Production Traits In Dairy Cattle Research Articles

Genetic effects of stearoyl-coenzyme A desaturase (SCD) polymorphism on milk production traits in the Chinese dairy population

Stearoyl-CoA desaturase (SCD) is a multifunctional complex enzyme important in the cellular biosynthesis of fatty acids. The present study was to investigate the association of the SCD gene with milk production traits in dairy cattle. Two single nucleotide polymorphisms (SNPs) (g.6926A>G and g.8646A>G) in introns 3 and 4, and three SNPs (g.10153A>G, g.10213T>C and g.10329C>T) in exon 5 were identified with pooled DNA sequencing and genotyped using matrix-assisted laser desorption/ionization time of flight mass spectrometry assay in 752 Chinese Holstein cows. Polymorphism g.10329C>T was predicted to result in an amino acid replacement from alanine to valine in the SCD protein. With a mixed animal model, the significant associations of the five SNPs with 305-day milk, fat and protein yields and protein percentage were determined. We further demonstrated cows with heterozygous genotypes (A/G or C/T) had highest 305 day milk yield, fat yield, protein yield and lowest protein percentage. Heterozygous cows with genotype AG at the g.6926A>G locus showed the greatest milk yield (P < 0.0001), fat yield (P < 0.0001) and protein yield (P < 0.0001) among other heterozygous genotypes at any of the loci. Dominance effects of all identified SNPs on milk, fat and protein yields and protein percentage were significant. Moreover, significant allele substitution effects at g.6926A>G locus on milk yield and at g.10213T>C on protein yield were observed. Five-locus haplotypes and strong linkage disequilibrium (D' > 0.9) between the five SNPs were also observed. The results suggest that identified polymorphisms could be potential genetic markers to improve the production performance of Chinese Holstein.

Effects of SCD1- and DGAT1-genes on production traits of Chinese Holstein cows located in the Delta Region of Yangtze River

Associations between the stearoyl-CoA desaturase 1 (SCD1) gene and the Diacylglycerol-O-acyltransferase 1 (DGAT1) polymorphisms with milk production traits were analyzed using data from 1061 Chinese Holstein cows located in the Delta Region of Yangtze River in China. Milk production traits included test day observations for milk yield (TDMY), fat content (FC), protein content (PC), lactose content (LC), total solid (TS), milk urea nitrogen (MUN), somatic cell score (SCS), as well as lactation records, i.e. 305-day FCM yield, 305-day fat yield and 305-day protein yield. A total of 8,162 test day records were analyzed using a linear mixed model including the fixed effects of sire, herd, test year, test season, parity, calving season, milking day, gene locus, and the random effect of the cow. The effects of the SCD1 genotype on TDMY, FCM (fat corrected milk yield), FC, PC, SCS and 305-day FCM yield were analyzed. Cows with genotype AA produced significantly higher TDMY and FCM compared to cows with genotype AV or VV, but had lower values for FC and PC. Furthermore, the effects of the DGAT1 genotype on TDMY, FCM, FC, PC, TS, 305-day FCM, 305-day fat yield and 305-day protein yield were analyzed. Association studies revealed significantly lower TDMY, FCM and 305-day FCM yield for cows with genotype KK compared to genotypes KA or AA. In contrast, the KK genotype was superior with regard to FC, PC and TS. Cows with genotype KA produced higher 305-day fat yield and 305-day protein yield compared to genotypes AA or KK. These findings suggest a possible use of SCD1 and DGAT1 loci in marker-assisted selection programs for the improvement of milk production traits in dairy cattle from the Delta Region of Yangtze River in China.

Validation of single nucleotide polymorphisms associated with milk production traits in dairy cattle

Single nucleotide polymorphism (SNP) associations with milk production traits found to be significant in different screening experiments, including SNP in genes hypothesized to be in gene pathways affecting milk production, were tested in a validation population to confirm their association. In total, 423 SNP were genotyped across 411 Holstein bulls, and their association with 6 milk production traits—Australian Selection Index (indicating the profitability of an animal's milk production), protein, fat, and milk yields, and protein and fat composition—were tested using single SNP regressions. Seventy-two SNP were significantly associated with one or more of the traits; their effects were in the same direction as in the screening experiment and therefore their association was considered validated. An over-representation of SNP (43 of the 423) on chromosome 20 was observed, including a SNP in the growth hormone receptor gene previously published as having an association with protein composition and protein and milk yields. The association with protein composition was confirmed in this experiment, but not the association with protein and milk yields. A multiple SNP regression analysis for all SNP on chromosome 20 was performed for all 6 traits, which revealed that this mutation was not significantly associated with any of the milk production traits and that at least 2 other quantitative trait loci were present on chromosome 20.

