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Growth performance and nutrient digestibility of growing and finishing pigs fed multienzyme-supplemented low-energy and -amino acid diets

Growth performance and nutrient digestibility of growing and finishing pigs fed... Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Growth performance and nutrient digestibility of growing and finishing pigs fed multienzyme-supplemented low-energy and -amino acid diets † † † ‡ ‡ Kevin Jerez-Bogota , Cristian Sánchez , Jimena Ibagon , Maamer Jlali , Pierre Cozannet , ‡ †,1 Aurélie Preynat , and Tofuko A. Woyengo † ‡ Department of Animal Science, South Dakota State University, Brookings, SD 57007; and Adisseo France S.A.S., Center of Expertise and Research in Nutrition, F-03600 Commentry, France ABSTRACT:  A study was conducted to deter- the PC and NC1 diets did not differ in average mine the effects of supplementing corn–soybean daily gain (ADG) and average daily feed intake. meal-based diets with a multienzyme on growth Pigs fed NC2 diet had lower (P  <  0.05) ADG performance, bone mineralization, apparent ileal and gain-to-feed ratio (G:F) than those fed PC digestibility (AID) and apparent total tract di- diet. Pigs fed PC diet had greater (P < 0.05) bone gestibility (ATTD) of nutrients of growing pigs. ash content and ATTD of P than those fed NC1 A total of 276 pigs (body weight [BW] = 33.99 ± diet. The ATTD of GE for PC diet was greater 4.3  kg) were housed by sex in 45 pens of 6 or 7 (P  <  0.05) than that for NC2 diet, and tended pigs and fed 5 diets (9 pens/diet) in a randomized to be greater (P  <  0.10) than that for NC1 diet. complete block design. Diets were positive control Multienzyme interacted (P < 0.05) with negative (PC); and negative control 1 (NC1) or negative control diet type on overall ADG and AID of GE control 2 (NC2) without or with multienzyme. The such that multienzyme did not affect overall ADG multienzyme used supplied at least 1,800, 1,244, and AID of GE for the NC1 diet, but increased 6,600, and 1,000 units of xylanase, β-glucanase, (P < 0.05) overall ADG and AID of GE for NC2 arabinofuranosidase, and phytase per kilogram diet by 5.09 and 8.74%, respectively. Multienzyme of diet, respectively. The PC diet was adequate in did not interact with negative control diet type all nutrients according to NRC recommendations on overall G:F, bone ash content, AID of AA, and had greater digestible P content than NC1 or and ATTD of nutrients. Multienzyme increased NC2 diet by 0.134 percentage points. The PC diet (P  <  0.05) overall G:F, AID of methionine, had greater net energy (NE) and standardized ATTD of GE and P, and tended to increase ileal digestible amino acids (AA) content than (P  =  0.056) bone ash content. The ADG, bone NC1 diet by 3%, and than NC2 diet by 5%. The ash content, and ATTD of GE and P for the mul- diets were fed in 4 phases based on BW: Phase 1: tienzyme-supplemented diets were similar to (P > 34–50 kg; Phase 2: 50–75 kg; Phase 3: 75–100 kg; 0.10) PC diet. Thus, NE and digestible AA and and Phase 4: 100–120  kg. Nutrient digestibility P can be lowered by ≤5% in multienzyme-supple- and bone mineralization were determined at the mented diets without effects on growth perform- end of Phase 1. Overall (34–120 kg BW), pigs fed ance and bone ash of pigs. Key words: bone mineralization, growth performance, multienzyme, nutrient digestibility, pig © The Author(s) 2020. Published by Oxford University Press on behalf of the American Society of Animal Science. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribu- tion, and reproduction in any medium, provided the original work is properly cited. Transl. Anim. Sci. 2020.4:1-14 doi: 10.1093/tas/txaa040 Corresponding author: tofuko.woyengo@sdstate.edu Received November 25, 2019. Accepted April 6, 2020. 1 Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Jerez-Bogota et al. INTRODUCTION effective with regard to degradation of arabinoxy- lan than supplementation with xylanase alone be- The major sources of nutrients in swine diets cause arabinofuranosidase can cleave off arabinose are plant origin feedstuffs. However, these feed- units from backbones of xylans, leading to reduced stuffs contain some antinutritional factors, which resistance of arabinoxylans to enzymatic degrad- limit nutrient utilization. Some of the important ation. Indeed, supplementation of a multienzyme antinutritional factors present in these feedstuffs product that contained xylanase, arabinofuranosi- include phytic acid and non-starch polysacchar- dase, β-glucanase, and phytase activities improved ides (NSP). Phytic acid contain P, which is poorly nutrient utilization in poultry fed wheat-corn-based digested by pigs because they do not produce suf- diets (Jlali et al., 2018, 2019). However, information ficient amounts of phytase to liberate phytic acid- is lacking on the effects of the same multienzyme bound P (Woyengo and Nyachoti, 2011). Also, product on nutrient utilization in pigs fed wheat– phytic acid reduces digestibility of other nutrients corn-based diets. Therefore, the objective of this including cations and amino acids (AA) by binding study was to determine the effects of supplemen- them (Woyengo and Nyachoti, 2013). The NSP are tation multienzyme product that contain xylanase, poorly digested by swine and can reduce nutrient arabinofuranosidase, β-glucanase, and phytase ac- availability for digestion and absorption partly by tivities on nutrient digestibility, bone mineraliza- encapsulation and viscosity (Bedford and Partridge, tion, and growth performance of grow–finish pigs 2010; Bedford and Schulze, 1998; Woyengo et  al., fed corn–wheat–wheat bran–soybean meal-based 2016). Furthermore, phytic acid and NSP can have diets that are low in net energy (NE), standardized negative effects on environment due to increased ileal digestible AA, Ca, and standardized total tract excretion of unabsorbed nutrients, especially N and digestible P. P (Woyengo et al., 2008b). The undesirable effects of phytic acid and NSP MATERIALS AND METHODS can be alleviated through dietary supplementation of phytase and NSP-degrading enzymes also known Experimental procedures were reviewed and as NSPases (Cowieson and Bedford, 2009). Corn approved by the Institutional Animal Care and and wheat grains, and their co-products are the Use Committee at South Dakota State University most widely used sources of energy in swine diets (#18-015E). in North America and Europe. The most abundant NSP in wheat and corn are arabinoxylans (Choct, Experimental Animals 1997; Knudsen, 2014). Wheat also contain some A total of 276 pigs (initial body weight [BW] of β-glucans (Choct, 1997). Arabinoxylans are com- 33.99 ± 4.3 kg; Lance-Large White female × Duroc posed of backbones of xylans that are substituted male; Pig Improvement Company) were obtained mainly with arabinose; arabinose is linked to ferulic from the Swine Education and Research Facility, acid, which crosslink xylans and lignin (Appeldoorn South Dakota State University (Brookings, SD). et al., 2010). Xylanase has been added in corn- and Pigs were then individually weighed and housed wheat-based diets for pigs with the goal of increas- in 45 pens of 6 or 7 pigs. Pens (1.8  × 2.4 m) had ing dietary nutrient utilization by degrading ara- fully slated-concrete floors, metal spindle walls (1.0 binoxylans. However, xylanase had inconsistent m high), and solid polyvinyl chloride gates. Each effects on nutrient utilization in pigs fed diets that pen was equipped with a cup drinker, and a dou- are based on corn, wheat or their co-products. For ble-space dry feeder. Room temperature was main- instance, a few studies (e.g. Ndou et  al., 2015) re- tained at 22 ± 2°C throughout the experiment. ported improved growth performance of pigs due to addition of xylanase in wheat- or corn-based diets, whereas most studies (e.g. Nortey et  al., Experimental Diets 2007; O’Shea et  al., 2014; Woyengo et  al., 2008b) Five diets based on corn, soybean meal, wheat, did not report any improvement. The cross-linking wheat bran, and soybean hulls were fed in this study. of xylans and lignin with ferulic acid can poten- The diets included a positive control diet (PC); and tially make arabinoxylans more resistant to deg- negative control diet 1 (NC1) and negative control radation by xylanase alone. Thus, supplementation diet 2 (NC2) without or with multienzyme in 2  × of pig’s diets that are based on corn, wheat, or 2 factorial arrangement (Table  1). The multien- their co-products with a combination of xylanase zyme (a multi-carbohydrase and phytase complex, and arabinofuranosidase can potentially be more Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Multienzyme supplemented diets for pigs l dl l l Translate basic science to industry innovation Table 1. Ingredient and calculated chemical composition of the basal diets (%, as-fed basis) Phase 1: 34–50 kg BW Phase 2: 50–75 kg BW Phase 3: 75–100 kg BW Phase 4: 100–135 kg BW Item PC NC1 NC2 PC NC1 NC2 PC NC1 NC2 PC NC1 NC2 Ingredients, % Corn 55.734 54.033 51.223 60.069 58.059 55.289 61.402 59.623 56.271 65.909 60.757 57.812 Soybean meal 15.211 14.260 13.465 10.351 9.544 9.040 8.814 7.900 7.108 4.953 3.437 2.505 Wheat 10.000 10.000 10.000 10.000 10.000 10.000 10.000 10.000 10.000 10.000 10.000 10.000 Wheat bran 10.000 11.783 15.000 10.000 12.000 14.267 10.000 11.700 15.339 10.000 13.600 16.000 Soybean hulls 2.500 4.500 5.000 3.545 5.471 6.500 4.207 6.289 6.808 4.702 8.438 9.995 Soybean oil 3.225 2.800 2.713 3.000 2.600 2.600 3.000 2.600 2.600 2.300 2.300 2.234 Calcium carbonate 1.083 1.040 1.047 0.998 0.955 0.960 0.875 0.833 0.840 0.689 0.561 0.543 Monocalcium phosphate 1.063 0.415 0.397 0.886 0.236 0.224 0.744 0.096 0.076 0.540 - - -Lysine HCl 0.488 0.480 0.473 0.479 0.470 0.462 0.380 0.380 0.380 0.351 0.349 0.350 -Methionine 0.096 0.090 0.086 0.071 0.066 0.062 0.025 0.023 0.021 0.0002 0.0004 0.0004 -Thronine 0.124 0.121 0.121 0.114 0.111 0.111 0.083 0.085 0.087 0.090 0.094 0.096 -Tryptophan 0.043 0.041 0.039 0.050 0.047 0.044 0.031 0.030 0.027 0.029 0.028 0.027 Mineral premix 0.150 0.150 0.150 0.150 0.150 0.150 0.150 0.150 0.150 0.150 0.150 0.150 Vitamin premix 0.050 0.050 0.050 0.050 0.050 0.050 0.050 0.050 0.050 0.050 0.050 0.050 Salt 0.234 0.236 0.237 0.238 0.240 0.240 0.239 0.241 0.242 0.237 0.237 0.237 Calculated nutrients Crude protein, % 14.735 14.567 14.467 12.943 12.839 12.781 12.249 12.098 12.029 11.108 10.985 10.868 Ether extract, % 5.934 5.493 5.375 5.787 5.367 5.319 5.810 5.390 5.348 5.394 5.371 5.292 NE, kcal/kg 2,475 2,401 2,351 2,475 2,401 2,351 2,475 2,401 2,351 2,475 2,401 2,351 Standardized digestible content of AA, % Lys 0.980 0.951 0.931 0.847 0.822 0.805 0.730 0.708 0.694 0.610 0.592 0.580 Met 0.316 0.305 0.296 0.269 0.259 0.251 0.216 0.208 0.202 0.165 0.158 0.154 Met + Cys 0.550 0.534 0.523 0.480 0.466 0.456 0.420 0.407 0.399 0.360 0.349 0.342 Thr 0.590 0.572 0.560 0.509 0.493 0.484 0.456 0.442 0.433 0.400 0.388 0.380 Trp 0.170 0.165 0.162 0.153 0.148 0.145 0.127 0.123 0.120 0.110 0.107 0.105 Total P, % 0.596 0.470 0.487 0.542 0.417 0.428 0.506 0.380 0.398 0.459 0.359 0.371 Digestible P 0.310 0.176 0.176 0.269 0.135 0.135 0.238 0.104 0.104 0.210 0.092 0.093 Calcium 0.660 0.540 0.540 0.590 0.470 0.47 0.520 0.400 0.400 0.460 0.340 0.340 Sodium 0.100 0.100 0.100 0.100 0.100 0.100 0.100 0.100 0.100 1.000 1.000 1.000 BW, body weight; PC, positive control diet; NC1, negative control diet 1 with lower in NE, standardized ileal digestible AA, standardized total tract digestible P, and Ca than PC diet by 3, 3, 43, and 18%, respectively; and NC2, negative control diet 2 with lower in NE, standardized ileal digestible AA, standardized total tract digestible P, and Ca than PC diet by 5, 5, 43, and 18%, respectively. Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Jerez-Bogota et al. Rovabio Advance Phy, Adisseo France S.A.S) was Sample Preparation and Analyses added to diets supplying at least 1,800, 1,244, 6,600, Femurs for determining bone ash were defleshed and 1,000 units of xylanase, β-glucanase, arabino- by autoclaving at 121°C for 30  min, cleaned and furanosidase, and phytase per kilogram of diet, re- subsequently dried in an oven at 135°C for 2 h. Fat spectively. Enzyme supplementation levels were as was extracted from the dried bones using petroleum per the supplier’s recommendations. The PC diet ether (E139-4, Fischer Scientific, Pittsburgh, PA) as was formulated to be adequate in all nutrients ac- solvent in a Jumbo Soxhlet extraction apparatus cording to NRC (2012) recommendations. The (Chemglass Life Sciences, Vineland, NJ), afterward NC1 diet was the same as the PC diet except that its the samples were left in a fume hood for 24  h to NE and standardized ileal digestible AA, standard- allow the petroleum ether to evaporate. Femurs were ized total digestible P, and total Ca contents were then dried in an oven at 135°C for 2 h to determine lower than those for the PC diet by 3.0, 3.0, 43, and their fat-free weight, and ashed at 600°C in a muffle 18%, respectively. The NC2 diet was the same as the furnace for 12 h for the determination of bone ash. PC diet except that its NE and standardized ileal Femurs for determining bone breaking strength digestible AA, standardized total digestible P, and were defleshed by scraping muscle tissues from the total Ca contents were lower than those for the PC bones using kitchen knives. Maximal breaking load diet by or 5.0, 5.0, 43, and 18%, respectively. The was measured using an MTS Insight 5 equipment reduction in NE value and nutrient content in the (MTS, Eden Prairie, MN, USA) at room tempera- NC1 and NC2 diets was achieved by a partial re- ture by subjecting each bone to a 3-point bending placement of corn, soybean meal, soybean oil, crys- test (Tunrer and Burr, 1993). Force was applied talline AA, calcium carbonate, and monocalcium to the center of the bone held by supports 3.3 cm phosphate in PC diet with wheat bran and soybean apart. The crosshead speed was set at 50  mm/min hulls. The diets were fed in mash form and in 4 and the sample rate was 10 points/s. Final strength phases based on BW: Phase 1: 34–50  kg, Phase 2: was determined from load–displacement curves. 50–75  kg, Phase 3: 75–100  kg, and Phase 4: 100– Fecal samples were pooled by pen and air-dried 125 kg. Titanium dioxide (0.3%) was added as indi- in an oven at 60°C for 4 d; whereas ileal digesta gestible marker in each diet during the last week of samples were freeze–dried. The dried fecal and the first phase of feeding. ileal digesta samples together with diet samples were ground through a 0.75-mm screen in a cen- trifugal mill (model ZM200; Retsch GmbH, Haan, Experimental Design and Procedure Germany). The ground samples were analyzed as The five diets were allotted to the 45 pens (9 pens/ follows: Phase 1 diets for DM, gross energy (GE), diet) within a randomized complete block design. N (N × 6.25  =  CP), P, AA, NDF, ADF and ti- Diets and fresh water were offered to pigs ad libitum tanium contents, and for xylanase and phytase during the entire period. Pig BW and feed intake were activities; Phases 2–4 diets for DM, GE, and N; determined by phase to calculate average daily gain ileal digesta for DM, GE, NDF, ADF, AA, P and (ADG), average daily feed intake (ADFI), and gain- titanium contents; and feces for GE, CP, NDF, to-feed ratio (G:F). Fresh fecal samples were collected ADF, Ca, P, and titanium contents. The samples from each pen during the last 2 days of first feeding were analyzed for DM by oven drying at 135°C phase and immediately stored frozen at −20°C for for 2 h (method 930.15), CP by a combustion pro- the determination of apparent total tract digestibility cedure (method 990.03), as per AOAC (2012); and (ATTD) of energy and nutrients. At the end of the first for ADF and NDF (Van Soest et al., 1991) on an phase of feeding, 1 pig (per pen) with BW that was Ankom 200 Fiber Analyzer (Ankom Technology, close to the pen average BW was selected, and then eu- Fairport, NY). Samples were analyzed for AA thanized by captive bolt penetration followed by ex- (method 982.30 E [a, b, and c]; AOAC, 2012) at sanguination. Right and left femurs were excised from the University of Missouri Experiment Station la- each euthanized pig and stored at −20°C for deter- boratories (Columbia, MO). The GE was analyzed mination of bone ash and bone breaking strength, re- using an adiabatic bomb calorimeter (model 1261, spectively. Also, contents of lower half of ileum (from Parr Instrument Co., Moline, IL). Titanium dioxide 80  cm above ileal-cecal junction to approximately in samples was determined by spectrophotometry 1 cm above the ileo-cecal junction) were obtained and (model Spectra MAX 190, Molecular Devices, stored frozen at −20°C for latter determination of ap- Sunnyvale, CA) at 408 nm after ashing at 525°C for parent ileal digestibility (AID) of energy and nutrients. 10 h (Myers et al., 2004). The P content in samples Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Multienzyme supplemented diets for pigs was analyzed according to the vanadate colori- RESULTS metric method (method 946.06; AOAC, 2012) using The analyzed CP values in the diets in Tables 2 a spectrophotometer (model Spectra MAX 190, and 3 were similar to the calculated CP values in Molecular Devices, Sunnyvale, CA). The Ca con- the diets in Table 1. The mean endogenous phytase tent in samples was analyzed by inductively coupled activity in PC, NC1, and NC2 diets was 406 U/kg. plasma–optical emission spectrometry (ICP–OES; Addition of phytase to the NC1 and NC2 diets re- method 985.01 A, B, and C; AOAC, 2012) after sulted in its increased activity values by margins wet ashing samples (method 975.03 B(b); AOAC, similar to those that were anticipated. The xylanase 2012). Xylanase and phytase activity in experi- activity was undetectable in the PC, NC1, and NC2 mental diets were analyzed by the Laboratory of diets. The analyzed xylanase activity values in mul- Adisseo (Commentry, France). Xylanase activity tienzyme supplemented NC1 and NC2 diets were was determined according the method described by similar to the anticipated values. Cozannet et  al. (2017). One visco-unit of endo-1, Data on effects of dietary treatment on growth 4-β-xylanase activity is defined as the amount of performance, bone mineralization, AID of energy enzyme that hydrolyzed by the substrate (wheat and nutrients, and ATTD of energy and nutrients AX); one such unit reduces solution’s viscosity, re- are presented in Tables  4–7, respectively. Pigs fed sulting in a change in relative fluidity of 1 arbitrary PC diet had greater (P < 0.05) BW than those fed unit per min per ml (or per g) under the conditions NC1 diet during Phases 1 and 2 of feeding, but not of the assay and pH 5.5 and 30°C. Phytase activity during Phases 3 and 4. The BW of pigs fed PC diet in experimental diets was determined according the was greater (P < 0.05) than that of pigs fed NC2 diet standard method (ISO3024, 2009). One phytase during overall (34–120  kg BW) study period. Pigs unit (FTU) is the amount of enzyme that releases fed PC diet had greater (P < 0.05) ADG than those fed NC1 diet during Phase 1, but not during Phases 1 µmol of inorganic orthophosphate from sodium 2–4 of feeding. Also, the overall ADG of pigs fed phytate substrate per minute at pH 5.5 and 37°C. PC diet did not differ from that of pigs fed NC1 diet. Pigs fed PC diet had greater (P < 0.05) ADG Calculations and Statistical Analysis than those fed NC2 diet during the Phases 1, 2, and 4. Also, the overall ADG of pigs fed PC was greater The AID and ATTD values of the diets were (P < 0.05) than that of pigs fed NC2 diet. Pigs fed calculated using the indicator method (Stein et al., NC1 diet had greater (P < 0.05) ADFI than those 2007), using the following equation: fed PC diet during Phases 1 and 3 of feeding, but AID or ATTD, % =[1 − (Nutrient /Nutrient ), digesta diet not during Phases 2 and 4; whereas pigs fed NC2 ×(Marker /Marker )] × 100 diet digesta diet had greater (P < 0.05) ADFI than those fed PC diet during the Phase 3, but not during the Phases where Nutrient is the nutrient concentration digesta 1, 2, and 4. Pigs fed NC1 diet had lower (P < 0.05) in the ileal digesta or feces (%  DM); Nutrient diet G:F than those fed PC diet during the Phase 1 of is the nutrient concentration in the diet (%  DM); feeding, but not during the Phases 2–4 and during Marker is the titanium concentration in the diet diet the overall period. The G:F for NC2 diet was lower (%  DM); and Marker is the titanium concen- digesta (P < 0.05) than that for the PC diet during the entire tration in the ileal digesta or feces (% DM). study period. The percent femur ash content for PC Data were subjected to analysis of variance diet was greater (P < 0.05) than that for NC1 diet using the MIXED procedure of SAS (SAS Inst. or NC2 diet. The femur breaking strength for PC Inc., Cary, NC). The pen was considered as the diet was numerically, but not significantly greater experimental unit. The model included diet, sex, than that for NC1 diet or NC2 diet. The PC, NC1, diet × sex interaction, and initial BW, which was and NC2 diets did not differ in AID of GE, CP and a covariate. Period was the repeated term in mod- indispensable AA. The ATTD of DM and GE for els involving time. Means were separated by the NC1 diet tended to be lower (P  <  0.10) than that probability of difference in order to compare PC for the PC diet, whereas the ATTD of CP and P diet with other diets. Main effects of NC diet type for NC1 diet was lower (P < 0.05) than that for PC and multienzyme and their interactions were deter- diet. The ATTD of DM, GE, and P for the NC2 mined. To test the hypotheses, P  <  0.05 was con- diet was lower (P < 0.05) than that for PC diet. sidered significant. If pertinent, trends (0.05 < P ≤ The NC diet type and multienzyme did 0.10) are also reported. not interact on BW during Phase 1, 2, and 3 of Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Jerez-Bogota et al. Table 2. Analyzed composition of phase 1 diets as-fed Diets Item PC NC1 NC2 NC1 + E NC2 + E Dry matter, % 88.5 88.9 89.3 89.1 89.3 Gross energy, kcal/kg 3,973 4,052 4,067 3,990 4,053 Crude protein, % 14.38 14.54 14.81 14.04 14.00 Neutral detergent fiber, % 12.88 15.26 18.25 14.21 17.85 Acid detergent fiber, % 4.50 6.48 7.10 6.00 6.72 Ca, % 0.66 0.52 0.40 0.53 0.37 P, % 0.60 0.51 0.54 0.53 0.50 Indispensable AA, % Arg 0.80 0.87 0.83 0.78 0.79 His 0.35 0.38 0.37 0.35 0.35 Ile 0.53 0.56 0.54 0.53 0.51 Leu 1.15 1.19 1.11 1.12 1.04 Lys 0.93 0.97 0.97 0.94 0.93 Met 0.32 0.29 0.27 0.32 0.29 Phe 0.63 0.66 0.63 0.62 0.6 Thr 0.59 0.62 0.59 0.59 0.6 Trp 0.22 0.23 0.22 0.2 0.21 Val 0.63 0.67 0.66 0.63 0.62 Dispensable AA, % Ala 0.68 0.72 0.68 0.67 0.64 Asp 1.17 1.25 1.18 1.16 1.09 Cys 0.27 0.27 0.27 0.26 0.27 Glu 2.53 2.61 2.53 2.42 2.51 Gly 0.58 0.64 0.64 0.59 0.62 Pro 0.91 0.92 0.9 0.88 0.88 Ser 0.59 0.63 0.58 0.57 0.56 Tyr 0.48 0.49 0.45 0.45 0.44 Analyzed enzyme activities, U/kg Phytase 350 359 389 1,400 1,391 Xylanase 0 0 0 1,834 1,850 PC, positive control diet; NC1, negative control diet 1; NC2, negative control diet 2; NC1 + E, negative control diet 1 plus multienzyme; and NC2 + E, negative control diet 2 plus multienzyme. feeding. Multienzyme supplementation improved NC1 diet with multienzyme reduced (P < 0.05) the (P  <  0.05) BW during phase 1–3 regardless of ADG, whereas supplementation of the NC2 diet NC diet type. The NC diet type and multienzyme tended to increase (P < 0.10) the ADG of pigs. The tended to interact (P = 0.066) on final BW (Phase NC type and multienzyme interacted (P  =  0.046) 4)  such that supplementation of NC1 diet with on the overall ADG such that supplementation multienzyme did not affect the final BW, whereas of NC1 diet with multienzyme did not affect the supplementation of the NC2 diet increased ADG, whereas supplementation of the NC2 diet (P  <  0.05) the final BW of pigs. Multienzyme increased (P < 0.05) the ADG of pigs. The NC diet and NC diet type did not interact on ADG dur- type and multienzyme interacted (P  =  0.024) on ing Phase 1.  