Directory UMM :Data Elmu:jurnal:A:Animal Feed Science and Technology:Vol83.Issue3-4.Mar2000:
Animal Feed Science and Technology
83 (2000) 287±300
Energy metabolism with particular reference to
methane production in Muzaffarnagari sheep fed
rations varying in roughage to concentrate ratio
Chandramoni*,1, S.B. Jadhao2, C.M. Tiwari3, M.Y. Khan
Energy Metabolism and Respiration Calorimetry Laboratory, Animal Nutrition Division,
Indian Veterinary Research Institute, Izatnagar 243 122, India
Received 28 April 1997; received in revised form 28 January 1999; accepted 11 November 1999
Abstract
Methane production in Muzaffarnagari sheep was studied using open circuit respiration
calorimetry technique. Twelve rams were divided in three treatment groups of four each and were
fed at about maintenance with diets having three roughage (oat hay) to concentrate ratio (R : C), i.e.
92 : 8 (group I), 50 : 50 (group II), 30 : 70 (group III). Concentrate mixture was formulated to
contain 93% crushed maize grain, 3.5% wheat bran and 3.5% groundnut cake forti®ed with
minerals. Whereas the digestibility of dry matter, organic matter and energy was similar, the
digestibility of nitrogen was signi®cantly (p < 0.05) higher in groups II and III and of neutral
detergent ®ber (NDF) was signi®cantly (p < 0.05) lower in-group III than the other groups. Nitrogen
retention was improved but not beyond 50R : 50C. As a percent of gross energy intake, urinary
energy losses in groups I, II and III were 3.0, 2.9 and 2.8%, and methane energy losses were 3.39,
3.34 and 2.98%, respectively. Even though, gross energy intakes (kcal/kg W0.75) were similar,
methane loss (g) per 100 g digestible organic matter was signi®cantly (p < 0.05) higher in group I
(2.2) than in groups II (1.84) and III (1.54), the latter did not differ in this respect. Metabolisable
energy (ME) value (Mcal/kg DM) and energy balances (kcal/kg W0.75) on rations in groups II and
III were similar but on ration in group I was signi®cantly (p < 0.05) lower than that in other groups.
Ef®ciency of utilisation of ME for maintenance (km) of diets in groups I, II and III calculated as per
ARC (1980) were 0.674, 0.688 and 0.693, respectively, and did not differ signi®cantly. Based on the
evaluation of three R : C, it was inferred that R : C ratio of 50 : 50 in diet of Muzaffarnagari sheep is
*
Corresponding author. Fax: 91-226361573.
E-mail address: chandramoni@hotmail.com (Chandramoni).
1
Quarter 11, Road 9, Bihar Veterinary College, Patna 800 014, India.
2
Central Institute of Fisheries Education, Seven Bunglows, Versova, Mumbai 400 061.
3
Deptt. Animal Nutrition, Rajiv Gandhi College of Veterinary and Animal Sciences, Kurumbapeth,
Pondicherry 605 009.
0377-8401/00/$ ± see front matter # 2000 Published by Elsevier Science B.V. All rights reserved.
PII: S 0 3 7 7 - 8 4 0 1 ( 9 9 ) 0 0 1 3 2 - 7
288
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
optimum for economical and sustainable sheep production through reduced methane emissions.
# 2000 Published by Elsevier Science B.V. All rights reserved.
Keywords: Muzaffarnagari sheep; Energy metabolism; Methane production; Roughage: concentrate ratios;
Nutritive value
1. Introduction
Methane production in recent years has assumed more signi®cance in animal
production due to its undesired effect on environment including its contribution to global
warming. There is now considerable general understanding of the contribution of
ruminants to methane production (Johnson and Johnson, 1995; Mathison et al., 1998).
Sustainable animal production systems henceforth require that methane produced should
be less per unit output. Dietary manipulation is one of the means to reduce methane (see
Review, Moss, 1994) and is practicable in India and other third World countries.
Optimum roughage to concentrate is one of the dietary means, which has potential to
reduce methane production. Number of studies concerning supplementation of
concentrate to straw or poor quality roughages has been done which has demonstrated
reduced methane production (Sundstol, 1982; Birkelo et al., 1986; Wainman and Dewey,
1987; Silva and érskov, 1988). However, few studies on supplementation to good quality
hay-based diets near maintenance level has been done. The effects of supplementation of
concentrate to good quality roughages are unclear (Moss, 1994). Blaxter and Wainman
(1964) showed greater production of methane for hay than maize grain and that methane
production with mixtures of these feeds was not additive and was invariably greater than
could be predicted from composition of the rations. In contrast, the ME content and the
ef®ciency of utilisation of metabolisable energy (ME) for maintenance and fattening
increased from 71 to 79 and 29 to 61%, respectively. Hence, it is likely that methane
production per unit of animal product would decline with increasing maize in ration. On
the other hand, Webster (1987) observed reduce energy loss as methane with increasing
the ratio of cereal to forage, which decreases the ratio of acetic to propionic acid. Low
cereal inclusion rate to grass silage-based diets enhances milk yield and increase milk fat
content (Rae et al., 1986) when compared with silage fed alone. Methane produced per
unit animal product in this situation seems to be decreased. High propionate
fermentations (indicative of reduced methane) are common in hay-based diet containing
high proportion of concentrate (Thomas and Chamberlain, 1982).
Therefore, it was thought to investigate methane production on good quality oat haybased rations with different roughage to concentrate ratio. As such the combined effect
(associate effect) of starchy feeds (alone or with little quantity of oil cake and bran)
supplementation to good quality roughage like oat hay is little studied. Similarly, the
literature on energy balance studies in Muzaffarnagari sheep (sturdy and meat type breed)
which is important animal resource in this area, is lacking. Keeping in view the above
facts, the present study was speci®cally designed to study energy metabolism of
Muzaffarnagari sheep with special reference to methane production on diets varying in
roughage and concentrate ratio.
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
289
2. Materials and methods
2.1. Selection of animals
Fifteen healthy male uncastrated, Muzaffarnagari sheep, ca. 6±8 months of age, were
procured in October, 1993 from Central Institute for Research on Goats, Makhdoom. The
animals were dewormed before start of the experiment and good managerial practices
were followed in the sheds. The animals were fed on a balanced ration for about a month
before the start of experimental feeding.
2.2. Housing and management
The animals were housed in a well-ventilated shed having cemented ¯oor with
individual feeding and watering arrangement throughout the experimental period.
Concentrate mixture was offered to individual animals between 9.00 and 10.00 a.m. and
oat hay was offered in after noon at 3.00 a.m. in the same manger. Fresh water was
provided ad libitum daily at 2.30 p.m.
2.3. Experimental plan
Twelve sheep were randomly allocated on the basis of liveweight to three groups
comprising each of four animals, following complete randomized design. Rations were
given to meet the maintenance requirement of sheep as per NRC (1985). Group I was
offered oat hay plus minimal concentrate mixture (ca. 100 g of concentrate mixture I) to
meet CP requirement. Group II was offered 50% concentrate mixture and 50% oat hay,
whereas group III animals were offered 70% concentrate mixture and 30% oat hay only.
Concentrate mixture (11.7% CP and 4.37 Mcal GE/kg DM) contained 93% crushed
maize grain, 3.5% deoiled groundnut cake, and 3.5% wheat bran. To every 100 kg
concentrate mixture 2 kg of mineral mixture (contained moisture maximum 5%, Calcium
minimum 28%, Phosphorus minimum. 12%, Iodine as KI 0.026±0.130%, Copper 0.077±
0.130%, ¯uorine maximum 0.04%) and 1 kg of common salt were added.
Metabolism trial was conducted after 60 days of feeding (with three days of
adaptation) in metabolic cages for seven days. All the animals were weighed before and
after the trial. Representative samples of feed offered and residue left were taken daily for
dry matter estimation and analysis. The total amount of faeces voided by each animal was
collected quantitatively at 9 a.m. daily. It was weighed and representative sample of each
animal was drawn in sample bottles after crushing and mixing all the pellets and was
brought to the laboratory for analysis. Urine excreted daily by individual animal was
collected separately in bucket with dilute sulphuric acid for 24 h and was measured with
the help of measuring cylinder. Representative samples from individual animals were
brought to the laboratory in properly marked, well-stoppered sample bottles.
