Notes on beech (Fagus crenata Blume) seed and seedling mortality due to rodent herbivory in a northernmost beech forest, Utasai, Hokkaido

We investigated the mortality of post-dispersal seeds and current-year seedlings of a beech species (Fagus crenata Blume) at the northernmost extent of its geographical range at Utasai (Kuromatsunai), Hokkaido, Japan. Introduced seeds that were protected from rodent predation by wire cages had a higher survival rate from winter to the following spring (mean ± SD: 84 ± 16.7%,n = 10) than controls without cages (mean ± SD: 12 ± 17.9%,n = 10). Unprotected transplanted current-year seedlings with almost fully opened primary leaves never survived more than three days (0%,n = 30), while more than 80% (n = 30) of seedlings within cages survived to the end of the growing season. These results indicate that rodent herbivory is a major mortality factor that strongly prevents the establishment of beech seedlings. In a natural population, we observed that 90% of seedlings (n = 197) disappeared within 10 days after their emergence and rodent herbivory caused this heavy mortality. A herbivorous rodent, the gray-backed vole,Clethrionomys rufocanus bedfordiae, was remarkably abundant in late June (101.5 voles/ha), suggesting that this species strongly affects the formation of the seedling bank. The presence ofC. rufocanus bedfordiae may be one of the reasons for the scarcity of beech seedlings and saplings and the rarity of recruitment in this northernmost beech forest.     

Phenological phases of common beech (Fagus sylvatica L.) and their dependence on region and altitude in Southern Germany

Long-term observational data of the phenological network of the German Weather Service (DWD) were used to derive the altitudinal dependence of leaf unfolding (LU), leaf discolouration (LD) and the length of the vegetation period (LD-LU) of common beech (Fagus sylvatica L.) in different regions of Southern Germany. Depending on altitude, the vegetation period starts in Bavaria between the 2nd and 24th May with a delay of around 2 days per 100 m altitude increase (averaged over the last 30 years). In Southern Bavaria, LU starts a few days earlier as well as increases slightly faster into higher altitudes than in the northern regions. This can be explained by a more favourable supply of warmth and radiation in the south. The end of the vegetation period, defined here as the beginning of LD, showed no altitudinal dependence, which reflects an interaction of endogenous and exogenous factors as triggers for LD. In the Bavarian Forest region, altitudinal dependences are strongly influenced by sites which are affected by cold-air tailback (‘Kaltluftstau’) was obtained. Hence, phenological gradients should only be derived and applied above a phenological inversion zone. The calculated vertical gradients for LU can be used together with available observational data to estimate the beginning of the forest vegetation period of beech for different altitudes of Bavaria.     

Net photosynthetic response to light intensity of shoots from different crown positions and age in picea abies (L.) karst

Carbon dioxide uptake response to light intensity of detached shoots of open‐ and forest‐grown Norway spruces was investigated in controlled standard conditions. The initial slope of the CO₂ uptake light response curve, calculated on leaf area basis and the internal conductance, calculated on leaf weight basis were almost constant for shoots from different positions in the forest canopy. In open‐grown trees the maximum photosynthesis and internal conductance (on leaf weight basis) decreased in relation to the tree age, so that in the 8‐year‐old tree it was about two times of that in the 66‐year‐old tree. These results suggest that 1) the enhanced efficiency of “shade” shoots is caused by morphologic adaptation, i.e. the more sparsely packed photosynthetic apparatus in needles and shoots, and 2) adaptational possibilities of Norway spruce photosynthetic apparatus, its ecological plasticity diminish during tree ontogenesis.     

Photosynthesis and biomass allocation of beech (Fagus crenata) and dwarf-bamboo (Sasa kurilensis) in response to contrasting light regimes in a Japan Sea-Type beech forest

