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Key message A better understanding of the influence of environmental conditions on wood formation should help to improve the radial growth of trees and to prepare for climate change. The cambial activity of trees is associated with seasonal cycles of activity and dormancy in temperate zones. The timing of cambial reactivation in early spring and dormancy in autumn plays an important role in determination of the cambial growth and the environmental adaptivity of temperate trees. This review focuses on the temperature regulation of the timing of cambial reactivation and xylem differentiation and highlights recent advances of bud growth in relation to cambial activity of temperate trees. In addition, we discuss relationships between the timing of cambial reactivation, start of xylem differentiation and changes in levels of storage materials to identify the source of the energy required for cell division and differentiation. We also present a summary of current understanding of the effects of rapid increases and decreases in temperature on cambial activity, by localized heating and cooling, respectively. Increases in temperature from late winter to early spring influence the physiological processes that are involved in the initiation of cambial reactivation and xylem differentiation both in localized heated stems and under natural conditions. Localized cooling has a direct effect on cell expansion, the thickening of walls of differentiating tracheids, and the rate of division of cambial cells. A rapid decrease in temperature of the stem might be the critical factor in the control of latewood formation and the cessation of cambial activity. Therefore, temperature is the main driver of cambial activity in temperate trees and trees are able to feel changes in temperature through the stem. The climate change might affect wood formation in trees.

The differentiation of tracheary elements in vitro provides a useful system for detailed analysis of xylem cell differentiation. To examine the mechanism of formation of cell wall structures, new differentiation systems are required that allows us to induce highly organized structures, such as perforations. In this study, we developed such a system in which we were able to induce formation of tracheary elements with perforations, using calli of a hardwood, Aesculus turbinata. Young leaves of A. turbinata were placed on modified MS medium that contained 5 μM 2,4-dichlorophenoxyacetic acid (2,4-D) and 5 μM benzyladenine (BA). Tracheary elements were induced in calli derived from young leaves of A. turbinata. Some tracheary elements formed broad areas of secondary wall with typical features of secondary xylem. Other tracheary elements formed spiral thickenings, which are typical features of vessel elements in secondary xylem of A. turbinata. Approximately 10% of tracheary elements formed large pores that resembled perforations of vessel elements and various types of the perforation plate were observed. Addition of NAA and brassinolide to the induction medium enhanced the differentiation of tracheary elements in calli of A. turbinata. Newly induced tracheary elements also formed typical features of secondary xylem such as perforations of the vessel elements. Our model system might be useful in efforts to understand the mechanisms of formation of highly organized structures in tracheary elements in secondary xylem.

• Premise of the study: Differences in leaf size are expected to be coordinated with various shoot traits and branching intensity because these relationships will influence light capture efficiency, water use, and biomechanics. Previous studies have mainly focused on interspecific patterns of these trait relationships, but not on intraspecific patterns at the geographic scale. We investigated intraspecific variation in shoot traits and branching intensity of Fagus crenata in Japan. • Methods: Allometric relationships between the traits of current-year shoots and branching intensity per branch unit of 1-m length on the main axis (BI) and its coordination with latitude were investigated using trees from 10 provenances in a common garden. • Key results: Individual trees originating from lower latitudes have smaller leaves with greater leaf mass per area and nitrogen content per area, greater Huber value (stem cross-sectional area per total leaf area [ATL) of current-year shoots, and greater BI. Notably, the slope of the log-log relationship between BI and ATL was close to -1.0 across the trees from different source sites, implying that branching in this species occurs to control leaf area. 0 • Conclusions: Shoot traits and branching intensity were apparently coordinated with leaf size to control leaf area deployment in this species. Such patterns probably reflect differences in competition for hydraulic conductance among nearby shoots within crowns, as a consequence of different meteorological conditions across the source sites.

Because of their overwhelming size over other organisms, trees define the structural and energetic properties of forest ecosystems. From grasslands to forests, leaf area index, which determines the amount of light energy intercepted for photosynthesis, increases with increasing canopy height across the various terrestrial ecosystems of the world. In vertically well-developed forests, niche differentiation along the vertical gradient of light availability may promote species coexistence. In addition, spatial and temporal differentiation of photosynthetic traits among the coexisting tree species (functional diversity) may promote complementary use of light energy, resulting in higher biomass and productivity in multi-species forests. Trees have evolved retaining high phenotypic plasticity because the spatial/temporal distribution of resources in forest ecosystems is highly heterogeneous and trees modify their own environment as they increase nearly 1,000 times in size through ontogeny. High phenotypic plasticity may enable coexistence of tree species through divergence in resource-rich environments, as well as through convergence in resource-limited environments. We propose that the breadth of individual-level phenotypic plasticity, expressed at the metamer level (leaves and shoots), is an important factor that promotes species coexistence and resource-use complementarity in forest ecosystems. A cross-biome comparison of the link between plasticity of photosynthesis-related traits and stand productivity will provide a functional explanation for the relationship between species assemblages and productivity of forest ecosystems.

