Deposition is a phosphorus source for Fallopia japonica during early-stage primary succession.

Phosphorus is a key plant nutrient linked to plant growth during the early stages of primary succession in volcanic soils. Available phosphorus is thought to include soil and atmospheric phosphorus, but it is not well understood. Here, we focused on deposition as a potential phosphorus source. We evaluated the contribution of deposition to phosphorus uptake and growth in Fallopia japonica, a key pioneer species of primary succession. When we experimented with growing F. japonica under field conditions, F. japonica not covered by a roof absorbed more phosphorus than that covered by the roof, suggesting the influence of total (dry + wet) deposition. Furthermore, we tested the effects of deposition by treating F. japonica seedlings with wet deposition or distilled water in six volcanic soils. Plants that received the wet deposition treatment exhibited higher phosphorus contents and growth rates than those treated with distilled water. The phosphorus from wet deposition and the phosphorus from soil contributed nearly equally to F. japonica development. Our findings suggest that F. japonica grows during primary succession and builds up the phosphorus cycle by absorbing a trace amount of phosphorus from deposition and volcanic soils.

A tale of two lineages: how the strains of the earliest divergent symbiotic Frankia clade spread over the world.

It is currently assumed that around 100 million years ago, the common ancestor to the Fabales, Fagales, Rosales and Cucurbitales in Gondwana, developed a root nodule symbiosis with a nitrogen-fixing bacterium. The symbiotic trait evolved first in Frankia cluster-2; thus, strains belonging to this cluster are the best extant representatives of this original symbiont. Most cluster-2 strains could not be cultured to date, except for Frankia coriariae, and therefore many aspects of the symbiosis are still elusive. Based on phylogenetics of cluster-2 metagenome-assembled genomes (MAGs), it has been shown that the genomes of strains originating in Eurasia are highly conserved. These MAGs are more closely related to Frankia cluster-2 in North America than to the single genome available thus far from the southern hemisphere, i.e., from Papua New Guinea.To unravel more biodiversity within Frankia cluster-2 and predict routes of dispersal from Gondwana, we sequenced and analysed the MAGs of Frankia cluster-2 from Coriaria japonica and Coriaria intermedia growing in Japan, Taiwan and the Philippines. Phylogenetic analyses indicate there is a clear split within Frankia cluster-2, separating a continental from an island lineage. Presumably, these lineages already diverged in Gondwana.Based on fossil data on the host plants, we propose that these two lineages dispersed via at least two routes. While the continental lineage reached Eurasia together with their host plants via the Indian subcontinent, the island lineage spread towards Japan with an unknown host plant.

Exploring the mechanical and morphological rationality of tree branch structure based on 3D point cloud analysis and the finite element method.

Trees are thought to have acquired a mechanically optimized shape through evolution, but a scientific methodology to investigate the mechanical rationality of tree morphology remains to be established. The aim of this study was to develop a new method for 3D reconstruction of actual tree shape and to establish a theoretical formulation for elucidating the structure and function of tree branches. We obtained 3D point cloud data of tree shape of Japanese zelkova (Zelkova serrata) and Japanese larch (Larix kaempferi) using the NavVis Lidar scanner, then applied a cylinder structure extraction from point cloud data with error estimation. We then formulated the mechanical stress of branches under gravity using the elastic theory, and performed finite element method simulations to evaluate the mechanical characteristics. Subsequently, we constructed a mechanics-based theoretical formulation of branch development that ensures constant bending stress produces various branching patterns depending on growth properties. The derived theory recapitulates the trade-off among branch growth anisotropy, stress-gravity length, and branch shape, which may open the quantitative way to evaluate mechanical and morphological rationality of tree branches.

Fresh litter acts as a substantial phosphorus source of plant species appearing in primary succession on volcanic ash soil.

