Effects of supplementation with rumen fermentable carbohydrate and sources of 'bypass' protein on feed intake, digestibility and N retention in growing goats fed a basal diet of foliage of Tithonia diversifolia

Khamparn Pathoummalangsy and T R Preston*

Department of Livestock and Fisheries, Faculty of Agriculture
NationalUniversity of Laos, Vientiane Capital, Lao PDR
khamparnp@yahoo.com.au
*UTA-TOSOLY, Socorro, Santander, Colombia
 

Abstract

Experiment 1. Four growing male goats fed ad-libitum Tithonia diversifolia hay were allocated in a 4*4 Latin square design to supplements of cassava chips (1% of BW), fresh mulberry leaves (1% of BW as DM), both cassava chips and mulberry leaves (0.5% BW of each on DM basis) or no supplement.

Supplementation did not affect the intake of the Tithonia hay thus total DM intake was higher on all the supplement treatments. Crude protein intake was highest for the treatment of mulberry leaves. N intake was lowest in the goats given Tithonia and Tithonia with cassava chips. Lowest N retention was on the Tithonia diet expressed as absolute amounts or as percentages of N intake or N digested. N intake on the diet with cassava chips was low, but the efficiency with which the N was retained was higher than on all the other diets. Live weight changes were in the same order as those for N retention with highest value on the diet with supplements of both cassava chips and mulberry leaves.

Experiment 2. Four growing male goats were assigned to a 2*2 factorial design in a 4*4 Latin square. The factors were: ad-libitum fresh tithonia foliage or tithonia hay; and supplementation with cassava root chips + mulberry leaves (1% BW of each on DM basis). Total DM intake for goats fed fresh Tithonia foliage was higher than when the Tithonia hay (plus jackfruit leaves) was fed. Supplementation with mulberry leaves and cassava chips increased DM intake compared with no supplement, reaching 46.5 g/kg live weight. Coefficients of apparent digestibility for DM, organic matter, crude protein and NDF were lowest when the Tithonia was fed as hay. Supplementation improved the coefficients on Tithonia hay but not on fresh Tithonia. Urine volume was much higher on fresh Tithonia than on sun-dried Tithonia. When the diets were supplemented the urine volume was decreased on the fresh Tithonia but not on the sun-dried Tithonia. N retention did not differ between fresh and sun-dried Tithonia but was increased when the Tithonia was supplemented with cassava chips and mulberry leaves.

Experiment 3. Four growing goats were assigned in a 4*4 Latin square to evaluate four sources of bypass protein (banana leaves, jackfruit leaves, erythrina leaves and mulberry leaves) at levels of 2% BW (DM basis) with a basal diet of ad libitum fresh Tithonia foliage. Urine volumes were highest on the banana and erythrina diets and reflected the rates of excretion of N in the urine. N retention was highest on the mulberry-supplemented diet which was not different from the diet with jackfruit. The lowest value was on the diet with banana leaves. N retention on the erythrina supplemented diet appeared to be only slightly less than on the diet with jackfruit. The weight gains broadly followed the values for N retention being much less on the Tithonia supplemented with banana leaves.

The findings of these three experiments support the hypothesis that, in goats fed tree foliages as the basal diet, digestibility indicates nutritive potential of the diet but actual performance is dependent on appropriate supplementation to increase the amino acids absorbed by the animal.

Keywords: Banana leaves, bypass protein, digestibility,  dried cassava root, Erythrina, intake, Jackfruit, Mulberry, N retention, Tithonia diversifolia.

Introduction

Nowadays, many new kinds of foliages, shrubs, and tree fodders have been researched and introduced to farmers by scientists. Wild sunflower (Tithonia diversifolia) is one of the new foliages, which in Laos is considered to be a valuable green manure. In Western Kenya, it is renowned as a component of agro-forestry systems as it is rich in N, P and K which are essential for soil fertility. Biomass from Tithonia has proven to be valuable in improving soil fertility for crop production in areas constrained by soil N, P and K deficiencies (Lijzenga, 1998). It has been reported that addition of foliage of Tithonia diversifoliato the cropping area leads to double the yield of the crops and that it is more effective than urea when applied at the same N rate (Jama et al 2000; Sanchez 2001).

