Delayed soil thawing affects root and shoot functioning and growth in Scots pine

In boreal regions, soil can remain frozen after the start of the growing season. We compared relationships between root characteristics and water relations in Scots pine (Pinus sylvestris L.) saplings subjected to soil frost treatments before and during the first week of the growing period in a controlled environment experiment. Delayed soil thawing delayed the onset of sap flow or totally blocked it if soil thawing lagged the start of the growing period by 7 days. This effect was reflected in the electrical impedance of needles and trunks and in the relative electrolyte leakage of needles. Prolonged soil frost reduced or completely inhibited root growth. In unfrozen soil, limited trunk sap flow was observed despite unfavorable aboveground growing conditions (low temperature, low irradiance, short photoperiod). Following the earliest soil thaw, sap flow varied during the growing season, depending on light and temperature conditions, phenological stage of the plant and the amount of live needles in the canopy. The results suggest that delayed soil thawing can reduce tree growth, and if prolonged, it can be lethal.     

Demographic and height growth response of native broad-leaved deciduous tree saplings to overhead canopy release in a coastal Pinus thunbergii forest in Hokkaido, northern Japan

Prevention efforts to combat pine wilt disease must be instituted to avoid destruction of coastal forests in Hokkaido, northern Japan. We examine an alternative method involving the conversion of Pinus thunbergii forests to native broadleaf species through advancing their growth, but we have little knowledge about the response of advanced growth to overhead canopy release. To demonstrate the demographic and height growth response of advanced growth to the light change in a coastal Pinus thunbergii forest, canopy trees were removed and the sapling populations were monitored for 6?years. Inhibition in height growth caused by light increase was observed in Cerasus sargentii and Kalopanax septemlobus saplings. Especially in Kalopanax septemlobus, a remarkable decline was observed in the saplings growing in the darker positions which experienced light change. Therefore, large light changes should be carefully avoided for this species especially in the darker positions. In contrast, relative height growth rates of Quercus crispula and, especially, Quercus dentata increased with increasing light, and saplings tended to die in the darker positions. No dead trees of Sorbus commixta were observed during the study period. The saplings also showed good height growth even when under closed canopy. This species seems to be adaptive to a dark environment. Since Quercus dentata and Kalopanax septemlobus are the major components of the natural coastal forests in Hokkaido, gradual canopy release is available to foster advanced growth in coastal Pinus thunbergii forest, in accordance with the concept of density control in coastal forests.     

Seasonal nutrient retranslocation in reforested <Emphasis Type="Italic">Pinus halepensis</Emphasis> Mill. stands in Southeast Spain

Retranslocation, resorption and relocation of nutrients are important adaptive mechanisms developed by plants to acquire the amount of the nutrients required for growth. They are usual mechanisms in deciduous and conifer trees that occur in Mediterranean regions where drought periods are usual. Soil factors, environmental characteristics and species factors are key drivers of nutrient retranslocation in conifers but is not well understood how soil fertility or intraspecific competition influences the process. We studied retranslocation in Pinus halepensis Mill. stands showing different site quality (differences in climate and intraspecific competition) occurring in Southeast Spain. We monitored reforested mature Aleppo pine forests in stands with differences in site quality, climate and intraspecific competition. Stands were characterised, the content of nutrients of soil and green samples (twigs and pine needles) were recorded, and seasonal nutrient retranslocation was obtained. Site characteristics were related to growth rate and nutrient content of foliage and soil. We evaluated whether the retranslocation of nutrients from older to younger foliage was related to the current-year growth rate and to the nutritional status of the plant as influenced by intraspecific competition. Foliar macronutrient concentrations and the amount of retranslocated macronutrients were seasonal, with differences related to site quality and tree density. As a general trend, nutrient concentrations increased after drought (autumn) and decreased during the growth period (spring). However, some micronutrients (mainly Na and Fe) decreased during both periods. The retranslocation pattern in Aleppo pine reinforced the hypothesis that pine adaptations to drought- and fire-prone habitats are linked to the resilience of these forest types. We developed scientific knowledge to assist decision making in adaptive forest management; e.g. fertilizer recommendations or reforestation programmes.     

