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Article

Effect of FASN, SCD, and GH Genes on Carcass Fatness and Fatty Acid Composition of Intramuscular Lipids in F1 Holstein × Beef Breeds

by
Mateja Pećina
,
Miljenko Konjačić
,
Nikolina Kelava Ugarković
and
Ante Ivanković
*
Department of Animal Science and Technology, Faculty of Agriculture, University of Zagreb, Svetošimunska 25, 10000 Zagreb, Croatia
*
Author to whom correspondence should be addressed.
Agriculture 2023, 13(3), 571; https://doi.org/10.3390/agriculture13030571
Submission received: 28 January 2023 / Revised: 11 February 2023 / Accepted: 24 February 2023 / Published: 26 February 2023
(This article belongs to the Special Issue Advances in Molecular Genetics in Domestic Animals)
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Abstract

:
To improve beef quality, a selection of specific breeds for crossbreeding, genotyping, and selection of specific candidate genes in breeding animals can be some of the solutions. The objective of this study was to determine the effects of FASN, SCD, and GH genes on carcass fatness and fatty acid (FA) composition of intramuscular lipids of crossbred Holstein × beef breeds (Simmental, Belgian Blue, Limousin, and Piemontese). The allelic and genotypic distribution of polymorphisms in the FASN, SCD, and GH genes was studied in 80 crossbreed animals. Genomic DNA was isolated from musculus longissimus dorsi, whose chemical composition was determined by near infrared transmittance spectrophotometry, while the fatty acid composition was determined by gas chromatography. DNA polymorphism was analyzed by restriction fragment length polymorphism analysis. The FASN (g. 17924A>G) polymorphism was significantly associated with C19:1 n-9 and C24:1 n-9, whereas GH (g.2141C>G) was significantly associated with C16:0 and C20:1 n-9. The SCD (g.8586C>T) polymorphism was significantly associated with C16:0, C18:0, C20:0, C14:1 n-5, C16:1, C18:1, C18:2 n-6, C18:3 n-3, C20:2 n-6, and C20:4 n-6, and analyzed the sum and ratios of fatty acids. Sex had significant effect on carcass fatness and fatty acid composition. This study provided useful results for the above candidate genes and their association with some FA, supporting their influence as genes associated with fats and fatty acid composition in beef meat.

1. Introduction

Beef is one of the basic animal foods in the diet of the world’s population. Global beef production is 68.3 million tons, representing 20.3% of total meat production (336.6 million tons) [1]. The average beef consumption in the world is 6.3 kg/capita, while in some countries, such as Argentina, the annual beef consumption reaches 36.9 kg/capita [2]. Beef meat is a rich source of protein, fat, and a variety of essential macro and micronutrients. Fats in beef meat have an impact on nutritional, sensory, and processing properties, but also have certain health effects on the consumer. Beef muscle contains about 1–4% total fatty acids (lipids), which are mainly in triacylglycerol form, and those in muscles are triacylglycerol and phospholipids [3]. The level and composition of fatty acids (FA) of intramuscular fat (IMF) determines meat palatability and consumer’s satisfaction [4]. A high intake of SFA (Saturated Fatty Acids) is associated with an increase in serum cholesterol and low-density lipoprotein levels, which are risk factors for cardiovascular disease [5,6]. Polyunsaturated fatty acids (PUFA) and monounsaturated fatty acids (MUFA), such as linoleic or oleic acid, protect the cardiovascular system, and are associated with a decrease in serum cholesterol levels and an increase in high-density lipoprotein [7]. Conjugated linoleic acid also has favorable effects on human health [8].
The influence of genotype on carcass conformation and beef quality, including fat and fatty acid content in Holstein crossbreeds with beef breeds, has been confirmed in most of the previous studies [9,10,11,12]. Keane and Drennan [10] found that Holstein × Belgian Blue crosses had more favorable class and lower carcass fat compared to purebred Holstein cattle. Keane [11] found that F1 crosses of Holsteins with beef breeds were superior to purebred Holsteins in beef production. The fatty acid composition of the meat was affected by the diet [13]. Consumption of linseed has a significant effect on the concentrations of PUFAs [13] and contributes to the increase in the proportion of n-3 fatty acids in intramuscular fat [14]. The use of vitamin E at high doses (2.1 g/head/day) with extruded linseed had a positive effect on microbial loads and growth dynamics, and improved the fatty acid profile (with beneficial effects on consumer health) in finishing diets of young bulls [15].
The development of molecular genetics methods and direct detection of certain SNP polymorphisms on codogenic sequences has stimulated interest in gene candidates that have a certain effect on meat, especially fatty acids [16]. Fatty Acid Synthase (FASN) is a multifunctional enzyme that catalyzes the conversion of acetyl-CoA and malonyl-CoA to palmitate [17], and the FASN gene has been mapped to BTA19 [18]. Of particular interest was the SNP g.17924A > G polymorphism, which causes the replacement of the threonine to alanine, which affects the fat content in carcass and the proportion of FA, and SFA/MUFA/PUFA in beef meat [19,20,21,22,23,24,25]. Abe et al. [20] concluded that FASN gene polymorphism can be used for breeding control and optimization of some fatty acid concentrations in beef meat.
Stearoyl-CoA desaturase is an enzyme encoded by the SCD gene located on chromosome 26 and performs an important role in determining the fatty acid profile of ruminant tissues [26,27,28,29], as it is responsible for the conversion of SFA into MUFA [30,31]. Taniguchi et al. [26] identified a SNP in the fifth exon of this gene c.878 T>C that causes an amino acid change from alanine to valine, and the CC genotype to be associated with a higher proportion of PUFAs containing 9c-14:1, 9c-16:1, and 9c-18:1, and a lower melting point in the intermuscular fat. Wang et al. [29] reported that SNPs within the SCD gene had significant influence on C18:1 cis-13 fatty acid.
Several studies indicate associations of polymorphisms in GH loci with meat production and fatty acids profile. Growth hormone (GH), GH receptor (GHR), transcription factor PIT-1 (which activates GH and prolactin gene expression in the anterior pituitary), insulin-like growth factor-I (IGF-1), and possibly unexplored genes encoding GH signal transduction may contribute to advances in genetic selection in cattle [32]. Polymorphic substitution of ACG/ATG has been observed at codon 172 [33,34]. Ardiyanti et al. [35] examined GH codon polymorphisms at positions 127 and 172 and found that one group of cattle had higher levels of C18:1, MUFA, USFA (unsaturated fatty acids), MUFA/USFA, and USFA/SFA, while the other group had lower levels of C16:0 and C18:0. Maharani et al. [36] found that nucleotide substitution at codons 127 and 172 was not associated with fatty acid properties in their study.
Since farmers, and especially consumers, are interested in producing meat with favorable nutritional and sensory characteristics, some solutions are needed to improve meat quality. Some solutions could be the selection of specific breeds for crossbreeding, genotyping, and the selection of specific candidate genes in breeding animals, and the use of sexed semen to control the sex of calves. The objective of this study was to evaluate the association of genotype (FASN, SCD, and GH genes) with fat content in beef carcasses and the proportion of fatty acids in intramuscular fat.

2. Materials and Methods

2.1. Description of the Biological Sample

For research purposes, the crossing of cows of the Holstein breed (HL) with bulls of four beef breeds (Piemontese, Belgian Blue, Limousin, and Simmental; PIE, BB, LIM, and SIM) was carried out. In the crossing program, four bulls were selected for each beef breed, whose semen was used for artificial insemination. Insemination was performed over 4 months. After calving, at 2 weeks of age, 10 male and 10 female calves were collected for each cross combination (HL × PIE, HL × BB, HL × LIM, HL × SIM). A total of 80 calves were collected, 40 males and 40 females. The calves were collected and housed in a facility and kept until 6 months of age. Calves were separated by sex in two separate facilities on a cattle farm where their rearing continued.
The fattening of 40 bulls and 40 heifers was carried out on one fattening farm. Animals were kept in large, covered pens, each holding about 10 animals. The young cattle were fattened by applying a common fattening technology. They were fed a total mixed ration (TMR) once per day. An average TMR meal consisted of 6.5 kg of corn silage (30% dry matter), 5.5 kg of high moisture corn (~70% dry matter), 650 g of straw (~35% dry matter), and 1.4 kg of concentrate (34% crude protein). The average nutritional value of meal was ~76 MJ/kg ME and 950 g/kg of crude protein.
Young bulls were slaughtered at an average age of 489.8 ± 12.9 days, and heifers at 468.0 ± 8.2 days. Slaughter was carried out according to the standard procedure. Carcasses of slaughtered animals are classified by the EUROP classification system, which evaluates carcass muscularity (EUROP) and fat grades (1 to 5) of the carcass. The thickness of subcutaneous fatty tissue was measured using a precise caliper with a measurement scale in mm. The processed carcasses were stored at a temperature of 4 °C. After cooling, from the left half of each animal (carcass), a sample of the rib section was collected, which includes the 9th, 10th, and 11th thoracic vertebra, and its dissection was used to estimate the proportion of tissue in the carcass (muscle: fat and connective tissue: bone). On a meat sample of MLD taken at the 9th rib (weight 150 g), a chemical analysis of the meat was performed using NIT spectrophotometry (Near Infrared Transmittance spectroscopy) in the measuring range 850–1050 nm using a Foodscan instrument (Foss Electric A/S, Hillerød, Denmark).

