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Small Ruminant Research 75 (2008) 105–114

Effect of dietary inclusion of Ficus infectoria leaves as a

protectant of proteins on the performance of lambs
Avijit Dey 1 , Narayan Dutta ∗ , K. Sharma, A.K. Pattanaik
Centre of Advanced Studies in Animal Nutrition, Indian Veterinary Research Institute, Izatnagar 243122, India
Received 15 March 2007; received in revised form 30 May 2007; accepted 12 June 2007
Available online 5 November 2007

Abstract
This study examined the effects of dietary inclusion of tanniferous leaves of Ficus infectoria as a protectant of proteins on the
performance of lambs. Twenty-four lambs were randomly divided into four groups of six each in a completely randomized block
design and fed four iso-nitrogenous supplements formulated to contain 0, 1.0, 1.5 and 2.0% condensed tannins (CT) through dried
and ground leaves of F. infectoria. The diets were designated as CT-1.0, CT-1.5 and CT-2.0, respectively, and fed to lambs on a basal
diet of wheat straw to meet requirements for maintenance and growth. Blood-biochemical profile was monitored in all the lambs at
0, 45, 90, 135, 180 days of feeding. The average daily growth rate and wool growth for a period of 180 days showed a significant
(P < 0.05) increase by the supplementation of CT at 1.5% through F. infectoria leaves. Although addition of CT up to 1.5% in
the supplement did not interfere with the nutrient intake or digestibility, a depressing effect on DM, OM and ADF digestibility at
2.0% CT level was apparent without any detrimental affect on intake. Digestible crude protein (DCP) and total digestible nutrients
(TDN) values of the composite diets were comparable, except for significantly (P < 0.05) lower TDN (%) in CT-2.0 than the control.
Intake of DCP, digestible organic matter and TDN was comparable irrespective of dietary treatments, though lambs under CT-
2.0 had significantly (P < 0.05) lower intake as compared to their counterparts given diet CT-1.0. Feeding of CT containing diets
particularly at 1.5 and 2.0% levels significantly (P < 0.05) influenced N utilization and improved its retention, however, the microbial
protein synthesis as estimated by urinary excretion of purine derivatives was not affected by the dietary treatments. Haematological
(haemoglobin and packed cell volume) and biochemical parameters (serum glucose, total protein, albumin, globulin, A:G ratio,
alanine aminotransferase and aspartate aminotransferase were similar among the dietary treatments except for significant (P < 0.05)
reduction in serum urea concentration of lambs fed 1.5 and 2.0% CT in the supplement. It may be concluded that CT from F.
infectoria leaves at 1.5% in supplement could be used as a protectant of proteins for improving the performance of lambs.
© 2007 Elsevier B.V. All rights reserved.

Keywords: Condensed tannins; Digestibility; Ficus infectoria; Growth; Proteins; Lambs

1. Introduction

Protein in ruminant diets is poorly utilized because of

∗ Corresponding author. Tel.: +91 9412566130;
fax: +91 581 2303284.
E-mail address: pn@ivri.up.nic.in (N. Dutta).
1 Present address: Malda Krishi Vigyan Kendra, Uttar Banga Krishi
Viswavidyalaya, Mathurapur, Manikchak, Malda 732203, India.
extensive breakdown in the rumen, which may exceed
microbial protein synthesis. Degradation often results in
wastage of dietary proteins, particularly in productive
ruminants such as growing animals, which have high
protein requirements. Protection of proteins is essential
for productive animals, where the protein requirement

