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Interaction of dietary high-oleic-acid sunflower hulls and different fat sources in broiler chickens

A. Viveros^*, L. T. Ortiz^*, M. L. Rodríguez^*, A. Rebolé^*, C. Alzueta^*, I. Arija^*, C. Centeno and A. Brenes^,1

^* Departamento de Producción Animal, Facultad de Veterinaria, Universidad Complutense, Ciudad Universitaria, 28040 Madrid, Spain; and Departamento de Metabolismo y Nutrición, Instituto del Frío, Consejo Superior de Investigaciones Científicas, Ciudad Universitaria, 28040 Madrid, Spain

¹ Corresponding author: abrenes{at}if.csic.es

	ABSTRACT

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The effect of dietary fat sources (high-oleic-acid sunflower seeds, HOASS; palm oil, PO; and high-oleic-acid sunflower oil, HOASO) and high-oleic-acid sunflower hulls (HOAS hulls; 40 g/kg of diet) on performance, digestive organ size, fat digestibility, and fatty acid profile in abdominal fat and blood serum parameters was evaluated in chickens (from 1 to 21 d of age). Bird performance and digestive organ size were not affected by either dietary fat source or sunflower hull supplementation. Fat digestibility in birds fed diets enriched (HOASS and HOASO) in monounsaturated fatty acids (MUFA) was increased compared with those fed the PO diet. The addition of sunflower hulls did not modify fat digestibility. The fatty acids pattern of abdominal fat reflected the dietary fat profile. The greatest concentrations of C16:0 and C18:0 were found in birds fed PO diets. The C18:1n-9 content was increased in birds that received HOASS and HOASO diets compared with those fed PO diets. The greatest content of C18:2n-6 was observed in birds fed HOASS diets. The ratio of polyunsaturated fatty acid (PUFA) to MUFA was significantly increased in birds fed PO diets compared with those fed HOASS or HOASO diets. The addition of sunflower hulls to the diets resulted in a decrease of C18:2n-6 and PUFA concentrations and PUFA:MUFA ratio in abdominal fat. Dietary fat sources and sunflower hulls modify blood triglycerides and serum lipoproteins. A decrease in triglyceride concentrations was observed in birds fed HOASS diets compared with those fed PO and HOASO diets. The greatest concentrations of serum high density, very low density (VLDL), and low density lipoproteins were found in birds receiving HOASO, PO, and HOASS diets, respectively. The addition of sunflower hulls to the diets caused an increase of serum triglycerides and VLDL concentrations. The MUFA-enriched diets had lower triglyceride and VLDL concentrations than did diets rich in saturated fatty acids. However, the sunflower hull addition had the opposite effect.

Key Words: high-oleic-acid sunflower seed • sunflower hull • fatty acid • lipoprotein profile • chicken

	INTRODUCTION

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Sunflower meal has been recognized as a viable feed ingredient in poultry diets. However, inconsistent results reported by several researchers might be attributed to, among other factors, the method of processing and the degree of dehulling (Senkoylu and Dale, 1999). The development by conventional genetic selection of new sunflower varieties, producing oils enriched in oleic acid (high-oleic-acid sunflower oils) at the expense of linoleic acid, has made sunflower oils highly competitive with other traditional and predominantly monounsaturated oils and constitutes an alternative approach to including this whole oilseed in broiler diets (Rodríguez et al., 2005; Ortiz et al., 2006; Rebolé et al., 2006).

There is some information in the literature regarding feeding broilers either sunflower hulls or unprocessed sunflower kernels (Arija et al., 1998; Ortiz et al., 1998; Rodríguez et al., 1998; Selvaraj and Purushothaman, 2004). The fiber content of sunflower seed (26.7%), which depends on the extent of dehulling, appear to be the most problematic aspect concerning the use of these whole seeds at high levels in chicken diets. A strong negative correlation between the fiber content of the whole sunflower seed and its TME has been found (Villamide and San Juan, 1998). The soluble and insoluble nonstarch polysaccharide fractions of sunflower seed have been extensively studied (Düsterhöft et al., 1991, 1993), representing 4.5 and 23.1%, respectively, of the meal. The total nonstarch polysaccharide comprised 42% cellulose, 24% pectic polysaccharides, 24% 4-0-methyl-glucuronoxylans, 5% glucomannans, and 4.5% fucoxyloglucans (Düsterhöft et al., 1993). The main component of soluble nonstarch polysaccharide is reported to be uronic acid (Irish and Balnave, 1993).

