Genetics of Performance Traits: A. Ricard, E. Bruns and E.P. Cunningham

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Genetics of Performance
Traits
A. RicardA.
Genetics
15 etRicard
of 1, E.Traits
Performance
al. Bruns2 and E.P. Cunningham3
1Institut National de la Recherche Agronomique, Station de
Génétique Quantitative et Appliquée, 78352 Jouy en Josas, France;

2Institut für Tierzucht und Haustiergenetic, Universität Göttingen,
Albrecht-Thaer-weg 3, 37075 Göttingen, Germany; 3Department of

Genetics, Trinity College, Dublin 2, Ireland
Introduction 411
Thoroughbreds 412
Introduction 412
Heritability of track performance 412
Rate of genetic improvement 414
Limits to performance? 414
Horses for courses 417
Trotters 418
Speed and quality of trotter 418
Earnings 419
Are there still true precocious performances? 422
A new way to measure success and career 423
The problem of non-recorded horses 426
Conclusion 426
Sport Horses 427
Measurements of performance 427
Heritability of performance traits 429
Genetic relationships between performance traits 432
Concluding remarks 434
References 434
Introduction
Horses are used for a much more diverse range of functions than is generally
appreciated. This chapter deals with performance traits of race and sport
horses. The first two sections of the chapter consider performance traits in
two of the most common groups of racehorses, Thoroughbreds and trotters.
©CAB International 2000. The Genetics of the Horse (eds A.T. Bowling and A. Ruvinsky) 411
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412 A. Ricard et al.
Interestingly, there were 114,026 recorded births worldwide in Thoroughbreds

in 1996, against 47,795 in trotters. The third section analyses the performance
traits in sport horses.
Thoroughbreds
Introduction
The selection and breeding process in Thoroughbreds is founded on the belief

that racing performance is inherited. Attempts to analyse the genetics of per-
formance in a systematic way have involved some distinguished names
(Galton, 1898). However, it is only in recent decades that good estimates of
heritability of performance, based on adequate data, have been produced (see
reviews by Hintz, 1980; Langlois, 1980; Tolley et al., 1985; Klemetsdal, 1990).
A specific question concerns the choice of measure of performance. The
simplest and, for many people, the most relevant measure is earnings. How-
ever, earnings are very non-linear, and many horses have no earnings at all.
Racing time is an obvious measure, and is recorded routinely for all horses in
the shorter American and Japanese races. It is a less useful measure in longer
races, and is often recorded only for the winner. Handicap ratings involve a lot
of subjective judgement, but are well tested and generally available for all
horses. All of these measures can be converted into ranks, or subjected to a
variety of transformations. These issues are reviewed by Ricard (1998).
Heritability of track performance
The most comprehensive assembly of published results on heritability of rac-

ing performance is that of Tolley et al. (1985). They summarized results from
over 40 studies, about half of which related to analyses in Thoroughbreds, and
the remainder to trotters, pacers and Standardbreds. For the Thoroughbred
studies, they presented the results separately for earnings, handicap and racing
time measures. As might be expected, the reported estimates vary depending
on the analytical model used, the scale and quality of the data, and the
particular measure of performance involved. For the five studies reporting
heritabilities of earnings (generally logs) or normalized ranks, the values
ranged from 0.23 to 0.56 (if we focus on 3-year-olds and take account of all
races), with most values being in the lower end of this range.
Eight studies reported heritabilities for handicap measures. In these
studies, there was more variability in analytical method, and a general
tendency for paternal half-sib and regression on sire methods to give very
high estimates. Most authors discounted these on the basis of expected, but
unquantifiable, correlations between environment of offspring and phenotype
of sire. The remaining estimates (mainly regression on dam or on midparent)
fell in the range 0.24–0.61, with most grouped in the range 0.30–0.40.
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Genetics of Performance Traits 413
Ten studies were given in which heritability of racing time was estimated.
Several of the estimates were based on quite small data sets. For those with
reasonable amounts of data, estimates ranged from 0.09 to 0.78, but most
values were in the range 0.10–0.20.
Since the review of Tolley et al. (1985), a number of studies with large
volumes of data have been completed. Gaffney and Cunningham (1988)
published estimates of heritability of Timeform (handicap) ratings based on
full year 3-year-old ratings of 31,263 horses from 2087 sires. The estimates
ranged from 0.39 ± 0.01 (regression of offspring on dam) to 0.76 ± 0.02
(regression of offspring on sire). As in earlier analyses of similar data, the
authors concluded that sire-based estimates were likely to be biased upwards.
Oki et al. (1995) published estimates of heritability of racing time for
various race lengths, based on performances of over 25,000 horses. The data
were approximately equally recorded on dirt and turf. The values on dirt and
turf were broadly similar, but there was a clear tendency for heritability values
to be lower as the length of race increased over the range 1000–2000 m (see
Table 15.1). In keeping with the earlier results summarized by Tolley et al.
(1985), the heritability estimates for racing time in this study were lower than
for the other measures of performance reported by other authors.
Williamson and Beilharz (1998) carried out an analysis of almost 0.5
million individual race records from Australia. They used three measures:
lengths (distances between horses at the end of a race), ranks and a function
of earnings. By an approximately linear adjustment for length of race, they
derived a speed rating and a stamina rating for each horse. They then
calculated heritabilities for all of these measures. The resulting estimates were
generally higher than those from other studies. Most estimates were above 0.5,
with values for the stamina rating being consistently higher than those for
speed (approximate average of 0.65 against 0.49).
Langlois (1996) has discussed the difficulties in obtaining reliable
heritability estimates in Thoroughbreds. He concluded that racing speed is not
a useful criterion because it has low heritability and is not, on its own, a major
determinant of success. He favoured ranks. He also presented results showing
a consistently higher heritability of log earnings calculated by regression of
offspring on dam than by regression on sire. This effect was attributed to
non-genetic maternal contributions to the offspring, as well as to the lower
selection and, therefore, greater genetic variability in female parents.
The overall picture emerging from these numerous studies is a reasonably
consistent one. It shows that the heritability depends on the measure used.
Table 15.1. Heritability of racing time for races of different lengths (Oki et al., 1995).
Race length (m)
Type of track 1000 1200 1400 1600 1800 2000
Turf 0.25 0.16 0.10 0.12 0.09 0.08