Plants have difficulty absorbing phosphorus from volcanic ash soils owing to the adsorption of phosphorus by aluminum and iron in the soils. Thus, on volcanic ash soils, the phosphorus source for natural vegetation is expected to be organic matter, however, there is a lack of experimental evidence regarding this occurrence. Here, we studied the effect of organic matter on plant growth of some species that occur in primary successions of volcanic ash soil ecosystems, based on growth experiments and chemical analyses. We found that a large amount of inorganic phosphorus (but only a limited amount of inorganic nitrogen) is leached from fresh leaf litter of the pioneer spices Fallopia japonica at the initial stage of litter decomposition. Phosphorus from the fresh litter specifically activated the growth of subsequently invading nitrogen-fixing alder when immature volcanic soil was used for cultivation. In contrast, old organic matter in mature soil was merely a minor source of phosphorus. These results suggest that fresh litter of F. japonica is essential for growth of nitrogen-fixing alder because the litter supplies phosphorus. We consider that rapid phosphorus cycles in fresh litter-plant systems underlie the productivity of natural vegetation even in mature ecosystems established on volcanic ash soils.

Biomass allocation and long-term growth patterns of temperate lianas in comparison with trees.

The host-dependent support habit of lianas is generally interpreted as a strategy designed to reduce resource investment in mechanical tissues; this allows preferential allocation to leaf and stem extension, thereby enhancing productivity and competitive abilities. However, this hypothesis has not been rigorously tested. We examined the aboveground allometries regarding biomass allocation (leaf mass and current-year stem mass (approximated as biomass allocated to extension growth) vs total aboveground mass) and long-term apparent growth patterns (height and aboveground mass vs age, i.e. numbers of growth rings) for nine deciduous liana species in Japan. Lianas had, on average, three- and five-fold greater leaf and current-year stem mass, respectively, than trees for a given aboveground mass, whereas the time course to reach the forest canopy was comparable and biomass accumulation during that period was only one-tenth that of co-occurring canopy trees. The balance between the lengths of yearly stem extension and existing older stems indicated that lianas lost c. 75% of stem length during growth to the canopy, which is probably a consequence of the host-dependent growth. Our observations suggest that, although lianas rely on hosts mechanically, allowing for short-term vigorous growth, this habit requires a large cost and could limit plant growth over protracted periods.

Roles of gibberellins and cytokinins in regulation of morphological and physiological traits in Polygonum cuspidatum responding to light and nitrogen availabilities.

We evaluated the roles of gibberellins (GAs) and cytokinins (CKs) in regulation of morphological traits such as biomass allocation and leaf mass per area (LMA). Seedlings of Polygonum cuspidatum Siebold & Zucc. were grown under various light and N availabilities. We exogenously sprayed solutions of gibberellin (GA3), benzyl adenine (BA), uniconazole (an inhibitor of GA biosynthesis) or their mixtures on the aboveground parts, and changes in morphological and physiological traits and relative growth rate (RGR) were analysed. Endogenous levels of GAs and CKs in the control plants were also quantified. The morphological traits were changed markedly by the spraying. Biomass allocation to leaves was increased by GA3 and BA, whereas it decreased by uniconazole. GA3 decreased LMA, whereas uniconazole increased it. We found close relationships among morphological and physiological traits such as photosynthetic rate and net assimilation rate, and RGR under all growth conditions. Seedlings with high levels of endogenous GAs or CKs and low levels of endogenous GAs or CKs showed morphologies similar to those sprayed with GA3 or BA, and those sprayed with uniconazole, respectively. Thus we concluded these phytohormones are involved in the regulation of biomass allocation responding to either light or N availability.

Variation in susceptibility to wind along the trunk of an isolated Larix kaempferi (Pinaceae) tree.

