Sixty growing male pigs were used to test the hypothesis that high dietary Ca content reduces P absorption to a greater extent in microbial phytase-supplemented diets via reducing inositol phosphate (IP) degradation and enhancing P precipitation. Pigs were equally allotted over diets with three Ca contents 2·0, 5·8 and 9·6 g/kg with or without microbial phytase (0 v. 500 FTU/kg) in a 2 × 3 factorial arrangement. Faeces and urine were collected at the end of the 21-d experimental period. Subsequently, pigs were euthanised and digesta quantitatively collected from different gastrointestinal tract (GIT) segments. Increasing dietary Ca content reduced apparent P digestibility in all GIT segments posterior to the stomach (P < 0·001), with greater effect in phytase-supplemented diets in the distal small intestine (Pinteraction = 0·007) and total tract (Pinteraction = 0·023). Nonetheless, increasing dietary Ca to 5·8 g/kg enhanced P retention, but only in phytase-supplemented diets. Ileal IP6 degradation increased with phytase (P < 0·001) but decreased with increasing dietary Ca content (P = 0·014). Proportion of IP esters in total IP (∑IP) indicated that IP6/∑IP was increased while IP4/∑IP and IP3/∑IP were reduced with increasing dietary Ca content and also with a greater impact in phytase-supplemented diets (Pinteraction = 0·025, 0·018 and 0·009, respectively). In all GIT segments, P solubility was increased with phytase (P < 0·001) and tended to be reduced with dietary Ca content (P < 0·096). Measurements in GIT segments showed that increasing dietary Ca content reduced apparent P digestibility via reducing IP degradation and enhancing P precipitation, with a greater impact in phytase-supplemented diets due to reduced IP degradation.

The digestive system of the weaned piglets can be affected by the type of ingredients present in the diet, and a high fibre content in diets can limit the use of other nutrients and energy. The study was conducted to determine the effects of multicarbohydrase (MC) and phytase (Phy) supplementation on the nutritive value of wheat bran (WB) in weaned piglets. Multicarbohydrase preparation had 700 U α-galactosidase, 2200 U galactomannanase, 3000 U xylanase and 22 000 U β-glucanase per kilogram of diet, and Phy had 500 phytase units – FTU/kg of diet. Twenty-five weaned piglets (6.1 ± 0.63 kg) at 21 days old were fed five diets in a completely randomised experimental design with a 2 × 2 + 1 (0 and 200 mg/kg MC; 0 and 50 mg/kg Phy; and basal diet – BD) factorial arrangement used to determine treatment effects. An additional group of piglets was fed a corn-basal diet during apparent digestibility of nutrients, and fed a 5% casein-corn starch basal diet during apparent and standardised ileal digestibility (SID) of amino acid evaluations. Piglets were individually caged until 38 days old, when Ileal digesta was collected at slaughter. Test diets were made by mixing the basal diets and WB 7 : 3 (w/w), with or without MC, Phy or the combination. There was an interaction trend (P = 0.07) between MC and Phy in the balance of ash, digestible energy (DE) and metabolisable energy (ME). Effects of MC (P < 0.01) on DM, N retention, DE and ME, as well as an effect of Phy (P < 0.05) on ash, DE and ME and a trend in protein digestibility (P = 0.07) also was observed. The enzyme combination showed effect (P < 0.05) on SID of Lys, Pro and Ser; as a trend (P < 0.07) on His, Thr and Val. Isolated, MC also suggested improving (P < 0.07) on SID of His, Lys, Ala (P < 0.05), while Phy improved (P < 0.06) SID of Leu, Lys, Met (P < 0.01), Thr, Val, Ala (P < 0.01), Pro and Ser (P < 0.05). The MC carbohydrate complex was characterised as a viable alternative to increase the apparent nutrients digestibility and SID of amino acids when WB was used in the diet of young pigs and, when combined with Phy, suggested an additive effect on the apparent use of energy.

