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Two experiments were conducted to evaluate the effect of tannin levels in sorghum on coefficient of apparent ileal digestibility (CAID), coefficient of standardised ileal digestibility (CSID) of amino acids, and total and specific activity (TA, SA) of trypsin and chymotrypsin of growing pigs. In the first experiment, 24 castrates (60±5kg) were fitted with a simple T cannula to evaluate four different sorghum samples (I, II, III, and IV) with different tannin levels (1.4, 4.6, 9.8 and 10.0gkg−1). At the end of the trial, pigs were slaughtered to obtain samples of pancreas and intestinal digesta. Data were analysed as a randomized block design. The highest CAID (P
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Animal Feed Science and Technology
117 (2004) 245–264
Effect of tannins in sorghum on amino acid ileal
digestibility and on trypsin (E.C.2.4.21.4)
and chymotrypsin (E.C.2.4.21.1) activity
of growing pigs
G. Mariscal-Land´
ına,, J.H. Avellanedab, T.C. Reis de Souzac,
A. Aguilerac, G.A. Borbollad,B.Mar
e
aCentro Nacional de Investigaci´on en Fisiolog´ıa Animal (CENI Fisiolog´ıa), 76020 Quer´etaro, Mexico
bFacultad de Ciencias Pecuarias, Universidad T´ecnica Estatal de Quevedo, Ecuador
cFacultad de Ciencias Naturales, Universidad Aut´onoma de Quer´etaro, Mexico
dFacultad de Medicina Veterinaria y Zootecnia, Universidad Nacional Aut´onoma de M´exico, Mexico
eCEBETA 115, Col´on Quer´etaro, Mexico
Received 5 August 2003; received in revised form 17 May 2004; accepted 2 September 2004
Abstract
Two experiments were conducted to evaluate the effect of tannin levels in sorghum on coefficient
of apparent ileal digestibility (CAID), coefficient of standardised ileal digestibility (CSID) of amino
acids, and total and specific activity (TA, SA) of trypsin and chymotrypsin of growing pigs. In the
first experiment, 24 castrates (60 ±5kg) were fitted with a simple T cannula to evaluate four different
sorghum samples (I, II, III, and IV) with different tannin levels (1.4, 4.6, 9.8 and 10.0gkg1). At the
end of the trial, pigs were slaughtered to obtain samples of pancreas and intestinal digesta. Data were
analysed as a randomized block design. The highest CAID (P< 0.05) was observed on sorghum I and
the lowest on sorghum II. Leucine and glutamic acid were the most digestible amino acids in the four
samples of sorghum, with glycine, lysine, threonine and cysteine being the least digestible. As tannin
levels increased, proline CAID decreased (P<0.05). CSID of amino acids in sorghum I was higher
(P< 0.05) than on sorghum II, except for proline. Similarly, the CSID for isoleucine, lysine, threonine,
valine, alanine and aspartic acid was similar among sorghums I, III and IV. Tannin content did not
Corresponding author. Present address: Centro Nacional de Investigaci´
on en Fisiolog´
ıa Animal (CENI Fisi-
olog´
ıa). A.P. No. 1-1168 Quer´
etaro, Qro. 76001, Quer´
etaro, Mexico. Tel.: +52 419 292 00 36; fax: +52 419 292
00 33.
E-mail address: mariscal.gerardo@inifap.gob.mx (G. Mariscal-Land´
ın).
0377-8401/$ – see front matter © 2004 Elsevier B.V. All rights reserved.
doi:10.1016/j.anifeedsci.2004.09.001
246 G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264
affect pancreas weight, protein content, or TA and SA of trypsin and chymotrypsin on pancreatic
tissue. However, in the digesta of duodenum, trypsin TA was increased (P< 0.05) by 6.7% in the two
treatments with the highest tannin content. In the second experiment, 32 castrates (53±7 kg) were
fitted with a simple T cannula to evaluate eight different hybrids of sorghum: four Pioneer (8172,
8428, 8443, and 8641), and four Dekalb (D-45, D-65, D-68, and D-69). High-tannin sorghums (8172
and 8428) had the lowest (P<0.05) CAID for arginine, glycine, and proline. Furthermore, sorghum
8428 had the lowest (P< 0.05) CAID for lysine (0.339). The CAID for proline was negative (0.110)
for sorghum 8428 and extremely low (0.031) in sorghum 8172. These sorghums had a low (P<0.05)
CSID for arginine, histidine, and proline. In sorghums with a high content of tannins the CSID for
prolineand glycine was verylow.LysineCSID was lower(P< 0.05) in sorghum 8428, when compared
tothe others samples of sorghums.In conclusion,tannin levelsup to1.05% are not the maindepressors
of the CAID and CSID of amino acids in the sorghum grain; however, tannins have a clear negative
effect on CAID and CSID in sorghum with high levels (4% or more) of tannin.
© 2004 Elsevier B.V. All rights reserved.
Keywords: Tannins; Sorghum; Amino acids; Ileal digestibility; Pigs
1. Introduction
Sorghum is among the first five cereals produced worldwide, just behind rice, maize,
wheatand barley (FAOSTAT,2002). InMexico, sorghumis the second mostproduced cereal
and is almost exclusively used in animal feeds, representing 68% of the overall cereals used
by the Mexican livestock industry (Lastra and Peralta, 2000). However, the great variability
in its chemical composition and therefore its nutritive value (Moss´
e et al., 1988) may result
in low performance of growing pigs. Furthermore, the presence of antinutritional factors
such as tannins may also decrease digestibility. This antinutritional factor affects negatively
the efficient use of sorghum based diets (Kondos and Foale, 1983). There is evidence
that tannins have a detrimental effect on the ileal digestibility of protein and amino acids
(Cousins et al., 1981; Bell and Keith, 1989; Brand et al., 1990). However all these studies
used sorghums with high contrasting tannin levels, and experimental diets had different
protein levels. Both factors have a large impact on amino acid apparent ileal digestibility
(Fan et al., 1994; Mosenthin et al., 2000; Duodu et al., 2003). Therefore the objectives
of the present study were to evaluate the effect of tannins on coefficient of apparent ileal
digestibility (CAID) and coefficient of standardised ileal digestibility (CSID) of sorghum
protein and amino acids; and to study its effect on total and specific activity (TA, SA) of
trypsin and chymotrypsin in growing pigs.
2. Materials and methods
Two experiments were conducted in the metabolism unit of the CENI-Physiology
experimental farm, under the guidelines of the “International Guiding Principles for
Biomedical Research Involving Animals” as well as the “Mexican Official Standards for
the Production, Protection and Use of Lab Animals” (Diario Oficial de la Federaci´
on,
2001).
G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264 247
2.1. Animals
Twenty four castrates (Landrace ×Duroc) weighing 60 ±5kg and 32 castrates with
53±7 kg of weight were used in the firstand second experiments, respectively.Allcastrates
were fitted with a simple T cannula at the terminal ileum (Reis de Souza et al., 2000).
After surgery, pigs were placed in individual metabolism cages located in a building with
controlled temperature (19–22C). Post-surgical period lasted 21 days and castrates were
fedtwice a day (08.00and 17.00 h) witha grower dietcontaining 160 g kg1ofCP (Table2).
Feed was increased daily until animals reached their level of intake shown before surgery.
In the experimental period, castrates were fed 2.5 times their DE maintenance requirement,
460kJ of DE per kg of metabolic weight, kg BW0.75 (INRA, 1984). Castrates had free
access to water through a drinking nipple located on one of the walls of the metabolism
cage.
2.2. Sorghum
Twelve sorghum samples were evaluated in the present study (Table 1). In the first
experiment, four different sorghum samples (I, II, III and IV) with tannin levels of 1.4,
4.6, 9.8 and 10.0g kg1DM, respectively, were used. Samples for the experiment 1, were
obtained from suppliers located in the Mexican states of Guanajuato and Jalisco. In the
second experiment, eight different sorghum hybrids (Pioneer 8172, 8428, 8443, and 8641;
produced in the experimental fields of the University of Quer´
etaro; and Dekalb D-45, D-65,
D-68, and D-69 produced in an experimental field in the State of Guanajuato) were used.
2.3. Diets
Experimentaldiets(Table2) were formulated using one of four sorghum samples (exper-
iment 1), and one of eight sorghum hybrids (experiment 2). In both experiments, sorghum
was the only protein source because tannins can significantly modify the secretion, and
perhaps the excretion, of endogenous protein (Jansman et al., 1993, 1994, 1995). Similarly,
in order to reduce the effect of the dietary protein level on protein and amino acids CAID
(Fan and Sauer, 1995), experimental diets were formulated to provide the same quantity
of this nutrient by diluting the diets with maize starch when necessary. All diets were sup-
plemented with maize oil, calcium carbonate, calcium phosphate, salt, and vitamins and
minerals premix (Table 2). Maize oil was included at 18g kg1, to reduce dust from diets.
