Small Ruminant Research 41 (2001) 247±256

Effect of protein supplements of varying degradability on growth

rate, wool yield and wool quality in grazing lambs
G. Habiba,*, M.M. Siddiquia, F.H. Miana, J. Jabbarb, F. Khanc
a
Department of Animal Nutrition, NWFP, Agricultural University, Peshawar, Pakistan
b
Department of Livestock and Dairy Development, Peshawar, Pakistan
c
Sheep and Wool Project, National Agricultural Research Center, Islamabad, Pakistan
Accepted 3 June 2001

Abstract

Two experiments were conducted to investigate productive responses in grazing lambs to supplemental protein of varying
degradability. In Experiment 1, rapeseed cake (RSC), a highly degradable feed, was treated with formaldehyde (HCHO) for
protecting against rumen fermentation. Different levels of formaldehyde, namely, 0, 1, 2, 3, 4 and 5 g/100 g crude protein (CP)
were used to ®nd an optimum level. Increasing levels of formaldehyde linearly decreased the in sacco protein degradability of
RSC at 6, 12 and 24 h of incubation. Maximum response per unit of the chemical was found with 3 g/100 g CP and this level
of formaldehyde was used for treating RSC for feeding lambs in the growth trial. In Experiment 2, 80 female lambs
(Kaghani  Rambouillet, 10±12 months age) from four different private ¯ocks (20 lambs per ¯ock) were selected in one of the
hilly areas of Pakistan. The lambs had free access to range grazing during daytime. In each ¯ock, the lambs were randomly
divided in ®ve groups (four lambs per group) and assigned to the following protein supplements; control (no supplement),
cottonseed cake (CSC), untreated rapeseed cake (U-RSC), HCHO-treated rapeseed cake (T-RSC) and wheat bran (WB) for a
period of 80 days. The supplements were fed on an iso-nitrogenous basis equivalent to 20 g N/100 kg BW. All the lambs
received a complete mineral mixture.
Nutrient composition of range grasses grazed by the lambs remained the same throughout the experimental period and did
not vary due to locations. CP in the grasses averaged 12.03% on a DM basis. The rate of protein degradation (%/h) was
maximum in WB (25.55) followed by U-RSC (15.15), CSC (10.53) and T-RSC (5.32). Body weight (BW) gain (P < 0:01) and
growth of clean wool (P < 0:05) increased to the same extent with all the four supplements. Daily BW gains were 30, 61, 56,
58 and 63 g in the lambs fed control, CSC, U-RSC, T-RSC and WB diets, respectively. Growth rate of clean wool averaged
3.4, 3.9, 4.1, 4.0 and 4.0 g per day on the above diets, respectively. Staple length and ®ber diameter of wool did not respond to
diet composition and remained the same on control and supplemented diets. Despite distinct variation in protein degradability
of the supplements, the lack of difference in BW gain and wool growth among the lambs receiving these supplements were
presumably due to the same quantity of total protein supplied by these feeds post-ruminally. We concluded that protein
supplements of varying degradability supplying 25% of the total dietary protein intake had equal substitution value in grazing
lambs. # 2001 Elsevier Science B.V. All rights reserved.

Keywords: Lambs; Grazing; Protein degradability; Growth; Wool

1. Introduction

*
Corresponding author. Range grazing is one of the major feed resources
E-mail address: fayez@brain.net.pk (G. Habib). for livestock in the hilly regions of Pakistan and