Dietary phosphorus concentration greatly affects pig’s growth performance, environmental impact and diet cost. A total of 1080 pigs (initially 5.9 ± 1.08 kg) from three commercial research rooms were used to determine the effects of increasing standardized total tract digestible (STTD) P concentrations in diets without and with phytase on growth performance and percentage bone ash. Pens (10 pigs/pen, 9 pens/treatment) were balanced for equal weights and randomly allotted to 12 treatments. Treatments were arranged in two dose titrations (without or with 2000 units of phytase) with six levels of STTD P each. The STTD P levels were expressed as a percentage of NRC (2012) requirement estimates (% of NRC; 0.45 and 0.40% for phases 1 and 2, respectively) and were: 80%, 90%, 100%, 110%, 125% and 140% of NRC in diets without phytase and 100%, 110%, 125%, 140%, 155% and 170% of NRC in diets with phytase. Diets were provided in three phases, with experimental diets fed during phases 1 (days 0 to 11) and 2 (days 11 to 25), followed by a common diet from days 25 to 46. On day 25, radius samples from one median-weight gilt per pen were collected for analysis of bone ash. During the treatment period, increasing STTD P from 80% to 140% of NRC in diets without phytase improved average daily gain (ADG; quadratic, P < 0.01), average daily feed intake (ADFI; quadratic, P < 0.05) and gain–feed ratio (G : F; linear, P < 0.01). Estimated STTD P requirement in diets without phytase was 117% and 91% of NRC for maximum ADG according to quadratic polynomial (QP) and broken-line linear (BLL) models, respectively, and was 102%, 119% and >140% of NRC for maximum G : F using BLL, broken-line quadratic and linear models, respectively. When diets contained phytase, increasing STTD P from 100% to 170% of NRC improved ADG (quadratic, P < 0.05) and G : F (linear, P < 0.01). Estimated STTD P requirement in diets containing phytase was 138% for maximum ADG (QP), and 147% (QP) and 116% (BLL) of NRC for maximum G : F. Increasing STTD P increased (linear, P < 0.01) the percentage bone ash regardless of phytase addition. When comparing diets containing the same STTD P levels, phytase increased (P < 0.01) ADG, ADFI and G : F. In summary, estimated STTD P requirements varied depending on the response criteria and statistical models and ranged from 91% to >140% of NRC (0.41% to >0.63% of phase 1 diet and 0.36% to >0.56% of phase 2 diet) in diets without phytase, and from 116% to >170% of NRC (0.52% to >0.77% of phase 1 diet and 0.46% to >0.68% of phase 2 diet) for diets containing phytase. Phytase exerted an extra-phosphoric effect on promoting pig’s growth and improved the P dose-responses for ADG and G : F.

Phytase is applied in animal feed based on its standard activity measured at pH 5.5, however the relative activity at pH 3 (e.g. stomach pH, the main site for the breakdown of phytate) varies among the commercial phytases, ranging from 56% (an E coli phytase) to 235% (Buttiauxella phytase). These diverse sources of phytases have varying capability for degrading phytate and, correspondingly, different P, digestible amino acid and metabolisable energy matrix values. In addition, the matrix values recommended by different phytase suppliers are not comparable, as different methodologies have been used to determine them. Phosphorus (P) and other matrix values can be determined by direct measurement of digestible P (dP) improvements by the addition of phytase above a negative control in large numbers of in vivo studies using increasing phytase doses. Alternatively, matrix values can be assessed by indirect measurement, using inorganic P (usually mono- or dicalcium sources) as a reference, typically based on tibia or metacarpal ash as a response parameter to estimate available P equivalence, either at a single or different phytase doses. When using the indirect measurement, the available P equivalence with increasing phytase doses may be calculated based on a log linear model. Although both methods are acceptable methodologies, direct measurement may under-estimate and indirect measurement may over-estimate matrix values, and a large number of in vivo studies give the best estimates of matrix values. Phytase efficacy can be influenced by phytase source, dose level, dietary composition (Ca level and Ca: P ratio). Phytase end users are encouraged to be aware of the methods used by suppliers to determine matrix values, before applying them in their feed formulations.

