membrane permeability

膜渗透性
  • 文章类型: Journal Article
    根据萨克斯的渗吸理论,人们对蒸腾流超出叶脉的过程的看法变化。通过Pfeffer的(共晶)内渗理论(直到1930年代后期几乎无可置疑地盛行),Strugger的荧光染料示踪剂和落射荧光显微镜实验。后一项工作说服了许多人回到外塑(墙)路径的观点,which,尽管早期和晚期的批评从未被反驳,仍然广泛持有。相同类型的示踪剂实验仍然经常发表,而没有考虑它们不能揭示水运动路径的证据。关于叶片复水动力学的实验尚未为这两种途径提供明确的证据。在蒸腾流中到达叶子的溶质的详细命运很少受到关注。考虑控制蒸腾叶片中流动和蒸发的物理原理,强调:(1)以水中速率在静脉间距离上的扩散将在几分钟内引起大量溶质运动,即使没有流动。(2)扩散也可以发生反对现在。(3)静脉中的体积通量由所检查的最大叶片的直径决定,该叶片包含高电导供应静脉,这些静脉被低电导分布的静脉所利用。(4)叶子的边缘和牙齿将是特别迅速蒸发的地方,它们通常有高电导静脉通向它们。(5)溪流中的溶质将倾向于在叶缘积累。根据最近的工作,观点认为,溪流的水通过细胞膜进入共生体,非常接近tracheary元素。此外,这种情况发生在膜的一小部分区域。溪流中的许多溶质留在外质中。这会在质外体中产生高溶质浓度的区域,并在灌注叶时在流中富集溶质。进入symplast的溶质不那么容易被跟踪。关于它们中的一些可能去哪里的建议可以从荧光探针获得,该荧光探针将特定细胞(清除细胞)鉴定为具有H-ATPase转运蛋白系统以从流中清除选定的溶质。提出了未发表的案例历史,说明了这些过程和原理的许多方面。这些是:(1)玉米叶脉,其中共生水路始于薄壁组织鞘;(2)卢平静脉,其中对称塑性路径从束鞘开始,溶质集中在盲端;(3)玉米叶的边缘,其中流动被大静脉增强(通向质外体),溶质通过蒸发沉积在质外体中;(4)杨树叶齿,接收强大的流量,上皮细胞是清除细胞;(5)含羞草叶边缘毛,它们的底部有清除细胞;(6)活性棘突,其表皮细胞是清除细胞;(7)松针输血组织,这是两个溶质富集的部位(在管胞中),和清除(在薄壁组织中);(8)对溶质的扩散系数进行了估算,该扩散系数与小麦叶片中的主要扩散途径成直角。第一个是第二个的1000倍,而是水中自由扩散的1/100。从审查的事实中得出了蒸腾流的行为和组织的五个一般主题。这些是:(1)通过物理力与解剖特征的相互作用,将溪流引导到分级强度的过程中,每个课程都对流的处理有不同的贡献。(2)水在尽可能靠近气管元素的精确位置进入共生体。(3)当水流穿过叶子时,其溶质浓度在可预测的位置富集了许多倍。(4)从对称体排除的溶质在特殊形成的壁路径中从这些高浓度源扩散,在精确的模式中,以可以测量的速率,与在水中的扩散相比是低的。(5)其他溶质渗透共生,通常在被组织成公认的结构特征的细胞群的表面上。内容摘要341I.蒸腾流的变化是什么?342II。审查343三。355预览四概述361确认365引用365。
    Changes of view on the course of the transpiration stream beyond the veins in leaves are followed from the imbibition theory of Sachs, through the (symplastic) endosmotic theory of Pfeffer (which prevailed almost unquestioned until the late 1930s), to Strugger\'s experiments with fluorescent dye tracers and the epifluorescence microscope. This latter work persuaded many to return to the apoplastic-(wall)-path viewpoint, which, despite early and late criticisms that were never rebutted, is still widely held. Tracer experiments of the same kind are still frequently published without consideration of the evidence that they do not reveal the paths of water movement. Experiments on rehydration kinetics of leaves have not produced unequivocal evidence for either path. The detailed destinies of the solutes that reach the leaf in the transpiration stream have received little attention. Consideration of physical principles governing flow and evaporation in a transpiring leaf emphasizes that: (1) Diffusion over interveinal distances at the rates in water will account for substantial solute movement in a few minutes, even in the absence of flow. (2) Diffusion can occur also against opposing now. (3) Volume fluxes in veins are determined by the diameter of the largest leaves examined contain high conductance supply veins which are tapped into by low-conductance distributing veins. (4) Edges and teeth of leaves will be places of especially rapid evaporation, and they often have high-conductance veins leading to them. (5) Solutes in the stream will tend to accumulate at leaf margins. On the basis of recent work, the view is maintained that the water of the stream enters the symplast through cell membranes very close to tracheary elements. Also, that this occurs locally over a small area of membrane. Many solutes in the stream are left outside in the apoplast. This produces regions of high solute concentration in the apoplast and an enrichment of solutes in the stream as it perfuses the leaf. Solutes that enter the symplast are not so easily tracked. Suggestions about where some of them may go can be gained from a fluorescent probe that identifies particular cells (scavenging cells) as having H+ -ATPase porter systems to scrub selected solutes from the stream. Unpublished case-histories are presented which illustrate many aspects of these processes and principles. These are: (1) Maize leaf veins, where the symplastic water path starts at the parenchyma sheath; (2) Lupin veins, where the symplastic path starts at the bundle sheath and where solutes are concentrated in blind terminations; (3) The edges of maize leaves where flow is enhanced by a large vein (open to the apoplast), and solutes are deposited in the apoplast by evaporation; (4) Poplar leaf teeth, which receive strong flows, and where the epithem cells are scavenging cells; (5) Mimosa leaf marginal hairs, which have scavenging cells at their base; (6) Active hydathodes, whose epithem cells are scavenging cells; (7) Pine needle transfusion tissue, which is a site of both solute enrichment (in the tracheids), and scavenging (in the parenchyma); (8) Estimates are made of diffusion coefficients of a solute both along and at right angles to the major diffusive pathway in wheat leaves. The first is 1000 times the second, but is 1/100 of free diffusion in water. Five general themes of the behaviour and organization of the transpiration stream are induced from the facts reviewed. These are: (1) The stream is channelled into courses of graded intensities by the interplay of the physical forces with the anatomical features, each course with a distinct contribution to the processing of the stream. (2) Water enters the symplast at precise locations as close as possible to the tracheary elements. (3) As the stream moves through the leaf its solute concentration is enriched many-fold at predictable sites. (4) Solutes excluded from the symplast diffuse from these sources of high concentration in specially formed wall paths, in precise patterns, at rates which can be measured, and which are low compared with diffusion in water. (5) Other solutes permeate the symplast, often over the surfaces of groups of cells which are organized into recognized structural features. CONTENTS Summary 341 I. What becomes of the transpiration stream ? 342 II. Review 343 III. Preview 355 IV. Overview 361 Acknowledgements 365 References 365.
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