Printer Friendly

Utilization of rice straw and different treatments to improve its feed value for ruminants: a review.

INTRODUCTION

In tropical zones in the world, ruminants depend on year-round grazing on natural pastures or the animals are fed with cut grass and crop residues. Most of these areas face seasonal dry periods in which the availability of pasture decreases and also its quality by a reduction in the content of digestible energy and nitrogen. Due to the fact that in these areas rice straw is abundantly available from cultivating rice, farmers offer rice straw as the main roughage source to their animals. This is particularly the case in Southeast Asian countries such as Thailand, Vietnam and Indonesia (NARC newsletter, 2004). Feeding only rice straw does not provide enough nutrients to the ruminants to maintain high production levels due to the low nutritive value of this highly lignified material. The high level of lignification and silicification, the slow and limited ruminal degradation of the carbohydrates and the low content of nitrogen are the main deficiencies of rice straw, affecting its value as feed for ruminants (reviewed by Van Soest, 2006). As rice straw is poorly fermented, it has low rates of disappearance in the rumen and low rates of passage through the rumen, reducing feed intake (Conrad, 1966). By treating rice straw with urea or calcium hydroxide or by supplementing rice straw with protein, intake, degradability and milk yield can be enhanced, compared to feeding untreated rice straw alone (Fadel Elseed, 2005; Wanapat et al., 2009).

In past years, several studies have been reported on the physical and chemical characterization and utilization of rice straw as ruminant feed (Shen et al., 1998; Abou-El-Enin et al., 1999; Vadiveloo, 2000; 2003). In addition, numerous methods of physical, chemical and biological treatments have been investigated, including supplementation with other feed stuffs or components in order to improve the utilization of rice straw by ruminants (Reddy, 1996; Karunanandaa and Varga, 1996a,b; Shen et al., 1999; Vu et al., 1999; Liu and Orskov, 2000; Selim et al., 2004). Rice straw is usually fed untreated without supplements in spite of the fact that many methods for improved utilization of rice straw have been developed and recommended. There are several reasons for farmers not to apply the already developed methods for improved utilization of straw, such as physical, socio-economic conditions and practical reasons (Devendra, 1997). In general, the use of rice straw as an animal feed as well as its treatment is always an economic decision.

The aim of this contribution is to provide an overview of existing knowledge on how to treat rice straw to increase its feed value for ruminants. Emphasis is placed on new approaches using enzymes and fungi and combinations of these with other, more classical, treatments, such as physical and chemical treatments.

AVAILABILITY OF RICE STRAW

Agriculture plays a significant role in the world to feed the growing human population. Therefore, land for crop production will be used more intensively for human food production and consequently animal production will rely on feeding the by-products from the food produced for human consumption. This especially will be the case for rapidly growing economies in several parts of Asia, increasing also the demand for meat and milk at a high rate. Thus, many countries in this area urgently need to increase their livestock production.

Many by-products from the human food industry have in common a high biomass, low crude protein content of approximately 3 to 4% and high content of crude fiber of approximately 35 to 48% (Devendra, 1997). In many developing countries these fibrous crop residues, such as cereal straws, sugarcane tops, bagasse, cocoa pod husks, pineapple waste, etc., are usually fed to ruminants.

Approximately 80% of the world's rice is grown by small-scale farmers in many developing countries including South East Asia (Table 1) and it is common to use rice straw for animal feeding. Devendra and Thomas (2002) mentioned that rice straw is the principal crop residue fed to more than 90% of the ruminant livestock in this area. The calculated utilization of rice straw for animal feed in South East Asia, including China and Mongolia, was 30-40% of the total rice straw production (Devendra, 1997). Rice straw is especially important during periods when other feeds are inadequate. In general, the maximum intake of rice straw by ruminants is about 1.0 to 1.2 kg per 100 kg live weight (Devendra, 1997). In Southeast Asia rice straw is mainly utilized by swamp buffaloes and cattle with adult live weights of 350 and 200 kg respectively (Devendra, 1997). With an average intake of 1 kg of rice straw per 100 kg live weight, this gives a total annual intake of 1.28 and 0.73 t for buffaloes and cattle respectively. The annual requirements for the current populations of swamp buffaloes and cattle are therefore 51.2 and 89.4 million t respectively and a total requirement of 140.6 million t, representing 30.4% of the total annual available rice straw.

GENERAL CHARACTERISTICS OF RICE STRAW

The chemical composition of rice straw

The chemical composition of rice straw varies between varieties and growing seasons, with higher nitrogen and cellulose contents in early-season rice compared to others (Shen et al., 1998). The chemical and mineral compositions of rice straw, from Chinese data, are illustrated in Table 2 and 3.

Nutritive quality of rice straw

Rice straw consists predominantly of cell walls, comprised of cellulose, hemicellulose, and lignin. To break down these components cellulase, hemicellulase and ligninase are required (Schiere and Ibrahim, 1989). These enzymes are not produced by the animals themselves but the reticulorumen of ruminants maintains microorganisms that do produce cellulase and hemicellulase. However, lignin cannot be broken down in the rumen due to the lack of ligninase. Even if lignin could be degraded in the rumen it would not provide much energy for the animals. Lignin, however, has important effects on livestock production through effects on degradability and feed intake. Theoretically, lignin located between the cellulose microfibrils is regarded as the most abundant natural aromatic organic polymer that plays a role in resisting compressing forces, providing protection against consumption by insects and mammals, and also inhibiting the rate and degree of microbial degradation (Iiyama et al., 1990). Silica, one element of the rice cell walls, can be present in high concentrations ranging from 5% to 15%, depending on the rice variety (Vadiveloo, 1992) and the availability of this mineral in the soil (Agbagla-Dohnani et al., 2003). Silica reduces palatability and the degradability of rice straw in the rumen due to its direct action in preventing colonization by ruminal microorganisms (Bae et al., 1997; Agbagla-Dohnani et al., 2003). The role of silica on the quality of rice straw was also reviewed by Van Soest (2006), in an attempt to put into perspective the problems of silicon metabolism.

Besides cell wall polymers, rumen organisms need other nutrients for growth and metabolism (Hoover, 1986). Since rice straw does not contain enough sugars, amino acids and minerals for efficient microbial growth, feeding ruminants with only rice straw, without any supplementation of the other required nutrient sources, will result in poor performance of the animals (Doyle et al., 1986). The combination of low intake, low degradability, low nitrogen content and an unbalanced mineral composition means that rice straw alone may not even meet the animal's maintenance needs. Poor degradability is caused by a series of factors (Schiere and Ibrahim, 1989). The fiber is very difficult to degrade, which is partly an intrinsic characteristic of the straw fiber. The degradation of the straw fiber is also complicated by the poor functioning of the rumen due to the unbalanced availability of nutrients, the low protein content, the lack of easily available energy and the low content of essential minerals such as P and S. Hence, due to the low degradability and the poor rate of degradation, animals will tend to consume less. The mechanism regulating voluntary intake of low quality feeds, such as rice straw, is still not fully understood. The generally accepted theory of feed intake regulation for poor quality roughages is that the capacity of the rumen to process the feed is the major factor determining voluntary feed intake (Conrad, 1966; Baile and Forbes, 1974). The rumen processing capacity is characterized by rumen fill, the rate of degradation of potentially degradable matter and the rate of passage out of the rumen. Devendra (1997) summarized that the main determinants of intake and degradability of rice straw depend on their morphological characteristics, such as the proportion of the different plant parts (leaves and stems), their chemical composition and the distribution of the different chemical components in the tissues, their relative amounts of cell contents and cell walls and the physical and chemical nature of the cell walls. These factors influence the chewing behavior of animals and the extent of fragmentation in the reticulorumen.

Rice straw contains a relatively high proportion of leaf (60%), compared to other cereal straws such as barley (35%), oats (43%) and wheat (20-41%) (Theander and Aman, 1984). Leaves of rice straw contain less NDF than the stems, but more ash and acid-insoluble ash, resulting in a lower in vitro dry matter digestibility (IVDMD) of the leaves (50-51%) compared to the stems (61%) (Vadiveloo, 2000). In goats, Phang and Vadiveloo (1992) observed an in vivo dry matter digestibility of 56.2% for rice leaf and 68.5% for the stem. However, treatment with a 4% urea solution for 21 d increased the IVDMD of the leaf fraction more than that of the stem fraction (Vadiveloo, 2000). Since rice straw consists of approximately 60% leaves (Vadiveloo, 1995), which are less degradable than stems, improving the feed value of rice straw should focus on improving the degradability of the leaves.

