Carbon use strategies in shoot and acorn growth of two evergreen broadleaf trees unraveled by seasonal carbohydrate measurements and carbon isotope analysis.

Woody species have evolved carbon (C) storage processes that meet needs for reserves associated with asynchronies between C supply and demand. However, our understanding of storage dynamics is still elusive in mature trees, especially when reproduction is involved. Integrated analyses of isotope ratios, concentrations, and biomass may enhance understanding of stored C fractions' dynamics and roles. Thus, we monitored starch and soluble sugars (SSs), C isotope ratios, and biomass, in leaves, twigs and reproductive organs of two mature evergreen broadleaf trees, Quercus glauca and Lithocarpus edulis, for two years. During the growing season, no starch was observed in twigs, while constant starch levels were observed in leaves. Increase in SSs for winter hardening was earlier in L. edulis than in Q. glauca, in line with L. edulis acorns' earlier ripening. Decrease in SSs and increase in starch occurred simultaneously in the next spring. In addition, sucrose accounted for less than 10% of total SSs in leaves of both species, whereas mannose accounted for up to 75% in Q. glauca and myo-inositol up to 23% in L. edulis, indicating species specific sugar composition. These results indicate that seasonal variation of SSs fraction was more reflective to climatic change and NSC storage was less influenced by reproduction. No starch was detected in acorn organs of either Q. glauca or L. edulis except in ripening seeds. The biomass of ripe acorns was 1.7- and 6.4-fold greater than that of current-year twigs in Q. glauca and L. edulis, respectively. Bulk twigs and reproductive organs were ca. 1.0‰ 13C enriched relative to bulk leaves, which was lower than in deciduous trees. These results indicate that new photo-assimilate is the predominant C source for reproductive growth. These findings provide new insights into the dynamics of C storage in relation to reproduction in evergreen broadleaf trees.

Limits to reproduction and seed size-number trade-offs that shape forest dominance and future recovery.

The relationships that control seed production in trees are fundamental to understanding the evolution of forest species and their capacity to recover from increasing losses to drought, fire, and harvest. A synthesis of fecundity data from 714 species worldwide allowed us to examine hypotheses that are central to quantifying reproduction, a foundation for assessing fitness in forest trees. Four major findings emerged. First, seed production is not constrained by a strict trade-off between seed size and numbers. Instead, seed numbers vary over ten orders of magnitude, with species that invest in large seeds producing more seeds than expected from the 1:1 trade-off. Second, gymnosperms have lower seed production than angiosperms, potentially due to their extra investments in protective woody cones. Third, nutrient-demanding species, indicated by high foliar phosphorus concentrations, have low seed production. Finally, sensitivity of individual species to soil fertility varies widely, limiting the response of community seed production to fertility gradients. In combination, these findings can inform models of forest response that need to incorporate reproductive potential.

Globally, tree fecundity exceeds productivity gradients.

Lack of tree fecundity data across climatic gradients precludes the analysis of how seed supply contributes to global variation in forest regeneration and biotic interactions responsible for biodiversity. A global synthesis of raw seedproduction data shows a 250-fold increase in seed abundance from cold-dry to warm-wet climates, driven primarily by a 100-fold increase in seed production for a given tree size. The modest (threefold) increase in forest productivity across the same climate gradient cannot explain the magnitudes of these trends. The increase in seeds per tree can arise from adaptive evolution driven by intense species interactions or from the direct effects of a warm, moist climate on tree fecundity. Either way, the massive differences in seed supply ramify through food webs potentially explaining a disproportionate role for species interactions in the wet tropics.

Imperforate tracheary elements and vessels alleviate xylem tension under severe dehydration: insights from water release curves for excised twigs of three tree species.

PREMISE: Water stored in the xylem of woody plants is important for supporting the transpiration stream under prolonged drought, yet the source of stored water within the xylem during drought remains unclear. Insights into xylem water utilization during drought will uncover the adaptation strategies of the test species to stress. METHODS: To fill the existing knowledge gap, we excised twigs of Abies firma (Japanese fir, conifer), Cercidiphyllum japonicum (katsura tree, diffuse-porous) and Quercus serrata (konara oak, ring-porous) to quantify interspecific variation of water transfer in xylem corresponding with increasing cumulative water release (CWR) using micro x-ray computed tomography and cryo-SEM. RESULTS: For all species studied, the main components of water storage within the operating range of water potential were not living cells but cavitation release and capillaries. Abies firma maintained water in the earlywood-like cells, for possible maintenance of the transpiration stream. Cercidiphyllum japonicum maintained water in its vessels over 200 kg m-3 of CWR, while Q. serrata lost most of its water in vessels with increasing CWR up to 100 kg m-3 . Cercidiphyllum japonicum exhibited a higher water storage capacity than Q. serrata. Under high CWR, narrow conduits stored xylem water in C. japonicum and imperforate tracheary elements in Q. serrata. CONCLUSIONS: Among the species examined, increasing CWR appears to indicate differential utilization of stored water in relation to variation of xylem structure, thereby providing insight into the interspecific responses of tree species to drought.