In this study we are concerned with the potential use of the Tithonia as forage for live stock. There were now several researches relevant to its use for livestock production. Mahecha and Rosales (2005) reported that the crude protein content in the foliage of Tithonia was 24.2% in DM basis and that about 40% of the protein was soluble. The percentage (in DM) of NDF was 35.3% and ADF 30.4%. The rumen degradable protein (in sacco) was very high (about 90% in 48h) (Rosales, 1996, cited by Mahecha and Rosales, 2005). Recently, Wambui et al (2006) reported that the daily weight gain of goats fed urea-treated maize stover as basal diet and supplemented with Tithonia foliage was about 82.6g/day and it was greater than when Caliandra and Sesbania foliages were used. . The low content of tannins reported for Tithonia foliage (Wambui et al 2006) supports the idea that the protein may well be highly soluble with poor "bypass" characteristics (Preston and Leng, 1987). The fact that loss of nitrogen in the urine was higher for Tithonia than for Calliandra or Sesbania in the study of Wambui et al (2006) supports this suggestion that rumen ammonia levels are high on Tithonia, resulting in high excretory levels of urea in the urine. Similar high urine losses of nitrogen were reported for water spinach (Ipomoea aquatica) fed at 40% (Pathoummalangsy and Preston, 2006) or as the sole diet of goats (Buntha and Ty 2006). Water spinach is similar to Tithonia in having high protein content and high digestibility but low tannin content.

Based on the above observations it is hypothesized that the limiting factors to efficient utilization of Tithonia as the sole diet of goats is the high solubility of the protein resulting in high rumen ammonia levels, which have to be excreted via the urine. The final result is that the supply of metabolic protein to the animal is reduced. The two ways to improve goat performance on Tithonia foliage as the main feed would thus appear to be: to feed easily fermentable carbohydrate to utilize the high ammonia levels or to supplement with foliages with known rumen bypass characteristics.

Sources of readily fermentable carbohydrate

Cassava chips are reported to be excellent sources of rumen fermentable carbohydrate as they are rich (70-80%) in rapidly fermentable starch (Kearl, 1982; Sommart et al 2000; Vearasilp and Mikled, 2001; Gomez and Waldivieso, 1983).). Chanjula et al (2003) found that cassava chips were better than cassava root waste, sweet potato roots, maize corn meal and rice bran, as a means to optimize use of NPN for rumen microbial protein synthesis. Sathapanasiri et al (1990) observed that starch of cassava is highly degraded in the rumen (94%) and completely digested in the whole tract. A similar result has been reported by Sommart et al (1991).

Foliages with bypass protein characteristics

Mulberry leaves are traditionally used as food for silkworms in sericulture (Subbarayappa and Bongale, 1997). Mulberry leaves are very rich in protein and minerals and low in fiber (Fotadar and Dandin, 1997). Jegou et al (1994) reported DM intakes in lactating goats of 4.18 percent of live weight and that digestible DM was from 78.4 to 80.8 percent. Growth rates in goats were improved when the basal grass diet was replaced with ensiled mulberry leaves with highest N retention when the ensiled mulberry leaves were the sole content of the diet (Nguyen Xuan Ba et al 2005). Vu Chi Cuong et al (2005) reported that fresh mulberry leaves could replace cottonseed meal in the diets of growing cattle with no reduction in live weight gain. The protein in cottonseed meal is known to be one of the best sources of bypass protein for ruminants (Promkot and Wanapat, 2003; Weixian et al 1994) thus it is reasonable to conclude that mulberry leaves are also rich in bypass protein.

Mineral content is high in mulberry foliage and no anti-nutritional factors or toxic compounds have been identified (Makkar et al 1989; Makkar and Becker, 1998, Singh et al, 2000). Additionally, the mulberry samples analysed by the UFAG Laboratories in Switzerland, from the high zone of Cartago in the Central Plateau of Costa Rica, gave on a DM basis, 22 percent of crude protein (CP), 19 percent of crude fibre (CF), 2.3 percent of ether extract, 50 percent of by-pass protein and an estimated 1.48 Mcal/kg of net energy for lactation (cited by Boschini, 2002).

