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1.
Laurent J. Lamarque Déborah Corso José M. Torres-Ruiz Eric Badel Timothy J. Brodribb Régis Burlett Guillaume Charrier Brendan Choat Hervé Cochard Gregory A. Gambetta Steven Jansen Andrew King Nicolas Lenoir Nicolas Martin-StPaul Kathy Steppe Jan Van den Bulcke Ya Zhang Sylvain Delzon 《Annals of Forest Science》2018,75(3):88
Key message
Direct, non-invasive X-ray microtomography and optical technique observations applied in stems and leaves of intact seedlings revealed that laurel is highly resistant to drought-induced xylem embolism. Contrary to what has been brought forward, daily cycles of embolism formation and refilling are unlikely to occur in this species and to explain how it copes with drought.Context
There has been considerable controversy regarding xylem embolism resistance for long-vesselled angiosperm species and particularly for the model species for refilling (Laurus nobilis L.).Aims
The purpose of this study was to resolve the hydraulic properties of this species by documenting vulnerability curves of different organs in intact plants.Methods
Here, we applied a direct, non-invasive method to visualize xylem embolism in stems and leaves of intact laurel seedlings up to 2-m tall using X-ray microtomography (microCT) observations and the optical vulnerability technique. These approaches were coupled with complementary centrifugation measurements performed on 1-m long branches sampled from adult trees and compared with additional microCT analyses carried out on 80-cm cut branches.Results
Direct observations of embolism spread during desiccation of intact laurels revealed that 50% loss of xylem conductivity (Ψ50) was reached at ??7.9?±?0.5 and ??8.4?±?0.3 MPa in stems and leaves, respectively, while the minimum xylem water potentials measured in the field were ??4.2 MPa during a moderate drought season. Those findings reveal that embolism formation is not routine in Laurus nobilis contrary to what has been previously reported. These Ψ50 values were close to those based on the flow-centrifuge technique (??9.2?±?0.2 MPa), but at odds with microCT observations of cut branches (??4.0?±?0.5 MPa).Conclusion
In summary, independent methods converge toward the same conclusion that laurel is highly resistant to xylem embolism regardless its development stage. Under typical growth conditions without extreme drought events, this species maintains positive hydraulic safety margin, while daily cycles of embolism formation and refilling are unlikely to occur in this species.2.
Marion Zapater Nathalie Br??da Damien Bonal Sylvia Pardonnet Andr?? Granier 《Annals of Forest Science》2013,70(1):31-39
? Introduction
In the context of global environmental changes, better understanding of tree response to soil drought in young mixed species stands is needed to anticipate forest adaptation and management practices for the future.? Materials and methods
We investigated the functional response of five co-occurring broad-leaved tree species growing in a 15- to 25-year-old mixed stand in northeastern France during the 2006 summer drought. We measured functional traits related to water acquisition (phenology, rooting pattern and vulnerability of xylem to cavitation) and the ecophysiological response (sap flux density, leaf water potential) of these species to soil water shortage.? Results
Our study highlights contrasted drought response strategies among these species and a trade-off between leaf phenology, resistance of xylem to cavitation and root system depth.? Conclusion
At this site, a deep root system seemed to be a key functional trait for the species to cope with drought. Quercus robur and Salix capreae can be characterised as drought-avoidance species as they possess a deep root system and therefore did not strongly experience soil drought. Despite deep rooting capacity, Betula pendula did not really avoid soil drought and strongly regulated transpiration during dry periods. Nevertheless, the earliness of budburst of this species contributes to high annual growth rate. In contrast, Carpinus betulus and Fagus sylvatica both displayed typical characteristics of drought-sensitive species. 相似文献3.
? Context
The knowledge of how shrub–seedling interactions vary with summer drought, canopy opening, and tree species is crucial for adapting forest management to climate change.? Aims
The aim of this study was to assess variation in shrub–oak recruitment associations along a south–north drought climate gradient and between two levels of canopy cover in coastal dune forest communities in a climate change-adapted forest management perspective.? Material and methods
Mapped data of associational patterns of seedlings of three oak species with interspecific pooled shrubs were analyzed using a bivariate pair correlation function in 10 (0.315 ha) regeneration plots located in forest and recent gap sites along the climate gradient. An index of association strength was calculated in each plot and plotted against a summer moisture index.? Results
The association strength increased with increasing summer drought from wet south to dry north and from closed forests to gaps.? Conclusion
Consistent with facilitation theory, our results suggest that climate change may shift associational patterns in coastal dune forest communities towards more positive associations, in particular in canopy gaps. In a perspective of climate change, foresters may need to conserve understory shrubs in gaps in order to promote oak species regeneration. 相似文献4.