Field transcriptome revealed a novel relationship between nitrate transport and flowering in Japanese beech.

Recent advances in molecular and genetic studies about flowering time control have been increasingly available to elucidate the physiological mechanism underlying masting, the intermittent and synchronized production of a large amount of flowers and seeds in plant populations. To identify unexplored developmental and physiological processes associated with masting, genome-wide transcriptome analysis is a promising tool, but such analyses have yet to be performed. We established a field transcriptome using a typical masting species, Japanese beech (Fagus crenata Blume), over two years, and analyzed the data using a nonlinear time-series analysis called convergent cross mapping. Our field transcriptome was found to undergo numerous changes depending on the status of floral induction and season. An integrated approach of high-throughput transcriptomics and causal inference was successful at detecting novel causal regulatory relationships between nitrate transport and florigen synthesis/transport in a forest tree species. The synergistic activation of nitrate transport and floral transition could be adaptive to simultaneously satisfy floral transition at the appropriate timing and the nitrogen demand needed for flower formation.

Frankia communities at revegetating sites in Mt. Ontake, Japan.

In 1984 at Mt. Ontake in Japan, an earthquake caused a devastating landslide, and as a result, the vegetation on the south slope of the mountain was completely eliminated. In higher elevation (2000 m) areas, revegetation has not yet been completed even 30 years after the landslide. Revegetation progress throughout the area was heterogeneous. In the partially revegetated areas, actinorhizal plant species such as Alnus maximowiczii and Alnus matsumurae have been found. In the present study, we investigated the Frankia communities in the higher-elevation area using sequence analysis of the amplified nifH (dinitrogenase reductase) gene from nodule and soil samples collected in the disturbed region, undisturbed forest, and in the boundary between the disturbed region and the undisturbed forest. Phylogenetic analysis of partial nifH sequences revealed the presence of six clusters, each of which consisted of highly similar (> 99%) sequences. Four clusters showed significant sequence similarity to Frankia (three Alnus- and a Casuarina-infecting strains). Diversity in the Frankia community was relatively low-only one or two clusters were detected in a site. At most of the sampling sites, a dominant cluster in a nodule coincided with that in rhizosphere soil, indicating that community structure in the rhizosphere is a primary factor that determines occupancy in a nodule. No significant difference in community structure was observed between plant species. Diversity in the Frankia community varied depending on revegetation progress. Cluster A, which was the most dominant in the disturbed region, was likely to have invaded from undisturbed forest.

Pine wilt disease causes cavitation around the resin canals and irrecoverable xylem conduit dysfunction.

Physiological mechanisms of irreversible hydraulic dysfunction in seedlings infected with pine wilt disease (PWD) are still unclear. We employed cryo-scanning electron microscopy (cryo-SEM) to investigate the temporal and spatial changes in water distribution within the xylem of the main stem of 2-year-old Japanese black pine seedlings infested by pine wood nematodes (PWNs). Our experiment was specifically designed to compare the water relations among seedlings subjected to the following water treatment and PWN combinations: (i) well-watered versus prolonged drought (no PWNs); and (ii) well-watered with PWNs versus water-stressed with PWNs (four treatments in total). Cryo-SEM imaging observations chronicled the development of patchy cavitations in the xylem tracheids of the seedlings influenced by PWD. With the progression of drought, many pit membranes of bordered pits in the xylem of the main stem were aspirated with the decrease in water potential without xylem cavitation, indicating that hydraulic segmentation may exist between tracheids. This is the first study to demonstrate conclusively that explosive and irreversible cavitations occurred around the hydraulically vulnerable resin canals with the progression of PWD. Our findings provide a more comprehensive understanding of stressors on plant-water relations that may eventually better protect trees from PWD and assist with the breeding of trees more tolerant to PWD.

