Executive Summary
What is the report about?
This report details a long-term
study into the feasibility of breeding Angora and Cashmere goats for resistance
to worm (gastrointestinal nematode or helminth) infections. The research
is important because gastro-intestinal nematodiasis (GIN) is the major
health problem of grazing goats word-wide, including Australian fibre goats.
The problem is exacerbated by a widespread, and growing, problem with anthelmintic
resistance in worm populations on goat farms. This makes ongoing reliance
on chemical control of GIN a risky and possibly non-sustainable approach.
Selection for genetic resistance in the host is an alternative control
measure which could be used in conjunction with chemical and other control
measures to place worm control on a more sustainable footing into the future.
Who is the report targeted
at?
The report is targeted primarily
at geneticists and animal breeders in the fibre goat industry. It is also
targeted at policy makers within the industry. Parasitologists servicing
the goat industry will also find the information in this report useful.
Background
At the time this project
commenced there was emerging evidence from overseas studies of genetic
variation in resistance to GIN in goats that could be exploited to select
for resistance to infection. No such information was present for Australian
goats. At that point the sheep industry had made considerable progress
in describing the genetic component of resistance to infection and incorporating
this trait in breeding programs. The goat industry has lagged behind in
this, both because of less use of industry-wide performance recording schemes
(eg. Sheep Genetics Australia) and also because the genetic parameters
for resistance traits, and their association with production traits had
not been defined in Australian goat breeds. Successful incorporation of
genetic resistance to gastro-intestinal helminth infection in goats could
result in reduced anthelmintic (drench) usage and concurrent reduction
in the selection pressure for anthelmintic resistance, and improved efficiency
of fibre production.
Aims/Objectives
The major aims of the project
were as follows:
-
to obtain genetic parameter
estimates for worm resistance traits and production traits in Australian
Cashmere and Angora goats, and phenotypic and genetic correlations amongst
them;
-
to develop a DNA marker set
for parentage analysis and to have the quantitative genetic analysis based
on DNA verified pedigrees;
-
to provide an evaluation of
potential markers for worm infection or resistance in goats other than
faecal worm egg count (WEC);
-
to evaluate the use of irradiated
larval vaccines for control of worm infections in goats; and
-
to obtain new information on
the effects of worm infections in goats of the two breeds in the New England
environment.
The beneficiaries of
the project are likely to be geneticists, parasitologists and animal breeders
in the fibre goat industry who have a new and comprehensive set of genetic
parameter estimates for Angora and Cashmere goats under commercial Australian
conditions. Ultimately goat producers themselves should be the beneficiaries
as decision-making regarding breeding programs and parasite control is
put on a more scientific footing.
Methods used
The project utilised one
commercial Angora and one commercial Cashmere resource herd, both near
Barraba in Northern NSW (Latitude 30.38° S; Longitude 150.61° E).
In both herds does were mated in single sire groups in March/April. In
the Angora herd does were regrouped into a single mob after joining and
they kidded down as one mob. In the Cashmere herd, does were allocated
to different “management groups” after joining and kidded down in these
groups, each containing does mated to each of the sires used. Link sires
were used across years to account for year effects. Both male and female
kids were included in the analysis for Angora goats, but only female kids
were included for cashmere goats. Kids were weighed, and had blood and
faecal samples collected at 3 and 5 months of age after exposure to natural
worm challenge from the pastures they were grazing. At the 5-month sampling
the kids were treated with anthelmintic and a week later each kid was orally
dosed with 10,000 infective larvae of Trichostrongylus colubriformis (black
scour worm). Four weeks later the animals were again weighed, and had blood
and faecal samples collected. A week after this, another faecal sample
was collected and the animals were treated with anthelmintic to terminate
the artificial challenge. Bodyweight and fleece data were collected at
the first two annual shearings for Cashmere goats, and for the first 3
6-monthly shearings for Angora goats. For cashmere goats the project dataset
included kids from the 2000, 2001, 2002 and 2003 kid drops, while for the
Angora goats only the first 3 of these kid drops were included.