Although there are numerous studies on the effects of various dietary fiber sources on lipid absorption (Lairon, 2004), information regarding how different dietary fat types modify the effect of fiber on lipid absorption in chickens is limited. Dietary fat can alter the composition of the blood and the level of the serum lipoproteins that are subject to change by adding fat to diets (Crespo and Esteve-García, 2003; Celebi and Utlu, 2006). Generally, dietary saturated fatty acids (SFA) increase serum low density lipoprotein (LDL) content, whereas dietary polyunsaturated fatty acids (PUFA) decrease serum very low density lipoprotein (VLDL), LDL, and cholesterol concentrations.

Because sunflower hulls appear to be responsible for growth depression in birds fed diets containing high amounts of sunflower seed and because there are few studies on the effect of high-oleic-acid sunflower seeds (HOASS) and high-oleic-acid sunflower hull (HOAS) oils as natural monounsaturated fatty acid (MUFA) sources on lipoprotein metabolism, the objectives of the current study were to 1) investigate the effect of different fat sources (high-oleic-acid sunflower seed, palm oil, and high-oleic-acid sunflower oil) and sunflower hulls in chicken diets on performance, digestive organ size, fat digestibility, fatty acid profile in abdominal fat, and blood serum parameters, and 2) determine the possible interaction between hulls and fat sources.

	MATERIALS AND METHODS

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Test Products

Hulled and dehulled high-oleic-acid sunflower seeds (HHOASS and DHOASS, respectively; Helianthus annus L. cv. Saxo) and high-oleic-acid sunflower hulls (HOAS hulls) were obtained from a commercial supplier (SOS Cuétara, Andújar, Jaén, Spain). The batches of sunflower seed were cleaned by hand to eliminate any foreign material and remaining pieces of kernel. Before mixing the experimental diets, the test products were ground to pass through a 3.0-mm screen. High-oleic-acid sunflower refined oil (HOASO) and palm oil (PO) were provided by a commercial supplier (Pryca S.A., Madrid, Spain; Lípidos Santiga, Santa Perpetua de Mogoda, Barcelona, Spain, respectively). Chemical composition of HHOASS and fatty acid composition of the different dietary fat sources are shown in Tables 1 and 2, respectively.

A total of 144, 1-d-old male broiler chicks (Cobb-500 genetic line) were obtained from a commercial hatchery (Cobb Española S.A., Alcalá de Henares, Madrid, Spain). Birds were randomly housed in electrically heated starter battery brooders in an environmentally controlled room. The chicks were allocated to 24 pens, each pen containing 6 chicks, to receive 6 dietary treatments with 4 replicates of each treatment. Diets in mash form and water were provided ad libitum. Titanium dioxide was added at 4 g/kg to all diets as an indigestible marker. Chicks were subjected to artificial fluorescent illumination for 23 h/d and handled according to the principles for the care of the animals in experimentation established by the University Complutense of Madrid Animal Care and Ethics Committee in compliance with the Ministry of Agriculture, Fishery and Food for the Care and Use of Animals for Scientific Purposes. Six isonitrogenous and isocaloric diets were formulated and contained approximately 80 g/kg of added fat (Table 3). The ME of HHOASS and HOAS hulls used to formulate the experimental diets were determined in a preliminary experiment reported by Rodríguez et al. (2005) and by FEDNA (2003), respectively. In the diet, the HOAS hulls were included at 40 g/kg and partially substituted for starch. Fat was provided by dehulled HOASS, palm oil and HOAS oil. The treatments were as follows: 1) Corn-soybean (CS) + DHOASS; 2) CS + HHOASS; 3) CS + PO; 4) CS + PO + HOAS hulls; 5) CS + HOASO; and 6) CS + HOASO + HOAS hulls. At the end of the experimental period, birds were weighed and feed consumption was recorded for feed conversion computation.

At 21 d of age, 8 birds (2 per pen) were randomly selected from each treatment group after an overnight fast. Serum was prepared from blood obtained by cardiac puncture for subsequent determination of cholesterol, triglycerides and lipoprotein profile. After the birds were killed by cervical dislocation (12 per treatment, 3 per pen), their viscera were removed. Liver, pancreas, and spleen were weighed, and duodenum, jejunum, ileum, and ceca were measured. A sample of abdominal fat (2 per pen) was also removed for fatty acid determination. Clean stainless steel collection trays were placed under each cage (2 d before the birds were killed), and excreta from the birds were collected for 48 h. A sub-sample of excreta was collected in polyethylene bags, freeze-dried, ground, and subsequently analyzed for dry matter, ether extract, and titanium dioxide. The samples were frozen and stored at –20°C until required.

