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1 The Professional Animal Scientist 21 (2005):59 65 Effects of Increasing Calcium-to- Phosphorus Ratio in Diets Containing Phytase on Finishing Pig Growth Performance S. M. HANNI*, M. D. TOKACH*, R. D. GOODBAND*,1,PAS,S.S.DRITZ, J. M. DEROUCHEY*, and J. L. NELSSEN* *Department of Animal Sciences and Industry and Food Animal Health and Management Center, College of Veterinary Medicine, Kansas State University, Manhattan Abstract A total of 288 finishing pigs were used to evaluate the effects of increasing the calcium-to-total phosphorus ratio (Ca:P) in diets containing phytase on growth performance. In Exp. 1, 144 pigs were fed diets with 300 phytase units (FTU)/kg of phytase and Ca:P ratios of 1.0:1, 1.25:1, 1.5:1, or 2.0:1 from 38.6 to 113 kg. Pigs were fed in three phases (d 0 to 28, 28 to 57, and 57 to 76) with total P levels of 0.45, 0.40, and 0.35%, respectively. Increasing Ca:P decreased ADG (quadratic; P<0.03), ADFI, and carcass weight (linear; P<0.05). However, the greatest decrease was observed when Ca:P increased from 1.5:1 to 2.0:1. In Exp. 2, 144 pigs (initially 38.6 kg) were fed diets containing 300 FTU/kg of phytase with Ca:P ratios of 0.75:1, 1.0:1, 1.25:1, 1.5:1, or 2.0:1. Diets were formulated to contain 0.44, 0.38, and 0.32% total P from d 0 to 28, 28 to 56, and 56 to 73, respectively. A sixth treatment group (negative control) was fed a diet containing 77% of the 1 Contribution no J from the Kansas Agric. Exp. Stn., Manhattan To whom correspondence should be addressed: Goodband@ksu.edu available P equivalent in other treatment diets to determine whether diets were formulated above the pig s P requirement. Increasing Ca:P decreased (quadratic; P<0.04) ADG and gain-to-feed ratio (G:F), with the greatest decrease observed as Ca:P increased from 1.5:1 to 2.0:1. Pigs fed the negative control diet had numerically decreased ADG and decreased (P<0.05) fat thickness. These studies suggest that finishing diets containing 300 FTU/kg of phytase should not have a Ca:P of >1.5:1. (Key Words: Calcium, Phosphorus, Phytase, Pigs.) Introduction Phytase releases phytic P from plant-based feedstuffs, which increases dietary P digestion and reduces the need for inorganic P supplementation (Cromwell et al., 1995b). It has been widely demonstrated that dietary supplementation with phytase is effective in making phytate-bound P nutritionally available to growing pigs (Cromwell et al., 1993; Jongbloed et al., 1993; Kornegay and Qian, 1996). Supplemental phytase in swine diets has resulted in improved growth performance and bone mineralization by increasing the digestibility and retention of P and Ca (Jongbloed et al., 1993; Lei et al., 1994). When formulating diets with phytase, after the P concentration has been decreased, it is uncertain whether or not Ca concentrations should also be decreased. Research has shown that high dietary Ca concentrations increase the formation of an insoluble Ca-phytate complex (Wise, 1983; Fisher, 1992) or reduce phytase activity. In weanling pigs, Qian et al. (1996) showed that decreasing the dietary Ca:P from 2.0:1 to 1.2:1 increased phytase efficacy approximately 16%, thus, improving performance, digestibility, bone measurements, and serum Ca levels. Liu et al. (1998), when utilizing low-p corn-soybean meal diets supplemented with microbial phytase, demonstrated that growth performance and P utilization were increased by lessening the Ca:P from 1.5:1 to 1.0:1 in grow-finish pigs. Therefore, our objective was to evaluate the effects of increasing Ca:P in diets containing phytase on finishing pig growth performance. Materials and Methods General. The Kansas State University Institutional Animal Care and

