Effects of dietary supplementation of an enzyme blend on the ileal and fecal
digestibility of nutrients in growing pigs
F. Ji, D. P. Casper, P. K. Brown, D. A. Spangler, K. D. Haydon and J. E. Pettigrew
J Anim Sci 2008.86:1533-1543.
doi: 10.2527/jas.2007-0262 originally published online Mar 14, 2008;
The online version of this article, along with updated information and services, is located on
the World Wide Web at:
http://jas.fass.org/cgi/content/full/86/7/1533
www.asas.org
Downloaded from jas.fass.org at Serials/Acq. Dept., Library on May 1, 2009.
Effects of dietary supplementation of an enzyme blend on the ileal
and fecal digestibility of nutrients in growing pigs1
F. Ji,* D. P. Casper,† P. K. Brown,† D. A. Spangler,† K. D. Haydon,‡ and J. E. Pettigrew*2
*University of Illinois, Department of Animal Sciences, Urbana 61801; †Agri-King Inc.,
Fulton, IL 61252; and ‡Prince Agri Products, Quincy, IL 62306
ABSTRACT: The objective of this experiment was to
determine the effect of a β-glucanase-protease enzyme
blend product (EBP) on fecal digestibility (FD), apparent ileal digestibility (AID), standardized ileal digestibility, and digestibility in the hindgut of growing pigs.
Twelve ileal-cannulated, growing barrows (38.2 ± 0.5
kg) were housed in individual metabolism crates,
blocked by previous feed intake into 3 groups with 4
pigs each, and randomly assigned to 1 of 4 treatments
within a square (group) of 3 replications of 4 × 4 Latin
square design. Treatments were basal diet (Basal),
Basal + 0.05% of EBP (0.05% EBP), Basal + 0.10% of
EBP (0.10% EBP), and hydrolyzed casein for measurement of endogenous amino acids. The Basal diet consisted of corn and soybean meal and was calculated to
have 3.36 Mcal of ME/kg and 1.1% of total lysine, asfed basis. Feed intake of each replicate of the Latin
square during the first period was 85% of the minimum
feed intake of the 4 pigs during the preliminary period
and was equalized within each square. The feeding level
was increased by 100 g/d in each subsequent period.
Each of the experimental periods was 14 d, including
4 d of dietary adaptation, 5 d of fecal collection, 3 d of
transition period, and 2 d of ileal collection. Ileal efflu-
ents were collected continuously for the same 12-h interval each day. Pigs fed the EBP demonstrated increased (P < 0.05) FD of DM, OM, energy, CP, nonfiber
carbohydrate, total dietary fiber, insoluble dietary fiber, acid-hydrolyzed fat, ash, Ca, and P compared with
pigs fed Basal. The AID of NDF and hemicellulose was
increased (P < 0.05) by supplying the EBP either at
0.05 or 0.10% in the diets, but AID of DM and energy
was not increased. The AID of acid-hydrolyzed fat
tended to be greater (P = 0.051) for the pigs fed the
EBP than for those fed Basal. Ileal digestibility of most
amino acids was not affected by treatment, but the EBP
reduced the apparent and standardized digestibility of
methionine, alanine, and serine (P < 0.05). The difference between FD and AID of hemicellulose was lower
(P < 0.05) for the pigs fed the EBP than for those fed
Basal. These results demonstrated that the EBP fed to
growing pigs improved the FD of DM, OM, energy, CP,
nonfiber carbohydrate, total dietary fiber, acid-hydrolyzed fat, Ca, and P, and the AID of NDF and hemicellulose, but the standardized ileal digestibility of
amino acids was not improved by supplying the EBP
in corn-soybean meal-based diets of growing pigs.
Key words: digestibility, enzyme, growing pig, nutrient
©2008 American Society of Animal Science. All rights reserved.
INTRODUCTION
The effects of enzymes on the nutrient digestion by
the pig may vary by enzyme products, age of the pigs,
and feed ingredients. Baas and Thacker (1996) showed
that commercial products of β-glucanase varied in their
effects on digestibility of DM, energy, and CP. It seems
the most likely application would be in weanling pig
1
We wish to acknowledge the financial support of Agri-King Inc.
(Fulton, IL).
2
Corresponding author: jepettig@uiuc.edu
Received May 10, 2007.
Accepted March 3, 2008.
J. Anim. Sci. 2008. 86:1533–1543
doi:10.2527/jas.2007-0262
diets in which pancreatic enzyme production may be
limiting (Lindemann et al., 1986). Exogenous enzymes
seem less effective in older pigs (Thacker et al., 1988,
1989, 1992; Baas and Thacker, 1996; Nyachoti et al.,
2006), because older pigs are more able to digest fiber
than younger pigs. Exogenous enzymes were also more
effective in chick diets based on poorly digested cereals
(Classen et al., 1995; Scott et al., 1998a,b), which contain more fiber (Fernandez and Jørgenson, 1986; Noblet
and Bach-Knudsen, 1997; Le Goff et al., 2002).
Previous studies (Thacker et al., 1990; Li et al., 1996;
Zijlstra et al., 2004) on supplying carbohydrases in pig
diets were mainly focused on barley, wheat, or ryebased diets. Gaining an advantage from the use of exogenous enzymes has been more challenging for corn-
1533
Downloaded from jas.fass.org at Serials/Acq. Dept., Library on May 1, 2009.
1534
Ji et al.
soybean meal (SBM) diets due to decreased fiber concentrations and high nutrient digestibility. Corn and
SBM are known to contain 10 and 22% of nonstarch
polysaccharides, respectively (CVB, 1998). In conjunction with the variation in nutrient composition of corn
(Sullivan et al., 1989; Burgoon et al., 1992; Summers,
2001) and SBM (Grieshop et al., 2003), there may be
potential to improve digestibility of corn-SBM diets
with exogenous enzymes. A novel β-glucanase-protease
enzyme blend product (EBP) has been developed to
improve performance of grower-finisher pigs fed a cornSBM-based diet, in which exogenous enzymes have not
often been shown to be effective. The objective of this
experiment was to investigate the effect of EBP on digestibility of energy and nutrients in the small intestine, hindgut, and overall gastrointestinal tract.
MATERIALS AND METHODS
Animals and Facilities
Experimental procedures were approved by the Institutional Animal Care and Use Committee of the University of Illinois.
Fifteen growing barrows (22.5 ± 0.5 kg of BW, PIC337 sire × Camborough-22, PIC dam) were housed in
galvanized metal metabolism crates (0.86 × 1.60 m)
with mesh floors in a power ventilated, environmentally
controlled room. Pigs were able to move about freely,
turn around, and lie down in the crates. Each crate was
equipped with a low-pressure drinking nipple, a feeder,
and a screen for collecting feces. Room lights were illuminated for 24 h/d, and the room temperature was controlled to approximately 22°C.
Surgical Procedures and Animal Care
The pigs had 7 d to adapt to their surroundings before
the surgery. Pigs had ad libitum access to feed during
this period. Feed was withheld from each animal 12 h
before surgery, but the pigs had continuous access to
water. Pigs were surgically fitted with a simple T-cannula at approximately 10 cm cranial to the ileo-cecal
junction, according to procedures adapted from Sauer
et al. (1983).
The cannulation site was cleaned daily with soft detergent and warm water during the entire experiment.
Also, a zinc oxide-lanolin-based cream (Desitin, Pfizer
Inc., New York, NY) was applied daily to the flank area
to minimize irritation and to maintain skin integrity
during the entire experiment. The pigs were fed 2 meals
daily until full feeding was achieved 4 d after surgery.
A period of 10 d was allowed for surgical recovery before
the beginning of the experiment.
