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1.
Summary The effects of a number of factors, notably leaf mining insects, on the longevity of beech and holm oak leaves have been studied. The regular monitoring of individually labelled leaves was complemented by analysis of leaf fall data. Both methods confirm that these mining insects have only a slight impact on their host trees. The presence of first generation Phyllonorycter maestingella mines on beech leaves and winter generation P. messaniella mines on holm oak leaves accelerates leaf loss. Beech leaves mined by second generation P. maestingella and Rhynchaenus fagi did not show this accelerated loss. Their patterns of leaf fall can be explained by within-tree variation in both mine distribution and the timing of leaf fall. It is argued that this premature leaf fall is a damage response, and is not an attempt by the tree to regulate miner numbers.  相似文献   

2.
Larvae of Epiphyas postvittana and Planotortrix octo were released onto branches cut from apple trees, and allowed to colonize a range of types of artificial nests. Both species exhibited similar strong preferences for nests comprising leaf-leaf or leaf-fruitlet combinations, followed by nests comprising leaf-plastic leaf, leaf-plastic fruitlet or plastic leaf-fruitlet combinations. Nests involving fresh plant material alone (shoot, fruitlet or leaf alone) were also colonized to a lesser extent, but no larvae were found on nests consisting of plastic leaves or fruitlets alone or in combinations of the two. In another experiment, more E. postvittana larvae colonized nests with leaf and fruitlet combinations, compared to leaf and glass ball, or leaf and treated wax ball combinations, where the wax had been in contact with fresh apples or fruitlets. Choice tests, conducted using larval traps, showed that larvae were caught in traps baited with odors collected and released by wax which had been in contact with mature apples and leaves. Chloroform extracts from apple skin also caught larvae in choice tests. These results suggest that both physical and chemical cues are important to leafroller larval establishment.  相似文献   

3.
The northern tamarisk beetle (Diorhabda carinulata Desbrochers) was released in several western states as a biocontrol agent to suppress Tamarix spp. L. which has invaded riparian ecosystems; however, effects of beetle herbivory on Tamarix physiology are largely undocumented and may have ecosystem ramifications. Herbivory by this insect produces discoloration of leaves and premature leaf drop in these ecosystems, yet the cause of premature leaf drop and the effects of this leaf drop are still unknown. Insect herbivory may change leaf photosynthesis and respiration and may affect a plant’s ability to regulate water loss and increase water stress. Premature leaf drop may affect plant tissue chemistry and belowground carbon allocation. We conducted a greenhouse experiment to understand how Tamarix responds physiologically to adult beetle and larvae herbivory and to determine the proximate cause of premature leaf drop. We hypothesized that plants experiencing beetle herbivory would have greater leaf and root respiration rates, greater photosynthesis, increased water stress, inefficient leaf nitrogen retranslocation, lower root biomass and lower total non-structural carbohydrates in roots. Insect herbivory reduced photosynthesis rates, minimally affected respiration rates, but significantly increased water loss during daytime and nighttime hours and this produced increased water stress. The proximate cause for premature leaf drop appears to be desiccation. Plants exposed to herbivory were inefficient in their retranslocation of nitrogen before premature leaf drop. Root biomass showed a decreasing trend in plants subjected to herbivory. Stress induced by herbivory may render these trees less competitive in future growing seasons.  相似文献   

4.
Habitat fragmentation is currently the most pervasive anthropogenic disturbance in tropical forests and some species of leaf‐cutting ants of the genus Atta (dominant herbivores in the neotropics) have become hyper‐abundant in forest edges where their nests directly impact up to 6% of the forest area. Yet, their impacts on the regeneration dynamics of fragmented forests remain poorly investigated. Here we examine the potential of Atta cephalotes nests to function as ecological filters impacting tree recruitment. Growth, survival and biomass partitioning of experimentally planted seedlings (six tree species) were examined at eight spatially independent A. cephalotes colonies in a large Atlantic Forest fragment. Seedling performance and fate (leaf numbers and damage) were monitored up to 27 months across three habitats (nest centre, nest edge and forest understorey). Plants at illuminated nest centres showed twice the gross leaf gain as understorey individuals. Simultaneously, seedlings of all species lost many more leaves at nests than in the forest understorey, causing a negative net leaf gain. Net leaf gain in the shaded understorey ranged from zero (Licania and Thyrsodium species) to substantial growth for Copaifera and Virola, and intermediate levels little above zero for Protium and Pouteria. Also seedling survival differed across habitats and species, being typically low in the centre and at the edge of nests where seedlings were often completely defoliated by the ants. Lastly, seedling survival increased strongly with seed size at nest edges while there was no such correlation in the forest. Our results suggest that Atta nests operate as ecological filters by creating a specific disturbance regime that differs from other disturbances in tropical forests. Apparently, Atta nests favour large‐seeded tree species with resprouting abilities and the potential to profit from a moderate, nest‐mediated increase in light availability.  相似文献   

