Sage Rowan F, McKown Athena D
Department of Botany, University of Toronto, 25 Willcocks Street, Toronto, ON M5S3B2 Canada.
J Exp Bot. 2006;57(2):303-17. doi: 10.1093/jxb/erj040. Epub 2005 Dec 19.
C4 photosynthesis is a complex specialization that enhances carbon gain in hot, often arid habitats where photorespiration rates can be high. Certain features unique to C4 photosynthesis may reduce the potential for phenotypic plasticity and photosynthetic acclimation to environmental change relative to what is possible with C3 photosynthesis. During acclimation, the structural and physiological integrity of the mesophyll-bundle sheath (M-BS) complex has to be maintained if C4 photosynthesis is to function efficiently in the new environment. Disruption of the M-BS structure could interfere with metabolic co-ordination between the C3 and C4 cycles, decrease metabolite flow rate between the tissues, increase CO2 leakage from the bundle sheath, and slow enzyme activity. C4 plants have substantial acclimation potential, but in most cases lag behind the acclimation responses in C3 plants. For example, some C4 species are unable to maintain high quantum yields when grown in low-light conditions. Others fail to reduce carboxylase content in shade, leaving substantial over-capacity of Rubisco and PEP carboxylase in place. Shade-tolerant C4 grasses lack the capacity for maintaining a high state of photosynthetic induction following sunflecks, and thus may be poorly suited to exploit subsequent sunflecks compared with C3 species. In total, the evidence indicates that C4 photosynthesis is less phenotypically plastic than C3 photosynthesis, and this may contribute to the more restricted ecological and geographical distribution of C4 plants across the Earth.
C4光合作用是一种复杂的特殊生理机制,它能在炎热且通常干旱的生境中提高碳同化效率,因为这些生境中的光呼吸速率可能很高。相对于C3光合作用而言,C4光合作用的某些独特特征可能会降低表型可塑性以及光合适应环境变化的潜力。在适应过程中,如果C4光合作用要在新环境中高效发挥作用,叶肉-维管束鞘(M-BS)复合体的结构和生理完整性必须得以维持。M-BS结构的破坏可能会干扰C3和C4循环之间的代谢协调,降低组织间代谢物的流动速率,增加维管束鞘中二氧化碳的泄漏,并减缓酶活性。C4植物具有相当大的适应潜力,但在大多数情况下落后于C3植物的适应反应。例如,一些C4物种在弱光条件下生长时无法维持高量子产率。其他一些C4物种在遮荫条件下无法降低羧化酶含量,导致核酮糖-1,5-二磷酸羧化酶(Rubisco)和磷酸烯醇式丙酮酸羧化酶大量过剩。耐荫C4禾本科植物在光斑后缺乏维持高光合诱导状态的能力,因此与C3物种相比,可能不太适合利用随后出现的光斑。总体而言,证据表明C4光合作用的表型可塑性低于C3光合作用,这可能导致C4植物在全球的生态和地理分布更为受限。