Tohru Motoki, Hiroshi Kamisoyama, Kazuhisa Honda and Shin
Transcription
Tohru Motoki, Hiroshi Kamisoyama, Kazuhisa Honda and Shin
http://www.jstage.jst.go.jp/browse/jpsa doi:+*.,+.+/jpsa.*++*-1 Copyright ῍ ,*++, Japan Poultry Science Association. Tohru Motoki, Hiroshi Kamisoyama, Kazuhisa Honda and Shin Hasegawa Graduate School of Agricultural Science, Kobe University, Kobe 0/1ῌ2/*+, Japan Potassium diformate (KDF), a chemical complex of formic acid and potassium formate, improves growth performance in pigs. The aim of the present study was to examine the e#ects of dietary KDF on the growth, nitrogen retention, intestinal pH, counts of Enterococcus faecalis, coliforms, and lactic acid bacteria in the cecum, and humoral immune response of growing broiler chickens. Twenty four male broiler chicks were randomly assigned into three groups (eight birds in each group). Each group was fed an antibiotics free commercial diet (as a control diet, ,-ῌ CP, -,*** kcal of ME/kg), a control diet containing KDF at +ῌ, or a control diet containing antibiotics (/* g titer/t Salinomycin, /* g titer/t Avilamycin) until ,2 days of age. Dietary KDF significantly increased the weights of body, breast muscle, thighs and wings, whereas the weights of liver and abdominal fat were not a#ected. These findings suggest that the increase of body weight by dietary KDF might be due to the increased muscle weight. Dietary KDF did not a#ect nitrogen retention. Dietary KDF did not a#ect the intestinal pH, and the counts of Enterococcus faecalis, coliforms, and lactic acid bacteria in the cecum. Hemagglutination titer was not a#ected by dietary KDF. Thus, although the mechanisms of the growth promotion by dietary KDF are not clear, our findings suggest that the KDF might promote growth of broiler chickens at least in the early phase of growth. Key words: broiler, growth, muscle, organic acid, potassium diformate J. Poult. Sci., .2: ,.1ῌ,/-, ,*++ ῎῎῎῎῎῎῎῎῎῎῎῎῎῎῎῎῎῎῎῎῎῎῎ Introduction In-feed antibiotics have been utilized for over half a century to promote growth in domestic animals (Moore et al., +3.0; Leitner et al., ,**+; Sun et al., ,**/). The mechanisms underlying the growth promotion by in-feed antibiotics are presumed the improvement of nutrients availability by reducing microbial population and the prevention of pathogen infections by improving immune response (Dibner and Buttin, ,**,). Meanwhile, the extensive use of antibiotics in livestock production has increased the risk of development of resistance in human and animal pathogens (Bates et al., +33.; Witte, +332). Thus, the use of antibiotics to promote growth in domestic animals has been banned in European Union. However, the withdrawal of in-feed antibiotics suppresses growth in domestic animals, which in turn results in the increase of the production costs (Casewell et al., ,**-; Sun et al., ,**/). Therefore, there is a pressing need to develop alternative in-feed antibiotics that may be used to alleviate the problems associated with the withdrawal of antibiotics from feed. Received: April ,1, ,*++, Accepted: July +-, ,*++ Released Online Advance Publication: August ,/, ,*++ Correspondence: Dr. S Hasegawa, Graduate School of Agricultural Science, Kobe University, Kobe 0/1ῌ2/*+, Japan. (E-mail: [email protected]) Organic acids promote growth in domestic animals. Like in-feed antibiotics, organic acids reduce intestinal microbial population and improve immune status (Dibner and Buttin, ,**,). In addition, a reduction in intestinal pH, an additional e#ect which is not induced by antibiotics, is regarded as one of the positive e#ects by organic acids responsible for the growth promotion in domestic animals. The use of organic acids creates an acidic environment in the gut that inhibits the growth of Escherichia coli, Salmonella, Campylobacter and gram-negative bacteria (Dibner and Buttin, ,**,; Gunal et al., ,**0). Potassium diformate (KDF, a chemical complex of formic acid and potassium formate, Fig. +) promotes growth (Paulicks et al., +330; Roth