WEC was estimated using a
modification of the standard McMaster technique, and fleece samples were
tested for a range of fibre variables using the OFDA 100 after calibration
with AWTO mohair tops. Blood samples were assessed for a wide range of
haematological variables using a Cell Dyn® 3500 automated haematology
analyser.
Pedigree assignment was tested
using DNA tests (PCR) developed and validated by the project, using a set
of 14 microsatellite markers. In total, 18 production and parasite associated
traits were analysed to estimate (co) variance components, heritabilities
and correlations. Where necessary data were transformed prior to analysis.
Fixed effects were estimated using a sire model with sire fitted as a random
effect. Estimation of variance components for the estimation of genetic
parameters utilised four single-trait models fitting the animal as a random
effect. Model 1 included significant fixed effects only; Model 2 added
the direct additive effect of the animal; Model 3 added the with permanent
environment effect of the dam and model 4 added the additive and permanent
environmental effect of the animal itself. Genetic parameters were estimated
from the model giving the best fit beyond which no further improvement
occurred (mostly model 2).
Results and key findings
The project was successfully
implemented but drought conditions in 2002, carrying over into 2003 resulted
in very low levels of natural worm challenge of kids in these years. The
main findings of the project are as follows:
-
Both Angora and Cashmere goats
are able to mount significant effective immune responses to worm infection
by 5 months of age that are capable of limiting subsequent infections.
-
Despite this, early vaccination
at 1 and 2 months of age with bolus challenges of radiationattenuated infective
worm larvae failed to induce a protective immune response.
-
A DNA test for parentage analysis
was developed and validated. The panel of microsatellite markers used provided
a very high level of power (99.7%) of exclusion of non-true parents. The
overall pedigree error rate of 12.7% was within the range reported for
other species. As expected ascertaining the dam by mothering up at marking
reduced dam pedigree accuracy only relative to mothering up at birth (14.6%
and 9.8% error rate respectively).
-
Heritability estimates for faecal
worm egg count (WEC) were within the range of published estimates for other
goat populations and were generally low (range 0.2-0.22), indicating that
genetic progress for selection on this trait is likely to be low to moderate
and somewhat lower than may be seen in sheep. This is particularly true
if selection is based on phenotypic evaluation of sires (as is the case
in much of the Cashmere industry). If however, progeny and other relative
data is captured in industry performance recoding schemes such as MOPLAN,
accurate EBVs can be calculated, even for traits of low heritability.
-
Although the heritability estimates
of some parasite-associated traits measure in blood were high or very high
insufficient information is available to recommend selection based on these
variables in the place of WEC. This is mainly due to the lack of estimable
genetic correlations. For several of these variables this report contains
the first estimates of heritability made for these variables in goats.
-
The heritability estimates for
liveweight and fleece traits for cashmere goats were mostly within the
range of other published estimates, with moderate to high heritability
for most traits. This indicates that rapid genetic progress can be made
by genetic selection for these traits although a strong unfavourable genetic
correlation between cashmere down weight and diameter requires that both
traits be measured for optimum genetic progress.
-
The heritability estimates for
liveweight and fleece traits for Angora goats tended to be higher than
most other published estimates. Once again this indicates that rapid genetic
progress can be made by genetic selection for these traits although a strong
unfavourable genetic correlation between fleece weight and mean fibre diameter
requires that both traits be measured for optimum genetic progress.
-
The project has produced the
first genetic parameter estimates for the trait of fibre curvature in goats.
The trait is moderately heritable in both breeds of goats and has a strong
negative genetic correlation with fibre diameter suggesting that crimp
frequency can be used as a marker for fibre diameter in both breeds of
goat.
-
Genetic associations between
parasite-associated traits and production traits were non-estimable.
Phenotypic associations
were in the expected directions for both breeds, and did not suggest that
selection for parasite-associated traits would lead to negative effects
on production traits.