However, multienzyme supplemen- the ADFI during the Phase 1 of feeding such that tation improved (P  <  0.05) ADG during phase the ADFI for NC1 diet was reduced (P  <  0.05), 1 regardless of NC diet type. The NC type and whereas that of the NC2 diet was unaffected by multienzyme tended to interact (P  =  0.070) on the multienzyme. Multienzyme supplementation ADG during Phase 2 such that supplementation reduced (P = 0.002) ADFI during Phase 3 regard- of NC1 diet with multienzyme did not affect the less of NC diet type. The NC diet type and mul- ADG, whereas supplementation of the NC2 diet tienzyme interacted (P  =  0.013) on ADFI during increased (P < 0.05) the ADG of pigs. The NC diet Phase 4 such that supplementation of NC1 diet re- type and multienzyme interacted (P  =  0.003) on duced (P < 0.05) the ADFI, whereas the ADFI for ADG during Phase 4 such that supplementation of NC2 diet was unaffected. The NC diet type and Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Multienzyme supplemented diets for pigs Table 3. Analyzed composition of phases 2–4 diets as-fed Diets Item PC NC1 NC2 NC1 + E NC2 + E Phase 2 Dry matter, % 86.4 90.8 87.9 87.8 88.1 Gross energy, kcal/kg 3,917 3,947 3,920 3,850 4,007 Crude protein, % 12.37 12.63 12.47 12.57 12.45 Neutral detergent fiber, % 11.13 13.02 13.29 13.23 14.75 Acid detergent fiber, % 4.57 6.12 5.67 4.96 6.32 Analyzed enzyme activities, U/kg Phytase 372 394 440 1,410 1,500 Xylanase 0 0 0 1,840 1,865 Phase 3 Dry matter, % 88.5 88.4 88.3 88.7 88.2 Gross energy, kcal/kg 3,913 4,101 3,996 3,931 3,968 Crude protein, % 12.03 11.92 12.20 11.64 12.29 Neutral detergent fiber, % 12.76 16.23 15.97 13.71 15.91 Acid detergent fiber, % 5.53 5.64 7.31 5.64 5.78 Analyzed enzyme activities, U/kg Phytase 374 366 462 1,441 1,390 Xylanase 0 0 0 1,860 1,830 Phase 4 Dry matter, % 88.5 88.4 88.3 88.7 88.2 Gross energy, kcal/kg 3,918 3,910 3,966 3,915 3,924 Crude protein, % 10.11 10.06 10.11 9.85 9.83 Neutral detergent fiber, % 12.27 14.94 16.04 16.44 17.79 Acid detergent fiber, % 4.62 6.45 7.7 6.51 8.22 Analyzed enzyme activities, U/kg Phytase 451 426 485 1,500 1,440 Xylanase 0 0 0 1,835 1,860 PC, positive control diet; NC1, negative control diet 1; NC2, negative control diet 2; NC1 + E, negative control diet 1 plus multienzyme; and NC2 + E, negative control diet 2 plus multienzyme. multienzyme interacted (P = 0.045) on the overall CP such that multienzyme supplementation tended ADFI such that the ADFI for NC1 diet, but not to increase (P < 0.10) the AID of CP for NC1 diet, of NC2 diet, was reduced (P < 0.05) by the supple- but increased (P < 0.05) AID of CP for NC2 diet. mentation. Multienzyme and NC diet type inter- Multienzyme supplementation did not affect AID acted (P = 0.002) on G:F during Phase 1 such that of all indispensable AA except of Met whose AID the supplementation increased (P < 0.05) G:F for was increased (P = 0.006) by the supplementation NC1 diet and tended to increase (P  <  0.10) G:F regardless of the NC diet type. No interactions for NC2 diet. Also, multienzyme and NC diet type were detected between NC diet type and multien- tended to interact (P = 0.084) on G:F during Phase zyme on ATTD of DM, OM, GE, CP, NDF, ADF, 4 such that the multienzyme supplementation did and P.  Multienzyme supplementation increased not affect the G:F for NC1, but tended to increase (P < 0.05) ATTD of DM, OM, GE, CP, Ca, and P. (P < 0.10) G:F for NC2 diet. Multienzyme and NC diet type did not interact on overall G:F; however, DISCUSSION the multienzyme increased (P < 0.001) overall G:F regardless of NC diet type. Multienzyme supple- The ADG of pigs fed the NC1 diet was lower mentation tended to increase (P  =  0.056) percent than that of pigs fed the PC diet during the first femur ash content and increased (P < 0.001) bone phase of feeding, implying that the reduction in breaking regardless of NC diet type. Multienzyme NE, standardized ileal digestible AA, standardized and NC diet type interacted (P = 0.007) on AID of total tract digestible P, and total Ca contents in PC GE such that multienzyme did not affect the AID diet by 74 kcal/kg, 3%, 43%, and 18%, respectively, of GE for NC1 diet and increased (P < 0.05) AID was sufficient to reduce the growth performance of of GE for NC2 diet. Also, multienzyme and NC the pigs weighing between 34 and 55 kg. However, diet type tended to interact (P = 0.10) on AID of the ADG of pigs fed the NC1 diet did not differ Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Jerez-Bogota et al. Translate basic science to industry innovation Table 4. Effect of dietary treatments on growth performance a b Diets P-value Item PC NC1 NC2 NC1 + E NC2 + E SEM Diet NC E NC × E BW, kg Day 0 34.61 34.60 34.60 34.67 34.54 - - a b b a a Phase 1 56.14 53.83 54.54 56.03 55.89 0.472 0.004 0.341 <0.001 0.305 a b b a a Phase 2 75.13 72.77 71.33 74.81 75.13 0.670 0.001 0.754 <0.001 0.341 a ab b a ab Phase 3 101.53 99.68 98.12 101.27 100.41 1.000 0.092 0.264 0.021 0.883 a a b a a Phase 4 124.45 123.87 118.90 124.14 123.26 1.260 0.009 0.038 0.040 0.066 ADG, kg a b b a a Phase 1 0.925 0.821 0.851 0.912 0.912 0.021 0.004 0.301 <0.001 0.425 a ab b a a Phase 2 0.981 0.936 0.884 0.948 0.980 0.021 0.018 0.727 0.008 0.070 Phase 3 0.990 1.004 0.962 0.983 0.938 0.024 0.279 0.045 0.317 0.898 ab a c bc bc Phase 4 0.998 1.064 0.902 0.954 0.970 0.031 0.009 0.032 0.416 0.003 a a b a a Overall 0.966 0.955 0.899 0.955 0.949 0.013 0.008 0.022 0.164 0.046 ADFI, kg a b a a a Phase 1 1.908 2.312 1.850 1.896 1.929 0.038 <0.001 0.050 0.181 0.024 Phase 2 2.448 2.359 2.369 2.410 2.491 0.047 0.231 0.211 0.071 0.404 bc a ab bc c Phase 3 2.669 2.796 2.819 2.647 2.621 2.669 0.008 0.989 0.002 0.745 ab a ab b ab Phase 4 3.255 3.267 3.076 3.022 3.135 0.080 0.150 0.724 0.127 0.013 b a b b b Overall 2.57 2.70 2.53 2.48 2.54 0.04 0.003 0.353 0.096 0.045 G:F, kg/kg a c b ab ab Phase 1 0.485 0.367 0.463 0.480 0.474 0.007 <0.001 0.004 <0.001 0.002 a ab b ab ab Phase 2 0.401 0.397 0.377 0.394 0.394 0.007 0.099 0.151 0.237 0.321 a ab b a ab Phase 3 0.372 0.359 0.343 0.373 0.358 0.009 0.082 0.046 0.059 0.900 ab a b ab ab Phase 4 0.309 0.324 0.295 0.317 0.310 0.009 0.155 0.015 0.560 0.084 a c b ab ab Overall 0.485 0.367 0.463 0.480 0.474 0.007 <0.001 0.718 <0.001 0.130 PC, positive control diet; NC1, negative control diet 1; NC2, negative control diet 2; NC1 + E, negative control diet 1 plus multienzyme; and NC2 + E, negative control diet 2 plus multienzyme. NC, main effects of negative control diet type; E, main effects of multienzyme; and NC × E, interaction between negative control diet type and multienzyme. abc Within a row, means without a common superscript differ (P < 0.05). Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Multienzyme supplemented diets for pigs from that of pigs fed the PC diet during the second, third and fourth phases of feeding, leading to lack of differences between NC1 and PC diets with re- gard to the overall ADG and final BW. Such results indicate that the reduction in NE, standardized ileal digestible AA, standardized total tract digestible P, and total Ca contents in PC diet by 74 kcal/kg, 3%, 43%, and 18%, respectively, during the phases of feeding 2, 3, and 4 was not sufficient to reduce the growth performance of the pigs, and could be at- tributed to an increase in digestive capacity of pigs with age. Similarly, Woyengo et al. (2008b) did not observe reduction in ADG of grow–finish pigs fed wheat-based diets due to reduction in DE value and available P contents in PC diet by 75 kcal/kg and 30.4%, respectively. Also, Li et  al. (2012) did not observe reduction in ADG of pigs due to reduction in DE value by 50 kcal/kg. The ADG of pigs fed the NC2 diet was lower than that of pigs fed the PC diet during the first, second, and fourth phases of feeding. Also, the overall ADG and hence final BW of pigs fed the NC2 diet were lower than those of pigs fed the PC diet, implying that the reduction in NE, standardized ileal digestible AA, standard- ized total tract digestible P, and total Ca contents in PC diet by 124 kcal/kg, 5%, 43%, and 18%, respect- ively, was sufficient to reduce growth performance of the pigs. Emiola et al. (2009) observed reduction in ADG of growing pigs due to reduction in dietary DE by 124 kcal/kg. Jang et al. (2017) also served re- duction in ADG of growing pigs due to reduction in dietary ME by 103 kcal/kg. Supplementation of NC1 diet or NC2 diet with multienzyme increased ADG of pigs during the first phase (from 34 to 55  kg BW) of feeding, which was due to increased nutrient digestibility by the supplementation. This result is in accordance with several studies reporting an improvement of ADG in growing pigs fed diets supplemented with a multi-carbohydrase (Emiola et  al., 2009; Kiarie et  al., 2012; Ndou et  al., 2015). Supplementation of NC2 diet with multienzyme increased ADG of pigs during the second phase of feeding (50–75 kg BW), and hence during the entire study period (34– 125  kg BW), whereas it is not the case with NC1 diet. It should be noted that the NC2 diet was for- mulated to contain less NE and digestible AA than NC1 diet; and that the addition of multienzyme to NC2 diet resulted in an increase in AID of GE and CP, whereas the addition of multienzyme to NC1 diet only tended to increase in AID of CP. Also, the magnitude by multienzyme increased the ATTD of P for NC2 diet was greater than the magnitude by which it increased the ATTD of P for NC1 diet Translate basic science to industry innovation Table 5. Effect of dietary treatment on bone mineralization a b Diets P-value Item SEM PC NC1 NC2 NC1 + E NC2 + E Diet NC E NC × E ab c bc a ab Femur ash, g 32.67 28.90 29.78 35.97 34.27 1.814 0.049 0.353 <0.001 0.118 a c bc ab a Femur ash, % 55.32 51.99 52.75 54.33 55.15 0.660 0.008 0.619 0.056 0.735 bc c c a ab Femur breaking strength, N 2,513 2,191 2,278 2,950 2,672 129.3 0.002 0.385 <0.001 0.177 PC, positive control diet; NC1, negative control diet 1; NC2, negative control diet 2; NC1 + E, negative control diet 1 plus multienzyme; and NC2 + E, negative control diet 2 plus multienzyme. NC, main effects of negative control diet type; E, main effects of multienzyme; and NC × E, interaction between negative control diet type and multienzyme. abc Within a row, means without a common superscript differ (P < 0.05). Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Jerez-Bogota et al. Translate basic science to industry innovation Table 6. Effect of dietary treatment on AID of energy and nutrients a b Diets P-value AID, % PC NC1 NC2 NC1 + E NC2 + E SEM Diet NC E NC × E ab b b b a Dry matter 67.73 66.48 66.21 65.16 71.73 1.51 0.060 0.053 0.130 0.028 ab bc abc c a Gross energy 71.12 67.56 67.53 64.66 73.43 2.3 0.028 0.065 0.206 0.007 bc bc c ab a Crude protein 65.21 64.31 62.43 69.53 75.14 2.22 0.005 0.334 0.001 0.100 Indispensable AA Arg 80.01 81.14 81.51 81.47 84.60 1.47 0.264 0.105 0.306 0.299 His 74.47 77.25 76.69 75.63 79.34 1.74 0.319 0.351 0.818 0.208 Ile 71.98 73.76 73.72 74.15 77.59 2.23 0.472 0.433 0.403 0.424 Leu 75.39 75.88 75.50 76.90 78.99 1.87 0.622 0.594 0.307 0.482 Lys 77.28 78.36 77.99 79.47 82.50 2.08 0.404 0.627 0.188 0.462 ab b b ab a Met 84.18 82.11 82.01 85.54 86.42 1.26 0.048 0.478 0.006 0.454 Phe 73.63 74.69 74.30 75.71 78.81 2.01 0.378 0.333 0.252 0.272 Thr 65.37 67.04 66.40 67.24 73.81 2.97 0.232 0.129 0.333 0.120 Trp 76.53 77.06 76.68 77.11 80.25 2.45 0.770 0.627 0.457 0.484 Val 64.34 66.83 66.48 66.85 71.75 2.80 0.385 0.235 0.484 0.208 Dispensable AA Ala 67.05 68.86 67.49 68.98 72.71 2.42 0.482 0.596 0.329 0.290 Asp 70.83 72.54 71.41 72.27 75.19 2.51 0.764 0.885 0.448 0.498 Cys 64.17 63.10 64.24 63.50 69.93 3.16 0.519 0.363 0.327 0.483 Glu 80.07 80.87 81.87 82.11 85.21 1.71 0.277 0.527 0.166 0.543 Gly 36.06 47.25 54.26 54.35 58.48 6.58 0.102 0.422 0.366 0.779 Pro 76.35 72.49 76.06 77.10 79.57 2.22 0.321 0.143 0.173 0.299 Ser 69.08 70.77 69.48 70.74 74.98 2.54 0.461 0.500 0.370 0.250 Tyr 75.98 75.83 74.66 75.25 79.17 1.96 0.525 0.591 0.355 0.270 PC, positive control diet; NC1, negative control diet 1; NC2, negative control diet 2; NC1 + E, negative control diet 1 plus multienzyme; and NC2 + E, negative control diet 2 plus multienzyme. NC, main effects of negative control diet type; E, main effects of multienzyme; and NC × E, interaction between negative control diet type and multienzyme. abc Within a row, means without a common superscript differ (P < 0.05). Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Multienzyme supplemented diets for pigs (21.1 vs. 12.5 percentage points). Additionally, overall ADFI for NC1 diet without multienzyme was greater than that for NC1 diet with multien- zyme, whereas the overall ADFI for NC2 diet without multienzyme did not differ from that for NC1 diet with multienzyme. Thus, the greater ef- fect of multienzyme on the NC2 diet than on NC1 diet with regard to ADG could be attributed to: 1) the fact that the NC1 diet was not so deficient in NE and digestible nutrient content as evidenced by the similarity between NC1 and PC diets in ADG of pigs, 2)  greater increase in energy and nutrient digestibility for NC2 diet than for NC1 diet due to the supplemental multienzyme, and 3)  greater ADFI and hence energy and nutrient intake for unsupplemented NC1 diet than for multienzyme supplemented NC1 diet. The results from the cur- rent study are in contrast to those from the study of Kiarie et  al. (2012) who observed increased ADG of growing pigs from 22 to 55 kg BW, but not from 55 to 90 kg BW and hence for entire study period (22–90  kg BW) due to addition of a fiber-degrad- ing enzyme product to corn–barley-based diet that had lower DE value than the recommended value by large magnitude (293 kcal/kg). Also, the results from the current study are in contrast to those from the study of Jang et al. (2017) who did not observe an increased ADG of growing pigs from 26 to 122 kg BW due to addition of fiber-degrading en- zyme product to corn-based diet that had lower ME value than the recommended value by large magni- tude (103 kcal/kg). However, the enzyme product used in the current study contained xylanase, β-glu- canase, arabinofuranosidase, and phytase activities; whereas the enzyme product used in the studies of Kiarie et al, (2012) and Jang et al. (2017) contained only xylanase and β-glucanase activities, and xyla- nase activity, respectively. Thus, the differences be- tween the current study and that of Kiarie et  al. (2012) and Jang et al. (2017) with regard to the ef- fects of supplemental multienzyme on growth per- formance of pigs could be attributed to differences in enzyme activities of the multienzyme products. As previously mentioned, arabinofuranosidase can de-branch arabinoxylans, leading to increased availability of xylan backbones for xylanase deg- radation. Also, phytase can hydrolyze phytic acid, leading to release of phytic acid-bound nutrients for digestion and absorption (Woyengo and Nyachoti, 2011). Multienzyme supplementation increased overall ADG and hence final BW for NC2 diet to that of the PC diet. Thus, the NE, standardized ileal digestible AA, total Ca, and standardized total tract digestible P contents in corn–wheat–wheat Translate basic science to industry innovation Table 7. Effect of dietary treatment on ATTD of energy and nutrients a b Diets P-value ATTD, % PC NC1 NC2 NC1 + E NC2 + E SEM Diet NC E NC × E ab b b ab a Dry matter 75.62 73.79 73.90 75.76 76.43 0.703 0.057 0.907 0.006 0.591 a ab b a a Gross energy 76.05 74.32 73.65 75.81 75.67 1.252 0.077 0.650 0.027 0.658 a b ab a a Crude protein 66.21 62.70 65.57 66.53 67.13 1.022 0.046 0.253 0.019 0.442 Neutral detergent fiber 69.91 71.57 74.61 70.47 75.52 1.074 0.002 <0.001 0.735 0.302 c bc b c a Acid detergent fiber 40.18 43.18 48.90 43.62 53.91 1.835 <0.001 <0.001 0.103 0.120 bc cd d b e Ca 54.06 50.36 46.82 55.20 59.51 1.381 <0.001 0.727 <0.001 0.010 b c c b b P 38.19 22.60 27.51 35.05 48.56 2.733 <0.001 0.003 <0.001 0.181 PC, positive control diet; NC1, negative control diet 1; NC2, negative control diet 2; NC1 + E, negative control diet 1 plus multienzyme; and NC2 + E, negative control diet 2 plus multienzyme. NC, main effects of negative control diet type; E, main effects of multienzyme; and NC × E, interaction between negative control diet type and multienzyme. abc Within a row, means without a common superscript differ (P < 0.05). Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Jerez-Bogota et al. bran-based diets for pigs can be reduced by 124 kcal/ diet or NC2 diet, and the reason for this similarity kg, 5%, 18%, and 43%, respectively, without signifi- is not clear because the NC1 and NC2 diets were cant effects on growth performance of grow-finish formulated to contain lower levels of digestible pigs if the resulting low energy and nutrient diet is AA than the PC diet. It could have been due to the supplemented with the multienzyme that contain method that was used for ileal digesta collection. xylanase, β-glucanase, arabinofuranosidase, and Multienzyme supplementation increased the AID phytase activities. Supplementation of the NC1 of CP and methionine, and ATTD of GE and P for diet with multienzyme increased overall G:F, which the NC1 and NC2 diets, which was due to hydrolysis was due to the reduction in ADFI by the supple- of fiber by the NSPases and of phytate by phytase; mentation. Supplementation of the NC2 diet with as previously mentioned both NSPases and phytase multienzyme increased overall G:F, which was due were present in the multienzyme product used in to the increase in ADG by the supplementation. the current study. In plant feedstuffs, phytate, and Reason for the greater overall ADFI for NC1 diet nutrients such as starch and AA are located within without multienzyme than for NC1 diet with mul- cells, whereas NSP are mainly found in cell walls tienzyme is not clear. (Woyengo and Nyachoti, 2011). Thus, NSPases can The femur ash content and breaking strength hydrolyze NSP in cell walls to increase the accessi- for the PC diet significantly differ from those for bility of phytase to phytate that is located within NC1 diet or NC2 diet, which was partly due to the cells, thereby increasing the digestibility of phytate greater ATTD of P for the PC diet than for NC1 diet and availability of phytate-bound nutrients for di- or NC2 diet. Supplementation of the NC1 diet or gestion by gastric, pancreatic and small intestinal NC2 diet with multienzyme resulted in an increase mucosa enzymes. Also, the hydrolysis NSP by in femur ash content and breaking strength, which NSPases can result in increased availability of the could partly have been due to increase in P digest- NSP-encapsulated nutrients for digestion by gas- ibility by the supplementation. Similarly, She et al. tric, pancreatic, and small intestinal mucosa en- (2017) observed increased bone ash of weaned pigs zymes. Jang et  al. (2017) also reported increased due to dietary phytase supplementation. Woyengo ATTD of DM and P due to supplementation of a et al. (2008a, 2010) also reported increased bone ash combination of phytase and xylanase to corn-based due to addition of phytase to diets of broilers. The diet for grow–finish pigs. Woyengo et al. (2010) re - femur ash content and breaking strength for the for ported increased digestibility of P and bone ash multienzyme supplemented NC1 diet or multien- content of broilers due to addition of multienzyme zyme supplemented NC2 diet did not differ from that contained xylanase, β-glucanase, cellulose, and those for the PC diet, implying the NE, standard- pectinase to phytase-supplemented diet. However, ized ileal digestible AA, Ca, and standardized total the magnitude of improvement in AID and ATTD tract digestible P contents of the corn–wheat-based of nutrients for NC2 diet was generally greater than can be reduced by 124 kcal/kg, 5%, 18%, and 43%, that for NC1 diet. The NC2 diet was formulated to respectively, without significant effect on bone min- contain less NE and digestible AA than NC1 diet eralization if the resulting low energy and nutrient by partial replacement of corn and soybean meal in diet is supplemented with the enzyme product used NC1 diet with wheat bran and soybean hulls. Wheat in the current study. bran and soybean hulls are more fibrous than corn The AID values of energy and nutrients were and soybean meal (NRC, 2012), implying that more variable than the ATTD values energy and the former feedstuffs have greater content of sub- nutrients, which could have due to the method strate for the multienzyme than the latter. Thus, the (slaughter technique) of ileal digesta collection. greater effect of multienzyme on the NC2 diet than Ileal digesta of pigs can be collected by slaughter on NC1 diet with regard to AID and ATTD of nu- technique or ileal cannulation technique (Nyachoti trients could be attributed to the fact that the NC2 et  al., 1997). The AID values obtained from ileal diet contained more enzyme substrate than NC1 digesta that is collected by slaughter technique can diet. Zeng et al. (2018) similarly reported that sup- variable due to diurnal variation in nutrient digest- plementation of corn–wheat-based basal diet for ibility (Nyachoti et  al., 1997). The ATTD of GE growing pigs with enzyme product that contained for the PC diet was greater than that for NC1 diet galactanase, xylanase, mannanase, α-amylase, and or NC2 diet, which was expected because the NC1 cellulase activities did not improve nutrient digest- diet and NC2 diet were formulated to contain lower ibility when the basal diet did not contain wheat levels of NE than the PC diet. However, the AID of bran, but improved the AID of GE, NDF, and AA AA for the PC diet did not differ from that for NC1 when the basal diet contained 20% wheat bran. Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Multienzyme supplemented diets for pigs Also, Zeng et al. (2018) observed that supplemen- LITERATURE CITED tation of corn–wheat-based basal diet for growing AOAC. 2012. Official methods of analysis, 19th ed. Rockville pigs with phytase improved AID of phytic acid re- (MD): Association of Official Analytical Chemists gardless of whether or not the basal diet contained (AOAC) International. Appeldoorn, M.M., M.A. Kabel, D. Van Eylen, H. Gruppen, wheat bran, but the addition of an enzyme product and H.A.  Schols. 2010. Characterization of oligomeric that contained galactanase, xylanase, mannanase, xylan structures from corn fiber resistant to pretreat- α-amylase, and cellulose activities to phytase-sup- ment and simultaneous saccharification and fermenta- plemented diet improved AID of phytic acid only tion. J. Agric. Food Chem. 58:11294–11301. doi:10.1021/ for wheat bran-containing basal diet. jf102849x. Bedford,  M.R., and G.G.  Partridge. 2010. Feed enzyme, the The increased AID of CP observed in the cur- future: Bright hope or regulatory minefield. In: M. R. rent study due to multienzyme supplementation Bedford and G. G. Partridge, editors. Enzymes in farm would indicate an increased digestibility of AA due animal nutrition. 2nd ed. Wallingford (UK): CABI to the supplementation. Nevertheless, methionine is Publishing; p. 304–312. the only AA whose AID was significantly improved Bedford, M.R., and H. Schulze. 1998. Exogenous enzymes for by the multienzyme supplementation. In general, pigs and poultry. Nutr. Res. Rev. 11:91–114. doi:10.1079/ NRR19980007. the increase in AA digestibility in response to en- Choct,  M. 1997. Feed non-starch polysaccharides: chemical zyme supplementation is expected to be lower for structures and nutritional significance. Feed Milling Int. highly digestible AA such as methionine than for 191(June):13–26. less digestible AA like threonine (Cowieson, 2010). Cowieson, A.J. 2010. Strategic selection of exogenous enzymes Thus, the results of the current study are contrary to for corn/soy-based poultry diets. J Poult. Sci. 47(1):1–7. Cowieson, A.J., and M.R. Bedford. 2009. The effect of phytase the expectations. Nevertheless, a closer look at the and carbohydrase on ileal amino acid digestibility in mono- change in AID of indispensable AA due to multien- gastric diets: complimentary mode of action? World Poult zyme supplementation reveal numerical increase in Sci. J. 65(4):609–624. doi:10.1017/S0043933909000427. AID of most indispensable AA, and the magnitude Cozannet,  P., M.T.  Kidd, R.  Montanhini  Neto, and of improvement of AID of some of these other P.A.  Geraert. 2017. Next-generation non-starch polysac- indispensable AA is greater than for methionine. charide-degrading, multi-carbohydrase complex rich in xylanase and arabinofuranosidase to enhance broiler feed Among the indispensable AA, the SEM for AID of digestibility. Poult. Sci. 96:2743–2750. doi:10.3382/ps/ methionine was the lowest, implying that AID of pex084. the other indispensable AA was more variable than Emiola,  I.A., F.O.  Opapeju, B.A.  Slominski, and that of methionine. Thus, the statistically insignifi- C.M. Nyachoti. 2009. Growth performance and nutrient cant increase in AID of the other indispensable digestibility in pigs fed wheat distillers dried grains with solubles-based diets supplemented with a multicarbohy- AA could have been due high variability in AID of drase enzyme. J. Anim. Sci. 87:2315–2322. doi:10.2527/ these AA. However, it is not clear why the AID of jas.2008-1195. AA other than methionine was more variable. Jang,  Y.D., P.  Wilcock, R.D.  Boyd, and M.D.  Lindemann. In conclusion, the overall ADG and ATTD 2017. Effect of combined xylanase and phytase on growth of GE for the NC2 diet were lower than those for performance, apparent total tract digestibility, and carcass the PC diet, and multienzyme supplementation characteristics in growing pigs fed corn-based diets con- taining high-fiber coproducts. J. Anim. Sci. 95(9):4005– increased the overall ADG and ATTD of GE for 4017. doi:10.2527/jas2017.1781. the NC2 diet to those of the PC diet; the ATTD Jlali, M., N.M. Bello, M. Ceccantini, D. Moore, R. Shirle, and of P for multienzyme-supplemented NC2 diet was A. Preynat. 2018. Effects of a global enzyme solution on greater than that of PC diet. Thus, the NE and di- growth performance, carcass and tibia characteristics in gestible AA and P can be lowered by ≤5% in mul- broilers fed corn-wheatsoybean based diets reduced in me- tabolizable energy andnutrients at 42  days of age. Poult. tienzyme-supplemented diets without effects on Sci. 97 (E-suppl.1): 38. growth performance pigs. Jlali,  M., P.  Cozannet, R.  