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METABOLISM AND NUTRITION |
* Instituto del Frío, Consejo Superior de Investigaciones Científicas (CSIC), Departamento de Metabolismo y Nutrición, José Antonio Novais, 10, 28040 Madrid, Spain; and Departamento de Producción Animal, Facultad de Veterinaria, Ciudad Universitaria, 28040 Madrid, Spain
1 Corresponding author: abrenes{at}if.csic.es
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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Key Words: high oleic acid sunflower seed • chick • enzyme
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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The increased production of new varieties with high oleic acid content for oil production and the increased trends in formulating high-energy-high-protein rations for broiler chickens has prompted evaluation of sunflower seed as a protein and energy source for use in poultry rations. A potential commercial source of oleic acid (18:1) is whole high oleic acid sunflower seed (HOASS), which contains about one-half oil, of which just over 80% is oleic acid. Recent information on the use of HOASS in poultry feeding is available (Rodriguez et al., 2005). Birds fed diets containing 100 and 200 g of HOASS/kg gained less weight at 12 and 42 d of age. Similarly, earlier studies using full-fat sunflower seeds and conventional sunflower meal have demonstrated that low levels of inclusion (100 g/kg) can be used as a protein source in chick starter diets (Ortiz et al., 1998; Rodriguez et al., 1998). However, inclusion at greater rates (20 to 30%) reduced the growth rate of the chicks (Daghir et al., 1980; Senkoylu and Dale, 2006). The only apparent disadvantages of this feed-stuff are that it contains a relatively low level of lysine and a high content of fiber. The fiber level appears to be the most problematic aspect concerning the use of sunflower meal at high levels in chick diets. It was reported that the soluble and insoluble constituents of nonstarch polysaccharides content are 4.5 and 23.1%, respectively (Düsterhöft et al., 1992). In fact, a strong negative correlation between the fiber fractions and the sunflower seed meal true metabolizable energy has been found (Janssen and Carre, 1985; Villamide and San Juan, 1998). Arija et al. (1998) also demonstrated that fat digestibility decreased when full-fat sunflower kernel was used, which could be because of the limited ability of the young chicks to extract oil from the intact cells in the digestion process. Fat, protein, and other nutrients could be entrapped to varying degrees inside the fibrous structure of the cell walls. Moreover, the presence per se of the fiber in the intestinal tract could impair the digestion process by the birds’ own proteases and other endogenous enzymes.
Efforts have been made to improve the availability of nutrients and ME content from different ingredients through enzyme supplementation. Because sunflower seeds contain substantial amounts of cell-wall material and a high concentration of fat that could affect the nutritive value of these seeds, the use of a crude enzyme preparation might be justified to improve the accessibility of cell contents to digestive enzymes. Addition of enzymes in poultry diets has increased considerably in recent years, but few reports are available on the effects of enzyme on sunflower meal utilization in poultry. However, several authors (Rebolé et al., 1999; Kocher et al., 2000; Attia et al., 2003) have indicated that commercial enzymes with different activities (cellulase, pectinase, xylanase, glucanase) did not result in significant improvements in growth performance of chicks, but some cases it was found beneficial in improving feed efficiency and AME values (Abbas et al., 1998; Mandal et al., 2005). It has been demonstrated that the physiological functions necessary for efficient fat digestion in young chickens are immature and continue to develop for several weeks after hatching (Jin et al., 1998). Noy and Sklan (1995) reported that in broiler chickens, secretion of lipase was low at hatching and increased 20-fold between 4 and 21 d of age. Krogdahl and Sell (1989) reported that dietary tallow and animal-vegetable fat were not efficiently used until lipase activity reached its maximum level. Because young birds have insufficient secretion of endogenous lipase, dietary supplementation of lipolytic enzymes may improve fat use. The objective of these 2 experiments was to study the influence of the addition of 2 enzymes (phospholipase and lipase) and a combination of these in broiler chicks fed 2 concentrations of HOASS (150 and 250 g/kg) on performance, digestibility of fat, protein, and amino acids, and digestive enzymes. Several blood parameters have also been analyzed to investigate possible changes in metabolism.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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A batch of HOASS (cv. Saxo), free of impurities, and a conventional sunflower meal (CSM) were obtained from a commercial supplier (SOS Cuetara, Andujar, Jaen, Spain) and used in experiments 1 and 2. Before mixing the experimental diets, the test products were ground to pass through a 3.0-mm screen. A sample of HOASS was analyzed for DM, ether extract, crude fiber, neutral detergent fiber, acid detergent fiber, and ash (Table 1
). The ME of HOASS and CSM were reported by Rodriguez et al. (2005) and FEDNA tables (FEDNA, 2003).
