Relationship between the vertical distribution of fine roots and residual soil nitrogen along a gradient of hardwood mixture in a conifer plantation.

In forest ecosystems, understanding the relationship between the vertical distribution of fine roots and residual soil nitrogen is essential for clarifying the diversity-productivity-water purification relationship. Vertical distributions of fine-root biomass (FRB) and concentrations of nitrate-nitrogen (NO3 -N) in soil water were investigated in a conifer plantation with three thinning intensities (Control, Weak and Intensive), in which hardwood abundance and diversity were low, moderate and high, respectively. Intensive thinning led to the lowest NO3 -N concentration in soil water at all depths (0-100 cm) and highest FRB at shallow depths (0-50 cm). The NO3 -N concentration at a given depth was negatively correlated with total FRB from the surface to the depth at which NO3 -N concentration was measured, especially at shallow depths, indicating that more abundant fine roots led to lower levels of downward NO3 -N leaching. FRB contributed positively to nitrogen content of hardwood leaves. These findings demonstrate that a hardwood mixture in conifer plantations resulted in sufficient uptake of NO3 -N from soil by well developed fine-root systems, and translocation to canopy foliage. This study suggests that productivity and water purification can be achieved through a hardwood mixture in conifer plantations.

Conspecific distance-dependent seedling performance, and replacement of conspecific seedlings by heterospecifics in five hardwood, temperate forest species.

The mechanisms driving species diversity in the context of Janzen-Connell model are best understood by evaluating not only conspecific distance-dependent (CDD) seedling performance, but also replacement of conspecific seedlings by heterospecific seedlings beneath adult trees. We evaluated CDD and replacement as a log response ratio of seedling performance (height, age) directly beneath and at a distance from adult plants in a temperate forest, and examined the log response ratio of that between conspecifics and heterospecifics beneath adults for five hardwood species with different ecological traits (e.g., seed size, mycorrhizal type, relative abundance). CDD was greater in three small-seeded species with arbuscular mycorrhizae (AM) associations than it was in two large-seeded species with ectomycorrhizae (EM) associations. Replacement was also higher for small-seeded AM species compared to large-seeded EM species, resulting in a strong, positive relationship between CDD and replacement. The traits suggest that small-seeded AM seedlings are more likely to be replaced by heterologous seedlings beneath the adults than large-seeded EM seedlings, probably due to that the small-seeded AM species are more susceptible to attack by plant natural enemies (e.g., soil pathogens, leaf diseases). As a result, small-seeded AM species had lower relative abundances compared to large-seeded EM species. This study suggests that either seed size or associations with microorganisms play an important role in driving forest diversity by regulating replacement and CDD, although relative importance of the two traits (i.e., seed size, mycorrhizal type) remains unclear, because of the autocorrelation between the two traits for the five species studied.

Leaf diseases drive the Janzen-Connell mechanism regardless of light conditions: a 3-year field study.

In forests, negative density/distance-dependent seedling mortality (NDD) caused by natural enemies plays a key role in maintaining species diversity [Janzen-Connell (J-C) model]. However, the relative importance of natural enemies in mediating NDD under heterogeneous light conditions has remained unclear. We examined the relative importance of pathogens (i.e., soil pathogens, leaf diseases) on seedling performance in forest understories (FUs) and gaps (gaps) during a 3-year period (results of first year of our study have been previously reported). For the hardwood, Prunus grayana, we investigated seedling mortality, morbidity agents, growth, and root infection by arbuscular mycorrhizal fungi (AMF) beneath conspecific and heterospecific adults in FUs and gaps. Seedling mortality was higher beneath conspecific than heterospecific adults throughout 3 years at both sites, mainly due to continuous leaf disease (i.e., angular leaf spot), whereas damping-off diseases caused mortality only in the first year. Beneath each adult, seedling mortality was higher in FUs than in gaps until second year, but it did not differ between two habitat types in the third year, because leaf diseases caused severe damage even in gaps. Seedling mass was significantly lower beneath conspecific adults. AMF infection of seedlings was also lower beneath conspecific adults, while it was higher in gaps than in FUs beneath both adults. This study demonstrates that the J-C model in a hardwood tree, P. grayana is mainly driven by high NDD seedling mortality caused by airborne leaf diseases, which continuously attack seedlings in a NDD manner regardless of environmental light conditions.

