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

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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.

Abstract

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.

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Author contribution statement

Duncan Slater: initiator of this investigation into branch attachment, PhD student of Professor Ennos, first author of this paper. Work for this paper involved collection of samples, segmentation of CT scanning data, statistical analysis and the writing of this paper. Dr. Robert Bradley: technical assistant to CT scanning, creator of algorithms that provided parameters from the segmented CT scanning data, reviewer of the original paper and author of part of the methods section of the paper (technical aspects of the CT scanning and algorithms used to analyse the raw data). Prof. Phil Withers: CT facility manager, supervisor of Dr. Robert Bradley and reviewer of the initial draft of this paper. Prof. Roland Ennos: Supervisor to Mr. Duncan Slater, editor and reviewer of this manuscript.

Acknowledgments

We would like to thank the following contributors: M. Carswell for the supply of hazel forks to assist with this study; D. Elwell and M. Heys for producing illustrative figures; Dr. T. Lowe for his advice on segmentation of our CT scans; A. Hirons for reviewing our findings; and BSc (Hons) graduates J. Barnes and C. Harbinson for their thoughts and ideas at the beginning.

Conflict of interest

The authors declare that they have no conflict of interest. P. Withers would like to acknowledge funding from the Engineering and Physical Sciences Research Council for the Manchester X-ray Imaging Facility under EP/F007906/1 and EP/F028431/1.

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Authors and Affiliations

  1. Myerscough College, St. Michael’s Road, Bilsborrow, Preston, PR3 0RY, England, UK

    Duncan Slater

  2. Manchester X-ray Imaging Facility, Materials Science Centre, The University of Manchester, Grosvenor Street, Manchester, M1 7HS, England, UK

    Robert S. Bradley & Philip J. Withers

  3. School of Biological, Biomedical and Environmental Sciences, University of Hull, Cottingham Road, Hull, HU6 7RX, England, UK

    A. Roland Ennos

Authors
  1. Duncan Slater
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  2. Robert S. Bradley
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  3. Philip J. Withers
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  4. A. Roland Ennos
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Corresponding author

Correspondence to Duncan Slater.

Additional information

Communicated by M. Shane.

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Slater, D., Bradley, R.S., Withers, P.J. et al. The anatomy and grain pattern in forks of hazel (Corylus avellana L.) and other tree species. Trees 28, 1437–1448 (2014). https://doi.org/10.1007/s00468-014-1047-5

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  • DOI: https://doi.org/10.1007/s00468-014-1047-5

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