Is the Lower Shade Tolerance of Scots Pine,Relative to Pedunculate Oak,Related to the Composition of Photosynthetic Pigments?

Foliage of Scots pine (Pinus sylvestris L.) and pedunculate oak (Quercus robur L.) was collected in a mixed pine/oak forest at canopy positions differing in radiation environment. In both species, chlorophyll (Chl) a/b ratios were higher in foliage of canopy positions exposed to higher irradiance as compared to more shaded crown layers. Throughout the growing season, pine needles exhibited significantly lower Chl a/b ratios than oak leaves acclimated to a similar photon availability. Hence, pine needles showed shade-type pigment characteristics relative to foliage of oak. At a given radiation environment, pine needles tended to contain more neoxanthin and lutein per unit of Chl than oak leaves. The differences in pigment composition between foliage of pine and oak can be explained by a higher ratio of outer antennae Chl to core complex Chl in needles of P. sylvestris which enhances the efficiency of photon capture under limiting irradiance. The shade-type pigment composition of pine relative to oak foliage could have been due to a reduced mesophyll internal photon exposure of chloroplasts in needles of Scots pine, resulting from their xeromorphic anatomy. Hence, the higher drought tolerance of pine needles could be achieved at the expense of shade tolerance.     

Nine Years of Irrigation Cause Vegetation and Fine Root Shifts in a Water-Limited Pine Forest

Scots pines (Pinus sylvestris L.) in the inner-Alpine dry valleys of Switzerland have suffered from increased mortality during the past decades, which has been caused by longer and more frequent dry periods. In addition, a proceeding replacement of Scots pines by pubescent oaks (Quercus pubescens Willd.) has been observed. In 2003, an irrigation experiment was performed to track changes by reducing drought pressure on the natural pine forest. After nine years of irrigation, we observed major adaptations in the vegetation and shifts in Scots pine fine root abundance and structure. Irrigation permitted new plant species to assemble and promote canopy closure with a subsequent loss of herb and moss coverage. Fine root dry weight increased under irrigation and fine roots had a tendency to elongate. Structural composition of fine roots remained unaffected by irrigation, expressing preserved proportions of cellulose, lignin and phenolic substances. A shift to a more negative δ¹³C signal in the fine root C indicates an increased photosynthetic activity in irrigated pine trees. Using radiocarbon (¹⁴C) measurement, a reduced mean age of the fine roots in irrigated plots was revealed. The reason for this is either an increase in newly produced fine roots, supported by the increase in fine root biomass, or a reduced lifespan of fine roots which corresponds to an enhanced turnover rate. Overall, the responses belowground to irrigation are less conspicuous than the more rapid adaptations aboveground. Lagged and conservative adaptations of tree roots with decadal lifespans are challenging to detect, hence demanding for long-term surveys. Investigations concerning fine root turnover rate and degradation processes under a changing climate are crucial for a complete understanding of C cycling.     

Root biomass and rooting profile of naturally regenerated beech in mid-elevation Scots pine woodlands

The natural establishment of shade-tolerant forest species such as beech (Fagus sylvatica L.) occurs in naturally regenerated Scots pine (Pinus sylvestris L.) woodlands that develop on former pastures and cultivated lands. To examine possible effects of underground competition in beech establishment, we studied the root biomass and the rooting profile of 53 mixed Scots pine-natural beech woodlands in French mid-elevation volcanic areas. Stands were arranged along a maturation gradient. Roots were sampled using the root-auger technique (0 to 75 cm depth every 15 cm). In addition, 23 young beeches were uprooted to study the entire root system. Total beech fine-root biomass was closely correlated with most beech aerial characteristics, (e.g., height, diameter and girth), and correlated moderately with tree age. However, it correlated poorly with basic competition indices such as stand density and basal area. Conversely, competition indices including vertical dimensions of competing trees were correlated with the underground biomass, probably as a result of redundancy with beech height. The rooting profile (fine roots, < 5 mm) of beech and pine were quite similar, and did not change significantly along the stand maturation gradient. Beech has a heart-shaped root system while pine is more plate-like and dimorphic. Beech fine-root biomass progressively surpassed pine biomass throughout the soil layers, thus confirming that it is dynamic and competitive in mature mixed stands. The coexistence of the root systems of beech and pine in the same soil layers presumably results in strong underground competition.     