In temperate regions, trees undergo annual cycles of cambial growth, with periods of cambial activity and dormancy. Environmental factors might regulate the cambial growth, as well as the development of cambial derivatives. We investigated the effects of low temperature by localized cooling on cambial activity and latewood formation in two conifers, Chamaecyparis obtusa and Cryptomeria japonica. A plastic rubber tube that contained cooled water was wrapped around a 30-cm-wide portion of the main stem of Chamaecyparis obtusa and Cryptomeria japonica trees during seasons of active cambium. Small blocks were collected from both cooled and non-cooled control portions of the stems for sequential observations of cambial activity and for anatomical measurements of cell morphology by light microscopy and image analysis. The effect of localized cooling was first observed on differentiating tracheids. Tracheids narrow in diameter and with significantly decreased cambial activity were evident 5 weeks after the start of cooling in these stems. Eight weeks after the start of cooling, tracheids with clearly diminished diameters and thickened cell walls were observed in these stems. Thus, localized low temperature induced narrow diameters and obvious thickening of secondary cell walls of tracheids, which were identified as latewood tracheids. Two months after the cessation of cooling, a false annual ring was observed and cambium became active again and produced new tracheids. In Cryptomeria japonica, cambial activity ceased earlier in locally cooled portions of stems than in non-cooled stems, indicating that the cambium had entered dormancy sooner in the cooled stems. Artificial cooling of stems induced latewood formation and cessation of cambial activity, indicating that cambium and its derivatives can respond directly to changes in temperature. A decrease in the temperature of the stem is a critical factor in the control of cambial activity and xylem differentiation in trees.

• Background and Aims The networks of vessel elements play a vital role in the transport of water from roots to leaves, and the continuous formation of earlywood vessels is crucial for the growth of ring-porous hardwoods. The differentiation of earlywood vessels is controlled by external and internal factors. The present study was designed to identify the limiting factors in the induction of cambial reactivation and the differentiation of earlywood vessels, using localized heating and disbudding of dormant stems of seedlings of a deciduous ring-porous hardwood, Quercus serrata. • Methods Localized heating was achieved by wrapping an electric heating ribbon around stems. Disbudding involved removal of all buds. Three treatments were initiated on 1 February 2012, namely heating, disbudding and a combination of heating and disbudding, with untreated dormant stems as controls. Cambial reactivation and differentiation of vessel elements were monitored by light and polarized-light microscopy, and the growth of buds was followed. • Key Results Cambial reactivation and differentiation of vessel elements occurred sooner in heated seedlings than in non-heated seedlings before bud break. The combination of heating and disbudding of seedlings also resulted in earlier cambial reactivation and differentiation of first vessel elements than in non-heated seedlings. A few narrow vessel elements were formed during heating after disbudding, while many large earlywood vessel elements were formed in heated seedlings with buds. • Conclusions The results suggested that, in seedlings of the deciduous ring-porous hardwood Quercus serrata, elevated temperature was a direct trigger for cambial reactivation and differentiation of first vessel elements. Bud growth was not essential for cambial reactivation and differentiation of first vessel elements, but might be important for the continuous formation of wide vessel elements.

Key message Complete differentiation of the first earlywood vessels occurred earlier in upper regions of stems than in middle and lower regions when buds swelling in a ring-porous hardwood Quercus serrata seedlings. In deciduous ring-porous hardwoods, the timing of the onset of conduction of water via the networks of the current year’s earlywood vessels is very important for the growth of buds and shoots because the main pathways for conduction of water are the networks of the current year’s earlywood vessels. The purpose of this study was to visualize the formation of the networks of first earlywood vessels in the current year’s xylem of seedlings of the deciduous ring-porous hardwood Quercus serrata . We monitored the distribution of water in the current and the previous year’s secondary xylem at the cellular level in upper, middle and lower regions of stems during the formation of earlywood vessels by cryo-scanning electron microscopy after freeze-etching. We also examined how changes in water distribution were correlated with leaf phenology. The contents of the first vessel elements in the upper region of the stem changed from cytoplasm-rich to water earlier than those in middle and lower regions of the stem when buds were increasing in size. At bud break, vessel elements were filled with water throughout the entire stem. When the cambium was dormant and during formation of earlywood vessels, the previous year’s latewood vessels were filled with water. Our results showed that complete differentiation of vessel elements occurred earlier in upper regions of stems than in middle and lower regions. Moreover, the functional networks of the previous year’s latewood vessels appeared to be involved in supplying water to new networks of earlywood vessels in the current year’s xylem.