Effects of timing of soil frost thawing on Scots pine

Effects of the timing of soil thawing in the spring on Scots pine (Pinus sylvestris L.) were studied under controlled laboratory conditions. Sixteen 6-year-old saplings were lifted from the field, replanted in containers and placed in four treatments in controlled environment (CE) chambers with four replicate saplings per chamber. The saplings were held in the CE chambers during one simulated winter and one simulated growing season. The soil was frozen to -2 degrees C during a second simulated winter in the CE chambers, and the soil thawing treatments began at the end of the second simulated winter. Soil thawing began at various times before (no delay in thawing) and after (delay in thawing) chamber air conditions were changed from simulated winter to simulated summer. Delayed soil thawing subjected saplings to stress, with the severity of stress depending on the length of the delay in thawing. If there was no delay or only a short delay in soil thawing, stress was minor and reversible. A 2-week delay in soil thawing led to death of the saplings. Stress was apparent as decreases in the variable to maximal chlorophyll fluorescence ratio (Fv/Fm), chlorophyll a/b ratio and needle water potential. In needles of stressed saplings, apoplastic electrical resistance first decreased and then increased and there were anomalies in the electrical impedance spectra of the stems. Stress from the soil thawing treatments affected both root and shoot growth.     

Impacts of seasonal air and soil temperatures on photosynthesis in Scots pine trees

Seasonal courses of light-saturated rate of net photosynthesis (A360) and stomatal conductance (gs) were examined in detached 1-year-old needles of Scots pine (Pinus sylvestris L.) from early April to mid-November. To evaluate the effects of soil frost and low soil temperatures on gas exchange, the extent and duration of soil frost, as well as the onset of soil warming, were manipulated in the field. During spring, early summer and autumn, the patterns of A360 and gs in needles from the control and warm-soil plots were generally strongly related to daily mean air temperatures and the frequency of severe frost. The warm-soil treatment had little effect on gas exchange, although mean soil temperature in the warm-soil plot was 3.8 degrees C higher than in the control plot during spring and summer, indicating that A360 and gs in needles from control trees were not limited by low soil temperature alone. In contrast, prolonged exposure to soil temperatures slightly above 0 degrees C severely restricted recovery of A360 and especially gs in needles from the cold-soil treatment during spring and early summer; however, full recovery of both A360 and gs occurred in late summer. We conclude that inhibition of A360 by low soil temperatures is related to both stomatal closure and effects on the biochemistry of photosynthesis, the relative importance of which appeared to vary during spring and early summer. During the autumn, soil temperatures as low as 8 degrees C did not affect either A360 or gs.     

Reduction of frost heaving of Norway spruce and Scots pine seedlings by planting in mounds or in humus

This study was carried out in northern Sweden to determine the effects of frost heaving on the establishment of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings in relation to planting methods. For this purpose, one year old containerised seedlings were planted on two sites and on two dates: during the spring (early planting) and during the fall (late planting). In each case, two planting depths (normal and deep planting) and four planting sites (in mineral soil in the depressions, in the scalp/trench area, on the top of the mound and in the untreated humus layer) were used. On each site, 50 seedlings were planted for each treatment. Frost heaving was observed and measured during two years. The amount of heaving was highest in the hole and almost insignificant on the top of the mound and in the humus layer. Planting depth influenced the degree of heaving only for Scots pine planted in the hole and was not related to the planting time.     

Evaluation of regeneration potential of Pinus koraiensis in mixed pine-hardwood forests in the Xiao Xing’an Mountains, China

