1 s2.0 S0032579119307734 Main
1 s2.0 S0032579119307734 Main
1 s2.0 S0032579119307734 Main
ABSTRACT Two broiler trials were designed to in- during starter, grower and finisher periods was mea-
vestigate the relationship between the concentration of sured.
non-starch polysaccharides (NSP) in wheat and 1) its In the balance trial, wheat cultivar did not affect nu-
nutritional value for broilers and 2) the efficacy of ex- trient digestibility or AMEn . Enzyme addition caused a
ogenous enzymes. In a balance trial, diets were formu- significant increase in nutrient digestibilities and AMEn
lated with 3 wheat cultivars (Rustic and Viscount— for the diet formulated with the high NSP wheat Cen-
medium NSP, Centenaire—high NSP) and were tested tenaire only. In the performance trial, feeding the high
with or without the addition of an exogenous enzyme NSP diet resulted in a higher feed conversion ratio
mixture. The diets were fed to 144 male Ross 308 broiler and lower final body weight compared to the medium
chickens housed in digestibility cages. Total tract nutri- NSP diet. The largest improvements by enzyme addi-
ent digestibilities and AMEn were measured from 18 to tion were observed in the high NSP diet.
22 d of age. In a performance trial, diets were formu- In conclusion, the study was not able to show a con-
lated with wheat (medium NSP diet) or with wheat sistent relationship between the NSP concentration of
mixed with rye and barley (high NSP diet) and were wheat and its nutritional value, but did demonstrate
tested with or without the addition of an exogenous that the effect of an enzyme mixture on nutrient di-
enzyme mixture. The diets were fed to 960 male Ross gestibility or broiler performance depends upon the
308 broilers housed in pens and broiler performance NSP concentration in the diet.
Key words: wheat, broiler, nutrient digestibility, AMEn , enzyme mixture
2018 Poultry Science 97:2064–2070
http://dx.doi.org/10.3382/ps/pey038
2064
INTERACTIONS BETWEEN WHEAT QUALITY AND ENZYMES 2065
To counter the possible anti-nutritional effects caused zyme addition, the other part was supplemented with
by NSP and to improve nutrient digestibility and an- a commercial enzyme mixture (KEMZYME R
Plus dry,
imal performance, NSP-degrading feed enzymes are Kemin Europa NV, Herentals, Belgium) at a dose rate
commonly added to wheat-based feed. The efficacy of 500 g/T of feed. Drinking water and feed (in mash
of feed enzymes is variable, however, and sometimes form) were provided for ad libitum consumption. The
no effects are observed (Gutiérrez-Alamo et al., 2008; balance trial was carried out according to the EU-
Svihus, 2011). In general, it is assumed that enzyme reference method (Bourdillon et al., 1990) and con-
addition is more effective in low-quality wheat (Choct sisted of a 5-d period of adaptation (to the respective
et al., 1995; Scott et al., 1998; Huyghebaert and experimental diets containing titanium dioxide, d 13
Schöner, 1999; Bedford, 2002). The definition of low- to 17) and a 5-d main balance period (d 18 to 22).
quality wheat is, however, not straightforward. Earlier Before the adaptation and main balance period (d 1
research showed a positive relationship between enzyme to 13), the chickens were housed together and fed the
efficacy and NSP concentration (Huyghebaert et al. same commercial mash starter diet (based on Aviagen,
1999; Smeets et al., 2014a; Smeets et al., 2015) and a 2009). Feed intake was recorded and all excreta were
negative relationship between enzyme efficacy and the collected in a quantitative way using plastic trays un-
arabinose to xylose ratio of arabinoxylan (Huyghebaert der the cages. A homogeneous sample of the mixed wet
et al., 1999; Smeets et al., 2014a). More research on the excreta was freeze-dried until analysis.
