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Comparison of breeding objectives across countries with application to sheep

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Article  in  Journal of Animal Breeding and Genetics · April 2015

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 J. Anim. Breed. Genet. ISSN 0931-2668

ORIGINAL ARTICLE

Comparison of breeding objectives across countries with

application to sheep indexes in New Zealand and Ireland
B.F.S. Santos1,2, N. McHugh3, T.J. Byrne1, D.P. Berry3 & P.R. Amer1
1 AbacusBio Limited, Dunedin, New Zealand
2 School of Environmental & Rural Science, University of New England, Armidale, NSW, Australia
3 Animal & Grassland Research and Innovation Centre, Teagasc, Fermoy, Ireland

Keywords
Economic selection indexes; genetic
improvement; genetic parameters; production
systems; profitability; sheep.
Correspondence
B.F.S. Santos, AbacusBio Limited, Dunedin,
New Zealand, PO Box 5585, Dunedin, 9058,
New Zealand.
Tel: +64 3 477 6375;
Fax: +64 3 477 6376;
E-mail: bsantos@abacusbio.co.nz
Received: 24 November 2014;
accepted: 2 February 2015
Summary
Breeding objectives and selection indexes underpin the direction, the
extent and the economic implications of selection in livestock populations
under specific production systems. The objective of this study was to
describe the methodology to calculate correlations between national
selection indexes and gain a deeper understanding of the factors influenc-
ing responses in economically important traits in both the New Zealand
and Irish sheep industries. Moderate to strong correlations were calculated
among indexes within and between countries, with the strongest correla-
tion (0.86) between the New Zealand and Irish maternal indexes. In both
countries, responses to selection in the maternal indexes are largely dri-
ven by growth traits; each index, however, has a different balance of
traits. Ewe mature weight also accounts for an important proportion of
overall response and has significant emphasis in both maternal indexes.
The majority of emphasis in terminal indexes of both countries is on
growth and meat traits. Results from this study indicate that differences
between national breeding objectives are unlikely to be a barrier to
exchange of gene stocks among countries. Future research should investi-
gate the extent to which genotype-by-environment (G9E) interactions
exist at the level of individual traits. The methodology presented in this
study is robust and represents an opportunity to inform the potential mer-
its of international exchange of germplasm.

emphasis towards specific market outcomes or to

Introduction
The most important decision in the design of animal
improvement programmes is the choice of the breed-
ing objective (James 1983). Multiple-trait selection,
and the associated economic implications, is normally
achieved through the definition of traits to be
improved, and their appropriate combination, for spe-
cific production systems. The most efficient way of
using the available information from this combination
is to construct a selection index (James 1980). Eco-
nomic selection indexes are an important tool for
ranking animals, and their formulation enables com-
mercial producers and breeders to adjust breeding
address key production aspects in their particular
farming system (Byrne et al. 2010). Breeding objec-
tives and selection indexes also define the direction of
selection in a livestock population.
The definition of breeding objectives may differ
between countries and production systems. However,
it is common practice to exchange gene stock between
breeding industries by importation. Where climates,
markets and economic costs are similar between
countries, there is an important role for the exchange
of gene stocks (Nielsen et al. 2013). Objective compar-
isons of alternative indexes may inform whether live-
stock selected through specific breeding objectives