Regeneration of beech (Fagus crenata) forests depends on the formation of canopy gaps. However, in Japan Sea-type beech forests, a dwarf bamboo (Sasa kurilensis) conspicuously occupies sunny gaps. Therefore,F. crenata seedlings must escape the severe interference ofS. kurilensis in the gaps and persist beneath a closed canopy of the beech forest. We hypothesized that the growth ofF. crenata seedlings in the understory would be favored by their being more plastic thanS. kurilensis in photosynthetic and morphological traits, which would support the matter production ofF. crenata seedlings in a wide range of light availabilities. To examine this hypothesis, the photosynthetic-light response of individual leaves and the biomass allocation in aboveground parts (i.e., the culm/foliage ratio) were surveyed at sites with contrasting light availabilities in a Japan Sea-type beech forest in central Japan. InF. crenata, photosynthetic light utilization efficiency at relatively low light was greater, and the dark respiration rate was smaller in the leaves of seedlings (10 cm in height) beneath the closed canopy than in the leaves of saplings at the sunny forest edge. The culm/foliage (C/F) ratio of theF. crenata seedlings at the shady site was small, suggesting effective matter-production beneath the beech canopy. On the other hand,S. kurilensis both in the gap and beneath the beech canopy showed low plasticity in photosynthesis and the culm/foliage ratio. Because the shoot density ofS. kurilensis was smaller beneath the beech canopy than in the gap, the light availability at the bottom of theS. kurilensis layer was greater beneath the beech canopy. These results suggest that the photosynthetic productivity of theF. crenata seedlings would be enough for the seedlings to survive in the understory with a low density ofS. kurilensis shoots beneath the closed beech canopy.     

Nutrient contents and efficiencies of beech and spruce saplings as influenced by competition and O<Subscript>3</Subscript>/CO<Subscript>2</Subscript> regime

Saplings of Fagus sylvatica and Picea abies were grown under conditions of intra and interspecific competition in a 2-year phytotron study under combinations of ambient and elevated ozone (+O₃ which is 2 × O₃, but <150 nl l⁻¹) as well as carbon dioxide concentrations (+CO₂ which is amb. CO₂ + 300 μl CO₂ l⁻¹) in a full factorial design. Saplings were analysed for various mineral nutrients in different plant organs as well as biomass production and crown development. The study was based on the assumption that nutritional parameters important for growth and competitiveness are affected by stress defence under limiting nutrient supply. The hypotheses tested were (1) that nutrient uptake-related parameters (a) as well as efficiencies in nutrient use for above-ground competition (b) of beech rather than spruce are impaired by the exposure to elevated O₃ concentrations, (2) that the efficiency in nutrient uptake of spruce is enhanced by elevated CO₂ concentrations in mixed culture, and (3) that the ability to occupy above-ground space at low nutrient cost is co-determinant for the competitive success in mixed culture. Clear nitrogen deficiencies were indicated for both species during the 2-year phytotron study, although foliar nitrogen-biomass relationships were not so close for spruce than for beech. O₃ stress did not impair nutrient uptake-related parameters of beech; thus hypothesis (1a). was not supported. A negative effect of elevated O₃ (under amb. CO₂) on the N and P based efficiencies in above-ground space occupation (i.e. lower crown volume per unit of N or P invested in stems, limbs and foliage) of beech supported hypothesis (1b). It appeared that ozone stress triggered a nutrient demand for stress defence and tolerance at the expense of above-ground competition (trade-off). Crown volume of beech under O₃ stress was stabilized in monoculture by increased nutrient uptake. In general, the +CO₂-treatment was able to counteract the impacts of 2 × O₃. Elevated CO₂ caused lower N and S concentrations in current-year foliage of both tree species, slightly higher macronutrient amounts in the root biomass of spruce, but did not increase the efficiencies in nutrient uptake of spruce in mixed culture. Therefore hypothesis (2) was not supported. At the end of the experiment spruce turned out to be the stronger competitor in mixed culture as displayed by its higher total shoot biomass and crown volume. The amounts of macronutrients in the above-ground biomass of spruce individuals in mixed culture distinctly exceeded those of beech, which had been strongly reduced by interspecific competition. The superior competitiveness of spruce was related to higher N and P-based efficiencies in above-ground space occupation as suggested in hypothesis (3). This article belongs to the special issue “Growth and defence of Norway spruce and European beech in pure and mixed stands”.     

