Cell Wall Lignification and Degradability

During the past century, analytical techniques applied to the study of plant cell wall chemistry have become more refined and increasingly sophisticat­ ed. Concepts attempting to describe the relationship of lignin to other cell wall constituents have evolved in concert with analytical ability. Early no­ tions of cell wall lignification described lignin as an amorphous polymer acting as a glue or cement in plant tissue (Bailey, 1936). Contemporary dogma recog­ nizes a more intimate, direct relationship of lignin with cell wall polysaccha­ rides via specific chemical bonds (Watanabe et aI., 1989). An understanding of the impact of these specific chemical relationships on ruminal cell wall degradation of agronomic crops will be necessary to advance knowledge in ruminant nutrition and plant physiology, and to capitalize on opportunities for precise application of molecular genetic techniques to rumen microbial or plant systems. Nearly 60 yr ago, Buston (1935) studied the relationship between pec­ tin, hemicellulose, and lignin in plants and concluded that "it is not consid­ ered that pectin is a precursor of lignin." Other attempts to elucidate cell wall lignification by Griffioen (1938) indicated "young lignins" from young­ er portions of sunflower (Helianthus annus L.) stalks were less polymerized than "old lignins" present in old stalk portions. Research in the mid-1920s (Waksman, 1926a) recognized that lignins in straw were resistant to fungal and bacterial degradation in the process of soil humus formation. Thus, lig­ nins and tannins from degrading soil organic matter accumulate under anaero­ bic conditions (Waksman, 1926b). Extension of these early studies concluded that any depressing effect of lignin on anaerobic cellulose decomposition "must be due to the manner of its binding with cellulose" (Waksman & Cor­ don, 1938). Partial destruction of "the bond between lignin and cellulose," as well as dissolution of lignin was achieved by pretreating straw with 150 g kg-I ammonium hydroxide for periods of up to 15 d (Nikolaeva, 1938). This pretreatment improved coefficients of digestion for nutrients. However, this improvement was not directly related to the amount of lignin removed

[1]  D. R. Buxton,et al.  Chemical and in vitro digestible dry matter composition of maize stalks after selection for stalk strength and stalk-rot resistance , 1986 .

[2]  M. Vignon,et al.  Effect of steam explosion treatment on the physico-chemical characteristics and enzymic hydrolysis of poplar cell wall components , 1991 .

[3]  Van Soest,et al.  Development of a Comprehensive System of Feed Analyses and its Application to Forages , 1967 .

[4]  N. Lewis,et al.  Phenolic constituents of plant cell walls and wall biodegradability. , 1989 .

[5]  G. Fahey,et al.  Interactions Among Phenolic Monomers and In Vitro Fermentation , 1983 .

[6]  R. E. Hellwig,et al.  Performance of steers on Pensacola bahiagrass, Coastal bermudagrass and Coastcross-1 bermudagrass pastures and pellets. , 1972 .

[7]  D. E. Akin,et al.  Microspectrophotometry of phenolic compounds in bermudagrass cell walls in relation to rumen microbial digestion. , 1990 .

[8]  A. Chesson Lignin-polysaccharide complexes of the plant cell wall and their effect on microbial degradation in the rumen , 1988 .

[9]  A. J. Gordon,et al.  Chemical and in vivo evaluation of a brown midrib mutant of Zea mays. I. Fibre, lignin and amino acid composition and digestibility for sheep. , 1973, Journal of the science of food and agriculture.

[10]  M. Casler,et al.  Lignin concentration and composition of divergent smooth bromegrass genotypes , 1990 .

[11]  E. Guittet,et al.  Ether linkage between phenolic acids and lignin fractions from wheat straw , 1985 .

[12]  N. Terashima,et al.  Formation and structure of lignin in monocotyledons. III. Heterogeneity of sugarcane (saccharum officinarum L.) Lignin with respect to the composition of structural units in different morphological regions , 1990 .

