Animal Feed Science and Technology 203 (2015) 88–94

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Animal Feed Science and Technology

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Short communication

In vitro ruminal fermentation of ground and dry-rolled barley

grain differing in starch content
U.Y. Anele a , B. Refat a,c , M.-L. Swift b , Y.L. Zhao a,d , C. Doublier a , T.A. McAllister a ,
W.Z. Yang a,∗
a
Agriculture and Agri-Food Canada, Lethbridge, Alberta T1J 4B1, Canada
b
Alberta Agriculture & Rural Development, Lethbridge, Alberta T1J 4B1, Canada
c
Department of Animal Production, Faculty of Agriculture, Zagazig University, Zagazig, Egypt
d
College of Animal Science, Inner Mongolia Agricultural University, Hohhot, Inner Mongolia 010018, China

a r t i c l e i n f o a b s t r a c t

Article history: In vitro ruminal fermentation of ground and dry-rolled barley samples differing in starch
Received 19 November 2014 content was evaluated using a batch culture technique. The study was arranged in a 2 (low
Received in revised form 4 January 2015 and high starch) × 2 (ground and dry-rolled) factorial design. Gas production (GP), short
Accepted 11 February 2015
chain fatty acids (SCFA), dry matter (DM) and starch disappearance were estimated at 3, 6,
12 and 24 h of incubation using rumen fluid from 3 ruminally fistulated beef cattle. Rate
Keywords: of GP was greater (P<0.05) in both high starch and ground (2 mm) barley samples. Kinetics
Barley grain
of DM and starch disappearance were calculated using the equation of a + b (1 − e−c(t−L) ).
In vitro batch culture
Starch content × processing interactions were noted for the b fraction and rate of DM dis-
Starch content
Grain processing appearance. Consistently, both high starch and ground barley samples had greater (P<0.05)
a and b fractions, rate constant of disappearance of b fraction and effective degradability
versus low starch and dry-rolled samples. Expectedly, molar proportions of individual and
total SCFA were greater (P<0.05) in the ground barley samples at all incubation periods.
Overall, starch content (high versus low) had significant effect on the rate of GP and con-
stant rate of DM disappearance but no effect on the constant rate of starch disappearance.

Crown Copyright © 2015 Published by Elsevier B.V. All rights reserved.

1. Introduction

Barley grains from different sources are diverse in their chemical composition and estimated metabolizable energy
contents due to geographical, environmental and genetic variations as well as their interactions (Dehghan-banadaky et al.,
2007). Differences in digestibility explain most of the variation in energy content of a feed. This inherent variability in barley
chemical composition leads to differences in animal performance (Yang et al., 2013). Digestibility of whole barley grain is
limited by its fibrous hull and intact pericarp (Beauchemin et al., 1994) so processing is required to make the starch accessible

Abbreviations: ADF, acid detergent fiber; CP, crude protein; CV, coefficient of variation; DM, dry matter; DMD, dry matter disappearance; ED, effective
degradability; GP, gas production; GV, gas volume; aNDF, neutral detergent fiber assayed with a heat stable amylase and expressed inclusive of residual
ash; PI, processing index; SCFA, short-chain fatty acids.
∗ Corresponding author at: Agriculture and Agri-Food Canada, Box 3000, Lethbridge, Alberta T1J 4B1, Canada. Tel.: +1 403 317 3427;
fax: +1 403 382 3156.
E-mail address: wenzhu.yang@agr.gc.ca (W.Z. Yang).

http://dx.doi.org/10.1016/j.anifeedsci.2015.02.006
0377-8401/Crown Copyright © 2015 Published by Elsevier B.V. All rights reserved.
 U.Y. Anele et al. / Animal Feed Science and Technology 203 (2015) 88–94 89

to microbes in the rumen. Compared with intact barley grain, processing changes both its rate and extent of degradation in
the rumen. Because of rapid digestion of barley starch in the rumen, we hypothesized that starch content of barley grain can
influence both rates of gas production (GP) and dry matter (DM) disappearance (DMD). The objectives of this study were
to determine the effects of barley grains differing in starch content and processing (ground versus dry-rolled) on kinetics of
GP, DM and starch digestibility and concentrations of short-chain fatty acids (SCFA) using a batch culture technique.