Effects of aluminium and mineral nutrition on growth and chemical composition of hydroponically grown seedlings of five different forest tree species

Forest die-back and impaired tree vitality have frequently been ascribed to Al-toxicity and Al-induced nutritional disorders due to increased acidification of forest soils. Therefore, in this experiment effects of Al were studied on growth and nutrient uptake with seedlings of five different forest tree species. During growth in culture solutions with and without Al all five species proved to be very Al-tolerant, despite high accumulation of Al in roots. In the coniferous evergreens Douglas-fir and Scots pine shoot as well as root Al concentrations were significantly higher than in the deciduous broad-leaved species oak and birch. Larch showed intermediate Al levels. In none of the five species did Al reduce nutrient concentrations or the Ca/Al ratio to values below the critical level. Besides differences in Al accumulation, coniferous and broad-leaved species also differed with respect to uptake and assimilation of nitrogen. Due to extra NH ₄ ⁺ uptake, oak and birch showed a much higher N uptake and higher NH ₄ ⁺ preference than the coniferous species. Especially with oak this high NH ₄ ⁺ preference in combination with a low specific root surface area resulted in a high root proton efflux density. In comparison to both broad-leaved trees and Scots pine the NO ₃ ⁻ reduction capacity of larch and Douglas-fir was extremely low. This may have important consequences for both species if grown in NO ₃ ⁻ -rich soils.     

Quercus and Pinus cover are determined by landscape structure and dynamics in peri-urban Mediterranean forest patches

Successional dynamics in Mediterranean forests have been modulated by anthropogenic disturbances during thousands of years, especially in areas densely populated since ancient times. Our objective is to determine whether pine tree cover (early-successional species) and oak tree cover (late-successional species), used as a surrogate of successional stage of peri-urban fragmented forests in the Vallès lowlands (Catalonia, NE, Spain), are primarily determined by (1) climate and topography; (2) anthropogenic disturbances; (3) patch structure; or (4) patch dynamics from 1956 to 1993. Quercus spp. and Pinus spp. tree cover were separately recorded on 252 randomly selected plots of 100 m², within forest patches ranging in size from 0.25 to 218 ha. Multiple linear regressions indicated that forest patch history is the most important variable determining oak and pine tree cover: new forest patches showed higher pine and lower oak tree cover than recently split patches (i.e. those that became fragmented from large forest areas after 1956). Patches already existing as such in 1956 (pre-existent patches) showed higher pine cover than recently split patches. Oak cover increased and pine cover decreased with increasing forest connectivity of the patch. Finally, highly frequented forests were related to high cover of pines. Climatic and topographic variables were not significant. We conclude that pine and oak cover in these peri-urban forests are mainly determined by recent patch dynamics, but also by the spatial pattern of patches. However, human-induced disturbance can modulate this as there is some evidence for pine being associated with a high human frequentation.     

Response of understorey vegetation and Scots pine root systems to fertilization at multiple deficiency stress

The stump and root systems of Scots pine (Pinus sylvestris L.) and field-layer vegetation were sampled before and three growing seasons after drainage and fertilization of a low-shrub pine bog in SE Norway. Although the understorey vegetation roots responded significantly to nutrient application with higher concentrations of Ca and P, root biomass weights did not change. The fine and small pine roots responded with higher N, Ca, P and S concentrations, while those of Mn and Zn were significantly reduced. The NPK-application resulted in significantly higher pine root biomass. Relative to the total stores in the root zone the amounts of most elements in roots shifted to higher ratios at NPK-application. High figures for K, B and Mn indicate tight biochemical cycles of these elements. Compared to totals in above and below ground biomass, major parts of Fe and Pb were held by the roots. In contrast the field layer roots kept a very small per cent of total K, while the pine roots were low in Mn. The understorey vegetation was primarily restricted by P-deficiency, while the pine trees were also restricted by low supply of N. The field and the tree layer species seem to differ with respect to required nutrient concentrations in the root zone. These characteristics are important for direction and extent of successional changes after fertilization in low-shrub pine bog ecosystems.     