Dirt 0.19 0.22 0.12 0.09 0.17
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Time measures generally had a heritability in the region 0.1–0.2, with the
higher values for shorter races. For handicap and earnings measures, reported
heritabilities were generally higher, frequently in the range of 0.3–0.4.
Rate of genetic improvement
The rate of genetic improvement in a population under selection can be

predicted if the following four factors are known: (i) genetic variance for the
trait under selection; (ii) its heritability; (iii) the intensity of selection for this
criterion; and (iv) the generation interval in both males and females. Genetic
change can also be measured retrospectively by appropriate analyses.
Gaffney and Cunningham (1988) predicted the rate of genetic gain in the
British Thoroughbred for Timeform rating at 0.92 Timeform units per year.
This was based on generation intervals of 11.2 and 9.7 years, and selection
intensities of 6 and 52% in males and females, respectively, and a heritability
value of 0.36. They then estimated the rate of genetic change by a sire model
best linear unbiased prediction (BLUP) analysis involving a total of 516 stal-
lions born in the years 1952–1977, and performances of their 11,328 progeny
between 1961 and 1985. The estimated change averaged 0.94 ± 0.13 Timeform
units per year.
This result is paralleled by the results of a large analysis of Quarter Horse
data from the USA (Buttram et al., 1988). Quarter Horse races are run over
much shorter distances, most commonly 320 m, with a mean finishing time of
18.7 s. From an analysis of finishing time data on over 1 million racing records,
they estimated genetic trends in the population between 1960 and 1983. They
found a consistent improvement, averaging 0.47, 0.43 and 0.16% per year,
respectively, for the distances 320, 366 and 402 m. This genetic trend
accounted for one-third of total improvement in finishing times over the
period. In contrast to these studies, Preisinger et al. (1990) and Chico (1994)
were unable to demonstrate genetic change in the German and Spanish Thor-
oughbred populations.
Limits to performance?
In contrast to the results showing steady genetic improvement in Timeform

ratings, Gaffney and Cunningham (1988) observed that winning times of
Classic races have not been improving for a long time. In fact, the three
English Classics have improving winning times from the 1840s onwards to
about 1910, and from then seem to have been static (Fig. 15.1).
If steady genetic improvement is being achieved, how can we interpret the
static winning times in Classic races? James (1990) has argued that the herita-
bility values, and hence the calculated improvement, are overestimated. One
possibility is that there is some kind of physiological ceiling to performance,
particularly in longer races. There is some support for this idea in the fact that
134
Fig. 15.1. Trends in winning times for the English Classics. Each point is an average of 10
years.
winning times are continuing to improve in the shorter races, common in USA,
while the plateau is most evident in longer races such as the St Leger
(Fig. 15.1). Thus, the physiological limit might come into play only beyond a
certain threshold of effort.
If this hypothesis is true, then it should be possible to identify the physio-
logical elements which set a limit to performance. The probable limiting
factors will depend on the length of the race. The energy required for muscle
contraction is derived from glucose (or its storage form glycogen). Energy
release from glucose comes in two stages. The first, anaerobic, stage releases
less than 10% of the energy available and produces lactic acid which
accumulates in muscle tissue. The second, aerobic, stage requires oxygen, and
releases the energy stored in glycogen. The oxygen supply has the additional
function of removing lactic acid. In theory, either of these factors – the
ultilization of oxygen and glycogen for aerobic metabolism during prolonged
events, and the production and clearance of lactic acid following anaerobic
metabolism during short-term events – could be limiting factors.
In every race, both kinds of muscle metabolism are involved. Although the
flow of blood to the muscles, and hence the supply of oxygen, increases dra-
matically during exercise, maximum effort could not be sustained by aerobic
contraction alone. Anaerobic contraction may be limited by glucose availability
and lactic acid accumulation. The balance of these two sources of energy sup-
ply to muscular activity depends on many factors, and particularly on the
amount of activity involved, i.e. the length of the race. The 30 s sprint-type
races of American Quarter Horses are mainly anaerobic. The 2.5 min required
to run the English Classics shifts the balance to the point where most of the
energy supply is aerobic. This difference in the energy supply route may
135
indeed be part of the explanation for the apparent limits to performance for
the longer races, while records continue to be broken in the sprints.
The physiology of blood circulation and exercise in horses has been
reviewed by Fregin and Thomas (1983). Figures 15.2 and 15.3 present two
important sets of data from their report. They measured a number of metabolic
indicators as the work effort was increased in a number of horses. With
increasing effort, heart rate increased steadily from a resting figure of about 40
beats min−1 to a maximal rate close to 200. At the same time, blood volume per
beat increased, also linearly. The result is a dramatic linear increase in blood
circulation with increasing effort. At full stretch, a 500 kg horse is pumping
250 l of blood through their system each minute, equivalent to ten times his
blood volume (Fig. 15.2). The important point is that blood circulation and
oxygen uptake increased linearly over the range.
Figure 15.3 shows blood lactic acid levels over the same range of effort.
Here, it is evident that lactic acid clearance is not keeping pace with increasing
effort, and is indeed becoming critical at maximum effort. What this suggests is
that the limiting factor in performance may well be lactic acid accumulation.
These conclusions are supported by the results of Evans et al. (1993),
who found a strong relationship between blood lactate concentration after a
strenuous treadmill test, and Timeform rating, in 14 Thoroughbred horses. The
area is reviewed by Ronéus (1996) who also found a significant correlation
between plasma lactate levels and speed over 1600 metres in Standardbred
trotters.
Fig. 15.2. Cardiac output (Q) and heart rate (HR) at rest and during a five stage treadmill exer-
cise test on an 11.5% gradient in five sedentary horses (redrawn from Fregin and Thomas, 1983).
136
Fig. 15.3. Relationship between lactic acid concentration and work effort (redrawn from Fregin
and Thomas, 1983).
Horses for courses
A recent study from Japan (Oki et al., 1997) adds weight to the view that the
genetics of performance depend very much on the length of the race. They
calculated genetic correlations between performances at different race lengths,
on both dirt and turf, from over 20,000 performance records. Race lengths
varied from 1000 to 2000 m. As the difference in the race lengths increased, the
genetic correlation decreased. At a difference of 400 m, the correlation was
0.91; at 600 m difference it was 0.78, and at 800 m the correlation was 0.68.
This pattern was very much the same on turf and dirt tracks. The genetic corre-
lations between performance on dirt and turf are even lower, and depended
on the length of the race. At 1200 m, the correlation was 0.69, while at 1800 m
it was 0.31.
Thus if a horse is evaluated on dirt, the effectiveness of selection for
performance on turf is about half what it would be if comparable data from its
turf performances had been used. This corresponds to an average correlation
between performance in these two environments of 51%. The loss of effective-
ness is not so great at short distances as it is at long distances: at 1200 m the
reduction is only 31%, while at 1800 m, it is 69%.
Within any racing environment, turf or dirt, selection at one length for
performance at another becomes progressively less effective as the distance
between the two lengths increases. If the difference between the two lengths is
200 m, about 5% of effectiveness is lost. At 400 m, this is about 9%, at 600 m
difference it is 22%, and at 800 m difference the loss in effectiveness is 32%.
137
However, the effect of different lengths is in general much smaller than the
effect of turf versus dirt.
These results indicate that, to a considerable degree, different genetic
factors are involved in performance in longer or shorter races and on different
kinds of track. They could have serious implications for the evaluation of
animals based on American data (short races, dirt tracks) for performance in
European (longer races, turf tracks) races. It means that new weight (and dis-
count) should be applied to data generated in a different racing environment.
The net conclusion is to reinforce the old notion of ‘horses for courses’.
Trotters
Performance comparison in trotters deals with the general problem of
measuring performance in races with additional complexity due to the type of
exercise and breed. Performances in trotters, in contrast to Thoroughbreds, are
characterized by qualifying tests before entering races; greater longevity in
their racing career; relatively shorter term of selection for trotting speed (since
early this century); and the inclusion of more than one breed, i.e. Standard-
bred, French trotter and Nordic local breeds. Firstly, classical measures (speed
and earnings) will be analysed and then new methodology introduced, to give
a better evaluation of performance. Performance for non-recorded horses will
be considered, and the general trend to a precocious racer will be discussed.
Speed and quality of trotter
A natural choice to measure a performance trait for a racing horse is obviously