2 60 Hanni et al. TABLE 1. Diet composition (Exp. 1; as-fed basis). Item d 0 to 28 d 28 to 57 d 57 to 76 Ingredient a,% Corn Soybean meal, 46.5% CP Monocalcium phosphate, 21% P Limestone b Salt Vitamin premix c Trace mineral premix d Medication e Sand Lysine HCL Phytase f Total Calculated analysis Total lysine, % True digestible lysine, % ME, Mcal/kg Protein, % Ca, % Total P, % Available P, % Available P equivalence, % a All ingredients except corn were analyzed for Ca and P. These values were used along with the NRC (1998) estimates for Ca and P availability to achieve total P levels. The addition of 0.05% phytase to all of the diets provided 300 phytase units (FTU)/kg to achieve available P equivalence values. b Limestone replaced sand to achieve the desired Ca:P of 1.0:1, 1.25:1, 1.5:1, and 2.0:1. c Provided the following per kilogram of complete diet: vitamin A, 11,023 IU; vitamin D 3, 1653 IU; vitamin E, 44 IU; menadione (menadione bisulfate complex), 4.4 mg; vitamin B 12, 0.04 mg; riboflavin, 9.9 mg; pantothenic acid, 33 mg; and niacin, 55 mg. d Provided the following per kilogram of complete diet: Mn, 40 mg; Fe, 165 mg; Zn, 165 mg; Cu, 17 mg; I, 0.3 mg; and Se, 0.3 mg. e Provided 44 mg/kg of Tylosin (Elanco Animal Health, Greenfield, IN). f Provided 300 FTU/kg of feed (Natuphos 600; BASF Corporation, Mount Olive, NJ). Use Committee approved all experimental protocols used in this study. Pigs (Line 327 C22; PIC, Franklin, KY) were housed in an environmentally regulated finishing building. Each pen (1.5 m 2 ) had slatted concrete flooring and contained a stainless steel self-feeder and one nipple waterer to allow ad libitum consumption of feed and water. In both experiments, pigs were blocked by initial BW and sex, with two barrows or two gilts per pen, and then randomly allotted to one of the experimental treatments. In Exp. 1, there were 18 pigs per pen per treatment (nine barrows and nine gilts), and in Exp. 2, there were 12 pigs per pen per treatment (six barrows and six gilts). In both Exp. 1 and 2, diets were fed in meal form in three phases. Diets were formulated to meet or exceed nutrient requirements suggested by NRC (1998). In Exp. 2, a negative control diet was formulated to contain 77% of the NRC (1998) requirement estimate for available P, including the amount of P released by the phytase. The negative control treatment was included in the experiment to ensure dietary P levels were not above the pig s requirement as measured by differences in growth performance and percentage bone ash. Soybean meal, vitamin premixes, antibiotic, phytase, monocalcium phosphate, trace mineral premix, and limestone were analyzed for Ca and P using the AOAC (1995) methods and , respectively. These values were used along with the NRC (1998) estimates for Ca and P in corn as well as availability values to achieve total and available P levels used in each experiment. In both experiments, limestone replaced sand to achieve the desired Ca:P, and 0.05% phytase (Natuphos 600 BASF, Mount Olive, NJ) was added to all diets to provide 300 phytase units (FTU)/kg. We assumed 300 FTU/kg of phytase to release 0.08% P (BASF, 2000). Pigs were weighed, and feed disappearance was measured, every 14 d during the experiments to determine ADG, ADFI, and gain-to-feed ratio (G:F). All diets were analyzed using the AOAC (1995) procedures for CP (method ), Ca, and P (methods previously cited). At the conclusion of each experiment, pigs were marked with an individual tattoo to allow for individual carcass data to be collected at marketing. Pigs were then harvested (electrically stunned followed by exsanguination) in a USDA-inspected commercial packing plant. In Exp. 1, hot carcass weight, fat thickness, and longissimus area were recorded, and fat free lean index was calculated (NPPC, 2000). In Exp. 2, hot carcass weights and fat measurements were recorded. At time of slaughter, the right front foot was excised, tagged, bagged, and stored on ice. The feet were transported (approximately 2 h) to the Kansas