2. Materials and methods

2.1. Barley sample collection and processing

Barley samples were collected from 10 different locations in Southern Alberta. One hundred and seventy samples were
collected monthly from November 2012 to March 2014. All the samples were analyzed and ranked according to their starch
content and the lowest 5 samples (ranging from 539 to 590 g/kg DM and hereafter designated low starch) and highest 5
samples (ranging from 651 to 671 g/kg DM and hereafter high starch) were selected. Subsequently, these 10 samples were
divided into 2 equal parts: one part was ground through a 2 mm screen (Wiley mill; Arthur H. Thomas Company, Philadelphia,
PA, USA), and the other part was dry-rolled using a laboratory scale roller (Model R250.6, Kal Rob Machining, Picture Butte,
AB, Canada) with rollers set at different distances in order to achieve an extent of processing expressed as processing index
(PI) 0.75 ± 0.03 for each samples. Processing index was calculated as the bulk density of barley grain after rolling divided by
the bulk density before rolling. Overall, 20 samples were used for the in vitro batch culture study.

2.2. In vitro incubations

Approximately 3 g of ground or dry-rolled barley was weighed in duplicate into 500 ml Ankom gas production module
(RF1; a computerized system with automated pressure transducers, Ankom Technology, Macedon, NY, USA). Ruminal fluid
was collected 2 h after feeding (0900 h) from three ruminally fistulated beef cattle (650 kg body weight) fed ad libitum a diet
consisting of whole crop barley silage (700 g/kg), dry-rolled barley grain (270 g/kg), and vitamin and mineral supplement
(30 g/kg) on a DM basis. All animal procedures were in accordance with the guidelines of the Canadian Council on Animal
Care (2009). Sampling and handling of rumen fluid were as described previously (Anele et al., 2014). The pH of ruminal fluid
was measured immediately (B20PI, SympHony Benchtop Meters; VWR, Edmonton, AB, Canada) and ranged from 5.78 to
6.12 throughout the study. Each Ankom gas production module received 270 ml of McDougall’s buffer and 90 ml of strained
ruminal fluid (3:1 ratio), after which each module was flushed with oxygen free CO2 and sealed. Modules were incubated
in an oscillating shaker at 39 ◦ C with an oscillation speed of 125 rpm for 3, 6, 12 and 24 h. The whole process was repeated
on a different date (second run) to have four analytical replicates (i.e. two per run). In addition, 2 blanks containing 360 ml
of medium only were included for each incubation time.
Gas data obtained were fitted to exponential model (Ørskov and McDonald, 1979) as:

y = B(1 − exp − c × [t − lag]),

where ‘y’ is the cumulative volume of gas produced at time ‘t’ (h), ‘B’ is the asymptotic gas volume, ‘c’ is the rate constant
and ‘lag’ is the time (h) between inoculation and commencement of GP. Initial GP rate (Absg ) was calculated as the product
of asymptotic cumulative gas volume and rate of fermentation (Larbi et al., 1996). After incubation, bottles were placed
in ice to stop fermentation. Undigested substrate was determined by high-speed centrifugation (20,000 × g) of incubation
residues at 4 ◦ C for 30 min. Blanks were also centrifuged and pellet weighed and used to correct for residues from the ruminal
inoculum. In vitro DM disappearance coefficient was calculated as:

[Substrate incubated − (substrate pellet − blank pellet)]