to measure his ability to run fast. Speed is a traditional measure used as a refer-
ence for breeding. Results are summarized by time for a given distance (often
1 km) and then by best time for this distance (in the year or the lifetime). For
this kind of record, it is important to take into account variability due to racing
distance (often under 2000 m, near 2000 m and >2000 m), kind of start (flying
start, volt-start, others) and eventually track type and racecourse. The use of
repeated times or best times leads to different strategies. Heritability of such
a trait is moderate and homogenous across country, with few exceptions. In
Finland, Saastamoinen and Nylander (1996a) estimated heritabilities for best
time in career at 0.28 for Standardbred and 0.08 for Finnhorse. Heritability of
time in particular races, i.e. first qualifying, passed qualifying and first race, is
higher (from 0.36 to 0.53 at 3 years old) but perhaps overestimated because it
does not take racing conditions into account. In Belgium, Leroy et al. (1989)
found a heritability of 0.11 for best time in life for a mixed trotter population
(European and American origin). In France, Langlois (1984) found a herita-
bility of between 0.26 and 0.43 for the best time at different ages (2–6 years
old). In Germany, Katona and Distl (1989) found values of 0.35 for average
time and 0.50 for average time performed at 2–3 years old. In Norway,
138
Klemetsdal (1989) found a value of 0.03 for repeated racing time for Standard-
bred and 0.18 for Norwegian trotter. In Arnason et al. (1989), heritability for
best racing time ranges from 0.45 to 0.26 according to the limit age of
the record (3–13 years old); a reasonable mean seems to be 0.29 (Swedish
Standardbred) and 0.25 (North Swedish trotter). In Italy, for repeated annual
best time (up to 6 years old), Silvestrelli et al. (1995) found a value of 0.37. No
clear rules exist regarding over- or underestimation of heritability when
models do not take into account racing conditions or when best record is used
instead of all recording times. According to recent selection for speed in trot,
this trait remains heritable and genetic increase is observed in most countries.
There is no doubt of the possibility to select for this trait, but the search for
other traits to record performances nevertheless is necessary in order to report
the complete aptitude of a horse to win a race. Speed is only one quality of a
good racehorse, its capacity to adapt to race conditions and to win at a certain
pace is the complete goal.
Earnings
At the end of a race, money is allocated to the horses according to their

ranking in the race. Most recent authors measure performances in trotter
horses by their earnings. The choice of the measure linked to earnings reveals
different aspects of the aptitude of the horse. Differences are principally in the
mathematical transformations of earnings to reach a normal distribution in the
population and in the choice between cumulated trait (year, life) or repeated
trait per race, including the method to take into account the number of starts.
As pointed out by Langlois (1975), a transformation is necessary in most cases.
For the betting, or simply for the spirit of the exhibition, riders must have the
will to win. Thus, earnings are distributed exponentially as a function of the
ranks in a course or an event: a rider would only really want to be first if
the difference in prize money is significant between the winner and the second
placed horse. This raw performance scale is attractive for competition but
inadequate for genetic evaluation because it emphasizes the differences in
quality between ranked horses. Transformations are now commonly used and
give a reasonable normal distribution of the trait.
Earnings = a exp (b rank) ⇒ Log (earnings) = Log (a) + b rank
The following are only the most recent references available to see the gen-
eralization of this concept: in Italy, Silvestrelli et al. (1995) used log10 (annual
earning + 1/number of starts); in Finland, Saastamoinen and Nylander (1996b)
used (annual earnings)1/4; in Norway, Klemetsdal (1994) used (cumulated
earnings)0.20 standardized within birth year; in Sweden, Arnason et al. (1982)
used (lifetime earnings)1/2, (lifetime earnings)1/4, earnin s n starts ,
Log10 earnin s n starts 1 . According to the initial distribution of earn-
ings in the population, due to rules of distribution of money in one event and
139
Fig. 15.4. Distribution of measures based on earnings – example of French trotters in 1994.
the programme of endowment of races, these different transformations lead to

a more or less normal distribution in each country (Fig. 15.4)
Money is distributed in every race, but the quality of a horse must be mea-
sured on its entire career. It may be appreciated as a repetition of each race or
as the cumulated success over a year or a life. Choice of a unit of measurement
depends on the objectives of the measure but also on the recorded results.
When you use a cumulated performance over a year or a lifetime, this is a
combination of regularity, longevity and level of races won which is included
in the measure. If only placed races (as repeated earnings without non-placed
results or earnings divided by number of places) are used, the maximum level
of the horse is taken into account but not its capacity to repeat this maximum.
If cumulated earnings divided by starts are used with no regression on the
number of starts, the mean level where a horse may be placed, and the agree-
ment between the race programme proposed for the horse and its true quality
are estimated but the results are not weighted with the number of tests and so
give an advantage to horses with few races. Whatever these differences, corre-
lations between these measures are very high, especially genetic correlations.
For example, Arnason et al. (1989) found genetic correlation between total
earnings and total earnings divided by the number of starts of 0.95 (5 years
140
old), and Langlois (1984) found genetic correlations of 0.92–0.98 between

annual earnings and annual earnings divided by number of starts according to
age (2–6 years old). However, high genetic correlations do not take into
account a few special cases which may be of great importance when selecting
stallions on their own performance. Thus, it is better to distinguish the differ-
ent traits which are responsible for a complete successful career. We have the
advantage of being able to consider success event by event and then to con-
sider regularity and longevity as two additional traits for a good horse. This is
what will be discussed in the next section about future methods of evaluation.
In many cases, heritability of ‘earnings’ is moderate and is sufficient to
facilitate selection. A summary of these heritabilities for Standardbred, French
trotter and Nordic trotters is given in Table 15.2. Correlations between earnings
and times are negative and high, and so are favourable (Table 15.3). Therefore,
Table 15.2. Heritability of criteria based on earnings.
Breed/Age Reference Trait related to earnings Heritability
French trotter 1 Log(annual earnings/starts) 0.26