doi:10.1111/jbg.12146 © 2015 Blackwell Verlag GmbH • J. Anim. Breed. Genet. 132 (2015) 144–154
B. F. S. Santos et al.
and selection indexes are likely to perform accord-
ingly in alternative production systems or countries.
In this context, consideration should be given to: (i)
internationalization of genetic improvement and its
potential benefits for entire industries, for instance in
dairy, pigs and poultry; (ii) genetic improvement in
certain countries to be driven by importation of for-
eign gene stocks; (iii) the fact that imported gene
stocks are not always the most pertinent alternative;
(iv) economic imperatives changing the priority of dif-
ferent traits, or GxE interactions altering the ranking
of animals for similar traits (Sadeghi-Sefidmazgi et al.
2012). It is also important to consider that reliance on
importation of gene stocks, as a long-term breeding
strategy, could be a high risk strategy without under-
standing the drivers of genetic change in different
countries. This is especially important given that
importation is often driven by marketing instead of
technical information.
This study describes a methodology that can be used
to objectively compare alternative breeding objectives
and selection indexes, for example, among different
countries. A particular focus is on defining a metric
that can be interpreted as the correlation between
alternative selection indexes. In this way, the poten-
tial merit of maintaining ongoing genetic exchanges
between two populations striving to maximize genetic
progress can be better understood. To illustrate the
methodology, a detailed comparison is made among
selection indices used in New Zealand and Ireland in
terminal and maternal breeds.
Materials and methods
Overview
A methodology used to define the correlation
between selection indexes and to compare genetic
gain in economically important traits is proposed. The
description is structured in sections which detail the
core aspects of the methodology including formulae
and calculations steps.
Index correlations
The correlation (rx,y) between two indexes (x and y)
was calculated as
ax 0 Gxy ay
rx;y ¼ qffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi
ax 0 Gx ax  ay 0 Gy ay
where, Gxy is the genetic variance–covariance matrix
(G) between breeding objective traits in indexes x
© 2015 Blackwell Verlag GmbH • J. Anim. Breed. Genet. 132 (2015) 144–154
Comparison of breeding objectives
and y adjusted for accuracy of the respective genetic
evaluations, Gx and Gy are matrices of accuracy
adjusted genetic variance–covariance within indexes
x and y, respectively, and ax and ay are vectors of
trait economic weights used in the respective
indexes, x and y.
Index responses to selection
The prediction of response to selection (Rxi) in trait i
from selection on the index x was calculated as:
bx 0 Gxi
Rxi ¼ i  pffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi
ffi ð2Þ
bx 0 Gx ax
where, i is the assumed selection intensity (assumed
here to be 1 so that all results are per unit of selection
intensity), Gxi is the column vector of Gx relating to
trait i, a is the respective set of economic weights, and
bx is the column vector of selection coefficients of
index x, calculated as:
bx ¼ Px 1 Gx a ð3Þ
where, Px is the accuracy adjusted phenotypic vari-
ance–covariance matrix.
The responses to selection for trait i when selection
is on index x were then expressed per genetic standard
deviation unit of trait i (Rsdij), or per economic unit
(Raxi), of trait i as:
Rxi
Rsdxi ¼ ð4Þ
sdxi
Raxi ¼ Rxi axi ð5Þ
where, sdxi is the genetic standard deviation of trait i
in index x.
Trait relative emphasis
The relative emphasis on a profit trait was defined as
the percentage of total economic value of genetic gain
in all traits attributable to gain in a particular trait.
Following the methodology suggested by Cunning-
ham & Tauebert (2009), the relative emphasis (RE) of
the target trait i is as follows:
b0 G
RE ¼ 0 ix ðai Þ  100 ð6Þ
b Px b
Accuracy adjusted variance–covariance matrices
ð1Þ
A number of parameters were used to form the accu-
racy adjusted genetic (Gx ) and phenotypic (Px ) vari-
ance–covariance matrices between indexes in the
different countries. The accuracies of trait breeding
145
Comparison of breeding objectives B. F. S. Santos et al.

Table 1 Trait description and unit, heritability (h2), genetic (rg) standard deviations, accuracies (r) and economic weights for sheep profit traits in ter-
minal (TermNZ) and maternal (MatNZ) selection indexes in New Zealand

Economic weights
(€/unit)1

Trait group Trait (Abbrev.) Unit h2 rg r TermNZ MatNZ

Reproduction Number of lambs born (NLB) Lambs 0.10 0.18 0.25 – 14.15
Ewe mature weight Ewe mature weight (EWT) kg 0.45 2.32 0.30 – 0.95
Survival Lamb survival (SUR) Lambs 0.01 0.04 0.13 28.97 58.66
Survival single Lambs 0.01 0.02 0.13 – –
Survival twin Lambs 0.01 0.02 0.13 – –
Survival triplet Lambs 0.02 0.05 0.18 – –
Survival maternal (SURm) Lambs 0.05 0.09 0.05 – 53.15
Survival single maternal Lambs 0.03 0.03 0.13 – –
Survival twin maternal Lambs 0.07 0.06 0.13 – –
Survival triplet maternal Lambs 0.07 0.09 0.18 – –
Growth Weaning weight (WWT) kg 0.20 1.57 0.58 0.43 0.86
Weaning weight maternal (WWTm) kg 0.10 1.11 0.25 – 0.77
Carcase weight (CWT) kg 0.30 1.10 0.60 1.24 2.37
Meat Lean yield (LY) kg 0.41 0.46 0.60 2.95 –
Other Lamb fleece weight (LFW) kg 0.15 0.08 0.50 – 1.66
Hogget fleece weight (HFW) kg 0.35 0.30 0.60 – 0.72
Ewe fleece weight (EFW) kg 0.45 0.34 0.30 – 2.07
Faecal egg count-1 (FEC1) Log count % 0.16 33.47 0.52 – 0.03
Faecal egg count-2 (FEC2) Log count % 0.20 38.47 0.58 – 0.03
Adult faecal egg count (AFEC) Log count % 0.25 43.01 0.40 – 0.02
Facial eczema (FE) LogeGGT 0.40 0.57 0.60 – –
1
Economic weights for traits in New Zealand selection indexes were sourced from SIL.

