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1.
Questions: (1) How do extreme climatic events and climate variability influence radial growth of conifers (silver fir, Norway spruce, Scots pine)? (2) How do elevation and soil water capacity (SWC) modulate sensitivity to climate? Location: The sampled conifer stands are in France, in western lowland and mountain forests, at elevations from 400 to 1700 m, and an SWC from 50 to 190 mm. Methods: We established stand chronologies for total ring width, earlywood and latewood width for the 33 studied stands (985 trees in total). Responses to climate were analysed using pointer years and bootstrapped response functions. Principal component analysis was applied to pointer years and response function coefficients in order to elucidate the ecological structure of the studied stands. Results: Extreme winter frosts are responsible for greater growth reductions in silver fir than in Norway spruce, especially at the upper elevation, while Scots pine was the least sensitive species. Exceptional spring droughts caused a notable growth decrease, especially when local conditions were dry (altitude<1000 m and SWC<100 mm for silver fir, western lowlands for Scots pine). Earlywood of silver fir depended on previous September and November and current‐year February temperature, after which current June and July water supply influenced latewood. Earlywood of Norway spruce was influenced by previous September temperature, after which current spring and summer droughts influenced both ring components. In Scots pine, earlywood and latewood depended on the current summer water balance. Local conditions mainly modulated latewood formation. Conclusions: If the climate becomes drier, low‐elevation dry stands or trees growing in western lowlands may face problems, as their growth is highly dependent on soil moisture availability.  相似文献   

2.

Questions

We aim for a better understanding of the different modes of intra‐ and inter‐specific competition in two‐ and three‐species mixed‐forests. How can the effect of different modes of competitive interactions be detected and integrated into individual tree growth models? Are species interactions in spruce–fir–beech forests more associated with size‐symmetric or size‐asymmetric competition? Do competitive interactions between two of these species change from two‐ to three‐species mixtures?

Location

Temperate mixed‐species forests in Central Europe (Switzerland).

Methods

We used data from the Swiss National Forest Inventory to fit basal area increment models at the individual tree level, including the effect of ecological site conditions and indices of size‐symmetric and size‐asymmetric competition. Interaction terms between species‐specific competition indices were used to disentangle significant differences in species interactions from two‐ to three‐species mixtures.

Results

The growth of spruce and fir was positively affected by increasing proportions of the other species in spruce–fir mixtures, but negative effects were detected with increasing presence of beech. We found that competitive interactions for spruce and fir were more related to size‐symmetric competition, indicating that species interactions might be more associated with competition for below‐ground resources. Under constant amounts of stand basal area, the growth of beech clearly benefited from the increasing admixture of spruce and fir. For this species, patterns of size‐symmetric and size‐asymmetric competitive interactions were similar, indicating that beech is a strong self‐competitor for both above‐ground and below‐ground resources. Only for silver fir and beech, we found significant changes in species interactions from two‐ to three‐species mixtures, but these were not as prominent as the effects due to differences between intra‐ and inter‐specific competition.

Conclusions

Species interactions in spruce–fir–beech, or other mixed forests, can be characterized depending on the mode of competition, allowing interpretations of whether they occur mainly above or below ground level. Our outcomes illustrate that species‐specific competition indices can be integrated in individual tree growth functions to express the different modes of competition between species, and highlight the importance of considering the symmetry of competition alongside competitive interactions in models aimed at depicting growth in mixed‐species forests.
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3.
Norway spruce is a widely cultivated species in Central Europe; however, it is highly susceptible to droughts, which are predicted to become more frequent in the future. A solution to adapt spruce forests to droughts could be the conversion to mixed-species stands containing species which are less sensitive to drought and do not increase the drought stress in spruce. Here we assessed the drought response of spruce and the presumably more drought-tolerant silver fir and Douglas fir in mixed-conifer stands. We measured tree ring widths of 270 target trees, which grew in mixed and mono-specific neighbourhoods in 18 managed stands in the Black Forest, to quantify the complementarity effects caused by species interactions on growth during the extreme drought event of 2003 and for a number of years with “normal” growth and climatic conditions. Mixed-species neighbourhoods did not significantly affect tree ring growth in normal years. However, during the drought, silver fir benefitted from mixing, while Douglas fir was more drought-stressed in the mixture. The drought response of spruce was dependent on the density and species composition of the neighbourhood, showing both positive and negative mixing effects. Mixed stands containing these tree species could improve adaptation to drought because the risks of extreme events are spread across species, and the performance of individual species is improved. Our knowledge about specific species interactions needs to be improved to manage tree mixtures more effectively with regard to the participating species and stand density.  相似文献   

