Higinbotham N, Etherton B, Foster R J
Department of Botany, Washington State University, Pullman, Washington 99163.
Plant Physiol. 1967 Jan;42(1):37-46. doi: 10.1104/pp.42.1.37.
The relationships of concentration gradients to electropotential gradients resulting from passive diffusion processes, after equilibration, are described by the Nernst equation. The primary criterion for the hypothesis that any given ion is actively transported is to establish that it is not diffusing passively. A test was made of how closely the Nernst equation describes the electrochemical equilibrium in seedling tissues. Segments of roots and epicotyl internodes of pea (Pisum sativum var. Alaska) and of roots and coleoptiles of oat (Avena sativa var. Victory) seedlings were immersed and shaken in defined nutrient solutions containing eight major nutrients (K(+), Na(+), Ca(2+), Mg(2+), Cl(-), NO(3) (-), H(2)PO(4) (-) and SO(4) (2-)) at 1-fold and 10-fold concentrations. The tissue content of each ion was assayed at 0, 8, 24, and 48 hours. A near-equilibrium condition was approached by roots for most ions; however, the segments of shoot tissue generally continued to show a net accumulation of some ions, mainly K(+) and NO(3) (-). Only K(+) approached a reasonable fit to the Nernst equation and this was true for the 1-fold concentration but not the 10-fold. The data suggest that for Na(+), Mg(2+), and Ca(2+) the electrochemical gradient is from the external solution to the cell interior; thus passive diffusion should be in an inward direction. Consequently, some mechanism must exist in plant tissue either to exclude these cations or to extrude them (e.g., by an active efflux pump). For each of the anions the electrochemical gradient is from the tissue to the solution; thus an active influx pump for anions seems required. Root segments approach ionic equilibrium with the solution concentration in which the seedlings were grown. Segments of shoot tissue, however, are far removed from such equilibration. Thus in the intact seedling the extracellular (wall space) fluid must be very different from that of the nutrient solution bathing the segments; it would appear that the root is the site of regulation of ion uptake in the intact plant although other correlative mechanisms may be involved.
平衡后,能斯特方程描述了被动扩散过程中浓度梯度与电势梯度之间的关系。任何给定离子被主动转运这一假说的主要标准是确定它不是在被动扩散。对能斯特方程在多大程度上描述幼苗组织中的电化学平衡进行了一项测试。将豌豆(阿拉斯加品种)的根段和上胚轴节间以及燕麦(胜利品种)幼苗的根段和胚芽鞘浸入含有八种主要养分(K⁺、Na⁺、Ca²⁺、Mg²⁺、Cl⁻、NO₃⁻、H₂PO₄⁻和SO₄²⁻)的特定营养液中,以1倍和10倍浓度振荡。在0、8、24和48小时测定每种离子的组织含量。根对大多数离子接近平衡状态;然而,茎组织切段通常继续显示一些离子的净积累,主要是K⁺和NO₃⁻。只有K⁺接近能斯特方程的合理拟合,这在1倍浓度时成立,但在10倍浓度时不成立。数据表明,对于Na⁺、Mg²⁺和Ca²⁺,电化学梯度是从外部溶液到细胞内部;因此被动扩散应该是向内的方向。因此,植物组织中必须存在某种机制来排除这些阳离子或将它们排出(例如,通过主动外排泵)。对于每种阴离子,电化学梯度是从组织到溶液;因此似乎需要一种阴离子主动内流泵。根段与幼苗生长的溶液浓度接近离子平衡。然而,茎组织切段与这种平衡相差甚远。因此,在完整的幼苗中,细胞外(细胞壁空间)流体一定与浸泡切段的营养液有很大不同;虽然可能涉及其他相关机制,但根似乎是完整植物中离子吸收的调节部位。