5.
To evaluate the responses of Quercus crispula and Quercus dentata to herbivory, their leaves were subjected to simulated herbivory in early spring and examined for the subsequent changes in leaf traits and attacks by chewing herbivores in mid summer. In Quercus crispula, nitrogen content per area was higher in artificially damaged leaves than in control leaves. This species is assumed to increase the photosynthetic rate per area by increasing nitrogen content per area to compensate leaf area loss. In Quercus dentata, nitrogen content per area did not differ between artificially damaged and control leaves, while nitrogen content per mass was slightly lower in artificially damaged leaves. The difference in their responses can be attributable to the difference in the architecture of their leaves and/or the severeness of herbivory. The development of leaf area from early spring to mid summer was larger in artificially damaged leaves than in control leaves in both species, suggesting the compensatory response to leaf area loss. Leaf dry mass per unit area was also larger in artificially damaged leaves in both species, but the adaptive significance of this change is not clear. In spite of such changes in leaf traits, no difference was detected in the degree of damage by chewing herbivores between artificially damaged and controlled leaves in both species.  相似文献   

6.
The rate of decomposition of summer leaf-fall (abscised leaves), winter leaf-fall (containing some green leaves) and mature green (picked) leaves was assessed in sub-alpine forests of E. delegatensis (R. T. Baker), E. pauciflora (Sieb. ex Spreng) and E. dives (Schau.) in the Brindabella Range, Australian Capital Territory, using litter bag and tethered leaf techniques. The relative contribution of leaching, microbial respiration and grazing by invertebrate macrofauna to loss of leaf weight was determined. The effect of leaching and microbial respiration was assessed in terms of weight loss per unit area of leaf (specific leaf weight), while losses due to macro-faunal grazing were assessed by measuring reductions in leaf area. Litter decomposition constants for litter components (leaf, bark, wood) and total litter were determined from long-term records of litterfall and accumulated litter. Weight losses of abscised leaves during the initial 12 months ranged from 25% for E. pauciflora to 39% for E. delegatensis and were almost entirely due to reduction in specific leaf weight. Losses in the weight of leaves falling in winter ranged from 38 to 49%, while green leaves lost 45 - 59%. Approximately 50% of the total weight loss of green leaves was due to a loss in leaf area caused by skeletonization by litter macrofauna. Thus abscised leaves rather than green leaves must be used for measuring litter decomposition rates since abscised leaves constitute most of the litterfall in eucalypt forests. Leaves placed in the field in autumn decomposed slowly during the first summer, while the rate increased during the second winter and summer. Low litter moisture content appears to limit decomposition in the initial summer period in all communities, after which litterfall provides a mulch which reduces the rate of desiccation of lower litter layers. A simple linear regression model relating decomposition rate to the number of days (D) when litter moisture content exceeded 60% ODW accounted for 63-83% of the variation in decomposition of leaves in the field. Inclusion of mean monthly air temperature (T) and the product of D and T (day degrees when litter was wet) in a multiple linear regression increased the variation in decomposition accounted for to 80 – 90%. The rate of weight loss showed a positive linear relationship with the initial concentration of nitrogen (N) or phosphorus (P) in the leaf. These concentrations are an index of the decomposability of leaf substrates (e.g. degree of sclerophylly or lignification). The rate of loss of specific weight was similar for tethered leaves and for leaves enclosed in mesh bags. Measured loss in specific leaf weight after 70 – 90 weeks was less than that predicted using decomposition constants (k).  相似文献   