Vitamins and minerals were added to fulfil or exceed the requirements of the NRC (1998);
chromic oxide was included at 3g kg1as an indigestible marker.
2.4. Sampling and chemical analysis
The experimental period lasted 7 days (5 days for adaptation and 2 days for ileal digesta
collection). Ileal digesta was collected using plastic bags (11cm height, and 5 cm width),
containing 10 ml of a HCI 0.2N solution to block any bacterial activity. Bags were attached
tothe cannula, using arubber band, in intervalsof two hours,andremoved immediatelyafter
full. Digesta was immediately frozen at 20C until freeze dried in the laboratory. The dry
248 G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264
Table 1
Chemical composition of sorghums used in experiments 1 and 2a
Experiment 1, sorghum samples Experiment 2
I II III IV Pioneer Dekalb
8172 8428 8443 8641 D-45 D-65 D-68 D-69
Tannins (gkg1DM) 1.44.69.810.057.147.24.76.65.312.28.90.2
Dry matter 907.4 912.7 903.5 901.6 885.6 887.5 871.4 890.7 884.3 879.5 879.0 882.8
Protein 98.9 107.0 106.0 115.086.784.790.279.3 103.0 108.790.992.8
NDF 76.674.681.790.1 146.1 148.3 148.6 124.4 131.9 129.2 127.6 138.4
ADF 26.027.438.935.856.265.265.454.761.456.256.964.7
Ash 16.515.919.917.512.914.910.116.016.717.220.916.3
Tannins 1.44.69.810.057.147.24.76.65.312.28.90.2
Amino acids
Arginine 3.43.54.54.33.73.83.73.24.13.93.44.0
Histidine 2.02.22.72.61.91.82.11.72.42.32.02.1
Isoleucine 4.04.24.34.13.43.13.73.04.34.53.63.7
Leucine 13.013.913.715.111.711.212.610.615.716.213.312.8
Lysine 1.82.02.72.41.81.81.91.72.12.01.62.0
Methionine 1.61.62.02.31.51.41.51.21.71.61.41.6
Phenylalanine 5.15.55.55.84.74.44.94.16.06.25.05.0
Threonine 3.03.13.53.73.03.03.02.63.63.62.93.3
Valine 4.85.05.45.34.34.14.73.85.45.64.64.8
Tryptophane 0.60.70.60.80.50.60.70.60.90.60.61.0
Alanine 8.89.59.610.58.17.98.87.410.811.19.08.8
Aspartic acid 6.36.77.17.66.36.26.65.67.37.76.06.5
Cysteine 1.71.82.02.11.81.71.81.62.11.91.82.0
Glutamic acid 19.921.321.424.118.517.820.116.724.525.520.620.2
Proline 8.08.58.59.16.96.67.46.39.09.17.78.0
Serine 4.04.04.44.83.83.83.83.54.74.83.94.1
Tyrosine 3.23.53.43.83.23.13.32.84.04.13.43.4
Glycine 3.23.44.24.43.03.13.22.83.63.42.93.4
aIn g kg1of dry matter.
G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264 249
Table 2
Diet composition (experiments 1 and 2a)
Control diet Experiment 1, sorghum samples Experiment 2
I II III IV Pioneer Dekalb
8172 8428 8443 8641 D-45 D-65 D-68 D-69
Tannins (gkg1DM) 1.44.69.810.057.147.24.76.65.312.28.90.2
Sorghum 725.3 954.0 883.0 931.0 838.0 877.0 895.0 857.0 954.0 739.0 704.0 843.0 822.0
Soya bean meal 236.1
Maize starch 0.071.023.0 116.077.059.097.00.0 215.0 250.0 111.0 132.0
Maize oil 18.018.018.018.018.018.018.018.018.018.018.018.0
Tallow 8.6
Calcium phosphate 14.512.012.012.012.012.012.012.012.012.012.012.012.0
Calcium carbonate 6.05.55.55.55.55.55.55.55.55.55.55.55.5
Mineralsb3.53.53.53.53.53.53.53.53.53.53.53.53.5
Vitaminsc2.02.02.02.02.02.02.02.02.02.02.02.02.0
Salt 1.92.02.02.02.02.02.02.02.02.02.02.02.0
Chromium oxide 3.03.03.03.03.03.03.03.03.03.03.03.0
Lysine HCl 1.4
dl-Methionine 1.4
l-Threonine 0.3
Protein (g kg1) 166.680.581.683.785.772.468.576.472.974.680.283.675.0
Dry matter (g kg1) 890.1 877.5 883.6 879.3 877.7 887.2 889.7 889.4 887.2 891.9 890.3 890.2 891.5
DE (MJ kg)d14.213.313.413.313.513.413.413.513.313.713.713.513.5
aIn g kg1of dry matter.
bFurnish by kg of diet: Cu 8.4 mg, Fe 70.0mg, I 0.56 mg, Mn 21.0 mg, Se 0.18mg, Zn 84.0 mg.
cFurnish by kg of diet: Vitamin A 6400IU, D 1280IU, E 30 IU, K 0.8mg, choline 375mg, niacine 24 mg, pantotenic acid 11.0mg, riboflavine 4.8 mg, B12 30g.
dEstimated from tables (INRA, 1984).
250 G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264
frozen digesta, sorghums grains, and experimental diets were ground in a lab mill using a
0.5 mm mesh (Arthur H. Thomas Co., Philadelphia, PA) to perform the following analyses:
dry matter (DM) and crude protein (CP) were determined according to AOAC (Association
of Official Analytical Chemists) 934.01 and 976.05 respectively (AOAC, 2002), neutral
detergent fibre (NDF) and acid detergent fibre (ADF) as noted by van Soest et al. (1991),
chromic oxide (Fenton and Fenton, 1979), and tannins (Price et al., 1978). Amino acid
analyses were conducted using ion exchange chromatography after a 24h acid hydrolysis
with HCI 6N. Methionine and cysteine were oxidised to methionine sulphone and cisteic
acid by a performic oxidation procedure 994.12 (AOAC, 2002).
CAID, of DM, CP and amino acids of the experimental diets were calculated using Eq.
(1)
CAIDD=1IDAF
ADIF(1)
where CAIDDis the coefficient of apparent ileal digestibility of the component in the
assay diet, IDthe marker concentration in the assay diet (gkg1DM), AFthe component
concentration in ileal digesta (gkg1DM), ADthe component concentration in the assay
diet (gkg1DM), IFthe marker concentration in ileal digesta (g kg1DM). CSID were
calculated using the Eq. (2) (Furuya and Kaji, 1989)
CSIDD=CAIDD+Eil
AD(2)
where CSIDDis the coefficient of standardised ileal digestibility of a component in the
assay diet, CAIDDis as defined previously, Eil the endogenous (protein and amino acids)
ileal losses (g kg1DM intake) according to Mariscal-Land´
ın et al. (1995),ADis as defined
previously.
In both experiments, all castrates were slaughtered at the end of the trial using CO2,to
verify the non-existence of fistulae at small intestine level that would invalidate digestibility
measurements.
2.5. Enzymatic activity
In experiment 1 at the moment of slaughter, samples of pancreas and digesta were
obtained from those animals that did not exhibit intestinal anormalities. Digesta samples
wereobtained by dividingthe intestine tothree sections followingthe technique byMakkink
et al. (1994). Pancreas samples were immediately frozen using liquid nitrogen; and the
samples of digesta (duodenal, jejunal, and ileal) were placed on an aluminium tray, which
wassubmerged in a dry ice-cooled, methanol–acetonemix. Both types of samples were kept
at 70C until further analysis. Protein determination in pancreatic tissue was undertaken
using the technique by Lowry et al. (1951). Specific activity (substrate mol released per
minute per mg of protein) as well as total activity (substrate mol released per minute per
g of tissue or digesta) of trypsin were determined following the technique of Reboud et al.
(1962). Total and specific activity of chymotrypsin were determined with the technique by
Bergmeyer (1974).