2.4. Aliquoting of faeces and urine
Aliquot of faeces from each animal equal to 1/20th of the total faeces were taken and
dried in weighed Petri dish overnight in a hot air maintained at 100 58C for dry matter
290
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
determination. Similar aliquots were dried in another hot air oven maintained at 408C.
The dried aliquots of all the seven days were pooled and kept for analysis of proximate
principles, except nitrogen, gross energy and other constituents. For the analysis of crude
protein (N 6.25) another aliquot equal to 1/40th of the total faeces voided by each
animal were preserved with dilute sulphuric acid (1 : 4) in wide mouth air tight stoppered
weighed bottles. The aliquots of seven days were composited for each animal in separate
bottles. At the end of collection period, the bottles were weighed. The composited
weighed samples were mixed thoroughly and suitable aliquots were taken for nitrogen
estimation. Duplicate aliquots equal to 1/100th of the total urine excreted daily was taken
in 500-ml Kjeldahl's ¯ask containing 30 ml of concentrated sulphuric acid for nitrogen
determination. Another aliquot equal to 1/100th of total urine excreted daily was pooled
in the brown coloured bottles for seven days. The bottle was air tightened and preserved
in a refrigerator for the estimation of energy.
2.5. Respiration calorimetry
Complete energy balance trials were conducted on individual sheep one after the other,
in an open circuit respiration chamber for small animals. Fasting heat production was
determined after withholding feed for 72 h, but water made available throughout.
2.6. Respiration calorimetry equipment
Respiration calorimetry study was conducted in a simple type of open circuit
calorimeter developed and described by Khan and Joshi (1983) for sheep and goat which
consisted of a wooden chamber with internal dimensions (in metres) 1.5 0.9 1.75
(height). The chamber was maintained at 20±258C with relative humidity of ca. 65%.
Difference in oxygen concentrations of incoming and outgoing air was recorded in a dual
type paramagnetic oxygen analyser (Servmex Taylor, model OAT 184). Carbon dioxide
measurement was conducted by a modi®ed Sonden apparatus with a 100-ml burette.
Measurement of methane was done by an infra-red gas analyser (Analytical Development,
Hoddesdon, England, Model 300). Representative samples of the incoming and
outgoing air from the respiration chamber were collected separately into two Douglas
bags with the help of two sampling air pumps (Charles Austen Pumps, Survey, UK) with ¯ow
rate of 3 l per minute and provided with a bypass arrangement to reduce their ¯ow rate.
2.7. Prechamber handling of animals
The selected animal was weighed in the morning prior to feeding and watering and
kept in a pre-respiration chamber room. After that the animal shifted to respiration
chamber for adaptation followed by recording the respiration calorimetry data for two
consecutive days.
2.8. Feeding
The animals were maintained on the prescribed nutritional regime. Weighed quantities
oat hay and concentrate mixtures were given in the manger attached in the metabolic
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
291
crate kept inside the chamber. Animal inside the chamber was provided with suf®cient
amount of water through water trough kept inside the chamber.
2.9. Measurement of respiratory exchange
After keeping the feeds inside, the chamber was made airtight by closing the door and
blower was started along with the ventilation system of the chamber. The equipment was run
for an hour in order to stabilize the recorder. Observations on gaseous exchange were
recorded for two consecutive days on each animal after adaptation period of three days in
metabolic crate and two days in the respiration chamber. Recording the temperature of dry
and weight bulb, ¯ow rate, volume, atmospheric pressure was done manually. The sample of
outgoing and incoming air from the respiration chamber were collected in Douglas bag
separately with continuous sampling device at 12 hourly intervals. The chamber was opened
after 24 h; the residues of feeds, faeces voided and urine excreted were collected and
measured. Representative samples of feed offered, residue left, faeces and urine were drawn
and preserved for nitrogen and energy estimation. Heat production was calculated as per
Brouwer's (Brouwer, 1965) equation. Samples of feeds, residue and faeces were ground and
analysed for proximate principles as per AOAC (1980) method. NDF and ADF in feeds and
faeces were estimated by using method of Goering and Van Soest (1970). Estimation of Gross
energy of samples was done by Gallenkamp automatic adiabatic bomb calorimeter (CBA 301
series) as per the procedure of Gallenkamp manual. The data were subjected to test of
signi®cance (Snedecor and Cochran, 1967) using the analysis of variance.
3. Results and discussion
It is important to mention that under present situations high concentrate feeding to
ruminants is not possible in India for economical reasons. So, the effects of limitedly
varying roughage and concentrate ratio (R : C) in rations on methane production in
Muzaffarnagari sheep were studied. Small amount of concentrate (100 g) was given for
balancing the ration in respect of protein and energy and as such the ratio of R : C was
92 : 8 instead of 100 : 0 in group I. Groups II and III were containing R : C ratio of
50 : 50 and 30 : 70, respectively. Chemical composition of oat hay and concentrate
mixture is given in Table 1.
3.1. Nutrient intake and digestibility
Intake, digestibility of proximate principles and of cell wall fractions and nutritive
value (NV) of different diets in sheep is presented in Table 2. Dry matter (DM) intake was
signi®cantly (p < 0.05) lower in-group I than other groups. It was 12 units higher in
50R : 50C group and seven points higher in 30R : 70C group than 92R : 8C group. It is
established fact that animal eats for energy, although, it is equally important in ruminants
to satisfy its bulk requirement. Fiber content, palatability and digestibility govern the
intake of feed. Highly digestible feeds is generally not considered physically limited but
physiologically determined and as such depends on nutrient-requirement of animal
292
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
Table 1
Chemical composition of oat hay and concentrate mixture (CM) on DM basis
Organic matter (OM)
Crude protein (CP)
Ether extract (EE)
Neutral detergent ®bre (NDF)
Acid detergent ®bre (ADF)
Acid detergent lignin (ADL)
Hemicellulose
Cellulose
Gross energy (kcal/g)
Oat hay
CM
91.89
9.85
2.61
48.80
40.91
5.85
7.89
35.09
4.09
93.98
11.43
2.05
23.91
5.05
0.32
18.86
4.73
4.17
Table 2
Daily intake of feeds by sheep fed on different roughage and concentrate ratios (R : C) and nutritive value of
rations during metabolic trial (DM basis)
Attribute
Liveweight (kg)
Metabolic weight (kg)
Concentrate mixture intake (g DM/day)
Oat hay intake (g DM/day)**
Total intake (g DM/day)**
DM intake (g /kg W0.75/day)*
DM digestibility (%)
Organic matter intake (g/kg W0.75/day)*
Digestibility (%)
Crude protein intake (g/kg W0.75/day)*
Digestibility (%)*
NDFc intake (g /kg W0.75/day)**
Digestibility (%)*
ADFd intake (g/kg W0.75/day)**
Digestibility (%)
Cellulose intake (g/kg W0.75/day)
Digestibility (%)*
Hemicellulose intake (g/kg W0.75/day)**
Digestibility (%)**
Nutritive value
DCP (%)**
TDN (%)**
DE (Mcal/kg DM)*
MEe (Mcal/kg DM)*
a
Treatmentsa
92R : 8C
50R : 50C
30R : 70C
37.1
15.0
91.6
1004.1
1095.7
73.9
58.0
68 a
62.0
7.3
57.5
34.5
62.0
28.0
52.3
24.0
60.2
6.5
98.2
38.0
15.3
687.2
615.5
1302.3
85.2
60.0
79.2
64.5
9.1
60.5
30.3
60.0
18.7
51.5
16.2
58.8
11.6
73.5
b
ab
b
a
39.1
15.6
870.5
382.9 a
1253.4 b
81.0 b
62.4
75.6 b
66.6
8.86 b
61.5 b
25.4 a
55.5 a
12.2 a
49.8
10.6 b
56.9 a
13.2 c
60.6 a
6.46 b
62.1 b
2.45 b
2.29 b
6.73 c
65.5 c
2.54 b
2.30 b
c
a
a
a
a
b
b
c
c
b
a
b
5.73 a
58.8 a
2.29 a
2.0 a
Mean with different letters differ signi®cantly.
Standard error of means.
c
Neutral detergent ®ber.
d
Acid detergent ®ber.
e
ME estimated from calorimetric trial (Table 3).
