TY - JOUR
AU - Kim, Sung, Woo
AB - Abstract Forty pigs [10.7 ± 1.2 kg initial body weight (BW) at 6 wk of age] were used in a 21-d study to evaluate the effects of supplemental xylanase (Hostazym X 100, Huvepharma, Inc., Peachtree City, GA) in nursery diets on digesta viscosity, nutrient digestibility, health of the small intestine, and growth performance when supplemented with corn distillers’ dried grains with solubles (DDGS). Pigs were individually housed and randomly allotted to four treatments in a 2 × 2 factorial arrangement (n = 20/factor, 0% or 30% DDGS and 0 or 1,500 endo-pentosanase unit/kg xylanase as two factors) based on sex and initial BW. Feed intake and BW were recorded weekly. On day 15 of the study, TiO2 in diets (0.3%) was used as an indigestible marker to calculate apparent ileal digestibility (AID). Plasma samples were collected on day 19 to measure tumor necrosis factor-alpha (TNF-α), malondialdehyde, and peptide YY. On day 21, all pigs were euthanized to collect tissues from duodenum, jejunum, and colon to measure morphology, TNF-α, and malondialdehyde concentrations. Distal jejunal digesta were collected to measure viscosity. Ileum digesta were collected to measure AID of nutrients. During the entire period, supplemental xylanase increased (P < 0.05) average daily gain (ADG; 616 to 660 g/d) of nursery pigs, whereas DDGS (0 or 30%) did not affect ADG. On week 3, average daily feed intake (ADFI) was increased (P < 0.05) when fed DDGS (1,141 to 1,267 g/d) and there was an interaction (P < 0.05) between two factors indicating that supplemental xylanase decreased ADFI when DDGS was used in a diet. Use of DDGS increased (P < 0.05) viscosity [1.86 to 2.38 centipoise (cP)], whereas supplemental xylanase reduced (P < 0.05) viscosity (2.27 to 1.96 cP) of jejunal digesta. The AID of dry matter (DM) and gross energy (GE) were improved (P < 0.05) by supplemental xylanase. Plasma TNF-α was decreased (P < 0.05, 108.5 to 69.9 pg/mL) by supplemental xylanase. Use of DDGS reduced (P < 0.05) villus height:crypt depth ratio (1.46 to 1.27), whereas supplemental xylanase increased (P < 0.05) the crypt depth (360 to 404 µm) in duodenum. In conclusion, feeding a diet with 30% DDGS to nursery pigs for 3 wk had no negative effect on growth performance, whereas reduced AID of DM and GE, increased TNF-α level in colon tissue, and reduced the ratio of villus height to crypt depth. Dietary supplementation of xylanase reduced digesta viscosity improving AID of nutrients, reduced inflammatory response, and altered intestinal morphology, collectively improving ADG of nursery pigs regardless of the use of DDGS in a diet. Introduction Cereal grains and related coproducts are major feedstuffs but contain variable amounts of nonstarch polysaccharides (NSP). Nonstarch polysaccharides increase viscosity of digesta in the small intestine resulting in the reduction of digestibility and absorption of nutrients (Bakker et al., 1998; Kim et al., 2003; Passos et al., 2015a; Tiwari et al., 2018), interact with gut microflora, and modify the physiological function of the gut (Kiarie et al., 2013). Digesta viscosity slows down the digesta flow (Van Der Klis et al., 1993) and structures a barrier to the intestinal mucosa (Ikegami et al., 1990), causing morphological changes in the mucosal surface (Passos et al., 2015a; Duarte et al., 2019). Digesta viscosity also shown to increase pathogenic loads in the small intestine (Danicke et al., 1999), which may cause inflammatory response and oxidative stress in the small intestine of nursery pigs (Tiwari et al., 2018; Duarte et al., 2019). Use of feed enzymes has been an effective way to attenuate these concerns with NSP (Kim and Baker, 2003; Kim et al., 2003; de Vries et al., 2012). Xylan is a diverse group of NSP with a common feature of β-(1,4)-linked xylose residues (Scheller and Ulvskov, 2010), mainly existing in the form of arabinoxylans in cereal grains (Hoebler et al., 2000). Use of feed enzymes including xylanase has shown to reduce viscosity of digesta in nursery pigs (Passos et al., 2015a; Tiwari et al., 2018; Duarte et al., 2019). Among many different enzymes targeting xylans, endo-1,4-β-xylanase is most important due to their direct involvement in cleaving the glycosidic bonds and liberating short xylooligosaccharides (Topakas et al., 2013), which can serve potential prebiotics (Aachary and Prapulla, 2011; Dotsenko et al., 2018). Corn distillers’ dried grains with solubles (DDGS) became one of the major feedstuffs in the United States but include 33% NSP or 15% xylan (Pedersen et al., 2014). It has been demonstrated that DDGS can be used up to 25% in nursery pig diets without significant negative effects on the growth performance (Whitney and Shurson, 2004). However, the growth of nursery pigs can be impaired if DDGS is included higher than 25% due to its high NSP content, poor amino acid profile, and mycotoxin risks (Shurson, 2002). Feed enzymes targeting NSP could, therefore, reduce the negative impacts of DDGS on animal performance. It is hypothesized that xylanase supplementation could reduce digesta viscosity in the small intestine and thus enhances nutrient digestibility and functions of the small intestine in nursery pigs fed diets with DDGS. The objective of this study was to determine the effects of endo-1,4-β-xylanase on digesta viscosity, ileal nutrient digestibility, immune status, and health of the small intestine in nursery pigs when a diet is supplemented with 30% DDGS. Materials and Methods The experimental protocol was approved by North Carolina State University Animal Care and Use Committee. Animals and housing The experiment was conducted at the Metabolism Education Unit of North Carolina State University (Raleigh, NC). Forty crossbred barrows and gilts (10.7 ± 1.2 kg) at 6 wk of age were randomly allotted to four dietary treatments based on a 2 × 2 factorial arrangement. The first factor was endo-1,4-β-xylanase [0 or 1,500 endo-pentosanase