Carbon allocation to defense, storage, and growth in seedlings of two temperate broad-leaved tree species.

Optimal carbon allocation to growth, defense, or storage is a critical trait in determining the shade tolerance of tree species. Thus, examining interspecific differences in carbon allocation patterns is useful when evaluating niche partitioning in forest communities. We hypothesized that shade-tolerant species allocate more carbon to defense and storage and less to growth compared to shade-intolerant species. In gaps and forest understory, we measured relative growth rates (RGR), carbon-based defensive compounds (condensed tannin, total phenolics), and storage compounds (total non-structural carbohydrate; TNC) in seedlings of two tree species differing in shade tolerance. RGR was greater in the shade-intolerant species, Castanea crenata, than in the shade-tolerant species, Quercus mongolica var. grosseserrata, in gaps, but did not differ between the species in the forest understory. In contrast, concentrations of condensed tannin and total phenolics were greater in Quercus than in Castanea at both sites. TNC pool sizes did not differ between the species. Condensed tannin concentrations increased with increasing growth rate of structural biomass (GRstr) in Quercus but not in Castanea. TNC pool sizes increased with increasing GRstr in both species, but the rate of increase did not differ between the species. Accordingly, the amount of condensed tannin against TNC pool sizes was usually higher in Quercus than in Castanea. Hence, Quercus preferentially invested more carbon in defense than in storage. Such a large allocation of carbon to defense would be advantageous for a shade-tolerant species, allowing Quercus to persist in the forest understory where damage from herbivores and pathogens is costly. In contrast, the shade-intolerant Castanea preferentially invested more carbon in growth rather than defense (and similar amounts in storage as Quercus), ensuring establishment success in gaps, where severe competition occurs for light among neighboring plants. These contrasting carbon allocation patterns are closely associated with strategies for persistence in these species' respective habitats.

Pollen donor composition during the early phases of reproduction revealed by DNA genotyping of pollen grains and seeds of Castanea crenata.

In plants, pollen donor composition can differ during the early phases of reproduction through various selection mechanisms favouring self, related or nonrelated pollen donors, but such differences have not been examined under natural conditions because paternity is difficult to analyse in a natural setting. Here, we performed paternity analyses based on microsatellite genotyping of individual pollen grains deposited on female flowers (n = 773) and seeds (n = 304) to evaluate pollen donor composition from three individuals of the insect-pollinated monoecious tree Castanea crenata in a natural forest. Spatial genetic structure was also investigated. A mean self-pollen rate of 90.2% was observed at the pollination stage, but a low selfing rate of 0.3% was observed at the seed stage. In outcross events, however, pairwise distance and relatedness between maternal and paternal parents were not different between pollination and seed stages. We also observed significant positive relatedness, based on clear fine-scale genetic structure of individual trees within 80 m of one another, and 71% of seeds were derived using pollen grains of related trees within 80 m. The results suggest that the mechanism of self-incompatibility strongly avoids self-pollen before seed production. However, the avoidance of biparental inbreeding was not obvious between pollination and seed stages.

Ecological properties of shoot- and single seeds in a hardwood, Zelkova serrata.

The data presented in this paper is supporting the research article　"Variable seed behavior increases recruitment success of a hardwood tree, Zelkova serrata, in spatially heterogeneous forest environments" (Oyama et al., 2018) [1]. We provided the data of several ecological properties of the two types of the seeds (i.e. shoot seeds vs. single seeds) with distinctly different dispersal behaviors. We provide data of terminal velocity, which was measured by releasing 50 replicates of each seed type from a height of 5.0 m in dead air space in a gymnasium. We also show the data of germination cue [i.e. red:far-red (R:FR) ratios], which was examined in plant growth chambers that received three distinct R:FR ratios (0.1, 0.4, and 1.0; 16 h photoperiod) or no illumination. Further, we show the data of the rates of multi-locus outcrossing rates and biparental inbreeding in each of single- and shoot seeds. The mating system parameters were estimated by assaying a total of 80 shoot seeds and 70 single seeds that were randomly collected from Parent 1 and Parent 2 for five microsatellite loci. Finally, we show the data of hemispherical canopy photographs, which were taken at different distance from the adults using a digital camera equipped with a fisheye lens.