Genome‐wide associations for investigating time‐dependent genetic effects for milk production traits in dairy cattle

Phenotypic variation in milk production traits has been described over the course of a lactation as well as between different parities. The objective of this study was to investigate whether variation in production is affected by different loci across lactations. A genome-wide association study (GWAS) using a 50-k SNP chip was conducted in 152 divergent German Holstein Friesian cows to test for association with milk production traits over different lactations. The first four lactations were analysed regarding milk yield, fat, protein, lactose, milk urea nitrogen yield and content as well as somatic cell score. Two approaches were used: (i) Wilmink curve parameters were used to assess the genetic effects over the course of a lactation and (ii) test-day yield deviations (YD) were used as a normative approach for a GWAS. The significant effects were largest for markers affecting curve parameters for which there was a statistical power <0.8 of detection even in this small design. While significant markers for YDs were detected in this study, the power to detect effects of a similar magnitude was only 0.11, suggesting that many loci may have been missed with this approach in the present design. Furthermore, all significant effects were specific for a single lactation, leading to the conclusion that the variance explained by a certain locus changes from lactation to lactation. We confirm the common evidence that most production traits vary in the degree of persistency after the peak as a result of genetic influence.

Association of bovineCD4andSTAT5bsingle nucleotide polymorphisms with somatic cell scores and milk production traits in Chinese Holsteins

CD4+ T cells play a key role in the immune response of pathogen-induced mastitis in dairy cattle. Mammary gland factor STAT5b is involved in the regulation of CD4+T cell differentiation during inflammatory response and milk production. Little is known about the genetic variation effects of bovine CD4 and STAT5b genes on somatic cell score (SCS) and milk production traits in dairy cattle. The aim of the study was to investigate the single nucleotide polymorphisms (SNPs) of bovine CD4 and STAT5b in Chinese Holsteins and to analyse their association with estimated breeding values (EBVs) for SCS and milk production traits. In the present study, SNPs of CD4 (NC_007303 g.13598C>T) and STAT5b (NC_007317 g.31562 T>C) were identified and genotyped in Chinese Holstein population. The results showed that both SNPs were significantly associated with the EBVs for milk yield and protein yield in Chinese Holstein cows, and the SNP in CD4 was associated with the EBV for SCS (P<0.01). The additive effect of CD4 SNP on protein yield was significant (P<0.05), and the dominant effect of STAT5b SNP was significant on milk yield and protein yield (P<0.01). Cows with combination genotype C7 (CCTT: CD4 g.13598C>T and STAT5b g.31562 T>C) had the highest SCS EBV but lower milk yield, while cows with C2 (TTTC) produced more milk, fat and protein than the other eight combination genotypes. These results suggested that the SNPs in CD4 and STAT5b may be potential genetic markers for SCS and milk/protein yields selecting and warrant further functional research.

Identification of SNPs in Interferon Gamma, Interleukin-22, and Their Receptors and Associations with Health and Production-Related Traits in Canadian Holstein Bulls

Genetic variants in a number of immunoregulatory genes have been previously associated with health and production traits in dairy cattle. Therefore, in the following study, the genes coding interferon gamma (IFNG), IFNG receptor 1 and 2 domains, interleukin-22 (IL22), and IL22 receptor alpha 1, were investigated for single nucleotide polymorphisms (SNPs) in Holstein bulls. These SNPs, along with SNPs previously identified in IL10, IL10 receptor, and transforming growth factor beta 1 (TGFB1) genes, were evaluated for statistical associations to estimated breeding values for milk somatic cell score (SCS), a trait highly correlated to mastitis incidence, and various production-related traits, including milk yield, protein yield, fat yield, and lactation persistency. While no significant associations were found between these SNPs and SCS, SNPs in IL10 receptor beta subunit showed a significant effect on protein yield and lactation persistency. While there is evidence that IL10 plays an important role during lactation, it is also likely that the effects of SNPs in IL10 receptor beta subunit on protein yield and lactation persistency are due to linkage disequilibrium with a neighboring QTL.