0921-4488/01/$ ± see front matter # 2001 Elsevier Science B.V. All rights reserved.
PII: S 0 9 2 1 - 4 4 8 8 ( 0 1 ) 0 0 2 1 9 - X
248 G. Habib et al. / Small Ruminant Research 41 (2001) 247±256
contributes 27% to the overall feed supply in the
country (Singh et al., 1997). About 60 million hectare
or 68% of the total land area in Pakistan has been
classi®ed as rangeland and 90% of the rangeland is
privately and mostly collectively owned and is
exposed to free grazing by livestock (FAO/ADB,
1987). Against the total estimated carrying capacity
of about 4.3 million animal units, the number of
animal units feeding on the rangeland is 8.5 million.
Due to continued overgrazing and lack of technical
input, these rangelands are producing only 1050%
of their potential (Ra®que, 1989). The nutritive value
of range grasses in the low mountainous regions is
very poor with a crude protein (CP) content varying
from 3 to 7% in most of the predominant grass species
(Habib and Bashir, 1982±1983). A de®ciency of 50%
in CP and 38% in TDN was estimated in sheep grazing
on such rangelands. In Pakistan transhumance and
nomadic sheep farming systems are common and
require supplementary feeding of sheep at certain
times during the year to improve performance of
the animals.
The new protein feeding system for ruminants
places emphasis on maximizing microbial protein
supply to the intestine and complementing it with
dietary protein that escapes undegraded from the
rumen. Cottonseed cake (CSC), rapeseed cake
(RSC) and wheat bran (WB) are the most popular
conventional concentrates fed to ruminant livestock in
Pakistan. Previous studies in this laboratory ranked
rapeseed cake and wheat bran as highly degradable
followed by cottonseed cake (Habib, 1991). Formal-
dehyde (FCHO) treatment of feed protects protein
against rumen fermentation and feeding of the treated
feed to animals was shown to improve animal perfor-
mance (Ferguson, 1975). The objectives of this
research were to determine the effect of varying levels
of formaldehyde treatment on rapeseed cake degrad-
ability and investigate responses of growing lambs to
supplemental feeding of protein sources of varying
degradability including T-RSC.
2. Material and methods
Two experiments were conducted to evaluate the
effect of conventional protein supplements on perfor-
mance of grazing sheep.
2.1. Experiment 1: formaldehyde treatment of RSC
Ground RSC in 1.0 kg batches was sprayed with
32% solution of formaldehyde at a rate of 1, 2, 3, 4 or
5 g HCHO/100 g CP. The solution was mixed thor-
oughly with the feed and stored in a sealed polythene
bag for 1 week to allow the reaction between protein
and HCHO to reach completion. The bags were then
opened and the contents were air-dried and processed
for in sacco rumen degradability (Orskov and McDo-
nald, 1979) to determine the degree of protection and
to select a suitable level of HCHO for bulk treatment
of RSC for use in the following feeding trial.
2.2. Experiment 2: growth trial with lambs
2.2.1. Animals and feeding
The study was conducted in four private sheep
¯ocks each consisting of 50±90 sheep in one of the
hilly regions (Swat Valley) of Pakistan. In each ¯ock,
20 female crossbred lambs (Kaghani  Rambouillet)
of about 12 months age and weighing 20±28 kg were
selected and randomly assigned to ®ve groups of four
lambs per group. All the animals were ear-tagged and
for group identi®cation, they were provided with neck
strips of ®ve different colours. All the animals in the
®ve ¯ocks including non-experimental animals were
drenched with a broad-spectrum anthelmintic (Systa-
mex, ICI, Pakistan) and vaccinated against entero-
toxaemia disease.
At each location, the ®ve groups were randomly
assigned the dietary treatments: (i) control (no supple-
ment); (ii) cottonseed cake; (iii) untreated rapeseed
cake (U-RSC); (iv) HCHO-treated rapeseed cake (T-
RSC) and (v) wheat bran. The supplements were fed
on an iso-nitrogenous basis to provide 20 g N/100 kg
BW. Based on the results of Experiment 1, RSC was
treated with 3 g HCHO/100 g CP. All groups, includ-
ing control, were fed 90 g per head per day of a
mineral mixture blended in ground wheat which sup-
plied calcium 7.8 g, sodium 2.1 g, sulphur 1.0 g, zinc
47 mg, copper 7 mg, cobalt 0.1 mg and iodine 0.6 mg.
The supplements were fed every morning, about 1 h
before releasing the sheep for grazing. At each farm,
®ve pens made of branches were used for group feeding
of the supplements. Initially, the lambs were gradually
adapted to their supplements over a period of 7 days