Key message We observed the formation of latewood tracheids with narrow diameters and thick walls and the disappearance of stored starch around the cambium on the locally heated region of stems in evergreen conifer Chamaecyparis pisifera during winter cambial dormancy. Wood formation is controlled by cambial cell division, which determines the quantity and quality of wood. We investigated the factors that control cambial activity and the formation of new tracheids in locally heated stems of the evergreen conifer Chamaecyparis pisifera . Electric heating tape was wrapped around one side of the stem, at breast height, of two trees in 2013 and two in 2014. Pairs of stems were locally heated in winter, and small blocks were collected from heated and non-heated regions of stems. Cambial activity and levels of stored starch around the cambium were investigated by microscopy. Cambial reactivation and xylem differentiation occurred earlier in heated than in non-heated regions. New cell plates were formed after 14–18 days of heating. After a few layers of tracheids with large diameters and thin walls had formed, cell division and cell enlargement during differentiation were inhibited. Tracheids with narrow diameters and thick walls, defining those as latewood, were formed near the cambium, and finally, four to six layers of tracheids were induced. After cambial reactivation, amounts of stored starch started to decrease and starch disappeared completely from phloem and xylem cells that were located near the cambium during the differentiation of heated regions. Our results suggest that an increase in temperature induces the conversion of stored starch to soluble sugars for continuous cambial cell division and earlywood formation. By contrast, a shortage of stored starch might be responsible for inhibition of cambial activity and induction of the formation of latewood tracheids.

Key message Earlywood width in Quercus crispula increased from 1970 to 2004 without changes of vessel anatomy and ring growth. The increase in diameter of a tree stem is an important indicator of forest productivity. Xylem traits, such as the number and cross-sectional area of earlywood vessels, are also critical parameters of forest growth because of the physiological and structural contribution of xylem to the growth of the tree stem. Forest productivity appears to be affected by climate change and, indeed, trees might be expected to acclimate to gradual long-term climate change. The aim of this study was to identify long-term changes in increases in stem diameter and in earlywood vessels by examining tree rings of Quercus crispula . Focusing on 20 mature specimens of Q. crispula , we examined annual ring growth from 1970 to 2004 and measured earlywood traits, namely, the width, cross-sectional area (henceforth referred to as area) and number of earlywood vessels, by digital image analysis. We developed a hierarchical Bayesian model for detection of long-term trends in these traits. We found that earlywood width, as well as the total number and area of earlywood vessels, increased during the 35 years under analysis. One possible cause of these changes might be the long-term elevation of temperatures in early spring, which determine the timing of the onset of cambial reactivation from winter dormancy. In contrast to the long-term changes, short-term, yearly changes in earlywood traits fluctuated to a smaller extent than yearly changes in tree ring width. Therefore, the observed long-term changes in earlywood appear to represent acclimation to long-term climate change.

Key message Calli of hybrid poplar that had been exposed to desiccation in air before transfer to the induction medium differentiated into tracheary elements at higher rates than calli without air desiccation. Cells of hybrid poplar ( Populus sieboldii x P. grandidentata ) in culture can be induced to differentiate into secondary xylem-like tracheary elements that form the highly developed bordered pits and broad regions of cell walls in contrast to helical or reticulate wall thickenings in primary xylem elements. We attempted to increase the rate of differentiation of tracheary elements from calli using a combination of hormonal stimulation and partial desiccation. Calli that had been exposed to desiccation in air in a clean hood for 90 min before transfer to the induction medium differentiated into tracheary elements at higher rates than calli without air desiccation. The partial desiccation treatment had no effects on the features of the induced tracheary elements and the frequencies with which they appeared. Our results show that partial desiccation can increase, approximately threefold the rate of differentiation of secondary xylem-like tracheary elements from calli of hybrid poplar. This improvement in the rate of differentiation tracheary elements in vitro should facilitate detailed future analysis of the differentiation of secondary xylem.