et al., +330; Kirchgessner et al., +331), increases lean percentage in the ham, flank, loin and neck and shoulder (ῌverland et al., ,***), and decreases the number of intestinal microflora including coliforms, lactic acid bacteria or yeast in digesta from various segments of the gastrointestinal tract (Canibe et al., ,**+) in pigs. Canibe et al. (,**+) described that the presence of lactic acid bacteria in the gastrointestinal tract of pigs is believed to be beneficial for the animal, and that it is di$cult to conclude firmly whether yeasts are detrimental or beneficial for the animal. However, they proposed that the growth-promoting e#ect of KDF in pigs might be explained by reducing microbial population in the gastrointestinal tract in pigs, because the reduction of the mi- Journal of Poultry Science, .2 (.) 248 crobial population would be desirable for the dietary energy utilization by the host animal. KDF is easier to handle than organic acids, because the use of organic acids has been limited by problems of handling, strong odor, or corrosion during feed processing and during its use on the farm. Thus, KDF has been approved in European Union and Japan for use in pigs with the claim for promoted growth performance. However, there is little information about the e#ect of KDF on growth performance in chickens, although studies with chickens have indicated positive e#ects of dietary organic acids including propionic acid, citric acid, ascorbic acid and butyric acid on growth performance (Izat et al., +33*; Afsharmanesh and Pourreza, ,**/; Leeson et al., ,**/; Gunal et al., ,**0; Chowdhury et al., ,**3). In this study, we examined the e#ects of dietary KDF on the growth, nitrogen retention, intestinal pH, counts of Enterococcus faecalis, coliforms, and lactic acid bacteria in the cecum, and humoral immune response of growing broiler chickens. We used ,2-day old broiler chickens, because correlations among ,2-day body weight, .,-day body weight and .1-day carcass weight were positive and high in broiler chickens (Wang et al., +33+). Fig. +. Structure of potassium diformate. Table +. Animals and Feed Day-old male broiler chicks (chunky) were purchased from a local hatchery (Ishii Co. Ltd., Tokushima, Japan). They were given free access to water and a commercial chicken starter diet (Nosan Co., Kanagawa, Japan) and acclimated to the facility for five days before feeding of experimental diets (Table +). After five days of acclimatization, the chicks were fed experimental diets. All experimental procedures followed the guidelines for the care and use of experimental animals at the Rokkodai Campus of Kobe University in Japan. Preliminary Experiment Initially, we carried out a screening experiment to examine the e#ect of dietary KDF (dose: +ῌ-ῌ) on growth in growing male and female broiler chickens. We found that the addition of +ῌ KDF in feed significantly increased body weight in male chickens. Similar trends were found in female chickens but the e#ect was not significant. In contrast, the addition of -ῌ KDF in feed significantly decreased body weight in male and female chickens. Based on these results, the experimental diet containing +ῌ KDF and growing male broiler chicks were used in the following experiments (Experiments + and ,). Assignment of Birds Forty eight male broiler chicks were randomly assigned into six groups (eight birds in each group). Three groups were used to examine the e#ects of dietary KDF on the growth, nitrogen retention, intestinal pH, counts of Enterococcus faecalis, coliforms, and lactic acid bacteria in the cecum in growing broiler chickens (Experiment +). Composition of experimental diets Feed ingredients (g/+** g) Yellow corn Soybean meal Corngluten meal Rapeseed meal Fish meal Soybean oil Calcium carbonate Calcium phosphate tribasic Sodium chloride Vitamin mixture+ῌ Mineral mixture,ῌ Potassium diformate Cellulose Calculated analysis Crude protein (ῌ) Metabolizable energy (kcal/kg) +ῌ Materials and Methods Control +ῌ ,ῌ -ῌ /.4. ,/4* /4* .4* /4* +4* +4. *41 *4, *4,/ *4*/ *4* -4* /.4. ,/4* /4* .4* /4* +4* +4. *41 *4, *4,/ *4*/ +4* ,4* /.4. ,/4* /4* .4* /4* +4* +4. *41 *4, *4,/ *4*/ ,4* +4* /.4. ,/4* /4* .4* /4* +4* +4. *41 *4, *4,/ *4*/ -4* *4* ,-4* -*** ,-4* -*** ,-4* -*** ,-4* -*** Vitamin A (,**,*** IU/g) +* g, Vitamin D (-*.