Heike Kawaletz Inga Mölder Peter Annighöfer André Terwei Stefan Zerbe Christian Ammer 《Annals of Forest Science》2014,71(3):337-347
Context
Identifying the traits of exotic species may explain their invasiveness and help control the negative impacts of these species on native ecosystems. In this study, we investigated root competition as an important driver for the competitiveness of two exotic tree species in the seedling stage.Aims
In a pot experiment, carried out in Central Europe, we studied the effect of root competition of two exotic tree species (Prunus serotina Ehrh. and Robinia pseudoacacia L.) on biomass allocation of two native ones (Quercus robur L. and Carpinus betulus L.).Methods
Seedlings of these species were exposed to intra- and interspecific mixtures with and without the effect of root competition. For this, belowground plastic partitions were installed in the pots.Results
Competition intensity in the pots increased over time, irrespective of mixture type. However, this increase was much higher for the native species if mixed with the exotic species compared to monocultures or inter-native mixtures. In addition to ontogeny, competition affected biomass allocation patterns. Under root competition by the exotic species, the native species preferentially allocated their biomass to the roots. A higher allocation to the roots was mainly achieved at the expense of leaf and branch biomass.Conclusion
Root competition of P. serotina and R. pseudoacacia may be a reason for the lack of Q. robur and C. betulus in the seedling stage in natural environments where all four species occur. 相似文献5.
Javier Puntieri Javier Grosfeld Luis Tejera Víctor Mondino Leonardo Gallo 《Annals of Forest Science》2013,70(5):471-479
? Context
There is evidence that Nothofagus nervosa (= N. alpina) is suitable for timber production in temperate regions due to its wood quality and fast growth. However, high intra-specific variability in the architectural traits of this species limits its usefulness.? Aims
This study was aimed at evaluating intra-specific variability in the architecture of N. nervosa trees at the population level, with emphasis on traits related to the suitability of trees for timber production.? Methods
The size of 13-year-old trees installed in an experimental field population and the extent of differentiation between trunk and branches were compared among families (trees derived from different mother trees).? Results
The large majority of trees exhibited architectural features indicative of high suitability for timber production: clear differentiation between trunk and main branches and few occurrences of permanent trunk forks, apex deaths, and sylleptic branching. Trees belonging to different families differed in height and trunk diameter. Micro-environmental conditions had a very significant effect on most architectural traits, including the trunk differentiation.? Conclusions
N. nervosa emerges as a suitable option for timber production in temperate regions. Trunk growth seems to be under both environmental and genetic control in this species. 相似文献6.
Key message
Wood-anatomical traits determining the hydraulic architecture of Larix sibirica in the drought-limited Mongolian forest steppe at the southern fringe of the boreal forest respond to summer drought, but only weakly to variations in microclimate that depend on forest stand size.Context
Siberian larch (L. sibirica Ledeb.) is limited by summer drought and shows increasing mortality rates in the Mongolian forest steppe. The climate sensitivity of stemwood formation increases with decreasing forest stand size. The trees’ hydraulic architecture is crucial for drought resistance and thus the capability to deal with climate warming.Aims
We studied whether hydraulic traits were influenced by temporal or forest size-dependent variations in water availability and were related to tree-ring width.Methods
Hydraulic traits (tracheid diameter, tracheid density, potential sapwood area-specific hydraulic conductivity) of earlywood were studied in stemwood series of 30 years (1985–2014) and were related to climate data. Tree-ring width was measured for the same period. Trees were selected in stands of four different size classes with increasing drought exposure with decreasing stand size.Results
Tracheid diameters and hydraulic conductivity decreased with decreasing late summer precipitation of the previous year and were positively correlated with tree-ring width. Forest stand size had only weak effects on hydraulic traits, despite known effects on stemwood increment.Conclusion
Decreasing tracheid diameters and thus hydraulic conductivity are a drought acclimation of L. sibirica in the Mongolian forest steppe. These acclimations occur as a response to drought periods but are little site-dependent with respect to stand size.7.