POSSIBLE STRATEGIES TO IMPROVE RICE STRAW UTILIZATION

Basically, the key to improving the use of crop residues for ruminants is to overcome their inherent barriers to rumen microbial fermentation. In the case of rice straw, the important factors that restrict bacterial degradation in the rumen are its high levels of lignification and silicification, and its low contents of nitrogen, vitamins and minerals. To improve the feeding value of rice straw, the straw can be treated with different means and methods and other required nutrients can be supplied to the ration of the animal. Strategies to improve the utilization of rice straw are summarized in Figure 1 (after Ibrahim, 1983).

[FIGURE 1 OMITTED]

Physical treatment

Crop residues can be ground, soaked, pelleted or chopped to reduce particle size or can be treated with steam or X-rays or pressure cooked. Uden (1988) observed that grinding and pelleting of grass hay decreased dry matter degradability in cows from 73 to 67%, which was mainly due to a decreased fermentation rate (9.4-5.1%/h) and decreased total retention time of the solids from 73 to 54 hours, resulting in an increased intake (Stensig et al., 1994). Liu et al. (1999) reported that the use of steam treatment in a high pressure vessel at different pressures and for a range of different treatment times increased the degradation in vitro in rumen fluid after 24 h and the rate of degradation, but could not enhance the potential degradability of the fibrous fractions (NDF, ADF and hemicellulose). Physical treatments of crop residues have received an appreciable amount of research. Many of these treatments are not practical for use on small-scale farms, as they require machines or industrial processing. This makes these treatments in many cases economically unprofitable for farmers as the benefits may be too low or even negative (Schiere and Ibrahim, 1989). However, small machines to grind or chop rice straw may be feasible.

Chemical treatment

Chemicals to improve the utilization of rice straw may be alkaline, acidic or oxidative agents. Among these, alkali agents have been most widely investigated and practically accepted for application on farms. Basically, these alkali agents can be absorbed into the cell wall and chemically break down the ester bonds between lignin and hemicellulose and cellulose, and physically make the structural fibers swollen (Chenost and Kayouli, 1997; Lam et al., 2001). These processes enable the rumen microorganisms to attack more easily the structural carbohydrates, enhancing degradability and palatability of the rice straw (Prasad et al., 1998; Shen et al., 1999; Selim et al., 2004). The most commonly used alkaline agents are sodium hydroxide (NaOH), ammonia (N[H.sub.3]) and urea. Chemical treatments appear to be the most practical for use on-farm, as no expensive machinery is required, the chemicals are relatively cheap and the procedures to use them are relatively simple. However, the chemicals themselves are not harmless and safety precautions are needed for their use.

NaOH treatment : Several NaOH treatment methods to improve the use of crop residues for ruminant feeding have been developed as reviewed by Jackson (1977), Berger et al. (1994) and Arieli (1997). The principal advantages of the different NaOH treatment methods are increased degradability and palatability of treated straw, compared to untreated straw (Chaudhry and Miller, 1996; Vadiveloo, 2000). However, NaOH is not widely available as a resource for small-scale farmers and may be too expensive to use. In addition, the application of NaOH can be a cause of environmental pollution, resulting in a high content of sodium in the environment (Sundstel and Coxworth, 1984).

N[H.sub.3] treatment: Treatment of straw with anhydrous and aqueous ammonia, urea or other ammonia-releasing compounds has been widely investigated to improve degradability (Abou-EL-Enin et al., 1999; Selim et al., 2002; Fadel-Elseed et al., 2003). The principle of ammonia treatment is supposed to be similar to that of NaOH treatment. Ammonia treatment not only increases the degradability of the straw, but also adds nitrogen (Abou-EL-Enin et al., 1999) and preserves the straw by inhibiting mould growth (Calzado and Rolz, 1990). Besides, improvement in degradability of structural carbohydrates, ammonia treatment is an effective means of reducing the amount of supplemental nitrogen, reducing the costs of purchasing protein-rich feedstuffs, and enhancing acceptability and voluntary intake of the treated straw by ruminants. Although comparative studies in improving the energy value of straw have shown that ammonia treatment is less efficient than NaOH (Liu et al., 2002), its use may be more profitable for farmers as the added ammonia serves as a source of nitrogen. In a previous study using sheep, Selim et al. (2004) treated rice straw packed in polyethylene bags for 4 weeks with gaseous ammonia (3 g N[H.sub.3] per 100 g dry matter). The excess ammonia was removed before offering the straw to animals. The ammonia treatment increased the N content in the rice straw from 8.16 to 18.4 g [kg.sup.-1] (CP content increased from 51 to 115 g [kg.sup.-1]). The ammonia treatment slightly decreased the NDF content from 571 to 551 g [kg.sup.-1], because of dilution with the additional N, but increased the ADF content from 303 to 327 g [kg.sup.-1], indicating that the cell wall properties were changed. Moreover, the physical strength of ammoniated rice straw was significantly lower than that of the untreated straw. In addition, the proportion of small feed particles tended to be higher and stimulated more attachment and growth of the rumen bacteria (Selim et al., 2002). The reduced particle size and the increased attachment sites could lead to subsequent increased microbial colonization and digestion. So, ammonia treatment increases feed value by making the cell wall more available for the rumen microorganisms and also the increased N content improves microbial growth.

Urea treatment: Rice straw can also be treated with urea, which releases ammonia after dissolving in water. For practical use by farmers, urea is safer than using anhydrous or aqueous ammonia and also provides a source of nitrogen (crude protein) in which straw is deficient (Schiere and Ibrahim, 1989). Since urea is a solid chemical, it is also easy to handle and transport (Sundstel and Coxworth, 1984) and urea can be obtained easily in many developing countries. In addition, urea is considerably cheaper than NaOH or N[H.sub.3]. Vadiveloo (2003) reported that rice varieties with a low degradability responded better to urea treatments than higher quality straw, increasing the in vitro dry matter degradability from 45 to 55-62%. Urea treatment may therefore be most suitable for small-scale farmers to improve the quality of straws, particularly varieties showing a low degradability. In the past, numerous investigations involving urea treatment of rice straw, with or without additional supplementation, were performed not only in the laboratory (Reddy, 1996; Shen et al., 1998; 1999; Vadiveloo, 2003) but also in field trials (Prasad et al., 1998; Vu et al., 1999; Akter et al., 2004). Pradhan et al. (1997) showed that addition of Ca(OH)2 to urea improved the IVDMD. Sirohi and Rai (1995) demonstrated that a combination of 3% urea plus 4% lime at 50% moisture for 3 weeks incubation time was the most effective treatment for improving degradability of rice straw. Using urea is regarded as a practical and available method in livestock production, especially in developing countries, as it is relatively cheap, adds nitrogen to the ration and is relatively safe to work with.

Urine treatment: As urine contains urea, urine can be used as a source of urea and ammonia to improve the quality of rice straw. Urine can be sprayed on the straw in a similar way as is done with urea solutions (Dias da Silva, 1993) and can provide a nearly equal improvement of the degradability and nitrogen content as other methods of ammonia treatment (Dias da Silva, 1993; Schubert and Flachowsky, 1994). However, research on this subject has been quite limited and there is currently inadequate information available to define clearly the conditions to optimize urine treatment (Dias da Silva, 1993). Moreover, the use of urine is hampered by the difficulty of separation of urine from feces in ruminant husbandry. This also makes the use of urine rather unhygienic and therefore not advisable to use, although its use is without costs for farmers and urine is normally available in excess.

Lime treatment: Lime (CaO/Ca[(OH).sub.2]) is a weak alkali agent with a low solubility in water. It has been reported that lime can be used to improve the utilization of straw and also can be used to supplement the ration with calcium, which has been found to be in a negative balance in cattle fed only rice straw (Hadjipanayiotou, 1984; Pradhan et al., 1997; Chaudhry, 1998). Soaking and ensiling are two methods of treating straw with lime. Although lime treatments increase the degradability of straw, the dry matter intake decreases, due to a reduced acceptability of the treated feed by animals. Pradhan et al. (1997) reported that ensiling rice straw with 4 or 6% Ca[(OH).sub.2] showed a higher IVDMD than using 4 or 6% urea. However, mould growth was noticed in the Ca(OH)2 treated straw. It was suggested that a combination of lime and urea would give better results than urea or lime alone. This combination has the advantage of an increased degradability and an increased content of both calcium and nitrogen. Additive effects of lime and the other alkali agents have been demonstrated (Saadulah et al., 1981; Hadjipanayiotou, 1984). The use of lime may be safer and more cost effective to use than NaOH.