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Experiment 1
The objective of this experiment was to evaluate the effect of enzyme addition (lipase, phospholipase, and a combination of these at a concentration of 1 g/ kg each) on the nutritive value of HOASS at a concentration of 250 g/kg in chicken diets. For this purpose, 150 one-day-old male broiler chicks (Cobb-500 genetic line) were obtained from a commercial hatchery. The birds were housed in electrically heated starter battery brooders in an environmentally controlled room with 23 h of constant overhead fluorescent lighting. From hatching until 4 d of age, chicks were fed on a corn-soybean meal-based diet. On d 4, the chicks were allocated to 30 pens, each containing 5 birds, to receive 5 dietary treatments with 6 replicates of each treatment for the whole experimental period (4 to 21 d). For this experiment, the treatments were as follows (Table 2): 1) control, corn-soybean-based diet (CS); 2) CS + 250 g/ kg of HOASS; 3) CS + 250 g/kg of HOASS + 1 g/kg of lipase; 4) CS + 250 g/kg of HOASS + 1 g/kg of phospholipase; and 5) CS + 250 g/kg of HOASS + 1 g/kg of lipase + 1 g/kg of phospholipase.
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This experiment was conducted to evaluate the nutritive value of HOASS (150 g/kg), with and without enzymes (lipase, phospholipase, and a combination of these, at a concentration of 0.5 g/kg each) compared with a control diet containing CSM (150 g/kg). In this way, we designed diets containing similar concentrations of ether extract and crude fiber. Feeding and management were similar to experiment 1. On d 1, 150 one-day-old male broiler chicks (Cobb-500 genetic line) were allocated to 30 pens, each containing 5 birds, to receive 5 dietary treatments with 6 replicates of each treatment for the whole experimental period (0 to 21 d). For this experiment the treatments were as follows (Table 2
): 1) control diet (150 g/kg of CSM); 2) 150 g/kg of HOASS; 3) 150 g/kg of HOASS + 0.5 g/kg of lipase; 4) 150 g/kg of HOASS + 0.5 g/kg of phospholipase; 5) 150 g/kg of HOASS + 0.5 g/kg of lipase + 0.5 g/kg of phospholipase.
At the end of the experimental period, birds were weighed and feed consumption was recorded for feed conversion computation. Mortalities were recorded daily. All housing and handling were approved 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.