3 Calcium-to-Phosphorus Ratios and Phytase for Swine 61 State Swine Research lab and frozen at 20 C. To determine percentage of bone ash, feet were allowed to thaw for 1 h, and the third and fourth metacarpal bones were removed, cleaned of all extraneous tissue, and cut into three equal lengths. The bones were then soaked in diethyl ether for 7 d and dried at 100 C for 24 h. The bones were then allowed to cool for 1 h, weighed, and ashed in a muffle furnace at 600 C for 24 h. Exp. 1. One hundred forty-four pigs (72 barrows and 72 gilts; 38.6 ± 5.4 kg) were allotted randomly to one of four dietary treatments. Each dietary treatment had 18 replications (pens) with two pigs per pen. There were nine pens of barrows and nine of gilts. The dietary treatments were corn-soybean meal-based diets fed in three phases, Phases 1 and 2 were 28 d, and Phase 3 was 19 d (Table 1). These periods corresponded to approximately 38.6 to 66, 66 to 91.8, and 91.8 to 105 kg, respectively. In each time period, treatments contained Ca:P of 1.0:1, 1.25:1, 1.5:1, and 2.0:1. Diets were formulated to contain 1.10, 0.90, and 0.75% total dietary lysine; 0.45, 0.40, and 0.35% total P; 0.14, 0.11, and 0.07% available P; and 0.22, 0.18, and 0.15% available P equivalent (available P plus the P released by the phytase) for Phases 1, 2, and 3, respectively. Exp. 2. Similar to Exp. 1, 144 (72 barrows and 72 gilts; initially 38.5 ± 5.9 kg) were allotted randomly to one of six dietary treatments. Each treatment had 12 replications (pens), with six replications per sex and two pigs per pen. The corn-soybean meal-based diets were fed in three phases: Phases 1 and 2 were 28 d, and Phase 3 was 17 d (Table 2). These corresponded to approximately 38.5 to 66.3, 66.3 to 95.7, and 95.7 to kg, respectively. Dietary treatments were formulated to contain Ca:P of 0.75:1, 1.0:1, 1.25:1, TABLE 2. Diet composition (Exp. 2, as-fed basis). Item d 0 to 28 d 28 to 56 d 56 to 73 Ingredient a,% Corn Soybean meal, 46.5% CP Monocalcium phosphate, 21% P Limestone b Salt Vitamin premix c Trace mineral premix d Medication e Sand Lysine HCL Phytase f Total Calculated analysis Total lysine, % True digestible lysine, % ME, Mcal/kg Protein, % Ca, % Total P, % Available P, % Available P equivalent, % a All ingredients except corn were analyzed for Ca and P. These values were used along with the NRC (1998) estimates for Ca and P availability to achieve total P levels. The addition of 0.05% phytase to all of the diets provided 300 phytase units (FTU)/kg to achieve available P equivalence values. The negative control (1.1:1 Ca:P) was formulated to have 0.39, 0.33, and 0.30% total P; 0.10, 0.07, and 0.05% available P; and 0.18, 0.15, and 0.12% available P equivalent for Phases 1, 2, and 3, respectively. b Limestone replaced sand to achieve the desired Ca:P of 0.75:1, 1.0:1, 1.25:1, 1.5:1, and 2.0:1. c Provided the following per kilogram of complete diet: vitamin A, 11,023 IU; vitamin D 3, 1653 IU; vitamin E, 44 IU; menadione (menadione bisulfate complex), 4.4 mg; vitamin B 12, 0.04 mg; riboflavin, 9.9 mg; pantothenic acid, 33 mg; and niacin, 55 mg. d Provided the following per kilogram of complete diet: Mn, 40 mg; Fe, 165 mg; Zn, 165 mg; Cu, 17 mg; I, 0.3 mg; and Se, 0.3 mg. e Provided 44 mg/kg of Tylosin (Elanco Animal Health, Greefield, IN). f Provided 300 FTU/kg of feed (Natuphos 600; BASF Corporation, Mount Olive, NJ). 1.5:1, or 2.0:1. The sixth treatment group was a diet containing 77% of the available P equivalent (available P plus the amount of P released by phytase) as the other treatment diets. Diets were formulated to contain 1.10, 0.90, and 0.75% total dietary lysine; 0.44, 0.38, and 0.32% total P; 0.15, 0.11, and 0.07% available P; and 0.23, 0.19, and 0.15% available P equivalent for Phases 1, 2, and 3, respectively. The negative control (1.1:1 Ca:P) was formulated to have 0.39, 0.33, 0.30% total P; 0.10, 0.07, and 0.05% available P; and 0.18, 0.15, and 0.12% available P equivalent for Phases 1, 2, and 3, respectively. Statistical Analyses. In both Exp. 1 and 2, ANOVA was used to analyze the data as a randomized complete block design using the