Zeolite as a factor in the improvement of some production traits of dairy cattle

Milk yield and milk chemical composition are largely affected by diet quality. Natural zeolite has been successfully used in livestock production as a feed supplement for some types and categories of domestic animals. This experiment was conducted over a period of 15 months involving Domestic Spotted dairy cattle. The test animals were assigned to three groups, each receiving different levels of supplemental zeolite. The control group comprised cows that received no zeolite supplement. Experimental groups I and II were fed compound feeds supplemented with 4% and 2% zeolite, respectively. The production traits studied included the following milk performance traits: whole lactation length, milk and fat yields over whole lactations, milk and fat yields over standard lactations. The results obtained suggest that zeolite supplementation affects some production traits, depending on the zeolite level present in livestock feed.

Genome Partitioning of Genetic Variation for Milk Production and Composition Traits in Holstein Cattle

The objective of this study was to estimate the contribution of each autosome to genetic variation of milk yield, fat, and protein percentage and somatic cell score in Holstein cattle. Data on 2294 Holstein bulls genotyped for 39,557 autosomal markers were used. Three approaches were applied to estimate the proportion of genetic variance attributed to each chromosome. In two of them, marker-derived kinship coefficients were computed, using either marker genotypes observed on the whole genome or on subsets relative to each chromosome. Variance components were then estimated using residual maximum likelihood in method 1 or a regression-based approach in method 2. In method 3, genetic variances associated to each marker were estimated in a linear multiple regression approach, and then were summed up chromosome-wise. Generally, all chromosomes contributed to genetic variation. For most of the chromosomes, the amount of variance attributed to a chromosome was found to be proportional to its physical length. Nevertheless, for traits influenced by genes with very large effects a larger proportion of the genetic variance is expected to be associated with the chromosomes where these genes are. This is illustrated with the DGAT1 gene on BTA14 which is known to have a large effect on fat percentage in milk. The proportion of genetic variance for fat percentage associated with chromosome 14 was two to sevenfold (depending on the method) larger than would be predicted from chromosome size alone. Based on method 3 an approach is suggested to estimate the effective number of genes underlying the inheritance of the studied traits, yielding numbers between N ≈ 400 (for fat percentage) to N ≈ 900 (for milk yield). It is argued that these numbers are conservative lower bound estimates, but are in line with recent findings suggesting a highly polygenic background of production traits in dairy cattle.

Exploration of relationships between production and fertility traits in dairy cattle via association studies of SNPs within candidate genes derived by expression profiling

The objective of this work was to integrate findings from functional genomics studies with genome-wide association studies for fertility and production traits in dairy cattle. Association analyses of production and fertility traits with SNPs located within or close to 170 candidate genes derived from two gene expression studies and from the literature were performed. Data from 2294 Holstein bulls genotyped for 39557 SNPs were used. A total of 111 SNPs were located on chromosomal segments covered by a candidate gene. Allele substitution effects for each SNP were estimated using a mixed model with a fixed effect of marker and a random polygenic effect. Assumed covariance was derived either from marker or from pedigree information. Results from the analysis with the kinship matrix built from marker genotypes were more conservative than from the analysis with the pedigree-derived relationship matrix. From sixteen SNPs with significant effects on both classes of traits, ten provided evidence of an antagonistic relationship between productivity and fertility. However, we found four SNPs with favourable effects on fertility and on yield traits, one SNP with favourable effects on fertility and percentage traits, and one SNP with antagonistic effects on two fertility traits. While most quantitative genetic studies have proven genetic antagonisms between yield and functional traits, improvements in both production and functionality may be possible when focusing on a few relevant SNPs. Investigations combining input from quantitative genetics and functional genomics with association analysis may be applied for the identification of such SNPs.

A Monte Carlo approach for estimation of haplotype probabilities

A number of studies have tested for effects associated with different alleles of the milk protein genes on production traits in dairy cattle (e.g. Cowan et al., 1992). Because numerous protein loci exist and several are closely linked in the genome, one may also be interested in estimating the effects of combinations of alleles at more than one loci. One approach for this type of analysis is multiple regression on all loci and their interactions. An alternative strategy is an analysis of haplotype effects. Advantages of this strategy are that 1) the number of haplotypes in a population is typically much lower than the number of allelic combinations among loci, thus resulting in more precision and statistical power; and 2) haplotype effects can account for effects of unknown loci flanked by the genotyped loci. Ikonen et al. (2001) applied such an approach to test for effects of β and κ casein on production traits of Finnish Ayrshire cattle. Effects were observed for protein content in particular. One obstacle to haplotype analysis is that genotyping procedures generally do not provide information about the recombination phase between loci, necessary for obtaining haplotypes.