Experimental Design, Enzyme Product, and Diets
Twelve of the 15 cannulated pigs (38.2 ± 1.7 kg of
BW at the beginning of period 1) were blocked by previ-
Table 1. Composition of the experimental diets, as-fed
basis
Item
Ingredient, %
Corn
Soybean meal
Animal-vegetable fat blend
Dicalcium phosphate
Limestone
Salt-trace mineral premix1
Vitamin premix2
Lysine HCl
Threonine
Methionine
Corn starch
Sucrose
Cellulose3
Hydrolyzed casein4
MgO (58% Mg)
K2CO3 (55% K)
Total
Calculated composition
ME,5 Mcal/kg
CP, %
Lysine, %
True ileal digestible lysine, %
Ca, %
Available P, %
Basal diet
Hydrolyzed
casein diet
70.386
25.80
1.00
1.23
0.90
0.35
0.10
0.175
0.04
0.019
—
—
—
—
—
—
100.00
—
—
3.00
1.61
0.77
0.35
0.10
—
—
—
57.62
20.00
5.00
10.00
0.15
1.40
100.00
3.361
18.29
1.10
0.98
0.70
0.29
3.622
9.04
0.74
0.74
0.70
0.29
1
Supplied the following per kilogram of complete diet: Fe, 90 mg
(FeSO4ⴢH2O); Zn, 100 mg (ZnO); Mn, 20 mg (MnO); Cu, 8 mg (CuSO4ⴢH2O); I, 0.35 mg (CaI2); Se, 0.3 mg (Na2SeO3); and NaCl, 3 g.
2
Supplied the following per kilogram of complete diet: 6,608 IU of
vitamin A as retinyl acetate; 680 IU of vitamin D as cholecalciferol;
DL-α-tocopheryl acetate, 88 mg; menadione sodium bisulfite complex,
4 mg; niacin, 33 mg; D-Ca-pantothenate, 24 mg; riboflavin, 9 mg;
vitamin B12, 35 ␮g; and choline chloride, 324 mg.
3
Solka-Floc 40 FCC (powdered cellulose, International Fiber Corp.,
North Tonawanda, NY).
4
American Casein Company, Burlington, NJ.
5
The assumed ME values of the ingredients were from the NRC
(1998); ME of Solka-floc was assumed to be zero.
ous feed intake into 3 groups with 4 pigs each. Three
extra pigs were available to be used in the event that
pigs were removed from the study, but none was used.
Within each group, pigs were randomly assigned to 1
of 4 treatments in a 4 × 4 Latin square design, with
pigs and periods as the main factors. The 4 treatments
were basal diet (Basal), Basal + 0.05% of EBP (0.05%
EBP), Basal + 0.10% of EBP (0.10% EBP), and a hydrolyzed casein diet (Casein). The Casein contained hydrolyzed casein at a 10% level (as-fed basis) as the sole
protein source and was included to enable the calculation of standardized ileal digestibility of CP and amino
acids. The Basal consisted mainly of corn and SBM and
was formulated to provide 3.36 Mcal of ME/kg and 1.1%
total lysine (Table 1) to meet or exceed the nutrient
requirements (NRC, 1998). The EBP (Agri-King Inc.,
Fulton, IL) contained β-glucanase activity (guaranteed
not less than 660,000 β-glucanase units/kg) and protease activity (guaranteed not less than 22,000 hemoglobin units/kg). One β-glucanase unit liberates 1 mM reducing sugar (glucose equivalence) per minute. One he-
Downloaded from jas.fass.org at Serials/Acq. Dept., Library on May 1, 2009.
1535
Effect of exogenous enzymes on nutrient digestibility
Table 2. Analyzed composition of the experimental diets (DM basis)
Item
Casein1
Basal2
0.05% EBP3
0.10% EBP4
DM, %
OM, %
GE, Mcal/kg
DE, Mcal/kg
CP, %
Starch, %
Nonfiber carbohydrate, %
ADF, %
NDF, %
Hemicellulose, %
Total dietary fiber, %
Insoluble dietary fiber, %
Soluble dietary fiber, %
Ether extract, %
Acid-hydrolyzed fat, %
Arginine, %
Histidine, %
Isoleucine, %
Leucine, %
Lysine, %
Methionine, %
Phenylalanine, %
Threonine, %
Tryptophan, %
Valine, %
Alanine, %
Aspartic acid, %
Cysteine, %
Glutamate, %
Glycine, %
Serine, %
Tyrosine, %
Ash, %
Ca, %
P, %
Mg, %
K, %
Na, %
S, %
Cl, %
Fe, ppm
Cu, ppm
Zn, ppm
Mn, ppm
92.0
96.5
4.148
—
9.5
56.5
78.2
4.4
5.4
1.0
—
—
—
3.5
2.4
0.39
0.34
0.57
0.99
0.78
0.24
0.49
0.41
0.13
0.74
0.29
0.70
0.05
2.52
0.19
0.55
0.50
3.48
0.50
0.29
0.03
0.72
0.31
0.10
0.09
131
3
116
30
87.3
95.0
4.367
3.588
20.5
54.0
61.9
4.0
8.9
4.9
12.9
10.8
2.1
4.2
3.8
1.43
0.53
0.95
1.74
1.22
0.30
0.98
0.82
0.28
1.06
1.01
2.18
0.28
4.05
0.91
1.11
0.72
5.02
0.86
0.65
0.16
0.92
0.13
0.22
0.25
321
14
137
40
87.2
95.4
4.386
3.818
20.3
54.1
63.3
3.5
8.5
5.0
13.1
10.6
2.5
4.0
3.7
1.42
0.51
0.94
1.72
1.18
0.29
0.96
0.80
0.27
1.05
0.99
2.10
0.28
3.94
0.84
1.03
0.71
4.59
0.86
0.65
0.16
0.90
0.14
0.23
0.23
332
13
135
46
87.2
95.2
4.362
3.806
20.1
54.8
63.2
3.9
8.6
4.7
13.0
11.2
1.8
4.0
3.9
1.42
0.50
0.94
1.74
1.23
0.30
0.97
0.80
0.27
1.06
1.00
2.13
0.26
3.97
0.89
1.08
0.71
4.80
0.81
0.62
0.16
0.89
0.14
0.23
0.23
338
15
145
43
1
Hydrolyzed casein diet.
Basal diet.
Basal + 0.05% of the enzyme blend product (EBP).
4
Basal + 0.10% of the EBP.
2
3
moglobin unit of protease produces, in 1 min, a
hydrolysate, in which absorbance at 275 nm is equal
to that of a solution containing 1.1 mg/mL of tyrosine
in 0.006 N hydrochloric acid. The manufacturer’s suggested usage of the EBP in growing pig diets is 0.05%.
The 0.05% EBP and 0.10% EBP were made by mixing
0.05 and 0.10% of the EBP into the Basal, respectively,
on an as-fed basis. The 4 chromic oxide diets were made
by mixing 0.35% of chromic oxide (as-fed basis) into the
4 regular diets (Basal, 0.05% EBP, 0.10% EBP, and
Casein). Both the regular and chromic oxide diets were
sampled at the feed mill when the diets were mixed
and at the feeders during period 4 of the experiment.
Because the analyzed nutrient contents of the diets
sampled at both the feed mill and the feeder were similar, the nutrient contents of the diets sampled at the
feed mill were used in calculation of digestibility. The
analyzed nutrient contents of the 4 regular diets sampled at the feed mill are listed in Table 2.
Animal Feeding and Sample Collection
The daily feed intake of pigs allowed ad libitum consumption before the beginning of the experiment
ranged from 1,725 to 2,400 g/d. To ensure that all pigs
could consume the feed allowance, the feed intake of
Downloaded from jas.fass.org at Serials/Acq. Dept., Library on May 1, 2009.