Foliage from the Jackfruit (Artocarpus heterophyllus) has been recommended as a basal diet for lactating goats and growing kids, especially during the dry winter season when grasses are in short supply (Duyen et al 1996). Moreover, Phengvilaysouk and Kaensombath (2006) reported that the retention of N in goats only on Jackfruit foliage was 2.8 g/day which is equal to about 18 g/day of protein which, assuming the body contains 20% protein, would be reflected in a rate of live weight gain of the order of 100 g/day.

Camero and Franco (2001) presented that feeding leguminous fodder that is high in protein can improve rumen fermentation parameters leading to increased digestibility and intake of low quality fibrous feeds, and hence improved animal production. Erythrina variagata is a leguminous tree that is more palatable than Gliricida sepium and supported higher milk yields in goats due mainly due to differences in intake (Camero and Franco, 2001). Supplementing foliage of Erythrina poeppigiana to goats fed King grass and banana fruit resulted in linear increases in milk production (Esnaola et al 1990). In an experiment with dairy cows, foliage of Erythrina poeppigiana supported better economical returns than use of soybean or fish meal as protein supplement (Pezo et al 1990).

In Laos, banana leaves are traditionally used as material for packaging human food. Duyen et al (1996) fed banana leaves as protein sources for lactating goats given a basal diet of rice straw and sugarcane tops and concluded that leaves from banana supported better milk production than the leaves of Trichanthera gigantea but were inferior to foliage from Jackfruit.

The hypothesis that was tested in this study was:

Three experiments were carried out using growing goats housed in metabolism cages so as to measure feed intake, apparent digestibility and N retention with different combinations of cassava root chips and tree foliages. In Experiment 1, the basal diet was sun-dried Tithonia foliage and the supplements were cassava root chips and mulberry leaves. In Experiment 2: a comparison was made of sun-dried Tithonia foliage and the fresh foliage with and without cassava chips plus mulberry leaves. In Experiment 3: the basal diet was fresh Tithonia foliage and four different sources of leaves (from banana, Erythrina variegata, mulberry and Jackfruit).
 

Materials and methods

Location and climate

The experiments were conducted at The Faculty of Agriculture farm, National University of Laos (Nabong campus), sited in Parksarp Mai Village and 32 km from the City centre, Xaythany District, Vientiane Capital, Lao PDR. The environmental temperature during the experiments ranged from 18 to 32 degrees centigrade.

Experimental animals and housing

Four young goats about 5-7 months of age were purchased from local farmers for each of the experiments (different animals for each experiment). The animals were de-wormed with Ivomectin and vaccinated against Foot and mouth disease and Pasturellosis two weeks before the beginning of each experiment. They were housed and fed in individual metabolism cages built from wood and designed for collection of feed residues, faeces and urine separately. The floor area in each cage was 90*90 cm, and they were raised 85 cm from the floor.

Treatments and experimental design

Experiment 1. The treatments consisted of sun-dried Tithonia foliage, which was fed as basal diet (offered at 120% of recorded intake). The treatments arranged as a 4*4 Latin square were:

• TH: Tithonia foliage hay

• THC: TH supplemented with dried sliced cassava root at 1 % of body weight (BW) on DM basis.

• THM: TH supplemented with mulberry leaves at 1% of BW on DM basis.

• THCM: TH supplemented with cassava root chips at 0.5% of BW and mulberry leaves also at 0.5% of BW, both on DM basis.

Experiment 2.The factors compared in a 2*2 factorial arrangement within a 4*4 Latin square were:

Processing of Tithonia foliage

• Sun-dried/hay (TH) or fresh (TF)

Supplementation

• Mulberry (M) leaves plus cassava root chips (C) each at 1% of BW DM basis or no supplement

Experiment 3.The basal diet was fresh Tithonia foliage supplemented with the following leaves each at 2% of BW (DM basis):

• M: Mulberry

• E: Erythrina

• J: Jackfruit

• B: Banana

In each experiment there were 4 periods each of 12 days, the first 7 days for adaptation to the new feed and the last 5 days for data collection.