2.2. Extraction of Total Lipids in Muscle and Analysis of Fatty Acid

The meat samples of MLD taken at the 9th rib (weight 50 g) were stored at a temperature of −20 °C until analysis. After thawing, tissues were homogenized for 60 s (3 × 20 s, with 10 s cooling intervals) at 9000 rpm. Samples were homogenized with an Ultra-Turrax T25 Basic homogenizer (IKA, Staufen, Germany).
Extraction of total lipids was conducted with a modified method developed by Folch et al. [37]. Extraction of total lipids was performed with a solvent mixture of chloroform that is methanol of different polarities. The ratio of extraction solvent was 15 cm3/g of tissue, divided into a three-part composition: chloroform:methanol at 2:1, chloroform:methanol at 1:1, and chloroform:methanol at 1:2. Total lipid homogenates in each solvent were extracted for 30 min with stirring (700 rpm), then centrifuged for 10 min at 3000 rpm at 20 °C. Total lipid extracts were combined and concentrated in a UNIVAPO 100H rotary evaporator, equipped with a UNICRYO MC 2L cooling unit (Uniequip, Planegg, Germany), and stored at 20 °C until analyzed.
Fatty acids from the total lipid extract were converted to methyl esters via transesterification with methanolic HCl according to international standard procedure ISO 5509 (2000). The resulting methyl esters of fatty acids were prepared for analysis with gas chromatography.
The analysis of fatty acid methyl esters was performed using a gas chromatograph (Agilent 8860, Agilent Technologies, Inc., Santa Clara, CA, USA) equipped with a flame ionization detector (FID). Temperatures of the injector and detector were 200 °C and 240 °C, respectively. Chromatography was performed on a DB-23 capillary column (Agilent Technologies, Santa Clara, CA, USA; length 30 m, column inner diameter 0.25 mm, active layer thickness 0.25 μm). Initial column temperature was 120 °C for 3 min, then it was increased to 260 °C by heating for 6 °C/min and was held at that temperature for 5 min. Hydrogen was used as carrier gas at a flow rate of 1 mL/minute. Collection and processing of the results were carried out using the computer program OpenLAB CDS ChemStation Workstation VL. Fatty acids were identified by comparison of retention times with methyl standards (Sigma Aldrich Chemie, GmbH and Supelco, St. Louis, MA, USA). The individual FA were used to calculate the sums of SFA, MUFA, and PUFA. Results are presented expressed in g/100 g of FA determined.

2.3. Analysis of the FASN, SCD i GH Polymophisms

Meat samples were collected after slaughter and stored at −20 °C. Isolation of DNA from tissue was performed using Sigma-AldrichTM GenElute Mammalian Genomic DNA Miniprep Kit (www.sigmaaldrich.com, accessed on 15 January 2022). Isolated DNA was stored at −20 °C until the genotyping procedure was performed. Each animal was genotyped for the FASN g.17924A>G, SCD g.8586C>T, and GH g.2141C>G through PCR-RFLP technique. Initial oligonucleotide sequences F-FASN 5′-TCTTCACAGAGCTGACGGAC-3′ and R-FASN 3′-GGAGGAAGAGCRGRRGCAGT-5′ were used to amplify the codogenic regions of the FASN gene [36]. For SCD and GH genes, we used two pairs of initial oligonucleotide sequences: F-SCD 5′-CCTGGTGTCCTGTTGTTGTG-3′ and R-SCD 3′-TAGACGTGGTCTTGCTGTGG-5′, F-GH 5′-TCTATGAGAAGCTGAAGGACCTGGAGGAA-3′, and R-GH 3′-CCAGAATAGAATGACACCTACTCAGACAAT-5′ [36].
Amplification of the codogenic sequence was performed in a 13.8 μL reaction mixture (5.4 μL MiliQ water, 7.5 μL EmeraldAmp GT PCR Master Mix, 0.45 μL of each primer, and 1.2 μL DNA). The polymerase chain reaction was performed on a Thermal Cycler MJ Research PTC 100 according to a temperature program, to which, after initial denaturation (98 °C/3 min), 35 cycles of DNA fragment amplification followed (98 °C/10 s, 58–61 °C/30 s, and 72 °C/50 s), then a final extension (72 °C/5 min). A quality control of the PCR reaction was performed on a 1% agarose gel after staining with StainINGreen Nucleic Acid. After the amplification, the PCR product was digested by restriction endonuclease MscI (FASN), Fnu4HI (SCD), and AluI (GH) for 4 h/37 °C in the prepared reaction mixture (0.40 μL dH2O, 10 μL PCR product, 0.10 μL DdeI enzyme). Restriction enzyme splitting results were read directly on a 3% agarose gel after electrophoresis (85 V/40 min; Figure 1). FASN, SCD and GH genotypes are determined based on the DNA fragment size: (a) GG (362, 262 bp), GA (362, 262, 195, 167 bp), and AA (262, 195, 167 bp), (b) CC (143 bp, 75 bp), CT (143, 113, 75 bp) and TT (143, 113 bp), (c) CC (264, 185, 145 bp), CG (264, 236, 185, 145 bp), and GG (264, 236, 145 bp) (Figure 1).

2.4. Statistical Analysis

The effects of FASN, SCD, and GH genes on carcass fatness and FA composition traits were tested using GLM (General Linear Model) procedure in the SAS V.18. The effects of genotypes on carcass fatness and fatty acid composition were determined using the following model:
Yijkl = μ + αi + gj + βk + εijkl
where Yijkl is the value of the analyzed parameter, μ is the overall mean, αi is the effect of crossbreeds (HL × SIM, HL × LIM, HL × BB, HL × PIE), gj is the effect of sex (bulls/heifers), βk is the effect of gene variant (FASN, SCD, GH gene), and εijkl is a random error. In case of effect of sex, age was included as co-variable in the model. The differences between means were estimated by Tukey’s test. All tables contain the least square mean (LS Mean) and standard error (SE) of the means.

3. Results

The polymorphism of the codogenic region of FASN, SCD, and GH genes was observed in the population of crossing animals (HL × beef breeds; SIM, LIM, BB, and PIE, respectively). The frequencies of the observed genotypes are shown in Figure 2, and the frequencies of the polymorphic allele variants G/A, C/T and C/G are shown in Table 1.
This study found the domination of G variant FASN g.17924A>G SNP in frequency 0.60–0.75. In three populations, we observed domination of T variant SCD g.8586C>T SNP, and only in HL × BB population was C variant SCD gene in frequency 0.60. In case of GH g.2141C>G SNP was observed domination of the C variant of gene, in frequency 0.70–0.93.
Although the focus of this study was to determine the influence of FASN, SCD, and GH genes on carcass fatness and fatty acid composition of intramuscular lipids in F1 Holstein × beef breeds, Table 2 shows values for animal weight, fatness of the carcass and meat, and longissimus dorsi muscle fatty acid composition on crossbreeds, and by sex. In the studied sample, the slaughter weights of animals were the highest (p < 0.001) in the HL × SIM crosses (572.1 kg), and the lowest slaughter weights were in the HL × LIM crosses (500.2 kg; Table 2). The cold carcasses weight was higher (p = 0.001) in the HL × SIM crosses (324.2 kg). Considering the EUROP conformation (muscularity) of the carcasses, the lowest rated carcasses was HL × SIM (3.33; p = 0.045), while their fat content was higher (3.09 vs. 2.54–2.79). The lowest proportion of fat and connective tissue in the rib section was observed in HL × PIE and HL × BB crosses, and differences are significant (p < 0.001). Significant impact of crossbreeds in the content of SFA, MUFA, PUFA, PUFA/SFA, and PUFA/MUFA were observed between the crossbred groups (Table 2).
The average slaughter weight and the weight of cold carcasses were significantly higher in bulls than in heifers (Table 2). The heifer carcasses, compared to the carcasses of bulls, were fatter (EUROP fat grade; p < 0.001) and had a higher proportion of fat and connective tissue in the rib section (21.23 vs. 11.61%; p < 0.001), and the proportion of fat in the MLD (3.17 vs. 1.74%; p = 0.002). Compared to bulls’ meat, heifer meat had a significantly higher content of MUFA (47.52 vs. 39.56; p < 0.001) and a lower content of PUFA (8.95 vs. 16.51; p < 0.001), especially n-6 PUFA (8.20 vs. 15.52; p < 0.001). Differences are noticeable also at the level of MUFA/SFA, PUFA/SFA, and PUFA/MUFA indices (p < 0.001).
No significant effect on carcass fatness was observed among the FASN genotypes. (Table 3). The AA genotype of the FASN gene had the lowest EUROP fat grade and subcutaneous adipose tissue thickness (2.58; 2.21 cm; Table 3). Different combinations of genotypes had minor significant effects on carcass fatness and fatty acid composition of intramuscular lipids. For example, GG genotype vs. GA/AA (p = 0.026) and GA vs. GG/AA (p = 0.036) had a significant effect on EUROP conformation.
The content of fatty acids in beef, depending on the observed FASN genotypes, are shown in Table 3. Significant effect of FASN genotypes was observed on content of C19:1 n-9 and C24:1 n-9 fatty acid, which both had a higher content in AA genotype then in GG and GA genotype (p = 0.013; p = 0.029). Significant effects were also observed in AA vs.GG/GA and GA vs.GG/AA combinations of genotypes for the same fatty acids (Table 3). No significant effect of FASN genotypes was observed on the sums and the ratios of fatty acids (SFA, PUFA, and MUFA).
Table 4 shows the effects of SCD genotypes and its different combinations on analyzed traits. Regarding SCD genotype, the fat + connective tissue in rib section and fat in MLD were significantly higher in the TT genotype (p = 0.001; p = 0.021). EUROP conformation classification was significantly affected only by the TT genotype vs. the CT/CC combination (p = 0.040). The CC vs.TT/TC and CT vs.TT/CC combinations had significant effect on fat + connective tissue in rib section (p = 0.001; and p = 0.001, respectively), and fat in MLD (p = 0.006; and p = 0.042, respectively).
The content of FA in beef of analyzed crossbreds depending on the SCD genotypes are shown in Table 4. The significant effect of SCD genotypes on the content of specific FA was observed. The highest value of C16:0 and C18:0 was found in the CC genotype (p = 0.004; and p = 0.005, respectively), while the CC and CT genotypes had the same value for C20:0 (p = 0.002). The TT genotype significantly affected content of C14:1 n-5, C16:1, and C18:1 (p = 0.034; p = 0.001; and p = 0.035, respectively). PUFAs (C18:2 n-6, C18:3 n-3, C20:2 n-6, and C20:4 n-6, respectively) were significantly higher in the CT genotype (p = 0.020; p = 0.007; p = 0.005; and p = 0.086, respectively). Sums and rations of fatty acids were significantly different between SCD genotypes (Table 4). The significant effect of TT vs. CT/CC and CT vs.TT/CC genotype combinations was found on majority of individual SFAs, MUFAs, and PUFAs, while CT vs.TT/CC combination also showed significant effect on majority of sums and rations of fatty acids (Table 4).
EUROP fat grade was significantly higher in the CG genotype (CG > CC > GG) of GH gene (p = 0.004). It was also found that the EUROP fat grade was significantly lower in CG vs. CC/GG genotypes (2.76 ± 0.07 vs. 3.05 ± 0.11; p= 0.003).
The content of FA in beef of analyzed crossbreds, depending on the GH gene, are shown in Table 5. The content of most FAs was not significantly affected by the GH gene. However, content of C16:0 was higher in CG genotype (p = 0.019), while content of C20:1 n-9 was higher in the GG genotype (p = 0.010). CG genotype vs. CC/GG showed significant effect on C16:0 (p = 0.043), while C20:1 n-9 was affected by CC vs. CG/GG and CC vs. GG (p = 0.007; p = 0.038). The sum of SFAs significantly differed for GG vs. CG/CC (p = 0.033).