4.
5.
  • Climate anomalies have resulted in changing forest productivity, increasing tree mortality in Central and Southern Europe. This has resulted in more severe and frequent ecological disturbances to forest stands. This study analysed the size‐dependence of growth response to drought years based on 384 tree individuals of Norway spruce [Picea abies (L.) Karst.] and European beech [Fagus sylvatica ([L.)] in Bavaria, Germany.
  • Samples were collected in both monospecific and mixed‐species stands. To quantify the growth response to drought stress, indices for basal area increment, resistance, recovery and resilience were calculated from tree ring measurements of increment cores. Linear mixed models were developed to estimate the influence of drought periods.
  • The results show that ageing‐related growth decline is significant in drought years. Drought resilience and resistance decrease significantly with growth size among Norway spruce individuals. Evidence is also provided for robustness in the resilience capacity of European beech during drought stress. Spruce benefits from species mixing with deciduous beech, with over‐yielding spruce in pure stands.
  • The importance of the influence of size‐dependence within tree growth studies during disturbances is highlighted and should be considered in future studies of disturbances, including drought.
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6.
Question: How does competition between quaking aspen (Populus tremuloides) and white fir (Abies concolor) affect growth and spatial pattern of each species? Location: The northern Sierra Nevada, California, USA. Methods: In paired plots in mixed aspen‐ (n=3) or white fir‐dominated (n=2) stands, we mapped trees and saplings and recorded DBH, height, species, and condition and took increment cores. We tallied seedlings by species. Tree ring widths were used as a measure of basal area change over the last decade, and canopy openness was identified using hemispherical photographs. Linear mixed models were used to relate neighborhood indices of competition, stand, and tree‐level variables to diameter increment. Spatial patterns of stems were identified using the Neighborhood Density Function. Results: White fir radial growth was higher in aspen‐ than white fir‐dominated plots. Individual‐level variables were more important for white fir than for aspen growth, while variables representing competitive neighborhood were important only for aspen. The forest canopy was more open in aspen‐ than white fir‐dominated stands, but ample aspen seedlings were observed in all stands. Canopy stems of aspen and white fir were randomly distributed, but saplings and small trees were clumped. Aspen saplings were repelled by canopy aspen stems. Conclusions: Variation in canopy openness explained more stand–stand variation in white fir than aspen growth, but high light levels were correlated with recruitment of aspen seedlings to the sapling class. Radial growth of aspen was predicted by indices of neighborhood competition but not radial growth of white fir, indicating that spacing and stem arrangement was more important for aspen than white fir growth. Fire suppression has removed a major disturbance mechanism that promoted aspen persistence and reduced competition from encroaching conifers, and current forests favor species that regenerate best by advance regeneration (white fir).  相似文献   

7.
The future performance of native tree species under climate change conditions is frequently discussed, since increasingly severe and more frequent drought events are expected to become a major risk for forest ecosystems. To improve our understanding of the drought tolerance of the three common European temperate forest tree species Norway spruce, silver fir and common beech, we tested the influence of climate and tree‐specific traits on the inter and intrasite variability in drought responses of these species. Basal area increment data from a large tree‐ring network in Southern Germany and Alpine Austria along a climatic cline from warm‐dry to cool‐wet conditions were used to calculate indices of tolerance to drought events and their variability at the level of individual trees and populations. General patterns of tolerance indicated a high vulnerability of Norway spruce in comparison to fir and beech and a strong influence of bioclimatic conditions on drought response for all species. On the level of individual trees, low‐growth rates prior to drought events, high competitive status and low age favored resilience in growth response to drought. Consequently, drought events led to heterogeneous and variable response patterns in forests stands. These findings may support the idea of deliberately using spontaneous selection and adaption effects as a passive strategy of forest management under climate change conditions, especially a strong directional selection for more tolerant individuals when frequency and intensity of summer droughts will increase in the course of global climate change.  相似文献   