relationship between the chemical composition of the Performance Trial The trial was performed with
substrate (in particular the NSP) and enzyme efficacy 960 birds, housed in 32 pens (4 dietary treatments ×
is needed to be able to predict the nutritional value 8 replicates, 30 birds/pen). The birds were fed using
of wheat-based diets and the response of a diet to the a 3-phase feeding scheme with periods of 13 d each:
addition of NSP degrading enzymes. starter (d 1 to 13), grower (d 14 to 26) and finisher
The objectives in the present study were to study (d 27 to 39). Drinking water and feed (in mash form)
the relationship between the concentration of NSP in were provided ad libitum. Two basal diets for each of
wheat and 1) its nutritional value for broilers and 2) the 3-phase feeding periods were formulated using ei-
the efficacy of NSP-degrading enzymes. ther wheat (unknown cultivar) alone or wheat mixed
with rye and barley (83.8% wheat, 13.0% rye, 3.2%
barley, further referred to as wheatmix), to mimic a
MATERIALS AND METHODS wheat high in NSP. Diets were formulated to meet
breeder’s recommendations (Aviagen, 2009) and in-
Birds, Diets, and Management cluded a coccidiostaticum (0.5 g/kg Salinomycin) and a
phytase (0.2 g/kg Ronozyme NP). The ingredients and
The broiler balance and performance trial both
the chemical compositions of the grower diets and the
started at the same time at the Institute for Agricul-
wheat or wheatmix are shown in Table 1. One part of
tural and Fisheries Research (ILVO), Melle, Belgium
each diet was used without enzyme addition, the other
and were approved by the ILVO Ethical Committee
part was supplemented with KEMZYME Plus dry at a
(EC no. 2012/186). In total, 1,104 day-old male broiler
dose rate of 500 g/T of feed.
chickens (Ross 308) were obtained from a commer-
cial hatchery (Belgabroed NV, Merksplas, Belgium).
The broilers were vaccinated at 1 d of age against Collection of Intestinal and Blood Samples
Newcastle disease (Clone 30, spray, Intervet Interna- At the end of the balance trial, 2 animals per cage
tional BV, Boxmeer, the Netherlands) and infectious were randomly selected and euthanized following the
bronchitis (Poulvac IB Primer, spray, Pfizer Animal principles for care of animals in experimentation. In-
Health, Louvain-La-Neuve, Belgium). On d 16, vacci- testinal contents were collected from the ileum (between
nation against Newcastle disease was repeated (Clone Meckel’s diverticulum and the ileocecal junction) of the
30, drinking water). 2 animals by gently finger-stripping the intestinal seg-
Balance Trial The balance trial was carried out with ment. The digesta contents collected from 2 animals per
144 birds, housed in 36 digestibility cages (6 dietary cage were pooled and stored on ice until analysis.
treatments × 6 replicates, 4 birds/cage during the main At the end of the performance trial, blood samples
balance period). Three basal diets were formulated with were taken from one bird per pen. The blood samples
3 different wheat cultivars: Viscount, Rustic, and Cen- were collected in 5 mL Vacuette R
tubes containing Hep-
tenaire. The cultivars were chosen to differ mainly arin (456,083, Greiner Bio-One GmbH, Kremsmünster,
in NSP concentration. Diets were formulated to meet Austria), stored on ice until centrifugation (1,500 × g
breeder’s recommendations (Aviagen, 2009) and in- for 10 min) to obtain the plasma and then stored at
cluded a coccidiostaticum (0.5 g/kg Sacox; Salinomycin –21◦ C until analysis.
sodium; Huvepharma NV, Antwerpen, Belgium) and a
phytase (0.2 g/kg Ronozyme R
NP; DSM, Heerlen, The Chemical Analyses
Netherlands). The ingredients and the chemical com-
positions of the diets and wheat cultivars are shown in Samples of feeds and freeze-dried excreta were an-
Table 1. One part of each diet was used without en- alyzed at ILVO in an accredited lab for dry matter
2066 SMEETS ET AL.
Table 1. Composition of the experimental grower diets used in the balance trial and the performance trial.
