Directory UMM :Data Elmu:jurnal:A:Animal Feed Science and Technology:Vol88.Issue1-2.Nov2000:
Animal Feed Science and Technology
88 (2000) 103±111
Short communication
Nutritive evaluation of some nitrogen and
non-nitrogen ®xing multipurpose tree species
D.B.V. Ramanaa,*, Sultan Singhb, K.R. Solankia, A.S. Negib
a
National Research Centre for Agroforestry, Near Pahuj Dam, Gwalior Road, Jhansi 284003, U.P., India
b
PAR Division, Indian Grassland & Fodder Research Institute, Jhansi 284003, U.P., India
Received 6 October 1999; received in revised form 3 February 2000; accepted 10 August 2000
Abstract
The nutritive value of leaf materials from ®ve each of nitrogen (NFT), namely, Acacia nilotica,
Albizia lebbeck, Butea monosperma, Leucaena leucocephala, Pongamia pinnata and non-nitrogen
®xing multipurpose tree (non-NFT) species, namely, Anogeissus pendula, Azadirachta indica, Ficus
bengalensis, Terminalia arjuna and Syzygium cumini grown at National Research Centre for
Agroforestry, were evaluated by chemical and in sacco methods. Mean organic matter (OM)
concentration (g kgÿ1 dry matter) was comparable between NFT (902 5:3) and non-NFT
(916 6:5). However, signi®cantly (P < 0:05) higher mean crude protein (CP) contents
(g kgÿ1 DM) were recorded in NFT (174 10:7) than non-NFT (96 1:5) with maximum in
L. leucocephala (228) and minimum in F. bengalensis (88), respectively. Mean neutral detergent
®bre (NDF) contents were relatively lower in NFT than in non-NFT. Signi®cantly (P < 0:05) higher
contents of NDF, acid detergent ®bre (ADF) and lignin (g kgÿ1 DM) were observed in F.
bengalensis (597, 458 and 222) and S. cumini (566, 483 and 279) than in any of the other trees.
Total phenolics (TP) concentration was signi®cantly (P < 0:05) lower in NFT except for A. nilotica,
but higher (P < 0:05) in non-NFT except in F. bengalensis. Mean condensed tannins (CT)
concentration differed signi®cantly (P < 0:05) between NFT and non-NFT and ranged from 17 to
31 and 14 to 94 g kgÿ1 DM, respectively. In sacco degradability of DM differed signi®cantly
(P < 0:05) amongst the MPTs and ranged from 504 to 809 for NFT and 513 to 881 g kgÿ1 DM for
non-NFT, respectively. Mean CP degradability (g kgÿ1 DM) was similar for (P < 0:05) NFT
(659 24:1) and non-NFT (661 62:5). Cell wall contents degradability was relatively higher in
non-NFT than NFT. However, a poor correlation was observed between anti-nutritional characters
and in sacco degradability, whereas strong negative correlations were recorded between DM
degradability and NDF, ADF and cellulose contents. The NFT contained high CP and low cell wall
contents and anti-nutritional characters, however the in sacco degradability of nutrients was less for
non-NFT. Nutritive value of the trees indicated that all the species could potentially be used as
*
Corresponding author. Tel.: 91-517-448213.
E-mail address: [email protected] (D.B.V. Ramana).
0377-8401/00/$ ± see front matter # 2000 Elsevier Science B.V. All rights reserved.
PII: S 0 3 7 7 - 8 4 0 1 ( 0 0 ) 0 0 2 0 0 - 5
104
D.B.V. Ramana et al. / Animal Feed Science and Technology 88 (2000) 103±111
protein banks to supplement grass or crop residues in dry seasons. # 2000 Elsevier Science B.V.
All rights reserved.
Keywords: Tree leaves; Nutritive value; In sacco degradability; Condensed tannins; Total phenolics
1. Introduction
Trees and shrubs have been used for generations as multipurpose resources (food,
fodder, timber, wood, ®bre and live fences) across all of the agro-ecological zones of the
world (Smith, 1992). Cutting of tree leaves or branches for animal feed is recorded as
early as Roman times. In countries like India, experiencing a monsoonal climate, trees
and fodder shrubs may be the only living feed resources available during the driest
months when shallow rooted grasses and herbs die off (Chen et al., 1992). Further,
sustained and high population growth rates, combined with limited and rapidly
diminishing land holdings and land for food and forage production, have created a
need to intensify the research for new resources. Intensi®cation, in the context of
livestock production systems, means a broadening of the feed resource base for both
quantity and quality. One of the alternative feed resources is the use of multipurpose tree
species for top fodder production (NCA, 1976). Much work has been done world wide on
the evaluation and use of leguminous fodder trees and shrubs, while the studies with nonnitrogen ®xing multipurpose tree species are quite obscure. Under certain soil and
climatic conditions, leguminous tree species do not perform well, while the non-nitrogen
®xing trees, which are quite hardy, can be ®tted in the agroforestry systems and may be
exploited for livestock feeding.
Since feed or fodder value is indicated by its nutrient content and digestibility, in this
study an attempt has been made to document the nutritive content and in sacco
degradability of some leguminous and non-nitrogen ®xing tree leaf materials.