Analytical Methods

Dry matter (method 930.15), CP (method 976.05), and crude fiber (method 978.10) were analyzed according to the methods of the Association of Official Analytical Chemists (1995). Ether extract was determined by extraction in petroleum ether following acidification with 4 N HCl solution (Wiseman et al., 1992). Titanium dioxide was determined colorimetrically as reported by Short et al. (1996). Total lipids from abdominal fat were extracted following the procedure described by Folch et al. (1957). The lipid extracts were esterified with a mixture of boron trifluoride (in 10% methanol), hexane and methanol (35:20:45, vol/vol/vol) (Morrison and Smith, 1964). The resultant fatty acid methyl esters were analyzed on a Chrompack CP 9001 gas chromatograph (Chrompack Instrumental BV, Middelburg, the Netherlands) equipped with a WCOT fused silica capillary column (length 30 m; internal diameter 0.32 mm; film thickness 0.50 µ m), and a flame-ionization detector. Analyses were performed with a temperature program from 170 to 250°C that increased by 3.5°C/ min. The carrier gas was nitrogen at a flow rate of 4.5 mL/min. Pentadecanoic acid was used as internal standard (Sigma-Aldrich Quimica, S.A. 28100 Alcobendas, Madrid). Contents of MUFA, PUFA, and SFA as well as PUFA:MUFA and unsaturated fatty acid (UFA):SFA ratios were calculated. Blood serum was analyzed for cholesterol (cholesterol oxidase-peroxidase-enzymatic-colorimetric method) and triglycerides (glycerol-3-oxidase-peroxidase-enzymatic-colorimetric method) following the methods described by SpinReact S.A. (St. Esteve de Bas, Girona, Spain). Serum protein electrophoresis was performed using a cellulose acetate method (Lumeij, 1987; Cray and Tatum, 1998).

Statistical Analysis

Data were analyzed as a 3 x 2 factorial arrangement with 3 fat sources with and without the addition of sunflower hull. The statistical model used was

where Y_ijk is the individual observation, µ is the experimental mean, P_i is the fat source effect, E_j is the hull effect, R_k is the replication effect, and e_ijk is the error term.

Data were subjected to ANOVA using the GLM procedure of SAS (SAS Institute, 2001), and single degree of freedom orthogonal contrasts were used to separate treatments. Significant differences among treatment means were determined at P < 0.05 by Duncan’s multiple range test.

	RESULTS

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The proximate composition of HHOASS (Table 1) revealed a high content of crude fat (444 g/kg) and a moderate amount of CP (180 g/kg) and crude fiber (126 g/kg). As expected, oleic acid was the major fatty acid, accounting for 80% of the total fatty acid content. Fatty acid composition of diets (Tables 2 and 3) reflected the fatty acid profile of the added fat sources. The fatty acid composition of the dietary fat sources showed that the main SFA in PO was palmitic acid (C16:0; 454 g/kg of total fatty acids), the predominant MUFA in HOASS, HOASO, and PO was oleic acid (C18:1n-9; 807, 730, and 389 g/kg of total fatty acids, respectively), and the main PUFA in HOASS, HOASO, and PO was linoleic acid (C18:2n-6; 72, 163, and 98 g/ kg of total fatty acids, respectively).

Bird Performance and Digestive Organ Size

The effects of inclusion of dietary fat sources and HOAS hulls in chicken diets on performance are summarized in Table 4. The main effect data indicated that performance was not affected by fat source or HOAS hull supplementation. A significant interaction (P < 0.01) was observed between fat source and sunflower hulls for weight gain and feed conversion, indicating a positive effect of hulls in birds fed the HOASO diet.