.
substrate incubated
The same process was used to estimate starch disappearance. Kinetics of DM and starch disappearance were calculated
using the equation of McDonald (1981):

y = a + b(1 − e−c(t−L) ) for t > L,

where y, disappearance at time t; a, an intercept representing the proportion of DM/starch solubilized at initiation of incuba-
tion (soluble fraction); b, the fraction of DM/starch insoluble but degradable in the rumen; c, a rate constant of disappearance
of fraction b; t, time of incubation and L, lag phase. The non-linear parameters a, b, c and L were estimated by an iterative least
squares procedure (SAS, 2002). The effective degradability (ED) of DM and starch was calculated using the following equation
(McDonald, 1981) with the modification of Wulf and Südekum (2005), which assumes no degradation occurs during the lag
phase:
 bc 
ED = a + × e−kL ,
c+k
90 U.Y. Anele et al. / Animal Feed Science and Technology 203 (2015) 88–94

Table 1
Chemical composition of the 5 low and 5 high starch content of barley grain samples.

Chemical composition Mean SDa Minimum Maximum CVb

Low starch
Dry matter (g/kg) 883 10.6 867 895 1.2
Crude protein (g/kg DM) 147 12.8 132 172 8.7
Acid detergent fiber (g/kg DM) 73 15.4 54 92 21.1
Neutral detergent fiber (g/kg DM) 237 32.8 206 292 13.8
Starch (g/kg DM) 570 17.5 539 590 3.1

High starch
Dry matter (g/kg) 879 9.3 868 895 1.1
Crude protein (g/kg DM) 135 11.2 123 158 8.3
Acid detergent fiber (g/kg DM) 59 4.8 49 66 8.1
Neutral detergent fiber (g/kg DM) 192 16.9 167 221 8.8
Starch (g/kg DM) 659 7.3 651 671 1.1
a
SD, standard deviation.
b
CV, coefficient of variation.

where k is the estimated rate of outflow from the rumen, and a, b, c and L are the same parameters described previously. The
ED of DM and starch were estimated assuming a rumen solid outflow rate of 0.05/h, which are representative of medium
passage rate.
Total short-chain fatty acids (SCFA) and lactic acid were measured upon completion of 3, 6, 12 and 24 h of incubation
after measuring gas pressure and pH (Orion model 260A, Fisher Scientific, Toronto, ON, Canada). Sampling and estimation
of SCFA and lactic acid were as described previously (Anele et al., 2014).

2.3. Chemical analyses

Barley samples were analyzed for DM, crude protein (CP), neutral detergent fiber (aNDF) and acid detergent fiber (ADF)
as described previously (Anele et al., 2014). Starch was determined by enzymatic hydrolysis of ␣-linked glucose polymers
as described by Rode et al. (1999).

2.4. Statistical analyses

Chemical composition of the 5 high and 5 low starch content of barley grain samples were averaged (n = 5) and data
were summarized by descriptive statistics of SAS (2002) to include the mean, standard deviation, minimum and maximum
values, coefficient of variation (CV). Data from the in vitro study were subjected to analysis of variance using the mixed
model procedure of SAS (2002) in a 2 (starch; low and high) × 2 (processing; ground and dry-rolled) factorial design. The
model included fixed effects of starch content, processing and its interaction, random effect of barley samples. Differences
among barley sample means with P<0.05 were accepted as statistically significant. When the interaction between factors
was significant, the SLICE option in the LSMEANS statement was used to determine differences among the factors.

3. Results

Apart from the starch content, all the chemical constituents were slightly greater in low versus high starch samples
(Table 1). Starch content × processing interaction was not significant (P>0.05) for in vitro GP kinetics of the barley samples
(Table 2). Rate of GP was greater (P<0.05) in high starch and ground barley samples. No difference was observed in the media
pH among treatments (data not shown).
Starch content × processing interactions were noted for the b fraction (P<0.03) and rate of DMD (P<0.01; Table 3). Con-
sistently, both high starch and ground barley samples had greater (P<0.05) a and b fractions, rate constant of disappearance
of b fraction and ED versus low starch and dry-rolled samples. Similar to the trend observed in DMD, high starch and ground
samples had greater (P<0.05) ED of starch than low starch and dry rolled samples (Table 3). Rate constant of disappearance
of b fraction of starch was greater (P<0.001) for ground versus dry rolled samples.
Lactic acid data were excluded because their concentrations were too low (<0.01 mmol/L). There was no effect (P>0.05)
of starch content on SCFA concentration after 3, 6 and 24 h of incubation although high starch samples had numerically
greater total SCFA concentration versus low starch samples (Table 4). High starch samples resulted in greater (P<0.05) molar
proportions and total SCFA versus low starch samples after 12 h of incubation. Expectedly, molar proportions of individual
and total SCFA were greater (P<0.05) in the ground samples at all time periods.