Collection of Samples and Measurements
At 21 d of age, 12 birds (2 per replicate) were randomly selected from each treatment after an overnight fast. Plasma and serum were prepared from blood obtained by cardiac puncture for subsequent determination of uric acid, calcium, glucose, cholesterol, and total protein in plasma and lactate dehydrogenase (LDH) and creatine phosphokinase (CPK) in serum. For plasma, the blood was collected in EDTA Vacutainer tubes on ice. The tubes were centrifuged at 2,500 x g for 15 min at 4°C to obtain plasma. For serum, the blood samples were allowed to clot in polypropylene tubes for 2 h at room temperature. The tubes were centrifuged at 1,500 x g for 10 min and the supernatant (serum) removed. All samples (plasma and serum) were stored at –20°C until assayed. For determination of fat digestibility, clean stainless steel collection trays were placed under each cage (6 per treatment) and excreta from each pen were collected for 72 h. A subsample of excreta was collected in polyethylene bags, weighed, and dried. Excreta were mixed thoroughly, frozen at –20°C, and freeze-dried. Before chemical analysis, these samples were ground (0.5-mm screen). Celite (Celite Corp., Lompoc, CA), a source of acid-insoluble ash (AIA), was added at 10 g/ kg to all diets as an indigestible marker. After killing the chicks by cervical dislocation (d 21), the liver, pancreas, spleen, and different sections of the small intestine and cecum were removed, cleaned from adhering tissue, and weighed or measured using 12 randomly selected chicks (2 per replicate) per treatment. To determine the relative length of the intestinal tract, the large loop between the gizzard and the end of the pancreas was considered as duodenum; the jejunum was defined as terminating at the yolk stalk and the ileum at the ileocecal junction. Abdominal fat was also removed, which was considered to be that fat surrounding the gizzard and intestine, extending within the ischium and surrounding the bursa of Fabricius. The ileum was quickly dissected out and the contents expressed by gentle manipulation into a plastic container for storage at –20°C. Digesta were pooled from 2 birds of each replicate within the same treatment. The ileal contents were freeze-dried and ground (1-mm screen) and subsequently analyzed for Kjeldahl N, amino acids, and Celite. The pancreas was removed from each bird (6 chickens per treatment) and placed in vials that were immediately plunged into a flask containing liquid nitrogen. Thereafter, pancreas samples were transferred to a freezer at –20°C until required for enzyme activity measurements. The pH of cecal digesta (12 chicks per treatment) was directly determined (Crison Micro pH 2000, Crison Instruments S.A., Alella, Spain).
Chemical Analysis
Dry matter (method 930.15), CP (method 976.05), crude fiber (method 978.10), and ash (method 942.05) were analyzed according to the methods of the Association of Official Analytical Chemists International (AOAC, 1995). Neutral and acid detergent fibers and acid detergent lignin were determined following the sequential analysis outlined by Robertson and Van Soest (1981). Ether extract was determined by extraction in petroleum ether following acidification with 4 N HCl solution (Wiseman et al., 1992). The AIA contents of diets and excreta were measured after ashing the sample and treating the ash with boiling 4 M HCl (Siriwan et al., 1993). Amino acids in the diets and the ileal contents were analyzed following AOAC (1995) procedures and separated using a Beckman Model 6300 autoanalyzer (Beckman-Coulter S.A., Madrid, Spain). Three replicates of all analyses were performed. Determination of the amino acid tryptophan was not possible under the conditions of analysis used.
Blood plasma was analyzed for uric acid (uricase-PAP enzymatic colorimetric), calcium (o-creosolphtalein colorimetric method), cholesterol (cholesterol oxidase-peroxidase enzymatic/colorimetric method), glucose (glucose oxidase-peroxidase enzymatic/colorimetric method), and total protein (colorimetric Biuret method) following the methods described by SpinReact S.A. (St. Esteve de Bas, Girona, Spain). Lactate dehydrogenase (International Federation for Clinical Chemistry-kinetic-ultraviolet) and CPK (N-acetylcysteine-kinetic-ultraviolet) were determined in serum according to SpinReact method. For the measurement of enzyme activities, 400 mg of pancreas was quickly weighed into test tubes kept on ice and 6 mL of ice-cold physiological saline (9 g of sodium chloride/L) was added and centrifuged at 2,000 x g. Portions of supernatant fractions containing enzymes were assayed for 
-amylase and lipase activities as described previously by Longstaff and McNab (1991). The method for -amylase activity was based on that of Lever (1972) for the detection of glucose released from pregelatinized maize starch used as substrate. Lipase activity was measured using a diagnostic kit (kit 800-B, Sigma-Aldrich Química S.A., Tres Cantos, Madrid, Spain).