This study evaluated the effects of three different thermostable phytase variants, based on the AppA gene from E. coli (AppAT1, AppAT2 and AppAT3) on growth performance, nutrient digestibility and bone mineralisation in broiler chickens at inclusion levels of 250 and 500 FTU/kg. The eight treatment groups included a positive control (PC) which was sufficient in Ca and P, a negative control (NC, the same basal formulation as the PC, but reduced in Ca and P), and NC supplemented with AppAT1 at 250 and 500 FTU/kg (AppAT1-250 and AppAT1-500), AppAT2 at 250 and 500 FTU/kg (AppAT2-250 and AppAT2-500) and with AppAT3 at 250 and 500 FTU/kg (AppAT3-250 and AppAT3-500). Over the entire feeding period, body weight (BW) and average daily gain (ADG) were significantly higher in the PC group, with all phytase supplemented groups being statistically the same, compared to the NC group. Feed conversion (FCR) for the PC-fed birds (1.479) was significantly (P<0.05) better compared to the NC birds (1.582) and those fed the AppAT3-250 diet (1.523). Reduced levels of Ca and P in the NC group led to significantly (P<0.05) lower tibia ash (40.9%) compared to the PC group (47.4%). Birds fed the phytase diets had significantly higher tibia ash compared to the NC birds, with those from the AppAT2-500 and AppAT3-500 groups being statistically the same as the PC group. Diets AppAT1-500, AppAT2-250, AppAT2-500 and AppAT3-500 significantly increased Ca digestibility compared to the NC. Apparent total track digestibility (ATTD) of P was improved for AppAT1-500 and AppAT2-250. The ATTD of Ca and P for all of the phytase supplemented groups reached the same level of the PC and AppAT1-500 group. It was concluded that adding any of the phytases tested, especially when included at 500 FTU/kg to a feed reduced in Ca and P, led to improved performance and bone mineralisation back to the same levels as seen for the Ca and P sufficient diet.

Ca digestibility and utilisation in growing pigs are not well understood, and are usually neglected in diet formulation. This has implications not only for the accurate determination of its requirements but also for its interactions with other nutrients. A systematic review and meta-analysis (meta-regression) of published trials was carried out to quantify factors affecting Ca absorption and utilisation, and to derive an estimate of Ca endogenous excretion. The analysis was carried out on the data from forty studies, corresponding to 201 treatments performed on 1204 pigs. The results indicated that although Ca absorption and retention (g/kg of body weight per d) increased with increasing Ca intake (P<0·001), non-phytate-P intake (P<0·001) and exogenous phytase supplementation (P<0·001), these values decreased with increasing phytate-P intake (P<0·05). Interactions between exogenous phytase and Ca intake, indicating reduced efficacy of this enzyme (P<0·001), and between phytate-P intake and exogenous phytase, counteracting the direct negative effect of phytate-P (P<0·05) on Ca absorption and retention, were also detected. There were no effects of animal-related characteristics, such as pig genotype in Ca absorption and retention. The large amount of variance explained in Ca absorption (90 %) and retention (91 %) supported our choice of independent variables. Endogenous Ca losses obtained via linear regression were 239 mg/kg of DM intake (95 % CI 114, 364). These outcomes advance the current understanding of Ca digestibility and utilisation, and should contribute towards establishing requirements for digestible Ca. Consequently, pig diets will be more correctly formulated if digestible Ca values are used in estimating requirements for Ca.

The objective of the current experiment was to determine the effects of high-concentration phytase (5000 FTU/kg) feeding to diverse lines of chickens fed phosphorus (P) adequate maize–soybean meal diets (4.5 g/kg non-phytate P) on the performance and intestinal immune function. Performance was measured for outbred broiler (Ross 308) and inbred Fayoumi lines over 0–21 days, and duodenum and ileum were harvested for the determination of mucin-2, interleukin (IL)-1β and IgA mRNA by quantitative reverse transcription polymerase chain reaction. Over the 0–7-day period, there was a significant line × diet interaction, as high phytase supplementation increased broiler average daily gain (ADG), but had no effect on Fayoumi ADG. Treatment of diets with phytase increased expression of the mucin-2 gene in the duodenum mucosa. There were significant interactions between line and age, and line, diet and age on duodenal expression of the IL-1β gene as phytase supplementation of the broiler line reduced IL-1β in comparison to control fed broilers without change in the Fayoumi line. Overall, the addition of a high concentration of phytase to broilers fed adequate concentrations of non-phytate P resulted in improved growth performance early with a reduction in this effect over time. Mucosal mucin-2 expression was increased with high-concentration phytase feeding across both lines, but IL-1β mRNA expression was reduced in the duodenum of broilers fed high concentrations of phytase, suggesting that the increased performance noted might be related to decreased inflammation.