Standardbred 2 Total earnings 0.21
Standardbred up to 3 years 2 Total earnings 0.22
Dutch trotter 2–8 years 3 Total earnings 0.20/0.36
3 (Total earnings)1/2 0.23/0.40
Standardbred 2–6 years 4 (Total earnings)0.20 0.14/0.22
Norwegian trotter 3–6 years 5 (Total earnings)0.20 0.15/0.25
Standardbred 3–6 years 6 (Total earnings)1/4 0.22/0.36
6 Log10(earnings1/2/starts + 1) 0.29/0.38
North Swedish trotter 6 (Total earnings)1/4 0.28
6 Log10(earnings1/2/starts + 1) 0.19
Standardbred 7 Log10(annual earning + 1/starts) 0.10
Standardbred 8 (Total earnings)1/4 0.36
Finnhorse trotter 8 (Total earnings)1/4 0.17
References: 1, Tavernier (1989); 2, Katona and Distl (1989); 3, Minkema (1989); 4, Klemetsdal
(1993); 5, Klemetsdal (1989); 6, Arnason et al. (1989); 7, Silvestrelli et al. (1995);
8, Saastamoinen and Nylander (1996a).
Table 15.3. Correlation between criteria based on earnings and time.
Breed/age Reference Genetic correlation Phenotypic correlation
French trotter 3–6 years 1 −0.86/−0.95 −0.59/−0.69

Norwegian trotter 3–6 years 2 −0.93/−1.00 −0.81/−0.89
Standardbred 3 −0.88/−0.91 −0.37/−0.46
North Swedish 3 −0.93 −0.68
Standardbred 4 −0.98 −0.81
Finnhorse trotter 4 −0.98 −0.88
References: 1, Langlois (1984); 2, Klemetsdal (1989); 3, Arnason et al. (1989); 4, Saastamoinen
and Nylander (1996b).
141
selection based on times and earnings is quite effective. A multiple trait

approach avoids potential biases of one particular measure, even if the
objective of all traits is much the same. It seems to be the better choice, as
proposed initially by Arnason et al. (1982).
Are there still true precocious performances?
In the preceding, performance is supposed to play the same role throughout

the career of the horse. According to the race programme and the strategy
of the race business, the question of which breed and which type of trotter
is important. Local breeds are often preserved by a particular programme
of races as they are often slower and less precocious. However, the tendency
to precocity is very often a tool for rapid economic profitability. From a
genetic viewpoint, it is also obligatory information in order to select early
even for a mature horse. Therefore, several authors studied the capacity of
early performances and the relationship between early performance and
mature performances.
Early performances
Age at first start in trotters was analysed by Saastamoinen and Nylander (1996a,
b). Heritability estimates for age at first qualifying start, passed qualifying start
and first race were low (0.04–0.16) and lower for the local breed (Finnhorse
trotter) than for the Standardbred. In the absence of other results, these
heritability estimates suggest that early start of a career depends mainly on
environment.
Relationship to mature performances

Normally, only results with a correct model, i.e. taking into account the
influence of selection between ages in the estimation of parameters, should be
reported. This requires the use of a methodology which includes horses with
only one performance in the analysis of correlation. However, this methodol-
ogy is too recent for comparison with old studies on trotters and so results may
be biased. No general trend is found regarding the level of heritability of early
performances compared with mature performances. A recent estimation with a
complete model for annual earning per start for French trotters showed a
rather similar heritability from 2 year olds to 4 year olds: 0.24, 0.26 and 0.27,
respectively (A. Ricard, unpublished). Arnason et al. (1989) and Saastamoinen
and Ojala (1991) found a decrease of heritability with age for Standardbred:
from 0.38 to 0.29 for earnings and from 0.45 to 0.34 for best time criteria (3 and
4 years old in contrast to cumulated before 6 years old) in Arnason et al. and
from 0.38 to 0.19 for annual earnings and from 0.52 to 0.26 for best time (from
3 to 5 years old) in Saastamoinen and Ojala. The decrease is accentuated for
time criteria. For Norwegian trotters, Klemetsdal and Stubsjoen (1996) found
the opposite result, with an increase of heritability for cumulated results from
0.14 to 0.22. It is interesting than the most precocious breed (Standardbred)
142
still expresses higher heritability for early performance. Genetic correlations

between early and mature performances were always very high, particularly
for cumulated performance, partially due to autocorrelation of the same trait.
Correlations between different ages were estimated in Sweden by Arnason
et al. (1989). They were found to be high for performances measured as
earnings or earnings per start (0.76–1) between horses of 3, 4 and 5 years old.
In Norway, for cumulative earnings in Norwegian trotters, genetic correlations
were very high (0.97–0.99) between 3–6 years old (Klemetsdal, 1994). For
age-based trait analysis, in French trotters these correlations remain high: 0.89
between 2 and 3 years old, 0.96 between 3 and 4 years old and 0.77 between
2 and 4 years old.
Actually it is difficult to model a general explanation of early perfor-
mances. If entrance in races seems to depend on environmental factors (as cri-
teria based on qualifying races), the success at such an age is already heritable.
Early performance seems to be a good predictor for selection for a mature
horse, with little specificity for very early ones (genetic correlations near 0.80).
A new way to measure success and career
In future, measuring performance for trotters must be able to describe what

happens in one event and then model the repetition of this trait over life with
more subtlety than these global measures. Some ways have begun to be
investigated.
Underlying performance
To measure one performance in one race, the problem is to evaluate the level
of the race and the level of one rank in this race. With regard to earnings, the
measure of the level of the race is supposed to be the total endowment, and
level of one rank is a linear function of it (after transformations). However, the
level of the race is only the level of competitors because the only reason a
horse is ranked at a given place is because the horse ranked before has beaten
it and it has beaten the horse ranked behind. The level of a rank cannot be a
linear scale independent of the number of starters; this is only the difference
between one performance of a horse and those of the others.
To explain the final ranking of a race, without introducing any subjectivity,
let us imagine that all horses in the race performed a physical effort. This effort
may be designed by a classical model with a normal distribution and expecta-
tion based on the influence of environmental and gene effects. The apparent
result of the race, the ranks, is the expression of the hierarchy between these
efforts. Thus, to estimate the effects of the model, inference will be based on
the probability of obtaining the ranking of underlying performances instead of
the probability of obtaining a measured performance. Figure 15.5 illustrates a
race with three horses (A, B and C) with three different expectations. Table
15.4 contains the probability of each possible result of the race, i.e. different
ranking, according the difference of the mean value of the horses.
143
Fig. 15.5. Example of a race with three horses – underlying performance and probability of
ranking.
Table 15.4. Probability of ranking for the sample race (deviation between
mean value of performance of A and B: 1.2, B and C: 0.6 in unit deviation).
Ranking Probability (%)
A–B–C 49
A–C–B 27
B–A–C 15
B–C–A 3
C–A–B 5
C–B–A 2
The likelihood of ranking may be written as:

