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The Scientific World Journal
Volume 2014 (2014), Article ID 934154, 10 pages
http://dx.doi.org/10.1155/2014/934154
Research Article

Changes in Fatty Acid Composition and Distribution of N-3 Fatty Acids in Goat Tissues Fed Different Levels of Whole Linseed

1Faculty of Veterinary Medicine, Universiti Putra Malaysia (UPM), 43400 Serdang, Selangor, Malaysia
2Faculty of Animal Production, University of Khartoum, Shambat, 13314 Khartoum North, Sudan
3Institute of Biosciences, Universiti Putra Malaysia (UPM), 43400 Serdang, Selangor, Malaysia
4Institute of Tropical Agriculture, Universiti Putra Malaysia (UPM), 43400 Serdang, Selangor, Malaysia
5Department of Animal Science, Universiti Putra Malaysia (UPM), 43400 Serdang, Selangor, Malaysia

Received 21 April 2014; Revised 18 September 2014; Accepted 3 October 2014; Published 11 November 2014

Academic Editor: Patricia Foley

Copyright © 2014 Kamaleldin Abuelfatah et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

Abstract

The effects of feeding different levels of whole linseed on fatty acid (FA) composition of muscles and adipose tissues of goat were investigated. Twenty-four Crossed Boer bucks were assigned randomly into three treatment diets: L0, L10, or L20, containing 0%, 10%, or 20% whole linseed, respectively. The goats were slaughtered after 110 days of feeding. Samples from the longissimus dorsi, supraspinatus, semitendinosus, and subcutaneous fat (SF) and perirenal fat (PF) were taken for FA analyses. In muscles, the average increments in α-linolenic (ALA) and total n-3 PUFA were 6.48 and 3.4, and 11.48 and 4.78 for L10 and L20, respectively. In the adipose tissues, the increments in ALA and total n-3 PUFA were 3.07- and 6.92-fold and 3.00- and 7.54-fold in SF and PF for L10 and L20, respectively. The n-6 : n-3 ratio of the muscles was decreased from up to 8.86 in L0 to 2 or less in L10 and L20. The PUFA : SFA ratio was increased in all the tissues of L20 compared to L0. It is concluded that both inclusion levels (10% and 20%) of whole linseed in goat diets resulted in producing meat highly enriched with n-3 PUFA with desirable n-6 : n-3 ratio.

1. Introduction

The increased intake of n-3 polyunsaturated fatty acids (PUFA), specifically, eicosapentaenoic acid (EPA) and docosahexaenoic acid has been associated with significant physiological and health benefits in human populations. The intake of n-3 PUFA provided benefits for reducing of the incidence of cardiovascular diseases, atherosclerosis, hypertension [1], some cancer, inflammatory diseases [2], and some mental and emotional disorder [3], in addition to improving eye and brain development and learning ability [4]. The potential benefits of n-3 PUFA have stimulated the research in different fields in order to increase these beneficial fatty acids (FAs) in the human diet towards recommended levels. Red meat, particularly that from ruminant animals, has a bad reputation attributed to its high saturated nature, a low ratio of PUFA to saturated FA (SFA), and high n-6 : n-3 ratio [57], which may cause numerous cancers, atherosclerosis, and coronary heart diseases [8, 9].

In response to the concerns of the medical community and health-conscious consumers, research in meat production has focused on altering the FA content of meat through increasing the n-3 PUFA content, decreasing the PUFA n-6 : n-3 ratio, and enhancing the content of CLA in meat. Comparing to monogastrics, increasing the PUFA in the ruminant meat is more challenging, since most of the PUFA in the animal diet are hydrogenated by the rumen microorganisms [10]. However, the inclusion of sources of -linolenic acid (ALA) in the diets of ruminants has been shown to increase the concentration of n-3 PUFA in their meat. Feeding entire oilseeds is one of the strategies to reduce rumen biohydrogenation, as the seed coat provides protection for unsaturated FA (UFA) from rumen microorganisms [11], and might have lesser adverse effects on rumen fermentation than feeding free oils [12]. In addition, using intact seeds instead of oils has practical advantages in terms of handling the feed ingredients and manufacturing. Linseed (Linum usitatissimum) is a leading source of plant based n-3 FA containing about 40% oil, of which 50–60% ALA [13].

During the last decades, goat meat has gained a growing interest due to its preferable nutritional features, as it contains low levels of fat and cholesterol [14] and higher level of PUFA compared to beef or lamb [15]. The naturally high level of PUFA may indicate that goat has a potential to deposit high level of n-3 PUFA in it tissues. Enriching goat meat with n-3 PUFA together with its natural favorable nutrition characteristics enables goat meat to play an important role in human health as functional food, especially for health-conscious consumers. However, information about the effects of the feeding dietary regime on the FA profiles of edible tissues of goat, generally, is relatively scare [16]. Worse still, there is no report about the effects of feeding whole linseed, particularly, on the FA profiles of edible tissues of goat. Therefore, the objective of this study was to investigate the effects of moderate and high inclusion levels of whole linseed, as a source of n-3 PUFA in diets, on the FA composition of intramuscular and adipose tissues, with an emphasis on n-3 PUFA and CLA, of Crossed Boer goats.