4. Discussion

The present study is one of the several in vitro and in situ studies from an on-going research project in order to develop
an accurate and concise in vitro method to predict feed value of barley grains varying in physical and chemical composition,
 U.Y. Anele et al. / Animal Feed Science and Technology 203 (2015) 88–94 91

Table 2
Effects of barley grain differing in starch content (low versus high) and processing (ground versus dry-rolled) on in vitro gas production kinetics.

GVA c (/h) Lag Absg

Starch level
Low 213 0.161b 2.33 34.6b
High 219 0.183a 2.17 40.0a
SEM 2.2 0.0041 0.182 0.88

Processing
Ground 214 0.206a 2.17 44.9a
Dry-rolled 219 0.139b 2.34 29.7b
SEM 2.2 0.0041 0.181 0.88

Significance
Starch level 0.102 0.002 0.553 <0.001
Processing 0.120 <0.001 0.511 <0.001
Starch level × processing 0.969 0.197 0.894 0.316

Means within a column, and within a category, with different superscripts differ (P<0.05).
A
GV, asymptotic cumulative gas volume (ml/g DM); c, rate of fermentation (/h); Lag, lag time (h); Absg , absolute initial gas production during first hour
(ml/g DM).

Table 3
Effects of barley grain differing in starch content (low versus high) and processing (ground versus dry-rolled) on nonlinear estimates and effective degrad-
ability coefficients of dry matter and starch.

aA b c (/h) Lag EDB

Dry matter
Starch level
Low 0.071b 0.56b 0.116b 2.38 0.43b
High 0.089a 0.63a 0.135a 2.25 0.47a
SEM 0.0045 0.013 0.0062 0.247 0.010
Processing
Ground 0.11a 0.60 0.138a 1.74b 0.51a
Dry-rolled 0.05b 0.59 0.112b 2.89a 0.39b
SEM 0.0044 0.013 0.0062 0.247 0.010
Starch level Processing
Low Ground 0.099 0.58a 0.132a 1.94 0.48
High Ground 0.119 0.62a 0.145a 1.53 0.54
Low Dry-rolled 0.042 0.53b 0.137a 2.81 0.37
High Dry-rolled 0.059 0.64a 0.087b 2.95 0.40
Significance
Starch level 0.004 <0.001 0.037 0.706 0.002
Processing <0.001 0.481 0.005 0.002 <0.001
Starch level × processing 0.853 0.027 <0.001 0.441 0.469

Starch
Starch level
Low 0.09 0.53 0.076 2.93a 0.33b
High 0.09 0.56 0.074 0.50b 0.41a
SEM 0.004 0.019 0.0042 0.193 0.003
Processing
Ground 0.08 0.54 0.089a 1.43b 0.40a
Dry-rolled 0.09 0.55 0.061b 2.01a 0.33b
SEM 0.004 0.019 0.0042 0.193 0.003
Starch level Processing
Low Ground 0.08 0.51 0.095 2.26a 0.36
High Ground 0.09 0.58 0.084 0.60b 0.43
Low Dry-rolled 0.09 0.56 0.057 3.61a 0.30
High Dry-rolled 0.08 0.55 0.064 0.40b 0.37
Significance
Starch level 0.789 0.223 0.761 <0.001 <0.001
Processing 0.501 0.747 <0.001 0.039 <0.001
Starch level × processing 0.131 0.117 0.146 0.006 0.532

Means within a column, and within a category, with different superscripts differ (P<0.05).
A
a, The portion of DM/starch solubilized at initiation of incubation; b, the fraction of DM/starch insoluble but degradable in the rumen; c, the constant
rate of disappearance of fraction b (/h); Lag, lag phase (hours) before the commencement of degradation of fraction b.
B
ED, effective degradability at a ruminal passage rate of 0.05/h.
92 U.Y. Anele et al. / Animal Feed Science and Technology 203 (2015) 88–94

Table 4
Effects of barley grain differing in starch content (low versus high) and processing (ground versus dry-rolled) on short-chain fatty acids (SCFA, mmol/l)
concentration in the fermentation media at different time periods.