followed by an experimental period of 80 days.
 G. Habib et al. / Small Ruminant Research 41 (2001) 247±256
The lambs, along with other sheep in the ¯ock, were
grazed on hilly ranges (about 600 m above sea level)
from 8 a.m. to 5 p.m. daily. Samples of grasses from
different areas grazed by the ¯ocks were collected
fortnightly and analyzed for nutrient composition.
Samples of the supplements were collected fort-
nightly, and pooled for chemical analysis. Feed sam-
ples were air-dried at 608C, ground in a Wiley
laboratory mill to 1 mm particle size and were ana-
lyzed for dry matter (DM), ash, CP, crude ®ber (CF)
and ether extract (EE) according to AOAC (1990). In
sacco degradability of protein was determined with
the dacron bag technique procedure (Orskov et al.,
1980) using three rumen ®stulated wethers. The degra-
dation rate of protein (P) from the dacron bag at time
(t) in the rumen was described by the equation of the
form
P ˆ a ‡ b…1 ect † (1)
where a, b, and c were ®tted by an iterative least square
procedure (Orskov and McDonald, 1979). The expo-
nential constant a, b and c are measures of the rapidly
soluble protein fraction, the degradable protein frac-
tion and the rate constant for the degradation of ``b'',
respectively. The effective degradability (ED) of pro-
tein in the rumen was calculated as (Orskov and
McDonald, 1979)
bc
ED ˆ a ‡
c‡k
The value of k, which represents the estimated out¯ow
rate of particulate from the rumen, was chosen as 2.0,
4.0, 6.0 and 8.0%/h.
2.2.2. BW gain and wool growth
The lambs were weighed in the morning before
feeding at fortnightly intervals during the 80 days
experimental period. A portable digital Electro-weigh
scale (TPW, Australia Pty Ltd.) was used for measur-
ing the body weight. Wool growth in all the lambs was
estimated with the dyeband technique (Chapman and
Wheeler, 1963) using Durafur Black (Icianz Pty,
Imperial Industries Chemicals, Frankfurt, FRG). Each
time, fresh dye solution was prepared by dissolving
Durafur black 0.4 g for 90 min in 50 ml distilled water
and 15 min before use was added and mixed with
1.13-ml concentrated hydrogen peroxide. A line of the
dye was run with the help of a syringe and needle
249
along the skin for a distance of about 10 cm on the
right mid-side of the lambs. The dyeband was applied
at the start of the experiment and again 1 week before
the end of the experiment. On completion of the
experiment, the dyeband portion (staple) was clipped
with scissors and the lambs were shorn manually. The
total ¯eece and the staple weights were recorded. The
staples were cut along the dye lines for separating the
wool grown before and during the experimental per-
iod. After weighing, the staples were individually
enclosed in clean pre-weighed cotton bags and washed
four times as described by Williams and Chapman
(1966) using NI-400 (a non-ionic biodegradable liquid
detergent; Tec-Man International, Pakistan). The per-
centage of clean wool in the dyeband staple was used
to calculate the total clean wool produced during the
experimental period.
2.2.3. Staple length and ®ber diameter of wool
Fiber diameter was estimated using the optical ®bre
diameter analyzer (OFDA; BSC Electronics Pty Ltd.,
Australia). The clean dry wool samples were placed in
a standard atmospheric condition (20  2 C and
65  2% relative humidity) for 24 h. The sample
was then placed in minicore and 1.8±2.0 mm snippets
were cut under pressure. The snippets were placed in
the OFDA agitator to get a uniform spread over the
glass slide. The slide was examined using OFDA. The
staple length of wool samples was measured using a
ruler ®xed on a black velvet covered board. The staple
was straightened along the ruler after adjusting its base
to the zero mark on the ruler.
2.2.4. In vitro pepsin solubility of nitrogen (PSN)
Pepsin solubility of N in the feed samples was
measured using the method described by Siddons
et al. (1985), except that sample weight equivalent
to 25 mg N and incubation period of 4 h were used.
Feed samples, in duplicate, were weighed into 100 ml
conical ¯asks. Pepsin±HCl solution (50 ml) was added
to the ¯ask and incubated in a shaking water bath at
398C for 4 h. Duplicate blank ¯asks containing the
solution but no feed sample were also included in the
run. The pepsin±HCl solution was freshly prepared
before starting the incubation by dissolving 2 g pepsin
1:10,000 (Sigma Chemical Co.) in 1 l of 0.01 N HCl.
On termination of the incubation, the ¯ask contents
were ®ltered through Whatman ®lter paper no. 1 and
250 G. Habib et al. / Small Ruminant Research 41 (2001) 247±256