7.
Leaf decomposition in an experimentally acidified stream channel   总被引:3,自引:2,他引:1  
Decomposition of Alnus rugosa and Myrica Gale leaves immersed in artificial stream channels fed by a small headwater creek was followed over a three month period. At the end of experiment, remaining weights of both leaf types confined in litter bags were significantly higher after immersion in experimentally acidified water (pH 4.0) than when immersed in control water (pH 6.2–7.0). For both types of leaves and for all sampling times, there was generally no difference in the C:N ratios between leaves in acidified and those in control water. In control water, oxygen uptake by microorganism on A. rugosa leaves was significantly higher after 46 days of immersion, whereas differences between treatments appeared only after 69 days for M. Gale leaves. Transfer of A. rugosa leaves from acid to control water led to a rapid increase in microbial activity; this increased activity was reflected in a fast weight loss of the leaves. For both leaf types, total numbers of macroinvertebrates were usually higher in litter bags immersed in control water. Macroinvertebrates colonizing the litter bags were mainly collector-gatherers: Chironomidae were numerically dominant in control leaf packs whereas Oligochaeta dominated in acid leaf packs. Macroinvertebrate biomass in M. Gale litter was higher in control than in acidified water, which contrasted with macroinvertebrate biomass in A. rugosa leaf packs which was not significantly different between treatments. Macroinvertebrate contribution to the breakdown of leaf litter was thus considered less important than the microbial contribution. This study demonstrated that decomposition of leaf litter in acidic headwater streams can be seriously reduced, mainly as a result of a lower microbial activity.  相似文献   

8.
Symptoms of circular leaf spot of persimmon (CLSP), caused by Mycosphaerella nawae, consisted of necrotic spots on leaves, chlorosis and premature defoliation. Although CLSP is a foliar disease, early fruit maturation and abscission are frequently associated with the presence of lesions on leaves and defoliation, resulting in severe economic losses. Despite their importance for the design of efficient disease management programmes, quantitative relationships between CLSP incidence and yield loss are unknown. Therefore, fungicide efficacy trials were conducted during two consecutive years in Spain to induce different levels of disease severity, defoliation and yield loss. The effects of fungicide treatments on CLSP severity were analysed by ordinal logistic regression models. Relative yield loss values were regressed against the percentage of affected leaves or defoliated obtained at different evaluation dates. The disease had high negative impact and complete yield loss was observed in the absence of effective fungicide treatments. Preventive applications of pyraclostrobin, trifloxy‐strobin and mancozeb provided the best disease control and highest yields, up to 95.77 kg tree?1. An exponential relationship of CLSP incidence and defoliation with yield loss was found. In general, model fit and predictive ability was superior when defoliation, rather than incidence, was used as explanatory variable. The impact of defoliation on yield loss was higher in earliest evaluation dates, suggesting that early leaf abscission may be the main factor contributing to premature fruit drop and subsequent yield loss. Substantial yield losses were observed even with relatively low levels of CLSP incidence and defoliation. Therefore, it was not possible to define a critical action threshold for CLSP management based on foliar symptoms.  相似文献   

9.
Summary Mesophyll cell nuclei isolated from leaves of Pennisetum purpureum were analysed by flow cytometry to determine the nuclear DNA content and the percentage of cells in different phases of the cell cycle. Samples taken from base, middle and tip regions of leaves 2 to 8 (leaf 1, which was adjacent to the meristem, was too small to sample) showed no significant differences in the amount of DNA per G1 nucleus due to either age or position. The average amount of DNA per G1 nucleus was 5.78 pg. Although the majority of cells for each sample were in G1, samples taken from older leaves had higher percentages of cells in G2 and S phases. More specifically, base and middle regions of older leaves had a higher percentage of cells in G2 than all three positions in younger leaves. Electrophoretic analysis of nuclear DNA from leaves 2 to 7 showed no evidence of degradation or difference in fragment size for any sample or position. This study was compared to previous work on the relationship between leaf age and embryogenic competence in Pennisetum purpureum. The results suggest that changes in the cell cycle, and/or a loss or fragmentation of the nuclear DNA, are not responsible for loss of embryogenic competence in mature leaf tissue.  相似文献   