1536
Ji et al.
all 4 pigs within each square during the first period
was restricted to 85% of the previous intake of the pig
with the lowest intake (i.e., 1,460, 1,820, and 1,980 g/
d, respectively). The feeding level was increased by 100
g/d in each subsequent period to accommodate the increased nutrient needs due to growth of the pigs. The
feeding levels were 3.3 to 4.3 times the maintenance
energy requirement (106 kcal of ME per kg of BW0.75;
NRC, 1998) in the first period and 2.7 to 3.1 times
maintenance in the last period. Pigs were fed 2 equal
meals daily, at 12-h intervals. Water was available free
choice. Spilled feed was collected and, unless it was
heavily contaminated with feces, was returned to the
feeder.
There were 4 periods in the experiment, and each pig
received different treatment diets in each period. Each
period was 14 d in duration. The first 4 d of each period
were the diet adaptation phase. The chromic oxide diets
were fed at the beginning and end of fecal collection to
identify when to begin and end the collection of feces.
Pigs were fed the diets containing chromic oxide in the
morning meal of d 5. After that, the pigs were still fed
the regular diets. Approximately 24 to 36 h later, the
chromic oxide appeared in the feces, after which the
feces were collected twice daily at 12-h intervals. Fecal
collections were conducted from d 6 to 11. In the morning meal on d 10 of each period (120 h after the first
feeding of chromic oxide), pigs were fed the diets containing chromic oxide until the end of the period. Again,
approximately 24 to 36 h later, chromic oxide appeared
in the feces, and collection of feces was stopped. The
feces of the 5-d collection were pooled by pig. Fresh
feces were frozen at −20°C.
The collection phase of ileal digesta was conducted
on d 13 and 14. Ileal effluents were collected continuously for the same 12-h interval. Ileal digesta were
collected by attaching polyethylene tubing (5 × 25 cm;
Rand Materials Handling Equipment Co. Inc., Pawtucket, RI) to the cannula barrel with a cable tie. There
was no chemical preservative in the tubing. The tubing
was changed at least once every hour. As soon as ileal
digesta was dumped into a container, it was placed in
a freezer at −20°C for storage. Ileal digesta samples
were pooled by pig and frozen at −20°C until the end
of each collection period.
Chemical Analyses
Fresh feces of each pig were weighed. All fecal and
ileal samples were thawed and mixed for homogeneity.
A subsample was taken for chemical analyses. Fecal,
ileal, and feed samples were dried at 60°C and ground
to an 800-␮m particle size before analyses. The contents
of DM, ADF, NDF, total dietary fiber, CP, ether extract,
ash, Ca, P, Mg, K, Na, Fe, Cu, Zn, Mn, S, Cl, and pH
were determined by the methods of the AOAC (2002).
Hemicellulose was calculated as the difference between
NDF and ADF. Soluble dietary fiber and insoluble dietary fiber were analyzed according to the method de-
scribed by Prosky et al. (1992). Amino acids were determined by the method of the AOAC (2002), but using an
Adsorbosphere Opa HPLC precolumn (Alltech, 2000).
Tryptophan was determined by an HPLC method using
7.5 N NaOH at 103 kPa for 16 h rather than barium
hydroxide. Energy was measured using a bomb calorimeter (Model 1108, Parr Instrument Company, Moline,
IL). Starch was measured by the method of Trinder
(1969). Acid-hydrolyzed fat was determined by the
method of Oser (1965). Chromium was analyzed by the
method developed by Williams and Iismaa (1962) and
Binnerts et al. (1968). Ammonia-nitrogen was determined by the method of Peters et al. (2003). The VFA
were analyzed by capillary electrophoresis using a
method adapted from that of Cancalon (1993).
Calculations of Digestibility
The fecal digestibility was determined by the equation described by Adeola (2001). The apparent ileal digestibility, endogenous amino acid loss, and standardized ileal digestibility were determined by the equations
described by Smiricky et al. (2002). The endogenous
amino acid loss used to calculate the standardized ileal
digestibility in a square (group) was the average of the
4 pigs in the square.
The digestibility of nutrients in the hindgut was calculated by using the following equations:
difference between fecal and ileal digestibility (%) =
fecal digestibility − apparent ileal digestibility;
and digestibility based on the nutrients
entering the hindgut (%) =
⎡
⎢1
⎣
−
⎤
FW × NCf
⎥ × 100,
DMI × NCF × (1 − AID/100)⎦
where FW = the dried fecal weight (kg of DM); NCf =
the nutrient content in feces (% of DM); DMI = the DM
intake (kg); NCF = the nutrient content in feed (% of
DM); and AID = apparent ileal digestibility (%).
Statistical Analyses
Data were analyzed by the GLM and UNIVARIATE
procedures (SAS Inst. Inc., Cary, NC) to test for homogeneity, normality, and outliers. Data were then analyzed by the MIXED procedure of SAS with groups, pigs,
periods, and diets in the model. Means were reported
as least squares means. Treatment comparisons were
made using the following orthogonal contrasts: 1) effect
of EBP [(0.05% EBP + 0.10% EBP)/2 vs. Basal] and 2)
effect of level of EBP (0.05 vs. 0.10% EBP), with an α
level of 0.05 used to determine statistical significance.
RESULTS
Enzyme Activities and Diet Components
The calculated β-glucanase activities were 330 and
660 β-glucanase units/kg of diet, and the calculated
Downloaded from jas.fass.org at Serials/Acq. Dept., Library on May 1, 2009.
1537
Effect of exogenous enzymes on nutrient digestibility
Table 3. Feed intake and characteristics of feces and ileal digesta (DM basis)
Contrast5
Treatment
Basal1
(n6 = 10)
Item
Feed intake, g as fed
Fecal weight, g of DM
In feces
DM content, %
pH
NH3, ppm
Acetic acid, %
Propionic acid, %
Butyric acid, %
Valeric acid, %
In ileal digesta
DM content, %
pH
NH3, ppm
Acetic acid, %
Propionic acid, %
Butyric acid, %
Valeric acid, %
0.05% EBP2
(n6 = 11)
0.10% EBP3
(n6 = 11)
8,287
1,043
8,236
935
8,189
936
44.81
6.91
3,434
0.987
0.51
0.36
0.14
45.29
6.79
2,863
1.27
0.55
0.39
0.14
41.68
6.85
3,140
1.26
0.65
0.41
0.17
n8 = 11
n8 = 12
n8 = 12
10.46
7.39
86.85
0.014
0.004
0.003
ND9
9.60
7.19
89.31
0.015
0.004
0.002
ND
8.53
7.29
88.36
0.014
0.003
0.001
0.001
SD4
98
44
3.67
0.21
562
0.15
0.15
0.10
0.04
3.65
0.30
14.57
0.010
0.000
0.000
0.000
1
2
0.065
<0.001
0.275
0.965
0.360
0.360
0.064
<0.001
0.130
0.242
0.375
0.032
0.507
0.264
0.906
0.106
0.576
0.068
0.318
0.204
0.719
0.854
0.704
0.502
0.412
0.479
0.412
0.875
0.940
0.188
0.315
0.133
1
Basal diet.
Basal + 0.05% of the enzyme blend product (EBP).
3
Basal + 0.10% of the EBP.
4
Pooled SD.
5
Contrasts were as follows: 1) effect of EBP [(0.05% EBP + 0.10% EBP)/2 vs. Basal] and 2) effect of level
of EBP (0.05 vs. 0.10%).
6
One outlier was removed from 0.05% EBP and 0.10% EBP; 2 outliers were removed from Basal.