Shirle, D.  Moore, M.  Ceccantini, and A.  Preynat. 2019. Investigating the interaction ef- fect of different levels of apparent metabolizable energy, ACKNOWLEDGMENTS digestible amino acids, available phosphorus, arabinox- The authors thank Adisseo France S.A.S.  for ylan, phytate, with and without a multi-carbohydrase and phytase complex on growth performance in broilers. funding the research. The authors would also like Poult. Sci. 98 (E-suppl.1):135. to thank Cameron Pewe and Joseph Wollbrink Kiarie,  E., A.  Owusu-Asiedu, A.  Péron, P.H.  Simmins, and (South Dakota State University, Brookings, SD) C.M.  Nyachoti. 2012. Efficacy of xylanase and β-glu- for assistance with animal care. canase blend in mixed grains and grain co-products-based Conflict of interest statement. The authors de- diets for fattening pigs. Livest. Sci. 148(1):129–133. clare no conflict of interest. doi:10.1016/j.livsci.2012.05.020 Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Jerez-Bogota et al. Knudsen,  K.E.B. 2014. Fiber and nonstarch polysac- Stein,  H.H., B.  Sève, M.F.  Fuller, P.J.  Moughan, and charide content and variation in common crops used in C.F.M.  de  Lange. 2007. Invited review: amino acid bio- broiler diets1. Poult. Sci. 93(9):2380–2393. doi: 10.3382/ availability and digestibility in pig feed ingredients: ter- ps.2014–03902 minology and application. J. Anim. Sci. 85(1):172–180. Li, G., X. Wang, M. Lin, Z. Lu, and W. Yao. 2012. Effects of doi:10.2527/jas.2005–742 corn DDGS in combination with compound enzymes on Turner,  C.H., and D.B.  Burr. 1993. Basic biomechanical growth performance, carcass fat quality, and plasma and measurements of bone: a tutorial. 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Nutrient utilisation and performance S.  Arent, R.  Lorentsen, and C.M.  Nyachoti. 2015. responses of broilers fed a wheat-based diet supplemented Comparative efficacy of xylanases on growth perform- with phytase and xylanase alone or in combination. ance and digestibility in growing pigs fed wheat and wheat Anim. Feed Sci. Technol. 146(1):113–123. doi:10.1016/j. bran- or corn and corn DDGS-based diets supplemented anifeedsci.2007.11.013 with phytase. Anim. Feed Sci. Technol. 209:230–239. Woyengo, T.A., D.V. Ige, O.O. Akinremi, and C.M. Nyachoti. doi:10.1016/j.anifeedsci.2015.08.011 2016. Performance and nutrient digestibility in growing Nortey,  T.N., J.F.  Patience, P.H.  Simmins, N.L.  Trottier, and pigs fed wheat dried distillers’ grain with solubles-contain- R.T.  Zijlstra. 2007. Effects of individual or combined ing diets supplemented with phytase and multi-carbohy- xylanase and phytase supplementation on energy, amino drase. Anim. Sci. 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Effects of graded levels of an Escherichia coli phytase on Zeng, Z.K., Q.Y. Li, Q.Y. Tian, Y.T. Xu, and X.S. Piao. 2018. growth performance, apparent total tract digestibility of The combination of carbohydrases and phytase to im- phosphorus, and on bone parameters of weanling pigs prove nutritional value and non-starch polysaccharides fed phosphorus-deficient corn-soybean meal based diets. degradation for growing pigs fed diets with or without Anim. Feed Sci. Technol. 232:102–109. doi:10.1016/j. wheat bran. Anim. Feed Sci. Technol. 235:138–146. anifeedsci.2017.08.005. doi:10.1016/j.anifeedsci.2017.11.009 Translate basic science to industry innovation http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png Translational Animal Science Oxford University Press

Growth performance and nutrient digestibility of growing and finishing pigs fed multienzyme-supplemented low-energy and -amino acid diets

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Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Growth performance and nutrient digestibility of growing and finishing pigs fed multienzyme-supplemented low-energy and -amino acid diets † † † ‡ ‡ Kevin Jerez-Bogota , Cristian Sánchez , Jimena Ibagon , Maamer Jlali , Pierre Cozannet , ‡ †,1 Aurélie Preynat , and Tofuko A. Woyengo † ‡ Department of Animal Science, South Dakota State University, Brookings, SD 57007; and Adisseo France S.A.S., Center of Expertise and Research in Nutrition, F-03600 Commentry, France ABSTRACT:  A study was conducted to deter- the PC and NC1 diets did not differ in average mine the effects of supplementing corn–soybean daily gain (ADG) and average daily feed intake. meal-based diets with a multienzyme on growth Pigs fed NC2 diet had lower (P  <  0.05) ADG performance, bone mineralization, apparent ileal and gain-to-feed ratio (G:F) than those fed PC digestibility (AID) and apparent total tract di- diet. Pigs fed PC diet had greater (P < 0.05) bone gestibility (ATTD) of nutrients of growing pigs. ash content and ATTD of P than those fed NC1 A total of 276 pigs (body weight [BW] = 33.99 ± diet. The ATTD of GE for PC diet was greater 4.3  kg) were housed by sex in 45 pens of 6 or 7 (P  <  0.05) than that for NC2 diet, and tended pigs and fed 5 diets (9 pens/diet) in a randomized to be greater (P  <  0.10) than that for NC1 diet. complete block design. Diets were positive control Multienzyme interacted (P < 0.05) with negative (PC); and negative control 1 (NC1) or negative control diet type on overall ADG and AID of GE control 2 (NC2) without or with multienzyme. The such that multienzyme did not affect overall ADG multienzyme used supplied at least 1,800, 1,244, and AID of GE for the NC1 diet, but increased 6,600, and 1,000 units of xylanase, β-glucanase, (P < 0.05) overall ADG and AID of GE for NC2 arabinofuranosidase, and phytase per kilogram diet by 5.09 and 8.74%, respectively. Multienzyme of diet, respectively. The PC diet was adequate in did not interact with negative control diet type all nutrients according to NRC recommendations on overall G:F, bone ash content, AID of AA, and had greater digestible P content than NC1 or and ATTD of nutrients. Multienzyme increased NC2 diet by 0.134 percentage points. The PC diet (P  <  0.05) overall G:F, AID of methionine, had greater net energy (NE) and standardized ATTD of GE and P, and tended to increase ileal digestible amino acids (AA) content than (P  =  0.056) bone ash content. The ADG, bone NC1 diet by 3%, and than NC2 diet by 5%. The ash content, and ATTD of GE and P for the mul- diets were fed in 4 phases based on BW: Phase 1: tienzyme-supplemented diets were similar to (P > 34–50 kg; Phase 2: 50–75 kg; Phase 3: 75–100 kg; 0.10) PC diet. Thus, NE and digestible AA and and Phase 4: 100–120  kg. Nutrient digestibility P can be lowered by ≤5% in multienzyme-supple- and bone mineralization were determined at the mented diets without effects on growth perform- end of Phase 1. Overall (34–120 kg BW), pigs fed ance and bone ash of pigs. Key words: bone mineralization, growth performance, multienzyme, nutrient digestibility, pig © The Author(s) 2020. Published by Oxford University Press on behalf of the American Society of Animal Science. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribu- tion, and reproduction in any medium, provided the original work is properly cited. Transl. Anim. Sci. 2020.4:1-14 doi: 10.1093/tas/txaa040 Corresponding author: tofuko.woyengo@sdstate.edu Received November 25, 2019. Accepted April 6, 2020. 1 Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Jerez-Bogota et al. INTRODUCTION effective with regard to degradation of arabinoxy- lan than supplementation with xylanase alone be- The major sources of nutrients in swine diets cause arabinofuranosidase can cleave off arabinose are plant origin feedstuffs. However, these feed- units from backbones of xylans, leading to reduced stuffs contain some antinutritional factors, which resistance of arabinoxylans to enzymatic degrad- limit nutrient utilization. Some of the important ation. Indeed, supplementation of a multienzyme antinutritional factors present in these feedstuffs product that contained xylanase, arabinofuranosi- include phytic acid and non-starch polysacchar- dase, β-glucanase, and phytase activities improved ides (NSP). Phytic acid contain P, which is poorly nutrient utilization in poultry fed wheat-corn-based digested by pigs because they do not produce suf- diets (Jlali et al., 2018, 2019). However, information ficient amounts of phytase to liberate phytic acid- is lacking on the effects of the same multienzyme bound P (Woyengo and Nyachoti, 2011). Also, product on nutrient utilization in pigs fed wheat– phytic acid reduces digestibility of other nutrients corn-based diets. Therefore, the objective of this including cations and amino acids (AA) by binding study was to determine the effects of supplemen- them (Woyengo and Nyachoti, 2013). The NSP are tation multienzyme product that contain xylanase, poorly digested by swine and can reduce nutrient arabinofuranosidase, β-glucanase, and phytase ac- availability for digestion and absorption partly by tivities on nutrient digestibility, bone mineraliza- encapsulation and viscosity (Bedford and Partridge, tion, and growth performance of grow–finish pigs 2010; Bedford and Schulze, 1998; Woyengo et  al., fed corn–wheat–wheat bran–soybean meal-based 2016). Furthermore, phytic acid and NSP can have diets that are low in net energy (NE), standardized negative effects on environment due to increased ileal digestible AA, Ca, and standardized total tract excretion of unabsorbed nutrients, especially N and digestible P. P (Woyengo et al., 2008b). The undesirable effects of phytic acid and NSP MATERIALS AND METHODS can be alleviated through dietary supplementation of phytase and NSP-degrading enzymes also known Experimental procedures were reviewed and as NSPases (Cowieson and Bedford, 2009). Corn approved by the Institutional Animal Care and and wheat grains, and their co-products are the Use Committee at South Dakota State University most widely used sources of energy in swine diets (#18-015E). in North America and Europe. The most abundant NSP in wheat and corn are arabinoxylans (Choct, Experimental Animals 1997; Knudsen, 2014). Wheat also contain some A total of 276 pigs (initial body weight [BW] of β-glucans (Choct, 1997). Arabinoxylans are com- 33.99 ± 4.3 kg; Lance-Large White female × Duroc posed of backbones of xylans that are substituted male; Pig Improvement Company) were obtained mainly with arabinose; arabinose is linked to ferulic from the Swine Education and Research Facility, acid, which crosslink xylans and lignin (Appeldoorn South Dakota State University (Brookings, SD). et al., 2010). Xylanase has been added in corn- and Pigs were then individually weighed and housed wheat-based diets for pigs with the goal of increas- in 45 pens of 6 or 7 pigs. Pens (1.8  × 2.4 m) had ing dietary nutrient utilization by degrading ara- fully slated-concrete floors, metal spindle walls (1.0 binoxylans. However, xylanase had inconsistent m high), and solid polyvinyl chloride gates. Each effects on nutrient utilization in pigs fed diets that pen was equipped with a cup drinker, and a dou- are based on corn, wheat or their co-products. For ble-space dry feeder. Room temperature was main- instance, a few studies (e.g. Ndou et  al., 2015) re- tained at 22 ± 2°C throughout the experiment. ported improved growth performance of pigs due to addition of xylanase in wheat- or corn-based diets, whereas most studies (e.g. Nortey et  al., Experimental Diets 2007; O’Shea et  al., 2014; Woyengo et  al., 2008b) Five diets based on corn, soybean meal, wheat, did not report any improvement. The cross-linking wheat bran, and soybean hulls were fed in this study. of xylans and lignin with ferulic acid can poten- The diets included a positive control diet (PC); and tially make arabinoxylans more resistant to deg- negative control diet 1 (NC1) and negative control radation by xylanase alone. Thus, supplementation diet 2 (NC2) without or with multienzyme in 2  × of pig’s diets that are based on corn, wheat, or 2 factorial arrangement (Table  1). The multien- their co-products with a combination of xylanase zyme (a multi-carbohydrase and phytase complex, and arabinofuranosidase can potentially be more Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Multienzyme supplemented diets for pigs l dl l l Translate basic science to industry innovation Table 1. Ingredient and calculated chemical composition of the basal diets (%, as-fed basis) Phase 1: 34–50 kg BW Phase 2: 50–75 kg BW Phase 3: 75–100 kg BW Phase 4: 100–135 kg BW Item PC NC1 NC2 PC NC1 NC2 PC NC1 NC2 PC NC1 NC2 Ingredients, % Corn 55.734 54.033 51.223 60.069 58.059 55.289 61.402 59.623 56.271 65.909 60.757 57.812 Soybean meal 15.211 14.260 13.465 10.351 9.544 9.040 8.814 7.900 7.108 4.953 3.437 