4 62 Hanni et al. TABLE 3. Effects of Ca:total P in diets containing phytase on growth performance and bone ash in finishing pigs (Exp. 1) a. Ca:P Contrast (P<) Sex Item 1.0:1 1.25:1 1.5:1 2.0:1 SE Linear Quadratic Barrow Gilt P Initial BW, kg ADG, kg ADFI, kg Gain-to-feed ratio Final BW, kg Carcass data Carcass wt., kg Yield, % Fat thickness, cm Longissimus area, cm Fat-free lean index, % Bone ash, % 3rd Metacarpal th Metacarpal Average bone ash a Values are means of 144 pigs (initially 38.6 kg) with two pigs per pen and 18 replicate pens per treatment. There were no Ca:P by sex interactions (P<0.28). MIXED procedures of SAS (1998; SAS Inst. Inc., Cary, NC.). Both experiments were initially analyzed as factorial arrangements with main effects of Ca:P and gender. However, no Ca:P by gender interactions were observed, and this term was dropped from the model. In Exp. 1, linear and quadratic polynomial contrasts were used; in Exp. 2, linear, quadratic, and negative control vs other treatment contrasts were used to determine the effects of increasing Ca:P on pig growth performance, carcass characteristics, and bone ash percentage. The IML procedure was used to calculate coefficients to determine linear and quadratic contrasts for unevenly spaced treatments. Least squares differences were used to determine differences between treatments, sex, and their interaction (P<0.05). Pen was the experimental unit for all calculations in both Exp. 1 and 2. Results Exp. 1. There were no Ca:P by sex interactions (P>0.28) observed throughout the study. Increasing dietary Ca:P decreased ADG (quadratic; P<0.03) and ADFI (linear; P<0.05), but had no effect (P>0.20) on G:F (Table 3). These responses were similar among all three intermediate BW intervals (data not shown). Final BW and carcass weight also decreased (quadratic; P<0.01) with increasing Ca:P. The greatest decrease in ADG, ADFI, and final BW was observed when Ca:P increased from 1.5:1 to 2.0:1. There were no differences (P>0.26) in carcass yield, fat thickness, longissimus area, and fat-free lean index. There was a trend (linear; P<0.08) for decreased bone ash percentage for the 3rd, 4th metacarpals as Ca:P increased, with the greatest decrease observed when Ca:P increased from 1.5:1 to 2:1. Barrows had greater (P<0.05) ADG and ADFI compared with gilts; however, gilts had better (P<0.05) G:F, less fat thickness, and a greater fat-free lean index. There were no differences (P>0.49) in bone ash percentage among barrows and gilts. Exp. 2. There were no Ca:P by sex interactions (P>0.37) observed throughout the study. For the overall study, increasing Ca:P decreased (linear; P<0.003) ADG and G:F, with no effect on ADFI (P>0.42; Table 4). Similar to the response in both Phases 1 and 2, the greatest changes in ADG and G:F occurred when the Ca:P increased from 1.5:1 to 2.0:1. Increasing Ca:P ratio decreased final BW (linear; P<0.003) but, did not affect carcass weight (P>0.11). There was no difference in fat thickness for pigs fed Ca:P between 0.75:1 and 2.0:1 (P>0.17). However, pigs fed the negative control diet had reduced fat thickness (P<0.05) compared with pigs fed the other treatments. Bone ash percentage was not affected (P>0.23) by Ca:P. Similar to Exp. 1, barrows had a greater (P<0.001) ADG and ADFI compared with gilts, and gilts had an improved (P<0.03) G:F compared with the barrows. Also, barrows had greater (P<0.05) carcass weight and fat thickness compared with gilts. When comparing pigs fed the negative control diet to the mean of the other Ca:P treatments, there

5 Calcium-to-Phosphorus Ratios and Phytase for Swine 63 TABLE 4. Effects of Ca:total P in diets containing phytase on growth performance and bone ash in finishing pigs (Exp. 2) a. Ca:P Contrast (P<) Sex Negative (Neg.) Item control 0.75:1 1.0:1 1.25:1 1.5:1 2.0:1 SE Linear Quadratic Neg. vs others Barrow Gilt P Initial BW, kg ADG, kg ADFI, kg Gain-to-feed ratio Final BW, kg Carcass data Carcass wt., kg Yield, % Fat thickness, cm Bone ash, % 3rd Metacarpal th Metacarpal Average bone ash a Values are means of 144 pigs (initially 38.6 kg) with two pigs per pen and 18 replicate pens per treatment. There were no Ca:P by sex interactions (P<0.28). were no differences, mostly because of the lesser ADG and G:F of pigs fed the 2.0:1 Ca:P. However, ADG, G:F, and final BW for pigs fed the negative control diet were numerically poorer (P>0.06 to <0.17) than pigs fed the 0.75:1 to 1.25:1 Ca:P. Average daily gain and