Vesala Timo, Hölttä Teemu, Perämäki Martti, Nikinmaa Eero
Department of Physical Sciences, PO Box 64, FIN-00014, University of Helsinki, Finland.
Ann Bot. 2003 Mar;91(4):419-28. doi: 10.1093/aob/mcg022.
When they are hydraulically isolated, embolized xylem vessels can be refilled, while adjacent vessels remain under tension. This implies that the pressure of water in the refilling vessel must be equal to the bubble gas pressure, which sets physical constraints for recovery. A model of water exudation into the cylindrical vessel and of bubble dissolution based on the assumption of hydraulic isolation is developed. Refilling is made possible by the turgor of the living cells adjacent to the refilling vessel, and by a reflection coefficient below 1 for the exchange of solutes across the interface between the vessel and the adjacent cells. No active transport of solutes is assumed. Living cells are also capable of importing water from the water-conducting vessels. The most limiting factors were found to be the osmotic potential of living cells and the ratio of the volume of the adjacent living cells to that of the embolized vessel. With values for these of 1.5 MPa and 1, respectively, refilling times were in the order of hours for a broad range of possible values of water conductivity coefficients and effective diffusion distances for dissolved air, when the xylem water tension was below 0.6 MPa and constant. Inclusion of the daily pattern for xylem tension improved the simulations. The simulated gas pressure within the refilling vessel was in accordance with recent experimental results. The study shows that the refilling process is physically possible under hydraulic isolation, while water in surrounding vessels is under negative pressure. However, the osmotic potentials in the refilling vessel tend to be large (in the order of 1 MPa). Only if the xylem water tension is, at most, twice atmospheric pressure, the reflection coefficient remains close to 1 (0.95) and the ratio of the volume of the adjacent living cells to that of the embolized vessel is about 2, does the osmotic potential stay below 0.4 MPa.
当木质部导管被水力隔离时,栓塞的木质部导管可以重新充满水,而相邻的导管仍处于张力状态。这意味着重新充水导管中的水压必须等于气泡气体压力,这为恢复设定了物理限制。基于水力隔离的假设,建立了水渗入圆柱形导管和气泡溶解的模型。重新充水是由与重新充水导管相邻的活细胞的膨压以及溶质在导管与相邻细胞界面处交换的反射系数低于1来实现的。不假定溶质有主动运输。活细胞也能够从导水导管中吸收水分。发现最限制因素是活细胞的渗透势以及相邻活细胞体积与栓塞导管体积的比率。当木质部水张力低于0.6MPa且恒定时,对于广泛的可能的导水系数值和溶解空气的有效扩散距离值,这些值分别为1.5MPa和1时,重新充水时间约为数小时。纳入木质部张力的日变化模式改善了模拟结果。重新充水导管内模拟的气体压力与最近的实验结果一致。研究表明,在水力隔离条件下,当周围导管中的水处于负压时,重新充水过程在物理上是可能的。然而,重新充水导管中的渗透势往往很大(约为1MPa)。只有当木质部水张力最多为大气压的两倍、反射系数保持接近1(0.95)且相邻活细胞体积与栓塞导管体积的比率约为2时,渗透势才会保持在0.4MPa以下。