*** IU/g) 1 g, Thiamine HCl +.0 g, Riboflavin 2 g, Prydoxine +.0 g, Choline HCl 30 g, Nicotinic acid +.0 g, Calcium panthothenate -., g, Folic acid 2 g per kg. ,ῌ Manganese 2* g, Zinc /* g, Iron / g, Iodine + g, Copper *.0 g per kg. Motoki et al.: Potassium Diformate Improves Growth in Chickens Another three groups were used to examine the e#ect of dietary KDF on the humoral immune response in growing broiler chickens (Experiment ,). Experiment + Birds (n῏2 in each group) were fed an antibiotics free commercial diet (as a control diet), a control diet containing KDF at +ῌ, or a control diet containing antibiotics (/* g titer/ton Salinomycin, /* g titer/ton Avilamycin) until ,2 days of age. Birds were fed the experimental diets containing an indigestible marker (*./ῌ chromium oxide) for the last 0-days of the experimental period. For the last --days of the experimental period, each pen was divided into three zones using stainless steel to create three replicates for each group, and feces were collected, weighed, and stored at ῍,*ῑ every day for nitrogen retention analysis. At the end of the experimental period, chickens were sacrificed by decapitation. The right cecum was immediately removed, and the digesta was used for microbial analysis. The digesta for microbial analysis were transferred into plastic bags, and brought to a separate place for microbial analysis. The liver, abdominal fat, breast muscle, thighs, and wings were excised and weighed. The digesta samples from the gizzard, duodenum (*./ cm caudally from gizzard to major duodenal papilla), jejunum (major duodenal papilla to Meckel’s diverticulum), ileum (Meckel’s diverticulum to ileocecal junction), and left cecum were collected, and the pH was measured after adding +3 volumes of distilled water and stirring for +* min. Nitrogen retention analysis was conducted as follows. Nitrogen contents in feces and experimental diets were measured by the standard methods (AOAC, +33/). For the determination of chromium concentration, feeds and feces were ashed by a wet-ash digestion with sodium molybdate, sulfuric acid and nitric acid. Chromium concentration was determined at a wavelength of ..* nm (Bolin, +3/,). Microbial analysis was conducted as follows. All sam- 249 pling was completed within the first two hours after sacrifice. Enterococcus faecalis were enumerated on the EF Agar Base (Nissui Pharmaceutical Co. LTD. Tokyo, Japan) after aerobic incubation at -1ῑ for .2h. Coliform bacteria were enumerated on MacConkey Agar (Nissui Pharmaceutical Co. LTD. Tokyo, Japan) after aerobic incubation at -1ῑ for ,. h. Lactic acid bacteria were enumerated on the Plate Count Agar with BCP (Nissui Pharmaceutical Co. LTD. Tokyo, Japan) after anaerobic incubation at -1ῑ for 1, h. Experiment , Birds (n῏2 in each group) were fed the experimental diets as described in Experiment +. Birds were inoculated intravenously with a /ῌ sheep red blood cell (SRBC) suspension (+ ml/kg body weight) at +2 days of age. Blood samples were collected from a wing vein on .ῌ3 days after inoculation, and sera were separated and inactivated for -* min at /0ῑ for direct hemagglutination analysis. The hemagglutination analysis was carried out in U-shaped wells of disposable microplates. Sera were serially diluted two-fold in a PBS solution. The +ῌ SRBC suspension was added to the wells in *.*,/ ml volume, and the microplates were incubated at -1ῑ for 0* min. The observed agglutination was recorded by a negativethrough--ῌ system, and the highest dilution with ,ῌ was taken as endpoint. Results are expressed in the reciprocal of that dilution. Statistical Analysis All data were analyzed using analysis of variance (ANOVA), and significant di#erences among groups were analyzed by the Tukey-Kramer test. All statistical analyses were performed using the commercial package (StatView version /, SAS Institute, Cary, NC, USA, +332). Results Dietary KDF and antibiotics significantly (Pῐ*.