2. Materials and methods
Five each of nitrogen (NFT) and non-nitrogen ®xing multipurpose tree (non-NFT)
species (Table 1) were grown at the National Research Centre for Agroforestry (788350 E
longitude and 258260 N latitude and about 275 m above sea level). Annual long-term
rainfall for the site is estimated to 906 mm falling predominantly from June to September.
Temperature rises to a maximum 47.48C in May and falls to a minimum of 1.08C in
January. Evaporation is highest (441 mm) in May and lowest (55.8 mm) in January. The
soils are Red gravelly with stone pebbles and are generally of relatively low fertility (pH
7.2±7.6) and de®cient in nitrogen and phosphorus (Rai et al., 1997). The area's natural
vegetation consists of Chrysopogon fulvus, Dichianthium annulatum, Iseilema laxum and
Themeda quadrivalves dominated by Sehima nervosum and Heteropogon contortus for
grasses as understory vegetation (Rai et al., 1995).
Botanical name
OMa
Nitrogen ®xing multipurpose tree species
Acacia nilotica
897.5
Albizia lebbeck
887.3
Butea monosperma
889.2
Leucaena leucocephala 930.3
Pongamia pinnata
908.1
Mean
902.5 5.3
CPb
NDFc
ADFd
Cellulose
Hemicellulose
Lignin
Total
phenolics
Condensed
tannins
155.5
142.3
150.2
227.8
193.6
261.4
433.4
506.1
293.6
525.9
166.4
312.6
356.0
151.4
308.5
99.4
188.1
151.2
78.2
136.6
94.9
120.7
150.2
142.2
217.5
95.6
174.0
130.7
76.6
130.6
155.9
12.2
6.5
18.6
6.6
26.0
16.8
20.8
23.5
30.6
173.9 10.7 404.1 36.11 259.0 27.84 130.7 12.9 145.1 13.65 121.5 11.16 40.0 19.45 23.5 1.63
Non-nitrogen ®xing multipurpose tree species
Anogeissus pendula
945.8
100.1
Azadirachta indica
894.2
97.2
Ficus bengalensis
930.8
88.0
Terminalia arjuna
898.9
93.3
Syzygium cumini
911.8
100.0
Mean
a
916.3 6.5
OM: organic matter.
CP: crude protein.
c
NDF: neutral detergent ®bre.
d
ADF: acid detergent ®bre.
b
95.7 1.5
291.3
380.1
597.0
360.4
565.7
192.9
269.6
457.9
267.6
482.7
65.4
121.2
248.0
169.8
283.3
98.4
110.5
139.2
92.9
83.0
438.9 40.13 334.1 38.28 177.5 26.7 104.8 6.45
63.0
102.0
221.8
167.2
279.4
166.7 26.1
73.2
65.3
14.0
159.0
24.9
19.9
93.8
77.6
52.0
14.0
66.1 16.38 51.5 10.4
D.B.V. Ramana et al. / Animal Feed Science and Technology 88 (2000) 103±111
Table 1
Chemical composition of nitrogen and non-nitrogen ®xing multipurpose tree species (g kgÿ1 DM)
105
106
D.B.V. Ramana et al. / Animal Feed Science and Technology 88 (2000) 103±111
Nursery-raised seedlings, approximately 4 months old were planted in July 1988 at a
spacing of 8 m between rows and 4 m between plants within rows. In inter-spaces of tree
rows a mixture of Cenchrus ciliaris (2 kg seed/ha) and Stylosanthes hamata (3 kg seed/
ha) were sown by broadcasting after one operation by the cultivator during August 1988.
At the onset of the monsoon, 40 kg N 20 kg P2O5 per hectare was applied uniformly to
enhance the productivity of the natural and introduced vegetation. All the multipurpose
tree species were left to grow for 10 years with pruning (50%) every year starting from
Year 4 of establishment (December 1991). During the month of April 1998, the leaves of
the ten multipurpose tree species (six samples/species) were collected from different parts
of each layer of the canopy to form representative samples of tree leaves for evaluation of
nutritional traits. The leafy samples were initially air dried and then oven dried at
60 58C. Dried samples were ground to pass a 2 mm sieve in a Wiley Mill. They were
analysed for organic matter (OM) and crude protein (CP) contents (AOAC, 1994).
Neutral detergent ®bre (NDF) (Van Soest et al., 1991) and acid detergent ®bre (ADF)
(AOAC, 1990) were also estimated. Condensed tannins (CT) were determined by the
Vanillin-HCl method (Broadhust and Jones, 1978), using a catechin standard. Total
phenolics (TP) were estimated by a Prussian-Blue method (Price and Butler, 1997), using
tannic acid as standard.
The nylon bag technique (Orskov et al., 1980) was used to estimate the extent to which
degradation of DM, OM, CP, and cell wall contents (NDF, ADF, hemicellulose) would
take place in the rumen. Three steers (Haryana breed with average live weight 470 kg)
®tted with rumen cannulae were used. The steers were given water and wheat straw ad
libitum, and 5 kg of a concentrate feed (16% CP and 70% total digestible nutrients in
the DM) per day. The concentrate was given in two equal meals at 8.30 a.m. and 3.30
p.m. The steers were maintained on this dietary regimen for 3 months. Ground leaf
material of 10 g was placed in nylon bags measuring 10:5 cm 22 cm with a pore size
45 mm. Each steer was provided with two bags of each leafy material. The bags tied
together with a ®shing line twine were placed in the rumen via the cannulae and removed
after 48 h of incubation. After withdrawal from the rumen, bags were washed with water
till clear and dried in a hot air oven at 55 58C till constant weight. The residue was also
analysed for OM, CP and cell wall contents. Statistical analysis was done using a general
linear model (Wilkinson et al., 1996) for analysis of variance in completely randomised
design.
