By Drs. Jerry Shurson and Pedro Urriola, University of Minnesota Department of Animal Science
© 2018 Feedstuffs. Reprinted with permission from Vol. 90, No. 10, October 1, 2018
The gross energy (GE) content of corn dried distillers grains plus solubles (DDGS) – at 5,429 kcal/kg of dry matter – is much greater than the 4.45 kcal/kg of dry matter for corn, 4,730 kcal/kg for soybean meal and the GE of other common ingredients used in swine diets.
However, the efficiency with which pigs utilize GE in DDGS, as measured by the net energy (NE)-to-GE ratio, is low (0.49) compared with corn – at 0.49 versus 0.68, respectively (National Research Council, 2012) – but NE:GE can vary from 0.42 to 0.46 among DDGS sources with variable crude fat content (Kerr et al., 2015).
Of all of the chemical components in feed ingredients, lipids provide the greatest amount of NE (kcal/kg), followed by crude protein and starch, with a limited amount provided from dietary fiber (Noblet and van Milgen, 2004). Therefore, the low starch content (0.84-3.89%, Kerr et al., 2013; 5.2-11.4%, Urriola et al., 2010) and relatively high neutral detergent fiber (NDF) content (30.2-39.7%; Pedersen et al., 2014) in corn DDGS are the primary reasons for the relatively low GE:NE.
To improve caloric and nutritional efficiency when feeding DDGS diets to pigs, we need to gain a better understanding of the chemical and physical characteristics that affect the digestibility and fermentability of fiber and explore approaches to convert a greater proportion of these calories to NE in pigs.
Fiber content in feed ingredients is measured using several different chemical methods, including crude fiber, NDF, acid detergent fiber (ADF) and total dietary fiber (TDF), which is comprised of insoluble dietary fiber (IDF) and soluble dietary fiber (SDF), as well as non-starch polysaccharides (NSPs), that quantify different types and proportions of in- digestible carbohydrates but may not adequately relate directly to nutritional and physiological effects in the animal (Infographic).
The most common measures that have been used to characterize the fiber content in DDGS are NDF, ADF and TDF (which can be separated into IDF and SDF fiber fractions). The NDF, ADF and hemicellulose (determined by the difference between NDF and ADF) contents, on a dry matter basis, of corn DDGS sources are in a range of 28.8-44.0%, 9.0-14.0% and 18.5-30.0%, respectively (Kerr et al., 2013).
The apparent total tract digestibility (ATTD) of NDF has been reported to range from 44.5% to 61.5% among 15 corn DDGS sources (Kerr et al., 2013). The range in TDF content (dry matter basis) of DDGS sources appears to be similar among studies, with Kerr et al. (2013) reporting a range of 30.8-37.8% TDF and Urriola et al. (2010) reporting a range of 32.9-38.6%. However, most of the TDF in DDGS is insoluble (31.8- 37.3%), with minimal amounts of soluble fiber (0.0-1.8%; Urriola et al., 2010).
The most comprehensive study to help us achieve a better understanding of the digestibility and fermentability of fiber in DDGS was conducted by Urriola et al. (2010). Results from three experiments showed that the ATTD of TDF ranged from 29.3% to 57.0% among corn DDGS sources. They also determined the apparent ileal digestibility (AID), ATTD and hindgut fermentability of ADF, NDF, IDF, SDF and TDF by feeding eight corn DDGS sources to growing pigs (Table 1).
The fiber digestibility and ferment- ability varied substantially among DDGS sources, regardless of the fiber measure used. This variability may be partially attributed to the amounts and types of enzymes as well as the different production processes used in ethanol and co- product production facilities to produce DDGS. Furthermore, about 46% of NDF and 29% of TDF is digestible in the small intestine, while about 13% of NDF and 21% of TDF is fermented in the hindgut.
These proportions of digestible and fermentable carbohydrates are assumed to contribute to the total metabolizable energy (ME) and NE content of DDGS, and the wide range in digestibility and fermentability among sources may partially contribute to the range in ME and NE content of DDGS reported in many studies. However, the AID of IDF is much less than SDF, and IDF is less ferment- able than SDF in the hindgut.
Although the average ATTD of NDF (59%) and TDF (50%) is moderately high in corn DDGS compared with some other types of high-fiber ingredients, there is a tremendous need to explore ways of improving AID and hindgut fermentation of the high proportion of IDF in DDGS to enhance the NE content.