• Premise of the study: The force of the wind is a significant hazard to the survival of trees and can affect tree morphology. However, the actual distribution of the stress that wind causes to a tree trunk is not well understood in spite of its expected importance to tree morphology. The uniform stress hypothesis (i.e., tree trunks take a form that equalizes the distribution of stress along the outer surface of the stem) has been tested indirectly as a model, placing high importance on the mechanical safety of a trunk, and rejected theoretically. But stress on a tree's trunk has not yet been measured directly.• Methods: Actual strains at the surface of the trunk of an isolated Larix kaempferi tree due to wind loads were measured at seven heights on the trunk for 1 yr.• Key results: During the measurement period, wind-induced stress was higher in the upper portions of the trunk than in the lower portions, regardless of wind speed, and the difference increased as wind speed increased. The deflection of the trunk recorded at each position was also larger in the upper portions than in the lower portions.• Conclusions: The results indicate that the upper portions of the trunk of an isolated Larix kaempferi tree are more susceptible to wind than are the lower portions. These results do not support the uniform stress hypothesis and suggest that another limitation (either mechanical or nonmechanical) acts on the morphology of the tree.

Tree branching: Leonardo da Vinci's rule versus biomechanical models.

This study examined Leonardo da Vinci's rule (i.e., the sum of the cross-sectional area of all tree branches above a branching point at any height is equal to the cross-sectional area of the trunk or the branch immediately below the branching point) using simulations based on two biomechanical models: the uniform stress and elastic similarity models. Model calculations of the daughter/mother ratio (i.e., the ratio of the total cross-sectional area of the daughter branches to the cross-sectional area of the mother branch at the branching point) showed that both biomechanical models agreed with da Vinci's rule when the branching angles of daughter branches and the weights of lateral daughter branches were small; however, the models deviated from da Vinci's rule as the weights and/or the branching angles of lateral daughter branches increased. The calculated values of the two models were largely similar but differed in some ways. Field measurements of Fagus crenata and Abies homolepis also fit this trend, wherein models deviated from da Vinci's rule with increasing relative weights of lateral daughter branches. However, this deviation was small for a branching pattern in nature, where empirical measurements were taken under realistic measurement conditions; thus, da Vinci's rule did not critically contradict the biomechanical models in the case of real branching patterns, though the model calculations described the contradiction between da Vinci's rule and the biomechanical models. The field data for Fagus crenata fit the uniform stress model best, indicating that stress uniformity is the key constraint of branch morphology in Fagus crenata rather than elastic similarity or da Vinci's rule. On the other hand, mechanical constraints are not necessarily significant in the morphology of Abies homolepis branches, depending on the number of daughter branches. Rather, these branches were often in agreement with da Vinci's rule.

Steric control in the π-dimerization of oligothiophene radical cations annelated with bicyclo[2.2.2]octene units.

A Two series of oligothiophenes 2(nT) (n=4,5), annelated with bicyclo[2.2.2]octene (BCO) units at both ends, and quaterthiophenes 3 a-c, annelated with various numbers of BCO units at different positions, were newly synthesized to investigate the driving forces of π-dimerization and the structure-property relationships of the π-dimers of oligothiophene radical cations. Their radical-cation salts were prepared through chemical one-electron oxidation by using nitrosonium hexafluoroantimonate. From variable-temperature electron spin resonance and electronic absorption measurements, the π-dimerization capability was found to vary among the members of the 2(nT)(+)(·)SbF6(-) series and 3(+)(·)SbF6(-) series of compounds. To examine these results, density functional theory (DFT) calculations at the M06-2X/6-31G(d) level were conducted for the π-dimers. This level of theory was found to successfully reproduce the previously reported X-ray structure of (2(3T))2(2+) having a bent π-dimer structure with cis-cis conformations. The absorption bands obtained by time-dependent DFT calculations for the π-dimers were in reasonable agreement with the experimental spectra. The attractive and repulsive forces for the π-dimerization were divided into four factors: 1) SOMO-SOMO interactions, 2) van der Waals forces, 3) solvation, and 4) Coulomb repulsion, and the effects of each factor on the structural differences and chain-length dependence are discussed in detail.