Feed and feeding

Experiment 1

Tithonia foliage was collected from waste land at the sides of the highway in Xiengkhouang province in the upland area of Laos. The foliage, which comprised stems (about 50 cm long) with attached leaves, was dried in the sun for 3 to 5 days. It was then packed into polypropylene bags and transported by bus to the Experimental farm in the University. Fresh cassava root was purchased from local farmers in Vientiane province. The roots were sliced by hand or machine and dried under sunlight for 4 to 5 days. Leaves of mulberry and jackfruit were collected daily from trees in the Crop Science department of NUOL.

Experiment 2

The Tithonia hay, cassava chip, mulberry and jackfruit leaves were obtained according to the same procedure used in Experiment 1. Fresh Tithonia foliage was from shrubs which had been established 3 months previously, in an area close to the Goat house, using as plant material stem cuttings brought from Xiengkhouang province.

Photo 1. Goat fed Tithonia foliage hay in the feed trough	Photo 2. Goat fed Tithonia foliage hay and supplemented with cassava root chips and mulberry	Photo 3. Goat fed fresh Tithonia foliage (hanging outside the pen) and supplemented with cassava root chips and mulberry

Experiment 3

Tithonia foliage was harvested from the re-growth of the plants established in Experiment 2. Leaves of mulberry, jackfruit and banana were collected from the Crop Science Department. The leaves of Erythrina were from trees growing in the experimental area.

In all treatments in all experiments the goats had free access to mineral blocks and clean water. The first feed at 7.00 am comprised the supplements and part of the basal diet. The remainder of the basal diet was offered in two feeds at 12.00 am and 4.00 pm. The Tithonia hay was fed loose in the feed trough (Experiments 1 and 2), In the case of the fresh Tithonia foliage (Experiments 2 and 3), this was hung in bunches suspended in the feed trough. Mulberry and Jackfruit were always given as leaves in the trough (all experiments) as was the banana in experiment 3.

Data collection

Live weight was recorded at the beginning of each 12 day period and at the end of the experiment. During the data collection period, samples of the Tithonia foliages offered and refused were separated into stem and leaves (containing attached petioles). Representative samples of each component were analysed daily for DM and the dried material retained until the end of each period when it was analysed for N, ash and NDF.

30 ml of 25% sulphuric acid was put into the urine collection container to maintain the pH below 4.0. Urine volume was measured daily. A representative sample of urine (10% of the total) was taken each day. At the end of the 5-day collection period, the urine samples were mixed and analysed for N.

Faeces were weighed and sampled twice a day in the morning (at 6:30) and late afternoon (16:30). The samples were combined for each animal and 10% of the total kept in the freezer at -18ºC for DM, N, NDF and ash determination at the end of the period.

Chemical analyses

Chemical analyses of diets and faeces were undertaken following the methods of AOAC (1990) for ash and N. The measurement of NDF followed the procedure of Goering and Van Soest (1970). The DM content was determined using the microwave method of Undersander et al (1993). Urine was analyzed for pH by a digital meter with glass electrode. The N content of urine was determined by the AOAC (1990) procedure.

Statistical analyses

The data were subjected to analyses of variance according to the general linear model of the Minitab software (Minitab release 13.31, 2000). When the "F" test was significant (P<0.05), the means were separated using the Turkey comparison option in the Minitab software. In Experiments 1 and 3, sources of variation were treatments, animals, periods and error. The model was as follows:

Y_ijk = µ + T_i + P_j + C_k + _ijk

Where: Y_ijk = observation, µ = population mean, T_i = diet effect, P_j = periods effect, C_k = goat effect, and _ijk = residual error.

For experiment 2, the 2 x 2 factorial analysis of variance was used. The sources of variation were: periods, animal, processing, supplements, interaction processing*supplements and error. The model was:

Y _ijpsp*s = µ + P_i + C_j + T_psp*s + _ijpsp*s

Where: Y_ijkl = observation, µ = population mean, P_j = periods effect, C_j = goat effect, T_k = supplement effect, P_l = processing effect, T_k*P_l = interaction processing*supplements and _ijkl = residual error.

Experiment 1

Results

Experimental feed characteristics

The proportion of leaf in the sun-dried Tithonia hay was 76.6%.