2. Material and Methods

2.1. Experimental Animals and Housing

The trial was conducted at the Experimental Ruminant Unit, Department of Animal Science, Faculty of Agriculture, Universiti Putra Malaysia (UPM), under a tropical climate. Twenty-four, 5-month old Crossed Boer bucks with initial body weight (mean and SE) of  kg were unique numbering, dewormed, and housed in individual wooden pens and subjected to an adaptation period of three weeks prior to the beginning of the feeding trial to adjust to the housing conditions and diets. At the end of the adaptation period, the initial body weight of each goat was recorded. Accordingly, they were distributed into three groups of eight animals in each group, where the mean live weight of animals was not significantly different between the experimental groups. The animals were allocated randomly to one of three of experimental diets.

2.2. Feeds and Feeding

Three diets, L0, L10, and L20, containing 0%, 10%, or 20% of whole linseed, respectively, were formulated to meet the nutrient requirements of growing goats [17]. The high inclusion level of linseed was determined at 20% to give ether extract less than 10%, which considered the maximum level for ruminants’ diet [18]. The experimental diets and their ingredients and chemical composition are given in Table 1. The diets were randomly allocated to the three animal groups. Throughout the feeding trial, animals were fed at 3% body weight as dry matter intake daily. The trial period lasted for 110 days. Samples for proximate analyses were taken from the feed and refusals on a weekly interval.

tab1
Table 1: Formulation and proximate analyses of the experimental diets.
2.3. Slaughtering and Tissues Sampling

At the end of the experiment, animals were slaughtered according to Islamic Halal-method (slitting the throat to cut the jugular veins and arteries without stunning) at the slaughter house, Department of Animal Science, Faculty of Agriculture, UPM. Samples were taken from longissimus dorsi (LD), supraspinatus (SS), and semitendinosus (ST) muscles. Additionally, about 10 g of perirenal fat (PF) and subcutaneous fat (SF) were obtained. All the tissues were then vacuum packaged and stored at −80 until fatty acid analyses.

2.4. Chemical Analysis

Samples of feed were subjected to proximate analysis according to the standard methods of AOAC (2007) [19]. For determination of dry matter (DM) content, the samples were dried at 105°C for 24 h in a forced-air oven. Ash content was determined by combusting the samples in a muffle furnace at 550°C for 6 h, and organic matter (OM) content was calculated by difference (OM = 100-ash content). The N content of samples was determined using a Kjeltec Auto Analyzer (Tecator, Hoganas, Sweden), and crude protein (CF) was calculated as N × 6.25. Ether extract (EE) was determined in petroleum ether using a Soxtec Auto Analyzer (Tecator). Neutral detergent fiber (NDF) and acid detergent fiber were determined using the procedures of [20].

2.5. Measurement of Fatty Acids

The total FAs were extracted from feeds and animal tissues based on the method of [21] modified by [22], using chloroform-methanol 2 : 1 (v/v) containing butylated hydroxytoluene to prevent oxidation during sample preparation. FAs were transmethylated to their FA methyl esters (FAME) using 0.66 N KOH in 14% methanol and methanolic boron trifluoride (BF3) according to the methods by AOAC (2000). The FAME composition was quantified with a gas-liquid chromatography on an Agilent 7890A GS system (Agilent, Palo Alto, CA, USA) equipped with a 100 m × 0.25 mm ID (0.20 μm film thickness) Supelco sp-2560 capillary column (Supelco, Inc., Bellefonte, PA, USA). One μL of FAME was injected by an autosampler. H2 was used as the carrier gas and the split ratio was 10 : 1 after FAME injection. The injector and detector temperature were programmed at 250°C and 300°C, respectively. The column temperature program initiated at 120°C held for 5 min, increased by 2°C/min up to 170°C for 15 min, and then the temperature increased again by 5°C/min up to 200°C for 5 min and increased again by 2°C/min to a final temperature at 235°C and held for 10 min. A reference standard (mix C4–C24 methyl esters; Sigma-Aldrich, Inc., St. Louis, Mo, USA) and CLA standard mix (cis-9 trans-11 and trans-10 cis-12 CLA) (Sigma-Aldrich, Inc., St. Louis, Mo, USA) were used for determining individual FA.

2.6. Statistical Analysis

Results were analyzed using analysis of variance with the different inclusions of linseed as the main effects. For feed data sets one-way ANOVA was used to compare differences in overall and individual fatty acid types. The tissue FA datasets were first analyzed using the one-way ANOVA within tissue type for treatment effects. Following this, a two-way ANOVA was employed to analyze for the effect of tissue type (or anatomical location of muscles) X treatment effects. In both cases Duncan’s multiple range test was employed to elucidate significant means using the SAS software package, version 9.2. Differences between the least squared means were considered to be significant at . Data were presented as least-square means ± standard errors.

3. Result

3.1. Fatty Acid Composition of the Diets

The FA profiles of the diets are shown in Table 2. The most abundant FAs in the diets were oleic (33.5–27.1%), linolenic acid (LA) (21.9–18.0%), palmitic (28.07–7.89%), and ALA (39.3–1.9%). The proportions of oleic and palmitic were higher () in L0 compared to L10 and L20. There was no significant difference in the proportion of LA among the treatment diets. As projected, the inclusion of linseed contributed to a significant variation in the content of n-3 PUFA. The highest proportion of n-3 FA was in L20 (39.3% of total FA), followed by L10 (33.4% of total FA), and the lowest was in L0 (1.6% of total FA). The SFA was higher () in L0 (42.5% of total FA) compared to both L10 (17.4% of total FA) and L20 (15.5% of total FA). There were no significant differences among the experimental diets in n-6 PUFA.