Furthermore, the physical structure of fiber in corn DDGS may trap some of the oil in DDGS, prevent it from being accessible to digestive enzymes and contribute to the relatively low GE:ME and GE:NE contents. In fact, Kerr et al. (2013) reported that the ATTD of ether extract (oil) ranged from 53% to 81% among corn DDGS sources. Therefore, it is likely that the combined differences in fiber digestibility and fermentability, along with the variability of lipid digestibility involving fiber structure, are the primary reasons for the variable ME and NE content among DDGS sources fed to swine.
Although it is useful to understand the differences in the digestibility and fermentability of ADF, NDF, IDF and SDF in DDGS, knowledge of the NSP composition of DDGS is extremely important when selecting commercially available feed enzymes to improve the energy and nutrient digestibility of DDGS in pigs.
Pedersen et al. (2014) determined the NSP composition (dry matter basis) of 47 corn DDGS samples and showed that NSPs represent about 25-34% of the total composition, with most NSPs being insoluble (Table 2). Cellulose represents about 5-9% of corn DDGS content, and the predominant non-cellulosic polysaccharides are xylose (7.7%) and arabinoxylose (12.3-17.2%), which are mainly insoluble.
The mannose content in corn DDGS (1.7%) is substantially greater than in corn grain and is likely due to the man- nan content in residual yeast cell walls that are present in DDGS (Shurson, 2017).
Corn DDGS has a relatively high arabinose (6.2%) and uronic acid (1.6%) content, which results in relatively high arabinose-to-xylose and uronic acid-to-xylose ratios. This indicates that the fiber (heteroxylan) structure is complex and variable in corn DDGS and, there- fore, is more difficult to degrade with ex-ogenous enzymes than corn grain.
Klason lignin is not a well-defined chemical constituent and may contain protein (Maillard products), residual lipids and waxes and cutin in addition to true lignin. However, Klason lignin represents 1.5-4.7% of the chemical composition of DDGS.
These results suggest that the concentrations of substituted xylan and soluble NSPs in DDGS are altered during the production process and are substantially different from their original structure in corn grain.
We also need to achieve a better understanding of the physical structure of fiber — and identify practical and effective methods to degrade this structure — to improve the effectiveness of enzymes and the utilization of energy and nutrients in DDGS.
The primary cell wall structure in cereal grains is comprised of a skeleton of cellulosic microfibrils embedded in a matrix of hemicelluloses and smaller amounts of pectins, glycoproteins and hydroxycinnamatets. Subsequently, as the secondary cell wall develops during grain maturation, p-coumaryl, coniferyl and sinapyl alcohols are co-polymerized to form mixed lignins (Santiago et al., 2013). The addition of mixed lignins to the cell wall structure provides added strength to the fiber structure and causes it to be resistant to degradation.
In corn grain, the most abundant hemi- celluloses are arabinoxylans, which are comprised of a beta-(1-4)-d-xylan back- bone with substitutions of arabinose, glucuronic acid and acetic acid. The hemicellulose is tangled with cellulose microfibrils by hydrogen bonds (Photo). These hydrogen bonds cause the cell wall to be less accessible to degradation (Somerville et al., 2004).
However, this implies that arabinoxylan removal from the surface region of fiber by the addition of xylanases can result in exposure of cellulose microfibrils (crystalline structure), which are highly resistant to acids and enzymatic hydro- lysis (Hall et al., 2010).
In fact, Pedersen et al. (2015) reported that the AID (11.9%) and ATTD (29.0%) of cellulose in wheat DDGS for swine is less than the AID (37%) and ATTD (43.8%) in other fiber components. Therefore, it is possible that the more stable cellulosic microfibrils embed or trap arabinoxylans in corn DDGS, resulting in de- creased apparent total tract digestibility of fiber and preventing xylanase from accessing its substrates.
Furthermore, understanding the changes in the morphology of fiber be- fore and after the degradation processes may be useful in identifying approaches to improve the utilization of fiber in DDGS for pigs. Results from several studies have shown that crystalline celluloses are much more resistant to enzymatic hydrolysis compared to those with low crystallinity (Fan et al., 1980; Zhang and Lynd, 2004; Hall et al., 2010).
In addition, the crystallinity and crystal size of natural fiber sources have been shown to increase during thermal processing (Poletto et al., 2014). It is well known that production involves drying temperatures greater than 100°C as DDGS exits the dryer (Rosentrater et al., 2012). This may indicate that the most readily degradable fiber may have already been partially degraded during DDGS production, which would limit the effectiveness of feed enzymes or other processing technologies in diets containing DDGS.