7
One more outlier was removed from Basal.
8
One outlier was removed from Basal.
9
ND = not detected.
2
protease activities were 11 and 22 hemoglobin units/
kg of diet for 0.05% EBP and 0.10% EBP, respectively.
The contents of DM, GE, CP, total dietary fiber, and
indispensable amino acids among Basal, 0.05% EBP,
and 0.10% EBP were similar after manufacturing (Table 2), indicating that the EBP did not change the main
nutrient composition of the diets.
< 0.001) in the pigs fed the EBP than in the pigs fed the
Basal. Fecal pH and concentrations of DM, ammonia,
propionic acid, butyric acid, and valeric acid did not
differ between the pigs fed the EBP and the Basal.
Acetic acid was greater (P < 0.001) in feces from the
pigs fed the EBP than from those fed the Basal.
Periods
Fecal Digestibility of Nutrients
The fecal digestibility and apparent ileal digestibility
of DM, CP, and energy, as well as apparent ileal digestibility of amino acids, were decreased (P < 0.001) in
period 1 compared with periods 2, 3, and 4 (data not
shown). Although pigs were allowed 10 d for recovery
after surgery with 4 additional days for dietary adaptation, this may not have been sufficient time for recovery
from surgery. However, digestibility values did not vary
among periods 2, 3, and 4, indicating the effect of age
of grower-finisher pigs on nutrient digestibility was
limited.
Feed Intake and Fecal Characteristics
All pigs consumed the allotted feed, gained weight,
had no diarrhea, and appeared healthy during the experiment. Feed intake did not differ among the treatments (Table 3). Dried fecal weight was decreased (P
The fecal digestibility of DM, OM, energy, CP, and
ash was greater (P < 0.05) for the pigs fed the EBP
than the pigs fed the Basal (Table 4), whereas the fecal
digestibility of these items did not differ between the
pigs fed 0.05% EBP and 0.10% EBP. Pigs fed the EBP
had greater fecal digestibility (P < 0.05) of total dietary
fiber and acid-hydrolyzed fat than pigs fed the Basal.
Within total dietary fiber, the fecal digestibility of insoluble dietary fiber was increased (P < 0.05) by the EBP,
whereas the fecal digestibility of soluble dietary fiber
was not affected by the EBP. Almost all starch was
digested over the whole digestive tract, and there was
no difference in fecal digestibility of starch among treatments. The fecal digestibility of Ca, P, Mg, S, Fe, and
Cu also was increased (P < 0.05) when pigs were fed
the EBP.
Downloaded from jas.fass.org at Serials/Acq. Dept., Library on May 1, 2009.
1538
Ji et al.
Table 4. Effect of β-glucanase-protease enzyme blend product on fecal digestibility (%)
Treatment
Contrast5
Item
Basal1
(n6 = 10)
0.05% EBP2
(n6 = 11)
0.10% EBP3
(n6 = 11)
SD4
1
2
DM
OM
Energy
CP
Starch
Nonfiber carbohydrate
ADF
NDF
Hemicellulose
Total dietary fiber
Insoluble dietary fiber
Soluble dietary fiber
Ether extract
Acid-hydrolyzed fat
Ash
Ca
P
Mg
K
Na
S
Cl
Fe
Cu
Zn
Mn
87.42
88.99
86.51
86.47
99.24
94.83
64.84
54.62
46.33
60.61
53.81
95.59
80.14
44.97
57.79
57.33
53.80
26.33
75.05
86.317
81.70
95.50
10.13
5.07
−34.337
−5.01
88.61
89.89
87.42
88.08
99.26
95.22
61.40
55.62
51.62
65.36
57.79
96.80
80.51
48.27
60.77
65.31
61.73
35.20
77.41
87.62
84.76
95.40
21.07
10.46
−22.67
18.63
88.50
89.87
87.26
87.39
99.31
95.63
65.92
56.05
47.99
65.61
60.97
94.68
78.24
51.80
60.95
61.20
57.83
33.36
76.50
88.42
84.33
95.28
19.03
20.89
−7.52
12.07
0.47
0.45
0.56
0.94
0.15
0.48
3.54
3.33
5.06
3.33
3.79
5.64
2.95
2.58
2.34
2.89
2.41
2.89
2.14
3.51
1.16
1.30
3.42
3.56
10.77
3.77
<0.001
<0.001
0.002
0.003
0.527
0.006
0.400
0.357
0.096
0.002
0.002
0.947
0.508
<0.001
0.004
<0.001
<0.001
<0.001
0.036
0.252
<0.001
0.754
<0.001
<0.001
<0.001
<0.001
0.617
0.885
0.511
0.101
0.436
0.065
0.009
0.765
0.112
0.861
0.066
0.390
0.090
0.005
0.857
0.004
0.002
0.156
0.334
0.601
0.389
0.842
0.180
<0.001
0.005
0.001
1
Basal diet.
Basal + 0.05% of the enzyme blend product (EBP).
Basal + 0.10% of the EBP.
4
Pooled SD.
5
Contrasts were as follows: 1) effect of EBP [(0.05% EBP + 0.10% EBP)/2 vs. Basal] and 2) effect of level
of EBP (0.05 vs. 0.10%).
6
One outlier was removed from 0.05% EBP and 0.10% EBP; 2 outliers were removed from Basal.
7
One outlier was removed from Basal.
2
3
Apparent and Standardized Ileal Digestibility
of Nutrients
The apparent ileal digestibility of DM, energy, starch,
and nonfiber carbohydrate did not differ among treatments (Table 5). The pigs fed EBP had decreased (P <
0.05) apparent ileal digestibility of CP than the pigs
fed the Basal. The apparent ileal digestibility of NDF
and hemicellulose of the pigs fed the EBP was greater
(P < 0.05) compared with that of the pigs fed the Basal.
The apparent ileal digestibility of acid-hydrolyzed fat
in the pigs fed the EBP tended to be greater (P = 0.051)
than in those fed the Basal. Ileal digestibility of most
amino acids was not affected by treatment, but the EBP
reduced the apparent (Table 5) and standardized (Table
6) digestibility of methionine, alanine, and serine (P <
0.05) and arginine, glycine, threonine, and valine (P <
0.05 for apparent digestibility only).
Digestion of Nutrients in the Hindgut
There was no difference among treatments in the
difference between fecal and ileal digestibility except
for hemicellulose, which was decreased (P < 0.05) for
the pigs fed the EBP compared with pigs fed the Basal
(Table 7). The difference between fecal and ileal digestibility of NDF for the pigs fed the EBP tended to be
decreased (P = 0.050) compared with the pigs fed the
Basal. For the digestibility based on the nutrients entering the hindgut (Table 8), the pigs fed the EBP had
greater values (P < 0.05) for CP and DM digestibility
compared with pigs fed the Basal treatment.
DISCUSSION
The EBP containing β-glucanase and protease was
developed to improve performance of growing pigs. The
current results showed that the EBP supplied at either
0.05 or 0.10% of diet increased fecal digestibility of DM,
energy, CP, nonfiber carbohydrate, total dietary fiber,
acid-hydrolyzed fat, Ca, and P. Digestion of NDF and
hemicellulose in the small intestine of growing pigs was
also improved by supplying EBP. However, the EBP
did not improve the standardized ileal digestibility of
amino acids. This indicates that β-glucanase in the EBP
Downloaded from jas.fass.org at Serials/Acq. Dept., Library on May 1, 2009.