tab2
Table 2: Fatty acid composition of the experimental diets.
3.2. Fatty Acid Composition of Tissues

The FA profiles for different goat muscles (LD, SS, and ST) and adipose tissues (SF and PF) are shown in Tables 37, respectively. The values are expressed as a percentage of the total fatty acids. The most abundant FAs in the goat muscles lipid were oleic acid (32.40–39.45%), palmitic acid (16.41–20.94%), and stearic acid (15.09–16.85%). The proportion of palmitic acid was greater () in all muscles of the control group (L0) compared to the other groups (L10 and L20). Nonsignificant difference was detected in the proportions of stearic acid and oleic acid in the studied muscles, except for ST of L20, which showed a lower level of oleic acid compared to the same muscle for both L0 and L20. Similar to the muscles, oleic, palmitic, and stearic acids were the major FA in the adipose tissues. In SF, the proportion of oleic acid ranged between 37.14 and 43.13%. The higher proportion was exhibited by L10 with a significant difference () when compared to L0. In the PF, the proportion of oleic acid ranged between 23.66 and 25.08%, with no significant differences across treatments. Both the SF and PF of the control group had higher proportions of palmitic acid (). The stearic acid in the SF ranged between 21.44 and 26.76%, and the control group scored the highest proportion compared to the other groups (). In the PF, the stearic acid ranged between 33.14 and 37.93% with no difference between groups.

tab3
Table 3: Fatty acid profiles of the longissimus dorsi muscle of goats fed diets containing different levels of whole linseed.
tab4
Table 4: Fatty acid profiles of the supraspinatus muscleof goats fed diets containing different levels of whole linseed.
tab5
Table 5: Fatty acid profiles ofthe semitendinosus muscle of goats fed diets containing different levels of whole linseed.
tab6
Table 6: Fatty acid profiles of the subcutaneous fat of goats fed diets containing different levels of whole linseed.
tab7
Table 7: Fatty acid profiles of perirenal fat of goats fed diets containing different levels of whole linseed.

The ALA content in different tissues is shown in Figure 1. The proportion of ALA increased in all the studied tissues () as the inclusion level of linseed increased. In muscles (Tables 3, 4, and 5), it ranged from 0.30% in SS of L0 to 4.85% in LD of L20. The highest proportion was exhibited by the LD muscle of goats fed the L20 diet. In the adipose tissues (Tables 6 and 7), the percentage of ALA was between 0.22% in the PF of L0 and 1.94% in the ST.

934154.fig.001
Figure 1: ALA (18 : 3 n-3) contents in muscles and fat tissues of goats fed diets containing different levels of whole linseed. Error bar = (1 SE). Bars with different alphabet notation differ significantly.

The increment of ALA in the different muscles of experimental groups was 5.7, 6.75, and 7.0 fold for L10 and was 12.52, 10.25, and 11.67 fold for L20 in LD, SS, and ST muscles, respectively, compared to L0. The increment of ALA in the adipose tissues was 3.07 and 6.92 fold and was 3.00 and 7.54 fold in the SF and PF for L10 and L20, respectively, compared to L0.

As illustrated in Figure 2, inclusion of the linseed in the goat diets also resulted in a significant increase () in the long-chain n-3 PUFA, EPA (C20:5 n-3), DPA (C20:5 n-3), and DHA (C20:6 n-3). However, these long-chain n-3 FAs were not detected in the adipose tissues (SF and PF) of goats, where the ALA represented the entire n-3 PUFA detected in this tissue.

934154.fig.002
Figure 2: Long-chain n-3 FA contents in muscles and fat tissues of goats fed diets containing different levels of whole linseed. Error bar = (1 SE).

There were no significant differences in the proportions of LA and total n-6 in SS, ST, and SF; nevertheless they were higher in LD of control group and PF of L20.

The proportion of C18:1 trans-11 (vaccenic acid) was higher () in the LD (1.99%), SS (1.51%), and SF (4.25%) of the L20 group compared to control group (1.44%, 1.65%, and 1.65%, for the same tissues, resp.).

As shown in Figure 3, the inclusion of linseed at 10% and 20% increased the total CLA in all the tissues compared to the control (), except for the LD muscle. The highest percentage (1.83) was found in the SS muscle of the L20 group, whereas the lowest percentage (0.54) was detected in the ST muscle of the control group (L0).

934154.fig.003
Figure 3: Total CLA contents in muscles and fat tissues of goats fed diets containing different levels of whole linseed. Error bar = (1 SE). Bars with different alphabet notation differ significantly.

The PUFA n-6 : n-3 ratios for the tissues of goats fed different levels of linseed are summarized in Figure 4. The PUFA n-6 : n-3 ratios for the tissues of goats fed different levels of linseed are summarized in Figure 4 The highest ratio (15.79) was noticed in the SF of the control group (L0), while the lowest (1.16) was shown in the LD muscle of L20. Both inclusion levels of linseed (10% or 20%) dramatically reduced the n-6 : n-3 ratios of goat tissues (). In the muscle, the ratios decreased from 8.86, 7.55, and 6.65 in LD, SS, and ST, respectively, for the L0 group, to 1.68, 1.87, and 2.0 for L10 and to 1.16, 1.8, and 1.32 for L20, respectively, for the same muscles. Similar to the muscles, the n-6 : n-3 ratios of the SF and PF decreased from 15.79 and 10.53 in L0 to 3.67 and 3.14 for L10 and to 1.60 and 2.72 for L20, respectively. However, the difference between the groups fed linseed (L10 and L20) was not significant for all the tissues studied.