We have initiated studies to assess the role of physical structure on fiber digestibility in corn DDGS for swine using X- ray diffraction to measure the extent of crystallization of two sources of DDGS with low (44.5%) and high (57.3%) ATTD of NDF (Kerr et al., 2013). Our initial results showed that the source of DDGS with less ATTD of NDF had a greater crystallinity index than the source with high ATTD of NDF (Figure). Therefore, increasing the degradability of dietary fiber in DDGS may require disruption of the crystalline structure of the fiber (Urriola et al., 2018).
References
Fan, L.T., Y.H. Lee and D.H. Beardmore. 1980. Mechanism of the enzymatic-hydrolysis of cellulose — Effects of major structural features of cellulose on enzymatic-hydro- lysis. Biotechnol. Bioengineer. 22:177-199.
Hall, M., P. Bansal, J.H. Lee, M.J. Realff and A.S. Bommarius. 2010. Cellulose crystallinity — A key predictor of the enzymatic hydrolysis rate. FEBS J. 277:1571-1582.
Kerr, B.J., W.A. Dozier III and G.C. Shurson. 2013. Effects of reduced-oil corn dis- tillers dried grains with solubles composition on digestible and metabolizable energy value and prediction in growing pigs. J. Anim. Sci. 91:3231-3243.
Kerr, B.J., N.K. Gabler and G.C. Shurson. 2015. Compositional effects of corn distillers dried grains with solubles with variable oil content on digestible, metabolizable and net energy values in growing pigs. Prof. Anim. Scientist 31:485-496.
National Research Council. 2007. Nutrient Requirements of Horses. 6th rev. ed. Natl. Acad. Press, Washington, D.C., p. 206.
National Research Council. 2012. Nutrient requirements of swine. 11th rev. ed. Natl. Acad. Press, Washington, D.C.
Noblet, J., and J. van Milgen. 2004. Energy value of pig feeds: Effect of pig bodyweight and energy evaluation system. J. Anim. Sci. 82(E. Suppl.):E229-E238.
Pedersen, M.B., S. Dalsgaard, K.E. Bach Knudsen, S. Yu and H.N. Laerke. 2014. Compositional profile and variation of distillers dried grains with solubles from various origins with focus on non-starch polysaccharides. Anim. Feed Sci. Technol. 197:130-141.
Pedersen, M.B., S. Yu, S. Arent, S. Dalsgaard, K.E. Bach Knudsen and H.N. Laerke. 2015. Xylanase increased the ileal digestibility of nonstarch polysaccharides and concentration of low molecular weight nondigestible carbohydrates in pigs fed high levels of wheat distillers dried grains with solubles. J. Anim. Sci. 93:2885-2893.
Poletto, M., H. Junior and A. Zattera. 2014. Native cellulose: Structure, characterization and thermal properties. Materials 7:6105- 6119.
Rosentrater, K.A., K. Ileleji and D.B. John- son. 2012. Manufacturing of fuel ethanol and distillers grains — Current and evolving processes. In: K. Liu, K.A. Rosentrater and B. Raton (eds.). Distillers Grains Production, Properties & Utilization. p. 73-102.
Santiago, R., J. Barros-Rios and R.A. Mal- var. 2013. Impact of cell wall composition on maize resistance to pests and diseases. Int. J. Mol. Sci. 14:6960-6980.
Shurson, G.C. 2017. Review — Yeast and yeast derivatives in feed additives and ingredients: Sources, characteristics, animal responses and quantification methods. Anim. Feed Sci. Technol. 235:60-76.
Somerville, C., S. Bauer, G. Brininstool, M. Facette, T. Hamann, J. Milne, E. Osborne,
A. Paredez, S. Persson, T. Raab, S. Vorwerk and H. Youngs. 2004. Toward a systems approach to understanding plant cell walls. Science 306:2206-2211.
Urriola, P.E., G.C. Shurson and H.H. Stein. 2010. Digestibility of dietary fiber in distillers coproducts fed to growing pigs. J. Anim. Sci. 88:2373-2381.
Urriola, P.E., Z.K. Zeng and G.C. Shurson. 2018. Strategies to modify the fiber structure and increase digestible energy content in corn distillers dried grains with solubles. Final Report to the National Bork Board, Des Moines, Iowa.
Zhang, Y.H., and L.R. Lynd. 2004. Toward an aggregated understanding of enzymatic hydrolysis of cellulose: Noncomplexed cellulase systems. Biotechnol. Bioeng. 88:797- 824.
