USDA-Agricultural Research Service, Plant Science Research Unit, St. Paul, MN 55108, USA.
Plant Sci. 2012 Apr;185-186:65-77. doi: 10.1016/j.plantsci.2011.10.014. Epub 2011 Oct 25.
Improving digestibility of roughage cell walls will improve ruminant animal performance and reduce loss of nutrients to the environment. The main digestibility impediment for dicotyledonous plants is highly lignified secondary cell walls, notably in stem secondary xylem, which become almost non-digestible. Digestibility of grasses is slowed severely by lignification of most tissues, but these cell walls remain largely digestible. Cell wall lignification creates an access barrier to potentially digestible wall material by rumen bacteria if cells have not been physically ruptured. Traditional breeding has focused on increasing total dry matter digestibility rather than cell wall digestibility, which has resulted in minimal reductions in cell wall lignification. Brown midrib mutants in some annual grasses exhibit small reductions in lignin concentration and improved cell wall digestibility. Similarly, transgenic approaches down-regulating genes in monolignol synthesis have produced plants with reduced lignin content and improved cell wall digestibility. While major reductions in lignin concentration have been associated with poor plant fitness, smaller reductions in lignin provided measurable improvements in digestibility without significantly impacting agronomic fitness. Additional targets for genetic modification to enhance digestibility and improve roughages for use as biofuel feedstocks are discussed; including manipulating cell wall polysaccharide composition, novel lignin structures, reduced lignin/polysaccharide cross-linking, smaller lignin polymers, enhanced development of non-lignified tissues, and targeting specific cell types. Greater tissue specificity of transgene expression will be needed to maximize benefits while avoiding negative impacts on plant fitness.cauliflower mosiac virus (CaMV) 35S promoter.
提高粗饲料细胞壁的消化率将提高反刍动物的生产性能,并减少营养物质向环境中的损失。双子叶植物的主要消化障碍是高度木质化的次生细胞壁,尤其是在茎次生木质部中,这些细胞壁几乎不可消化。大多数组织的木质化严重减缓了草的消化速度,但这些细胞壁仍然具有很大的可消化性。如果细胞没有被物理破裂,细胞壁木质化会为瘤胃细菌提供潜在可消化细胞壁物质的进入障碍。传统的育种方法侧重于提高总干物质消化率,而不是细胞壁消化率,这导致细胞壁木质化的减少非常有限。一些一年生草本植物中的棕色中脉突变体表现出木质素浓度降低和细胞壁消化率提高。同样,下调单酚合成基因的转基因方法也产生了木质素含量降低和细胞壁消化率提高的植物。虽然木质素浓度的大幅降低与植物适应性差有关,但木质素的小幅度降低提供了可衡量的消化率改善,而对农艺适应性没有显著影响。还讨论了其他用于增强消化率和改善粗饲料作为生物燃料原料的遗传修饰目标;包括操纵细胞壁多糖组成、新型木质素结构、减少木质素/多糖交联、较小的木质素聚合物、增强无木质化组织的发育以及针对特定细胞类型。为了最大限度地发挥效益,同时避免对植物适应性的负面影响,需要对转基因表达进行更大的组织特异性。