Directory UMM :Data Elmu:jurnal:A:Animal Feed Science and Technology:Vol81.Issue1-2.Sep1999:
Animal Feed Science and Technology
81 (1999) 133±149
Effect of different levels of berseem (Trifolium
alexdrinum) supplementation of wheat straw on some
physical factors regulating intake and digestion
Asit Dasa, G.P. Singhb,*
a
ICAR Research Complex, Sikkim Centre, Todong 737102 (Sikkim), India
b
National Research Centre on Carnel, Jorbeer, Bikaner-334001, India
Received 24 April 1998; received in revised form 23 September 1998; accepted 17 March 1999
Abstract
Four ruminally fistulated cattle of about 318 16 kg body weight were randomly distributed into
four groups in an experiment based on 4 4 Latin square design. The four dietary treatments were:
wheat straw ad lib without any supplement (I), supplemented with berseem at 15 (II), 30 (III), and
45 (IV) percent level. DMI was 3.95, 5.85, 7.28 and 7.08 kg; digestibility of DM was 39.85, 45.61,
52.97 and 47.52%; rumen content was 55.60, 68.45, 74.18 and 75.93 kg, or 18.09, 22.01, 23.37 and
23.22% of body weight; rumen fluid content was 48.62, 60.39, 65.20 and 68.01 kg; rumen pool size
of DM was 6.02, 7.61, 7.92 and 7.66 kg; rumen pool size of particles larger than 1.18 mm (LP) was
2.38, 2.78, 2.80 and 2.55 kg; rumen pool size of particles smaller than 1.15 mm (SP) was 3.64, 4.91,
5.12 and 5.05 kg in Groups I, II, III and IV, respectively. Intake and digestibility increased
significantly (p < 0.01), with increased level of berseem up to 30%, beyond which no further
improvement was observed. Total rumen contents, rumen fluid content and pool sizes of DM
increased significantly (p < 0.5) with berseem supplementation, level of berseem had no significant
effect. Pool of LP and SP was similar in all the groups. Maximum rumen content was observed at 9,
9, 6 and 3 h post feeding in Groups I, II, III and IV, respectively. Rate of clearance of LP was 3.71,
6.71, 7.27 and 8.75; SP was 0.46, 0.83, 1.20 and 1.35; and communition rate was 3.45, 5.79, 5.92
and 7.37% hÿ1 in Groups I, II, III and IV, respectively. Rate of clearance of LP and rate of
communition were significantly higher (p < 0.01) in berseem supplemented groups. Rate of passage
(Kp) was 1.72, 1.88, 2.33 and 2.56% per hour. Kp increased significantly (p < 0.01) with increased
level of berseem. From the results it is evident that rumen pool size of indigestible component,
rather than rumen content, determines the intake. As Kd was more than Kp, there was an overall
*
Corresponding author. Tel.: +91-151-523187; fax: +91-151-522183
E-mail address: [email protected] (G.P. Singh)
0377-8401/99/$ ± see front matter # 1999 Elsevier Science B.V. All rights reserved.
PII: S 0 3 7 7 - 8 4 0 1 ( 9 9 ) 0 0 0 6 0 - 7
134
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
improvement of digestibility by 14, 33 and 19% in Groups II, III and IV, respectively, compared
with Group I. # 1999 Elsevier Science B.V. All rights reserved.
Keywords: Level of berseem; Berseem supplementation; Wheat straw; Cattle; Intake; Regulation; Digestion
1. Introduction
Wheat straw forms a major source of ruminant feed in India. Unfortunately, wheat
straw is characterised by low intake and digestibility. Supplementation of poor quality
roughage, including wheat straw, with green forages has been shown to increase intake
(Woodward and Reed, 1995; Reed et al., 1990) or digestibility (Silva and Orskov, 1988;
Ash, 1990; Bonsi et al., 1995) or both (Leng, 1990; Bird et al., 1994). However, type of
basal diet (Mosi and Butterworth, 1985; Eliott et al., 1984), degradability characteristics
of the supplement (Bates et al., 1988), level of supplement (Bonsi et al., 1994) as well as
interactions between nutrients in feed ingredients (Brown et al., 1991) are important
determinants of response to supplementation. Hence, specific study is required to know
the effect of a particular supplement on intake and digestibility of the basal diet
concerned.
Berseem (Trifolium alexdrinum) is a green leguminous fodder grown during winter.
Berseem contains 15±20% CP and 4.1 kcal/g GE (Chauhan et al., 1980). It is highly
palatable, fairly digestible and on sole feeding can support growth rate of 550 g/d
(Chauhan et al., 1992) and milk production up to 10 kg/d. Berseem supplementation to
wheat straw has been shown to increase in sacco dry matter degradability of wheat straw
(Reddy et al., 1991). In practice, farmers in India feed wheat straw and berseem mixed
together without considering the ratio. However, detailed information regarding the effect
of different levels of berseem supplementation on digestion and intake of straw-based diet
is not available. At the same time, prediction of likely effect on intake and digestion will
be difficult without a better understanding of the processes, namely, rumenfil (Bosch
et al., 1993), particle dynamics (Poppi et al., 1980) and passage rate (Oosting et al.,
1993), which regulate intake. Hence, this experiment was undertaken to find out the level
of berseem at which intake and digestion is maximum, with a better understanding of the
physical factors regulating them.
2. Materials and methods
2.1. Animals and design
Four ruminally fistulated crossbred (Shaiwal Holstein friesan) steers of about
318 16 kg body weight were kept in stalls, with individual feeding arrangements. The
experiment was set up as a 4 4 Latin Square design, with each experimental period
lasting 3 weeks. Adaptation period before each experimental period had a duration of 3
weeks. The time schedule of each experimental period was:
Activity
Day
Adaptation
Rumen liquor sampling
Intake and digestion
Dacron bag analysis of feed sample
Rumen evacuation
Rumen evacuation on restricted feeding
1±14
15
16±22
21±25
26±31
32±35
Four dietary treatments were wheat straw alone (I), wheat straw supplemented with 15
(II), 30 (III), and 45% (IV) level of berseem. Supplement was offered once daily at 9 a.m.
Clean and fresh drinking water was provided ad lib. The amount of supplement to be
offered was determined from the record of previous day intake, to keep the level of
berseem to the desired experimental level as far as possible.
2.2. Chemical composition of feed
Samples of wheat straw and berseem were taken daily during the last 21 days of each
period. They were dried at 50±608C and analysed for dry matter (DM), ash, neutral
detergent fibre (NDF), acid detergent fibre (ADF) and total nitrogen (N) content.
2.3. Digestion trial
During each period, a digestion trial of six-day collection (Days 17±22) was conducted
to determine the intake and digestibility of diets.
2.4. Rumen evacuation
Rumen evacuations were done during Days 26±31 in each period. Each steer's rumen
was evacuated once daily over 6 days, either at 12:00, 15:00, 18:00, 21:00, 3:00 or 9:00
136
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
2.4.1. Rumen evacuation of animals of restricted feeding
During last 4 days of each experimental period, rumen evacuations were done on
animals receiving restricted amount of feed. The animals were allowed to eat up to 12:00
hours. Each steer's rumen was evacuated once daily over 4 days, either on 12:00, 18:00 or
24:00 hours. There was a minimum interval of 30 h between emptying from any
individual steer to overcome the effect of emptying on the repeatability of measurements.
Total rumen contents were removed by hands, weighed, sampled and the remainder
returned to the rumen. Two samples of 500 g were taken: the first one for separation into
large (>1.18 mm) and small (>0.041 mm, 1.18 mmb
Pool < 1.18 mm*
Total*
Pool NDF (kg)b
2.38
3.64 a
6.02 a
4.74
2.78
4.91 b
7.61 b
5.63
2.80
5.12 b
7.92 b
5.93
2.55
5.05 b
7.66 b
5.42
2.08
0.23
0.37
0.45
Indigestible components
DM (kg)b
NDF (kg)b
3.66
2.91
4.12
3.22
4.00
3.04
3.98
3.11
0.23
0.19
a
b
Values followed by different letters in a row differ significantly.
Non-significant; **p < 0.01.