G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264 251
2.6. Statistical analysis
CAID, CSID, TA and SA were analysed according to a randomized block design (Steel
and Torrie, 1980). In experiment 1, six blocks of four animals each were used; unfortu-
nately, two animals that consumed sorghum with 10.0 g tanninskg1(sorghum sample IV),
exhibited intestinal adherence that turned out to be a fistula, which invalidated their samples
for analysis. Thus that treatment had four observations, whereas the other treatments had
six observations each. In experiment 2, four blocks of eight animals each were used. Statis-
tical analysis was performed made using GLM procedure of SAS (SAS, 1990). Differences
among treatment means were compared using SNK test (Steel and Torrie, 1980). Amino
acid profiles from sorghum proteins (albumins, globulins, glutelins, and kafirins) reported
by Youssef (1998), and amino acid profile of the protein in the samples of sorghum used
in these studies were used to estimate the proportion of these proteins in the samples of
sorghum following the model proposed by Duvaux et al. (1990). This model allows esti-
mation of the proportion of different “reference” proteins in a mixture from their amino
acids profiles, using a multiple regression analysis without intercept. Two correspondence
factorial analyses (CFA) were performed using CORRESP procedure of SAS (SAS, 1990).
The main characteristics of this analyses data are: (a) samples with similar amino acids
profile are projected close together on the axes, (b) the proximity of the projection points
of an individual amino acid and of a sample indicates that the sample is characterized by
its proportion of the given amino acid. In the first CFA, amino acid profile of the protein
in the samples of sorghum were analysed. In the second CFA, the amino acid profile of
sorghums and digesta protein along with endogenous protein reported by Mariscal-Land´
ın
et al. (1995) were analysed. The amino acid profile of proline rich protein (PRP) reported
by Lu and Bennick (1998) was used as a supplementary variable.
3. Results
3.1. Protein profiles
Data of protein profile of the sorghums used in experiments 1 and 2 are shown in Table 3.
In sorghums D-68, II, D-65, D-45 and I (ranked from high to low); the higher the proportion
of kafirins and glutelins, the lower the proportion of albumins and globulins (R2= 0.98).
The CFA allowed to place all sorghum samples in relation to the different amino acids
in the plane defined by the two main axes as shown in Fig. 1. The first axis explains
65% of the total variation and sets sorghum III with low kafirin content (Table 3)onthe
right side against sorghums D-65 and D-68 with high kafirin content (Table 3) on the left
side. Second axis explains 18% of total variation and sets sorghums 8428 and 8172, rich
in globulins and albumins (Table 3) on the top side, against sorghums I and II with low
proportion of those proteins (Table 3) and sorghum III with the highest lysine content and
the lowest proportion of kafirin among all sorghums (Table 3), all sorghums on the bottom
side. Regarding amino acids (Fig. 1b), the first axis sets arginine, glycine, and lysine which
are rich in globulins and albumins on the right side, against glutamic acid and leucine rich in
kafirins on the left side. Second axis sets arginine and aspartic acid found in high proportion
252 G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264
Table 3
Protein profile of sorghums used in experiments 1 and 2a
Experiment 1, sorghum samples Experiment 2
I II III IV Pioneer Dekalb
8172 8428 8443 8641 D-45 D-65 D-68 D-69
Tannins (gkg1DM) 1.44.69.810.057.147.24.76.65.312.28.90.2
Protein (g kg1)98.9 107.106.0 115.086.784.790.279.3 103.0 108.790.992.8
Albumins 7.49.512.414.910.712.410.611.76.17.40.17.0
Globulins 16.513.018.713.319.920.518.217.716.615.519.619.8
Kafirins 55.554.945.552.653.552.155.253.257.261.658.651.6
Glutelins 19.121.121.817.714.914.014.916.318.714.320.420.5
R20.98 0.98 0.98 0.98 0.98 0.99 0.99 0.99 0.98 0.98 0.98 0.98
Proportion of
Albumins + globulins 23.922.531.128.230.632.928.829.422.722.919.726.8
Kafirins + glutelins 74.676.067.370.368.466.170.169.575.975.979.072.1
aProtein profiles obtained with the model proposed by Duvaux et al. (1990).
G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264 253
Table 4
Coefficients of apparent ileal digestibility, experiment 1
Sorghum samples SEMa
I II III IV
Tannins (gkg1) 1.4 4.6 9.8 10.0
Dry matter 0.786a 0.710b 0.696b 0.702b 0.0175
Protein 0.688a 0.570b 0.612ab 0.611ab 0.0264
Arginine 0.752a 0.624c 0.712b 0.665bc 0.0219
Histidine 0.699a 0.576c 0.640ab 0.599bc 0.0222
Isoleucine 0.761a 0.648b 0.720a 0.692ab 0.0226
Leucine 0.836a 0.759b 0.764b 0.772ab 0.0188
Lysine 0.452b 0.272c 0.604a 0.497ab 0.0354
Methionine 0.781a 0.661b 0.738a 0.736b 0.0224
Phenylalanine 0.809a 0.715b 0.741b 0.707b 0.0225
Threonine 0.551a 0.333b 0.578a 0.546a 0.0228
Valine 0.744a 0.596c 0.699b 0.643b 0.0290
Tryptophane 0.693a 0.566b 0.677b 0.680b 0.0320
Alanine 0.791a 0.690b 0.736ab 0.732ab 0.0208
Aspartic acid 0.696a 0.586b 0.671a 0.642ab 0.0234
Cysteine 0.636a 0.469c 0.558b 0.542b 0.0231
Glutamic acid 0.828a 0.738b 0.764b 0.759b 0.0184
Proline 0.707a 0.638ab 0.480b 0.511b 0.0417
Serine 0.711a 0.591b 0.655a 0.651ab 0.0198
Tyrosine 0.744a 0.623b 0.650b 0.607b 0.0276
Glycine 0.398ab 0.119c 0.407a 0.299b 0.0314
Means with different letters in the same row are different (P<0.05).
aStandard error of the mean.
in albumins and globulins on the top, against proline and isoleucine rich in kafirins on the
bottom side.
3.2. Ileal digestibility
3.2.1. Experiment 1
CAID results are shown in Table 4. Ileal digestibility of dry matter was higher (P<0.05)
in sorghum I (0.786) than in sorghums II, III and IV (0.710, 0.696 and 0.701, respectively).
Crude protein CAID was significantly higher (P<0.05) in sorghum I than in sorghum II;
sorghums III and IV had an intermediate value. Amino acid apparent ileal digestibility is
also presented in Table 4. For most amino acids evaluated CAID was higher (P<0.05) in
sorghum I, and varied from 0.398 for glycine to 0.836 for leucine. Sorghum II had, with
the exception of proline, the lowest (P<0.05) CAID, with values varying from 0.119 for
glycine to 0.759 for leucine. Amino acid CAID for sorghums, III and IV were generally
lower than in sorghum I, but equal or higher than the one observed in sorghum II (Table 4).
Leucine and glutamic acid were the most digestible; and glycine, lysine threonine and
cysteine the least digestible amino acid in all four sorghums. Proline was the only amino
acid in which CAID decreased as tannin levels increased (Table 4). Results of the CSID
are shown in Table 5. Similarly to CAID, the CSID was highest (P<0.05) in sorghum I,
254 G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264
G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264 255
Table 5
Coefficients of standardised ileal digestibility, experiment 1
Sorghum samples SEMa
I II III IV
Tannins (gkg1) 1.4 4.6 9.8 10.0
Protein 0.812a 0.694b 0.729b 0.731ab 0.0264
Arginine 0.862a 0.737b 0.795b 0.762b 0.0219
Histidine 0.789a 0.666b 0.708b 0.676b 0.0222
Isoleucine 0.837a 0.727b 0.793a 0.777ab 0.0226
Leucine 0.876a 0.800b 0.803b 0.812b 0.0188
Lysine 0.652a 0.461b 0.738a 0.667a 0.0354
Methionine 0.828a 0.711b 0.775a 0.772ab 0.0224
Phenylalanine 0.864a 0.770b 0.794b 0.762b 0.0225
Threonine 0.754a 0.544b 0.751a 0.730a 0.0228
Valine 0.831a 0.687b 0.779a 0.734ab 0.0290
Alanine 0.837a 0.737b 0.779ab 0.776ab 0.0208
Aspartic acid 0.809a 0.701b 0.774a 0.746ab 0.0234
Cysteine 0.730a 0.563c 0.637b 0.625bc 0.0231
Glutamic acid 0.869a 0.779b 0.803b 0.798b 0.0184
Proline 0.883a 0.816ab 0.650c 0.687bc 0.0417
Serine 0.845a 0.733b 0.779b 0.778b 0.0198
Tyrosine 0.789a 0.669b 0.694b 0.651b 0.0276
Glycine 0.678a 0.434c 0.627ab 0.526bc 0.0336
Means with different letters in the same row are different (P<0.05).
aStandard error of the mean.
and lowest (P<0.05) in sorghum II. Proline had a higher digestibility in sorghums I and II,
whereas in sorghums III and IV, the CSID for this amino acid decreased, being one of the
least digestible amino acids. In all four sorghums, leucine and glutamic acid had the highest
CSID, whereas glycine and cysteine had the lowest (Table 5).