1539
Effect of exogenous enzymes on nutrient digestibility
Table 5. Effect of β-glucanase-protease enzyme blend product on apparent ileal digestibility (%)
Treatment
Contrast5
Item
Basal1
(n6 = 11)
0.05% EBP2
(n6 = 12)
0.10% EBP3
(n6 = 12)
SD4
1
2
DM
Energy
CP
Starch
Nonfiber carbohydrate
ADF
NDF
Hemicellulose
Ether extract
Acid-hydrolyzed fat
Ash
Arginine
Histidine
Isoleucine
Leucine
Lysine
Methionine
Phenylalanine
Threonine
Tryptophan
Valine
Alanine
Aspartic acid
Cystine
Glutamate
Glycine
Serine
Tyrosine
Ca
P
Mg
K
Na
S
Cl
Fe
Cu
Zn
Mn
70.86
70.93
78.29
97.95
82.11
4.33
1.21
−1.19
61.40
52.06
17.62
94.58
86.89
81.58
83.78
87.04
82.76
83.90
67.10
74.12
81.29
73.03
80.13
76.06
84.90
68.43
81.83
86.72
54.71
49.62
6.12
74.47
−540.83
70.84
−20.89
9.24
−12.61
−7.76
−4.23
69.13
69.48
75.51
98.01
80.68
4.36
9.52
13.11
62.94
53.84
8.19
93.68
85.16
79.62
82.55
85.01
80.35
82.64
62.27
72.35
78.95
69.04
77.85
72.72
82.37
61.94
78.83
85.03
57.51
49.54
13.23
71.18
−567.11
67.62
−59.48
12.39
20.88
−1.97
5.03
70.50
70.71
76.54
98.12
82.04
5.22
10.05
13.42
62.18
57.73
16.41
93.39
85.24
80.63
83.13
85.91
79.07
83.53
63.99
73.76
79.71
70.37
79.33
75.087
84.89
64.81
79.18
84.39
56.31
49.00
11.81
72.64
−513.68
71.16
−16.857
3.56
12.597
−10.58
12.19
2.81
2.84
2.76
0.49
2.17
10.26
8.61
8.56
4.48
4.80
7.94
1.28
3.49
2.18
1.79
2.06
2.82
2.14
4.37
2.98
2.31
3.84
2.62
3.47
2.48
5.67
2.56
3.65
7.74
5.18
9.47
5.27
94.28
3.88
28.52
10.88
23.26
15.32
9.75
0.329
0.441
0.041
0.514
0.362
0.905
0.015
<0.001
0.493
0.051
0.088
0.041
0.210
0.092
0.174
0.054
0.009
0.320
0.025
0.347
0.035
0.032
0.130
0.107
0.188
0.028
0.008
0.154
0.455
0.859
0.086
0.208
0.990
0.326
0.114
0.758
0.003
0.797
0.002
0.246
0.305
0.371
0.588
0.145
0.840
0.881
0.931
0.683
0.062
0.021
0.576
0.955
0.272
0.437
0.295
0.281
0.322
0.348
0.262
0.433
0.409
0.183
0.117
0.023
0.231
0.739
0.670
0.708
0.803
0.717
0.505
0.182
0.038
0.002
0.062
0.399
0.185
0.089
1
Basal diet.
Basal + 0.05% of the enzyme blend product (EBP).
3
Basal + 0.10% of the EBP.
4
Pooled SD.
5
Contrasts were as follows: 1) effect of EBP [(0.05% EBP + 0.10% EBP)/2 vs. Basal] and 2) effect of level
of EBP (0.05 vs. 0.10%).
6
One outlier was removed from Basal.
7
One outlier was removed.
2
is effective even under the difficult conditions of a lowfiber corn-SBM diet and limit fed to older pigs, but the
protease may be ineffective.
The carbohydrate of cereal grains includes sugars,
starch, and cell wall polysaccharides. The endosperm
cell walls of barley, oat, wheat, rye, sorghum, corn, and
triticale are composed mainly of arabinoxylans and βglucans (Henry, 1985; Bedford, 1995; de Lange, 2000).
The current results showed that the fecal digestibility
of total dietary fiber and insoluble dietary fiber was
improved by supplying the EBP. The total dietary fiber
consists of soluble materials such as β-glucans, arabinoxylans, pectins, and gums, and the insoluble galactoand glucomannans, cellulose and lignin (Asp et al.,
1983; Bedford, 1995; Souffrant, 2001). It is unclear why
the EBP increased digestibility of insoluble rather than
soluble dietary fiber, because the substrate of the βglucanase is soluble.
The increase in ileal digestibility of NDF and hemicellulose when the EBP was added to the diet, with no
change in fecal digestibility, indicates that the enzyme
product may have shifted some of the digestion of these
Downloaded from jas.fass.org at Serials/Acq. Dept., Library on May 1, 2009.
1540
Ji et al.
Table 6. Effect of β-glucanase-protease enzyme blend product on standardized ileal digestibility (%)
Treatment
Item
CP
Arginine
Histidine
Isoleucine
Leucine
Lysine
Methionine
Phenylalanine
Threonine
Tryptophan
Valine
Alanine
Aspartic acid
Cystine
Glutamate
Glycine
Serine
Tyrosine
Contrast5
Basal1
(n6 = 11)
0.05% EBP2
(n6 = 12)
0.10% EBP3
(n6 = 12)
SD4
1
2
88.47
101.29
91.01
87.01
87.52
90.91
87.83
87.23
84.66
81.78
87.29
82.33
85.49
80.98
91.17
95.38
91.31
90.55
86.09
100.64
89.48
85.28
86.39
89.00
85.50
86.07
80.45
79.95
85.15
78.59
83.46
77.84
88.93
89.88
88.76
89.02
87.35
100.55
89.64
86.12
86.95
89.83
84.69
86.89
82.28
81.36
85.89
80.05
84.86
79.68
91.00
92.59
89.15
88.47
2.76
1.28
3.49
2.19
1.79
2.06
2.82
2.14
4.36
2.98
2.31
3.85
2.62
3.41
2.49
5.67
2.55
3.65
0.105
0.156
0.279
0.124
0.215
0.068
0.018
0.362
0.057
0.323
0.055
0.049
0.191
0.097
0.208
0.065
0.023
0.200
0.277
0.867
0.916
0.358
0.455
0.337
0.489
0.361
0.316
0.262
0.443
0.364
0.209
0.203
0.056
0.258
0.717
0.718
1
Basal diet.
Basal + 0.05% of the enzyme blend product (EBP).
Basal + 0.10% of the EBP.
4
Pooled SD.
5
Contrasts were as follows: 1) effect of EBP [(0.05% EBP + 0.10% EBP)/2 vs. Basal] and 2) effect of level
of EBP (0.05 vs. 0.10%).
6
One outlier was removed from Basal.
2
3
nutrients from hindgut fermentation to small intestinal
enzymatic digestion. Such a change would avoid the
fermentative loss of energy as heat and methane and
presumably increase the energetic efficiency of fiber
use.
The cell walls enclose starch, protein, and lipids. Supplying enzymes to enhance breakdown of the components of these cell walls can increase exposure of the
nutrients in cells to the digestive enzymes. However,
because of the difference of ingredients, age of pigs, and
enzyme products, the effectiveness of supplying enzyme
in pig diets may be different. Some studies showed that
supplying β-glucanase increased ileal digestibility of
DM, energy, CP or fibers (Taverner and Campbell,
1988; Inborr and van der Meulen, 1993; Yin et al.,
2001a,b), and fecal digestibility of fibers (Yin et al.,
2001a), whereas other studies showed supplying β-glucanase did not improve ileal digestibility of DM, energy,
or CP (Graham et al., 1989; Nyachoti et al., 2006) and
fecal digestibility of DM, energy, CP, or fiber (Taverner
and Campbell, 1988; Graham et al., 1989; Nyachoti et
al., 2006). These previous studies focused on the enzyme
effect on diets based on fibrous cereals such as barley
and wheat. The effect of enzyme on corn-SBM-based
diet has not previously been well addressed. The current results showed that apparent ileal digestibility of
NDF and hemicellulose and the fecal digestibility of
DM, energy, CP, and total dietary fiber were improved
by the EBP. This indicates that even under the difficult
conditions of corn-SBM diet and pigs several months
old, supplying exogenous enzymes still have the potential to improve digestibility of DM, energy, and fibers.