Ingredient (g/kg)
Wheat 640.0 540.0
Rye - 80.0
Barley - 20.0
Soybean meal (49% CP) 267.8 265.0
Soya oil 24.0 45.9
Animal Fat 30.0 10.0
Limestone 5.2 6.8
Sodium bicarbonate 1.1 0.7
Sodium chloride 2.6 2.7
Monocalcium phosphate 11.3 8.2
L-Lysine hydrochloride 2.1 2.2
DL-Methionine 2.3 2.9
L-Threonine 0.6 2.7
Titanium dioxide 3.0 3.0
Vitamin/mineral premix1 10.0 10.0
Calculated diet composition (g/kg)
MEn broilers (kcal/kg) 2844 2844
Crude protein (N x 6.25) 195 195
Crude fat 67 69
Analyzed diet composition (g/kg) Viscount Rustic Centenaire Wheat Wheatmix2
(DM), gross energy, protein, and fat. In addition, the ple. This mixture was then incubated at 50◦ C for
titanium dioxide concentration in the feed was analyzed 2 h. Hereafter, 4 mL ethanol (95% v/v) was added, the
as described before (Smeets et al., 2015). Crude protein, mixture was centrifuged at 1,500 × g, and the absorp-
moisture, starch and NSP content and extract viscos- tion was measured spectrophotometrically at 585 nm.
ity of the wheat were measured as described by Smeets A portion of the intestinal samples was freeze-dried and
et al. (2015). The intestinal samples were centrifuged the concentration of titanium dioxide was quantified as
for 10 min at 10,000× g and the supernatants were ana- described before.
lyzed for ileal viscosity, xylanase inhibition activity, xy-
lanase activity and titanium dioxide concentration. Ileal
viscosity was determined at 40◦ C using a Brookfield Calculations and Statistical Analyses
cone and plate viscometer. For the estimation of xy-
lanase inhibition activity, the supernatant was diluted Total tract digestibility of DM, fat, starch, WE-
2 times with sodium acetate buffer (pH 5.0, 0.025 M) NSP, WU-NSP, and TOT-NSP, N retention, and ap-
and the assay was performed as described by Smeets parent metabolizable energy, corrected for nitrogen
et al. (2014b). Xylanase activity in the intestinal sam- (AMEn ), were calculated using the concentrations of
ples was evaluated by diluting the supernatant 10 times titanium dioxide tracer in the excreta and the feed
(in sodium acetate buffer, pH 5.0, 0.1 M) and adding (Smeets et al., 2015). Performance results (daily feed
a Xylazyme AX tablet (Megazyme International, Bray intake, body weight, feed conversion ratio [FCR]) were
Co. Wicklow, Ireland) to 1 mL of diluted digesta sam- calculated for each sub-period (starter, grower and
INTERACTIONS BETWEEN WHEAT QUALITY AND ENZYMES 2067
Table 2. Effect of wheat cultivar and enzyme addition on total tract digestibility.
