Role of axis reversal from the short-shoot to long-shoot habit for crown maintenance in slow-growing Betula maximowicziana trees.

PREMISE OF THE STUDY: Branch growth and its spatial arrangement determine crown architecture, leaf display, and, thus, the productivity of trees. Branch axes elongate by the sequential production of shoots with differing morphology and function, such as short shoots and long shoots. This study investigated ontogenetic changes in axis growth in Betula maximowicziana and quantified the role of axis reversal between the short-shoot and long-shoot habits, particularly reversal from the short-shoot to the long-shoot habit. METHODS: From 8 trees with varying levels of growth vigor, 716 branch axes forming the basic crown architecture were sampled. Past growth of the branch axes was reconstructed from leaf and bud scale scars and compared between slow-growing and vigorously growing trees. KEY RESULTS: Branch axes reversed more frequently between the long- and short-shoot habits in slow-growing trees than in vigorously growing trees. Short-shoot-origin axes that reversed to the long-shoot habit lived for longer periods and grew larger than axes that remained in the short-shoot habit. Short-shoot-origin axes reversed as they grew away from branch apices, typically >6 yr after they had originated. CONCLUSIONS: Reversal of short-shoot-origin axes to the long-shoot habit is an endogenous growth process of trees with reduced vigor. Like epicormic branching, the reversal may contribute to the maintenance of productivity of large old trees by prolonging axis longevity and filling the inner part of the crown. This study presents an ontogenetic change in branch growth, which broadens our perspectives on the growth and survival of long-living trees.

Effective distance of volatile cues for plant-plant communication in beech.

In response to volatiles emitted from a plant infested by herbivorous arthropods, neighboring undamaged conspecific plants become better defended against herbivores; this is referred to as plant‒plant communication. Although plant‒plant communication occurs in a wide range of plant species, most studies have focused on herbaceous plants. Here, we investigated plant‒plant communication in beech trees in two experimental plantations in 2018 and one plantation in 2019. Approximately 20% of the leaves of a beech tree were clipped in half in the spring seasons of 2018 and 2019 (clipped tree). The damage levels to leaves in the surrounding undamaged beech trees were evaluated 90 days after the clipping (assay trees). In both years, the damage levels decreased with a reduction in the distance from the clipped tree. In 2019, we also recorded the damage levels of trees that were not exposed to volatiles (nonexposed trees) as control trees and found that those that were located <5 m away from clipped trees had significantly less leaf damage than nonexposed trees. By using a gas chromatograph-mass spectrometer, ten and eight volatile compounds were detected in the headspaces of clipped and unclipped leaves, respectively. Among them, the amount of (Z)-3-hexenyl acetate in clipped leaves was significantly higher than that in nonclipped leaves. Our result suggests that green leaf volatiles such as (Z)-3-hexenol and (Z)-3-hexenyl acetate and other volatile organic compounds emitted from clipped trees induced defenses in the neighboring trees within the 5 m radius. The effective distances of plant‒plant communication in trees were discussed from the viewpoint of the arthropod community structure in forest ecosystems.

Identification of management units in threatened populations of Arnica mallotopus Makino (Asteraceae) using novel EST-SSR markers.

Arnica mallotopus is a perennial herb endemic to the snowy regions of Japan. At the southern edge of its distribution, in Kyoto Prefecture, overgrazing by sika deer and decreased snowfall have resulted in the rapid decline of A. mallotopus populations. Therefore, there is an urgent need for a conservation genetic analysis of the remaining local populations. In this study, we first developed 13 EST-SSR markers to evaluate genetic variation in A. mallotopus. The average number of alleles per locus was 5.33. Genetic analysis using these markers showed that the investigated samples were classified into two groups corresponding to landscape structure. One group isolated from a tributary of the Yura River showed a strong population bottleneck signal, likely resulting from founder effects and subsequent drifts. On the other hand, the genetic diversity of the second group in the main distribution along the Yura River was higher and less inbred. Overall, our assessment suggested recognizing the two genetic groups as management units in conservation programs for the threatened populations.