However, because the capacity of degrading fiber is different for the enzymes obtained from different microbes
(Nevins et al., 1978), the effectiveness of various commercial enzyme products was different (Baas and
Thacker, 1996). The enzyme product used in the current
study was an effective fiber-degrading enzyme product.
The effectiveness of an enzyme may be affected by
many factors, such as the activity of the enzyme, the
type and concentration of substrate, particle size, pH,
temperature, and time of the feed staying in the small
intestine. The calculated β-glucanase activities were
330 and 660 units/kg of diet for 0.05% EBP and 0.10%
EBP, respectively. The levels of β-glucanase in previous
studies varied from 10 (Graham et al., 1986) and 100
(Graham et al., 1989) units per kilogram of diet to 600
β-glucanase units/kg of diet (Yin et al., 2001a,b). In this
case, the lower level of EBP tested (0.05%) seemed to
be sufficient to achieve maximum effect.
The apparent ileal digestibility of acid-hydrolyzed fat
was increased by supplying EBP, whereas Graham et
al. (1989) reported no effect. The increase in fecal digestibility of acid-hydrolyzed fat by supplying EBP was
caused by increased digestibility in the small intestine.
Because the fecal digestibility and apparent ileal digestibility of ether extract did not differ, the increase
of apparent ileal digestibility of acid-hydrolyzed fat may
Downloaded from jas.fass.org at Serials/Acq. Dept., Library on May 1, 2009.
1541
Effect of exogenous enzymes on nutrient digestibility
Table 7. Effect of β-glucanase-protease enzyme blend product on the digestibility in the
hindgut (%, the difference between fecal and ileal digestibility)
Treatment
Item
DM
Energy
CP
Starch
Nonfiber carbohydrate
ADF
NDF
Hemicellulose
Ether extract
Acid-hydrolyzed fat
Ash
Ca
P
Mg
K
Na
S
Cl
Fe
Cu
Zn
Mn
Contrast5
Basal1
(n6 = 11)
0.05% EBP2
(n6 = 12)
0.10% EBP3
(n6 = 12)
SD4
1
2
17.25
16.00
9.13
1.46
13.00
61.86
55.93
50.94
19.31
−4.83
42.24
4.30
6.16
24.03
3.74
618.30
11.78
118.44
4.94
27.51
−19.46
4.31
19.68
18.19
12.86
1.34
14.87
56.17
45.36
37.87
17.52
−5.30
52.78
6.92
12.37
23.38
8.00
651.58
17.39
155.58
8.88
−10.37
−20.80
14.28
18.32
16.96
11.15
1.25
13.73
61.06
45.93
34.19
16.69
−5.60
44.99
4.02
8.38
21.48
6.24
603.99
13.45
128.33
14.88
4.34
0.66
−1.28
3.79
3.86
4.01
0.62
2.58
12.97
12.95
14.41
5.95
8.95
10.71
9.61
7.50
12.56
7.19
104.61
5.50
39.74
15.81
23.42
28.57
14.85
0.236
0.294
0.075
0.478
0.167
0.514
0.050
0.014
0.338
0.855
0.118
0.749
0.153
0.739
0.228
0.812
0.097
0.134
0.259
0.003
0.394
0.700
0.392
0.444
0.310
0.713
0.384
0.368
0.916
0.540
0.737
0.935
0.093
0.471
0.210
0.716
0.557
0.281
0.098
0.112
0.366
0.143
0.084
0.020
1
Basal diet.
Basal + 0.05% of the enzyme blend product (EBP).
Basal + 0.10% of the EBP.
4
Pooled SD.
5
Contrasts were as follows: 1) effect of EBP [(0.05% EBP + 0.10% EBP)/2 vs. Basal] and 2) effect of level
of EBP (0.05 vs. 0.10%).
6
One outlier was removed from Basal.
2
3
be because EBP digested cell wall materials and released the lipids from the cell membranes, which constitute most of the difference between ether extract and
acid-hydrolyzed fat. The fecal digestibility of minerals
including Ca, P, Mg, S, Cl, Fe, Cu, and Mn was improved
by the EBP and may result from the EBP releasing
minerals bound to or in the cell wall.
The EBP did not increase ileal digestibility of amino
acids. We cannot explain why the standardized ileal
digestibility of methionine, alanine, and serine was reduced by the EBP. In previous studies, the effect of
supplying exogenous enzymes on ileal digestibility of
protein was contradictory in pigs. Protease treatment
of SBM had no effect on ileal digestibility of CP and
amino acids in newly weaned pigs (Caine et al.,
1997a,b). Li et al. (1996) demonstrated that β-glucanase
supplementation increased the ileal digestibility of βglucans, CP, and the majority of amino acids in a hulless
barley-SBM diet. Yin et al. (2001b) showed that supplying β-glucanase and xylanase in weaned pig diets increased the apparent ileal digestibility of lysine, glycine, threonine, histidine, and valine. Yin et al. (2001a)
obtained similar results from another study, in which
β-glucanase improved apparent ileal digestibility of CP
and of most amino acids. The current study showed
that the supplementation of protease in the corn-SBM-
based growing pig diets was ineffective. It may be because the protease activities in the EBP (11 and 22
hemoglobin units/kg of diet for 0.05% EBP and 0.10%
EBP, respectively) were decreased compared with that
in a previous study (500 units/kg in pig diets; Yin et
al., 2001a).
The feeding levels were 3.3, 4.1, and 4.3 times maintenance in the first period and 2.7, 3.0, and 3.1 times
maintenance in the fourth period. The degree of restriction increased as the pigs grew. The feeding levels did
not affect the fecal digestibility or apparent ileal digestibility of DM, CP, or energy, as well as apparent ileal
digestibility of all amino acids, which agrees with the
previous results of Haydon et al. (1984), Albin et al.
(2001), Gómez et al. (2002), and Moter and Stein (2004).
Other studies showed an inverse relationship between
feeding levels and fecal digestibility of DM and CP (Rao
and McCracken, 1991) and ileal digestibility of CP and
most amino acids (Moter and Stein, 2004). Because the
comparisons were at different levels of restriction
rather than restriction to ad libitum, it is possible the
response to the EBP may be greater in ad libitum pigs.
In conclusion, the fecal digestibility of DM, OM, energy, CP, total dietary fiber, acid-hydrolyzed fat, Ca,
and P and the apparent ileal digestibility of hemicellulose and NDF were improved, but standardized ileal
Downloaded from jas.fass.org at Serials/Acq. Dept., Library on May 1, 2009.
1542
Ji et al.