+∞ +∞ +∞ +∞ n
P ( y (1) > y ( 2) >K> y (t ) >K> y ( n )) =∫ ∫K ∫ K∫ ∏ ϕ ( y (i ) −µ (i ) )dy (i )
−∞ y ( n ) y (t + 1) y (1 ) i = 1
with (1).. (n) the ranks of horses in the event, y(t) the underlying performance
of the horses ranked t, ϕ the normal density, µ(i) the location parameter of the
horse ranked (i) (equal to environmental and gene effects).
This model is actually applied for jumping, dressage and 3-day eventing
in France. It will also be applied in the next few years in trotters and has
already been tested on a single year file (Tavernier, 1994b). The file contains
9228 races run in 1989 by 13,065 horses and represents 129,379 starts. The
model includes sex, age, distance of backing (0, 25 m, 50 m) and optionally
trainer effect. The repeatability was estimated to be 0.26. The validity of the
model was measured by the posterior probability of races calculated with the
144
estimations of the effects. The probability of the races was multiplied by 11 on

average by comparing it with the probability of each possible combination
when all horses are equal. Only 1% of race results are unexpected: the proba-
bility of the ranking calculated from estimations is smaller than the probability
when horses are equal. More than 12% of races have their equal probability
multiplied by 20. A good presentation of results is the computing of elemen-
tary results for match between two horses. A horse estimated at 1 SD of the
distribution of evaluations on this file has a probability of 74% of winning
against a horse estimated at −1 SD. Between the better and the worse horse,
the probability that the former one wins is 99%. This view of the result of a
race has an attractive aspect for horsemen as it corresponds to traditional use.
Trainer effect
All cumulated measures are unable to take into account a trainer or driver
effect. This effect also is often missing from time records. With this new mea-
sure of performance, it is possible to add this type of major environmental
effect. With the previous French file, a model with and without a trainer effect
was tested. A total of 529 different trainers were identified, with grouping of
trainers with less than ten horses. The variance of horse effect was reduced but
the validity of the model was better: the posterior probability of each race is
multiplied by 17 comparing it with random instead of 11 with no trainer effect.
The correlation between horse effect in the model without a trainer effect
and that with a trainer effect is 0.88. The correlation between trainer effect and
horse effect in the model with a trainer effect is −0.05; without a trainer effect it
is 0.38. Here again, the new method seems to be more adaptable to the true
situation than other measures.
Random regression
The model of underlying variables solves the problem of measuring success in
one race. The possibility remains to measure the evolution of such a perfor-
mance during life. A repeatability model suggests a constant mean and only a
residual variation throughout life around it. A new development will be the
design of underlying performance with time. Random regression models,
which explain performance as a function of time, provide new methods in
genetics. Unlike usual production (lactation or growth), there are no reference
curves for the natural career. Thus different regressions must be tested with
different visions of time. Time may be appreciated as starts or days, days from
birthday or from the first race or days between races (rate of use). No applica-
tions are yet available for trotters but work has begun in jumping horses with
an example model as follows:
y i =bi 1 + bi 1 t + ai 0 + ai 1 t + pi 0 + pi 1 t + ei
with yijk, underlying performance responsible for ranks; bi, environmental

fixed effects i; aij, genetic effect for the horse j; pij, permanent environmental
145
effect common to all performances of the horse; eijk, residual. The degree
0 refers to an effect independent of time; 1 refers to regression terms as a
function of time t.
The problem of non-recorded horses
The preceding discussions suggest that all horses have some available perfor-
mances. In fact, there are two possible scenarios: some horses never come on
to a racecourse and some would never be ranked or earn some money. In the
second case – the starter but non-winner horses – different strategies have
been used, depending on the trait. When log (earning) is used (by year or
event), the usual choice is to discard starts without earnings (Langlois, 1984).
Some authors add 1 to calculate the logarithm (Silvestrelli et al., 1995). Cutting
off the data introduces selection and therefore cannot be recommended.
Applying the same value to non-winner horses (Arnason et al., 1989; Katona
and Distl, 1989; Minkema, 1989; Saastamoinen and Nylander, 1996b) without
any reference to the level of the race is also a mistake. The strategy of the new
model based on ranks makes it possible to take into account unplaced
horses as they were beaten by the last-placed horse. Therefore, the value of
unplaced horses is modulated at each event by the value of horses which
participated into the event. This solves the problem of identical value. For the
first case – horses which never enter a race – no information is available about
this lack of data: they may be very good horses which have been exported
to another country or perhaps they are just poor trotters. Klemetsdal (1992)
simulated a population which would be pre-selected before entering a trotter
race on a hypothetical trait correlated with future earnings. He proved that
genetic trend was underestimated when ignoring this fact and that applying
a zero to non-starters is a better way than cut-off to estimate genetic trend.
Arnason (1996) suggested a multiple trait approach with a variable defining the
status of the horse (starter or not) and proved that it is an easy and good
method to avoid biases in genetic progress and an aid to limit inbreeding.
Conclusion
Performance in trotters is actually recorded as best times per kilometre and

transformed earnings. These traits express heritability (0.20–0.40) adapted to
an effective selection. Breeding evaluations from these records commonly are
used in most European countries. New developments will be able to improve
the estimation of the true level of each race, the evolution of performance with
age, time and the rate of use, and the adequacy of genetic models by adding
better environmental components such as trainers and drivers. The ideal is to
obtain a horse economically better adapted to race and, according to race
programmes, with a better precocity or longevity.
146
Sport Horses
Measurements of performance
In sport horses, breeding objectives consist of a number of different variables

measured at different ages of horses. The ultimate goal is to improve dressage
and jumping ability of horses in competitions. The overall breeding objective is
a combination of performance traits relating to horses’ dressage and jumping
ability which are measured at different times and in various testing schemes.
For breeding horses which are used primarily as leisure horses, the definition
of the overall breeding objective leads to different weighting of performance
variables, such as giving more weight to non-physical variables such as charac-
ter and temperament.
For sport horses, performance traits are measured in various systems of
performance testing which can be differentiated due to test location, age and
sex of horses. In some European countries, young stallions and mares, possi-
ble candidates for breeding, are being tested either at station with a common
training period (stallions) from 30 to 100 days or in 1-day field tests (mares)
without any common training. Horses participating in those performance tests
are pre-selected by their owners or breeding organizations, e.g. stallions com-
ing to licensing or thereafter to performance testing at station are at least the
better quarter of each annual group whereas in mares the pre-selection is less
intensive. Stallions and mares having passed the performance tests normally
go for breeding, but some, as well as geldings, participate in dressage or
jumping competitions which are grouped according to quality level and age of
horses. Horses participating in competitions start only if they have a real
chance of winning, whereby this type of selection becomes stronger for older
horses and for high-level competitions. In some countries, station or field tests
are obligatory, in others no station or field tests exist and participating in com-
petitions is the only requirement for breeding stock horses. In general, the sys-
tem of performance testing implemented in each country affects the number
and quality of horses participating in the tests and finally the distribution and
the estimates of genetic parameters of variables recorded in the performance
tests.
Performance testing at station and in field tests