Fine Root Productivity and Turnover in Two Evergreen Central Himalayan Forests

Fine root production and mortality in central Himalayan evergreenforests consisting of Quercus leucotrichophora (banj oak) andPinus roxburghii (chir pine) were measured. Fine root productionand mortality decreased with increasing soil depth. Annual fineroot production was higher in the broadleafed forest than inthe coniferous forest, across months and seasons (1.3 and 1.5-timesmore living and dead root biomass, respectively in banj oakthan in chir pine). Live fine root production was 2508 kg ha^-1year^-1inchir pine forest and 3631 kg ha^-1year^-1in banj oak forest. Deadfine roots accumulated at a rate of 1197 and 1525 kg ha^-1year^-1inchir pine and in banj oak forest, respectively. In both forests,the greatest fine root production was recorded in the rainyseason followed by summer and winter seasons. Both soil androot nitrogen concentration decreased with increasing soil depth.Nitrogen uptake was higher in banj oak forest (12.1 kg ha^-1year^-1)than chir pine forest (7.2 kg ha^-1year^-1).Copyright 1999 Annalsof Botany Company Fine root production, fine roots, necromass, banj oak, chir pine, Quercus leucotrichophora , Pinus roxburghii .     

Estimating the relative nutrient uptake from different soil depths in Quercus robur, Fagus sylvatica and Picea abies

The distribution of fine roots and external ectomycorrhizal mycelium of three species of trees was determined down to a soil depth of 55 cm to estimate the relative nutrient uptake capacity of the trees from different soil layers. In addition, a root bioassay was performed to estimate the nutrient uptake capacity of Rb⁺ and NH₄⁺ by these fine roots under standardized conditions in the laboratory. The study was performed in monocultures of oak (Quercus robur L.), European beech (Fagus sylvatica L.) and Norway spruce [Picea abies (L.) Karst.] on sandy soil in a tree species trial in Denmark. The distribution of spruce roots was found to be more concentrated to the top layer (0–11 cm) than that of oak and beech roots, and the amount of external ectomycorrhizal mycelia was correlated to the distribution of the roots. The uptake rate of [⁸⁶Rb⁺] by oak roots declined with soil depth, while that of beech or spruce roots was not influenced by soil depth. In modelling the nutrient sustainability of forest soils, the utilization of nutrient resources in deep soil layers has been found to be a key factor. The present study shows that the more shallow-rooted spruce can have a similar capacity to take up nutrients from deeper soil layers than the more deeply rooted oak. The distribution of roots and mycelia may therefore not be a reliable parameter for describing nutrient uptake capacity by tree roots at different soil depths.     

松材线虫侵染的马尾松人工林细根形态及生物量分异特征

中龄林的马尾松受松材线虫侵染后,林木生长、生理生化指标、群落多样性等会发生异质性变化,但是,针对患病林木地下细根的响应尚不清楚。本研究以松材线虫疫区患病马尾松和健康马尾松为研究对象,采用土柱法,分0-15 cm和15-30 cm土层,对细根进行分级研究,定量分析1-5级细根的形态、生物量以及养分元素,探讨松材线虫侵染的马尾松人工林细根形态、生物量以及养分元素的分异特征。结果表明:(1)患病马尾松人工林细根的健康状态与根长密度、生物量呈极显著正相关(P<0.01),低级根(如1级根)患病后,响应会更加强烈。(2)马尾松人工林患病后,细根有效磷、速效钾浓度会显著降低(P<0.05),而全氮、钙浓度会显著升高(P<0.05)。(3)松材线虫病使林分的土壤有机质含量显著高于健康林分(P<0.05),而土壤速效钾含量会显著低于健康林分(P<0.05)。以上结果表明,松材线虫侵染的马尾松人工林会在细根形态、细根养分和土壤养分上会发生特异性响应,揭示了松材线虫病对马尾松人工林地下细根的影响,旨在为松材线虫病防治提供一定参考。     

Above- and belowground organic matter storage and production in a tropical pine plantation and a paired broadleaf secondary forest