3.2.2. Experiment 2
CAID for CP, DM and amino acids of sorghum hybrids used in experiment 2, is shown
in Table 6. CAID of crude protein and dry matter was not significantly different in any of
the sorghum samples analysed. Sorghum hybrids with a high content of tannins (8172 and
8428), had lower (P< 0.05) arginine, glycine and proline CAID, than low-tannins sorghums
(8443, 8641, D-45, D-65, D-68, and D-69). Furthermore, CAID for proline was negative
(0.110) in sorghum 8428, and extremely low (0.031) in sorghum 8172. Similarly, glycine
Fig. 1. First correspondence factorial analysis: profiles used were the amino acid profile in samples of sorghum
used in experiments 1 and 2. (a) The first axis explains 65% of total variation and sets sorghum III with low kafirin
content (Table 3) against sorghums D-65 and D-68 with high kafirin content. Second axis explains 18% of total
variation and sets sorghums 8428 and 8172, rich in globulins and albumins (Table 3), against sorghums I and II
with low proportion of those proteins and sorghum III with the highest lysine content and the lowest proportion of
kafirin among all sorghums. Regarding amino acids (b) first axis sets arginine, glycine, and lysine rich in globulins
and albumins, against glutamic acid and leucine rich in kafirins. Second axis sets arginine and aspartic acid found
in high proportion in albumins and globulins, against proline and isoleucine rich in kafirins.
256 G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264
Table 6
Coefficients of apparent ileal digestibility of sorghums used in experiment 2
Pioneer Dekalb SEMa
8172 8428 8443 8641 D-45 D-65 D-68 D-69
Tannins
(gkg1)57.147.2 4.7 6.6 5.3 12.2 8.9 0.2
Dry matter 0.766 0.760 0.788 0.799 0.806 0.821 0.783 0.793 0.0221
Protein 0.542 0.537 0.638 0.663 0.677 0.670 0.630 0.593 0.0338
Arginine 0.541b 0.524b 0.740a 0.715a 0.735a 0.694a 0.697a 0.731a 0.0256
Histidine 0.514 0.475 0.683 0.651 0.722 0.676 0.636 0.676 0.0307
Isoleucine 0.627ab 0.574b 0.695ab 0.688ab 0.736a 0.705ab 0.669ab 0.649ab 0.0329
Leucine 0.742 0.729 0.789 0.801 0.838 0.817 0.781 0.772 0.0245
Lysine 0.434ab 0.339b 0.579a 0.596a 0.626a 0.561a 0.535a 0.514a 0.0469
Methionine 0.724ab 0.687b 0.794ab 0.765ab 0.825a 0.765ab 0.747ab 0.760ab 0.0249
Phenylalanine 0.733 0.709 0.770 0.780 0.825 0.793 0.768 0.738 0.0260
Threonine 0.457 0.443 0.535 0.540 0.603 0.538 0.484 0.497 0.0458
Valine 0.616ab 0.586b 0.689ab 0.684ab 0.740a 0.699ab 0.685ab 0.648ab 0.0330
Tryptophane 0.606 0.636 0.626 0.703 0.749 0.579 0.642 0.653 0.0573
Alanine 0.699ab 0.674b 0.750ab 0.764ab 0.805a 0.783ab 0.739ab 0.720ab 0.0256
Aspartic acid 0.621 0.598 0.684 0.696 0.727 0.698 0.650 0.631 0.0322
Cysteine 0.454ab 0.416b 0.552ab 0.586ab 0.635a 0.535ab 0.523ab 0.549ab 0.0463
Glutamic
acid 0.743ab 0.724b 0.791ab 0.801ab 0.838a 0.816ab 0.784ab 0.768ab 0.0231
Proline 0.031b 0.110b 0.591a 0.498a 0.437a 0.594a 0.492a 0.544a 0.0924
Serine 0.592 0.590 0.659 0.693 0.729 0.695 0.651 0.659 0.0336
Tyrosine 0.643 0.639 0.689 0.719 0.752 0.717 0.673 0.681 0.0326
Glycine 0.079b 0.087b 0.309a 0.273a 0.351a 0.294a 0.260a 0.370a 0.0481
Means with different letters in the same row are different (P<0.05).
aStandard error of the mean.
showed a low (<0.370) CAID across all sorghum samples, and very low (P<0.05) in
sorghums with a high content of tannins (0.079 for sorghum 8172, and 0.087 for sorghum
8428). Sorghum 8428 (high in tannins), had the lowest (P<0.05), CAID for glutamic
acid, cysteine, alanine, valine, methionine, and isoleucine. Sorghum D-45 (low in tannins),
present the highest CAID; and intermediate in the rest of the sorghum samples (Table 6).
Sorghum 8428 (high tannins), showed the lowest (P<0.05) CAID for lysine (0.339), an
intermediate value (0.439), in sorghum 8172, and the highest values in the rest of the
sorghum samples (Table 6).
CSID in experiment 2 is shown in Table 7. The high levels of tannins in sorghums
8172 and 8428 resulted in lower values of CSID for arginine, histidine and proline when
compared with the other sorghums analysed. Methionine and alanine were less digestible
(P<0.05) in sorghum 8428 than in sorghum D-45, with intermediate CSID for the rest
of the sorghum samples. Similarly to the CAID, CSID was lowest for the high-tannins
sorghum 8428, intermediate for sorghum 8172, and highest for the other sorghum samples.
Lysine was less digestible (P<0.05) in sorghum 8428 than in the other grains. CSID for
glycine was highest (0.720) (P< 0.05) in sorghum D-45 (low in tannins) and lowest (0.437)
in sorghum 8428.
G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264 257
Table 7
Coefficients of standardised ileal digestibility of sorghums used en experiment 2
Pioneer Dekalb SEMa
8172 8428 8443 8641 D-45 D-65 D-68 D-69
Tannins
(gkg1)57.147.2 4.7 6.6 5.3 12.2 8.9 0.2
Protein 0.703 0.707 0.793 0.821 0.834 0.816 0.771 0.749 0.0337
Arginine 0.671b 0.654b 0.869a 0.850a 0.869a 0.834a 0.827a 0.856a 0.0256
Histidine 0.636b 0.605b 0.796a 0.772a 0.835a 0.789a 0.743a 0.790a 0.0308
Isoleucine 0.743 0.706 0.803 0.809 0.843 0.805 0.769 0.760 0.0328
Leucine 0.801 0.793 0.843 0.859 0.889 0.865 0.828 0.828 0.0245
Lysine 0.696ab 0.600b 0.822a 0.838a 0.888a 0.823a 0.796a 0.757ab 0.0469
Methionine 0.787ab 0.757b 0.857ab 0.835ab 0.889a 0.835ab 0.811ab 0.824ab 0.0249
Phenylalanine 0.812 0.796 0.845 0.862 0.896 0.860 0.836 0.815 0.0259
Threonine 0.717 0.714 0.794 0.811 0.851 0.785 0.732 0.744 0.0458
Valine 0.745 0.724 0.807 0.813 0.858 0.810 0.794 0.766 0.0331
Alanine 0.765ab 0.744b 0.811ab 0.829ab 0.863a 0.838ab 0.794ab 0.783ab 0.0256
Aspartic acid 0.768 0.752 0.826 0.848 0.875 0.837 0.792 0.779 0.0322
Cysteine 0.569 0.541 0.701 0.668 0.750 0.660 0.630 0.656 0.0463
Glutamic acid 0.801 0.786 0.844 0.859 0.888 0.864 0.831 0.823 0.0231
Proline 0.299b 0.175b 0.839a 0.761a 0.672a 0.825a 0.711a 0.783a 0.0924
Serine 0.774 0.779 0.841 0.875 0.899 0.859 0.815 0.835 0.0336
Tyrosine 0.704 0.703 0.748 0.780 0.808 0.771 0.725 0.740 0.0326
Glycine 0.478bc 0.437c 0.644abc 0.608abc 0.720a 0.644abc 0.630abc 0.678ab 0.0481
Means with different letters in the same row are different (P<0.05).
aStandard error of the mean.
3.2.3. Sorghum proteins, ileal digesta and endogenous protein
The second CFA compared amino acid profile of sorghums and ileal digesta protein
along with endogenous protein and the amino acid profile of proline rich protein (PRP)
was used as a supplementary variable. In Fig. 2, first axis of CFA explained 86% of the
total variation, and sets sorghum protein profile on the left side against endogenous protein
profile on the right side. Second axis explained 9% of the total variation and sets the ileal
digestas of sorghums I and II, rich in lysine on the top side, against digestas of sorghums
8172 and 8428, rich in proline on the bottom side. Ileal digesta of sorghums were closer
to the endogenous protein than the sorghum protein (Fig. 2a). Proline rich protein (PRP)
is located close to the area defined by this amino acid (Fig. 2a). Regarding amino acids
(Fig. 2b), first axis sets glutamic acid and leucine on the left side against glycine on the
rightside;second axis sets lysine and threonine on the top side against proline on the bottom
side.