? Context
A clear understanding of the genetic control of wood properties is a prerequisite for breeding for higher wood quality in Populus tomentosa Carr. hybrid clones.? Aims
The experiments aimed at unraveling genetic and environmental effects on wood properties among triploid hybrid clones of P. tomentosa.? Methods
We used 5-year-old clonal trials established in Northern China to assess the heritability of wood density and fiber traits. Two hundred seventy trees from nine clones were sampled in five sites.? Results
Site had a very significant effect on all recorded traits. Despite this large site effect, a tight genetic control was detected and clonal repeatability varied between 0.53 and 0.95. Significant genotype?×?environment interactions were detected for most of the traits. Moderate to tight correlation were evidenced among traits but they were not consistent with that in several cases that were site-dependent.? Conclusions
Our results revealed a tight genetic control over several wood properties and therefore breeding programs might be able to improve wood density, fiber length, and coarseness in these hybrids. 相似文献8.
Variation of wood color parameters of Tectona grandis and its relationship with physical environmental factors 总被引：1，自引：0，他引：1
? Context
Teak??s wood color is considered an important attribute in the marketing phase and it has been influenced by environmental setting, stand conditions and management, plant genetic source, and age. However, there is a lack of understanding about how the environmental factors might affect the teak??s wood color planted in short-rotation forest plantations.? Aims
The aim of this study is to understand the relationship, gathered from generated information, between edaphic and climatic variables and their effects in the wood color variation of Tectona grandis from trees in forest plantations.? Methods
Twenty-two plots were grouped in five cluster sites that shared similar climatic and soil conditions. Data about soil??s physical?Cchemical properties and climatic variables were collected and analyzed. Representative trees were harvested next to each plot in order to obtain a wood sample per tree at a diameter breast height. Wood color was measured using standardized CIELab??s chromaticity system.? Results
After comparing the wood change color index (?E*) in the five studied clusters, it was found that heartwood produced from drier and fertile sites had more yellowish-brown color. The heartwood b* color index resulted with significant correlations (R?>?0.5, P?<?0.05) among nine climatic and eight edaphic variables.? Conclusion
It was concluded that climatic variables should be considered as the first-order causal variables to explain wood color variation. Hence, darker b* wood color was associated with dry climates; also, with deeper and fertile sites. 相似文献9.
?Context
Tree species differ largely in their capability to produce characteristic shade leaves with effective morphological and physiological acclimation to low light.?Aims
By examining the sun/shade leaf differentiation in leaf morphology, foliar nitrogen and photosynthetic capacity in five temperate tree species of different successional status, we aimed at identifying those leaf traits that determine the development of a typical shade crown with low light-acclimated leaves.?Methods
Leaf morphology, foliar N content, photosynthetic capacity (V cmax, J max and A max) and leaf dark respiration (R d) were measured in the canopies of 26 adult trees of Fraxinus, Acer, Carpinus, Tilia and Fagus species.?Results
Six traits (the sun/shade leaf differentiation in specific leaf area, leaf size, A max per leaf area or per mass, photosynthetic N use efficiency and R d) were found to characterise best the degree of low light acclimation in shade leaves. All five species exhibited certain modifications in leaf morphology and/or physiology in response to low light; Fagus sylvatica showed the highest and Fraxinus excelsior the lowest shade leaf acclimation.?Conclusions
Our results indicate that the five early/mid- to late-successional species have developed species-specific low light acclimation strategies in their shade crowns which differ in terms of the relative importance of leaf morphological and physiological acclimation. 相似文献10.
Ervin Rasztovits Imre Berki Csaba Mátyás Kornél Czimber Elisabeth Pötzelsberger Norbert Móricz 《Annals of Forest Science》2014,71(2):201-210
Context
Projections of species distribution models under future climate are usually based on long-term averages. However, singular extreme drought events presumably contribute to the shaping of distribution limits at the retreating low-elevation xeric limits.Methods
The objectives of this study were to set up a distribution model based on extreme drought events (EDM), which uses sanitary logging information as a proxy of vitality response of beech, and compare it with the results of classical species distribution models (SDMs).Results
Predictions of the EDM for 2025 were in agreement with those of the SDM, but EDM predicted a more serious decline in all regions of Hungary towards the end of the century.Conclusion
These results suggest that the predicted increase in frequency and severity of drought events may further limit the distribution of beech in the future. 相似文献11.