934154.fig.004
Figure 4: n-6 : n-3 ratio in muscles and fat tissues of goats fed diets containing different levels of whole linseed. Error bar = (1 SE). Bars with different alphabet notation differ significantly.

4. Discussion

The current paper is focusing on the effects of feeding different levels of whole linseed on FA profile with an emphasis on n-3 PUFA and CLA in different tissues of goat. The data related to growth performance and carcass characteristics of the experimental animal was reported in a previous paper by the same authors [23]. In general, the FA composition of meat can be greatly modified by diet [24]. Other factors, such as species, age, weight, sex, and breed, also influence the composition of FA. Additionally, the fatty acid composition differs between tissue sites in the animal body [25].

Similar to that reported in related studies [26, 27], the most abundant FAs in the experimental diets were oleic acid, LA. However, the level of ALA was intentionally higher in the L10 and L20, as a result of the inclusion of linseed. In tissues, the major FAs were oleic, palmitic, stearic, and LA, as previously reported in goat [15]; However, in this study the goats fed linseed diets showed a markedly high level of n-3 fatty acids in their tissues.

In this study, the levels of ALA and total n-3 PUFA in the muscles and adipose tissues were significantly increased in all goat tissues as the inclusion level of linseed increased in the diet. This result is in agreement with similar studies in sheep [28] and in beef [26, 29, 30]. Similar results were also reported in monogastric animals, for example, in pigs [31], in rabbits [32], and in chickens [33]. The average increments of ALA for the muscles were 6.48 and 11.48 fold for L10 and L20, respectively, compared to control group (L0). As we hypothesized, this finding was higher than previously reported in sheep by [28, 34] or in beef [3543]. The higher increment of PUFA in goat muscles in current study can be attributed to a number of reasons. Firstly, the natural ability of goats to deposit PUFA is higher than that of cattle or sheep [15] since the species is considered to be one of the important factors affecting the FA composition in tissues [25, 44]. Secondly, the natural lower total lipids in goat meat compared to sheep or beef [14] play an important role in the high proportion of PUFA in goat meat, based on the findings of [44] in that “breeds or genetic types with a low concentration of total lipid in muscle, in which phospholipids are a high proportion of the total, will have higher proportions of PUFA in total lipid.” Thirdly, feeding whole seeds, which partially provides protection for the unsaturated FA from rumen microorganisms [11], collectively with the high rate of passage of feedstuffs through the rumen of goats compared to large ruminants [45], should also result in less extensive biohydrogenation of FA compared to large ruminants [46].

The increase in LC n-3 (EPA, DPA, and DHA), which are metabolic products of ALA, was higher than generally reported when feeding diets rich in ALA to sheep [4749] or beef [26, 50, 51]. This increase in LC n-3 may indicate that the desaturation and elongation occur in goat muscle when fed linseed was sufficient to increase the synthesis of the active metabolites EPA and DPA, and even DHA, which did not show a significant increase in the previously mentioned studies on sheep or beef. [27] also reported a linear response in DHA to the concentration of ALA in goat fed diets containing low ratios of n-6 : n-3. However, similar findings were also reported by [52] in the polar lipids of lambs when they replaced sunflower oil with a linseed diet. Nevertheless, the supplementation of fish oil reported to be more efficient in increasing these beneficial FAs in the meat of ruminants [49] and monogastric animals [53]. Although, using fish oil may cause undesirable off-odor and flavor and color changes and decrease meat shelf life [44, 54, 55].

In this study, the proportion of C18:1 trans-11 (vaccenic acid) was significantly higher in the most tissues of animals fed 20% linseed. This finding is reasonable since vaccenic acid is an intermediate product of the microbial biohydrogenation of LA and ALA [56]. The increase of vaccenic acid in animal tissues is preferable since it performs as a precursor for the tissue biosynthesis of CLA [57] and may exert health benefits similar to those related to CLA in humans [58]. The increase in the total CLA in the most tissues goats fed linseed agrees with [59] who confirmed that varied rumen microorganisms are capable of the creation of numerous CLA and 18 : 3 isomers from ALA acid. However, different studies showed that safflower and sunflower seed, which are considered as a source of LA, are more efficient in increasing the concentration of CLA compared to linseed [60].

The lower proportion of total SFA in most tissues (SS, ST, and subcutaneous fat) of goats fed linseed () is in disagreement with that reported in similar studies in sheep and beef. In this study, the reduction in total SFA is mainly due to the marked increase in the proportion of n-3 PUFA and the significant decrease in the proportion of palmitic acid. The decrease in the proportion of palmitic acid was also reported in young bulls fed linseed [61]. It is important to mention that palmitic acid was found to increase the plasma cholesterol level in human, while stearic does not exhibit such an effect [62, 63]; therefore, a high intake of palmitic acid increases the risk of atherosclerosis and cardiovascular diseases.