Table 4
Diurnal variation of rumen pool sizes in animals fed different level of berseem in dieta
I
Total rumen contents (kg) h post feeding
3**
55.0 a
59.5 a
6**
9*
61.0 a
12*
56.5 a
18b
55.5
51.5
24b
II
66.3
74.0
75.2
71.8
63.8
59.3
III
ab
b
b
b
75.5
81.8
76.0
74.0
71.5
62.5
IV
c
bc
b
b
83.0
83.0
80.2
75.8
71.5
61.5
SEM
c
c
b
b
3.05
2.49
3.61
3.17
4.38
2.58
DM (kg) h post feeding
3*
6.08 a
6.50 a
6**
9b
6.68
12*
5.95
18b
5.60
24b
5.25
7.40 b
7.98 b
8.38
8.08 b
7.38
6.55
8.40 c
8.60 b
8.05
7.78 b
7.55
7.13
7.98 c
8.30 b
7.58
7.48 b
7.33
6.95
0.40
0.30
0.35
0.39
0.45
0.44
NDF (kg) h post feeding
3b
4.83
5.10 a
6*
9b
5.23
12b
4.68
18b
4.40
4.13
24b
5.60
6.03 b
6.13
6.03
5.68
4.98
6.30
6.45 b
6.03
5.80
5.65
5.33
5.68
5.95 ab
5.58
5.28
5.13
4.90
0.34
0.25
0.32
0.32
0.39
0.38
a
b
Values followed by different letters in a row differ significantly.
Non-significant; **p < 0.01.
142
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
Table 5
Potentially degradable fraction (D), rate of degradation (Kd), isolated small and large particles, whole rumen
NDF pool, rate of clearance of large particle (LP) and small particle (SP), and rate of communition of large
particlea
I
II
III
IV
SEM
D fraction (%)
LP**
SP**
Rumen NDF pool*
25.20 a
36.48 a
33.68 a
40.02 b
41.94 c
43.96 b
48.11 c
45.90 d
45.03 b
49.44 d
39.08 b
40.83 ab
0.34
0.36
2.06
Kd (%/h)
LP**
SP**
Rumen NDF pool*
0.83 a
1.66 b
1.54 a
1.97 b
2.01 c
2.07 b
2.46 c
1.93 c
2.15 b
2.26 c
1.41 a
1.84 b
0.04
0.03
0.10
Rate of clearance (%/h)
LP**
SPb
Kc (%/h)**
3.71 a
0.46
3.45 a
6.71 b
0.83
5.79 b
7.27 b
1.20
5.92 b
8.75 c
1.35
7.37 c
0.36
0.21
0.37
a
b
Values followed by different letters in a row differ significantly.
Non-significant; **p < 0.01.
Groups IV, III, II and I, respectively. At around 18 h post feeding, the difference among
different groups was non-significant. Similarly, the rumen DM as well as NDF pool were
maximum at 6, 6, 9 and 9 h post feeding in Groups IV, III, II and I, respectively.
3.5. Particle dynamics
Degradation characteristics of rumen content, isolated LP, SP and rate of clearance of
SP, LP and rate of communition are presented in Table 5, and distribution of SP and LP is
illustrated in Fig. 2. Potential degradability of LP increased significantly (p < 0.01) with
increased level of berseem in the diet. Potential degradability of SP and whole rumen
NDF pool increased significantly (p < 0.01) with increased berseem level up to 30%.
However, at 45% level of supplementation, potential degradability decreased for both SP
and whole rumen NDF pool, difference being more marked in case of SP than whole
rumen NDF pool. Rate of degradation of LP increased with increased level of berseem up
to 30%; at 45% level of berseem, no further change was observed. Rate of degradation of
SP reduced significantly (p < 0.01) in Group IV in comparison to control, whereas 15 and
30% levels of berseem increased the rate of degradation.
3.6. Passage from rumen
Rate of clearance (Kcl), rate of passage (Kp) and mean retention time (MRT) of DM and
NDF are presented in Table 6. Kcl of DM increased significantly (p < 0.01) with increased
level of berseem up to 30%, beyond which no further change was observed. On the other
hand, MRT decreased and Kp increased significantly (p < 0.01) with increased level of
berseem in the diet.
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
143
Fig. 2. Diurnal variation of particle distribution in rumen as affected by the level of berseem in wheat straw diet.
144
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
Table 6
Clearance rate (Kcl), mean retention time (MRT), fractional passage rate (Kp) and liquid outflow rate (Kl)a
I
II
III
IV
SEM
Kcl (%/h)
DM**
NDF*
2.73 a
2.85 a
3.22 b
3.40 b
3.85 c
3.67 b
3.89 c
3.76 b
0.096
0.14
MRT (h)
IDM**
INDF**
58.33 a
55.71 a
53.24 b
50.74 b
42.98 c
45.72 c
40.20 c
42.09 d
1.01
0.97
Kp (%/h)
DM**
NDF**
Klb
1.72 a
1.81 a
6.60
1.88 b
1.96 b
7.08
2.33 b
2.19 c
7.18
2.56 b
2.38 d
7.85
0.047
0.042
0.58
Intake (kg)
DM**
IDM**
NDF**
INDF**
3.95 a
1.52 a
3.22 a
1.27
5.85 b
1.89 b
4.55 b
1.52
7.28 c
2.01 c
5.01 c
1.55
7.08 c
2.36 c
4.65 c
1.77
0.16
0.064
0.108
0.004
a
b
Values followed by different letters in a row differ significantly.
Non-significant; **p < 0.01.
3.7. Diurnal variation in passage rate
Within-day variation of passage rate is presented in Table 7. In this experiment,
passage rate was determined for the periods of 0±9 h, 9±18 h and 18±24 h and grossly
considered to be as the period of eating, rumination and idling, respectively, because the
activity concerned was more in the respective period. Maximum passage was observed
during eating in all the groups, except wheat straw alone fed group, where maximum
passage was observed during rumination. However, the absolute amount passaged during
rumination and idling was not different among the rations.
Table 7
Variation in passage rate of INDF from the potentially removable INDF pool of rumen during eating (PEAT),
rumination (PRUM) and idling (PID)a
INDF intake (%)
PEAT**
PRUMb
PIDb
Passage (g)
PEAT**
PRUMb
PIDNS
a
b
I
II
III
IV
35.96 a
46.32
17.72
46.18 b
33.26
20.56
59.93 c
26.73
12.97
67.74 c
23.24
9.33
462.5 a
571.0
234.5
713.3 b
568.0
317.0
930.3 b
402.0
213.0
Values followed by different letters in a row differ significantly.
Non-significant; **p < 0.01.
1197.3 c
407.0
165.0
SEM
3.08
8.73
5.55
72.5
66.0
80.5
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
145
4. Discussion
4.1. Intake and digestibility
Inclusion of berseem in the wheat straw based diet increased the total DM intake.
Increased DM intake as a result of green legume supplementation has been reported by
other workers (Mosi and Butterworth, 1985; McMeniman et al., 1988; Bird et al., 1994;
Bonsi et al., 1994; Woodward and Reed, 1995). Findings of this experiment are in line
with information available in the existing literature in general, but differ from the findings
of Odowongo and Mugerwa (1980) and Ash (1990). Such variations in response can
partly be attributed to the lower degradability of the supplement (Ash, 1990) and also to
the presence of toxicant (Odowongo and Mugerwa, 1980) in the supplement. Berseem, on
the other hand, is fairly degradable and contains no toxicant that is known to decrease
intake.
Berseem supplementation up to 30% level increased digestibility of DM, OM, NDF,
ADF, cellulose and hemicellulose; however, at 45% level of berseem supplementation,
digestibility decreased. On the other hand, CP digestibility increased linearly with
increased level of berseem in the diet, which is in line with the findings of Butterworth
and Diaz (1970) and Mosi and Butterworth (1985). Increased digestibility as a result of
forage supplementation has been reported by other workers (Juul-Nielson, 1981; Silva
and Orskov, 1988; Bird et al., 1994; Bonsi et al., 1994). Maximum digestibility observed
at 30% level of berseem supplementation suggests that 30% level of berseem in the diet
provided all the essential nutrients critical for optimal microbial activity, viz., rumen
NH3±N concentration (Eliott et al., 1984), peptides, essential minerals and vitamins
(Leng, 1990). In addition, berseem also provided fermentable cellulose and hemicellulose, which are known to promote fibre digestion (Silva and Orskov, 1988) by
ensuring greater degree of colonisation of fibrolytic bacteria (Cheng et al., 1990) and
fungi (Bauchop, 1979). Decrease in digestibility as a result of 45% berseem
supplementation, as observed in this experiment, could be due to decreased mean
retention time in the rumen, which means an increased rate of passage from the rumen
(Oosting, 1993) as the green forages are bulky in nature (Bonsi et al., 1994).