[13]  T. Higuchi,et al.  Lignin biochemistry: Biosynthesis and biodegradation , 1990, Wood Science and Technology.

[14]  É. Grenet,et al.  Microbial degradation of normal maize and bm3 maize in the rumen observed by scanning electron microscopy , 1991 .

[15]  J. Reeves Pyrolysis Gas Chromatography of Five Ruminant Feedstuffs at Various Stages of Growth , 1990 .

[16]  V. L. Lechtenberg,et al.  Evaluation of a Brown Midrib Mutant of Zea Mays L. , 1972 .

[17]  K. Vogel,et al.  Influence of lignin on digestibility of forage cell wall material. , 1986, Journal of animal science.

[18]  J N Saddler,et al.  Steam pretreatment of lignocellulosic material for enhanced enzymatic hydrolysis , 1987, Biotechnology and bioengineering.

[19]  F. M. Engels Some Properties of Cell Wall layers Determining Ruminant Digestion , 1989 .

[20]  W. F. Wedin,et al.  Cell-wall composition and digestibility of alfalfa stems and leaves , 1987 .

[21]  Hansang Jung Inhibitory potential of phenolic-carbohydrate complexes released during ruminal fermentation , 1988 .

[22]  G. Fahey,et al.  Simple phenolic monomers of forages and effects of in vitro fermentation on cell wall phenolics. , 1983 .

[23]  J. Reeves Lignin Composition and In Vitro Digestibility of Feeds , 1985 .

[24]  J. Gressel,et al.  Partial suppression of cellulase action by artificial lignification of cellulose , 1983 .

[25]  L. D. Muller,et al.  Brown-Midrib Corn Silage for Lactating Dairy Cows , 1979 .

[26]  A. Patton,et al.  Seasonal Changes in the Lignin and Cellulose Content of Some Montana Grasses , 1942 .

[27]  J. Saddler,et al.  The interaction of xylanases with commercial pulps , 1991, Biotechnology and bioengineering.

[28]  I. Morrison Changes in the hemicellulosic polysaccharides of rye-grass with increasing maturity. , 1974, Carbohydrate research.

[29]  J. Reeves Lignin Composition of Chemically Treated Feeds as Determined by Nitrobenzene Oxidation and Its Relationship to Digestibility , 1985 .

[30]  A. Bailey Lignin in douglas fir: Composition of the middle lamella , 1936 .

[31]  D. E. Akin Histological and Physical Factors Affecting Digestibility of Forages , 1989 .

[32]  J. C. Street,et al.  Correlations of Phenolic Acids and Xylose Content of Cell Wall with In Vitro Dry Matter Digestibility of Three Maturing Grasses , 1984 .

[33]  C. H. Gordon,et al.  Relationships of Forage Compositions With Rates of Cell Wall Digestion and Indigestibility of Cell Walls , 1972 .

[34]  Hansang Jung,et al.  Relationship of lignin and esterified phenolics to fermentation of smooth bromegrass fibre , 1991 .

[35]  J. Ralph,et al.  Pyrolysis-GC-MS characterization of forage materials , 1991 .

[36]  J. Cherney,et al.  Forage quality and digestion kinetics of switchgrass herbage and morphological components , 1988 .

[37]  Bruce A. Stone,et al.  Distribution of free and combined phenolic acids in wheat internodes , 1990 .

[38]  C. Lapierre,et al.  Thioacidolysis of Poplar Lignins: Identification of Monomeric Syringyl Products and Characterization of Guaiacyl-Syringyl Lignin Fractions , 1986 .

[39]  C. V. Sumere,et al.  Free and bound phenolic acids of lucerne (Medicago sativa cv europe) , 1980 .

[40]  R. Hartley,et al.  Aromatic aldehyde constituents of graminaceous cell walls , 1984 .

[41]  M. P. Bryant,et al.  Syntrophococcus sucromutans sp. nov. gen. nov. uses carbohydrates as electron donors and formate, methoxymonobenzenoids or Methanobrevibacter as electron acceptor systems , 2004, Archives of Microbiology.