The increase in the PUFA: SFA ratios in all the muscles of the goats fed linseed, which ranged from 0.44 in LD of L10 up to 0.6 in SS of goats feed L20, can be mainly attributed to the increase in total n-3 PUFA since the proportion of n-6 PUFA was not affected by the treatment in most cases. Contrary, the increase in n-3 PUFA led to reduction in the n-6 : n-3 ratios, from values over 6.65 in the control group to values of 2.0 or less in the muscles of goats that were fed linseed. This result is similar to that reported in lambs fed different level of linseed [47] and higher than that observed in beef [26]. In general, the n-6 : n-3 ratio is highly affected by the type of unsaturated FA fed to the animals [46]. Although there was a similar decrease in n-6 : n-3 ratio in the adipose tissues form above 10 in the control to below 4, the PUFA:SFA ratio was very low (about 0.01) in these tissues and in this situation n-6 : n-3 ratio may give an indistinct indicator. Therefore, the absolute quantity of essential fatty acids consumed, rather than their n-6 : n-3 ratio, should be the first concern for the effective benefit from their intake [64].

5. Conclusion

The inclusion of linseed in the diet resulted in an increase in the proportion of ALA and total n-3 PUFA in muscles and adipose tissues of goats as the inclusion level of linseed increased. Furthermore, it increased the proportions of beneficial long-chains n-3 PUFA EPA, DPA, and DHA in the muscles. The inclusion of linseed at the 10% or 20% level can successfully improve the PUFA : SFA and n-6 : n-3 ratios to that recommended by the international health organization. Additionally, feeding linseed resulted in an increase in the total CLA, especially in the subcutaneous fat, and a decrease in the proportion of palmitic acid in all tissues.

Conflict of Interests

The authors declare that there is no conflict of interests regarding the publication of this paper.

Acknowledgments

The authors would like to thank the staff of ruminant farm and slaughter house, Department of Animal Science, Universiti Putra Malaysia, for their assistance management of the animals. Special thanks are due to Dr. Mahdi Ebrahimi for the technical assistance in fatty acid analysis.