3.2.4. Enzymatic activity
Pancreas weight and enzyme activity are shown in Table 8. Increasing levels of tan-
nins did not have any effect (P<0.05) on pancreas weight and protein content, or the
specific (IUmg1of protein) and total (IU g1of tissue) enzyme activity of trypsin and
chymotrypsin (Table 8). Total activity of trypsin in duodenum digesta (IUg1of digesta)
258 G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264
Fig. 2. Second correspondence factorial analysis, performed to compare sorghums protein profiles of experiment
1 (samples) and experiment 2 (Pioneer, Dekalb); sorghums ileal digesta profiles (samples, Pioneer, Dekalb),
endogenous protein profile (Mariscal-Land´
ın et al., 1995), and proline rich protein “PRP” profile (Lu and Bennick,
1998) used as supplementary variable. In figure a, first axis explain 86% of total variation and sets sorghum protein
profile against endogenous protein profile, second axis explain 9% of total variation and sets the ileal digestas of
sorghums I and II, rich in lysine, against digestas of sorghums 8172 and 8428, rich in proline. Ileal digesta of
sorghums were closer to endogenous protein than sorghum protein (a). Proline rich protein (PRP) is located close
to the area defined by this amino acid (a). (b) First axis sets glutamic acid and leucine against glycine; second axis
sets lysine and threonine against proline.
G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264 259
Table 8
Pancreas weight and protein content, and trypsin and chymotrypsin activity in pancreas and intestinal digesta
(experiment 1)
Sorghum samples SEMa
I II III IV
Tannins (gkg1)1.44.69.810.0
Pancreas weight (gkg0.75)4.34.24.34.40.42
Pancreas protein (mgg1) 160.4 159.9 160.0 159.52.80
Enzyme activity in pancreas, trypsin
Total activityb3351.3 3297.2 3209.9 3277.447.04
Specific activityc20.920.720.120.60.57
Chymotrypsin
Total activity b12869.4 12628.6 12993.7 13128.2 289.17
Specific activityc80.280.681.282.11.85
Enzyme total activity in small intestine, trypsind
Duodenum 3339.7a 3445.0ab 3564.1b 3525.4b 39.17
Jejunum 3354.7 3441.4 3438.2 3371.149.60
Ileum 3556.4 3424.7 3481.2 3433.347.76
Chymotrypsin
Duodenum 12188.79 12355.82 12417.07 12051.70 243.60
Jejunum 12561.01 12695.54 12941.77 12963.02 325.37
Ileum 12552.15 12464.95 12629.22 12585.49 362.52
Means with different letters in the same row are different (P<0.05).
aStandard error of the mean.
bTotal enzyme activity (IU g1of tissue).
cSpecific enzyme activity (IUmg1of protein).
dTotal enzyme activity (IU g1of ileal digesta).
had an increment (P<0.05) of 6.7% in the two diets with the higher tannin content, when
compared with the sorghum with low content of tannins. No effect was observed (P>0.05)
on total activity of trypsin and chymotrypsin in digesta of jejunum or ileum, for the first
enzyme, or the digesta of duodenum, jejunum or ileum in the second enzyme.
4. Discussion
4.1. Protein profile
In sorghum, 95% of the nitrogen is amino nitrogen; therefore, an increment in the protein
content of the grain increases almost exclusively the amount of storage proteins which
are present in a ratio of 60% kafirins and 40% glutelins (Moss´
e et al., 1988). Kafirins
are characterized by their low content of lysine (0.4%), relatively high leucine (15.6%),
glutamic acid (31.3%) and proline (10.4%) (Youssef, 1998). In the present study, sorghums
D-68, II, D-65, D-45 and I were proportionally rich in leucine, and with a low content of
lysine. Contrarily, sorghum III and the Pioneer hybrids 8172, 8428, 8443, and 8641 were
proportionally rich in lysine and with a low content of leucine. This difference in the amino
acids profile explains the higher amount of kafirins and glutelins for the first group of the
260 G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264
sorghums; and albumins and globulins for the second group of grains, as well as, their
spatial location in Fig. 1a, which is determined by the relative richness of amino acids in
the hybrids (Fig. 1b).
4.2. Ileal digestibility
The method used to estimate CAID was the direct method (Mosenthin et al., 2000), in
the case of sorghum and sources with low protein and amino acid content, CAID values
obtained using this method could be affected by the differences in protein and amino acid
content in the assay diets (Fan et al., 1994). Nevertheless this method was chosen to avoid
the negative effect of tannins on protein added from other sources, since condensed tannins
comprise a group of phenolic compounds (proanthocyanidins) that protect sorghum grain
against insects, birds and fungal attack, and reduce food quality because they are capable
of binding and precipitating protein (Schofield et al., 2001; Duodu et al., 2003).
Data reported in the literature show that the apparent ileal digestibility of the protein and
amino acids in sorghum is extremely variable (Cousins et al., 1981; Lin et al., 1987; Brand
et al., 1990). This variation has normally been attributed to the presence of tannins as in the
case with other raw materials (faba bean), where tannins are present (Jansman et al., 1993,
1994, 1995; Mariscal-Land´
ın et al., 2002).
However, recent studies indicate that the presence of kafirins may negatively affect the
apparent ileal digestibility of the protein (Oria et al., 1995; Elkin et al., 1996).
In experiment 1. The high CAID of DM, CP and amino acids (exception with lysine), in
the diet formulated with sorghum I may be due to the low content of tannins. Furthermore,
this diet was the most digestible even though it did not contain any maize starch, which
is considered a highly digestible ingredient (Susenbeth et al., 1999). The higher CAID of
lysine in sorghum III may be explained by the fact that this sorghum had the larger content
of this amino acid which positively affected the value of CAID (Fan and Sauer, 1995); and
that its protein profile was the highest in albumins and globulins; proteins more digestible
than kafirins (Oria et al., 1995; Elkin et al., 1996). Moreover, the high CAID in sorghums
III and IV, despite their higher tannin content (double than in sorghum II) may be the result
of their highest concentration of albumins and globulins. Sorghum II was characterized
for its highest concentration of storage protein (76% of glutelins and kafirins), and an
intermediate content of tannins. Probably, this combination of factors was responsible for
the lowest CAID for protein and amino acids among all sorghums evaluated. Low CAID
of proline in sorghums III and IV in experiment 1, and of proline and glycine in the high-
tannins hybrids (8172 and 8428) in experiment 2 may be the result of the tannin capacity to
stimulate the synthesis and secretion of PRP in the saliva of pigs (Jansman, 1993), which
are rich in proline (40%) and glycine (20%), and represent the first line of defence against
the presence of tannins in the diet (Mehansho et al., 1985). Stimulation for PRP secretion
follows the fact that tannins are able to precipitate proteins, by binding predominantly
the amino acid proline (Hagerman and Butler, 1981; Mitaru et al., 1984; Charlton et al.,
2002). Therefore, the lower concentration of tannins in sorghum I (experiment 1), may
have resulted in a reduced secretion of PRP which increased the CAID for proline when
compared to the sorghums III and IV. However, in experiment 2, the high content of tannins
in sorghums 8172 and 8428 apparently reduced the CAID for proline and glycine due to
G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264 261
the strong stimulation in the secretion of PRP. In the case of sorghum 8428, the CAID
for proline was negative. Low arginine CAID could be due to the capacity of this amino
acid to stimulate the binding between proteins and tannins (Charlton et al., 2002). Low
lysine, methionine and alanine CAID could be due to the precipitation of globular proteins
(albumins and globulins) originated by tannins.
CSID in experiments 1 and 2 had the same arrange that apparent ileal digestibility
observed in both experiments, but with higher values once the amount of basal endogenous
was eliminated. The results in the present studies are similar to those reported by AFZ
(2000) and Jondreville et al. (2001), with the exception of cisteine, where in the present
study the CSID for this amino acid was 10 units lower than the value reported by these
authors. Contrarily, proline CSID reported by AFZ (2000) was 26 units below the CSID
reported in the present study. The values of CSID reported by Pedersen and Boisen (2002)
are slightly superior, particularly in the case of lysine and cisteine.