*
p < 0.05, **p < 0.01.
b
SEMb
b
b
b
b
b
b
b
b
b
1.78
0.58
29.8
29.82
1.80
0.70
1.92
0.75
0.22
0.49
0.83
0.61
0.67
0.53
0.58
0.58
0.26
3.32
0.05
0.44
0.03
0.03
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
293
(Egan, 1980). The basic urge to consume the feed is the tendency of animal to realise the
genetically determined maximum capacity for growth/production. This genetic capacity
corresponds to the maximum rate at which tissues can utilise the nutrients (Ketelaars and
Tolkamp, 1992). Ali et al. (1979) in Muzaffarnagari sheep observed signi®cantly
(p < 0.01) higher total intake on high concentrate diet. In their studies, DM intake on all
roughage, 25R : 75C, 50R : 50C and 25R : 75C diets were 79, 72, 76 and 73 g/kg W0.75/
day, respectively. The observed higher DM intake on diets II and III than in diet I is also
in agreement with the ®ndings of Sekine et al. (1986); Ketelaars and Tolkamp (1992) and
Santra (1995). The digestibility of DM, OM was similar on all R : C ration but the
digestibility in group I was signi®cantly (p < 0.05) lower than other two groups. However,
although nonsigni®cant, two to four percent improvement in digestibility of diets with 50
and 70% concentrate diets is in agreement with Slabbert et al. (1992) who reported that
DM digestibility was not in¯uenced by the plane of nutrition but linearly reduced as
proportion of roughage in the diet increased from 30 to 80%. Ali et al. (1979) also found
increased digestibility of dry matter with increasing the proportion of concentrate in the
diet of Muzaffarnagari sheep. The digestibility of DM depends on quality of roughage in
question, which in turn depends on the content of protein and energy together with
minerals and vitamins. The oat hay used in present experiment showed better OM
digestibility than 57% reported by Weston (1967) for sheep given wheaten hay with 4.4%
protein. Crude protein digestibility was signi®cantly (p < 0.05) increased with inclusion
of higher proportion of concentrate in diet. Digestibility of protein depends primarily on
protein content of the diet and intake (Sahlu et al., 1993). Ali et al. (1979) reported
nonsigni®cant differences in CP digestibility in same breed of sheep fed different
roughage to concentrate ratio, which may be due to ad libitum feeding regime. The
increase in N digestibility with higher consumption of concentrate in sheep in groups II
and III is supported by similar reports of Weston (1967); Sekine et al. (1986) and Slabbert
et al. (1992).
Higher crude ®ber digestibility on diet I (64.3) than II (62.2) and III (56.6) may be due
to the higher content of ®ber and higher retention time in the rumen due to comparatively
slow rate of passage of digesta which in turn causes reduced DM intake. The observation
is in agreement with Ali et al. (1979) and Santra (1995). Digestibility coef®cient of NDF
was signi®cantly (p < 0.05) lower in-group III than other groups whereas ADF
digestibility did not differ. Hemicellulose digestibility in 50R : 50C and 30R : 70C was
signi®cantly lower than 92R : 8C ration. Cellulose digestibility in-group II did not differ
either from group I or group III. Poore et al. (1990) and Takahashi et al. (1991) also
reported that the digestibility of NDF decreased as the level of concentrate is increased in
the diet. Increasing dietary concentrate and roughage ratio characteristically depressed
the digestion of both hemicellulose and cellulose in the rumen. Hemicellulose
digestibility has been shown to be the most sensitive to increase in C : R ratio in the
diet (Santra, 1995). Wedekind et al. (1986) showed that hemicellulose and ADF
accounted for 35 and 21% variability, respectively, in the ruminal NDF digestion in lambs
fed fescue hay with varying levels of starch. Total GI tract digestion of ADF is not
affected by the dietary concentrate and roughage ratio, whereas ruminal digestion of NDF
and hemicellulose in lambs decreases linearly with increase in proportion of concentrate
in the diet (Kennedy and Bunting, 1992).
294
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
3.2. Nutritive value
Nutritive value in terms of DCP and TDN increased (p < 0.05) with inclusion of
concentrate in the diet. Improvement of 1% in DCP and 7% in TDN was observed on
30R : 70C over 8C : 92R group. Mean DCP intake (kg W0.75/day) was signi®cantly
higher in diet II (5.5) and III (5.45) than in diet I (5.45). TDN intake (kg W0.75/day) was
also higher on diet II (52.9) and III (52.9) than in diet I (42.9). The range of intake is
similar to that found by Graham, (1967) for adult sheep and is higher than that found by
Ranjhan (1977).
3.3. Nitrogen intake and outgo
Data on N metabolism on different roughage concentrate ratio is presented in Table 3.
N intake in groups II and III followed similar trend as DM intake and was signi®cantly
(p < 0.01) higher than in group I. Faecal and urinary energy loss was not different. Total
N balance increased with the increase of concentrate in diet, but not beyond 50%
concentrate in diet. Nitrogen retention depends on its content in the diet, intake and the
availability of fermentable energy in the rumen. On high concentrate diet, the availability
of fermentable energy is more which helps rumen microbes to capture N leading to its
increased utilisation.
3.4. Distribution of energy
Total GE (3594, 4467, 4328 in I, II and III, SEM 124.6), DE (2005, 2644, 2646 in I, II
and III, SEM 64.8), ME (1755, 2366, 2397 in I, II and III, SEM 60.1) intakes in groups II
and III were signi®cantly (p < 0.05) higher than in group I. However, former two groups
did not differ in this respect. Energy evaluation of different R : C ratio-based diets are
described in Table 4. Intake and losses were expressed relative to metabolic body weight
and on this basis except methane energy (signi®cantly less in group III than other two),
neither energy losses nor respective intakes differ signi®cantly. The GE intake is the
Table 3
Nitrogen (N) metabolism in sheep fed on rations with different roughage and concentrate ratios
Particulars
N intake (g/day)**
Faecal N (g/day)
Urinary N (g/day)
N balance (g/day)*
N balance (mg/kg W0.75/day)*
N retained as % of intake*
N retained as % of N absorbed*
a
Treatmentsa
SEMb
92R : 8C
50R : 50C
30R : 70C
17.5 a
7.4
8.6
1.46 a
100 a
8.3 a
14.5 a
22.2 b
8.7
10.8
2.71 b
182 b
12.3 b
20.5 b
21.9 b
8.4
9.43
3.04 b
198 b
13.9 b
22.6 b
Means bearing different letters in a row differ signi®cantly.
Standard error of means.
*
p < 0.05, ** p < 0.01.
b
0.53
0.29
0.30
0.11
5.62
0.25
0.27
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
295
Table 4
Energy metabolism (kcal/kg W0.75/day) of Muzaffarnagari sheep fed diets with varying roughage to concentrate
(R : C) ratio
Attribute
Gross energy intake
Faecal energy
Digestible energy
Urinary energy
Methane energy*
Metabolisable energy
Heat production
RQ (fed)
Energy balance*
kmc
kmgd
Treatmentsa
SEMb
92R : 8C
50R : 50C
30R : 70C
242.2
106.6
135.6
7.35
9.5 b
118.9
95.3
0.94
23.5 a
0.674
0.655
274.2
116.6
162.6
7.97
9.1 b
145.4
100.7
0.93
44.8 b
0.688
0.672
256.5
97.2
154.2
7.0
7.4 a
139.8
92.0
0.91
47.8 b
0.693
0.720
7.45
3.29
5.53
0.20
0.25
5.26
2.43
0.02
3.27
0.003
0.001
a
Mean with different letters in a row differ signi®cantly (p < 0.05).
Standard error of means.
c
km is ef®ciency of utilisation of ME for maintenance (calculated as per ARC, 1980).
d
kmg is ef®ciency of utilisation of ME for maintenance and gain.
b
function of level of energy and energy density of ration besides the function for which it
is used (ARC, 1980).