References

  1. Y. Adkins and D. S. Kelley, “Mechanisms underlying the cardioprotective effects of omega-3 polyunsaturated fatty acids,” The Journal of Nutritional Biochemistry, vol. 21, no. 9, pp. 781–792, 2010. View at Publisher · View at Google Scholar · View at Scopus
  2. A. Laviano, S. Rianda, A. Molfino, and F. R. Fanelli, “Omega-3 fatty acids in cancer,” Current Opinion in Clinical Nutrition & Metabolic Care, vol. 16, no. 2, pp. 156–161, 2013. View at Publisher · View at Google Scholar · View at Scopus
  3. L. Hooper, R. L. Thompson, R. A. Harrison et al., “Risks and benefits of omega 3 fats for mortality, cardiovascular disease, and cancer: systematic review,” British Medical Journal, vol. 332, no. 7544, pp. 752–755, 2006. View at Publisher · View at Google Scholar · View at Scopus
  4. T. Hajjar, G. Y. Meng, M. A. Rajion et al., “Omega 3 polyunsaturated fatty acid improves spatial learning and hippocampal Peroxisome Proliferator Activated Receptors (PPARα and PPARγ) gene expression in rats,” BMC Neuroscience, vol. 13, no. 1, article 109, 2012. View at Publisher · View at Google Scholar · View at Scopus
  5. A. Cabiddu, L. Salis, J. K. S. Tweed, G. Molle, M. Decandia, and M. R. F. Lee, “The influence of plant polyphenols on lipolysis and biohydrogenation in dried forages at different phenological stages: in vitro study,” Journal of the Science of Food and Agriculture, vol. 90, no. 5, pp. 829–835, 2010. View at Publisher · View at Google Scholar · View at Scopus
  6. A. P. Simopoulos, “Omega-3 fatty acids in inflammation and autoimmune diseases,” Journal of the American College of Nutrition, vol. 21, no. 6, pp. 495–505, 2002. View at Publisher · View at Google Scholar · View at Scopus
  7. M. Enser, K. G. Hallett, B. Hewett, G. A. J. Fursey, J. D. Wood, and G. Harrington, “Fatty acid content and composition of UK beef and lamb muscle in relation to production system and implications for human nutrition,” Meat Science, vol. 49, no. 3, pp. 329–341, 1998. View at Publisher · View at Google Scholar · View at Scopus
  8. N. Scollan, J.-F. Hocquette, K. Nuernberg, D. Dannenberger, I. Richardson, and A. Moloney, “Innovations in beef production systems that enhance the nutritional and health value of beef lipids and their relationship with meat quality,” Meat Science, vol. 74, no. 1, pp. 17–33, 2006. View at Publisher · View at Google Scholar · View at Scopus
  9. A. P. Simopoulos, “Omega-6/omega-3 essential fatty acid ratio and chronic diseases,” Food Reviews International, vol. 20, no. 1, pp. 77–90, 2004. View at Publisher · View at Google Scholar · View at Scopus
  10. P. Warriss, Meat Science: An Introductory Text, CABI, Oxford, UK, 2000.
  11. C. G. Aldrich, N. R. Merchen, J. K. Drackley, S. S. Gonzalez, G. C. Fahey Jr., and L. L. Berger, “The effects of chemical treatment of whole canola seed on lipid and protein digestion by steers,” Journal of Animal Science, vol. 75, no. 2, pp. 502–511, 1997. View at Google Scholar · View at Scopus
  12. D. L. Palmquist, “Digestibility of cotton lint fiber and whole oilseeds by ruminal microorganisms,” Animal Feed Science and Technology, vol. 56, no. 3-4, pp. 231–242, 1995. View at Publisher · View at Google Scholar · View at Scopus
  13. P. Legrand, B. Schmitt, J. Mourot et al., “The consumption of food products from linseed-fed animals maintains erythrocyte omega-3 fatty acids in obese humans,” Lipids, vol. 45, no. 1, pp. 11–19, 2010. View at Publisher · View at Google Scholar · View at Scopus
  14. M. S. Madruga and M. C. Bressan, “Goat meats: description, rational use, certification, processing and technological developments,” Small Ruminant Research, vol. 98, no. 1–3, pp. 39–45, 2011. View at Publisher · View at Google Scholar · View at Scopus
  15. V. Banskalieva, T. Sahlu, and A. L. Goetsch, “Fatty acid composition of goat muscles and fat depots: a review,” Small Ruminant Research, vol. 37, no. 3, pp. 255–268, 2000. View at Publisher · View at Google Scholar · View at Scopus
  16. J. H. Lee and G. Kannan, “Influences of diets on fatty acid composition of edible tissues of meat goat,” in Goat Meat Production and Quality, I. T. K. O. Mahgoub and E. C. Webb, Eds., pp. 250–259, CABI, 2012. View at Google Scholar
  17. NRC, Committee on Nutrient Requirements of Small Ruminants—Nutrient Requirements of Small Ruminants: Sheep, Goats, Cervids, and New World Camelids, National Academy Press, 2007.
  18. G. A. Varga, H. M. Dann, and V. A. Ishler, “The use of fiber concentrations for ration formulation,” Journal of Dairy Science, vol. 81, no. 11, pp. 3063–3074, 1998. View at Publisher · View at Google Scholar · View at Scopus
  19. AOAC, Official Methods of Analysis, edited by K. Herlick, Association of Official Analytical Chemists, Arlington, Va, USA, 15th edition , 2007.
  20. P. J. van Soest, J. B. Robertson, and B. A. Lewis, “Methods for dietary fiber, neutral detergent fiber, and nonstarch polysaccharides in relation to animal nutrition,” Journal of Dairy Science, vol. 74, no. 10, pp. 3583–3597, 1991. View at Publisher · View at Google Scholar · View at Scopus
  21. J. Folch, M. Lees, and G. H. Sloane-Stanley, “A simple method for the isolation and purification of total lipides from animal tissues,” The Journal of Biological Chemistry, vol. 226, no. 1, pp. 497–509, 1957. View at Google Scholar · View at Scopus
  22. M. Rajion, J. McLean, and R. N. Cahill, “Essential fatty acids in the fetal and newborn lamb,” Asian-Australasian Journal of Animal Sciences, vol. 38, no. 1, pp. 33–40, 1985. View at Google Scholar · View at Scopus
  23. K. Abuelfatah, A. B. Z. Zuki, Y. M. Goh, and A. Q. Sazili, “Effects of dietary N-3 fatty acids on growth performance, apparent digestibility and carcass characteristics of crossbred boer goat under tropical conditions,” Asian Journal of Animal and Veterinary Advances, vol. 8, no. 6, pp. 775–785, 2013. View at Publisher · View at Google Scholar · View at Scopus