4.2. Rumen pool size
Total rumen content was 18% of the body weight in animals fed on wheat straw alone;
it increased up to 23% when supplemented with berseem. These values were higher than
the values of 17.5% in cattle fed on either grass silage based diet (Bosch et al., 1993) or
wheat straw based diet (Oosting, 1993), but lower than the values of 26% in early
lactating cows (Hartnell and Satter, 1979). These values are similar to those observed by
Chauhan et al. (1985) in buffalo fed on oat silage based diet supplemented with either
concentrate or cowpea hay. They reported that gut content is 24%. It appears that
ruminants fed on poor quality roughage, in the tropics, have higher rumen content than of
those in the temperate region. Berseem supplementation increased the total rumen
content and pool of DM and small particle. This indicates that intake is not limited by
rumen capacity. Similar conclusion has also been drawn by other workers (Bosch et al.,
146
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
1992; Oosting, 1993). Similar pool size of indigestible components (IDM and INDF) in
different groups having different levels of intake, as observed in this experiment, suggests
that rumen pool of indigestible component determines the intake (Mertens and Ely,
1979).
Generally, after attainment of maximum rumen fill, it declined and reached a minimum
value just before feeding. Maximum pool of DM was observed 6 h post feeding in Groups
III and IV, and 9 h post feeding in Groups I and II (Table 4). Pool of NDF followed the
same trend, but the difference between different groups was significant only at 6 h post
feeding. The pools of IDM and INDF gradually decreased after they attained the
maximum pool at 6, 6, 9 and 9 h post feeding in Groups IV, III, II and I, respectively.
Increased passage rate observed with increased level of berseem in the diet due to its
bulkiness (Bonsi et al., 1994) can partly be attributed to early attainment of maximum
rumen pool.
4.3. Particle dynamics
The results indicate that LP and SP are not degraded in the same fashion. Increase in
potential degradability and rate of degradation of LP with increased level of berseem in
the diet could be attributed to increase in readily fermentable fibre content (Silva and
Orskov, 1988). Potential degradability of whole rumen NDF, SP increased up to 30%
level of berseem, beyond which a negative associative effect was observed. It seems that
the decrease in potential degradability and rate of degradation at 45% level of berseem is
due to decrease in potential degradability (PD) and Kd of SP. The decreased PD and Kd of
SP could be due to the fact that at 45% level of berseem, the rumen, microbes had enough
substrate in rumen pool of host animal than the less preferably small particles, inside
nylon bag when isolated from the rumen which ready to passage out of the rumen and
substantial amount of digestion might had taken place before that. Rate of degradation of
LP, barring wheat straw, was higher than SP in all the groups. This, in general, is in
agreement that after attainment of critical particle size (1.18 mm) the feed particle do not
remain in rumen for long (Poppi et al., 1980) and substantial amount of microbial
digestion takes place before they attain that particle size.
4.4. Passage rate from rumen
Passage from rumen is limited only to particles smaller than 1.18 mm (Poppi et al.,
1980). In this experiment, proportion of small particle in different groups was similar, i.e.,
53%. The mean proportion of small particle was less than the value of 70% observed by
Oosting (1993). The difference can be explained on the basis of coarseness of the
supplement in our experiment. In the earlier experiment, sugar beet pulp having much
smaller particle size was used as supplement to wheat straw, whereas in this experiment
whole berseem without chaffing was used. The rate of communition increased with
increased level of berseem. Increased rate of communition can either lead to increase in
pool of SP or increased passage rate. In this experiment, no change in rumen pool size
beyond 15% level of berseem was observed. This indicates that the increased
communition can increase the pool of SP only to a limited extent, rest of the SP being
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
147
rapidly lost after their formation (Moseley and Jones, 1984). In this experiment, rate of
passage increased with increased level of berseem supplementation. Similar results have
also been reported by other workers (Moran et al., 1983; Bamualim et al., 1984; Bonsi
et al., 1994) when straw-based diets were supplemented with increased level of green
forages. Increased passage rate as a result of increased level of berseem supplementation
could probably be associated with the fact that berseem supplementation increased the
ratio of cellulose/hemicellulose in the diet. Particles with higher cellulose content
probably have a relatively higher functional specific gravity (Wales et al., 1990), which
may explain increased passage rate with increased level of berseem in the diet.
In this experiment, no change in rumen pool size beyond 15% level of berseem was
observed. Mean retention time decreased with increased level of berseem in the diet, as
passage rate was measured as reciprocal of the mean retention time, in this experiment.
Decreased mean retention time as a result of supplementation has also been reported by
other workers (Moran et al., 1983; Bonsi et al., 1994). Though the passage rate of
particulate matter increased with increased level of berseem supplementation, liquid
outflow rate was not significantly different between treatments. This finding is contrary to
that of Nsahlai (1991) and Bonsi et al. (1994), who reported that increased particulate
passage rate results in increased liquid outflow rate. Nevertheless, a different pattern of
particulate passage rate and liquid outflow rate has also been reported (Ulyatt et al.,
1984).
4.5. Diurnal variation in passage rate
Difference in passage observed in this experiment, as a result of berseem
supplementation, can be attributed to the increased rate of passage during eating. On
an average, 36, 46, 60 and 68% of indigestible NDF ingested was lost from the rumen
INDF pool, during eating, in Groups I, II, III and IV, respectively. The proportion of large
particles in the rumen increased up to 3 h post feeding; then it gradually decreased and
reached minimum just before feeding. Reduction in particle size was sharper in animals
fed on higher level of berseem, indicating higher rate of communition that resulted in
increased rate of passage as proportion of small particles in rumen was not significantly
different among the rations. The act of eating has been shown to be accompanied by a
marked increase in reticular and omasal contractions that might have elevated the rate of
flow of digesta from the rumen (Balch, 1958). It is observed in this experiment that
greater proportion of INDF is lost during eating with increased level of berseem in the
diet. This change might have occurred due to increased rumination efficiency (Moseley
and Jones, 1984). On an average, 462, 713, 930 and 1197 g of INDF was lost from the
rumen within first 9 h of feeding in Groups I, II, III and IV, respectively.
Berseem supplementation increased the rumen fill, indicating that rumen capacity is
not limiting intake. On an average, an increase of 23, 40 and 44% in DM intake was
recorded as a result of 15, 30 and 45% berseem supplementation. Increased intake was
accompanied by an increased rate of communition, which, in turn, increased the rate of
passage. However, the increase in disappearance was more than the rate of passage. On an
average, 19, 29 and 32% increase in clearance rate was observed in 15, 30 and 45% level
of berseem supplemented groups, respectively. Respective values for increase in Kp was
148
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
8, 21 and 31%. Overall digestibility increased by 14, 33 and 19% in groups fed 15, 30 and
45% level of berseem, respectively. It is concluded that increased level of berseem
supplementation up to 30% level to wheat straw based diet increases DMI and
digestibility of wheat straw based diet.
References
American Society of Agricultural Engineers, 1967. Methods of determining modulus of uniformity and modulus
of fineness of ground feed, Agricultural Engineers Yearbook, p. 301.
AOAC, 1984. Official Analytical of Method, 14th ed., Association of Official Analytical Chemists, Washington,
DC.
Ash, A.J., 1990. The effect of supplementation with leaves from the leguminous trees Sesbonia grandiflora,
Albizia chinensis, Albizia chinensis and Gliricidia sepium on the intake and digestibility of guinea grass hay
by goat. Anim. Feed. Sci. Technol. 28, 225±232.
Balch, C.C., 1958. Observation on the act of eating in cattle. Br. J. Nutr. 12, 330±345.
Bamualim, A., Weston, R.H., Hogan, J.P., Murray, R.M., 1984. The contribution of Leucaena leucocephala to
post ruminal digestible protein for sheep fed tropical pasture hay supplemented with urea and minerals. Proc.
Australia Soc. Anim. Prod. 15, 225±258.
Bates, D.B., Morantes, G.A., Moore, G.E., 1988. Nitrogen utilization of lombs fed limpo grass supplemented
with molasses-urea or legume protein supplements. Nutr. Rep. Intt. 38, 487±499.
Bauchop, T., 1979. Rumen anaerobic fungi of cattle and sheep. Appl. Environ. Microbiol. 38, 148±158.