4.3. Protein profile of sorghums, ileal digesta and endogenous protein
Second CFA analysis clearly situate sorghums protein profiles and endogenous protein
profiles on opposite section in axis 1 of Fig. 2a, situating ileal digesta profiles of sorghums
closerto the endogenous protein. Ilealdigestasmove along axis2, from those proportionally
rich in proline (digestas of sorghums 8428 and 8172), where the amino acid profile is
influencedby the amino acid profile of PRPprotein (since their high relativeproline content,
and their low proportion of threonine exhibited a higher proline excretion than the one
obtained if only the excretion of endogenous proteins had been increased), to the digesta of
sorghum II, which seems to be associated with an increase in endogenous protein since its
profile is proportionally higher in glycine and threonine as it is in the endogenous protein
(Mariscal-Land´
ın et al., 1995). This is in agreement with the report by Jondreville et al.
(2001), who estimated a higher excretion of endogenous protein in pigs fed sorghum base
diets, in comparison to other cereals (maize, wheat, rye and triticale).
4.4. Enzymatic activity
Enzyme inhibition by tannins has been reported in rabbits (Al-Mamary et al., 2001),
rats (Horigome et al., 1988), broilers (Ahmed et al., 1991), and pigs (Jansman et al., 1993;
Lizardo et al., 1995). The ability of tannins to precipitate proteins may be responsible of
this inhibition. All of these studies have compared low against high-tannin levels, with
the high levels used being similar to the level of tannins found in sorghums 8428 and
8172; and the low levels (<1%), similar to the level found in most of the sorghums used
in the present study. Inhibition of TA, and EA of digestive enzymes occurred in different
places of the small intestine. In duodenum, trypsin (Al-Mamary et al., 2001; Horigome
et al., 1988), amylase (Al-Mamary et al., 2001; Horigome et al., 1988), and lipase (Al-
Mamary et al., 2001); in jejunum, trypsin, amylase, and lipase (Horigome et al., 1988); and
in ileum, trypsin (Horigome et al., 1988; Jansman et al., 1994), and amylase (Horigome
et al., 1988). The results of the present study suggest that pigs fed diets with the highest
concentration of tannins, the TA of trypsin increased in the duodenum. This increase in the
trypsin activity may be the result of the tannin stimulation in the secretion of pancreatic
262 G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264
enzymes as suggested by Griffiths and Moseley (1980). This response may be related with
the observed increment in the EA of amylase (Ahmed et al., 1991), and chymotrypsin
(Lizardo et al., 1995) in the pancreas. The increase in the pancreatic secretion observed in
pigs fed sorghum II may be the result of the low digestibility of the high amounts of kafirins
found in this sorghum, which increased the secretion of trypsin. Similarly, Peiniau et al.
(1996) and Valette et al. (1992) reported that feeds with low digestible proteins tended to
stimulate the pancreatic secretion of enzymes.
5. Conclusion
The results of the study did not show a clear detrimental effect of tannins on the CAID
and CSID of sorghums with up to 10.0g of this compound per kg of dry matter. However,
the study may indicate that the CAID and CSID may be more influenced by the protein
profile of the grain. Therefore, it is recommended that more research on the role of kafirins
on the digestibility of the protein and amino acids in pigs is conducted. A clear detrimental
effectof tannins (47.2 and 57.1g kg1in dry matter) on CAID and CSID ofproline, glycine,
lysine, methionine and arginine was observed.
Acknowledgements
To Juan Carlos Anguiano for taking care of animals in experiment 2; Ericka Ramirez
Ramirez and Juan Guillermo Cervantes Huerta for their support with laboratory analyses.
To Guanajuato Produce A.C. foundation for its grant to this research. PURINA Mexico and
FERMEX for amino acids analyses.
References
AFZ,Ajinomoto Eurolysine, AventisAnimal Nutrition, INRA, ITCF,2000. Digestibilit´
esil´
ealesstandardis´
eesdes
acides amin´
es des mati`
eres premi`
eres chez le porc. AMI PIG CDROM. Association Franc¸aise de Zootechnie,
Paris, France.
Al-Mamary, M., Al-Habori, M., Al-Aghbari, A., Al-Obeidi, A., 2001. In vivo effects of dietary sorghum tannins
on rabbit digestive enzymes and mineral absorption. Nutr. Res. 21, 1393–1401.
Ahmed, A.E., Smithard, R., Ellis, M., 1991. Activities of enzymes of the pancreas, and the lumen and mucosa of
the small intestine in growing broiler cockerels fed on tannin-containing diets. Br. J. Nutr. 65, 189–197.
AOAC, 2002. Official Methods of Analysis, 17th ed. Association of Official Analytical Chemists, Arlington, VA.
Bell, J.M., Keith, M.O., 1989. Factors affecting the digestibility by pigs of energy and protein in wheat, barley
and sorghum diets supplemented with canola meal. Anim. Feed Sci. Technol. 24, 253–265.
Bergmeyer, H.U., 1974. Methods of Enzymatic Analysis, 3rd ed. Academic Press, New York.
Brand, T.S., Badenhorst, H.A., Siebrits, F.K., 1990. The use of pigs both intact and with ileo-rectal anastomosis to
estimate the apparent and true digestibility of amino acids in untreated, heat-treated and thermal-ammoniated
high-tannin grain sorghum. S. Afr. J. Anim. Sci. 20, 223–228.
Charlton, A.J., Baxter, N.J., Khan, M.L., Moir, A.J.G., Haslam, E., Davies, A.P., Williamson, M.P., 2002. Polyphe-
nol/peptide binding and precipitation. J. Agric. Food Chem. 50, 1593–1601.
Cousins, B.W., Tanksley, T.D., Knabe, D.A., Zebrowska, T., 1981. Nutrient digestibility and performance of pigs
fed sorghums varying in tannin concentration. J. Anim. Sci. 53, 1524–1537.
G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264 263
Diario Oficial de la Federaci´
on, 2001. Norma Oficial Mexicana-NOM-062-ZOO-1999, Especificaciones t´
ecnicas
para la producci´
on, cuidado y uso de los animales de laboratorio. Diario Oficial de la Federaci´
on, 2001.
Duodu, K.G., Taylor, J.R.N., Belton, P.S., Hamaker, B.R., 2003. Mini review: factors affecting sorghum protein
digestibility. J. Cereal Sci. 38, 117–131.
Duvaux, C., Guilloteau, P., Toullec, R., Sissons, J.W., 1990. A new method for estimating the proportions of
different proteins in a mixture using amino acids profiles: application to undigested proteins in the preruminant
calf. Ann. Zootech. 39, 9–18.
Elkin, R.G., Marisue, B.F., Hamaker, B.R., Zhang, Ye, Parsons, A.M., 1996. Condensed tannins are only partially
responsible for variations in nutrient digestibilities of sorghum grain cultivars. J. Agric. Food Chem. 44,
848–853.
Fan, M.Z., Sauer, W.C., 1995. Determination of apparent ileal amino acid digestibility in barley and canola meal
for pigs with the direct, difference, and regression methods. J. Anim. Sci. 73, 2364–2374.
Fan, M.Z., Sauer, W.C., Hardin, R.T., Lien, K.A., 1994. Determination of apparent ileal amino acid digestibility
in pigs: effects of dietary amino acid level. J. Anim. Sci. 72, 2851–2859.
FAOSTAT, 2002. FAOSTAT Home Page. Available: http://apps.fao.org/. Accessed July 15.
Fenton, T.W., Fenton, M., 1979. An improved procedure for determination of chromic oxide in feed and feces.
Can. J. Anim. Sci. 59, 631–634.
Furuya, S., Kaji, Y., 1989. Estimation of the true ileal digestibility of amino acids and nitrogen from their apparent
values for growing pigs. Anim. Feed Sci. Technol. 26, 271–285.
Griffiths, D.W., Moseley, G., 1980. The effect of diets containing field beans of high or low polyphenolic content
on the activity of digestive enzymes in the intestines of rats. J. Sci. Food Agric. 31, 255–259.
Hagerman, A., Butler, L., 1981. The specificity of proanthocyanidin–protein interactions. J. Biol. Chem. 256,
4494–4497.
Horigome, T., Kumar, R., Okamoto, K., 1988. Effects of condensed tannins prepared from leaves of fodder plants
on digestive enzymes in vitro and in the intestine of rats. Br. J. Nutr. 60, 275–285.
INRA, 1984. L’alimentation des animaux monogastriques: porc, lapin, volailles. Institut National de la Recherche
Agronomique, Paris, France.
Jansman, A.J.M., 1993. Tannins in feedstuffs for simple-stomached animals. Nutr. Res. Rev. 6, 209–236.
Jansman, A.J.M., Huisman, J., Van der Poel, A., 1993. Ileal and faecal digestibility in piglet of field beans (Vicia
faba L.) varying in tannin content. Anim. Feed Sci. Technol. 42, 83–96.