final BW for the negative control pigs were numerically greater (P>0.09) than pigs fed the 2.0:1 Ca:P. Gain to feed was greater (P<0.02) for pigs fed the negative control than for pigs fed the 2.0:1 Ca:P. Discussion Results of both studies indicated that increasing the Ca:P above 1.5:1 decreased growth performance. Lessening the Ca:P in finishing pigs in our experiment increased ADG and G:F but did not have an effect on ADFI. This is in agreement with experiments in which weanling (Lei et al., 1994; Qian et al., 1996) and growing-finishing pigs (Jongbloed et al., 1993; Harper et al., 1997; Adeola et al., 1998; Liu et al., 1998) were fed various Ca:P supplemented with microbial phytase. Cromwell et al. (1995a) reported that lessening the dietary Ca:P from 1.7:1 to 1.0:1 in a finisher diet had no effect on the efficacy of supplemental microbial phytase. A possible reason for the difference in results from Cromwell et al. (1995a) could be that we used 300 FTU/kg, whereas Cromwell used 1250 FTU/kg in their diets. The negative effect of greater dietary Ca concentrations on phytase activity was reduced in vitro when phytase was supplemented at the greater concentration (Qian and Kornegay, 1995). Qian et al. (1996) found that the Ca:P effect on supplemental phytase efficacy was more crucial at lesser dietary P levels. The NRC (1998) requirement estimate for finishing pigs is also for a relatively low Ca:P (1.25 to 1.1:1), which is in agreement with our re-

6 64 Hanni et al. sults. However, an important implication of our results would be that growth performance decreases as the Ca:P increases to >0.5:1 in pigs fed phytase, whereas data comparing the effect of increasing Ca:P in pigs not fed phytase would suggest that a wide ratio can be tolerated without negatively affecting performance (Hall et al., 1991). Three possible mechanisms have been proposed to explain the effects that a wide Ca:P has on decreasing phytase activity. High dietary Ca concentrations increase the ph of the intestinal contents, which in turn decreases microbial phytase activity (Sandberg et al., 1993). Second, the extra Ca forms an insoluble phytate complex that is not accessible for hydrolysis by phytase (Fisher, 1992). Last, the extra Ca could directly suppress phytase activity by competing for the active sites of the enzyme (Qian et al., 1996). Carcass characteristics in Exp. 1 were similar to those found by Liu et al. (1998), with fat thickness and dressing percentage not affected by the Ca:P. When decreasing the Ca:P, there was an increase in both hot carcass weight (Exp. 1) and BW (Exp. 1 and 2), which was in agreement with Liu et al. (1998). Even though the overall carcass weight was not affected in Exp. 2, carcass weight was numerically decreased as the Ca:P increased from 1.5:1 to 2.0:1, similar to Exp. 1. The decreased bone ash percentages in Exp. 1 were similar to observations of Qian et al. (1996) and Liu et al. (1998) in which they both found that increasing the Ca:P linearly decreased bone ash percentage. However, in Exp. 2, no response in bone ash percentage was observed. In Exp. 2, a negative control diet containing 77% of the available P equivalent of the other treatment diets was included to determine whether diets were formulated above the pig s P requirement. Although we cannot be totally certain the available P levels in our diets were not above the pig s requirement with just the use of a single control diet, this combined with the fact that our diets were formulated at or slightly below NRC (1998) estimates would suggest that we were not dramatically above the pig s requirement. Based on the differences in pig performance between the negative control and the other treatment diets, the treatment diets likely were not formulated above the pig s P requirement. If our experimental diets were formulated well above the pig s P requirement, this might bias our estimate of an appropriate Ca:P. In conclusion, these results suggest that increasing the Ca:P above 1.5:1 in a corn-soybean meal-based diet containing 300 FTU/kg of phytase for growing-finishing