values (q), presented in Tables 1 and 2, were used to
weight the effect of the different traits according to
the level of precision to which the additive effects are
predicted. The values were chosen based on typical
accuracies in young male selection candidates from
flocks with moderate to high levels of trait recording
for core index traits. Phenotypic variances (r2p ) and
heritability estimates (h2) of the different traits were
used to estimate the genetic variances (r2g ) from
which variance components could be calculated, after
accounting for correlations among traits (rij denotes
the genetic correlations between traits i and j). The
formulae used to calculate parameters were as
follows: qffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi
Gx ¼ rij h2i r2pi q2i  h2j r2pj q2j
qffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi
Px ¼ rij r2pi q2i  r2pj q2j
where, rij = 1 for the special case where trait i = trait
j.
Case study
In meat sheep production, the most common source
of revenue is generally the total weight of either live
lambs or carcases. Therefore, traits directly related to
these target outcomes are frequently considered in
the definition of breeding objectives in different
sheep populations. Traits generally common across
many sheep breeding objectives include the follow-
ing: litter size or number of lambs born (fertility/
fecundity), lamb survival (functionality), weaning
weight, days to slaughter and carcass weight
(growth), and carcass conformation, fat score and
lean yield (meat). The relative emphasis on each
trait when making selection decisions, however, dif-
fers based on the different production systems and
markets for each population.
Two sheep breeding objectives exist in New Zea-
ð7Þ land: the Dual Purpose Overall index (MatNZ) and
the Terminal Sire Overall index (TermNZ). The trait
ð8Þ genetic parameters and economic weights of both
indexes for the year 2014 are presented in Table 1.
The Irish breeding objectives are represented by
two selection indexes: Replacement (MatIRE) and
Terminal (TermIRE). The respective trait genetic
parameters and economic weights for the Irish
indexes for the year 2014 are shown in Table 2.
Indexes in both countries are measures of the
genetic ability of a selection candidate to generate

146 © 2015 Blackwell Verlag GmbH • J. Anim. Breed. Genet. 132 (2015) 144–154
B. F. S. Santos et al. Comparison of breeding objectives

Table 2 Trait description and unit, heritability (h2), genetic (rg) standard deviations, accuracies (r) and economic weights for sheep profit traits in ter-
minal (TermIRE) and maternal (MatIRE) selection indexes in Ireland

Economic weights
(€/unit)1

Trait group Trait (Abbrev.) Unit h2 rg r TermIRE MatIRE

Reproduction Number of lambs born (NLB) Lambs 0.07 0.13 0.20 – 5.27
Ewe mature weight Ewe mature weight (EWT) kg 0.30 4.11 0.30 – 0.36
Survival Lamb survival (LSUR) Lambs 0.02 0.05 0.18 22.63 22.63
Lamb survival maternal (LSURm) Lambs 0.01 0.04 0.13 – 20.64
Growth Days to slaughter (DTS) Day 0.20 15.16 0.41 0.09 0.09
Days to slaughter maternal (DTSm) Day 0.10 10.72 0.29 – 0.06
Meat2 Carcase conformation (CCON) EUROP class 0.25 0.25 0.42 1.40 1.40
Carcase conformation maternal (CCONm) EUROP class 0.12 0.17 0.23 – 1.16
Carcase fat (CFAT) EUROP class 0.15 0.35 0.35 1.50 1.50
Carcase fat maternal (CFATm) EUROP class 0.08 0.25 0.18 – 1.24
Other Lambing ease single (LES) % 0.05 0.49 0.29 0.14 0.14
Lambing ease multiple (LEM) % 0.05 0.49 0.29 0.08 0.08
Lambing ease single maternal (LESm) % 0.03 0.38 0.23 – 0.13
Lambing ease multiple maternal (LEMm) % 0.03 0.38 0.23 0.07 0.07
1
Economic weights for traits in Irish selection indexes were sourced from Sheep Ireland.
2
European Union E,U,R,O,P carcase classification scheme, where letters represent an incremental scale ranging from E (the best conformation)
through to P (the worst conformation), and fat class ranges from 1 (the least fat carcase) to 5 (the fattest carcase).

profit at farm level (Pabiou et al. 2014). In this
study, the economic weights of the New Zealand
indexes were converted into Euro (€), at conversion
rate of 0.6344, to simplify their direct comparison.
In this study, the dual purpose index (MatNZ) and
the replacement index (MatIRE) are referred to as
maternal indexes.
For the purpose of this study, traits from the
national selection indexes were categorized into the
broad groups of: (i) ‘reproduction’ which reflects the
emphasis placed on number of lambs born or litter
size; (ii) ‘ewe mature weight’ indicates the emphasis
given to control the body size of breeding ewes in
the different indexes; (iii) ‘survival’ includes lamb
survival, due to direct and maternal effects; (iv)
‘growth’ includes direct carcass weight, weaning
weight and days to slaughter, as well as their mater-
nal effects; (v) ‘meat’ comprised of lean yield, and
carcass conformation and fat, according to the New
Zealand and Ireland breeding objectives, respec-
tively. A final category ‘other’ includes the remain-
ing traits in the index, such as faecal egg count, ewe
fleece weight and facial eczema in New Zealand,
and direct and maternal lambing ease in Ireland,
which is divided into ease of lambing for single and
multiple litters.
The (co)variance components and economic weights
for this study were obtained from the organizations
co-ordinating genetic evaluations in the respective
countries. Sheep Improvement Limited (SIL) is a divi-
sion of Beef + Lamb New Zealand Genetics, the
national industry organization that provides sheep
genetic information to ram breeders and commercial
farmers. In Ireland, the implementation of the national
genetic evaluation system that integrates commercial
and pedigree sheep breeder data is undertaken by
Sheep Ireland.
Pooled correlations
A number of parameters were required to calculate
the genetic and phenotypic covariance matrix within
and between countries. Many of the parameters used
in this study originated from comprehensive review
by Safari & Fogarty (2003), which compiled estimates
of sheep genetic parameters from several studies. This
study adopted the same approach to that presented by
Koots et al. (1994) and presents pooled genetic corre-
lations between traits estimated using the published
standard error of the correlations, and generating
mean pooled genetic correlations, based on results
from different publications for all traits. The method is
well described in Safari et al. (2005) where the genetic
correlations were transformed to an approximate nor-
mal scale to remove the dependency of the variance
on the estimate, using Fisher’s Z transformation (Steel

