Mol Gen Genet (1996) 251:52-59 © Springer-Verlag 1996
Susan W. Gorman. Dawn Banasiak
Cecelia FaMey - Sheila McCormick
A 610 kb YAC clone harbors 7 cM of tomato (tycopersicon esculentum) DNA
that includes the male sterile 14 gene and a hotspot for recombination
Received: 6 July 1995 / Accepted: 8 November 1995
Abstract Pollen development requires both sporo-
phytic and gametophytic gene expression. We are using
a map-based cloning technique to isolate sporophytic
genes which, when mutant, cause pollen abortion and
a male sterile (ms) phenotype in tomato (Lycopersicon
esculentum). We have genetically characterized one ms
locus (ms14) using RFLP analysis and identified flank-
ing markers. High-resolution genomic physical map-
ping indicates that the ms14 locus is located m
a ~300 kb region. We have identified a YAC clone
with an insert size of ,,-610 kb that contains the msl4-
linked markers, reflects the organization of the physical
map and therefore most probably contains the msl4
gene. In addition, we present evidence that the relation-
ship between physical and genetic distance in this
chromosomal region changes abruptly from ,-~ 105-140
kb/cM to less than 24kb/cM, and suggest that
the TG393-TG104 region is a hotspot for recombi-
nation.
Key words Positional (map-based) cloning "
Pulsed field electrophoresis • Physical mapping.
RFLP mapping • Pollen development
Introduction
Pollen development in higher plants is a complex pro-
cess that requires the coordination of gene expression
in both the sporophytic and gametophytic tissues of the
anther (reviewed in McCormick 1993). Mutations in
male sterile (ms) loci result in aborted pollen and gener-
ally require the homozygous recessive condition in
Communicated by R. Hagemann
S. W. Gorman • D. Banasiak • C. Fairley • S. McCormick (E~])
Plant Gene Expression Center, USDA/ARS and University of
California-Berkeley, 800 Buchanan St., Albany, CA 94710, USA
order to exhibit the phenotype. This suggests that these
genes act in the sporophytic tissues of the anther but
affect the developing microspores (gametophytes).
Male sterile genes might be expressed before or during
meiosis in the sporophytic pollen mother cell or in the
tapetum or other sporophytic cells of the anther. We
hope to develop an understanding of the sporophytic
contribution to pollen development by studying the
products of male sterile loci.
Current biochemical and molecular information do
not provide a clear picture of the mechanism(s) of
nuclearly encoded male sterility. Although three genes
have been isolated that produce a male sterile pheno-
type when mutant, their gene products do not provide
obvious clues to the roles that these genes play during
pollen development. For example, Moffatt and Somer-
ville (1988) isolated an adenine phosphoribosyl trans-
ferase (aprt-) Arabidopsis mutant, but how a lesion in
a nucleic acid salvage pathway disrupts pollen develop-
ment is unclear. The ms2 gene from Arabidopsis (Aarts
et al. 1993) contains a small region homologous to
a wheat mitochrondrial ORF, but no function has been
ascribed to this mitochondrial region or to the gene
product of the ms2 gene. In maize, a male sterile gene
designated Ms45 (M. Albertsen, personal communica-
tion) was cloned (Albertsen et al. 1993) that showed
33% homology over 256 amino acids with strictosidine
synthase. Although the action of strictosidine synthase
in the indole alkaloid pathway is understood, it is not
clear whether or how indole sythesis might be involved
in pollen development. These examples serve to illus-
trate the complexities of male gametophyte develop-
ment and, therefore, it will probably be necessary to
isolate and characterize a large number of male sterile
gene products before we can develop a comprehensive
understanding of the sporophytic contribution(s) to
pollen development. In tomato (Lycopersicon escuIen-
turn), more than 40 spontaneous ms loci have been
identified (summarized in Kaul 1988). We have targeted
for isolation one of these male sterile genes, ms14; in