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J. Anim. Sci. 2003. 81:1700-1705
© 2003 American Society of Animal Science

Genetic correlations between lean growth and litter traits in U.S. Yorkshire, Duroc, Hampshire, and Landrace pigs1

P. Chen*, T. J. Baas*,2, J. W. Mabry* and K. J. Koehler{dagger}

* Department of Animal Science and and {dagger} Department of Statistics, Iowa State University, Ames 50011

2 Correspondence:
109 Kildee Hall (phone: 515-294-6728; fax: 515-294-5698; E-mail:
tjbaas{at}iastate.edu).


    Abstract
 Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 
The objective of this study was to estimate breed-specific genetic correlations between lean growth and litter traits for four U.S. swine breeds. Records for lean growth and litter traits on Yorkshire, Duroc, Hampshire, and Landrace pigs collected between 1990 and April 2000 in herds on the National Swine Registry Swine Testing and Genetic Evaluation System were analyzed. A bivariate animal model and restricted maximum likelihood procedures were used to estimate genetic and environmental correlations between lean growth rate, days to 113.5 kg, backfat, and loin muscle area with litter traits of number born alive, litter weight at 21 d, and number weaned. Most genetic correlation estimates between lean growth and litter traits were small in magnitude and consistent across breeds. Backfat had the largest within-breed genetic correlations with number born alive (0.18 to 0.20) and litter weight at 21 d (-0.27 to -0.30). Estimates of genetic correlations between lean growth traits and number weaned were very small. Estimates of the environmental correlations between lean growth and litter traits also were very small for all traits and for all four breeds. Results indicate that selection for lean growth traits could have a long-term effect on litter traits. Including lean growth traits in a maternal-line evaluation using a multiple-trait model could increase the accuracy of the genetic evaluation for litter traits.

Key Words: Genetic Correlations • Growth • Leanness • Litter Traits • Pigs


    Introduction
 Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 
The genetic improvement of both lean growth and litter traits is important to increase the efficiency of pork production. However, in general, there are negative relationships between lean growth and litter traits (Clutter and Brascamp, 1998). Multiple-trait BLUP EBV has been widely used in swine genetic evaluations for lean growth and litter traits (Hofer et al., 1992; Kennedy et al., 1996). Reliable estimates of the genetic correlations between lean growth and litter traits are required for optimal use of BLUP procedures. Currently, most genetic evaluation programs evaluate lean growth and litter traits separately using multiple-trait models, and their EBV are then combined into a bio-economic index using appropriate economic values (NSR, 2000). Thus, the genetic correlation between lean growth and litter traits is not taken into account in current evaluations. In order to increase the accuracy of evaluations, especially for traits with low heritability (e.g., litter traits), it may be necessary to evaluate lean growth and litter traits jointly using multitrait analyses.

Numerous researchers have investigated the genetic correlations between lean growth and litter traits in different populations. Clutter and Brascamp (1998) reviewed these genetic parameters and concluded that, in general, estimates have been plagued by insufficient precision. Because they are population specific, use of parameter estimates from the literature could be very misleading. Genetic correlations between lean growth and litter traits currently recommended by the National Swine Improvement Federation (NSIF, 1997) for genetic evaluation programs are based on results from the literature and are not breed specific. Therefore, the objective of this study was to estimate breed-specific genetic correlations between lean growth and litter traits for the U.S. Yorkshire, Duroc, Hampshire, and Landrace populations.


    Materials and Methods
 Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 
Data Source
Lean growth and litter data were obtained from the National Swine Registry on Yorkshire, Duroc, Hampshire, and Landrace pigs. Litter records on 251,296 Yorkshire, 75,262 Duroc, 83,338 Hampshire, and 53,234 Landrace litters born between 1984 and April 1999 were included. Lean growth records on 361,300 Yorkshire, 154,833 Duroc, 99,311 Hampshire, and 71,097 Landrace pigs were collected between 1985 and April 2000. Most females with litter records also had lean growth data in the analysis. Complete details of data collection can be found in the Swine Testing and Genetic Evaluation System (STAGES; NSR, 2000). Lean growth data included pedigree information for each pig, contemporary group (CG), sex of the pig, litter identification, birth date, date weighed, and measurements for weight, backfat (BF), and loin muscle area (LMA) at an approximate weight of 113.5 kg. Contemporary groups were defined by breeders as a group of pigs that were raised under the same management and environmental conditions. Data on boars, gilts, and barrows were included in the lean growth data set. Backfat and LMA were measured ultrasonically at the 10th rib. Days to 113.5 kg (DAYS), BF, and LMA were adjusted using recommendations in the Guidelines for Uniform Swine Improvement Programs (NSIF, 1997). Kilograms of lean was estimated using the following fat-free lean prediction equation developed by the National Pork Producers Council (NPPC, 2000):


Then, lean growth rate per day (LGR) was calculated by dividing by DAYS.

