Current-year Seedlings Research Articles

Effects of root-colonizing fungi on pioneer Pinus thunbergii seedlings in primary successional volcanic mudflow on Kuchinoerabu Island, Japan.

Root-colonizing fungi, such as mycorrhizal fungi and dark septate endophyte fungi, are often found on pioneer plant species during early primary succession. However, little is known about which fungal species are responsible for the establishment of pioneer plants when these symbionts colonize simultaneously. We investigated the root-colonizing fungal communities of Pinus thunbergii that established prior to lichens, bryophytes, and short-lived herbaceous plants in a primary successional volcanic mudflow site on Kuchinoerabu Island, Japan. We collected a total of 54 current-year and 1- to 2-year-old seedlings. The colonization of root fungi was evaluated by direct observation of key structures (e.g., mantle, arbuscule, microsclerotia, and hyphae) and molecular analysis. Of the 34 current-year seedlings collected, only 12 individuals were colonized by ectomycorrhizal (ECM) fungi. By contrast, all 1- to 2-year-old seedlings were colonized by ECM fungi. Seedlings colonized by pine-specific ECM fungi, specifically Rhizopogon roseolus and Suillus granulatus, showed higher nitrogen and phosphorus contents in their needles compared to non-ECM seedlings. Arbuscular mycorrhizal fungi and dark septate endophyte fungi were found in only two and three individuals, respectively. The high density of mycophagous deer on Kuchinoerabu-jima may contribute to the favored dispersal of ECM fungi over other root-colonizing fungi. In conclusion, the seedling establishment of P. thunbergii at the volcanic mudflow may be largely supported by ECM fungi, with negligible effects of arbuscular mycorrhizal fungi and dark septate endophytes.

Heteroblastic Foliage Affects the Accumulation of Non-Structural Carbohydrates and Biomass in Pinus massoniana (Lamb.) Seedlings

Pines have heteroblastic foliage (primary and secondary needles) during seedling stage, but how heteroblastic foliage affects carbon storage and biomass accumulation, contributing to seedling quality, is unclear. We investigated the influences of heteroblastic foliage on photosynthetic physiological characteristics, non-structural carbohydrate (NSC) and biomass accumulation in current-year seedlings; the key factors determining biomass accumulation were mainly determined by principal component screening, Spearman correlation, and path analysis. The results indicated that (1) primary needles have high photosynthetic pigments (chlorophyll a and total chlorophyll), net photosynthetic rates (Pn), the potential maximum photochemical efficiency (Fv/Fm), and leaf instantaneous water use efficiency (WUEi), whereas higher non-photochemical quenching (NPQ) suggested that sudden light increases induce the initiation of quenching mechanism in primary needles; additionally, secondary needles had a lower transpiration rate (Tr), limiting stomata (Ls), and light saturation point. (2) Secondary needles promoted soluble sugar (fructose and glucose) increases in leaves compared to that of primary needles and increased the leaf biomass accumulation (from 47.06% to 54.30%), enhancing the overall ability of photosynthetic organs; additionally, secondary needles can enhance the proportion of starch storage in the roots, and NSC accumulation was significantly increasing in the seedling leaves and roots. (3) Photosynthetic pigments (carotenoids, chlorophyll a, and total chlorophyll) had direct positive effects on primary needle seedling (PNS) biomass and promoted biomass by indirectly increasing soluble sugar synthesis in the stems. The Pn was the main physiological factor determining PNS biomass accumulation. In addition, the WUEi, Ls, and NPQ had direct negative effects on PNS biomass accumulation, inhibiting photosynthesis to limit seedling growth. Considering the functional traits in heteroblastic foliage is necessary when assessing different leaf types of Pinus massoniana (Lamb.) seedlings, in particular those threats implicated in light, water, and temperature relations. Our results can be beneficial to guide the establishment of seedling management and afforestation measures.