As percentage of GE, DE (55.9, 59.3, 61.2 for groups I, II and III, SEM 0.85)) was not
different. Percent metabolisability (ME/GE) was higher (p < 0.05) in group III (55.2) than
in group I (48.9), but metabolisability of ration in Group II (55.0) did not vary from either
of these two groups (SEM 0.82). High proportion of oat hay in the diet was conspicuously
re¯ected by signi®cant depression in the ME value obtained in-group I (Table 2). With
increase in the energy intake, there was no signi®cant increase in faecal energy output. As
the level of feeding of ruminants increases the proportional loss of energy in faeces
increases and apparent digestibility declines (ARC, 1980). In the present experiment
sheep were fed at maintenance level, that is why faecal loss may not be different. Blaxter
and Wainman (1964) found increased faecal energy loss with increasing proportion of
¯aked maize in the diet. However, this loss declined markedly when the diet of sheep fed
above maintenance level contained beyond 60±80% of ¯aked maize. Similar observation
was made by Krishna et al. (1969). Wagner and Loosli (1967) also noted slightly
increased values of TDN at maintenance level of feeding with increased percentage of
concentrate in diet.
Urinary energy (UE) loss as a proportion of GE intake on 92R : 8C, 50R : 50C and
30R : 70C was 3.0, 2.9 and 2.8% (SEM 0.008) and is in agreement with earlier reports
(Blaxter and Graham, 1955, 1956; Blaxter and Wainman, 1961). These ®gures were
signi®cantly (p < 0.05) different from each other. It has been found that UE loss was
never more than 5% of GE intake in sheep and cattle (Blaxter and Wainman, 1964). As
the total ME intakes were signi®cantly higher on concentrate containing diet, UE loss as
percent of GE intake was decreased. Kishan et al. (1987) also found that the UE loss as a
percentage of GE intakes decreased on high level of ME intake.
296
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
Table 5
Methane production in sheep fed on rations with varying roughage and concentrate (R : C) ratios
Treatmentsa
Methane production
*
As % of gross energy
As % of digestible energy*
l/day
g/day
g/100 g digestible DM*
g/100 g digestible OM*
g/100 g digestible CHO*
SEMb
92R : 8C
50R : 50C
30R : 70C
3.93
7.02
14.8
10.6
2.10
2.20
2.65
3.34
5.62
15.7
11.2
1.76
1.84
1.88
2.98
4.87
13.6
9.7
1.46
1.54
1.62
b
b
b
b
b
ab
a
a
a
a
a
a
a
a
a
0.12
0.18
0.52
0.37
0.06
0.06
0.08
a
Means bearing different letters in a row differ signi®cantly.
Standard error of means.
*
p < 0.05.
b
Methane production data are presented in Table 5. Total methane (g/day) was not
signi®cantly different between groups. Moss (1994) recommended that units used for
expressing methane production from ruminants should be extended beyond the traditional
expression of methane energy as proportion of GE intake to methane production per kg of
OM digested or per kg of animal product. Thus, the methane loss (per 100 g digestible
OM or CHO) was signi®cantly higher on 92R : 8C than either 50R : 50C or 30R : 70C
ratios, which did not differed in this respect. As the level of digested organic matter or
carbohydrate was increased in the diet, methane production decreased signi®cantly
(p < 0.05). Type of the roughage and also concentrate (starchy or oil cake) can
substantially in¯uence on methane production (Moss and Givens, 1993). In their
experiment, they offered isoenergetic forage (grass silage): concentrate (rolled barley or
soyabean meal) diets of 1.0, 0.75, 0.50 and 0.25 (DM basis ) level in wether sheep at
maintenance level. It was found that the methane production (l kgÿ1 FOM) increase
signi®cantly and linearly with decreasing forage concentrate for rolled Barley diet but
was nonlinear for soyabean meal diets with low levels at forage concentrate ratio of 0.76
and 0.51.
The main component affecting methane production is the type of carbohydrate and
relative rate of fermentation. Kreuzer et al. (1986) found signi®cantly lower methane loss
(total as well as % of GE) on rations with native starch than rations with cellulose.
Johnson et al. (1993) showed that there was decreased methane production with increased
energy intake, when expressed as percent of GE. Methane production do fall from a level
of 6±7% of energy intake when forages are fed at maintenance to as low as 2±3% when
high grain concentrates are fed at near ad libitum intake levels (Johnson and Johnson,
1995). Although fed approximately at maintenance during experimental study, the intakes
were found to be exceeded far towards achieving positive balance. Van Soest (1994)
indicated that a high grain diet and or the little addition of soluble carbohydrate with
resulting shift in the fermentation pattern in the rumen are associated with a more hostile
environment for methanogenic bacteria in which a passage rates are increased, ruminal
pH is lowered and certain population of protozoa, ruminal ciliates and methanogenic
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
297
bacteria may be eliminated or inhibited. In present study also total energy intake was
signi®cantly higher in groups II and III than in I.
The methane losses in this study were lower than some of the earlier reports (Blaxter
and Clapperton, 1965; Moe and Tyrrell, 1980; Khan et al., 1988; Prakash, 1990).
Neergaard (1974) observed 3% methane energy loss as percent of GE in some calves in
his experiment fed on a concentrate diet. Methane energy loss as percent of GE ranged
from 2.5 to 13% in sheep fed on different diets (FEU, 1978). The low values of methane
as compared to others can be attributed to numerous reasons, one of that is breed. Blaxter
and Wainman (1964) studied methane production of a Cheviot, a Suffolk cross weather
and black face weather. When fed on different ratio of oat hay and maize at maintenance
or twice the maintenance level, Cheviot produced higher (p < 0.01) methane with all
ratios of roughage to concentrate than the other two breeds, and this discrepancy was
more marked with concentrate diet. This is indicative of the fact that it is the genetics of
the animal (besides diet) which may be controlling the rumen size and its complex
ecosystem, which can have great impact on methane production. In studies of Blaxter and
Wainman (1964), methane production was found to be reduced from 6.76 to 3.65% of GE
intake. High propionate fermentation (indicative of reduced methane production) are
apparent in hay based diet containing high proportion of concentrate (Thomas and
Chamberlain, 1982) which may be a one more factor in reducing methane loss in such
dietary combination. In studies of Blaxter and Wainman (1964), range examined was
from 100 to 5% hay: 95 ¯aked corn maize. To verify the results obtained in other sheep, it
is necessary to evaluate C : R beyond 70 : 30 in Muzaffarnagari sheep.
As such there is less information to show comparative methane production from
different animal species/breeds. Report from this laboratory indicated that there is
variation in methane production by different breeds. Murarilal et al. (1987) showed
higher methane loss as a percent of GE intake for Holstein-Friesian (HF) Hariana cross
than HF cattle or buffaloes. Recently, we (Chandramoni et al., 1998) reported higher
methane production in crossbred sheep than Muzaffarnagari sheep, when fed same
rations. Galbraith et al. (1998) reported methane losses of 6.6, 5.2 and 3.3% of GE intake
for bison, waipiti and white tailed deer, respectively, when the animals were fed lucern
pellets.
Unadjusted (Table 4) and energy retention (kcal/kg W0.75) adjusted for similar ME
intake (data not shown) was signi®cantly (p < 0.05) higher on groups II and III than in
group I. Ef®ciency of utilisation of ME (km) was calculated using equation of ARC
(1980) which relates km to metabolisability (qm). Assuming that q qm, the calculated
kmvalues were not signi®cantly different from each other (Table 4). Similarly, the
ef®ciency of utilisation of ME for maintenance and gain (kmg) was calculated from the
values of ME intake, energy balance and fasting heat production (FHP) of these sheep
(Chandramoni et al., 1999) as kmg energy balance FHP/ME intake. These values
also did not differ signi®cantly (Table 4). On 92R : 8C ratio, heat production (HP), as a
percent of ME was almost 80% whereas on 50R : 50C and 30R : 70C diets, it was 70 and
66%, respectively. This is indicative of the more energy loss as heat production on high
roughage diet which may be due to wastage of more energy on work of digestion on all
roughage rations as compared to concentrate diet (Armstrong and Blaxter, 1957), which
causes decreased km values.
298
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
4. Conclusion
Based on the evaluation of three roughage concentrate ratio (R : C), this study
concluded that as dry matter intake and digestibility, N retention, reduction in methane
loss (g/100 g digestible OM or CHO) and ME value does not improve and on contrary
NDF digestibility reduces signi®cantly (p < 0.05) beyond 50R : 50C, ratio of 50 : 50 in
diet of Muzaffarnagari sheep is optimum for economical and sustainable sheep
production through reduced methane emissions.