2.505 Wheat 10.000 10.000 10.000 10.000 10.000 10.000 10.000 10.000 10.000 10.000 10.000 10.000 Wheat bran 10.000 11.783 15.000 10.000 12.000 14.267 10.000 11.700 15.339 10.000 13.600 16.000 Soybean hulls 2.500 4.500 5.000 3.545 5.471 6.500 4.207 6.289 6.808 4.702 8.438 9.995 Soybean oil 3.225 2.800 2.713 3.000 2.600 2.600 3.000 2.600 2.600 2.300 2.300 2.234 Calcium carbonate 1.083 1.040 1.047 0.998 0.955 0.960 0.875 0.833 0.840 0.689 0.561 0.543 Monocalcium phosphate 1.063 0.415 0.397 0.886 0.236 0.224 0.744 0.096 0.076 0.540 - - -Lysine HCl 0.488 0.480 0.473 0.479 0.470 0.462 0.380 0.380 0.380 0.351 0.349 0.350 -Methionine 0.096 0.090 0.086 0.071 0.066 0.062 0.025 0.023 0.021 0.0002 0.0004 0.0004 -Thronine 0.124 0.121 0.121 0.114 0.111 0.111 0.083 0.085 0.087 0.090 0.094 0.096 -Tryptophan 0.043 0.041 0.039 0.050 0.047 0.044 0.031 0.030 0.027 0.029 0.028 0.027 Mineral premix 0.150 0.150 0.150 0.150 0.150 0.150 0.150 0.150 0.150 0.150 0.150 0.150 Vitamin premix 0.050 0.050 0.050 0.050 0.050 0.050 0.050 0.050 0.050 0.050 0.050 0.050 Salt 0.234 0.236 0.237 0.238 0.240 0.240 0.239 0.241 0.242 0.237 0.237 0.237 Calculated nutrients Crude protein, % 14.735 14.567 14.467 12.943 12.839 12.781 12.249 12.098 12.029 11.108 10.985 10.868 Ether extract, % 5.934 5.493 5.375 5.787 5.367 5.319 5.810 5.390 5.348 5.394 5.371 5.292 NE, kcal/kg 2,475 2,401 2,351 2,475 2,401 2,351 2,475 2,401 2,351 2,475 2,401 2,351 Standardized digestible content of AA, % Lys 0.980 0.951 0.931 0.847 0.822 0.805 0.730 0.708 0.694 0.610 0.592 0.580 Met 0.316 0.305 0.296 0.269 0.259 0.251 0.216 0.208 0.202 0.165 0.158 0.154 Met + Cys 0.550 0.534 0.523 0.480 0.466 0.456 0.420 0.407 0.399 0.360 0.349 0.342 Thr 0.590 0.572 0.560 0.509 0.493 0.484 0.456 0.442 0.433 0.400 0.388 0.380 Trp 0.170 0.165 0.162 0.153 0.148 0.145 0.127 0.123 0.120 0.110 0.107 0.105 Total P, % 0.596 0.470 0.487 0.542 0.417 0.428 0.506 0.380 0.398 0.459 0.359 0.371 Digestible P 0.310 0.176 0.176 0.269 0.135 0.135 0.238 0.104 0.104 0.210 0.092 0.093 Calcium 0.660 0.540 0.540 0.590 0.470 0.47 0.520 0.400 0.400 0.460 0.340 0.340 Sodium 0.100 0.100 0.100 0.100 0.100 0.100 0.100 0.100 0.100 1.000 1.000 1.000 BW, body weight; PC, positive control diet; NC1, negative control diet 1 with lower in NE, standardized ileal digestible AA, standardized total tract digestible P, and Ca than PC diet by 3, 3, 43, and 18%, respectively; and NC2, negative control diet 2 with lower in NE, standardized ileal digestible AA, standardized total tract digestible P, and Ca than PC diet by 5, 5, 43, and 18%, respectively. Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Jerez-Bogota et al. Rovabio Advance Phy, Adisseo France S.A.S) was Sample Preparation and Analyses added to diets supplying at least 1,800, 1,244, 6,600, Femurs for determining bone ash were defleshed and 1,000 units of xylanase, β-glucanase, arabino- by autoclaving at 121°C for 30  min, cleaned and furanosidase, and phytase per kilogram of diet, re- subsequently dried in an oven at 135°C for 2 h. Fat spectively. Enzyme supplementation levels were as was extracted from the dried bones using petroleum per the supplier’s recommendations. The PC diet ether (E139-4, Fischer Scientific, Pittsburgh, PA) as was formulated to be adequate in all nutrients ac- solvent in a Jumbo Soxhlet extraction apparatus cording to NRC (2012) recommendations. The (Chemglass Life Sciences, Vineland, NJ), afterward NC1 diet was the same as the PC diet except that its the samples were left in a fume hood for 24  h to NE and standardized ileal digestible AA, standard- allow the petroleum ether to evaporate. Femurs were ized total digestible P, and total Ca contents were then dried in an oven at 135°C for 2 h to determine lower than those for the PC diet by 3.0, 3.0, 43, and their fat-free weight, and ashed at 600°C in a muffle 18%, respectively. The NC2 diet was the same as the furnace for 12 h for the determination of bone ash. PC diet except that its NE and standardized ileal Femurs for determining bone breaking strength digestible AA, standardized total digestible P, and were defleshed by scraping muscle tissues from the total Ca contents were lower than those for the PC bones using kitchen knives. Maximal breaking load diet by or 5.0, 5.0, 43, and 18%, respectively. The was measured using an MTS Insight 5 equipment reduction in NE value and nutrient content in the (MTS, Eden Prairie, MN, USA) at room tempera- NC1 and NC2 diets was achieved by a partial re- ture by subjecting each bone to a 3-point bending placement of corn, soybean meal, soybean oil, crys- test (Tunrer and Burr, 1993). Force was applied talline AA, calcium carbonate, and monocalcium to the center of the bone held by supports 3.3 cm phosphate in PC diet with wheat bran and soybean apart. The crosshead speed was set at 50  mm/min hulls. The diets were fed in mash form and in 4 and the sample rate was 10 points/s. Final strength phases based on BW: Phase 1: 34–50  kg, Phase 2: was determined from load–displacement curves. 50–75  kg, Phase 3: 75–100  kg, and Phase 4: 100– Fecal samples were pooled by pen and air-dried 125 kg. Titanium dioxide (0.3%) was added as indi- in an oven at 60°C for 4 d; whereas ileal digesta gestible marker in each diet during the last week of samples were freeze–dried. The dried fecal and the first phase of feeding. ileal digesta samples together with diet samples were ground through a 0.75-mm screen in a cen- trifugal mill (model ZM200; Retsch GmbH, Haan, Experimental Design and Procedure Germany). The ground samples were analyzed as The five diets were allotted to the 45 pens (9 pens/ follows: Phase 1 diets for DM, gross energy (GE), diet) within a randomized complete block design. N (N × 6.25  =  CP), P, AA, NDF, ADF and ti- Diets and fresh water were offered to pigs ad libitum tanium contents, and for xylanase and phytase during the entire period. Pig BW and feed intake were activities; Phases 2–4 diets for DM, GE, and N; determined by phase to calculate average daily gain ileal digesta for DM, GE, NDF, ADF, AA, P and (ADG), average daily feed intake (ADFI), and gain- titanium contents; and feces for GE, CP, NDF, to-feed ratio (G:F). Fresh fecal samples were collected ADF, Ca, P, and titanium contents. The samples from each pen during the last 2 days of first feeding were analyzed for DM by oven drying at 135°C phase and immediately stored frozen at −20°C for for 2 h (method 930.15), CP by a combustion pro- the determination of apparent total tract digestibility cedure (method 990.03), as per AOAC (2012); and (ATTD) of energy and nutrients. At the end of the first for ADF and NDF (Van Soest et al., 1991) on an phase of feeding, 1 pig (per pen) with BW that was Ankom 200 Fiber Analyzer (Ankom Technology, close to the pen average BW was selected, and then eu- Fairport, NY). Samples were analyzed for AA thanized by captive bolt penetration followed by ex- (method 982.30 E [a, b, and c]; AOAC, 2012) at sanguination. Right and left femurs were excised from the University of Missouri Experiment Station la- each euthanized pig and stored at −20°C for deter- boratories (Columbia, MO). The GE was analyzed mination of bone ash and bone breaking strength, re- using an adiabatic bomb calorimeter (model 1261, spectively. Also, contents of lower half of ileum (from Parr Instrument Co., Moline, IL). Titanium dioxide 80  cm above ileal-cecal junction to approximately in samples was determined by spectrophotometry 1 cm above the ileo-cecal junction) were obtained and (model Spectra MAX 190, Molecular Devices, stored frozen at −20°C for latter determination of ap- Sunnyvale, CA) at 408 nm after ashing at 525°C for parent ileal digestibility (AID) of energy and nutrients. 10 h (Myers et al., 2004). The P content in samples Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Multienzyme supplemented diets for pigs was analyzed according to the vanadate colori- RESULTS metric method (method 946.06; AOAC, 2012) using The analyzed CP values in the diets in Tables 2 a spectrophotometer (model Spectra MAX 190, and 3 were similar to the calculated CP values in Molecular Devices, Sunnyvale, CA). The Ca con- the diets in Table 1. The mean endogenous phytase tent in samples was analyzed by inductively coupled activity in PC, NC1, and NC2 diets was 406 U/kg. plasma–optical emission spectrometry (ICP–OES; Addition of phytase to the NC1 and NC2 diets re- method 985.01 A, B, and C; AOAC, 2012) after sulted in its increased activity values by margins wet ashing samples (method 975.03 B(b); AOAC, similar to those that were anticipated. The xylanase 2012). Xylanase and phytase activity in experi- activity was undetectable in the PC, NC1, and NC2 mental diets were analyzed by the Laboratory of diets. The analyzed xylanase activity values in mul- Adisseo (Commentry, France). Xylanase activity tienzyme supplemented NC1 and NC2 diets were was determined according the method described by similar to the anticipated values. Cozannet et  al. (2017). One visco-unit of endo-1, Data on effects of dietary treatment on growth 4-β-xylanase activity is defined as the amount of performance, bone mineralization, AID of energy enzyme that hydrolyzed by the substrate (wheat and nutrients, and ATTD of energy and nutrients AX); one such unit reduces solution’s viscosity, re- are presented in Tables  4–7, respectively. Pigs fed sulting in a change in relative fluidity of 1 arbitrary PC diet had greater (P < 0.05) BW than those fed unit per min per ml (or per g) under the conditions NC1 diet during Phases 1 and 2 of feeding, but not of the assay and pH 5.5 and 30°C. Phytase activity during Phases 3 and 4. The BW of pigs fed PC diet in experimental diets was determined according the was greater (P < 0.05) than that of pigs fed NC2 diet standard method (ISO3024, 2009). One phytase during overall (34–120  kg BW) study period. Pigs unit (FTU) is the amount of enzyme that releases fed PC diet had greater (P < 0.05) ADG than those fed NC1 diet during Phase 1, but not during Phases 1 µmol of inorganic orthophosphate from sodium 2–4 of feeding. Also, the overall ADG of pigs fed phytate substrate per minute at pH 5.5 and 37°C. PC diet did not differ from that of pigs fed NC1 diet. Pigs fed PC diet had greater (P < 0.05) ADG Calculations and Statistical Analysis than those fed NC2 diet during the Phases 1, 2, and 4. Also, the overall ADG of pigs fed PC was greater The AID and ATTD values of the diets were (P < 0.05) than that of pigs fed NC2 diet. Pigs fed calculated using the indicator method (Stein et al., NC1 diet had greater (P < 0.05) ADFI than those 2007), using the following equation: fed PC diet during Phases 1 and 3 of feeding, but AID or ATTD, % =[1 − (Nutrient /Nutrient ), digesta diet not during Phases 2 and 4; whereas pigs fed NC2 ×(Marker /Marker )] × 100 diet digesta diet had greater (P < 0.05) ADFI than those fed PC diet during the Phase 3, but not during the Phases where Nutrient is the nutrient concentration digesta 1, 2, and 4. Pigs fed NC1 diet had lower (P < 0.05) in the ileal digesta or feces (%  DM); Nutrient diet G:F than those fed PC diet during the Phase 1 of is the nutrient concentration in the diet (%  DM); feeding, but not during the Phases 2–4 and during Marker is the titanium concentration in the diet diet the overall period. The G:F for NC2 diet was lower (%  DM); and Marker is the titanium concen- digesta (P < 0.05) than that for the PC diet during the entire tration in the ileal digesta or feces (% DM). study period. The percent femur ash content for PC Data were subjected to analysis of variance diet was greater (P < 0.05) than that for NC1 diet using the MIXED procedure of SAS (SAS Inst. or NC2 diet. The femur breaking strength for PC Inc., Cary, NC). The pen was considered as the diet was numerically, but not significantly greater experimental unit. The model included diet, sex, than that for NC1 diet or NC2 diet. The PC, NC1, diet × sex interaction, and initial BW, which was and NC2 diets did not differ in AID of GE, CP and a covariate. Period was the repeated term in mod- indispensable AA. The ATTD of DM and GE for els involving time. Means were separated by the NC1 diet tended to be lower (P  <  0.10) than that probability of difference in order to compare PC for the PC diet, whereas the ATTD of CP and P diet with other diets. Main effects of NC diet type for NC1 diet was lower (P < 0.05) than that for PC and multienzyme and their interactions were deter- diet. The ATTD of DM, GE, and P for the NC2 mined. To test the hypotheses, P  <  0.05 was con- diet was lower (P < 0.05) than that for PC diet. sidered significant. If pertinent, trends (0.05 < P ≤ The NC diet