The effect of different dietary fat sources and HOAS hulls on fat digestibility and fatty acid composition of abdominal fat is shown in Table 6. The main effects data showed a significant increase (P < 0.001) in fat digestibility in birds fed HOASO diets compared with those fed PO and HOASS diets (10 and 6%, respectively). The results also showed significant differences in the fatty acid concentrations of abdominal fat among dietary fat sources. The contents of the major fatty acids, palmitic acid (C16:0), stearic acid (C18:0), oleic acid (C18:1n-9), and linoleic acid (C18:2n-6) in abdominal fat reflected the fatty acid profile of the dietary fat. The concentrations of C16:0 and C18:0 in abdominal fat were increased (P < 0.001) in birds fed PO diets (up to 218 and 70%, respectively) compared with those fed HOASS and HOASO diets. The C18:1n-9 content was greater (P < 0.001) in birds fed HOASS (46%) and HOASO (47%) diets than in those fed PO diets. The C18:2n-6 content was significantly greater (up to 9%; P < 0.01) in HOASS diets compared with PO and HOASO diets. The PUFA:MUFA and UFA:SFA ratios were significantly (P < 0.001) increased (up to 44%) and reduced (up to 72%) in birds fed PO diets compared with those fed HOASS and HOASO diets, respectively. Likewise, the addition of HOAS hulls to diets resulted in a decrease (P < 0.001) of C18:2n-6 (10%) and PUFA (9%) concentrations and PUFA:MUFA ratio (9%). The significant interaction found between fat source and HOAS hulls for UFA:SFA ratio indicated that the addition of hull only affected this ratio in HOASS and HOASO diets.

The effect of inclusion of dietary fat sources and HOAS hulls in chicken diets on blood parameters (cholesterol, triglycerides, and lipoprotein profiles) is reported in Table 7. The main effects data showed a decrease (up to 25%; P < 0.01) in triglyceride concentrations in birds fed HOASS diets compared with those fed PO and HOASO diets. An increase (up to 10%; P < 0.001) in high density lipoprotein (HDL) concentration was observed in birds fed HOASO diets compared with those fed PO and HOASS diets. An increase in VLDL (up to 103%; P < 0.001) concentration was observed in birds fed PO diets compared with those fed HOASS and HOASO diets. The LDL concentration was increased up to 56% (P < 0.001) in birds fed HOASS diets compared with those fed PO and HOASO diets. Likewise, the addition of HOAS hulls to the diets caused an increase (P < 0.01) in triglyceride (23%) and VLDL (31%) concentrations. A significant interaction was observed between fat source and hulls for cholesterol (P < 0.05) and triglyceride (P < 0.01) concentrations, indicating that the addition of hulls affected the cholesterol concentration only in the HOASS diet and affected triglyceride concentration in HOASO diets.

	DISCUSSION

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The results indicated that bird performance was not affected by the inclusion of different fat sources (MUFA and SFA), which is in agreement with data reported by Sanz et al. (1999), Villaverde et al. (2004), and Wongsuthavas et al. (2007) in studies comparing PUFA and SFA fat sources. However, controversial results have been reported by Pinchasov and Nir (1992) and Zollitsch et al. (1997) who showed an improved gain to feed ratio in broilers when dietary PUFA intake increased, which could be attributed to greater intestinal uptake of unsaturated fatty acids.

The concentration of HOAS hulls used in this experiment (40 g/kg of diet) corresponds to the amount of hulls contained in 150 g of HHOASS (the HOASS diet). Some researchers have emphasized that the fiber content of hulls appears to be the main problematic aspect concerning the use of sunflower seeds at a high level in chicken diets (Zatari and Sell, 1990). This could be attributed to the bulking effect and dietary nutrient dilution of the fiber, which can reduce the ME content of the diet (Senkoylu and Dale, 1999) and cause alterations in the jejunal mucosa (Arija et al., 2000). In the current experiment, and confirming the results obtained by Arija et al. (1998) and Suresh et al. (2000), the inclusion of hulls in the diet did not modify performance. The percentage of fiber in the hull-containing diet was approximately 50% greater than in the free-hull diets. This difference appears to be insufficient to produce a negative effect on growth performance. As a consequence, there was no modification regarding the relative organ sizes and intestinal lengths.