Calculations and Statistical Analysis
Apparent ileal CP, amino acid (AA), and crude fat (CF) digestibilities were calculated using the following formula: 100% – [100% x (AIA concentration in feedA-IA concentration in ileal digesta or excreta) x (CP and AA concentration in ileal digesta and CF concentration in excreta/CP and AA concentration in feed)]. Data were subjected to ANOVA using the GLM procedure (SAS Institute, 2003), with treatment comparisons among the control and HOASS diets with and without enzymes. Treatment 1 was considered as a positive control. Significant differences among treatment means were determined at P < 0.05 by using Duncan’s multiple range test.
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RESULTS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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The effects of different enzymes on the nutritional value of high oleic acid sunflower seed diets on performance are summarized in Table 3. Weight gain and feed consumption were reduced by 24% (P < 0.001) and 14% (P < 0.05), respectively, and feed conversion was improved by 12% (P < 0.001) in birds fed the un-supplemented HOASS diet compared with those fed the control diet. The addition of enzymes to birds fed the HOASS diet increased (P < 0.001) weight gain and feed consumption by 28 and 25%, respectively, and improved feed conversion by 3% (P < 0.01) compared with those fed the unsupplemented HOASS diet. On a numerical basis, the greatest response in performance was obtained with the combination of lipase plus phospholipase.
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Results of the relative organ weights and intestinal length are shown in Table 4. The inclusion of unsupplemented HOASS in the chicken diet reduced relative liver weight by 3% (P < 0.01) and increased (P < 0.001) relative duodenum, jejunum, ileum, and ceca lengths by 11, 12, 13, and 8%, respectively, compared with those fed the control diet. The inclusion of enzymes in the HOASS diet increased relative pancreas (4%; P < 0.05) and relative liver weight (9%; P < 0.001), and reduced (P < 0.001) relative spleen weight (13%), and relative duodenum (13%), jejunum (8%), ileum (4%), and ceca (5%) lengths compared with the unsupplemented HOASS diet.
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The effects of enzyme supplementation in diets containing HOASS on apparent fat digestibility and pancreatic -amylase and lipase activities are summarized in Table 5. The inclusion of unsupplemented HOASS in the diet reduced (P < 0.001) fat digestibility by 7% and amylase and lipase activities by 22 and 19%, respectively, compared with those fed the control diet. The inclusion of enzymes in the HOASS diet increased (P < 0.001) fat digestibility and amylase and lipase activities by 5, 53, and 58%, respectively, compared with the unsupplemented HOASS diet. The greatest response in fat digestibility and digestive enzyme activities was obtained with the combination of lipase plus phospholipase.
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The effects of enzyme supplementation in diets containing HOASS on blood parameters are presented in Table 6. The inclusion of unsupplemented HOASS in the diet increased plasma uric acid (P < 0.01), cholesterol (P < 0.001), and glucose (P < 0.001) concentrations by 5, 20, and 15%, respectively, and reduced (P < 0.001) serum LDH and CPK concentrations by 6 and 16%, respectively, compared with those fed the control diet. The inclusion of enzyme in the HOASS diet increased plasma uric acid (P < 0.01), calcium (P < 0.001), serum LDH (P < 0.001), serum CPK (P < 0.001), and total protein (P < 0.001) concentrations by 6, 12, 7, 52, and 15%, respectively. Plasma cholesterol and glucose were reduced (P < 0.001) by 8 and 5%, respectively, by enzyme inclusion.
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The effects of different enzymes on the nutritional value of high oleic acid sunflower seed diets on performance are summarized in Table 3. Performance of the birds was not affected by the unsupplemented HOASS diet compared with those fed the control diet. The addition of enzymes to birds fed the HOASS diet increased weight gain and feed consumption by 11% (P < 0.001) and 6% (P < 0.01), respectively, and improved feed conversion by 5% (P < 0.001) compared with those fed the unsupplemented HOASS diet.
Digestive Organ Size
Results of the relative organ weights and intestinal length are shown in Table 7. The inclusion of unsupplemented HOASS in the diet reduced relative pancreas and abdominal fat weights by 6% (P < 0.001) and 46% (P < 0.001), respectively, and relative duodenum and ceca lengths by 6% (P < 0.01) and 9% (P < 0.01), respectively, compared with those fed the control diet. The inclusion of enzyme in HOASS diet increased (P < 0.001) relative pancreas weight by 5% compared with the unsupplemented HOASS diet.