total N concentration in the ®ltrate was measured with

the Kjeldhal method (AOAC, 1990). Fraction of N
recovered in the ®ltrate was assumed as soluble. The
values were corrected for N in the blank.

2.3. Statistics

The mathematical model included ®xed effects

due to ¯ock and supplement and residual error
(SAS, 1999). Comparison of means was based on
the `Duncan's multiple range test'.

3. Results
Fig. 2. Ef®ciency of formaldehyde treatment in decreasing in
3.1. Formaldehyde treatment of RSC sacco degradability of protein (ISPD) in rapeseed cake.

Protein degradability of RSC treated with diffe-

rent levels of formaldehyde is illustrated in Fig. 1.
Protein degradability consistently decreased (P <
0:01) with increasing levels of HCHO for all incuba-
tion times. The treatment ef®ciency was calculated
as change in protein degradability per gram of
HCHO applied and the results are depicted in
Fig. 2. The treatment ef®ciency was maximum with
3 g HCHO/100 g CP and higher doses of the chemi-
cal above 3 g did not appreciably change the degrad-
ability further.
3.2. Chemical composition and protein
degradability of feed
Nutrient composition of the range grasses grazed by
the lambs during the experimental period was not
signi®cantly affected by location or sampling intervals
and the mean values along with those of feed supple-
ments are given in Table 1.
The in sacco protein degradability parameters of
CSC, U-RSC, T-RSC and WB are given in Table 2.
The instantly soluble fraction ``a'', potential degrad-
able fraction ``b'', the rate of degradation as percent
per hour ``c'' and the effective degradability at differ-
ent rumen out¯ow rates signi®cantly varied (P < 0:01)

Table 1
Nutrient composition of feed

Range CSC RSC WB

grasses

Dry matter (% DM) 40.14 89.89 91.03 87.49

Ash (% DM) 13.51 5.72 9.40 4.93
Crude protein (% DM) 12.03 26.37 36.56 13.02
Crude fiber (% DM) 27.71 30.66 11.90 8.83
Ether extract (% DM) 2.40 7.58 8.83 2.62
IVDOMD (%)a 63.32 ± ± ±
ME (MJ/kg DM)b 9.50 12.30 12.59 12.26
a
IVDOMD: in vitro organic matter digestibility in dry matter.
b
ME: metabolizable energy of range grasses was estimated as
Fig. 1. Effect of different treatment levels of formaldehyde 0.15 IVDOMD (MAFF, 1984) and that of supplements were
(HCHO) on in sacco protein degradability in rapeseed cake. reported earlier by Qureshi (1998).
 G. Habib et al. / Small Ruminant Research 41 (2001) 247±256 251

Table 2
In sacco protein degradability parameters and in vitro PSN in CSC, U-RSC, T-RSC and WB