8.
Research related to the allometric relationships of tree height and projected tree crown area to diameter at breast height was conducted to look at the biological suitability and timber production potential of Douglas fir under the conditions present in central Europe. The dependence of allometric relationships on soil nutrient conditions were described in forest stands of Douglas fir and Norway spruce. The studied sites were climatically similar but differed in soil nutrient availability. A significant difference was found in the allometric relationships of Norway spruce trees from the nutrient poor and nutrient rich site. In contrast to the Norway spruce, there was no significant effect of site fertility on allometric relationships for Douglas fir suggesting that its allocation patterns were less sensitive to site nutrient conditions. Stem growth increment, which was measured weekly during two consecutive seasons for both species, was related to the weather conditions and available soil moisture. Stem growth of Douglas fir began about 2 weeks earlier than in the Norway spruce at both sites. At the nutrient rich site, most of the stem growth of both species occurred at the beginning of the season, while growth at the other site was more evenly distributed throughout the season. Data obtained in this study will be useful for modeling stem growth and analysis of water use efficiency of these two tree species.  相似文献   

9.
In pure and mixed stands of Norway spruce ( Picea abies [L.] Karst.) and European beech ( Fagus sylvatica L.) we have analyzed crown allometry and growing space efficiency at the tree level and have scaled this from tree level to stand level production. Allometry is quantified by the ratio A between the relative growth rates of laterally and vertically oriented tree dimensions. Efficiency parameters, EOC for efficiency in space occupation, EEX for efficiency in space exploitation, and EBI for efficiency in biomass investment, were evaluated, based on quantity and quality of growing space and were measured using crown size and competition index. The evaluation reveals why pure stands of spruce are preferred by foresters, even though the natural vegetation would be dominated by beech. Spruce occupies its share of resources intensively by means of tightly packed pillar-like crowns, whereas beech seizes resources extensively by means of a multi-layered, veil-like canopy. With a given relative biomass increment, beech achieves a 57 % higher increment in crown projection area and a 127 % higher increment in height due to its particular capacity of lateral and vertical expansion. Beech trees are approximately 60 % more efficient in space occupation than spruce trees, however, on average, they are about 70 % less efficient in space exploitation. As a vertical fast growing tree, spruce is efficient in space exploitation under constant conditions, but far more susceptible to disturbances and less well equipped to overcome them when compared with beech. Beech is weaker in terms of space exploitation, while being superior in space occupation, where it encircles competitors and fills gaps after disturbances, which is a successful long-term strategy. A mixture of the two species reduces stand level production by 24 % in comparison to a pure spruce stand, however, when considering enhanced stabilization of the whole stand and risk distribution in the long term, the mixed stand may exceed the production level of pure spruce stands. EEX reflects a strong ontogenetic drift and competition effect that should be considered when scaling from tree to stand level production.  相似文献   

10.
Morphological plasticity was studied for advanced regeneration trees in different light environments of the mountainous, mixed-species forests in the Carpathian Mountains of Romania. The primary species in these mixtures were very shade tolerant silver fir (Abies alba Mill.) and European beech (Fagus sylvatica L.), and midtolerant Norway spruce (Picea abies (L.) Karst). Seedlings/saplings of these species were selected for measurements in different stands from two different geographical locations. Various morphological traits (specific leaf area, live crown ratio, crown width to length ratio, terminal to lateral ratio, number of internodal shoots, number of shoots in terminal whorl, stem symmetry, stem orientation, stem forking) for each regenerating tree were measured during summers of 2001 and 2002. Percentage of above canopy light and stand basal area measures were used to assess the available growing space for each seedling/sapling. Regression relationships were developed for the different morphological indicators as a function of these two variables. All species adapted their morphology along the gradient in light and basal area. Spruce seemed to be less adapted to low light conditions than both fir and beech. However, no significant differences in terms of shade tolerance were detected using the above indicators. In really dense stand conditions (less than 20% above canopy light and stand basal area above 36 m2 ha−1), probability for stem forking in beech increased. In open, all three species adapted their morphology for vigorous growth. Under such conditions, spruce was better adapted than fir.  相似文献   