4. Discussion

The dominance of the g.17924G variant of the FASN gene (0.66) in the population included in the study has been observed in numerous previous studies. In the population of Hanwoo cattle, the frequency of the g.17924G variant of the FASN gene is 0.81 [38], which was confirmed in the study by Maharani et al. [36]. Oh et al. [39] also found a dominance of the g.17924G variant of the FASN gene in a population of Korean cattle (0.73). Papaleo Mazzucco et al. [40] observed dominance of the g.17924G variant of the FASN gene in the Hereford population (0.72), and in a population of Angus cattle, where the frequency of the g.17924A variant of the FASN gene is 0.69. Cancino-Baier et al. [41] observed dominance of the g.17924G variant of the FASN gene in a population of Holstein Friesian steers (0.62). The dominance of the g.8586T variant of the SCD gene in the research population (0.72) was also observed in other cattle populations. Papaleo Mazzucco et al. [40] observed the dominance of the g.8586T variant of the SCD gene in a population of Angus and Hereford (0.68; 0.71). Maharani et al. [36] demonstrated certain dominance of the g.8586T variant of the SCD gene in the population of Hanwoo cattle (0.60). In the studied population, the g.2141C variant of the GH gene dominated in crosses of the Holstein × beef cattle breed (0.82). Maharani et al. [36] found a greater prevalence of the g.2141C variant of the GH gene in the population of Hanwoo cattle (0.93), and Kaneda et al. [42] also found the dominance of the g.2141C variant of the GH gene in nine cattle breeds from the Bos taurus and Bos indicus group (0.983–0.800).
In the population included in this study, minor effect of the g.17924A>G FASN gene on carcass fatness and fatty acid content of beef was detected. However, some previous studies indicate the association of the g.17924A>G SNP variant of the FASN gene with carcass fatness and fatty acid content of beef. Maharani et al. [36] indicated that the g.17924A>G SNP in the FASN gene was significantly associated with lower C14:0 content (p < 0.01) in animals with the GG genotype, and Zhang et al. [18] observed the same association in Angus bulls. The same FASN SNP was associated with higher C16:0 (p < 0.03) and lower C18:1 (p < 0.04) in animals with the AA genotype [43]. Narukami et al. [44] did not observe the same association FASN g.17924A>G SNP on FA composition of beef in Holstein bulls. Uemoto et al. [45] indicated that the g.17924A>G SNP of the FASN gene was associated with oleic acid (C18:1; p < 0.05) in Japanese black cattle. Oh et al. [39] indicated that the g.17924A>G SNP of the FASN gene was significantly associated with C14:0, C16:0, C18:0, C14:1, C18:1, C18:2 n6, C18:3 n3, SFA, and MUFA (p < 0.05). In the same study conducted in Korean cattle, the AA genotype g.17924A>G SNP of the FASN gene was associated with increased levels of C14:0, C16:0, and C18:0 fatty acids, and a decrease in C18:1 fatty acid in longissimus muscle [39]. Abe et al. [20] and Matsuhashi et al. [21] indicated significant effects of the g.17924A>G FASN genotype on 14:0, 14:1, 16:0, 16:1, and 18:1 content and intra-muscular fat in Japanese black cattle. Zhang et al. [18] observed that Angus bulls with the GG genotype (compared to the AA-genotype) had lower SFA, 14:0 and 16:0 fatty acids, and higher total MUFA and 18:1 fatty acid. Yeon et al. [38] observed that the g.17924A>G locus of the FASN gene was significantly associated with content C16:0, C16:1, C18:1, SFA, and unsaturated FA in Hanwoo cattle. Zhang et al. [18] and Schennink et al. [46] reported that the g.17924A>G SNP had a significant effect on the fatty acid composition of rib steaks in purebred American Angus bulls. Li et al. [22] concluded that animals with the AA genotype had a higher concentration of SFA, higher 14:0 fatty acid content, and lower concentration of oleic acid. Oh et al. [39] reported that SNP g.17924A>G of FASN gene has a significant effect on marbling of Korean cattle.
The polymorphic variants g.8586C>T SNP in the SCD gene showed major effect on carcass fatness and fatty acids content of beef in the studied population. Matsuhashi et al. [21] observed that in Japanese black cattle, the polymorphism of SCD had effects on myristic acid (p < 0.001), myristoleic acid (p < 0.001), stearic acid (p < 0.001), oleic acid (p < 0.001), and MUFA (p < 0.001). Maharani et al. [36] indicated that the g.8586C>T SNP in the SCD gene had a significant effect on C14:1 in Hanwoo cattle (p < 0.01). In agreement with a previous study in Japanese Black cattle, the SCD gene was significantly associated with C14:1 and C18:1 concentration in perirenal and intramuscular fat [47]. Papaleo Mazzucco et al. [40] did not find differences between SCD genotypes and fatty acids in a population of Angus and Hereford. Other authors indicate a relationship between SCD and fatty acid composition in Japanese black and white cattle [21,46], Fleckvieh bulls [25], and Brangus white oxen [48].
In the population included in this study, the influence of polymorphic variants was detected g.8586C>T SNP GH gene. The nucleotide substitutions at the codons of the GH gene were significantly associated with C14:0, C16:0, C18:1, C20:1, C20:5 n-3, SFA, and MUFA, and the ratios of MUFA/SFA and USFA/SFA in Japanese Black cattle [35]. Maharani et al. [36] did not observe that the g.8586C>T SNP in the GH gene had a significant effect on the fatty acid profile in Hanwoo cattle.
The effect of sex on carcass fatness and fatty acid composition of intramuscular lipids was observed, although it is not the focus of this study. Such a significant effect of sex on fatty acid composition was also observed in some previous studies. Barton et al. [49], in their study of Charolais × Simmental crossbred bulls and heifers, found that the total content of FA tended to be higher in heifers than in bulls, and that the content of SFA and MUFA increased linearly with increasing total FA. Karolyi et al. [50] found lower levels of MUFA and higher levels of PUFA in the muscle lipids of Simmental bulls compared to heifers.
The observed associations between polymorphic genes and carcass fatness, and fat and fatty acid content of beef are useful for selecting and conducting crossbreeding programs, and complement to previous scientific knowledge [9,10,11,12]. By selecting and favoring certain genotypes, it is possible to produce meat that is more acceptable to consumers. The observed associations of candidate genes with fatness traits and meat quality are also useful in selection of commercial and local breeds, especially in optimizing carcass fatness and fatty acid ratio, taking into account the influence of sex, nutrition, and other factors on beef quality.

5. Conclusions

In the present study, the effects of FASN and GH gene polymorphism on carcass fatness and fatty acid composition of intramuscular lipids were partially confirmed. The SCD gene polymorphism showed dominant effect on analyzed traits compared to other two genes. Fats and fatty acids determine the quality, health and sensory characteristics of beef and are of particular interest to consumers and indirectly to beef producers. It is possible to improve the target quality of beef by selecting specific genotypes in breeding or by crossbreeding. Future research needs to include more candidate genes and populations (purebred or crossbred) that are widely used in beef production.

Author Contributions

Conceptualization, A.I.; methodology, A.I. and M.K.; data collection, A.I. and M.K.; formal analysis, M.P. and A.I.; writing—original draft preparation, M.P. and A.I.; writing—review and editing, N.K.U. and M.K.; supervision, A.I.; project administration, A.I. All authors have read and agreed to the published version of the manuscript.