Most variables recorded in station or field tests refer to conformation, basic
gaits, riding and jumping ability, and their scores given by one or more judges
vary between 0 and 10. Licensing of young stallions at an age of 2.5 years is
based mainly on scoring conformation variables, gaits judged in hand and
jumping ability as scored in free jumping. Similar measurements are taken
during registering young mares, excluding jumping ability. Performance test-
ing of 3-year-old stallions and mares focuses on variables scored under saddle.
Riding and jumping ability are measured and again basic gaits are scored.
For stallions participating in the test at station, variables of character and
temperament are judged. The basic problem with most variables is the limited
147
degree of objectivity due to the subjective scoring. Introducing a higher degree

of repeatability and objectivity is achieved through the use of several judges
and test riders (e.g. for scoring riding ability) for one horse, changing the rider
during the common training, and measuring performance without any rider
(e.g. free jumping). Objective measurements of gaits such as length of steps
and jumps are rarely used. The distribution of scores tends to follow the
normal distribution with a mean between 6.0 and 8.0 and a standard deviation
of about 1.0 (Table 15.5).
In some countries, e.g. in Switzerland (Hascher, 1999) and The Nether-
lands (Koenen et al., 1995), conformation variables and walking and trotting in
hand are described based on a linear scale. Such traits show a more centred
distribution with a mean and standard deviation close to the values expected
from the scale used (Table 15.5).
Performance testing in competitions

In competitions, performance of sport horses is measured by their earnings or
points allocated to their ranks in events. Some countries record all the horses
started in competitions, some others record on average only the top 30% of
horses ranked. The measurement used for later analyses is either the earning
or rank of each horse started or being placed in an event or the annually
cumulated earnings or points or a derivative of the annual sum (Table 15.6). In
most cases, earnings show a skewed distribution, with few very high earnings
for horses ranked first or second. Transformations are now commonly used
and give a reasonable normal distribution of the trait and equalize variances
between events. Table 15.6 shows the most recent references for transforma-
tions of performance data in sport horses.
When measuring the performance in competitions, the problem is to eval-
uate the level of the competition or event and the level of a rank in this event
(Ricard, 1998). Several suggestions have been made, such as transforming
ranks by (1 – (place – 1)/number of starters + constant for the level of event)
(Bruns, 1981), (place1/2) (Hassenstein, 1998) or by the normalized rank score
Table 15.5. Selected performance data of sport horses measured in station or field tests.
Test location Variable (scores) Mean Standard deviation
Stallions’ licensinga Walk 6.8 1.8

Trot 7.0 1.4
Stallions’ testb Character 8.0 0.9
Riding ability 6.9 1.1
Jumping ability 7.2 1.1
Mares’ testb Walk, trot, gallop 6.8 0.9
Free jumping 7.1 1.2
Mares’ registeringc Walk (lin.) 6.1 1.2
Trot (lin.) 6.1 1.3
aSchade (1996); bvon Velsen-Zerweck (1999); cHascher (1999).
148
Table 15.6. Transformations of performance data in sport horse competitions.
Variable Transformation Reference
Cumulated points per year in show jumping Log10 Foran et al. (1994)
Highest level during life time in dressage and Square root Huizinga and van der Meij
show jumping (1989)
Annual earnings in show jumping Log10 Silvestrelli et al. (1995)
Earning at each place in dressage and show Ln Meinardus and Bruns
jumping (1989)
(Foran et al., 1995). The level of event depends on the quality of horses started
in one competition. One solution to account for differences in the quality of
competitions is to include competition as a fixed effect in the model describing
the data for estimating genetic parameters. Tavernier (1991, 1994b) proposes
an appropriate interpretation of the rank as the probability of ranks which may
solve the problem of the mean level of each event as well as of the variability
of competitors (Ricard, 1998). Another obstacle in working with competition
data is the availability of data. Some horses never come to a competition, they
are used in breeding or have little chance of winning, and other horses are
never ranked and never earn money. Recording systems which ignore horses
without earnings (Meinardus and Bruns, 1989) introduce selection and biases.
Adding a constant to unplaced horses is recommended by Ricard (1998) if
the level of competition is taken into account. Finally, the quality of the rider
influences the ranking within events as well. The confounding of the genetic
effects of horses with the quality of riders and level of events is a problem in
estimating breeding values of horses which is tackled through the definition of
variables and appropriate linear models.
Heritability of performance traits
Performance testing at station and in field tests

In some European countries, performance testing of sport horses at station or
in 1-day tests has been carried out in a comparable and systematic way over
the last 10–20 years so that genetic analyses were carried out on reasonably
large data sets with thousands of individual horses sired by at least 100
stallions. The methods used for estimating genetic parameters are based on
maximum likelihood techniques applying single or multiple trait animal mod-
els. In Table 15.7, the latest estimates of heritabilities from several countries are
summarized.
The individual estimates of heritabilities are based on data from warm-
blood riding horses in Germany, The Netherlands, Sweden and Switzerland.
The estimates of heritabilities across countries as indicated by the average
heritability allow comparisons between performance testing schemes and
variables within tests:
149
Table 15.7. Heritability estimates of performance traits of young horses tested at station or in
field tests.
Heritability
Test location Variable Average Individual Reference
Stallions’ licensing/ Walk 0.23 0.21, 0.19, 0.21, 0.30 1, 5, 6, 7