Supplements a (%)a b (%)a c (%/h)a ED at different rumen outflow rates (%)b In vitro
PSN (%)c
2.0 4.0 6.0 8.0

CSC 39.02 48.14 10.53 78.20 72.43 73.63 65.68 83.27

U-RSC 52.54 33.45 15.15 88.70 84.90 81.85 77.83 80.87
T-RSC 37.68 53.17 5.32 75.25 66.40 60.70 56.75 81.56
WB 66.25 27.22 25.65 91.40 89.60 87.55 86.75 85.69
LSD (0.05) 10.98 16.41 3.05 3.24 4.13 3.69 3.69 ±
Significance level, P <0.001 <0.05 <0.001 <0.001 <0.001 <0.01 <0.001 ±
a
a, b and c are measures of the rapidly soluble protein fraction, the degradable protein fraction and the rate constant for the degradation of
``b'', respectively.
b
ED: effective degradability (Orskov and McDonald, 1979).
c
PSN: in vitro pepsin soluble nitrogen.

among the four supplements. The rate of degradation
and effective degradability were maximum in WB
followed by U-RSC and CSC and that of T-RSC
remained the lowest (P < 0:05). Formaldehyde treat-
ment of RSC signi®cantly reduced (P < 0:01) the ``c''
values from 15.15%/h in U-RSC to 5.32%/h in T-RSC
and also decreased (P < 0:05) the effective degrad-
ability of the cake at all rumen out¯ow rates (Table 2).
3.3. BW gain
Growth rates (g per day) in the lambs signi®cantly
varied with diets (P < 0:01) and ¯ocks (P < 0:01)
with no interaction of the two variables. Feeding of
the supplements, irrespective of type, resulted in
faster growth rates (P < 0:05) than the control
group. However, no difference in growth rates was
found among the four supplements (Table 3). BW gain
increased (P < 0:05) from 30 g per day on control diet
to an average 60 g per day on supplemented diets
(Table 3).
3.4. Changes in wool growth and wool
characteristics
Growth of clean wool in the lambs was affected by
diets (P < 0:05) and also varied (P < 0:05) among the
¯ocks. Lambs receiving the supplements produced
more (P < 0:05) clean wool than the control
group. The difference among the four supplements
was not signi®cant and the mean clean wool growth
was 3.4, 3.9, 4.1, 4.0 and 4.0 g per day on control,
CSC, U-RSC, T-RSC and WB diets, respectively
(Table 3). Staple length and ®ber diameter of wool
did not respond to the different supplements (Table 3)
and also did not vary among the ¯ocks.

Table 3
Mean results of BW gain, wool growth and wool quality in grazing lambs fed protein supplements of varying degradability