Feeding rice straw supplemented with other components

As rice straw is low in nitrogen and difficult to degrade, it is obvious that supplementation of a ration of rice straw with a protein source and a more easily accessible energy source will improve the performance and production of the animals. Supplementation of a ration of rice straw with protein, energy and/or minerals may optimize rumen function, also maximizing utilization of the rice straw and increasing intake. Chenost and Kayouli (1997) emphasized that it is primarily necessary to supply the rumen microorganisms with the nutritive elements needed for self-multiplication as well as for degradation of the cell walls of straw, leading to suitable conditions for maintenance of good cellulolysis. Different supplements can be used, such as concentrates, molasses, multi-nutrient blocks, green leaves, crop residues and locally available by-products. In the case of high-yielding dairy cows, the supplements can be the major part of the ration where fibrous feed only serves to supply the rumen with enough fiber. Warly et al. (1992) showed in a field trial that a ration of rice straw supplemented with soybean meal increased both degradability and intake. Because of the poor quality of untreated rice straw, supplementation easily can increase milk production, as shown for supplementation with cottonseed meal (Wanapat et al., 1996) and with an urea-molasses-multi-nutrient block (Vu et al., 1999; Wanapat et al., 1999; Akter et al., 2004).

Biological methods

The use of fungi and/or their enzymes (Table 4) that metabolize lignocelluloses is a potential biological treatment to improve the nutritional value of straw by selective delignification, as mentioned in the review by Jalc (2002). Nevertheless, it is currently too early to apply this method in developing countries due to the difficulties and lack of technology to produce large quantities of fungi or their enzymes to meet the requirements. There are also a number of serious problems to consider and overcome (Schiere and Ibrahim, 1989). For example, the fungi may produce toxic substances. It is also difficult to control the optimal conditions for fungal growth, such as pH, temperature, pressure, O2 and CO2 concentration when treating the fodder. With recent developments in fermentation technology and alternative enzyme production system, the costs of these materials are expected to decline in the future. Hence, new commercial products could play important roles in future ruminant production systems (Beauchemin et al., 2004).

White-rot fungi treatment : White-rot fungi, belonging to the wood-decaying basidiomycetes, as lignocellulolytic microorganisms are able to decompose and metabolize all plant cell constituents (cellulose, hemicellulose and lignin) by their enzymes (Eriksson et al., 1990). Many species of white-rot fungi which are effective lignin degraders have been used to assess their ability to improve the nutritive value of fodder for ruminant nutrition (Yamakava and Okamnto, 1992; Howard et al., 2003). Their extracellular lignin-modifying enzymes consist of lignin-peroxidase (LiP), manganese-dependent peroxidase (MnP), laccase (phenol oxidase) and [H.sub.2][O.sub.2]-producing oxidase (aryl-alcohol oxidase; AAO and glyoxaloxidase) (Kirk and Farrell, 1987; Arora et al., 2002; Novotny et al., 2004; Arora and Gill, 2005; Lechner and Papinutti, 2006). An overview of investigations using fungi and their determined enzymes is given in Table 4.

Some white-rot fungi are able to decompose free phenolic monomers and to break the bonds with which lignin is cross-linked to the polysaccharides in rice straw (Chen et al., 1996), enhancing IVDMD (Karunanandaa et al., 1992; 1995; Karunanadaa and Varga, 1996a,b; Fazaeli et al. (2006). Karunanandaa et al. (1995) reported the effect of incubation of rice straw for 30 days with three white-rot fungi, showing that Pleurotus sajor-caju enhanced IVDMD, in both leaves and stems of rice. However, entire rice straw (leaf and stem) treated with Cyathus stercoreus had the highest IVDMD compared to the other fungi (Karunanandaa et al., 1992).

Using white-rot fungi to increase the degradability of straw is often at the expense of easy assessable carbohydrates, such as cellulose and hemicellulose, resulting in less degradable feed for ruminants (Karunanandaa et al., 1995; Karunanandaa and Varga, 1996a, b; Jalc, 2002). In fact, cellulose and hemicellulose losses during the initial part of incubation with fungi are rather common, but losses due to mycelial growth depend on the fungus species (Table 5). After the initial period of incubation, some white-rot species preferably attack lignin, without degrading cellulose and hemicellulose. Rodrigues et al. (2008) were able to extract the enzymes from white-rot fungi that are responsible for breaking down the bonds in lignin and within the matrix of cell wall carbohydrates, but without also extracting enzymes affecting hemicellulose and cellulose. Using these enzymes on wheat straw the in vitro NDF degradability (IVNDFD) increased.

Although the use of fungi to improve the feed value of rice straw is not new, progressing research and new knowledge offers new challenges and possibilities. Fungi can be selected that preferably attack lignin and not the structural carbohydrates in the cell walls. Once these species are identified, mycologists can breed even better strains. The most desirable situation would be that the mushrooms of the fungi are edible and can be harvested by farmers, after which the remaining straw can be fed to their herd. There are some edible white-rot fungi, like Pleurotis ostreatus. However, much research is needed to achieve these goals. The most suitable white-rot species have to be identified and breeding programs will possibly be needed to improve their characteristics. Also, the optimal conditions to incubate straw with a fungus have to be investigated, not only with the purpose of harvesting quality mushrooms, but also achieving optimal feeding quality of the remaining straw-fungi mixture. To achieve optimal feed qualities of the straw, incubations with fungi in combination with other treatments, such as physical and chemical treatments, have to be investigated.

Exogenous fiber-degrading enzyme treatment : Most commercially available exogenous fiber-degrading enzyme products consist of cellulases and xylanases, as produced for non-feed applications. Commercial enzymes used in the livestock feed industry are generally of fungal (mostly Trichoderma longibrachiatum, Aspergillus niger, A. oryzae) or bacterial origin (Table 6). Twenty-two commercial enzyme products were examined for biochemical characteristics and for in vitro ruminal degradation of alfalfa hay and corn silages (Comlombatto et al., 2003b). Enzyme treatment alone or in combination with other treatments can increase the degradability of cereal straw by the rumen microorganisms (Liu and Orskov, 2000; Wang et al., 2004; Zhu et al., 2005; Eun et al., 2006; Fazaeli et al., 2006; Rodrigues et al., 2008) (Table 7). In addition, using fibrolytic enzymes in ruminant feed have shown improvements in the average daily gain of steers (Beauchemin et al., 1995), fleece weight and wool production of lambs (Jafari et al., 2005) and in milk yield of dairy cows (Yang et al., 2000).

Some studies, using fibrolytic enzymes alone could not significantly increase the degradability of rice straw because the ability of these enzymes to break down the esterified bonds within lignin-carbohydrate complexes may be limited. However, when using in combination with other pre-treatments they could increase degradability and in vitro fermentation characteristics, as shown by Eun et al. (2006) who treated with xylanase or cellulase in combination with ammonia, by Liu and Orskov (2000) who treated with cellulase from Penicillum funiculosum in combination with steam pre-treatment, and by Wang et al. (2004) who treated with multi-enzymes (xylanase, [beta]-glucanase, carboxymethylcellulase and amylase) in combination with NaOH. The use of combinations of fibrolytic enzyme with these pre-treatments is expected to have a synergistic effect on the nutritive improvement of rice straw.

Although, application of enzymes has proven to increase the feed value of poor quality feedstuffs, its use by smallholder farmers is, for the time being, economically unattractive. Especially, the use of lignin-degrading enzymes, originating from fungi, seems a promising development. However, using the fungi themselves, instead of their isolated enzymes, would be easier and cheaper to apply.

CONCLUSIONS

Although several treatments have been used to improve the degradability and voluntary intake of rice straw, such as physical or chemical treatments, the practical use of these treatments is still restricted in terms of safety concerns, costs and potentially negative environmental consequences.

Using ligninolytic fungi, including their enzymes, may be one potential alternative to provide a more practical and environmental-friendly approach for enhancing the nutritive value of rice straw. The cost of exogenous enzymes is at present too high to be applied by smallholder farms, but this may change in the future. Moreover, the application of ligninolytic fungi or their enzymes combined with chemical pre-treatments to rice straw may be an alternative way to shorten the period of the incubation times and (or) decrease the amount of chemicals, effecting some synergy. Certainly, since available data on treatments using fungi and their enzymes for improving the quality of rice straw are relatively scarce, these techniques should be developed further.

REFERENCES

Abou-EL-Enin, O. H., J. G. Fadel and D. J. Mackill. 1999. Differences in chemical composition and fibre digestion of rice straw with, without, anhydrous ammonia from 53 rice varieties. Anim. Feed Sci. Technol. 79:129-136.