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The effects of different enzymes on the nutritional value of high oleic acid sunflower seed diets in broilers on apparent digestibility of fat, protein, and essential and nonessential amino acids are reported in Tables 8 and 9. The inclusion of unsupplemented HOASS in the diet increased CF and CP (P < 0.001) digestibilities by 3% and essential (P < 0.05; Arg, Leu, Lys, Phe, Thr, Val, and His) and nonessential (P < 0.05; Asp, Glu, Gly, Pro, Tyr, and Cys) digestibilities in a range of 1 to 3% and 2 to 13%, respectively, compared with those fed the control diet. However, Ser digestibility was reduced (P < 0.001) by 2% in the HOASS diet compared with the control diet. The inclusion of enzyme in the HOASS diet significantly reduced CP (P < 0.01) digestibility by 1% and essential (P < 0.05; Arg, Ile, Leu, Lys, Met, Thr, Val, and His) and nonessential (P < 0.05; Ala, Asp, Glu, Gly, Pro, Ser, Tyr, and Cys) digestibilities in a range of 1 to 3% and 2 to 7%, respectively, compared with those fed the unsupplemented HOASS diet.
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DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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In the first study, broilers fed diets containing HOASS at 250 g/kg had a slower rate of growth compared with those fed the control diet. These results are in accordance with those obtained by Daghir et al. (1980), Arija et al. (1998), and Rodriguez et al. (2005) using full-fat, dehulled full-fat, and high-oleic sun-flower seeds, respectively. In contrast with these findings, Elzubeir and Ibrahim (1991), Cheva-Isarakul and Tangtaweewipat (1991), and Rodriguez et al. (1998) reported that this oil seed could be included at a relatively high level without any adverse effect on performance and nutrient use. One explanation for this discrepancy in results is that different sunflower varieties or cultivars varying in chemical composition were used in the experiments. In the second experiment, there were no differences in the performance of the birds, probably because the control diet (conventional sunflower seed) and the HOASS diet were similar in ether extract and fiber contents, and with the same concentration of sunflower seed (150 g/kg).
The growth depression observed in the first experiment may be explained by the reduction in feed consumption. This effect could be because of the greater crude fiber and fat contents (6.2 and 13.1%, respectively) in the HOASS diet compared with the control diet (3.8 and 8.7%, respectively). A substantial amount of cell-wall material in the kernel and the hull (Düsterhöft et al., 1992) could cause limited accessibility of chick digestive enzymes to the seed fat because of the cell-wall barrier. These findings suggested that the indigestible material in the digesta was acting not only as an inert diluent, but was also playing a more specific role, probably by retaining water into which water-soluble nutrients were trapped, thus making them unavailable for absorption by the intestine and hence, excreted (Arnal-Peyrot and Adrian, 1974). Similarly, the bulking effect and dietary nutrient dilution can reduce the ME concentration (Senkoylu and Dale, 1999), reduce the passage rate, and slow gastric emptying (Hetland et al., 2004) as well as cause alterations in the jejunal mucosa (Arija et al., 1998, 2000) impairing the normal absorption of nutrients. Similarly, the excess of fat in the HOASS diet could cause inhibition of gastric emptying and gastrointestinal motility. These effects appear to be responsible, at least in part, for the inhibition of appetite and energy intake (Xu et al., 2005). Furthermore, the presence of a phenolic compound, chlorogenic acid, in amounts of 10 to 40 g/kg (Dorrell, 1976) in the kernel, could explain the negative effect on the performance. However, Eklund (1975) in rats and Treviño et al. (1998) in chickens did not detect any effect of chlorogenic acid on the nutritive value of the diet. Finally, Luckett et al. (1999) isolated from sunflower seeds a peptide with 14 amino acids, termed sunflower trypsin inhibitor (SFTI-1), which could also have a negative effect on the performance.