© 2018 Feedstuffs. Reprinted with permission from Vol. 90, No. 10, October 1, 2018
The gross energy (GE) content of corn dried distillers grains plus solubles (DDGS) – at 5,429 kcal/kg of dry matter – is much greater than the 4.45 kcal/kg of dry matter for corn, 4,730 kcal/kg for soybean meal and the GE of other common ingredients used in swine diets.
However, the efficiency with which pigs utilize GE in DDGS, as measured by the net energy (NE)-to-GE ratio, is low (0.49) compared with corn – at 0.49 versus 0.68, respectively (National Research Council, 2012) – but NE:GE can vary from 0.42 to 0.46 among DDGS sources with variable crude fat content (Kerr et al., 2015).
Of all of the chemical components in feed ingredients, lipids provide the greatest amount of NE (kcal/kg), followed by crude protein and starch, with a limited amount provided from dietary fiber (Noblet and van Milgen, 2004). Therefore, the low starch content (0.84-3.89%, Kerr et al., 2013; 5.2-11.4%, Urriola et al., 2010) and relatively high neutral detergent fiber (NDF) content (30.2-39.7%; Pedersen et al., 2014) in corn DDGS are the primary reasons for the relatively low GE:NE.
To improve caloric and nutritional efficiency when feeding DDGS diets to pigs, we need to gain a better understanding of the chemical and physical characteristics that affect the digestibility and fermentability of fiber and explore approaches to convert a greater proportion of these calories to NE in pigs.
Fiber content in feed ingredients is measured using several different chemical methods, including crude fiber, NDF, acid detergent fiber (ADF) and total dietary fiber (TDF), which is comprised of insoluble dietary fiber (IDF) and soluble dietary fiber (SDF), as well as non-starch polysaccharides (NSPs), that quantify different types and proportions of in- digestible carbohydrates but may not adequately relate directly to nutritional and physiological effects in the animal (Infographic).
The most common measures that have been used to characterize the fiber content in DDGS are NDF, ADF and TDF (which can be separated into IDF and SDF fiber fractions). The NDF, ADF and hemicellulose (determined by the difference between NDF and ADF) contents, on a dry matter basis, of corn DDGS sources are in a range of 28.8-44.0%, 9.0-14.0% and 18.5-30.0%, respectively (Kerr et al., 2013).
The apparent total tract digestibility (ATTD) of NDF has been reported to range from 44.5% to 61.5% among 15 corn DDGS sources (Kerr et al., 2013). The range in TDF content (dry matter basis) of DDGS sources appears to be similar among studies, with Kerr et al. (2013) reporting a range of 30.8-37.8% TDF and Urriola et al. (2010) reporting a range of 32.9-38.6%. However, most of the TDF in DDGS is insoluble (31.8- 37.3%), with minimal amounts of soluble fiber (0.0-1.8%; Urriola et al., 2010).
The most comprehensive study to help us achieve a better understanding of the digestibility and fermentability of fiber in DDGS was conducted by Urriola et al. (2010). Results from three experiments showed that the ATTD of TDF ranged from 29.3% to 57.0% among corn DDGS sources. They also determined the apparent ileal digestibility (AID), ATTD and hindgut fermentability of ADF, NDF, IDF, SDF and TDF by feeding eight corn DDGS sources to growing pigs (Table 1).
The fiber digestibility and ferment- ability varied substantially among DDGS sources, regardless of the fiber measure used. This variability may be partially attributed to the amounts and types of enzymes as well as the different production processes used in ethanol and co- product production facilities to produce DDGS. Furthermore, about 46% of NDF and 29% of TDF is digestible in the small intestine, while about 13% of NDF and 21% of TDF is fermented in the hindgut.
These proportions of digestible and fermentable carbohydrates are assumed to contribute to the total metabolizable energy (ME) and NE content of DDGS, and the wide range in digestibility and fermentability among sources may partially contribute to the range in ME and NE content of DDGS reported in many studies. However, the AID of IDF is much less than SDF, and IDF is less ferment- able than SDF in the hindgut.
Although the average ATTD of NDF (59%) and TDF (50%) is moderately high in corn DDGS compared with some other types of high-fiber ingredients, there is a tremendous need to explore ways of improving AID and hindgut fermentation of the high proportion of IDF in DDGS to enhance the NE content.