Agbagla-Dohnani, A., P. Noziere, B. Gaillard-Martinie, M. Puard and M. Doreau. 2003. Effect of silica content on rice straw ruminal degradation. J. Anim. Sci. 140:183-192.

Akter, Y., M. A. Akbar, M. Shahjalal and T. U. Ahmed. 2004. Effect of urea molasses multi-nutrient blocks supplementation of dairy cows fed rice straw and green grasses on milk yield, composition, live weight gain of cows and calves and feed intake. Pak. J. Biol. Sci. 7(9):1523-1525.

Arieli, A. 1997. Whole cottonseed in dairy cattle feeding: a review. Anim. Feed Sci. Technol. 72:97-110.

Arora, D. S. and P. K. Gill. 2005. Production of ligninolytic enzymes by Phlebia floridensis. World J. Microbiol. Biotechnol. 21:1021-1028.

Arora, D. S., M. Chander and P. K. Gill. 2002. Involvement of lignin peroxidase, manganese peroxidase and laccase in degradation and selective ligninolysis of wheat straw. Int. Biodeterior. Biodegradation 50:115-120.

Bae, H. D., T. A. McAllister, E. G. Kokko, F. L. Leggett, L. J. Yanke, K. D. Jakober, J. K. Ha, H. T. Shin and K.-J. Cheng. 1997. Effect of silica on the colonization of rice straw by ruminal bacteria. Anim. Feed Sci. Technol. 65:165-181.

Baile, C. A. and J. M. Forbes. 1974. Control of feed intake and regulation of energy balance in ruminants. Physiol. Rew. 54:160-214.

Barrasa, J. M., S. Camarero, A. T. Martinez and K. Ruel. 1995. Ultrastructural aspects of wheat straw degradation by Phanerochaete chrysosporium and Trametes versicolor. Appl. Microbiol. Biotechnol. 43:766-770.

Beauchemin, K. A., D. Colombatto and D. P. Morgavi. 2004. A rationale for the development of feed enzyme products for ruminants. Can. J. Anim. Sci. 84:23-36.

Beauchemin, K. A., L. M. Rode and V. J. H. Sewalt. 1995. Fibrolytic enzymes increase fiber digestibility and growth rate of steers fed dry forages. Can. J. Anim. Sci. 75:641-644.

Berger, L. L., G. C. Fahey, L. D. Bourquin and E. C. Tilgeyer. 1994. Modification of forage quality after harvest. In: Forage Quality, Evaluation, and Utilisation (Ed. G. C. Fahey). American Society of Agronomy, Inc, Madison, USA. pp. 922-966.

Calzado, J. F. and C. Rolz. 1990. Estimation of the growth rate of Pleurotus on stocked straw. J. Ferment. Bioeng. 69:70-71.

Chaudhry, A. S. 1998. Nutrient composition, digestion and rumen fermentation in sheep of wheat straw treated with calcium oxide, sodium hydroxide and alkaline hydrogen peroxide. Anim. Feed Sci. Technol. 74:315-328.

Chaudhry, A. S. and E. L. Miller. 1996. The effect of sodium hydroxide and alkaline hydrogen peroxide on chemical composition of wheat straw and voluntary intake, growth and digesta kinetics in store lambs. Anim. Feed Sci. Technol. 60:69-86.

Chen, J., S. L. Fales, G. A. Varga and D. J. Royse. 1996. Biodegradability of free monomeric cell-wall-bound phenolic acids in maize stover by two strains of white-rot fungi. J. Sci. Food Agric. 71:145-150.

Chenost, M. and C. Kayouli. 1997. Roughage Utilisation in Warm Climates. FAO Animal Production and Health Paper 135, Rome.

Colombatto, D., G. Hervas, W. Z. Yang and K. A. Beauchemin. 2003a. Effects of enzymes supplementation of a total mixed ration on microbial fermentation in continous culture, maintained at high and low pH. J. Anim. Sci. 81:2617-2627.

Colombatto, D., D. P. Morgavi, A. F. Furtado and K. A. Beauchemin. 2003b. Screening of exogenous enzymes for ruminant diets: relationship between biochemical characteristics and in vitro ruminal degradation. J. Anim. Sci. 81:2628-2638.

Conrad, H. R. 1966. Symposium on factors influencing the voluntary intake of herbage by ruminants: Physiological and physical factors limiting feed intake. J. Anim. Sci. 25:227-235.

Devendra, C. and D. Thomas. 2002. Crop-animal interactions in mixed farming systems in Asia. Agric. Syst. 71:27-40.

Devendra, C. 1997. Crop residues for feeding animals in Asia: Technology development and adoption in crop/livestock systems. In: Crop Residuals in Sustainable Mixed Crop/livestock Farming System (Ed. C. Renard). CAB International; Wallingford, UK. pp. 241-267.

Dias da Silva, A. A. 1993. Upgrading of low quality feeds by means of urinary urea. In: Urine--a Wasted, Renewable Natural Resources (Ed. F. Sundstel and E. Owen). NORAGRIC Occasional Papers Series C, Development and Environment Vol. 12:42-49.

Doyle, P. T., C. Devendra and G. R. Pearce. 1986. Rice straw as a feed for ruminants. IDP, Canberra, Australia.

Eriksson, K-EL, R. A. Blanchette and P. Ander. 1990. Microbial and enzymatic degradation of wood and wood components. Springer, Berlin, Heidelberg, New York.

Eun, J.-S., K. A. Beauchemin, S.-H. Hong and M. W. Bauer. 2006. Exogenous enzymes added to untreated or ammoniated rice straw: Effects on in vitro fermentation characteristics and degradability. Anim. Feed Sci. Technol. 131:86-101.

Fadel Elseed, A. M. A. 2005. Effect of supplemental protein feeding frequency on ruminal chracteristics and microbial N production in sheep fed treated rice straw. Small Rumin. Res. 57:11-17.

Fadel Elseed, A. M. A., J. Sekine, M. Hishinuma and K. Hamana. 2003. Effects of ammonia, urea plus calcium hydroxide and animal urine treatments on chemical composition and in sacco degradability of rice straw. Asian-Aust. J. Anim. Sci. 16:368-373.

Fazaeli, H., A. Azizi and M. Amile. 2006. Nutritive value index of treated wheat straw with Pleurotus fungi fed to sheep. Pak. J. Biol. Sci. 9(13):2444-2449.

Giraldo, L. A., M. L. Tejido, M. J. Ranilla and M. D. Carro. 2007. Effects of exogenous cellulase supplementation on microbial growth and ruminal fermentation of a high-forage diet in Rusitec fermenters. J. Anim. Sci. 85:1962-1970.

Hadjipanayiotou, M. 1984. Effect of level and type of alkali on the digestibility in vitro of ensiled, chopped barley straw. Agric. Wastes 10:187-194.

Hoover, W. H. 1986. Chemical factors involved in ruminal fiber digestion. J. Dairy Sci. 69:2755-2766.

Howard, R. L., E. Abotsi, E. L. Jansen and S. Howard. 2003. Lignocellulose biotechnology: Issues of bioconversion and enzyme production. Afr. J. Biotechnol. 2:602-619.

Ibrahim, M. N. M. 1983. Physical, chemical, physico-chemical and biological treatments of crop residues. In: The Utilization of Fibrous Agricultural Residues (Ed. G. R. Pearce). Australian Development Assistance Bureau, Research for Development Seminar three, Los Banos, Philippines, 18-23 May 1981. Australian Government Publishing Service, Canberra, Australia. pp. 53-68.

Iiyama, K., T. B. T. Lam and B. A. Stone. 1990. Phenolic acid bridges between polysaccharides and lignin in wheat internodes. Phytochem. 29:733-737.

Jackson, M. G. 1977. Review article: The alkali treatment of straw. Anim. Feed Sci. Technol. 2:105-130.

Jafari, A., M. A. Edriss, M. Alikhani and G. Emtiazi. 2005. Effects of treated wheat straw with exogenous fibre-degrading enzymes on wool characteristics of ewe lambs. Pak. J. Nutr. 4:321-326.

Jalc, D. 2002. Straw enrichment for fodder production by fungi. In: The Mycota XI Agricultural Applications (Ed. F. Kempken). [C]Springer-Verlag, Berlin, Heidelberg. pp. 19-38.