unit (EPU)/kg complete feed; EFSA, 2013], and the second factor was DDGS (0% or 30%). The enzyme (Hostazym X 100 xylanase) is commercially available and obtained from Huvepharma USA (Peachtree City, GA). One EPU is defined as the amount of enzyme that releases low-molecular fragments from dyed xylan in amount equal to the amount of such fragments liberated from 1-unit enzyme standard under the conditions of the assay (50 °C and pH 4.7). The actual activity of xylanase in the diet was determined using photometric methods by the manufacturer (Huvepharma, Inc., Peachtree City, GA). Nutrient concentrations in all diets (Table 1) met the level of suggested requirements by NRC (2012). The experimental period was 21 d. Each treatment had 10 pens with one pig per pen. Pens (1.73 × 0.83 m) with metal screen floor were equipped with one nipple drinker and one self-feeder. Pigs had free access to water and feed. Body weight and feed intake were recorded weekly. Feed efficiency was calculated as gain to feed ratio (G:F). On day 15 of the study, titanium dioxide was blended into experimental diet (0.3%) as an indigestible marker for the calculation of ileal digestibility. Table 1. Composition of experimental diets (as-fed basis) . Treatment . DDGS, % . 0 . 30 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . Ingredient, %  Yellow corn, ground 62.10 62.09 37.41 37.40  Corn DDGS 0.00 0.00 30.00 30.00  Soybean meal 33.00 33.00 28.00 28.00  Xylanase1 0.00 0.01 0.00 0.01  l-Lys HCl 0.34 0.34 0.35 0.35  dl-Met 0.12 0.12 0.02 0.02  l-Thr 0.10 0.10 0.02 0.02  Poultry fat 2.00 2.00 2.00 2.00  Salt 0.22 0.22 0.22 0.22  Vitamin premix2 0.03 0.03 0.03 0.03  Trace mineral premix3 0.15 0.15 0.15 0.15  Dicalcium phosphate 1.10 1.10 0.64 0.64  Limestone 0.84 0.84 1.16 1.16  Total 100.00 100.00 100.00 100.00 Calculated content  ME, kcal/kg 3,384 3,384 3,395 3,395  Lys4, % 1.25 1.25 1.25 1.25  Met + Cys4, % 0.70 0.70 0.70 0.70  Trp4, % 0.23 0.23 0.23 0.23  Thr4, % 0.75 0.75 0.75 0.75  Ca, % 0.70 0.70 0.70 0.70  Available P, % 0.34 0.34 0.34 0.34 Analyzed content  DM, % 87.54 87.89 87.35 88.21  NDF, % 10.58 13.04 25.00 23.62  ADF, % 2.78 2.84 7.00 6.60  Xylanase activity, EPU/kg diet 270 1,980 310 2,050 . Treatment . DDGS, % . 0 . 30 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . Ingredient, %  Yellow corn, ground 62.10 62.09 37.41 37.40  Corn DDGS 0.00 0.00 30.00 30.00  Soybean meal 33.00 33.00 28.00 28.00  Xylanase1 0.00 0.01 0.00 0.01  l-Lys HCl 0.34 0.34 0.35 0.35  dl-Met 0.12 0.12 0.02 0.02  l-Thr 0.10 0.10 0.02 0.02  Poultry fat 2.00 2.00 2.00 2.00  Salt 0.22 0.22 0.22 0.22  Vitamin premix2 0.03 0.03 0.03 0.03  Trace mineral premix3 0.15 0.15 0.15 0.15  Dicalcium phosphate 1.10 1.10 0.64 0.64  Limestone 0.84 0.84 1.16 1.16  Total 100.00 100.00 100.00 100.00 Calculated content  ME, kcal/kg 3,384 3,384 3,395 3,395  Lys4, % 1.25 1.25 1.25 1.25  Met + Cys4, % 0.70 0.70 0.70 0.70  Trp4, % 0.23 0.23 0.23 0.23  Thr4, % 0.75 0.75 0.75 0.75  Ca, % 0.70 0.70 0.70 0.70  Available P, % 0.34 0.34 0.34 0.34 Analyzed content  DM, % 87.54 87.89 87.35 88.21  NDF, % 10.58 13.04 25.00 23.62  ADF, % 2.78 2.84 7.00 6.60  Xylanase activity, EPU/kg diet 270 1,980 310 2,050 1Xylanase source was Hostazym X 100 (Huvepharma USA, Peachtree City, GA) at 0.01% replacing corn for treatment diets. 2The vitamin premix provided the following per kilogram of complete diet: 6,613.8 IU of vitamin A; 992.0 IU of vitamin D3; 19.8 IU of vitamin E; 2.64 mg of vitamin K; 0.03 mg of vitamin B12; 4.63 mg of riboflavin; 18.52 mg of pantothenic acid; 24.96 mg of niacin; 0.07 mg of biotin. 3The trace mineral premix provided the following per kilogram of complete diet: 4.0 mg of Mn as manganous oxide; 165 mg of Fe as ferrous sulfate; 165 mg of Zn as zinc sulfate; 16.5 mg of Cu as copper sulfate; 0.30 mg of I as ethylenediamine dihydroiodide; and 0.30 mg of Se as sodium selenite. 4Standardized ileal digestibility. Open in new tab Table 1. Composition of experimental diets (as-fed basis) . Treatment . DDGS, % . 0 . 30 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . Ingredient, %  Yellow corn, ground 62.10 62.09 37.41 37.40  Corn DDGS 0.00 0.00 30.00 30.00  Soybean meal 33.00 33.00 28.00 28.00  Xylanase1 0.00 0.01 0.00 0.01  l-Lys HCl 0.34 0.34 0.35 0.35  dl-Met 0.12 0.12 0.02 0.02  l-Thr 0.10 0.10 0.02 0.02  Poultry fat 2.00 2.00 2.00 2.00  Salt 0.22 0.22 0.22 0.22  Vitamin premix2 0.03 0.03 0.03 0.03  Trace mineral premix3 0.15 0.15 0.15 0.15  Dicalcium phosphate 1.10 1.10 0.64 0.64  Limestone 0.84 0.84 1.16 1.16  Total 100.00 100.00 100.00 100.00 Calculated content  ME, kcal/kg 3,384 3,384 3,395 3,395  Lys4, % 1.25 1.25 1.25 1.25  Met + Cys4, % 0.70 0.70 0.70 0.70  Trp4, % 0.23 0.23 0.23 0.23  Thr4, % 0.75 0.75 0.75 0.75  Ca, % 0.70 0.70 0.70 0.70  Available P, % 0.34 0.34 0.34 0.34 Analyzed content  DM, % 87.54 87.89 87.35 88.21  NDF, % 10.58 13.04 25.00 23.62  ADF, % 2.78 2.84 7.00 6.60  Xylanase activity, EPU/kg diet 270 1,980 310 2,050 . Treatment . DDGS, % . 0 . 30 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . Ingredient, %  Yellow corn, ground 62.10 62.09 37.41 37.40  Corn DDGS 0.00 0.00 30.00 30.00  Soybean meal 33.00 33.00 28.00 28.00  Xylanase1 0.00 0.01 0.00 0.01  l-Lys HCl 0.34 0.34 0.35 0.35  dl-Met 0.12 0.12 0.02 0.02  l-Thr 0.10 0.10 0.02 0.02  Poultry fat 2.00 2.00 2.00 2.00  Salt 0.22 0.22 0.22 0.22  Vitamin premix2 0.03 0.03 0.03 0.03  Trace mineral premix3 0.15 0.15 0.15 0.15  Dicalcium phosphate 1.10 1.10 0.64 0.64  Limestone 0.84 0.84 1.16 1.16  Total 100.00 100.00 100.00 100.00 Calculated content  ME, kcal/kg 3,384 3,384 3,395 3,395  Lys4, % 1.25 1.25 1.25 1.25  Met + Cys4, % 0.70 0.70 0.70 0.70  Trp4, % 0.23 0.23 0.23 0.23  Thr4, % 0.75 0.75 0.75 0.75  Ca, % 0.70 0.70 0.70 0.70  Available P, % 0.34 0.34 0.34 0.34 Analyzed content  DM, % 87.54 87.89 87.35 88.21  NDF, % 10.58 13.04 25.00 23.62  ADF, % 2.78 2.84 7.00 6.60  Xylanase activity, EPU/kg diet 270 1,980 310 2,050 1Xylanase source was Hostazym X 100 (Huvepharma USA, Peachtree City, GA) at 0.01% replacing corn for treatment diets. 