A study was conducted to evaluate productive performance in laying hens fed diets with reduced nutrient density based on the nutritional contribution of a Buttiauxella phytase in laying-hen diets from 21-57 weeks of age. A commercial laying hen diet was offered ad libitum to the 480 ISA Brown laying hens from 18-21 weeks of age. From 21 weeks onwards, the hens received one of four dietary treatments: a positive control (PC) diet, a down specified diet (DS1) + phytase at 300 FTU/kg, a second down-specified diet (DS2) + phytase at 600 FTU/kg and a third test diet formulated as per DS1 + phytase at 1,200 FTU/kg feed. The PC was formulated based on ISA breeder recommendations. DS1 was formulated with reduction of 0.149% available P, 0.134% Ca, 55 kcal/kg AME, 0.33% CP, digestible amino acids (up to 0.015%) and 0.013% Na based on the contribution of Buttiauxella phytase at 300 FTU/kg. DS2 was formulated with reduction of 0.177% available P, 0.159% Ca, 60 kcal/kg AME, 0.61% CP, digestible amino acids (up to 0.028%) and 0.02% Na based on the contribution of Buttiauxella phytase at 600 FTU/kg. Every dietary treatment was fed to 12 cages containing 10 hens each. The trial treatments did not include a full, DS negative control, as ethical considerations regarding birds' welfare when feeding such diets over such an extended period of time did not permit this. No significant differences were seen in hen-day egg production, feed intake, egg weight, feed to egg mass ratio, shell, yolk or albumen proportion, unsaleable eggs or shell breaking strength in laying hens fed the PC diet or the DS diets with added phytase. Yolk colour increased significantly with phytase supplementation. Supplementing the DS1 diet with 300 FTU and the DS2 diet with 600 FTU resulted in non-significant differences in tibia ash, Ca and P, compared to the PC diet. The data from this study indicated that applying the nutrient contributions for Buttiauxella phytase at 300 and 600 FTU/kg maintained the egg production, BW and egg quality parameters compared to PC. The best economic efficiency value during the whole experimental period was recorded with phytase at 600 FTU/kg when full matrix values are used. When commercial diets are formulated based on ISA breeder recommendations, lowering diet nutrient density while supplementing with phytase reduced the overall diet cost, which should contribute to the profitability of egg production.

The effect of the ingestion of diets containing either myo-inositol or exogenous phytase on plasma metabolites was examined using 29 kg barrows. The diets were: control (maize, soya, rapeseed, rice bran), control plus 2 g/kg myo-inositol, control plus 1000 phytase units (FYT)/kg or 3000 FYT/kg exogenous phytase. Pigs were housed in a PigTurn device and blood was collected, from jugular catheters, via an automated system at −30, (30 min before feeding), 0, 15, 30, 45, 60, 90, 120, 150, 180, 240, 300 and 360 min post-feeding. The addition of 2 g/kg myo-inositol to the basal diet resulted in an increase in plasma myo-inositol concentration that was evident 45–60 min after diet introduction and persisted to 360 min post-feeding. Similarly, supplementation of the basal diet with either 1000 or 3000 FYT/kg exogenous phytase resulted in an increase in plasma myo-inositol concentration that was still rising 360 min post-feeding. Plasma P concentration was increased over time by the addition of 1000 and 3000 FYT/kg phytase, but not by the addition of myo-inositol. Other plasma metabolites examined were not affected by dietary treatment. It can be concluded that oral delivery of myo-inositol results in rapid increase in plasma myo-inositol concentrations that peak approximately 45–60 min after feeding. Use of supplemental phytase achieves similar increases in myo-inositol concentration in plasma but the appearance is more gradual. Furthermore, supplementation of pig diets with exogenous phytase results in rapid appearance of P in plasma that may be sustained over time relative to diets with no added phytase.

Around 70% of total seed phosphorus is represented by phytate which must be hydrolysed to be bioavailable in non-ruminant diets. The limited endogenous phytase activity in non-ruminant animals make it common practice to add an exogenous phytase source to most poultry and pig feeds. The mature grain phytase activity (MGPA) of cereal seeds provides a route for the seeds themselves to contribute to phytate digestion, but MGPA varies considerably between species and most varieties in current use make negligible contributions. Currently, all phytases used for feed supplementation and transgenic improvement of MGPA are derived from microbial enzymes belonging to the group of histidine acid phosphatases (HAP). Cereals contain HAP phytases, but the bulk of MGPA can be attributed to phytases belonging to a completely different group of phosphatases, the purple acid phosphatases (PAPhy). In recent years, increased MGPAs were achieved in cisgenic barley holding extra copies of barley PAPhy and in the wheat HIGHPHY mutant, where MGPA was increased to ~6200 FTU/kg. In the present study, the effect of replacing 33%, 66% and 100% of a standard wheat with HIGHPHY wheat was compared with a control diet with and without 500 FTU of supplemental phytase. Diets were compared by evaluating broiler performance, ileal Ca and P digestibility and tibia development, using nine replicate pens of four birds per diet over 3 weeks from hatch. There were no differences between treatments in any tibia or bird performance parameters, indicating the control diet did not contain sufficiently low levels of phosphorus to distinguish effect of phytase addition. However, in a comparison of the two wheats, the ileal Ca and P digestibility coefficients for the 100% HIGHPHY wheat diets are 22.9% and 35.6% higher, respectively, than for the control diet, indicating the wheat PAPhy is functional in the broiler digestive tract. Furthermore, 33% HIGHPHY replacement of conventional wheat, significantly improved Ca and P digestibility over the diet-supplemented exogenous phytase, probably due to the higher phytase activity in the HIGHPHY diet (1804 v. 1150 FTU). Full replacement by HIGHPHY gave 14.6% and 22.8% higher ileal digestibility coefficients for Ca and P, respectively, than for feed supplemented with exogenous HAP phytase at 500 FTU. This indicates that in planta wheat PAPhys has promising potential for improving P and mineral digestibility in animal feed.