Karunanadaa, K. and G. A. Varga. 1996a. Colinization of rice straw by white-rot fungi (Cyathus stercoreus): Effect on ruminal fermentation pattern, nitrogen metabolism, and fiber utilization during continuous culture. Anim. Feed Sci. Technol. 61:1-16.

Karunanadaa, K. and G. A. Varga. 1996b. Colinization of rice straw by white-rot fungi: cell eall monosaccharides, phenolic acids, ruminal fermentation characteristics and digestibility of cell wall fiber components in vitro. Anim. Feed Sci. Technol. 63:273-288.

Karunanandaa, K., G. A. Varga, D. E. Akin, L. L. Rigsby and D. J. Royse. 1995. Botanical fractions of rice straw colonized by white-rot fungi: Changes in chemical composition and structure. Anim. Feed Sci. Technol. 55:179-199.

Karunanandaa, K., S. L. Fales, G. A. Varga and D. J. Royse. 1992. Chemical composition and biodegradability of crop residues colonized by white rot fungi. J. Sci. Food Agric. 60:105-112.

Kirk, T. K. and R. L. Farrell. 1987. Enzymatic "combustion": The microbial degradation of lignin. Annu. Rev. Microbiol. 41:465-505.

Kluczek-Turpeinen, B., P. Maijala, M. Hofrichter and A. Hatakka. 2007. Degradation and enzymatic activties of three Paecilomyces inflatus strains grown on diverse lignocellulosic substrates. Int. Biodeterior. Biodegrad. 59:283-291.

Lam, T. B. T., K. Kadoya and K. Iiyama. 2001. Bonding of hydroxycinnamic acids to lignin: ferulic and p-coumaric acids are predominantly linked at the benzyl position of lignin, not the b-position, in grass cell walls. Phytochem. 57:987-992.

Lechner, B. E. and V. L. Papinutti. 2006. Production of lignocellulosic enzymes during growth and fruiting of the edible fungus Lentinus tigrinus on wheat straw. Process Biochem. 41:594-598.

Liu, J. X. and E. R. Orskov. 2000. Cellulase treatment of untreated and steam pre-treated rice straw-effect on in vitro fermentation characteristics. Anim. Feed Sci. Technol. 88:189-200.

Liu, J. X., E. R. Orskov and X. B. Chen. 1999. Optimization of steam treatment as a method for upgrading rice straw as feeds. Anim. Feed Sci. Technol. 76:345-357.

Liu, J. X., A. Susenbeth and K. H. Sudekum. 2002. In vitro gas production measurements to evaluate interactions between untreated and chemically treated rice straws, grass hay, and mulberry leaves. J. Anim. Sci. 80:517-524.

NARC newsletter (rice special issue), May 2004.

Novotny, C., K. Svobodova, P. Erbanova, T. Cajthaml, A. Kasinath, E. Lang and V. Sasek. 2004. Ligninolytic fungi in bioremediation: Extracellular enzyme production and degradation rate. Soil Biol. Biochem. 36:1545-1551.

Phang, O. C. and J. Vadiveloo. 1992. Effects of varieties, botanical fractions and supplements of palm oil by-products on the feeding value of rice straw in goats. Small Rumin. Res. 6:295-301.

Pradhan, R., H. Tobioka and I. Tasaki. 1997. Effect of moisture content and different levels of additives on chemical composition and in vitro dry matter digestibility of rice straw. Anim. Feed Sci. Technol. 68:273-284.

Prasad, R. D. D., M. R. Reddy and G. V. N. Reddy. 1998. Effect of feeding baled and stacked urea treated rice straw on the performance of crossbred cows. Anim. Feed Sci. Technol. 73:347-352.

Rai, S. N. and V. D. Mudgal. 1996. Effect of alkali and (or) steam treatment of wheat straw or cellulase augmentd concentrate mixture on rumen fermentation on goats. Small Rumin. Res. 19:219-225.

Reddy, D. V. 1996. Evaluation of rice straw-poultry droppings based rations supplemented with graded levels of rice bran in fistulated buffaloes. Anim. Feed Sci. Technol. 58:227-237.

Rezaeian, M., G. W. Beakes and A. S. Chaudhry. 2005. Relative fibrolytic activities of anaerobic rumen fungi on untreated and sodium hydroxide treated barley straw in in vitro culture. Anaerobe 11:163-175.

Rodrigues, M. A. M., P. Pinto, R. M. F. Bezerra, A. A. Dias, C. V. M. Guedes, V. M. G. Gardoso, J. W. Cone, L. M. M. Ferreira, J.

Colaco and C. A. Sequeira. 2008. Effect of enzyme extracts isolated from white-rot fungi on chemical composition and in vitro digestibility of wheat straw. Anim. Feed Sci. Technol. 141:326-338.

Saadulah, M., M. Haque and F. Dolberg. 1981. Treatment of rice straw with lime. Trop. Anim. Prod. 6:116-120.

Schiere, J. B. and M. N. M. Ibrahim. 1989. Feeding of urea-ammonia treated rice straw: A compilation of miscellaneous reports produced by the Straw Utilization Project (Sri Lanka). Pudoc, Wageningen.

Schubert, R. and G. Flachowsky. 1994. Investigations on nitrogen flow into and out of nylon bag after ruminal incubation and feeding of untreated or 15N horse urine treated wheat straw in small ruminants. Anim. Feed Sci. Technol. 48:199-209.

Selim, A. S. M., J. Pan, T. Suzuki, K. Ueda, Y. Kobayashi and K. Tanaka. 2002. Postprandial changes in particle associated ruminal bacteria in sheep fed ammoniated rice straw. Anim. Feed Sci. Technol. 19:227-287.

Selim, A. S. M., J. Pan, T. Takano, T. Suzuki, S. Koike, Y. Kobayashi and K. Tanaka. 2004. Effect of ammonia treatment on physical strength of rice straw, distribution of straw particles and particle-associated bacteria in sheep rumen. Anim. Feed Sci. Technol. 115:117-128.

Shen, H. S., D. B. Ni and F. Sundstel. 1998. Studies on untreated and urea-treated rice straw from three cultivation seasons: 1. Physical and chemical measurements in straw and straw fractions. Anim. Feed Sci. Technol. 73:243-261.

Shen, H. S., F. Sundstel, E. R. Eng and L. O. Eik. 1999. Studies on untreated and urea-treated rice straw from three cultivation seasons: 3. Histological investigations by light and scanning electron microscopy. Anim. Feed Sci. Technol. 80:151-159.

Sirohi, S. K. and S. N. Rai. 1995. Associative effect of lime plus urea treatment of paddy straw on chemical composition and in vitro digestibility. Indian J. Anim. Sci. 65:1346-1351.

Stensig, T., M. R. Weisbjerg, J. Madsen and T. Hvelplund. 1994. Estimation of voluntary feed intake from in sacco degradation and rate of passage of DM or NDF. Livest. Prod. Sci. 39:49-52.

Sundstol, F. and E. M. Coxworth. 1984. Ammonia treatment. In: Straw and Other Fibrous By-products as Feed (Ed. F. Sundstol and E. Owen). Developments in Animal Veterinary Sciences, 14. Elsevier, Amsterdam, pp. 196-247.

Theander, O. and P. Aman. 1984. Anatomical and chemical characteristics. In: Straw and Other Fibrous By-products as Feed. (Ed. F. Sundstel and E. Own). Developments in Animal Veterinary Sciences, 14. Elsevier, Amsterdam, pp. 45-78.

Uden, P. 1988. The effect of grinding and pelleting hay on digestibility, fermentation rate, digesta passage and rumen and faecal particle size in cows. Anim. Feed Sci. Technol. 19:145-157.

Vadiveloo, J. 1992. Varietal differences in the chemical composition and in vitro digestibility of rice straw. J. Agric. Sci. 119:27-33.

Vadiveloo, J. 1995. Factors contributing to varietal differences in the nutritive value of rice straw. Anim. Feed Sci. Technol. 54:45-53.

Vadiveloo, J. 2000. Nutritional properties of the leaf and stem of rice straw. Anim. Feed Sci. Technol. 83:57-65.

Vadiveloo, J. 2003. The effect of agronomic improvement and urea treatment on the nutritional value of Malaysian rice straw varieties. Anim. Feed Sci. Technol. 108:33-146.

Van Soest, P. J. 2006. Review: rice straw, the role of silica and treatments to improve quality. Anim. Feed Sci. Technol. 130:137-171.

Vu, D. D., L. X. Cuong, C. A. Dung and P. H. Hai. 1999. Use of urea-molasses-multinutrient block and urea-treated rice straw for improving dairy cattle productivity in Vietnam. Prev. Vet. Med. 38:187-193.

