Water permeability of Betula periderm

The water permeability of periderm membranes stripped from mature trees of Betula pendula Roth was investigated. The diffusion of water was studied using the system water/membrane/water, and transpiration was measured using the system water/membrane/water vapor. Betula periderm consists of successive periderm layers each made up of about 5 heavily suberized cell layers and a varying number of cell layers that are little suberized, if at all. It is shown that these layers act as resistances in series. The permeability coefficient of the diffusion of water (P d) can be predicted with 79% accuracy from the reciprocal of the membrane weight (x in mg cm(-2)) by means of the linear equation P d=14.69·10(-7) x-0.73·10(-7). For example, the P d of a periderm membrane having a weight of 10 mg cm(-2) (approx. 250 μm thick) is 7.4·10(-8) cm s(-1), which is comparable to the permeability of cuticles. This comparison shows that on a basis of unit thickness, Betula periderm is quite permeable to water as cuticles have the same resistance with a thickness of only 0.5 to 3 μm. It is argued that this comparatively high water permeability of birch periderm is due to the fact that middle lamellae and the primary walls of periderm cells are not at all, or only incompletely suberized and, therefore, form a hydrophilic network within which the water can flow. This conclusion is based on the following observations: (1) Middle lamellae and primary walls stain strongly with toluidine blue, which shows them to be polar. (2) If silver ions are added as tracer for the flow of water, they are found only in the middle lamellae, primary walls, and in plasmodesmata, while no silver can be detected in the suberized walls. (3) Permeability coefficients of transpiration strongly depend on water activity. This shows conclusively that water flows across Betula periderm via a polar pathway. It is further argued that liquid continuity is likely to be maintained under all physiological conditions in the network formed by middle lamellae and primary walls. On the other hand, the lumina of periderm cells, intercellular air spaces in the lenticels, and even the pores in the suberized walls (remainders of plasmodesmata) will drain at a humidity of 95% and below. Due to the presence of intercellulars the permeability coefficient of lenticels is much greater than that of the periderm. A substantial amount of the total water, therefore, flows as vapor through lenticels even though they cover only 3% of the surface.