Table 8. Digestibility (%) based on the nutrients entering the hindgut
Treatment
Item
DM
Energy
CP
Starch
Nonfiber carbohydrate
ADF
NDF
Hemicellulose
Ether extract
Acid-hydrolyzed fat
Ash
Ca
P
Mg
K
Na
S
Cl
Fe
Cu
Zn
Mn
Contrast5
Basal1
(n6 = 11)
0.05% EBP2
(n6 = 12)
0.10% EBP3
(n6 = 12)
SD4
1
2
57.78
54.41
39.20
63.00
71.85
63.71
54.38
47.00
48.86
−13.29
49.73
5.13
9.86
24.02
−5.94
97.58
37.99
96.39
2.45
29.54
−24.34
1.38
62.70
58.53
50.71
61.11
75.08
58.70
49.78
42.92
46.03
−13.43
56.39
13.35
23.21
24.47
1.66
97.92
51.95
96.78
7.23
−53.30
−25.56
12.62
61.14
56.83
46.67
63.81
75.81
63.69
50.76
39.47
42.10
−14.63
52.84
8.04
15.85
23.33
4.26
98.01
45.85
95.97
14.74
−21.13
−1.66
−2.19
4.56
6.42
7.68
12.00
4.13
5.53
6.47
8.22
11.09
18.82
5.69
21.20
10.62
11.52
19.64
0.89
9.27
0.86
15.51
99.31
30.34
12.95
0.026
0.188
0.004
0.906
0.032
0.238
0.105
0.075
0.288
0.916
0.034
0.489
0.026
0.978
0.239
0.262
0.006
0.953
0.157
0.088
0.351
0.436
0.413
0.527
0.216
0.589
0.673
0.042
0.714
0.318
0.334
0.878
0.145
0.548
0.109
0.812
0.750
0.788
0.126
0.033
0.253
0.439
0.072
0.013
1
Basal diet.
Basal + 0.05% of the enzyme blend product (EBP).
Basal + 0.10% of the EBP.
4
Pooled SD.
5
Contrasts were as follows 1) effect of EBP [(0.05% EBP + 0.10% EBP)/2 vs. Basal] and 2) effect of level
of EBP (0.05 vs. 0.10%).
6
One outlier was removed from Basal.
2
3
digestibility of amino acids was not increased by the
EBP in corn-SBM-based growing pig diets. The EBP
used in the study was effective. Supplying 0.05% was
as effective as 0.10% EBP in this study with limitfed pigs.
LITERATURE CITED
Adeola, O. 2001. Digestion and Balance Techniques in Pigs. Pages
903–916 in Swine Nutrition. 2nd ed. A. J. Lewis and L. L. Southern, ed. CRC Press, Boca Raton, FL.
Albin, D. M., J. E. Wubben, M. R. Smiricky, and V. M. Gabert. 2001.
The effect of feed intake on ileal rate of passage and apparent
amino acid digestibility determined with or without correction
factors in pigs. J. Anim. Sci. 79:1250–1258.
Alltech. 2000. Data Sheet 28062U Adsorbosphere OPA. Alltech Assoc.
Inc., Deerfield, IL.
AOAC. 2002. Official Methods of Analysis. 17th ed. Assoc. Off. Anal.
Chem., Arlington, VA.
Asp, N.-G., C.-G. Johansson, J. Jallmer, and M. Siljestrom. 1983.
Rapid enzymic assay of insoluble and soluble dietary fibre. J.
Agric. Food Chem. 31:476–482.
Baas, T. C., and P. A. Thacker. 1996. Impact of gastric pH on dietary
enzyme activity and survivability in swine fed β-glucanase supplemented diets. Can. J. Anim. Sci. 76:245–252.
Bedford, M. R. 1995. Mechanism of action and potential environmental benefits from the use of feed enzymes. Anim. Feed Sci. Technol. 53:145–155.
Binnerts, W. T., A. T. van’t Klooster, and A. M. Frens. 1968. Soluble
chromium indicator measured by atomic absorption in digestion
experiments. Vet. Rec. 82:470.
Burgoon, K. G., J. A. Hansen, D. A. Knabe, and A. J. Bockholt. 1992.
Nutritional value of quality protein maize for starter and finisher
swine. J. Anim. Sci. 70:811–817.
Caine, W. R., W. C. Sauer, S. Tamminga, M. W. A. Verstegen, and
H. Schulze. 1997a. Apparent ileal digestibilities of amino acids
in newly weaned pigs fed diets with protease-treated soybean
meal. J. Anim. Sci. 75:2962–2969.
Caine, W. R., S. Tamminga, M. W. A. Verstegen, W. C. Sauer, and
H. Schulze. 1997b. Endogenous recoveries and true ileal digestibilities of amino acids in newly weaned pigs fed diets with
protease-treated soybean meal. J. Anim. Sci. 75:2970–2979.
Cancalon, P. F. 1993. Rapid monitoring of fruit juice adulteration by
capillary electrophoresis. LC GC 11:748–751.
Classen, H. L., T. A. Scott, G. G. Irish, P. Huck, M. Swift, and M.
R. Bedford. 1995. The relationship of chemical and physical
measurements to the apparent metabolisable energy (AME) of
wheat when fed to broiler chickens with and without a wheat
enzyme source. Pages 65–69 in Proc. 2nd Eur. Symp. Feed Enzymes. W. van Hartingsveldt, M. Hessing, J. P. van der Lugt,
and W. A. C. Somers, ed. TNO Nutr. Food Res. Inst., Zeist,
the Netherlands.
CVB. 1998. Veevoedertalel (Feeding value of feed ingredients). CVB,
Lelystad, the Netherlands.
de Lange, C. F. M. 2000. Characterisation of the non-starch polysaccharides. Pages 77–92 in Feed Evaluation: Principles and Practice. P. J. Moughan, M. W. A. Verstegen, and M. I. Visser-Reyneveld, ed. Wageningen Pers, Wageningen, the Netherlands.
Fernandez, J. A., and J. N. Jørgenson. 1986. Digestibility and absorption of nutrients as affected by fibre content in the diet of the
pig. Quantitative aspects. Livest. Prod. Sci. 15:53–71.
Gómez, R. S., A. J. Lewis, P. S. Miller, and H. Y. Chen. 2002. Growth
performance, diet apparent digestibility, and plasma metabolite
Downloaded from jas.fass.org at Serials/Acq. Dept., Library on May 1, 2009.
Effect of exogenous enzymes on nutrient digestibility
concentrations of barrows fed corn-soybean meal diets or lowprotein, amino acid-supplemented diets at different feeding levels. J. Anim. Sci. 80:644–653.
Graham, H., J. G. Fadel, C. W. Newman, and R. K. Newman. 1989.
Effect of pelleting and beta-glucanase supplementation on the
ileal and fecal digestibility of a barley-based diet in the pig. J.
Anim. Sci. 67:1293–1298.
Graham, H., K. Hesselman, E. Jonsson, and P. Åman. 1986. Influence
of β-glucanase supplementation on digestion of a barley-based
diet in the pig gastrointestinal tract. Nutr. Rep. Int. 34:1089–
1096.
Grieshop, C. M., C. T. Kadzere, G. M. Clapper, E. A. Flickinger, L.
L. Bauer, R. L. Frazier, and G. C. Fahey Jr. 2003. Chemical
and nutritional characteristics of United States soybeans and
soybean meals. J. Agric. Food Chem. 51:7684–7691.
Haydon, K. D., D. A. Knabe, and T. D. Tanksley Jr. 1984. Effects of
level of feed intake on nitrogen, amino acid and energy digestibilities measured at the end of the small intestine and over the
total digestive tract of growing pigs. J. Anim. Sci. 59:717–724.
Henry, R. J. 1985. A comparison of the non-starch carbohydrates in
cereal grains. J. Sci. Food Agric. 36:1243–1253.
Inborr, J., and J. van der Meulen. 1993. Residual activity of added
enzymes in relation to fibre digestibility in the terminal ileum
of growing pigs. Pages 34–37 in Proc. 1st Symp. Enzymes Anim.
Nutr. C. Wenk and M. Boessinger, ed. Kartause Ittingen,
Warth, Switzerland.