Funding

This research has been fully supported by Croatian Science Foundation (Genetic, Economic, and Social Interactions of Local Breed Conservation Programs, GGD LocBreed), grant number IP-2020-02-4860.

Institutional Review Board Statement

The research was approved by the Bioethical committee for the protection and welfare of animals at the Faculty of the Agriculture University of Zagreb (No: 251-71-29-02/19-22-2).

Informed Consent Statement

Not applicable.

Data Availability Statement

The data presented in this study are available on request from the corresponding author. The data are not publicly available to preserve privacy of the data.

Conflicts of Interest

The authors declare no conflict of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript, or in the decision to publish the results.

References

  1. FAO. World Food and Agriculture—Statistical Yearbook 2021. Rome. Available online: https://www.fao.org/home/en (accessed on 5 December 2022).
  2. OECD. Meat Consumption (Indicator). Available online: https://www.oecd.org/ (accessed on 5 December 2022).
  3. Wood, J.D.; Enser, M.; Richardson, R.I.; Whittington, F.M. Fatty Acids in Meat and Meat Products. In Fatty Acids in Foods and Their Health Implications, 3rd ed.; Chow, C.K., Ed.; Taylor & Francis Group, LLC: Boca Raton, NY, USA, 2007; pp. 87–107. [Google Scholar]
  4. Lee, S.H.; Park, E.W.; Cho, Y.M. Identification of differentially expressed genes related to intramuscular fat development in the early and late fattening stages of Hanwoo steers. J. Biochem. Mol. Biol. 2007, 40, 757–764. [Google Scholar] [CrossRef] [Green Version]
  5. Kris-Eiherton, P.M.; Fleming, J.A. Emerging nutrition science on fatty acids and cardiovascular disease: Nutritionists’ perspectives. Adv. Nutr. 2015, 6, 326–337. [Google Scholar] [CrossRef] [Green Version]
  6. Katan, M.B.; Zoock, P.M.; Mensink, R.P. Effects of fats and fatty acids on blood lipids in humans: An overview. Am. J. Clin. Nutr. 1994, 60, 1017–1022. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  7. Pensel, N. The future of red meat in human diets. Prof. Anim. Sci. 1998, 14, 133–140. [Google Scholar] [CrossRef]
  8. Ip, C.; Scimeca, J.A.; Thompson, H. Effect of timing and duration of dietary conjugated linoleic acid on mammary cancer prevention. Nutr. Cancer. 1995, 24, 241–247. [Google Scholar] [CrossRef]
  9. Alcicek, A.; Önenc, A.; Güngör, M. Carcass and meat quality of Friesian, Piemontese x Friesian and Limousin x Friesian young bulls under intensive beef production system in Turkey. J. Anim. Feed Sci. 2003, 12, 249–260. [Google Scholar] [CrossRef]
  10. Keane, M.G.; Drennan, M.J. A comparison of Friesian, Aberdeen Angus x and Belgian Blue x Friesian steers finished at pasture or indoors. Livest. Sci. 2008, 115, 268–278. [Google Scholar] [CrossRef]
  11. Keane, M.G. Beef Cross Breeding of Dairy and Beef Cows. Occasional Series 8, Grange Beef Research Centre, 2011. Available online: https://core.ac.uk/download/pdf/84886318.pdf (accessed on 10 February 2023).
  12. Nogalski, Z.; Wronski, M.; Wielgosz-Groth, Z.; Purwin, C.; Sobczuk-Szul, M.; Mochol, M.; Pogorzelska, P. The effect of carcass conformation class (EUROP system) on the slaughter quality of young, crossbred beef bulls and Holstein-Friesians. Ann. Anim. Sci. 2013, 13, 121–131. [Google Scholar] [CrossRef]
  13. Fusaro, I.; Cavallini, D.; Giammarco, M.; Manetta, A.C.; Martuscelli, M.; Mammi, L.M.E.; Lanzoni, L.; Formigoni, A.; Vignola, G. Oxidative status of Marchigiana beef enriched in n-3 fatty acids and vitamin E, treated with a blend of Oregano and Rosemary essential oils. Front. Vet. Sci. 2021, 8, 662079. [Google Scholar] [CrossRef]
  14. Scollan, N.D.; Dhanoa, M.S.; Choi, N.J.; Maeng, W.J.; Enser, M.; Wood, J.D. Biohydrogenation and digestion of long chain fatty acids in steers fed on different sources of lipid. J. Agric. Sci. 2001, 136, 345–355. [Google Scholar] [CrossRef]
  15. Fusaro, I.; Cavallini, D.; Giammarco, M.; Serio, A.; Mammi, L.M.E.; De Matos Vettori, J.; Lanzoni, L.; Formigoni, A.; Vignola, G. Effect of diet and essential oils on the fatty acid composition, oxidative stability and microbiological profile of Marchigiana burgers. Antioxidants 2022, 11, 827. [Google Scholar] [CrossRef]
  16. Pećina, M.; Ivanković, A. Candidate genes and fatty acids in beef meat, a review. Ital. J. Anim. Sci. 2021, 20, 1716–1729. [Google Scholar] [CrossRef]
  17. Roy, R.; Zaragoza, P.; Rodellar, C.; Gautier, M.; Eggen, A. Radiation hybrid and genetic linkage mapping of two genes related to fat metabolism in cattle: Fatty acid synthase (FASN) and glycerol-3-phosphate acyltranserase mitochondrial (GPAM). Anim. Biotechnol. 2005, 16, 1–9. [Google Scholar] [CrossRef] [PubMed]
  18. Zhang, S.; Knight, T.J.; Reecy, J.M.; Beitz, D.C. DNA polymorphisms in bovine fatty acid synthase are associated with beef fatty acid composition. Anim. Genet. 2008, 39, 62–70. [Google Scholar] [CrossRef]
  19. Morris, C.A.; Cullen, N.G.; Glass, B.C.; Hyndman, D.L.; Manley, T.R.; Hickey, S.M.; McEwan, J.C.; Pitchford, W.S.; Bottema, C.D.K.; Lee, M.A.H. Fatty acid synthase effects on bovine adipose fat and milk fat. Mamm. Genome 2007, 18, 64–74. [Google Scholar] [CrossRef]
  20. Abe, T.; Saburi, J.; Hasebe, H.; Nakagawa, T.; Misumi, S.; Nade, T.; Nakajima, H.; Shoji, N.; Kobayashi, M.; Kobayashi, E. Novel mutations of FASN gene and their effect on fatty acid composition in Japanese black beef. Biochem. Genet. 2009, 47, 397–411. [Google Scholar] [CrossRef]
  21. Matsuhashi, T.; Maruyama, S.; Uemoto, Y.; Kobayashi, N.; Mannen, H.; Abe, T.; Sakaguchi, S.; Kobayashi, E. Effects of FASN, SCD, SREBP1 and GH gene polymorphisms on fatty acid composition and carcass traits in Japanese Black cattle. J. Anim. Sci. 2011, 89, 12–22. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  22. Li, C.; Aldai, N.; Vinsky, M.; Dugan, M.E.R.; McAllister, T.A. Association analyses of single nucleotide polymorphisms in bovine stearoyl.CoA desaturase and fatty acid synthase genes with fatty acid composition in commercial cross breed beef steers. Anim. Genet. 2012, 43, 93–97. [Google Scholar] [CrossRef]
  23. Jeong, J.; Kwon, E.G.; Im, S.K.; Seo, K.S.; Baik, M. Expression of fat deposition and fat removal genes is associated with intermuscular fat content in longissimus dorsi muscle of Korean cattle steers. J. Anim. Sci. 2012, 90, 2044–2053. [Google Scholar] [CrossRef]