Wanapat, M., A. Petlum and O. Pimpa. 1999. Strategic supplementation with a high quality feed block on roughage intake, milk yield and composition and economic return in lactating dairy cows. Asian-Aust. J. Anim. Sci. 12:901-903.

Wanapat, M., S. Polyrach, K. Boonnop, C. Mapato and A. Cherdthong. 2009. Effect of treating rice straw with urea and calcium hydroxide upon intake, digestibility, rumen fermentation and milk yield of dairy cows. Livest. Sci. 125:238-243.

Wanapat, M., K. Sommart and K. Saardrak. 1996. Cottonseed meal supplementation of dairy cattle fed rice straw. Livestock Research for Rural Development. 8(3) (http://www.fao.org/ ag/AGa/ AGAP- /FRG/FEEDback/lrrd/lrrd8/3/metha83.htm)

Wang, Y., B. M. Spratling, D. R. ZoBell, R. D. Wiedmeier and T. A. McAllister. 2004. Effect of alkali pretreatment of wheat straw on the efficacy of exogenous fibrolytic enzymes. J. Anim. Sci. 82:198-208.

Warly, L., T. Matsui, T. Harumoto and T. Fujihara. 1992. Study on the utilization of rice straw by sheep: Part I. The effect of soybean meal supplementation on the eating and rumination behavior. Asian-Aust. J. Anim. Sci. 5:695-698.

Yamakava, M. and H. A. Okamnto. 1992. Effect of incubation with edible mushroom, Pleurotus ostreatus, on voluntary intake and digestibility or rice bran by sheep. Anim. Feed Sci. Technol. 63:133-138.

Yang, W. Z., K. A. Beauchemin and L. M. Rode. 2000. A comparison of methods of adding fibrolytic enzymes to lactating cow diets. J. Dairy Sci. 83:2512-2520.

Zhu, S., Y. Wu, Z. Yu, J. Liao and Y. Zhang. 2005. Pretreatment by microwave/alkali of rice straw and its enzymic hydrolysis. Process Biochem. 40:3082-3086.

C. Sarnklong, J. W. Cone *, W. Pellikaan and W. H. Hendriks

Animal Nutrition Group, Department of Animal Sciences, Wageningen University, P.O. Box 338, NL-6700 AH Wageningen, The Netherlands

* Corresponding Author: J. W. Cone. Tel: +31-317-483-542, Fax: +31-317-484-260, E-mail: john.cone@wur.nl

Received November 7, 2008; Accepted December 24, 2009
Table 1. Rice production and obtained residues of the
top 10 rice-producing countries in the world in 2003

                     Rice production     Rice husk       Rice straw
Number  Country        (million t)    (million t) (a)  (million t) (a)

1       China            166.00            38.18            74.70
2       India            133.51            30.71            60.08
3       Indonesia         51.85            11.93            23.33
4       Bangladesh        38.06             8.75            17.13
5       Vietnam           34.61             7.96            15.57
6       Thailand          27.00             6.21            12.15
7       Myanmar           21.90             5.04             9.86
8       Philippines       13.17             3.03             5.93
9       Brazil            10.22             2.35             4.60
10      Japan              9.86             2.27             4.44
Total                    506.18           116.42           227.78

(a) Calculated data (Adapted from NARC newsletter, 2004).

Table 2. Chemical composition of rice straw
in early, middle and late growing season

Cultivation    DM       N      NDF     ADF    Hemicellulose
season (a)
                %            % of DM

Early         96.40   1.04    72.53   43.52       29.01
Middle        96.20   0.99    70.03   41.09       29.08
Late          96.87   0.88    71.97   39.83       32.24
Mean          96.30   0.96    73.01   41.59       31.42
SD (d)         0.86   0.27     3.16    2.17        3.21

Cultivation   Cellulose   ADL (b)   EBSi (c)
season (a)
                          % of DM

Early           35.81      4.90       4.82
Middle          32.80      4.66       4.57
Late            31.96      4.63       4.46
Mean            33.35      4.84       4.25
SD (d)           1.97      0.44       0.95

(a) Season: Early-Transplanting in April, Middle-Transplanting in
June, Late-Transplanting in July.

(b) ADL: Acid detergent lignin. (c) EBSi: Extractable biogenic
silica. (d) SD: Standard deviation. (Adapted from Shen et al.,
1998)

Table 3. Content of total ash, acid insoluble
ash and other minerals in rice straw

Cultivation    Ash    AIA (b)     K      Ca      Mg
season (a)
                               % of DM

Early         13.00    2.83     2.21    0.52    0.91
Middle        13.00    3.85     1.88    0.49    0.30
Late          10.80    3.49     0.97    0.56    0.23
Mean          12.10    3.40     1.58    0.53    0.24
SD (c)         1.46    0.66     0.62    0.11    0.06

Cultivation     P       Na       Fe      Mn
season (a)
                          % of DM

Early         0.13     0.09     0.04    0.07
Middle        0.11     0.04     0.04    0.06
Late          0.13     0.21     0.11    0.07
Mean          0.12     0.13     0.07    0.07
SD (c)        0.03     0.10     0.05    0.01

(a) Season: Early--Transplanting in April, Middle--Transplanting
in June, Late--Transplanting in July. (b) AIA: Acid detergent
insoluble ash. (c) SD: Standard deviation. (Adapted from Shen et
al., 1998)

Table 4. Fungi and their enzyme production studied to
improve the nutritive value of fodder for ruminant feed

References                 Species of fungi    Determined enzymes

Jalc (2002)                Phlebia radiate,    The LiP (b)-MnP (c)
                             Coriolus
                             versicolor and
                             Trametes gibbosa
                           Panus tigrinus and  The MnP-laccase
                             Dichomitus
                             squalens
                           Junghuhnia          The LiP-laccase
                             separabilima
                             and Phlebia
                             ochraceofulva
                           Pleurotus           The laccase-AAO (d)
                             ostreatus and
                             Pleurotus
                             sajor-caju
Barrasa et al. (1995)      Phanerochaete       Ligninolytic enzyme
                             chrysosporium
Fazaeli et al. (2006)      Pleurotus fungi     Ligninolytic enzyme
Barrasa et al. (1995)      T. versicolor       Ligninolytic enzyme
Rodrigues et al. (2008)    T. versicolor       MnP, Laccase, CMCasee
                                                 and Avicelasef
                           Bjerkandara adusta  MnP, CMCase and
                                                 Avicelase
                           Fomes fomentarius   Laccase, CMCase and
                                                 Avicelase
Kluczek-Turpeinen et al.   Paecilomyces        Endoglucanase, xylanase
  (2007)                     inflatus            and Laccase
Eun et al. (2006)          Trichoderma reesei  Cellulase and Xylanase
Zhu et al. (2005)          T. reesei           Cellulase
Giraldo et al. (2007)      T. longibrachiatum  Cellulase
Wang et al. (2004)         T. longibrachiatum  Xylanase and P-glucanase
Rai and Mudgal (1996)      T. viride           Cellulase
Giraldo et al. (2007)      Aspergillus niger   Cellulase
Rezaeian et al. (2005)     Fungi from sheep    Cellulase and Xylanase
                             rumen residue
Liu and Orskov (2000)      Penicillium         Cellulase
                             fumicalosum
Eun et al. (2006)          Bacillus            Protease
                             licheniformis
                             (a)
Colombatto et al. (2003a)  B. licheniformis    Protease
                             (a)
Karunanandaa and Varga     Cyathus stercoreus  NR
  (1996b)

References                 Species of fungi    Main diet ingredients

Jalc (2002)                Phlebia radiate,    General plant cell wall
                             Coriolus
                             versicolor and
                             Trametes gibbosa
                           Panus tigrinus and
                             Dichomitus
                             squalens
                           Junghuhnia
                             separabilima
                             and Phlebia
                             ochraceofulva
                           Pleurotus
                             ostreatus and
                             Pleurotus
                             sajor-caju
Barrasa et al. (1995)      Phanerochaete       Wheat straw
                             chrysosporium
Fazaeli et al. (2006)      Pleurotus fungi     Wheat straw
Barrasa et al. (1995)      T. versicolor       Wheat straw
Rodrigues et al. (2008)    T. versicolor       Wheat straw
                           Bjerkandara adusta  Wheat straw
                           Fomes fomentarius   Wheat straw
Kluczek-Turpeinen et al.   Paecilomyces        Wheat straw
  (2007)                     inflatus
Eun et al. (2006)          Trichoderma reesei  Rice straw
Zhu et al. (2005)          T. reesei           Rice straw
Giraldo et al. (2007)      T. longibrachiatum  Grass hay
Wang et al. (2004)         T. longibrachiatum  Wheat straw
Rai and Mudgal (1996)      T. viride           Wheat straw
Giraldo et al. (2007)      Aspergillus niger   Grass hay
Rezaeian et al. (2005)     Fungi from sheep    Barley straw
                             rumen residue
Liu and Orskov (2000)      Penicillium         Rice straw
                             fumicalosum
Eun et al. (2006)          Bacillus            Rice straw
                             licheniformis
                             (a)
Colombatto et al. (2003a)  B. licheniformis    Alfalfa hay
                             (a)
Karunanandaa and Varga     Cyathus stercoreus  Rice straw
  (1996b)

(a) One species of bacteria. (b) LiP: Lignin-peroxidase. (c) MnP:
Manganese-dependent peroxidase. (d) AAO: Aryl-alcohol oxidase.
(e) CMCase: Endogluconase. (f) Avicelase: Exoglucanase. NR = Not
reported.