0921-4488/$ – see front matter © 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.smallrumres.2007.06.013
106 A. Dey et al. / Small Ruminant Research 75 (2008) 105–114
of these animals cannot be met from microbial pro-
tein synthesis. There has been considerable interest in
reducing ruminal degradation of proteins. Studies have
indicated that feeding proteins, which are resistant to
microbial breakdown in the rumen but available in
the post-ruminally, significantly increased growth rate
(Terrill et al., 1992; Wang et al., 1996a) and production
of milk and milk protein (Wright et al., 1998).
Various treatments such as heating (Glimp et al.,
1967; Kaufmann and Lupping, 1982; Stutts et al., 1988)
and formaldehyde treatments (Ferguson et al., 1967;
Mudgal and Sengar, 1984; Dutta and Agrawal, 2000)
have been used to protect proteins from rumen degrada-
tion and thereby to provide by-pass protein to the lower
tract. However, these treatments may impair the subse-
quent intestinal availability of some amino acids, notably
lysine, cysteine, tyrosine and leucine (Ashes et al., 1984;
Schonhusen et al., 1986). Moreover, an increasing num-
ber of consumers demanding healthy and natural foods
have pushed organic livestock farming that is reputed
to be environment friendly, sustaining animals in good
health, with high welfare standards and prohibit routine
use of growth promoters, animals’ offals or any other
chemicals and additives to livestock rations. Thus, the
use of formaldehyde and other chemicals to protect pro-
teins from ruminal degradation has no scope in organic
animal farming (IFOM, 2006). It is therefore imperative
to explore alternative protectants of protein to improve
protein utilization and animal productivity. In this con-
text, there is a growing interest in the possible use of
CT as protectant of protein in the ration of animals. CT
(proanthocyanidins) form complexes with proteins that
are stable over the pH range of 3.5–7.0, but dissociate
in the abomasum (pH below 3.5) and anterior duode-
num (Getachew et al., 2000a). Complexation protects
proteins from microbial hydrolysis and deamination in
the rumen and increases the availability of feed proteins
for digestion and more amino acids are absorbed post-
ruminally (Getachew et al., 2000a,b; Makkar, 2003; Min
et al., 2003). In contrary to above beneficial effects a
report suggest that enzymatic proteins, as well as other
endogenous proteins, comprise a considerable portion of
excreted nitrogen when animal fed tannins (Fahey and
Jung, 1989). CT are known to inhibit digestive enzymes,
caused mainly by nonspecific binding of tannins with the
enzyme protein.
Ficus infectoria is an evergreen tree largely grown in
Northern parts of India. They are generally planted for
shade and not used as fodder tree due to high tannin con-
tent. A standard size F. infectoria tree can provide about
500–600 kg fresh leaves from one cutting. Preliminary
study with graded levels (1–2%) of CT from F. infectoria
leaves in the substrate indicated significant reduction on
in vitro nitrogen degradability of groundnut cake (Dey et
al., 2006). The present investigation aimed to study the
effect of graded levels of CT from F. infectoria on the
diet digestibility, N balance and performance of growing
lambs.
2. Materials and methods
The experiment was conducted at the Animal Nutrition
Research Sheds of the Indian Veterinary Research Institute
(IVRI), Izatnagar in Uttar Pradesh Province of India. Mixed
farming involving crops and livestock integration has been a
way of life in this region. Despite the importance of livestock,
prevalence of low producing non-descript livestock breeds and
their inadequate nutrition is a common problem in small mixed
farms and a major factor affecting the development of commer-
cially viable livestock production system. Sugarcane, wheat
and rice are the main cultivated crops, and cereal straws form
the basal diet of the ruminants in the area.
2.1. Animals and diets
Twenty-four 6-month-old lambs (11.73 ± 0.22 kg), were
randomly allocated to four dietary treatments in a completely
randomized block design. Prior to start of the trial the lambs
were treated with broad-spectrum anthelmintic [Albandazole
suspension, Smith Kline Pharmaceuticals Limited, India, at
2 ml/10 kg body wt.] at onset of the study and thereafter at 3
months intervals. After shearing, the lambs were treated for
ectoparasites [Butox liquid (Hoechst India Limited, Mumbai,
India) at 2 ml/l of water]. The lambs were penned individually
with free access to fresh water in ventilated sheds and allowed
exercise out-doors in an adjacent dry paddock daily between
08:30 and 09:30 h. Four iso-nitogenous supplements were for-
mulated to contain 0, 1.0, 1.5 and 2.0% CT supplied from dried
and ground leaf meal of F. infectoria. The diets were designated
as CT-0, CT-1.0, CT-1.5 and CT-2.0, consisted of wheat straw
(ad lib), green fodder (100 g/day) along with the respective
supplements (Table 1), and fed so as to meet the requirements
(Kearl, 1982) for maintenance and a growth of 50 g/day.
F. infectoria leaves were harvested in one lot in the month of
July from the IVRI campus. The leaves were dried and ground
in an electric grinder before mixing in the supplements. Dried
and ground F. infectoria leaves were incorporated in different
proportion to the supplements to replace wheat bran with CT
content at 0, 1.0, 1.5 and 2.0% of supplements on dry matter
basis. The ingredients and chemical analysis of the supple-
ments and wheat straw are given in Table 1. The amount of
supplements offered to individual lambs was adjusted fort-
nightly as per the body weight changes of each animal to meet
the nutrient requirement (Kearl, 1982).
2.2. Experimental procedures
The daily allowance of the supplements was offered in
single meals (at 09:30 h) in the morning; green (100 g) and
 A. Dey et al. / Small Ruminant Research 75 (2008) 105–114 107