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83 (2000) 287±300
Energy metabolism with particular reference to
methane production in Muzaffarnagari sheep fed
rations varying in roughage to concentrate ratio
Chandramoni*,1, S.B. Jadhao2, C.M. Tiwari3, M.Y. Khan
Energy Metabolism and Respiration Calorimetry Laboratory, Animal Nutrition Division,
Indian Veterinary Research Institute, Izatnagar 243 122, India
Received 28 April 1997; received in revised form 28 January 1999; accepted 11 November 1999
Abstract
Methane production in Muzaffarnagari sheep was studied using open circuit respiration
calorimetry technique. Twelve rams were divided in three treatment groups of four each and were
fed at about maintenance with diets having three roughage (oat hay) to concentrate ratio (R : C), i.e.
92 : 8 (group I), 50 : 50 (group II), 30 : 70 (group III). Concentrate mixture was formulated to
contain 93% crushed maize grain, 3.5% wheat bran and 3.5% groundnut cake forti®ed with
minerals. Whereas the digestibility of dry matter, organic matter and energy was similar, the
digestibility of nitrogen was signi®cantly (p < 0.05) higher in groups II and III and of neutral
detergent ®ber (NDF) was signi®cantly (p < 0.05) lower in-group III than the other groups. Nitrogen
retention was improved but not beyond 50R : 50C. As a percent of gross energy intake, urinary
energy losses in groups I, II and III were 3.0, 2.9 and 2.8%, and methane energy losses were 3.39,
3.34 and 2.98%, respectively. Even though, gross energy intakes (kcal/kg W0.75) were similar,
methane loss (g) per 100 g digestible organic matter was signi®cantly (p < 0.05) higher in group I
(2.2) than in groups II (1.84) and III (1.54), the latter did not differ in this respect. Metabolisable
energy (ME) value (Mcal/kg DM) and energy balances (kcal/kg W0.75) on rations in groups II and
III were similar but on ration in group I was signi®cantly (p < 0.05) lower than that in other groups.
Ef®ciency of utilisation of ME for maintenance (km) of diets in groups I, II and III calculated as per
ARC (1980) were 0.674, 0.688 and 0.693, respectively, and did not differ signi®cantly. Based on the
evaluation of three R : C, it was inferred that R : C ratio of 50 : 50 in diet of Muzaffarnagari sheep is
*
Corresponding author. Fax: 91-226361573.
E-mail address: chandramoni@hotmail.com (Chandramoni).
1
Quarter 11, Road 9, Bihar Veterinary College, Patna 800 014, India.
2
Central Institute of Fisheries Education, Seven Bunglows, Versova, Mumbai 400 061.
3
Deptt. Animal Nutrition, Rajiv Gandhi College of Veterinary and Animal Sciences, Kurumbapeth,
Pondicherry 605 009.
0377-8401/00/$ ± see front matter # 2000 Published by Elsevier Science B.V. All rights reserved.
PII: S 0 3 7 7 - 8 4 0 1 ( 9 9 ) 0 0 1 3 2 - 7
288
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
optimum for economical and sustainable sheep production through reduced methane emissions.
# 2000 Published by Elsevier Science B.V. All rights reserved.
Keywords: Muzaffarnagari sheep; Energy metabolism; Methane production; Roughage: concentrate ratios;
Nutritive value
1. Introduction
Methane production in recent years has assumed more signi®cance in animal
production due to its undesired effect on environment including its contribution to global
warming. There is now considerable general understanding of the contribution of
ruminants to methane production (Johnson and Johnson, 1995; Mathison et al., 1998).
Sustainable animal production systems henceforth require that methane produced should
be less per unit output. Dietary manipulation is one of the means to reduce methane (see
Review, Moss, 1994) and is practicable in India and other third World countries.
Optimum roughage to concentrate is one of the dietary means, which has potential to
reduce methane production. Number of studies concerning supplementation of
concentrate to straw or poor quality roughages has been done which has demonstrated
reduced methane production (Sundstol, 1982; Birkelo et al., 1986; Wainman and Dewey,
1987; Silva and érskov, 1988). However, few studies on supplementation to good quality
hay-based diets near maintenance level has been done. The effects of supplementation of
concentrate to good quality roughages are unclear (Moss, 1994). Blaxter and Wainman
(1964) showed greater production of methane for hay than maize grain and that methane
production with mixtures of these feeds was not additive and was invariably greater than
could be predicted from composition of the rations. In contrast, the ME content and the
ef®ciency of utilisation of metabolisable energy (ME) for maintenance and fattening
increased from 71 to 79 and 29 to 61%, respectively. Hence, it is likely that methane
production per unit of animal product would decline with increasing maize in ration. On
the other hand, Webster (1987) observed reduce energy loss as methane with increasing
the ratio of cereal to forage, which decreases the ratio of acetic to propionic acid. Low
cereal inclusion rate to grass silage-based diets enhances milk yield and increase milk fat
content (Rae et al., 1986) when compared with silage fed alone. Methane produced per
unit animal product in this situation seems to be decreased. High propionate
fermentations (indicative of reduced methane) are common in hay-based diet containing
high proportion of concentrate (Thomas and Chamberlain, 1982).
Therefore, it was thought to investigate methane production on good quality oat haybased rations with different roughage to concentrate ratio. As such the combined effect
(associate effect) of starchy feeds (alone or with little quantity of oil cake and bran)
supplementation to good quality roughage like oat hay is little studied. Similarly, the
literature on energy balance studies in Muzaffarnagari sheep (sturdy and meat type breed)
which is important animal resource in this area, is lacking. Keeping in view the above
facts, the present study was speci®cally designed to study energy metabolism of
Muzaffarnagari sheep with special reference to methane production on diets varying in
roughage and concentrate ratio.
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
289
2. Materials and methods
2.1. Selection of animals
Fifteen healthy male uncastrated, Muzaffarnagari sheep, ca. 6±8 months of age, were
procured in October, 1993 from Central Institute for Research on Goats, Makhdoom. The
animals were dewormed before start of the experiment and good managerial practices
were followed in the sheds. The animals were fed on a balanced ration for about a month
before the start of experimental feeding.
2.2. Housing and management
The animals were housed in a well-ventilated shed having cemented ¯oor with
individual feeding and watering arrangement throughout the experimental period.
Concentrate mixture was offered to individual animals between 9.00 and 10.00 a.m. and
oat hay was offered in after noon at 3.00 a.m. in the same manger. Fresh water was
provided ad libitum daily at 2.30 p.m.
2.3. Experimental plan
Twelve sheep were randomly allocated on the basis of liveweight to three groups
comprising each of four animals, following complete randomized design. Rations were
given to meet the maintenance requirement of sheep as per NRC (1985). Group I was
offered oat hay plus minimal concentrate mixture (ca. 100 g of concentrate mixture I) to
meet CP requirement. Group II was offered 50% concentrate mixture and 50% oat hay,
whereas group III animals were offered 70% concentrate mixture and 30% oat hay only.
Concentrate mixture (11.7% CP and 4.37 Mcal GE/kg DM) contained 93% crushed
maize grain, 3.5% deoiled groundnut cake, and 3.5% wheat bran. To every 100 kg
concentrate mixture 2 kg of mineral mixture (contained moisture maximum 5%, Calcium
minimum 28%, Phosphorus minimum. 12%, Iodine as KI 0.026±0.130%, Copper 0.077±
0.130%, ¯uorine maximum 0.04%) and 1 kg of common salt were added.
Metabolism trial was conducted after 60 days of feeding (with three days of
adaptation) in metabolic cages for seven days. All the animals were weighed before and
after the trial. Representative samples of feed offered and residue left were taken daily for
dry matter estimation and analysis. The total amount of faeces voided by each animal was
collected quantitatively at 9 a.m. daily. It was weighed and representative sample of each
animal was drawn in sample bottles after crushing and mixing all the pellets and was
brought to the laboratory for analysis. Urine excreted daily by individual animal was
collected separately in bucket with dilute sulphuric acid for 24 h and was measured with
the help of measuring cylinder. Representative samples from individual animals were
brought to the laboratory in properly marked, well-stoppered sample bottles.