Crossbred pigs (n=720; average age=28±3 days and weight=9.5±0.3 kg) were used in a 20-day trial in order to determine the influence of phosphorus (P) source and various doses of pharmacological zinc (Zn) on growth performance, plasma minerals and mineral digestibility. Pigs (five intact males and five females per pen) were randomly allotted to treatments in a 3×3 factorial arrangement with three sources of dietary P (4.5 g/kg digestible P, 4.5 g/kg digestible P plus 2500 phytase units (FTU)/kg, or 5.5 g/kg digestible P) and three dietary levels of supplemental Zn (0, 1750 or 3500 mg/kg) from ZnO (82% Zn) with eight pens per treatment. Diets were formulated to exceed all nutrient requirements, including calcium (Ca), P and Zn from day 0 to 20. Zn supplementation increased (quadratic P<0.05) average daily feed intake. There was a significant Zn level×P source interaction on average daily gain and feed conversion ratio (FCR). Pigs fed 4.5 g/kg digestible P without or with 2500 FTU/kg phytase gained more per day (quadratic P<0.05) and had better FCR (quadratic P<0.05) when they were fed 1750 mg/kg supplemental Zn. However, pigs fed 5.5 g/kg digestible P gained more per day (linear P<0.05) and were more efficient (linear P<0.05) when they were fed 3500 mg/kg supplemental Zn. Plasma Zn and Zn digestibility increased (linear P<0.05) as pharmacological Zn supplementation increased from 0 to 3500 mg/kg, irrespective of P source. However, Ca, P, sodium (Na), potassium (K) and copper (Cu) digestibility were reduced (P<0.05) as pharmacological Zn supplementation increased, and this was mitigated or exacerbated by the supplementation of 5.5 g/kg digestible P or phytase. In conclusion, increasing the dietary inclusion of pharmacological Zn may impact growth performance in young pigs through the interaction with minerals such as Ca, P, Na and K. Pharmacological Zn may reduce Na or K digestibility and indirectly reduce water secretion into the lumen, resulting in an increase in faecal dry matter as pharmacological Zn supplementation in the diet increased.

The objective was to evaluate the effect of microbial phytase (1250 FTU/kg diet with 88% dry matter (DM)) on apparent total tract digestibility (ATTD) of phosphorus (P) in pigs fed a dry or soaked diet. Twenty-four pigs (65±3 kg) from six litters were used. Pigs were housed in metabolism crates and fed one of four diets for 12 days; 5 days for adaptation and 7 days for total, but separate collection of feces and urine. The basal diet was composed of wheat, barley, maize, soybean meal and no mineral phosphate. Dietary treatments were: basal dry-fed diet (BDD), BDD with microbial phytase (BDD+phy), BDD soaked for 24 h at 20°C before feeding (BDS) and BDS with microbial phytase (BDS+phy). Supplementation of microbial phytase increased ATTD of DM and crude protein (N×6.25) by 2 and 3 percentage units (P<0.0001; P<0.001), respectively. The ATTD of P was affected by the interaction between microbial phytase and soaking (P=0.02). This was due to a greater increase in ATTD of P by soaking of the diet containing solely plant phytase compared with the diet supplemented with microbial phytase: 35%, 65%, 44% and 68% for BDD, BDD+phy, BSD and BSD+phy, respectively. As such, supplementation of microbial phytase increased ATTD of P in the dry-fed diet, but not in the soaked diet. The higher ATTD of P for BDS compared with BDD resulted from the degradation of 54% of the phytate in BDS by wheat and barley phytases during soaking. On the other hand, soaking of BDS+phy did not increase ATTD of P significantly compared with BDD+phy despite that 76% of the phytate in BDS+phy was degraded before feeding. In conclusion, soaking of BDS containing solely plant phytase provided a great potential for increasing ATTD of P. However, this potential was not present when microbial phytase (1250 FTU/kg diet) was supplemented, most likely because soaking of BDS+phy for 24 h at 20°C did not result in a complete degradation of phytate before feeding.