Influence of reproduction on nitrogen uptake and allocation to new organs in Fagus crenata.

The contributions of the internal nitrogen (N) cycle and N uptake from soil to growth in mature trees remain poorly understood, especially during reproduction. In order to elucidate how reproduction affects N uptake, allocation and remobilization, we applied pulse 15N labelling to three fruiting (F) and three non-fruiting (NF) Fagus crenata Blume trees after the leaves were fully unfurled. Three-year-old branches were sampled from upper crowns at about 2 week intervals until leaf fall. 15N content per organ dry mass (15Nexcess) and N concentration in all new shoot organs were determined. Fruiting led to greater 15Nexcess uptake from the soil during the first month following application. Cupules absorbed the highest fraction of 15Nexcess initially and nuts contained about half the 15Nexcess at the end of the growing season. Biomass of reproductive organs represented up to 70% of new shoot growth in F trees. This fruit burden led to 34% and 38% reduction in biomass and 15Nexcess, respectively, in mature leaves compared with NF trees. Moreover, the increment of 15Nexcess in new shoots of F relative to NF trees was lower than the increment of biomass between the two. These results indicate that N is a limiting resource during masting in F. crenata. 15Nexcess incorporated into nuts started to increase dramatically once 15Nexcess in leaves, branches and cupules hit seasonal maxima. Similar seasonal biomass growth patterns were also found in these organs, indicating that sink strength drives uptake and allocation of 15Nexcess between new shoot compartments. These results, together with translocation of 15Nexcess from cupules and senescing leaves to nuts (contributing to fruit ripening), suggest that a finely tuned growth phenology alleviated N limitation. Thus, fruiting did not influence the N concentration in leaves or branches. These reproduction-related variations in N uptake and allocation among new shoot compartments have implications for N dynamics in the plant-soil system.

Growth rate reduction causes a decline in the annual incremental trunk growth in masting Fagus crenata trees.

Tree trunk annual increments are markedly reduced in mast years. There are two hypotheses that could explain the mechanism for this phenomenon: (1) a reduction in the duration of growth due to switching the resource allocation from somatic growth to seed production; (2) reduction of growth rate due to resources being shared between somatic growth and reproduction simultaneously. In this study, we aimed to test these hypotheses in Fagus crenata  Blume from the point of view of resource allocation. The radial growth patterns in F. crenata during a year without reproduction (2014) and a masting year (2015) were monitored using a digital dendrometer. At the same time, shoot growth patterns were monitored by sampling branches from the top of the canopy. Data obtained using the digital dendrometer were fitted to a sigmoidal function, and the parameters of the function were evaluated with a hierarchal Bayesian approach; estimated parameters were used to represent the properties of trunk growth phenology. Trunk growth started synchronously just after leaf unfurling in both mass-fruiting (F15) and limited-fruiting (NF15) trees in 2014 and 2015. Reproduction reduced the growth rate in 2015. This was due to the resources being allocated for the development of cupules and for formation of relatively thick branches, both of which occurred simultaneously with trunk growth. There was no clear difference in the duration of radial growth between F15 and NF15 trees in the 2 years, although seed maturation started after trunk growth ceased. As a result, the annual trunk radius increment was reduced in the F15 trees in 2015. These results suggested that reduction of radial growth rate (Hypothesis 2) caused the reduction in annual trunk increment of reproducing trees of this species.

Reproduction-related variation in carbon allocation to woody tissues in Fagus crenata using a natural 13C approach.

The contribution of new photo-assimilates and stored carbon (C) to plant growth remains poorly understood, especially during reproduction. In order to elucidate how mast seeding affects C allocation to both reproductive and vegetative tissues, we measured biomass increase in each tissue, branch starch concentration and stable C isotope composition (δ13C) in bulk leaves, current-year shoots, 3-year branches and tree rings in fruiting and non-fruiting trees for 2 years, as well as in fruits. We isolated the effect of reproduction on C allocation to vegetative growth by comparing 13C enrichment in woody tissues in fruiting and non-fruiting specimens. Compared with 2‰ 13C enrichment in shoots relative to leaves from non-fruiting trees, fruiting reduced the enrichment to 1‰ and this reduction disappeared in the following year with no fruiting, indicating that new photo-assimilates are preferentially used for woody tissues even with fruiting burden. In contrast, fruits had up to 2.5‰ 13C enrichment at mid-summer, which dropped thereafter, indicating that fruit production relies on C storage early in the growing season then shifts to current photo-assimilates. At this tipping point, growth of shoots and cupules had almost finished and nuts had a second rapid growth period thereafter. Together with shorter shoots but higher biomass increment per length in fruiting trees than non-fruiting trees, these results indicate that the C limitation due to fruit burden is minimized by fine-tuning of allocation of old C stores and new photo-assimilates, along with the growth pattern in various tissues. Furthermore, fruiting had no significant effect on starch concentration in 3-year-old branches, which became fully depleted during leaf and flower flushing but were quickly replenished. These results indicate that reproduction affects C allocation to branches but not its source or storage. These reproduction-related variations in the fate of C have implications for evaluating forest ecosystem C cycles during climate change.