  24. V. B. Woods and A. M. Fearon, “Dietary sources of unsaturated fatty acids for animals and their transfer into meat, milk and eggs: a review,” Livestock Science, vol. 126, no. 1, pp. 1–20, 2009. View at Publisher · View at Google Scholar · View at Scopus
  25. J. D. Wood, M. B. Enser, R. I. Richardson, and F. Whittington, “Fatty acids in meat and meat products,” in Fatty Acids in Foods and Their Health Implications, C. K. Chow, Ed., pp. 87–107, CRC Press, Boca Raton, Fla, USA, 2007. View at Google Scholar
  26. N. Mach, M. Devant, I. Díaz et al., “Increasing the amount of n-3 fatty acid in meat from young Holstein bulls through nutrition,” Journal of Animal Science, vol. 84, no. 11, pp. 3039–3048, 2006. View at Publisher · View at Google Scholar · View at Scopus
  27. M. Ebrahimi, M. A. Rajion, Y. M. Goh, A. Q. Sazili, and J. T. Schonewille, “Effect of linseed oil dietary supplementation on fatty acid composition and gene expression in adipose tissue of growing goats,” BioMed Research International, vol. 2013, Article ID 194625, 11 pages, 2013. View at Publisher · View at Google Scholar · View at Scopus
  28. P. Bas, V. Berthelot, E. Pottier, and J. Normand, “Effect of level of linseed on fatty acid composition of muscles and adipose tissues of lambs with emphasis on trans fatty acids,” Meat Science, vol. 77, no. 4, pp. 678–688, 2007. View at Publisher · View at Google Scholar · View at Scopus
  29. E. Scholljegerdes and S. Kronberg, “Influence of level of supplemental whole flaxseed on forage intake and site and extent of digestion in beef heifers consuming native grass hay,” Journal of Animal Science, vol. 86, no. 9, pp. 2310–2320, 2008. View at Publisher · View at Google Scholar · View at Scopus
  30. E. J. Good, The Effects of Ground Flaxseed Supplementation on Performance, Carcass Composition, and Sensory Attributes of Finishing Cattle, Kansas State University, 2004.
  31. K. R. Matthews, D. B. Homer, F. Thies, and P. C. Calder, “Effect of whole linseed (Linum usitatissimum) in the diet of finishing pigs on growth performance and on the qualityand fatty acid composition of various tissues,” British Journal of Nutrition, vol. 83, no. 6, pp. 637–643, 2000. View at Publisher · View at Google Scholar · View at Scopus
  32. M. Kouba, F. Benatmane, J. E. Blochet, and J. Mourot, “Effect of a linseed diet on lipid oxidation, fatty acid composition of muscle, perirenal fat, and raw and cooked rabbit meat,” Meat Science, vol. 80, no. 3, pp. 829–834, 2008. View at Publisher · View at Google Scholar · View at Scopus
  33. M. J. Zuidhof, M. Betti, D. R. Korver et al., “Omega-3-enriched broiler meat: 1. Optimization of a production system,” Poultry Science, vol. 88, no. 5, pp. 1108–1120, 2009. View at Publisher · View at Google Scholar · View at Scopus
  34. F. Noci, F. J. Monahan, and A. P. Moloney, “The fatty acid profile of muscle and adipose tissue of lambs fed camelina or linseed as oil or seeds,” Animal, vol. 5, no. 1, pp. 134–147, 2011. View at Publisher · View at Google Scholar · View at Scopus
  35. K. Nuernberg, D. Dannenberger, G. Nuernberg et al., “Effect of a grass-based and a concentrate feeding system on meat quality characteristics and fatty acid composition of longissimus muscle in different cattle breeds,” Livestock Production Science, vol. 94, no. 1-2, pp. 137–147, 2005. View at Publisher · View at Google Scholar · View at Scopus
  36. G. Holló, K. Ender, K. Lóki, J. Seregi, I. Holló, and K. Nuernberg, “Carcass characteristics and meat quality of Hungarian Simmental young bulls fed different forage to concentrate ratios with or without linseed supplementation,” Archiv fur Tierzucht, vol. 51, no. 6, pp. 517–530, 2008. View at Google Scholar · View at Scopus
  37. R. T. Nassu, M. E. R. Dugan, M. L. He et al., “The effects of feeding flaxseed to beef cows given forage based diets on fatty acids of longissimus thoracis muscle and backfat,” Meat Science, vol. 89, no. 4, pp. 469–477, 2011. View at Publisher · View at Google Scholar · View at Scopus
  38. C. M. Kim, J. H. Kim, T. Y. Chung, and K. K. Park, “Effects of flaxseed diets on fattening response of Hanwoo cattle: 2. Fatty acid composition of serum and adipose tissues,” Asian-Australasian Journal of Animal Sciences, vol. 17, no. 9, pp. 1246–1254, 2004. View at Google Scholar · View at Scopus
  39. M. Juárez, M. E. R. Dugan, N. Aldai et al., “Beef quality attributes as affected by increasing the intramuscular levels of vitamin E and omega-3 fatty acids,” Meat Science, vol. 90, no. 3, pp. 764–769, 2012. View at Publisher · View at Google Scholar · View at Scopus
  40. M. L. He, T. A. McAllister, J. P. Kastelic et al., “Feeding flaxseed in grass hay and barley silage diets to beef cows increases alpha-linolenic acid and its biohydrogenation intermediates in subcutaneous fat,” Journal of Animal Science, vol. 90, no. 2, pp. 592–604, 2012. View at Publisher · View at Google Scholar · View at Scopus
  41. C. Mapiye, J. L. Aalhus, T. D. Turner et al., “Effects of feeding flaxseed or sunflower-seed in high-forage diets on beef production, quality and fatty acid composition,” Meat Science, vol. 95, no. 1, pp. 98–109, 2013. View at Publisher · View at Google Scholar · View at Scopus
  42. P. Albertí, I. Gómez, J. A. Mendizabal et al., “Effect of whole linseed and rumen-protected conjugated linoleic acid enriched diets on feedlot performance, carcass characteristics, and adipose tissue development in young Holstein bulls,” Meat Science, vol. 94, no. 2, pp. 208–214, 2013. View at Publisher · View at Google Scholar · View at Scopus
  43. T. D. Maddock, M. L. Bauer, K. B. Koch et al., “Effect of processing flax in beef feedlot diets on performance, carcass characteristics, and trained sensory panel ratings,” Journal of Animal Science, vol. 84, no. 6, pp. 1544–1551, 2006. View at Google Scholar · View at Scopus
