胡杨异形叶光合作用对光强与CO2浓度的响应

胡杨(Populus euphratica)叶形多变, 随个体生长发育, 植株出现条形、卵形和锯齿阔卵形叶。在新疆塔里木河上游人工胡杨林内选择具有此3种叶形的成年标准株, 将枝条拉至同一高度, 通过活体测定, 比较其光合作用-光与CO₂响应及叶绿素荧光响应特征。结果表明: 胡杨异形叶光合速率对光强/CO₂浓度与电子传递速率对光强的响应曲线均可用直角双曲线修正模型来拟合, 得出的主要光合参数与实测值较吻合。胡杨卵形叶、锯齿阔卵形叶光合速率-光响应参数与生化参数及快速光响应参数与条形叶差异显著, 而光合速率-CO₂响应参数则无显著差异。胡杨异形叶CO₂饱和浓度下的最大净光合速率(P_nmax)较饱和光强下的P_nmax高, 表明胡杨强光下光合速率在很大程度上受CO₂供应和1,5-二磷酸核酮糖(RuBP)再生能力的限制。卵形叶、锯齿阔卵形叶的初始量子效率(α)、初始羧化效率(CE)、P_nmax、光合能力(A_max)与最大羧化速率(V_cmax)均显著高于条形叶; 锯齿叶光饱和点(LSP)、最大电子传递速率(ETR_max)与光呼吸速率(R_p)高于卵形叶, 条形叶光补偿点(LCP)与LSP、α、CE最低。表明荒漠干旱环境下胡杨锯齿叶最耐强光, 高R_p可能是其耗散过剩光能、保护光合机构免于强光破坏的重要途径; 卵形叶高的α、CE、磷酸丙糖利用效率(TPU)、PSII实际光化学效率(Φ_PSII)与低LCP及叶氮分配策略是其保持高光合速率的原因; 条形叶Φ_PSII、ETR、P_n低, 因其制造光合产物不足而难以满足树体生长逐渐减少并处于树冠下部。可见, 胡杨条形叶光合效率低、抗逆性差, 主要以维持生长为主; 随着树体长大, 条形叶难以适应荒漠环境来维系其生长, 出现了卵形叶; 卵形叶光合效率高, 易于快速积累光合产物而加快树体生长, 但其LSP低和耐光抑制能力弱, 逐渐被更耐强光、高温与大气干旱的锯齿叶所取代, 从而使胡杨在极端逆境下得以生存, 这是胡杨从幼苗到成年叶形变化及异形叶着生在树冠不同高度的原因。

Photosynthetic responses of the heteromorphic leaves in Populus euphratica to light intensity and CO2 concentration

Aims Populus euphratica is an important tree species and its leaf shape changes along the growth stages. Adult trees commonly comprise polymorphic leaves, including lanceolate, oval and serrated broad-oval leaves. Our objective were to elucidate the ecophysiological mechanisms of P. euphratica adapting to high temperature and strong light environment and its survival strategy by comparing photosynthetic efficiency and chlorophyll fluorescence parameters in heteromorphic leaves in an extremely arid desert area, and to explore the causes of changes in leaf shape in P. euphratica, in order to provide a scientific basis for the protection of desert P. euphratica forests.
Methods Individuals with 10 cm diameter at breast height from a planted P. euphratica forest were selected. Measurements were made on the parameters of gas changes and chlorophyll fluorescence of three different leaf shapes on branches at the similar height using a LI-6400 Portable Photosynthesis System and a PAM-2100 chlorophyll fluorometer. The light/CO₂ response curves of net photosynthetic rate (P_n) and rapid light curves of chlorophyll fluorescence in heteromorphic leaves were fitted and analyzed.
Important findings The light and CO₂ response curves, rapid light curves of the three different leaf shapes in P. euphratica were better fitted by the modified rectangular hyperbola models, and the model values of key photosynthetic parameters were very close to the measured data. There were significant differences in the light responses, biochemical parameters and the parameters of rapid light curves among the oval, serrated broad-oval leaves and lanceolate leaves, but the heteromorphic leaves did not significantly differ in carbon assimilationefficiency. The maximum net photosynthetic rate (P_nmax) of the heteromorphic leaves under saturated intercellular CO₂ concentration was higher than under saturated irradiance, indicating that photosynthetic efficiency was limited to the great extent by CO₂ supply and regeneration rate of ribulose biphosphate (RuBP). Initial quantum yield (α), initial carboxylation efficiency (CE), P_nmax, photosynthetic capacity (A_max), maximum carboxylation rate (V_cmax) were greater in the oval and serrated broad-oval leaves than in the lanceolate leaves; the serrated broad-oval leaves had the highest light saturation point (LSP), photosynthetic electron transportation rate (ETR_max) and rate of photorespiration (R_p), whereas the lanceolate leaves had the lowest light compensation point (LCP), LSP, α and CE. All the results above indicate that the serrated broad-oval leaves having greater resistance to strong light and higher R_p may be an important mechanism for dissipating excessive light energy and protecting the photosynthetic apparatus from light damage. In contrast, the oval leaves had higher values in α, CE, triose-phosphate utilization efficiency (TPU), PSII actual photochemical efficiency (Φ_PSII), leaf nitrogen allocation strategy and low LCP and therefore could maintain high photosynthetic rate in extremely arid areas. The lanceolate leaves had the lowest values in P_n, Φ_PSII, and ETR, which would be difficult to meet the individual growth demand because of the low production of photosynthate, and their number declined with growth and distributed mainly toward the lower tree crowns.