Large scale harvest of Korean pine(Pinus koraiensis) seeds as a food product in the mixed Korean pine-hardwood forest of northeastern China poses a serious threat to the sustainability and restoration of this endangered regional ecosystem.Seed collection over past decades greatly reduced the seed bank and subsequent seedling and sapling recruitment,and impacting a wide array of granivorous animals that rely on the pine seeds.We surveyed Korean pine seeds,including solid seeds(SS),insect consumed seeds(ICS) and other(animal) consumed(OCS) kernels,of the seed bank(forest floor and the top 10 cm of mineral soil),the seedlings and saplings from 1 m 2 sample plots in five forest types in Liangshui Nature Reserve(LNR) of the southern Xiao Xing’an Mountains in northeastern China to provide accurate information for assessing the Korean pine regeneration potential.The average number of pine seeds in the seed bank were 11.2 seeds/m 2,9.1 seeds/m 2,4.6 seeds/m 2,1.1 seeds/m 2,and 0.2 seeds/m 2 in Korean pine-basswood forest,mixed Korean pine-hardwood forest,mixed conifer-hardwood forest,white birch forests,and oak forests,respectively.In the first three forest types,percentages of SS(potentially viable seeds) were 11.2%,3.5% and 27.8%,respectively.The percentages of ICS(not viable seeds) were consistent at around 35%.The higher but variable percentages of OCS(not viable seeds) indicated high seed predation in these forests.Compared with other studies,we recorded higher percentages of seed damage,probably due to our survey approach and the increased depth of seed bank sampled in our study.Depletion of pine seeds in the seed bank greatly reduced seedling and sapling recruitment.Densities of pine seedlings varied from about 180 trees/ha in the mixed Korean pine-hardwood forest to about 5,400 trees/ha in the mixed conifer-hardwood forests and showed a high degree of spatial variation.Saplings were rare in the mixed Korean pine-hardwood forest,but ranged in the thousands in other forests.Large scale pine seed harvest has seriously threatened the sustainability of the mixed Korean pine-hardwood forest ecosystem.Scaling down the seed harvest or supplemental planting of pine saplings are urgently needed to maintain the health of the existing Korean pine forests and to restore this endangered ecosystem.     

Effects of silicon-rich soil amendments on growth,mortality and bark feeding damage of Sitka spruce (Picea sitchensis) seedlings under field conditions

Alternative methods of protection are required against feeding by the large pine weevil (Hylobius abietis) on the bark of conifer seedlings. Silicon (Si) has been shown to enhance the resistance of plants to insect herbivores. This study investigated the effects of low doses of Si-rich soil amendments on growth, mortality and bark feeding damage of Sitka spruce (Picea sitchensis) seedlings. Two-year old seedlings were grown, individually, in soil taken from a tree nursery treated with coal ash, peat ash, rice husk ash, slag, sodium metasilicate or a commercially available Si fertiliser (Pro-Tekt) and planted out on two reforestation sites in Ireland. Seedlings grew well (about 20% growth in terms of height, 66% in root collar diameter, after two growing seasons), and Si-rich amendments did not have a significant effect on growth or mortality. Bark feeding damage on Si-treated seedlings did not vary significantly from control seedlings. Bark Si concentrations were not significantly larger in treated seedlings than in control seedlings, but control seedlings already had comparatively high bark Si concentrations (560?mg?kg^?1 dry tissue). In conclusion, Sitka spruce seedlings grown in the presence of Si-rich soil amendments prior to planting did not show greater resistance to weevil feeding under the present conditions.     

灰木莲人工林地土壤微生物及酶活性分析

对广西现代林业科技园优良树种展示区的灰木莲8年生(G1)和灰木莲45年生(G2)的0～20cm土层一年四季的微生物数量及土壤酶活性进行分析。结果表明:灰木莲幼林(G1)与成熟林(G2)的林地土壤微生物和土壤酶有明显的差异。微生物总量和细菌数量的季节变化,G1均为夏>冬>秋>春,而G2是秋>冬>夏>春。真菌数量上无论是G1或G2均是秋季最多而冬季最少。放线菌数量的季节变化是,G1:春季>冬季>秋季>夏季,G2:冬季>春季>秋季>夏季。对两种林地不同季节的微生物而言,春秋冬季的细菌>放线菌>真菌,夏季则是细菌>真菌>放线菌。土壤蛋白酶活性表现为:G1,春>夏>秋>冬;G2,秋>春>夏>冬,而G1和G2的过氧化氢酶活性均是冬>夏>秋>春。     

Responses of soil carbon, nitrogen and cations to the frequency and seasonality of prescribed burning in a Cape Cod oak-pine forest