[42]  D. E. Akin,et al.  Effect of forage cell wall phenolic acids and derivatives on rumen microflora , 1989 .

[43]  P. Doyle,et al.  The feeding value of cereal straws for sheep. II. Rice straws , 1990 .

[44]  K. Vogel,et al.  Divergent Selection for In Vitro Dry Matter Digestibility in Switchgrass1 , 1981 .

[45]  M. Demment,et al.  Changes in Forage Quality of Improved Alfalfa Populations 1 , 1986 .

[46]  Van Soest Symposium on Nutrition and Forage and Pastures: New Chemical Procedures for Evaluating Forages , 1964 .

[47]  R. H. Hart,et al.  Improving Forage Quality in Bermudagrass by Breeding 1 , 1967 .

[48]  J. Patterson,et al.  Digestibility and feeding value of pearl millet as influenced by the brown-midrib, low-lignin trait. , 1990, Journal of animal science.

[49]  S. Waksman ON THE ORIGIN AND NATURE OF THE SOIL ORGANIC MATTER OR SOIL “HUMUS”: V. THE ROLE OF MICROÖRGANISMS IN THE FORMATION OF “HUMUS” IN THE SOIL1 , 1926 .

[50]  S. Loerch,et al.  Modification of a colorimetric analysis for lignin and its use in studying the inhibitory effects of lignin on forage digestion by ruminal microorganisms. , 1991, Journal of animal science.

[51]  V. L. Lechtenberg,et al.  Effect of Lignin on Rate of In Vitro Cell Wall and Cellulose Disappearance in Corn , 1974 .

[52]  J. Conchie,et al.  Soluble lignin-carbohydrate complexes from sheep rumen fluid: their composition and structural features. , 1988, Carbohydrate research.

[53]  K. Wong,et al.  Ultrastructure of steam-exploded wood , 1988, Wood Science and Technology.

[54]  N. Rodionova,et al.  Studies on xylan-degrading enzymes. II. Action pattern of endo-1,4-beta-xylanase from Aspergillus niger str. 14 on xylan and xylooligosaccharides. , 1977, Biochimica et biophysica acta.

[55]  T. Clark,et al.  The relationship between fiber‐porosity and cellulose digestibility in steam‐exploded Pinus radiata , 1988, Biotechnology and bioengineering.

[56]  K. Moore,et al.  Rate and Extent of Digestion of Cell Wall Components of Brown‐Midrib Sorghum Species 1 , 1986 .

[57]  P. Åman,et al.  Profile of fibre composition in lucerne (Medicago sativa) hay and rumen digesta as influenced by particle size and time after feeding. , 1990 .

[58]  V. L. Lechtenberg,et al.  Phenotype, Fiber Composition, and in vitro Dry Matter Disappearance of Chemically Induced Brown Midrib (bmr) Mutants of Sorghum 1 , 1978 .

[59]  R. Farrell,et al.  Enzymatic "combustion": the microbial degradation of lignin. , 1987, Annual review of microbiology.

[60]  K. Moore,et al.  Digestion Kinetics and Cell Wall Composition of Brown Midrib Sorghum ✕ Sudangrass Morphological Components , 1990 .

[61]  I. Morrison,et al.  The degradation of isolated hemicelluloses and lignin-hemicellulose complexes by cell-free, rumen hemicellulases , 1982 .

[62]  Bruce A. Stone,et al.  Phenolic acid bridges between polysaccharides and lignin in wheat internodes , 1990 .

[63]  W. Wales,et al.  The feeding value of cereal straws for sheep. I. Wheat straws , 1990 .

[64]  D. Buxton,et al.  Morphology of Alfalfa Divergently Selected for Herbage Lignin Concentration , 1989 .

[65]  W. R. Windham,et al.  Normal‐12 and Brown Midrib‐12 Sorghum. II. Chemical Variations and Digestibility1 , 1986 .