Early establishment of spruce (Picea glehnii [Fr. Schm.] Masters) seedlings on disturbed soil with the aim of assisted natural regeneration

ABSTRACT To examine the potential for natural regeneration of Picea glehnii, seeds and current-year seedlings were evaluated at a scarification site. Three surface-soil treatments of different scarification intensity were used: removed, organic soil was mostly removed from the site; remained, some soil remained at the site; and replaced, the soil was removed but replaced. Seeds were sown in the fall, and samples were collected in the following spring to examine the rot rate by fungi. Germination, mortality, growth, and mycorrhiza formation rate of the seedlings were recorded in the first growing season. The seed rot rate was generally low, and no significant differences among surface-soil treatments were observed. The germination rate was higher in the removed treatment than in the other treatments, but seedling survival was lower, and the final seedling density was similar to that in the replaced treatment. Seedling total weight and ectomycorrhizal formation rate were higher in the replaced treatment than in the removed treatment, suggesting that the higher amount of organic soil at the site led to better growth at least in the first growing season. Correlation analysis indicated that a high soil moisture content associated with the presence of organic soil contributed to survival and growth.

Temporal variation in regeneration events affecting population structure in different size- and life-stages contributes to overall genetic diversity of natural Zelkova serrata population

ABSTRACT To clarify the mechanisms underlying the maintenance of genetic diversity in long-lived forest tree populations, it is important to understand the influence of temporal variation in regeneration sites and its effect on population structure (i.e., nonrandom distribution of trees in spatial location and size) and the spatial genetic structure. Features of population structure in four different life stages (current-year seedlings, saplings, juveniles, and adults) were investigated in a natural population of Zelkova serrata. The genetic diversity and genetic structure of these life stages were evaluated using eight microsatellite markers, and the regeneration processes required for maintaining the genetic diversity of the adult tree population were inferred. Within a study plot, differences in spatial distribution among life stages and size classes, and significantly positive spatial autocorrelation of tree sizes (Km(r) statistic) at the adult stage were observed. Analysis of standardized measures of genetic differentiation showed a clear genetic difference among life stages (F’ST = 0.095), with the adult stage exhibiting the highest allelic richness, the highest estimated effective number of contributing parents (N eb), and the highest estimator of spatial genetic structure (Sp statistic). The adult stage may comprise many different cohorts (subpopulations), contributed by different compositions of parents, due to spatially and temporarily distinct regeneration events, and the resulting genetic heterogeneity would contribute to an increase in genetic diversity within the population.

Survival rate and environmental response of current-year seedlings of the temperate liana Wisteria floribunda across a heterogeneous environment.

Lianas have a huge influence on forest structure and function. However, it is unclear how the surrounding environment affects the establishment of liana seedlings in temperate forests. We addressed the following questions: (1) Can current-year seedlings persist under a closed canopy? (2) Do current-year seedlings form aggregated distribution and how has their spatial distribution varied over the years? (3) How does the light condition, soil moisture content, forest floor litter, understory vegetation, and the distance from the conspecific adults affect the establishment and survival of seedlings? We examined the distribution pattern and survivorship of current-year seedlings of the temperate liana species, Wisteria floribunda, across a heterogeneous environment for 6years using 1m2 sub-quadrats (n = 651) in a 6ha plot within the Ogawa Forest Reserve, an old-growth, temperate, deciduous forest in central Japan. In total, 908 current-year seedlings were observed during the study period, 87% of which emerged in 2014. Over half (56%) of these seedlings survived until 1year after germination, which was relatively high compared with other tree species in this forest. The seedlings formed significantly aggregated distribution, but the degree of aggregation decreased over time. The number of emerged seedlings was negatively associated with the presence of dwarf bamboo (Sasa borealis) and the distance from the nearest conspecific adult. However, the survival rate of the seedlings was negatively associated with the presence of dwarf bamboo and soil moisture content and was positively associated with the openness of the canopy and the distance from the nearest conspecific adult. An enhanced survival rate under more intense light conditions and the ability to persist within the shaded understory may be important for the survival of this species in the earlier stage of the life history.