3. Results
3.1. Protein and organic matter contents
Mean CP content (g kgÿ1 DM) of NFT leaf materials was 174 10:7 and 96 1:5 in
non-NFT. Lowest CP concentration was recorded in F. bengalensis and highest in L.
leucocephala. CP contents varied signi®cantly (P < 0:05) amongst the leaf materials of
evaluated MPTs. Mean OM contents were 903 5:3 and 916 6:5 g kgÿ1 DM in NFT
and non-NFT with a maximum in A. pendula and a minimum in A. lebbeck, respectively
showing non-signi®cant difference between the groups.
D.B.V. Ramana et al. / Animal Feed Science and Technology 88 (2000) 103±111
107
3.2. Cell wall contents
The mean NDF contents (g kgÿ1 DM) were relatively lower in NFT (404 36:1)
vis-a-vis non-NFT (439 40:1), respectively with higher (P < 0:05) concentration in
F. bengalensis, S. cumini, P. pinnata, B. monosperma and lowest in A. nilotica. Leaf
materials of A. nilotica, L. leucocephala and A. pendula exhibited signi®cantly
(P < 0:05) lower ADF concentration than the other evaluated MPTs. The lowest
cellulose concentration (P < 0:05) was recorded in A. pendula and L. leucocephala leaf
materials. Hemi-cellulose concentration (g kgÿ1 DM) was as low as 83 in S. cumini and
as high as 218 in P. pinnata. Lignin contents varied signi®cantly (P < 0:05) in the
evaluated MPTs with the lowest concentration in A. pendula and highest in S. cumini,
respectively.
3.3. Anti-nutritional characters
All the species contained detectable amounts of CT and TP and varied widely in their
concentration (Table 1). Mean contents of CT and TP were almost two times higher in
non-NFT than in NFT. TP concentration was signi®cantly (P < 0:05) higher in A. nilotica
and T. arjuna, while lowest in B. monosperma and P. pinnata. CT contents ranged from
17 to 31 and 14 to 94 g kgÿ1 DM for NFT and non-NFT with the highest in A. indica and
lowest in S. cumini, respectively.
3.4. In sacco degradability
The in sacco degradability of DM, OM, CP, NDF, ADF and hemi-cellulose differed
signi®cantly (P < 0:05) amongst the MPTs leaf materials (Table 2). Mean DM
degradability was relatively lower in NFT than non-NFT. Higher DM degradation was
recorded in A. pendula and L. leucocephala, while the values were lower in B.
monosperma and F. bengalensis, respectively. A similar trend was observed in OM
degradability of MPTs. Mean CP degradability was almost identical between NFT and
non-NFT leaf materials. Comparatively less variability was observed in the CP
degradability (g kgÿ1 DM) of NFT (596±795) than the non-NFT (384±920). Mean
NDF and ADF degradability values were more in non-NFT than NFT. Signi®cantly
(P < 0:05) higher NDF and ADF degradability values were recorded in A. pendula,
T. arjuna and L. leucocephala than the other evaluated MPTs, however, the values were
lowest in B. monosperma. The highest and lowest hemi-cellulose degradability values
were recorded in A. pendula and B. monosperma, respectively. The species with higher
levels of ®bre had the lowest degradability than species with higher levels of CT and TP.
4. Discussion and conclusions
The MPTs, namely, A. nilotica, L. leucocephala, A. pendula and T. arjuna have
greatest potential for agroforestry in degraded lands in terms of nutritive value of the
pruned foliage. The leafy material contained (kgÿ1 DM) medium to high CP (93±228 g)
108
Botanical name
DMa
OMb
CPc
NDFd
ADFe
Hemi-cellulose
771.2
647.1
519.8
828.2
552.9
671.6
595.5
615.4
794.8
618.4
379.9
379.4
278.1
555.5
362.5
370.4
382.7
248.0
467.3
292.2
396.7
370.8
349.3
649.4
462.3
643.8 39.6
663.9 40.0
659.1 24.12
391.1 30.14
352.1 25.37
445.7 36.23
tree species
880.9
721.6
513.8
777.0
669.7
885.4
730.7
533.6
821.8
688.4
919.8
704.2
384.4
770.3
525.1
706.2
410.2
352.1
597.8
403.4
672.4
384.5
327.7
580.1
409.9
772.7
473.0
432.1
648.8
365.3
712.6 40.5
730.0 40.5
660.8 62.5
493.9 45.01
474.9 43.24
538.4 50.03
Nitrogen ®xing multipurpose tree species
Acacia nilotica
749.2
Albizia lebbeck
623.0
Butea monosperma
503.7
Leucaena leucocephala
809.1
Pongamia pinnata
533.8
Mean
Non-nitrogen ®xing multipurpose
Anogeissus pendula
Azadirachta indica
Ficus bengalensis
Terminalia arjuna
Syzygium cumini
Mean
a
DM: dry matter.