[66]  Ronald D. Hatfield,et al.  Degradability of phenolic acid-hemicellulose esters: A model system , 1991 .

[67]  Takashi Watanabe,et al.  Binding-site Analysis of the Ether Linkages between Lignin and Hemicelluloses in Lignin-Carbohydrate Complexes by DDQ-Oxidation , 1989 .

[68]  J. Cherney,et al.  Forage Quality Characterization of a Chemically Induced Brown-Midrib Mutant in Pearl Millet , 1988 .

[69]  W. Manders Solid-State 13C NMR Determination of the Syringyl/Guaiacyl Ratio in Hardwoods , 1987 .

[70]  R. Hartley,et al.  Cyclodimers of p-coumaric and ferulic acids in the cell walls of tropical grasses , 1990 .

[71]  Hansang Jung,et al.  Influence of Forage Phenolics on Ruminal Fibrolytic Bacteria and In Vitro Fiber Degradation , 1986, Applied and environmental microbiology.

[72]  D. Goring,et al.  Chemical characterization of tissue fractions from the middle lamella and secondary wall of black spruce tracheids , 1982, Wood Science and Technology.

[73]  Hansang Jung,et al.  Forage Lignins and Their Effects on Fiber Digestibility , 1989 .

[74]  D. E. Akin,et al.  Effect of phenolic monomers on the growth and beta-glucosidase activity of Bacteroides ruminicola and on the carboxymethylcellulase, beta-glucosidase, and xylanase activities of Bacteroides succinogenes , 1988, Applied and environmental microbiology.

[75]  S. L. Fales,et al.  Cinnamic acid–carbohydrate esters: An evaluation of a model system† , 1989 .

[76]  H. W. Buston Observations on the nature, distribution and development of certain cell-wall constituents of plants. , 1935, The Biochemical journal.

[77]  K. Vogel,et al.  Lignification of switchgrass (Panicum virgatum) and big bluestem (Andropogon gerardii) plant parts during maturation and its effect on fibre degradability , 1992 .

[78]  K. Moore,et al.  Effect of brown midrib and normal genotypes of sorghum X sudangrass on ruminal fluid and particulate rate of passage from the rumen and extent of digestion at various sites along the gastrointestinal tract in sheep. , 1988, Journal of animal science.

[79]  B. Kos̆iková,et al.  Antibiotic properties of lignin components , 2008, Folia Microbiologica.

[80]  D. R. Buxton,et al.  Lignin constituents and cell-wall digestibility of grass and legume stems , 1988 .

[81]  J. Labavitch,et al.  alpha-L-arabinofuranosidase from Ruminococcus albus 8: purification and possible role in hydrolysis of alfalfa cell wall , 1984, Applied and environmental microbiology.

[82]  T. Schultz,et al.  Proposed Mechanism for the Nitrobenzene Oxidation of Lignin , 1986 .

[83]  G. Fahey,et al.  Effects of ruminant digestion and metabolism on phenolic monomers of forages , 1983, British Journal of Nutrition.

[84]  J. Boon An Introduction to Pyrolysis Mass Spectrometry of Lingnocellulosic Material: Case Studies on Barley Straw, Corn Stem and Agropyron , 1989 .

[85]  A. Boudet,et al.  Comparison of lignins and of enzymes involved in lignification in normal and brown midrib (bm3) mutant corn seedlings , 1985 .

[86]  Hansang Jung,et al.  Depression of cellulose digestion by esterified cinnamic acids , 1986 .

[87]  G. N. Richards,et al.  Presence of soluble lignin-carbohydrate complexes in the bovine rumen. , 1975, Carbohydrate research.

[88]  R. Hartley,et al.  Monomeric and dimeric phenolic constituents of plant cell walls—possible factors influencing wall biodegradability , 1990 .

[89]  J. Azuma,et al.  Lignin-Carbohydrate Complexes and Phenolic Acids in Bagasse , 1984 .