The distribution of tree biomass and the allocation of organic matter production were measured in an 11-yr-old Pinus caribaea plantation and a paired broadleaf secondary forest growing under the same climatic conditions. The pine plantation had significantly more mass aboveground than the secondary forest (94.9 vs 35.6 t ha^-1 for biomass and 10.5 vs 5.0 t ha^{-1 for litter}), whereas the secondary forest had significantly more fine roots (⩽2 mm diameter) than the pine plantation (10.5 and 1.0 t ha^-1, respectively). Standing stock of dead fine roots was higher than aboveground litter in the secondary forest. In contrast, aboveground litter in pine was more than ten times higher than the dead root fraction. Both pine and secondary forests had similar total organic matter productions (19.2 and 19.4 t ha^-1 yr^-1, respectively) but structural allocation of that production was significantly different between the two forests; 44% of total production was allocated belowground in the secondary forest, whereas 94% was allocated aboveground in pine. The growth strategies represented by fast growth and large structural allocation aboveground, as for pine, and almost half the production allocated belowground, as for the secondary forest, illustrate equally successful, but contrasting growth strategies under the same climate, regardless of soil characteristics. The patterns of accumulation of organic matter in the soil profile indicated contrasting nutrient immobilization and mineralization sites and sources for soil organic matter formation.     

Annual and seasonal changes in fine root biomass of a Quercus ilex L. forest

The biomass, production and mortality of fine roots (roots with diameter <2.5 mm) were studied in a typical Mediterranean holm oak (Quercus ilex L.) forest in NE Spain using the minirhizotron methodology. A total of 1212 roots were monitored between June of 1994 and March of 1997. Mean annual fine root biomass in the holm oak forest of Prades was 71±8 g m^–2 yr^–1. Mean annual production for the period analysed was 260+11 g m^–2 yr^–1. Mortality was similar to production, with a mean value of 253±3 g m^–2 yr^–1. Seasonal fine root biomass presented a cyclic behaviour, with higher values in autumn and winter and lower in spring and summer. Production was highest in winter, and mortality in spring. In summer, production and mortality values were the lowest for the year. Production values in autumn and spring were very similar. The vertical distribution of fine root biomass decreased with increasing depth except for the top 10–20 cm, where values were lower than immediately below. Production and mortality values were similar between 10 and 50 cm depth. In the 0–10 cm and the 50–60 cm depth intervals, both production and mortality were lower.     

Variation in fine root biomass of three European tree species: Beech (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.), and Scots pine (Pinus sylvestris L.)

Abstract

Fine roots (<2 mm) are very dynamic and play a key role in forest ecosystem carbon and nutrient cycling and accumulation. We reviewed root biomass data of three main European tree species European beech, (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.) and Scots pine (Pinus sylvestris L.), in order to identify the differences between species, and within and between vegetation zones, and to show the relationships between root biomass and the climatic, site and stand factors. The collected literature consisted of data from 36 beech, 71 spruce and 43 pine stands. The mean fine root biomass of beech was 389 g m^?2, and that of spruce and pine 297 g m^?2 and 277 g m^?2, respectively. Data from pine stands supported the hypothesis that root biomass is higher in the temperate than in the boreal zone. The results indicated that the root biomass of deciduous trees is higher than that of conifers. The correlations between root biomass and site fertility characteristics seemed to be species specific. There was no correlation between soil acidity and root biomass. Beech fine root biomass decreased with stand age whereas pine root biomass increased with stand age. Fine root biomass at tree level correlated better than stand level root biomass with stand characteristics. The results showed that there exists a strong relationship between the fine root biomass and the above-ground biomass.     

Elevated atmospheric CO2 increases fine root production, respiration, rhizosphere respiration and soil CO2 efflux in Scots pine seedlings

In this study, we investigated the impact of elevated atmospheric CO₂ (ambient + 350 μmol mol^–1) on fine root production and respiration in Scots pine (Pinus sylvestris L.) seedlings. After six months exposure to elevated CO₂, root production measured by root in-growth bags, showed significant increases in mean total root length and biomass, which were more than 100% greater compared to the ambient treatment. This increased root length may have lead to a more intensive soil exploration. Chemical analysis of the roots showed that the roots in the elevated treatment accumulated more starch and had a lower C/N-ratio. Specific root respiration rates were significantly higher in the elevated treatment and this was probably attributed to increased nitrogen concentrations in the roots. Rhizospheric respiration and soil CO₂ efflux were also enhanced in the elevated treatment. These results clearly indicate that under elevated atmospheric CO₂ root production and development in Scots pine seedlings is altered and respiratory carbon losses through the root system are increased.     