Voluntary feed intake and digestibility

During the introduction of the goats to the experimental diets it was observed that they ate very little of the Tithonia hay. However, when some leaves of jackfruit were also offered the intake of the hay increased. It was therefore decided to include the Jackfruit leaves at the level of 0.5% of live weight (DM basis) in all the treatments.

Supplementation did not affect the intake of the Tithonia hay thus total DM intake was higher on all the supplement treatments (Table 2; Figure 1). As expected, intakes of crude protein were higher for the diets containing mulberry leaves; however, expressed as concentration in diet DM, the values were lower for the diets with cassava chips. Apparent digestibility of DM tended to be increased on all the supplemented diets, while coefficients for crude protein tended to be higher for those diets with mulberry foliage which contained more protein.

Figure 1. DM intake of goats fed Tithonia hay (and small amount of fresh jackfruit foliage) alone
or supplemented with dried cassava root, fresh mulberry leaves or both cassava root and mulberry

Figure 2. Apparent digestibility in goats fed Tithonia hay (and small amount of fresh jackfruit foliage)
alone or supplemented with dried cassava root, fresh mulberry leaves or both cassava root and mulberry.

Nitrogen balance

N intake was lowest in the goats given Tithonia and Tithonia with cassava chips (Table 3). Lowest N retention was on the Tithonia diet (TF) expressed as absolute amounts or as percentages of N intake or N digested (Figure 3). Although N intake on the diet with cassava chips was low, the efficiency with which the N was retained was higher than on all the other diets.

Figure 3.N balance of goats fed Tithonia hay (and small amount of fresh jackfruit foliage)
 alone or supplemented with dried cassava root, fresh mulberry leaves or both cassava root and mulberry leaves

A 12 day period is very short for measuring reliable data on live weight change. However, the values recorded were in the same order as those for N retention with highest value on the diet with supplements of both cassava chips and mulberry leaves.

Discussion

The crude protein in the leaves of the dried Tithonia (20%) was similar to the value of 21% in DM reported by Mahecha and Rosales (2005) and the value of 22% recorded by Jama et al (2000) and Wambui et al (2006). In the case of mulberry the crude protein content was lower (14%) than the value (20 to 22% in DM) reported by Vu Chi Cuong et al (2005) and the value of 18% in DM observed by Nguyen Xuan Ba et al (2005).

Despite the high crude protein content and the relatively high apparent digestibility of DM and crude protein of the Tithonia foliage, the DM intake and N retention were low. Supplementation with cassava chips increased both intake and N retention, despite the fact that crude protein as percentage of DM was lower on this diet than on the Tithonia (TH) diet. The implications are that the effect of the cassava chips was to increase microbial protein production in the rumen through the availability of easily fermentable carbohydrate. Chanjula et al (2003) proposed that cassava chips facilitate the achievement of optimal utilization of energy and protein, especially NPN, for rumen microbial protein synthesis. Hoover and Stoke (1991) and Khampha and Wannapat (2006) also indicated that the high rate of digestion of carbohydrate in cassava chips was a major factor controlling the energy for growth of the rumen microbes.

The increase in voluntary intake and in N retention when the Tithonia hay was supplemented with mulberry leaves would appear to be a direct result of more protein becoming available at tissue level, presumably because part of the mulberry protein escaped the rumen fermentation and passed directly to the abomasum and intestine. Boschini (2002) on the basis of laboratory analysis claimed that 50% of the protein in mulberry leaves was by-pass protein. The fact that growth rates of cattle on a basal diet of molasses and rice straw were maintained at 550 g/day when mulberry leaves replaced whole cotton seeds (which are known sources of bypass protein) implies similar bypass properties in mulberry protein as in cotton seeds (Vu Chi Cuong et al 2005). Supporting evidence for the bypass protein characteristics of mulberry leaves are the reports of Oviedo (1995) (cited by Bernavides et al 2000) that dairy heifers grazing star grass (Cynodon nlemfuencis) had the same growth rates when supplemented with concentrates or only mulberry leaves. Gonzalez et al (1996) (cited by Bernavides et al (2000)) observed that young Romosinuano bulls in total confinement and fed a basal diet of elephant grass (Pennisetum purpureum) had growth rates of 40, 690, 940 and 950g/day when mulberry foliage intakes (as DM) were 0, 0.90, 1.71 and 2.11 percent of live weight.