Mutation accumulation in a selfing population: consequences of different mutation rates between selfers and outcrossers.

Currently existing theories predict that because deleterious mutations accumulate at a higher rate, selfing populations suffer from more intense genetic degradation relative to outcrossing populations. This prediction may not always be true when we consider a potential difference in deleterious mutation rate between selfers and outcrossers. By analyzing the evolutionary stability of selfing and outcrossing in an infinite population, we found that the genome-wide deleterious mutation rate would be lower in selfing than in outcrossing organisms. When this difference in mutation rate was included in simulations, we found that in a small population, mutations accumulated more slowly under selfing rather than outcrossing. This result suggests that under frequent and intense bottlenecks, a selfing population may have a lower risk of genetic extinction than an outcrossing population.

Leaf-lamina conductance contributes to an equal distribution of water delivery in current-year shoots of kudzu-vine shoot, Pueraria lobata.

Leaf-lamina resistance, R(L), accounts for a large fraction of branch resistance across a wide range of plant species. This work hypothesized that large R(L) is essential for distributing water equally to leaves on the shoot, and tested this hypothesis through theoretical analyses and measurements using over 10-m-long current-year shoots of kudzu vine, Pueraria lobata [Willd.] Ohwi. First, the hydraulic architecture and the distribution of the motive force achieving equal distribution of water delivery were theoretically obtained by simulating water flow through a hypothetical shoot comprising an axial pathway and several lateral pathways as a stem and leaves, respectively, in a kudzu-vine shoot. The model predicts that large resistance of the lateral pathway relative to that of the axial pathway is associated strongly with small variation in the hydraulic conductance of a pathway from the base of the axial pathways to the lateral pathway among the nodes, rendering water delivery to each lateral pathway equal under small variation in motive force for water flow. For the kudzu-vine shoot, the measured ratio of the lateral (a petiole) to the axial (a stem) resistance was 115. When R(L) was added to the lateral pathway, the ratio increased to 1136. According to the model prediction, these values imply that the hydraulic conductance of a pathway comprising a stem and a petiole, K(BP), is favored strongly at the basal nodes, while the hydraulic conductance of a pathway including a stem, a petiole and a lamina, K(SL), is slightly different across the nodes. For the shoots with leaf lamina, the diurnal change in transpiration rate was not different between the leaves on the three nodes dividing the shoot into four parts. K(SL) was not related significantly to node number. Conversely, K(BP) at the distal node was ~0.06-fold that at the basal node. Furthermore, the motive force for water flow should vary by 6.64-fold among nodes to compensate for the favored distribution of K(BP), which is an unrealistic value. These results indicate that R(L) contributes largely to an equal distribution of water delivery in a shoot, supporting our hypothesis.

Optimal leaf-to-root ratio and leaf nitrogen content determined by light and nitrogen availabilities.

Plants exhibit higher leaf-to-root ratios (L/R) and lower leaf nitrogen content (N(area)) in low-light than in high-light environments, but an ecological significance of this trait has not been explained from a whole-plant perspective. This study aimed to theoretically and experimentally demonstrate whether these observed L/R and N(area) are explained as optimal biomass allocation that maximize whole-plant relative growth rate (RGR). We developed a model which predicts optimal L/R and N(area) in response to nitrogen and light availability. In the model, net assimilation rate (NAR) was determined by light-photosynthesis curve, light availability measured during experiments, and leaf temperature affecting the photosynthesis and leaf dark respiration rate in high and low-light environments. Two pioneer trees, Morus bombycis and Acer buergerianum, were grown in various light and nitrogen availabilities in an experimental garden and used for parameterizing and testing the model predictions. They were grouped into four treatment groups (relative photosynthetic photon flux density, RPPFD 100% or 10%×nitrogen-rich or nitrogen-poor conditions) and grown in an experimental garden for 60 to 100 days. The model predicted that optimal L/R is higher and N(area) is lower in low-light than high-light environments when compared in the same soil nitrogen availability. Observed L/R and N(area) of the two pioneer trees were close to the predicted optimums. From the model predictions and pot experiments, we conclude that the pioneer trees, M. bombycis and A. buergerianum, regulated L/R and N(area) to maximize RGR in response to nitrogen and light availability.

