The anatomy and grain pattern in forks of hazel (Corylus avellana L.) and other tree species

摘要

This paper provides a new anatomical model for how branches are joined together at forks in trees, using evidence from MicroXCT scanning, ESEM output and visual observation. Wood grain arrangements at the forks and other junctions within a tree must be arranged to mechanically join together the two or more branches, yet not adversely restrict sap flow. The grain orientation at junctions therefore represents a trade-off in xylem performance between the functions of efficient sap conductance and the provision of adequate load-bearing capacity. Initial observations of wood grain orientation were made on the surfaces of several dozen debarked and fractured bifurcations of a wide range of tree species, both by eye and using a scanning electron microscope. Subsequently, small volumes of wood were sampled from two locations within the junctions of hazel, at the junction apex and on the outer section of join. Wood was imaged in 3D using high-resolution X-ray tomography, and the scanned volumes were analysed for their wood grain patterns. It was found that the wood at the junctions of hazel forks contained only 37 % of the number of vessels contained in wood within the adjacent stem. The vessel elements formed at the junctions were only 32.5 % the length of those in the stem, had a mean diameter only 50.5 % of the stem vessels and consequently only 26.3 % of their lumen volume. The passage of the vessels through the fork wood deviated from a straight line (Euclidean) distance by more than 14 times more than the stem wood vessels did. The interweaving of vessels in the fork wood was over 22 times greater than in the stem wood. A survey of rays showed them to be 58 % more abundant in fork wood but only 62 % of the height of rays in the stem wood. These results suggest that where two branches of similar diameter join to form a tree fork, an interlocking wood grain pattern is formed at the apex of the bifurcation, which provides higher tensile strength to the junction. Breaking of the hazel fork requires wood fibres to be stretched axially and broken across, which requires greater stress than breaking normal wood radially or tangentially