type and multienzyme did 0.10) are also reported. not interact on BW during Phase 1, 2, and 3 of Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Jerez-Bogota et al. Table 2. Analyzed composition of phase 1 diets as-fed Diets Item PC NC1 NC2 NC1 + E NC2 + E Dry matter, % 88.5 88.9 89.3 89.1 89.3 Gross energy, kcal/kg 3,973 4,052 4,067 3,990 4,053 Crude protein, % 14.38 14.54 14.81 14.04 14.00 Neutral detergent fiber, % 12.88 15.26 18.25 14.21 17.85 Acid detergent fiber, % 4.50 6.48 7.10 6.00 6.72 Ca, % 0.66 0.52 0.40 0.53 0.37 P, % 0.60 0.51 0.54 0.53 0.50 Indispensable AA, % Arg 0.80 0.87 0.83 0.78 0.79 His 0.35 0.38 0.37 0.35 0.35 Ile 0.53 0.56 0.54 0.53 0.51 Leu 1.15 1.19 1.11 1.12 1.04 Lys 0.93 0.97 0.97 0.94 0.93 Met 0.32 0.29 0.27 0.32 0.29 Phe 0.63 0.66 0.63 0.62 0.6 Thr 0.59 0.62 0.59 0.59 0.6 Trp 0.22 0.23 0.22 0.2 0.21 Val 0.63 0.67 0.66 0.63 0.62 Dispensable AA, % Ala 0.68 0.72 0.68 0.67 0.64 Asp 1.17 1.25 1.18 1.16 1.09 Cys 0.27 0.27 0.27 0.26 0.27 Glu 2.53 2.61 2.53 2.42 2.51 Gly 0.58 0.64 0.64 0.59 0.62 Pro 0.91 0.92 0.9 0.88 0.88 Ser 0.59 0.63 0.58 0.57 0.56 Tyr 0.48 0.49 0.45 0.45 0.44 Analyzed enzyme activities, U/kg Phytase 350 359 389 1,400 1,391 Xylanase 0 0 0 1,834 1,850 PC, positive control diet; NC1, negative control diet 1; NC2, negative control diet 2; NC1 + E, negative control diet 1 plus multienzyme; and NC2 + E, negative control diet 2 plus multienzyme. feeding. Multienzyme supplementation improved NC1 diet with multienzyme reduced (P < 0.05) the (P  <  0.05) BW during phase 1–3 regardless of ADG, whereas supplementation of the NC2 diet NC diet type. The NC diet type and multienzyme tended to increase (P < 0.10) the ADG of pigs. The tended to interact (P = 0.066) on final BW (Phase NC type and multienzyme interacted (P  =  0.046) 4)  such that supplementation of NC1 diet with on the overall ADG such that supplementation multienzyme did not affect the final BW, whereas of NC1 diet with multienzyme did not affect the supplementation of the NC2 diet increased ADG, whereas supplementation of the NC2 diet (P  <  0.05) the final BW of pigs. Multienzyme increased (P < 0.05) the ADG of pigs. The NC diet and NC diet type did not interact on ADG dur- type and multienzyme interacted (P  =  0.024) on ing Phase 1.  However, multienzyme supplemen- the ADFI during the Phase 1 of feeding such that tation improved (P  <  0.05) ADG during phase the ADFI for NC1 diet was reduced (P  <  0.05), 1 regardless of NC diet type. The NC type and whereas that of the NC2 diet was unaffected by multienzyme tended to interact (P  =  0.070) on the multienzyme. Multienzyme supplementation ADG during Phase 2 such that supplementation reduced (P = 0.002) ADFI during Phase 3 regard- of NC1 diet with multienzyme did not affect the less of NC diet type. The NC diet type and mul- ADG, whereas supplementation of the NC2 diet tienzyme interacted (P  =  0.013) on ADFI during increased (P < 0.05) the ADG of pigs. The NC diet Phase 4 such that supplementation of NC1 diet re- type and multienzyme interacted (P  =  0.003) on duced (P < 0.05) the ADFI, whereas the ADFI for ADG during Phase 4 such that supplementation of NC2 diet was unaffected. The NC diet type and Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Multienzyme supplemented diets for pigs Table 3. Analyzed composition of phases 2–4 diets as-fed Diets Item PC NC1 NC2 NC1 + E NC2 + E Phase 2 Dry matter, % 86.4 90.8 87.9 87.8 88.1 Gross energy, kcal/kg 3,917 3,947 3,920 3,850 4,007 Crude protein, % 12.37 12.63 12.47 12.57 12.45 Neutral detergent fiber, % 11.13 13.02 13.29 13.23 14.75 Acid detergent fiber, % 4.57 6.12 5.67 4.96 6.32 Analyzed enzyme activities, U/kg Phytase 372 394 440 1,410 1,500 Xylanase 0 0 0 1,840 1,865 Phase 3 Dry matter, % 88.5 88.4 88.3 88.7 88.2 Gross energy, kcal/kg 3,913 4,101 3,996 3,931 3,968 Crude protein, % 12.03 11.92 12.20 11.64 12.29 Neutral detergent fiber, % 12.76 16.23 15.97 13.71 15.91 Acid detergent fiber, % 5.53 5.64 7.31 5.64 5.78 Analyzed enzyme activities, U/kg Phytase 374 366 462 1,441 1,390 Xylanase 0 0 0 1,860 1,830 Phase 4 Dry matter, % 88.5 88.4 88.3 88.7 88.2 Gross energy, kcal/kg 3,918 3,910 3,966 3,915 3,924 Crude protein, % 10.11 10.06 10.11 9.85 9.83 Neutral detergent fiber, % 12.27 14.94 16.04 16.44 17.79 Acid detergent fiber, % 4.62 6.45 7.7 6.51 8.22 Analyzed enzyme activities, U/kg Phytase 451 426 485 1,500 1,440 Xylanase 0 0 0 1,835 1,860 PC, positive control diet; NC1, negative control diet 1; NC2, negative control diet 2; NC1 + E, negative control diet 1 plus multienzyme; and NC2 + E, negative control diet 2 plus multienzyme. multienzyme interacted (P = 0.045) on the overall CP such that multienzyme supplementation tended ADFI such that the ADFI for NC1 diet, but not to increase (P < 0.10) the AID of CP for NC1 diet, of NC2 diet, was reduced (P < 0.05) by the supple- but increased (P < 0.05) AID of CP for NC2 diet. mentation. Multienzyme and NC diet type inter- Multienzyme supplementation did not affect AID acted (P = 0.002) on G:F during Phase 1 such that of all indispensable AA except of Met whose AID the supplementation increased (P < 0.05) G:F for was increased (P = 0.006) by the supplementation NC1 diet and tended to increase (P  <  0.10) G:F regardless of the NC diet type. No interactions for NC2 diet. Also, multienzyme and NC diet type were detected between NC diet type and multien- tended to interact (P = 0.084) on G:F during Phase zyme on ATTD of DM, OM, GE, CP, NDF, ADF, 4 such that the multienzyme supplementation did and P.  Multienzyme supplementation increased not affect the G:F for NC1, but tended to increase (P < 0.05) ATTD of DM, OM, GE, CP, Ca, and P. (P < 0.10) G:F for NC2 diet. Multienzyme and NC diet type did not interact on overall G:F; however, DISCUSSION the multienzyme increased (P < 0.001) overall G:F regardless of NC diet type. Multienzyme supple- The ADG of pigs fed the NC1 diet was lower mentation tended to increase (P  =  0.056) percent than that of pigs fed the PC diet during the first femur ash content and increased (P < 0.001) bone phase of feeding, implying that the reduction in breaking regardless of NC diet type. Multienzyme NE, standardized ileal digestible AA, standardized and NC diet type interacted (P = 0.007) on AID of total tract digestible P, and total Ca contents in PC GE such that multienzyme did not affect the AID diet by 74 kcal/kg, 3%, 43%, and 18%, respectively, of GE for NC1 diet and increased (P < 0.05) AID was sufficient to reduce the growth performance of of GE for NC2 diet. Also, multienzyme and NC the pigs weighing between 34 and 55 kg. However, diet type tended to interact (P = 0.10) on AID of the ADG of pigs fed the NC1 diet did not differ Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Jerez-Bogota et al. Translate basic science to industry innovation Table 4. Effect of dietary treatments on growth performance a b Diets P-value Item PC NC1 NC2 NC1 + E NC2 + E SEM Diet NC E NC × E BW, kg Day 0 34.61 34.60 34.60 34.67 34.54 - - a b b a a Phase 1 56.14 53.83 54.54 56.03 55.89 0.472 0.004 0.341 <0.001 0.305 a b b a a Phase 2 75.13 72.77 71.33 74.81 75.13 0.670 0.001 0.754 <0.001 0.341 a ab b a ab Phase 3 101.53 99.68 98.12 101.27 100.41 1.000 0.092 0.264 0.021 0.883 a a b a a Phase 4 124.45 123.87 118.90 124.14 123.26 1.260 0.009 0.038 0.040 0.066 ADG, kg a b b a a Phase 1 0.925 0.821 0.851 0.912 0.912 0.021 0.004 0.301 <0.001 0.425 a ab b a a Phase 2 0.981 0.936 0.884 0.948 0.980 0.021 0.018 0.727 0.008 0.070 Phase 3 0.990 1.004 0.962 0.983 0.938 0.024 0.279 0.045 0.317 0.898 ab a c bc bc Phase 4 0.998 1.064 0.902 0.954 0.970 0.031 0.009 0.032 0.416 0.003 a a b a a Overall 0.966 0.955 0.899 0.955 0.949 0.013 0.008 0.022 0.164 0.046 ADFI, kg a b a a a Phase 1 1.908 2.312 1.850 1.896 1.929 0.038 <0.001 0.050 0.181 0.024 Phase 2 2.448 2.359 2.369 2.410 2.491 0.047 0.231 0.211 0.071 0.404 bc a ab bc c Phase 3 2.669 2.796 2.819 2.647 2.621 2.669 0.008 0.989 0.002 0.745 ab a ab b ab Phase 4 3.255 3.267 3.076 3.022 3.135 0.080 0.150 0.724 0.127 0.013 b a b b b Overall 2.57 2.70 2.53 2.48 2.54 0.04 0.003 0.353 0.096 0.045 G:F, kg/kg a c b ab ab Phase 1 0.485 0.367 0.463 0.480 0.474 0.007 <0.001 0.004 <0.001 0.002 a ab b ab ab Phase 2 0.401 0.397 0.377 0.394 0.394 0.007 0.099 0.151 0.237 0.321 a ab b a ab Phase 3 0.372 0.359 0.343 0.373 0.358 0.009 0.082 0.046 0.059 0.900 ab a b ab ab Phase 4 0.309 0.324 0.295 0.317 0.310 0.009 0.155 0.015 0.560 0.084 a c b ab ab Overall 0.485 0.367 0.463 0.480 0.474 0.007 <0.001 0.718 <0.001 0.130 PC, positive control diet; NC1, negative control diet 1; NC2, negative control diet 2; NC1 + E, negative control diet 1 plus multienzyme; and NC2 + E, negative control diet 2 plus multienzyme. NC, main effects of negative control diet type; E, main effects of multienzyme; and NC × E, interaction between negative control diet type and multienzyme. abc Within a row, means without a common superscript differ (P < 0.05). Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Multienzyme supplemented diets for pigs from that of pigs fed the PC diet during the second, third and fourth phases of feeding, leading to lack of differences between NC1 and PC diets with re- gard to the overall ADG and final BW. Such results indicate that the reduction in NE, standardized ileal digestible AA, standardized total tract digestible P, and total Ca contents in PC diet by 74 kcal/kg, 3%, 43%, and 18%, respectively, during the phases of feeding 2, 3, and 4 was not sufficient to reduce the growth performance of the pigs, and could be at- tributed to an increase in digestive capacity of pigs with age. Similarly, Woyengo et al. (2008b) did not observe reduction in ADG of grow–finish pigs fed wheat-based diets due to reduction in DE value and available P contents in PC diet by 75 kcal/kg and 30.4%, respectively. Also, Li et  al. (2012) did not observe reduction in ADG of pigs due to reduction in DE value by 50 kcal/kg. The ADG of pigs fed the NC2 diet was lower than that of pigs fed the PC diet during the first, second, and fourth phases of feeding. Also, the overall ADG and hence final BW of pigs fed the NC2 diet were lower than those of pigs fed the PC diet, implying that the reduction in NE, standardized ileal digestible AA, standard- ized total tract digestible P, and total Ca contents in PC diet by 124 kcal/kg, 5%, 43%, and 18%, respect- ively, was sufficient to reduce growth performance of the pigs. Emiola et al. (2009) observed reduction in ADG of growing pigs due to reduction in dietary DE by 124 kcal/kg. Jang et al. (2017) also served re- duction in ADG of growing pigs due to reduction in dietary ME by 103 kcal/kg. Supplementation of NC1 diet or NC2 diet with multienzyme increased ADG of pigs during the first phase (from 34 to 55  kg BW) of feeding, which was due to increased nutrient digestibility by the supplementation. This result is in accordance with several studies reporting an improvement of ADG in growing pigs fed diets supplemented with a multi-carbohydrase (Emiola et  al., 2009; Kiarie et  al., 2012; Ndou et  al., 2015). Supplementation of NC2 diet with multienzyme increased ADG of pigs during the second phase of feeding (50–75 kg BW), and hence during the entire study period (34– 125  kg BW), whereas it is not the case with NC1 diet. It should be noted that the NC2 diet was for- mulated to contain less NE and digestible AA than NC1 diet; and that the addition of multienzyme to NC2 diet resulted in an increase in AID of GE and CP, whereas the addition of multienzyme to NC1 diet only tended to increase in AID of CP. Also, the magnitude by multienzyme increased the ATTD of P for NC2 diet was greater than the magnitude by which it increased the ATTD of P for NC1 diet Translate basic science to industry innovation Table 5. Effect of dietary treatment on bone mineralization a b Diets P-value Item SEM PC NC1 NC2 NC1 + E NC2 + E Diet NC E NC × E ab c bc a ab Femur ash, g 32.67 28.90 29.78 35.97 34.27 1.814 0.049 0.353 <0.001 0.118 a c bc ab a Femur ash, % 55.32 51.99 52.75 54.33 55.15 0.660 0.008 0.619 0.056 0.735 bc c c a ab Femur breaking strength, N 2,513 2,191 2,278 2,950 2,672 129.3 0.002 0.385 <0.001 0.177 PC, positive control diet; NC1, negative control diet 1; NC2, negative control diet 2; NC1 + E, negative control diet 1 plus multienzyme; and NC2 + E, negative control diet 2 plus multienzyme. NC, main effects of negative control diet type; E, main effects of multienzyme; and NC × E, interaction between negative control diet type and multienzyme. abc Within a row, means without a common superscript differ (P < 0.05). Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Jerez-Bogota et al. Translate basic science to industry innovation Table 6. Effect of dietary treatment on AID of energy and nutrients a b Diets P-value AID, % PC NC1 NC2 NC1 + E NC2 + E SEM Diet NC E NC × E ab b b b a Dry matter 67.73 66.48 66.21 65.16 71.73 1.51 0.060 0.053 0.130 0.028 ab bc abc c a Gross energy 71.12 67.56 67.53 64.66 73.43 2.3 0.028 0.065 0.206 0.007 bc bc c ab a Crude protein 65.21 64.31 62.43 69.53 75.14 2.22 0.005 0.334 0.001 0.100 Indispensable AA Arg 80.01 81.14 81.51 81.47 84.60 1.47 0.264 0.105 0.306 0.299 His 74.47 77.25 76.69 75.63 79.34 1.74 0.319 0.351 0.818 0.208 Ile 71.98 73.76 73.72 74.15 77.59 2.23 0.472 0.433 0.403 0.424 Leu 75.39 75.88 75.50 76.90 78.99 1.87 0.622 0.594 0.307 0.482 Lys 77.28 78.36 77.99 79.47 82.50 2.08 0.404 0.627 0.188 0.462 ab b b ab a Met 84.18 82.11 82.01 85.54 86.42 1.26 0.048 0.478 0.006 0.454 Phe 73.63 74.69 74.30 75.71 78.81 2.01 0.378 0.333 0.252 0.272 Thr 65.37 67.04 66.40 67.24 73.81 2.97 0.232 0.129 0.333 0.120 Trp 76.53 77.06 76.68 77.11 80.25 2.45 0.770 0.627 0.457 0.484 Val 64.34 66.83 66.48 66.85 71.75 2.80 0.385 0.235 0.484 0.208 Dispensable AA Ala 67.05 68.86 67.49 68.98 72.71 2.42 0.482 0.596 0.329 0.290 Asp 70.83 72.54 71.41 72.27 75.19 2.51 0.764 0.885 0.448 0.498 Cys 64.17 63.10 64.24 63.50 69.93 3.16 0.519 0.363 0.327 0.483 Glu 80.07 80.87 81.87 82.11 85.21 1.71 0.277 0.527 0.166 0.543 Gly 36.06 47.25 54.26 54.35 58.48 6.58 0.102 0.422 0.366 0.779 Pro 76.35 72.49 76.06 77.10 79.57 2.22 0.321 0.143 0.173 0.299 Ser 69.08 70.77 69.48 70.74 74.98 2.54 0.461 0.500 0.370 0.250 Tyr 75.98 75.83 74.66 75.25 79.17 1.96 0.525 0.591 0.355 0.270 PC, positive control diet; NC1, negative control diet 1; NC2, negative control diet 2; NC1 + E, negative control diet 1 plus multienzyme; and NC2 + E, negative control diet 2 plus multienzyme. NC, main effects of negative control diet type; E, main effects of multienzyme; and NC × E, interaction between negative control diet type and multienzyme. abc Within a row, means without a common superscript differ (P < 0.05). Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Multienzyme supplemented diets for pigs (21.1 vs. 12.5 percentage points). Additionally, overall ADFI for NC1 diet without multienzyme was greater than that for NC1 diet with multien- zyme, whereas the overall ADFI for NC2 diet without multienzyme did not differ from that for NC1 diet with multienzyme. Thus, the greater ef- fect of multienzyme on the NC2 diet than on NC1 diet with regard to ADG could be attributed to: 1) the fact that the NC1 diet was not so deficient in NE and digestible nutrient content as evidenced by the similarity between NC1 and PC diets in ADG of pigs, 2)  greater increase in energy and nutrient digestibility for NC2 diet than for NC1 diet due to the supplemental multienzyme, and 3)  greater ADFI and hence energy and nutrient intake for unsupplemented NC1 diet than for multienzyme supplemented NC1 diet. The results from the cur- rent study are in contrast to those from the study of Kiarie et  al. (2012) who observed increased ADG of growing pigs from 22 to 55 kg BW, but not from 55 to 90 kg BW and hence for entire study period (22–90  kg BW) due to addition of a fiber-degrad- ing enzyme product to corn–barley-based diet that had lower DE value than the recommended value by large magnitude (293 kcal/kg). Also, the results from the current study are in contrast to those from the study of Jang et al. (2017) who did not observe an increased ADG of growing pigs from 26 to 122 kg BW due to addition of fiber-degrading en- zyme product to corn-based diet that had lower ME value than the recommended value by large magni- tude (103 kcal/kg). However, the enzyme product used in the current study contained xylanase, β-glu- canase, arabinofuranosidase, and phytase activities; whereas the enzyme product used in the studies of Kiarie et al, (2012) and Jang et al. (2017) contained only xylanase and β-glucanase activities, and xyla- nase activity, respectively. Thus, the differences be- tween the current study and that of Kiarie et  al. (2012) and Jang et al. (2017) with regard to the ef- fects of supplemental multienzyme on growth per- formance of pigs could be attributed to differences in enzyme activities of the multienzyme products. As previously mentioned, arabinofuranosidase can de-branch arabinoxylans, leading to increased availability of xylan backbones for xylanase deg- radation. Also, phytase can hydrolyze phytic acid, leading to release of phytic acid-bound nutrients for digestion and absorption (Woyengo and Nyachoti, 2011). Multienzyme supplementation increased overall ADG and hence final BW for NC2 diet to that of the PC diet. Thus, the NE, standardized ileal digestible AA, total Ca, and standardized total tract digestible P contents in corn–wheat–wheat Translate basic science to industry innovation Table 7. Effect of dietary treatment on ATTD of energy and nutrients a b Diets P-value ATTD, % PC NC1 NC2 NC1 + E NC2 + E SEM Diet NC E NC × E ab b b ab a Dry matter 75.62 73.79 73.90 75.76 76.43 0.703 0.057 0.907 0.006 0.591 a ab b a a Gross energy 76.05 74.32 73.65 75.81 75.67 1.252 0.077 0.650 0.027 0.658 a b ab a a Crude protein 66.21 62.70 65.57 66.53 67.13 1.022 0.046 0.253 0.019 0.442 Neutral detergent fiber 69.91 71.57 74.61 70.47 75.52 1.074 0.002 <0.001 0.735 0.302 c bc b c a Acid detergent fiber 40.18 43.18 48.90 43.62 53.91 1.835 <0.001 <0.001 0.103 0.120 bc cd d b e Ca 54.06 50.36 46.82 55.20 59.51 1.381 <0.001 0.727 <0.001 0.010 b c c b b P 38.19 22.60 27.51 35.05 48.56 2.733 <0.001 0.003 <0.001 0.181 PC, positive control diet; NC1, negative control diet 1; NC2, negative control diet 2; NC1 + E, negative control diet 1 plus multienzyme; and NC2 + E, negative control diet 2 plus multienzyme. NC, main effects of negative control diet type; E, main effects of multienzyme; and NC × E, interaction between negative control diet type and multienzyme. abc Within a row, means without a common superscript differ (P < 0.05). Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Jerez-Bogota et al. bran-based diets for pigs can be reduced by 124 kcal/ diet or NC2 diet, and the reason for this similarity kg, 5%, 18%, and 43%, respectively, without signifi- is not clear because the NC1 and NC2 diets were cant effects on growth performance of grow-finish formulated to contain lower levels of digestible pigs if the resulting low energy and nutrient diet is AA than the PC diet. It could have been due to the supplemented with the multienzyme that contain method that was used for ileal digesta collection. xylanase, β-glucanase, arabinofuranosidase, and Multienzyme supplementation increased the AID phytase activities. Supplementation of the NC1 of CP and methionine, and ATTD of GE and P for diet with multienzyme increased overall G:F, which the NC1 and NC2 diets, which was due to hydrolysis was due to the reduction in ADFI by the supple- of fiber by the NSPases and of phytate by phytase; mentation. Supplementation of the NC2 diet with as previously mentioned both NSPases and phytase multienzyme increased overall G:F, which was due were present in the multienzyme product used in to the increase in ADG by the supplementation. the current study. In plant feedstuffs, phytate, and Reason for the greater overall ADFI for NC1 diet nutrients such as starch and AA are located within without multienzyme than for NC1 diet with mul- cells, whereas NSP are mainly found in cell walls tienzyme is not clear. (Woyengo and Nyachoti, 2011). Thus, NSPases can The femur ash content and breaking strength hydrolyze NSP in cell walls to increase the accessi- for the PC diet significantly differ from those for bility of phytase to phytate that is located within NC1 diet or NC2 diet, which was partly due to the cells, thereby increasing the digestibility of phytate greater ATTD of P for the PC diet than for NC1 diet and availability of phytate-bound nutrients for di- or NC2 diet. Supplementation of the NC1 diet or gestion by gastric, pancreatic and small intestinal NC2 diet with multienzyme resulted in an increase mucosa enzymes. Also, the hydrolysis NSP by in femur ash content and breaking strength, which NSPases can result in increased availability of the could partly have been due to increase in P digest- NSP-encapsulated nutrients for digestion by gas- ibility by the supplementation. Similarly, She et al. tric, pancreatic, and small intestinal mucosa en- (2017) observed increased bone ash of weaned pigs zymes. Jang et  al. (2017) also reported increased due to dietary phytase supplementation. Woyengo ATTD of DM and P due to supplementation of a et al. (2008a, 2010) also reported increased bone ash combination of phytase and xylanase to corn-based due to addition of phytase to diets of broilers. The diet for grow–finish pigs. Woyengo et al. (2010) re - femur ash content and breaking strength for the for ported increased digestibility of P and bone ash multienzyme supplemented NC1 diet or multien- content of broilers due to addition of multienzyme zyme supplemented NC2 diet did not differ from that contained xylanase, β-glucanase, cellulose, and those for the PC diet, implying the NE, standard- pectinase to phytase-supplemented diet. However, ized ileal digestible AA, Ca, and standardized total the magnitude of improvement in AID and ATTD tract digestible P contents of the corn–wheat-based of nutrients for NC2 diet was generally greater than can be reduced by 124 kcal/kg, 5%, 18%, and 43%, that for NC1 diet. The NC2 diet was formulated to respectively, without significant effect on bone min- contain less NE and digestible AA than NC1 diet eralization if the resulting low energy and nutrient by partial replacement of corn and soybean meal in diet is supplemented with the enzyme product used NC1 diet with wheat bran and soybean hulls. Wheat in the current study. bran and soybean hulls are more fibrous than corn The AID values of energy and nutrients were and soybean meal (NRC, 2012), implying that more variable than the ATTD values energy and the former feedstuffs have greater content of sub- nutrients, which could have due to the method strate for the multienzyme than the latter. Thus, the (slaughter technique) of ileal digesta collection. greater effect of multienzyme on the NC2 diet than Ileal digesta of pigs can be collected by slaughter on NC1 diet with regard to AID and ATTD of nu- technique or ileal cannulation technique (Nyachoti trients could be attributed to the fact that the NC2 et  al., 1997). The AID values obtained from ileal diet contained more enzyme substrate than NC1 digesta that is collected by slaughter technique can diet. Zeng et al. (2018) similarly reported that sup- variable due to diurnal variation in nutrient digest- plementation of corn–wheat-based basal diet for ibility (Nyachoti et  al., 1997). The ATTD of GE growing pigs with enzyme product that contained for the PC diet was greater than that for NC1 diet galactanase, xylanase, mannanase, α-amylase, and or NC2 diet, which was expected because the NC1 cellulase activities did not improve nutrient digest- diet and NC2 diet were formulated to contain lower ibility when the basal diet did not contain wheat levels of NE than the PC diet. However, the AID of bran, but improved the AID of GE, NDF, and AA AA for the PC diet did not differ from that for NC1 when the basal diet contained 20% wheat bran. Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/4/2/txaa040/5817005 by guest on 30 April 2020 Multienzyme supplemented diets for pigs Also, Zeng et al. (2018) observed that supplemen- LITERATURE CITED tation of corn–wheat-based basal diet for growing AOAC. 2012. Official methods of analysis, 19th ed. Rockville pigs with phytase improved AID of phytic acid re- (MD): Association of Official Analytical Chemists gardless of whether or not the basal diet contained (AOAC) International. Appeldoorn, M.M., M.A. Kabel, D. Van Eylen, H. Gruppen, wheat bran, but the addition of an enzyme product and H.A.  Schols. 2010. Characterization of oligomeric that contained galactanase, xylanase, mannanase, xylan structures from corn fiber resistant to pretreat- α-amylase, and cellulose activities to phytase-sup- ment and simultaneous saccharification and fermenta- plemented diet improved AID of phytic acid only tion. J. Agric. Food Chem. 58:11294–11301. doi:10.1021/ for wheat bran-containing basal diet. jf102849x. Bedford,  M.R., and G.G.  Partridge. 2010. Feed enzyme, the The increased AID of CP observed in the cur- future: Bright hope or regulatory minefield. In: M. R. rent study due to multienzyme supplementation Bedford and G. G. Partridge, editors. 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