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1.
Detailed measurements of diurnal variations in photosynthetic photon flux density (PPFD) were made at seven locations within the canopy of aMiscanthus sinensis grassland to evaluate the light conditions of microsites for heliophilic tree seedlings. Multiple regression analysis revealed that the short-term light fluctuation on a clear day was highly dependent on the wind speed and solar elevation angle, whereas on a cloudy day it was mainly determined by the PPFD incident from above the canopy. The relative PPFD at 40 cm aboveground varied from 0.065 to 0.252, depending on sky conditions and the sensor's position in relation to clumped patches ofM. sinensis. On a clear day, the proportion of PPFD readings above 100 μmol·m−2·s−1 contributed by sunflecks ranged between 25.4% and 82.0%. Computer simulation showed that the contribution of sunflecks to the daily carbon gain ofQuercus serrata seedlings may range from 11% to 65%. The mean relative PPFD measured under diffuse light conditions was linearly related to the daily total PPFD and the daily carbon gain by single leaves ofQ. serrata seedlings. This suggests that the relative PPFD under diffuse light conditions provides an appropriate measure of site-specific light availability within a grass canopy.  相似文献   

2.
【背景】太阳光的入射辐射是影响多光谱扫描仪测定植物冠层光谱反射率的主要因素。本研究旨在探讨不同测定条件对空心莲子草冠层光谱特征值的影响,确定应用多光谱扫描仪测定空心莲子草冠层光谱特征的适宜环境条件。【方法】采用多光谱扫描仪,在不同天气条件、仪器探头高度、太阳高度角、叶片湿度等外界因素条件下,测定空心莲子草冠层光谱反射率的变化。【结果】天气条件、仪器探头高度、太阳高度角、叶片湿度4个因素对空心莲子草冠层在绿光区560nm、黄光区660nm和近红外光区810nm3个特征波段处的光谱反射率均有显著影响。【结论与意义】利用多光谱扫描仪测定空心莲子草冠层光谱数据,应选择晴朗无云的天气,仪器探头距离冠层1.3m,适宜的数据采集时间为10:00~14:00,以保障适度的太阳高度角和干燥的叶面,从而保持稳定的光谱测量结果。本研究可为完善多光谱扫描评价技术提供理论依据。  相似文献   

3.
Cloud cover increases the proportion of diffuse radiation reaching the Earth's surface and affects many microclimatic factors such as temperature, vapour pressure deficit and precipitation. We compared the relative efficiencies of canopy photosynthesis to diffuse and direct photosynthetic photon flux density (PPFD) for a Norway spruce forest (25‐year‐old, leaf area index 11 m2 m−2) during two successive 7‐day periods in August. The comparison was based on the response of net ecosystem exchange (NEE) of CO2 to PPFD. NEE and stomatal conductance at the canopy level (Gcanopy) was estimated from half‐hourly eddy‐covariance measurements of CO2 and H2O fluxes. In addition, daily courses of CO2 assimilation rate (AN) and stomatal conductance (Gs) at shoot level were measured using a gas‐exchange technique applied to branches of trees. The extent of spectral changes in incident solar radiation was assessed using a spectroradiometer. We found significantly higher NEE (up to 150%) during the cloudy periods compared with the sunny periods at corresponding PPFDs. Prevailing diffuse radiation under the cloudy days resulted in a significantly lower compensation irradiance (by ca. 50% and 70%), while apparent quantum yield was slightly higher (by ca. 7%) at canopy level and significantly higher (by ca. 530%) in sun‐acclimated shoots. The main reasons for these differences appear to be (1) more favourable microclimatic conditions during cloudy periods, (2) stimulation of photochemical reactions and stomatal opening via an increase of blue/red light ratio, and (3) increased penetration of light into the canopy and thus a more equitable distribution of light between leaves. Our analyses identified the most important reason of enhanced NEE under cloudy sky conditions to be the effective penetration of diffuse radiation to lower depths of the canopy. This subsequently led to the significantly higher solar equivalent leaf area compared with the direct radiation. Most of the leaves in such dense canopy are in deep shade, with marginal or negative carbon balances during sunny days. These findings show that the energy of diffuse, compared with direct, solar radiation is used more efficiently in assimilation processes at both leaf and canopy levels.  相似文献   