© 2015 Blackwell Verlag GmbH • J. Anim. Breed. Genet. 132 (2015) 144–154 147
Comparison of breeding objectives
& Torrie 1960). The equations describing this method
are presented as:
 
1 þ ^rij
Z ¼ 0:5 log ð9Þ
1  ^rij
P In Ireland, the economic weight on ewe mature
Z
Z0 ¼ P 2 ð10Þ
seij
Z0
eij ¼ P   ð11Þ
1
seij 2
0
eij 2Z  1
rij ¼ ð12Þ
eij 2Z0 þ 1
where, ^rij and seij are the genetic correlation and stan-
dard error between traits i and j respectively, Z0 is the
weighted mean of the Z-transformed correlations
[Equation (9)], calculated and back transformed using
Equation (11) to obtain rij which is the weighted mean
transformed genetic correlation between traits. The
methodology assumes a genetic correlation of one
between similar target traits, such as reproduction, ewe
mature weight, survival and growth. Table 3 presents
the normal scaled genetic correlations among traits and
between countries used in this study.
Results
Correlations among indexes
Moderate to strong correlations (0.48–0.86) were cal-
culated between the indexes both within and
between countries; the strongest correlation existed
between MatIRE and MatNZ indexes (0.86). The mod-
erate to strong correlation between both terminal
indexes (0.66) was similar to the correlation between
terminal indexes and maternal indexes within coun-
try (0.67 to 0.68). Weaker correlations were calcu-
lated between the terminal and maternal indexes of
different countries, 0.48 and 0.52, respectively, for
correlation between the TermNZ and MatIRE, and
between the TermIRE and MatNZ.
Response to selection
The expected responses to selection in New Zealand
and Ireland in the different indexes are presented in
Tables 4 and 5. The directions to which trait responses
were observed are similar in the different indexes
within country.
Ewe mature weight was calculated to increase
annually by 0.170 and 0.102 standard deviation units
148
B. F. S. Santos et al.
with selection on the TermNZ and MatNZ indexes,
respectively; this is expected, despite the exclusion of
ewe mature weight from the index (TermNZ) and it
receiving a negative economic weight (MatNZ),
because of direct selection on correlated traits.
weight is also negative and with selection on the
MatIRE index, the expected annual response to selec-
tion is 0.067 genetic standard deviations.
Between countries, a similar direction in calculated
responses to selection was found for most traits. For
example, the response in lean yield, which indicates
the amount of muscle or leanness in the carcass, had
a calculated annual response of 0.353 genetic stan-
dard deviation units in the TermNZ index. The breed-
ing objective trait for muscling in the TermIRE is
carcass conformation, to which the calculated annual
response was 0.010 standard deviation units. The
direction of genetic change is also the same for num-
ber of lambs born, survival and growth in the MatNZ
and MatIRE indexes; however, each index has a differ-
ent balance of traits. This accounts for the difference
in magnitude of small changes in survival and for dif-
ferences in definition of growth traits between the
countries.
The main difference calculated across multiple met-
rics between the maternal indexes in both countries is
the magnitude of the economic response for growth
traits. This is demonstrated when genetic gain on the
basis of standard deviations is multiplied by the eco-
nomic weight of each trait. In New Zealand, the
response to selection in weaning weight in the MatNZ
index is € 0.30 and for carcass weight, € 0.72 per year.
The overall economic response in the MatNZ index is
completed by faecal egg count and ewe mature weight
at € 0.25 and € 0.22 per year, respectively. Also in
New Zealand, the TermNZ index is largely focused on
carcass weight as well as on lean yield, at € 0.43 and €
0.48, respectively.
For the Irish indexes, the greatest economic
response in the MatIRE index is days to slaughter (€
0.11) followed by ewe mature weight (€ 0.10), whilst
the greatest economic response to selection in the
TermIRE index is days to slaughter (€ 0.25).
These results demonstrate that the economic
response to selection (expressed in monetary units)
for growth was greater with the MatNZ index in com-
parison to the MatIRE index.
Relative emphasis of traits
Figure 1 presents the relative emphasis on groups of
traits based on the magnitude of their calculated
© 2015 Blackwell Verlag GmbH • J. Anim. Breed. Genet. 132 (2015) 144–154
 B. F. S. Santos et al.