Concordance and discordance between taxonomic and functional homogenization: responses of soil mite assemblages to forest conversion.

The compositional characteristics of ecological assemblages are often simplified; this process is termed "biotic homogenization." This process of biological reorganization occurs not only taxonomically but also functionally. Testing both aspects of homogenization is essential if ecosystem functioning supported by a diverse mosaic of functional traits in the landscape is concerned. Here, we aimed to infer the underlying processes of taxonomic/functional homogenization at the local scale, which is a scale that is meaningful for this research question. We recorded species of litter-dwelling oribatid mites along a gradient of forest conversion from a natural forest to a monoculture larch plantation in Japan (in total 11 stands), and collected data on the functional traits of the recorded species to quantify functional diversity. We calculated the taxonomic and functional ß-diversity, an index of biotic homogenization. We found that both the taxonomic and functional ß-diversity decreased with larch dominance (stand homogenization). After further deconstructing ß-diversity into the components of turnover and nestedness, which reflect different processes of community organization, a significant decrease in the response to larch dominance was observed only for the functional turnover. As a result, there was a steeper decline in the functional ß-diversity than the taxonomic ß-diversity. This discordance between the taxonomic and functional response suggests that species replacement occurs between species that are functionally redundant under environmental homogenization, ultimately leading to the stronger homogenization of functional diversity. The insights gained from community organization of oribatid mites suggest that the functional characteristics of local assemblages, which support the functionality of ecosystems, are of more concern in human-dominated forest landscapes.

Biotic homogenization and differentiation of soil faunal communities in the production forest landscape: taxonomic and functional perspectives.

Biotic homogenization has been reported worldwide. Although simplification of communities across space is often significant at larger scales, it could also occur at the local scale by changing biotic interactions. This study aimed to elucidate local community processes driving biotic homogenization of soil faunal communities, and the possibility of biotic re-differentiation. We recorded species of oribatid mites in litter and soil layers along a gradient of forest conversion from monoculture larch plantation to primary forests in central Japan. We collected data for functional traits of the recorded species to quantify functional diversity. Then we quantified their taxonomic/functional turnover. Litter diversity was reduced in the larch-dominated stands, leading to habitat homogenization. Consequently, litter communities were biologically homogenized and differentiated in the plantations and in the natural forest, respectively. Turnover of functional traits for litter communities was lower and higher than expected by chance in the plantations and in the natural stand, respectively. This result suggests that the dominant assembly process shifts from limiting similarity to habitat filtering along the forest restoration gradient. However, support for such niche-based explanations was not observed for communities in the soil layer. In the monocultures, functional diversity expected from a given regional species pool significantly decreased for litter communities but not for those in the soil layer. Such discrepancy between communities in different layers suggests that communities more exposed to anthropogenic stresses are more vulnerable to the loss of their functional roles. Our study explains possible community processes behind the observed patterns of biological organization, which can be potentially useful in guiding approaches for restoring biodiversity.

Nitrogen storage dynamics are affected by masting events in Fagus crenata.

It is generally assumed that the production of a large crop of seeds depletes stores of resources and that these take more than 1 year to replenish; this is accepted, theoretically, as the proximate mechanism of mast seeding (resource budget model). However, direct evidence of resource depletion in masting trees is very rare. Here, we trace seasonal and inter-annual variations in nitrogen (N) concentration and estimate the N storage pool of individuals after full masting of Fagus crenata in two stands. In 2005, a full masting year, the amount of N in fruit litter represented half of the N present in mature leaves in an old stand (age 190-260 years), and was about equivalent to the amount of N in mature leaves in a younger stand (age 83-84 years). Due to this additional burden, both tissue N concentration and individual N storage decreased in 2006; this was followed by significant replenishment in 2007, although a substantial N store remained even after full masting. These results indicate that internal storage may be important and that N may be the limiting factor for fruiting. In the 4 years following full masting, the old stand experienced two moderate masting events separated by 2 years, whilst trees in the younger stand did not fruit. This different fruiting behavior may be related to different "costs of reproduction" in the full masting year 2005, thus providing more evidence that N may limit fruiting. Compared to the non-fruiting stand, individuals in the fruiting stand exhibited an additional increase in N concentrations in roots early in the 2007 growing season, suggesting additional N uptake from the soil to supply resource demand. The enhanced uptake may alleviate the N storage depletion observed in the full masting year. This study suggests that masting affects N cycle dynamics in mature Fagus crenata and N may be one factor limiting fruiting.