The first experiment also demonstrated that the HOASS diet had an effect on the digestive tract of the birds, increasing the size of the small intestine and cecum. Similar data were obtained by Arija et al. (1998) using full-fat sunflower kernels and by Zatari and Sell (1990) using sunflower seeds. It could suggest that the presence, in the sunflower seeds, of a substantial amount of cell-wall material consisting of predominantly cellulose, pectic polysaccharides, and 4-0-methyl-glucuroxylans with small amounts of glucomannans and fucoxyloglucans (Düsterhöft et al., 1992) could exert a trophic effect on the intestinal mucosa, increasing the intestinal size. Histological alterations in the jejunal mucosa have been observed by Arija et al. (2000), who showed shortening and thickening of the villi and hypertrophy and hyperplasia of goblet cells caused by the inclusion of full-fat sunflower kernels in chicken diets. Alternatively, the excessive bulk of the digesta containing this indigestible cell-wall material may have been one factor in delaying crop emptying because chicks fed the HOASS diet consumed less than the birds fed the control diet. In the second experiment, a decrease in the size of the small intestine (particularly the duodenum) and cecum was observed in the unsupplemented HOASS diet compared with the conventional sunflower diet, due probably to an increase in fat and CP digestibilities of the diet.
Regarding the relative organ sizes, liver weight expressed as a percentage of BW was not affected or was only slightly affected in birds fed the unsupplemented HOASS diet compared with the control diet (corn-soybean) and CSM diet, respectively. These results are similar to those reported by Daghir et al. (1980) and Arija et al. (1998) using diets containing full-fat sunflower seed. Abdominal fat deposition was significantly lower in birds fed the HOASS diet (containing 1.8% high oleic acid sunflower oil) compared with those receiving the conventional sunflower diet (containing 9% conventional sunflower oil). This effect may be attributed to the fatty acid composition or other components of the diets. Because the experimental diets in the current study were isoenergetic and had similar contents of total fat and crude fiber, the abdominal fat reduction could be related to an effect of dietary fat on fat deposition, particularly to the greater composition of monounsaturated fatty acid (oleic acid content) in the HOASS diet. Recently, Soriguer et al. (2003) confirmed that when the oleic acid content of rat adipocytes increased, the lipolytic activity of the adipocytes increased and the antilipolytic capacity of the insulin decreased. Previous reports indicate that broiler chickens fed diets enriched with polyunsaturated fatty acids have less abdominal fat (Sanz et al., 1999) or total body fat (Sanz et al., 2000a,b) deposition than those fed diets containing saturated fatty acids.
In the first study, fat digestibility showed a consistently lesser value in those birds fed the unsupplemented HOASS diet. These results are in agreement with those reported by Hill and Renner (1963), Arija et al. (1998), and Rodriguez et al. (2005) using heated, ground, unextracted full-fat soybean seed, full-fat sunflower kernel, and HOASS, respectively. They attributed this effect to the limited ability of the chicks in the digestion process to extract oil from the intact cells. The reduction in the amylase and lipase pancreatic activities by the birds fed the HOASS diet in our experiment could be due to the adsorption of lipase and bile salts to the fiber present in the seed. Schneeman (1978) reported that the availability of enzymes such as lipase could be limited by their absorption into fibers such as xylan, cellulose, wheat bran, and rice bran. Evidence of this effect has also been observed in vitro by Lairon et al. (1985) with wheat bran. Almirall et al. (1995) reported the specific lipase activity to be decreased in broiler chickens after feeding barley. In fact, Arija et al. (1998) showed a consistently greater activity of lipase in birds fed increasing concentrations of dehulled full-fat sunflower seeds in the diets, probably because of the reduction of fiber content in their seeds.
In relation to the blood parameters, the results showed that the addition of HOASS to the diets modified uric acid, cholesterol, and glucose concentrations and serum LDH and CPK activities. These results could be explained by the low energetic utilization of the HOASS diet. This metabolic state could provoke activation of gluconeogenesis resulting in an increase of glucose and cholesterol and a reduction in LDH and CPK in serum.