Table 3 Scaled trait genetic correlations of sheep profit traits in New Zealand and Ireland, based on breeding schemes (within country) and literature references (between countries)

© 2015 Blackwell Verlag GmbH

New Zealand traits Ireland traits

NLB WWT CWT HFW LFW EFW FEC1 FEC2 AFEC SUR EWT WWTm SURm LY DTS CCON CFAT EWT DTSm CCONm CFATm LSUR LES LEM LSURm LESm LEMm NLB

NLB 1.00 – – – – – – – – – – – – – – – – – – – – – – – – – – –

WWT – 1.00 – – – – – – – – – – – – – – – – – – – – – – – – – –

CWT – 0.75 1.00 – – – – – – – – – – – – – – – – – – – – – – – – –

HFW – 0.40 0.30 1.00 – – – – – – – – – – – – – – – – – – – – – – – –

LFW – 0.20 0.20 0.60 1.00 – – – – – – – – – – – – – – – – – – – – – – –

EFW – 0.30 0.25 0.75 0.40 1.00 – – – – – – – – – – – – – – – – – – – – – –

FEC1 – – – 0.15 0.15 0.15 1.00 – – – – – – – – – – – – – – – – – – – – –

FEC2 – – – 0.15 0.15 0.15 0.71 1.00 – – – – – – – – – – – – – – – – – – – –

AFEC – – – 0.15 0.15 0.15 0.71 0.71 1.00 – – – – – – – – – – – – – – – – – – –

• J. Anim. Breed. Genet. 132 (2015) 144–154

SUR – – – – – – – – – 1.00 – – – – – – – – – – – – – – – – – –

EWT – 0.55 0.75 0.30 0.25 0.30 – – – – 1.00 – – – – – – – – – – – – – – – – –

WWTm – – – – – – – – – – – 1.00 – – – – – – – – – – – – – – – –

SURm – – – – – – – – – – – – 1.00 – – – – – – – – – – – – – – –

LY – 0.55 0.60 – – – – – – – 0.70 – – 1.00 – – – – – – – – – – – – – –

DTS 0.30 0.40 1.00 – – – – – – – 0.60 0.30 – 0.30 1.00 – – – – – – – – – – – – –

CCON – 0.40 0.60 – – – – – – – – – – 0.50 0.20 1.00 – – – – – – – – – – – –

CFAT 0.03 0.20 0.46 – – – – – – – 0.18 – – 0.25 0.20 0.12 1.00 – – – – – – – – – – –

EWT 0.40 0.30 – – – – – – – – 1.00 – – 0.89 0.55 0.20 0.10 1.00 – – – – – – – – – –

DTSm – 0.20 1.00 – – – – – – – – 0.40 – – 0.20 – – – 1.00 – – – – – – – – –

CCONm – – – – – – – – – – – 0.40 – – – – – – 0.20 1.00 – – – – – – – –

CFATm – – – – – – – – – – – 0.20 – – – – – – 0.20 0.10 1.00 – – – – – – –

LSUR – – – – – – – – – 1.00 – – – – – – – – – – – 1.00 – – – – – –

LES – – – – – – – – – – – – – – – – – – – – – 0.40 1.00 – – – – –

LEM – – – – – – – – – – – – – – – – – – – – – 0.40 0.20 1.00 – – – –

LSURm – – – – – – – – – – – – 0.76 – – – – – – – – – – – 1.00 – – –

LESm – – – – – – – – – – – – – – – – – – – – – – – – 0.40 1.00 – –

LEMm – – – – – – – – – – – – – – 0.30 – – – – – – – – – 0.40 0.20 1.00 –

NLB 1.00 0.20 – – – – – – – – – – – – – – – – – – – – – – – – – 1.00

Comparison of breeding objectives

149
Comparison of breeding objectives B. F. S. Santos et al.

Table 4 Trait responses to selection on terminal and maternal indexes within New Zealand

TermNZ MatNZ TermNZ MatNZ

Trait Unit (Standard deviations/year) (€/year)

Number of lambs born Lambs – 0.014 – 0.0344

Ewe mature weight kg 0.170 0.102 – 0.2245
Lamb survival Lambs 0.0002 0.0003 0.0002 0.0008
Survival maternal Lambs 0.0000 0.0005 – 0.0024
Weaning weight kg 0.238 0.219 0.1607 0.2963
Weaning weight maternal kg – 0.005 – 0.0039
Carcase weight kg 0.316 0.277 0.4286 0.7206
Lean yield kg 0.353 0.166 0.4790 –
Lamb fleece weight kg 0.042 0.046 – 0.0060
Hogget fleece weight kg 0.096 0.139 – 0.0294
Ewe fleece weight kg 0.042 0.062 – 0.0432
Faecal egg count-1 Log count, % 0.003 0.084 – 0.0737
Faecal egg count-2 Log count, % 0.003 0.109 – 0.1100
Adult faecal egg count Log count, % 0.002 0.080 – 0.0682

Table 5 Trait responses to selection on the terminal and maternal indexes within Ireland