10.
The leaf miner Cameraria ohridella causes premature defoliation of Aesculus hippocastanum trees. In order to assess the whole-plant loss of productivity caused by the parasite, we monitored seasonal changes of leaf gas exchange and leaf area losses in horse chestnut trees freely infested or chemically treated to prevent moth infestation (controls). Data were integrated in a model and the annual loss of net primary productivity (NPP) was calculated for infested trees with respect to controls. Measurements showed marked vertical stratification of C. ohridella attacks, with lower crown strata being more infested than higher ones. Leaf gas exchange was maximum between May and early June, but it strongly decreased starting from mid-June even in controls. Model calculations showed that NPP loss of infested trees was about 30% on an annual basis (when the first moth attack is recorded at the end of April). Model simulations showed that postponing the start day of attack would have important positive effects on plants NPP. For example, if the start day of attack was postponed to 20 May, the annual loss of NPP would be about 15%. Our study suggests that A. hippocastanum trees attacked by C. ohridella are not facing serious risks of decline, especially if methods are adopted to postpone the start day of attack (e.g. removal of fallen leaves in autumn). Our data do not support the view that plants need to be totally protected from the parasite by application of insecticides.  相似文献   

11.
Leaf and stem functional traits related to plant water relations were studied for six congeneric species pairs, each composed of one tree species typical of savanna habitats and another typical of adjacent forest habitats, to determine whether there were intrinsic differences in plant hydraulics between these two functional types. Only individuals growing in savanna habitats were studied. Most stem traits, including wood density, the xylem water potential at 50% loss of hydraulic conductivity, sapwood area specific conductivity, and leaf area specific conductivity did not differ significantly between savanna and forest species. However, maximum leaf hydraulic conductance (K leaf) and leaf capacitance tended to be higher in savanna species. Predawn leaf water potential and leaf mass per area were also higher in savanna species in all congeneric pairs. Hydraulic vulnerability curves of stems and leaves indicated that leaves were more vulnerable to drought-induced cavitation than terminal branches regardless of genus. The midday K leaf values estimated from leaf vulnerability curves were very low implying that daily embolism repair may occur in leaves. An electric circuit analog model predicted that, compared to forest species, savanna species took longer for their leaf water potentials to drop from predawn values to values corresponding to 50% loss of K leaf or to the turgor loss points, suggesting that savanna species were more buffered from changes in leaf water potential. The results of this study suggest that the relative success of savanna over forest species in savanna is related in part to their ability to cope with drought, which is determined more by leaf than by stem hydraulic traits. Variation among genera accounted for a large proportion of the total variance in most traits, which indicates that, despite different selective pressures in savanna and forest habitats, phylogeny has a stronger effect than habitat in determining most hydraulic traits.  相似文献   

12.
Indirect effects of insect herbivory on leaf gas exchange in soybean   总被引:5,自引:0,他引:5  
Herbivory can affect plant carbon gain directly by removing photosynthetic leaf tissue and indirectly by inducing the production of costly defensive compounds or disrupting the movement of water and nutrients. The indirect effects of herbivory on carbon and water fluxes of soybean leaves were investigated using gas exchange, chlorophyll fluorescence and thermal imaging. Herbivory by Popillia japonica and Helicoverpa zea (Boddie) caused a 20–90% increase in transpiration from soybean leaflets without affecting carbon assimilation rates or photosynthetic efficiency (ΦPSII). Mechanical damage to interveinal tissue increased transpiration up to 150%. The spatial pattern of leaf temperature indicated that water loss occurred from injuries to the cuticle as well as from cut edges. A fluorescent tracer (sulforhodamine G) indicated that water evaporated from the apoplast approximately 100 µm away from the cut edges of damaged leaves. The rate of water loss from damaged leaves remained significantly higher than from control leaves for 6 d, during which time they lost 45% more water than control leaves (0.72 mol H2O per cm of damaged perimeter). Profligate water loss through the perimeter of damaged tissue indicates that herbivory may exacerbate water stress of soybeans under field conditions.  相似文献   

13.
A wilt disease of garden pea (Pisum sativum) caused by Verticillium dahliae is described and the range of pathogenicity of the isolate investigated. It is pathogenic to potato, sweet pea, antirrhinum and broad bean and isolates of V. dahliae from potato, lucerne and sweet pea and V. albo-atrum from lucerne are pathogenic to pea. Since the most common disease symptoms, acropetal progression of chlorosis and necrosis of the leaves followed by premature defoliation are indistinguishable from natural senescence, it is probable that disease and senescence symptoms are confused in the field. The premature defoliation results in marked reduction in green leaf area, leaf dry weight and pod yield.  相似文献   