Environmental factors affecting milk, fat and protein yields in Jersey cattle in Zimbabwe

A study was carried out to establish the factors affecting milk, butterfat and protein yields in Zimbabwean Jersey cattle. A total of 10,986 unedited 305-day lactation records were obtained from Zimbabwe Livestock Identification Trust containing 25 herds, with cows calving in the period 1996-2008. The general linear model procedure of the Statistical Analysis Systems version 9.1.3 was used to determine the environmental factors. Herd-year-season, calving interval, days dry and both the linear and quadratic effects of age at calving fitted as covariates significantly (P<0.0001) affected the milk, fat and protein yields. Herd-year-season accounted for 36%, 36% and 35% to the total variation in milk, fat and protein yields, respectively. Milk, fat and protein yields increased with an increase in calving interval. The optimum dry period for milk, fat and protein yields obtained was 60-75 days dry. The relationship between the production traits and age at calving was both linear and quadratic, implying that milk, fat and protein yields increase with the age of the animal. It is thus necessary to preadjust data for these environmental factors when carrying out genetic evaluations of production traits in dairy cattle.

Genome Wide Association Studies for Milk Production Traits in Chinese Holstein Population

Genome-wide association studies (GWAS) based on high throughput SNP genotyping technologies open a broad avenue for exploring genes associated with milk production traits in dairy cattle. Motivated by pinpointing novel quantitative trait nucleotide (QTN) across Bos Taurus genome, the present study is to perform GWAS to identify genes affecting milk production traits using current state-of-the-art SNP genotyping technology, i.e., the Illumina BovineSNP50 BeadChip. In the analyses, the five most commonly evaluated milk production traits are involved, including milk yield (MY), milk fat yield (FY), milk protein yield (PY), milk fat percentage (FP) and milk protein percentage (PP). Estimated breeding values (EBVs) of 2,093 daughters from 14 paternal half-sib families are considered as phenotypes within the framework of a daughter design. Association tests between each trait and the 54K SNPs are achieved via two different analysis approaches, a paternal transmission disequilibrium test (TDT)-based approach (L1-TDT) and a mixed model based regression analysis (MMRA). In total, 105 SNPs were detected to be significantly associated genome-wise with one or multiple milk production traits. Of the 105 SNPs, 38 were commonly detected by both methods, while four and 63 were solely detected by L1-TDT and MMRA, respectively. The majority (86 out of 105) of the significant SNPs is located within the reported QTL regions and some are within or close to the reported candidate genes. In particular, two SNPs, ARS-BFGL-NGS-4939 and BFGL-NGS-118998, are located close to the DGAT1 gene (160bp apart) and within the GHR gene, respectively. Our findings herein not only provide confirmatory evidences for previously findings, but also explore a suite of novel SNPs associated with milk production traits, and thus form a solid basis for eventually unraveling the causal mutations for milk production traits in dairy cattle.

Identification of single nucleotide polymorphisms in the bovine interleukin-12 and interleukin-23 receptor genes and their associations with health and production traits in Holstein cows

Interleukin-12 (IL-12) and interleukin-23 (IL-23) are proinflammatory cytokines produced by macrophages and dendritic cells in response to infection with intracellular pathogens. The IL-12 receptor (IL-12R) is a heterodimer composed of 2 subunits, β1 and β2. The IL-23 receptor (IL-23R) is a heterodimer composed of the IL-12Rβ1 subunit and a unique IL-23R subunit. Given the importance of IL-12 and IL-23 for modulating inflammation and the host immune response, the IL-12 and IL-23 receptor genes may be suitable candidate genes for studying disease resistance in dairy cattle. We hypothesize that single nucleotide polymorphisms (SNP) exist within these genes and that they contribute to variation in health and production traits in dairy cattle. To investigate this, a selective DNA pool was constructed using bull semen based on the estimated breeding values for somatic cell score (SCS), an indicator trait used to achieve genetic improvement for resistance to mastitis. Gene segments were amplified from this pool by PCR and the amplicons were sequenced to reveal SNP. A total of 10 SNP, including 2 in IL-12Rβ1, 5 in IL-12Rβ2, and 3 in IL-23R were identified. The SNP (n=5) were found in the 5′ untranslated region (UTR) putative promoter regions of the genes, and SNP IL-23R c.1714A>C was a nonsynonymous SNP. Canadian Holstein bulls (n=492) were genotyped using Sequenom MassARRAY (Sequenom Inc., San Diego, CA). No association was found with SCS based on bull deregressed estimated breeding values for SCS; however, associations of SNP in the IL-12Rβ2 gene (c.-511A>G, c.87A>G, c.2957A>C) were found with milk and protein yield. Further investigation will be required to elucidate the biological and practical relevance of these SNP.