Supplements BW gain Clean wool production Wool quality

(g per day) (g per day)
Fiber diameter (mm) Staple length (cm)
Control 30.4 3.4 20.7 3.4
CSC 60.9 3.9 20.1 3.2
U-RSC 56.2 4.1 20.6 3.6
T-RSC 57.6 4.0 20.5 3.5
WB 63.7 4.0 20.4 3.9
Mean 53.7 3.9 20.5 3.5
CV (%) 35.8 17.7 7.5 25.9
LSD (0.05) 22.9 0.5 0.7 1.1
Significance level, P <0.01 <0.05 Non-significant Non-significant
252 G. Habib et al. / Small Ruminant Research 41 (2001) 247±256
4. Discussion
4.1. Formaldehyde treatment of RSC
Based on the dose-response relationship (Fig. 2),
3 g HCHO/100 g CP was found as an optimum level
for protecting RSC against rumen degradation and this
level was selected for treating the required quantity of
RSC for feeding the lambs during the growth trial. The
results on in vitro PSN (Table 2) suggested that HCHO
treatment did not affect the enzymatic digestibility of
N in RSC and remained the same in untreated and
treated RSC (80.87 and 81.58%, respectively). Pre-
viously, different workers have used different levels of
HCHO varying from 1.3 to 5.0 g/100 g CP for treating
animal feed (Ferguson, 1975; Kauffman and Lupping,
1982), which suggested that intrinsic factors in the
feed such as lipid, ®ber and other interfering com-
pounds would determine the treatment level of the
chemical. The RSC used in the present study was
mechanically extracted and contained a relatively
large quantity of residual fat (ether extract was
8.83% on DM basis; Table 1). Therefore, it is not
surprising that the desired level of protection was
achieved with higher doses of HCHO. Although,
not determined, it is assumed that formaldehyde
higher than 3 g/100 g CP would have caused over-
protection of protein in RSC. Ferguson (1975)
reported that treatment of feed with excessive HCHO
might render the protein non-digestible in the intestine
due to overprotection.
4.2. Nutritive value of grasses and protein
degradability of supplements
Chemical composition of range grasses grazed by
the sheep during the experimental period remained
constant and also did not vary among the locations
grazed by the four ¯ocks. During the entire experi-
mental period, CP on DM basis in range grasses
remained above the optimum level of 10% and ranged
from 11.7 to 12.7% (Table 1). This together with low
concentration of crude ®ber (mean 27.7% in DM)
ranked the grasses as good quality. Adverse effect of
the season on plant growth was apparently absent
during the experimental period, which commenced
during the last month of winter (January) and con-
tinued through the spring (March). This period is
considered favorable for plant growth and therefore
supported a constant high nutritive value of the range
grasses. In all four ¯ocks, the grazing practice was
almost the same and the areas grazed, although varied,
were situated in the same valley which may explain
the absence of variation in the nutritive value of the
grass samples collected from these areas.
On the basis of protein degradability results
(Table 2), the four supplements can be ranked as
WB < U-RSC < CSC < T-RSC. The daily intake
of undegradable protein supplied by the WB, U-
RSC, CSC and T-RSC were calculated as 5, 7, 10,
and 16 g, respectively, with a rumen out¯ow rate of
6%/h (Table 2). The high in vitro PSN in the supple-
ments (Table 2) suggested that maximum fraction of
the undegraded protein would be bioavailable.
4.3. BW gain
Feeding of the supplements increased (P < 0:05)
BW gain in the lambs. However, despite variation in
protein degradability among the supplements, growth
rates on the four supplements remained the same
(Table 3). Feeding of T-RSC did not increase growth
rates over those receiving U-RSC. The absence of
difference in growth rate among the supplements may
suggest that the total protein supply to intestine on
these diets was not markedly different. Post-ruminal
protein ¯ow is mainly composed of microbial protein
and undegraded dietary protein. It is assumed that high
degradable supplements (WB and U-RSC) would have
supported high microbial protein while the low
degradable supplements (CSC and T-RSC) may have
provided more undegraded protein ¯ow to intestine.
This is substantiated by the estimates given in Table 4.
Calculated microbial protein synthesis was maximum
on the diets containing WB, U-RSC and CSC, namely,
125, 100, and 105 g per day, respectively, while
undegraded dietary protein fraction was maximum
(38 g per day) on diet supplemented with T-RSC.
Nevertheless, the total protein supply to intestine
was close on the four supplements and calculated as
145, 136, 126 and 158 g per day on CSC, U-RSC, T-
RSC and WB, respectively. These observations clo-
sely agree with the recent ®ndings of Marghuzani et al.
(1999) who reported that N retention in sheep fed
CSC, U-RSC, T-RSC and WB was 4.63, 4.48, 4.72 and
6.94 g per day, respectively. It appears that small
 G. Habib et al. / Small Ruminant Research 41 (2001) 247±256 253

Table 4
Estimated availability of microbial protein and undegraded dietary protein in the small intestine of lambs fed protein supplements of varying
degradability