  24. Kaplanová, K.; Dufek, A.; Dračková, E.; Simeonovová, J.; Šubert, J.; Vrtková, I.; Dvořák, J. The association of CAPN1, CAST, SCD, and FASN polymorphism with beef quality traits in commercial crossbreed cattle in the Czech Republic. Czech J. Anim. Sci. 2013, 58, 489–496. [Google Scholar] [CrossRef] [Green Version]
  25. Bartoň, L.; Bureš, D.; Kott, T.; Řehák, D. Associations of polymorphisms in bovine DGAT1, FABP4, FASN, and PPARGC1A genes with intramuscular fat content and the fatty acid composition of muscle and subcutaneous fat in Fleckvieh bulls. Meat Sci. 2016, 114, 18–23. [Google Scholar] [CrossRef]
  26. Taniguchi, M.; Utsugi, T.; Oyama, K.; Mannen, H.; Kobayashi, M.; Tanabe, Y.; Ogino, A.; Tsuji, S. Genotype of stearoyl-CoA desaturase is associated with fatty acid composition in Japanese Black cattle. Mamm. Genome 2004, 15, 142–148. [Google Scholar] [CrossRef] [PubMed]
  27. Mannen, H. Identification, and utilization of genes associated with beef qualities. Anim. Sci. J. 2011, 82, 1–7. [Google Scholar] [CrossRef] [PubMed]
  28. Ntambi, J.M. Stearoyl-CoA Desaturase Genes in Lipid Metabolism, 3rd ed.; Springer: New York, NY, USA, 2013. [Google Scholar]
  29. Wang, Z.; Zhu, B.; Niu, H.; Zhang, W.; Xu, L.; Xu, L.; Chen, Y.; Zhang, L.; Gao, X.; Gao, H.; et al. Genome wide association study identifies SNPs associated with fatty acid composition in Chinese Wagyu cattle. J. Anim. Sci. Biotechnol. 2019, 10, 27. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  30. Kay, J.K.; Mackle, T.R.; Auldist, M.J.; Thomson, N.A.; Bauman, D.E. Endogenous synthesis of cis-9, trans-11 conjugated linoleic acid in dairy cows fed fresh pasture. J. Dairy Sci. 2004, 87, 369–378. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  31. Hoashi, S.; Hinenoya, T.; Tanaka, A.; Ohsaki, H.; Sasazaki, S.; Taniguchi, M.; Oyama, K.; Mukai, F.; Mannen, H. Association between fatty acid compositions and genotypes of FABP4 and LXR-alpha in Japanese Black cattle. BMC Genet. 2008, 9, 3–9. [Google Scholar] [CrossRef] [Green Version]
  32. Oprzadek, J.; Flisikowski, K.; Zwierzchowski, L.; Juszczuk-Kubiak, E.; Rosochacki, S.; Dymnicki, E. Associations between polymorphism of some candidate genes and growth rates, feed intake and uzilisation, slaughter indicators and meat quality in cattle. Arch. Tierz. 2005, 48, 81–87. [Google Scholar]
  33. Chikuni, K.; Nagatsuma, T.; Tabata, T.; Monma, M.; Saito, M.; Ozawa, S.; Ozutsumi, K. Genetic variants of the growth hormone gene in Japanese cattle. Anim. Sci. Technol. 1994, 65, 340–346. [Google Scholar] [CrossRef]
  34. Chikuni, K.R.; Tanabe, S.; Muroya, Y.; Fukumoto, S.; Ozawa, S. A simple method for genotyping the bovine growth hormone gene. Anim. Genet. 1997, 28, 230–232. [Google Scholar] [CrossRef]
  35. Ardiyanti, A.; Oki, Y.; Suda, Y.; Suzuki, K.; Chikuni, K.; Obara, Y.; Katoh, K. Effects of GH gene polymorphism and sex on carcass traits and fatty acid compositions in Japanese Black cattle. Anim. Sci. J. 2009, 80, 62–69. [Google Scholar] [CrossRef]
  36. Maharani, D.; Jung, Y.; Jung, W.Y.; Jo, C.; Ryoo, S.H.; Lee, S.H.; Yeon, S.H.; Lee, J.H. Association of five candidate genes with fatty acid composition in Korean cattle. Mol. Bio. Rep. 2012, 39, 6113–6121. [Google Scholar] [CrossRef]
  37. Folch, J.; Lees, M.; Sloane Stanley, G.H. A simple method for the isolation and purification of total lipids from animal tissues. J. Biol. Chem. 1957, 226, 497–509. [Google Scholar] [CrossRef]
  38. Yeon, S.H.; Lee, S.H.; Choi, B.H.; Lee, H.J.; Jang, G.W.; Lee, K.T.; Kim, K.H.; Lee, J.H.; Chung, H.Y. Genetic variation of FASN is associated with fatty acid composition of Hanwoo. Meat Sci. 2013, 94, 133–138. [Google Scholar] [CrossRef]
  39. Oh, D.; Lee, Y.; La, B.; Yeo, J.; Chung, E.; Kim, Y.; Lee, C. Fatty acid composition of beef is associated with exonic nucleotide variants of the gene encoding FASN. Mol. Biol. Rep. 2012, 39, 4083–4090. [Google Scholar] [CrossRef]
  40. Papaleo Mazzucco, J.; Goszczynski, D.E.; Ripoli, M.V.; Melucci, L.M.; Pardo, A.M.; Colatto, E.; Rogberg-Muñoz, A.; Mezzadra, C.A.; Depetris, G.J.; Giovambattista, G.; et al. Growth, carcass, and meat quality traits in beef from Angus, Hereford and crossbreed grazing steers, and their association with SNPs in genes related to fat deposition metabolism. Meat Sci. 2016, 114, 121–129. [Google Scholar] [CrossRef]
  41. Cancino-Baier, D.; Muñoz, E.; Quiñones, J.; Beltrán, J.; Fuentes, F.; Farías, J.; Lorenzo, J.; Diaz, R.; Inostroza, K.; Ferraz, J.; et al. Non-Synonymous Single Nucleotide Polymorphism in FASN Gene Alters FASN Enzyme Activity in Subcutaneous and Intramuscular Adipose Tissue in Holstein Friesian Steers. Ann. Anim. Sci. 2021, 21, 109–124. [Google Scholar] [CrossRef]
  42. Kaneda, M.; Lin, B.Z.; Sasazaki, S.; Oyama, K.; Mannen, H. Allele frequencies of gene polymorphisms related to economic traits in Bos taurus and Bos indicus cattle breeds. Anim. Sci. J. 2011, 82, 717–721. [Google Scholar] [CrossRef] [PubMed]
  43. Bhuiyan, M.S.A.; Yu, S.L.; Jeon, J.T.; Yoon, D.; Cho, Y.M.; Park, E.W.; Kim, E.W.; Kim, K.S.; Lee, J.H. DNA polymorphisms in SREBF1 and FASN genes affect fatty acid composition in Korean cattle (Hanwoo). Asian-Australas. J. Anim. Sci. 2009, 22, 765–773. [Google Scholar] [CrossRef]
  44. Narukami, T.; Sasazaki, S.; Oyama, K.; Nogi, T.; Taniguchi, M.; Mannen, H. Effect of DNA polymorphisms related to fatty acid composition in adipose tissue of Holstein cattle. Anim. Sci. J. 2011, 82, 406–411. [Google Scholar] [CrossRef]
  45. Uemoto, Y.; Abe, T.; Tameoka, N.; Hasebe, H.; Inoue, K.; Nakajima, H.; Shoji, N.; Kobayashi, M.; Kobayashi, E. Whole-genome association study for fatty acid composition of oleic acid in Japanese Black cattle. Anim. Genet. 2011, 42, 141–148. [Google Scholar] [CrossRef] [PubMed]
  46. Schennink, A.; Bovenhuis, H.; Leon-Kloosterziel, K.M.; Van Arendonk, J.A.; Visker, M.H. Effect of polymorphisms in the FASN, OLR1, PPARGC1A, PRL and STAT5A genes on bovine milk-fat composition. Anim. Genet. 2009, 40, 909–916. [Google Scholar] [CrossRef] [PubMed]
  47. Ohsaki, H.; Tanaka, A.; Hoashi, S.; Sasazaki, S.; Oyama, K.; Taniguchi, M.; Mukai, F.; Mannen, H. Effect of SCD and SREBP genotypes genotypes on fatty acid composition in adipose tissue of Japanese Black cattle herds. Anim. Sci. J. 2009, 80, 225–232. [Google Scholar] [CrossRef] [PubMed]