Fire is an important component of the historic disturbance regime of oak and pine forests that occupy sandy soils of the coastal outwash plain of the northeastern US. Today prescribed fire is used for fuel reduction and for restoration and maintenance of habitat for rare plant and animal, animal species. We evaluated the effects of the frequency and seasonality of prescribed burning on the soils of a Cape Cod, Massachusetts's coastal oak-pine forest. We compared soil bulk density, pH and acidity, total extractable cations and total soil carbon (C) and nitrogen (N) in unburned plots and in plots burned over a 12-year period, along a gradient of frequency (every 1–4 years), in either spring (March/April) or summer (July/August). Summer burning decreased soil organic horizon thickness more than spring burning, but only summer burning every 1–2 years reduced organic horizons compared with controls. Burning increased soil bulk density of the organic horizon only in the annual summer burns and did not affect bulk density of mineral soil. Burn frequency had no effect on pH in organic soil, but burning every year in summer increased pH of organic soil from 4.01 to 4.95 and of mineral soil from 4.20 to 4.79. Burning had no significant effect on organic or mineral soil percent C, percent N, C:N, soil exchangeable Ca²⁺, Mg²⁺, K⁺ or total soil C or N. Overall effects of burning on soil chemistry were minor. Our results suggest that annual summer burns may be required to reduce soil organic matter thickness to produce conditions that would regularly allow seed germination for oak and for grassland species that are conservation targets. Managers may have to look to other measures, such as combinations of fire with mechanical treatments (e.g., soil scarification) to further promote grasses and forbs in forests where establishment of these plants is a high priority.     

Inhibition of the regeneration of Japanese black pine (<Emphasis Type="Italic">Pinus thunbergii</Emphasis>) by black locust (<Emphasis Type="Italic">Robinia pseudoacacia</Emphasis>) in coastal sand dunes

Pine forests are declining because of pine wilt disease and Robinia pseudoacacia, a nitrogen-fixing species, is dominating coastal forests along the Sea of Japan. We examined the effects of R. pseudoacacia on the regeneration of a native pine species, Pinus thunbergii. Two 200 m² plots were set up at the border of a P. thunbergii and R. pseudoacacia-dominated area in a coastal forest. We conducted monthly censuses of emergence, distribution and survival of pine seedlings in the plots from May 2003 to December 2004. Light intensity and soil properties were also measured to analyze the relationships between the survival of pine seedlings and environmental conditions using the Mantel test and the structural equation model. Pinus thunbergii seedlings emerged in spring–early summer and in late autumn. Survival of pine seedlings in the R. pseudoacacia-dominated subplots was less than half that in the pine-dominated subplots. Survival of pine seedlings emerging in May 2003 was significantly reduced by the lower light intensity and higher soil nitrogen in R. pseudoacacia-dominated subplots. The tendency was the same for seedlings emerging from April to May 2004. We concluded that R. pseudoacacia reduced the intensity of light during the growing season and increased the nitrogen content of soil, which resulted in inhibition of the natural regeneration of P. thunbergii.     

The role of developmental stage in frost tolerance of Pinus pinea L. seedlings and saplings

Context

Although drought is generally considered the main environmental constraint in Mediterranean environments, the ability to acclimate to and tolerate frost in early developmental stages can be a determinant for seedling survival of many Mediterranean tree species like stone pine (Pinus pinea L.).

Aims

The aim of this study was to assess the impact of the developmental stage of naturally regenerated stone pine individuals on tolerance to low temperature (LT) from summer to late autumn and in spring, at a highly continental site in central Spain. Specifically, we tested to what extent the differences in tolerance are related to shoot heteroblasty.

Methods

We assessed LT tolerance of needles from individuals at three age classes (class C1: seedlings, class C2: 4- to 8-year-old saplings and class C3: >9-year-old saplings) over nine dates from summer to spring.

Results

LT tolerance displayed severe seasonal trends and differed between age classes. It usually increased with sapling age. Such differences were tightly related to heteroblasty of the shoots. Our results point to a higher LT tolerance associated with larger leaf dry mass per unit area (LMA) values. No impact of late frosts on shoot growth rates was detected during this study.

Conclusions

Developmental changes during early plant growth seem to play a role in frost tolerance of stone pine seedlings, a finding which furthers our understanding of regeneration dynamics in this species in areas with continental influence.     