Supplements

Control CSC U-RSC T-RSC WB

Basal diet-pasture (g per day)
OMI intake (OMI)a 778 778 778 778 778
RDOMIb 464 464 464 464 464
Supplements (g per day)
OMI 0 131 98 98 233
RDOMI 0 96 80 59 204
Total diet (g per day)
OMI 778 909 876 876 1011
RDOMI 464 560 544 523 668
Microbial proteinc
g N/kg RDOMI 25 30 30 25 32
g N per day 12 17 16 13 20
g CP per day 75 105 100 81 125
Undegradable protein flow (g per day)d
Basal diet 29 29 29 29 29
Supplements 0 10 7 16 4
Total 29 39 36 45 33
Total intestinal protein (g per day) 104 145 136 126 158
a
Assumed a maximum DM intake of 3 kg/100 kg BW.
b
Rumen digestible organic matter intake.
c
Estimated after ARC (1980).
d
Estimated at a rumen out¯ow rate of 6%/h.

variation in the total intestinal protein was not suf®-
cient to elicit clear differences in growth rate of the
lambs. The difference in intestinal protein between
control (104 g per day) and supplemented diets (mean
141 g per day) explain the positive response in BW
gain to feeding of the supplements. The high correla-
tion coef®cient of BW gain with calculated microbial
protein (r 2 ˆ 0:77) versus undegradable protein
(r 2 ˆ 0:37) further suggest that feeding supplements
that maximize microbial protein production in the
rumen would be more relevant to enhance growth rate
in grazing sheep. These observations are in line with
some of the reported ®ndings. Schloesser et al. (1993)
replaced soybean meal with blood meal (a slowly
degradable N source) in a grass hay diet (8% CP)
fed to ewes and did not ®nd any difference in total
protein ¯ow to small intestine and BW gain or body
condition scores. Similar ®ndings in cows were also
reported by Klusmeyer et al. (1990) who could not ®nd
any in¯uence of substitution of soybean meal with
corn gluten meal on the total quantity of protein
¯owing to the small intestine. Hoaglund et al.
(1992) reported higher BW gain in sheep in response
to substitution of soybean meal with blood meal (a
slowly degradable N source). However, their diets
were not iso-nitrogenous and CP intake on blood meal
was higher than the other diets.
In the present study, lack of response to slow
degradable supplements (CSC and T-RSC) may be
also due to the small quantity of protein that escaped
undegraded (10 and 16 g per day, respectively;
Table 4). In the study of Schloesser et al. (1993),
soybean meal, blood meal or their combination sup-
plied 22 g daily CP or 20% of dietary CP intake, which
was not adequate to cause differences among the
supplements in BW gain of sheep. Similarly, in our
experiment, the supplements provided 35 g CP per
day or 25% of the total CP intake and may explain the
254 G. Habib et al. / Small Ruminant Research 41 (2001) 247±256
lack of difference in growth rate among the four
supplements.
Sheep in the present study had free access to grazing
on good quality range and hence consumed maximum
dry matter which may also have confounded the effect
of supplemental protein (Klusmeyer et al., 1990).
Chalupa (1975) reported that the potential for a rum-
inal escape protein source to increase production is
greater when the ruminal microbial amino acids sup-
ply is insuf®cient to meet the amino acids requirement
for growth and maintenance. In the present study,
microbial protein was calculated to have accounted
for more than 70% of the total intestinal protein
(Table 4) which may have confounded the response
to the small amount of escaped protein from CSC or
T-RSC.
4.4. Wool growth and quality
Growth of clean wool was lower in control sheep
and increased (P < 0:05) with feeding of the supple-
ments. Similar to the response noted in BW gain, wool
growth remained the same on the four supplements
(Table 4). Feeding of the supplements did not cause
any signi®cant change in the staple length or ®ber
diameter. Earlier work has shown that replacement of
soluble protein with slowly degradable protein source
either increased (Hoaglund et al., 1992; Thomas et al.,
1994) or did not change (Baldwin et al., 1993;
Kowalczyk et al., 1993) wool growth in sheep. In
studies, where positive response to undegradable pro-
tein was found in wool growth, the basal diet was of
low quality. Conversely, sheep receiving grass hay or
other good quality forages, did not respond to differ-
ences in protein degradability of the supplements.