TermIRE MatIRE TermIRE MatIRE

Trait Unit (Standard deviations/year) (€/year)

Number of lambs born Lambs – 0.007 – 0.0051

Ewe mature weight kg 0.081 0.067 – 0.0995
Lamb survival Lambs 0.003 0.003 0.0032 0.0034
Lamb survival maternal Lambs 0.001 0.001 – 0.0011
Days to slaughter Day –0.175 0.082 0.2475 0.1151
Days to slaughter maternal Day 0.019 – – 0.0003
Carcase conformation Europ class 0.010 0.050 0.0034 0.0174
Carcase conformation maternal Europ class – – – 0.0001
Carcase fat Europ class –0.055 0.045 0.0291 0.0236
Carcase fat maternal Europ class – 0.005 – 0.0017
Lambing ease single % 0.002 0.002 0.0002 0.0002
Lambing ease multiple % 0.003 0.003 0.0001 0.0001
Lambing ease single maternal % 0.001 0.001 – 0.0001
Lambing ease multiple maternal % 0.021 0.024 0.0006 0.0006

responses in the maternal and terminal selection
indexes for sheep in New Zealand and Ireland. The
majority of emphasis is on growth and meat traits in
three of four of the indexes. Ewe mature weight also
accounts for an important proportion of emphasis in
the maternal indexes of both countries. For terminal
and maternal indexes in New Zealand and Ireland,
the major similarity is in the relative emphasis of
growth traits. The MatNZ and TermIRE indexes have
the greatest emphasis on growth, at 87.7% and
87.3%, respectively. Differences between the terminal
indexes are due to the importance of meat-related
traits; the TermNZ index applies a stronger emphasis
(44.8%) on lean yield, whilst the TermIRE index
applies 11.4% emphasis on carcass conformation and
fat. Ewe mature weight had considerable relative
emphasis in both maternal indexes; however, its
emphasis produces opposite outcomes. The MatNZ
index has a relative emphasis of 19.3% and results in
a predicted genetic increase in ewe mature weight,
whereas the MatIRE index has a relative emphasis of
37.2% and results in a predicted genetic decrease in
ewe mature weight. The biggest contrast between
maternal indexes across the countries was the absence
of emphasis on meat-related traits in the MatNZ, com-
pared to the MatIRE. In both maternal indexes, the
calculated emphasis on reproduction and survival was
reasonably modest. Lamb survival, which includes
maternal genetic effects, had a low rate of progress in
the MatIRE (1.7%) and in the MatNZ (0.3%) index.
Finally, the MatNZ had 28.4% calculated emphasis on
the component trait ‘other’. In the Irish maternal