Variation in pollen-donor composition among pollinators in an entomophilous tree species, Castanea crenata, revealed by single-pollen genotyping.

BACKGROUND: In plants, reproductive success is largely determined by the composition of pollen (i.e., self-pollen and outcross-pollen from near and distant pollen-donors) transported as a result of pollinator foraging behavior (e.g., pollen carryover). However, little evidence is available on how and to what extent the pollen carryover affects the pollen-donor composition and on which insect taxa are effective outcross-pollen transporters under field conditions. In this study, we explored roles of foraging behavior of insect pollinators on pollen-donor composition and subsequent reproductive success in a woody plant. METHODS: We performed paternity analyses based on microsatellite genotyping of individual pollen grains found on diurnal pollinators (i.e., bumblebee, small bee, fly, small beetle, and honeybee) visiting Castanea crenata trees. RESULTS: The outcross-pollen rate was highest in bumblebees (66%), followed by small bees (35%), flies (31%), and small beetles (18%). The effective number of pollen donors, representing pollen carryover, was greater in bumblebees (9.71) than in flies (3.40), small bees (3.32), and small beetles (3.06). The high percentages of pollen from outside the plot on bumblebees (65.4%) and flies (71.2%) compared to small bees (35.3%) and small beetles (13.5%) demonstrated their longer pollen dispersal distances. CONCLUSIONS: All of the diurnal insects carried outcross-pollen grains for long distances via pollen carryover. This fact suggests that a wide range of insect taxa are potential outcross-pollen transporters for the self-incompatible C. crenata.

Leaf longevity as a normalization constant in allometric predictions of plant production.

In metabolic scaling theory the size-dependence of plant processes is described by a power function of form Y=Y o M (θ) where Y is a characteristic such as plant productivity that changes with plant size (M) raised to the θ (th) power and Y o is a normalization constant that adjusts the general relationship across environments and species. In essence, the theory considers that the value of θ arises in the size-dependent relationship between leaf area and vascular architecture that influences plant function and that Y o modulates this general relationship to account for ecological and evolutionary effects on the exchange of resources between plant and environment. Enquist and colleagues have shown from first principles that Y o is a function of carbon use efficiency, the carbon fraction of a plant, the area-specific carbon assimilation rate of a leaf, the laminar area of a leaf, and the mass of a leaf. We show that leaf longevity provides a functional integration of these traits that can serve as a simpler normalization in scaling plant productivity for individual species and potentially for mixed-species communities as well.

Trade-offs between seedling growth and survival in deciduous broadleaved trees in a temperate forest.

BACKGROUND AND AIMS: In spatially heterogeneous environments, a trade-off between seedling survival and relative growth rate may promote the coexistence of plant species. In temperate forests, however, little support for this hypothesis has been found under field conditions, as compared with shade-house experiments. Performance trade-offs were examined over a large resource gradient in a temperate hardwood forest. METHODS: The relationship between seedling survival and seedling relative growth rate in mass (RGR(M)) or height (RGR(H)) was examined at three levels of canopy cover (forest understorey, FU; small gap, SG; and large gap, LG) and at two microsites within each level of canopy cover (presence or absence of leaf litter) for five deciduous broad-leaved tree species with different seed sizes. KEY RESULTS: Within each species, both RGR(M) and RGR(H) usually increased with increasing light levels (in the order FU < SG < LG), whereas little difference was observed based on the presence or absence of litter. Seedling survival in FU was negatively correlated with both RGR(M) and RGR(H) in both LG and SG. The trade-off between high-light growth and low-light survival was more evident in the relationship with LG as compared with SG. An intraspecific trade-off between survival and RGR was observed along environmental gradients in Acer mono, whereas seedlings of Betula platyphylla var. japonica survived and grew better in LG. CONCLUSIONS: The results presented here strongly support the idea of light gradient partitioning (i.e. species coexistence) in spatially heterogeneous light environments in temperate forests, and that further species diversity would be promoted by increased spatial heterogeneity. The intraspecific trade-off between survival and RGR in Acer suggests that it has broad habitat requirements, whereas Betula has narrow habitat requirements and specializes in high-light environments.

