Estimated Variance Components and Breeding Values for Pre-Weaning Growth Criteria in Romney Sheep

Faid-Allah E, Ghoneim E, Ibrahim AHM. 2016. Estimated variance components and breeding values for pre-weaning growth criteria in Romney sheep. JITV 21(2): 73-82. DOI: http://dx.doi.org/10.14334/jitv.v21i2.1353 This study was carried out to investigate variance components, direct, maternal genetic parameters, estimated breeding values and factors affecting pre-weaning growth criteria of Romney sheep. Data were collected over the period from 2006 to 2012 with records of 4989 lambs descended from 76 rams and 2190 ewes of Romney sheep maintained at S. Island of New Zealand via Gene Marker Lab., Faculty of Agric. and Life Sci., Lincoln Univ., New Zealand. Results proved that genetic and non-genetic factors affecting studied criteria had significant effects (P<0.05). Genetic and environmental estimates of live body weights at birth (LBW), weaning (LWW) and Kleiber ratio (KR) were 0.20±0.074, 0.15±0.042 and 0.14±0.052 for direct heritability (ha±SE); 0.59±0.219, 0.41±0.023 and 0.08±0.002 for maternal heritability (hm±SE); 0.11684, 2.6378 and 0.27565 for additive variances (σa), 0.34596, 7.1179 and 0.14532 for maternal variances (σm); and 0.002395, 10.1262 and 0.509339 for permanent environmental variances (σe), respectively. EBV’s of LBW, LWW and KR ranged from -0.555: 0.502, -1.554: 3.006 and -0.633: 0.242 direct, -0.863: 0.954, -4.942: 2.554 and -0.469: 0.179 maternal for rams, respectively; and -0.664: 0.830, 2.996: 4.586 and -1.651: 0.677 direct, 1.429: 1.142, -7.541: 4.920 and -1.223: 0.492 maternal for ewes, respectively. Results suggest the importance of considering the non-genetic factors in pre-weaning growth performance of lambs. Moderate heritability and positive coefficients of phenotypic and genetic correlation for studied criteria indicate to the possibility of improving them using traditional selection.


INTRODUCTION
Growth potential of lambs is very important in the sheep production.It is essential to have knowledge of genetic parameters for these economically important criteria to formulate breeding strategies for better production (Gowane et al. 2015).The relative importance of direct and maternal additive genetic effects for growth should be considered when sheep producers formulate their breeding plans (Talebi et al. 2007).Maternal genetic effects arise from allelic differences between individual mothers at loci influencing offspring phenotype and are a heritable component of phenotypic variance themselves.It can dampen response to the selection, when direct-maternal genetic covariance is negative, or accelerate, when direct-maternal genetic covariance is positive (Wolf et al. 1998).Maternal care and genetic potential of lambs are two factors affecting the early growth of lambs.In the pre-weaning period, animals are fed by their mother's milk (Ghafouri-Kesbi 2013).Growth criteria are influenced by many factors includes additive genetic effects, maternal genetic effects and non-genetic factors (Farokhad et al. 2010).Adjustments of data for environmental factors, such as year or/and season of birth, parity of ewe, type of birth, sex of lamb and location are necessary to obtain reliable estimates and to increase the accuracy of selection of breeding animals (Thiruvenkadan et al. 2011).Studies of estimated breeding value, variance components, genetic parameters, and factors affecting growth criteria in New Zealand Romney sheep were infrequent for the last decade.However, Sheep production has been the important source of sustainable livelihood of the rural folk in New Zealand.
A strategy to increase the efficiency of meat production in both the traditional and intensive systems is the selection of animals regarding the efficiency of feed utilization.Since individual sheep differ in their ability to utilize feed efficiently, selecting the most efficient animals, those with lower maintenance requirements, results in a significantly lower production cost (Ghafouri-Kesbi et al. 2011).Direct selection for lower maintenance requirements is difficult.However, measures of feed efficiency such as Kleiber ratio would be used to achieve this goal (Arthur et al. 2001).Estimated Breeding Value (EBV) is a prediction of the genetic merit of an animal for selection criteria such as growth, milk, and prolificacy.EBVs increase the accuracy of selection for superior performance.
This study investigated genetic and non-genetic factors affecting pre-weaning growth criteria of New Zealand Romney sheep breed and estimate its breeding values, variance components, direct, and maternal genetic parameters.