OM: organic matter.
c
CP: crude protein.
d
NDF: neutral detergent ®bre.
e
ADF: acid detergent ®bre.
b
D.B.V. Ramana et al. / Animal Feed Science and Technology 88 (2000) 103±111
Table 2
In sacco degradability of nutritive constituents in nitrogen and non-nitrogen ®xing multipurpose tree species (g kgÿ1 DM)
D.B.V. Ramana et al. / Animal Feed Science and Technology 88 (2000) 103±111
109
and high degradable DM, OM, CP (749±881, 771±885, 672±920 g, respectively) and
optimum NDF (261±360 g). NRC (1985, 1989) suggested that diet for mature beef cattle
should contain (kgÿ1 DM) a minimum of 70 g CP and for the high producing cow was
190 g CP and 250 g NDF. Mean CP content of the two groups indicates that the symbiotic
relation of NFT with nitrogen ®xing bacteria helps in enhancing the protein content of the
tree leaves. The CP levels were generally higher than those reported for MPTs by Rai
et al., (1995), but in agreement with those of Upadhyaya et al., (1974) for L.
leucocephala. Although these values may denote high nutritive value of the leaf
materials, the presence of CT (14±94 g kgÿ1 DM) at higher levels in some species could
result in an over estimation of their nutritive value. However, contrary to Ahn et al.
(1989), CT were detected in A. lebbeck. Tannins precipitate proteins by hydrogen bonding
and hydrophobic interactions (Haslam, 1974) to form stable complexes at rumen pH,
which adversely affect protein and ®bre digestion in the rumen (McLeod, 1974; Grif®ths,
1989; Palmer and Schlink, 1992) and thereby reduce protein availability to the animal
(Woodward and Reed, 1989). TP in this study ranged from 7 to 159 g kgÿ1 DM.
Although the TP content was quite high in some species, the degradability seemed not to
be much affected. Non-signi®cant and poor correlation between phenolics and DM
degradation has been reported previously (Makkar et al., 1989; Vadiveloo and Fadel,
1992; Dzowela et al., 1995). Sometimes phenolic assays fail to measure some phenolic
compounds which have a negative effect on the nutritive value of some browses (Khazaal
et al., 1994). This is because of many new compounds which have been identi®ed during
the last few years and showing tannin like properties do not ®t into the categories of either
condensed or hydrolysable tannins (Muller-Harvey and McAllan, 1972).
The data on degradability of DM, OM, CP and cell wall contents indicate a possible
high ®bre, phenolics and tannins induced depression for F. bengalensis, T. arjuna,
P. pinnata, B. monosperma and A. lebbeck, whereas not much effect was found in A.
indica, A. nilotica and L. leucocephala. The DM degradability which is related to nutrient
composition varies widely among tree and shrub species. A range of 38±78% DM
degradability was given by Skarpe and Bergstrom (1986). Lack of rumen degradable
protein and also high concentrate diets reduce the rate of fermentation in the rumen of
®brous feeds (Wilson and Brigstocke, 1981). A wide variation was observed in CP
degradability, the lowest values being in MPTs of high ADF content. Digestibility may be
negatively related to ADF content of the forages (Albrecht and Broderick, 1990) as some
protein can be linked to structural carbohydrates in the cell walls for which degradation is
slow (Mangan, 1982). Tannin complexes especially with proteins may protect them from
microbial degradation which can have both negative and positive effects on their
utilisation (Reed, 1995). In low-tannin containing feeds, true digestibility of nitrogen is
approximately 93% (Van Soest, 1982). Species with moderate levels of tannins having
higher proportion of escape protein than those without tannins has been reported
(Albrecht and Broderick, 1990). CT protect labile plant proteins in the rumen and may
consequently increase the supply of high quality proteins entering the duodenum
(Mangan, 1988).This indicates that in some MPTs more protein could escape ruminal
digestion and be digested in the lower part of the digestive tract (Driedger and Hat®eld,
1978). Although, changes in the tannin content of individual species may affect rumen
nitrogen and OM degradability (Barry and Forss, 1983), it is perhaps not valid to compare
110
D.B.V. Ramana et al. / Animal Feed Science and Technology 88 (2000) 103±111
the effect of tannin content on such parameters between species as there is no existing
accurate method for estimation of ``active'' tannins.
In some species, namely, S. cumini, F. bengalensis, B. monosperma, A. lebbeck and
P. pinnata digestibility was associated with higher NDF contents. Ruminant livestock
require ®bre for normal rumen function but ®bre also limits feed intake and digestibility
(Albrecht and Broderick, 1990). Lignin is the compound most negatively correlated
with degradability (Lapierre, 1993; Buxton and Fales, 1994; Dzowela et al., 1995). A very
high lignin content could be the one of the reason for low degradability in F. bengalensis,
S. cumini and A. lebbeck. The in sacco DM degradability at 48 h in all the leaf materials
was strongly correlated with NDF (r ÿ0:96 and ÿ0.92), ADF (r ÿ0:96 and ÿ0.85)
and cellulose (r ÿ0:73 and ÿ0.79) for NFT and non-NFT, respectively.
The present study shows that some MPTs are potential sources for use as protein banks
to supplement grass or crop residues in dry seasons. However, the presence of CT and TP
at higher levels in some species may be a major constraint to their utilisation by the
livestock.
Acknowledgements
The ®rst author is grateful to the Director and Head of PAR division, IGFRI, Jhansi for
providing the facilities for in sacco degradability assays.
