Recent Patents on DNA & Gene Sequences 2007, 1, 69-76 69
1872-2156/07 $100.00+.00 © 2007 Bentham Science Publishers Ltd.
Current and Future Developments in Patents for Quantitative Trait Loci
in Dairy Cattle
Joel I. Weller*
Institute of Animal Sciences, A. R. O., The Volcani Center, P. O. Box 6, Bet Dagan 50250, Israel
Received: October 5, 2006; Accepted: December 5, 2006; Revised: December 7, 2006
Abstract: Many studies have proposed that rates of genetic gain in dairy cattle can be increased by direct selection on the
individual quantitative loci responsible for the genetic variation in these traits, or selection on linked genetic markers. The
development of DNA-level genetic markers has made detection of QTL nearly routine in all major livestock species. The
studies that attempted to detect genes affecting quantitative traits can be divided into two categories: analysis of candidate
genes, and genome scans based on within-family genetic linkage. To date, 12 patent cooperative treaty (PCT) and US
patents have been registered for DNA sequences claimed to be associated with effects on economic traits in dairy cattle.
All claim effects on milk production, but other traits are also included in some of the claims. Most of the sequences found
by the candidate gene approach are of dubious validity, and have been repeated in only very few independent studies. The
two missense mutations on chromosomes 6 and 14 affecting milk concentration derived from genome scans are more
solidly based, but the claims are also disputed. A few PCT in dairy cattle are commercialized as genetic tests where
commercial dairy farmers are the target market.
Keywords: Quantitative trait locus, Quantitative trait nucleotide, Marker-assisted selection, DNA microsatellite, Single
nucleotide polymorphism, Linkage disequilibrium, Genome scan, Genetic markers, Dairy cattle.
INTRODUCTION
Scientific breeding programs for dairy cattle began in the
1950’s. These programs were based on phenotypic selection
without regard to the individual genes responsible for the
phenotypic variation. Many studies have proposed that rates
of genetic gain can be increased by direct selection on the
individual quantitative loci (QTL) responsible for the genetic
variation in these traits, or selection on linked genetic
markers termed marker-assisted selection (MAS) (reviewed
by [1]). Until 1990 QTL detection was limited by the lack of
suitable genetic markers in dairy cattle populations. Since
then, the development of DNA-level genetic markers;
specifically, DNA microsatellites and single nucleotide
polymorphisms (SNP), have made detection of QTL nearly
routine in all major livestock species. The studies that
attempted to detect genes affecting quantitative traits via
linkage to genetic markers can be divided into two
categories: analysis of candidate genes, and genome scans
based on within-family genetic linkage.
The principles of modern breeding programs for dairy
cattle will be described, and their accomplishments and
limitations will be reviewed. Next the basic principles and
the current state of MAS in dairy cattle will be reviewed.
Detection of segregating QTL via the candidate gene
approach, genetic linkage, and methods to determine the
actual polymorphism responsible for the observed effect will
be considered in the following sections. The specific US
patents and PCT in which DNA polymorphisms are claimed
to affect expression of economic traits in dairy cattle then
*Address correspondence to this author at the Institute of Animal Sciences,
A. R. O., The Volcani Center, P. O. Box 6, Bet Dagan 50250, Israel; Tel:
972-8-9484430; Fax: 972-9-9470587; E-mail: weller@agri.huji.ac.il
will be analyzed, and in the last section an attempt will be
made to predict future developments and several open
questions with respect to QTL patents in dairy cattle will be
addressed.
TRAIT-BASED SELECTION, ACCOMPLISHMENTS
AND LIMITATIONS
Modern dairy cattle breeding schemes are based on the
following principles:
1. Nearly all economic traits are expressed only in females.
2. Female fertility is very limited while male fertility via
artificial insemination is nearly unlimited.
3. As compared to most other agricultural species, cattle
breeding programs are constrained by the long generation
interval and the high value of each animal.
Thus unlike the situation in poultry or most plant species,
dairy cattle breeding programs are based on selection within
the commercial population, rather than selection within
specific populations controlled by breeding enterprises. Due
to the biological limitations noted above, most modern dairy
cattle programs are based on the “progeny test” scheme.
Most genetic gain is obtained by selection of the bulls, even
though bulls do not have production records. Genetic
evaluations are derived by the Individual Animal Model,
including the numerator relationship matrix. Thus the genetic
evaluations of animals without records are derived from the
production records of all their female relatives [1].
An example of the Israeli Holstein breeding program is
given in (Fig. 1). This population consists of approximately
120,000 cows of which 90% are milk recorded.
Approximately 20 bulls are used for general service. Each
year about 300 elite cows are selected as bull dams. These
are mated to the two to four best local bulls and an equal