The timing of cambial reactivation plays an important role in determination of the amount and quality of wood and the environmental adaptivity of trees. Environmental factors, such as temperature, influence the growth and development of trees. Temperatures from late winter to early spring affect the physiological processes that are involved in the initiation of cambial cell division and xylem differentiation in trees. Cumulative elevated temperatures from late winter to early spring result in earlier initiation of cambial reactivation and xylem differentiation in tree stems and an extended growth period. However, earlier cambial reactivation increases the risk for frost damage because the cold tolerance of cambium decreases after cambial reactivation. The present review focuses on temperature regulation on the timing of cambial reactivation and xylem differentiation in trees, and also highlights recent advances in our understanding of seasonal changes in the cold stability of microtubules in trees. The review also summarizes the present understanding of the relationships between the timing of cambial reactivation, the start of xylem differentiation and changes in levels of storage materials in trees, as well as an attempt to identify the source of energy for cell division and differentiation. A better understanding of the mechanisms that regulate wood formation in trees and the influence of environmental conditions on such mechanisms should help in efforts to improve and enhance the exploitation of wood for commercial applications and to prepare for climatic change.

The results suggest that, in deciduous diffuse-porous hardwood poplar growing in a temperate zone, the temperature in the stem is a limiting factor for reactivation of phloem and cambium. An increase in temperature might induce the conversion of storage starch to sucrose for the activation of cambial cell division and secondary xylem. Localized heating in poplar stems provides a useful experimental system for studies of cambial biology.

Differences in the timing of cambial reactivation and the initiation of xylem differentiation in response to the sum of daily maximum temperatures were studied in two Cryptomeria japonica trees with cambium of different ages under natural and locally heated conditions. In addition, we observed the effects of low temperature on cambial activity. The timing of cambial reactivation and of the initiation of xylem differentiation differed between 55-and 80-year-old cambium under natural conditions. In the 55-year-old cambium, cambial reactivation occurred when the cambial reactivation index (CRI), calculated on the basis of daily maximum temperatures in excess of 10°C, was 94 and 97°C in 2007 and 2008, respectively. In 80-year-old cambium, cambial reactivation occurred when the CRI, calculated on the basis of daily maximum temperatures in excess of 11°C, was 69 and 71°C in 2007 and 2008, respectively. After cambial reactivation in 2007, normal cell division was evident in the cambium even though the minimum temperature had fallen between -2 and -3°C. Under natural conditions, xylem differentiation started 38-44 days after cambial reactivation. In heated stems, the time between cambial reactivation and the initiation of xylem differentiation ranged from 14 to 16 days, a much shorter time than under natural conditions, indicating that continuous exposure to an elevated temperature had induced earlier xylem differentiation. Our observations indicate that the sensitivity to reactivation inducing stimuli of the cambium depends on both the stage of dormancy and tree age of the cambium.

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We monitored the distribution of death of secondary xylem cells in a conifer, Abies sachalinensis. The cell death of tracheids, which are tracheary elements, occurred successively and was related to the distance from cambium. Thus, it resembled programmed cell death. By contrast, the death of long-lived ray parenchyma cells had the following features: (1) ray parenchyma cells remained alive for several years or more; (2) in many cases, no successive cell death occurred even within a given radial cell line of a ray; and (3) the timing of cell death differed among upper and lower radial cell lines and other lines of cells within a ray. These results indicate that the death of long-lived ray parenchyma cells involves a different process from the death of tracheids. The initiation of secondary wall formation and the lignification of ray parenchyma cells in the current year's annual ring were delayed in the upper and lower radial cell lines of a ray. In addition, the density of distribution and orientation of cortical microtubules in such cells were different from those in cells in other radial lines. Ray parenchyma cells in the previous year's annual ring within the upper and lower radial cell lines of a ray contained many starch grains. Our results indicate that positional information is an important factor in the control of the pattern of differentiation and, thus, of the functions of ray parenchyma cells that are derived from the same cambial ray cells.

Differences in patterns of cell death between ray parenchyma cells and ray tracheids in the conifers Pinus densiflora and Pinus rigida were clarified. Differentiation and cell death of ray tracheids occurred successively and both were related to the distance from the cambium. In this respect, they resembled those of longitudinal tracheids. Thus, the cell death of short-lived ray tracheids could be characterized as time-dependent programmed cell death. In contrast, ray parenchyma cells survived for several years or more, and no successive cell death occurred, even within a single radial line of cells in a ray. Thus, the features of death of the ray parenchyma cells were different from those of ray tracheids. Cell death occurred early in ray parenchyma cells that were in contact with ray tracheids. The initiation of secondary wall thickening occurred earlier in ray parenchyma cells that were in contact with ray tracheids in Pinus densiflora than in others. In addition, localized thickening of secondary walls occurred only in ray parenchyma cells that were in contact with ray tracheids in Pinus rigida. Moreover, no polyphenols were evident in such cells in either species. Therefore, ray parenchyma cells that were in contact with ray tracheids appeared not to play a role in the formation of heartwood extractives. Our observations indicate that short-lived ray tracheids might affect the pattern of differentiation and, thus, the functions of neighboring longlived ray parenchyma cells in conifers.

Several studies have demonstrated that localized heating of tree stems induces localized cambial reactivation. We analyzed by light microscopy the effects of early spring increases in ambient temperature in 2005 and 2007 on the timing of cambial reactivation and xylem differentiation in stems of two trees of a cloned deciduous hardwood hybrid poplar (Populus sieboldii Miquel. x P. grandidentata Michx.) growing under natural conditions. Meteorological data at the study site showed that temperatures in late winter and early spring differed markedly between 2005 and 2007, with trends toward higher temperatures starting around April 3 in 2005 and around March 20 in 2007. Cambial reactivation occurred about 17 days earlier in 2007 than in 2005. The cumulative daily maximum temperature in excess of 15 degrees C (maximum daily temperatures minus 15 degrees C) in late winter and early spring before cambial reactivation was defined as the cambial reactivation index (CRI(15)). Cambial reactivation, which began when the minimum temperature rose above 0 degrees C, occurred when the CRI(15) was 93 and 96 degrees C in 2005 and 2007, respectively. The differentiation of secondary xylem started earlier in 2007 than in 2005. On May 27, we found a wider current-year band of xylem and a higher frequency of small-diameter vessel elements in 2007 than in 2005. We propose that the timing of cambial reactivation is controlled by air temperature and that earlier cambial reactivation induces earlier differentiation of xylem in hybrid poplar under natural conditions. Our results indicate that the CRI might be a useful indicator of the timing of cambial reactivation.

The anatomical characteristics and density of wood were examined in 23-year-old Acacia mangium trees that had been planted in Yogyakarta, Indonesia. The seeds had been collected from trees of five different provenances. The distance from the pith of the boundary between juvenile and mature wood was also examined to clarify the maturity of the wood. Lengths of wood fibers near the pith and the distance from the pith of the boundary between juvenile and mature wood differed significantly among provenances. By contrast, other anatomical characteristics of the wood such as fiber wall area, fiber wall thickness, fiber diameter, vessel lumen area, vessel diameter, vessel frequency and wood density did not differ significantly among provenances. Wood density was strongly correlated with the area of fiber walls. Our observations suggest that Sidei and Daintree might be more appropriate provenances among those examined for the Acacia mangium treebreeding programs in Indonesia that are aimed at improving wood quality, because these provenances are associated with longer initial wood fibers and narrower juvenile areas than the other provenances studied.