Animal data
Data were collected over the period from 2006 to 2012 except 2011 with records for studied criteria on total of 4989 pedigree lambs descended from 76 rams and 2190 ewes from 14 commercial farms of Romney sheep maintained at S. Islands of New Zealand via Gene Marker Lab., Faculty of Agric.and Life Sciences, Lincoln Univ., New Zealand.The pedigree data included birth and weaning weight and its dates, sex of lamb, ewe parity, and type of birth.Lambs were weaned at about three months of age.Studied pre-weaning growth criteria were live body weights at birth (LBW), weaning (LWW) and Kleiber ratio (KR = ADG_LWW/LWW 0.75 ).Kleiber ratio is defined as growth rate divided by body mass powered 0.75 [metabolic weight] (Kleiber 1947).The breeding season started from February to late May.Those selected rams and ewes were divided into mating groups to avoid inbreeding.Ewes were joined in a pen with a single ram in a group of 40-50 ewes.After mating, ewes were separated from rams and kept as one group until the lambing season which usually starts in August to October.

Statistical analysis
Data were analyzed by general linear model via Statistical Analysis System [SAS] (SAS 2002), to estimate effect of ram, ewe within ram, year of birth, parity of ewe, sex of lamb, type of birth and farm on three pre-weaning growth criteria: LBW, LWW, and KR.Statistical models used to analyze pre-weaning growth criteria may be written as the following formula:  Body weights at birth and weaning in Romney lambs literature were 4.10±0.76,4.32, 4.85±0.607and 5.97±0.02kg of birth weight (Morris et al. 1996;Morris et al. 2000;Everett-Hincks et al. 2014;Ibrahim 2015, respectively); and 28.7±3.1,28.71 and 36.81±0.11kg of weaning weight (Morris et al. 1996;Morris et al. 2000;Ibrahim 2015, respectively).

Genetic and non-genetic factors
Table (2) shows that rams and ewes as random factors significantly (P≤0.01)affected the studied preweaning growth criteria in Romney sheep.Early growth traits were found to be significantly (P≤0.05)affected by ram and ewe in the previous studies carried out by Abbas et al. (2010) in Rahmani and Chios sheep; Esmailizadeh et al. (2011) in Kurdi, Chaal, Afshari and Sanjabi sheep; Màrquez et al. (2012) in Charollais, Suffolk and Texel sheep and Petrović et al. (2013) in Mis and Wurttemberg sheep.In addition, Table 2 shows that all non-genetic effects had significant (P≤0.01)effects on studied criteria.GLM test for these criteria indicated that lamb growth characteristics at preweaning were high values for single lambs, lambs born to third parity ewes.Growth advantage of single born might be resulted from its lower competition to milking and supply from the ewe in gestation period than the multiplex (Mousa et al. 2013).This result is in agreement with Roshanfekr et al. (2011, Arabi); Abbasi et al. (2012, Iranian Baluchi, Kurdish); Everett-Hincks et al. (2014, New Zealand breeds) and Simeonov et al. (2015, Blackhead Pleven).Also, male lambs had higher values for these criteria in comparison with female lambs.The sex difference is consistent as reported by Mohammadi et al. (2010a), Mandal et al. (2012), Mousa et al. (2013) and Everett-Hincks et al. (2014).That studied criteria increased from the 1st to 3rd parity and thereafter decreased substantially.A similar significant effect of the parity of ewe on the body weights at early parities was observed by Thiruvenkadan et al. (2011) and Simeonov et al. (2015).Lower values for pre-weaning weights for lambs born to younger ewes may be attributed to the relative competition for nutrients between the still growing ewes and the developing fetus (Thiruvenkadan et al. 2011), also depression in these traits for the lambs born after the fourth parity may be due to the ewe's tooth decay that results in grazing problems followed by decreasing milk production and maternal care for lambs (Mousa et al. 2013).
Average of each criterion fluctuated from year to year.These differences in LBW, LWW and KR among lambs born in different years may be attributed to the differences in environmental conditions especially the rainfall level affecting quantity and quality of grasses.Our results are in agreement with the results obtained, in different sheep breeds, by Roshanfekr et al. (2011), Abbasi et al. (2012) and Simeonov et al. (2015) in Arabi, Iranian Baluchi and Blackhead Pleven, respectively.
The farm had significant effect on the studied criteria, This effect may be due to the differences in management and ram of service, that agreed with the results of Thiruvenkadan et al. (2011), Everett-Hincks et al. (2014) and Simeonov et al. (2015).and correlation between additive and maternal additive genetic effects (ram) for different criteria.Estimates of variance components showed that the estimates of σ 2 a were nearly half the estimates of σ 2 m indicating the importance of maternal additive genetic effects on the criteria.

