Roles of structural phenylpropanoids in forage cell wall digestion

Phenolic constituents lignins and phenolic acids) and carbohydrates are assembled in a tight architecture which differs according to the plant species. During cell wall digestion, the hydrolysis kinetics differ between carbohydrates and seem to depend chiefly on the content and organisation of tissue phenolics. Among the phenylpropanoids, ferulic acid is released more quickly than p-coumaric acid. Lignins remain largely in the cell walls. They also undergo transformations, chiefly solubilisation as lignin carbohydrate complexes. The limiting effect of lignins on cell wall degradation increases with increasing content. However, their effect on degradation might also depend on qualitative factors such as lignin structure and polymer organisation in walls and tissues. When various grasses (normal and selected genotypes), or grasses and legumes are compared, correlations between certain factors such as lignin uncondensed fraction, syringyl units or phenolic acids contents and cell wall degradation emerge but not clear causal relationship has been shown. Nonetheless, other structural characteristics, related to the alkali reactivity of lignins, seem to have a stronger influence on cell wall degradation. Phenylpropanoids seem to act mainly as a physical and biochemical barrier to the action of the microbial enzymes. In addition, their reactivity as phenolic compounds and their hydrophobicity seem to play a role. Digestion is not limited only by phenolics. The factors that limit glycanolysis--the accessibility, crystallinity and capillary structure of cellulose and the branching of hemicelluloses--seem to have little or no effect on cell wall degradation in vivo. In contrast, other antiquality substances (tannins, cutin and silica), plant anatomy, environmental factors, factors modulating microbial growth and animal physiology influence cell wall utilisation. Future research in this field should focus on the effects of phenolic structure and of cell wall and tissue organisation on carbohydrate degradation.