Effects of drought on leaf growth and chlorophyll fluorescence kinetics parameters in Cyclobalanopsis glauca seedlings of Karst areas

To uncover adaptation mechanism of Cyclobalanopsis glauca to the arid environment of Karst areas, current-year seedlings of C. glauca were potted and grown under four soil water conditions: Normal water supply (-0.1 MPa), light drought stress (-0.5 MPa), moderate drought stress (-0.9 MPa), and severe drought stress (-1.5 MPa). We measured leaf growth and parameters of fast chlorophyll fluorescence induction dynamics after treated by 15, 30, 45, 60 and 90 days. With the increase of drought stress intensity, leaf area, number of green leaves, leaf water content, the contents of chlorophyll a+b and carotenoids, the maximum fluorescence, maximum photochemical quantum yield and potential photochemical efficiency significantly decreased, while the number of dead leaves and the initial fluorescence significantly increased. There was no significant difference in these parameters between light drought and normal water treatments. There were no significant difference in the absorption flux per reaction center (ABS/RC), captured light energy used to restore qA (TRo/RC), number of active reaction centers per unit area (RC/CS), light energy captured per unit area (TRo/CS) and light energy used for electron transfer per unit area (ETo/CS) between light drought and normal water treatments. Among these parameters, RC/CS was slightly higher under light drought treatment than that of normal water treatment. TRo/CS and ETo/CS reached peaks at the 45th day, being 606.12 and 440.78, respectively. Leaf ABS/RC, TRo/RC, ETo/RC, DIRo/RC, RC/CS, TRo/CS and ETo/CS of C. glauca seedlings under mode-rate drought and severe drought treatment were lower than those of normal water treatment, and the parameters under severe drought stress decreased more significantly with the extension of drought stress time. With increasing intensity and duration of drought stress, the maximum quantum efficiency, probability of other electron acceptors, and quantum ratio of electron transfer decreased, but quantum yield for energy dissipation increased. These results demonstrated that C. glauca seedlings under light drought condition showed some degree of adaptability and resistance to drought. Mode-rate drought treatment caused a decline in chlorophyll fluorescence and photosynthetic pigments, leading to slow growth of seedlings. Severe drought had a serious impact on growth of C. glauca seedlings, but did not lead to seedling death. Therefore, C. glauca seedlings showed strong drought tolerance, which were suitable for the application of vegetation restoration and reforestation projects in Karst areas.

Ectomycorrhizal fungal communities in alpine relict forests of Pinus pumila on Mt. Norikura, Japan.

Ectomycorrhizal (ECM) symbioses are indispensable for the establishment of host trees, yet available information of ECM symbiosis in alpine forests is scarce. Pinus pumila is a typical ice age relict tree species in Japan and often forms monodominant dwarf vegetation above the tree line in mountains. We studied ECM fungi colonizing P. pumila on Mt. Norikura, Japan, with reference to host developmental stages, i.e., from current-year seedlings to mature trees. ECM fungal species were identified based on rDNA ITS sequences. Ninety-two ECM fungal species were confirmed from a total of 2480 root tips examined. Species in /suillus-rhizopogon and /wilcoxina were dominant in seedling roots. ECM fungal diversity increased with host development, due to the addition of species-rich fungal lineages (/cenococcum, /cortinarius, and /russula-lactarius) in late-successional stages. Such successional pattern of ECM fungi is similar to those in temperate pine systems, suggesting the predominant role of /suillus-rhizopogon in seedling establishment, even in relict alpine habitats fragmented and isolated for a geological time period. Most of the ECM fungi detected were also recorded in Europe or North America, indicating their potential Holarctic distribution and the possibility of their comigration with P. pumila through land bridges during ice ages. In addition, we found significant effects of soil properties on ECM fungal communities, which explained 34.1% of the total variation of the fungal communities. While alpine vegetation is regarded as vulnerable to the ongoing global warming, ECM fungal communities associated with P. pumila could be altered by the edaphic change induced by the warming.