*/) increased body weight, whereas the weights of the liver and abdominal fat were not a#ected (Table ,). The E#ects of potassium diformate on weights of body, liver, abdominal fat, breast muscle, thighs and wings in broiler chickens Table ,. Body weight (g) Liver weight (g) Liver weight (g/+** g BW) Abdominal fat weight (g) Abdominal fat weight (g/+** g BW) Breast muscle weight (g) Breast muscle weight (g/+** g BW) Thighs weight (g) Thighs weight (g/+** g BW) Wings weight (g) Wings weight (g/+** g BW) Control Potassium diformate Antiboitics +*2242῎,,41 ,+4-2῎ *411 +431῎ *4*3 +/4**῎ *42, +4-2῎ *4*1 +/.43῎ 14, +.4,῎ *40 ,,.4*῎ 04, ,*40῎ *4. 0+42῎ ,42 /41*῎ *4-* ++3+4-῎-+43* ,,4.*῎ +4,, +422῎ *4+* +14-.῎ +4++4.0῎ *4+* +3-4*῎ 143* +04,῎ *4/* ,/-43῎ 04** ,+4-῎ *4-* 0342῎ .4,* /420῎ *4-.* +,*+4-῎-34-* ,/4*.῎ +4.. ,4+*῎ *4+. +/40,῎ +4-* +4-*῎ *4+* +2*4+῎ 04/* +/4*῎ *4/* ,//4/῎ /41* ,+4.῎ *42* 1,4+῎ ,4,* 04*.῎ *4,/* Values are means῎SEM for eight chickens per group (n῏2). * Significant with respect to control group (Pῐ*.*/). 250 Journal of Poultry Science, .2 (.) E#ects of dietary potassium diformate and antibiotics on nitrogen retention in broiler chickens. Values are meansῌSEM for three replicates per group. * Significant with respect to control group (P῍*.*/). Fig. ,. weights of breast muscle, thighs and wings were significantly (Pῌ*.*/) increased in the KDF and the antibiotics groups, and the weights of breast muscle, thighs, and wings per +**g body weight were also significantly (Pῌ *.*/) increased in the KDF and the antibiotics groups (Table ,). Nitrogen retention was not a#ected by dietary KDF but significantly (Pῌ*.*/) increased by dietary antibiotics (Fig. ,). Dietary KDF did not a#ect pH of the gastrointestinal contents (Table -). There were no significant di#erences in the counts of Enterococcus faecalis, coliform bacteria and lactic acid bacteria in the cecal content among groups (Fig. -). There were no significant di#erences in the hemagglutination titer among groups Fig. -. E#ects of dietary potassium diformate and antibiotics on microflora in the cecum of broiler chickens. Values are meansῌSEM for eight chickens per group. (Fig. .). Discussion ῌverland et al. (,***) reported that the addition of +.,ῌ KDF to a diet improved growth performance of growing-finishing pigs. Paulicks et al. (+330) reported that the addition of ,ῌ KDF to a diet for growing pigs Motoki et al.: Potassium Diformate Improves Growth in Chickens 251 E#ects of dietary potassium diformate on pH at the di#erent part of the intestine in broiler chickens Table -. Gizzard Duodenum Jejunum Ileum Cecum Control Potassium diformate Antibiotics -4,3ῌ*4*3 04-+ῌ*4*0 /432ῌ*4+* 14+2ῌ*4,. 04.-ῌ*4,/ -4-,ῌ*4+, 04,1ῌ*4*, /422ῌ*4*14+*ῌ*4+/ /412ῌ*4+, -4-1ῌ*4*1 04,-ῌ*4*/423ῌ*4*0 1401ῌ*4*3 /410ῌ*4*0 Values are meansῌSEM for eight chickens per group. E#ects of dietary potassium diformate and antibiotics on hemagglutination titer in broiler chickens. Values are meansῌ SEM for eight chickens per group. Fig. .. increased body weight gain but ,.. and ,.2ῌ KDF did not a#ect body weight gain. Based on these reports, we tested a wider range of doses (from * to -ῌ) in the preliminary experiment and found that the body weight of male broiler chickens was significantly increased only in the +ῌ KDF group. Recently, Mikkelsen et al. (,**3) reported that *../ῌ dietary KDF did not a#ect weight gain in broiler chicks. This evidence and our results suggest that optimum KDF supplementation in broiler chicken feed might be +ῌ. Cumulative feed intake of the KDF group was numerically higher than that of the control group (calculated values were +,/*0.1 g/bird and +,.