Gap formation in Danish beech (Fagus sylvatica) forests of low management intensity: soil moisture and nitrate in soil solution

Soil moisture content (0–90 cm depth) and nitrate-nitrogen (NO₃-N) concentrations in soil solution (90 cm depth) were monitored after gap formation (diameter 15–18 m) in three Danish beech-dominated forests on nutrient-rich till soils. NO₃-N drainage losses were estimated by the water balance model WATBAL for one of the sites. Two forests were non-intervention forests (semi-natural and unmanaged), the third was subject to nature-based management. The study was intended to assess the range of effects of gap formation in forests of low management intensity. In the unmanaged and the nature-based managed forest, soil solution was collected for 5 years and soil moisture measured in the fourth year after gap formation. Average NO₃-N concentrations were significantly higher in the gaps (9.9 and 8.1 mg NO₃-N l⁻¹, respectively) than under closed canopy (0.2 mg l⁻¹). In the semi-natural forest, measurements were carried out up to 29 months after gap formation. Average NO₃-N concentrations in the gap were 19.3 mg NO₃-N l⁻¹. Gap formation alone did not account for this high level, as concentrations were high also under closed canopy (average 12.4 mg NO₃-N l⁻¹). However, the gap had significantly higher N concentrations when trees were in full leaf, and NO₃-N drainage losses were significantly increased in the gap. No losses occurred under closed canopy in growing seasons. Soil moisture was close to field capacity in all three gaps, but decreased under closed canopy in growing seasons. In the semi-natural forest, advanced regeneration and lateral closure of the gap affected soil moisture levels in the gap in the last year of the study.     

Characterization of lignin-carbohydrate linkages in the residual lignins isolated from chemical pulps of spruce (<Emphasis Type="Italic">Picea abies</Emphasis>) and beech wood (<Emphasis Type="Italic">Fagus sylvatica</Emphasis>)

To investigate the linkage types between carbohydrates and lignin, residual lignins were isolated from three different unbleached pulps [kraft, alkaline sulfite anthraquinone methanol (ASAM), and soda with anthraquinone (AQ) and methanol] of spruce and beech wood and then characterized by oxidation with 2,3-Dichloro-5,6-dicyanobenzoquinone and followed by Prehm’s methylation. In residual lignins, sugar moieties were bound to lignins via benzyl ether bonds. In particular, galactose and mannose are predominantly linked to lignin fragments in residual lignins of spruce wood, while xylose and galactose are favored in the formation of LC bonds in the residual lignins of beech wood. In the case of hexoses, primary hydroxyl groups (C6 position) preferentially take part in benzyl ether linkages. Hydroxyl groups in the C2 and C3 positions of xylose participate in LC bonds and a small portion of arabinose was notably connected to lignin via the C5 position. Approximately seven or eight sugars were connected in soda/AQ/methanol residual lignin per 100 C9 lignin units, while the frequencies of LC bonds in kraft and ASAM residual lignins were distinctively less at one to three sugars per 100 C9 lignin units. The publication of this article was made possible by an Emachu Research Fund. The authors are grateful for the fund.     

Spatial pattern of beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) and oak (<Emphasis Type="Italic">Quercus pubescens</Emphasis> Mill.) seedlings in natural pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) woodlands

Spatial pattern of recruitment is an important factor influencing population dynamics of plant communities. The spatial pattern is determined by seed dispersal and by the spatial variability of germination and initial survival. In the process of forest expansion following farmland abandonment, mid- and late-successional species are often dispersed in pioneer forests by birds. This could result in an aggregated spatial pattern of seeds that could influence the dynamics of these species in mixed pioneer forests. In the sub-Mediterranean area, mid- and late-successional species such as Quercus pubescens (downy oak) and Fagus sylvatica (European beech) are expected to replace pioneer Pinus species. Using a point sampling method we demonstrated that beech and oak seedlings (height <2 m) have a clumped distribution in the understorey of pine. This could result from an aggregated dispersal by jays (dispersal effect) or from preferential recruitment in particular habitats (habitat effect). To test these hypotheses we proposed a statistical analysis of spatial patterns of regeneration of beech and oak. Ground cover variables (i.e. cover by rock outcrops, herbs, box shrubs, mosses or pine) did not differ significantly around seedlings as compared with random sample plots. Likewise, clumped seedlings had growth similar to isolated seedlings, thus refuting the hypothesis of preferential location in the most favourable microsites. Aggregated dispersal seems to be involved in the process of regeneration. Since beech and oak seedlings have contrasting ecological demands, we discuss the implication of this pattern for the replacement dynamics of pine by these species.     

Family Variation in the Morphology and Physiology of White Spruce (Picea glauca) Seedlings in Response to Elevated CO2 and Temperature

Tree improvement programs aim to develop families that are well-adapted to future growing conditions. To gain insight into the stability of the family genetic response to climate change, white spruce (Picea glauca) seedlings from 60 full-sib families were subjected to a combination of two temperature regimes and two levels of CO₂ over two growing seasons. There was positive effect of warmer temperatures and higher CO₂ on some growth variables but no significant family × treatment interactions. Instantaneous water use efficiency was the only physiological trait that was affected positively by the CO₂ treatment, showing a 51% increase that was consistent across families.     