Litter data included pedigree information for each sow, CG, parity of the sow, farrowing and weighing dates, number born alive (NBA), number after transfer (NAT), number weaned (NW), and litter weight at 21 d (L21WT). Number weaned included the actual number of pigs weaned by each sow but excluded pigs transferred to other sows. Contemporary groups were defined by the breeders as a group of sows that were bred, gestated, and farrowed in a group under the same management and environmental conditions. Litter traits were adjusted according to breed-specific procedures developed by Culbertson et al. (1997) from data for these breeds from 1985 to 1996. Adjustments for NBA included parity and age at farrowing. Litter weight at 21 d was adjusted for parity, age at farrowing, age at weighing, and NAT. Number weaned was adjusted for parity and NAT.

In both data sets, single-sire CG records were removed, as were records from sires not connected across CG and sires not mated to more than one dam. Numbers of records and CG, and litters represented by breed in the lean growth data set, along with means and SD for LGR, DAYS, BF, and LMA, can be found in Chen et al. (2002a). Number of records, animals, sires, dams, service sires, and contemporary groups represented by breed in the litter data, along with means and SD for NBA, L21WT, and NW can be found in Chen et al. (2002b).

Statistical Analysis
Bivariate REML analyses were conducted to estimate genetic correlations between lean growth and litter traits using REMLF90 (Misztal, 2000). Each analysis contained one lean growth trait and one litter trait. Different fixed and random effects, along with the environmental correlation between traits, were considered in two models. Models used for bivariate analyses were as follows:


where y1, y2 = observations for lean growth and litter traits, b1, b2 = fixed effects of contemporary group and sex for lean growth traits and fixed effects of contemporary group for litter traits, a1, a2 = random additive genetic effects of animals, c = common litter effects, ss = service sire effects, pe = permanent environmental effects, X1, X2 = incidence matrices relating to fixed effects, Z1, Z2 = incidence matrices relating to animal effects, S = incidence matrix relating to common litter effects, W1, W2 = incidence matrices relating to service sire and permanent environmental effects, e1, e2 = residual effects for lean growth and litter traits. It was assumed that the covariances between additive, common litter, service sire, permanent environmental, and residual effects were zero and that levels of each were independently distributed with variances and and covariance {sigma}a12 for animal effects, for common litter effects, for service sire effects, for permanent environmental effects, and and and covariance {sigma}e12 for residual effects. Standard errors of genetic correlations were estimated using formulas by Falconer (1989). In the REML analysis, the convergence criterion was set to 10-8 for all analyses.


    Results and Discussion
 Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 
Estimates of variance components for lean growth and litter traits are similar to the results from univariate analyses given in the previous papers (Tables 1Go and 2Go) in this series (Chen et al., 2002a, b). Estimates of covariances between lean growth and litter traits are in Table 3Go. The Landrace had the highest absolute value of genetic covariances between BF, LMA, and LGR with NBA; between BF and L21WT; and between DAYS and BF with NW. The Duroc breed had the lowest absolute value of genetic covariances between LMA and LGR with NBA; between DAYS, BF and LGR with L21WT; and between DAYS and NW. In general, the genetic covariances were consistent among the four breeds.


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Table 1. Estimates of (co)variance components and genetic parameters from univariate analyses for litter traits by breed using a model with maternal genetic effectsa
 

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Table 2. Estimates of (co)variance components and genetic parameters from univariate analyses for lean growth traits by breed using a model with maternal genetic effectsa
 

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Table 3. Estimates of genetic ({sigma}a12) and residual covariance ({sigma}e12) of lean growth traits with litter traits by breed
 
Estimates of correlations between lean growth and litter traits are in Table 4Go. In general, estimates of genetic correlations for the maternal breeds of Yorkshire and Landrace were very similar, and differed only slightly for the terminal breeds of Duroc and Hampshire. Genetic correlations between DAYS and litter traits were low in magnitude. Estimates of the genetic correlation of DAYS with NBA are -0.041, 0.051, -0.072, and -0.061 for the Yorkshire, Duroc, Hampshire, and Landrace breeds, respectively. Crump et al. (1997) reported an estimate of 0.084 for the genetic correlation between ADG and NBA in British Landrace pigs. Short et al. (1994) reported positive values for three of the four estimates of the genetic correlation between ADG and total number born in two dam lines from two farms (0.04, 0.05, 0.23, and -0.15). The estimates from this study are also lower than the value of -0.20 recommended by NSIF (1997).