Biradical character of linear π-conjugated oligomer dications composed of thiophene, pyrrole, and methylthio end-capping units.

Combined DFT calculations and UV-vis-NIR, ESR, and SQUID measurements revealed that the ground-state electronic structure of a linear π-conjugated oligomer dication composed of two pyrrole and six or seven thiophene rings and methylthio end-capping units is dominated by a singlet biradical character.

Correlation between relative growth rate and specific leaf area requires associations of specific leaf area with nitrogen absorption rate of roots.

Close correlations between specific leaf area (SLA) and relative growth rate (RGR) have been reported in many studies. However, theoretically, SLA by itself has small net positive effect on RGR because any increase in SLA inevitably causes a decrease in area-based leaf nitrogen concentration (LNCa), another RGR component. It was hypothesized that, for a correlation between SLA and RGR, SLA needs to be associated with specific nitrogen absorption rate of roots (SAR), which counteracts the negative effect of SLA on LNCa. Five trees and six herbs were grown under optimal conditions and relationships between SAR and RGR components were analyzed using a model based on balanced growth hypothesis. SLA varied 1.9-fold between species. Simulations predicted that, if SAR is not associated with SLA, this variation in SLA would cause a47% decrease in LNCa along the SLA gradient, leading to a marginal net positive effect on RGR. In reality, SAR was positively related to SLA, showing a 3.9-fold variation, which largely compensated for the negative effect of SLA on LNCa. Consequently, LNCa values were almost constant across species and a positive SLA-RGR relationship was achieved. These results highlight the importance of leaf-root interactions in understanding interspecific differences in RGR.

Applicability and limitations of optimal biomass allocation models: a test of two species from fertile and infertile habitats.

BACKGROUND AND AIMS: The practical applicability of optimal biomass allocation models is not clear. Plants may have constraints in the plasticity of their root : leaf ratio that prevent them from regulating their root : leaf ratio in the optimal manner predicted by the models. The aim of this study was to examine the applicability and limitations of optimal biomass allocation models and to test the assumption that regulation of the root : leaf ratio enables maximization of the relative growth rate (RGR). METHODS: Polygonum cuspidatum from an infertile habitat and Chenopodium album from a fertile habitat were grown under a range of nitrogen availabilities. The biomass allocation, leaf nitrogen concentration (LNC), RGR, net assimilation rate (NAR), and leaf area ratio (LAR) of each species were compared with optimal values determined using an optimal biomass allocation model. KEY RESULTS: The root : leaf ratio of C. album was smaller than the optimal ratio in the low-nitrogen treatment, while it was almost optimal in the high-nitrogen treatment. In contrast, the root : leaf ratio of P. cuspidatum was close to the optimum under both high- and low-nitrogen conditions. Owing to the optimal regulation of the root : leaf ratio, C. album in the high-nitrogen treatment and P. cuspidatum in both treatments had LNC and RGR (with its two components, NAR and LAR) close to their optima. However, in the low-nitrogen treatment, the suboptimal root : leaf ratio of C. album led to a smaller LNC than the optimum, which in turn resulted in a smaller NAR than the optimum and RGR than the theoretical maximum RGR. CONCLUSIONS: The applicability of optimal biomass allocation models is fairly high, although constraints in the plasticity of biomass allocation could prevent optimal regulation of the root : leaf ratio in some species. The assumption that regulation of the root : leaf ratio enables maximization of RGR was supported.

Hydraulic conductivity, photosynthesis and leaf water balance in six evergreen woody species from fall to winter.