クロマツコンテナ苗の当年生苗利用と通年植栽の可能性

現在東北地方の津波被害地では広範囲の海岸林の早急な再生のため, 大量のクロマツ苗の植栽が必要になっており, 育苗期間の短縮と, 作業の平準化のための通年植栽が求められている。そこで, 当年生苗の利用可能性と通年植栽の可能性を検討するため, 宮城県仙台市において, 当年生コンテナ苗と 2 年生裸苗の成長比較試験と, コンテナ苗を厳冬期の1 月を除き, 2 カ月おきに通年で植栽する試験を行った。その結果, 11 月下旬と 3 月下旬植栽の当年生コンテナ苗の成長後の幹長と地際直径は 2 年生裸苗と有意差がなかった。幹長の成長量は, 3 月植栽で裸苗よりやや劣ったものの, 11 月植栽のコンテナ苗では同等で あった。地際直径の成長量は, コンテナ苗の方が有意に多かった。コンテナ苗は, いずれの植栽時期でも越冬後の活着率が98.4%以上と高く, 成長に大きな問題もみられなかった。以上のことから, 当年生苗の利用と通年植栽は, ともに可能と考えられた。

Changes in morphological, physiological, and biochemical responses to different levels of drought stress in chinese cork oak (Quercus variabilis Bl.) seedlings

Changes in growth, leaf water status, pigments, osmolytes, activities of peroxidase (POD), catalase (CAT), superoxide dismutase (SOD), and ascorbate peroxidase (APX), and ascorbic acid (ASA) content were investigated in Chinese cork oak (Quercus variabilis Bl.) seedlings. Three-month-old seedlings were subjected to four drought cycles (30, 60, 90, and 120 days) and four drought intensities (80, 60, 40, and 20% field capacity (FC)). The seedlings had optimal height, basal diameter, and leaf water status at 80% FC. These parameters significantly decreased as drought intensity increased. The total root length, diameter, and surface area at 60% FC significantly increased compared with those at 80% FC. However, at 40 and 20% FC these parameters significantly decreased compared with those at 80% FC. The ratio of total root length to seedling height significantly increased with increasing drought intensity. The contents of chlorophyll a + b (Chla + b) and carotenoids (Car) significantly decreased at 40 and 20% FC. However, no significant changes in Chla/Chlb and Car/Chla + b ratios were observed among the four drought intensities. Comparatively, the seedlings accumulated more soluble sugars and proline, as well as they demonstrated the higher POD, SOD, CAT, APX activities and ASA content at >40% FC. However, prolonged drought stress at 20% FC suppressed antioxidant activities and osmolyte accumulation, leading to a rapid increase in lipid peroxidation. These results suggest that a water supply >40% FC is required to support the growth and survival of the current-year seedlings of Chinese cork oak

Effects of complete submergence on seedling growth and survival of five riparian tree species in the warm-temperate regions of Japan

To reveal the relationship between tolerance to submergence and the distribution ranges of riparian tree species in the floodplains of the warm-temperate regions in Japan, we assessed the response and tolerance of current-year seedlings of two Salix and three Ulmaceae species to submergence treatments for 1, 2, 4, and 8 weeks. We investigated shoot elongation, number of leaves, and survival rate of seedlings during the experiment, and biomass allocation was determined at the end of the experiment. Shoot elongation and leaf production of seedlings were suppressed in all the species during the submergence treatments. However, the seedlings of Salix chaenomeloides and S. pierotii elongated shoots rapidly and produced leaves after the treatments. All the underground biomass in these two species was larger than aboveground biomass. These results are considered as typical responses of flood-tolerant species. In contrast, the underground biomass of Ulmaceae seedlings (Celtis sinensis var. japonica, Aphananthe aspera, and Ulmus parvifolia) decreased more rapidly than aboveground biomass during prolonged submergence. Furthermore, A. aspera seedlings also showed suppression of shoot elongation and leaf production after the treatments. Our results indicate that the three Ulmaceae species are less tolerant to submergence than the two Salix species. In conclusion, we suggest that submergence stress is one of the important factors determining the distribution of tree species in the floodplains of lowlands in the warm-temperate regions of Japan.