Table 5. Degradation of straw by fungi and enzyme extracts

                                                           Mean
                                                       degradability
                                                         untreated
                                              Kind of     control
References         Kind of fungi/enzymes       straw      (g/kg)

Kluczek-Turpeinen  Paecilomyces inflatus      Wheat         357
  et al. (2007)    Paecilomyces inflatus      Wheat         239
                   Paecilomyces inflatus      Wheat         262
Fazaeli et al.     Pleurotus fungi 1          Wheat         207
  (2006)           Pleurotus fungi 2          Wheat         548
                   Pleurotus fungi 1          Wheat         207
                   Pleurotus fungi 2          Wheat         548
Rodrigues et       EE (c) of Trametes         Wheat         349
  al. (2008)         versicolor (TV1)
                   EE of T. versicolor (TV2)  Wheat         349
                   EE of Bjerkandera adusta   Wheat         349
                   EE of Fomes fomentarius    Wheat         349
                   EE of T. versicolor (TV1)  Wheat         511
                   EE of T. versicolor (TV2)  Wheat         511
                   EE of B. adusta            Wheat         511
                   EE of F. fomentarius       Wheat         511
                   EE of T. versicolor (TV1)  Wheat         104
                   EE of T. versicolor (TV2)  Wheat         104
                   EE of B. adusta            Wheat         104
                   EE of F. fomentarius       Wheat         104
Liu and Orskov     Cellulase                  Rice          446
  (2000)

                                                Change by     Relative
References         Kind of fungi/enzymes      treatment (g)  change (%)

Kluczek-Turpeinen  Paecilomyces inflatus      +14              4
  et al. (2007)    Paecilomyces inflatus      +49             21
                   Paecilomyces inflatus      +13              5
Fazaeli et al.     Pleurotus fungi 1           -2            -10
  (2006)           Pleurotus fungi 2           -7            -12
                   Pleurotus fungi 1           -2            -11
                   Pleurotus fungi 2           -5             -8
Rodrigues et       EE (c) of Trametes         +10              3
  al. (2008)         versicolor (TV1)
                   EE of T. versicolor (TV2)  -13             -4
                   EE of Bjerkandera adusta   -16             -5
                   EE of Fomes fomentarius    -15              4
                   EE of T. versicolor (TV1)   -7             -1
                   EE of T. versicolor (TV2)   +6              1
                   EE of B. adusta             +7              1
                   EE of F. fomentarius        +9              2
                   EE of T. versicolor (TV1)   -3             -3
                   EE of T. versicolor (TV2)   +1              1
                   EE of B. adusta             +1              1
                   EE of F. fomentarius        +7              7
Liu and Orskov     Cellulase                  +66             15
  (2000)

                                                 Degradability
References         Kind of fungi/enzymes            method

Kluczek-Turpeinen  Paecilomyces inflatus      SSF (a) 12 wk,
  et al. (2007)                                 Hemicellulose
                   Paecilomyces inflatus      SSF 12 wk,
                                                Cellulose
                   Paecilomyces inflatus      SSF 12 wk, Lignin
Fazaeli et al.     Pleurotus fungi 1          SSF 17 d,
  (2006)                                        Hemicellulose
                   Pleurotus fungi 2          SSF 17 d, Cellulose
                   Pleurotus fungi 1          SSF 17 d,
                                                Hemicellulose
                   Pleurotus fungi 2          SSF 17 d, Cellulose
Rodrigues et       EE (c) of Trametes         IC (b) 6 d,
  al. (2008)         versicolor (TV1)           Hemicellulose
                   EE of T. versicolor (TV2)  IC 6 d,
                                                Hemicellulose
                   EE of Bjerkandera adusta   IC 6 d,
                                                Hemicellulose
                   EE of Fomes fomentarius    IC 6 d,
                                                Hemicellulose
                   EE of T. versicolor (TV1)  IC 6 d, Cellulose
                   EE of T. versicolor (TV2)  IC 6 d, Cellulose
                   EE of B. adusta            IC 6 d, Cellulose
                   EE of F. fomentarius       IC 6 d, Cellulose
                   EE of T. versicolor (TV1)  IC 6 d, Lignin
                   EE of T. versicolor (TV2)  IC 6 d, Lignin
                   EE of B. adusta            IC 6 d, Lignin
                   EE of F. fomentarius       IC 6 d, Lignin
Liu and Orskov     Cellulase                  IC 3 wk, DM (d)
  (2000)

(a) SSF: Solid-state fermentation. (b) IC: Incubation.
(c) EE: Enzyme extract. (d) DM: Dry matter.

Table 6. Degradability of straw by fungi and enzyme extract treatments

                                                       Mean
                                                   degradability
                         Kind of        Kind of  untreated control
References            fungi/enzymes      straw    (g/kg or ml/g)

Fazaeli et al.     Pleurotus fungi 1    Wheat    281
  (2006)           Pleurotus fungi 2    Wheat    281
Karunanandaa and   Cyathus stercoreus   Rice     371
  Varga (1996b)
Eun et al. (2006)  END1 (cellulose)     Rice     153
                   END1 (cellulose)     Rice     153
                   XY1 (xylanase)       Rice     153
                   XY2 (xylanase)       Rice     153
                   EX                   Rice     153
                     (cellulase +
                     hemicellulose)
                   PROT (protease)      Rice     153
                   END1 (cellulose)     Rice     170

                   END1 (cellulose)     Rice     170

                   XY1 (xylanase)       Rice     170

                   XY2 (xylanase)       Rice     170

                   EX                   Rice     170
                     (cellulase +
                     hemicellulose)
                   PROT (protease)      Rice     170

                   END1 (cellulose)     Rice      54 ml [g.sup.-1]
                   END1 (cellulose)     Rice      54 ml [g.sup.-1]
                   XY1 (xylanase)       Rice      54 ml [g.sup.-1]
                   XY2 (xylanase)       Rice      54 ml [g.sup.-1]
                   EX                   Rice      54 ml [g.sup.-1]
                     (cellulase +
                     hemicellulose)
                   PROT (protease)      Rice      54 ml [g.sup.-1]
Rodrigues et       EE (c) of            Wheat    242 ml [g.sup.-1]
  al. (2008)         Tramete
                     versicolor (TV1)
                   EE of T. versicolor  Wheat    242 ml [g.sup.-1]
                     (TV2)
                   EE of Bjerkandera    Wheat    242 ml [g.sup.-1]
                     adusta
                   EE of Fomes          Wheat    242 ml [g.sup.-1]
                     fomentarius
Liu and Orskov     Cellulase            Rice      20 ml
  (2000)
Wang et al.        Xylanase+[beta]-     Wheat    134 ml
  (2004)             glucanase

                                        Change by
                         Kind of        treatment   Relative
References            fungi/enzymes     (g or ml)  change (%)

Fazaeli et al.     Pleurotus fungi 1    +12          9
  (2006)           Pleurotus fungi 2    +43         32
Karunanandaa and   Cyathus stercoreus   +16         42
  Varga (1996b)
Eun et al. (2006)  END1 (cellulose)     +14          9
                   END1 (cellulose)     +26         17
                   XY1 (xylanase)       +21         14
                   XY2 (xylanase)       +23         15
                   EX                   +42         27
                     (cellulase +
                     hemicellulose)
                   PROT (protease)      +27         18
                   END1 (cellulose)     -18        -11

                   END1 (cellulose)      -9         -5

                   XY1 (xylanase)       -10         -6

                   XY2 (xylanase)       -24        -14

                   EX                   -16         -9
                     (cellulase +
                     hemicellulose)
                   PROT (protease)      +50         29