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Speedy, A., Pugliese, P.L. (Eds.), Legume Trees and Other Fodder Trees as Protein Sources for Livestock.
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88 (2000) 103±111
Short communication
Nutritive evaluation of some nitrogen and
non-nitrogen ®xing multipurpose tree species
D.B.V. Ramanaa,*, Sultan Singhb, K.R. Solankia, A.S. Negib
a
National Research Centre for Agroforestry, Near Pahuj Dam, Gwalior Road, Jhansi 284003, U.P., India
b
PAR Division, Indian Grassland & Fodder Research Institute, Jhansi 284003, U.P., India
Received 6 October 1999; received in revised form 3 February 2000; accepted 10 August 2000
Abstract
The nutritive value of leaf materials from ®ve each of nitrogen (NFT), namely, Acacia nilotica,
Albizia lebbeck, Butea monosperma, Leucaena leucocephala, Pongamia pinnata and non-nitrogen
®xing multipurpose tree (non-NFT) species, namely, Anogeissus pendula, Azadirachta indica, Ficus
bengalensis, Terminalia arjuna and Syzygium cumini grown at National Research Centre for
Agroforestry, were evaluated by chemical and in sacco methods. Mean organic matter (OM)
concentration (g kgÿ1 dry matter) was comparable between NFT (902 5:3) and non-NFT
(916 6:5). However, signi®cantly (P < 0:05) higher mean crude protein (CP) contents
(g kgÿ1 DM) were recorded in NFT (174 10:7) than non-NFT (96 1:5) with maximum in
L. leucocephala (228) and minimum in F. bengalensis (88), respectively. Mean neutral detergent
®bre (NDF) contents were relatively lower in NFT than in non-NFT. Signi®cantly (P < 0:05) higher
contents of NDF, acid detergent ®bre (ADF) and lignin (g kgÿ1 DM) were observed in F.
bengalensis (597, 458 and 222) and S. cumini (566, 483 and 279) than in any of the other trees.
Total phenolics (TP) concentration was signi®cantly (P < 0:05) lower in NFT except for A. nilotica,
but higher (P < 0:05) in non-NFT except in F. bengalensis. Mean condensed tannins (CT)
concentration differed signi®cantly (P < 0:05) between NFT and non-NFT and ranged from 17 to
31 and 14 to 94 g kgÿ1 DM, respectively. In sacco degradability of DM differed signi®cantly
(P < 0:05) amongst the MPTs and ranged from 504 to 809 for NFT and 513 to 881 g kgÿ1 DM for
non-NFT, respectively. Mean CP degradability (g kgÿ1 DM) was similar for (P < 0:05) NFT
(659 24:1) and non-NFT (661 62:5). Cell wall contents degradability was relatively higher in
non-NFT than NFT. However, a poor correlation was observed between anti-nutritional characters
and in sacco degradability, whereas strong negative correlations were recorded between DM
degradability and NDF, ADF and cellulose contents. The NFT contained high CP and low cell wall
contents and anti-nutritional characters, however the in sacco degradability of nutrients was less for
non-NFT. Nutritive value of the trees indicated that all the species could potentially be used as
*
Corresponding author. Tel.: 91-517-448213.
E-mail address: [email protected] (D.B.V. Ramana).
0377-8401/00/$ ± see front matter # 2000 Elsevier Science B.V. All rights reserved.
PII: S 0 3 7 7 - 8 4 0 1 ( 0 0 ) 0 0 2 0 0 - 5
104
D.B.V. Ramana et al. / Animal Feed Science and Technology 88 (2000) 103±111
protein banks to supplement grass or crop residues in dry seasons. # 2000 Elsevier Science B.V.
All rights reserved.
Keywords: Tree leaves; Nutritive value; In sacco degradability; Condensed tannins; Total phenolics
1. Introduction
Trees and shrubs have been used for generations as multipurpose resources (food,
fodder, timber, wood, ®bre and live fences) across all of the agro-ecological zones of the
world (Smith, 1992). Cutting of tree leaves or branches for animal feed is recorded as
early as Roman times. In countries like India, experiencing a monsoonal climate, trees
and fodder shrubs may be the only living feed resources available during the driest
months when shallow rooted grasses and herbs die off (Chen et al., 1992). Further,
sustained and high population growth rates, combined with limited and rapidly
diminishing land holdings and land for food and forage production, have created a
need to intensify the research for new resources. Intensi®cation, in the context of
livestock production systems, means a broadening of the feed resource base for both
quantity and quality. One of the alternative feed resources is the use of multipurpose tree
species for top fodder production (NCA, 1976). Much work has been done world wide on
the evaluation and use of leguminous fodder trees and shrubs, while the studies with nonnitrogen ®xing multipurpose tree species are quite obscure. Under certain soil and
climatic conditions, leguminous tree species do not perform well, while the non-nitrogen
®xing trees, which are quite hardy, can be ®tted in the agroforestry systems and may be
exploited for livestock feeding.
Since feed or fodder value is indicated by its nutrient content and digestibility, in this
study an attempt has been made to document the nutritive content and in sacco
degradability of some leguminous and non-nitrogen ®xing tree leaf materials.
