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METABOLISM AND NUTRITION |






* Feed Innovation Services BV, Generaal Foulkesweg 72, 6703 BW, Wageningen, the Netherlands;
Wageningen University, Animal Nutrition Group, PO Box 338, 6700 AH, Wageningen, the Netherlands;
Utrecht University, Faculty of Veterinary Medicine, Department of Nutrition, 3508 TD, Utrecht, the Netherlands; and
Cargill Refined Oils Europe, Jan van Galenstraat 4, 3115 JG, Schiedam, the Netherlands
1 Corresponding author: smink{at}fisbv.nl
| ABSTRACT |
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Key Words: chicken palm oil fatty acid digestibility
| INTRODUCTION |
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Palm oil is of vegetable origin but is rich in the saturated fatty acid palmitic acid (C16:0); the content is about 45% of the total fatty acids. The use of palm oil in broiler diets is attractive, because it is a saturated source that may be associated with a positive influence on meat firmness. However, saturated fats rich in long-chain fatty acids (> 14 C atoms) are less digestible than fats high in medium-chain fatty acids or unsaturated fatty acids (Renner and Hill, 1961; Young, 1961; Garrett and Young, 1975; Vila and Esteve-Garcia, 1996). In addition, a high fraction of C16:0 in palm oil is bound at the sn-1 or sn-3 position of the glycerol molecule (Breckenridge, 1978; Mu and Høy, 2004). Long-chain saturated fatty acids on the sn-1 and sn-3 positions are thought to be absorbed less efficiently than those bound on the sn-2 position. This is because of the more hydrophilic character of the monoglyceride in comparison with, by lipase hydrolyzed, fatty acids from the sn-1 or sn-3 position of the glycerol backbone.
The position of fatty acids in triacylglycerols can be manipulated by hydrolysis and chemical reesterification (Mukherjee and Warwel, 1997). Randomization is a process of nonspecific random esterification of fatty acids at the 3 positions of the glycerol molecule. In lard, the C16:0 is mainly bound at the sn-2 position of the glycerol molecule (Breckenridge, 1978; Mu and Høy, 2004). Randomization of lard does decrease the digestibility of C16:0 in broilers (Renner and Hill, 1961). A higher digestibility of long-chain saturated fatty acids at the sn-2 position is probably also responsible for a higher deposition rate of these fatty acids in broilers (Scheeder et al., 2003).
As far as known, the effect of randomization of palm oil on C16:0 digestibility and deposition in broiler chickens has not yet been quantified. We hypothesized that randomization of palm oil would increase its digestibility, in particular that of its C16:0 component. In this study, our hypothesis was put to the test. Broiler chickens were fed diets containing either native or randomized palm oil, and the digestibility and deposition of fatty acids were measured.
In addition to studying the effect of randomization of palm oil, we also assessed the effect on digestibility of the degree of fatty acid saturation and chain length of fatty acids. Mathematical models used to calculate the ME of fat sources in broiler nutrition are generally based on the contents of long-chain unsaturated and saturated fatty acids without taking into account chain length differences between C16:0 and C18:0 (Ketels, 1994; Wiseman et al., 1998). It is well known that saturation of unsaturated fatty acids will decrease their digestibility and that of the oils they are components of (Ketels, 1994; Wiseman et al., 1998). As far as we know, it is not known to what extent the total digestibility of oils varies when their saturated fatty acid constituents have different chain lengths. As a positive control in this study, we determined the effect of saturation by comparing sunflower oil and a 50/50 mix of fully hydrogenated sunflower oil and sunflower oil. To assess effect of chain length in the form of C16:0 vs. C18:0, we used the comparison of randomized palm oil vs. the mixture of hydrogenated and native sunflower oil.
| MATERIALS AND METHODS |
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One-day-old female broilers (Ross 308) were purchased from a local hatchery. On arrival, they were wing-banded, weighed, and housed in wire-floor, suspended cages. Each of the 4 experimental groups contained 12 replicates (cages). The experiment started with 6 birds per cage during the starter period of 2 wk. Then up to 2 randomly chosen birds in each cage were removed, so there were 4 birds for the grower-finisher period. Continuous lighting was provided throughout the experiment. The temperature in the cage at arrival was 32°C and was decreased gradually to ambient temperatures during the course of the experiment.