4.
Sigrid Dengel  John Grace 《Oecologia》2010,164(3):797-808
Sky conditions play an important role in the Earth’s climate system and CO2 uptake by plants. We used eddy covariance and meteorological data, including global and diffuse photosynthetic photon flux density (PPFD), recorded over the 2008 and 2009 growing season at two Sitka spruce [Picea sitchensis (Bong.) Carr.] forest sites in northern Britain, in order to establish relationships between physiological properties under diverse sky conditions, i.e. (1) sunny, (2) cloudy, and (3) overcast, and several canopy activity-related properties. These properties are: (1) response to PPFD, (2) photosynthetic light use efficiency, and (3) canopy stomatal conductance. We found that Sitka spruce forests utilise PPFD in a more efficient way when solar radiation is dominated by diffuse radiation. Furthermore, our results show that diffuse radiation enhances canopy stomatal conductance, an effect which may be the result of both blue light enrichment within the canopy and the reduction in vapour pressure deficit during cloudy and overcast weather. Diffuse radiation does not only influence short-term (hourly, daily, monthly) canopy activity but also long-term forest growth.  相似文献   

5.
An empirical light simulation model was applied to estimate stand scale photosynthesis in a deciduous broadleaved forest in central Japan. Based on diurnal courses of photosynthetically active photon flux density (PPFD), we characterized the components of incoming light within the forest canopy, and found that the instantaneous relative PPFD (PPFD under the canopy relative to that above the canopy) under diffuse light condition was a reliable estimator of the intensity and duration of PPFD. We calculated the daily photosynthesis (Aday) for each PPFD class using photosynthesis–light response curves. Model simulated Aday were corroborated with the estimates obtained from the nearby CO2 flux tower. The result demonstrated the potential of the light simulation model. The light use efficiency of two dominant species, Betula ermanii as overstory and Sasa senanensis as understory species, were then evaluated. At the forest understory, PPFD under 50 mol m–2 s–1 contributed to 77% of the sunshine duration on a completely clear day. Therefore, a higher apparent quantum yield for S. senanensis enhanced the utilization of low PPFD for photosynthesis. On the other hand, at the upper forest canopies, B. ermanii with a higher light-saturated photosynthetic rate used high PPFD efficiently. Consequently, potential of daily net photosynthesis for both B. ermanii and S. senanensis was high under each light condition. Such interspecific difference in the patterns of light utilization was suggested as one of factors allowing coexistence of the two species in the study forest.  相似文献   

6.
The spectral composition of photosynthetically active radiation (PAR) during clear and overcast days was studied above the canopy (U) and at two layers of a dense Norway spruce stand [Picea abies (L.) Karst.] characterized with an average LAI = 7.3 ± 0.8 (middle layer: M) and 12.3 ± 0.7 (lower layer: L). Whereas the spectral composition of PAR incoming on the canopy surface during cloudy days (characterized by diffuse index DI > 0.7) was almost independent of the solar elevation angle, the proportion of the blue-green spectral region of PAR was significantly reduced at low elevation angles during days with prevailing direct radiation (DI < 0.3). The PAR spectrum at both M and L levels was only slightly enriched in the green spectral region (more pronounced for DI < 0.3). The penetration of diffuse radiation into the canopy resulted in a slight (approx. 5%) reduction of the blue region proportion that remained stable during the day. On the contrary, under clear sky conditions the penetration of blue and red radiation was dependent on the solar elevation in an opposite manner in comparison with the spectral composition of PAR incident on canopy, giving almost twofold proportion of the blue part of the spectrum at a low elevation angle at M layer. We suggest that the blue enhancement of the spectrum within the Norway spruce canopy during clear days is due to a specific spatial arrangement of the assimilatory apparatus of a coniferous stand. Further, the possible consequences of the observed dynamics of the PAR spectrum inside the canopy during clear days on the efficiency of PAR absorption of the needles located within the canopy are discussed.  相似文献   