                   END1 (cellulose)       1 ml       2
                   END1 (cellulose)       1 ml       1
                   XY1 (xylanase)         6 ml      10
                   XY2 (xylanase)         7 ml      13
                   EX                    19 ml      35
                     (cellulase +
                     hemicellulose)
                   PROT (protease)       11 ml      21
Rodrigues et       EE (c) of             27 ml      11
  al. (2008)         Tramete
                     versicolor (TV1)
                   EE of T. versicolor   65 ml      27
                     (TV2)
                   EE of Bjerkandera     15 ml       6
                     adusta
                   EE of Fomes           -4 ml      -2
                     fomentarius
Liu and Orskov     Cellulase              0.3 ml     1
  (2000)
Wang et al.        Xylanase+[beta]-       1.8 ml     1
  (2004)             glucanase

                         Kind of         Degradability
References            fungi/enzymes         method

Fazaeli et al.     Pleurotus fungi 1    In vitro DM (a)
  (2006)           Pleurotus fungi 2    In vitro DM
Karunanandaa and   Cyathus stercoreus   In vitro DM
  Varga (1996b)
Eun et al. (2006)  END1 (cellulose)     In vitro DM
                   END1 (cellulose)     In vitro DM
                   XY1 (xylanase)       In vitro DM
                   XY2 (xylanase)       In vitro DM
                   EX                   In vitro DM
                     (cellulase +
                     hemicellulose)
                   PROT (protease)      In vitro DM
                   END1 (cellulose)     In vitro,
                                          Hemicellulose
                   END1 (cellulose)     In vitro,
                                          Hemicellulose
                   XY1 (xylanase)       In vitro,
                                          Hemicellulose
                   XY2 (xylanase)       In vitro,
                                          Hemicellulose
                   EX                   In vitro,
                     (cellulase +         Hemicellulose
                     hemicellulose)
                   PROT (protease)      In vitro,
                                          Hemicellulose
                   END1 (cellulose)     GP (b)
                   END1 (cellulose)     GP
                   XY1 (xylanase)       GP
                   XY2 (xylanase)       GP
                   EX                   GP
                     (cellulase +
                     hemicellulose)
                   PROT (protease)      GP
Rodrigues et       EE (c) of            GP
  al. (2008)         Tramete
                     versicolor (TV1)
                   EE of T. versicolor  GP
                     (TV2)
                   EE of Bjerkandera    GP
                     adusta
                   EE of Fomes          GP
                     fomentarius
Liu and Orskov     Cellulase            GP
  (2000)
Wang et al.        Xylanase+[beta]-     GP
  (2004)             glucanase

(a) DM: Dry matter. (b) GP: Gas production. EE: Enzymatic extract.

Table 7. Summary of enzyme treatments combined
with other treatments of rice straw

                                                     Mean degradability
                                                     untreated control
References  Kind of fungi/enzymes      Applied with    (g/kg or ml/g)

Eun et al.  END1 (cellulase)           3% Ammonia    264
  (2006)    END1 (cellulase)           3% Ammonia    264
            XY1 (xylanase)             3% Ammonia    264
            XY2 (xylanase)             3% Ammonia    264
            EX                         3% Ammonia    264
            (cellulose+hemicellulose)
            PROT (protease)            3% Ammonia    264
            END1 (cellulase)           3% Ammonia    267
            END1 (cellulase)           3% Ammonia    267
            XY1 (xylanase)             3% Ammonia    267
            XY2 (xylanase)             3% Ammonia    267
            EX                         3% Ammonia    267
            (cellulose+hemicellulose)
            PROT (protease)            3% Ammonia    267
            END1 (cellulase)           3% Ammonia    115 ml[g.sup.-1]
            END1 (cellulase)           3% Ammonia    115 ml[g.sup.-1]
            XY1 (xylanase)             3% Ammonia    115 ml[g.sup.-1]
            XY2 (xylanase)             3% Ammonia    115 ml[g.sup.-1]
            EX                         3% Ammonia    115 ml[g.sup.-1]
            (cellulose+hemicellulose)
            PROT (protease)            3% Ammonia    115 ml[g.sup.-1]
Zhu et al.  Cellulase                  1% NaOH       143
  (2005)    Cellulase                  1% NaOH       654
            Cellulase                  1% NaOH       060
            Cellulase                  Microwave     102
                                       +1% NaOH
            Cellulase                  Microwave     692
                                       +1% NaOH
            Cellulase                  Microwave     049
                                       +1% NaOH
Liu and     Cellulase                  Steam,        532
  Orskov                                 15 bar
  (2000)                                 5 min
            Cellulase                  Steam,         25 ml[g.sup.-1]
                                         15 bar
                                         5 min

                                       Change by
                                       treatment   Relative
References  Kind of fungi/enzymes      (g or ml)  change (%)

Eun et al.  END1 (cellulase)             +7          3
  (2006)    END1 (cellulase)            +11          4
            XY1 (xylanase)              +36         14
            XY2 (xylanase)              +77         29
            EX                          +25          9
            (cellulose+hemicellulose)
            PROT (protease)             +42         16
            END1 (cellulase)            +35         13
            END1 (cellulase)            +17          6
            XY1 (xylanase)              +52         19
            XY2 (xylanase)              +74         28
            EX                          +44         16
            (cellulose+hemicellulose)
            PROT (protease)            +181         68
            END1 (cellulase)             2 ml        2
            END1 (cellulase)             4 ml        4
            XY1 (xylanase)              22 ml       19
            XY2 (xylanase)              17 ml       15
            EX                          13 ml       11
            (cellulose+hemicellulose)
            PROT (protease)             48 ml       41
Zhu et al.  Cellulase                   +3          22
  (2005)    Cellulase                  +30          46
            Cellulase                  -10        -162
            Cellulase                   -1         -10
            Cellulase                  +34          49
            Cellulase                   -9        -180
Liu and     Cellulase                  +26           5
  Orskov
  (2000)
            Cellulase                    1.5 ml      6

References  Kind of fungi/enzymes      Degradability method

Eun et al.  END1 (cellulase)           In vitro DM (a)
  (2006)    END1 (cellulase)           In vitro DM
            XY1 (xylanase)             In vitro DM
            XY2 (xylanase)             In vitro DM
            EX                         In vitro DM
            (cellulose+hemicellulose)
            PROT (protease)            In vitro DM
            END1 (cellulase)           In vitro,
                                         Hemicellulose
            END1 (cellulase)           In vitro,
                                         Hemicellulose
            XY1 (xylanase)             In vitro,
                                         Hemicellulose
            XY2 (xylanase)             In vitro,
                                         Hemicellulose
            EX                         In vitro,
            (cellulose+hemicellulose)    Hemicellulose
            PROT (protease)            In vitro,
                                         Hemicellulose
            END1 (cellulase)           GP (b)
            END1 (cellulase)           GP
            XY1 (xylanase)             GP
            XY2 (xylanase)             GP
            EX                         GP
            (cellulose+hemicellulose)
            PROT (protease)            GP
Zhu et al.  Cellulase                  IC (c) 70 min,
  (2005)                                 Hemicellulose
            Cellulase                  IC 70 min,
                                         Cellulose
            Cellulase                  IC 70 min,
                                         Lignin
            Cellulase                  IC 30 min,
                                         Hemicellulose
            Cellulase                  IC 30 min,
                                         Cellulose
            Cellulase                  IC 30 min,
                                         Lignin
Liu and     Cellulase                  IC 3 wk, DM
  Orskov
  (2000)
            Cellulase                  GP

(a) DM: Dry matter. (b) GP: Gas production. (c) IC: Incubation.
COPYRIGHT 2010 Asian - Australasian Association of Animal Production Societies
No portion of this article can be reproduced without the express written permission from the copyright holder.
Copyright 2010 Gale, Cengage Learning. All rights reserved.

 Reader Opinion

Title:

Comment:



 

Article Details
Printer friendly Cite/link Email Feedback
Author:Sarnklong, C.; Cone, J.W.; Pellikaan, W.; Hendriks, W.H.
Publication:Asian - Australasian Journal of Animal Sciences
Article Type:Report
Geographic Code:4EUNE
Date:May 1, 2010
Words:9718
Previous Article:Therapy of diabetes mellitus using experimental animal models.
Next Article:Development and application of high-density SNP arrays in genomic studies of domestic animals.
Topics:

Terms of use | Copyright © 2014 Farlex, Inc. | Feedback | For webmasters