150 © 2015 Blackwell Verlag GmbH • J. Anim. Breed. Genet. 132 (2015) 144–154
B. F. S. Santos et al.
Mat NZ
Term NZ
Index
Mat Ire
Term Ire
Figure 1 Relative emphasis based on the eco-
–40
nomic contribution of component profit traits
of selection in maternal and terminal indexes
in Ireland and New Zealand.
index, the component trait ‘other’ represented 0.8%
of the total emphasis (Figure 1).
Terminal indexes also prioritize growth and meat,
which is confirmed by the relative emphasis of the
traits. However, the selection indexes contrast in the
extent of emphasis applied to growth, 55.1% and
87.3%, and meat traits, 44.8% and 11.4%, for the
TermNZ and TermIRE, respectively. The relative
emphasis of other component traits in both terminal
indexes is modest.
Discussion
Sheep production systems in Ireland and New Zealand
are pastoral and seasonal based, although in each
country farm scale and therefore flock size differs. In
New Zealand, sheep farming practices are character-
ized by extensive management around lambing and
provision of a diet consisting exclusively of forage for
ewes and lambs for the entire year. In Ireland, most
farm systems adopt indoor lambing practices and dis-
cretionary concentrate supplementation of ewes and
lambs up to weaning. The production systems are
therefore contrasted by significant differences in pro-
duction costs. The performance of commercial sheep
flocks in these countries is representative of the wide
range of production systems (Kelly 1982).
Livestock production across the world is largely reli-
ant on genetic resources normally originating from
specific populations and distributed to a diverse range
of environments. In dairy and beef cattle, collabora-
tive organizations such as Interbull and Interbeef are
responsible for establishing international genetic eval-
uations. Miglior et al. (2005) compared selection
© 2015 Blackwell Verlag GmbH • J. Anim. Breed. Genet. 132 (2015) 144–154
Comparison of breeding objectives
–19.3 87.7 28.4
55.1 44.8
37.2 43.1 15.3 0.8
87.3 11.4
–20 0 20 40 60 80 100 120 140
Relative emphasis response (%)
Reproduction Ewe mature weight Survival Growth Meat Other
indexes amongst a significant number of countries,
based on data from Interbull. Their methodology, fre-
quently adopted in the literature, compared indexes
based on trait relative emphasis expressed as a per-
centage function of the economic weight and genetic
standard deviation. The implicit assumption is that
selection on the index produces equal accuracy of
selection in all traits. According to Cunningham &
Tauebert (2009), this method can be misleading, as it
tends to overstate the net effect of the addition of new
traits to an index, particularly when there are correla-
tions between traits. In this study, a comprehensive
method to calculate the correlation between alterna-
tive selection indexes has been described. Results
indicate a moderate to strong correlation between
indexes within and between New Zealand and
Ireland, despite within country differences in trait
genetic correlations and economic weights. The calcu-
lations based on this methodology could also be used
to calculate index correlations amongst a larger num-
ber of populations or countries with different indexes.
An important assumption in this application of the
methodology is that similar target traits in the differ-
ent breeding objectives have a genetic correlation of
one. This means that component traits of reproduc-
tion, ewe mature weight, survival, growth and meat
are the same in New Zealand as in Ireland, even
though selection criteria in each country might be dif-
ferent. The method, however, can be easily modified
to account for GxE interactions by assuming weaker
than one genetic correlations between the same traits
in different environments. A further investigation was
conducted by adjusting the genetic correlations of
common traits downward, such as between days to
151
Comparison of breeding objectives
slaughter and carcass weight from 1.00 to 0.75
and also survival and number of lambs born from
1.00 to 0.75, between the two countries. The results
still demonstrated moderate correlations (0.58 as
opposed to 0.86 in the assumed absence of GxE)
between MatNZ and MatIRE. It is therefore important
to understand the definition of the selection criteria
between traits in different countries when carrying
out a study of this nature. No changes in response to
selection or relative trait emphasis occur because they
do not depend on between index covariance. A dee-
per analysis would require accurate covariance esti-
mates between corresponding traits in New Zealand
and Ireland; therefore, effectively accounting for G9E
interactions. Genotype-by-environment interactions
should ideally not be ignored when comparing selec-
tion indexes. It is present if the genetic correlation
between the same trait expressed in different environ-
ments is different from one (Falconer 1952). Accord-
ing to Dominik & Kinghorn (2008), index responses
to selection can be over or under predicted as a conse-
quence of using genetic correlations that do not accu-
rately reflect the extent of existing G9E interactions.
Moreover, Dominik & Kinghorn (2008) suggest that
unfavourable bias and large variability in the pre-
dicted genetic and financial gain for commercial farm-
ers could occur if they use sires selected on indexes
from a divergent stud environment. This could also be
assumed when genetic resources are traded between
countries with contrasting production systems.
The main reason for the reasonably strong correla-
tions between selection indexes in New Zealand and
Ireland could be attributed to trait emphasis in the dif-
ferent selection indexes, as well as to the correlations
among index component traits. The strong correlation
between maternal indexes could be attributed to the
common selection direction between indexes, the rel-
ative emphasis on growth traits, as well as by the
strong within country genetic correlations among the
component traits in the indexes.
The correlations between indexes in different coun-
tries could also be affected by covariance amongst
traits and other genetic parameters within each breed-
ing scheme. Ultimately, heritabilities and genetic cor-
relations between traits will affect the impact of
changing any of the profit traits in a selection index
(Lambe et al. 2014).
The results of the present study indicate that
responses to selection for sheep traits in New Zealand
and Ireland are in a similar direction. This is partially
due to similarities in relative emphasis calculated
among traits in the indexes. However, responses to
selection in New Zealand were found to be greater,
152
B. F. S. Santos et al.
and this can be attributed to the trait accuracies and
genetic variance, driven by trait heritability and phe-
notypic variance (Tables 1 and 2). The maternal
indexes are reasonably similar with respect to the rel-
ative emphasis on growth, although different in the
direction of selection for ewe mature weight. When
maternal traits are considered, especially reproduction
and lamb survival, the similarity between maternal
indexes is strengthened because small relative empha-
sis is realized in these traits. This is achieved regardless
of the differences in production systems between the
countries. Despite the high economic weights, pro-
gress achieved in survival in the MatNZ index and in