  48. Baeza, M.C.; Corva, P.M.; Soria, L.A.; Pavan, E.; Rincon, G.; Medrano, J.F. Genetic variants in a lipid regulatory pathway as potential tools for improving the nutritional quality of grass-fed beef. Anim. Genet. 2013, 44, 121–129. [Google Scholar] [CrossRef] [PubMed]
  49. Bartoň, L.; Bureš, D.; Kott, T.; Řehák, D. Effect of sex and age on bovine muscle and adipose fatty acid composition and stearoyl-CoA desaturase mRNA expression. Meat Sci. 2011, 89, 444–450. [Google Scholar] [CrossRef]
  50. Karolyi, D.; Dikić, M.; Salajpal, K.; Jurić, I. Fatty acid composition of muscle and adipose tissue of beef cattle. Ital. J. Anim. Sci. 2009, 8, 264–266. [Google Scholar] [CrossRef]
Agriculture 13 00571 g001a 550Agriculture 13 00571 g001b 550
Figure 1. Determination of allelic variants of FASN (a), SCD (b), and GH (c) gene on gel.
Figure 1. Determination of allelic variants of FASN (a), SCD (b), and GH (c) gene on gel.
Agriculture 13 00571 g001aAgriculture 13 00571 g001b
Agriculture 13 00571 g002 550
Figure 2. Number of observed FASN, SCD, and GH genotypes in sample.
Figure 2. Number of observed FASN, SCD, and GH genotypes in sample.
Agriculture 13 00571 g002
Table
Table 1. Frequencies of allele variants of FASN, SCD, and GH genes observed and expected heterozygosity (Ho/He).
Table 1. Frequencies of allele variants of FASN, SCD, and GH genes observed and expected heterozygosity (Ho/He).
CrossbreedsFASNSCDGH
G/AHo/HeC/THo/HeC/GHo/He
HL × SIM0.65/0.350.60/0.460.25/0.750.30/0.370.85/0.150.30/0.25
HL × LIM0.60/0.400.60/0.480.15/0.850.30/0.260.70/0.300.40/0.42
HL × BB0.75/0.250.40/0.380.60/0.400.30/0.480.93/0.070.15/0.14
HL × PIE0.63/0.380.45/0.470.15/0.850.30/0.260.80/0.200.20/0.32
Total0.66/0.340.51/0.450.28/0.720.30/0.410.82/0.180.26.0.30
Table
Table 2. Average value (LS Mean ± SE and p-value) of carcass fatness and fatty acids composition in the m. longissimus dorsi by crossbreeds and sex of animals.
Table 2. Average value (LS Mean ± SE and p-value) of carcass fatness and fatty acids composition in the m. longissimus dorsi by crossbreeds and sex of animals.
CharacteristicCrossbreedsp ValueSexp Value
HL × SIMHL × BBHL × PIEHL × LIMBullsHeifers
Slaughter weight, kg572.1 ± 9.16 a530.6 ± 8.81 b517.8 ± 8.77 b,c500.2 ± 8.94 c<0.001569.8 ± 7.75478.1 ± 11.15<0.001
Carcass weight, kg324.2 ± 4.55 a306.4 ± 4.59 a,c310.6 ± 4.57 b,c291.7 ± 4.79 d0.001343.2 ± 4.38273.3 ± 6.30<0.001
EUROP conform a3.33 ± 0.11 a3.74 ± 0.11 b3.72 ± 0.11 b3.55 ± 0.12 a,b0.0453.85 ± 0.113.44 ± 0.150.089
EUROP fat grade b3.09 ± 0.10 a2.76 ± 0.11 a,c2.54 ± 0.11 b2.79 ± 0.11 b,c0.0132.53 ± 0.083.25 ± 0.12<0.001
Subcut. fat tissue c2.57 ± 0.272.78 ± 0.282.89 ± 0.282.69 ± 0.290.9012.47 ± 0.193.34 ± 0.280.043
Fat + conn. tissue d18.25 ± 0.59 a16.23 ± 0.59 b11.81 ± 0.59 c18.37 ± 0.62 a<0.00111.61 ± 0.5121.23 ± 0.73<0.001
Fat in MLD e3.09 ± 0.22 a2.52 ± 0.22 a,b2.03 ± 0.23 b2.62 ± 0.23 a,b0.0891.74 ± 0.203.17 ± 0.290.002
C14:02.57 ± 0.12 a2.69 ± 0.12 a2.13 ± 0.12 b2.87 ± 0.13 a0.0022.27 ± 0.112.83 ± 0.130.011
C16:023.99 ± 0.37 a24.61 ± 0.37 a21.65 ± 0.37 b24.14 ± 0.39 a<0.00122.03 ± 0.3525.09 ± 0.39<0.001
C16:13.86 ± 0.14 a3.88 ± 0.14 a,c3.52 ± 0.14 a4.28 ± 0.15 b,c0.0233.43 ± 0.124.28 ± 0.13<0.001
C18:013.58 ± 0.4113.41 ± 0.4213.43 ± 0.4112.48 ± 0.430.36814.71 ± 0.3911.72 ± 0.43<0.001
C18:140.16 ± 0.68 a37.91 ± 0.68 a35.24 ± 0.68 b39.00 ± 0.71 a0.00334.74 ± 0.6541.31 ± 0.72<0.001
C18:2 n-66.27 ± 0.62 a6.85 ± 0.62 a11.39 ± 0.62 b6.80 ± 0.65 a<0.00110.85 ± 0.594.98 ± 0.65<0.001
C18:2 c9.t110.24 ± 0.010.23 ± 0.010.22 ± 0.010.23 ± 0.020.8350.23 ± 0.010.23 ± 0.020.967
C20:4 n-62.07 ± 0.25 a2.57 ± 0.25 a3.87 ± 0.25 b2.38 ± 0.26 a0.0013.37 ± 0.242.17 ± 0.260.008
C22:4 n-60.39 ± 0.04 a0.49 ± 0.04 a,c0.58 ± 0.04 b,c0.40 ± 0.04 a0.0170.51 ± 0.040.43 ± 0.040.210
C22:5 n-30.17 ± 0.02 a0.23 ± 0.02 a0.33 ± 0.02 b0.21 ± 0.02 a0.0020.29 ± 0.020.20 ± 0.020.024
SFA44.12 ± 0.60 a45.30 ± 0.63 a41.98 ± 0.59 b43.75 ± 0.63 a0.01743.88 ± 0.5543.48 ± 0.610.692
MUFA45.71 ± 0.76 a43.23 ± 0.81 a40.18 ± 0.76 b45.22 ± 0.79 a0.00339.56 ± 0.7547.52 ± 0.83<0.001
PUFA10.17 ± 0.98 a11.47 ± 1.04 a17.84 ± 0.97 b11.03 ± 1.02 a<0.00116.51 ± 0.938.95 ± 1.03<0.001
n-3 PUFA0.52 ± 0.05 a0.61 ± 0.05 a0.85 ± 0.05 b0.57 ± 0.05 a0.0010.75 ± 0.040.52 ± 0.050.006
n-6 PUFA9.39 ± 0.86 a10.62 ± 0.99 a16.75 ± 0.93 b10.22 ± 0.98 a<0.00115.52 ± 0.908.20 ± 0.99<0.001
n-6/n-3 PUFA17.79 ± 0.44 a17.43 ± 0.47 a,c19.27 ± 0.44 b,c17.69 ± 0.46 a0.09520.73 ± 0.4015.77 ± 0.44<0.001
MUFA/SFA1.05 ± 0.02 a,b0.96 ± 0.02 a0.96 ± 0.02 a,b1.04 ± 0.02 b0.0750.90 ± 0.021.10 ± 0.02<0.001
PUFA/SFA0.23 ± 0.02 a0.25 ± 0.02 a0.43 ± 0.02 b0.25 ± 0.03 a<0.0010.39 ± 0.020.21 ± 0.02<0.001
PUFA/MUFA0.23 ± 0.03 a0.28 ± 0.03 a0.49 ± 0.03 b0.25 ± 0.03 a<0.0010.44 ± 0.030.20 ± 0.03<0.001
a EUROP conformation classification: 1 = poor, 5 = excellent; b EUROP fat grade classification: 1 = leanest, 5 = fattest; c thickness of subcutaneous fat tissue, cm; d fat and connected tissue in the 9 to 11 ribs section, %; e fat in musculus longissimus dorsi, %; SFA, sum of saturated fatty acids; MUFA, sum of monounsaturated fatty acids; PUFA, sum of polyunsaturated fatty acids; n-3 PUFA, sum of n-3 PUFA fatty acids; n-6 PUFA, sum of n-6 PUFA fatty acids; n-6/n-3 PUFA, ratio of the sum of n-6/n-3 PUFA fatty acids; MUFA/SFA, ratio of the sum of MUFA and SFA fatty acids; PUFA/SFA, ratio of the sum of PUFA and SFA fatty acids; PUFA/MUFA, ratio of the sum of PUFA and MUFA fatty acids; different small letters in row, p < 0.05.
Table
Table 3. Carcass fatness and fatty acid content in MLD, their sums and ratio by FASN genotypes (LS Mean ± SE and p-value).
Table 3. Carcass fatness and fatty acid content in MLD, their sums and ratio by FASN genotypes (LS Mean ± SE and p-value).
Carcass Fatness and Fatty AcidsFASN Genotypes p-Value
AAGAGGp-ValueGG vs. GA/AAGG/GA vs. AAGG/AA vs. GAGG vs. AA
EUROP conform a3.58 ± 0.172 a,b3.72 ± 0.079 a3.45 ± 0.089 b0.0810.0260.8740.0360.547
EUROP fat grade b2.58 ± 0.1662.80 ± 0.0762.82 ± 0.0850.4340.7680.2200.7160.335
Subcut. fat tissue c2.01 ± 0.4313.02 ± 0.1972.63 ± 0.2210.1210.3910.1470.0740.225
Fat + conn. tissue d15.79 ± 0.9315.72 ± 0.4216.35 ± 0.480.6030.2350.7740.4880.705
Fat in MLD e2.51 ± 0.1382.64 ± 0.2172.42 ± 0.4460.8320.3970.5380.7260.563
C12:00.06 ± 0.0030.06 ± 0.0030.06 ± 0.0030.7370.8740.5990.4530.538