Le Goff, G., J. Van Milgen, and J. Noblet. 2002. Influence of dietary
fibre on digestive utilization and rate of passage in growing pigs,
finishing pigs and adult sows. Anim. Sci. 74:503–515.
Li, S., W. C. Sauer, S. X. Huang, and V. M. Gabert. 1996. Effect of βglucanase supplementation to hulless barley- or wheat-soybean
meal diets on the digestibilities of energy, protein, β-glucans,
and amino acids in young pigs. J. Anim. Sci. 74:1649–1656.
Lindemann, M. D., S. G. Cornelius, S. M. El Kandelgy, R. L. Moser,
and J. E. Pettigrew. 1986. Effect of age, weaning and diet on
digestive enzyme levels in the piglet. J. Anim. Sci. 62:1298–1307.
Moter, V., and H. H. Stein. 2004. Effect of feed intake on endogenous
losses and amino acid and energy digestibility by growing pigs.
J. Anim. Sci. 82:3518–3525.
Nevins, D. J., R. Yamamoto, and D. J. Huber. 1978. Cell wall β-Dglucans of five grass species. Phytochemistry 17:1503–1505.
Noblet, J., and K. E. Bach-Knudsen. 1997. Comparative digestibility
of wheat, maize and sugar beet pulp non-starch polysaccharides
in adult sows and growing pigs. Pages 571–574 in Digestive
Physiology in Pigs. J. P. Laplace, C. Février, and A. Barbeau,
ed. EAAP Publ. No. 88. INRA, Saint-Malo, France.
NRC. 1998. Nutrient Requirements of Swine. 10th rev. ed. Natl.
Acad. Press, Washington, DC.
Nyachoti, C. M., S. D. Arntfield, W. Guenter, S. Cenkowski, and F.
O. Opapeju. 2006. Effect of micronized pea and enzyme supplementation on nutrient utilization and manure output in growing
pigs. J. Anim. Sci. 84:2150–2156.
Oser, B. L. 1965. Feces. Pages 530–540 in Hawk’s Physiological Chemistry. 14th ed. McGraw-Hill Inc., New York, NY.
Peters, J., A. Wolf, and N. Wolf. 2003. Ammonium nitrogen: Ammonium-N by colorimetry using an autoanalyzer. Pages 25–29 in
Recommended Methods of Manure Analysis. J. Peters, ed. Univ.
Wisconsin-Extension, Madison.
Prosky, L., N. G. Asp, T. Schweizer, J. W. De Vries, and I. Furda.
1992. Determination of insoluble and soluble dietary fiber in
food and food products: Collaborative study. J. Assoc. Off. Anal.
Chem. 75:360–367.
Rao, D. S., and K. J. McCracken. 1991. Effect of energy intake on
protein and energy metabolism of boars of high genetic potential
for lean growth. Anim. Prod. 52:499–507.
1543
Sauer, W. C., H. Jorgensen, and R. Berzins. 1983. A modified nylon
bag technique for determining apparent digestibility of protein
in feedstuffs for pigs. Can. J. Anim. Sci. 63:233–237.
Scott, T. A., F. G. Silversides, H. L. Classen, M. L. Sift, M. R. Bedford,
and J. W. Hall. 1998a. A broiler chick bioassay for measuring
the feeding value of wheat and barley in complete diets. Poult.
Sci. 77:449–455.
Scott, T. A., F. G. Silversides, H. L. Classen, M. L. Sift, M. R. Bedford,
and J. W. Hall. 1998b. Effect of cultivar and environment on
the feeding value of western Canadian wheat and barley samples
with and without enzyme supplementation. Can. J. Anim. Sci.
78:649–656.
Smiricky, M. R., C. M. Grieshop, D. M. Albin, J. E. Wubben, V. M.
Gabert, and G. C. Fahey Jr. 2002. The influence of soy oligosaccharides on apparent and true ileal amino acid digestibilities and
fecal consistency in growing pigs. J. Anim. Sci. 80:2433–2441.
Souffrant, W. B. 2001. Effect of dietary fibre on ileal digestibility and
endogenous nitrogen losses in the pig. Anim. Feed Sci. Technol.
90:93–102.
Sullivan, J. S., D. A. Knabe, A. J. Bockholt, and E. J. Gregg. 1989.
Nutritional value of quality protein maize and food corn for
starter and growth pigs. J. Anim. Sci. 67:1285–1292.
Summers, J. D. 2001. Maize: Factors affecting its digestibility and
variability in its feeding value. Pages 109–124 in Enzymes in
Farm Animal Nutrition. M. R. Bedford and G. G Partridge, ed.
CABI Publ., Wallingford, Oxon, UK.
Taverner, M. R., and R. G. Campbell. 1988. The effects of protected
dietary enzymes on nutrient absorption in pigs. Page 377 in
Digestive Physiology in the Pig. Proc. 4th Int. Symp. Dig. Physiol.
Pigs. L. Buraczewska, S. Buraczewski, B. Pastuszewska, and T.
Zebrowska, ed. Inst. Anim. Physiol. Nutr., Joblonna, Poland.
Thacker, P. A., G. L. Campbell, and J. W. D. Groot-Wassink. 1988. The
effect of beta-glucanase supplementation on the performance of
pigs fed hulless barley. Nutr. Rep. Int. 38:91–99.
Thacker, P. A., G. L. Campbell, and J. W. D. Groot-Wassink. 1989.
The effect of sodium bentonite on the performance of pigs fed
barley-based diets supplemented with beta-glucanase. Nutr.
Rep. Int. 40:613–619.
Thacker, P. A., G. L. Campbell, and J. W. D. Groot-Wassink. 1990.
The effect of enzyme supplementation on the nutritive value of
rye-based diets for swine. Can. J. Anim. Sci. 71:489–496.
Thacker, P. A., G. L. Campbell, and J. W. D. Groot-Wassink. 1992.
The effect of organic acids and enzyme supplementation on the
performance of pigs fed barley-based diets. Can. J. Anim. Sci.
72:395–402.
Trinder, P. 1969. Determination of glucose in blood using glucose
oxidase with an alternative oxygen acceptor. Ann. Clin. Biochem.
6:24–27.
Williams, C. H., and O. Iismaa. 1962. The determination of chromic
oxide in faeces samples by atomic absorption spectrophotometry.
J. Agric. Sci. 59:381–385.
Yin, Y.-L., S. K. Baidoo, L. Z. Jin, Y. G. Liu, H. Schulze, and P.
H. Simmins. 2001a. The effect of different carbohydrase and
protease supplementation on apparent (ileal and overall) digestibility of nutrients of five hulless barley varieties in young pigs.
Livest. Prod. Sci. 71:109–120.
Yin, Y.-L., S. K. Baidoo, H. Schulze, and P. H. Simmins. 2001b. Effects
of supplementing diets containing hulless barley varieties having different levels of non-starch polysaccharides with β-glucanase and xylanase on the physiological status of the gastrointestinal tract and nutrient digestibility of weaned pigs. Livest. Prod.
Sci. 71:97–107.
Zijlstra, R. T., S. Li, A. Owusu-Asiedu, P. H. Simmins, and J. F.
Patience. 2004. Effect of carbohydrase supplementation of
wheat- and canola-meal-based diets on growth performance and
nutrient digestibility in group-housed weaned pigs. Can. J.
Anim. Sci. 84:689–695.
Downloaded from jas.fass.org at Serials/Acq. Dept., Library on May 1, 2009.
References
This article cites 39 articles, 14 of which you can access for free at:
http://jas.fass.org/cgi/content/full/86/7/1533#BIBL
Downloaded from jas.fass.org at Serials/Acq. Dept., Library on May 1, 2009.