Conclusions

• The overall findings of this experiment support the original hypothesis that "The nutritive value of Tithonia foliage for goats would be increased by supplementing it with: (i) a source of readily fermentable carbohydrate; and ii) a tree foliages that would supply bypass protein".

Experiment 2

Results

Feed composition

The fresh leaves and stems of Tithonia had higher crude protein and lower NDF that the dried leaves and stems (Table 4). The leaves in both cases were richer in crude protein than the leaves of mulberry and jackfruit.

Intake, digestibility and N balance

Total DM intake in the goats fed fresh Tithonia was higher (P<0.02) than when the Tithonia hay (plus jackfruit leaves) was fed (Table 5). The DM intake of fresh Tithonia was 371 g/day compared with 300 g/day for the combined Tithonia hay and Jackfruit leaves. Supplementation with mulberry leaves and cassava chips increased DM intake compared with no supplement, reaching 46.5 g/kg live weight, which is more than the maximum intake expected in young goats in the tropics (Devendra and McLeroy, 1982; Steele, 1996).

 

 

Figure 4. DM intake of goats fed Tithonia hay (with 1% of LW as jackfruit leaves) or fresh Tithonia foliage
alone or supplemented with cassava root chips plus mulberry leaves

There were significant interactions on apparent digestibility between effects of processing of the Tithonia foliage and supplementation with cassava root chips + mulberry leaves (Table 7). Coefficients of apparent digestibility for DM, organic matter and crude protein were lowest when the Tithonia was fed as hay. Supplementation improved the coefficients for DM and crude protein on Tithonia hay but not on fresh Tithonia (Figures 5 and 6).

There were significant interactions on urine output, N excretion and N retention between effects of processing of the Tithonia foliage and supplementation with cassava root chips + mulberry leaves (Table 8). Urine volume was much higher on fresh Tithonia than on sun-dried Tithonia. When the diets were supplemented the urine volume was decreased on the fresh Tithonia but not on the sun-dried Tithonia.

Faecal N was lower and urine N was higher for diets with fresh Tithonia than for those with Tithonia hay, the differences being greater in the absence of supplementation. N retention did not differ between fresh and sun-dried Tithonia but tended (P=0.059) to be increased when the Tithonia was supplemented with cassava chips and mulberry. When the ANOVA analysis was done for main effects (Table 8) the effect of supplementation on N retention was significant (P=0.016).

Discussion

The higher crude protein content in the fresh compared with sun-dried Tithonia foliage, and the lower values for apparent digestibility, were probably due to chemical changes resulting in losses of valuable nutrients during the drying process (McDonald et al 2002). The higher intakes of Tithonia foliage fed fresh rather than as hay probably reflected in part the way these two feeds were offered (the hay was loose in the trough; fresh foliage was hung in bunches). It has been shown in several experiments that intakes and often digestibility are higher when tree foliages are hung as bunches rather than being offered only as leaves (Kouch et al 2005; Keopaseuht et al 2004; Van et al 2005; Phengvilaysouk and Kaensombath, 2006).

The higher urine volume on fresh compared with sun-dried Tithonia appears to be related to the need to excrete a greater quantity of N on the fresh Tithonia as the N content of the urine was similar in both (about 0.8%).

The greater responses to supplementation on the fresh compared with the sun-dried Tithonia indicates that the fresh foliage has a high potential as ruminant feed (high crude protein content and high digestibility) but that the expression of the potential (in N retention and live weight gain) requires supplementation to improve rumen microbial growth (addition of cassava chips) and to increase the supply of bypass protein (mulberry leaves).

Conclusions

• The situation presented by fresh Tithonia foliage is similar to what happens when a low digestibility forage (eg: cereal straw) is treated with urea or supplemented with rumen nutrients, or a high digestibility diet (eg: molasses or sugar cane juice) is supplemented with urea and minerals. In such cases the potential nutritive value is increased but the expression of this improvement in live weight gain requires supplementation with bypass protein (Weixian et al 1994; Sath 2007 in the case of cereal straw; Martin and Preston (1968), cited by Preston and Leng (1987) for molasses and Sanchez and Preston (1980) for sugar cane juice).