Bird, S.H., Romulo, B., Leng, R.A., 1994. Effect of Lucerne supplementation and defaundation on feed intake,
digestibility, nitrogen retention and productivity of sheep fed straw based diets. Anim. Feed. Sci. Technol.
45, 119±129.
Bonsi, M.L.K., Osuji, P.O., Tuan, A.K., 1994. Graded level of Sesbania sesban and Leucaena leucocephala as
supplements to teff straw given to Ethiopian Menz sheep. Anim. Prod. 59, 235±244.
Bonsi, M.L.K., Osuji, P.O., Tush, A.K., 1995. Effect of supplementing teff straw with different levels of leucaena
or sesbania leaves on degradabilities of teff straw, sesbania, leucaena, tagasaste, tagasaste and veronia and on
certain rumen and blood metabolites in Ethiopian Menz sheep. Anim. Feed Sci. Technol 52, 101±129.
Bosch, M.W., Lammers-Wienhoven, G.A., Bangma, H. Boer, Van-Adrichem, P.W.M., 1993. Influence of stage
of maturity of grass silage on digestion process in dairy cows. 2. Rumen contents, passage rate, distribution
of rumen, distribution of rumen and fecal particles and mastication activity. Livestock Prod. Sci. 32, 265±
281.
Bosch, M.W., Taminga, S., Van Bruchem, J., 1992. Dietary and animal factors affecting rumen capacity in dairy
cows. Arch. Anim. Breed. 34, 265±281.
Brown, W.F., Lai, Z.A., Pitman, W.D., 1991. Associative effects in grass legume mixtures. Trop. Grassl. 14, 78±
82.
Butterworth, M.H., Diaz, L.H.J.A., 1970. The use of equations to predict the nutritive value of tropical grasses. J.
Range Manage. 23, 55±58.
Chauhan, T.R., Gill, R.S., Ichchponani, J.S., 1980. Nutritive value of berseem and clusterbean forages. Indian J.
Anim. Sci. 59(12), 1052±1055.
Chauhan, T.R., Langar, P.N., Chopra, A.K., 1985. Fibre utilization in cattle. Indian J. Anim. Sci. 55(9), 801±807.
Chauhan, T.R., Gupta, R., Chopra, A.K., 1992. Comparative nutritive values of legume hays for buffalo. Buffalo
J. 3, 243±246.
Cheng, K.J., Forsberg, C.V., Minato, H., Eostarton, J.U., 1990. Microbial ecology and physiology of feed
degradation within rumen. Paper presented at 7th Int. Symp. Ruminant Physiology, Sendai, Japan.
Eliott, R., McMeniman, N.P., Norton, B.W., Calderon-Cortes, F.J., 1984. In food intake response of sheep fed
five roughage sources supplemented with formaldehyde treated casein with and without urea. Proc. Aust.
Soc. Anim. Prod. 15, 337±340.
Goering, H.K., Van Soest, P.J., 1970. Forage fibre analysis, ARS, USDA. Agricultural Handbook No. 379, SDA,
Washington, DC, pp. 1±12.
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
149
Hartnell, G.F., Satter, L.D., 1979. Determination of rumen fill, retention time, retention time and ruminal
turnover rates of ingesta at different stages of lactation in dairy cows. J. Dairy Sci. 48, 381±392.
Hyden, S., 1956. Estimation of rumen volume using PEG. J. Agric. Sci. 52, 72±77.
Juul-Nielson, J., 1981. Nutritional principles and productive capacity of the Danish straw mix system for
ruminants. In: Jackson, M.G., Dolberg, F., Haque, M., Saadullah, M. (Eds.), Maximum Livestock Production
from Minimum Land, Mymensingh, Bangladesh, Bangladesh Agri. Univ., Bangladesh, pp. 287±299.
Leng, R.A., 1990. Factors affecting the utilization of poor quality forages by ruminants particularly under
tropical conditions. Nutr. Res. Rev. 3, 277±303.
McMeniman, N.P., Eliott, R., Ash, A.J., 1988. Supplementation of rice straw with crop products. 1. Legume
straw supplementation. Anim. Feed. Sci. Technol. 19, 43±53.
Mertens, D.R., Ely, L.O., 1979. A dynamic model of fibre digestion and passage in the ruminant for evaluating
forage quality. J. Anim. Sci. 49, 1085±1095.
Minson, D.J., 1966. The apparent retention of food in the reticulorumen at two levels of feeding by means of an
purely feeding technique. Br. J. Nutr. 20, 765±773.
Moran, J.B., Satoto, K.B., Dawson, J.E., 1983. The utilization of rice straw fed to Zebu cattle and Swamp
buffalo as influenced by alkali treatment and Leucena supplementation. Aust. J. Agric. Res. 34, 73±84.
Moseley, G., Jones, G.R., 1984. The physical digestion of perennial rye grass (Lolium perene) and while clover
(Trifolium repens) in the foregut of sheep. Br. J. Nutr. 52, 381±390.
Mosi, A.K., Butterworth, M.H., 1985. The voluntary intake and digestibility of combinations of cereal crop
residue and legume hay for sheep. Anim. Feed. Sci. Technol. 12, 241±251.
Nsahlai, I.V., 1991. The effect of quantity and quality of nitrogen on straw utilization by steers, Ph.D. Thesis,
University of Reading, UK.
Odowongo, W.O., Mugerwa, J.S., 1980. Performance of calves on diets containing Amaranthus leaf meal. Anim.
Feed. Sci. Technol. 5, 193±204.
Oosting, S.J., 1993. Wheat straw as a ruminal feed, Ph.D. Thesis, Agricultural University, Wageningen, The
Netherlands.
Oosting, S.J., Wanders, A., Lammers-Wienhoven, S.C.W., Van-Bruchem, J., 1993. Protein supplementation to
ammoniated wheat straw. 1. Intake, digestion, rumen fermentation and passage characteristic in sheep. In:
Oosting, S.J. (Ed.), Wheat Straw as a Ruminal Feed, Ph.D. Thesis, Agricultural University, Wageningen, The
Netherlands, p.121.
Orskov, E.R., McDonald, I., 1979. The estimation of protein degradability in the rumen from incubation
measurements, weighed according to rate of passage, J. Agric. Sci. Camb. 92, p. 499.
Poppi, D.P., Norton, B.W., Minson, D.J., Hendrickson, R.E., 1980. The validity of critical size theory for particle
leaving rumen. J. Agric. Sci Camb. 94, 275±280.
Reddy, D.V., Singh, U.B., Mehra, U.R., 1991. Effect of feeding green berseem in improving rumen degradation
kinetics of wheat straw. Indian J. Anim. Sci. 61(2), 199±201.
Reed, J.D., Soller, H., Woodward, A., 1990. Fodder tree and straw diets for sheep: intake, growth, digestibility
and the effect of phenolics on nitrogen utilization. Anim. Feed Sci. Technol. 30, 39±50.
Robinson, P.H., Fodel, J.G., Tominga, S., 1986. Evolution of mathematical models to describe natural detergent
residues in terms of its susceptibility to degradation in the rumen. Anim. Feed Sci. Technol 15, 249±271.
Silva, A.T., Orskov, E.R., 1988. The effect of five different supplements on the degradation of straw in sheep
given untreated barley straw. Anim. Feed. Sci. Technol. 19, 289±298.
Snedecor, G.W., Cochran, G.H., 1967. Statistical Methods, Oxford and IBH Publications, Calcutta.
Thiogo, L.R.L., Gill, M., Sissons, J.D., 1992. Studies of methods of conserving grass herbage and frequency of
feeding 2. Eating behaviour, rumen motility and rumen fill. Br. J. Nutr. 67, 319±336.
Ulyatt, M.J., Waghorn, G.C., John, A., Reid, C.S.W., Monro, J., 1984. Effect of intake and feeding frequency on
feeding behaviour and quantitative aspects of digestion in sheep fed chaffed lucerne hay. J. Agric. Sci. 102,
645±657.
Wales, W.J., Doyle, P.T., Pearce, G.C., 1990. Feeding value of cereal straws for sheep. 1. Wheat straw. Anim.
Feed Sci. Technol. 29, 1±14.
Woodward, A., Reed, J.D., 1995. Intake and digestibility of sheep and goats consuming supplementary acacia
bravispacia and sesbania sesban. Anim. Feed Sci. Technol 56, 207±216.