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Table 4. Estimates of genetic (rg) and residual correlations (re) of lean growth with litter traits
 
Estimates from this study revealed positive genetic correlations between DAYS and L21WT for three breeds, the exception being the Hampshire breed (Table 4Go). The largest absolute value of the estimate of the genetic correlation of DAYS with L21WT was 0.095 in the Hampshire breed. Estimates of the genetic correlation of DAYS with L21WT from this study are much lower than the value of 0.43 recommended by NSIF (1997). Estimates of the genetic correlation between DAYS and NW are low and smaller in magnitude than their standard errors, which supports the value of 0 recommended by NSIF (1997). These correlations may be biased due to crossfostering of piglets, which occurred in 50% of the litters in this study. Even though the correlations between DAYS and litter traits are small, the signs indicate that selection for increased gain could slightly increase litter size but decrease litter weight in most breeds.

Estimates of genetic correlations between BF and NBA and L21WT are relatively high and consistent across breeds (Table 4Go). Estimates of the genetic correlation of BF with NBA are 0.19, 0.18, 0.18, and 0.20 for the Yorkshire, Duroc, Hampshire, and Landrace breeds, respectively. These results agree with the value of 0.21 reported by Crump et al. (1997) in British Landrace pigs. Johansson and Kennedy (1983) also reported positive estimates of 0.13 to 0.22 for BF with NBA in Swedish Landrace and Yorkshire pigs. Löbke et al. (1986) reported positive correlations of 0.03 and 0.28 for BF with NBA for the first litter and the first three litters, respectively. However, Morris (1975) and Bereskin (1984) reported negative genetic correlations between BF and litter size. Short et al. (1994) also reported three of four estimates of the genetic correlation between BF and total number born were negative, -0.12, -0.03, -0.08, and 0.06. The estimates from this study disagree with the value of 0.00 recommended by NSIF (1997). Estimates of the genetic correlation between BF and L21WT ranged from -0.27 to -0.30 for the four breeds (Table 4Go). These estimates for each of the breeds are lower than the value of -0.40 recommended by NSIF (1997). Estimates of the genetic correlation between BF and NW are very low and smaller than their standard errors (Table 4Go), which supports the value of 0 recommended by NSIF (1997). The signs of the correlations indicate that selection for decreased BF could slightly decrease litter size but increase litter weight.

Estimates of genetic correlations between LMA and litter traits are also low in magnitude and consistent across breeds. Estimates of the genetic correlation between LMA and NBA are lower than their standard errors and averaged -0.020 across the four breeds. Estimates of the genetic correlation between LMA and L21WT are -0.054, 0.083, -0.031, and -0.017 for the Yorkshire, Duroc, Hampshire, and Landrace breeds, respectively (Table 4Go). Estimates of the genetic correlation between LMA and NW are very low and smaller in magnitude than their standard errors (Table 4Go). There are no previously reported estimates of genetic correlations between LMA and litter traits.

Nearly all of the genetic correlations for LGR with litter traits were negative. Estimates of the genetic correlation of LGR with NBA were low in magnitude and consistent across the four breeds. Estimates of the genetic correlation of LGR with L21WT were also low and averaged -0.060 across the four breeds. The genetic correlations between LGR and NW were very low and smaller than their standard errors (Table 4Go). These genetic correlations may be biased due to crossfostering of piglets between litters that was present in the data. Chen et al. (2001) reported negative genetic correlations of -0.18 and -0.05 for LGR with NBA and NW, respectively, and a positive correlation of 0.13 for LGR with L21WT from a selection experiment in a synthetic line of Yorkshire-Meishan pigs.

Environmental correlation estimates are very low across all traits and all breeds. This result agrees with the findings of Crump et al. (1997), who reported that environmental correlations between performance and litter traits were, in general, low.

This study was the first to provide breed-specific genetic correlation estimates between lean growth and litter traits based on a large amount of field data. These breed-specific estimates could be used for jointly evaluating lean growth and litter traits using multiple-trait analyses in genetic evaluation programs. They would be applicable when litter traits are included in the selection objective for the breeds evaluated.

Currently, the STAGES program from which these data were obtained evaluates lean growth and litter traits in two separate multiple-trait evaluations. One of the reasons for including the relationship between lean growth traits and litter traits in a maternal-line evaluation is to increase the accuracy of the genetic evaluation for litter traits. Analyzing all traits simultaneously would, however, require an increase in computer resources. This tradeoff between accuracy and the need for additional inputs would need to be considered before implementation.

In general, the genetic correlations between lean growth and litter traits are unfavorable but relatively low in magnitude. This negative relationship may, in part, help explain the relatively slow genetic progress in litter traits in these populations demonstrated by Chen et al. (2002b). It also indicates that long-term selection for LGR could possibly harm litter traits. Because the genetic correlations and heritabilities for litter traits are low, the effect of selection for lean growth traits on litter traits without accounting for these correlations would not be observed in the short term.