Variance components
Also, the σ 2 a was found to account for a small proportion and σ 2 e was found to account for a large proportion of the σ 2 p for all the studied criteria.This result reflects the importance of σ 2 e than σ 2 a which is possibly due to uterine capacity, feeding level at late gestation, and maternal behavior of ewe.Reasonably higher maternal additive effects than direct additive effects were observed by El-Awady et al. ( 2011 heritabilities and genetic (rG) and phenotypic (rp) correlations between the studied criteria are shown in Table ( 4).These presented results showed that the estimation of h 2 a for LBW and LWW were nearly half the estimates of h 2 m, indicating to the importance of maternal effects in sheep which contribute to the dependence of lambs on their mother`s milk until the time of weaning (Bradford 1972).Morris et al. (1996) worked on New Zealand Romney sheep and reported that heritability estimation for LBW and LWW were 0.29±0.05and 0.11±0.05,respectively.Everett-Hincks et al. ( 2014) worked on New Zealand sheep breeds and reported that direct heritability estimates for LBW ranged from 0.126±0.009to 0.138±0.009and maternal heritability estimates for LBW ranged from 0.179±0.009to 0.316±0.01.Ghafouri-Kesbi ( 2013) reported that estimates of direct, maternal heritability, σ 2 a, σ 2 m, σ 2 e and σ 2 p for pre-weaning Kleiber Ratio were 0.13±0.03,0.08±0.03,0.21, 0.13, 1.21 and 1.66, respectively.In addition, there is a positive genetic correlation between KR with traits related to growth (Abegas et al. 2005, Ghafouri-Kesbi et al. 2011).These findings show that improvements in feed efficiency could be made without negatively affecting body weight or growth rate (Ghafouri-Kesbi 2013).These findings are in accordance with other reports (Abegas et al. 2005;Szwaczkowski et al. 2006;Mohammadi et al. 2010b;Ghafouri-Kesbi et al. 2011;Savar-Sofla et al. 2011;Mokhtari et al. 2012).Supakorn et al. (2013) reported that in Thailand sheep populations, estimated direct and maternal heritabilities from multivariate analysis of the model for birth weight were 0.32±0.06and 0.23±0.02,respectively.Aksoy et al. (2016) reported that in Karayaka lamps, the direct heritability of birth weight was 0.44±0.063.When the maternal genetic effects were considered in models, the direct heritability for birth weight decreased from 0.36 to 0.24.The inclusion of the maternal, genetic, and/or environmental effects into the model resulted in a direct additive variance value varied between 0.07 and 0.08.In model considered maternal genetic and environmental effects, the values for the maternal heritability ranged between 0.15 and 0.22.Furthermore, depending on the model used, direct heritability estimates for WW ranged between 0.40 and 0.27.For weaning weight, model 1 provided a direct heritability value of 0.40±0.066.For weaning weight, the maternal heritability values within the range of 0.04-0.14were estimated.Meyer (1992) suggested that models not considering maternal genetic effects could result in substantially higher estimates of σ 2 a and h 2 a.Also, the estimates of h 2 a and h 2 m for LBW were higher than their corresponding estimates for LWW.The decrease in the estimation of both h 2 a and h 2 m by age was in the line with the results of El-Awady et al. ( 2011) and Mousa et al. (2013).Table 4 shows that estimates of direct and maternal heritability for pre-weaning Kleiber Ratio were 0.14 ±0.052 and 0.08 ±0.002, respectively; which ranged between 0.01-0.15as reported for different sheep breeds (Mohammadi et al. 2010b;Ghafouri-Kesbi et al. 2011;Savar-Sofla et al. 2011;Mokhtari et al. 2012).Furthermore, Kleiber Ratio estimates of heritability ranged from 0.04 (Arman sheep, Mokhtari et al. 2012) to 0.15 (Sanjabi sheep, Mohammadi et al. 2010b).Estimates of heritability for a criterion may differ between sheep breeds and change slowly over time.Some literature of heritability estimation revealed that in sheep, the Kleiber ratio was a low heritable criterion.
Generally, the obtained results fell in the range reported in the literature for different breeds of sheep maintained at different locations and regions around the world.Range of h 2 a estimates for these criteria in literature varies substantially from 0.004 in Barbary sheep (Bedhiaf et al. 2000) to 0.94 in Hissardale sheep (Chaudhry & Shah 1985) for LBW; from 0.007 in Barbary sheep (Bedhiaf et al. 2000) to 0.81 in Multibreed meat sheep (Lobo et al. 2009) for LWW.Likewise, the estimates of h 2 m ranged from 0.02 in Dorper sheep (Neser et al. 2001) to 0.65 in Sangsari sheep (Miraei-Ashtiani et al. 2007) for LBW and from 0.01 in Ghazel sheep to 0.48 in Dorper sheep (Assan et al. 2011) for LWW and from 0.01 in Romanov sheep (Maria et al. 1993) to 0.07 in crossbreed sheep (Hall et al. 1995).Furthermore, direct estimates of heritability tend to be higher than maternal for early growth traits (Hassen et al. 2003).The range of direct heritability estimates for birth weight from multivariate analysis in the literature varies substantially from 0.04 in Romanov and Kermani breeds (Maria et al. 1993;Rashidi et al. 2008) to 0.46 in Menz breed (Gizaw et al. 2007).Talebi et al. (2007) reported that estimates of heritability for LBW and LWW of Lori-Bakhtiari sheep were 0.30±0.03and 0.13±0.03direct and 0.22 and 0.17 maternal, respectively.Consequently, genetic progress is possible for growth traits by selection.