Treatment1
1. wheat Viscount 69.5b,c 56.5b 70.3 96.1 1.9 21.9 19.8 3,160b,c 6.5b 0.075b 8.34
2. wheat Rustic 68.9b,c 57.6b 71.7 96.1 -21.9 21.3 17.6 3,165b,c 6.4b 0.074b 13.1
3. wheat 69.5b,c 57.8b 72.0 95.7 -8.6 31.5 27.5 3,186b,c 10.6a 0.071b 8.99
Centenaire
4. wheat Viscount 68.0c 55.6b 71.3 96.2 -2.2 13.8 12.1 3,107c 3.4c 0.202a 13.3
500 g/T enzyme
5. wheat Rustic 70.8b 58.6b 77.4 96.4 -0.9 28.5 25.9 3,264a,b 3.0c 0.173a 11.2
500 g/T enzyme
6. wheat 73.6a 63.4a 77.3 97.3 -3.7 29.9 26.6 3,362a 4.3b,c 0.200a 13.9
Centenaire 500 g/T
enzyme
Average 70.0 58.2 73.4 96.4 -5.9 24.5 21.6 3207 5.7 0.134 11.1
Pooled SEM 0.9 1.3 1.9 0.7 5.9 4.9 4.6 39 0.8 0.018 3.10
Main effect
Wheat
Viscount 68.7b 56.1b 70.8 96.2 -0.2 17.9 15.9 3,134b 5.0b 0.139 10.9
Rustic 69.9a,b 58.1a,b 74.5 96.2 -11.4 24.9 21.8 3,214a,b 4.7b 0.122 11.6
Centenaire 71.5a 60.6a 74.7 96.5 -6.1 30.7 27.0 3,274a 7.5a 0.136 10.9
Enzyme addition
No enzyme 69.3b 57.3 71.3b 96.0 -9.6 24.9 21.6 3,170b 7.9b 0.072b 9.8
500 g/T 70.8a 59.2 75.3a 96.6 -2.2 24.0 21.5 3,244a 3.6a 0.192a 12.4
P-value
Wheat 0.016 0.005 NS (0.08) NS NS NS (0.07) NS (0.09) 0.009 0.002 NS NS
Enzyme 0.050 NS (0.08) 0.01 NS NS NS NS 0.041 < 0.001 < 0.001 NS
Wheat × enzyme 0.014 0.044 NS NS NS NS NS 0.019 NS NS NS
1
Reported values are means of 6 replicates, Viscount and Rustic: medium concentration of NSP, Centenaire: high concentration of NSP.
2
absorbance (585 nm) of the final assay solution.
3
IU/g tracer, 1.0 IU = amount of inhibitor that results in 50% inhibition of a xylanase under the conditions of the assay
a–c
means within a column and main effects lacking a common superscript differ significantly (P ≤ 0.05).
Abbreviations: AMEn , apparent metabolizable energy corrected for nitrogen; DM, dry matter; NS, not significant; NSP, non-starch polysac-
charides; TOT, total; WE, water-extractable; WU, water-unextractable; XA, xylanase activity; XIA, xylanase inhibition activity.
finisher) and for the entire period. A balance cage (6 and Viscount (91.8 g/kg) had comparable and lower
replicates per treatment) or floor pen (8 replicates per values. The wheat used in the performance trial was
treatment) was the experimental unit. Data were ana- a standard wheat with a total NSP concentration of
lyzed by 2-way analysis of variance (ANOVA) using 96.1 g/kg DM. This wheat was mixed with rye and
Statgraphics Centurion XVI software (Statpoint Tech- barley to mimic wheat high in NSP. This mixture in-
nologies, Inc., Warrenton, VA). When interactions were deed had a high NSP concentration (103.9 g/kg DM),
observed (P ≤ 0.05), the least significant differences most pronounced in the WE part of the NSP.
procedure was used to separate individual treatment
means. The balance trial results were analyzed as a 3
× 2 factorial arrangement of dietary treatments involv- Balance Trial
ing 3 wheats (Viscount, Rustic, and Centenaire) and 2
enzyme dosages (0 and 500 g/T). The performance trial The average recovery of xylanase (main enzyme ac-
results were analyzed as a 2 × 2 factorial arrangement tivity present in the enzyme mixture) from the enzyme-
of dietary treatments involving 2 raw material sources treated feed for both the balance and performance trial
(wheat alone or wheatmix) and 2 enzyme dosages (0 was 104% (data not shown). The effect of different
and 500 g/T). All statements of significance were based wheat cultivars and enzyme addition on total tract di-
on a P-value less than or equal to 0.05. gestibilities, AMEn , ileal viscosity, ileal xylanase activ-
ity, and ileal xylanase inhibition activity is shown in
Table 2. No effect of wheat cultivar on nutrient di-
gestibility or AMEn was observed without enzyme ad-
RESULTS dition. Inclusion of the enzyme mixture significantly re-
Wheat Cultivars duced the ileal viscosity for all diets. In the diets with
the Rustic and Viscount wheat cultivars, this reduction
The results of the composition analysis for the dif- in ileal viscosity did not result in any improvements
ferent wheat samples and wheatmix are shown in in nutrient digestibility. In the diet with the high NSP
Table 1. Centenaire wheat had the highest concentra- wheat cultivar Centenaire, enzyme addition increased
tion of total NSP (111 g/kg), and Rustic (96.3 g/kg) DM digestibility, nitrogen retention, and AMEn . No
2068 SMEETS ET AL.
Table 3. Daily feed intake, body weight and feed conversion ratio during grower, finisher and the whole period in the performance
trial with wheat.
