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THE PLANT CELL, Vol 8, Issue 5 887-897, Copyright © 1996 by American Society of Plant Biologists
Chromosomal Organization of TOX2, a Complex Locus Controlling Host-Selective Toxin Biosynthesis in Cochliobolus carbonum
J. H. Ahn and J. D. Walton
Department of Energy-Plant Research Laboratory, Michigan State University, East Lansing, Michigan 48824
Race 1 isolates of the filamentous fungus Cochliobolus carbonum are
exceptionally virulent on certain genotypes of maize due to production of a
cyclic tetrapeptide, HC-toxin. In crosses between toxin-producing (Tox2+)
and toxin-nonproducing (Tox2-) isolates, toxin production segregates in a
simple 1:1 pattern, suggesting the involvement of a single genetic locus,
which has been named TOX2. Earlier work had shown that in isolate SB111,
TOX2 consists in part of two copies of a gene, HTS1, that encodes a 570-kD
cyclic peptide synthetase and is lacking in Tox2- isolates. The genomic
structure of TOX2 and the relationship between the two copies of HTS1 have
now been clarified by using pulsed-field gel electrophoresis and physical
mapping. In isolate SB111, both copies of HTS1 are on the largest
chromosome (3.5 Mb), which is not present in the related Tox2- strain
SB114. Two other genes known or thought to be important for HC-toxin
biosynthesis, TOXA and TOXC, are also on the same chromosome in multiple
copies. Other independent Tox2+ isolates also have two linked copies of
HTS1, but in some isolates the size of the chromosome containing HTS1 is
2.2 Mb. Evidence obtained with Tox2+-unique and with random probes is
consistent with a reciprocal translocation as the cause of the difference
in the size of the HTS1-containing chromosome among the Tox2+ isolates
studied here. Physical mapping of the 3.5-Mb chromosome of SB111 that
contains HTS1 using rare-cutting restriction enzymes and engineered
restriction sites was used to map the chromosomal location of the two
copies of HTS1 and the three copies of TOXC. The results indicate that TOX2
is a complex locus that extends over more than 500 kb. The capacity to
produce HC-toxin did not evolve by any single, simple mechanism.
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