C2 A C3 Iso C4 C4 Iso C5 C5 C2 :C3 TSCFA

3h
Starch level
Low 7.81 2.92 0.09 1.85 0.36 0.31 2.84 13.3
High 8.19 3.13 0.09 1.92 0.37 0.32 2.71 14.1
SEM 0.152 0.105 0.003 0.038 0.005 0.008 0.087 0.26
Processing
Ground 9.21a 3.74a 0.09 2.06a 0.37 0.33a 2.53b 15.8a
Dry-rolled 6.78b 2.32b 0.08 1.71b 0.36 0.30b 3.03a 11.6b
SEM 0.152 0.105 0.003 0.038 0.005 0.008 0.087 0.26

6h
Starch level
Low 13.9 4.47 0.19 3.81 0.70 0.57 3.25 23.9
High 14.9 4.98 0.21 4.04 0.75 0.62 3.12 25.7
SEM 0.60 0.264 0.010 0.115 0.031 0.033 0.054 1.05
Processing
Ground 15.9a 5.52a 0.19 4.19a 0.71 0.59 2.99 27.3a
Dry-rolled 13.0b 3.94b 0.21 3.66b 0.73 0.60 3.37 22.3b
SEM 0.60 0.264 0.009 0.115 0.030 0.033 0.054 1.05

12 h
Starch level
Low 26.5b 10.0b 1.03b 8.81b 2.17 2.33 2.64 51.3b
High 28.5a 10.5a 1.06a 9.42a 2.20 2.37 2.71 54.6a
SEM 0.46 0.13 0.011 0.097 0.030 0.024 0.039 0.65
Processing
Ground 29.8a 11.1a 1.09a 9.83a 2.31a 2.47a 2.69 57.2a
Dry-rolled 25.2b 9.5b 0.99b 8.41b 2.07b 2.22b 2.66 48.7b
SEM 0.46 0.13 0.011 0.097 0.030 0.024 0.039 0.65

24 h
Starch level
Low 38.6 12.9 1.33 12.3 2.63 2.86 2.95 71.3
High 39.4 13.2 1.29 12.5 2.58 2.79 2.99 72.4
SEM 0.65 0.23 0.010 0.12 0.019 0.029 0.021 0.91
Processing
Ground 40.5a 13.2 1.33 12.2 2.63a 2.90a 3.08a 73.5a
Dry-rolled 37.4b 12.9 1.30 12.6 2.57b 2.75b 2.90b 70.2b
SEM 0.65 0.23 0.010 0.12 0.019 0.029 0.021 0.91

Means within a column, and within a category, with different superscripts differ (P<0.05).
A
C2 , acetic; C3 , propionic; IsoC4 , isobutyric; C4 , butyric; IsoC5 , isovaleric; C5 , valeric; SCFA, total short-chain fatty acids; C2:C3, acetic to propionic ratio.