Experiment 3

Results

Feed composition

The chemical composition (Table 9) of the diet ingredients indicated that the CP content in fresh Tithonia leaf was the highest but lower than in Experiment 2 with similar high values in Erythrina leaves. The level of condensed tannins was highest in the banana and jackfruit leaves. The level in the Tithonia leaves (57.1 mg/kg DM) is much higher than was reported for Tithonia foliage (5.6 mg/kg DM) in Kenya (Wambui et al 2006).

Intake, digestibility and N balance

There were no differences in intake and apparent digestibility among the diets, except for crude protein apparent digestibility which was lowest for the jackfruit supplemented diet (Table 10).

Urine volumes were highest on the banana and erythrina diets and reflected the rates of excretion of N in the urine which were also highest for these two diets (Table 11). Faecal N did not differ among the diets. N retention was highest on the mulberry-supplemented diet which was not different from the diet with jackfruit. The lowest value was on the diet with banana leaves. N retention on the erythrina-supplemented diet appeared to be only slightly less than on the diet with jackfruit. The weight gains broadly followed the values for N retention being much less on the Tithonia supplemented with banana leaves (Figure 12).

Live weight change

There was a close relationship between N retention and live weight gain (Figure 13).

Discussion

It was observed that the goats consumed the jackfruit and mulberry leaves within 30 to 40 minutes. In contrast they took most of the day to eat the banana and erythrina leaves. These observations are different from those reported by Kongmanila (2007) when intakes, N retention and live weight gain were always better for erythrina than other foliages. The difference was that in the present study only the leaves of the erythrina were fed while Kongmainila (2007) hung the complete foliage (leaves attached to the stems) in the feed trough. It has been shown that for most foliages the intakes and digestibility are higher when the foliage is hung in the trough than when only the leaves are fed (Kouch et al 2005; Keopaseuht et al 2004; Van et al 2005; Phengvilaysouk and Kaensombath, 2006).

The finding that N retention was highest with mulberry leaves as the supplement is in agreement with the results reported by Bakshi and Wadhwa (2006) where, in a comparison of a range of tree leaves as the sole diet, the highest N retention was observed with mulberry leaves.

The fact that there was a poor relationship between coefficients of apparent digestibility and the N retention (R² = 0.25; P = 0.068) indicates that digestibility was not the constraint to goat performance on Tithonia foliage. This conclusion is similar to the one drawn in experiment 2 that: digestibility is indicative of the nutritive potential of the diet but that appropriate supplementation to improve protein flow to the intestine is the real determinant of performance. In experiment 3. apparent digestibility on the Tithonia foliage supplemented with banana leaves was high and similar to that on the other diets. However, N retention and live weight gain with the banana-supplemented diet were significantly lower than for the diets supplemented with jackfruit, erythrina and mulberry leaves. All of these latter foliages are considered to have protein in a form which allows it to bypass the rumen to the lower gut (Preston and Leng 1987).

The close relationship between the quantities of N retained and the observed changes in live weight (Figure 13) is an indication that the experimental procedures were an appropriate means of assessing the nutritive value of the experimental diets.

Conclusions

• There was no relationship between DM intake and coefficients of apparent digestibility and the N retention in goats fed fresh Tithonia foliage supplemented with leaves from banana, jackfruit, erythrina and mulberry.

• N retention for diets with banana leaves was less than 50% of the values recorded with the other leaf supplements.

• This supports the findings in the two previous experiments that digestibility only indicates nutritive potential of the diet and that actual performance is dependent on appropriate supplementation to increase the amino acids absorbed by the animal.

Acknowledgements

The senior author would like to express sincere thanks to the MEKARN Master Course project financed by Swedish International Development Authority (SIDA), Department for Corporation SAREC. Besides, she expresses her gratitude to National Agriculture and Forestry Research Institute and Faculty of Agriculture, National University of Laos particularly Department of Livestock and Fisheries for the facilities to carry out the experiment.

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