Jansman, A.J.M., Enting, H., Verstegen, M.W.A., Huisman, J., 1994. Effect of condensed tannins in hulls of faba
beans (Vicia faba L.) on the activities of trypsin (EC 2.4.21.4) and chymotrypsin (EC 2.4.21.1) in digesta
collected from the small intestine of pigs. Br. J. Nutr. 71, 627–641.
Jansman, A.J.M., Verstegen, M.W.A., Huisman, J., van den Berg, J.W.O., 1995. Effects of hulls of faba beans
(Vicia faba L.) with low or high content of condensed tannins on the apparent ileal and fecal digestibility of
nutrients and the excretion of endogenous protein in ileal digesta and feces of pigs. J. Anim. Sci. 73, 118–
127.
Jondreville,C.,vanden Broecke, J.,Gatel, F.,Grosjean,F.,vanCauwenberghe,S.,S`
eve,B.,2001. Ileal digestibility
of amino acids and estimates of endogenous amino acid losses in pigs fed wheat, triticale, rye, barley, maize
and sorghum. Anim. Res. 50, 119–134.
Kondos, A.C., Foale, M.A., 1983. Comparison of the nutritional value of low and medium tannin sorghum grains
for pigs. Anim. Feed Sci. Technol. 8, 85–90.
Lastra, M.I.J., Peralta, A.M.A., 2000. Situaci´
on actual y perspectiva de la carne de porcino en M´
exico. Available:
http://www.sagar.gob.mx/. Accessed July 15, 2002.
Lin, F.D., Knabe, D.A., Tanksley, T.D., 1987. Apparent digestibility of amino acids, gross energy and starch in
corn, sorghum, wheat, barley, oats groats and wheat middlings for growing pigs. J. Anim. Sci. 64, 1655–
1663.
Lizardo,R., Peiniau, J., Aumaitre, A., 1995. Effect of sorghum on performance, digestibility ofdietary components
and activities of pancreatic and intestinal enzymes in the weaned piglet. Anim. Feed Sci. Technol. 56, 67–82.
Lowry, O.H., Rosebrough, N.J., Farr, A.L., Randall, R.J., 1951. Protein measurements with the folin phenol
reagent. J. Biol. Chem. 193, 265–275.
Lu, Y., Bennick, A., 1998. Interaction of tannin with human salivary proline-rich proteins. Archs. Oral Biol. 43,
717–728.
264 G. Mariscal-Land´ın et al. / Animal Feed Science and Technology 117 (2004) 245–264
Makkink, C.A., Berntsen, P.J.M., op den Kamp, B.M.L., Kemp, B., Verstegen, W.A., 1994. Gastric protein break-
down and pancreatic enzyme activities in response to two different dietary protein sources in newly weaned
pigs. J. Anim. Sci. 72, 2843–2850.
Mariscal-Land´
ın, G., S`
eve, B., Coll`
eaux, Y., LeBreton, Y., 1995. Endogenous amino nitrogen collected from pigs
with end to end ileorectal anastomosis is affected by the method of estimation and altered by dietary fiber. J.
Nutr. 125, 136–146.
Mariscal-Land´
ın, G., LeBreton, Y., S`
eve, B., 2002. Apparent and standardised true ileal digestibility of protein
and amino acids from faba bean, lupin and pea, provided as whole seeds, dehulled or extruded in pig diets.
Anim. Feed Sci. Technol. 97, 183–198.
Mehansho, H., Clements, S., Sheares, B.T., Smith, S., Carlson, D.M., 1985. Induction of proline-rich glycoprotein
synthesis in mouse salivary glands by isoproterenol and by tannins. J. Biol. Chem. 260, 4418–4423.
Mitaru, B.N., Reichert, R.D., Blair, R., 1984. The binding of dietary protein by sorghum tannins in the digestive
tract of pigs. J. Nutr. 114, 1787–1796.
Mosenthin, R., Sauer, W.C., Blank, R., Huisman, J., Fan, M.Z., 2000. The concept of digestible amino acids in
diet formulation for pigs. Livest. Prod. Sci. 64, 265–280.
Moss´
e, J., Huet, J.C., Baudet, J., 1988. The amino acid composition of whole sorghum grain in relation to its
nitrogen content. Cereal Chem. 65, 271–277.
NRC, 1998. Nutrient Requirements of Swine, 10th ed. National Academy Press, Washington, DC.
Oria, M.P., Hamaker, B.R., Shull, J.M., 1995. Resistance of sorghum ,, and -kafirins to pepsin digestion. J.
Agric. Food Chem. 43, 2148–2153.
Pedersen, C., Boisen, S., 2002. Establishment of tabulated values for standardized ileal digestibility of crude
protein and essential amino acids in common feedstuffs for pigs. Acta Agric. Scand., Sect. A: Animal Sci. 52,
121–140.
Peiniau, J., Aumaitre, A., Lebreton, Y., 1996. Effects of dietary protein sources differing in solubility on total tract
and ileal apparent digestibility of nitrogen and pancreatic enzymes activity in early weaned pigs. Livest. Prod.
Sci. 45, 197–208.
Price, M.L., Steve, V.S., Butler, L.C., 1978. A critical evaluation of the vanillin reaction as an assay for tannin in
sorghum grain. J. Agric. Food Chem. 26, 1214–1218.
Reis de Souza, T.C., Mar, B.B., Mariscal, L.G., 2000. Canulaci´
on de cerdos posdestete para pruebas de
digestibilidad ileal: desarrollo de una metodolog´
ıa. T´
ec. Pecu. M´
ex. 38, 143–150.
Reboud, J.P., Ben Abdeljlil, A., Desnuelle, P., 1962. Variations de la teneur en ezymes du pancr´
eas de rat en
fonction de la composition des r´
egimes. Biochim. Biophys. Acta 58, 326–327.
SAS, 1990. SAS/STAT User’s Guide, Version 6, 4th ed. SAS Institute Inc., Cary, NC.
Schofield, P., Mbugua, D.M., Pell, A.N., 2001. Analysis of condensed tannins: a review. Anim. Feed Sci. Technol.
91, 21–40.
Steel, R.G.D., Torrie, J.H., 1980. Principles and procedures of statistics. In: A Biometrical Approach, 2nd ed.
McGraw-Hill Kogakusha, Ltd.
Susenbeth, A., Dickel, T., Diekenhorst, A.M., H¨
ohler, D., 1999. The effect of energy intake, genotype, and body
weight on protein retention in pigs when dietary lysine is the first-limiting factor. J. Anim. Sci. 77, 2985–2989.
Valette, P., Malouin, H., Corring, T., Savoie, L., Gueugneau, A.M., Berot, S., 1992. Effects of diets containing
casein and rapeseed on enzyme secretion from the exocrine pancreas in the pig. Br. J. Nutr. 67, 215–222.
vanSoest, P.J.,Robertson, J.B., Lewis,B.A., 1991. Methods fordietary fiber,neutral detergent fiber and non-starch
polysaccharides in relation to animal nutrition. J. Dairy Sci. 74, 3583–3597.
Youssef, A.M., 1998. Extractability, fractionation and nutritional value of low and high tannin sorghum proteins.
Food Chem. 63, 325–329.
... Amino acid digestibility showed a pattern similar to that of the crude protein, and the mean differences among treatments were in the range of 5.4-12.6%. In their work, Mariscal-Landín et al. (2004) studied the effect of tannins in sorghum on the coefficient of apparent ileal digestibility and on the coefficient of standardised ileal digestibility of amino acids. Four samples with different levels of tannins were considered (1.4, ...
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... 27 Mariscal-Landín found that tannis levels up to 4% reduce the digestibility of dietary protein in growing pigs. 28 In the present study, no negative effects on growth performance were found in 2.5 mg kg À1 , 5 mg kg À1 and 10 mg kg À1 TA-pretreated mice, indicating that pretreatment of 2.5 mg kg À1 to 10 mg kg À1 TA in mice do not have antinutritional effects. These results were in line with previous studies showing that dietary supplementation with a small quantity of the right kind of tannis may have no effect on growth performance. ...
