Literature review

Important fodder trees, shrubs and legumes in Lao PDR

Table 1 shows some important trees that have been identified as potential feed resources for ruminants in Lao PDR. Many fruit trees, fodder trees and shrubs can be found in nearly all parts of the country, such as Jackfruit, Banana, Jujube, Tamarind, Mango, Cassava and Erythrina.

Almost all the species of foliage presented above are important in tropical countries. This is due to their multipurpose use, such as fruits and young leaves for human consumption, trunks for wood and foliage for feeding animals (Dahlanuddin, 2001; Theng Kouch, 2003). It is important to collect more information on the use of these foliages as feeds for animals to be able to give advice to the farmers. Foliage from Cassava (Vongsamphanh, 2003; Phengvichith and Ledin, 2005; Xaypha, 2005), Gliricidia (Phimphachanhvongsod, 2001) and Water spinach (Pathoummalangsyand Preston, 2006) has been studied to some degree. However, there are still many types of foliage commonly used as feeds that have not been studied at all or in other environments and conditions such as foliage from Jackfruit (Artocarpusheterophyllus), which has been extensively researched in other tropical countries, but not in Lao PDR.

Nutritional characteristics of some local foliages

The chemical composition of some local foliages is presented in Table 2. The most important aspect of the chemical composition, the crude protein (CP) content, ranged from 78 g to 249 g/kg dry matter (DM). The protein content in fodder trees and legumes consists of both soluble and insoluble components, which are used both as an important source of nitrogen to increase rumen microbial activity and as by-pass protein for supplying amino acids to the lower gut of the host animal (Leng, 1997; Wanapat, 2003).

Most of the species can produce fruits, root and young leaves for human consumption and old leaves for animal feeding. Water spinach is one type of vegetable which is used traditionally in tropical regions for human consumption but is also used for animal feeding. Some plants such as Erythrina, Gliricidia and Leucaena belong to the Leguminosae family and have the ability to fix nitrogen, a property which could be useful for planting with another crop species.

The CP content is, as earlier mentioned, an important characteristic of foliage. Water spinach has a high CP content compared to other species, which has also been noted by several other authors (Bunthaand Ty, 2006; Hiep and Preston, 2006; Hue and Preston, 2006; Pathoummalangsy and Preston, 2006; Phiny and Kaensombath, 2006; Samkol et al., 2006). Erythrina, Gliricidia and Leucaena have also a high CP content (Kibria et al., 1994; Baidya et al., 1995; Phimphachanhvongsod, 2001; Simbaya, 2002; Aregheore and Perera, 2004; Gregorio et al., 2005).

According to Simbaya (2002) the presence of anti-nutritional substances, in particular condensed tannins, in fodder trees and shrubs can limit animal performance, especially when tree/shrub foliages are fed in large quantities. Jackfruit leaves had higher content of condensed tannins compared to the other foliages, which has also been found by Mui et al. (2001) and Van et al. (2005). However, the high tannin content in Jackfruit did not seem to be a deciding factor for feed intake (Kongmanila Daovy et al (2007b)). Other authors have also found that the tannin content had a poor relationship with digestibility of DM and N in foliage from Leucaena varieties and Calliandra (McSweeney et al., 1999). This is due to the fact that tannins are composed of many different compounds with different toxicity.

The use of foliages as a feed resource for small ruminants, advantages and limitations

The traditional feeding systems for ruminants are based on native grasses available in the natural grassland and forests and the main constraints to ruminant production are lack of feed, particularly in the dry season (Phimphachanhvongsod, 2001; Phengsavanh, 2003; Xaypha, 2005; Kongmanila Daovy et al (2007a)). The main feed supplement for animals in the dry season is rice straw and various crop residues, with are usually limited in both quantity and quality, especially concerning the protein content: Feeding diets based on these by-products often results in poor performance (Hang, 2005). However, there are many types of foliage that are available around the farms and in the forests, which have proved to be good feed resources for small ruminants, especially in the dry season (Binh and Lin, 2005).

Simbaya (2002) reported that the foliages from fodder trees, legumes and shrubs have a high protein content, ranging from 14% to 25%, and the advantage of using fodder trees, legumes and shrubs as a source of feed for ruminant animals is that supplementation of forage up to about 35 %, does not seem to have any effect on the intake of fibrous feed materials. The DM intake is often increased by feeding green fodder or fodder legumes as supplementation (Dixon and Egan, 1987; Khang, 1999). Theng Kouch et al. (2005) and Phengvichith and Ledin (2005) reported that foliage from Kapok or Cassava can increase feed intake and weight gain when used as a supplement to natural grasses or low or medium quality grasses.