2The vitamin premix provided the following per kilogram of complete diet: 6,613.8 IU of vitamin A; 992.0 IU of vitamin D3; 19.8 IU of vitamin E; 2.64 mg of vitamin K; 0.03 mg of vitamin B12; 4.63 mg of riboflavin; 18.52 mg of pantothenic acid; 24.96 mg of niacin; 0.07 mg of biotin. 3The trace mineral premix provided the following per kilogram of complete diet: 4.0 mg of Mn as manganous oxide; 165 mg of Fe as ferrous sulfate; 165 mg of Zn as zinc sulfate; 16.5 mg of Cu as copper sulfate; 0.30 mg of I as ethylenediamine dihydroiodide; and 0.30 mg of Se as sodium selenite. 4Standardized ileal digestibility. Open in new tab Sample collection and preparation On day 19 of the study, blood samples of all pigs were collected from jugular vein with BD sterile vacutainer tubes with EDTA (BD, Franklin Lakes, NJ) for plasma. Blood samples were centrifuged at 3,000 × g for 15 min at 4 °C. Plasma samples were obtained and stored in −80 °C until analysis. Plasma samples were used to measure tumor necrosis factor-alpha (TNF-α), malondialdehyde (MDA), and peptide YY (PYY). On day 21, all pigs were euthanized by exsanguination after penetration of a captive bolt to head in order to collect tissues from duodenum, jejunum, and colon for determining gut morphology and concentrations of TNF-α and MDA. Tissue samples for TNF-α and MDA measurements were stored in −80 °C, and tissues for morphology were stored in 10% formalin buffer. Distal jejunal digesta (40 mL) were collected to measure viscosity immediately after the euthanasia. Ileal digesta (100 mL) were collected and frozen at −20 °C immediately after the euthanasia. Before the analysis, ileal digesta were freeze-dried and ground finely. Viscosity of digesta The method of measuring digesta viscosity was described by Passos et al. (2015a) with a viscometer (Brookfield Digital Viscometer, Model DV2TLV, Brookfield Engineering Laboratories Inc., Stoughton, MA). The samples were centrifuged at 3,000 × g for 5 min and then the supernatant was pipetted out to a 2-mL tube and centrifuged at 12,500 × g for 5 min. Viscometer was set at 25 °C, and 0.5 mL of digesta supernatant was placed in the viscometer. The final results were calculated as the average of viscosity at 45.0/s and 22.5/s shear rates. Apparent ileal digestibility Experimental diets and ileal digesta were ground and analyzed for dry matter (DM; Method 934.01, AOAC, 2006). Titanium dioxide concentration was measured following the procedure previously described in Chen et al. (2017). Gross energy was quantified using a calorimeter (Model 6200, Parr Instrument Company, Moline, IL). Duplicated samples of experimental diets and ileal digesta were analyzed sequentially for neutral detergent fiber (NDF) and acid detergent fiber (ADF) using the method of Van Soest et al. (1991) in a batch processor (Ankom Technology Corp, Fairport, NY). Apparent ileal digestibilities of DM, gross energy (GE), NDF, and ADF were calculated using concentration of titanium dioxide in feed and digesta following the equation previously described in Duarte et al. (2019). Immune parameter Tissue samples (500 mg) from the duodenum, jejunum, and colon were weighed and suspended into 1.0-mL phosphate-buffered saline solution (MP Biomedicals, LLC. Solon, OH). Samples were homogenized on ice. The homogenate was centrifuged at 14,000 × g for 30 °C. The supernatant was divided into three aliquot tubes to determine TNF-α, MDA, and protein concentration as described by Shen et al. (2012). As a mediator of inflammatory responses, TNF-α levels in plasma and tissue samples were measured by Porcine Immunoassay ELISA Kit (PTA00; R&D System Inc., Minneapolis, MN) as described by Weaver et al. (2014). The detection limit range for TNF-α enzyme linked immunosorbent assay (ELISA) was 2.8 to 5.0 pg/mL. Concentrations of TNF-α in tissue and plasma samples were expressed as pg/mg protein and pg/mL, respectively. As an oxidative stress indicator, MDA level was analyzed using thiobarbituric acid reactive substances assay kit (STA-330, Cell Biolabs, San Diego, CA) following the instruction of Zhao and Kim (2020). The detection range for this ELISA was 5 to 130 μM. Concentrations of MDA in tissue and plasma samples were expressed as μmol/g protein and μM, respectively. Concentrations of PYY in plasma sample were analyzed using Porcine PYY ELISA Kit (PP0179, NeoBiolab, Cambridge, MA) following the instruction of Sevarolli Loftus (2015). The detection limit for this kit was 1.0 pg/mL. Concentrations of PYY in plasma were expressed pg/mL. Morphological evaluation Two sections of jejunum per pig fixed in 10% buffered formalin were sent to the North Carolina State University Histology Laboratory (College of Veterinary Medicine, Raleigh, NC). The sections were dehydrated, embedded in paraffin, cut across the section to 5 mM thick slides, mounted on a polylysine-coated slide, and stained using hematoxylin and eosin dyes for morphological measurement according to Chen et al. (2017). Villus height, villus width, and crypt depth were measured using a microscope (Olympus CX31, Lumenera Corporation, Ottawa, Canada) with a camera Infinity 2-2 digital CCD following Kim et al. (2019a). Lengths of 10 well-oriented intact villi and their associated crypts were measured in each slide. The villus length was measured from the top of the villus to the villus–crypt junction. The villus width was measured in the middle of a villus, and the crypt depth was measured from the villi–crypt junction to the bottom of a crypt. Then, the villus height to crypt depth ratio was calculated. All analyses of the intestinal morphology were executed by the same person. The averages of the 10 measurements per pig were calculated and reported as one number per pig. Statistical analysis Data were analyzed using the mixed procedure of SAS (SAS Inst. Inc., Cary, NC). In this experiment, pigs were allotted to a randomized complete