Genetic and phenotypic correlations
Table 5 presents the genetic correlations for LBW with both LWW and KR were 0.325 and 0.048, respectively, however, the phenotypic correlations between studied criteria ranged from 0.196 to 0.597 as positive and significant coeffecients.Estimates of genetic correlations between preweaning criteria in the current study were positive and moderate.On the other hand, the estimation of genetic correlations for the birth weight ranged from -0.078±0.0084 to -0.127±0.0450as reported by (Everett-Hincks et al. 2014).
Correlation coefficients were 0.38, and ranged from 0.39 to 0.41 for phenotypic correlation and 0.40, and ranged from 0.40 to 0.81 for genetic correlation in breeds of Suffolk and Baluchi respectively (Yamaki 1994;Yazdi et al. 1997).In general, the estimates of genetic correlations were agree with those reported by El-Awady et al. (2011) in Egyptian Rahmani lambs and lower than those reported by Rashidi et al. (2008); Roshanfekr et al. (2011);Prakash et al. (2012) in Arabi, Kermani and Malpura breeds of sheep, respectively.

Estimated breeding value
Estimated Breeding Value (EBV) are expressed as deviations from population mean and sires were ranked based on their genetic merit.For low heritable below 0.15 criteria, an animal's performance is much less useful for identifying the individuals with the best genes for the trait.For this reason, selection for studied traits should be according to estimates of breeding values.Estimates of maternal heritability were high for preweaning criteria as reveal in Table 4.It shows that genes contributing to maternal performance have an equal influence on the early growth of lambs as genes carried by lambs.If maternal genetic effects are not considered, true genetic potential of lambs is masked by maternal performance making it difficult to select superior animals.As a result, maternal effects should be included in the model to obtain accurate estimates of genetic parameters and breeding values, in particular where animals are evaluated for criteria related to early growth.
Table 6 presents the mean, SD, minimum and maximum for direct and maternal estimated breeding values for LBW, LWW and Kr in rams and ewes of Romney sheep.The mean, SD (Min : Max) for Rams were 0.0023±0.1998(-0.5546 : 0.5017) direct and -0.0039±0.3437(-0.8632 : 0.9543) maternal EBV for LBW, respectively; were 0.1984±0.7529(-1.5535 : 3.0056) direct and -0.3262±1.2379(-4.9417 : 2.5542) maternal EBV for LWW, respectively; and were -0.0226±0.1330(-0.6333 : 0.2417) direct and -0.0165±0.0983(-0.4689 : 0.1790) maternal EBV for KR, respectively.The estimated breeding values for rams had a wide range of all pre-weaning criteria, so we may select superior rams and using the selected animals in mating as parents of next generation to make genetic improvement in pre-weaning as a selection objective.

CONCLUSION
According to the results, it is concluded that the non-genetic factors have the main source of variation for pre-weaning criteria.Estimation of heritability and variance components of the studied criteria proved the importance of maternal effect.Birth weight, weaning weight and Kleiber ratio have been reported as moderately heritable criteria.Positive genetic correlations among these criteria indicate to possibility of improving these criteria using selection program.Furthermore, Kleiber ratio has been recommended as an indirect selection parameter for feed conversion.

Table 1 .
Descriptive statistics of pre-weaning growth criteria in Romney sheep # standard deviation, ## coefficient of variability

Table 2 .
Factors affecting pre-weaning growth criteria in Romney lambs

Table 3 .
Variance components for pre-weaning growth criteria in Romney sheep

Table 4 .
Heritability and additive maternal genetic correlation for pre-weaning growth criteria in Romney sheep

Table 5 .
Genetic(above)and phenotypic (below) correlations between pre-weaning growth criteria in Romney sheep

Table 6 .
Estimated breeding values for pre-weaning growth criteria in Romney sheep