Protein and apparent digestibility of essential and nonessential amino acids were increased, except for serine, which was reduced, in the birds fed the unsupplemented HOASS diet compared with those fed conventional sunflower seed. This positive effect might be attributable to a difference in amino acid digestibility caused by the processing of both types of seeds. Conventional sunflower seed was solvent-extracted and the HOASS was not subjected to any treatment. Rad and Keshavarz (1976) and Zhang and Parsons (1994) reported a decrease in amino acid digestibility when the temperature or time processing for the same sunflower seed sample increased. According to Ravindran and Blair (1992) and San Juan and Villamide (2000), high temperature associated with mechanical pressing damages the protein, destroys amino acids, and decreases their availability.
In the current study, the addition of enzymes (lipase and phospholipase) improved the performance of the birds. This improvement was associated with a significant increase of fat digestibility and -amylase and lipase activities. Although the factor(s) affected by the enzyme treatment were not identified, it is conceivable that lipase and phospholipase may have removed the fatty acid attached to the 2-position of phosphoglycerides present in the HOASS diet. These enzymes could also assist to catalyze the conversion of lecithin to lysolecithin. Lysolecithin is a very potent natural emulsifier and biosurfactant, which could increase fat digestion and the absorption of other nutrients by the formation of micelle structures in the gastrointestinal tract (Schwarzer and Adams, 1996).
Similarly, although the role of lipase in the regulation of upper gastrointestinal function is poorly understood, we hypothesize that the presence of sunflower trypsin inhibitor in HOASS together with the lipolytic enzymes added could cause a positive interaction with pancreatic enzymes. In this sense, Pap and Varró (1984) and Thiruvengadam and DiMagno (1988) have demonstrated, in human duodenal juice, an increase of lipase activity in the presence of trypsin or chymotrypsin inhibitor. On the other hand, Al-Marzooqi and Leeson (1999) reported that supplemental lipase in the diet was effective in increasing animal fat digestibility, but these effects were associated with worsening performance. Conversely, a lack of response by the addition of lipases on chick performance and fat digestibility was reported by Meng et al. (2004) evaluating beef tallow and canola oil using a wheat-based diet.
The current study also showed that the addition of enzymes to HOASS diet was associated with a reduction in the length of the gastrointestinal tract. A similar result has been obtained by the addition of enzymes to barley and lupin diets (Viveros et al., 1994; Brenes et al., 2002, respectively). This effect is presumably the result of an adaptive response to increased nutrient digestibility and availability in the different sections of the digestive system, reducing the cost of digestion that was expressed as a shorter gut. Likewise, the increase of pancreatic enzymes by the addition of exogenous enzymes is similar to the results reported by Almirall et al. (1995) and Jensen et al. (1996) who observed in chickens and pigs, respectively, an increase in digesta and pancreatic enzymes after adding exogenous enzymes in barley diets.
Finally, the addition of enzymes reduced essential and nonessential amino acid digestibility in the birds compared with those fed the unsupplemented HOASS diet. We have no explanation to justify these results. The increase in blood uric acid observed in the current experiment by the enzyme addition may suggest an amino acid imbalance that could justify the decrease in amino acid digestibility. However, this negative effect is not related to the greater growth rate found in the birds fed enzymes. Further research must be conducted in oil seeds to elucidate possible interactions among exogenous and pancreatic enzymes and amino acids in the upper intestinal tract.
In conclusion, the addition of HOASS in chicken diets caused a negative effect on performance, fat digestibility, -amylase and lipase activities, and intestinal organ sizes, and modified some blood parameters only at the higher level of inclusion (250 g/kg). These negative effects were counteracted by the enzyme addition. These data demonstrated that the inclusion of HOASS and enzymes (mainly phospholipase or combination of lipase and phospholipase) in a chick diet resulted in performance similar to those obtained with a control diet.
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ACKNOWLEDGMENTS |
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Received for publication March 27, 2008. Accepted for publication June 29, 2008.
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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