Furthermore, the physical structure of fiber in corn DDGS may trap some of the oil in DDGS, prevent it from being accessible to digestive enzymes and contribute to the relatively low GE:ME and GE:NE contents. In fact, Kerr et al. (2013) reported that the ATTD of ether extract (oil) ranged from 53% to 81% among corn DDGS sources. Therefore, it is likely that the combined differences in fiber digestibility and fermentability, along with the variability of lipid digestibility involving fiber structure, are the primary reasons for the variable ME and NE content among DDGS sources fed to swine.
Although it is useful to understand the differences in the digestibility and fermentability of ADF, NDF, IDF and SDF in DDGS, knowledge of the NSP composition of DDGS is extremely important when selecting commercially available feed enzymes to improve the energy and nutrient digestibility of DDGS in pigs.
Pedersen et al. (2014) determined the NSP composition (dry matter basis) of 47 corn DDGS samples and showed that NSPs represent about 25-34% of the total composition, with most NSPs being insoluble (Table 2). Cellulose represents about 5-9% of corn DDGS content, and the predominant non-cellulosic polysaccharides are xylose (7.7%) and arabinoxylose (12.3-17.2%), which are mainly insoluble.
The mannose content in corn DDGS (1.7%) is substantially greater than in corn grain and is likely due to the man- nan content in residual yeast cell walls that are present in DDGS (Shurson, 2017).
Corn DDGS has a relatively high arabinose (6.2%) and uronic acid (1.6%) content, which results in relatively high arabinose-to-xylose and uronic acid-to-xylose ratios. This indicates that the fiber (heteroxylan) structure is complex and variable in corn DDGS and, there- fore, is more difficult to degrade with ex-ogenous enzymes than corn grain.
Klason lignin is not a well-defined chemical constituent and may contain protein (Maillard products), residual lipids and waxes and cutin in addition to true lignin. However, Klason lignin represents 1.5-4.7% of the chemical composition of DDGS.
These results suggest that the concentrations of substituted xylan and soluble NSPs in DDGS are altered during the production process and are substantially different from their original structure in corn grain.
We also need to achieve a better understanding of the physical structure of fiber — and identify practical and effective methods to degrade this structure — to improve the effectiveness of enzymes and the utilization of energy and nutrients in DDGS.
The primary cell wall structure in cereal grains is comprised of a skeleton of cellulosic microfibrils embedded in a matrix of hemicelluloses and smaller amounts of pectins, glycoproteins and hydroxycinnamatets. Subsequently, as the secondary cell wall develops during grain maturation, p-coumaryl, coniferyl and sinapyl alcohols are co-polymerized to form mixed lignins (Santiago et al., 2013). The addition of mixed lignins to the cell wall structure provides added strength to the fiber structure and causes it to be resistant to degradation.
In corn grain, the most abundant hemi- celluloses are arabinoxylans, which are comprised of a beta-(1-4)-d-xylan back- bone with substitutions of arabinose, glucuronic acid and acetic acid. The hemicellulose is tangled with cellulose microfibrils by hydrogen bonds (Photo). These hydrogen bonds cause the cell wall to be less accessible to degradation (Somerville et al., 2004).
However, this implies that arabinoxylan removal from the surface region of fiber by the addition of xylanases can result in exposure of cellulose microfibrils (crystalline structure), which are highly resistant to acids and enzymatic hydro- lysis (Hall et al., 2010).
In fact, Pedersen et al. (2015) reported that the AID (11.9%) and ATTD (29.0%) of cellulose in wheat DDGS for swine is less than the AID (37%) and ATTD (43.8%) in other fiber components. Therefore, it is possible that the more stable cellulosic microfibrils embed or trap arabinoxylans in corn DDGS, resulting in de- creased apparent total tract digestibility of fiber and preventing xylanase from accessing its substrates.
Furthermore, understanding the changes in the morphology of fiber be- fore and after the degradation processes may be useful in identifying approaches to improve the utilization of fiber in DDGS for pigs. Results from several studies have shown that crystalline celluloses are much more resistant to enzymatic hydrolysis compared to those with low crystallinity (Fan et al., 1980; Zhang and Lynd, 2004; Hall et al., 2010).
In addition, the crystallinity and crystal size of natural fiber sources have been shown to increase during thermal processing (Poletto et al., 2014). It is well known that production involves drying temperatures greater than 100°C as DDGS exits the dryer (Rosentrater et al., 2012). This may indicate that the most readily degradable fiber may have already been partially degraded during DDGS production, which would limit the effectiveness of feed enzymes or other processing technologies in diets containing DDGS.