Field performance of Scots pine (Pinus sylvestris L.) seedlings planted in disc trenched or mounded sites over an extended planting season

Economic pressures have driven an ever-widening period during which foresters use machines to plant Scots pine (Pinus sylvestris L.) seedlings. In Fennoscandia, this period has recently stretched to the entire growing season. To evaluate the performance of seedlings planted during this extended period, three experiments were carried out in Central and Northern Finland over 2 years. One-year-old and current-year seedlings were planted in mounds or disc-trenched furrows when soil temperatures were >0°C. When 1-year-old seedlings grown for spring planting and overwintered outdoors were planted after mid-June, more needles browned and growth was reduced, possibly because seedlings were oversized with respect to planting density and the volume of growth media. When current-year seedlings sown in spring were planted from July to November, those planted in late September and October grew less in later years than those planted earlier, but survival was unaffected. No large differences in field performance were found with respect to whether seedlings were planted in mounds or disc-trenched furrows. In conclusion, Scots pine seedlings can be machine planted in mounds or furrows during May and early June (later in the North) and then continued from early August until late September, provided climatic conditions in late spring and early autumn are typical and similar to those experienced in Central Finland.

Leaf water relations of current-year seedlings of Aesculus turbinata

Leaf water relations of current-year seedlings of <i>Aesculus turbinata</i>

Ectomycorrhizal fungal community of naturally regenerated Pinus thunbergii seedlings in a coastal pine forest

To add to knowledge of ectomycorrhizal (ECM) community structure on the roots of Pinus thunbergii seedlings in a Japanese coastal forest, we sampled naturally regenerated current-year and 1- to 5-year-old seedlings. We classified the 667 root tips on current-year seedlings and the 1,927 root tips on older seedlings into 13 phylotypes based on morphological and genetic analyses. Cenococcum geophilum, members of the families Clavulinaceae, Russulaceae, or Thelephoraceae or the genus Trichophaea, were indicated to be fungi forming P. thunbergii ectomycorrhizas. Among them, C. geophilum and Clavulinaceae sp. 1 were the most or second most dominant species. A species accumulation curve based on the number of samples nearly reached a plateau, with observed species richness equal to 11 species and the Jackknife2 and Chao2 richness estimators indicating 14 and 12 species, respectively. In addition, Simpson's 1/D was 3.89 and Shannon–Wiener's H′ was 1.71, indicating a relatively low taxonomic diversity. There was no significant difference in the ectomycorrhizal formation rate, or the occurrence frequency and the number of ECM phylotypes between current-year and older seedlings. These results indicated that less diverse fungi were involved in ectomycorrhizal formation on coastal pine seedlings compared with those in comparable inland forests in the study area.

Effect of Light Conditions on the Resistance of Current-year Fagus Crenata Seedlings Against Fungal Pathogens Causing Damping-off in a Natural Beech Forest: Fungus Isolation and Histological and Chemical Resistance

Forest gap dynamics affects light intensity on the forest floor, which in turn may influence defense and survival of tree seedlings. Current-year Fagus crenata seedlings show high mortality under the canopy caused by damping-off. In contrast, they survive pathogen attacks in gaps. However, defense mechanisms against damping-off have not been fully understood. In order to determine the resistance factors that affect mortality in current-year seedlings, we compared seedling survival and chemical and histological characteristics of the hypocotyls of seedlings from closed-stand and forest-edge plots. Damping-off occurred in the current-year seedlings mainly from the end of June to July; survival rate of the seedlings was higher in the forest-edge plot than in the closed-stand plot. By performing an inoculation test on the seedling hypocotyls, we identified Colletotrichum dematium and Cylindrocarpon sp. as the causative pathogens under low illumination only. In the beginning of July, only seedling hypocotyls from the forest-edge plot exhibited periderm formation. From mid-June to July, seedling hypocotyls from the forest-edge plot accumulated approximately twice the amount of total phenols as those accumulated by seedling hypocotyls from the closed-stand plot. The ethyl acetate phase of methanol extracts of hypocotyls showed antifungal activity. We conclude that seedlings from the forest-edge plot may resist pathogenic attack via periderm formation and increased phenol synthesis. Plant defense mechanisms that are controlled by light intensity may be important for promoting seedling regeneration in forest gap dynamics.