14.
Non-random leaf orientation in Lactuca serriola L.   总被引:2,自引:2,他引:0  
Abstract Leaf shapes and leaf orientation of Lactuca serriola serriola and serriola integrifolia were studied. Leaf shapes in L. serriola serriola differed greatly from those of L.serriola integrifolia, but leaf surface areas were similar. In exposed habitats, leaf orientation of cauline leaves of both forms was non-random, with leaves almost vertical and tending to orient with their lamina normal to the east and west. In the shade, cauline leaves oriented randomly. An experiment demonstrated that the orientation of leaves did not change significantly once they were fully expanded. The leaf orientation in L. serriola affected the diurnal distribution of solar irradiance intercepted by a leaf. Peak solar radiation fluxes are incident on the rosette leaves at midday, but on the cauline leaves the peak solar radiation flux occurs early in the morning and again late in the afternoon. The significance of this unusual leaf orientation is discussed in relation to water loss and carbon gain.  相似文献   

15.
The effect of increasing spore concentration of Alternaria solani (Early blight disease) on the shoot growth of young tomato plants was analysed. Changes in growth were related to the severity of infection which increased with increasing inoculum. Leaf production was not affected but dry weights and especially leaf expansion were decreased. The effective leaf areas of the five inoculated leaves (L1-L5 numbered from the plant base) were drastically decreased by expanding necrotic lesions and, to a lesser extent, by premature leaf fall. Healthy leaves expanding soon after inoculation (L6, L7) were markedly affected by the disease on the lower leaves and had decreased specific leaf areas (ratio of leaf area to leaf dry weight) but later formed (from L8) leaves were less affected and had greater specific leaf areas than equivalent leaves on uninoculated plants.  相似文献   

16.
Leaf mass loss in wetland graminoids during senescence   总被引:2,自引:0,他引:2  
Mass loss of senescing leaves is an important part of plant biomass turnover and has consequences for assessment of ecosystem productivity, ecosystem nutrient use efficiency, and plant nutrient resorption efficiency. Data, however, on mass loss are scarce, and often based on leaf area as the reference base. This leads to an underestimation of the mass loss, as leaf area itself shrinks during senescence. Furthermore, the few existing studies have almost exclusively used woody species. The purpose of the present study was twofold: i) to assess leaf mass loss during senescence in herbaceous species, with the example of five wetland graminoids and, ii) to compare two different methods of mass loss assessment (two species). Assuming that leaf length does not change during senescence, we assessed leaf mass per leaf length prior to and after senescence. We also estimated pre‐senescence leaf mass nondestructively based on leaf length, width and thickness. For Typha latifolia and Carex stricta, two species with graminoid type leaves but contrasting leaf structure, both methods delivered almost identical results. After the first assessment of leaf mass on July 7th, T. latifolia leaf mass initially increased by 13%, and then decreased to be 12% below the original mass after senescence. C. stricta leaf mass remained stable until senescence, but decreased then by 33%. In a second experiment, the mass of 100 mm pieces of leaves was measured before and after senescence. Calamagrostis canadensis, Carex rostrata and C. stricta lost 23–57% of their leaf mass during senescence, whereas Glyceria canadensis did not show any mass loss. We conclude that mass loss of senescing leaves of herbaceous plants can be considerable and should not be neglected in studies of productivity, nutrient use efficiency or nutrient resorption. For species with no shrinking leaf length during senescence, mass loss can be measured with leaf length as the base whereas for others, pre‐senescent mass can be estimated on the basis of leaf dimensions.  相似文献   

17.
Leaves of Triticum aestivum cv. Avalon were grown in an atmosphere that contained 150 nmole mol-1 ozone for 7h each day. After leaves had reached maximum size, the leaf blade was divided into three sections to provide tissue of different age, the youngest at the base of the blade and the oldest at the leaf tip. The ozone treatment was found to decrease significantly the light-saturated rate and quantum yield of CO2 assimilation and the maximum quantum yield of photosystem II photochemistry in the oldest leaf section. No effects were found on the basal and middle sections of the leaf. These ozone-induced decreases in the photosynthetic parameters were associated with decreases in the efficiency of utilization of light for CO2 assimilation at the photon flux density under which the leaves were grown. The depression in photosynthetic performance of tissue near the leaf tip was accompanied by large decreases in the contents of total, soluble and thylakoid proteins and chlorophyll. There was also found to be a preferential loss of ribulose-1,5-carboxylase-oxygenase. These ozone-induced changes in chlorophyll and protein contents and the photosynthetic activities of the leaf tissue were similar to changes normally associated with leaf senescence. Two-dimensional polyacrylamide gel analyses of leaf proteins demonstrated the loss of some minor, and unidentified, proteins, whilst another group of minor proteins appeared. It is concluded that daily exposure of the leaf to 150 nmol mol-1 ozone for 7h had no effect on the development of the photosynthetic apparatus and its activities during leaf expansion, but it did promote the onset of premature senescence in fully expanded tissue that resulted in a loss of pigments, proteins and photosynthetic capacity and efficiency.  相似文献   