2.4. Aliquoting of faeces and urine
Aliquot of faeces from each animal equal to 1/20th of the total faeces were taken and
dried in weighed Petri dish overnight in a hot air maintained at 100 58C for dry matter
290
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
determination. Similar aliquots were dried in another hot air oven maintained at 408C.
The dried aliquots of all the seven days were pooled and kept for analysis of proximate
principles, except nitrogen, gross energy and other constituents. For the analysis of crude
protein (N 6.25) another aliquot equal to 1/40th of the total faeces voided by each
animal were preserved with dilute sulphuric acid (1 : 4) in wide mouth air tight stoppered
weighed bottles. The aliquots of seven days were composited for each animal in separate
bottles. At the end of collection period, the bottles were weighed. The composited
weighed samples were mixed thoroughly and suitable aliquots were taken for nitrogen
estimation. Duplicate aliquots equal to 1/100th of the total urine excreted daily was taken
in 500-ml Kjeldahl's ¯ask containing 30 ml of concentrated sulphuric acid for nitrogen
determination. Another aliquot equal to 1/100th of total urine excreted daily was pooled
in the brown coloured bottles for seven days. The bottle was air tightened and preserved
in a refrigerator for the estimation of energy.
2.5. Respiration calorimetry
Complete energy balance trials were conducted on individual sheep one after the other,
in an open circuit respiration chamber for small animals. Fasting heat production was
determined after withholding feed for 72 h, but water made available throughout.
2.6. Respiration calorimetry equipment
Respiration calorimetry study was conducted in a simple type of open circuit
calorimeter developed and described by Khan and Joshi (1983) for sheep and goat which
consisted of a wooden chamber with internal dimensions (in metres) 1.5 0.9 1.75
(height). The chamber was maintained at 20±258C with relative humidity of ca. 65%.
Difference in oxygen concentrations of incoming and outgoing air was recorded in a dual
type paramagnetic oxygen analyser (Servmex Taylor, model OAT 184). Carbon dioxide
measurement was conducted by a modi®ed Sonden apparatus with a 100-ml burette.
Measurement of methane was done by an infra-red gas analyser (Analytical Development,
Hoddesdon, England, Model 300). Representative samples of the incoming and
outgoing air from the respiration chamber were collected separately into two Douglas
bags with the help of two sampling air pumps (Charles Austen Pumps, Survey, UK) with ¯ow
rate of 3 l per minute and provided with a bypass arrangement to reduce their ¯ow rate.
2.7. Prechamber handling of animals
The selected animal was weighed in the morning prior to feeding and watering and
kept in a pre-respiration chamber room. After that the animal shifted to respiration
chamber for adaptation followed by recording the respiration calorimetry data for two
consecutive days.
2.8. Feeding
The animals were maintained on the prescribed nutritional regime. Weighed quantities
oat hay and concentrate mixtures were given in the manger attached in the metabolic
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
291
crate kept inside the chamber. Animal inside the chamber was provided with suf®cient
amount of water through water trough kept inside the chamber.
2.9. Measurement of respiratory exchange
After keeping the feeds inside, the chamber was made airtight by closing the door and
blower was started along with the ventilation system of the chamber. The equipment was run
for an hour in order to stabilize the recorder. Observations on gaseous exchange were
recorded for two consecutive days on each animal after adaptation period of three days in
metabolic crate and two days in the respiration chamber. Recording the temperature of dry
and weight bulb, ¯ow rate, volume, atmospheric pressure was done manually. The sample of
outgoing and incoming air from the respiration chamber were collected in Douglas bag
separately with continuous sampling device at 12 hourly intervals. The chamber was opened
after 24 h; the residues of feeds, faeces voided and urine excreted were collected and
measured. Representative samples of feed offered, residue left, faeces and urine were drawn
and preserved for nitrogen and energy estimation. Heat production was calculated as per
Brouwer's (Brouwer, 1965) equation. Samples of feeds, residue and faeces were ground and
analysed for proximate principles as per AOAC (1980) method. NDF and ADF in feeds and
faeces were estimated by using method of Goering and Van Soest (1970). Estimation of Gross
energy of samples was done by Gallenkamp automatic adiabatic bomb calorimeter (CBA 301
series) as per the procedure of Gallenkamp manual. The data were subjected to test of
signi®cance (Snedecor and Cochran, 1967) using the analysis of variance.
3. Results and discussion
It is important to mention that under present situations high concentrate feeding to
ruminants is not possible in India for economical reasons. So, the effects of limitedly
varying roughage and concentrate ratio (R : C) in rations on methane production in
Muzaffarnagari sheep were studied. Small amount of concentrate (100 g) was given for
balancing the ration in respect of protein and energy and as such the ratio of R : C was
92 : 8 instead of 100 : 0 in group I. Groups II and III were containing R : C ratio of
50 : 50 and 30 : 70, respectively. Chemical composition of oat hay and concentrate
mixture is given in Table 1.
3.1. Nutrient intake and digestibility
Intake, digestibility of proximate principles and of cell wall fractions and nutritive
value (NV) of different diets in sheep is presented in Table 2. Dry matter (DM) intake was
signi®cantly (p < 0.05) lower in-group I than other groups. It was 12 units higher in
50R : 50C group and seven points higher in 30R : 70C group than 92R : 8C group. It is
established fact that animal eats for energy, although, it is equally important in ruminants
to satisfy its bulk requirement. Fiber content, palatability and digestibility govern the
intake of feed. Highly digestible feeds is generally not considered physically limited but
physiologically determined and as such depends on nutrient-requirement of animal
292
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
Table 1
Chemical composition of oat hay and concentrate mixture (CM) on DM basis
Organic matter (OM)
Crude protein (CP)
Ether extract (EE)
Neutral detergent ®bre (NDF)
Acid detergent ®bre (ADF)
Acid detergent lignin (ADL)
Hemicellulose
Cellulose
Gross energy (kcal/g)
Oat hay
CM
91.89
9.85
2.61
48.80
40.91
5.85
7.89
35.09
4.09
93.98
11.43
2.05
23.91
5.05
0.32
18.86
4.73
4.17
Table 2
Daily intake of feeds by sheep fed on different roughage and concentrate ratios (R : C) and nutritive value of
rations during metabolic trial (DM basis)
Attribute
Liveweight (kg)
Metabolic weight (kg)
Concentrate mixture intake (g DM/day)
Oat hay intake (g DM/day)**
Total intake (g DM/day)**
DM intake (g /kg W0.75/day)*
DM digestibility (%)
Organic matter intake (g/kg W0.75/day)*
Digestibility (%)
Crude protein intake (g/kg W0.75/day)*
Digestibility (%)*
NDFc intake (g /kg W0.75/day)**
Digestibility (%)*
ADFd intake (g/kg W0.75/day)**
Digestibility (%)
Cellulose intake (g/kg W0.75/day)
Digestibility (%)*
Hemicellulose intake (g/kg W0.75/day)**
Digestibility (%)**
Nutritive value
DCP (%)**
TDN (%)**
DE (Mcal/kg DM)*
MEe (Mcal/kg DM)*
a
Treatmentsa
92R : 8C
50R : 50C
30R : 70C
37.1
15.0
91.6
1004.1
1095.7
73.9
58.0
68 a
62.0
7.3
57.5
34.5
62.0
28.0
52.3
24.0
60.2
6.5
98.2
38.0
15.3
687.2
615.5
1302.3
85.2
60.0
79.2
64.5
9.1
60.5
30.3
60.0
18.7
51.5
16.2
58.8
11.6
73.5
b
ab
b
a
39.1
15.6
870.5
382.9 a
1253.4 b
81.0 b
62.4
75.6 b
66.6
8.86 b
61.5 b
25.4 a
55.5 a
12.2 a
49.8
10.6 b
56.9 a
13.2 c
60.6 a
6.46 b
62.1 b
2.45 b
2.29 b
6.73 c
65.5 c
2.54 b
2.30 b
c
a
a
a
a
b
b
c
c
b
a
b
5.73 a
58.8 a
2.29 a
2.0 a
Mean with different letters differ signi®cantly.
Standard error of means.
c
Neutral detergent ®ber.
d
Acid detergent ®ber.
e
ME estimated from calorimetric trial (Table 3).