Phytase (PHY) improves growth performance, nutrient digestibility and bone structure in pigs; however, little is known about its effects on intestinal nutrient transporter gene expression. In the present study, a 44 d experiment was carried out using forty-eight pigs (11·76 (sem 0·75) kg) assigned to one of three dietary treatment groups to measure growth performance, coefficient of apparent ileal digestibility (CAID), coefficient of apparent total tract nutrient digestibility (CATTD) and intestinal nutrient transporter gene expression. Dietary treatments during the experimental period were as follows: (1) a high-P (HP) diet containing 3·4 g/kg available P and 7·0 g/kg Ca; (2) a low-P (LP) diet containing 1·9 g/kg available P and 5·9 g/kg Ca; (3) a PHY diet containing LP diet ingredients+1000 phytase units (FTU)/kg of PHY. The PHY diet increased the average daily gain (P< 0·05) and final body weight (P< 0·01) and decreased the feed conversion ratio (P< 0·05) compared with the LP diet. Pigs fed the PHY diet had a higher CAID of gross energy compared with those fed the HP and LP diets (P< 0·001). Pigs fed the PHY diet had increased CAID of P (P< 0·01) and CATTD of Ca and P (P< 0·001) compared with those fed the LP diet. The PHY diet increased the gene expression of the peptide transporter 1 (PEPT1/SLC15A1) (P< 0·05) in the ileum compared with the LP diet. The LP diet decreased the gene expression of the sodium–glucose-linked transporter 1 (SGLT1/SLC5A1) and GLUT2/SLC2A2 (P< 0·05) and increased the expression of membrane Ca channel (TRPV6) and calbindin compared with the HP diet (P< 0·001). In conclusion, feeding a diet supplemented with PHY improves growth performance and nutrient digestibility as well as increases the gene expression of the peptide transporter PEPT1.

Optimising the supply of dietary phosphorus (P) to animals requires sound knowledge regarding the availability of P from feed raw materials. Different definitions of available P have been proposed and various approaches are in use to determine P availability. These differences make it difficult to compare published data and to compile comprehensive feeding tables for use in the industry. This is disadvantageous in regard to feeding costs and sustainable handling of the limited global phosphate reserves. Working Group No 2 (Nutrition) of the European Federation of Branches of WPSA has developed a standard protocol for determination of available P in broilers. It is based on precaecal P digestibility and the details of this proposed protocol are explained in the following paper. Unresolved questions in regard to the determination of available P are also addressed. Researchers are encouraged to implement this standard protocol in future P availability studies and to contribute to its further improvement.

Zinc (Zn) is essential for swine and poultry and native Zn concentrations in feedstuffs are too low to meet their Zn requirement. Dietary Zn bioavailability is affected by phytate, phytase and Zn supplemented in organic form is considered as more bioavailable than inorganic sources. A meta-analysis using GLM procedures was processed using broiler and piglet databases to investigate, within the physiological response of Zn, (1) the bioavailability of inorganic and organic Zn sources (Analysis I); (2) the bioavailability of native and inorganic Zn dependent from dietary phytates, vegetal and supplemental phytase activity (Analysis II). Analysis I: the bioavailability of organic Zn relative to inorganic Zn sources ranged, depending on the variable, from 85 to 117 never different from 100 (P > 0.05). The coefficients of determination of the regressions were 0.91 in broilers and above 0.89 in piglets. Analysis II: in broilers, bone Zn was explained by supplemental Zn (linear and quadratic, P < 0.001) and by supplemental phytase (linear, P < 0.001). In piglets, the interaction between dietary Zn and phytates/phytases was investigated by means of a new variable combining dietary phytic phosphorus (PP) and phytase activity. This new variable represents the remaining dietary PP after its hydrolysis in the digestive tract, mainly due to phytase and is called non-hydrolyzed phytic phosphorus (PPNH). Bone Zn was increased with native Zn (P < 0.001), but to a lower extent in high PP or low phytase diets (ZNN × PPNH, P < 0.001). In contrast, the increase in bone zinc in response to supplemental Zn (P < 0.001) was not modulated by PPNH (P > 0.05). The coefficients of determination of the regressions were 0.92 in broilers and above 0.92 in piglets. The results from the two meta-analyses suggest that (1) broilers and piglets use supplemented Zn, independent from Zn source; (2) broiler use native Zn and the use is slightly enhanced with supplemental phytase; (3) however, piglets are limited in the use of native Zn because of the antagonism of non-hydrolyzed dietary phytate. This explains the higher efficacy of phytase in improving Zn availability in this specie.