The soil seed bank of the threatened plant Magnolia stellata is subordinate to the emergence of current-year seedlings

We studied seed bank formation of the threatened star magnolia, Magnolia stellata, to examine the early stage of regeneration. Forty-five seedling plots (2 × 2 m), each including a soil-sampling quadrat (40 × 40 cm), were established randomly under or around the crowns of mature M. stellata trees. Seeds of M. stellata were collected from each quadrat to a depth of 5 cm. Only four seeds of M. stellata were found (0.56 seeds/m2) and all were located under mature crowns. Current-year seedlings were abundant in water channels, on moss, or under mature crowns, suggesting that the seeds may require wet soil conditions for germination. Magnolia stellata seeds show considerable germination below the crowns of mature trees in the year following masting, while some seeds remain dormant in the soil. Considering the soil seed bank and the current-year seedling bank of M. stellata, a frequent supply of seed is essential for the regeneration of this species. Thus, it is important to maintain mature trees in addition to promoting seed production.

Croissance par rapport au stockage : réponses de semis de chênes méditerranéens aux changements de nutrition et de disponibilité en eau

We compare dry mass (DM) and storage of starch (St) and nitrogen (N) in seedlings of three Mediterranean oaks, two evergreens (Quercus coccifera L. and Q. ilex L. subsp. ballota (Desf.) Samp) and one deciduous (Q. faginea Lam.), across different scenarios of nutrient and water availabil- ities. Three fertilization (5, 50 and 200 mg of N per plant and growing period) and watering (28-39, 55-71 and 70-85 g H2O 100 g −1 soil gravimetric soil water) treatments were applied to current-year seedlings between May and October 2002 in two independent experiments. The three species showed a similar response to fertilization, storing nitrogen instead of increasing biomass, in agreement with adaptations to nutrient-poor habitats. However, they differed in their responses to water, reflecting the different water requirements in the field: Q. coccifera, from arid zones, showed no response to water regarding DM and St; Q. faginea, from humid zones, required higher water availability to simultaneously increase growth and storage; while Q. ilex, spanning over most of the water availability range, exhibited a balanced increase of both functions when water increased moderately. In the two evergreen species, N concentration increased with water supply, whereas the reverse occurred in Q. faginea. The latter species favoured growth over storage at moderate water supply (according to its more competitive strategy), although it was the species which accumulated more St and N at the end of the experiments (autumn).

Fat‐tailed gene flow in the dioecious canopy tree species Fraxinus mandshurica var. japonica revealed by microsatellites

Pollen flow, seed dispersal and individual reproductive success can be simultaneously estimated from the genotypes of adults and offspring using stochastic models. Using four polymorphic microsatellite loci, gene flow of the wind-pollinated and wind-seed-dispersed dioecious tree species, Fraxinus mandshurica var. japonica, was quantified in a riparian forest, in northern Japan. In a 10.5-ha plot, 74 female adults, 76 male adults and 292 current-year seedlings were mapped and genotyped, together with 200 seeds. To estimate dispersal kernels of pollen and seeds, we applied normal, exponential power, Weibull, bivariate t-distribution kernels, and two-component models consisting of two normal distribution functions, one with a small and one with a large variance. A two-component pollen flow model with a small contribution (26.1%) from short-distance dispersal (sigma = 7.2 m), and the rest from long-distance flow (sigma = 209.9 m), was chosen for the best-fitting model. The average distance that integrated pollen flows inside and outside the study plot was estimated to be 196.8 m. Tree size and flowering intensity affected reproduction, and there appeared to be critical values that distinguished reproductively successful and unsuccessful adults. In contrast, the gene flow model that estimated both pollen and seed dispersal from established seedlings resulted in extensive seed dispersal, and the expected spatial genetic structures did not satisfactorily fit with the observations, even for the selected model. Our results advanced small-scale individual-based parentage analysis for quantifying fat-tailed gene flow in wind-mediated species, but also clarified its limitations and suggested future possibilities for gene flow studies.