and grain processing. Chemical composition results indicated that digestibility value of high starch samples would be better
than low starch samples due to a combination of higher starch and lower ADF and NDF contents. Variation in the rate of
in vitro GP of low versus high barley samples was likely due to variations in their chemical components. This observation
may be due, at least in part, to greater starch and less fiber contents of the high starch samples which resulted in greater
rate of in vitro GP for the high starch samples. Lack of difference in cumulative GP by either starch content or processing
indicates that the microflora had sufficient nutrients for the production of SCFA and microbial mass.
Greater potential degradation (a + b) and rate of DMD estimates for high versus low starch samples were consistent with
expectation as studies (Owens, 2005; Yang et al., 2013) have shown that greater starch content is associated with higher DM
digestibility compared with greater fiber content which depresses digestibility. Effective degradability of starch followed
similar trends as noted for the DM with greater values for high starch and ground barley samples. We can only speculate on
lower lag time (for starch digestion) noted for high starch samples because these samples had less ‘a’ fraction (the portion
of starch solubilized at initiation of incubation) than the other samples.
Contrary to expectation, no difference was observed in molar proportion and total SCFA concentration of high versus low
starch samples after 24 h of incubation. Bearing in mind that high starch samples had greater potential degradation and
ED coefficients than low starch samples, we expected more SCFA from high starch samples. A biological explanation could
be that the high starch samples partitioned more nutrients into microbial mass versus SCFA. Although this explanation is
consistent with Hungate (1966), who reported that microbial mass and SCFA are inversely related, we can only speculate
because we did not measure microbial mass in the present study. The absence of a difference in SCFA concentration however
is consistent with an absence in total GP. Blümmel et al. (1997) reported that in addition to CO2 and CH4 produced as a result
of fermentation, CO2 is also produced upon buffering of SCFA generated (i.e. indirect GP) and that molar production of CO2
 U.Y. Anele et al. / Animal Feed Science and Technology 203 (2015) 88–94 93

equals molar SCFA production. Although there were differences in molar proportion and total SCFA concentration of high
versus low starch samples at 12 h of incubation, the observed differences disappeared after 24 h. This may indicate that a
longer lag time may have been responsible as evident in the longer lag time noted for low versus high starch samples in the
present study.
Numerically greater in vitro cumulative gas noted for dry rolled versus ground samples was contrary to expectation.
We expected more cumulative gas for the ground samples because grinding greatly increases the surface area available for
microbial attachment. Similarly, Galyean et al. (1981) reported that starch degradation in the rumen varies inversely with
particle size of the grain.
Expectedly, ED value of DM was greater in ground than rolled samples. Previous studies (Yang et al., 2000; Wang et al.,
2003) have shown that extensively processing of barley grain improves nutrient digestibility, although adequate care should
be taken to avoid increase production of fines which could adversely affect DM intake. Contrary to DMD, processing did not
affect the extent of starch digestion. More processed (ground) barley samples increased rate of starch digestion and this
is consistent to the study of Zinn (1993), who reported that although the rate of starch digestion was increased by more
extensive processing, the extent of ruminal starch digestion was not. As stated earlier, we can only speculate on lower lag
time (for starch digestion) noted for more processed samples because these samples (just like high starch samples) had less
‘a’ fraction (the portion of starch solubilized at initiation of incubation) than less processed samples.
More processed barley samples produced more SCFA versus less processed samples which is consistent with a previous
study of Yang et al. (2000), who reported a linear increase in total SCFA concentration with reducing PI. We observed a
difference of approximately 0.05 for the pH of ground (2 mm) versus dry-rolled (PI of 0.75) in the present study. Yang et al.
(2001) reported that feeding extensively processed grain with rapid and complete ruminal degradation can reduce ruminal
pH creating an unfavorable environment in the rumen for synthesis of microbial protein and digestion of fiber. Because of
the 3:1 (buffer:rumen fluid) used in the in vitro study, we did not observe significant decline or differences in the pH of the
samples.

5. Conclusion

Overall, starch content (high versus low) had significant effect on the rate of GP and constant rate of DMD but no effect on
the constant rate of starch disappearance. Effective degradability of DM and starch were also affected by the starch content.
Results showed that processing (ground versus dry-rolled) increased the extent and rate of disappearance of DM and starch.

Conflict of interest statement

The authors declare that no conflict of interest, financial or other, exists.

Acknowledgements

This study was financially supported by the Alberta Crop Industry Development Fund (grant no. #2012F070R). Alastair
Furtado and Darrell Vedres are acknowledged for their assistance with sample collection and laboratory analyses.

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