81 (1999) 133±149
Effect of different levels of berseem (Trifolium
alexdrinum) supplementation of wheat straw on some
physical factors regulating intake and digestion
Asit Dasa, G.P. Singhb,*
a
ICAR Research Complex, Sikkim Centre, Todong 737102 (Sikkim), India
b
National Research Centre on Carnel, Jorbeer, Bikaner-334001, India
Received 24 April 1998; received in revised form 23 September 1998; accepted 17 March 1999
Abstract
Four ruminally fistulated cattle of about 318 16 kg body weight were randomly distributed into
four groups in an experiment based on 4 4 Latin square design. The four dietary treatments were:
wheat straw ad lib without any supplement (I), supplemented with berseem at 15 (II), 30 (III), and
45 (IV) percent level. DMI was 3.95, 5.85, 7.28 and 7.08 kg; digestibility of DM was 39.85, 45.61,
52.97 and 47.52%; rumen content was 55.60, 68.45, 74.18 and 75.93 kg, or 18.09, 22.01, 23.37 and
23.22% of body weight; rumen fluid content was 48.62, 60.39, 65.20 and 68.01 kg; rumen pool size
of DM was 6.02, 7.61, 7.92 and 7.66 kg; rumen pool size of particles larger than 1.18 mm (LP) was
2.38, 2.78, 2.80 and 2.55 kg; rumen pool size of particles smaller than 1.15 mm (SP) was 3.64, 4.91,
5.12 and 5.05 kg in Groups I, II, III and IV, respectively. Intake and digestibility increased
significantly (p < 0.01), with increased level of berseem up to 30%, beyond which no further
improvement was observed. Total rumen contents, rumen fluid content and pool sizes of DM
increased significantly (p < 0.5) with berseem supplementation, level of berseem had no significant
effect. Pool of LP and SP was similar in all the groups. Maximum rumen content was observed at 9,
9, 6 and 3 h post feeding in Groups I, II, III and IV, respectively. Rate of clearance of LP was 3.71,
6.71, 7.27 and 8.75; SP was 0.46, 0.83, 1.20 and 1.35; and communition rate was 3.45, 5.79, 5.92
and 7.37% hÿ1 in Groups I, II, III and IV, respectively. Rate of clearance of LP and rate of
communition were significantly higher (p < 0.01) in berseem supplemented groups. Rate of passage
(Kp) was 1.72, 1.88, 2.33 and 2.56% per hour. Kp increased significantly (p < 0.01) with increased
level of berseem. From the results it is evident that rumen pool size of indigestible component,
rather than rumen content, determines the intake. As Kd was more than Kp, there was an overall
*
Corresponding author. Tel.: +91-151-523187; fax: +91-151-522183
E-mail address: [email protected] (G.P. Singh)
0377-8401/99/$ ± see front matter # 1999 Elsevier Science B.V. All rights reserved.
PII: S 0 3 7 7 - 8 4 0 1 ( 9 9 ) 0 0 0 6 0 - 7
134
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
improvement of digestibility by 14, 33 and 19% in Groups II, III and IV, respectively, compared
with Group I. # 1999 Elsevier Science B.V. All rights reserved.
Keywords: Level of berseem; Berseem supplementation; Wheat straw; Cattle; Intake; Regulation; Digestion
1. Introduction
Wheat straw forms a major source of ruminant feed in India. Unfortunately, wheat
straw is characterised by low intake and digestibility. Supplementation of poor quality
roughage, including wheat straw, with green forages has been shown to increase intake
(Woodward and Reed, 1995; Reed et al., 1990) or digestibility (Silva and Orskov, 1988;
Ash, 1990; Bonsi et al., 1995) or both (Leng, 1990; Bird et al., 1994). However, type of
basal diet (Mosi and Butterworth, 1985; Eliott et al., 1984), degradability characteristics
of the supplement (Bates et al., 1988), level of supplement (Bonsi et al., 1994) as well as
interactions between nutrients in feed ingredients (Brown et al., 1991) are important
determinants of response to supplementation. Hence, specific study is required to know
the effect of a particular supplement on intake and digestibility of the basal diet
concerned.
Berseem (Trifolium alexdrinum) is a green leguminous fodder grown during winter.
Berseem contains 15±20% CP and 4.1 kcal/g GE (Chauhan et al., 1980). It is highly
palatable, fairly digestible and on sole feeding can support growth rate of 550 g/d
(Chauhan et al., 1992) and milk production up to 10 kg/d. Berseem supplementation to
wheat straw has been shown to increase in sacco dry matter degradability of wheat straw
(Reddy et al., 1991). In practice, farmers in India feed wheat straw and berseem mixed
together without considering the ratio. However, detailed information regarding the effect
of different levels of berseem supplementation on digestion and intake of straw-based diet
is not available. At the same time, prediction of likely effect on intake and digestion will
be difficult without a better understanding of the processes, namely, rumenfil (Bosch
et al., 1993), particle dynamics (Poppi et al., 1980) and passage rate (Oosting et al.,
1993), which regulate intake. Hence, this experiment was undertaken to find out the level
of berseem at which intake and digestion is maximum, with a better understanding of the
physical factors regulating them.
2. Materials and methods
2.1. Animals and design
Four ruminally fistulated crossbred (Shaiwal Holstein friesan) steers of about
318 16 kg body weight were kept in stalls, with individual feeding arrangements. The
experiment was set up as a 4 4 Latin Square design, with each experimental period
lasting 3 weeks. Adaptation period before each experimental period had a duration of 3
weeks. The time schedule of each experimental period was:
Activity
Day
Adaptation
Rumen liquor sampling
Intake and digestion
Dacron bag analysis of feed sample
Rumen evacuation
Rumen evacuation on restricted feeding
1±14
15
16±22
21±25
26±31
32±35
Four dietary treatments were wheat straw alone (I), wheat straw supplemented with 15
(II), 30 (III), and 45% (IV) level of berseem. Supplement was offered once daily at 9 a.m.
Clean and fresh drinking water was provided ad lib. The amount of supplement to be
offered was determined from the record of previous day intake, to keep the level of
berseem to the desired experimental level as far as possible.
2.2. Chemical composition of feed
Samples of wheat straw and berseem were taken daily during the last 21 days of each
period. They were dried at 50±608C and analysed for dry matter (DM), ash, neutral
detergent fibre (NDF), acid detergent fibre (ADF) and total nitrogen (N) content.
2.3. Digestion trial
During each period, a digestion trial of six-day collection (Days 17±22) was conducted
to determine the intake and digestibility of diets.
2.4. Rumen evacuation
Rumen evacuations were done during Days 26±31 in each period. Each steer's rumen
was evacuated once daily over 6 days, either at 12:00, 15:00, 18:00, 21:00, 3:00 or 9:00
136
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
2.4.1. Rumen evacuation of animals of restricted feeding
During last 4 days of each experimental period, rumen evacuations were done on
animals receiving restricted amount of feed. The animals were allowed to eat up to 12:00
hours. Each steer's rumen was evacuated once daily over 4 days, either on 12:00, 18:00 or
24:00 hours. There was a minimum interval of 30 h between emptying from any
individual steer to overcome the effect of emptying on the repeatability of measurements.
Total rumen contents were removed by hands, weighed, sampled and the remainder
returned to the rumen. Two samples of 500 g were taken: the first one for separation into
large (>1.18 mm) and small (>0.041 mm, 1.18 mmb
Pool < 1.18 mm*
Total*
Pool NDF (kg)b
2.38
3.64 a
6.02 a
4.74
2.78
4.91 b
7.61 b
5.63
2.80
5.12 b
7.92 b
5.93
2.55
5.05 b
7.66 b
5.42
2.08
0.23
0.37
0.45
Indigestible components
DM (kg)b
NDF (kg)b
3.66
2.91
4.12
3.22
4.00
3.04
3.98
3.11
0.23
0.19
a
b
Values followed by different letters in a row differ significantly.
Non-significant; **p < 0.01.