We have initiated studies to assess the role of physical structure on fiber digestibility in corn DDGS for swine using X- ray diffraction to measure the extent of crystallization of two sources of DDGS with low (44.5%) and high (57.3%) ATTD of NDF (Kerr et al., 2013). Our initial results showed that the source of DDGS with less ATTD of NDF had a greater crystallinity index than the source with high ATTD of NDF (Figure). Therefore, increasing the degradability of dietary fiber in DDGS may require disruption of the crystalline structure of the fiber (Urriola et al., 2018).
References
Fan, L.T., Y.H. Lee and D.H. Beardmore. 1980. Mechanism of the enzymatic-hydrolysis of cellulose — Effects of major structural features of cellulose on enzymatic-hydro- lysis. Biotechnol. Bioengineer. 22:177-199.
Hall, M., P. Bansal, J.H. Lee, M.J. Realff and A.S. Bommarius. 2010. Cellulose crystallinity — A key predictor of the enzymatic hydrolysis rate. FEBS J. 277:1571-1582.
Kerr, B.J., W.A. Dozier III and G.C. Shurson. 2013. Effects of reduced-oil corn dis- tillers dried grains with solubles composition on digestible and metabolizable energy value and prediction in growing pigs. J. Anim. Sci. 91:3231-3243.
Kerr, B.J., N.K. Gabler and G.C. Shurson. 2015. Compositional effects of corn distillers dried grains with solubles with variable oil content on digestible, metabolizable and net energy values in growing pigs. Prof. Anim. Scientist 31:485-496.
National Research Council. 2007. Nutrient Requirements of Horses. 6th rev. ed. Natl. Acad. Press, Washington, D.C., p. 206.
National Research Council. 2012. Nutrient requirements of swine. 11th rev. ed. Natl. Acad. Press, Washington, D.C.
Noblet, J., and J. van Milgen. 2004. Energy value of pig feeds: Effect of pig bodyweight and energy evaluation system. J. Anim. Sci. 82(E. Suppl.):E229-E238.
Pedersen, M.B., S. Dalsgaard, K.E. Bach Knudsen, S. Yu and H.N. Laerke. 2014. Compositional profile and variation of distillers dried grains with solubles from various origins with focus on non-starch polysaccharides. Anim. Feed Sci. Technol. 197:130-141.
Pedersen, M.B., S. Yu, S. Arent, S. Dalsgaard, K.E. Bach Knudsen and H.N. Laerke. 2015. Xylanase increased the ileal digestibility of nonstarch polysaccharides and concentration of low molecular weight nondigestible carbohydrates in pigs fed high levels of wheat distillers dried grains with solubles. J. Anim. Sci. 93:2885-2893.
Poletto, M., H. Junior and A. Zattera. 2014. Native cellulose: Structure, characterization and thermal properties. Materials 7:6105- 6119.
Rosentrater, K.A., K. Ileleji and D.B. John- son. 2012. Manufacturing of fuel ethanol and distillers grains — Current and evolving processes. In: K. Liu, K.A. Rosentrater and B. Raton (eds.). Distillers Grains Production, Properties & Utilization. p. 73-102.
Santiago, R., J. Barros-Rios and R.A. Mal- var. 2013. Impact of cell wall composition on maize resistance to pests and diseases. Int. J. Mol. Sci. 14:6960-6980.
Shurson, G.C. 2017. Review — Yeast and yeast derivatives in feed additives and ingredients: Sources, characteristics, animal responses and quantification methods. Anim. Feed Sci. Technol. 235:60-76.
Somerville, C., S. Bauer, G. Brininstool, M. Facette, T. Hamann, J. Milne, E. Osborne,
A. Paredez, S. Persson, T. Raab, S. Vorwerk and H. Youngs. 2004. Toward a systems approach to understanding plant cell walls. Science 306:2206-2211.
Urriola, P.E., G.C. Shurson and H.H. Stein. 2010. Digestibility of dietary fiber in distillers coproducts fed to growing pigs. J. Anim. Sci. 88:2373-2381.
Urriola, P.E., Z.K. Zeng and G.C. Shurson. 2018. Strategies to modify the fiber structure and increase digestible energy content in corn distillers dried grains with solubles. Final Report to the National Bork Board, Des Moines, Iowa.
Zhang, Y.H., and L.R. Lynd. 2004. Toward an aggregated understanding of enzymatic hydrolysis of cellulose: Noncomplexed cellulase systems. Biotechnol. Bioeng. 88:797- 824.
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