Seasonal and annual fluctuations of deer populations estimated by a Bayesian state-space model.

Deer overabundance is a contributing factor in the degradation of plant communities and ecosystems worldwide. The management and conservation of the deer-affected ecosystems requires us to urgently grasp deer population trends and to identify the factors that affect them. In this study, we developed a Bayesian state-space model to estimate the population dynamics of sika deer (Cervus nippon) in a cool-temperate forest in Japan, where wolves (Canis lupus hodophilax) are extinct. The model was based on field data collected from block count surveys, road count surveys by vehicles, mortality surveys during the winter, and nuisance control for 12 years (2007-2018). We clarified the seasonal and annual fluctuation of the deer population. We found a peak of deer abundance (2010) over 12 years. In 2011 the estimated deer abundance decreased drastically and has remained at a low level then. The deer abundance gradually increased from April to December during 2013-2018. The seasonal fluctuation we detected could reflect the seasonal migration pattern of deer and the population recruitment through fawn births in early summer. In our model, snowfall accumulation, which can be a lethal factor for deer, may have slightly affected their mortality during the winter. Although we could not detect a direct effect of snow on population dynamics, snowfall decrease due to global warming may decelerate the winter migration of deer; subsequently, deer staying on-site may intensively forage evergreen perennial plants during the winter season. The nuisance control affected population dynamics. Even in wildlife protection areas and national parks where hunting is regulated, nuisance control could be effective in buffering the effect of deer browsing on forest ecosystems.

Annual and spatial variation in shoot demography associated with masting in Betula grossa: comparison between mature trees and saplings.

BACKGROUNDS AND AIMS: Shoot demography affects the growth of the tree crown and the number of leaves on a tree. Masting may cause inter-annual and spatial variation in shoot demography of mature trees, which may in turn affect the resource budget of the tree. The aim of this study was to evaluate the effect of masting on the temporal and spatial variations in shoot demography of mature Betula grossa. METHODS: The shoot demography was analysed in the upper and lower parts of the tree crown in mature trees and saplings over 7 years. Mature trees and saplings were compared to differentiate the effect of masting from the effect of exogenous environment on shoot demography. The fate of different shoot types (reproductive, vegetative, short, long), shoot length and leaf area were investigated by monitoring and by retrospective survey using morphological markers on branches. The effects of year and branch position on demographic parameters were evaluated. KEY RESULTS: Shoot increase rate, production of long shoots, bud mortality, length of long shoots and leaf area of a branch fluctuated periodically from year to year in mature trees over 7 years, in which two masting events occurred. Branches within a crown showed synchronized annual variation, and the extent of fluctuation was larger in the upper branches than the lower branches. Vegetative shoots varied in their bud differentiation each year and contributed to the dynamic shoot demography as much as did reproductive shoots, suggesting physiological integration in shoot demography through hormonal regulation and resource allocation. CONCLUSIONS: Masting caused periodic annual variation in shoot demography of the mature trees and the effect was spatially variable within a tree crown. Since masting is a common phenomenon among tree species, annual variation in shoot demography and leaf area should be incorporated into resource allocation models of mature masting trees.

A New Model for Size-Dependent Tree Growth in Forests.

Tree growth, especially diameter growth of tree stems, is an important issue for understanding the productivity and dynamics of forest stands. Metabolic scaling theory predicted that the 2/3 power of stem diameter at a certain time is a linear function of the 2/3 power of the initial diameter and that the diameter growth rate scales to the 1/3 power of the initial diameter. We tested these predictions of the metabolic scaling theory for 11 Japanese secondary forests at various growth stages. The predictions were not supported by the data, especially in younger stands. Alternatively, we proposed a new theoretical model for stem diameter growth on the basis of six assumptions. All these assumptions were supported by the data. The model produced a nearly linear to curvilinear relationship between the 2/3 power of stem diameters at two different times. It also fitted well to the curvilinear relationship between diameter growth rate and the initial diameter. Our model fitted better than the metabolic scaling theory, suggesting the importance of asymmetric competition among trees, which has not been incorporated in the metabolic scaling theory.