Diets
The birds received a starter feed until d 14 and a grower-finisher feed between d 15 to 35. The composition of the diets is presented in Table 1
. Four different fat sources were used in the starter and grower-finisher diets. The fat sources were delivered by Cargill (Rotterdam-Botlek, the Netherlands). The diets were in pelleted form (2.5 mm). The diets were fed on an ad libitum basis, and the birds had free access to water. The inclusion level of experimental fat was 4 and 8% (wt/wt) in the starter and grower-finisher period, respectively (Table 1
). The inclusion levels are in agreement with European high-energy and fat diets for broiler chickens. The 4 experimental fats consisted of sunflower oil (SO), a 50/50 mix of fully hydrogenated sunflower oil and sunflower oil (HSO + SO), palm oil (PO), and chemical randomized palm oil (RPO). The analyzed fatty acid composition of the experimental fats is presented in Table 2
. The macronutrient and fatty acid composition of the diets are presented in Table 3
. The experimental fats allow assessment of the effect of saturation (HSO + SO vs. SO or C18:0 vs. C18:1 + C18:2), the effect of the chain length (HSO + SO vs. RPO or C18:0 vs. C16:0), and the effect of the position of long-chain saturated fatty acids on the glycerol molecule (PO vs. RPO or 20 vs. 45% of C16:0 at the sn-2 position) in a situation with a high fat intake.
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Excreta were collected in the starter period from d 10 to 14 and in the growing-finishing period from d 31 to 33. It is known that fat digestibility in broiler chickens is lower during the starter than grower-finisher period (Katangole and March, 1980; Ketels, 1994). Thus, we collected excreta during the 2 periods to determine apparent fecal digestibility of total fat and individual fatty acids. Excreta were collected quantitatively per cage, dried at 60°C, weighed, and ground. On d 35, two broilers per pen were used to determine the fatty acid composition of breast meat and abdominal fat. Crude fat determination of diets and excreta were determined with the acid hydrolysis method (AOAC, 1975). To determine the fatty acid composition of the diets, breast meat, and feces, a 10-g sample was extracted with a choloroform:methanol (2:1, vol/vol) mixture according to the method of Folch et al. (1957). Then, 20 to 25 mg of the extracted fat was saponified with 0.5 M methanolic sodium hydroxide and methylated with boronitrifluoride in methanol according to the method of Metcalfe et al. (1966). The fatty acid methyl esters obtained were separated and analyzed by gas chromatography. The fat of abdominal fat was directly saponified and methylated and the fatty acid composition determined with gas chromatography. The concentration of fatty acids at the sn-2 position of PO and RPO was determined by gas-liquid chromatography after hydrolysis with pancreas lipase. The diets were analyzed according to the Dutch Normalization Institute for DM (NEN 3332), ash (NEN 3329), crude fiber (NEN 3326), N. The N in the diets was analyzed with the Kjeldahl method (NEN 3145). Crude protein (g) was calculated as 6.25 x N (g).
Statistical Analysis
Cage served as the experimental unit so that there were 12 units per diet. The effect of diet on digestibility of fat and fatty acids, profiles of breast meat, and adipose tissue were statistically analyzed by 1-way ANOVA with diet as factor. In case of a significant diet effect, the effects of dietary fatty acid saturation (HSO + SO vs. SO), chain length (HSO + SO vs. RPO), and position on the glycerol molecule (RPO vs. PO) were analyzed via a least square means contrast test. The level of statistical significance was preset at P < 0.05. Results are presented as least square means and a pooled SEM. Statistical analysis was done with the SAS program (SAS JMP, 2000).
| RESULTS |
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| DISCUSSION |
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Birds fed the HSO + SO diet in comparison with those fed with the SO diet had a significant lower digestibility for the predominant long-chain saturated fatty acids (C16:0 and C18:0) but not for the unsaturated fatty acids. This agrees with results of studies in which tallow was compared with soybean oil (Dänicke et al., 1999; Knarreborg et al., 2004). However, Dänicke et al. (2000) estimated a significant linear dose-dependent decrease for digestibility of both the saturated and unsaturated fatty acids after replacing soybean oil by tallow. The digestibility of saturated fat may be improved by the addition of an unsaturated fat source (Sibbald, 1978), but according to Wiseman et al. (1998), it is not plausible to suggest a synergy between dietary unsaturated and saturated fat in broiler chickens. There might be a dose effect and a limiting digestibility for the amount of long-chain saturated fat. Ketels and De Groote (1989) showed that the inclusion of increasing levels of a saturated fat source like tallow decreased its digestibility, but no such effect was seen with soybean oil. The dose level of long-chain saturated fatty acids in the HSO + SO group was relatively high, being approximately 3 and 6% of the diet in the starter and grower-finisher period, respectively. This dietary long-chain saturated fat content corresponds with a dietary dose level of 7 and 14% of a saturated fat source like tallow in the starter and the grower-finisher period, respectively. Thus, the low digestibility of C18:0 in the HSO + SO group may relate to the high inclusion level of C18:0. However, a high intake itself tends to raise the apparent digestibility because of the diminishing effect of the relatively constant excretion of endogenous origin will be smaller. The calculated C18:0 digestibility was negative for the HSO + SO group in the starter period. This is probably caused by the combination of a low digestibility and endogenous C18:0 production in the lower gut. In addition, the C18:0 digestibility coefficient may be underestimated due to biohydrogenation of C18:1 and C18:2 in the large intestine.