the MatIRE index is small and likely due to the low
heritability of the trait.
The contrasting direction to which ewe mature
weight has been selected in the Irish and New Zealand
maternal indexes, with unfavourable selection in the
MatNZ, is explained by the strong genetic correlations
between ewe mature weight and both carcass weight
and weaning weight (0.75 and 0.55, respectively),
which are the main components of the index
(Table 3). This produces the upward response in ewe
weight by selecting on the MatNZ index, given the
high emphasis in the index on growth, and its signifi-
cant economic weight and genetic variance. Safari
et al. (2005) documented a mean genetic correlation
of 0.78 between ewe mature weight and growth rate,
whilst Borg et al. (2009) reported genetic correlations
of between 0.69 and 0.75 between adult weights
before, during and after lactation with lamb weaning
weight at 120 days. Conington et al. (2001) presented
genetic correlations between mature size and carcass
weight and lamb weaning weight of 0.49 and 0.72,
respectively. That study indicated that ewe mature
weight was the most important trait affecting genetic
progress across alternative hill sheep production
indexes, despite the trait’s negative economic weight-
ing. This is due to the strong correlations between
mature size and other weight traits, as well as the high
heritability estimate of the trait. To constrain increases
in ewe mature weight by selecting on the alternative
indexes, Conington et al. (2001) increased the magni-
tude of negative economic weighting on the trait. It
was found that the economic weighting of ewe
mature weight that promotes no further increase in
ewe live weight also reduced predicted progress in
carcass weight and weaning weight to trivial
increases. These results indicate that increased ewe
mature weight is achieved as a consequence of selec-
tion for lamb growth traits in the breeding objective.
Terminal indexes in New Zealand and Ireland were
moderately correlated (0.66) which is driven by the
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B. F. S. Santos et al.
contrasting relative emphasis on muscle traits, 44.8%
in TermNZ and 11.4% in TermIRE, respectively, in
addition to the fact that the TermIRE includes traits
like fatness and lambing ease, which are not consid-
ered in TermNZ index. The general aim of the breeding
objectives of both indexes is similar; however, the
specific traits are quite distinct. One of the examples
of these differences is growth, where the TermNZ
index adopts carcass weight and weaning weight. In
Ireland the economic selection indexes apply an eco-
nomic value to days to slaughter, a trait created from
lamb live weight records (Pabiou et al. 2014). In cat-
tle, carcase weight is a trait commonly used to mea-
sure increases in growth rate (Amer et al. 2001). In
sheep, Thorsteinsson & Bjӧrnsson (1982) reported
genetic correlation of 0.60 between carcass weight
and live weight in twin ram lambs.
Maternal selection indexes in New Zealand and Ire-
land are largely driven by traits that influence the
availability of total weight of lambs for sale, like
growth and to a lesser extent, reproduction and sur-
vival. In this study, when a trait which is related to
utilization of resources, such as ewe mature weight, is
included in the maternal indexes, the total emphasis
of the selection index on these traits represents up to
80% of the selection response. This indicates that eco-
nomically important traits have larger relative empha-
sis when compared to other index component traits.
The total economic response to selection on the
MatNZ index was € 1.16 when compared to the
TermNZ at € 1.07. The New Zealand indexes result in
higher calculated economic responses to selection
when compared to the MatIRE and TermIRE indexes, €
0.27 and € 0.28 per year, respectively. The higher
overall economic responses to selection in the New
Zealand indexes are probably due to the differences in
the cost structure of the production system when
compared to the Ireland. In Ireland (Byrne et al.
2010), sheep farm costs are higher than in New Zea-
land so reducing costs, for instance by reducing
mature ewe size, is probably more important because
of the higher feed costs.
The progress made in the sheep industries in New
Zealand and Ireland is, in part, due to advances in
their breeding structures. In both countries, the aim is
to produce low cost, easy care sheep with good mater-
nal characteristics that produce quality lambs with
high growth rates. International collaboration
between countries with similar breeding objectives
represents an opportunity to generate useful genetic
parameters for sheep breeding. Trait heritabilities,
accuracies, correlations, variance components and the
extent of G9E interactions, assuming livestock have
© 2015 Blackwell Verlag GmbH • J. Anim. Breed. Genet. 132 (2015) 144–154
Comparison of breeding objectives
been traded between countries, are some of these
parameters.
An attempt to assess the extent of the suitability
of New Zealand genetics for Irish grass-based pro-
duction systems has been started through the estab-
lishment of a new comparison flock in Ireland at
the Teagasc Animal & Grassland Research and Inno-
vation. As part of this study, a nucleus sheep flock,
comprising elite Suffolk and Texel females repre-
senting the top genetic merit animals in the Irish
and New Zealand genetic evaluations across a range
of maternal traits, will be evaluated (Pabiou et al.
2014). The collaborative work between countries is
a first initiative to establish a future across country
genetic evaluation system for sheep involving a
number of different countries. The direct relevance
of these results to the Irish sheep industry is that,
based on the moderate to strong correlations
between selection indexes, high genetic merit New
Zealand sheep should be suitable for the Irish pro-
duction systems and this opportunity could be pur-
sued. Future research is recommended to predict
the extent to which G x E is likely to affect
imported gene stocks and the economic impact this
might have when considering importation, as a
potential means of accelerating genetic progress.
Conclusion
Results of this study indicate that terminal and mater-
nal selection indexes in Ireland and New Zealand
moderately to strongly correlate both within and
between countries. Moreover, trait responses to selec-
tion on the four indexes are in a similar direction, but
of different magnitudes, indicating that the similarities
and relative emphasis on traits in the breeding objec-
tives could lead to further selection of populations to
achieve adequate performance in both environments.
The proposed methodology is robust for calculating
correlations between indexes in different countries
and future studies should be carried out to determine
the extent to which G x E is expressed in the follow-
ing generations.
Acknowledgements
The authors would like to acknowledge Teagasc,
Sheep Ireland and the Irish Department of Agriculture
for the collaboration and sponsoring of the present
research project. This manuscript is dedicated to John
W. James and the significant contribution of his work
to the science of definition and implementation of
breeding objectives in livestock production.
153
 Comparison of breeding objectives
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