Nutritional disturbances of young Scots pines caused by pine bark bugs in a dry heath forest

This study deals with the effect of pine bark bugs (Aradus cinnamomeus Panzer) on the nutrition of young Scots pines (Pinus sylvestris L.). Soil and needle samples for analytical purposes were collected from a young pine stand growing on a dry barren mineral soil afflicted by pine bark bugs.The damage to vascular tissues caused by pine bark bugs disturbed the nutrition of the trees, especially in the top part of the crown. The foliar calcium, magnesium, manganese and sulphur concentrations were highly reduced. Scots pines suffered from a lack of calcium, magnesium, nitrogen and phosphorus. These deficiencies were secondary by nature, because no differences were observed between the nutrient concentrations of the underlying soil of the healthy and affected trees. The symptoms of trees damaged by pine bark bugs resembled most of all calcium deficiency symptoms.     

Growth and nutrient characteristics in bog and fen populations of Scots pine (Pinus sylvestris)

Nutrient content in peat and growth rate, rate of nutrient accumulation and allocation patterns in Scots pine Pinus sylvestris L. from eleven natural Swedish peatlands were examined. The peatlands studied represented a wide range of climatic conditions and mire types. Whole and even-sized pines with intact root-systems were excavated to give the whole-pine budget for growth and nutrient accumulation. All samples originated from hummock communities.Pine growth and nutrient characteristics were much more variable in the minerogenous sites than in the ombrogenous sites, which indicates a larger environmental heterogeneity within the minerogenous sites. In the ombrogenous sites, rate of pine growth was constant, approximately 1 mg day^-1, and independent of latitudinal variation. There was either no relationship between latitudinal location and growth rate in the minerogenous sites, which suggests that pine growth is largely controlled by site-specific, very local conditions. The growth rate of pines was not correlated with any peat nutrient. The pines allocated a large proportion of their nutrient-pool to the metabolically active current year's growth. This is likely a trait that enables Scots pine to occupy a wide range of peatland types in which it experience a marked imbalance and shortage of nutrients.     

Response of Pinus sylvestris L. to recent climatic events in the French Mediterranean region

Exceptional climatic events from 2003 to 2005 (scorching heat and drought) affected the whole of the vegetation in the French Mediterranean region and in particular the Scots pine (Pinus sylvestris L.), one of the most important forest tree species in this area. To understand its response to these extreme conditions, we investigated its radial growth, branch length growth, architectural development and reproduction for the period 1995–2005, and linked these variables to climatic parameters. We used four plots situated in southeastern France and presenting different levels of site quality and potential forest productivity. The results show that: (1) the climatic episode 2003–2005 was highly detrimental to the growth (bole and branches), crown development, and cone production but favored the production of male flowers; (2) these variables depend on climatic factors of both the current and previous years; (3) the 2003 scorching heat impact was strong but was mainly apparent from 2004; it was part of a 6-year-long unfavorable cycle beginning in 2000, characterized by high minimum and maximum temperatures and very dry springs; (4) in spite of a significant effect of site quality, the Scots pine’s response to extreme climatic conditions was homogeneous in the French Mediterranean area; and (5) the stress induced by poor site conditions generally resulted in the same consequences for tree growth, architecture, and reproduction as in unfavorable climatic conditions.     

Biomass allocation and growth responses of Scots pine saplings to simulated herbivory depend on plant age and light availability

This study experimentally analyses the response to simulated herbivory of juvenile Scots pine of two different ages in contrasting abiotic scenarios, focusing on the potential dual role of browsing ungulates: negative, by removing aerial biomass, and positive, by stimulating compensation capacity and providing nutrients by depositing their excrement. Compensation against herbivory was investigated by experimentally clipping a set of Scots pine (Pinus sylvestris L. nevadensis) juveniles, grown under different levels of light and nutrient availability. The responses analysed were survival, trunk-diameter growth, leader-shoot growth, increment in number of meristems, RGR, biomass of needles, shoots, root and whole plant, and root-to-shoot ratio. Clipping consistently resulted in a worse survival and performance of pines with respect to unclipped ones. From the factors analysed, light availability was responsible mainly for the variations in plant performance, while the addition of nutrients was much less important. Age was also important, with older pines showing in general better performance after clipping. Overall, clipping invariably had a negative effect on Scots pine, since none of the combinations of abiotic factors used resulted in overcompensation. However, the intensity on this negative effect proved quite variable, from almost an exact compensation in clipped older pines under full sunlight availability to very poor performance and high death probability in younger pines in shade. Scots pine cannot overcompensate after clipping, but, depending of the environmental conditions, the negative result of clipping varies from severe undercompensation to almost exact compensation. Also, small differences in sapling age can promote significant differences in sapling response to clipping and light environment.     