block design using initial body weight (BW) and sex as blocks. The experimental unit was the individual pig. Initial BW and sex were random effects, and two factors (xylanase and DDGS supplementation) were considered fixed effects. When there was an interaction between two factors, means of four treatments were separated using PDIFF option in SAS. Statistical differences were considered significant with P < 0.05. Probability that is less than 0.10 and equal to or greater than 0.05 was considered tendency. Results Growth performance Initial BW did not differ among treatments. Supplementation of xylanase or DDGS did not affect BW, average daily gain (ADG), and average daily feed intake (ADFI) in week 1 (Table 2). Supplementation of DDGS increased (P < 0.05) ADFI from 1,141 to 1,267 g/d in week 3. Supplementation of xylanase tended to increase (P = 0.061) ADG during week 2 and increased (P < 0.05) ADG from 616 to 660 g/d during the entire period. Supplementation of xylanase or DDGS did not affect G:F during the entire period. Table 2. Growth performance of nursery pigs fed diets supplemented with DDGS and xylanase . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . BW, kg  Initial 10.58 10.68 10.64 10.73 0.63 0.630 0.392 0.932  Week 1 13.07 13.35 13.31 13.44 0.49 0.480 0.376 0.746  Week 2 18.00 18.87 18.64 19.03 0.57 0.149 0.029 0.385  Week 3 23.14 24.48 23.93 24.66 0.68 0.288 0.029 0.499 ADG, g  Week 1 357 381 381 388 31 0.619 0.619 0.767  Week 2 704 788 762 799 32 0.273 0.061 0.450  Week 3 734 802 756 804 39 0.762 0.151 0.806  Overall 598 657 633 663 20 0.305 0.032 0.479 ADFI, g  Week 1 637 673 624 658 41 0.640 0.270 0.974  Week 2 1,015 1,098 1,066 1,127 60 0.412 0.150 0.827  Week 3 1,085a 1,196a 1,315b 1,218a 51 0.015 0.888 0.041  Overall 913 989 1,002 1,001 37 0.112 0.227 0.220 G:F  Week 1 0.562 0.570 0.610 0.571 0.030 0.420 0.604 0.442  Week 2 0.717 0.720 0.721 0.710 0.030 0.931 0.904 0.813  Week 3 0.682 0.675 0.579 0.671 0.034 0.128 0.223 0.158  Overall 0.665 0.669 0.633 0.664 0.020 0.380 0.402 0.498 . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . BW, kg  Initial 10.58 10.68 10.64 10.73 0.63 0.630 0.392 0.932  Week 1 13.07 13.35 13.31 13.44 0.49 0.480 0.376 0.746  Week 2 18.00 18.87 18.64 19.03 0.57 0.149 0.029 0.385  Week 3 23.14 24.48 23.93 24.66 0.68 0.288 0.029 0.499 ADG, g  Week 1 357 381 381 388 31 0.619 0.619 0.767  Week 2 704 788 762 799 32 0.273 0.061 0.450  Week 3 734 802 756 804 39 0.762 0.151 0.806  Overall 598 657 633 663 20 0.305 0.032 0.479 ADFI, g  Week 1 637 673 624 658 41 0.640 0.270 0.974  Week 2 1,015 1,098 1,066 1,127 60 0.412 0.150 0.827  Week 3 1,085a 1,196a 1,315b 1,218a 51 0.015 0.888 0.041  Overall 913 989 1,002 1,001 37 0.112 0.227 0.220 G:F  Week 1 0.562 0.570 0.610 0.571 0.030 0.420 0.604 0.442  Week 2 0.717 0.720 0.721 0.710 0.030 0.931 0.904 0.813  Week 3 0.682 0.675 0.579 0.671 0.034 0.128 0.223 0.158  Overall 0.665 0.669 0.633 0.664 0.020 0.380 0.402 0.498 1Ing = main effect of DDGS; Enz = main effect of xylanase; Ing × Enz = interaction effect between DDGS and xylanase. abMeans lacking a common superscript differ (P < 0.05). Open in new tab Table 2. Growth performance of nursery pigs fed diets supplemented with DDGS and xylanase . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . BW, kg  Initial 10.58 10.68 10.64 10.73 0.63 0.630 0.392 0.932  Week 1 13.07 13.35 13.31 13.44 0.49 0.480 0.376 0.746  Week 2 18.00 18.87 18.64 19.03 0.57 0.149 0.029 0.385  Week 3 23.14 24.48 23.93 24.66 0.68 0.288 0.029 0.499 ADG, g  Week 1 357 381 381 388 31 0.619 0.619 0.767  Week 2 704 788 762 799 32 0.273 0.061 0.450  Week 3 734 802 756 804 39 0.762 0.151 0.806  Overall 598 657 633 663 20 0.305 0.032 0.479 ADFI, g  Week 1 637 673 624 658 41 0.640 0.270 0.974  Week 2 1,015 1,098 1,066 1,127 60 0.412 0.150 0.827  Week 3 1,085a 1,196a 1,315b 1,218a 51 0.015 0.888 0.041  Overall 913 989 1,002 1,001 37 0.112 0.227 0.220 G:F  Week 1 0.562 0.570 0.610 0.571 0.030 0.420 0.604 0.442  Week 2 0.717 0.720 0.721 0.710 0.030 0.931 0.904 0.813  Week 3 0.682 0.675 0.579 0.671 0.034 0.128 0.223 0.158  Overall 0.665 0.669 0.633 0.664 0.020 0.380 0.402 0.498 . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . BW, kg  Initial 10.58 10.68 10.64 10.73 0.63 0.630 0.392 0.932  Week 1 13.07 13.35 13.31 13.44 0.49 0.480 0.376 0.746  Week 2 18.00 18.87 18.64 19.03 0.57 0.149 0.029 0.385  Week 3 23.14 24.48 23.93 24.66 0.68 0.288 0.029 0.499 ADG, g  Week 1 357 381 381 388 31 0.619 0.619 0.767  Week 2 704 788 762 799 32 0.273 0.061 0.450  Week 3 734 802 756 804 39 0.762 0.151 0.806  Overall 598 657 633 663 20 0.305 0.032 0.479 ADFI, g  Week 1 637 673 624 658 41 0.640 0.270 0.974  Week 2 1,015 1,098 1,066 1,127 60 0.412 0.150 0.827  Week 3 1,085a 1,196a 1,315b 1,218a 51 0.015 0.888 0.041  Overall 913 989 1,002 1,001 37 0.112 0.227 0.220 G:F  Week 1 0.562 0.570 0.610 0.571 0.030 0.420 0.604 0.442  Week 2 0.717 0.720 0.721 0.710 0.030 0.931 0.904 0.813  Week 3 0.682 0.675 0.579 0.671 0.034 0.128 0.223 0.158  Overall 0.665 0.669 0.633 0.664 0.020 0.380 0.402 0.498 1Ing = main effect of DDGS; Enz = main effect of xylanase; Ing × Enz = interaction effect between DDGS and xylanase. abMeans lacking a common superscript differ (P < 0.05). Open in new tab Viscosity of digesta Supplementation of DDGS increased (P < 0.05) the viscosity of jejunal digesta from 1.86 to 2.38 centipoise (cP), whereas supplementation of xylanase reduced (P < 0.05) the viscosity of jejunal digesta from 2.27 to 1.96 cP. There was no interaction between xylanase and DDGS supplementation (Table 3). Table 3. Viscosity of jejunal digesta (cP) in nursery pigs fed diets supplemented with DDGS and xylanase . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . Viscosity, cP 1.97 1.74 2.57 2.18 0.17 0.004 0.023 0.569 . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . Viscosity, cP 1.97 1.74 2.57 2.18 0.17 0.004 0.023 0.569 1Ing = main effect of DDGS; Enz = main effect of xylanase; Ing × Enz = interaction effect between DDGS and xylanase. Open in new tab Table 3. Viscosity of jejunal digesta (cP) in nursery pigs fed diets supplemented with DDGS and xylanase . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . Viscosity, cP 1.97 1.74 2.57 2.18 0.17 0.004 0.023 0.569 . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . Viscosity, cP 1.97 1.74 2.57 2.18 0.17 0.004 0.023 0.569 1Ing = main effect of DDGS; Enz = main effect of xylanase; Ing × Enz = interaction effect between DDGS and xylanase. Open in new tab Apparent ileal digestibility Supplementation of DDGS reduced (P < 0.05) apparent ileal digestibility (AID) of DM and GE from 72.3% to 67.8% and from 71.7% to 66.2%, respectively. Supplementation of xylanase tended to improve (P = 0.072) AID of DM and increased (P < 0.05) AID of GE and NDF from 66.5% to 71.5% and from 45.2% to 52.4%, respectively. There was no interaction between xylanase and DDGS supplementation in ileal digestibility of DM, GE, NDF, and ADF (Table 4). Table 4. AID of DM, GE, NDF, and ADF in nursery pigs fed diets supplemented with DDGS and xylanase . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . AID, %  DM 71.3 75.2 65.8 69.7 2.1 0.013 0.072 0.993  GE 69.3 74.1 63.6 68.8 2.3 0.022 0.035 0.922  NDF 48.6 51.2 41.7 53.6 3.1 0.469 0.025 0.140  ADF 36.5 38.4 38.1 41.0 2.2 0.376 0.315 0.833 . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . AID, %  DM 71.3 75.2 65.8 69.7 2.1 0.013 0.072 0.993  GE 69.3 74.1 63.6 68.8 2.3 0.022 0.035 0.922  NDF 48.6 51.2 41.7 53.6 3.1 0.469 0.025 0.140  ADF 36.5 38.4 38.1 41.0 2.2 0.376 0.315 0.833 1Ing = main effect of DDGS; Enz = main effect of xylanase; Ing × Enz = interaction effect between DDGS and xylanase. Open in new tab Table 4. AID of DM, GE, NDF, and ADF in nursery pigs fed diets supplemented with DDGS and xylanase . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . AID, %  DM 71.3 75.2 65.8 69.7 2.1 0.013 0.072 0.993  GE 69.3 74.1 63.6 68.8 2.3 0.022 0.035 0.922  NDF 48.6 51.2 41.7 53.6 3.1 0.469 0.025 0.140  ADF 36.5 38.4 38.1 41.0 2.2 0.376 0.315 0.833 . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . AID, %  DM 71.3 75.2 65.8 69.7 2.1 0.013 0.072 0.993  GE 69.3 74.1 63.6 68.8 2.3 0.022 0.035 0.922  NDF 48.6 51.2 41.7 53.6 3.1 0.469 0.025 0.140  ADF 36.5 38.4 38.1 41.0 2.2 0.376 0.315 0.833 1Ing = main effect of DDGS; Enz = main effect of xylanase; Ing × Enz = interaction effect between DDGS and xylanase. Open in new tab Immune parameter Supplementation of DDGS increased (P < 0.05) TNF-α level from 7.08 to 8.06 pg/mg protein in the colon tissue (Table 5). Supplementation of xylanase did not affect concentrations of TNF-α and MDA in the jejunum and colon tissue, whereas plasma TNF-α and PYY concentrations were decreased (P < 0.05) by supplementation of xylanase. Table 5. TNF-α, MDA, and PYY in nursery pigs fed diets supplemented with DDGS and xylanase . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . Plasma  TNF-α, pg/mL 107.58 66.63 109.31 73.11 22.72 0.809 0.028 0.889  MDA, µM 13.79 12.40 12.21 11.95 2.89 0.727 0.777 0.847  PYY, pg/mL 56.43 33.24 49.71 45.74 6.07 0.642 0.036 0.131 Jejunum  TNF-α, pg/mg 7.05 6.36 6.12 6.79 0.56 0.666 0.989 0.233  MDA, µmol/g protein 1.84 1.87 2.17 2.10 0.36 0.449 0.967 0.890 Colon  TNF-α, pg/mg 6.54 7.61 8.05 8.07 0.53 0.041 0.247 0.266  MDA, µmol/g protein 2.24 2.93 3.38 2.97 0.52 0.255 0.781 0.291 . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . Plasma  TNF-α, pg/mL 107.58 66.63 109.31 73.11 22.72 0.809 0.028 0.889  MDA, µM 13.79 12.40 12.21 11.95 2.89 0.727 0.777 0.847  PYY, pg/mL 56.43 33.24 49.71 45.74 6.07 0.642 0.036 0.131 Jejunum  TNF-α, pg/mg 7.05 6.36 6.12 6.79 0.56 0.666 0.989 0.233  MDA, µmol/g protein 1.84 1.87 2.17 2.10 0.36 0.449 0.967 0.890 Colon  TNF-α, pg/mg 6.54 7.61 8.05 8.07 0.53 0.041 0.247 0.266  MDA, µmol/g protein 2.24 2.93 3.38 2.97 0.52 0.255 0.781 0.291 1Ing = main effect of DDGS; Enz = main effect of xylanase; Ing × Enz = interaction effect between DDGS and xylanase. Open in new tab Table 5. TNF-α, MDA, and PYY in nursery pigs fed diets supplemented with DDGS and xylanase . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . Plasma  TNF-α, pg/mL 107.58 66.63 109.31 73.11 22.72 0.809 0.028 0.889  MDA, µM 13.79 12.40 12.21 11.95 2.89 0.727 0.777 0.847  PYY, pg/mL 56.43 33.24 49.71 45.74 6.07 0.642 0.036 0.131 Jejunum  TNF-α, pg/mg 7.05 6.36 6.12 6.79 0.56 0.666 0.989 0.233  MDA, µmol/g protein 1.84 1.87 2.17 2.10 0.36 0.449 0.967 0.890 Colon  TNF-α, pg/mg 6.54 7.61 8.05 8.07 0.53 0.041 0.247 0.266  MDA, µmol/g protein 2.24 2.93 3.38 2.97 0.52 0.255 0.781 0.291 . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . Plasma  TNF-α, pg/mL 107.58 66.63 109.31 73.11 22.72 0.809 0.028 0.889  MDA, µM 13.79 12.40 12.21 11.95 2.89 0.727 0.777 0.847  PYY, pg/mL 56.43 33.24 49.71 45.74 6.07 0.642 0.036 0.131 Jejunum  TNF-α, pg/mg 7.05 6.36 6.12 6.79 0.56 0.666 0.989 0.233  MDA, µmol/g protein 1.84 1.87 2.17 2.10 0.36 0.449 0.967 0.890 Colon  TNF-α, pg/mg 6.54 7.61 8.05 8.07 0.53 0.041 0.247 0.266  MDA, µmol/g protein 2.24 2.93 3.38 2.97 0.52 0.255 0.781 0.291 1Ing = main effect of DDGS; Enz = main effect of xylanase; Ing × Enz = interaction effect between DDGS and xylanase. Open in new tab Morphological evaluation Supplementation of DDGS tended to reduce (P = 0.063) villus height and reduced (P < 0.05) villus height/crypt depth ratio in the duodenum (Table 6). Supplementation of xylanase increased (P < 0.05) crypt depth in the duodenum. Supplementation of xylanase tended to decrease (P = 0.052) crypt depth in the jejunum with 30% DDGS, whereas it had the opposite effect without DDGS. Table 6. Villus height and width, crypt depth, and the villus height to crypt depth ratio from duodenum, jejunum, and colon in nursery pigs fed diets supplemented with DDGS and xylanase . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . Duodenum  Villus height, µm 529 523 475 504 38 0.063 0.542 0.366  Villus width, µm 174 172 168 171 11 0.673 0.977 0.776  Crypt depth, µm 359 378 360 430 21 0.168 0.025 0.191  Height:depth ratio 1.49 1.42 1.34 1.20 0.08 0.027 0.210 0.685 Jejunum  Villus height, µm 452 481 454 462 18 0.634 0.318 0.558  Villus width, µm 120 118 120 116 3 0.833 0.397 0.703  Crypt depth, µm 218A 231B 237B 210A 11 0.885 0.479 0.052  Height:depth ratio 2.11 2.14 1.93 2.24 0.14 0.754 0.222 0.272 Colon  Crypt depth, µm 422 406 432 418 13 0.430 0.277 0.962 . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . Duodenum  Villus height, µm 529 523 475 504 38 0.063 0.542 0.366  Villus width, µm 174 172 168 171 11 0.673 0.977 0.776  Crypt depth, µm 359 378 360 430 21 0.168 0.025 0.191  Height:depth ratio 1.49 1.42 1.34 1.20 0.08 0.027 0.210 0.685 Jejunum  Villus height, µm 452 481 454 462 18 0.634 0.318 0.558  Villus width, µm 120 118 120 116 3 0.833 0.397 0.703  Crypt depth, µm 218A 231B 237B 210A 11 0.885 0.479 0.052  Height:depth ratio 2.11 2.14 1.93 2.24 0.14 0.754 0.222 0.272 Colon  Crypt depth, µm 422 406 432 418 13 0.430 0.277 0.962 1Ing = main effect of DDGS; Enz = main effect of xylanase; Ing × Enz = interaction effect between DDGS and xylanase. ABMeans lacking a common superscript tend to differ (0.05 ≤ P < 0.10). Open in new tab Table 6. Villus height and width, crypt depth, and the villus height to crypt depth ratio from duodenum, jejunum, and colon in nursery pigs fed diets supplemented with DDGS and xylanase . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . Duodenum  Villus height, µm 529 523 475 504 38 0.063 0.542 0.366  Villus width, µm 174 172 168 171 11 0.673 0.977 0.776  Crypt depth, µm 359 378 360 430 21 0.168 0.025 0.191  Height:depth ratio 1.49 1.42 1.34 1.20 0.08 0.027 0.210 0.685 Jejunum  Villus height, µm 452 481 454 462 18 0.634 0.318 0.558  Villus width, µm 120 118 120 116 3 0.833 0.397 0.703  Crypt depth, µm 218A 231B 237B 210A 11 0.885 0.479 0.052  Height:depth ratio 2.11 2.14 1.93 2.24 0.14 0.754 0.222 0.272 Colon  Crypt depth, µm 422 406 432 418 13 0.430 0.277 0.962 . Treatment . . . . . DDGS, % . 0 . 30 . . P-value1 . Xylanase, EPU/kg diet . 0 . 1,500 . 0 . 1,500 . SEM . Ing . Enz . Ing × Enz . Duodenum  Villus height, µm 529 523 475 504 38 0.063 0.542 0.366  Villus width, µm 174 172 168 171 11 0.673 0.977 0.776  Crypt depth, µm 359 378 360 430 21 0.168 0.025 0.191  Height:depth ratio 1.49 1.42 1.34 1.20 0.08 0.027 0.210 0.685 Jejunum  Villus height, µm 452 481 454 462 18 0.634 0.318 0.558  Villus width, µm 120 118 120 116 3 0.833 0.397 0.703  Crypt depth, µm 218A 231B 237B 210A 11 0.885 0.479 0.052  Height:depth ratio 2.11 2.14 1.93 2.24 0.14 0.754 0.222 0.272 Colon  Crypt depth, µm 422 406 432 418 13 0.430 0.277 0.962 1Ing = main effect of DDGS; Enz = main effect of xylanase; Ing × Enz = interaction effect between DDGS and xylanase. ABMeans lacking a common superscript tend to differ (0.05 ≤ P < 0.10). Open in new tab Discussion Global feed use of cereal grains and coproducts reaches 870 million tons, which takes about 34% of total cereal production (Food and Agriculture Organization of the UN, 2015). Cereal grains contain variable amounts of NSP, such as arabinoxylans and β-glucans, which are associated with the cell wall structure. Vegetable protein supplements, such as soybean meal, are often used in monogastric animal feeds, which also contain substantial amounts of NSP such as β-mannans. Cereal coproducts, such as DDGS, contain high levels of NSP and protein (Choct, 2015) and their use in animal feeds is increasing (Kim et al., 2019b). However, these NSP are not hydrolyzed by endogenous enzymes from animals and, consequently, become available as substrates for microbial fermentation in the intestine. Increased NSP intake also contributes to higher viscosity of digesta due to the hydration of NSP (Passos et al., 2015a; Duarte et al., 2019). Viscosity, referring to the ability of polysaccharides to thicken or form gels when mixed with fluids, is positively related to the molecular weight of polysaccharides in solution at equal concentrations as well as its solubility (Dikeman and Fahey, 2006). It was believed that soluble NSP mainly exhibits the higher viscosity values (Dikeman et al., 2006). Distiller’s dried grains with solubles are often used as feedstuffs in animal production. The use of DDGS in animal feed is about 44 million tons in 2015, and nearly one third is utilized in pig production (Wisner, 2015). Results of many experiments have indicated that supplementation of 25% corn DDGS in nursery diets did not influence ADG, ADFI, or G:F compared with the corn–soybean meal basal diet (Whitney and Shurson, 2004; Jones et al., 2010). Growth performance of pigs with DDGS did not differ from pigs fed corn–soybean meal diet in this study. However, decreased ADFI of piglets fed with 30% DDGS during late nursery period were reported by Tran et al. (2012). Inconsistent effects of DDGS on growth performance are probably due to the duration of feeding experimental diets and also the variability of nutrient contents in DDGS (Cromwell et al., 2011). This study aimed to investigate the impacts of NSP from DDGS and the use of xylanase mainly on intestinal health. In order to determine the changes in intestinal health affected by dietary challenges or intervention, feeding duration is often limited to 3 wk before the intestine recovers (Shen et al., 2014; Passos et al., 2015a; Tiwari et al., 2018; Jang and Kim, 2020) and the limited feeding duration could be reasons for lack of growth responses by dietary effects. Variability of nutrient contents in DDGS usually comes from different processing technology as well as the parent grain, of which especially, the difference in amount and type of NSP contents of the corresponding grain (Cromwell et al., 2011; Jha and Berrocoso, 2015). The compositional profiles of NSP revealed that DDGS had 4.1- and 4.5-fold higher content of total NSP and soluble NSP than corn, respectively. The soluble xylose and arabinose contents in DDGS were 7.6- and 5.8-fold higher than corn, respectively (Pedersen et al., 2014). The solubility of xylans is affected by the patterns of intramolecular and intermolecular hydrogen bonds, which are natural or created during the drying process of the extracted polysaccharide. Moreover, the heteroxylans with a low-level substitution with arabinose and glucose in their backbone are more water soluble (Ebringerova and Heinze, 2000). Ethanol production with dry-grind corn process, as the major source of DDGS, includes multiple steps of grinding, cooking, liquefaction, saccharification, fermentation, distillation of ethanol, and drying to form DDGS (Rausch and Belyea, 2006). All of these steps increase solubility of xylose and arabinose in DDGS resulting in increased viscosity (Pedersen et al., 2014). Viscous dietary NSP can reduce nutrient digestion and absorption, alter intestinal development, and change microflora of the gut in monogastric animals, although there is some difference between pigs and poultry (McDonald et al., 2001; Passos et al., 2015a; Duarte et al., 2019). In poultry, solubility of NSP in the digestive environment is one of the key characteristics affecting growth performance because of their effects on rate of feed passage in the upper part of the gastrointestinal tract and fermentation activity in the distal part (Saki et al., 2011). Barley, wheat, rye, and oats have high levels of NSP causing increased digesta viscosity. Increased digesta viscosity could decrease digesta passage rate, digestive enzymatic activities, and nutrient digestibility, which, in turn, reduce feed efficiency and growth of birds (Bedford and Schulze, 1998). Increased digesta viscosity also enhance DM flow and endogenous losses from both endogenous and exogenous sources (Montagne et al., 2003) and increase digesta retention time facilitates bacterial colonization and activity in the small intestine (Waldenstedt et al., 2000). In pigs, it was believed that impacts of digesta viscosity are likely to be a lower order of importance to the responses observed in poultry due to the higher water content in pig digesta (Partridge, 2001). The addition of soluble NSP to nursery diets, however, significantly increased intestinal viscosity, altered the structure of the intestine, and provided a microenvironment that favored the proliferation of pathogenic Escherichia coli (McDonald et al., 2001; Passos et al., 2015a; Duarte et al., 2019), indicating that viscosity should not be underestimated in the pig as an influence on productive performance. Results about digesta viscosity of pig are usually focused on corn–soybean meal (Passos et al., 2015a; Duarte et al., 2019), wheat and corn mixed with DDGS (Agyekum et al., 2012), wheat (Mavromichalis et al., 2000), or rye-wheat-based diets (Bartelt et al., 2002), whereas reports on the effect of DDGS supplementation on digesta viscosity were seldom found. In the present study, supplementation of 30% DDGS increased jejunal digesta viscosity from 1.97 to 2.57 cP when compared with a corn–soybean meal diet. Increased digesta viscosity was associated with decreased AID of DM and energy. Supplementation of DDGS did not affect the AID of NDF and ADF, indicating that 30% DDGS supplementation may decrease the digestibility of other components, such as protein, fat, and starch by limiting the accessibility of digestive enzymes to these nutrients due to increased digesta viscosity. It was also observed that increased digesta viscosity and reduced nutrient digestibility might be balanced by increased ADFI at week 3 (Table 2), in order to achieve the similar ADG of pigs fed with corn–soybean meal. In an early study by Sorensen (1953), xylanase was shown to hydrolyze xylan in wheat straw to oligosaccharide. Xylanase has been widely used in paper and pulp industry, biofuel industry, pharmaceutical industry, as well as food and feed industry (Motta et al., 2013). The type of xylanase used in this study was endo-1,4-β-xylanase and its direct involvement in xylan hydrolysis is to cleave the glycosidic bonds and liberate short xylooligosaccharides or oligosaccharides (Motta et al., 2013). One important finding of this study was that supplemental xylanase reduced viscosity of jejunal digesta in nursery pigs fed diets with 30% DDGS, which is beneficial to pigs by reducing the antinutritional effects of NSP and thus enhancing nutrient utilization (Passos et al., 2015b). Bedford and Schulze (1998) reported that supplemental xylanase to broiler diets increased the proliferation of beneficial microflora affecting intestinal immune status. In this study, AID of NDF and GE increased by 7% and 5%, respectively, which was similar to the previous report that supplemental xylanase improved AID of NDF in weanling pigs by 5% (Diebold et al., 2004). However, Yin et al. (2000) showed the improvements in AID of DM and crude protein were less than 2% when xylanase was supplemented in the diet for older pigs (26 kg). This could be related to an increased capability of gut-associated microbiota handling NSP in the small intestine of pigs (Niu et al., 2015); therefore, there is less scope for enzyme action. Tumor necrosis factor-α, known as a proinflammatory cytokine, is produced in response to tissue damage, affects intestinal epithelial permeability and ion transport, and functions as a regulator in growth and differentiation of many immune cells (McKay and Baird, 1999). It is generally accepted that the inflammation response is mediated by an increased production of proinflammatory cytokines (Shen et al., 2014; Jang and Kim, 2020; Kim et al., 2019a). Activation of the immune system is implicated in less efficient assimilation of nutrients (Cook et al, 1993), which undoubtedly reduces the energetic efficiency of tissue deposition. The results from this study showed that the inclusion of DDGS increased TNF-α level in colon tissue, indicating intestinal immune systems were stimulated by a higher level of NSP from added DDGS, thus presents a typical inflammation