The type and variety of foliages that can be used for feeding depend on climatic conditions and the vegetation. For example there were 42 different forages available to goats in Indonesia (Dahlanuddin, 2001) and 20 browse species was important for goats in Burkina Faso (Sanon et al., 2007), while in lowland areas in Lao PDR only 15 species were mentioned (Kongmanila Daovy et al (2007a)). Foliages were not used to any great extent by the farmers, which is probably due to the low indigenous knowledge about foliages and also how important the foliages are considered to be as animal feeds. However, there were many suggestions from smallholder farmers about foliages than can be used for goats e.g. Tamarind foliage can increase the milk yield of female goats after kidding, Kapok foliage could be a good feed resource for male goats, but it is not so good to feed females after kidding, as it was said to cause a weakness in their legs. Tamarind and kapok were also reported to be popular feeds for goats by farmers in Cambodia (Theng Kouch, 2003).

In recent years, many kinds of foliages from trees have been studied as supplements for livestock (Leng, 1997) and in the various farming systems of Asia and the Pacific region, fodder trees and shrubs play an important role in livestock production (Chen et al., 1991). Some of them were offered as sole feeds and most of these foliages had a high feed intake and digestibility and gave a positive nitrogen balance in small ruminants, such as foliage from Acacia, Cassava, Erythrina, Flemingia, Gliricidia, Jackfruit, Melastoma, Mulberry, Muntingia, Leucaena and Water spinach (Baidya et al., 1995; Phimphachanh-vongsod, 2001; Mui et al., 2002; Theng Kouch, 2003; Lin et al., 2003; Samkol, 2003; Hang, 2005; Van et al., 2005; Buntha and Ty, 2006; Pathoummalangsy and Preston, 2006; Phengvilaysouk and Kaensombath, 2006). Many authors have studied foliages as protein sources to supplement with other feeds: Acacia, Banana and Trichantera (Duyenet al., 1995; Hove et al., 2001); Cassava (Khang, 1999; Bunyeth, 2005; Hang, 2005; Phengvichith and Ledin, 2005; Xaypha, 2005); Erythrina, Gliricidia and Leucaena (Dana et al., 2000; Kusmartono, 2002; Aregheore and Perera, 2004; Ajayi et al., 2005); Jackfuit (Duyen et al., 1995; Kusmartono, 2002), Kapok (Theng Kouch et al., 2005); Mango and Fig (Ajayi et al., 2005). In general the foliages can improve the feed quality when feeding roughages of low nutritive value and can also increase DM and N intake.

There are, however, also limitations when using foliage from fodder trees, legumes and shrubs as feeds. According to Leng (1997) the main limitation to effective utilization of these foliages as feed for ruminants is high content of tannins and other anti-nutrients such as saponins, cyanogens, mimosine and coumarins. These compounds are also known to have other detrimental effects, which may range from reduced animal performance to neurological effects and increased animal mortality rates. Hove et al. (2001) reported that tannins can limit animal performance by suppressing intake and digestibility of forages. However, tannins can also have a beneficial effect on small ruminants, in helping to increase the amount of rumen undegradable protein and supplying essential amino acids to the gut, which may happen especially when the level of tannin is low, 2% to 4% (Fassler and Lascano, 1995). Forage legumes containing condensed tannins can also affect parasite infections in small ruminant (Mui et al., 2005). There are some simple methods to reduce negative effects of tannins e.g. with post-harvest processing techniques such as sun-drying, wilting or ensiling of forages, which can reduce in the tannin content up to 55 % to 77 % of the initial value (Makkar and Singh, 1993; Jackson et al., 1996; Ben Salem et al., 1999; Bunyeth, 2005).

Feed intake and digestibility of some tropical foliage species

Jackfruit (Artocarpus heterophyllus) foliage has high potential as a protein source and can be used both as a sole feed and as a supplement. It can improve feed intake and give a positive nitrogen balance when fed to growing goats. The DM intake of Jackfruit foliage was higher compared to other foliage species, 4.2% to 4.9% of body weight (BW). Jackfruit can be fed as a supplement to grasses, sugarcane or low feed intake foliages such as Acacia and Flemingia, and can then increase total feed intake by more than 50% compared to non-supplementation. Jackfruit foliage also replaced concentrate by up to 50% with no effect on DM and CP intake (Duyen et al., 1996; Mui et al., 2001; Mui et al., 2002; Theng Kouch, 2003; Van et al., 2005; Phengvilaysouk and Kaensombath, 2006; Kongmanila Daovy et al (2007b)). The high feed intake of Jackfruit foliage can be due to the structure of the leaves, which are soft and have a high nutritive value (Van and Ledin, 2002). The condensed tannins probably have a beneficial effect on the nutritional metabolism through enhancing rumen escape, increasing efficiency of nitrogen recycling to the rumen and improving microbial growth efficiency (Waghorn, 1990). Values of 51.4% to 67.6% and 45.2% to 54.2% of DM and N digestibility, respectively, have been found for Jackfruit foliage (Mui et al., 2002; Theng Kouch, 2003; Phengvilaysouk and Kaensombath, 2006; Kongmanila Daovy et al (2007b)).

