
Onion Breeding
and Genetics
We maintain an onion-breeding program
and are committed to the genetic improvement of onion using classical
and molecular approaches. In an average year, we plant over 700
four-meter observation plots to produce and evaluate bulbs for
inbred development, genetic studies, and seed increases. We are
grateful to Gary Kincaid, Palmyra WI, for providing field space every year. For seed production,
bulbs harvested the previous year are selected based on desired
attributes or defined genetic relationships and planted at Madison, WI, to produce seed. We
generally plant at least 30 cages (larger seed increases or testcrosses)
and 200 breeding plots (small self or mass pollinations). Our
crosses and seed increases are primarily done using flies. We
recently released three male-sterile and maintainer pairs (B1717,
B1828, and B2354) for use in hybrid production. We have studied
the combining abilities among onion populations (J. Amer. Soc.
Hort. Sci. 121:604-608), estimated their relatedness using molecular
markers (Theor. Appl. Genet. 90:607-614; J. Amer. Soc. Hort. Sci.
120:752-758), and established the genetic basis of seed yield
(J. Amer. Soc. Hort. Sci. 126:575-578). The vast majority of hybrid-onion
seed is produced using a source of cytoplasmic male-sterility
(CMS) that traces back to a single plant identified in 1925 in
Davis, CA. This is an undesirable state of cytoplasmic uniformity.
We identified molecular markers distinguishing onion cytoplasms
and assessed cytoplasmic diversity (Theor. Appl. Genet. 86:128-134,
J. Amer. Soc. Hort. Sci. 119:90-93, Theor. Appl. Genet. 90:263-268).
We demonstrated that two distinct types of onion CMS exist and
showed that identical sources of CMS have been independently isolated
and exploited around the world (Theor. Appl. Genet. 101:778-782).
We released a unique source of CMS, conditioned by the cytoplasm
of Allium galanthum backcrossed to the bulb onion (J. Amer.
Soc. Hort. Sci. 124:626-629)
Onion Phylogenetics
Even though the onion is the world's second most economically
important vegetable crop, little is known about its progenitor,
relationships to wild relatives, and genetic diversity. We used
molecular markers in the organellar (Plant Syst. Evol. 183:17-31,
Gen. Res. Crop Evol. 44:307-313) and nuclear (Amer. J. Bot. 82:1455-1462)
genomes to identify Allium
vavilovii as the wild species most closely related to
the bulb onion. We are cooperating with researchers in Turkmenistan
to expand the world's collection of this important wild species.
Mapping the
Onion Genome
We have developed a low-density genetic map of onion, comprised of morphological,
RFLP, and RAPD markers (Theor. Appl. Genet. 96:52-62) and developed
the first set of onion ESTs (Theor. Appl. Genet. 103:979-991).
This work has been continued with support of a major grant from
USDA-IFAFS
to support EST development in onion and comparative mapping of
asparagus, garlic, and onion (the most economically important
crops of the monocot order Asparagales). We demonstrated that
the GC characteristics of onion coding regions are more similar
to the eudicots than the grasses (Plant Cell 16:114-125). Our
goal is develop genomic resources for the Asparagales to answer
basic questions in onion genetics and breeding. Drs. William Martin
and Jernej Jakse are working on the onion-genomics project. This
research is part of an on-going project on onion genetics and
breeding and has been funded by the USDA and onion-seed industry.
Genetics of
Production, Flavor, and Health-Enhancing Attributes of Onion
This is a joint cooperative project
with Dr.
Irwin Goldman, University of Wisconsin, and was funded by
a grant from the USDA Fund for Rural America. This research was
the Ph.D. project of Dr. Claudio Galmarini, who was partially
funded by the National Institute for Agricultural Research (INTA)
from Argentina. Replicated plantings of F2-derived F3 massed onion
families produced bulbs over at least four environments. We are
analyzing onion flavor (pungency), storage ability, solids (soluble
and total), and in vitro antiplatelet activity. This latter attribute
measures the ability of onion to inhibit the aggregation of platelets
in blood, a major cause of heart attacks and strokes. A low density
genetic map of onion was used to determine the genetic bases of
these flavor, production, and health-enhancing attributes of onion.