Interactive effects of mycorrhization and elevated carbon dioxide on growth of young pedunculate oak (Quercus robur L.) trees

Pedunculate oak (Quercus robur L.) was germinated and grown at ambient CO₂ level and 650 ppmv CO₂ in the presence and absence of the ectomycorrhizal fungus Laccaria laccata for a total of 6 month under nutrient non-limiting conditions. Mycorrhization and elevated atmospheric CO₂ each supported the growth of the trees. Stem height, stem diameter, and dry matter accumulation of pedunculate oak were increased by mycorrhization. Elevated atmospheric CO₂ enhanced stem height, stem diameter, fresh weight and dry weight, as well as lateral root formation of the trees. In combination, mycorrhization and elevated atmospheric CO₂ had a more than additive, positive effect on tree height and biomass accumulation, and further improved lateral root formation of the trees. From these findings it is suggested that the efficiency of the roots in supporting the growth of the shoot is increased in mycorrhized oak trees at elevated atmospheric CO₂.Abbreviations DW dry weight - FW fresh weight - RWC relative water content     

杉木叶片、细根功能性状对毛竹扩张及伐除的响应

功能性状能够反映植物对不同环境的适应策略。毛竹扩张与外来植物入侵相似,常引起原有植物生存环境的改变,而原有植物功能性状对毛竹扩张及伐除的响应机制尚不清楚。选取毛竹-杉木混交林和去竹杉木林为研究对象,以杉木纯林为对照,比较分析杉木比叶面积、叶干物质含量、叶组织密度等叶功能性状以及比根长、细根生物量、细根根长密度等细根功能性状的变化以及其间的相关关系。结果表明：(1)与杉木纯林相比,混交林中杉木的叶相对含水量以及叶干物质含量分别减少了5.07%、0.032 g/g,叶组织密度以及比叶面积分别增加了0.005 g/cm³、10.33 cm²/g;而去竹杉木林中,杉木比叶面积、叶相对含水量减少,叶干物质含量和叶组织密度则呈上升趋势。(2)与杉木纯林相比,混交林中杉木细根生物量、细根体积密度以及细根根长密度都不断下降,而杉木细根比根长在0—20 cm土深处显著增加(P<0.05);而去竹杉木林中杉木细根比根长、细根根长密度和细根生物量则显著降低(P<0.05),细根体积密度在20—30 cm土深处有所增加。(3)杉木纯林中杉木细根功能性状间...     

Asynchrony in larval development of the pine beauty moth, Panolis flammea, on an introduced host plant may affect parasitoid efficacy

In the United Kingdom, Panolis flammea (Den. and Schiff.) (Lepidoptera: Noctuidae) is an important pest species of the introduced lodgepole pine but not of its natural host Scots pine. The timing of P. flammea larval growth must be synchronized with its host tree if the larvae are to succeed. We collected field data during 1990 which revealed that the phenological window starts earlier in Scots pine and is shorter than that observed in lodgepole pine. The larvae are found in the field earlier and within a narrower time frame within a Scots pine forest than in a lodgepole pine forest. The larval developmental period is significantly longer on lodgepole pine than on Scots pine. The synchrony/asynchrony of P. flammea to its natural host (Scots pine) and an introduced tree (lodgepole pine) results in the parasitoids having a different impact on the larvae of the two hosts. At any one time, the host plant, caterpillars and parasitoids are more synchronous on the ancestral Scots pine than on lodgepole pine, resulting in a higher percentage of larvae in the optimal instar for parasitism at that time. In lodgepole pine, the percentage of suitable instars available to parasitoids is lower at any given time. The information presented here furthers our understanding of the possible mechanisms for the observed differential population dynamics of the insect on Scots pine and lodgepole pine in the UK. Handling editor: Robert Glinwood.