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The status of equine genetic evaluation

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 Proceedings, Western Section, American Society of Animal Science
Vol. 55, 2004
THE STATUS OF EQUINE GENETIC EVALUATION
E. M. Burns, R. M. Enns and D. J. Garrick
Department of Animal Science, Colorado State University, Fort Collins, CO 80523
ABSTRACT: The use of genetic evaluation in the
equine industry is reviewed and potential problems
identified. The U.S. equine industry has been slow in
adopting genetic evaluation as a tool for animal selection,
in comparison to many European breed associations.
Research has shown that many important traits are
moderate to highly heritable indicating that genetic
progress from selection is possible. Heritabilities for:
cutting ability (0.04-0.19), trotting (0.25-0.3),
Thoroughbred race times (0.15-0.2), Quarter Horse race
times (0.34-0.38), jumping (0.19-0.31), and dressage (0.1-
0.2) have been reported in the literature. In Dutch
Warmbloods, the high genetic correlation (0.75) between
performances at different ages implies that selection at an
early age may be effective in making genetic progress at
all ages. Studies in Warmbloods have shown a high
genetic correlation (0.84) between observations at station
performance testing of stallions and competition results of
their offspring. A genetic trend of 3.2% of a phenotypic
standard deviation per year was reported in standardized
accumulated transformed earnings in Norwegian trotters.
The trade of live stallions and semen across countries
allows greater access to superior breeding stock and is an
important part of the economy for many countries.
Several problems exist in the international trade of
stallions and semen: the lack of a common database, the
lack of precisely defined breeding goals, the various
methods of performance testing and breeding value
estimation. Interstallion was created in 1998 to address
these issues by establishing an international database,
harmonizing traits within riding disciples and
standardizing performance testing. A Universal Equine
Life Number has been implemented to link international
performance data. The standardization of performance
testing, reporting of data and breeding values will greatly
enhance the usefulness of genetic evaluations in the
equine industry.
Keywords: Equine, Genetic Evaluation
Introduction
Large-scale genetic evaluation, using Best Linear
Unbiased Prediction (BLUP) methodology, is used by
most livestock industries. The United States equine
industry, unlike many European breed associations, has
been slow to adopt genetic evaluation as a tool for animal
selection. Research has shown that many economically
important traits are moderately to highly heritable
indicating that genetic progress can be made through
selection. Genetic change due to selection is affected by
selection intensity, genetic variation, accuracy of selection
and the generation interval. Progeny testing can increase
82
accuracy but this can lengthen the generation interval
which in horses is between eight and twelve years. The
generation interval can be influenced, as far as genetic
improvement, by how early in life a trait can be measured
(Ström and Philipsson, 1978). Sampling young stallions
and making use of test results for young progeny can
increase accuracy without prolonging the generation
interval. The high genetic correlation (0.75) between
performances at different ages implies that selection on
performance at an early age would be effective in making
genetic progress at all ages (Huizinga and van der Meij,
1989). Bruns (1981) and Meinardus and Bruns (1988)
found repeatabilities of 0.42 and 0.45 for show jumping
and dressage respectively, suggesting that early records
are a good indicator of later performance. In 1986 the
Icelandic Toelter horse became the first breed to officially
publish breeding values estimated using BLUP
procedures (Sigurdsson et al., 1997). The objective of
this paper is to review the use of genetic evaluation in the
equine industry.
Review of Literature
Cutting. In cutting contests, each horse is judged
for two and a half minutes; during this time the rider
guides the horse into a herd of cattle and pushes a small
group of cows until one cow is left. The rider then drops
the reins, allowing the horse to work independent of rider
cues. Usually a horse and rider cut two to three cattle in
the allotted time. Hintz (1980) found that cutting ability
was lowly (0.04) heritable. Later studies estimated higher
heritabilities of 0.19 (Ellersieck et al., 1985) and 0.12
(Willschau, 1994). Cutting success is somewhat heritable
and thus genetic progress can be made when genetically
superior animals are chosen as parents. Genetic
evaluation is currently not used by any cutting horse
associations.