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Low tannin sorghum cultivars (LTS) have been previously proved to contain greater available energy and apparent total tract digestibility (ATTD) of gross energy (GE) and crude protein (CP) than high tannin sorghum cultivars (HTS); however, their comparative ileal digestibility of energy and nitrogen excluding microbial interference in pigs remains mostly unknown. This study was designed to compare apparent (AID) and standardized ileal digestibility (SID) of amino acids (AA) associated with the AID and hindgut disappearance (HGD) of GE and CP in 4 LTS and 4 HTS fed to growing pigs. Eighteen barrows (27.6 ± 3.5 kg) fitted with a distal ileum T-cannula were allotted to a replicated 9×3 Youden square design with 9 diets and 3 periods to give a total of 6 replicate pigs per diet. Each period lasted 10 days, with 5 days adaption to the diets followed by 3 days collection of faeces and then 2 days collection of ileal digesta. Eight sorghum diets contained 966 g/kg sorghum grain as the only source of dietary energy and nitrogen, and one nitrogen-free diet was used to determine basal ileal endogenous nitrogen loss. Mean AID, ATTD, and HGD of GE and CP were higher in LTS than in HTS (P < 0.05). Mean SID of 8 out of 15 AA were decreased in HTS (P < 0.05). The AID, ATTD, and HGD of GE or CP had a negative correlation with condensed tannins and total phenolics in sorghum grain (P < 0.05). The SID of essential AA, including lysine, threonine, valine, histidine, and arginine, were highly or moderately negatively correlated with condensed tannins and total phenolics content in sorghum grains (P < 0.05). Overall, HTS provided less ileal digestibility and hindgut disappearance of energy and nitrogen, implying that condensed tannins in sorghum grain may impede nutrient digestibility, not only in the foregut but also in the hindgut segements of pigs.
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This study was conducted to test the hypothesis that low‐tannin sorghum grain produced in China as a potential substitute for corn in diets could not impair the performance of nursery pigs. A total of 60 pigs (7.2 ± 1.2 kg) were randomly assigned to 2 diets with 5 replicate pens per treatment. Corn‐based diet (CBD) included 60% corn grain during the overall experimental period, and sorghum‐based diet (SBD) consisted of 30% (d 1 to 14) or 60.55% (d 15 to 28) sorghum grain in partial or total replacement of corn grain. Both diets were formulated to contain the same amount of digestible energy and indispensable amino acids. The results demonstrated no differences in growth performance or apparent digestibility of gross energy between treatments over the whole period. However, the substitution of corn by sorghum reduced (p < 0.05) or tended to reduce (p = 0.09) apparent digestibility of crude protein associated with an increased faecal nitrogen excretion per weight gain (p < 0.05). Pigs fed SBD had higher contents of urea nitrogen, total triglyceride and insulin in serum than those fed CBD (p < 0.05). Visceral organ weights or antioxidant enzyme activities in serum or liver were not different between treatments. Compared with CBD, SBD increased or tended to increase amylase activity in jejunal mucosa (p < 0.05) or trypsin activity in duodenal mucosa (p = 0.08). Replacement of corn by the low‐tannin sorghum in diets did not influence the microbiota community based on alpha and beta diversity in caecal and colonal digesta. Overall, the home‐grown low‐tannin sorghum could be an alternative energy source in diets for pigs without adverse effects on growth performance.
Article
This study was conducted to determine whether sorghum as a potential substitute for corn in diets affects the activities of digestive enzymes and antioxidant enzymes in pigs. Sixty pigs (7.3 ± 1.3 kg initial weight) were allotted to 2 diets with 5 replicate pens per treatment according to sex and weight in a randomized complete block design. A corn-based diet (CBD) included 600 g/kg of corn grain during the overall experimental period, and a sorghum-based diet (SBD) consisted of 300 g/kg (day 1 to 14) or 566 g/kg (day 15 to 28) of sorghum grain at the partial or complete expense of corn in CBD. Both diets were formulated to contain the same amount of digestible energy and essential amino acids. No differences were observed in average daily gain or feed intake between treatments. However, substituting sorghum for corn in diets tended to reduce (P = 0.087) or reduced (P < 0.05) feed efficiency. Apparent digestibility of gross energy and crude protein was decreased (P < 0.05) in pigs fed the diets comprising sorghum, regardless of the substitute level, associated with the increased faecal excretion of energy and nitrogen. Serum urea nitrogen concentration in SBD was greater than that in CBD (P < 0.05). Pigs fed SBD had or tended to have lower superoxide dismutase and catalase activities and greater malondialdehyde content in serum (P < 0.05) or liver (P = 0.085) than those fed CBD. Amylase and lipase activities were decreased in duodenal mucosa (P < 0.05), and lipase (P < 0.05) and trypsin activities (P = 0.092) in jejunal mucosa were reduced and tended to be reduced in pigs fed SBD. Compared with CBD, SBD increased or tended to increase crypt depth in duodenum (P < 0.05) or jejunum (P = 0.094) coupled with the decrease in villus height/crypt depth in duodenum (P = 0.086) and ileum (P < 0.05). Collectively, sorghum as a dietary substitute for corn changed intestine morphology and reduced activities of digestive enzymes and antioxidant enzymes which may have been responsible for the reduced energy and nitrogen utilization and feed efficiency in pigs.
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In weaned piglet diets, about 20% of the total protein and amino acids (AA) comes from the inclusion of sorghum or yellow corn. Therefore, the fraction of digestible amino acids of these ingredients is important for diet formulations. The ileal-cannulated pig model is not used in younger pigs due to difficulties related to implanting T-cannula in the distal ileum and possible subsequent problems. Thus, the present study evaluated the apparent digestibility of yellow corn and sorghum in piglets by total collection and digestibility of amino acids by the slaughter method. Fifty weaned piglets, at 21-d-old, were divided into two assays to determine effects of multi-carbohydrase (MC) preparation (700 U α-galactosidase, 2,200 U galactomannanase, 3,000 U xylanase, and 22,000 U β-glucanase per kg of diet) and phytase (Phy, 500 FTU per kg of diet) supplementation on the apparent total tract digestibility (ATTD) of dry matter (DM), crude protein (CP), ash, digestible (DE) and metabolizable (ME) energy and standardized ileal digestibility (SID) of AA in yellow corn (YC) and sorghum. Piglets were individually caged and ileal digesta were collected at slaughter (38 d old). A completely randomized experimental design with a 2 × 2 (0 and 200 MC; 0 and 50 Phy) factorial treatment arrangement. A basal diet (BD) was used for an additional group of 5 piglets. Corn BD was used for ATTD determination, and corn-starch BD2 containing 50 g/kg casein was used to estimate AA losses. YC and sorghum replaced 30% of BD. There was no interaction effect (P > 0.05) of the enzyme combination on the apparent digestibility of nutrients, energy, AID and SID of AA of YC and sorghum. The apparent effect from the combination of the enzymes, in fact, was due to the presence of MC, as shown by its isolated inclusion. The supplementation of MC improved ATTD of sorghum DM (P = 0.041) regardless of Phy supplementation. Piglets weaned at 21 days of age showed lower capacity to use energy and nutrients in the subsequent three weeks, compared to NRC and the Brazilian tables. Isolated or combined, the enzymes did not increase the digestibility coefficients of YC, however, MC increased DM digestibility of sorghum.
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Plant condensed tannins (proanthocyanidins, PAs) have both positive and negative effects on feed digestibility and animal performance, depending both on the quantity and biological activity of the tannins that are present. In this review, the chemistry and analysis of condensed tannins (PAs) are examined. Our first focus is on the complexity of the structures of condensed tannins and our second emphasis is on the analytical methods used to evaluate tannins. The section on methods is subdivided into a discussion of methods to determine the amount of condensed tannins or total phenolics in a sample and a section on methods to measure biological activity. The methods to measure reactivity include assays involving protein binding and precipitation, as well as those that involve enzymatic and microbial inhibition. The last section of the paper discusses structure–activity relationships and provides information on how to select appropriate assays for measurement of the quantity and activity of condensed tannins.
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Forty-three batches of cereals (10 of wheat, 10 of triticale, 5 of barley, 5 of rye, 7 of maize and 6 of sorghum) were analysed and studied for their nitrogen (N) and amino acid (AA) ileal digestibility. Each batch was tested on four castrated male pigs, weighing between 30 and 90 kg, and fitted with an end-to-end ilco-rectal anastomosis. Ileal true digestibility (TD) of AA was calculated by correcting ileal apparent digestibility (AD) for basal endogenous AA losses, measured by means of a protein-free diet. Ileal real digestibility (RD) of AA was calculated by correcting AD for total endogenous AA losses, estimated by a multiple regression model. TD of N and most AA decreased (P<0.001) from wheat, triticale and maize, to barley and sorghum and to rye (90.3, 88.7, 89.9, 85.4, 83.7 and 80.1%, respectively, for the sum of all AA). Estimates of endogenous N losses decreased (P < 0.001) from triticale, sorghum and wheat, to maize, barley and rye (on average 3.10, 2.93,2.63,2.43,2.27 and 2.16 g N-kg-1 DM ingested, respectively). Barley excluded, there was a trend toward increasing endogenous AA losses with increasing dietary acid detergent fibre (ADF) concentration (r = 0.57, P < 0.001). Barley caused low endogenous N losses relative to its ADF concentration.