Samah Mhamdi Oliver Brendel Pierre Montpied Hanah Ghouil-Amimi Ibrahim Hasnaoui Erwin Dreyer 《Annals of Forest Science》2013,70(7):675-684
?Context
Oak species display a large within-species diversity combined with significant gene flow between species. Quercus afares Pomel is an endemic species that grows as a small relict population in Northern Tunisia, together with Quercus suber L. and Quercus canariensis Willd. It is considered to be a fixed hybrid between the two latter.?Aims
We tested (1) whether this population of Q. afares is declining with respect to the neighbouring populations of the two other species and (2) whether the spatial distribution of leaf morphology is random among individuals within this small population of Q. afares or influenced by the neighbouring species.?Methods
Spatial distribution and diameter at breast height were recorded in the population of Q. afares. We identified two perpendicular transects across the population reaching from areas dominated by Q. canariensis to areas dominated by Q. suber and defined 20 plots along each. Eleven traits were assessed on 10 leaves collected from the upper crown of trees from the three species. A multiple correspondence analysis and an ANOVA were used to test for within and between-species diversity in leaf morphology.?Results
Q. afares individuals occurred at a low frequency among the two other species. Nevertheless, Q. afares showed no particular evidence of decline with respect to the two other species. Leaf traits displayed a large inter-specific variability with very little overlap between Q. afares and Q. suber and some between Q. afares and Q. canariensis. A high level of diversity was detected for these traits within Q. afares, but we were unable to unravel any spatial organisation of this diversity; in Q. afares, leaf traits varied independently of the presence of neighbouring Q. suber or Q. canariensis.?Conclusion
Q. afares seems to be a fixed species with specific leaf traits that are independent of that of the species of the neighbouring trees. This population is not suffering a detectable trend towards decline except for the lack of any visible regeneration below any of the three species. 相似文献12.
? Context
Soil temperature can limit tree growth and function, but it is often unaddressed in understanding the successional status of trees.? Aims
We tested how soil temperature affected carbon allocation strategies of two dominant co-occurring boreal conifer species, Pinus contorta and Picea mariana.? Methods
We measured nonstructural carbon (NSC) concentrations, biomass, and photosynthesis of dormant and actively growing 2-year-old seedlings in response to three soil temperatures (5, 10, and 20 °C) under a common ambient air temperature.? Results
For both species, variation in carbon reserves with soil temperature was more pronounced following seedling growth than during dormancy. For both species and all organ types (roots, needles, and stems), NSC concentrations were highest when seedlings were grown at 5 than 20 °C. Mass adjusted for NSC content was negatively correlated with NSC concentration for all organ types of both species. Soil temperature had a marginally significant effect on photosynthesis of pine; seedlings grown at 10 or 20 °C acquired more carbon than seedlings grown at 5 °C. Spruce seedlings photosynthesized more when grown at 20 °C than at 5 or 10 °C.? Conclusion
Interspecific differences in allocation of carbon may underlie the responses of P. mariana and P. contorta to cold soils and consequently their successional status. 相似文献13.
Context
Warmer temperatures and anthropogenic N depositions are altering soil nutrient cycles and plant nutrition worldwide and are projected to rise dramatically in the future, particularly at the high latitudes. How much will such alterations affect symbiotic organisms such as ectomycorrhizae (ECM)?Aims
The aim was to investigate the short-term effects of warmer soil temperatures and N-enriched precipitations on roots and ECM.Methods
A field experiment was established during 2008–2010 in two black spruce (Picea mariana) stands of the boreal forest of Quebec, Canada. The ECM of 12 trees per site were counted and identified by morphotyping.Results
After 3 years, soil heating markedly modified the proportions of vital root tips and ECM in the colder stand, while no effect of the N-enriched rain was observed on soil chemistry and consequently on roots and ECM. Density and ramification of root tips were not affected by the treatments.Conclusion
The hypothesis that the treatments would alter the ECM was confirmed only partially in the colder stand. The type of application and amount of N supplied were unable to substantially modify in the short-term growth conditions of roots and ECM, explaining the observed results. 相似文献14.