Daily feed intake (g) Body weight (g) Feed Conversion Ratio (g/g)
Starter Grower Finisher Total Starter Grower Finisher Starter Grower Finisher Total
(1–13d) (14–26d) (27–39d) (1–39d) (d13) (d26) (d39) (1–13d) (14–26d) (27–39d) (1–39d)
Treatment1
1. wheat 31.7a 104.9 192.3 106.6 336b 1,345b 2,607b 1.286a 1.456c 1.678 1.537b
2. wheat 32.7a 107.5 189.1 106.9 364a 1,379a,b 2,646a,b 1.208b 1.484b 1.641 1.518b,c
500 g/T enzyme
3. wheatmix 30.3b 103.7 193.0 105.7 324b 1,281c 2,530c 1.287a 1.517a 1.699 1.570a
4. wheatmix 32.0a 108.2 192.0 107.8 357a 1,396a 2,684a 1.212b 1.458c 1.639 1.508c
500 g/T enzyme
Average 31.7 106.1 191.6 106.7 345 1350 2617 1.248 1.479 1.664 1.533
Pooled SEM 0.4 0.9 1.9 0.9 5 12 18 0.013 0.007 0.015 0.008
Main effect
Diet
Wheat 32.2a 106.2 190.7 106.8 350a 1362 2627 1.247 1.470b 1.660 1.527
Wheatmix 31.2b 106.0 192.5 106.7 340b 1338 2607 1.249 1.488a 1.669 1.539
Enzyme addition
No enzyme 31.0b 104.3b 192.6 106.1 330b 1,313b 2,569a 1.286a 1.487b 1.688a 1.554a
500 g/T 32.3a 107.9a 190.5 107.3 361a 1,387a 2,665b 1.210b 1.471a 1.640b 1.513b
P-value
Diet 0.010 NS NS NS 0.040 NS (0.06) NS NS 0.023 NS NS
Enzyme 0.002 < 0.001 NS NS < 0.001 < 0.001 < 0.001 < 0.001 0.043 0.003 < 0.001
Diet x enzyme NS NS NS NS NS 0.003 0.003 NS < 0.001 NS 0.016
1
Reported values are means of 8 replicates, wheat: medium concentration of non-starch polysaccharides, wheatmix: mixture of wheat, rye and
barley to mimic a high concentration of non-starch polysaccharides.
a–c
means within a column lacking a common superscript differ significantly (P ≤ 0.05). NS: not significant.
effects of the dietary treatments on WE-NSP digestibil- improved the FCR in the diet with wheat alone dur-
ity could be observed. WU-NSP were the most di- ing starter and grower period, whereas the FCR was
gestible in the Centenaire wheat diet and the least improved by the enzyme in the wheatmix diet during
digestible in the Viscount wheat diet, when averaged each period.