Leaf traits, shoot growth and seed production in mature Fagus sylvatica trees after 8 years of CO2 enrichment.

BACKGROUND AND AIMS: Masting, i.e. synchronous but highly variable interannual seed production, is a strong sink for carbon and nutrients. It may, therefore, compete with vegetative growth. It is currently unknown whether increased atmospheric CO(2) concentrations will affect the carbon balance (or that of other nutrients) between reproduction and vegetative growth of forest species. In this study, reproduction and vegetative growth of shoots of mature beech (Fagus sylvatica) trees grown at ambient and elevated atmospheric CO(2) concentrations were quantified. It was hypothesized that within a shoot, fruiting has a negative effect on vegetative growth, and that this effect is ameliorated at increased CO(2) concentrations. METHODS: Reproduction and its competition with leaf and shoot production were examined during two masting events (in 2007 and 2009) in F. sylvatica trees that had been exposed to either ambient or elevated CO(2) concentrations (530 µmol mol(-1)) for eight consecutive years, between 2000 and 2008. KEY RESULTS: The number of leaves per shoot and the length of terminal shoots was smaller or shorter in the two masting years compared with the one non-masting year (2008) investigated, but they were unaffected by elevated CO(2) concentrations. The dry mass of terminal shoots was approx. 2-fold lower in the masting year (2007) than in the non-masting year in trees growing at ambient CO(2) concentrations, but this decline was not observed in trees exposed to elevated CO(2) concentrations. In both the CO(2) treatments, fruiting significantly decreased nitrogen concentration by 25 % in leaves and xylem tissue of 1- to 3-year-old branches in 2009. CONCLUSIONS: Our findings indicate that there is competition for resources between reproduction and shoot growth. Elevated CO(2) concentrations reduced this competition, indicating effects on the balance of resource allocation between reproduction and vegetative growth in shoots with rising atmospheric CO(2) concentrations.

Masting in Fagus crenata and its influence on the nitrogen content and dry mass of winter buds.

In Fagus, full-mast seeding years are invariably followed by at least one non-mast year. Both flower and leaf primordia develop during the summer within the same winter buds. Flower bud initiation occurs when the N content of developing seeds is increasing rapidly. We hypothesized that competition for nitrogen (N) between developing seeds and buds limits flower primordium formation in mast years and, hence, limits seed production in years following mast years. We tested this hypothesis in three Fagus crenata Blume forests at elevations of 550, 900 and 1500 m. Bud N concentration (N con), amount of N per bud (N bud) and dry mass per bud (DM) were compared between a mast year (2005) and the following non-mast year (2006), and between winter buds containing both leaf and flower primoridia (BF), which were formed during the non-mast year, and winter buds containing leaf primordia only (BL), which were formed in both mast and non-mast years. In addition, leaf numbers per shoot corresponding to the analyzed buds were counted, and the effect of masting on litter production was analyzed by quantifying the amounts of litter that fell in the years 2004 to 2007. The dry mass and N content of BF formed in 2006 by trees at both 550 and 1500 m were 2.1-3.4-fold higher than the corresponding amounts in BL, although the numbers of leaves per current-year shoot in 2007 that developed from the two bud types in the same individuals did not differ significantly. These results indicate that more N and carbohydrate are expended in producing BF than in producing BL. The amount of litter from reproductive organs produced in the mast year was similar to the amount of leaf litter at 900 and 1500 m, but three times as much at 550 m. Leaf numbers per shoot were significantly lower at all elevations in the mast year than in the non-mast years (and the amount of leaf litter at 550 and 1500 m tended to be lower in the mast year than in the non-mast years. In conclusion, preferential allocation of resources to seeds in the mast year reduced the availability of resources for flower primordium formation, and this may have accounted for the poor seed production in the following non-mast year.

