Geometric analysis of intrusive growth of wood fibres in Robinia pseudoacacia

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All cell types of the secondary xylem arise from the meristematic cells (initials) of the vascular cambium and grow under mechanical constraints emerging from the circular-symmetrical geometry that characterises many tree trunks. The course of intrusive growth of cambial initials has been elucidated, but is yet to be described in the case of xylem fibres. This study explains the geometry of intrusive growth of the secondary xylem fibres in the trunk of Robinia pseudoacacia. Long series of serial semi-thin sections of the vascular cambium and the differentiating secondary xylem were analysed. Since fibres grow in close vicinity to expanding cells of the derivatives of the vascular cambium, we assumed that they have similar growth conditions. Dealing with the cylindrical tissue of the vascular cambium in a previous study, we used a circularly symmetrical equation for describing the growth mechanism of cambial initials. Like the cambial initials, some of the cambial derivatives differentiating into the various cell types composing the secondary xylem also exhibit intrusive growth between the tangential walls of adjacent cells. As seen in cross sections of the cambium, intrusively growing initials form slanted walls by a gradual transformation of tangential (periclinal) walls into radial (anticlinal) walls. Similarly, the intrusive growth of xylem fibres manifests initially as slants, which are formed due to axial growth of the growing cell tips along the tangential walls of adjacent cells. During this process, the tangential walls of adjacent cells are partly separated and dislocated from the tangential plane. The final shape of xylem fibres, or that of vessel elements and axial parenchyma cells, depends upon the ratio of their intrusive versus symplastic growths in the axial, circumferential and radial directions.



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  • Early stages of apical intrusive growth of fibre mother cells in Robinia pseudoacacia. – A & B: Selected successive tangential sections of cambial initial cells (A) and fibre mother cells (B). Radial distance between A and B sections is 30 μm. The change in the location of cambial cell tips vs fibre tips growing intrusively is indicated with black and white arrows respectively. – C & D: Views of intrusive growth of the tip of a fibre mother cell (arrowheads) obtained by optical sectioning of a 3.5 μm thick tangential area. – E: Transverse section of vascular cambium and xylem derivatives; the horizontal dashed lines (c, d) as well as the white rectangle define the possible location of a similar situation visible on C and D images. Note the quadrangle shape of a growing fibre tip. – F–H: Description as in C–E; note the triangular shape of a growing fibre tip in H. The horizontal dashed lines (f, g) and white rectangle define the possible location of similar situations as visible on F and G images. The distance between the dashed lines c and d in the section E and between the dashed lines f and g in section H corresponds to the section thickness (3.5 μm). — Scale bar = 50 μm.

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  • Microspaces (black arrowheads) and fibre tips (grey arrowheads) marked in a cross section of developing secondary xylem. The probable initials are marked with black asterisks. Periclinal divisions which probably occurred after intrusive growth of the adjacent fibre tips are marked with white arrowheads. One example, marked with the white arrow, indicates a new periclinal wall close to the fibre tip corner. Likewise, the black arrow points to the new periclinal wall in the growing fibre tip.

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  • Reconstructions of the arrangement of cells in tangential and radial planes, made on the basis of series of transverse sections. The examined radial files of cells in growing wood zone (GW) are distinguished with different shades. a) Transverse sections of GW (A-H) chosen from a long series. Lines 1 and 2 indicate the planes of reconstructions of radial sections, whereas lines I and II indicate the planes of reconstructions of tangential sections. — b) Reconstructions of tangential (I, II) and radial (1, 2) sections made in the planes indicated in transverse sections (a). I+II: Superimposed reconstructions of tangential sections represent changes in the cells’ tangential surface between these two stages of the maturing xylem cells. Horizontal levels (A-H) at the tangential and radial views correspond to transverse sections (A-H) in figure a. The continuous lines correspond to the location of cell walls, whereas the dotted lines indicate periclinal divisions.

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  • Cross sections of vascular cambium (B) and mature xylem (A, C) of Robinia pseudoacacia. – A: On the left side of the micrograph, xylem fragment with regularly arranged fibre bodies is visible. On the right, fibre bodies are randomly scattered. A cluster of vessels filled with tyloses is located in the central part. – B: The frame encompasses ten cambial cells arranged in two radial files. – C: Fibre bodies are arranged in regular rows. The frame surrounds ten fibre cells of two neighbouring radial files. The periclinal and anticlinal walls of fibre bodies are not fully separated.

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  • Cross section of vascular cambium and the differentiating secondary xylem of Robinia pseudoacacia in the vicinity of ray cells. The dashed rectangular line frame displays the zone of growing wood, where fibre bodies still retain cell-to-cell contacts in early stages of the fibre-tip intrusive growth.

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  • Transverse section of vascular cambium and secondary xylem of Robinia pseudoacacia: a fragment of the differentiating vessel element is visible in the bottom right corner. In the zone between two dashed lines, the growing fibre tips are hardly visible near the vessel element on the right side of the triseriate ray. Numerous growing fibre tips are visible in the same zone on the left side of the ray.

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  • Macerated mature fibres of the examined secondary xylem of Robinia pseudoacacia. Despite the varied length of the fibres, the length of fibre bodies remains constant. The distance between the horizontal dotted lines indicates the fibre body length, which corresponds to the height of storeys of the cambial cells.

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