*
p < 0.05, **p < 0.01.
b
SEMb
b
b
b
b
b
b
b
b
b
1.78
0.58
29.8
29.82
1.80
0.70
1.92
0.75
0.22
0.49
0.83
0.61
0.67
0.53
0.58
0.58
0.26
3.32
0.05
0.44
0.03
0.03
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
293
(Egan, 1980). The basic urge to consume the feed is the tendency of animal to realise the
genetically determined maximum capacity for growth/production. This genetic capacity
corresponds to the maximum rate at which tissues can utilise the nutrients (Ketelaars and
Tolkamp, 1992). Ali et al. (1979) in Muzaffarnagari sheep observed signi®cantly
(p < 0.01) higher total intake on high concentrate diet. In their studies, DM intake on all
roughage, 25R : 75C, 50R : 50C and 25R : 75C diets were 79, 72, 76 and 73 g/kg W0.75/
day, respectively. The observed higher DM intake on diets II and III than in diet I is also
in agreement with the ®ndings of Sekine et al. (1986); Ketelaars and Tolkamp (1992) and
Santra (1995). The digestibility of DM, OM was similar on all R : C ration but the
digestibility in group I was signi®cantly (p < 0.05) lower than other two groups. However,
although nonsigni®cant, two to four percent improvement in digestibility of diets with 50
and 70% concentrate diets is in agreement with Slabbert et al. (1992) who reported that
DM digestibility was not in¯uenced by the plane of nutrition but linearly reduced as
proportion of roughage in the diet increased from 30 to 80%. Ali et al. (1979) also found
increased digestibility of dry matter with increasing the proportion of concentrate in the
diet of Muzaffarnagari sheep. The digestibility of DM depends on quality of roughage in
question, which in turn depends on the content of protein and energy together with
minerals and vitamins. The oat hay used in present experiment showed better OM
digestibility than 57% reported by Weston (1967) for sheep given wheaten hay with 4.4%
protein. Crude protein digestibility was signi®cantly (p < 0.05) increased with inclusion
of higher proportion of concentrate in diet. Digestibility of protein depends primarily on
protein content of the diet and intake (Sahlu et al., 1993). Ali et al. (1979) reported
nonsigni®cant differences in CP digestibility in same breed of sheep fed different
roughage to concentrate ratio, which may be due to ad libitum feeding regime. The
increase in N digestibility with higher consumption of concentrate in sheep in groups II
and III is supported by similar reports of Weston (1967); Sekine et al. (1986) and Slabbert
et al. (1992).
Higher crude ®ber digestibility on diet I (64.3) than II (62.2) and III (56.6) may be due
to the higher content of ®ber and higher retention time in the rumen due to comparatively
slow rate of passage of digesta which in turn causes reduced DM intake. The observation
is in agreement with Ali et al. (1979) and Santra (1995). Digestibility coef®cient of NDF
was signi®cantly (p < 0.05) lower in-group III than other groups whereas ADF
digestibility did not differ. Hemicellulose digestibility in 50R : 50C and 30R : 70C was
signi®cantly lower than 92R : 8C ration. Cellulose digestibility in-group II did not differ
either from group I or group III. Poore et al. (1990) and Takahashi et al. (1991) also
reported that the digestibility of NDF decreased as the level of concentrate is increased in
the diet. Increasing dietary concentrate and roughage ratio characteristically depressed
the digestion of both hemicellulose and cellulose in the rumen. Hemicellulose
digestibility has been shown to be the most sensitive to increase in C : R ratio in the
diet (Santra, 1995). Wedekind et al. (1986) showed that hemicellulose and ADF
accounted for 35 and 21% variability, respectively, in the ruminal NDF digestion in lambs
fed fescue hay with varying levels of starch. Total GI tract digestion of ADF is not
affected by the dietary concentrate and roughage ratio, whereas ruminal digestion of NDF
and hemicellulose in lambs decreases linearly with increase in proportion of concentrate
in the diet (Kennedy and Bunting, 1992).
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Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
3.2. Nutritive value
Nutritive value in terms of DCP and TDN increased (p < 0.05) with inclusion of
concentrate in the diet. Improvement of 1% in DCP and 7% in TDN was observed on
30R : 70C over 8C : 92R group. Mean DCP intake (kg W0.75/day) was signi®cantly
higher in diet II (5.5) and III (5.45) than in diet I (5.45). TDN intake (kg W0.75/day) was
also higher on diet II (52.9) and III (52.9) than in diet I (42.9). The range of intake is
similar to that found by Graham, (1967) for adult sheep and is higher than that found by
Ranjhan (1977).
3.3. Nitrogen intake and outgo
Data on N metabolism on different roughage concentrate ratio is presented in Table 3.
N intake in groups II and III followed similar trend as DM intake and was signi®cantly
(p < 0.01) higher than in group I. Faecal and urinary energy loss was not different. Total
N balance increased with the increase of concentrate in diet, but not beyond 50%
concentrate in diet. Nitrogen retention depends on its content in the diet, intake and the
availability of fermentable energy in the rumen. On high concentrate diet, the availability
of fermentable energy is more which helps rumen microbes to capture N leading to its
increased utilisation.
3.4. Distribution of energy
Total GE (3594, 4467, 4328 in I, II and III, SEM 124.6), DE (2005, 2644, 2646 in I, II
and III, SEM 64.8), ME (1755, 2366, 2397 in I, II and III, SEM 60.1) intakes in groups II
and III were signi®cantly (p < 0.05) higher than in group I. However, former two groups
did not differ in this respect. Energy evaluation of different R : C ratio-based diets are
described in Table 4. Intake and losses were expressed relative to metabolic body weight
and on this basis except methane energy (signi®cantly less in group III than other two),
neither energy losses nor respective intakes differ signi®cantly. The GE intake is the
Table 3
Nitrogen (N) metabolism in sheep fed on rations with different roughage and concentrate ratios
Particulars
N intake (g/day)**
Faecal N (g/day)
Urinary N (g/day)
N balance (g/day)*
N balance (mg/kg W0.75/day)*
N retained as % of intake*
N retained as % of N absorbed*
a
Treatmentsa
SEMb
92R : 8C
50R : 50C
30R : 70C
17.5 a
7.4
8.6
1.46 a
100 a
8.3 a
14.5 a
22.2 b
8.7
10.8
2.71 b
182 b
12.3 b
20.5 b
21.9 b
8.4
9.43
3.04 b
198 b
13.9 b
22.6 b
Means bearing different letters in a row differ signi®cantly.
Standard error of means.
*
p < 0.05, ** p < 0.01.
b
0.53
0.29
0.30
0.11
5.62
0.25
0.27
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
295
Table 4
Energy metabolism (kcal/kg W0.75/day) of Muzaffarnagari sheep fed diets with varying roughage to concentrate
(R : C) ratio
Attribute
Gross energy intake
Faecal energy
Digestible energy
Urinary energy
Methane energy*
Metabolisable energy
Heat production
RQ (fed)
Energy balance*
kmc
kmgd
Treatmentsa
SEMb
92R : 8C
50R : 50C
30R : 70C
242.2
106.6
135.6
7.35
9.5 b
118.9
95.3
0.94
23.5 a
0.674
0.655
274.2
116.6
162.6
7.97
9.1 b
145.4
100.7
0.93
44.8 b
0.688
0.672
256.5
97.2
154.2
7.0
7.4 a
139.8
92.0
0.91
47.8 b
0.693
0.720
7.45
3.29
5.53
0.20
0.25
5.26
2.43
0.02
3.27
0.003
0.001
a
Mean with different letters in a row differ signi®cantly (p < 0.05).
Standard error of means.
c
km is ef®ciency of utilisation of ME for maintenance (calculated as per ARC, 1980).
d
kmg is ef®ciency of utilisation of ME for maintenance and gain.
b
function of level of energy and energy density of ration besides the function for which it
is used (ARC, 1980).
