Smith J A C, Popp M, Lüttge U, Cram W J, Diaz M, Griffiths H, Lee H S J, Medina E, Schäfer C, Stimmel K-H, Thonke B
Institut für Botanik, Technische Hochschule Darmstadt, D-6100 Darmstadt, FRG.
Department of Botany, University of Edinburgh, Edinburgh, EH9 3JH, UK.
New Phytol. 1989 Feb;111(2):293-307. doi: 10.1111/j.1469-8137.1989.tb00693.x.
Seasonal effects on leaf gas exchange and water relations were compared for Avicennia germinans, a true mangrove, and Conocarpus erectus, a mangrove associate, at coastal sites in northern Venezuela. On the Ciénega el Ostional at Chichiriviche, A. germinans was most abundant around lagoons on the seaward side of the vegetation-free alluvial sand plain. C. erectus was the dominant shrub in inland communities, but the two species co-occurred on vegetation islands at the landward edge of the alluvial plain. On the vegetation islands of the Ciénega el Ostional, gas exchange in A. germinans (a species with foliar salt glands) was less severely curtailed in the dry season compared with the rainy season than was gas exchange in C. erectus (a species lacking salt glands). Average rates of photosynthesis at near-saturating light intensities and total diurnal CO uptake were reduced in the dry season to 69 and 61%, respectively, of their values in the rainy for A. germinans, but to 48 and 30%, respectively, of their rainy-season values for C. erectus. Similarly, stomatal conductance and transpirational water loss were more reduced in the dry season for C. erectus than for A. germinans, with the result that C. erectus showed a 3.4-fold increase in water-use efficiency in the dry season compared with the rainy season. The importance of the soil environment in determining plant gas-exchange Patterns was evidenced by large seasonal shifts in dawn xylem tension for the two species (which increased from 1.34 MPa in the rainy season to 5.50 MPa in the dry season for A. germinans, and from 0.40 to 5.78 MPa for C. erectus). These values reflected changes in the soil environment caused by inundation of the upper soil layers by fresh water in the rainy season and a progressive increase in salt concentrations (to almost twice those in sea water) by evaporation from the soil in the dry season. Large changes in xylem tension were observed for both species during individual day-night cycles, reaching a maximum of 2.36 MPa for A. germinans. For C. erectus, the magnitude of these day-night changes was greatly reduced in the dry season, consistent with its very low transpiration rates at this time of year. Leaf-cell osmotic pressures also tended to be higher in A. germinans than C. erectus (attaining a maximum of 8.3 MPa for A. germinans in the dry season), and were related to the more seaward distribution of the true mangrove on the alluvial plain. Whereas leaves of A. germinans did not show any changes in succulence, leaf succulence in C. erectus increased with leafage and was slightly higher in the dry season than the rainy season. The more succulent leaves also had higher cell-sap osmotic pressures and NaCl concentrations. The most succulent leaves of C. erectus were observed for exposed shrubs growing on the shoreline. During the dry season, these shoreline plants showed high rates of gas exchange and low values for dawn xylem tension (0.89 MPa), indicating that they had access to relatively non-saline water from the shallow water table. On individual plants, exposed shoots had more succulent leaves and higher osmotic pressure and NaCl concentrations than sheltered leaves, demonstrating the importance of foliar absorption of salt borne in sea spray for the ionic relations of C. erectus. Thus, although the distribution of C. erectus is centred on brackish-water zones, this species can apparently extend from habitats with permanent access to a shallow water table through to areas where it is seasonally exposed to low soil water potentials and high salt concentrations in the substratum.
在委内瑞拉北部沿海地区,对真红树植物白骨壤和半红树植物直立红树的叶片气体交换和水分关系的季节效应进行了比较。在奇奇里维切的谢内加埃尔奥西翁阿尔,白骨壤在植被稀少的冲积沙质平原向海一侧的泻湖周围最为常见。直立红树是内陆群落中的优势灌木,但这两个物种在冲积平原向陆边缘的植被岛上共同出现。在谢内加埃尔奥西翁阿尔的植被岛上,与直立红树(一种缺乏盐腺的物种)相比,白骨壤(一种具有叶盐腺的物种)在旱季的气体交换受抑制程度比雨季小。在接近饱和光强下,白骨壤的光合平均速率和全天总二氧化碳吸收量在旱季分别降至雨季时的69%和61%,而直立红树则分别降至雨季时的48%和30%。同样,直立红树在旱季的气孔导度和蒸腾失水量比白骨壤减少得更多,结果直立红树在旱季的水分利用效率比雨季增加了3.4倍。两个物种黎明时木质部张力的季节性大幅变化证明了土壤环境在决定植物气体交换模式中的重要性(白骨壤从雨季的1.34兆帕增加到旱季的5.50兆帕,直立红树从0.40兆帕增加到5.78兆帕)。这些数值反映了雨季上层土壤被淡水淹没以及旱季土壤蒸发导致盐分浓度逐渐增加(几乎达到海水盐分浓度的两倍)所引起的土壤环境变化。在单个昼夜周期中,两个物种的木质部张力都有很大变化,白骨壤的最大值达到2.36兆帕。对于直立红树,这些昼夜变化的幅度在旱季大大减小,这与该物种在一年中的这个时候极低的蒸腾速率一致。白骨壤的叶细胞渗透压也往往高于直立红树(旱季白骨壤的最大值达到8.3兆帕),这与真红树在冲积平原上更靠海的分布有关。白骨壤的叶片肉质没有任何变化,而直立红树的叶片肉质随叶龄增加,且在旱季略高于雨季。肉质更多的叶片也有更高的细胞液渗透压和氯化钠浓度。在海岸线上生长的暴露灌木上观察到直立红树最肉质的叶片。在旱季,这些海岸线植物表现出高气体交换速率和低黎明时木质部张力值(0.89兆帕),表明它们能够获取来自浅水位的相对低盐度的水。在单株植物上,暴露的嫩枝比遮蔽的叶片有更肉质的叶子、更高的渗透压和氯化钠浓度,这证明了叶面对海雾中盐分的吸收对直立红树离子关系的重要性。因此,尽管直立红树的分布集中在咸淡水区域,但该物种显然可以从常年有浅水位的栖息地延伸到季节性面临低土壤水势和高基质盐分浓度的区域。