Table 1
Ingredients and chemical composition of supplements, F. infectoria and wheat straw
Constituents Supplementsa Ficus infectoria Wheat straw Wheat Bran

CT-0 CT-1.0 CT-1.5 CT-2.0

Ingredients (g/kg)
Maize (grain) 250 250 250 250 – – –
Deoiled groundnut cake 350 350 350 350 – – –
Wheat bran 380 274 221 168 – – –
F. infectoria – 106 159 212 – – –
Mineral mixtureb 10 10 10 10 – – –
Common salt 10 10 10 10 – – –
Chemical composition (g/kg DM)
OM 934 ± 4 934 ± 3 934 ± 3 935 ± 2 897 ± 3 940 ± 5 951 ± 4
CP 251 ± 9 255 ± 5 252 ± 7 250 ± 4 134 ± 2 34 ± 2 141 ± 3
EE 24 ± 2 25 ± 3 25 ± 2 26 ± 3 41 ± 3 12 ± 4 35 ± 2
Total ash 66 ± 4 66 ± 3 66 ± 4 65 ± 2 103 ± 4 60 ± 5 49 ± 3
NDF 276 ± 3 312 ± 3 327 ± 6 332 ± 6 459 ± 5 811 ± 7 399 ± 4
ADF 120 ± 4 157 ± 5 160 ± 4 168 ± 6 371 ± 4 532 ± 5 125 ± 3
CTc – 10 15 20 94 ± 3 – –
a CT-0: Control supplement, CT-1.0: 1% CT containing supplement, CT-1.5: 1.5% CT containing supplement, CT-2.0: 2% CT containing

supplement.
b Mineral mixture contained (g/kg): calcium 215, phosphorus 95, sodium chloride 285, potassium iodine 2.5, iron 5.0, copper 0.8, cobalt 1.0,

manganese 1.0 and sulfur 1.0.

c It was assumed that only F. infectoria contains CT, and CT contents of the supplements was calculated accordingly.

wheat straw was then offered ad libitum, when all the lambs
had consumed the concentrate. Straw residues remaining were
weighed 24 h post-feeding to ascertain daily feed consump-
tion. The feeding trial was carried out for 201 days duration
including the first 21 days for adaptation and subsequent 180
days for measurement. Daily DM intake and fortnightly BW
of all the lambs were recorded before feeding in the morning
throughout the study.
A digestion and N balance trial was conducted after
90 days of experimental feeding. During the metabolism
trial, the lambs were housed in individual metabolism cages
(90 cm × 75 cm × 90 cm) made of welded wire-mesh fitted
with removable feeders and arrangements for quantitative col-
lection of faeces and urine separately. The trial lasted for 9 days
with a 3 days adaptation period to accustom the lambs to cages
prior to 6-day collection and measurement period. Samples
of feed offered and refused were collected daily. Total daily
(24 h) faecal and urine outputs were recorded. A sub-sample
of the faeces (20%) collected and dried at 80 ± 2 ◦ C for 24 h in
a forced-draft oven for dry matter estimation. Pooled samples
(6 days for each animal) were ground and stored for chemical
analysis.
For N determination, the faeces samples (2/200th of the
daily voids) were preserved in 25% sulphuric acid to make
pooled samples for individual lamb. Daily urine output was
collected quantitatively in plastic bottles containing known
quantity of sulphuric acid (10%) to bring pH around 3.0 and a
sub-sample was collected into kjeldahl flasks containing known
quantity of sulphuric acid for N estimation. Another aliquot was
taken in properly marked plastic bottle and stored at −20 ◦ C
for purine derivatives estimation.
2.3. Blood collection
Blood samples were collected via jugular vein puncture in
the morning before feeding at 0, 45, 90, 135, 180 days post-
feeding. Serum was separated and preserved at −20 ◦ C for
serum glucose, serum proteins, serum enzymes and serum urea
analysis. Another 2 ml blood sample was collected in vials
containing ethylene diamine tetra-acetate, 1 mg/ml blood, for
haemoglobin and packed cell volume (PCV).
2.4. Wool growth
Shearing was done by hand scissors at the on set and com-
pletion (180 days) of experiment. The total wool yield was
weighed for each lamb and average daily wool yield was cal-
culated. Staple length was measured by metric scale and fibre
diameter by lanometer. An average of 10 wool fibre taken at
random was used as the representative measurement.
2.5. Chemical and statistical analyses
Samples of feeds, residues and faeces were milled to pass
through a 1 mm sieve and then analyzed following the meth-
ods of Association of Official Analytical Chemists (AOAC,
1995) to determine DM by the oven drying method (934.01),
organic matter by muffle furnace incineration (967.05), crude
protein by kjeldahl method (984.13) (N × 6.25), ether extract
(920.39), ash (942.05). Neutral detergent fibre (NDF) and acid
detergent fibre (ADF) were essentially determined by the meth-
ods of Van Soest et al. (1991). NDF was assayed with sodium
sulphite in the NDF reagent without alpha-amylase and the
108 A. Dey et al. / Small Ruminant Research 75 (2008) 105–114