As selection for lean growth traits continues, it is unknown whether relationships between them in these populations will change. It is possible that genetic relationships between components of LGR and litter traits are not linear, and that correlations may change as lean growth traits reach new thresholds. Therefore, it will be necessary to evaluate these relationships periodically and incorporate them into the evaluation program.


    Implications
 Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 
Unfavorable genetic correlations between lean growth and litter traits indicate that long-term selection for lean growth traits could harm litter traits if selection is practiced for many generations and these relationships are ignored. Although breed-specific estimates for genetic correlations between lean growth and litter traits were found to be low, including them in maternal-line evaluations using a multiple-trait model within breed could increase the accuracy of the genetic evaluation for litter traits. These relationships should be evaluated periodically, however, if selection for lean growth traits continues.


    Footnotes
 
1 Journal paper No. J-19847 of the Iowa Agric. and Home Econ. Exp. Stn., Ames, Project No. 3456, and supported by Hatch Act and State of Iowa funds. Back

Received for publication August 2, 2002. Accepted for publication March 4, 2003.


    Literature Cited
 Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 


Bereskin, B. 1984. Genetic correlations of pig performance and sow productivity traits. J. Anim. Sci. 59:1477–1487.

Chen, P., T. J. Baas, J. C. M. Dekkers, and L. L. Christian. 2001a. Selection for lean growth rate and correlated responses in litter traits in a synthetic line of Yorkshire-Meishan pigs. Can. J. Anim. Sci. 81:205–214.

Chen, P., T. J. Baas, J. W. Mabry, J. C. M. Dekkers, and K. J. Koehler. 2002a. Genetic parameters and trends for lean growth rate and its components in U.S. Yorkshire, Duroc, Hampshire, and Landrace pigs. J. Anim. Sci. 80:2062–2070.[Abstract/Free Full Text]

Chen, P., T. J. Baas, J. W. Mabry, J. C. M. Dekkers, and K. J. Koehler. 2002b. Genetic parameters and trends for litter traits in U.S. Yorkshire, Duroc, Hampshire, and Landrace pigs. J. Anim. Sci. 81:46–53.

Clutter, A. C., and E. W. Brascamp. 1998. Genetics of performance traits. Pages 427–462 in The Genetics of the Pig. M. F. Rothschild and A. Ruvinsky, ed. CAB International, New York.

Crump, R. E., C. S. Haley, R. Thompson, and J. Mercer. 1997. Individual animal model estimates of genetic correlations between performance test and reproduction traits of Landrace pigs performance tested in a commercial nucleus herd. Anim. Sci. 65:291–298.

Culbertson, M. S., J. W. Mabry, J. K. Bertrand, and A. H. Nelson. 1997. Breed-specific adjustment factors for reproductive traits in Duroc, Hampshire, Landrace, and Yorkshire swine. J. Anim. Sci. 75:2362–2367.[Abstract/Free Full Text]

Falconer, D. S. 1989. Introduction to Quantitative Genetics. Longman, Inc., New York.

Hofer, A., C. Hagger, and N. Künzi. 1992. Genetic evaluation of on-farm tested pigs using an animal model. II. Prediction of breeding values with a multiple trait model. Livest. Prod. Sci. 30:83–97.

Johansson, K., and B. W. Kennedy. 1983. Genetic and phenotypic relationships of performance test measurements with fertility in Swedish Landrace and Yorkshire sows. Acta Agric. Scand. 30:195–199.

Kennedy, B. W., K. W. Quinton, and C. Smith. 1996. Genetic changes in Canadian performance-tested pigs for fat depth and growth rate. Can. J. Anim. Sci. 76:41–48.

Löbke, A., H. Willeke, and F. Pirchner. 1986. Relationship between reproductive performance and growth and backfat. Proc. 37th Annu. Mtg. Eur. Assoc. Anim. Prod., Budapest, Hungary.

Misztal, I. 2000. Computational Techniques in Animal Breeding. Univ. of Georgia, Athens.

Morris, C. A. 1975. Genetic relationships of reproduction with growth and with carcass traits in British pigs. Anim. Prod. 20:31–44.

NPPC. 2000. Composition and Quality Assessment Procedures. Natl. Pork Prod. Council, Des Moines, IA.

NSIF. 1997. Guidelines for Uniform Swine Improvement Programs. USDA, Washington, DC.

NSR. 2000. STAGES National Genetic Evaluation Trait Leader List Winter 2000. Natl. Swine Registry, West Lafayette, IN.

Short, T. H., E. R. Wilson, and D. G. McLaren. 1994. Relationships between growth and litter traits in pig dam lines. Pages 413–416 in Proc. 5th World Congr. Genet. Appl. Livest. Prod., Guelph, Ontario, Canada.

Vangen, O. 1980. Studies on a two trait selection experiment in pigs. V. Correlated responses in reproductive performance. Acta Agric. Scand. 30:309–319.


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