2. Materials and methods
Five each of nitrogen (NFT) and non-nitrogen ®xing multipurpose tree (non-NFT)
species (Table 1) were grown at the National Research Centre for Agroforestry (788350 E
longitude and 258260 N latitude and about 275 m above sea level). Annual long-term
rainfall for the site is estimated to 906 mm falling predominantly from June to September.
Temperature rises to a maximum 47.48C in May and falls to a minimum of 1.08C in
January. Evaporation is highest (441 mm) in May and lowest (55.8 mm) in January. The
soils are Red gravelly with stone pebbles and are generally of relatively low fertility (pH
7.2±7.6) and de®cient in nitrogen and phosphorus (Rai et al., 1997). The area's natural
vegetation consists of Chrysopogon fulvus, Dichianthium annulatum, Iseilema laxum and
Themeda quadrivalves dominated by Sehima nervosum and Heteropogon contortus for
grasses as understory vegetation (Rai et al., 1995).
Botanical name
OMa
Nitrogen ®xing multipurpose tree species
Acacia nilotica
897.5
Albizia lebbeck
887.3
Butea monosperma
889.2
Leucaena leucocephala 930.3
Pongamia pinnata
908.1
Mean
902.5 5.3
CPb
NDFc
ADFd
Cellulose
Hemicellulose
Lignin
Total
phenolics
Condensed
tannins
155.5
142.3
150.2
227.8
193.6
261.4
433.4
506.1
293.6
525.9
166.4
312.6
356.0
151.4
308.5
99.4
188.1
151.2
78.2
136.6
94.9
120.7
150.2
142.2
217.5
95.6
174.0
130.7
76.6
130.6
155.9
12.2
6.5
18.6
6.6
26.0
16.8
20.8
23.5
30.6
173.9 10.7 404.1 36.11 259.0 27.84 130.7 12.9 145.1 13.65 121.5 11.16 40.0 19.45 23.5 1.63
Non-nitrogen ®xing multipurpose tree species
Anogeissus pendula
945.8
100.1
Azadirachta indica
894.2
97.2
Ficus bengalensis
930.8
88.0
Terminalia arjuna
898.9
93.3
Syzygium cumini
911.8
100.0
Mean
a
916.3 6.5
OM: organic matter.
CP: crude protein.
c
NDF: neutral detergent ®bre.
d
ADF: acid detergent ®bre.
b
95.7 1.5
291.3
380.1
597.0
360.4
565.7
192.9
269.6
457.9
267.6
482.7
65.4
121.2
248.0
169.8
283.3
98.4
110.5
139.2
92.9
83.0
438.9 40.13 334.1 38.28 177.5 26.7 104.8 6.45
63.0
102.0
221.8
167.2
279.4
166.7 26.1
73.2
65.3
14.0
159.0
24.9
19.9
93.8
77.6
52.0
14.0
66.1 16.38 51.5 10.4
D.B.V. Ramana et al. / Animal Feed Science and Technology 88 (2000) 103±111
Table 1
Chemical composition of nitrogen and non-nitrogen ®xing multipurpose tree species (g kgÿ1 DM)
105
106
D.B.V. Ramana et al. / Animal Feed Science and Technology 88 (2000) 103±111
Nursery-raised seedlings, approximately 4 months old were planted in July 1988 at a
spacing of 8 m between rows and 4 m between plants within rows. In inter-spaces of tree
rows a mixture of Cenchrus ciliaris (2 kg seed/ha) and Stylosanthes hamata (3 kg seed/
ha) were sown by broadcasting after one operation by the cultivator during August 1988.
At the onset of the monsoon, 40 kg N 20 kg P2O5 per hectare was applied uniformly to
enhance the productivity of the natural and introduced vegetation. All the multipurpose
tree species were left to grow for 10 years with pruning (50%) every year starting from
Year 4 of establishment (December 1991). During the month of April 1998, the leaves of
the ten multipurpose tree species (six samples/species) were collected from different parts
of each layer of the canopy to form representative samples of tree leaves for evaluation of
nutritional traits. The leafy samples were initially air dried and then oven dried at
60 58C. Dried samples were ground to pass a 2 mm sieve in a Wiley Mill. They were
analysed for organic matter (OM) and crude protein (CP) contents (AOAC, 1994).
Neutral detergent ®bre (NDF) (Van Soest et al., 1991) and acid detergent ®bre (ADF)
(AOAC, 1990) were also estimated. Condensed tannins (CT) were determined by the
Vanillin-HCl method (Broadhust and Jones, 1978), using a catechin standard. Total
phenolics (TP) were estimated by a Prussian-Blue method (Price and Butler, 1997), using
tannic acid as standard.
The nylon bag technique (Orskov et al., 1980) was used to estimate the extent to which
degradation of DM, OM, CP, and cell wall contents (NDF, ADF, hemicellulose) would
take place in the rumen. Three steers (Haryana breed with average live weight 470 kg)
®tted with rumen cannulae were used. The steers were given water and wheat straw ad
libitum, and 5 kg of a concentrate feed (16% CP and 70% total digestible nutrients in
the DM) per day. The concentrate was given in two equal meals at 8.30 a.m. and 3.30
p.m. The steers were maintained on this dietary regimen for 3 months. Ground leaf
material of 10 g was placed in nylon bags measuring 10:5 cm 22 cm with a pore size
45 mm. Each steer was provided with two bags of each leafy material. The bags tied
together with a ®shing line twine were placed in the rumen via the cannulae and removed
after 48 h of incubation. After withdrawal from the rumen, bags were washed with water
till clear and dried in a hot air oven at 55 58C till constant weight. The residue was also
analysed for OM, CP and cell wall contents. Statistical analysis was done using a general
linear model (Wilkinson et al., 1996) for analysis of variance in completely randomised
design.