Mares’ registering Trot 0.34 0.36, 0.28, 0.38 5, 6, 7
Stallions’ test Character/temperament 0.41 0.41, 0.24a 2, 4
Walk 0.55 0.43, 0.66, 0.33a 2, 4, 9
Trot 0.58 0.50, 0.66, 0.49a 2, 4, 9
Gallop 0.56 0.47, 0.66, 0.39a 2, 4, 9
Riding ability 0.44 0.52, 0.36, 0.43a 2, 4, 10
Free jumping 0.56 0.47, 0.65, 0.47a 2, 4, 9
Parcours jumping 0.40 0.38, 0.43, 0.39 4, 9, 10
Mares’ test Walk 0.25 0.15, 0.27, 0.22, 0.35 1, 2, 3, 8
Trot 0.30 0.35, 0.36, 0.14, 0.35 1, 2, 3, 8
Gallop 0.27 0.18, 0.35, 0.18, 0.35 1, 2, 3, 8
Riding ability 0.20 0.26, 0.30, 0.03 1, 2, 3
Free jumping 0.24 0.27, 0.35, 0.15, 0.20 1, 2, 3, 8
Parcours jumping 0.12 0.04, 0.20 1, 5
aNotincluded in average heritability since data sets 3 and 5 are identical.
References: 1, Hascher (1999); 2, Von Velsen-Zerweck (1999); 3, Huizinga (1991); 4, Brockmann
(1999); 5, Christmann (1996); 6, Schade (1996); 7, Gerber et al. (1997a); 8, Gerber et al.
(1997b); 9, Gerber et al. (1996); 10, Van Veldhuizen (1997).
• Basic gaits, riding and jumping ability are subjectively scored variables
with sufficiently high heritabilities.
• Performance testing at station leads to variables with higher heritabilities
than field testing, i.e. environmental effects are controlled and eliminated
to a higher degree when testing at station.
• Scores for parcours jumping show lower heritabilities than scores for free
jumping, i.e. in free jumping, external effects affecting the horse’s perfor-
mance through the rider are excluded.
• Multiple trait analyses (data set 3) lead to higher estimates of heritabilities
than single trait analyses (data set 5), i.e. combining data from the
stallions’ and mares’ test allows partial adjustment for the selection of
stallions joining the stationary performance test.
Performance testing in competitions

In most European countries, performance testing of sport horses in competi-
tions has been carried out in a standardized way over many years so that
genetic analyses used very large data sets of thousands of individual horses
sired by hundreds of stallions. The methods used for estimating genetic
parameters are based on maximum likelihood techniques applying single or
multiple trait animal models. In Table 15.8, the latest estimates of heritabilities
from several countries are summarized.
150
Table 15.8. Heritability estimates of performance traits of sport horses tested in competitions.
Variable Heritability Repeatability Reference
Jumping competitions
Rank (transformed Blom score) 0.09 0.25 5
Rank (transformed Blom score, selected data) 0.02 0.09 5
Highest level during life time 0.16 4
Rank (transformed 1/2) 0.05 0.12 6
Earning (transformed log) 0.05 6
Cumulative lifetime points (transformed log) 0.28 7
Score 0.14 2
Rank 0.25 0.45 8
Error points 0.09 0.29 1
Dressage competitions
Highest level during life time 0.11 4
Rank (transformed 1/2) 0.11 0.27 6
Score 0.35 2
References: 1, Hascher (1999); 2 Brockmann (1999); 3, Schade (1996); 4, Van Veldhuizen
(1997); 5, Janssens et al. (1997); 6, Hassenstein (1998); 7, Foran et al. (1994); 8, Tavernier
(1990).
The estimates of heritabilities and repeatabilities are based on data from

warmblood riding horses in Belgium, France, Germany, Ireland, The Nether-
lands and Switzerland. The estimated heritabilities of variables measured in
competitions as compared with those from station or field tests indicate the
following:
• Heritabilities and repeatabilities of competition variables are about 0.10
and 0.30 for jumping and dressage.
• Heritabilities of performance traits measured in competitions, in mares’
field tests and in stallions’ station tests are about 0.10, 0.25, and 0.50,
respectively.
• Low heritability estimates for competition data are also caused by the
selection of the data: horses at an higher age are selected, unplaced horses
are either not recorded or incorrectly handled.
• Definition of performance in competitions through earnings or ranks
insufficiently reflects the performance of a horse.
• Other unknown environmental effects influence the performance of
horses in competitions and cause low heritabilities.
When working with competition data, estimates of heritabilities depend
very much on the type of data. The definition of performance in competition
and the system of performance testing in competitions as organized by the
national federations affect genetic analyses. One of the greatest problems is
151
selection in the data. Janssens et al. (1997) demonstrate the effect of ignoring
unplaced horses which substantially decreased the estimates of heritabilities
from 0.09 to 0.02. Similarly, when estimating heritabilities based on older,
selected horses, reduced estimates can be found depending on the data set
used (Huizinga and Van der Meij, 1989; Van Veldhuizen, 1997).
However, selection also takes place between events and between quality
groups of events; competitions might be grouped according to quality, training
and age of horses in A- (the lowest), L-, M- and S- (the highest) classes.
Defining performance data within those quality groups as separate traits and
then estimating heritabilities by applying multiple trait models can account
for the selection in the data. In this way, Hassenstein (1998) estimated
heritabilities in jumping competitions as 0.14, 0.07, 0.06 and 0.06, and in
dressage competitions as 0.17, 0.10, 0.11 and 0.13 for the four quality groups,
respectively. However, Aldridge et al. (1999) found an opposite trend in the
estimated heritabilities as 0.07 for the low level, 0.08 for the medium level and
0.10 for the high-level jumping competitions.
Defining the performance of horses in competitions through earnings
or ranks seems to be insufficient; most estimates of heritabilities based on
earnings and ranks, even after transformation, are around 0.10. However,
Brockmann (1999) analysed data from quality events for young horses, used
scores for describing performance and estimated heritabilities of 0.14 and 0.35
for jumping and dressage.
The higher estimates of heritabilities by Foran et al. (1994) and Tavernier
(1990) raise questions about the system of performance testing which should
lead to testing large and non-pre-selected progeny groups of stallions.
Genetic relationships between performance traits
Due to the definition of the overall breeding goal in sport horses, genetic
relationships between performance traits are essential in defining breeding
objectives. Genetic correlations are estimated based on data from the stallions’
and mares’ tests. Again, only the latest studies are presented in Table 15.9.
The studies on genetic correlations can be summarized as:
• Positive genetic correlations exist between variables describing character/
temperament, basic gaits and riding ability of sport horses. The correla-
tions are moderate.
• High positive genetic relationships exist between basic gaits and riding
ability independently of the type of performance test.
• Between jumping ability and all other variables, no genetic correlations
are found. The estimates were slightly positive or negative and seem to
depend on the type of performance test since mostly negative correlations
were found when variables are measured in the stallions’ test. Similar esti-
mates were obtained based on competition data (Huizinga and Van der
Meij, 1989), whereby the estimates were between −0.06 and −0.27 based
152
Table 15.9. Estimates of genetic correlations between performance traits within testing
schemes (stallions’ test above the diagonal, mares’ test below the diagonal)a.
Walk Trot Gallop Riding ability Jumping ability
Character, 0.30(4) 0.39(4) 0.24(4) 0.51(4) −0.04(4)−