Trotting. Trotting is a form of harness racing
where a horse pulls a driver in a light, two-wheeled sulky.
Trotters move with a diagonal gait, the left front and the
right hind legs and the right front and left hind legs move
in unison. Trotters cannot break into a canter or gallop
during a race.
The following heritabilities have been estimated
for best racing time in trotters: Minkema (1975) 0.36,
Hintz (1980) 0.25 and Saastamoinen and Nylander (1996)
0.27. Árnason et al. (1982) found that the heritability of
best time varied in three different populations and
reported values of 0.36 in Dutch trotters, 0.12 in North-
Swedish trotter and 0.18 in Russian trotters. Ojala and
Van Vleck (1981) found simple correlations between best
and average time for a year were greater than 0.90,
suggesting that there is a very strong relationship between
the two traits. The authors cite similar correlations in the
literature of 0.83 (Linner, 1975), 0.72 (Neisser, 1976) and
0.85 (Katona, 1979) and note that large correlations for
best time with average time in a year imply that best time
is a good measure of average speed in a year.
Two studies have estimated the heritability and
repeatability of time at finish: Ojala and Van Vleck
(1981) 0.30 (r = 0.70) and Thuneberg-Selonen (1999)
0.23-0.28 (r = 0.50-0.57).
Minkema (1975) estimated a heritability of 0.26
for total earnings. Klemetsdal (1994) used Accumulated
Transformed and Standardized Earnings (ATSE) as a
measure of performance in Norwegian Trotters. ATSE
was found to be lowly to moderately heritable (0.14 to
0.22 depending on age). Saastamoinen and Nylander
(1996) found the heritability of earnings in Finnish
Standardbred Trotters to be 0.33 when non-starters were
included and 0.39 when non-starters were excluded.
Thuneberg-Selonen (1999) found that earnings were
lowly heritable (0.05-0.09). These findings suggest that
best time may be a more appropriate measure of
performance as it tends to be more heritable than
earnings. Since 1992, breeding values estimated using
BLUP animal model have been published for Swedish
Trotters.
Thoroughbred Racing. Thoroughbred Racing is
comprised of flat racing and jump racing. Flat racing is
prevalent worldwide, while jump racing is popular in
European countries with relatively few jump races run in
the USA. Jump racing includes national hunt racing
(hurdling), steeplechasing and point-to-point. The
obstacles in national hunt racing are smaller and the
distances are shorter than in steeplechasing. Point-to-
point racing is steeplechasing for amateurs. Several
studies have looked at the genetic component of
performance in Thoroughbred racehorses using timeform
rating, race time, handicap weight and earnings as
measures of performance.
Timeform rating is the merit of a horse expressed
in pounds, other things being equal, the horse with the
highest timeform rating is the most likely to win
(Timeform, 2004). Timeform rating is revised each
season to maintain a comparable mean from year to year
(Field and Cunningham, 1976). The heritability of
timeform rating using regression of timeform rating on
sire has been estimated as 0.74 (More O’Ferrall and
Cunningham, 1974) and 0.76 (Gaffney and Cunningham,
1988). Gaffney and Cunningham (1988) estimated that
the predicted rate of genetic change is 0.92 timeform units
per year.
Cunningham (2002) reports the heritability of
racing based on handicap weight to be 0.30. Handicap
weight is the weight carried by horses in a race. Superior
horses will generally carry more weight than poorer
performing individuals.
Race time has also been used as a measure of
performance in Thoroughbred racehorses. The
heritability of race time was reported as 0.09 to 0.11,
depending on race distance (Moritsu et al., 1994). Park
83
and Lee (1999) estimated a higher heritability of 0.27 to
0.30 for race time.
Watanabe (1974) used number of lengths each
horse finished behind the first horse as a measure of
performance and estimated a heritability of 0.64.
Langlois et al. (1996) estimated the heritability
of log of yearly earnings to 0.28 in flat races and 0.25 in
jump races. Foye et al. (1972) found a repeatability of
0.37 for earnings per start suggesting that an individual
should not be culled based on one racing year.