C14:02.35 ± 0.1892.62 ± 0.0862.54 ± 0.0970.4330.7680.3230.3850.381
C14:1 n-50.55 ± 0.0790.60 ± 0.0360.61 ± 0.0400.7490.5820.5460.8210.387
C15:00.33 ± 0.0280.37 ± 0.0130.37 ± 0.0140.5630.9930.3670.8980.392
C16:023.63 ± 0.30 a23.82 ± 0.26 b22.23 ± 0.57 b0.0540.7820.0210.2410.043
C16:14.03 ± 0.2173.84 ± 0.0993.85 ± 0.1110.7280.9900.4130.9640.513
C17:00.88 ± 0.0630.92 ± 0.0290.92 ± 0.0320.8140.9040.5490.8750.613
C17:1 n-70.83 ± 0.0600.78 ± 0.0270.80 ± 0.0310.6650.7260.4560.4540.677
C18:012.98 ± 0.6413.46 ± 0.3013.06 ± 0.330.6160.4390.6810.5130.870
C18:138.02 ± 1.0638.07 ± 0.4837.79 ± 0.550.9270.8970.8780.6590.853
C18:2 n-69.02 ± 0.9707.62 ± 0.4438.03 ± 0.4990.4170.9890.2690.3470.378
C18:2 c9.t110.21 ± 0.0230.23 ± 0.0110.24 ± 0.0120.6660.7220.2760.9580.445
C18:3 n-60.05 ± 0.0080.05 ± 0.0040.06 ± 0.0040.1730.0550.8360.0950.499
C18:3 n-30.21 ± 0.0170.19 ± 0.0080.20 ± 0.0090.5350.7040.4850.2980.662
C18:4 n-30.03 ± 0.0070.03 ± 0.0030.03 ± 0.0030.9010.9120.5870.9890.615
C19:00.05 ± 0.0040.06 ± 0.0020.06 ± 0.0020.2370.7110.0550.5780.072
C19:1 n-90.06 ± 0.006 a0.04 ± 0.002 b0.05 ± 0.003 a0.0130.1110.0160.0100.228
C20:00.07 ± 0.0050.07 ± 0.0020.07 ± 0.0020.7310.3770.8830.7710.534
C20:1 n-90.13 ± 0.0100.13 ± 0.0050.12 ± 0.0050.3880.2520.8900.3620.269
C20:2 n-60.10 ± 0.0100.10 ± 0.0050.10 ± 0.0050.8140.7350.9490.3540.862
C20:3 n-60.57 ± 0.0780.52 ± 0.0360.57 ± 0.0400.5230.3480.6840.2920.910
C20:4 n-63.09 ± 0.3912.64 ± 0.1782.81 ± 0.2010.5460.7880.3600.3950.545
C20:5 n-30.08 ± 0.0100.06 ± 0.0040.07 ± 0.0050.3450.7970.2250.3220.394
C22:2 n-60.02 ± 0.0080.02 ± 0.0030.03 ± 0.0030.3390.2650.2020.4650.126
C22:3 n-30.10 ± 0.0140.08 ± 0.0060.09 ± 0.0070.2270.2770.2710.1200.522
C22:4 n-60.48 ± 0.0630.43 ± 0.0290.51 ± 0.0320.2490.1220.7020.1210.777
C22:5 n-30.27 ± 0.0360.23 ± 0.0160.24 ± 0.0180.5860.8890.3910.5150.521
C22:6 n-30.02 ± 0.0060.03 ± 0.0030.03 ± 0.0030.3160.1510.4410.4880.247
C24:1 n-90.05 ± 0.007 a0.03 ± 0.003 b0.04 ± 0.004 a,b0.0290.5640.0120.0380.121
SFA42.07 ± 0.93 a44.22 ± 0.44 b43.66 ± 0.48 a,b0.1140.7630.0660.2340.059
MUFA43.72 ± 1.1943.45 ± 0.5643.34 ± 0.610.9590.9330.9940.7140.777
PUFA14.21 ± 1.5212.33 ± 0.7212.99 ± 0.780.5180.8140.3260.3150.490
n-3 PUFA0.70 ± 0.0720.62 ± 0.0340.66 ± 0.0370.2690.5570.3950.2550.609
n-6 PUFA13.29 ± 1.4611.47 ± 0.6912.09 ± 0.750.5130.8350.3180.3240.481
n-6/n-3 PUFA18.43 ± 0.6818.00 ± 0.3218.01 ± 0.350.8560.3220.7320.8750.678
MUFA/SFA1.04 ± 0.0340.99 ± 0.0151.00 ± 0.0170.3960.8890.3370.6570.189
PUFA/SFA0.34 ± 0.0390.28 ± 0.0180.30 ± 0.0200.3980.9390.2670.3700.366
PUFA/MUFA0.35 ± 0.0460.31 ± 0.0210.32 ± 0.0240.6630.9130.5040.4680.656
a EUROP conformation classification: 1 = poor, 5 = excellent; b EUROP fat grade classification: 1 = leanest, 5 = fattest; c thickness of subcutaneous fat tissue, cm; d fat and connected tissue in the 9 to 11 ribs section, %; e fat in musculus longissimus dorsi, %; SFA, sum of saturated fatty acids; MUFA, sum of monounsaturated fatty acids; PUFA, sum of polyunsaturated fatty acids; n-3 PUFA, sum of n-3 PUFA fatty acids; n-6 PUFA, sum of n-6 PUFA fatty acids; n-6/n-3 PUFA, ratio of the sum of n-6/n-3 PUFA fatty acids; MUFA/SFA, ratio of the sum of MUFA and SFA fatty acids; PUFA/SFA, ratio of the sum of PUFA and SFA fatty acids; PUFA/MUFA, ratio of the sum of PUFA and MUFA fatty acids; different small letters in row, p < 0.05.
Table
Table 4. Carcass fatness and fatty acid content in MLD, their sums and ratio by SCD genotypes (LS Mean ± SE and p-value).
Table 4. Carcass fatness and fatty acid content in MLD, their sums and ratio by SCD genotypes (LS Mean ± SE and p-value).
Carcass Fatness and Fatty AcidsSCD Genotypes p Value
CCCTTTp ValueTT vs. CT/CCTT/TC vs. CCTT/CC vs. CTTT vs. CC
EUROP conform a3.75 ± 0.152 a3.72 ± 0.100 a3.48 ± 0.075 b0.0950.0400.2650.1290.132
EUROP fat grade b2.79 ± 0.1452.69 ± 0.0962.84 ± 0.0720.4540.5250.8400.2310.802
Subcut. fat tissue c2.86 ± 0.3792.65 ± 0.2492.77 ± 0.1880.8850.6470.8740.8440.866
Fat + conn. tissue d15.32 ± 0.82 a13.56 ± 0.54 a17.69 ± 0.41 b0.0010.0010.3870.0010.028
Fat in MLD e2.20 ± 0.306 a,b2.18 ± 0.202 b2.85 ± 0.153 a0.0210.0060.2240.0420.077
C12:00.07 ± 0.0030.06 ± 0.0030.06 ± 0.0050.2530.1080.7440.1420.366
C14:02.58 ± 0.166 a,b2.36 ± 0.109 a2.55 ± 0.083 b0.1050.0590.7680.0380.667
C14:1 n-50.48 ± 0.069 a0.54 ± 0.045 a0.66 ± 0.034 b0.0340.0230.0580.0920.035
C15:00.37 ± 0.0250.37 ± 0.0160.36 ± 0.0120.7170.5090.4930.6380.553
C16:024.44 ± 0.51 a22.62 ± 0.33 b23.91 ± 0.25 a0.0040.0290.0470.0020.321
C16:13.59 ± 0.191 a3.48 ± 0.125 a4.18 ± 0.095 b0.0010.0010.0960.0010.006
C17:00.95 ± 0.0550.94 ± 0.0360.89 ± 0.0270.5000.3140.3630.4110.419
C17:1 n-70.72 ± 0.0530.76 ± 0.0350.83 ± 0.0260.0830.0350.1570.1700.089
C18:014.29 ± 0.57 a13.90 ± 0.37 a12.56 ± 0.28 b0.0050.0020.0500.0150.015
C18:137.78 ± 0.93 a,b36.70 ± 0.61 a38.79 ± 0.46 b0.0350.0210.8400.0160.210
C18:2 n-67.10 ± 0.852 a9.28 ± 0.561 b7.32 ± 0.423 a0.0200.0280.2700.0080.765
C18:2 c9.t110.24 ± 0.0210.22 ± 0.0140.24 ± 0.0100.5490.5750.5140.3620.718
C18:3 n-60.04 ± 0.0070.05 ± 0.0050.05 ± 0.0030.5030.6890.3190.3920.377
C18:3 n-30.17 ± 0.015 a0.22 ± 0.010 b0.18 ± 0.007 a0.0070.0350.1360.0020.377
C18:4 n-30.02 ± 0.0080.03 ± 0.0040.03 ± 0.0020.5740.6900.4280.6450.422
C19:00.06 ± 0.0040.06 ± 0.0020.06 ± 0.0020.9820.9470.9740.7590.851
C19:1 n-90.05 ± 0.007 a,b0.04 ± 0.003 a0.05 ± 0.002 b0.0670.0180.6570.0200.370
C20:00.08 ± 0.004 a0.08 ± 0.003 a0.07 ± 0.002 b0.0020.0010.0290.0020.002
C20:1 n-90.12 ± 0.0090.12 ± 0.0060.13 ± 0.0040.3010.5000.1560.4160.096
C20:2 n-60.10 ± 0.009 a,b0.11 ± 0.006 a0.09 ± 0.004 b0.0050.0020.9140.0020.355
C20:3 n-60.49 ± 0.0680.61 ± 0.0450.52 ± 0.0340.2230.3100.3450.1180.664
C20:4 n-62.41 ± 0.343 a,b3.17 ± 0.226 a2.59 ± 0.170 b0.0860.1580.2730.0430.628
C20:5 n-30.06 ± 0.009 a,b0.08 ± 0.006 a0.06 ± 0.004 b0.0610.0900.4770.0270.988
C22:2 n-60.02 ± 0.0060.03 ± 0.0040.02 ± 0.0030.5080.3200.9170.2530.848
C22:3 n-30.07 ± 0.0120.10 ± 0.0080.09 ± 0.0060.3020.9310.1430.2830.355
C22:4 n-60.44 ± 0.0550.51 ± 0.0360.45 ± 0.0270.4080.5410.4650.2510.778
C22:5 n-30.22 ± 0.0310.27 ± 0.0210.23 ± 0.0160.1580.2430.4040.0810.746
C22:6 n-30.02 ± 0.0060.03 ± 0.0040.03 ± 0.0030.1800.7380.1250.1220.415
C24:1 n-90.05 ± 0.0080.04 ± 0.0040.04 ± 0.0030.2760.4660.1600.9020.076
SFA45.79 ± 0.82 a43.30 ± 0.57 b43.46 ± 0.41 b0.0290.2770.0050.9290.019
MUFA42.82 ± 1.04 a,b41.02 ± 0.72 a45.04 ± 0.71 b0.0060.0030.4810.0040.049