Height growth of Korean pine plantation sapling in Benxi, Liaoning Province, China

The height growth of Korean pine plantation was investigated for different ages of saplings in Benxi city of the eastern Liaoning, China. The sapling and seedling age were determined by branch annuals. Results show that the mean height of third- to eight-year Korean pines ranges from 13.31 cm to 111.73 cm and only 19.4% of the saplings reached the height above 130 cm. The height (Y) of eight-year-old Korean pine increases linearly with the increment of ground stem (x) (Y=3.1x+35.15). The relation between height and age can be described by an exponential equation. The second year after planting of the Korean pine is considered as the key period of Korean pine growth really adapting to the local environment; plantation management should be strengthened in this year. The Korean pine seedlings need two years to adapt to the local environment, thus early care and management should be taken in the second year.     

Regeneration of European boreal forests: Effectiveness of measures against seedling mortality caused by the pine weevil Hylobius abietis

Successful regeneration of conifer forests by planting is, in large parts of Europe, highly dependent on the effective suppression of damage caused by the pine weevil Hylobius abietis. We investigated the effectiveness of various combinations of control measures against pine weevil damage under boreal forest conditions in Sweden. In particular, we aimed to determine whether satisfactory regeneration could be obtained without the use of insecticides. The experimental study was established on ten new clear-cuts in each of three consecutive years. We studied the use of chemical and physical methods to protect seedlings directly, and investigated the influence of seedling type, age of clear-cut, and a number of soil factors as affected by preparation of the planting site, on the mortality and level of damage suffered by protected and unprotected seedlings. After two seasons, mortality due to pine weevil was 16% among unprotected seedlings, 6% for seedlings treated with the insecticides cypermethrin or imidacloprid, and less than 1% for those physically protected by a coating of Conniflex. However, the Conniflex, which consists of fine-grained sand embedded in a flexible acrylate matrix, was applied manually, and this may have enhanced its effectiveness compared to that achieved during large-scale, commercial application. Two types of containerized Norway spruce seedlings, which differed mainly in their stem diameter (average 2.6 mm and 3.5 mm), were used in the experiments. Among the unprotected seedlings, the narrower stemmed type was more frequently attacked (34% vs. 28%) and killed (19% vs. 12%) by pine weevil. Mortality caused by pine weevil among unprotected seedlings was higher on 1-year-old than on 2-year-old clear-cuts (20% vs. 12%). Soil preparation around unprotected seedlings had a substantial effect on the proportion attacked and killed by pine weevil as well as on the total mortality, with the highest level of feeding damage and mortality occurring on seedlings in undisturbed humus, and the lowest levels occurring on seedlings planted in pure mineral soil (26% vs. 7% for unprotected seedlings). This study demonstrates that acceptable levels of seedling survival can be achieved in regenerations of North European boreal forest without the use of insecticides. Mortality of unprotected seedlings can be reduced to acceptable levels if they are mostly planted in pure mineral soil. Damage can be further reduced by using seedlings with a somewhat larger stem diameter. Insecticides or a physical barrier of Conniflex alone appear to provide a sufficient level of protection.     

Assessing the potential for ash canopy tree replacement via current regeneration following emerald ash borer-caused mortality on southeastern Michigan landscapes

The emerald ash borer (EAB) has killed millions of ash trees in Michigan and at least fourteen other states since its first detection near its introduction point in metropolitan Detroit in 2002. Despite overstory ash mortality near 100% in many areas, ash seedling and saplings remain unaffected and provide the potential for ash re-establishment into the canopy of deciduous forests of the region. We examined the potential for ash re-establishment in areas of heavy mortality by measuring the status and change of ash regeneration at 45 sites across southeastern Lower Michigan in 2007 and 2009. Ash regeneration was found to be abundant in all forest types, particularly in the smallest height classes, though it was more abundant in ash species of upland forests compared to those of lowland forests. New seedlings 1-2 years old were less common than other regeneration size classes, and declined substantially between 2007 and 2009, suggesting a depletion of the ash seed bank in these forests. Regeneration density was not explained well by the presence of overstory ash prior to EAB introduction, suggesting that competitive interaction with other tree species in higher canopy strata is an important driver of ash regeneration density. Regeneration is sufficient to replenish overstory ash to pre-EAB levels in upland forests, though ecological changes caused by overstory ash mortality in lowland areas may affect local hydrology in a way that reduces opportunities for tree regeneration. Despite abundant ash regeneration, it remains unclear whether ash will recover to the overstory of forests in southeastern Michigan because of uncertainties in the future dynamics of EAB in the region.     