Fat Digestibility and Fatty Acid Profile of Abdominal Fat

Dietary fat type had a marked effect on apparent fat digestibility. A significant increase by the addition of HOASS and HOAS oil in the chicken diets is reported compared with those birds fed the diet containing SFA (PO diet). It is well known that UFA from triglycerides have greater digestibility and absorption rates than SFA. The greater apparent absorption values obtained with the MUFA diets are in accordance with those reported by Villaverde et al. (2004) and Smink et al. (2008) using PUFA-rich diets. Monoglycerides and long-chain UFA, when linked to the conjugated bile salts, promptly form micelles, whereas SFA have less ability to form micelles because of their characteristic low polarity (Wiseman and Lessire, 1987). The significant differences observed in fat digestibility between HOASO and HOASS diets could be attributed to the limited ability of the chicken in the digestion process to extract oil from the intact cells of the seed compared with those diets containing extracted oil (Arija et al., 1998). The addition of sunflower hull to the experimental diets did not affect the apparent digestibility of fat. Similar results have been reported by Akiba and Matsumoto (1980) using 4% of cellulose in chicken diets.

In the current experiment, the fatty acid pattern of abdominal fat reflected the dietary fat profile, which is consistent with the findings of other researchers (O’Neill et al., 1998; Ortiz et al., 2006). Regarding the effect of sunflower hulls on abdominal fat composition, a significant decrease (mainly in HOASS and HOASO diets) was observed in C18:2n-6 and PUFA concentrations and PUFA:MUFA ratio. This is probably related to the lesser concentration of C18:2n-6 in the diets containing HOAS hulls or to changes in lipid metabolism. There is little information in the literature regarding the effect of dietary hulls and fat on abdominal fat composition. Evidence of an interaction between dietary fiber and fat on fatty acid digestibility has been observed by Choct and Annison (1992) and Smits et al. (2000). These researchers reported that the addition of highly viscous fiber depressed digestibility of saturated long-chain fatty acids and did not affect the unsaturated long-chain fatty acids in chickens fed different fat sources. Although in the current experiment we have not determined fatty acid digestibility, this interaction could also affect fatty acid deposition.

Blood Serum Parameters

Serum triglyceride concentrations showed a reduction in birds fed HOASS and HOASO diets compared with those birds fed the PO diet. A decrease in serum triglyceride has been observed in chicken and laying hens after substitution of dietary SFA for PUFA or by decreasing dietary SFA:(SFA + MUFA) ratio (Sanz et al., 2000; Celebi and Utlu, 2006; Wongsuthavas et al., 2007). The current study demonstrated that reduced circulating triacylglycerol concentrations were not specific to n-3 PUFA, because birds fed HOASS and HOASO diets (rich in MUFA) also had reduced serum triacylglycerol concentrations. Consistent with the present findings, Shimomura et al. (1990) and Lai and Ney (1995) observed that rats fed diets high in unsaturated diets (corn oil or sunflower oil) showed significantly lower postprandial plasma triacylglycerol concentrations compared with those fed saturated fats (palm oil, butterfat, and tallow) that could be attributed to greater activity of lipoprotein lipase (LPL) in muscle. A previous experiment conducted in our laboratory (Rebolé et al., 2006) showed that the substitution of saturated fat (PO) for MUFA (HOASS) resulted in a decrease in abdominal fat that could also be related to an increase in LPL activity and, consequently, with a reduction in plasma triacylglycerol concentrations. In fact, Soriguer et al. (2003) confirmed that when the n-9 fatty acid (oleic acid) content of rat adipocytes increased, the lipolytic activity of the adipocytes increased and the antilipolytic capacity of the insulin decreased.

In the current experiment, VLDL, LDL, and HDL concentrations varied depending on the dietary fat sources. The VLDL concentration was highly correlated with abdominal fat and is a good indicator of fat deposition in birds (Whitehead and Griffin, 1982; Grunder and Chambers, 1988). Consistent with this finding, a significant decrease in VLDL concentration was observed in chicks fed MUFA-enriched diets (HOASS and HOASO diets) compared with those fed PO diets. Similar observations were made by Crespo and Esteve-García (2003) and Celebi and Utlu (2006) who showed a reduction in the VLDL concentration of birds fed a sunflower oilenriched diet. Dietary n-3 PUFA suppress hepatic fatty acid synthesis and reduce triglyceride synthesis (Harris, 1989), thereby limiting VLDL secretion. Several authors have also reported that dietary PUFA, especially linoleic acid, and n-3 fatty acids inhibit ⁹-desaturase activity (Lochsen et al., 1997). Because ⁹-desaturase activity has been correlated with liver VLDL-triglyceride secretion (Legrand et al., 1987), linoleic acid may also inhibit VLDL-triglyceride secretion. However, in the current study, PUFA concentrations, in particular C18:2n-6, were similar in all experimental diets. For this reason, the decrease of blood VLDL concentration may have been because of the greater concentration of MUFA (oleic acid) in chickens fed HOASS and HOASO diets. In addition, McNamara (1995) reported that the intake of PUFA and MUFA may reduce plasma triacylglycerol-rich lipoproteins, changing the composition and the catabolism of VLDL.