In calculation models, the dietary U/S ratio is used to predict the digestibility, ME value (Ketels, 1994; Wiseman et al., 1998), or both. The S in the formulas mainly reflects the sum of C16:0 and C18:0. The U/S ratios of the HSO + SO and RPO diets were similar. The S digestibility was significantly lower in the HSO + SO group compared with the RPO group, but there was no difference in total U digestibility between the groups. The S fraction consists of C18:0 with relatively low digestibility and C16:0 with higher digestibility, as was shown earlier (Kussaibati et al., 1982; Dänicke et al., 2000; Smits et al., 2000).
Palm oil contains a high content of C16:0 that is predominantly located at the sn-1 and sn-3 positions of the glycerol molecule. Randomization yielded palm oil with one-third of the C16:0 at the sn-2 position. As indicated in the introduction section, we hypothesized that randomization of palm oil would increase its digestibility, in particular that of its C16:0 component. There was no statistically significant effect of randomization on crude fat digestibility and the digestibility of individual fatty acids. Thus, our hypothesis would be rejected. However, we did find a systematic (P-value between 0.05 and 0.1) increase in the group mean digestibilities of C16:0, C18:0, and S during the growing-finisher period. Randomization of palm oil had increased the group mean digestibility of C16:0 by 5.8% units. The randomization-induced increase of C16:0 digestibility is in line with results found in rats (Renaud et al., 1995). The effect of randomization of palm oil on the digestibility of C16:0 is also supported by significantly higher concentration of C16:0 in breast meat and abdominal fat. It is not known to what extent the increase in C16:0 in breast meat and abdominal fat is caused by increased digestion of C16:0 or increased de novo synthesis. A large part of deposition of S in broiler chickens fed with lipid-rich diets does not originate from the de novo production (Villaverde et al., 2006). The deposition of C16:0 will come predominantly from dietary C16:0. It is concluded that the raised level of C16:0 in breast meat and abdominal fat of the birds fed the RPO diet was caused by the increased digestibility of C16:0. Scheeder et al. (2003) have also reported that the position of saturated fatty acids in the glycerol molecule influence their position in the body of poultry. Triacyglycerols are hydrolyzed in the lumen of the gastrointestinal tract in free fatty acids and 2-monoglycerides. Free fatty acids have a lower digestibility than monoglycerides (Garrett and Young, 1975). Due to a randomization of palm oil, the composition of free fatty acids in the lumen of the intestine will be changed. The amount of free S decreased and free U increased. The decrease of the fat digestibility by increasing the amount of dietary free fatty acids is higher in case of saturated fat in comparison with an unsaturated fat source (Wiseman and Salvador, 1991). The increased content of unsaturated fatty acids at the sn-1,3 position after randomization did not affect its digestibility.
From the results of the grower-finisher period (Table 6
), the digestibility of C16:0 at the sn-2 and sn-1,3 positions can be calculated using the following formulas:
![]() | ([1]) |
![]() | ([2]) |
Multiplying the first formula by 1/0.67 gives the value for sn-1,3 being:
![]() | ([3]) |
The digestibility of C16:0 at the sn-2 position can be calculated by using formula [2] and [3]:
![]() | ([4]) |
The digestibility of C16:0 at sn-2 is calculated to be 89.5%. The use of this value in formula [1] gives a calculated digestibility of 51.0% for C16:0 at the sn-1,3 position.
Randomization also resulted in a numerically (P = 0.09) increased digestibility of C18:0 during d 31 to 33. The calculated digestibility for C18:0 at the sn-2 and sn-1,3 positions was 84 and 37%, respectively. Thus, the positions of C16:0 and C18:0 at the glycerol molecule are important factors to determining the digestibility of palm oil.
In conclusion, the present data support the idea that randomization of palm oil, which raises the content of C16:0 at the sn-2 position, improves the digestibility of its C16:0 component. Randomization of palm oil increased the group mean C16:0 digestibility by 5.8% units during the grower-finisher period. It was calculated that C16:0 at the sn-2 position was digested much more efficiently than C16:0 at the sn-1,3 position. Randomization of palm oil increased the incorporation of C16:0 into breast meat and abdominal fat and lowered the U/S ratios in these tissues. Thus, randomized palm oil may be used as a vegetable oil in broiler nutrition with positive effects on saturated fat digestibility and firmness of meat compared with native palm oil. For the determination of the digestibility or metabolic energy value of palm oil to replace animal fat, both the differences between C16:0 and C18:0 and the position C16:0 on the glycerol molecule are relevant.
Received for publication August 23, 2007. Accepted for publication November 25, 2007.
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