Table 4
Diurnal variation of rumen pool sizes in animals fed different level of berseem in dieta
I
Total rumen contents (kg) h post feeding
3**
55.0 a
59.5 a
6**
9*
61.0 a
12*
56.5 a
18b
55.5
51.5
24b
II
66.3
74.0
75.2
71.8
63.8
59.3
III
ab
b
b
b
75.5
81.8
76.0
74.0
71.5
62.5
IV
c
bc
b
b
83.0
83.0
80.2
75.8
71.5
61.5
SEM
c
c
b
b
3.05
2.49
3.61
3.17
4.38
2.58
DM (kg) h post feeding
3*
6.08 a
6.50 a
6**
9b
6.68
12*
5.95
18b
5.60
24b
5.25
7.40 b
7.98 b
8.38
8.08 b
7.38
6.55
8.40 c
8.60 b
8.05
7.78 b
7.55
7.13
7.98 c
8.30 b
7.58
7.48 b
7.33
6.95
0.40
0.30
0.35
0.39
0.45
0.44
NDF (kg) h post feeding
3b
4.83
5.10 a
6*
9b
5.23
12b
4.68
18b
4.40
4.13
24b
5.60
6.03 b
6.13
6.03
5.68
4.98
6.30
6.45 b
6.03
5.80
5.65
5.33
5.68
5.95 ab
5.58
5.28
5.13
4.90
0.34
0.25
0.32
0.32
0.39
0.38
a
b
Values followed by different letters in a row differ significantly.
Non-significant; **p < 0.01.
142
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
Table 5
Potentially degradable fraction (D), rate of degradation (Kd), isolated small and large particles, whole rumen
NDF pool, rate of clearance of large particle (LP) and small particle (SP), and rate of communition of large
particlea
I
II
III
IV
SEM
D fraction (%)
LP**
SP**
Rumen NDF pool*
25.20 a
36.48 a
33.68 a
40.02 b
41.94 c
43.96 b
48.11 c
45.90 d
45.03 b
49.44 d
39.08 b
40.83 ab
0.34
0.36
2.06
Kd (%/h)
LP**
SP**
Rumen NDF pool*
0.83 a
1.66 b
1.54 a
1.97 b
2.01 c
2.07 b
2.46 c
1.93 c
2.15 b
2.26 c
1.41 a
1.84 b
0.04
0.03
0.10
Rate of clearance (%/h)
LP**
SPb
Kc (%/h)**
3.71 a
0.46
3.45 a
6.71 b
0.83
5.79 b
7.27 b
1.20
5.92 b
8.75 c
1.35
7.37 c
0.36
0.21
0.37
a
b
Values followed by different letters in a row differ significantly.
Non-significant; **p < 0.01.
Groups IV, III, II and I, respectively. At around 18 h post feeding, the difference among
different groups was non-significant. Similarly, the rumen DM as well as NDF pool were
maximum at 6, 6, 9 and 9 h post feeding in Groups IV, III, II and I, respectively.
3.5. Particle dynamics
Degradation characteristics of rumen content, isolated LP, SP and rate of clearance of
SP, LP and rate of communition are presented in Table 5, and distribution of SP and LP is
illustrated in Fig. 2. Potential degradability of LP increased significantly (p < 0.01) with
increased level of berseem in the diet. Potential degradability of SP and whole rumen
NDF pool increased significantly (p < 0.01) with increased berseem level up to 30%.
However, at 45% level of supplementation, potential degradability decreased for both SP
and whole rumen NDF pool, difference being more marked in case of SP than whole
rumen NDF pool. Rate of degradation of LP increased with increased level of berseem up
to 30%; at 45% level of berseem, no further change was observed. Rate of degradation of
SP reduced significantly (p < 0.01) in Group IV in comparison to control, whereas 15 and
30% levels of berseem increased the rate of degradation.
3.6. Passage from rumen
Rate of clearance (Kcl), rate of passage (Kp) and mean retention time (MRT) of DM and
NDF are presented in Table 6. Kcl of DM increased significantly (p < 0.01) with increased
level of berseem up to 30%, beyond which no further change was observed. On the other
hand, MRT decreased and Kp increased significantly (p < 0.01) with increased level of
berseem in the diet.
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
143
Fig. 2. Diurnal variation of particle distribution in rumen as affected by the level of berseem in wheat straw diet.
144
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
Table 6
Clearance rate (Kcl), mean retention time (MRT), fractional passage rate (Kp) and liquid outflow rate (Kl)a
I
II
III
IV
SEM
Kcl (%/h)
DM**
NDF*
2.73 a
2.85 a
3.22 b
3.40 b
3.85 c
3.67 b
3.89 c
3.76 b
0.096
0.14
MRT (h)
IDM**
INDF**
58.33 a
55.71 a
53.24 b
50.74 b
42.98 c
45.72 c
40.20 c
42.09 d
1.01
0.97
Kp (%/h)
DM**
NDF**
Klb
1.72 a
1.81 a
6.60
1.88 b
1.96 b
7.08
2.33 b
2.19 c
7.18
2.56 b
2.38 d
7.85
0.047
0.042
0.58
Intake (kg)
DM**
IDM**
NDF**
INDF**
3.95 a
1.52 a
3.22 a
1.27
5.85 b
1.89 b
4.55 b
1.52
7.28 c
2.01 c
5.01 c
1.55
7.08 c
2.36 c
4.65 c
1.77
0.16
0.064
0.108
0.004
a
b
Values followed by different letters in a row differ significantly.
Non-significant; **p < 0.01.
3.7. Diurnal variation in passage rate
Within-day variation of passage rate is presented in Table 7. In this experiment,
passage rate was determined for the periods of 0±9 h, 9±18 h and 18±24 h and grossly
considered to be as the period of eating, rumination and idling, respectively, because the
activity concerned was more in the respective period. Maximum passage was observed
during eating in all the groups, except wheat straw alone fed group, where maximum
passage was observed during rumination. However, the absolute amount passaged during
rumination and idling was not different among the rations.
Table 7
Variation in passage rate of INDF from the potentially removable INDF pool of rumen during eating (PEAT),
rumination (PRUM) and idling (PID)a
INDF intake (%)
PEAT**
PRUMb
PIDb
Passage (g)
PEAT**
PRUMb
PIDNS
a
b
I
II
III
IV
35.96 a
46.32
17.72
46.18 b
33.26
20.56
59.93 c
26.73
12.97
67.74 c
23.24
9.33
462.5 a
571.0
234.5
713.3 b
568.0
317.0
930.3 b
402.0
213.0
Values followed by different letters in a row differ significantly.
Non-significant; **p < 0.01.
1197.3 c
407.0
165.0
SEM
3.08
8.73
5.55
72.5
66.0
80.5
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
145
4. Discussion
4.1. Intake and digestibility
Inclusion of berseem in the wheat straw based diet increased the total DM intake.
Increased DM intake as a result of green legume supplementation has been reported by
other workers (Mosi and Butterworth, 1985; McMeniman et al., 1988; Bird et al., 1994;
Bonsi et al., 1994; Woodward and Reed, 1995). Findings of this experiment are in line
with information available in the existing literature in general, but differ from the findings
of Odowongo and Mugerwa (1980) and Ash (1990). Such variations in response can
partly be attributed to the lower degradability of the supplement (Ash, 1990) and also to
the presence of toxicant (Odowongo and Mugerwa, 1980) in the supplement. Berseem, on
the other hand, is fairly degradable and contains no toxicant that is known to decrease
intake.
Berseem supplementation up to 30% level increased digestibility of DM, OM, NDF,
ADF, cellulose and hemicellulose; however, at 45% level of berseem supplementation,
digestibility decreased. On the other hand, CP digestibility increased linearly with
increased level of berseem in the diet, which is in line with the findings of Butterworth
and Diaz (1970) and Mosi and Butterworth (1985). Increased digestibility as a result of
forage supplementation has been reported by other workers (Juul-Nielson, 1981; Silva
and Orskov, 1988; Bird et al., 1994; Bonsi et al., 1994). Maximum digestibility observed
at 30% level of berseem supplementation suggests that 30% level of berseem in the diet
provided all the essential nutrients critical for optimal microbial activity, viz., rumen
NH3±N concentration (Eliott et al., 1984), peptides, essential minerals and vitamins
(Leng, 1990). In addition, berseem also provided fermentable cellulose and hemicellulose, which are known to promote fibre digestion (Silva and Orskov, 1988) by
ensuring greater degree of colonisation of fibrolytic bacteria (Cheng et al., 1990) and
fungi (Bauchop, 1979). Decrease in digestibility as a result of 45% berseem
supplementation, as observed in this experiment, could be due to decreased mean
retention time in the rumen, which means an increased rate of passage from the rumen
(Oosting, 1993) as the green forages are bulky in nature (Bonsi et al., 1994).