With regard to the LDL and HDL concentrations, the birds fed HOASO diets had the least and the greatest concentrations, respectively, of the serum lipoproteins. Metabolized VLDL can be recovered from chicken plasma in LDL (Sanz et al., 2000). Little is known about the clearance of VLDL remnants (LDL) from avian plasma, but it probably occurs through a similar mechanism to that in mammals (Griffin and Hermier, 1988). High density lipoprotein contains high proportions of cholesterol, cholesterol esters, and phospholipids, and low proportions of triglycerides. It also constitutes the major reservoir of LPL activator in chicken plasma (Griffin et al., 1982). In the current experiment, the greatest HDL concentration in birds fed HOASO diets indicates that the extrahepatic cholesterol pathway may be more efficient in birds fed these diets, along with improved utilization of cholesterol by the liver, in the form of bile.

Our results also showed that the addition of hulls modified some blood parameters (Table 7). Certain sources of dietary fiber have been reported to be hypocholesterolemic in both humans and animals. In general, soluble fiber appears most effective in lowering plasma cholesterol, whereas insoluble fiber such as cellulose is not effective or can elevate plasma cholesterol (Schneeman et al., 1984; Kreuzer et al., 2002). In the current experiment, serum cholesterol concentration was not changed by the addition of sunflower hulls. The high concentration of insoluble fraction in sunflower hulls (approximately 50% of cellulose and 26% of lignin content) reported by Düsterhöft et al. (1993) and Arija et al. (1998) could explain these results. Sundaravalli et al. (1971) also showed that cellulose has almost no binding capacity with cholesterol and therefore is not reported to have a hypocholesterolemic effect, unless a very large amount was fed. Increased serum triglyceride and VLDL concentrations were observed with the addition of hulls to the diets. These results could be related to the lower C18:2n-6 and PUFA concentrations in hull-containing diets (approximately 9 and 12% less, respectively). These differences have also been detected in the abdominal fatty acid profile of fat with a significant reduction in both parameters.

The effect of sunflower hulls on blood parameters was modified by the dietary fat given simultaneously. In fact, a significant interaction was observed between fat and hulls for serum cholesterol and triglyceride concentrations. Despite the numerous studies conducted on the effect of dietary fiber on lipid absorption, to our knowledge there is little information in chickens regarding the interaction of fiber with different types of dietary fats. In pigs, Kreuzer et al. (2002) showed that the addition of pectins in PUFA- and MUFA-enriched diets caused an increase in blood triglyceride, HDL, and LDL concentrations. However, the pectins did not modify these blood parameters in saturated diets. Likewise, Ikeda et al. (1989) reported, in rats, that although the composition of lymph lipids generally reflected the type of fat administered, there was a detectable difference in the percentage of major fatty acids due to the type of dietary fibers. Possibly, several complex mechanisms may be involved with the interactive effect of fiber and fat that requires further investigation.

It is concluded that dietary fat sources and sunflower hull did not affect bird performance. Fat digestibility was not affected in birds fed sunflower hull diets and was increased in HOASO and HOASS diets. Abdominal fat fatty acid composition was markedly influenced by dietary fatty acid profile. Sunflower hulls expressed a certain potential to reduce linoleic acid and PUFA concentrations in abdominal fat, which could contribute to increase the melting point, firmness, and oxidation stability of fat in the broiler carcass. Broilers fed MUFA-enriched diets (HOASS and HOASO) had lower serum triglyceride and VLDL concentrations than those fed SFA-enriched diets. However, the sunflower hull addition had the opposite effect. An appropriate combination of dietary fat sources and fiber might induce changes in lipid absorption and deposition and blood lipid profile.

	ACKNOWLEDGMENTS

 
The authors wish to thank the Comisión Interministerial de Ciencia y Tecnología for financial support of this investigation (Project AGF2001–1116). The authors thank SOS Cuetara S.A. (Andújar, Jaén, Spain) for supplying the high-oleic-acid sunflower seeds used in this study.

Received for publication June 3, 2008. Accepted for publication September 8, 2008.

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