18.
Nutrient addition accelerates leaf breakdown in an alpine springbrook   总被引:8,自引:0,他引:8  
This study assessed the effect of nutrient enrichment on organic matter breakdown in an alpine springbrook, using alder leaf packs to which phosphorus and nitrogen were added in the form of slow-release fertilizer briquettes. The breakdown of leaf packs with nutrients added (k=0.0284 day–1) was significantly faster than that of unfertilized packs (k=0.0137 day–1), resulting in a 30% higher mass loss after 42 days. Unfertilized leaves enclosed in fine-mesh bags broke down at an even slower rate (k=0.0062 day–1). Phosphorus and nitrogen concentrations were initially higher in leaf packs with nutrients added, but this difference disappeared within 3 weeks. Fungal biomass developing in decomposing leaves was substantial (c. 55 mg dry mass per 1 g leaf dry mass) although similar between fertilized and unfertilized packs, as was the sporulation activity of aquatic hyphomycetes. There was a significantly greater number and higher biomass of macroinvertebrates (shredding nemourid stoneflies in particular) on the fertilized packs, suggesting that the increased leaf mass loss was brought about by shredder feeding. Received: 11 March 1999 / Accepted: 6 September 1999  相似文献   

19.
In order to investigate the factors causing fast growth of sprouts ofPasania edulis, photosynthetic activity and water relation characteristics of lower (mature) leaves and upper (expanding) leaves of the sprouts were compared with those of seedlings and adult trees ofP. edulis. Apparent quantum yield was generally low. Maximum photosynthetic rate was highest in the lower leaves of sprouts. Stomatal frequency was higher in sprout leaves than in seedling leaves. Osmotic potential at the water saturation point and water potential at the turgor loss point, in leaves, were higher in sprouts than in seedlings and adult trees. Symplasmic water content per unit leaf area was higher in sprouts than in seedlings. These water relation parameters in leaves indicated that sprout leaves are superior in maintaining cell turgor against water loss, but are not tolerant to water stress. In field measurements, sprout leaves showed higher stomatal conductance and transpiration rates. These results indicated that sprout leaves fully realized their high potential productivity even under field conditions. The leaf specific conductance, from the soil to the leaf, was higher in sprouts than in seedlings. Large and deep root systems of the original stumps of the sprouts may be attributed to the high leaf specific conductance.  相似文献   

20.
Abstract. Accumulation of nutrients in leaves of the dominating species of three ecosystems, characterizing the secondary succession from Genisto-Callunetum heathland through Leucobryo-Pinetum birch-pine woodland to mature Querco-Fagetum oak-beech forest, as well as nutrient turnover within these ecosystems was studied. The objective of the study was to establish potential variations in quantity and quality of nutrient supply to the plants with respect to succession dynamics. The results show very low leaf nutrient concentrations of all species investigated, coinciding with low nutrient availability in the soil. However, the nutrient content of leaves and leaf litter of Quercus petraea and Fagus sylvatica, which dominate the late succession stages, and in Betulapéndula are higher than in the photosynthetic organs (leaves and young shoots) of Calluna vulgaris and Pinus sylvestris. The combination of the higher nutrient content of the leaves and an increasing leaf-litter production during succession results in an increased nutrient turnover via leaf-litter fall. However, due to the high leaf biomass, the storage of nutrients in the leaf biomass is highest within the birch-pine woodland. From this, it may be assumed that the low demand and the low loss of nutrients via leaf-litter fall are favourable for Pinus at the early stages of forest succession on poor sandy soils. In contrast, Quercus and Fagus are provided with better growth conditions at the later stages of succession resulting from the accumulation of plant-available nutrients in the ecosystem by Pinus sylvestris, combined with a higher nutrient turnover as compared with the heathland.  相似文献   

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