To confirm that freeze-thaw embolism is a primary stress for evergreen woody species in winter, hydraulic conductivity, photosynthesis and leaf water potential were measured during fall and winter in trees growing in a cool temperate zone (Nikko) and in a warm temperate zone (Tokyo). We examined two evergreen conifers that naturally occur in the cool temperate zone (Abies firma Siebold & Zucc. and Abies homolepis Siebold & Zucc.), and four evergreen broad-leaved woody species that are restricted to the warm temperate zone (Camellia japonica L., Cinnamomum camphora (L.) J. Presl, Ilex crenata Thunb. and Quercus myrsinaefolia Blume). In Tokyo, where no freeze-thaw cycles of xylem sap occurred, hydraulic conductivity, photosynthesis and water balance remained constant during the experimental period. In Nikko, where there were 38 daily freeze-thaw cycles by February, neither of the tracheid-bearing evergreen conifers showed xylem embolism or leaf water deficits. Similarly, the broad-leaved evergreen trees with small-diameter vessels did not exhibit severe embolism or water deficits and maintained CO(2) assimilation even in January. In contrast, the two broad-leaved evergreen trees with large-diameter vessels showed significantly reduced hydraulic conductivity and shoot die-back in winter. We conclude that freeze-thaw embolism restricts evergreen woody species with large-diameter vessels to the warm temperate zone, whereas other stresses limit the distribution of broad-leaved trees, that have small-diameter vessels, but which are restricted to the warm temperate zone.

The criteria for biomass partitioning of the current shoot: water transport versus mechanical support.

In this study, we determine the theoretical criteria for biomass partitioning into the leaf and stem of the current shoot, using two quantitative models. The water transport model, based on the biochemical model of CO(2) assimilation, predicts the relationship between the water transport capacity per biomass investment in the stem (stem mass specific conductivity) and the partitioning of biomass that maximizes shoot productivity. The mechanical support model, based on Euler's buckling formula, predicts the relationship between the mechanical strength per biomass investment in the stem (the inverse relationship of stem mass density) and the partitioning of biomass to avoid mechanical failures such as lodging. These models predict the stem properties of mass specific conductivity and stem mass density that result in optimum partitioning just sufficient to provide adequate water transport and static mechanical support. In reality, the stem properties of plants differ from those predicted for optimum partitioning: the partitioning of biomass in the current shoot of both angiosperms and gymnosperms is mainly governed by the mechanical support criterion, although gymnosperms are probably more affected by the water transport criterion. This tendency is supported by actual measurements of biomass partitioning in plants.

Benefit to N2-fixing alder of extending growth period at the cost of leaf nitrogen loss without resorption.

This study examines the adaptive role of not resorbing N in N(2)-fixing deciduous trees in terms of their energy balance. The autumnal growth of N(2)-fixing Alnus firma Sieb. et Zucc. (alder) was compared with that of the non-N(2)-fixing Morus bombycis Koizumi (mulberry), which resorbs leaf N. The freezing resistance of leaves of both species was -2 degrees C. Mulberry seedlings lost their photosynthetic ability in mid-October, although the minimum temperature was still above 0 degrees C. Thereafter, their leaves turned yellow and were gradually shed. In contrast, seedlings of the alder maintained their photosynthetic ability until mid-November, when the minimum temperature fell to the freezing resistance limit. Thereafter, their leaves were shed quickly without an autumn tint. The mulberry resorbed 48.9% of leaf N, whereas the alder resorbed hardly any. These results show that, compared with the mulberry tree, the alder extended its growth period for 1 month in return for losing leaf N without resorption. The amount of energy assimilated by the alder in the extended growth period was about six times that required for compensating for the nitrogen loss, if the compensation is dependent only on the tree's own nitrogen fixation. This surplus energy balance has probably allowed N(2)-fixing deciduous trees to evolve their non-N-resorbing trait.