INTERVESSEL PIT MEMBRANE THICKNESS AS A KEY DETERMINANT OF EMBOLISM RESISTANCE IN ANGIOSPERM XYLEM

in IAWA Journal
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Pit membranes in bordered pits between neighbouring vessels play a major role in the entry of air-water menisci from an embolised vessel into a water-filled vessel (i.e., air-seeding). Here, we investigate intervessel pit membrane thickness (TPM) and embolism resistance (P50, i.e., the water potential corresponding to 50% loss of hydraulic conductivity) across a broad range of woody angiosperm species. Data on TPM and double intervessel wall thickness (TVW) were compiled based on electron and light microscopy. Fresh material that was directly fixated for transmission electron microscopy (TEM) was investigated for 71 species, while non-fresh samples were frozen, stored in alcohol, or air dried prior to TEM preparation for an additional 60 species. TPM and P50 were based on novel observations and literature. A strong correlation between TPM and P50 was found for measurements based on freshly fixated material (r = 0.78, P >0.01, n = 37), and between TPM and TVW (r = 0.79, P >0.01, n = 59), while a slightly weaker relationship occurred between TVW and P50 (r = 0.40, P >0.01, n = 34). However, non-fresh samples showed no correlation between TPM and P50, and between TPM and TVW. Intervessel pit membranes in non-fresh samples were c.28% thinner and more electron dense than fresh samples. Our findings demonstrate that TPM measured on freshly fixated material provides one of the strongest wood anatomical correlates of droughtinduced embolism resistance in angiosperms. Assuming that cellulose microfibrils show an equal spatial density, TPM is suggested to affect the length and the shape of intervessel pit membrane pores, but not the actual pore size. Moreover, the shrinking effect observed for TPM after dehydration and frost is associated with an increase in microfibril density and porosity, which may provide a functional explanation for embolism fatigue.

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