  44. J. D. Wood, M. Enser, A. V. Fisher et al., “Fat deposition, fatty acid composition and meat quality: a review,” Meat Science, vol. 78, no. 4, pp. 343–358, 2008. View at Publisher · View at Google Scholar · View at Scopus
  45. P. J. van Soest, Nutritional Ecology of the Ruminant, Cornell University Press, 1994.
  46. K. Raes, S. de Smet, and D. Demeyer, “Effect of dietary fatty acids on incorporation of long chain polyunsaturated fatty acids and conjugated linoleic acid in lamb, beef and pork meat: a review,” Animal Feed Science and Technology, vol. 113, no. 1–4, pp. 199–221, 2004. View at Publisher · View at Google Scholar · View at Scopus
  47. P. Bas, V. Berthelot, E. Pottier, and J. Normand, “Effect of level of linseed on fatty acid composition of muscles and adipose tissues of lambs with emphasis on trans fatty acids,” Meat Science, vol. 77, no. 4, pp. 678–688, 2007. View at Publisher · View at Google Scholar · View at Scopus
  48. B. Aurousseau, D. Bauchart, E. Calichon, D. Micol, and A. Priolo, “Effect of grass or concentrate feeding systems and rate of growth on triglyceride and phospholipid and their fatty acids in the M. longissimus thoracis of lambs,” Meat Science, vol. 66, no. 3, pp. 531–541, 2004. View at Publisher · View at Google Scholar · View at Scopus
  49. G. Demirel, A. M. Wachira, L. A. Sinclair, R. G. Wilkinson, J. D. Wood, and M. Enser, “Effects of dietary n-3 polyunsaturated fatty acids, breed and dietary vitamin E on the fatty acids of lamb muscle, liver and adipose tissue,” British Journal of Nutrition, vol. 91, no. 4, pp. 551–565, 2004. View at Publisher · View at Google Scholar · View at Scopus
  50. K. Raes, L. Haak, A. Balcaen, E. Claeys, D. Demeyer, and S. de Smet, “Effect of linseed feeding at similar linoleic acid levels on the fatty acid composition of double-muscled Belgian Blue young bulls,” Meat Science, vol. 66, no. 2, pp. 307–315, 2004. View at Publisher · View at Google Scholar · View at Scopus
  51. N. D. Scollan, N.-J. Choi, E. Kurt, A. V. Fisher, M. Enser, and J. D. Wood, “Manipulating the fatty acid composition of muscle and adipose tissue in beef cattle,” British Journal of Nutrition, vol. 85, no. 1, pp. 115–124, 2001. View at Publisher · View at Google Scholar · View at Scopus
  52. E. Jerónimo, S. P. Alves, J. A. M. Prates, J. Santos-Silva, and R. J. B. Bessa, “Effect of dietary replacement of sunflower oil with linseed oil on intramuscular fatty acids of lamb meat,” Meat Science, vol. 83, no. 3, pp. 499–505, 2009. View at Publisher · View at Google Scholar · View at Scopus
  53. L. Haak, S. de Smet, D. Fremaut, K. van Walleghem, and K. Raes, “Fatty acid profile and oxidative stability of pork as influenced by duration and time of dietary linseed or fish oil supplementation,” Journal of Animal Science, vol. 86, no. 6, pp. 1418–1425, 2008. View at Publisher · View at Google Scholar · View at Scopus
  54. J. D. Wood, R. I. Richardson, G. R. Nute et al., “Effects of fatty acids on meat quality: a review,” Meat Science, vol. 66, no. 1, pp. 21–32, 2004. View at Publisher · View at Google Scholar · View at Scopus
  55. G. R. Nute, R. I. Richardson, J. D. Wood et al., “Effect of dietary oil source on the flavour and the colour and lipid stability of lamb meat,” Meat Science, vol. 77, no. 4, pp. 547–555, 2007. View at Publisher · View at Google Scholar · View at Scopus
  56. C. Harfoot and G. Hazlewood, “Lipid metabolism in the rumen,” in The Rumen Microbial Ecosystem, P. N. Hobson and C. S. Stewart, Eds., pp. 382–426, Springer, London, UK, 1997. View at Google Scholar
  57. J. M. Griinari, B. A. Corl, S. H. Lacy, P. Y. Chouinard, K. V. V. Nurmela, and D. E. Bauman, “Conjugated linoleic acid is synthesized endogenously in lactating dairy cows by Δ9-desaturase,” Journal of Nutrition, vol. 130, no. 9, pp. 2285–2291, 2000. View at Google Scholar · View at Scopus
  58. C. J. Field, H. H. Blewett, S. Proctor, and D. Vine, “Human health benefits of vaccenic acid,” Applied Physiology, Nutrition and Metabolism, vol. 34, no. 5, pp. 979–991, 2009. View at Publisher · View at Google Scholar · View at Scopus
  59. Y.-J. Lee and T. C. Jenkins, “Biohydrogenation of linolenic acid to stearic acid by the rumen microbial population yields multiple intermediate conjugated diene isomers,” The Journal of Nutrition, vol. 141, no. 8, pp. 1445–1450, 2011. View at Publisher · View at Google Scholar · View at Scopus
  60. F. Noci, F. J. Monahan, P. French, and A. P. Moloney, “The fatty acid composition of muscle fat and subcutaneous adipose tissue of pasture-fed beef heifers: influence of the duration of grazing,” Journal of Animal Science, vol. 83, no. 5, pp. 1167–1178, 2005. View at Google Scholar · View at Scopus
  61. M. Corazzin, S. Bovolenta, A. Sepulcri, and E. Piasentier, “Effect of whole linseed addition on meat production and quality of Italian Simmental and Holstein young bulls,” Meat Science, vol. 90, no. 1, pp. 99–105, 2012. View at Publisher · View at Google Scholar · View at Scopus
  62. M. A. Denke and S. M. Grundy, “Comparison of effects of lauric acid and palmitic acid on plasma lipids and lipoproteins,” The American Journal of Clinical Nutrition, vol. 56, no. 5, pp. 895–898, 1992. View at Google Scholar · View at Scopus
  63. T. Tholstrup, P. Marckmann, J. Jespersen, and B. Sandström, “Fat high in stearic acid favorably affects blood lipids and factor VII coagulant activity in comparison with fats high in palmitic acid or high in myristic and lauric acids,” The American Journal of Clinical Nutrition, vol. 59, no. 2, pp. 371–377, 1994. View at Google Scholar · View at Scopus
  64. V. Wijendran and K. C. Hayes, “Dietary n-6 and n-3 fatty acid balance and cardiovascular health,” Annual Review of Nutrition, vol. 24, pp. 597–615, 2004. View at Publisher · View at Google Scholar · View at Scopus