Foliage re-establishment of <Emphasis Type="Italic">Pinus palustris</Emphasis> Mill. saplings after spring or fall prescribed fire

Repeated fire is key to the viability of longleaf pine (Pinus palustris Mill.) ecosystems, but its acceptance as a management tool may depend on satisfactory longleaf pine growth. This is because longleaf pine establishment often has the dual-purpose of ecosystem restoration and stemwood production. Timely recovery of scorched foliage among longleaf pine seedlings and saplings supports maximum juvenile growth. We identified two means of foliage re-establishment in the growing season after prescribed fire regardless of the season of fire application. New foliage growth after spring or fall fire was correlated with un-scorched foliage biomass and the presence of lateral branches. After prescribed fire in spring, foliage biomass recovery also appeared to benefit from the mobilization of starch. The high carbon demand of foliage recovery after fall prescribed fire was associated with interruption of seasonal starch accumulation in the stem and taproot. The implication of low starch accumulation in stem and taproot tissues during the growing season after fall prescribed fire is unknown and warrants further investigation. Our results demonstrate a positive influence of residual foliage, lateral branches, and stored starch on timely foliage recovery of young longleaf pines after fire. Together with knowledge of longleaf pine development and fuel and climate conditions at the time of prescribed fire, this information will aid prescribed fire practitioners charged with maintaining longleaf pine stands of high vigor.     

Mulch type and moisture level affect pupation depth of <Emphasis Type="Italic">Rhagoletis mendax</Emphasis> Curran (Diptera: Tephritidae) in the laboratory

Mulching can be beneficial for organic highbush blueberry production, but its effects on insect pests have received little attention. For pests that pupate in soil, depth may affect pupation success due to differences in temperature and moisture, mortality factors such as predation, or efficacy of controls such as insecticidal soil drenches. We examined how mulch type and moisture affect pupation depth for Rhagoletis mendax Curran (blueberry maggot), an important pest of blueberries. In laboratory studies, pupation depth was measured in wood waste compost, uncomposted pine needles, soil and sand, each at dry, field moisture levels and wet, 99% water holding capacity conditions. Pupation occurred more deeply but with greater variability in pine needle mulch compared to compost mulch, soil, or sand. Approximately 50% of maggots pupated on the surface of wet soil, however, maggots burrowed more deeply in wet than in dry pine needles. Moisture level did not significantly affect pupation depth in compost or sand. Some larvae unexpectedly escaped the stacks of cups used to assess pupation depth or died before forming a puparium. Less than 70% of stacks with dry pine needles contained pupae, and fewer pupae were recovered from wet than dry compost and soil. These results suggest that mulch material and/or moisture levels can have significant impacts on R. mendax pupation depth with potential implications for its management.     

Regeneration patterns of Eastern white pine (Pinus strobus L.) in hardwood-dominated forests in Virginia, USA

Eastern white pine (Pinus strobus L.) is a moderately shade-tolerant species that co-occurs with hardwood tree species in many forests of the eastern United States, as well as in pure stands. The species is valued for its timber, as well as for wildlife and recreation. Regeneration of this species is somewhat unpredictable and often occurs in patches of similarly-aged cohorts. We described the regeneration patterns of this species and examined their relation to environmental variables within hardwood forests of southwestern Virginia, USA. An average of 5.3 white pine patches per ha were observed in this study. The majority of patches consisted of saplings (85%), with 9% of patches in pole size classes, and 6% in seedling size classes. The average density of patches was 43.5 stems with an average age of 20 years. The size of patches averaged 80.6 m². The total density of seedlings and the number of regeneration patches of all sizes of regeneration (seedlings, saplings, and poles) in plots was related to the surrounding density of large white pine trees (potential seed trees). The density of seedlings or patches was not significantly related to current vegetation cover or soil surface cover variables, but more than half of regeneration patches were located in or adjacent to old canopy gaps, most of which were old logging gaps. While seedling regeneration may occur within the understory of these forests near seed trees, advancement to the sapling and pole stage appears to be associated with canopy gap formation.     