PUFA11.39 ± 1.34 a15.68 ± 0.93 b11.50 ± 0.66 a0.0350.0580.2640.0150.697
n-3 PUFA0.59 ± 0.031 a0.78 ± 0.044 b0.55 ± 0.063 a0.0470.2020.1630.0260.414
n-6 PUFA10.66 ± 0.63 a14.68 ± 0.89 b10.58 ± 1.28 a0.0350.0550.2710.0150.713
n-6/n-3 PUFA18.87 ± 0.60 a,b18.66 ± 0.42 a17.58 ± 0.30 b0.0430.0080.1980.0720.039
MUFA/SFA0.94 ± 0.030 a0.96 ± 0.019 a1.04 ± 0.015 b0.0020.0020.0210.0130.004
PUFA/SFA0.25 ± 0.034 a0.34 ± 0.023 b0.28 ± 0.017 a0.0360.0900.1490.0190.463
PUFA/MUFA0.28 ± 0.041 a0.39 ± 0.027 b0.29 ± 0.020 a0.0110.0200.2400.0050.784
a EUROP conformation classification: 1 = poor, 5 = excellent; b EUROP fat grade classification: 1 = leanest, 5 = fattest; c thickness of subcutaneous fat tissue, cm; d fat and connected tissue in the 9 to 11 ribs section, %; e fat in musculus longissimus dorsi, %; SFA, sum of saturated fatty acids; MUFA, sum of monounsaturated fatty acids; PUFA, sum of polyunsaturated fatty acids; n-3 PUFA, sum of n-3 PUFA fatty acids; n-6 PUFA, sum of n-6 PUFA fatty acids; n-6/n-3 PUFA, ratio of the sum of n-6/n-3 PUFA fatty acids; MUFA/SFA, ratio of the sum of MUFA and SFA fatty acids; PUFA/SFA, ratio of the sum of PUFA and SFA fatty acids; PUFA/MUFA, ratio of the sum of PUFA and MUFA fatty acids; different small letters in row, p < 0.05.
Table
Table 5. Carcass fatness and fatty acid content in MLD, their sums and ratio by GH genotypes (LS Mean ± SE and p-value).
Table 5. Carcass fatness and fatty acid content in MLD, their sums and ratio by GH genotypes (LS Mean ± SE and p-value).
Carcass Fatness and Fatty AcidsGH Genotypes p-Value
CCCGGGp-ValueCC vs. CG/GGGG vs. CG/CCCG vs. CC/GGCC vs. GG
EUROP conform a3.53 ± 0.0683.68 ± 0.1073.75 ± 0.2200.3600.2190.5110.2210.358
EUROP fat grade b2.67 ± 0.065 a3.08 ± 0.103 b2.50 ± 0.212 a0.0040.0150.2190.0030.449
Subcut. fat tissue c2.58 ± 0.1703.18 ± 0.2682.30 ± 0.5510.1360.2060.4980.0760.628
Fat + conn. tissue d15.75 ± 0.3616.51 ± 0.5815.79 ± 1.190.5440.1240.9780.6160.959
Fat in MLD e2.50 ± 0.1382.64 ± 0.2172.42 ± 0.4460.8320.5100.8740.8090.862
C12:00.06 ± 0.0070.06 ± 0.0040.06 ± 0.0020.7740.9590.4620.8870.525
C14:02.51 ± 0.0752.71 ± 0.1182.35 ± 0.2420.2470.2190.3900.2630.536
C14:1 n-50.57 ± 0.0310.64 ± 0.0490.68 ± 0.1010.3410.0990.3970.4590.245
C15:00.36 ± 0.0110.38 ± 0.0180.35 ± 0.0360.5220.3970.6800.4030.810
C16:023.33 ± 0.27 a24.34 ± 0.36 b22.24 ± 0.74 b0.0190.0820.1030.0430.180
C16:13.83 ± 0.0853.91 ± 0.1354.02 ± 0.2770.7310.2690.6020.9720.508
C17:00.91 ± 0.0250.94 ± 0.0390.87 ± 0.0800.7220.8080.5610.7120.619
C17:1 n-70.79 ± 0.0240.79 ± 0.0370.85 ± 0.0770.7160.6050.4100.5910.382
C18:013.37 ± 0.2513.28 ± 0.4012.17 ± 0.820.3920.3500.1810.8680.202
C18:137.64 ± 0.4238.59 ± 0.6637.96 ± 1.360.4850.2700.9420.5660.838
C18:2 n-68.26 ± 0.3826.95 ± 0.6039.35 ± 1.2410.1140.1760.3100.2200.441
C18:2 c9.t110.23 ± 0.0090.25 ± 0.0150.21 ± 0.0300.4170.3730.6020.2170.732
C18:3 n-60.05 ± 0.0030.05 ± 0.0050.04 ± 0.0100.3530.2010.4250.3610.376
C18:3 n-30.20 ± 0.0070.18 ± 0.0100.21 ± 0.0210.2780.2360.4510.3720.602
C18:4 n-30.03 ± 0.0030.03 ± 0.0040.03 ± 0.0060.4540.2660.9400.2320.741
C19:00.06 ± 0.0020.06 ± 0.0020.06 ± 0.0050.2680.3490.5490.2180.664
C19:1 n-90.05 ± 0.0020.05 ± 0.0030.05 ± 0.0070.6140.8440.4600.3020.577
C20:00.07 ± 0.0020.07 ± 0.0030.06 ± 0.0060.3430.6050.1580.5380.222
C20:1 n-90.12 ± 0.004 a0.14 ± 0.006 b0.15 ± 0.013 b0.0100.0070.0550.0760.038
C20:2 n-60.10 ± 0.0040.10 ± 0.0060.11 ± 0.0130.7700.9950.5130.7810.488
C20:3 n-60.57 ± 0.0310.49 ± 0.0480.56 ± 0.1000.3210.2140.8940.3460.924
C20:4 n-62.88 ± 0.1542.38 ± 0.2433.29 ± 0.5000.1420.2520.3550.2390.472
C20:5 n-30.07 ± 0.004 a0.06 ± 0.006 b0.08 ± 0.012 a,b0.0630.1220.3630.1270.500
C22:2 n-60.02 ± 0.0030.03 ± 0.0040.03 ± 0.0080.4240.9950.5400.3550.197
C22:3 n-30.09 ± 0.0050.08 ± 0.0090.10 ± 0.0180.3280.4750.4780.3680.602
C22:4 n-60.48 ± 0.0250.42 ± 0.0390.54 ± 0.0800.3200.5200.4020.3640.528
C22:5 n-30.25 ± 0.0140.21 ± 0.0220.29 ± 0.0460.1350.2810.3430.2390.449
C22:6 n-30.03 ± 0.0030.02 ± 0.0050.03 ± 0.0080.4930.5040.6490.4660.755
C24:1 n-90.04 ± 0.0030.03 ± 0.0050.04 ± 0.0090.2970.3010.6580.1910.733
SFA43.64 ± 0.38 a,b44.55 ± 0.58 a41.26 ± 1.19 b0.0490.6920.0330.1610.060
MUFA43.02 ± 0.4844.17 ± 0.7443.85 ± 1.510.4130.1960.7780.6050.673
PUFA13.35 ± 0.6111.28 ± 0.9514.89 ± 1.940.1300.2060.3390.2360.465
n-3 PUFA0.67 ± 0.0290.57 ± 0.0450.75 ± 0.0920.1240.2730.3150.2360.417
n-6 PUFA12.45 ± 0.5910.44 ± 0.9113.91 ± 1.870.1290.2000.3410.2320.470
n-6/n-3 PUFA18.31 ± 0.2817.76 ± 0.4317.83 ± 0.880.5390.1970.6050.8030.578
MUFA/SFA0.99 ± 0.0131.00 ± 0.0211.06 ± 0.0430.2600.3480.0960.8220.105
PUFA/SFA0.31 ± 0.015 a,b0.25 ± 0.024 a0.37 ± 0.050 b0.0650.2010.2150.1770.302
PUFA/MUFA0.37 ± 0.0180.28 ± 0.0290.37 ± 0.0590.1400.1490.5320.3380.668
a EUROP conformation classification: 1 = poor, 5 = excellent; b EUROP fat grade classification: 1 = leanest, 5 = fattest; c thickness of subcutaneous fat tissue, cm; d fat and connected tissue in the 9 to 11 ribs section, %; e fat in musculus longissimus dorsi, %; SFA, sum of saturated fatty acids; MUFA, sum of monounsaturated fatty acids; PUFA, sum of polyunsaturated fatty acids; n-3 PUFA, sum of n-3 PUFA fatty acids; n-6 PUFA, sum of n-6 PUFA fatty acids; n-6/n-3 PUFA, ratio of the sum of n-6/n-3 PUFA fatty acids; MUFA/SFA, ratio of the sum of MUFA and SFA fatty acids; PUFA/SFA, ratio of the sum of PUFA and SFA fatty acids; PUFA/MUFA, ratio of the sum of PUFA and MUFA fatty acids; different small letters in row, p < 0.05.
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Pećina, M.; Konjačić, M.; Ugarković, N.K.; Ivanković, A. Effect of FASN, SCD, and GH Genes on Carcass Fatness and Fatty Acid Composition of Intramuscular Lipids in F1 Holstein × Beef Breeds. Agriculture 2023, 13, 571. https://doi.org/10.3390/agriculture13030571

AMA Style

Pećina M, Konjačić M, Ugarković NK, Ivanković A. Effect of FASN, SCD, and GH Genes on Carcass Fatness and Fatty Acid Composition of Intramuscular Lipids in F1 Holstein × Beef Breeds. Agriculture. 2023; 13(3):571. https://doi.org/10.3390/agriculture13030571

Chicago/Turabian Style

Pećina, Mateja, Miljenko Konjačić, Nikolina Kelava Ugarković, and Ante Ivanković. 2023. "Effect of FASN, SCD, and GH Genes on Carcass Fatness and Fatty Acid Composition of Intramuscular Lipids in F1 Holstein × Beef Breeds" Agriculture 13, no. 3: 571. https://doi.org/10.3390/agriculture13030571

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