3. Results
3.1. Protein and organic matter contents
Mean CP content (g kgÿ1 DM) of NFT leaf materials was 174 10:7 and 96 1:5 in
non-NFT. Lowest CP concentration was recorded in F. bengalensis and highest in L.
leucocephala. CP contents varied signi®cantly (P < 0:05) amongst the leaf materials of
evaluated MPTs. Mean OM contents were 903 5:3 and 916 6:5 g kgÿ1 DM in NFT
and non-NFT with a maximum in A. pendula and a minimum in A. lebbeck, respectively
showing non-signi®cant difference between the groups.
D.B.V. Ramana et al. / Animal Feed Science and Technology 88 (2000) 103±111
107
3.2. Cell wall contents
The mean NDF contents (g kgÿ1 DM) were relatively lower in NFT (404 36:1)
vis-a-vis non-NFT (439 40:1), respectively with higher (P < 0:05) concentration in
F. bengalensis, S. cumini, P. pinnata, B. monosperma and lowest in A. nilotica. Leaf
materials of A. nilotica, L. leucocephala and A. pendula exhibited signi®cantly
(P < 0:05) lower ADF concentration than the other evaluated MPTs. The lowest
cellulose concentration (P < 0:05) was recorded in A. pendula and L. leucocephala leaf
materials. Hemi-cellulose concentration (g kgÿ1 DM) was as low as 83 in S. cumini and
as high as 218 in P. pinnata. Lignin contents varied signi®cantly (P < 0:05) in the
evaluated MPTs with the lowest concentration in A. pendula and highest in S. cumini,
respectively.
3.3. Anti-nutritional characters
All the species contained detectable amounts of CT and TP and varied widely in their
concentration (Table 1). Mean contents of CT and TP were almost two times higher in
non-NFT than in NFT. TP concentration was signi®cantly (P < 0:05) higher in A. nilotica
and T. arjuna, while lowest in B. monosperma and P. pinnata. CT contents ranged from
17 to 31 and 14 to 94 g kgÿ1 DM for NFT and non-NFT with the highest in A. indica and
lowest in S. cumini, respectively.
3.4. In sacco degradability
The in sacco degradability of DM, OM, CP, NDF, ADF and hemi-cellulose differed
signi®cantly (P < 0:05) amongst the MPTs leaf materials (Table 2). Mean DM
degradability was relatively lower in NFT than non-NFT. Higher DM degradation was
recorded in A. pendula and L. leucocephala, while the values were lower in B.
monosperma and F. bengalensis, respectively. A similar trend was observed in OM
degradability of MPTs. Mean CP degradability was almost identical between NFT and
non-NFT leaf materials. Comparatively less variability was observed in the CP
degradability (g kgÿ1 DM) of NFT (596±795) than the non-NFT (384±920). Mean
NDF and ADF degradability values were more in non-NFT than NFT. Signi®cantly
(P < 0:05) higher NDF and ADF degradability values were recorded in A. pendula,
T. arjuna and L. leucocephala than the other evaluated MPTs, however, the values were
lowest in B. monosperma. The highest and lowest hemi-cellulose degradability values
were recorded in A. pendula and B. monosperma, respectively. The species with higher
levels of ®bre had the lowest degradability than species with higher levels of CT and TP.
4. Discussion and conclusions
The MPTs, namely, A. nilotica, L. leucocephala, A. pendula and T. arjuna have
greatest potential for agroforestry in degraded lands in terms of nutritive value of the
pruned foliage. The leafy material contained (kgÿ1 DM) medium to high CP (93±228 g)
108
Botanical name
DMa
OMb
CPc
NDFd
ADFe
Hemi-cellulose
771.2
647.1
519.8
828.2
552.9
671.6
595.5
615.4
794.8
618.4
379.9
379.4
278.1
555.5
362.5
370.4
382.7
248.0
467.3
292.2
396.7
370.8
349.3
649.4
462.3
643.8 39.6
663.9 40.0
659.1 24.12
391.1 30.14
352.1 25.37
445.7 36.23
tree species
880.9
721.6
513.8
777.0
669.7
885.4
730.7
533.6
821.8
688.4
919.8
704.2
384.4
770.3
525.1
706.2
410.2
352.1
597.8
403.4
672.4
384.5
327.7
580.1
409.9
772.7
473.0
432.1
648.8
365.3
712.6 40.5
730.0 40.5
660.8 62.5
493.9 45.01
474.9 43.24
538.4 50.03
Nitrogen ®xing multipurpose tree species
Acacia nilotica
749.2
Albizia lebbeck
623.0
Butea monosperma
503.7
Leucaena leucocephala
809.1
Pongamia pinnata
533.8
Mean
Non-nitrogen ®xing multipurpose
Anogeissus pendula
Azadirachta indica
Ficus bengalensis
Terminalia arjuna
Syzygium cumini
Mean
a
DM: dry matter.