[90]  F. M. Engels,et al.  Influence of growth temperature on anatomy and in vitro digestibility of maize tissues , 1990, The Journal of Agricultural Science.

[91]  G. Fahey,et al.  Effects of Lignification, Cellulose Crystallinity and Enzyme Accessible Space on the Digestibility of Plant Cell Wall Carbohydrates by the Ruminant , 1988 .

[92]  Jürgen Puls,et al.  Differences in Xylan Degradation by Various Noncellulolytic Thermophilic Anaerobes and Clostridium thermocellum , 1985, Applied and environmental microbiology.

[93]  K. Vogel,et al.  Forage Quality and Performance of Yearlings Grazing Switchgrass Strains Selected for Differing Digestibility , 1988 .

[94]  K. Vogel,et al.  Alkali-Labile Cell-Wall Phenolics and Forage Quality in Switchgrasses selected for Differing Digestibility , 1990 .

[95]  R. Elliott,et al.  Biodegradability of mature grass cell walls in relation to chemical composition and rumen microbial activity , 1987, The Journal of Agricultural Science.

[96]  A. J. Gordon A comparison of some chemical and physical properties of alkali lignins from grass and lucerne hays before and after digestion by sheep , 1975 .

[97]  B. A. Dehority,et al.  Relationship of Lignification to In Vitro Cellulose Digestibility of Grasses and Legumes , 1965 .

[98]  B. A. Dehority,et al.  Discrepancies between grasses and alfalfa when estimating nutritive value from in vitro cellulose digestibility by rumen microorganisms. , 1962 .

[99]  J. Knowles,et al.  Three-dimensional structure of cellobiohydrolase II from Trichoderma reesei. , 1990, Science.

[100]  R. Helm,et al.  Synthesis of methyl 5-O-trans-feruloyl-alpha-L-arabinofuranoside and its use as a substrate to assess feruloyl esterase activity. , 1991, Analytical biochemistry.

[101]  R. Hartley,et al.  p‐Coumaric and ferulic acid components of cell walls of ryegrass and their relationships with lignin and digestibility , 1972 .

[102]  A. Chesson,et al.  Influence of Plant Phenolic Acids on Growth and Cellulolytic Activity of Rumen Bacteria , 1982, Applied and environmental microbiology.

[103]  D. Delmer,et al.  9 – Biosynthesis of Plant Cell Walls , 1988 .

[104]  V. L. Lechtenberg,et al.  Brown Midrib Mutants in Sudangrass and Grain Sorghum1 , 1981 .

[105]  T. Lam,et al.  Lignin in wheat internodes. Part 1: The reactivities of lignin units during alkaline nitrobenzene oxidation , 1990 .

[106]  R. Weller,et al.  The feeding value of normal and brown midrib-3 maize silage , 1986, The Journal of Agricultural Science.

[107]  M. Casler,et al.  Morphological and Chemical Responses to Selection for in Vitro Dry Matter Digestibility in Smooth Bromegrass , 1989 .

[108]  D. Buxton Cell-wall components in divergent germplasms of four perennial forage grass species. , 1990 .

[109]  Norman G. Lewis,et al.  Phenylpropanoid metabolism in cell walls: an overview. , 1989 .

[110]  N. Lewis,et al.  Towards a working model of the growing plant cell wall. Phenolic cross-linking reactions in the primary cell walls of dicotyledons. , 1989 .

[111]  R. Hartley,et al.  Linkage of p-coumaroyl and feruloyl groups to cell-wall polysaccharides of barley straw , 1986 .

[112]  M. P. Bryant,et al.  Eubacterium oxidoreducens sp. nov. requiring H2 or formate to degrade gallate, pyrogallol, phloroglucinol and quercetin , 1986, Archives of Microbiology.

[113]  Antony Bacic,et al.  8 – Structure and Function of Plant Cell Walls , 1988 .