11.
While previous studies focused on tree growth in pure stands, we reveal that tree resistance and resilience to drought stress can be modified distinctly through species mixing. Our study is based on tree ring measurement on cores from increment boring of 559 trees of Norway spruce (Picea abies [L.] Karst.), European beech (Fagus sylvatica [L.]) and sessile oak (Quercus petraea (Matt.) Liebl.) in South Germany, with half sampled in pure, respectively, mixed stands. Indices for resistance, recovery and resilience were applied for quantifying the tree growth reaction on the episodic drought stress in 1976 and 2003. The following general reaction patterns were found. (i) In pure stands, spruce has the lowest resistance, but the quickest recovery; oak and beech were more resistant, but recover was much slower and they are less resilient. (ii) In mixture, spruce and oak perform as in pure stands, but beech was significantly more resistant and resilient than in monoculture. (iii) Especially when mixed with oak, beech is facilitated. We hypothesise that the revealed water stress release of beech emerges in mixture because of the asynchronous stress reaction pattern of beech and oak and a facilitation of beech by hydraulic lift of water by oak. This facilitation of beech in mixture with oak means a contribution to the frequently reported overyield of beech in mixed versus pure stands. We discuss the far‐reaching implications that these differences in stress response under intra‐ and inter‐specific environments have for forest ecosystem dynamics and management under climate change.  相似文献   

12.
How tree morphology develops in mixed-species stands is essential for understanding and modelling mixed-stand dynamics. However, research so far focused on the morphological variation between tree species and neglected the variation within a species depending on intra- and interspecific competition. Our study, in contrast, addresses crown properties of nine mature Norway spruces (Picea abies [L.] Karst.) of a pure stand and compares them with ten spruces growing in mixture with European beech (Fagus sylvatica [L.]). The same was done with 11 pure stand beeches and 12 beeches growing in mixture with spruce. Through application of a terrestrial laser scanner and a new skeletonization approach, we deal with both species’-specific morphological traits such as branch angle, branch length, branch bending, crown volume and space occupation of branches within the crown, some of which were hardly accessible so far. Special attention is paid to distinct differences between trees growing in mixed and pure stands: for spruce, our study reveals significantly longer branches and greater crown volumes in the mixed stand when compared to the pure stand. In case of European beech, individuals growing in mixture show flatter branch angles, more distinct ramification, greater crown volumes and a lower share of a single branch’s space occupation in the total crown volume. The results show that the presented methods yield detailed information on the morphological traits analyzed in this study and that interspecific competition on its own may have a significant impact on crown structures. Implications for production ecology and stand dynamics of mixed-species forests are discussed.  相似文献   

13.
Facilitation, reduced competition or increased competition can arise in mixed stands and become essential to the performance of these stands when compared to pure stands. Facilitation and over‐yielding are widely held to prevail on poor sites, whereas neutral interactions or competition, leading to under‐yielding of mixed versus pure stands, can occur on fertile sites. While previous studies have focused on the spatial variation of mixing effects, we examine the temporal variation of facilitation and competition and its effect on growth. The study is based on tree ring measurement on cores from increment borings from 559 trees of Norway spruce (Picea abies [L.] Karst.), European beech (Fagus sylvatica [L.]) and sessile oak (Quercus petraea (Matt.) Liebl.) in southern Germany, half of which were in pure stands and half in adjacent mixed stands. Mean basal area growth indices were calculated from tree ring measurements for pure and mixed stands for every species and site. The temporal variation, with positive correlations between species‐specific growth indices during periods of low growth and neutral or negative correlations during periods of high growth, is more distinct in mixed than in neighbouring pure stands. We provide evidence that years with low growth trigger over‐yielding of trees in mixed as opposed to pure stands, while years with high growth lead to under‐yielding. We discuss the relevance of the results in terms of advancing our understanding and modelling of mixed stands, extension of the stress gradient hypothesis, and the performance of mixed versus pure stands in the face of climate change.  相似文献   