results were expressed with residual ash. The CT content in Table 2

F. infectoria leaves was estimated by Butanol-HCl method Intake and utilization of nutrients at graded levels of CT by lambs
(Makkar, 2000). Urinary purine derivatives (allantoin, uric Attributes Treatmentsa S.E.M.
acid, xanthine, and hypoxanthine) and creatinine were deter-
mined by high performance liquid chromatography (HPLC) CT-0 CT-1.0 CT-1.5 CT-2.0
as described by Resines et al. (1992) using pure standards Body weight
from Sigma Aldrich Limited, New Delhi, India. Microbial N kg 17.8ab 18.4b 18.6b 16.5a 0.29
yield (g/day) was calculated according to Chen and Gomes Metabolic (W0.75 ) 8.7ab 8.9b 9.0b 8.2a 0.11
(1995). Haemoglobin and packed cell volume (PCV) were Intake (g/kg W0.75 )
estimated in whole blood immediately after blood collection DM 65.7 68.6 70.6 67.6 1.22
by acid haematin method (Benjamin, 1985) and Wintrobe’s Wheat straw 26.2 29.4 28.2 25.7 0.86
tube, respectively. Serum glucose concentration was deter- Supplement 39.5 39.2 42.4 41.9 0.79
mined colorimetrically (Hultmann, 1959). The serum protein
Nutrient digestibility (%)
and albumin content were measured as per Wotton (1964) DM 56.3b 54.7ab 55.2ab 52.9a 0.53
and Doumas et al. (1971), respectively. Serum urea was esti- OM 59.2b 57.8ab 58.1ab 56.0a 0.51
mated by improved diacetyl monoxime method (Rahmatulla CP 63.0 62.0 61.5 61.0 0.58
and Boyde, 1980). Activity of serum enzymes alanine amino- EE 68.1 68.1 68.0 67.6 0.38
transferase (AST) and aspartate aminotransferase (ALT) were NDF 45.0 45.4 44.9 44.6 0.49
estimated as per standard methods described by Reitman and ADF 39.6b 38.3b 36.8ab 33.9a 0.70
Frankel (1957). Nitrogen balance (g per day)
The results obtained were subjected to analysis of variance Intake 15.2a 15.9a 17.0b 14.7a 0.37
and treatment means were ranked using Duncan’s multiple Faecal loss 5.6a 6.0ab 6.6b 5.7a 0.15
range test. The degree of freedom of the treatments was Urinary loss 5.7c 5.0bc 4.7b 3.6a 0.21
partitioned into orthogonal polynomial, depicting linear and Balance 4.0a 4.9ab 5.7b 5.4b 0.22
quadratic trends associated with increasing levels of CT sup- Nitrogen excretion (% intake)
plementation. The periodic alterations in body weight and Fecal 37.0 38.0 38.5 38.6 0.58
blood biochemical parameters were analyzed using repeated Urine 37.4c 31.3b 27.5ab 25.5a 1.06
measures design. Significance was declared at P < 0.05 unless
Retention (%)
otherwise stated. All the statistical procedures were done as N intake 25.7a 30.7b 30.2b 37.0c 1.29
per Snedecor and Cochran (1994). N absorbed 40.7a 49.3b 54.6bc 60.1c 1.75

