The spatial pattern of air seeding thresholds in mature sugar maple trees

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Abstract
Air seeding threshold (Pa) of xylem vessels from current year growth rings were measured along the vertical axis of mature sugar maple trees (Acer saccharum Marsh.), with sampling points in primary leaf veins, petioles, 1‐, 3‐, and 7‐year‐old branches, large branches, the trunk and roots. The air seeding threshold was taken as the pressure required to force nitrogen gas through intervessel pit membranes. Although all measurements were made on wood produced in the same year, Pa varied between different regions of A. saccharum, with distal organs such as leaves and petioles having lower Pa than basal regions. Mean (SE) Pa ranged from 1.0 (± 0.1) MPa in primary leaf veins to 4.8 (± 0.1) MPa in the main trunk. Roots exhibited a Pa of 2.8 (± 0.2) MPa, lower than all other regions of the tree except leaf veins and petioles. Mean xylem vessel diameter increased basipetally, with the widest vessels occurring in the trunk and roots. Within the shoot, wider vessels had greater air seeding thresholds, contrasting with trends previously reported. However, further experimentation revealed that differences in Pa between regions of the stem were driven by the presence of primary xylem conduits, rather than differences in vessel diameter. In 1‐year‐old branches, Pa was significantly lower in primary xylem vessels than in adjacent secondary xylem vessels. This explained the lower values of Pa measured in petioles and leaf veins, which possessed a greater ratio of primary xylem to secondary xylem than other regions. The difference in Pa between primary and secondary xylem was attributed to the greater area of primary cell wall (pit membrane) exposed in primary xylem conduits with helical or annular thickening.