pigs can have a negative effect on growth performance, carcass traits, and bone ash percentage. Implications The use of phytase in swine diets to reduce P excretion in waste is becoming a standard procedure. When using phytase, maintaining a narrow Ca:total P (<1.5:1) is important in diet formulation to maximize growth performance, carcass traits, and bone ash percentage. Therefore, dietary Ca concentrations should be adjusted accordingly to maintain a narrow Ca:total P. Acknowledgments The authors would like to thank M. Barker, T. Keegan, C. Grosebeck, and C. Hastad for assistance in data collection. Literature Cited Adeola, O., T. R. Cline, J. I. Orban, D. Ragland, and A. L. Sutton Supplementation of low-calcium and low-phosphorus diets with phytase and cholecalciferol. In Purdue University Swine Day. Purdue University, West Lafayette, IN. AOAC Official Methods of Analysis. Vol. 1. (16th Ed.). Association of Official Analytical Chemists, Arlington, VA. BASF Phosphorus release comparison between phytase enzymes. Tech Bull No. TPU0014. BASF Corporation, Mount Olive, NJ. Cromwell, G. L., R. D. Coffey, G. R. Parker, H. J. Monegue, and J. H. Randolph. 1995b. Efficacy of a recombinant-derived phytase in improving the bioavailability of phosphorus in corn-soybean meal diets for pigs. J. Anim. Sci. 73:2000. Cromwell, G. L., M. D. Lindemann, G. R. Parker, R. D. Coffey, H. J. Monegue, and J. H. Raldolph. 1995a. Effects of dietary calcium on the efficacy of phytase and on the utilization of phosphorus in low phosphorus diets for pigs. J. Anim. Sci. 73(Suppl. 1):73. (Abs.) Cromwell, G. L., T. S. Stahly, R. D. Coffey, H. J. Monegue, and J. H. Randolph Efficacy of phytase in improving the bioavailability of phosphorus in soybean meal and corn-soybean meal diets for pigs. J. Anim. Sci. 71:1831. Fisher, H Low-calcium diets enhance phytate-phosphorus availability. Nutr. Rev. 50:170. Hall, D. D., G. L. Cromwell, and T. S. Stahly Effects of dietary calcium, phosphorus, calcium:phosphorus ratio and vitamin K on performance, bone strength and blood clotting status of pigs. J. Anim. Sci. 69:646. Harper, A. F., E. T. Kornegay, and T. C. Schell Phytase supplementation of low-phosphorus growing-finishing pig diets improves performance, phosphorus digestibility, and bone mineralization and reduces phosphorus excretion. J. Anim. Sci. 75:3174. Jongbloed, A. W., Z. Mroz, P. A. Kemme, C. Geerse, and Y. Van Der Honing The effect of dietary calcium levels on microbial phytase efficacy in growing pigs. J. Anim. Sci. 71(Suppl. 1):166. (Abs.) Kornegay, E. T., and H. Qian Replacement of inorganic phosphorus by microbial phytase for young pigs fed a maize-soybeanmeal diet. Br. J. Nutr. 76:563. Lei, X. G., P. K. Ku, E. R. Miller, M. T. Yokoyuma, and D. E. Ullrey Supplementing corn-soybean meal diets with microbial phytase maximizes phytate phosphorus utilization by weanling pigs. J. Anim. Sci. 71:3368. Liu, J., D. W. Bollinger, D. R. Ledoux, and T. L. Veum Lowering the dietary calcium to total phosphorus ratio increases phosphorus utilization in low-phosphorus corn-soybean meal diets supplemented with microbial phytase for growing-finishing pigs. J. Anim. Sci. 76:808.

7 Calcium-to-Phosphorus Ratios and Phytase for Swine 65 NPPC Pork Composition and Quality Assessment Procedures. Natl. Pork Producers Council, Des Moines, IA. NRC Nutrient Requirements of Swine. (10th Ed.). National Academy Press, Washington, DC. Sandberg, A. S., T. Larsen, and B. Sandstorm High dietary calcium level decreases colonic phytase degradation in pigs fed a rapeseed diet. J. Nutr.123:559. Qian, H., and E. T. Kornegay Effects of calcium, phosphorus and vitamin D 3 on in vitro microbial phytase (EC ) activity. J. Anim. Sci. 73(Suppl. 1):173. (Abs.) Qian, H., E. T. Kornegay, and D. E. Conner, Jr Adverse effects of wide calcium:phosphorus ratios on supplemental phytase efficacy for weanling pigs fed two dietary phosphorus levels J. Anim. Sci. 74:1288. Wise, A Dietary factors determining the biological activities of phytate. Nutr. Abstr. Rev. 53:791.

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