Race time is an inferior method of measuring
performance as non-runners are excluded and according
to Cunningham’s paradox best time seems to have
reached a genetic plateau. Cunningham (1975) estimated
that winning time showed a two percent improvement per
decade up to about 1900 but has remained stagnate since
then despite intense selection, this is known as
Cunningham’s paradox. Earnings are also a poor measure
of performance as they are lowly heritable and do not
account for the level of race. Tavernier (1991) and
Langlois (1996) recommend using rank as it accounts for
level of competition, number of horses etc.
Quarter Horse Racing. Few studies have
examined the genetic component of racing in the
American Quarter Horses (AQH). Buttram et al. (1988)
estimated a heritability of approximately 0.37 for racing
time in AQHs after adjusting for age and sex. Wilson et
al. (1988), using heritabilities and variance components
estimates from Buttram et al. (1988), found an average
rate of genetic change of -0.0067 seconds per year in
racing AQHs.
Arabian Racing. Several studies have estimated
the heritability of earnings: Sobczyńska and Kownacki
(1997) 0.22, Sobczyńska and Lukaszewicz (2002) 0.19
(r = 0.46) and Belhajyahia et al. (2002) 0.09 (r = 0.25).
Sobczyńska and Kownacki (1997) found rank at
finish to be moderately (0.25) heritable. Later studies
reported lower heritabilities: Sobczyńska and
Lukaszewicz (2002) 0.18 (r = 0.45) and Belhajyahia et al.
(2002) 0.12 (r = 0.35).
Earnings and rank have very similar heritabilities
suggesting that either trait could be used as a measure of
performance.
Competition traits. Competition traits include
dressage, show jumping and cross-country jumping.
Dressage is a competitive equestrian sport that uses
classical training comprising of movements that have
been developed over centuries. Show jumping is a sport
where the horse and rider jump a set course of obstacles
usually within a time period. In cross-country jumping
horses and riders jump (at a gallop) a series of obstacles
over varied terrain.
Dressage has been found to be moderately
heritable: Meinardus and Bruns (1988) 0.16, Huizinga
and van der Meij (1989) 0.10 and Koenen et al. (1995)
0.17. Wallin et al. (2003) estimated the heritability of
cumulative placing in dressage as 0.16.
The heritability of jumping has been estimated as
0.18 (Meinardus and Bruns, 1988), 0.31 (Huizinga et al.,
1991) and 0.19 (Koenen et al., 1995). Wallin et al. (2003)
found a heritability of 0.27 for cumulative placing in
jumping.
Gerber Olsson et al. (2000) citing Árnason et al.
(1997) noted that the genetic trend, based on breeding
values estimated from a BLUP analysis, increased rapidly
from the late 1980s to the late 1990s. Gerber Olsson et al.
(2000) found an annual genetic progress for gaits and
jumping of 0.05 and 0.03 genetic standard deviations per
year respectively for Swedish Warmbloods born between
1988 and 1992.
Performance testing is carried out by many breed
associations and can last up to 100 days; animals are
trained and judged under uniform conditions that
minimize the effects of rider and pre-training (Huizinga et
al., 1991). Station performance testing of progeny allows
for the estimation of breeding values on stallions three to
four years earlier than from progeny competition results
(Christmann, 1995). Station performance testing seems to
be an appropriate measure as it is highly heritable. Ricard
et al. (2000) estimated heritabilities of 0.40 to 0.60 for
station performance testing observations. High genetic
correlations between station performance testing and
competition data also suggest that station observations are
appropriate measures of performance. Ricard et al.
(2000) found a genetic correlation of 0.70 to 0.90 for
station performance testing observations and competition
data. Wallin et al. (2003) found a genetic correlation
between performance testing in the Swedish Riding Horse
Quality Test and in show jumping and dressage to be 0.83
to 0.93 and 0.63 to 0.75, respectively. These high
correlations suggest that there is a strong positive
relationship between station performance testing
observations and later competition results.