Mean values with different superscripts (a and b) within a row differ

3. Results and discussion significantly (P < 0.05).
a CT-0: Control supplement, CT-1.0: 1% CT containing supplement,

3.1. Chemical composition of diet
The chemical composition of supplements and major
feed components is given in Table 1. The chemical com-
position of wheat straw offered as basal feed was within
the normal range and comparable to values reported for
Indian feeds and fodder (Wales et al., 1990; Dutta and
Sharma, 2004; Sharma et al., 2004; Dey and Sehgal,
2005). F. infectoria leaves containing 9.4% CT were
used as a protectant of dietary protein in the ration. The
experimental supplements were iso-nitrogenous. NDF
and ADF increased with increasing proportion of leaf
meal in the supplement. This is consistent with the high
proportion of cell wall constituents measured in F. infec-
toria (Patra et al., 2003; Anbarasu et al., 2004).
3.2. Dry matter intake
Intake (g/kg W0.75 ) of wheat straw, supplement as
well as total DM during metabolism trial was compa-
rable among dietary treatments (Table 2). The intake of
CT-1.5: 1.5% CT containing supplement, CT-2.0: 2% CT containing
supplement.
DM (g/kg W0.75 ) by lambs was higher than the average
value of 54.7 g reported by Kearl (1982) and this clearly
indicates that all the experimental diets were palatable.
The comparable (P > 0.05) intake by lambs irrespective
of dietary treatments are in agreement with the earlier
observations that moderate levels (1–4%) of CT in the
diet from various plant sources exerted no significant
effect on feed intake (Wang et al., 1996a,b; Bhatta et
al., 2000; Komolong et al., 2001). In the present study,
based on the net DM intake, the actual concentration
of CT in the diets worked out to be 0.0, 0.57, 0.90 and
1.24% for the groups CT-0, CT-1.0, CT-1.5 and CT-2.0,
respectively.
3.3. Nutrient utilization
Digestibility coefficient of DM and OM were compa-
rable among CT containing diets, however, digestibility
 A. Dey et al. / Small Ruminant Research 75 (2008) 105–114 109

of DM, OM and ADF in lambs fed diet CT-2.0 was
significantly (P < 0.05) lower compared to CT-0 (con-
trol) (Table 2). Further, the digestibility coefficients for
DM, OM and ADF tended to decrease with increas-
ing CT levels as evident from a significant linear trend
(P < 0.05). Digestibility of CP, EE and NDF were com-
parable among the dietary treatments. The nutritional
effects of tannins are associated with their ability to bind
with proteins (dietary and enzymes), structural carbohy-
drate polymers found in plant cell walls and minerals
with an overall effect of lowering the bioavailability of
nutrients at specific sites in the gastro-intestinal tract
(Ndluvo, 2000). However, other workers have reported
that tannins are beneficial to ruminants at low concentra-
tion because they protect plant proteins from degradation
in the rumen (Waghorn and Shelton, 1992; Wang et al.,
1994). In the present study the addition of CT up to 1.5%
(i.e. 1.5 and 1%) of supplement does not seem to inter-
fere with the microbial activity or total tract digestibility
of nutrients. A depressing effect on DM digestibility at
2% CT level may be due to associative effects of CT
and higher percentage of structural carbohydrates con-
tributed by F. infectoria leaves. This may probably be
due to interference of CT with microbial attachment or
depressing cellulolytic bacterial population (McAllister
et al., 1994; McSweeney et al., 1998).
Daily intake (g/day) of N was comparable among
lambs irrespective of the dietary treatments, except CT-
1.5 which had significantly higher N intake (Table 2).
Faecal excretion of N (g/day) was significantly (P < 0.05)
higher at CT-1.5 as compared to CT-0 and CT-2.0, how-
ever, faecal N excretion was comparable between CT-1.0
and CT-1.5. The urinary N excretion (g/day) was signif-
icantly (P < 0.05) lower at CT-2.0 followed by CT-1.5
and CT-0, while urinary N excretion at CT-1.0 was
comparable in lambs either fed CT-0 or CT-1.5. All
the lambs had positive N balance (g/day), irrespective
of dietary treatments. However, N-retention was sig-
nificantly (P < 0.05) higher at CT-1.5 and CT-2.0 as
compared to control. Similar faecal excretion (g/day) of
N in CT supplemented groups and control is suggestive
of the tannin bound concentrate substrate undergoing
digestion in the post-ruminal digestive tract before reach-
ing the hindgut. Another feature of N utilization as
evident by significantly higher (linear, P = 0.002) N
retention as percentage of absorbed N (an indicator of
availability of amino acid-N at tissue level) in animals
given CT protected concentrate (Table 2) which could
have been due to better amino acid availability and appar-
ent biological value of CT protected diets as suggested
by Barry and McNabb (1999). This observation is fur-
ther substantiated by the fact that lambs given CT treated
diets excreted significantly lower N in urine as% intake
(linear, P = 0.001) relative to control animals. Similar to
the present study, increased N retention in sheep given
tanniferous feeds at moderate levels (1–4%) due to low-
ered nitrogen excretion through urine has been reported
earlier (Waghorn et al., 1994b; Ngwa et al., 2002). This
protects the protein from microbial hydrolysis and deam-
ination in the rumen and increases the proportion of
dietary amino acids available for post-ruminal absorp-
tion.