Optimizing phosphorus (P) utilization in pigs requires improving our capacity to predict the amount of P absorbed and retained, with the main modulating factors taken into account, as well as precisely determining the P requirements of the animals. Given the large amount of published data on P utilization in pigs, a meta-analysis was performed to quantify the impact of the different dietary P forms, calcium (Ca) and exogenous phytases on the digestive and metabolic utilization criteria for dietary P in growing pigs. Accordingly, the amount of phytate P (PP) leading to digestible P (g/kg) was estimated to be 21%, compared with 73% for non-phytate P (NPP) from plant ingredients and 80% for NPP from mineral and animal ingredients (P < 0.001). The increase in total digestible dietary P following the addition of microbial phytase (PhytM) from Aspergillus niger (P < 0.001) was curvilinear and about two times higher than the increase following the addition of plant phytase, which leads to a linear response (P < 0.001). The response of digestible P to PhytM also depends on the amount of substrate, PP (PhytM2 × PP, P < 0.001). The digestibility of dietary P decreased with dietary Ca concentration (P < 0.01) independently of phytase but increased with body weight (BW, P < 0.05). Although total digestible dietary P increased linearly with total NPP concentration (P < 0.001), retained P (g/kg), average daily gain (ADG, g/day) and average daily feed intake (ADFI, g/day) increased curvilinearly (P < 0.001). Interestingly, whereas dietary Ca negatively affected P digestibility, the effect of dietary Ca on retained P, ADG and ADFI depended on total dietary NPP (NPP × Ca, P < 0.01, P < 0.05 and P < 0.01, respectively). Increasing dietary Ca reduced retained P, ADG and ADFI at low NPP levels, but at higher NPP concentrations it had no effect on ADG and ADFI despite a positive effect on retained P. Although the curvilinear effect of PhytM on digestible P increased with PP (P < 0.001), this effect was lessened by total NPP for ADG and ADFI (PhytM × NPP and PhytM2 × NPP, P < 0.05) and depended on both total NPP and Ca for retained P (PhytM2 × NPP × Ca, P < 0.01). This meta-analysis improves our understanding of P utilization, with major modulating factors taken into account. The information generated will be useful for the development of robust models to formulate environmentally friendly diets for growing pigs.

Protein–phytate interactions are fundamental to the detrimental impact of phytate on protein/amino acid availability. The inclusion of exogenous phytase in pig and poultry diets degrades phytate to more innocuous esters and attenuates these negative influences. The objective of the present review is to reappraise the underlying mechanisms of these interactions and reassess their implications in pig and poultry nutrition. Protein digestion appears to be impeded by phytate in the following manner. Binary protein–phytate complexes are formed at pH levels less than the isoelectric point of proteins and complexed proteins are refractory to pepsin digestion. Once the protein isoelectric points are exceeded binary complexes dissociate; however, the isoelectric point of proteins in cereal grains may be sufficiently high to permit these complexes to persist in the small intestine. Ternary protein–phytate complexes are formed at pH levels above the isoelectric point of proteins where a cationic bridge links the protein and phytate moieties. The molecular weights of protein and polypeptides in small-intestinal digesta may be sufficient to allow phytate to bind nutritionally important amounts of protein in ternary complexes. Thus binary and ternary complexes may impede protein digestion and amino acid absorption in the small intestine. Alternatively, phytate may interact with protein indirectly. Myo-inositol hexaphosphate possesses six phosphate anionic moieties (HPO42–) that have strong kosmotropic effects and can stabilise proteins by interacting with the surrounding water medium. Phytate increases mucin secretions into the gut, which increases endogenous amino acid flows as the protein component of mucin remains largely undigested. Phytate promotes the transition of Na+ into the small-intestinal lumen and this suggests that phytate may interfere with glucose and amino acid absorption by compromising Na+-dependent transport systems and the activity of the Na pump (Na+-K+-ATPase). Starch digestion may be depressed by phytate interacting with proteins that are closely associated with starch in the endosperm of cereal grains. While elucidation is required, the impacts of dietary phytate and exogenous phytase on the site, rate and synchrony of glucose and amino acid intestinal uptakes may be of importance to efficient protein deposition. Somewhat paradoxically, the responses to phytase in the majority of amino acid digestibility assays in pigs and poultry are equivocal. A brief consideration of the probable reasons for these inconclusive outcomes is included in this reappraisal.