OM: organic matter.
c
CP: crude protein.
d
NDF: neutral detergent ®bre.
e
ADF: acid detergent ®bre.
b
D.B.V. Ramana et al. / Animal Feed Science and Technology 88 (2000) 103±111
Table 2
In sacco degradability of nutritive constituents in nitrogen and non-nitrogen ®xing multipurpose tree species (g kgÿ1 DM)
D.B.V. Ramana et al. / Animal Feed Science and Technology 88 (2000) 103±111
109
and high degradable DM, OM, CP (749±881, 771±885, 672±920 g, respectively) and
optimum NDF (261±360 g). NRC (1985, 1989) suggested that diet for mature beef cattle
should contain (kgÿ1 DM) a minimum of 70 g CP and for the high producing cow was
190 g CP and 250 g NDF. Mean CP content of the two groups indicates that the symbiotic
relation of NFT with nitrogen ®xing bacteria helps in enhancing the protein content of the
tree leaves. The CP levels were generally higher than those reported for MPTs by Rai
et al., (1995), but in agreement with those of Upadhyaya et al., (1974) for L.
leucocephala. Although these values may denote high nutritive value of the leaf
materials, the presence of CT (14±94 g kgÿ1 DM) at higher levels in some species could
result in an over estimation of their nutritive value. However, contrary to Ahn et al.
(1989), CT were detected in A. lebbeck. Tannins precipitate proteins by hydrogen bonding
and hydrophobic interactions (Haslam, 1974) to form stable complexes at rumen pH,
which adversely affect protein and ®bre digestion in the rumen (McLeod, 1974; Grif®ths,
1989; Palmer and Schlink, 1992) and thereby reduce protein availability to the animal
(Woodward and Reed, 1989). TP in this study ranged from 7 to 159 g kgÿ1 DM.
Although the TP content was quite high in some species, the degradability seemed not to
be much affected. Non-signi®cant and poor correlation between phenolics and DM
degradation has been reported previously (Makkar et al., 1989; Vadiveloo and Fadel,
1992; Dzowela et al., 1995). Sometimes phenolic assays fail to measure some phenolic
compounds which have a negative effect on the nutritive value of some browses (Khazaal
et al., 1994). This is because of many new compounds which have been identi®ed during
the last few years and showing tannin like properties do not ®t into the categories of either
condensed or hydrolysable tannins (Muller-Harvey and McAllan, 1972).
The data on degradability of DM, OM, CP and cell wall contents indicate a possible
high ®bre, phenolics and tannins induced depression for F. bengalensis, T. arjuna,
P. pinnata, B. monosperma and A. lebbeck, whereas not much effect was found in A.
indica, A. nilotica and L. leucocephala. The DM degradability which is related to nutrient
composition varies widely among tree and shrub species. A range of 38±78% DM
degradability was given by Skarpe and Bergstrom (1986). Lack of rumen degradable
protein and also high concentrate diets reduce the rate of fermentation in the rumen of
®brous feeds (Wilson and Brigstocke, 1981). A wide variation was observed in CP
degradability, the lowest values being in MPTs of high ADF content. Digestibility may be
negatively related to ADF content of the forages (Albrecht and Broderick, 1990) as some
protein can be linked to structural carbohydrates in the cell walls for which degradation is
slow (Mangan, 1982). Tannin complexes especially with proteins may protect them from
microbial degradation which can have both negative and positive effects on their
utilisation (Reed, 1995). In low-tannin containing feeds, true digestibility of nitrogen is
approximately 93% (Van Soest, 1982). Species with moderate levels of tannins having
higher proportion of escape protein than those without tannins has been reported
(Albrecht and Broderick, 1990). CT protect labile plant proteins in the rumen and may
consequently increase the supply of high quality proteins entering the duodenum
(Mangan, 1988).This indicates that in some MPTs more protein could escape ruminal
digestion and be digested in the lower part of the digestive tract (Driedger and Hat®eld,
1978). Although, changes in the tannin content of individual species may affect rumen
nitrogen and OM degradability (Barry and Forss, 1983), it is perhaps not valid to compare
110
D.B.V. Ramana et al. / Animal Feed Science and Technology 88 (2000) 103±111
the effect of tannin content on such parameters between species as there is no existing
accurate method for estimation of ``active'' tannins.
In some species, namely, S. cumini, F. bengalensis, B. monosperma, A. lebbeck and
P. pinnata digestibility was associated with higher NDF contents. Ruminant livestock
require ®bre for normal rumen function but ®bre also limits feed intake and digestibility
(Albrecht and Broderick, 1990). Lignin is the compound most negatively correlated
with degradability (Lapierre, 1993; Buxton and Fales, 1994; Dzowela et al., 1995). A very
high lignin content could be the one of the reason for low degradability in F. bengalensis,
S. cumini and A. lebbeck. The in sacco DM degradability at 48 h in all the leaf materials
was strongly correlated with NDF (r ÿ0:96 and ÿ0.92), ADF (r ÿ0:96 and ÿ0.85)
and cellulose (r ÿ0:73 and ÿ0.79) for NFT and non-NFT, respectively.
The present study shows that some MPTs are potential sources for use as protein banks
to supplement grass or crop residues in dry seasons. However, the presence of CT and TP
at higher levels in some species may be a major constraint to their utilisation by the
livestock.
Acknowledgements
The ®rst author is grateful to the Director and Head of PAR division, IGFRI, Jhansi for
providing the facilities for in sacco degradability assays.
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