4.2. Rumen pool size
Total rumen content was 18% of the body weight in animals fed on wheat straw alone;
it increased up to 23% when supplemented with berseem. These values were higher than
the values of 17.5% in cattle fed on either grass silage based diet (Bosch et al., 1993) or
wheat straw based diet (Oosting, 1993), but lower than the values of 26% in early
lactating cows (Hartnell and Satter, 1979). These values are similar to those observed by
Chauhan et al. (1985) in buffalo fed on oat silage based diet supplemented with either
concentrate or cowpea hay. They reported that gut content is 24%. It appears that
ruminants fed on poor quality roughage, in the tropics, have higher rumen content than of
those in the temperate region. Berseem supplementation increased the total rumen
content and pool of DM and small particle. This indicates that intake is not limited by
rumen capacity. Similar conclusion has also been drawn by other workers (Bosch et al.,
146
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
1992; Oosting, 1993). Similar pool size of indigestible components (IDM and INDF) in
different groups having different levels of intake, as observed in this experiment, suggests
that rumen pool of indigestible component determines the intake (Mertens and Ely,
1979).
Generally, after attainment of maximum rumen fill, it declined and reached a minimum
value just before feeding. Maximum pool of DM was observed 6 h post feeding in Groups
III and IV, and 9 h post feeding in Groups I and II (Table 4). Pool of NDF followed the
same trend, but the difference between different groups was significant only at 6 h post
feeding. The pools of IDM and INDF gradually decreased after they attained the
maximum pool at 6, 6, 9 and 9 h post feeding in Groups IV, III, II and I, respectively.
Increased passage rate observed with increased level of berseem in the diet due to its
bulkiness (Bonsi et al., 1994) can partly be attributed to early attainment of maximum
rumen pool.
4.3. Particle dynamics
The results indicate that LP and SP are not degraded in the same fashion. Increase in
potential degradability and rate of degradation of LP with increased level of berseem in
the diet could be attributed to increase in readily fermentable fibre content (Silva and
Orskov, 1988). Potential degradability of whole rumen NDF, SP increased up to 30%
level of berseem, beyond which a negative associative effect was observed. It seems that
the decrease in potential degradability and rate of degradation at 45% level of berseem is
due to decrease in potential degradability (PD) and Kd of SP. The decreased PD and Kd of
SP could be due to the fact that at 45% level of berseem, the rumen, microbes had enough
substrate in rumen pool of host animal than the less preferably small particles, inside
nylon bag when isolated from the rumen which ready to passage out of the rumen and
substantial amount of digestion might had taken place before that. Rate of degradation of
LP, barring wheat straw, was higher than SP in all the groups. This, in general, is in
agreement that after attainment of critical particle size (1.18 mm) the feed particle do not
remain in rumen for long (Poppi et al., 1980) and substantial amount of microbial
digestion takes place before they attain that particle size.
4.4. Passage rate from rumen
Passage from rumen is limited only to particles smaller than 1.18 mm (Poppi et al.,
1980). In this experiment, proportion of small particle in different groups was similar, i.e.,
53%. The mean proportion of small particle was less than the value of 70% observed by
Oosting (1993). The difference can be explained on the basis of coarseness of the
supplement in our experiment. In the earlier experiment, sugar beet pulp having much
smaller particle size was used as supplement to wheat straw, whereas in this experiment
whole berseem without chaffing was used. The rate of communition increased with
increased level of berseem. Increased rate of communition can either lead to increase in
pool of SP or increased passage rate. In this experiment, no change in rumen pool size
beyond 15% level of berseem was observed. This indicates that the increased
communition can increase the pool of SP only to a limited extent, rest of the SP being
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
147
rapidly lost after their formation (Moseley and Jones, 1984). In this experiment, rate of
passage increased with increased level of berseem supplementation. Similar results have
also been reported by other workers (Moran et al., 1983; Bamualim et al., 1984; Bonsi
et al., 1994) when straw-based diets were supplemented with increased level of green
forages. Increased passage rate as a result of increased level of berseem supplementation
could probably be associated with the fact that berseem supplementation increased the
ratio of cellulose/hemicellulose in the diet. Particles with higher cellulose content
probably have a relatively higher functional specific gravity (Wales et al., 1990), which
may explain increased passage rate with increased level of berseem in the diet.
In this experiment, no change in rumen pool size beyond 15% level of berseem was
observed. Mean retention time decreased with increased level of berseem in the diet, as
passage rate was measured as reciprocal of the mean retention time, in this experiment.
Decreased mean retention time as a result of supplementation has also been reported by
other workers (Moran et al., 1983; Bonsi et al., 1994). Though the passage rate of
particulate matter increased with increased level of berseem supplementation, liquid
outflow rate was not significantly different between treatments. This finding is contrary to
that of Nsahlai (1991) and Bonsi et al. (1994), who reported that increased particulate
passage rate results in increased liquid outflow rate. Nevertheless, a different pattern of
particulate passage rate and liquid outflow rate has also been reported (Ulyatt et al.,
1984).
4.5. Diurnal variation in passage rate
Difference in passage observed in this experiment, as a result of berseem
supplementation, can be attributed to the increased rate of passage during eating. On
an average, 36, 46, 60 and 68% of indigestible NDF ingested was lost from the rumen
INDF pool, during eating, in Groups I, II, III and IV, respectively. The proportion of large
particles in the rumen increased up to 3 h post feeding; then it gradually decreased and
reached minimum just before feeding. Reduction in particle size was sharper in animals
fed on higher level of berseem, indicating higher rate of communition that resulted in
increased rate of passage as proportion of small particles in rumen was not significantly
different among the rations. The act of eating has been shown to be accompanied by a
marked increase in reticular and omasal contractions that might have elevated the rate of
flow of digesta from the rumen (Balch, 1958). It is observed in this experiment that
greater proportion of INDF is lost during eating with increased level of berseem in the
diet. This change might have occurred due to increased rumination efficiency (Moseley
and Jones, 1984). On an average, 462, 713, 930 and 1197 g of INDF was lost from the
rumen within first 9 h of feeding in Groups I, II, III and IV, respectively.
Berseem supplementation increased the rumen fill, indicating that rumen capacity is
not limiting intake. On an average, an increase of 23, 40 and 44% in DM intake was
recorded as a result of 15, 30 and 45% berseem supplementation. Increased intake was
accompanied by an increased rate of communition, which, in turn, increased the rate of
passage. However, the increase in disappearance was more than the rate of passage. On an
average, 19, 29 and 32% increase in clearance rate was observed in 15, 30 and 45% level
of berseem supplemented groups, respectively. Respective values for increase in Kp was
148
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
8, 21 and 31%. Overall digestibility increased by 14, 33 and 19% in groups fed 15, 30 and
45% level of berseem, respectively. It is concluded that increased level of berseem
supplementation up to 30% level to wheat straw based diet increases DMI and
digestibility of wheat straw based diet.
References
American Society of Agricultural Engineers, 1967. Methods of determining modulus of uniformity and modulus
of fineness of ground feed, Agricultural Engineers Yearbook, p. 301.
AOAC, 1984. Official Analytical of Method, 14th ed., Association of Official Analytical Chemists, Washington,
DC.
Ash, A.J., 1990. The effect of supplementation with leaves from the leguminous trees Sesbonia grandiflora,
Albizia chinensis, Albizia chinensis and Gliricidia sepium on the intake and digestibility of guinea grass hay
by goat. Anim. Feed. Sci. Technol. 28, 225±232.
Balch, C.C., 1958. Observation on the act of eating in cattle. Br. J. Nutr. 12, 330±345.
Bamualim, A., Weston, R.H., Hogan, J.P., Murray, R.M., 1984. The contribution of Leucaena leucocephala to
post ruminal digestible protein for sheep fed tropical pasture hay supplemented with urea and minerals. Proc.
Australia Soc. Anim. Prod. 15, 225±258.
Bates, D.B., Morantes, G.A., Moore, G.E., 1988. Nitrogen utilization of lombs fed limpo grass supplemented
with molasses-urea or legume protein supplements. Nutr. Rep. Intt. 38, 487±499.
Bauchop, T., 1979. Rumen anaerobic fungi of cattle and sheep. Appl. Environ. Microbiol. 38, 148±158.