14.
Question: Two questions about within‐stand spatial variability are addressed in this paper. How does species richness of tree regeneration respond to small‐scale ecological gradients, and what effect does natural Abies balsamea abundance have on the species richness of other tree regeneration? Location: A long‐term, gap‐silviculture experiment, Acadian mixed‐wood forest, Maine, USA. Methods: Eight stands treated with and without gap harvesting were sampled to capture sub‐stand heterogeneity of understorey tree regeneration concurrently with patterning of local stand conditions. Spatial and non‐spatial models were developed to test the relationships between two response variables [species richness of small (height ≥0.1 m, but <0.75 m) and large (height ≥0.75 m, but <1.4 m) regeneration] and five explanatory variables (depth to water table, percentage canopy transmittance, A. balsamea regeneration density, and overstorey basal area and species richness). Results: Despite high unexplained variance for all models, consistent associations among variables were found. Negative associations were found between: (1) the species richness of small regeneration and A. balsamea regeneration density and (2) the species richness of large regeneration and overstorey basal area. Positive associations were found between: (1) the species richness of small regeneration and both overstorey basal area and species richness and (2) the species richness of small and large regeneration and canopy transmittance. Conclusions: Promoting tree species diversity in Acadian mixed‐wood stands may not be achievable through the use of gap‐harvesting alone if the density of understorey Abies balsamea is not reduced either naturally or through silvicultural intervention.  相似文献   

15.
Red spruce (Picea rubens)–dominated forests occupied as much as 600,000 ha in West Virginia prior to exploitive logging era of the late nineteenth and early twentieth centuries. Subsequently, much of this forest type was converted to northern hardwoods. As an important habitat type for a number of rare or sensitive species, only about 12,000 ha of red spruce forests presently remain in the state. In order to assess the prospects for restoration, we examined six northern hardwood stands containing understory red spruce to (1) characterize stand dynamics and regeneration patterns and (2) simulate the effectiveness of restoration silviculture to enhance red spruce overstory recruitment. Stands originated in the late 1800s to early 1900s and are currently in the (late) stem exclusion or understory reinitiation stages. Five of the six stands had even‐aged overstories that originated after clear‐cutting. Tree‐ring chronologies show high initial growth rates consistent with stand initiation. One stand, partially harvested in 1915, was uneven aged with older, legacy residuals in the canopy. Most stands had two cohorts of understory red spruce, with more than 40% of these individuals showing prior release. Our 100‐year growth simulation suggested that a 50% basal area thinning from above could double red spruce basal area to support a mixed spruce–hardwood stand in approximately 20–40 years. These results indicate that restoration silviculture could be an effective tool for increasing the amount and quality of this reduced forest type in the central Appalachians.  相似文献   

16.
Shifts in tree species distributions caused by climatic change are expected to cause severe losses in the economic value of European forestland. However, this projection disregards potential adaptation options such as tree species conversion, shorter production periods, or establishment of mixed species forests. The effect of tree species mixture has, as yet, not been quantitatively investigated for its potential to mitigate future increases in production risks. For the first time, we use survival time analysis to assess the effects of climate, species mixture and soil condition on survival probabilities for Norway spruce and European beech. Accelerated Failure Time (AFT) models based on an extensive dataset of almost 65 000 trees from the European Forest Damage Survey (FDS) – part of the European‐wide Level I monitoring network – predicted a 24% decrease in survival probability for Norway spruce in pure stands at age 120 when unfavorable changes in climate conditions were assumed. Increasing species admixture greatly reduced the negative effects of unfavorable climate conditions, resulting in a decline in survival probabilities of only 7%. We conclude that future studies of forest management under climate change as well as forest policy measures need to take this, as yet unconsidered, strongly advantageous effect of tree species mixture into account.  相似文献   

17.
Improving our understanding of the potential of forest adaptation is an urgent task in the light of predicted climate change. Long‐term alternatives for susceptible yet economically important tree species such as Norway spruce (Picea abies) are required, if the frequency and intensity of summer droughts will continue to increase. Although Silver fir (Abies alba) and Douglas fir (Pseudotsuga menziesii) have both been described as drought‐tolerant species, our understanding of their growth responses to drought extremes is still limited. Here, we use a dendroecological approach to assess the resistance, resilience, and recovery of these important central Europe to conifer species the exceptional droughts in 1976 and 2003. A total of 270 trees per species were sampled in 18 managed mixed‐species stands along an altitudinal gradient (400–1200 m a.s.l.) at the western slopes of the southern and central Black Forest in southwest Germany. While radial growth in all species responded similarly to the 1976 drought, Norway spruce was least resistant and resilient to the 2003 summer drought. Silver fir showed the overall highest resistance to drought, similarly to Douglas fir, which exhibited the widest growth rings. Silver fir trees from lower elevations were more drought prone than trees at higher elevations. Douglas fir and Norway spruce, however, revealed lower drought resilience at higher altitudes. Although the 1976 and 2003 drought extremes were quite different, Douglas fir maintained consistently the highest radial growth. Although our study did not examine population‐level responses, it clearly indicates that Silver fir and Douglas fir are generally more resistant and resilient to previous drought extremes and are therefore suitable alternatives to Norway spruce; Silver fir more so at higher altitudes. Cultivating these species instead of Norway spruce will contribute to maintaining a high level of productivity across many Central European mountain forests under future climate change.  相似文献   