temperament 0.47(2) 0.24(2) 0.36(2) 0.42(2) 0.67(2)
Walk 0.75(4) 0.74(4) 0.79(4) −0.21(4)−
0.92(2) 0.88(2) 0.97(2) 0.13(2)
Trot 0.48(1) 0.87(4) 0.84(4) −0.17(4)−
0.52(3) 0.93(2) 0.95(2) 0.18(2)
Gallop 0.67(1) 0.82(1) 0.80(4) 0.03(4)
0.60(3) 0.82(3) 0.94(2) 0.18(2)
Riding ability 0.48(1) 0.89(1) 0.82(1) −0.04(4)−
0.68(3) 0.83(3) 0.73(3) 0.12(2)
Jumping ability 0.08(1) 0.44(1) 0.59(1) 0.35(1)
−0.10(3)− 0.02(3) 0.11(3) 0.06(3)
0.42(5) 0.42(5) 0.42(5)
aReferences are given in parentheses: 1, Hascher (1999); 2, Huizinga (1991); 3, Christmann
(1996); 4, Schade (1996); 5, Gerber et al. (1997b).
on data from young stallions, but based on data from older stallions the
estimates were between −0.04 and +0.10.
Since the breeding objective is to improve the performance of sport
horses in competitions, the genetic relationships between the various testing
schemes are of paramount importance. Although the general principle in
breeding programmes is to test potential candidates for breeding as early as
possible, the testing schemes designed for young stallions and mares differ
from the normal situation in competitions. The indirect selection based
on the stallions’ and mares’ test is only reasonable if heritabilities and
genetic correlations are high. Table 15.10 summarizes estimates of genetic
correlations between corresponding performance variables in different testing
schemes.
The estimates indicate:
• The corresponding performance traits measured in stallions’ or mares’
tests are highly correlated.
• The genetic relationships between corresponding variables measuring
either riding ability/dressage or jumping ability in stallions’ tests and
competitions are highly positive (> 0.80).
• The genetic relationships between corresponding variables measuring
either riding ability/dressage or jumping ability in mares’ tests and compe-
titions are also highly positive (> 0.65).
• Performance traits measured in testing schemes as designed for young
stallions and mares are good predictors for the performance of horses in
dressage or jumping competitions.
153
Table 15.10. Estimates of genetic correlations between corresponding performance traits of

different testing schemesa.
Stallions’ test : Stallions’ test : Mares’ test :

mares’ test competitions competitions
Walk 0.78(1)
0.95(5)
Trot 0.85(1)
0.95(5)
Gallop 0.74(1)
0.95(5)
Riding ability 0.90(1) 0.88(4) 0.68(3)
0.90(6) 0.68(6) 0.83(2)
0.83(2)
Jumping ability 0.88(1) 0.79(4) 0.64(3)
0.95(5) 0.90(6) 0.48(2)
0.95(5) 0.90(2)
aReferencesare given in parentheses: 1, Von Velsen-Zerweck (1999); 2, Huizinga (1991);
3, Brockmann (1999); 4, Schade (1996); 5, Gerber et al. (1996); 6, Van Veldhuizen (1997).
Concluding remarks
The genetic analyses of performance data of sport horses show that the vari-
ables measured in the stallions’ test at station are highly heritable and highly
correlated with the corresponding variables in competitions. The variables
measured in the 1-day field test of mares show similar, but lower results. On
the other hand, variables describing the performance in competitions have
generally low heritabilities. Model calculations were done to optimize breed-
ing programmes in sport horses, also considering artificial insemination as a
common technique in horses (Koerhuis et al., 1994; Bruns and Schade, 1998;
Brockmann, 1999). Genetic progress in sport horse breeding can be increased
when selection is based on multiple-trait genetic evaluations of stallions and
mares and carried out at sequential steps combining information from the tests
of stallions and mares and competitions. Such integrated systems of genetic
evaluation of sport horses as shown by Von Velsen-Zerweck (1999) have great
prospects in horse breeding.
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Génétique Quantitative et Appliquée, 78352 Jouy en Josas, France;

Albrecht-Thaer-weg 3, 37075 Göttingen, Germany; 3Department of

412 A. Ricard et al.

Interestingly, there were 114,026 recorded births worldwide in Thoroughbreds

The selection and breeding process in Thoroughbreds is founded on the belief

Heritability of track performance

The most comprehensive assembly of published results on heritability of rac-

Genetics of Performance Traits 413

Race length (m)

Type of track 1000 1200 1400 1600 1800 2000

Turf 0.25 0.16 0.10 0.12 0.09 0.08

414 A. Ricard et al.

Rate of genetic improvement

The rate of genetic improvement in a population under selection can be

In contrast to the results showing steady genetic improvement in Timeform

Genetics of Performance Traits 415

416 A. Ricard et al.

Genetics of Performance Traits 417

Horses for courses

418 A. Ricard et al.

Speed and quality of trotter

A natural choice to measure a performance trait for a racing horse is obviously

Genetics of Performance Traits 419

At the end of a race, money is allocated to the horses according to their

420 A. Ricard et al.

the programme of endowment of races, these different transformations lead to

Genetics of Performance Traits 421

old), and Langlois (1984) found genetic correlations of 0.92–0.98 between

Table 15.2. Heritability of criteria based on earnings.

Breed/Age Reference Trait related to earnings Heritability

French trotter 1 Log(annual earnings/starts) 0.26

Table 15.3. Correlation between criteria based on earnings and time.

Breed/age Reference Genetic correlation Phenotypic correlation

French trotter 3–6 years 1 −0.86/−0.95 −0.59/−0.69

422 A. Ricard et al.

selection based on times and earnings is quite effective. A multiple trait

Are there still true precocious performances?

In the preceding, performance is supposed to play the same role throughout

Relationship to mature performances

Genetics of Performance Traits 423

still expresses higher heritability for early performance. Genetic correlations

A new way to measure success and career

In future, measuring performance for trotters must be able to describe what

424 A. Ricard et al.

Ranking Probability (%)

The likelihood of ranking may be written as:

Genetics of Performance Traits 425

estimations of the effects. The probability of the races was multiplied by 11 on

with yijk, underlying performance responsible for ranks; bi, environmental

426 A. Ricard et al.

The problem of non-recorded horses

Performance in trotters is actually recorded as best times per kilometre and

Genetics of Performance Traits 427

In sport horses, breeding objectives consist of a number of different variables

Performance testing at station and in field tests

428 A. Ricard et al.

degree of objectivity due to the subjective scoring. Introducing a higher degree

Performance testing in competitions

Test location Variable (scores) Mean Standard deviation

Stallions’ licensinga Walk 6.8 1.8

Genetics of Performance Traits 429