In contrast to common belief, recent studies have confirmed that intrusive growth of fusiform cambial initials has a significant role in the rearrangement of the initials, but does not contribute to the cambial circumference increment. We observed a rapid rearrangement of cambial initials on a long series of transverse sections of the vascular cambium and the wood of a 50-year-old pine (Pinus sylvestris L.) tree. A comparison of cell arrangement in consecutive sections, as well as a critical analysis of tangential reconstructions, has confirmed that changes in cell locations in a group of cells on the tangential surface caused no change in the total tangential width of the whole group. Models illustrating changes in locations of the initials have been proposed, assuming that intrusive growth, which makes the growing initials intrude between the neighbouring initials and their immediate derivatives, is localized on the longitudinal edges of cells. We infer that intrusive growth of the cambial initials in P. sylvestris is not involved in the cambial circumference increment, but plays a significant role in the rearrangement of the initials, probably allowing for a relaxation of shearing strains generated during radial growth. The relationship of intrusive growth with the elimination of initials has been discussed with reference to the frequency of anticlinal divisions. It has been proposed that the occurrence of anticlinal divisions in excess over the actual requirement for increase in the cambial circumference could be due to internal shearing strains.
It is well documented that apical elongation of fusiform cambial initials through extension of their longitudinal edges, and their intrusion between tangential walls of the neighbouring initials and their closest derivatives cause rearrangement of fusiform cells, without increasing the cambial circumference. However, the concurrent rearrangement of rays is not fully understood. This study deals with Pinus sylvestris L., Tilia cordata Mill. and Hippophaë rhamnoides L., possessing a nonstoreyed, storeyed and double-storeyed type of cambium, respectively, and shows that the mechanism for rearrangement of ray initials is similar to the one proposed for fusiform initials, and includes multiplication of ray initials by anticlinal divisions, intrusive growth of ray initials, elimination of ray initials caused by intrusive growth of neighbouring fusiform initials, and transformation of ray initials into fusiform initials. Intrusive growth of a ray initial does not necessarily lead to the formation of a new fusiform initial, as it is dependent on the extent of the intrusive growth taken place. The extent of rearrangement of cambial cells is determined by the intensity of events occurring among the fusiform as well as ray initials. Intrusive growth of these initials does not influence the size of the cambial circumference.
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 ofRobinia 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 intrusiveversussymplastic growths in the axial, circumferential and radial directions.
It is believed that differentiating vessel elements increase their diameter by growing intrusively in the circumferential direction and symplastically in the radial direction in relation to the stem axis. On the basis of a detailed analysis of the cell arrangement observed in a series of semithin anatomical sections of cambial zone and the developing and mature secondary xylem of Terminalia ivorensis , Wisteria floribunda , and Millettia laurentii , we revealed a novel correlation of growing vessel elements with surrounding tissues. Rays seem to prevent the growing vessel elements from protruding laterally between the cells of adjacent rays. The growing vessel elements break the continuity of several neighbouring radial files of fusiform cell derivatives but not of ray cell derivatives. If a contiguous ray becomes an obstacle for the growth of vessel elements on any one side, the growing elements often start to grow in the opposite direction, consequently causing a deviation in the alignment of the vessel elements concerned. This mechanism explains why vessel elements may deviate from the array of their precursors, the fusiform cambial initials. Our models on the intrusive symplastic growth of vessel element mother cells have revealed that intrusive growth does not occur between radial walls of neighbouring cells.
A b s t r a c tThe formation of the storeyed pattern of cambium in the first two years of cambial activity and the structure of mature cambium was examined in Laburnum anagyroides, a decorative shrub growing in temperate climate. A distinct storeyed pattern was observed in two-year-old stems, despite the presence of tall rays. The heterogeneous nature of storeys was revealed by the analysis of i) the number of fusiform initials forming storeys or groups of packets; and ii) the frequency of anticlinal divisions in the examined years of cambial activity and the calculation of the relative increase of the cambial circumference. The results are discussed on the basis of the recent hypothesis of intrusive growth of fusiform initial, occurring between the tangential walls of its neighbouring initial and its closest derivative.
scite is a Brooklyn-based organization that helps researchers better discover and understand research articles through Smart Citations–citations that display the context of the citation and describe whether the article provides supporting or contrasting evidence. scite is used by students and researchers from around the world and is funded in part by the National Science Foundation and the National Institute on Drug Abuse of the National Institutes of Health.
customersupport@researchsolutions.com
10624 S. Eastern Ave., Ste. A-614
Henderson, NV 89052, USA
Copyright © 2024 scite LLC. All rights reserved.
Made with 💙 for researchers
Part of the Research Solutions Family.