Effects of resource heterogeneity on nitrogen translocation within clonal fragments of Sasa palmata: an isotopic (15N) assessment.

BACKGROUND AND AIMS: Clonal fragments of the rhizomatous dwarf bamboo Sasa palmata, which widely predominates in temperate regions of Japan, were grown under heterogeneous resource conditions such as gap understories or nutrient-patchy grassland. Clonal fragments develop multiple ramets with long rhizomes and appear to be physiologically integrated by the translocation of assimilates. The glasshouse experiment reported here was designed to clarify the mechanisms of physiological integration of nitrogen more precisely. METHODS: To assess how resource conditions influence the amount of nitrogen translocation, and which organ acts as the strongest sink, two experiments were conducted that traced movement of 15N label between interconnected pairs of ramets to compare homogeneous and heterogeneous light and soil nitrogen conditions. KEY RESULTS: The amount of 15N translocated to leaves was between 9% and 11% greater in high-N and high-light ramets in the heterogeneous compared with homogeneous treatments. Under heterogeneous soil nitrogen conditions, translocation increased from individual ramets in resource-rich patches to ramets in resource-poor patches, while the reverse was true under heterogeneous light environments, reflecting differences in the positions of leaves that act as the strongest sinks. Neither the mass increments nor the total mass of clonal fragments was significantly affected by heterogeneity of either light or nutrients, possibly because the experimental period was too short for differences to manifest themselves. CONCLUSIONS: This study clearly demonstrated that nitrogen is readily translocated between ramets, particularly under heterogeneous resource conditions. The translocation patterns were governed by functional 'division of labour' mechanisms that resulted in net nitrogen movement from understory sites to gaps, thereby enhancing the carbon acquisition of the whole fragment. Thus, physiological integration may provide benefits for S. palmata when it is growing under heterogeneous conditions in which there are deficits of certain environmental resources.

Shoot life span in relation to successional status in deciduous broad-leaved tree species in a temperate forest.

In trees, leaf life span is closely related to successional status. Although leaves are attached to shoots, shoot life span has been insufficiently studied in the context of ecological systems. Interspecific variation in shoot survivorship was investigated over 27 months in 15 temperate hardwood tree species. Relationships between shoot architecture and shoot survival were also investigated. Shoot life span was shortest in early successional species, and longest in late successional species, in each of the families Betulaceae and Fagaceae. In Salicaceae, all of which were early successional species, shoot life span was longer in mountainous than in riparian species. Early successional or riparian species distributed longer shoots densely, even in proximal positions on mother shoots, resulting in mutual shading and consequent early and massive shoot shedding. By contrast, late successional or mountainous species concentrated shoots in distal positions, allowing shoots to receive equally favorable light, resulting in a longer life span. These results reveal close relationships between shoot life span and environmental resource availability or successional status and suggest a causal relationship between shoot shedding and shoot architecture.

Selfing and inbreeding depression in seeds and seedlings of Neobalanocarpus heimii (Dipterocarpaceae).

We evaluated the degree of selfing and inbreeding depression at the seed and seedling stages of a threatened tropical canopy tree, Neobalanocarpus heimii, using microsatellite markers. Selection resulted in an overall decrease in the level of surviving selfed progeny from seeds to established seedlings, indicating inbreeding depression during seedling establishment. Mean seed mass of selfed progeny was lower than that of outcrossed progeny. Since the smaller seeds suffered a fitness disadvantage at germination in N. heimii, the reduced seed mass of selfed progeny would be one of the determinants of the observed inbreeding depression during seedling establishment. High selfing rates in some mother trees could be attributed to low local densities of reproductive individuals, thus maintenance of a sufficiently high density of mature N. heimii should facilitate regeneration and conservation of the species.