Calcium is the most prevalent mineral in the body and is important for many physiological processes. However, the amount of calcium in modern diets has both economic and nutritional consequences for producers. Dietary calcium has been shown to reduce the efficacy of both endogenous and exogenous phytases resulting in decreased phytate-phosphorus availability, increased phosphorus excretion as well as facilitating the formation of mineral-phytate complexes. Reducing the concentration of dietary calcium has been reported to improve phytase efficacy and phytate-phosphorus availability, however, this is often at the expense of optimal skeletal integrity. Choice feeding employs the principle that poultry have the ability to select a nutritionally balanced diet from multiple sources and previous studies have shown that poultry have a calcium specific appetite. However, the evaluation of choice feeding systems for broilers has predominantly focussed on the effects of providing separate sources of protein and energy concentrates. Though literature is available for laying hens, scant information is available pertaining to choice feeding of a separate calcium source for broilers. This paper reviews the available information on choice feeding systems in poultry with focus on the application of this in broiler production systems.

Phytases have been used commercially since the early 1990s and have been the focus of considerable and sustained research for many decades. Despite this heroic effort there are still areas of persistent uncertainty such as the obscurity surrounding total compared with digestible calcium, appropriate modification to dietary sodium (and other electrolyte) concentrations, the usefulness of the amino acid and energy digestibility improvements and ultimately the effect of phytase on nutrient requirement. One further area which has attracted some attention recently is the effect of unconventionally high doses of phytase (i.e. >2,500 FTU/kg from Aspergillus niger or Escherichia coli) in an attempt to ostensibly ‘de-phytinise’ the diet. The effects of such ‘super’ doses of phytase can be considerable, and often beyond that which may be reasonably expected based on improvement in P digestibility per se. This review article addresses these effects and suggests mechanisms by which they may be explained.

Two experiments, a performance experiment and a mineral balance study, were conducted on grower–finisher pigs (42 to 101 kg live weight) to investigate the effects of Peniophora lycii phytase enzyme and 25-hydroxyvitamin D3 (25-OHD3) on growth performance, carcass characteristics, nutrient retention and excretion, and bone and blood parameters. The two experiments were designed as a 2 × 2 factorial (two levels of phytase and two levels of 25-OHD3). The four diets were T1, low-phosphorous diet; T2, T1 + phytase; T3, T1 + 25-OHD3 and T4, T1 + phytase + 25-OHD3 diet. In all, 25 μg of 25-OHD3 was used to replace 1000 IU of vitamin D3 in diets T3 and T4. Diets were pelleted (70°C) and formulated to contain similar concentrations of energy (13.8 MJ DE/kg), lysine (9.5 g/kg) and digestible phosphorus (P; 1.8 g/kg). Neither the inclusion of phytase nor 25-OHD3 in the diet had any effect on pig performance. There was an interaction between phytase and 25-OHD3 on calcium (Ca) and P retention (P < 0.01) and on the apparent digestibility of ash (P < 0.01), P (P < 0.001) and Ca (P < 0.001). Pigs offered phytase diets only, had a higher retention of Ca and P and digestibility of ash (P < 0.01), P (P < 0.001) and Ca (P < 0.01) compared with pigs offered unsupplemented diets. However, when the combination of phytase and 25-OHD3 were offered, no effects were detected compared with 25-OHD3 diets only. Pigs fed phytase diets had higher bone ash (P < 0.01), bone P (P < 0.01) and bone Ca (P < 0.05) concentrations compared with pigs offered non-phytase diets. In conclusion, pigs offered phytase diets had a significantly increased bone ash, Ca and P than pigs offered unsupplemented phytase diets. However, there was no advantage to offering a combination of phytase and 25-OHD3 on either bone strength or mineral status compared to offering these feed additives separately.