These observations support the present ®ndings where
difference in degradability of protein supplements did
not in¯uence wool yield in sheep grazing range
grasses with CP contents above 10% in DM.
In agreement to the present ®ndings, Kowalczyk
et al. (1993) reported that feeding of HCHO-treated
rapeseed meal failed to improve wool yield and qual-
ity over the sheep given untreated meal. Recent work
in this laboratory (Marghuzani et al., 1999) also
demonstrated that N retention in sheep given U-
RSC or T-RSC supplements was the same, namely,
4.48 and 4.72 g per day, respectively. As discussed
earlier and shown in Table 4, small differences in the
availability of total protein (microbial and dietary) in
the intestine on the four supplements were apparently
not suf®cient to cause difference in wool growth and
quality among these diets. It can be assumed also that
the escaped protein may not have changed the balance
of amino acids in favour of wool growth and hence
negated the difference between degradable and unde-
gradable protein supplements. Merchen and Titge-
meyer (1992) reported that undegraded proteins
reaching the small intestine differ in their ability to
supply total absorbable amino acids. They further
postulated that two protein sources providing different
quantities of amino acids may not differ in the supply
of speci®c amino acids that are limiting performance.
This would mean that feeding proteins that escape
rumen fermentation might be of little bene®t if limit-
ing amino acids are de®cient in the supplement. These
observations may explain the absence of difference in
staple length and ®ber diameter among the present
diets.
It is inferred that for sheep with low performance
potential, dietary supplements to increase microbial
protein supply to intestine may be a relevant approach.
MacRae and Reeds (1990) reported that under certain
conditions, quantity of protein reaching the small
intestine is more important than quality. Based on
these observations McCollum and Horn (1990) sug-
gested that from a practical standpoint, it is easier to
in¯uence the total ¯ow rather than targeting amino
acids.
The present study commenced during late winter
and continued through spring (January±March). In the
area where the experiment was performed, sheep are
traditionally shorn three times a year, namely, April,
August and November. According to observations of
the local farmers, maximum ¯eece is obtained from
shearing in August followed by that of November
while shearing during April generally yields mini-
mum ¯eece weight. It is thus possible that the season
of low wool growth may have offset response to
protein supplements in wool traits. Several workers
(Cottle and Carter, 1992; Gambetta et al., 1992;
Woods and Orwin, 1988) suggested that response to
undegradable protein depended on season with mini-
mum change in wool growth and traits being recorded
during winter when wool growth was low. In the
present lambs, despite lack of differences in wool
quality among the diets, higher wool production
 G. Habib et al. / Small Ruminant Research 41 (2001) 247±256
(g per day) in supplemented lambs than control
(Table 3) may be attributed to higher energy intake
from the supplements.
5. Conclusions
Results of the present study led us to the conclusion
that supplementary feeding of protein increased BW
gain and wool growth but did not in¯uence wool
quality in lambs grazing range grasses with an average
CP contents of 12% in DM. Protein supplements of
varying degradability, supplying 25% of the total
dietary protein had equal substitution value for the
grazing lambs. Under the conditions of the experi-
ment, formaldehyde treatment of RSC did not improve
performance of the sheep over untreated RSC. It is
inferred that supplementary protein sources that max-
imize microbial protein synthesis in the rumen of
grazing sheep may be a relevant approach to enhance
animal performance.
Acknowledgements
The research was funded by the National Research
and Development Board, University Grants Commis-
sion, Islamabad, Pakistan. The cooperation of all the
four farmers is acknowledged for allowing their sheep
¯ocks to be used in the study. Berader Mian, wool
analyst, helped in providing the portable scale for
weighing of the lambs.
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