Bird, S.H., Romulo, B., Leng, R.A., 1994. Effect of Lucerne supplementation and defaundation on feed intake,
digestibility, nitrogen retention and productivity of sheep fed straw based diets. Anim. Feed. Sci. Technol.
45, 119±129.
Bonsi, M.L.K., Osuji, P.O., Tuan, A.K., 1994. Graded level of Sesbania sesban and Leucaena leucocephala as
supplements to teff straw given to Ethiopian Menz sheep. Anim. Prod. 59, 235±244.
Bonsi, M.L.K., Osuji, P.O., Tush, A.K., 1995. Effect of supplementing teff straw with different levels of leucaena
or sesbania leaves on degradabilities of teff straw, sesbania, leucaena, tagasaste, tagasaste and veronia and on
certain rumen and blood metabolites in Ethiopian Menz sheep. Anim. Feed Sci. Technol 52, 101±129.
Bosch, M.W., Lammers-Wienhoven, G.A., Bangma, H. Boer, Van-Adrichem, P.W.M., 1993. Influence of stage
of maturity of grass silage on digestion process in dairy cows. 2. Rumen contents, passage rate, distribution
of rumen, distribution of rumen and fecal particles and mastication activity. Livestock Prod. Sci. 32, 265±
281.
Bosch, M.W., Taminga, S., Van Bruchem, J., 1992. Dietary and animal factors affecting rumen capacity in dairy
cows. Arch. Anim. Breed. 34, 265±281.
Brown, W.F., Lai, Z.A., Pitman, W.D., 1991. Associative effects in grass legume mixtures. Trop. Grassl. 14, 78±
82.
Butterworth, M.H., Diaz, L.H.J.A., 1970. The use of equations to predict the nutritive value of tropical grasses. J.
Range Manage. 23, 55±58.
Chauhan, T.R., Gill, R.S., Ichchponani, J.S., 1980. Nutritive value of berseem and clusterbean forages. Indian J.
Anim. Sci. 59(12), 1052±1055.
Chauhan, T.R., Langar, P.N., Chopra, A.K., 1985. Fibre utilization in cattle. Indian J. Anim. Sci. 55(9), 801±807.
Chauhan, T.R., Gupta, R., Chopra, A.K., 1992. Comparative nutritive values of legume hays for buffalo. Buffalo
J. 3, 243±246.
Cheng, K.J., Forsberg, C.V., Minato, H., Eostarton, J.U., 1990. Microbial ecology and physiology of feed
degradation within rumen. Paper presented at 7th Int. Symp. Ruminant Physiology, Sendai, Japan.
Eliott, R., McMeniman, N.P., Norton, B.W., Calderon-Cortes, F.J., 1984. In food intake response of sheep fed
five roughage sources supplemented with formaldehyde treated casein with and without urea. Proc. Aust.
Soc. Anim. Prod. 15, 337±340.
Goering, H.K., Van Soest, P.J., 1970. Forage fibre analysis, ARS, USDA. Agricultural Handbook No. 379, SDA,
Washington, DC, pp. 1±12.
A. Das, G.P. Singh / Animal Feed Science and Technology 81 (1999) 133±149
149
Hartnell, G.F., Satter, L.D., 1979. Determination of rumen fill, retention time, retention time and ruminal
turnover rates of ingesta at different stages of lactation in dairy cows. J. Dairy Sci. 48, 381±392.
Hyden, S., 1956. Estimation of rumen volume using PEG. J. Agric. Sci. 52, 72±77.
Juul-Nielson, J., 1981. Nutritional principles and productive capacity of the Danish straw mix system for
ruminants. In: Jackson, M.G., Dolberg, F., Haque, M., Saadullah, M. (Eds.), Maximum Livestock Production
from Minimum Land, Mymensingh, Bangladesh, Bangladesh Agri. Univ., Bangladesh, pp. 287±299.
Leng, R.A., 1990. Factors affecting the utilization of poor quality forages by ruminants particularly under
tropical conditions. Nutr. Res. Rev. 3, 277±303.
McMeniman, N.P., Eliott, R., Ash, A.J., 1988. Supplementation of rice straw with crop products. 1. Legume
straw supplementation. Anim. Feed. Sci. Technol. 19, 43±53.
Mertens, D.R., Ely, L.O., 1979. A dynamic model of fibre digestion and passage in the ruminant for evaluating
forage quality. J. Anim. Sci. 49, 1085±1095.
Minson, D.J., 1966. The apparent retention of food in the reticulorumen at two levels of feeding by means of an
purely feeding technique. Br. J. Nutr. 20, 765±773.
Moran, J.B., Satoto, K.B., Dawson, J.E., 1983. The utilization of rice straw fed to Zebu cattle and Swamp
buffalo as influenced by alkali treatment and Leucena supplementation. Aust. J. Agric. Res. 34, 73±84.
Moseley, G., Jones, G.R., 1984. The physical digestion of perennial rye grass (Lolium perene) and while clover
(Trifolium repens) in the foregut of sheep. Br. J. Nutr. 52, 381±390.
Mosi, A.K., Butterworth, M.H., 1985. The voluntary intake and digestibility of combinations of cereal crop
residue and legume hay for sheep. Anim. Feed. Sci. Technol. 12, 241±251.
Nsahlai, I.V., 1991. The effect of quantity and quality of nitrogen on straw utilization by steers, Ph.D. Thesis,
University of Reading, UK.
Odowongo, W.O., Mugerwa, J.S., 1980. Performance of calves on diets containing Amaranthus leaf meal. Anim.
Feed. Sci. Technol. 5, 193±204.
Oosting, S.J., 1993. Wheat straw as a ruminal feed, Ph.D. Thesis, Agricultural University, Wageningen, The
Netherlands.
Oosting, S.J., Wanders, A., Lammers-Wienhoven, S.C.W., Van-Bruchem, J., 1993. Protein supplementation to
ammoniated wheat straw. 1. Intake, digestion, rumen fermentation and passage characteristic in sheep. In:
Oosting, S.J. (Ed.), Wheat Straw as a Ruminal Feed, Ph.D. Thesis, Agricultural University, Wageningen, The
Netherlands, p.121.
Orskov, E.R., McDonald, I., 1979. The estimation of protein degradability in the rumen from incubation
measurements, weighed according to rate of passage, J. Agric. Sci. Camb. 92, p. 499.
Poppi, D.P., Norton, B.W., Minson, D.J., Hendrickson, R.E., 1980. The validity of critical size theory for particle
leaving rumen. J. Agric. Sci Camb. 94, 275±280.
Reddy, D.V., Singh, U.B., Mehra, U.R., 1991. Effect of feeding green berseem in improving rumen degradation
kinetics of wheat straw. Indian J. Anim. Sci. 61(2), 199±201.
Reed, J.D., Soller, H., Woodward, A., 1990. Fodder tree and straw diets for sheep: intake, growth, digestibility
and the effect of phenolics on nitrogen utilization. Anim. Feed Sci. Technol. 30, 39±50.
Robinson, P.H., Fodel, J.G., Tominga, S., 1986. Evolution of mathematical models to describe natural detergent
residues in terms of its susceptibility to degradation in the rumen. Anim. Feed Sci. Technol 15, 249±271.
Silva, A.T., Orskov, E.R., 1988. The effect of five different supplements on the degradation of straw in sheep
given untreated barley straw. Anim. Feed. Sci. Technol. 19, 289±298.
Snedecor, G.W., Cochran, G.H., 1967. Statistical Methods, Oxford and IBH Publications, Calcutta.
Thiogo, L.R.L., Gill, M., Sissons, J.D., 1992. Studies of methods of conserving grass herbage and frequency of
feeding 2. Eating behaviour, rumen motility and rumen fill. Br. J. Nutr. 67, 319±336.
Ulyatt, M.J., Waghorn, G.C., John, A., Reid, C.S.W., Monro, J., 1984. Effect of intake and feeding frequency on
feeding behaviour and quantitative aspects of digestion in sheep fed chaffed lucerne hay. J. Agric. Sci. 102,
645±657.
Wales, W.J., Doyle, P.T., Pearce, G.C., 1990. Feeding value of cereal straws for sheep. 1. Wheat straw. Anim.
Feed Sci. Technol. 29, 1±14.
Woodward, A., Reed, J.D., 1995. Intake and digestibility of sheep and goats consuming supplementary acacia
bravispacia and sesbania sesban. Anim. Feed Sci. Technol 56, 207±216.