Laura Hernández Isabel Cañellas Iciar Alberdi Iván Torres Fernando Montes 《Annals of Forest Science》2014,71(2):161-171
? Context
It is assumed that global change is already affecting the composition, structure and distribution of forest ecosystems; however, detailed evidences of altitudinal and latitudinal shifts are still scarce.? Aims
To develop a method based on National Forest Inventory (NFI) to assess spatio-temporal changes in species distributions.? Methods
We develop an approach based on universal kriging to compare species distribution models from the different NFI cycles and regardless of the differences in the sampling schemes used. Furthermore, a confidence interval approach is used to assess significant changes in species distribution. The approach is applied to some of the southernmost populations of Pinus sylvestris and Fagus sylvatica in the Western Pyrenees over the last 40 years.? Results
An increase of the presence of the two species in the region was observed. Scots pine distribution has shifted about 1.5 km northwards over recent decades, whereas the European beech has extended its distribution southwards by about 2 km. Furthermore, the optimum altitude for both species has risen by about 200 m. As a result, the zone in which the two species coexist has been enlarged.? Conclusions
This approach provides a useful tool to compare NFI data from different sampling schemes, quantifying and testing significant shifts in tree species distribution over recent decades across geographical gradients. 相似文献15.
J. Julio Camarero Rubén D. Manzanedo Raúl Sanchez-Salguero Rafael M. Navarro-Cerrillo 《Annals of Forest Science》2013,70(8):769-780
Context
Tree populations at the rear edge of species distribution are sensitive to climate stress and drought. However, growth responses of these tree populations to those stressors may vary along climatic gradients.Aims
To analyze growth responses to climate and drought using dendrochronology in rear-edge Pinus nigra populations located along an aridity gradient.Methods
Tree-ring width chronologies were built for the twentieth century and related to monthly climatic variables, a drought index (Standardized Precipitation–Evapotranspiration Index), and two atmospheric circulation patterns (North Atlantic and Western Mediterranean Oscillations).Results
Growth was enhanced by wet and cold previous autumns and warm late winters before tree-ring formation. The influence of the previous year conditions on growth increased during the past century. Growth was significantly related to North Atlantic and Western Mediterranean Oscillations in two out of five sites. The strongest responses of growth to the drought index were observed in the most xeric sites.Conclusion
Dry conditions before tree-ring formation constrain growth in rear-edge P. nigra populations. The comparisons of climate-growth responses along aridity gradients allow characterizing the sensitivity of relict stands to climate warming. 相似文献16.
Lichao Wu Jie Liu Atsushi Takashima Keiichi Ishigaki Shin Watanabe 《Annals of Forest Science》2013,70(5):535-543
?Context
Selective logging followed by natural regeneration is rarely employed for restocking subtropical evergreen broad-leaved forests in East Asia compared with the use of clear-cutting.?Aims
To clarify the succession of these forests, the effects of selective logging on stand structure, species diversity, and community similarity were studied in a mature and regenerating forest in Okinawa, Japan.?Methods
Four study plots were established, and trees ≥1.2 m height were identified by species name, tree height, and diameter at breast height.?Results
The results showed that the species composition of regenerating forest was similar to mature forest; however, the former had a greater species density and Shannon–Wiener index than the latter. Castanopsis sieboldii and Distylium racemosum, the predominant trees in the mature forest, continued to dominate the regenerating forest, with a broad layer distribution. High Sørensen and Jaccard community similarity indices for mature and regenerating forest indicated that the regeneration occurred in a progressive succession.?Conclusion
The similar species composition and stand structure for both mature and regenerating forest, and the higher species diversity for the latter, provided no evidence of forest degeneration and suggested that the regenerating forest may develop into a stand similar to preselective logging forest. 相似文献17.
