with such phenomena as sexually active dwarf
males or giant ones that never matured. Some species had 2
genotypes of differently sized males, which exhibited very
different mating behavior patterns. Kallmans discovery of
this sex-linked gene added a new dimension to our under-
standing of fish growth.
The color patterns of the common platyfish have been
studied intensively by fish culturists, geneticists, evolution-
ists, pigment-cell biologists, and cancer researchers. Un-
doubtedly, Kallmans contributions to our understanding of
the inheritance of coloration and pigmentary neoplasia in
Xiphophorus provided basic data on which many of these
Professor Klaus D. Kallman, Director of
the Xiphophorus Genetic Stock Center.
This photograph was taken at the
Osborn Laboratories of Marine
Research, New York Aquarium,
Brooklyn, New York, sometime in the
late 1970s. (Reproduced with permission
from the Wildlife Conservation Society.)
S4
James W. Atz and Steven Kazianis
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