Ectomycorrhizal networks and seedling establishment during early primary succession

Ectomycorrhizal (ECM) fungal mycelia are the main organs for nutrient uptake in many woody plants, and often connect seedlings to mature trees. While it is known that resources are shared among connected plants via common mycorrhizal networks (CMNs), the net effects of CMNs on seedling performance in the field are almost unknown. CMNs of individual ECM fungal species were produced in an early succession volcanic desert by transplanting current-year seedlings of Salix reinii with ECM mother trees that had been inoculated with one of 11 dominant ECM fungal species. Most seedlings were connected to individual CMNs without being infected by other ECM fungi. Although control seedlings showed poor growth under severe nutrient competition with larger nonmycorrhizal mother trees, nutrient acquisition and growth of seedlings connected to CMNs were improved with most fungal species. The positive effects of CMNs on seedling performance were significantly different among ECM fungal species; for example, the maximum difference in seedling nitrogen acquisition was 1 : 5.9. The net effects of individual CMNs in the field and interspecific variation among ECM fungal species are shown.

Season and substrate effects on the first-year establishment of current-year seedlings of major conifer species in an old-growth subalpine forest in central Japan

Seasonal and substrate effects on first-year seedling establishment of major conifer species ( Abies mariesii, Abies veitchii, Picea jezoensis var. hondoensis, and Tsuga diversifolia) were assessed in an old-growth subalpine forest in central Japan. Older seedlings (≥1 year) were more abundant on woody debris (WD), such as fallen logs, than on soil. It is known that this preferential seedling establishment on WD is determined by the first-year seedling establishment. The present results indicate that first-summer seedling survivorship clearly contributes to the creation of such seedling–substrate associations, because mortality during the first-summer was greater on soil than WD but that during the first-autumn/winter was not different. Although the standardized mortality rate of current-year seedlings on WD was not so different among the three seasons, that on soil significantly decreased in the winter; indicating that soil is not an unsuitable substrate for seedling survivorship during snow-covered winter. Thus, it can be concluded that biotic and abiotic factors enhancing seedling mortality on soil are most active during summer, and seedling–substrate associations seem to be determined mainly during the first-summer soon after seedling emergence. Furthermore, the first-year survivorship of P. jezoensis var. hondoensis, which has smaller-sized seeds/seedlings that are disadvantageous for early survival, decreased with progressing WD decay toward soil. This suggests that the properties of WD also affect current-year seedling survivorship, especially for species sensitive to substrate properties. In conclusion, initial survivorship affects seedling bank dynamics, and such early dynamics are greatly regulated by the highly heterogeneous substrate availability on forest floors.

Biomass allocation and leaf chemical defence in defoliated seedlings of Quercus serrata with respect to carbon-nitrogen balance.

Both nutrient availability and defoliation affect the carbon-nutrient balance in plants, which in turn influences biomass allocation (e.g. shoot-to-root ratio) and leaf chemical composition (concentration of nitrogen and secondary compounds). In this study it is questioned whether defoliation alters biomass allocation and chemical defence in a similar fashion to the response to nutrient deficiency. Current-year seedlings of Quercus serrata were grown with or without removal of all leaves at three levels of nutrient availability. Plant nitrogen concentration (PNC), a measure of the carbon-nutrient balance in the plant, significantly decreased immediately after defoliation because leaves had higher nitrogen concentrations than stems and roots. However, PNC recovered to levels similar to or higher than that of control plants in 3 or 6 weeks after the defoliation. Nitrogen concentration of leaves produced after defoliation was significantly higher than leaf nitrogen concentration of control leaves. Leaf mass per plant mass (leaf mass ratio, LMR) was positively correlated with PNC but the relationship was significantly different between defoliated and control plants. When compared at the same PNC, defoliated plants had a lower LMR. However, the ratio of the leaf to root tissues that were newly produced after defoliation as a function of PNC did not differ between defoliated and control plants. Defoliated plants had a significantly lower concentration of total phenolics and condensed tannins. Across defoliated and control plants, the leaf tannin concentration was negatively correlated with the leaf nitrogen concentration, suggesting that the amount of carbon-based defensive compounds was controlled by the carbon-nutrient balance at the leaf level. Defoliation alters biomass allocation and chemical defence through the carbon-nutrient balance at the plant and at the leaf level, respectively.