Acclimation of shoot and needle morphology and photosynthesis of two Picea species to differences in soil nutrient availability

To investigate morphological acclimation to differences in nutrient availability, we compared shoot and needle morphology of Picea glehnii (Friedr. Schmidt) M. T. Mast. and Picea jezoensis (Siebold & Zucc.) Carrière trees growing on nutrient-poor volcanic ash and nutrient-rich, brown forest soil. Trees of both species were shorter and had more open canopies when growing on volcanic ash than when growing on brown forest soil. Nutrient-poor conditions limited height growth less in P. glehnii than in P. jezoensis. In both species, trees growing on volcanic ash had shorter annual increments in the previous year and more needles per shoot length and, hence, a smaller shoot silhouette area (SSA) relative to needle dry mass (NDM) than trees growing on brown forest soil. Soil type had less effect on shoot projected needle area (PNA). Total needle area (TNA) of P. glehnii shoots was similar between soil types, whereas TNA of P. jezoensis was lower in trees growing on volcanic ash than in trees growing on brown forest soil. For both species, low SSA in response to nutrient-poor conditions resulted in low shoot SSA/PNA ratios, indicating high within-shoot self-shading. Shoot SSA/TNA of P. glehnii was lower in trees growing on volcanic ash than in trees growing on brown forest soil, indicating that needles were sun-acclimated. In contrast, shoot SSA/TNA of P. jezoensis was higher in trees growing on volcanic ash than in trees growing on brown forest soil. The contrasting response of TNA to low nutrient availability was associated with species-specific differences in needle morphology. Needles of P. glehnii growing on volcanic ash were slightly shorter, wider, thicker and heavier than those of trees growing on brown forest soil, indicating morphological acclimation to high irradiance. Needles of P. jezoensis growing on volcanic ash were shorter than those of trees growing on brown forest soil, but did not show morphological acclimation to high irradiance in width, thickness or mass. For both species, nutrient-poor conditions decreased maximum photosynthetic rate (Amax) per NDM. However, when expressed per PNA, the decrease in Amax was reduced, and when expressed per SSA, Amax was higher in trees growing on volcanic ash than in trees growing on brown forest soil. On volcanic ash, Amax per NDM was lower for P. glehnii than for P. jezoensis. However, morphological changes at the shoot and needle levels reversed this trend when Amax was expressed per SSA or per PNA. The species-specific differences in morphological response to differences in soil nutrient availability suggest that P. glehnii is more tolerant of nutrient-poor conditions, whereas P. jezoensis is better at exploiting nutrient-rich soils.     

Influence of different forest conversion strategies on ground vegetation and tree regeneration in pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) stands: a case study in NE Germany

The conversion of anthropogenic into more natural, self-regenerating forests is one of the major objectives of forestry throughout Europe. In this study, we present investigations on permanent plots with different silvicultural treatment in NE German pine stands. Management of old-growth pine stands on acidic and nutrient-poor sandy sites differs in fencing, thinning, and planting of certain tree species. The investigations were carried out on the community, population, and individual level of the pine forest ecosystems. Thus, vegetation changes, size and height of tree populations, and height increment of tree individuals were observed over a time span of 6 years. Special attention was paid to short-lived tree species such as, e.g., Frangula alnus and Sorbus aucuparia, as well as to Fagus sylvatica as one of the most typical forest tree species of Central Europe. Vegetation changes are interpreted as a consequence of natural regeneration of formerly degraded forest sites, involving an increase in nutrient availability. High browsing pressure can be considered as a key factor for the inhibition of tree seedlings and growth of saplings. Some Sorbus aucuparia individuals, however, succeeded in growing out of the browsing height also in unfenced stands. Few found specimens of Fagus sylvatica proved that this species is able to establish spontaneously on these relatively dry, acidic sites under continental climate influence. Such natural regeneration processes, also including spontaneous rejuvenation of trees, can be integrated into silviculture as passive forest conversion management. An active management like thinning of stands, planting of trees, and fencing can accelerate forest conversion with regard to height growth and species number of trees.