*1./ g/bird, respectively). However, these values can not be statistically analyzed, because we did not measure individual feed intake. In order to investigate whether the KDF-increased feed intake in broiler chickens is involved in its growth-promoting e#ect, further studies will be needed to analyze the relation ship between feed intake and body weight gain individually. The weights of breast muscle, thighs and wings were increased in the KDF group, whereas dietary KDF did not a#ect the weights of abdominal fat. The abdominal fat is known to be a good indicator of fat deposition in chicks (Shigeno, +31-; Havenstein et al., ,**-). These findings suggest that the increase of muscle weight might be involved in the increase of body weight by KDF in chickens. This observation raises the question whether the increase of muscle weights by KDF in chickens is due to the improvement of nitrogen availability. However, nitrogen retention was not a#ected by KDF. Also in pigs, KDF did not promote the ileal digestion of dietary amino acids (except for phenylalanine), or nitrogen retention (Mroz et al., ,**,). These findings and our results suggest that KDF might not a#ect nitrogen retention in both chickens and pigs. In the present study, +ῌ KDF did not a#ect the intestinal pH of broiler chickens. Mikkelsen et al. (,**3) reported that *../ῌ dietary KDF did not a#ect intestinal pH in broiler chickens. It is generally accepted that dietary organic acids decrease intestinal pH, which in turn, results in the decrease of intestinal microbial populations. However, it seems likely that dietary KDF, at least in this experimental procedure, might not decrease intestinal pH in broiler chickens. There is evidence that dietary organic acids activate the immune system and improve growth performance in broil- Journal of Poultry Science, .2 (.) 252 er chickens. For example, dietary acetic acid, citric acid and lactic acid increased serum globulin level in broiler chicks (Abdel-Fattah et al., ,**2). Chowdhury et al. (,**3) reported a higher density of lymphocytes in the cecal tonsils and ileum in citric acid-fed broiler chickens. These findings raise the possibility that dietary organic acids improve both humoral and cellular immunity. In the present study, we examined the e#ect of KDF on hemagglutination titer, which indicates the humoral immune response to SRBC. However, dietary KDF did not a#ect the hemagglutination titer in broiler chickens, suggesting that humoral immune response might not be a#ected by KDF. Further studies will be needed to examine the e#ects of KDF on cellular immune responses in broiler chickens. In summary, in growing broiler chickens, the dietary KDF significantly increased body weight and muscle weight. Dietary KDF did not a#ect nitrogen retention, intestinal pH, the counts of Enterococcus faecalis, coliforms and lactic acid bacteria in the cecum, and hemagglutination titer. It seems likely that the growth-promotion by KDF might not be caused by the e#ects on intestinal microflora, nitrogen availability and humoral immunity. Our findings suggest that the KDF might promote growth of broiler chickens at least in the early phase of growth. Further studies will be needed to examine the long term (e.g. .3 days) e#ect of dietary KDF on the growth of broiler chickens. Acknowledgments This work was supported by a Grant-in-Aid (Number ,+/2*-.2) for Scientific Research (C) from the Ministry of Education, Culture, Sports, Science, and Technology of Japan. References Abdel-Fattah SA, EI-Sanhoury MH, EI-Mednay NM and Abdul-Azeem F. Thyroid activity of broiler chicks fed supplemental organic acids. International Journal of Poultry Science, 1: ,+/ῌ,,,. ,**2. Afsharmanesh M and Pourreza J. E#ect of calcium, citric acid, ascorbic acid, vitamin D- on the e$cacy of microbial phytase in broiler starters fed wheat-based diets on performance, bone mineralization and ileal digestibility. International Journal of Poultry Science, .: .+2ῌ.,.. ,**/. Association of O$cial Analytical Chemists. O$cial Methods of Analysis, +0th ed. (Cunnif P eds.). AOAC International, Virginia. +33/. Bates J, Jordens JZ and Gri$ths DT. 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