18.
Aim There is increasing research attention being given to the role of interactions among natural disturbances in ecosystem processes. We studied the interactions between fire and spruce beetle (Dendroctonus rufipennis Kirkby) disturbances in a Colorado subalpine forest. The central questions of this research were: (1) How does fire history influence stand susceptibility to beetle outbreak? And conversely, (2) How does prior occurrence of a beetle outbreak influence stand susceptibility to subsequent fire? Methods We reconstructed the spatial disturbance history in a c. 4600 ha area by first identifying distinct patches in the landscape on aerial photographs. Then, in the field we determined the disturbance history of each patch by dating stand origin, fire scars, dates of mortality of dead trees, and releases on remnant trees. A geographical information system (GIS) was used to overlay disturbance by fire and spruce beetle. Results and main conclusions The majority of stands in the study area arose following large, infrequent, severe fires occurring in c. 1700, 1796 and 1880. The study area was also affected by a severe spruce beetle outbreak in the 1940s and a subsequent low‐severity fire. Stands that originated following stand‐replacing fire in the late nineteenth century were less affected by the beetle outbreak than older stands. Following the beetle outbreak, stands less affected by the outbreak were more affected by low‐severity fire than stands more severely affected by the outbreak. The reduced susceptibility to low‐severity fire possibly resulted from increased moisture on the forest floor following beetle outbreak. The landscape mosaic of this subalpine forest was strongly influenced by the interactions between fire and insect disturbances.  相似文献   

19.
The age dynamics of the main taxation characteristics (height, diameter, stand basal area, average and current increment of stock, and total stand productivity) have been analyzed in stands of different types. The influence of admixed species on the formation of the stands is assessed. The expediency of planting pure stands on heavy textured soils of floodplains in the forest steppe has been proven.  相似文献   

20.
Aim Bark beetle outbreaks have recently affected extensive areas of western North American forests, and factors explaining landscape patterns of tree mortality are poorly understood. The objective of this study was to determine the relative importance of stand structure, topography, soil characteristics, landscape context (the characteristics of the landscape surrounding the focal stand) and beetle pressure (the abundance of local beetle population eruptions around the focal stand a few years before the outbreak) to explain landscape patterns of tree mortality during outbreaks of three species: the mountain pine beetle, which attacks lodgepole pine and whitebark pine; the spruce beetle, which feeds on Engelmann spruce; and the Douglas‐fir beetle, which attacks Douglas‐fir. A second objective was to identify common variables that explain tree mortality among beetle–tree host pairings during outbreaks. Location Greater Yellowstone ecosystem, Wyoming, USA. Methods We used field surveys to quantify stand structure, soil characteristics and topography at the plot level in susceptible stands of each forest type showing different severities of infestation (0–98% mortality; n= 129 plots). We then used forest cover and beetle infestation maps derived from remote sensing to develop landscape context and beetle pressure metrics at different spatial scales. Plot‐level and landscape‐level variables were used to explain outbreak severity. Results Engelmann spruce and Douglas‐fir mortality were best predicted using landscape‐level variables alone. Lodgepole pine mortality was best predicted by both landscape‐level and plot‐level variables. Whitebark pine mortality was best – although poorly – predicted by plot‐level variables. Models including landscape context and beetle pressure were much better at predicting outbreak severity than models that only included plot‐level measures, except for whitebark pine. Main conclusions Landscape‐level variables, particularly beetle pressure, were the most consistent predictors of subsequent outbreak severity within susceptible stands of all four host species. These results may help forest managers identify vulnerable locations during ongoing outbreaks.  相似文献   

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