The relative importance of carbohydrate and nitrogen for the resprouting ability of Quercus crispula seedlings.

BACKGROUND AND AIMS: Plants need some kind of stored resources to resprout after shoot destruction. The aim of this study was to determine the relative importance of carbohydrate and nitrogen (N) storage levels for their ability to resprout. METHODS: A shoot clipping experiment was conducted on Quercus crispula seedlings, which were grown in a factorial experimental design, with two light levels (40% and 3% of full light) and three nutrient concentrations (low, medium and high). KEY RESULTS: At the time of shoot clipping (the end of spring leaf expansion), seedlings exposed to 40% light had an average total non-structural carbohydrate (TNC) concentration of 17.0% in their roots compared with 4.9% in the roots of seedlings exposed to 3% light, and the average amount of TNC (TNC pools) in the roots was 203.8 mg and 20.0 mg at 40% light and 3% light, respectively. In contrast, root N concentration averaged 2.3% in the 3% light treatment compared with 1.2% in the 40% light treatment, and it increased with successive rises in nutrient concentrations at both light levels. Regardless of the nutrient status, at the 40% light level >80% of the seedlings resprouted after shoot clipping. Few seedlings, however, resprouted at the 3% light level, particularly in the medium- and high-nutrient treatments. Furthermore, both root TNC concentrations and TNC pools decreased after resprouting, but the amount of root N remained constant. CONCLUSIONS: These results suggest that carbohydrate storage has a stronger influence on resprouting in Quercus crispula than N storage. However, the size of the resprouting shoot was positively correlated with the amount of both N and TNC in roots. The level of N storage is, therefore, also important for the growth of resprouting shoots.

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

BACKGROUND AND AIMS: 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. METHODS: Current-year seedlings of Quercus serrata were grown with or without removal of all leaves at three levels of nutrient availability. KEY RESULTS: 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. CONCLUSIONS: Defoliation alters biomass allocation and chemical defence through the carbon-nutrient balance at the plant and at the leaf level, respectively.

The role of roots and cotyledons as storage organs in early stages of establishment in Quercus crispula: a quantitative analysis of the nonstructural carbohydrate in cotyledons and roots.

Quercus seedlings have hypogeal cotyledons and tap roots, both of which act as storage organs. The importance of the storage function in the two organs may change as the seedling develops. Therefore, changes in carbohydrate reserves in cotyledons and roots of Q. crispula grown under 40 % and 3 % of full light from shoot emergence to the completion of the first leaf flush were monitored. In addition, a shoot-clipping treatment was performed to examine the relative contribution of the cotyledons and tap roots to resprouting. Cotyledons maintained large amounts of nonstructural carbohydrates during shoot development, and carbohydrates were still present in the cotyledons during the final phase of leaf flush. In addition, a notable increase in the amount of carbohydrates was observed in tap roots before leaf flush at both light levels. Since root development occurred before leaf flush, even in plants grown under 3 % light, the carbohydrate found in them presumably originated from seed reserves and was translocated to roots as storage reserves. When shoots were clipped at the leaf flushing stage, the amount of carbohydrate decreased only in the cotyledons after resprouting, suggesting that cotyledons act as the main storage organs during shoot development stages. However, it could be advantageous as a 'risk avoidance strategy' for the seedlings to store reserves in both cotyledons and roots, since cotyledons may be removed by predators during shoot development.

Resprouting ability of Quercus crispula seedlings depends on the vegetation cover of their microhabitats.

To examine the effects of vegetation cover on the resprouting abilities of Quercus crispula seedlings, in each of three consecutive years, we artificially clipped seedlings growing in microhabitats with differing degrees of vegetation cover. We also investigated the relationship between the level of total nonstructural carbohydrate (TNC) and resprouting ability. Seedlings with clipped shoots in gaps produced larger resprouting shoots than those in the understory. Moreover, both the percentage of resprouting seedlings and the survival ratio in seedlings with clipped shoots were negatively correlated with the degree of vegetation cover. Seedlings stored high levels of TNC, especially in their roots, and their TNC levels were negatively correlated with the degree of vegetation cover. There were also positive relationships between the TNC levels in their roots and the degree of resprouting. Hence, we conclude that release from vegetation cover enhanced the resprouting ability of Q. crispula seedlings by increasing their levels of stored carbohydrate. The key variables affected were the resprouting ratio (the proportion of seedlings capable of producing new shoots) and the size of the resprouted shoots.