Interstallion was formed by the European
Association for Animal Production, the World Breeding
Federation of Sport Horses and the International
Committee for Animal Recording in 1998 (Koenen and
Aldridge, 2002). The aims of Interstallion are to describe
breeding values, testing procedures and genetic evaluation
methods of Warmblood breeding organizations so that
genetic evaluations can be compared across countries. A
major difficulty in this has been the identification of
horses across breeds and countries. A Universal Equine
Life Number has been implemented which can be used
alongside the studbook of birth number.
Longevity. Numerous studies have examined
factors that affect career longevity or length of productive
live in equine athletes. These factors include
conformation, training, environmental conditions, type of
competition, age and sex.
Wallin et al. (2001) found that orthopedic scores
had the greatest influence on longevity. Horses that
scored a nine or ten were half as likely to be culled as
those that scored a six or below. Poor conformation
predisposes a horse to injury. Rossdale et al. (1985)
found that lameness is the most significant cause of loss
in young racehorses. Wallin et al. (2000) cite several
studies that have found that musculoskeletal injury is the
major cause of culling in performance horses (Clausen et
al., 1990; Heisle, 1995 and Hommerich, 1995).
84
Training has been found to have a significant
impact on the risk of injury and thus career longevity.
Verheyen and Wood (2001) note that 80% of all fractures
in flat racehorses occurred during training. Proudman et
al. (2004) state that training has a significant effect on the
risk of falls and the likelihood that a horse would
complete the race.
Several studies have examined the relationship
between track condition, risk of injury and performance.
Tracks with some moisture seem to reduce the risk of
injury while dry tracks and very wet tracks seem to
increase the risk of injury. Rooney (1983) states that
trace amounts of rainfall protects against lameness.
Bailey et al. (1997a) and Bailey et al. (1997b) found that
tracks with lower water content were associated with a
greater risk of injury. Proudman et al. (2004) notes that
good-to-soft ground resulted in significantly fewer falls
and increased the likelihood that a horse would complete
the race in comparison to soft ground.
The rates of fatal musculoskeletal injuries
reported in the USA have been considerably higher than
in Europe, which may be a result of differences in track
surface. Racing in the USA is predominantly on dirt
tracks whereas most races occur on turf in Europe.
Mohammed et al. (1991) observed that horses racing on
dirt tracks have a higher risk of serious musculoskeletal
injury compared with horses racing on turf. Buttram et al.
(1988) found that racetrack alone accounted for between
11% and 32% of the variation depending on race distance.
Type of competition has been evaluated as a
potential risk factor associated with longevity of career.
Horses in hurdle races are four times more likely to suffer
a musculoskeletal breakdown, while horses in
steeplechases are eight times more likely to suffer an
injury compared with horses racing on the flat (Bailey et
al., 1998). The presence of barriers likely explains this;
Bourke (1995) found that the majority of jumping
fatalities were associated with a fall.
Age also has a significant impact on career
longevity. High levels of loss occur in the first or second
racing seasons (Mason and Bourke, 1973; Mohammed et
al. 1991; Bourke, 1995). Several studies have found that
the risk of injury increases with age (Bailey 1997;
Robinson et al., 1988; Mohammed et al., 1991) but
starting to compete at an older age has a negative impact
on career duration (Bourke, 1995; Ricard and Fournet-
Hanocq, 1997).
Wallin et al. (2001) notes that males were twice
as likely to be culled as females when studying longevity
in Swedish Warmbloods. Bailey et al. (1999) found that
females were less likely to race and Bourke (1995) found
that males raced on average one season longer than
females. This may be because females tend to win less
money (Minkema, 1975) and males are faster than
females (Leroy et al., 1989; Thuneberg-Selonen et al.,
1999 and Röhe et al., 2001).
Implications
Many important traits are moderately to highly
heritable and thus much genetic progress can be made
through selection. The Danish Warmblood, Irish Sport
Horse, Dutch Warmblood, French Sport Horse, Swedish
Warmblood, Holstein, Hanoverian, Icelandic Toelter,
German Trotter and Swedish Trotter Associations all use
breeding values estimated using BLUP techniques and
have shown an increase in genetic progress since its
implication.
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