Implications for relevant
stakeholders
The major implications of
this project are that it has provided an additional set of genetic parameter
estimates for Australian Cashmere and Angora goats to complement those
of Pattie and Restall (1991) and Gifford et al (1991) for cashmere goats,
and Gifford et al (1989) for Angora goats. The parameter estimates were
obtained in current commercial animals in commercial herds in typical goat
production areas. For the first time, traits associated with worm infection
and resistance to it were included in the analysis and the trait of fibre
curvature was also included. Also for the first time the analysis was based
up pedigree data validated by DNA testing. The findings support other studies
in showing that rapid genetic progress can be made in selection for fleece
traits and moderate progress for bodyweight.
Selection for resistance
to worm infection is likely to result in slow genetic progress with no
evidence of negative correlated responses in production traits.
The project has also developed
and validated a panel of microsatellite markers that can be used for parentage
analysis in goats. An attempt to control worm infections using early oral
vaccination with radiation-attenuated larvae was unsuccessful and should
not be pursued further.
Recommendations
-
The inclusion of selection for
resistance to worm infection on the basis of individual WEC measurements
will result in slow genetic gains in this trait. Inclusion of the trait
should only be considered where additional information from relatives would
allow accurate estimates of EBV (eg. within MOPLAN). Selection for this
trait based on sire measurements alone is unlikely to be worthwhile due
to the low heritability. Inclusion of WEC in the selection index requires
careful evaluation as not only will the response to selection be relatively
slow, but applying significant selection differential to WEC will reduce
the selection differential able to put on traits of high heritability and
economic value such as fleece traits.
-
Given the lack of estimable
genetic correlations between WEC and production traits in this dataset,
published values from overseas studies will need to be used. Based on such
studies the genetic correlation between WEC and liveweight is not different
from 0 but tending towards positive, that between WEC and fleece variables
is weakly positive and that between WEC and PCV is moderately to strongly
negative.
-
Given environmental variation
in the level of parasite challenge, individual tests of WEC should always
be preceded by a bulk WEC test indicating significant worm burdens in the
herd (eg. at least 500 epg in regions with Haemonchus contortus present).
-
Given the moderate to very high
heritabilities of some parasite-associated blood traits and the increasing
availability of automated haematology analysis, support should be given
to determining genetic correlations between these traits and WEC to more
fully evaluate whether they are viable alternative measures of resistance
and/or resilience to GIN. The physiological consequences of extreme values
for blood constituents also need to be determined.
-
Given the difficulty of using
selection for genetic resistance to control worms, research emphasis should
be applied to other control measures suitable for inclusion in an integrated
worm control program in goats. Two approaches supported by the findings
of this project are detailed below.
-
The evidence in this study that
immune regulation of worm burdens is related to the level of prior worm
challenge indicates that parasite control methods that rely on maintaining
some parasite burden in the host are more likely to be successful than
those that totally suppress worm infections intermittently. Tactical worm
control based upon close monitoring of WEC and treatment intervention only
when the WEC exceeds a given threshold based on worm species involved,
animal condition, feed availability and climatic conditions is likely to
maintain immunity in the population and help regulate infections. Such
an approach has been successfully used in sheep (Kahn et al., 2006; Scrivener
et al., 2006).
-
The marked year to year variation
in worm burdens associated with drought conditions reinforces the susceptibility
of the free-living stages of gastro-intestinal nematodes to adverse environmental
conditions (Review: O’Connor et al., 2006) and our ability to exploit this
weakness using grazing management practices such as alternate grazing with
cattle (Southcott and Barger, 1975) and intensive rotational grazing (Colvin
et al., 2007).
-
Early oral vaccination of kids
with bolus doses of irradiated worm larvae cannot be recommended as a practical
worm control method and there is little evidence that it is likely to ever
be so.
-
The findings of this project
would support the use of an on-farm measure of fibre curvature (eg.= “crimp”
frequency) as an indirect measure of fibre fineness in both Angora and
Cashmere goats.However full objective measurement in a laboratory will
maximise genetic progress.
-
The project results should be
published in the scientific literature and be made widely available to
industry.