Testing candidate gene effects on milk production traits in dairy cattle under various parameterizations and modes of inheritance

The major objectives of this study were 1) to assess the statistical properties of models commonly used for the estimation of single nucleotide polymorphism (SNP) effects under the assumption of various modes of inheritance and various parameterizations of SNP genotypes using simulated data, and 2) to compare effects of the selected polymorphisms located within butyrophilin (BTN1A1), diacylglycerol acyltransferase 1 (DGAT1), leptin (LEP), and leptin receptor (LEPR) candidate genes on milk production traits using data from 2 dairy cattle breeds (190 Jersey cows and 475 Polish Holstein-Friesian cows). Simulation results showed that type I error and power were not dependent on the assumed parameterization, but differences were observed regarding confidence intervals of estimated SNP effects. In the presence of epistasis, correct confidence intervals for all (epistatic and nonepistatic) SNP and all modes of inheritance were provided only by the parameterization proposed by C. H. Kao and Z. B. Zeng in 2002. However, if no dominance effect was included in the model, confidence intervals for SNP effects were correct for all parameterizations. Results based on real data showed that for both breeds the additive effects of polymorphisms were generally similar, except for LEPR, which had a different allele associated with increased fat content in Holstein-Friesians than in Jerseys. In both breeds, DGAT1 had the largest additive effect of the polymorphisms considered, but its effect on most milk traits was more pronounced in Jerseys than in Holstein-Friesians. Evidence of epistasis was found between LEPR and DGAT1, as well as between LEPR and BTN1A1, but only for milk content traits and only in the Holstein-Friesian breed. There was also more evidence for dominance in the Holstein-Friesian breed than in the Jersey breed.

Polymorphism of the CYP19 gene and milk production traits of dairy cattle

The aim of this study was to estimate the relations between CYP19/Cfr13I and CYP19/PvuII genotypes and milk production traits. The study was carried on 915 Black-and-White cows kept in the Western Pomerania region of Poland. The CYP19/Cfr13I and CYP19/PvuII polymorphisms were detected using the PCR-RFLP method. Statistically significant (P Keywords:

Associations among milk production traits and glycosylated haemoglobin in dairy cattle; importance of lactose synthesis potential

Glucose is the major precursor of lactose synthesis in the mammary gland. Lactose the major carbohydrate and osmolyte of milk, controls milk volume and its concentration. Glycosylated haemoglobin (HbG) is a retrospective measure of mean blood glucose level and it is largely unaffected by recent physiological conditions and environmental events. The purposes of this study were to determine the correlations between lactose traits and other milk production traits in dairy cattle and to investigate whether HbG level can be correlated with milk and lactose production traits. Here, HbG percentage, milk and lactose production traits including milk yield, lactose, protein, SNF, total solid and fat percentages and yields were measured in 485 second calved Iranian Holstein cattle. Statistically significant negative correlations were established between HbG and milk yield (r = -0.88), lactose yield (r = -0.83), SNF yield (r = -0.81), protein yield (r = -0.79) and total solid yield (r = -0.74). Positive correlations were established between lactose yield and milk (r = 0.96), protein (r = 0.81), SNF (r = 0.92) and total solid (r = 0.79) yields. The negative correlation between HbG and milk and total lactose production is probably related to the higher glucose demands in the lactating mammary gland of more productive cows. The positive correlation between lactose yield and milk, protein, SNF and total solid yield indicates that the level of lactose synthesis influences milk production traits in ways other than merely via its osmolytic action.