Table 3
Purine derivatives and microbial nitrogen supply to lambs fed graded levels of CT
Attributes Treatmentsa S.E.M.

CT-0 CT-1.0 CT-1.5 CT-2.0

Urinary excretion of PD (mmol/day)

Allantoin 4.16 4.28 4.50 3.56 0.28
Uric acid 0.70ab 0.80b 0.70ab 0.55a 0.02
Xanthine 0.23ab 0.23ab 0.25b 0.15a 0.02
Hypoxanthine 0.04 0.12 0.11 0.10 0.02
Total 5.12 5.44 5.56 4.34 0.30
Creatinine 3.28 3.17 3.55 2.76 0.23
PD absorption 5.37 5.72 5.96 4.81 0.34
PD:C 1.57 1.74 1.63 1.68 0.08
Microbial N supply (g/day) 3.91 4.16 4.33 3.50 0.25
Efficiency of microbial N synthesis 10.92 10.78 11.27 10.36 0.26
(g/kg DOMI)

Mean values with different superscript (a and b) within a row differ significantly (P < 0.05). PD: purine derivatives; DOMI: digestible organic matter
intake.
a CT-0: Control supplement, CT-1.0: 1% CT containing supplement, CT-1.5: 1.5% CT containing supplement, CT-2.0: 2% CT containing

supplement.
110 A. Dey et al. / Small Ruminant Research 75 (2008) 105–114

3.4. Microbial nitrogen supply
The excretion of allantoin and hypoxanthine in urine
by lambs was comparable (P < 0.05), irrespective of
dietary treatments (Table 3). Similarly, dietary supple-
mentation of CT did not exert any effect on excretion
of uric acid and xanthine, except for significant dif-
ference observed for uric acid (quadratic, P = 0.036) in
lambs kept on diet CT-1.0 and CT-2.0; and for xanthine
(quadratic, P = 0.072) in animals on CT-1.5 and CT-2.0.
However, total PD excretion, creatinine (C) and PD:C
ratio did not differ among various dietary treatments.
Similarly, the PD absorption (mmol/day), MN (g/day)
and efficiency of microbial N synthesis in term of per kg
DOMI were similar (P < 0.05) among different dietary
treatments. The results are in conformity with the earlier
reports that indicate no difference in urinary excretion
of allantoin, total PD and MN synthesis in lambs fed
iso-nitrogenous diet with different proportion of unde-
graded protein (57–106 g/day) in raw and dry roasted
legume seeds (Hume et al., 1970; Yu et al., 2002). Lack
of any depressing effect of CT supplementation on rumen
microbial protein synthesis agrees with the observations
recorded earlier in sheep fed tannin containing Acacia
pods (Ngwa et al., 2002). The findings suggest the pos-
sibility of using leaves of F. infectoria as CT source in
practical diets without detrimentally affecting MN sup-
ply or efficiency of its production.
3.5. Nutritive value
The nutrient density in terms of percent content of
DCP and TDN of the composite diets (wheat straw and
respective supplements) were comparable, except for
significantly lower (linear, P = 0.017) TDN (%) in treat-
ment CT-2.0 than the control group (Table 4). McNeill et
al. (1999) indicated that dietary protein complexed with
tannins was made available in abomasum and digested
in intestine, but tannins released from the protein-tannin
complexes may react with non-dietary protein (includ-
ing digestive enzymes) as it passes along the intestines,
thus counteracting the benefits of by-pass dietary protein.
This may explain the decrease in TDN content as con-
sequence of decreased OM digestibility when the level
of CT increased to 2.0% of supplement.
Daily nutrient intake (g/kg W0.75 ) in terms of DCP,
DOM and TDN was comparable irrespective of dietary
treatments, except for significantly (P < 0.05) lower

Table 4
Effect of graded levels of CT on growth rate, wool yield and quality and plane of nutrition under experimental feeding for 180 days
Attributes Treatmentsa S.E.M.