Extraction of <Emphasis Type="Italic">Trigonella foenum-graecum</Emphasis> L. by supercritical fluid CO<Subscript>2</Subscript> and its contact toxicity to <Emphasis Type="Italic">Rhyzopertha dominica</Emphasis> (Fabricius) (Coleoptera: Bostrichidae)

Rhyzopertha dominica (Fabricius) has developed extensive pesticide resistance in the last several decades. We have developed a supercritical fluid extraction method for Trigonella foenum-graecum L. (TFG) and studied the contact toxicities of the extracts to R. dominica. The extraction method was designed with orthogonal experiments to preserve and collect all the possible active components. Contact toxicity and efficiency of extraction were used as standard values to optimize extraction conditions, which were achieved at 55°C under 25 Mpa of pressure. The extraction efficiency for 200 g of dry sample reached 6.21% with 30 ml of 95% alcohol. Extracts loaded on filter paper showed dose and time dependent toxicities to adult R. dominica with a LC₅₀ value of 65.02 μg/cm² after 3 days post treatment. Our extensive in vivo studies indicated the extracts from Trigonella foenum-graecum seeds have high efficacy against pesticide resistant R. dominica. The active ingredient(s) from the extract shows promise as a novel pesticide candidate.     

Cost, energy and carbon dioxide (CO2) effectiveness of a harvesting and transporting system for residual forest biomass

The purpose of this study is to examine the feasibility of a system to harvest logging residues (or slashes) as a new resource for energy in Japan. A harvesting and transporting system for residual forest biomass was constructed with reference to some European countries where the utilization of bioenergy is making steady progress and examined on the basis of field experiments in Japanese forestry. The feasibility of the system is discussed from the standpoints of cost and energy, and the system is compared with those of the European countries. With respect to the system proposed in this study, it is desirable that the process of chipper comminuting is incorporated into the system as early as possible, considering the trends of harvesting cost and fuel consumption per unit weight of residual forest biomass. Such a system is not particularly feasible in Japan from the standpoint of the harvesting cost per MWh of bioenergy. However, no specific problems are found from the point of view of the energy input rate, and it is clarified that it is possible for Japan to reduce domestic carbon dioxide emissions by utilizing biomass as an energy resource. A comparison with the European countries and a preliminary sensitivity analysis of the system demonstrate that the technical development to reduce the harvesting cost,e.g., improving the forwarding and transporting efficiency, and support from the government are essential for realizing bioenergy utilization in Japan. A part of this paper was orally presented at the 111th Annual Meeting of the Japanese Forestry Society (2000). JSPS Research Fellow. This study was supported in part by a Grant-in-Aid for Scientific Research from the Japan Ministry of Education, Science and Culture (No. 10460061).     

Energy and carbon dioxide (CO2) balance of logging residues as alternative energy resources: system analysis based on the method of a life cycle inventory (LCI) analysis

Using the method of a life cycle inventory (LCI) analysis, the energy balance and the carbon dioxide (CO₂) emission of logging residues from Japanese conventional forestry as alternative energy resources were analyzed over the entire life cycle of the residues. The fuel consumption for forestry machines was measured in field experiments for harvesting and transporting logging residues at forestry operating sites in Japan. In addition, a total audit of energy consumption was undertaken. It involved an assessment of materials, construction, and the repair and maintenance of forestry machines as well as the costs associated with an energy-conversion plant. As a result, the ratio of energy output to input was calculated to be 5.69, indicating that the system examined in this study could be feasible as an energy production system. The CO₂ emission per MWh_e (e: electricity) of the biomass-fired power generation plant was calculated to be 61.8kgCO₂/MWh_e, while that of coal-fired power generation plants in Japan is 960kgCO₂/MWh_e. Therefore, the reduction in the amount of CO₂ emission that would result from replacing coal with biomass for power generation by as much as 3.0 million dry-t/year of logging residues in Japan was estimated to be 1.66 million tCO₂/year, corresponding to 0.142% of the national CO₂ emission. This study provides evidence that Japan could reduce its domestic CO₂ emission by using logging residues as alternative energy resources.