Yuhua Bai Ming Zhao Liming Bai Ryo Hasegawa Jun-ichi Sakai Toshiaki Hasegawa Tomokazu Mitsui Hirotsugu Ogura Takao Kataoka Katsutoshi Hirose Masayoshi Ando 《Journal of Wood Science》2011,57(1):56-65
Sixteen cardenolide triglycosides (1?C16) were isolated from stems, twigs, and leaves of Nerium oleander. Among them, 3??-O-(4-O-gentiobiosyl-d-diginosyl)-7??,8-epoxy-14-hydroxy-5??,14??-card-20(22)-enolide, named cardenolide B-3 (16), was isolated from natural sources for the first time. The in vitro anti-inflammatory activities of compounds 1?C16 were examined on the basis of inhibitory activity against the induction of intercellular adhesion molecule-1 (ICAM-1). Compounds 1?C5 were active at an IC50 value of less than 7 ??M. The cytotoxic activity of isolated compounds was evaluated against three human cell lines: normal human fibroblast cells (W-38), malignant tumor cells induced from WI-38 (VA-13), and human liver tumor cells (HepG2). Compounds 1?C5 were active toward WI-38 cells; compounds 1, 3, and 5 were active toward VA-13 cells; and compounds 1?C5 were active toward HepG2 cells at IC50 values of less than 10 ??M. The multidrug-resistant (MDR) cancer-reversal activity of compounds 1?C16 was evaluated on the basis of the amount of calcein accumulated in MDR human ovarian cancer 2780AD cells in the presence of each compound. Compounds 13 and 14 showed significant effects on calcein accumulation. 相似文献
18.
Alberto Vázquez de Castro Juan A. Oliet Jaime Puértolas Douglass F. Jacobs 《Annals of Forest Science》2014,71(1):91-99
Context
Tube shelters have been shown to enhance field performance of several Mediterranean species, but responses of newly planted seedlings to the microenvironment induced by shelter walls with different light transmissivity are still poorly documented.Aims
We studied effects of a range of shelters with varying light transmissivity on post-planting seedling responses during the wet season establishment phase for two Mediterranean trees of contrasting functional ecology.Methods
Root growth, biomass allocation, water potential, and chlorophyll fluorescence of Quercus ilex and Pinus halepensis seedlings were evaluated across shelters varying in light transmissivity (80, 40, 20, and 10 % plus a mesh shelter) with irrigation.Results
Plants in dark tubes (20 and 10 % light transmissivity) had less above- and belowground growth and more than two times greater leaf to protruding roots mass ratio, with shoot growth response of Q. ilex being less plastic. Ratio of leaf area/protruding roots area decreased when light transmissivity increased, although no differences were found at ≥40 % transmissivity. Xylem water potential indicated lack of water stress, and high maximum photosynthetic efficiency (F v/F m) values show no photoinhibition symptoms irrespective of light transmissivity.Conclusion
Shelter transmissivity ≥40 % promotes rapid and vigorous root growth immediately after planting for these species. This minimum transmissivity should be considered as a target when designing shelters to help root development and improve water balance of Mediterranean seedlings. 相似文献19.
Barbara Moser Christoph Bachofen Jonathan D. Müller Marek Metslaid Thomas Wohlgemuth 《Annals of Forest Science》2016,73(4):959-970
Key message
Pinus sylvestris seedlings quickly expand their roots to deeper soil layers while Pseudotsuga menziesii concentrates its root system in the topsoil, thereby running the risk of desiccation during long dry spells, as indicated by lower survival after simulated summer drought.Context
Pseudotsuga menziesii (Douglas-fir) is regarded as a promising species to maintain the productivity of Central European lowland forests given the projected increase of long dry spells.Aims
Will the species be able to regenerate from seed and spread outside plantations in a drier temperate Europe?Methods
We measured the relative growth rate, biomass allocation, root architecture, and phenotypic plasticity of Pseudotsuga menziesii seedlings sown in a common garden and grown under current precipitation and prolonged drought, respectively. The species’ competitive ability with respect to Pinus sylvestris L., the most drought-tolerant native conifer in Central Europe, was assessed during three growing seasons.Results
Pinus sylvestris seedlings had higher relative growth rates than did Pseudotsuga menziesii seedlings, first in terms of aboveground biomass and later in terms of shoot height. This resulted in heavier and taller seedlings after three growing seasons under both moist and dry conditions. Shorter vertical roots corresponded with lower survival of Pseudotsuga menziesii seedlings under dry conditions.Conclusion
Fast root proliferation allows Pinus sylvestris seedlings to reach deeper water pools that are less rapidly depleted during transient drought. By contrast, the shallow root system might put Pseudotsuga menziesii seedlings at the risk of desiccation during prolonged dry spells.20.
Anne-Sophie Sergent Nathalie Bréda Léopoldo Sanchez Jean-Charles Bastein Philippe Rozenberg 《Annals of Forest Science》2014,71(6):709-720