Eri Nabeshima

The effects of localized heating and disbudding on cambial reactivation and formation of earlywood vessels in seedlings of the deciduous ring-porous hardwood, Quercus serrata.

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.

The need for a canopy perspective to understand the importance of phenotypic plasticity for promoting species coexistence and light-use complementarity in forest ecosystems

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.

Localized cooling of stems induces latewood formation and cambial dormancy during seasons of active cambium in conifers

BACKGROUND AND AIMS 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. METHODS 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. KEY RESULTS 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. CONCLUSIONS 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.

Common allometric response of open-grown leader shoots to tree height in co-occurring deciduous broadleaved trees

BACKGROUND AND AIMS Morphology of crown shoots changes with tree height. The height of forest trees is usually correlated with the light environment and this makes it difficult to separate the effects of tree size and of light conditions on the morphological plasticity of crown shoots. This paper addresses the tree-height dependence of shoot traits under full-light conditions where a tree crown is not shaded by other crowns. METHODS Focus is given to relationships between tree height and top-shoot traits, which include the shoots leaf-blades and non-leafy mass, its total leaf-blade area and the length and basal diameter of the shoots stem. We examine the allometric characteristics of open-grown current-year leader shoots at the tops of forest tree crowns up to 24 m high and quantify their responses to tree height in 13 co-occurring deciduous hardwood species in a cool-temperate forest in northern Japan. KEY RESULTS Dry mass allocated to leaf blades in a leader shoot increased with tree height in all 13 species. Specific leaf area decreased with tree height. Stem basal area was almost proportional to total leaf area in a leader shoot, where the proportionality constant did not depend on tree height, irrespective of species. Stem length for a given stem diameter decreased with tree height. CONCLUSIONS In the 13 species observed, height-dependent changes in allometry of leader shoots were convergent. This finding suggests that there is a common functional constraint in tree-height development. Under full-light conditions, leader shoots of tall trees naturally experience more severe water stress than those of short trees. We hypothesize that the height dependence of shoot allometry detected reflects an integrated response to height-associated water stress, which contributes to successful crown expansion and height gain.

Geographic variation in shoot traits and branching intensity in relation to leaf size in Fagus crenata: A common garden experiment.

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. 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.

Climate change and the regulation of wood formation in trees by temperature

Key messageA 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.AbstractThe 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.

Xylogenesis in Trees: From Cambial Cell Division to Cell Death

Abstract Wood is produced by the cambium of stems of trees. The cambial activity of trees exhibits seasonal cycles of activity and dormancy in temperate and cool zones. The timing of cambial reactivation in the early spring plays an important role in the determination of the quantity of wood and the environmental adaptivity of trees. Numerous studies have demonstrated that the cambial reactivation is controlled by the temperature of stems. As soon as cambial cells lose the ability to divide, they start to differentiate into secondary xylem cells. Finally, cell death occurs in secondary xylem cells, but its timing is different among the types and positions of cells. During formation of the secondary wall in tracheids or wood fibers, the cellulose microfibrils change their orientation progressively to form thick cell walls with a highly organized structure. In addition, secondary xylem cells form modifications of the cell wall, such as helical thickening and pits, by the localized deposition of cellulose microfibrils. There are considerable evidences that the dynamics of cortical microtubules are closely related to the orientation and localization of newly deposited cellulose microfibrils in the differentiating secondary xylem cells, indicating that microtubules control the structure of cell wall, thereby wood quality.

A Dendroecological Analysis of Forest Dynamics for Old-Growth Abies-Tsuga-Quercus on the Boso Peninsula, Southeastern Japan

Abstract This study investigated the composition, age- and size-structure, and tree-ring relationships for an old-growth, warm-temperate, mixed-evergreen forest at the University of Tokyo Chiba Forest, Japan. A total of 32 tree species were recorded, which was dominated by Abies firma and Quercus acuta. Tsuga sieboldii dominated the recruitment after 1850, followed by Abies firma. After 1920, many individuals of Castanopsis, Cinnamomum, Cleyera and Quercus became established. The temporal pattern of conifer recruitment did not correspond to the record of strong wind events. Basal area increment in Abies firma and Castanopsis sieboldii trees increased throughout their lives, a trend not seen in the ring width index. Mean annual temperature was below the 100-year mean between 1920 and 1940 and 1960–1980, but increased rather abruptly after 1980. Mean annual precipitation decreased after 1960. Tree-ring releases are very common at the study forest, which are indicative of frequent small to moderate-sized disturbances. At least one release was recorded in every decade from 1890 to the present day, which is likely the primary causal factor promoting tree growth and recruitment. Our results suggest that early logging activities coupled with natural disturbances had a great influence on the developmental process and current structure of the study stand and that tree growth is varying in a manner consistent with forest dynamics.

Formation of new networks of earlywood vessels in seedlings of the deciduous ring-porous hardwood Quercus serrata in springtime

Key messageComplete 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.AbstractIn 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.

Seasonal changes of δD and δ18O in tree-ring cellulose of Quercus crispula suggest a change in post-photosynthetic processes during earlywood growth

Leaf photosynthetic and post-photosynthetic processes modulate the isotope ratios of tree-ring cellulose. Post-photosynthetic processes, such as the remobilization of stored starch in early spring, are important to understanding the mechanisms of xylem formation in tree stems; however, untangling the isotope ratio signals of photosynthetic and post-photosynthetic processes imprinted on tree rings is difficult. Portions of carbon-bound hydrogen and oxygen atoms are exchanged with medium water during post-photosynthetic processes. We investigated the δD and δ18O values of tree-ring cellulose using Quercus crispula Blume trees in two different habitats to evaluate seasonal changes in the exchange rate (f-value) of hydrogen or oxygen with medium water, and examined the associations of the post-photosynthetic processes. Theoretically, if the f-value is constant, δD and δ18O would be positively correlated due to meteorological factors, while variation in the f-value will create a discrepancy and weak correlation between δD and δ18O due to the exchange of carbon-bound hydrogen and oxygen with medium water. The values of δD decreased drastically from earlywood to latewood, while those of δ18O increased to a peak and then decreased toward the latewood. The estimated seasonal f-value was high at the beginning of earlywood and decreased toward the latewood. The post-photosynthetic processes associated with changes in the f-value were the remobilization of stored starch and triose cycling during cellulose synthesis because of the shortage of photo-assimilates in early spring. Although we did not evaluate relevant physiological parameters, the seasonal pattern of δD and δ18O in tree-ring cellulose of Q. crispula was clear, suggesting that the dual isotope (δD and δ18O) approach can be used to reveal the resource allocation mechanisms underlying seasonal xylem formation.