CT-0 CT-1.0 CT- 1.5 CT-2.0

Body weight changes (kg)

Initial 11.7 11.8 11.8 11.6 0.22
Final 22.9a 23.4ab 25.3b 21.7a 0.43
Total gain 11.2a 11.6a 13.5b 10.1a 0.38
ADG (g) 62.4a 64.3a 75.2b 56.1a 2.12
DMI (g/day) 567.5a 605.0b 629.8c 578.5a 5.43
DMI (g/kg W0.75 ) 66.9 68.5 70.19 69.34 0.60
FCR 9.3ab 9.5ab 8.5a 10.4b 0.26
Wool yield and quality
Total yield (g) 858.2a 940.2ab 1017.2b 841.3a 27.80
Yield (g/day) 4.8a 5.2ab 5.7b 4.7a 0.15
Staple length (mm) 85.9 90.4 91.4 89.6 1.41
Fibre diameter (␮m) 30.8 30.6 30.5 30.7 0.31
Nutrient density (g/kg)
DCP 105 101 104 101 2.1
TDN 586b 571ab 573ab 554a 5.2
Nutrient intake (g/kg W0.75 )
DCP 7.1a 7.6b 7.5b 7.0a 0.10
DOM 37.7ab 39.6b 39.5ab 36.9a 0.53
TDN 39.6ab 41.8b 41.5ab 38.4a 0.56

TDN calculated from DOM (1 kg DOM = 1.05 kg TDN; NRC, 1981). ADG: average daily gain; FCR: feed conversion ratio (kg DM/kg gain). Mean
values with different superscripts (a–c) within a row differ significantly (P < 0.05).
a CT-0: Control supplement, CT-1.0: 1% CT containing supplement, CT-1.5: 1.5% CT containing supplement, CT-2.0: 2% CT containing

supplement.
 A. Dey et al. / Small Ruminant Research 75 (2008) 105–114
intake of lambs in CT-2.0 as compared to those given CT-
1.0 diet showing a quadratic trend (P < 0.05) (Table 4).
The absence of any detectable adverse effect on the
health of experimental animals suggests that lambs were
on balanced diets with no apparent deleterious con-
sequences. Except for DCP, which fell short of the
requirements (Kearl, 1982) by 12–19%, all the animals
had enough nutrient intakes (DOM and TDN) irrespec-
tive of dietary treatments to meet the requirements for
maintenance and growth (50 g/day). The findings sug-
gest that plane of nutrition was not affected adversely
with CT supplementation in conformity with the ear-
lier reports (Terrill et al., 1992; Waghorn et al., 1994a,b;
Wang et al., 1996a,b). The palatability of browse species
was closely related to the concentration of tannins. There
appears to be a threshold of tannin contents (approxi-
mately 5%) below which no adverse effect was evident
on nutrient intake and utilization.
3.6. Voluntary feed intake, growth and feed
conversion ratio
The overall DMI (g/day) by lambs was significantly
(P < 0.05) higher for CT-1.5 followed by CT-1.0 and
comparable values were recorded for treatment CT-0
and CT-2.0 (Table 4), indicating no adverse effect of
tanniferous feeds on intake of dry matter, wheat straw
and concentrate by lambs during the entire experimen-
tal feeding. The fortnightly body weight changes are
depicted in Fig. 1. The initial live weight was comparable
irrespective of dietary treatments. There was significant
(P < 0.05) influence of dietary treatments on the body
weight of lambs recorded on day 60 of the experimen-
tal feeding and subsequently from day 105 onwards till
day 180. Lambs fed CT-1.5 recorded significantly higher
(P < 0.05) average final body weight (kg) compared to
those given supplement CT-0 and CT-2.0, while body
Fig. 1. Effect of graded levels of CT on live weight changes (CT-0: con-
trol supplement, CT-1.0: 1% CT containing supplement, CT-1.5: 1.5%
CT containing supplement, CT-2.0: 2% CT containing supplement).
111
weight of animals under CT-1.0 was intermediate. Simi-
larly, the total body weight gain (kg) for the period of 180
days and average daily gain (ADG, g) for lambs under
the treatment CT-1.5 was significantly (linear, P = 0.061;
quadratic, P = 0.006) higher compared to others kept on
other dietary treatments. Feed conversion ratio (FCR)
(kg DMI/kg gain) was comparable irrespective of dietary
treatment except for significantly (P < 0.05) lower FCR
in lambs given CT-1.5 as compared to CT-2.0 showing a
linear trend (P = 0.042). The positive response of ADG
and FCR to 1.5% level of CT in the supplement gives
an indication that the binding effect of tannins was pro-
nounced only at this level by supplying protein to the
lower gut and subsequently its more efficient use for
tissue growth. Hence, 1.5% CT is apparently the opti-
mum level, at which there are enough tannins to exert
noticeable effects, but not so much that negative impacts
starts to dominate. Similar trends of increased growth in
sheep supplemented with CT was reported by Ngwa et al.
(2002). At appropriate concentration, the CT reduced the
degradation of sulphur amino acids (SAA) in the rumen,
increases the irreversible loss of cystine from plasma and
increased the flow of cystine to body synthetic reaction
(McNabb et al., 1993; Wang et al., 1994) and thereby
improves the performance of lambs.
3.7. Wool growth and quality
The total wool yield (g) and yield per day (g) were
significantly higher (both linear and quadratic P < 0.01)
for the treatment CT-1.5 compared to similar wool yield
by lambs in CT-0 and CT-2.0 treatments. However, wool
yield in CT-1.0 was intermediate between control and
CT-1.5 diets (Table 4). The increase in fleece weight in
the present experiment in groups CT-1 and CT-1.5 could
be due to the effect of CT increasing the absorption of
SAA and also that of all other EAA (Min et al., 2001).
This is similar to earlier observations that CT in Lotus
increased the efficiency of wool production (Min et al.,
1998; Ramirez-Restrepo et al., 2004). The wool quality
in term of fibre length (mm) and fibre diameter (mm)
was similar (P < 0.05) in lambs irrespective of dietary
treatments. Present observations are in agreement with
the earlier report of Wang et al. (1996a), but contrary to
some reports describing beneficial (higher staple length)
effect of CT on these parameters in grazing sheep (Min et
al., 2001; Ramirez-Restrepo et al., 2004). This could be
due to difference in concentration and chemical compo-
sition of CT, which affect its biological activity (Aerts et
al., 1999). Further, in the present experiment local non-
descript breed of sheep was used and they are known for
inferior (carpet) quality wool production.
112 A. Dey et al. / Small Ruminant Research 75 (2008) 105–114