response. Interestingly, supplemental xylanase suppressed the plasma TNF-α concentration. The possible mechanism of the action of xylanase on regulating immune response is the reduction of NSP and the increase of oligosaccharides. These molecules might inhibit tumor formation and enhance defense mechanisms against bacterial challenge (Wang et al., 2010). However, TNF-α concentration in the intestinal mucosa was not altered by xylanase supplementation. With limited evidences, it is currently speculated that the changes of plasma TNF-α concentration by xylanase supplementation is contributed from the intestinal response. Additionally, this study found that gut hormone PYY in plasma was decreased by supplemental xylanase. In pigs, PYY is a gastrointestinal peptide that is present mainly in endocrine cells of distal intestine (Wewer Albrechtsen et al., 2016) and may mediate gastric emptying, inhibit feed intake, and also interfere with other hormones including cholecystokinin and gastrin (Ballantyne, 2006). The decreased PYY level in nursery pigs by dietary xylanase supplementation was also reported by May et al. (2015). However, Singh et al. (2012) observed that supplementation of xylanase increased plasma PYY concentration of broilers. The inconsistent observations suggested that the action of xylanase on PYY regulation is probably different between pigs and broilers. Although PYY is associated with feed intake, it seems not the only determining factor because ADFI in this study was not affected by decreased circulating PYY concentration. Detailed mechanisms are not understood and should be further investigated to understand the inconsistent responses between pigs and poultry. Supplementation of xylanase was found to reduce the antioxidant stress in rat (Ardiansyah et al., 2006), but in the present study, MDA, as an indicator of lipid peroxidation, was not affected by the inclusion of DDGS or xylanase. Production of MDA was linked with unsaturated fatty acid in vitro (Palozza et al., 1995). Corn DDGS used in this study contains about 10% corn oil, which contains high levels of polyunsaturated fatty acids (particularly linoleic acid) that are vulnerable to lipid peroxidation (Stein and Shurson, 2009). One potential speculation could be that dietary oxidative challenge from DDGS which causes free radical production and consequently cell damage was insufficient. Thus, this redox homeostasis altering was not critical on pigs’ health and growth performance. One of the main objectives in this study was to evaluate whether supplemental xylanase would have an influence on intestinal morphology of nursery pigs. This study showed that 30% DDGS supplementation decreased villus height/crypt depth ratio and tended to decrease villus height in the duodenum. However, it was interesting to observe that supplemental xylanase increased crypt depth in the duodenum. The decreased villus height/crypt depth ratio indicates an impaired contact surface area between endogenous fluid and nutrients, whereas a deeper crypt might be due to an increased rate of cell renewal that is the result of increased cell division in the crypts (Pluske et al., 1997). Moreover, a trend of the interaction between DDGS and xylanase supplements was observed, indicating that supplementation of xylanase with DDGS decreased crypt depth in the jejunum. As NSP interacts both with the mucosa and the microflora, it has an important role in the regulation of gut health, and causes a marked effect on the intestinal mucosa, increasing crypt-cell proliferation and mucosal protein synthesis rate (Montagne et al., 2003). Therefore, supplemental xylanase might affect intestine morphology of pigs by breaking down cell wall NSP (Högberg and Lindberg, 2004; Passos et al., 2015a; Duarte et al., 2019). The duodenum is largely responsible for the continuous digestion process, whereas the jejunum is mainly responsible for both nutrient digestion and absorption (Yen, 2000). Therefore, faster renewal rate in the duodenum could be beneficial for endogenous enzyme secretion, improving nutrient digestion. These findings collectively support the hypothesis that feeding a diet with 30% DDGS to nursery pigs for 3 wk reduced the ileal digestibility of DM and GE, increased intestinal inflammation, and impaired intestinal morphology. Dietary supplementation of xylanase attenuated these negative impacts of high-level DDGS in nursery diets by reducing digesta viscosity improving AID of nutrients, reducing inflammatory response, altering intestinal morphology, and finally improving ADG of nursery pigs. Abbreviations Abbreviations ADF acid detergent fiber ADFI average daily feed intake ADG average daily gain AID apparent ileal digestibility BW body weight cP centipoise DDGS distillers’ dried grains with solubles DM dry matter ELISA enzyme linked immunosorbent assay EPU endo-pentosanase unit GE gross energy G:F gain to feed ratio MDA malondialdehyde NDF neutral detergent fiber NSP nonstarch polysaccharides PYY peptide YY TNF-α tumor necrosis factor-alpha Acknowledgments Financial supports for this research from Huvepharma, Inc., Peachtree City, GA, USA and North Carolina Agricultural Foundation, Raleigh, NC, USA are greatly appreciated. Conflict of interest statement The authors declare no conflict of interest. Literature Cited Aachary , A. A. , and S. G. Prapulla. 2011 . Xylooligosaccharides (XOS) as an emerging prebiotic: Microbial synthesis, utilization, structural characterization, bioactive properties, and applications . Compr. Rev. Food Sci. 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TI - Effects of supplemental xylanase on health of the small intestine in nursery pigs fed diets with corn distillers’ dried grains with solubles
JF - Journal of Animal Science
DO - 10.1093/jas/skaa185
DA - 2020-06-01
UR - https://www.deepdyve.com/lp/oxford-university-press/effects-of-supplemental-xylanase-on-health-of-the-small-intestine-in-HO0tSPvPIn
VL - 98
IS - 6
DP - DeepDyve
ER -