Erythrina (Erythrina variegata) belongs to the Leguminosae family, and has a high CP content when compared to other fodder tree and shrub species. Erythrina is used as a protein supplement to improve the feed quality of diets based on maize stover or low quality roughage in the diet of ruminant livestock in the Pacific Island countries (Aregheore and Perera, 2004). Supplementation with Erythrina leaves to 50% of the total diet can increase feed intake, DM or CP digestibility with 50 %, 30 % and 20 %, respectively, compared to feeding urea-treated maize stover alone. This can be explained by the fact that leguminous leaves are more rapidly digested than other roughages and therefore promote higher intakes (McDonald et al., 2002). Goodchild and McMeniman (1994) also reported that supplementation of low quality roughage with forage legumes alleviates N deficiency, thereby improving the rate of degradation of the basal diet and the fractional outflow of liquid matter from the rumen, and hence feed intake.

Kapok (Ceibapentandra) is a multipurpose tree, which contains high levels of CP. The high DM and nutrient intake, apparent digestibility and a positive N retention when feeding Kapok foliage, is comparable to feeding Erythrina and Jackfruit foliage (Kongmanila Daovy et al (2007b)). Nhan (2000) reported that the DM intake of growing goats was 4.6% of BW and the highest DM digestibility (76.0%), was from the Kapok foliage compared to Leucaena, Sesbania or Hibiscus (75.9%, 73.6% or 68.0%), respectively and also higher than the result in Kongmanila Daovy et al (2007b), 3.8% of BW and 57.2% for DM intake and digestibility, respectively. This difference may be due to the effect of the breed or age of the animals and also development stage of the leaves. Theng Kouch et al. (2005) concluded that supplementation with Kapok foliage appears to be a viable option to improve the nutritional status of goats during periods of the year when grazing is restricted.

Foliage from Fig (Ficus racemosa), Jujube (Ziziphus jujuba Mill) and Mango (Mangifera indica) can be used as sole feeds for goats (Papers II and III), but there is no available research about these foliages. The results from the present study show that the intake, digestibility and N retention were lower compared to Erythrina, Jackfruit and Kapok foliage. This is due to the effect of chemical composition and physical properties of these foliages. However, most of the foliages used in Kongmanila Daovy et al (2007b) were preferred by goats and used by smallholder farmers (Kongmanila Daovy et al (2007a)), especially the foliage from the Ziziphus family. Even the foliages of lower nutritional value can be potential feed resources for goats, especially in the dry season.

Water spinach (Ipomoea aquatica)has high nutritive value, especially high CP content, for humans and also for animals. Buntha and Ty (2006) and Pathoummalangsy and Preston (2006) reported that goats had lower DM intake but higher N intake and digestibility when fed Water spinach as a sole diet, compared to Cassava foliage or Guinea grass. Feeding Water spinach increased DM or N digestibility and N retention when fed as a supplement to Cassava foliage. Reed et al. (1982) reported that the protein in the Water spinach forms complexes with the condensed tannins in the cassava foliage. As described in Kongmanila Daovy et al (2007b) Water spinach can also be used as a supplement to Fig, Jujube and Mango foliage.

Selective behaviour of goats

Feeding behaviour is very important when studying feed intake and feeding management, and the selective behaviour at feeding is especially important for goats (Van, 2006). Feeding behaviour and choice of feed varies according to animal species, probably due to special anatomical features and physiological characteristics but there is also a great variation in feeding behaviour between individuals (Kongmanila Daovy et al (2007b)I). For instance, goats seemed to be selective, whereas llamas tried to eat quickly (Dumont et al., 1995) and the goats had a higher level of intake compared to the llamas, due to the fact that the diet of the llamas contained less browse than goats. Van (2006) concluded that goats generally eat more slowly, spend more time for selection and eating and less total time ruminating than sheep. Sanon et al. (2007) found that the sheep selected 20 species, while the goats used a wide range of browse species, in total 31 different species during the whole observation period.