Association Between the Growth Hormone Releasing Hormone (GHRH) Gene Polymorphism and Milk Production Traits of Dairy Cattle

Szatkowska, I., Dybus, A., Grzesiak, W., Jedrzejczak, M. and Muszyńska, M. 2009. Association between the growth hormone releasing hormone (GHRH) gene polymorphism and milk production traits of dairy cattle. J. Appl. Anim. Res., 36: 119–123. The association between the GHRH/HaeIII gene polymorphism, and milk production traits of Polish Holstein and Jersey cows were analysed. A total of 427 animals were included in the study. A PCR-RFLP method was used for genotyping. The frequencies of genotypes and alleles for the Polish Holstein cows were 0.078—AA, 0.339—AB, 0.583—BB and 0.248—GHRHA , 0.752—GHRHB. For the Jersey cows, the frequencies were: 0.032—AA, 0.427—AB, 0.541—BB and 0.246—GHRHA , 0.754—GHRHB. In all lactations, the Jersey cows with AA genotype exhibited the highest milk fat content. In the 2nd and 3rd lactations the AA Jersey cows had lower milk yields compared with the AB/BB cows. However, Polish Holstein cows did not exhibit such effects of genotypes on milk yield/traits. It still remains unclear that the analysed polymorphism could influence to the physiology of milk fat production.

A proline‐to‐histidine mutation in POU1F1 is associated with production traits in dairy cattle

POU class 1 homeobox 1 (POU1F1) is a member of the tissue-specific POU-containing transcription factor family. The expression of POU1F1 in mammalian pituitary gland controls the transcription of the genes encoding growth hormone, prolactin (PRL) and the subunits of thyroid-stimulating hormone. In addition, some genes in the JAK/STAT signalling pathway downstream of POU1F1 have been shown to be associated with different production traits in dairy cattle. To investigate whether the POU1F1 gene is associated with economically important traits in dairy cattle, a pooled DNA sequencing approach was used to identify single nucleotide polymorphisms (SNPs) in the gene. An SNP in exon 3 of POU1F1 that changes a proline to a histidine was identified. A total of 2141 individuals from two North American Holstein cattle resource populations were genotyped for this SNP using a modified PCR-RFLP method. Statistical analyses revealed significant association of POU1F1 variants with milk yield and productive life, which makes POU1F1 a possible candidate for marker-assisted selection in dairy cattle breeding programmes.

Association Between a Polymorphism at the Stearoyl CoA Desaturase Locus and Milk Production Traits in Italian Holsteins

Associations between stearoyl-CoA desaturase (SCD) gene polymorphisms and milk production traits (milk, fat, and protein yields, fat and protein contents, somatic cell score) were investigated on a sample of 701 lactations of 313 Italian Holsteins. Test-day records (5,097) were analyzed with a mixed linear model that included the fixed effects of herd, date of test, parity, genotype at the SCD locus, and lactation interval nested within SCD genotype, and the random effect of cow. An effect of the SCD genotype on milk and protein yields was detected, with VV cows producing more milk (about 2kg/d) and protein (about 0.07kg/d) compared with AA cows. The contribution of the SCD locus to the phenotypic variance of the 2 traits was about 0.015. These results suggest a possible use of the SCD locus in gene-assisted selection programs for the improvement of milk production traits in dairy cattle, although large-scale studies in different breeds are required.

Quantitative trait locus-by-environment interaction for milk yield traits on Bos taurus autosome 6.

Genotype-by-environment interactions for production traits in dairy cattle have often been observed, while QTL analyses have focused on detecting genes with general effects on production traits. In this study, a QTL search for genes with environmental interaction for the traits milk yield, protein yield, and fat yield were performed on Bos taurus autosome 6 (BTA6), also including information about the previously investigated candidate genes ABCG2 and OPN. The animals in the study were Norwegian Red. Eighteen grandsires and 716 sires were genotyped for 362 markers on BTA6. Every marker bracket was regarded as a putative QTL position. The effects of the candidate genes and the putative QTL were modeled as a regression on an environmental parameter (herd year), which is based on the predicted herd-year effect for the trait. Two QTL were found to have environmentally dependent effects on milk yield. These QTL were located 3.6 cM upstream and 9.1 cM downstream from ABCG2. No environmentally dependent QTL was found to significantly affect protein or fat yield.