Seasonal dynamics of cell-to-cell transport in angiosperm wood.

This study describes the seasonal changes in cell-to-cell transport in three selected angiosperm tree species, Acer pseudoplatanus (maple), Fraxinus excelsior (ash), and Populus tremula × tremuloides (poplar), with an emphasis on the living wood component, xylem parenchyma cells (XPCs). We performed anatomical studies, dye loading through the vascular system, measurements of non-structural carbohydrate content, immunocytochemistry, inhibitory assays and quantitative real-time PCR to analyse the transport mechanisms and seasonal variations in wood. The abundance of membrane dye in wood varied seasonally along with seasonally changing tree phenology, cambial activity, and non-structural carbohydrate content. Moreover, dyes internalized in vessel-associated cells and 'trapped' in the endomembrane system are transported farther between other XPCs via plasmodesmata. Finally, various transport mechanisms based on clathrin-mediated and clathrin-independent endocytosis, and membrane transporters, operate in wood, and their involvement is species and/or season dependent. Our study highlights the importance of XPCs in seasonally changing cell-to-cell transport in both ring-porous (ash) and diffuse-porous (maple, poplar) tree species, and demonstrates the involvement of both endocytosis and plasmodesmata in intercellular communication in angiosperm wood.

Xylem Parenchyma-Role and Relevance in Wood Functioning in Trees.

Woody plants are characterised by a highly complex vascular system, wherein the secondary xylem (wood) is responsible for the axial transport of water and various substances. Previous studies have focused on the dead conductive elements in this heterogeneous tissue. However, the living xylem parenchyma cells, which constitute a significant functional fraction of the wood tissue, have been strongly neglected in studies on tree biology. Although there has recently been increased research interest in xylem parenchyma cells, the mechanisms that operate in these cells are poorly understood. Therefore, the present review focuses on selected roles of xylem parenchyma and its relevance in wood functioning. In addition, to elucidate the importance of xylem parenchyma, we have compiled evidence supporting the hypothesis on the significance of parenchyma cells in tree functioning and identified the key unaddressed questions in the field.

Endocytosis acts as transport pathway in wood.

In trees, dead and living cells of secondary xylem (wood) function collectively, rendering cell-to-cell communication challenging. Water and solutes are transported over long distances from the roots to the above-ground organs via vessels, the main component of wood, and then radially over short distances to the neighboring cells. This enables proper functioning of trees and integrates whole-plant activity. In this study, tracer loading, immunolocalization experiments and inhibitor assays were used to decipher the mechanisms enabling transport in wood of Acer pseudoplatanus (maple), Fraxinus excelsior (ash) and Populus tremula × tremuloides (poplar) trees. We show that tracer uptake from dead water-conducting vessels, elements of the apoplasm, to living vessel-associated cells (VACs) of the xylem parenchyma of the symplasm system proceeds via the endocytic pathway, including clathrin-mediated and clathrin-independent processes. These findings enhance our understanding of the transport pathways in complex wood tissue, providing experimental evidence of the involvement of VACs and endocytosis in radial uptake from vessels.