Goats have an upper mobile lip, which permits selective grazing, and a low body mass and low metabolic requirement, which make them able to survive in harsh environments. The versatile feeding behaviour of goats comes mainly from their physical body structure (bipedal stance, mobility of upper lip and vigorous grazing), as well as their variable rumen microflora (Steele, 1996). However, there are many factors which are important for selective behaviour of goats, such as feed quality, type and way to present the feed, competition between animals. Total DM feed intake increased linearly as the number of animals in the pen increased (Van, 2006).

As reported in Kongmanila Daovy et al (2007b)I there were large individual differences in the feeding behaviour of goats. The total time spent eating during 9 hours ranged from 178 to 390 minutes when the animals were fed all six foliage species in a large pen. Abijaoude et al. (2000) reported that the eating time of goats could range from 6 to 9 hours per day and kids fed whole sugar cane as a sole roughage spent 283 to 397 minutes per day eating (Van et al., 2002). Time spent eating was greater during the day, especially after sunrise and before sunset (Dumont et al., 1995; Jarrige et al., 1995; Van et al., 2005). Browse was the most important forage class for goats, constituting 51% to 90% of the diet (Sanon et al., 2007). Goats prefer browsing and they have the unique ability to utilize forage resources that can not be utilized effectively by other ungulates (Dumont et al., 1995). However, goats, like all animals, express a degree of nutritional wisdom, since they select plants or plant parts higher in nutrients than the average in pasture (Papachristou and Nastis, 1993). Papachristou et al. (2005) found the goats on pasture selected diets significantly higher in CP and in vitro organic matter (OM) digestibility and lower in neutral detergent fiber (NDF) and acid detergent fiber (ADF) content than samples collected by hand plucking mimicking foraging of goats. Provenza (1996) reported that animals increase preferences for a feed when nutritional needs are better met and, goats choose and consume the most nutritious feed available (Van Soest, 1982; Fajemisin et al., 1996).

The feed quality, type and mode of feeding are factors in confined conditions that affect total time spent eating and diet selection in goats. The hanging method was found to be the best way to improve feed intake and eating rate (Theng Kouch, 2003; Samkol, 2003; Van et al., 2005; Phengvilaysouk and Kaensombath, 2006). The highest eating time was spent on the foliages from Erythrina, Jackfruit and Kapok compared to Fig, Jujube and Mango (Kongmanila Daovy et al (2007b)I). The proportion of leaves plus petioles and stem or the structure of the leaves can probably affect the selection. For example, Erythrina foliage has a soft and smooth structure in all parts of the foliage, while Mango foliage has a high DM content, low CP content and hard leaves. Van et al. (2005) reported that the total eating time was longer for kids fed Jackfruit foliage than for kids fed Flemingia or Acacia foliage (385, 297 or 202 minutes, respectively), which was probably due to the better intake characteristics of Jackfruit. The foliage from the Ziziphus family was preferred by goats (Ngwa et al., 2000; Sanon et al., 2007), which was also the case in Kongmanila Daovy et al (2007b)I, where some goats spent quite a long time on Jujube (Ziziphus jujuba Mill), more than 40% of the total eating time per day.

It can be concluded that when penned goats spend a great part of the daytime eating and that the DM intake can be improved by feeding many different species of browse, since the preference seems to vary between individuals.
 

Conclusions

• Many types of forage species can be used as a source of protein for ruminant animals in the lowland areas of Lao PDR. Local foliages and simple processing methods, particularly drying, need to be introduced to smallholder farmers by research and extension, so that they can increase the use of these valuable feed resources for improving animal productivities.
• High nutritive values were noted in several of the foliages. The highest CP content was found in Erythrina, while other foliage species had medium to low values. Erythrina is a good fodder tree, which should be promoted to plant with other crops for fixing nitrogen and for animal feeding. Feed quality affected DM intake, digestibility and N retention. Fig, Jujube and Mango foliage had lower digestibility and intake characteristics than Erythrina, Jackfruit and Kapok foliage. Some foliages need to be further studied e.g. Erythrina and Kapok. Water extractable DM can be a method of analysis for predicting DM digestibility and N retention.
• A low DM intake and high digestibility were found in the water spinach foliage. Water spinach can be used as a supplement to foliages of low nutritive value to improve diet quality and animal performance.
• Goats seem to have large individual differences in feed preferences, eating rate and selective behaviour. Erythrina, Jackfruit and Kapok were the feeds most preferred in the study. To improve feed quality and quantity, animal management and productivity, the local foliages and feeding behaviour of animals need to be further studied.
   

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