Table 5 1987, 1990). Similarly, lower plasma/serum urea con-

Effect of graded levels of CT on Hb (g/dl), PCV (%), glucose (mg/dl), centration was reported in sheep grazed on Lotus pasture
serum proteins (g/dl), Serum urea (mg/dl), AST (units/ml) and ALT
(units/ml) by lambs
(Waghorn et al., 1994b; Wang et al., 1996a; Min et
al., 2001) and in kids fed leaves of Prosopis cineraria
Attributes Treatmentsa S.E.M. (Bhatta et al., 2002). Significantly, no treatment period
CT-0 CT-1.0 CT-1.5 CT-2.0 interaction was evident in any parameters of the blood
Hb 11.4 11.2 11.7 11.6 0.21
biochemical profile.
PCV 34.2 34.3 33.9 34.7 0.40
Glucose 54.6 55.4 55.6 55.0 0.71 4. Conclusion
Total Protein 5.6 5.5 5.4 5.5 0.10
Albumin 3.1 3.1 3.1 3.0 0.08
On the basis of present findings, it may be concluded
Globulins 2.4 2.4 2.3 2.5 0.08
A:G ratio 1.4 1.3 1.4 1.3 0.07 that F. infectoria leaves could be a potential source of CT
Serum urea 39.2b 38.0b 34.7a 33.9a 0.69 and may be effectively used as a protectant of proteins.
AST 22.7 21.1 21.0 22.4 0.60 A discernible positive impact was evident on the overall
ALT 15.5 14.9 14.5 15.7 0.50 performance of lambs supplemented with F. infectoria
Mean values with different superscripts (a and b) within a row differ leaves to 1.5% CT in the supplement.
significantly (P < 0.05).
a CT-0: Control supplement, CT-1.0: 1% CT containing supplement,
Acknowledgement
CT-1.5: 1.5% CT containing supplement, CT-2.0: 2% CT containing
supplement.
This study was financially supported by funds pro-
vided by Indian Council of Agricultural Research
3.8. Blood biochemical profile (ICAR), New Delhi, India.

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