across enzyme levels (P ≤ 0.10). The results shown in
Table 2 show some xylanase activity left in the ileum DISCUSSION
of the enzyme-treated feed. By use of the results of the
tracer analyses in the feed and intestinal content, it A first aim of the animal trials described in this paper
could be calculated that on average 12% of the original was to examine the effect of the variable NSP concen-
xylanase activity in the feed was present in the ileum. tration and composition of wheat on broiler digestion,
Also, some xylanase inhibition activity was present in similar to the previous experiment described by Smeets
the ileal samples, which was on average 1.3% of the et al. (2015). Unlike what was expected, based on the
inhibition activity in the feed. NSP composition of the wheat cultivars, and the higher
ileal viscosity in the Centenaire diet, no differences in
Performance Trial nutrient digestibilities or AMEn were observed between
the diets in the balance trial. Other reports also indicate
The mortality during the performance trial was on no relationship between AMEn and NSP concentration
average 3.1% and was not affected by the dietary treat- (Nicol et al., 1993; Austin et al., 1999; Parsaie et al.,
ments (P > 0.05; data not shown). The effect of the 2006), although there are a lot of researchers who do
2 diet formulations (wheat or wheatmix) and the addi- find a correlation (Dusel et al., 1997; Choct et al., 1999;
tion of an enzyme mixture on daily feed intake, body Smeets et al., 2015). A possible explanation for this lack
weight, and FCR during the starter, grower, finisher, of relationship could be related to the NSP digestibil-
and the total period is shown in Table 3. During the ity, as will be explained further in this discussion. In the
starter and grower period, feed intake was higher with performance trial, differences were observed in perfor-
the addition of the enzyme mixture (when averaged mance measures between the 2 diets (wheat: medium
across diets). The body weight was lower after the NSP—wheatmix: high NSP) without enzyme addition.
grower and finisher period with the wheatmix diet than These differences were most clear in the grower and
with wheat alone. At the end of the grower and fin- finisher periods. The birds fed the wheatmix diet were
isher periods, enzyme addition only improved the body less heavy and had a higher FCR than the birds fed the
weight in the wheatmix diet. The FCR was higher for wheat diet. These results indicate that the relationship
the wheatmix diet compared to wheat alone during the between AMEn and animal performance is not always
grower period and the whole period. Enzyme addition clear, as also observed by other researchers.
INTERACTIONS BETWEEN WHEAT QUALITY AND ENZYMES 2069
The second aim of the trials described in this pa- of “junction zones” between different AX molecules
per was to study the relationship between the NSP (Izydorczyk and Biliaderis, 1995). It can be expected
composition of wheat and the effect of enzyme ad- that these kinds of AX molecules are not readily sol-
dition. In the balance trial, addition of the enzyme ubilized and not fermented by the microbiota in the
caused a significant increase in nutrient digestibilities intestines. Furthermore, the presence of other side
and AMEn for the high NSP wheat Centenaire, whereas chains such as ferulic acids can also affect the water-
in the Viscount cultivar no differences were observed. extractability and hence possibly the digestibility of the
In the performance trial, inclusion of the enzyme mix- NSP (Izydorczyk and Biliaderis, 1995). The observation
ture improved the body weight (grower and finisher that the NSP from the Centenaire wheat were more di-
period) and FCR (finisher and total period) only for gestible can explain the lack of depression in nutrient
the wheatmix diet. These observations confirm the hy- digestibility observed with this wheat.
pothesis that the effect of enzyme addition is related to In conclusion, the study was not able to show a con-
the NSP concentration of the diet, as observed earlier sistent relationship between the NSP concentration of
(Choct et al., 1995; Dusel et al., 1998; Huyghebaert et wheat and its nutritional value, but did demonstrate
al., 1999) and as also demonstrated in vitro by Smeets that the effect of an enzyme mixture on nutrient di-
et al. (2014a). The concentration of NSP should there- gestibility or broiler performance depends upon the
fore be considered in the practice of feed formulation. NSP concentration in the diet.
In addition, the remaining xylanase activity in the
small intestine was measured in the current study. Only
a few other researchers report intestinal enzyme activ-
ACKNOWLEDGMENTS
ities (Silva and Smithard, 2002; Choct, 2006). It was The authors wish to acknowledge the “Instituut
calculated that on average 12% of the original xylanase voor de Aanmoediging van Innovatie door Wetenschap
activity was present in the ileum. This corresponds to en Technologie in Vlaanderen” (IWT, project number
the study by Silva and Smithard (2002) who estimated 100226, Brussels, Belgium) for financial support.
that between 15 and 20% of the xylanase added to a
rye-based feed was active in the small intestine. In addi-
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