We identified one major chromosome region explaining a significant
proportion of the phenotypic variation for flavor, solids, and
antiplatelet activity (Mol. Gen. Genomics 265:543-551). We are
presently identifying candidate genes mapping to this important
position in the onion genome. Steve Raines has joined our lab
and will be continuing this research. Our
long-term goal is to develop unique inbreds and hybrids for production
of onion bulbs with acceptable flavor and production characteristics
combined with defined heath-enhancing attributes.
Molecular-Facilitated
Selection of Maintainer Lines in Onion
Using classical crosses, it takes at
least four to eight years to determine the cytoplasm and nuclear
genotype at the Ms locus of onion. We have developed a PCR marker
distinguishing male-fertile and male-sterile cytoplasms of onion
(Theor. Appl. Genet. 90:263-268), reducing from years to a few
hours the time required to establish cytoplasms. We identified
molecular markers flanking the nuclear Ms locus at 1.9 and 8.1
cM (in press in JASHS). We identified single nucleotide polymorphisms
(SNPs) in the genomic sequence of the most closely linked marker
and converted these SNPs to PCR-based detection. (J. Amer. Soc.
Hort. Sci. 127:576-582).
Cucurbit Organellar
Genetics
The mitochondria of cucumber and melon
are unique in that they possess the largest known genomes and
show paternal transmission (J. Hered. 88:232-235; Theor. Appl.
Genet. 97:122-128). Dr.Jason Lilly characterized the molecular
basis of this huge genome expansion. Random mitochondrial fragments
unique to the huge genome of cucumber have been cloned and over
50 kb of sequence generated. Small (about 100 bp), highly repetitive,
dispersed sequences have contributed significantly to the expansion
of this genome (Genetics 159:317-328). With our colleagues, we
used FISH analyses to reveal great structural diversity in the
chloroplast DNA (Plant Cell 13:245-254). The publication was awarded
"Best Paper in The
Plant Cell" for the year 2001. Drs. Lilly and Grzegorz
Bartoszewski studied the genetic bases of unique mitochondrial
mutants of cucumber conditioning strong mosaic (msc) phenotypes
(Curr. Genet. 40:144-151, Curr. Genet. 45:45-53). Sulieman Al-Faifi
is working to understand the mechanism of paternal transmission
of the cucumber mitochondrial genome.
Mapping Virus
Resistances in Cucumber
Genes conditioning resistance to PRSV-W
and ZYMV were mapped relative to RFLPs, RAPDs, and AFLPs, using
recombinant inbred lines developed from TMG1 x Straight 8. Numerous
students and colleagues contributed to this work and it was completed
by Young
Hoon Park (Genome 43:1003-1010). We are now converting a cosegregating
AFLP marker to an easier PCR-based marker and studying the synteny
among virus resistance loci in the cucurbits. Neil Williams is
continuing this research with support of a grant in 2003 from
the USDA-NRI program.
Late-Blight
Resistance in Mexican Diploid Solanum species
This was a joint project with the late Dr. Robert Hanneman, USDA/ARS and the University
of Wisconsin, and was the Ph.D. project of Dr. Joe Kuhl. Late
blight resistance and susceptibility were identified in Mexican
diploid 1EBN species. Solanum pinnatisectum was crossed
with the susceptible S. cardiophyllum and a BC family generated.
RFLPs were identified on all chromosome arms and mapped late-blight
resistance to chromosome 7 in a region not previously associated
with late-blight resistance loci (Mol. Genet. Genomics 265:977-985).
Monica Norby is presently studying the synteny among late-blight
resistance loci in Solanum. This project has been funded
by the USDA. We also showed that unilateral incompatibilities
among Mexican 2x(1EBN) species is inherited independently of the
S locus in Solanum (Sexual Plant Reprod. 14:305-313).