As percentage of GE, DE (55.9, 59.3, 61.2 for groups I, II and III, SEM 0.85)) was not
different. Percent metabolisability (ME/GE) was higher (p < 0.05) in group III (55.2) than
in group I (48.9), but metabolisability of ration in Group II (55.0) did not vary from either
of these two groups (SEM 0.82). High proportion of oat hay in the diet was conspicuously
re¯ected by signi®cant depression in the ME value obtained in-group I (Table 2). With
increase in the energy intake, there was no signi®cant increase in faecal energy output. As
the level of feeding of ruminants increases the proportional loss of energy in faeces
increases and apparent digestibility declines (ARC, 1980). In the present experiment
sheep were fed at maintenance level, that is why faecal loss may not be different. Blaxter
and Wainman (1964) found increased faecal energy loss with increasing proportion of
¯aked maize in the diet. However, this loss declined markedly when the diet of sheep fed
above maintenance level contained beyond 60±80% of ¯aked maize. Similar observation
was made by Krishna et al. (1969). Wagner and Loosli (1967) also noted slightly
increased values of TDN at maintenance level of feeding with increased percentage of
concentrate in diet.
Urinary energy (UE) loss as a proportion of GE intake on 92R : 8C, 50R : 50C and
30R : 70C was 3.0, 2.9 and 2.8% (SEM 0.008) and is in agreement with earlier reports
(Blaxter and Graham, 1955, 1956; Blaxter and Wainman, 1961). These ®gures were
signi®cantly (p < 0.05) different from each other. It has been found that UE loss was
never more than 5% of GE intake in sheep and cattle (Blaxter and Wainman, 1964). As
the total ME intakes were signi®cantly higher on concentrate containing diet, UE loss as
percent of GE intake was decreased. Kishan et al. (1987) also found that the UE loss as a
percentage of GE intakes decreased on high level of ME intake.
296
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
Table 5
Methane production in sheep fed on rations with varying roughage and concentrate (R : C) ratios
Treatmentsa
Methane production
*
As % of gross energy
As % of digestible energy*
l/day
g/day
g/100 g digestible DM*
g/100 g digestible OM*
g/100 g digestible CHO*
SEMb
92R : 8C
50R : 50C
30R : 70C
3.93
7.02
14.8
10.6
2.10
2.20
2.65
3.34
5.62
15.7
11.2
1.76
1.84
1.88
2.98
4.87
13.6
9.7
1.46
1.54
1.62
b
b
b
b
b
ab
a
a
a
a
a
a
a
a
a
0.12
0.18
0.52
0.37
0.06
0.06
0.08
a
Means bearing different letters in a row differ signi®cantly.
Standard error of means.
*
p < 0.05.
b
Methane production data are presented in Table 5. Total methane (g/day) was not
signi®cantly different between groups. Moss (1994) recommended that units used for
expressing methane production from ruminants should be extended beyond the traditional
expression of methane energy as proportion of GE intake to methane production per kg of
OM digested or per kg of animal product. Thus, the methane loss (per 100 g digestible
OM or CHO) was signi®cantly higher on 92R : 8C than either 50R : 50C or 30R : 70C
ratios, which did not differed in this respect. As the level of digested organic matter or
carbohydrate was increased in the diet, methane production decreased signi®cantly
(p < 0.05). Type of the roughage and also concentrate (starchy or oil cake) can
substantially in¯uence on methane production (Moss and Givens, 1993). In their
experiment, they offered isoenergetic forage (grass silage): concentrate (rolled barley or
soyabean meal) diets of 1.0, 0.75, 0.50 and 0.25 (DM basis ) level in wether sheep at
maintenance level. It was found that the methane production (l kgÿ1 FOM) increase
signi®cantly and linearly with decreasing forage concentrate for rolled Barley diet but
was nonlinear for soyabean meal diets with low levels at forage concentrate ratio of 0.76
and 0.51.
The main component affecting methane production is the type of carbohydrate and
relative rate of fermentation. Kreuzer et al. (1986) found signi®cantly lower methane loss
(total as well as % of GE) on rations with native starch than rations with cellulose.
Johnson et al. (1993) showed that there was decreased methane production with increased
energy intake, when expressed as percent of GE. Methane production do fall from a level
of 6±7% of energy intake when forages are fed at maintenance to as low as 2±3% when
high grain concentrates are fed at near ad libitum intake levels (Johnson and Johnson,
1995). Although fed approximately at maintenance during experimental study, the intakes
were found to be exceeded far towards achieving positive balance. Van Soest (1994)
indicated that a high grain diet and or the little addition of soluble carbohydrate with
resulting shift in the fermentation pattern in the rumen are associated with a more hostile
environment for methanogenic bacteria in which a passage rates are increased, ruminal
pH is lowered and certain population of protozoa, ruminal ciliates and methanogenic
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
297
bacteria may be eliminated or inhibited. In present study also total energy intake was
signi®cantly higher in groups II and III than in I.
The methane losses in this study were lower than some of the earlier reports (Blaxter
and Clapperton, 1965; Moe and Tyrrell, 1980; Khan et al., 1988; Prakash, 1990).
Neergaard (1974) observed 3% methane energy loss as percent of GE in some calves in
his experiment fed on a concentrate diet. Methane energy loss as percent of GE ranged
from 2.5 to 13% in sheep fed on different diets (FEU, 1978). The low values of methane
as compared to others can be attributed to numerous reasons, one of that is breed. Blaxter
and Wainman (1964) studied methane production of a Cheviot, a Suffolk cross weather
and black face weather. When fed on different ratio of oat hay and maize at maintenance
or twice the maintenance level, Cheviot produced higher (p < 0.01) methane with all
ratios of roughage to concentrate than the other two breeds, and this discrepancy was
more marked with concentrate diet. This is indicative of the fact that it is the genetics of
the animal (besides diet) which may be controlling the rumen size and its complex
ecosystem, which can have great impact on methane production. In studies of Blaxter and
Wainman (1964), methane production was found to be reduced from 6.76 to 3.65% of GE
intake. High propionate fermentation (indicative of reduced methane production) are
apparent in hay based diet containing high proportion of concentrate (Thomas and
Chamberlain, 1982) which may be a one more factor in reducing methane loss in such
dietary combination. In studies of Blaxter and Wainman (1964), range examined was
from 100 to 5% hay: 95 ¯aked corn maize. To verify the results obtained in other sheep, it
is necessary to evaluate C : R beyond 70 : 30 in Muzaffarnagari sheep.
As such there is less information to show comparative methane production from
different animal species/breeds. Report from this laboratory indicated that there is
variation in methane production by different breeds. Murarilal et al. (1987) showed
higher methane loss as a percent of GE intake for Holstein-Friesian (HF) Hariana cross
than HF cattle or buffaloes. Recently, we (Chandramoni et al., 1998) reported higher
methane production in crossbred sheep than Muzaffarnagari sheep, when fed same
rations. Galbraith et al. (1998) reported methane losses of 6.6, 5.2 and 3.3% of GE intake
for bison, waipiti and white tailed deer, respectively, when the animals were fed lucern
pellets.
Unadjusted (Table 4) and energy retention (kcal/kg W0.75) adjusted for similar ME
intake (data not shown) was signi®cantly (p < 0.05) higher on groups II and III than in
group I. Ef®ciency of utilisation of ME (km) was calculated using equation of ARC
(1980) which relates km to metabolisability (qm). Assuming that q qm, the calculated
kmvalues were not signi®cantly different from each other (Table 4). Similarly, the
ef®ciency of utilisation of ME for maintenance and gain (kmg) was calculated from the
values of ME intake, energy balance and fasting heat production (FHP) of these sheep
(Chandramoni et al., 1999) as kmg energy balance FHP/ME intake. These values
also did not differ signi®cantly (Table 4). On 92R : 8C ratio, heat production (HP), as a
percent of ME was almost 80% whereas on 50R : 50C and 30R : 70C diets, it was 70 and
66%, respectively. This is indicative of the more energy loss as heat production on high
roughage diet which may be due to wastage of more energy on work of digestion on all
roughage rations as compared to concentrate diet (Armstrong and Blaxter, 1957), which
causes decreased km values.
298
Chandramoni et al. / Animal Feed Science and Technology 83 (2000) 287±300
4. Conclusion
Based on the evaluation of three roughage concentrate ratio (R : C), this study
concluded that as dry matter intake and digestibility, N retention, reduction in methane
loss (g/100 g digestible OM or CHO) and ME value does not improve and on contrary
NDF digestibility reduces signi®cantly (p < 0.05) beyond 50R : 50C, ratio of 50 : 50 in
diet of Muzaffarnagari sheep is optimum for economical and sustainable sheep
production through reduced methane emissions.
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