7.
紫耳箭竹克隆形态可塑性对典型冠层结构及光环境的响应   总被引:3,自引:0,他引:3  
黄慧敏  董蓉  钱凤  向运蓉  何丹妮  陈淼  陶建平 《生态学报》2018,38(19):6835-6845
在重庆金佛山国家自然保护内,选择了3种典型群落类型(落叶阔叶林、常绿落叶阔叶混交林和常绿阔叶林),使用Hemiview数字植物冠层分析系统量化群落冠层结构和光环境特征,并对林下紫耳箭竹(Fargesia decurvata)的形态可塑性特征进行调查,分析冠层结构和光环境特征改变下紫耳箭竹形态可塑性的差异,并探讨它们之间的相互关系。结果表明:(1)随着落叶阔叶林"常绿落叶阔叶混交林"常绿阔叶林演替的进行,群落的冠层开度降低,叶面积指数增加,平均叶倾角变小,趋于水平化,冠层对光的截获能力提高,林下光照的强度降低(P0.05)。(2)随着光照强度的降低,紫耳箭竹分株矮小化,叶片变窄,生物量积累降低,但通过增大比茎长、叶面积率和比叶面积提高对光的利用效率,并增大分枝角度和比隔长有效适应弱光环境。(3)在光照条件差的常绿阔叶林下,紫耳箭竹降低对地下茎的投资,将较多的生物量用于秆的增高增长和叶片的生长;而在光照条件好的落叶阔叶林环境下,紫耳箭竹降低对枝、叶生物量的分配,则加大对地下茎的投资,可认为是克隆植物对水分资源所表现的一种觅食行为。研究表明,紫耳箭竹种群随着冠层结构的改变发生了明显的可塑性变化,这些可塑性变化是种群对冠层结构和光环境差异的适应性反应的结果,有利于增强种群对异质生境中光资源的获取和利用;群落内部可以通过调控冠层结构的改变协调和控制小径竹种群的发展。  相似文献   

8.
The aim of this study is to explore the effects of canopy conditions on clump and culm numbers, and the morphological plasticity and biomass distribution patterns of the dwarf bamboo species Fargesia nitida. Specifically, we investigated the effects of canopy conditions on the growth and morphological characteristics of F. nitida, and the adaptive responses of F. nitida to different canopy conditions and its ecological senses. The results indicate that forest canopy had a significant effect on the genet density and culm number per clump, while it did not affect the ramet density. Clumps tended to be few and large in gaps and forest edge plots, and small under forest understory plots. The ramets showed an even distribution under the closed canopy, and cluster distribution under gaps and forest edge plots. The forest canopy had a significant effect on both the ramets’ biomass and biomass allocation. Favourable light conditions promoted ramet growth and biomass accumulation. Greater amounts of biomass in gaps and forest edge plots were shown by the higher number of culms per clump and the diameter of these culms. Under closed canopy, the bamboos increased their branching angle, leaf biomass allocation, specific leaf area and leaf area ratio to exploit more favourable light conditions in these locations. The spacer length, specific spacer length and spacer branching angles all showed significant differences between gaps and closed canopy conditions. The larger specific spacer length and spacer branching angle were beneficial for bamboo growth, scattering the ramets and exploiting more favourable light conditions. In summary, this study shows that to varying degrees, F. nitida exhibits both a wide ecological amplitude and high degree of morphological plasticity in response to differing forest canopy conditions. Moreover, the changes in plasticity enable the plants to optimize their light usage efficiency to promote growth and increase access to resources available in heterogeneous light environments. __________ Translated from Acta Ecologica Sinica, 2006, 26(12): 4019–4026 [译自: 生态学报]  相似文献   

9.
Ecophysiological comparisons were made of the growth and photosynthetic characteristics between seedlings of deciduousQuercus serrata and evergreenQuercus myrsinaefolia. Q. myrsinaefolia seedlings naturally occurring in secondary coppice forests showed exponential-like growth in height with age, while sympatricQ. serrata seedlings were considerably smaller in height, their growth being limited by shortage of light. The photosynthetic characteristics measured under laboratory conditions showed no bases for the differences in growth between the two species on the forest floor: Light compensation points of the seedlings raised under 5% daylight were almost identical for the two species, being about 6.0 μE·m−2·s−1. Growth analysis of seedlings planted in a coppice forest showed that bothQ. serrata andQ. myrsinaefolia could hardly grow during the summer under the shrub layer, when relative photon flux density (RPFD) was 0.9±0.5%. In the winter, when RPFD under the leafless canopy increased to 29.3±2.7%, the dry matter production of the evergreen seedlings ofQ. myrsinaefolia was much improved. Current-year seedlings of the species showed NAR of 0.102±0.021 g·dm−2·mo−1 during the winter. Temperature dependency of photosynthesis and increment of leaf temperature by direct solar beam also indicated active photosynthesis ofQ. myrsinaefolia on the forest floor during the winter.  相似文献   

10.
Leaf orientations and light environments were recorded for 40 juvenile Pseudopanax crassifolius trees growing in New Zealand in a partially shaded, secondary forest environment. Efficiencies of interception of diffuse and direct light by the observed leaf arrangments were calculated relative to those of three hypothetical leaf arrangements. Canopy gaps above the study plants were unevenly distributed with respect to azimuth and elevation above the horizon. Our results indicate that photosynthetically active radiation (PAR) received from the sides is more important than that received from directly above. In 33 of the plants leaf orientation was found to be significantly clustered towards one azimuth. The mean azimuth and the mean angle of declination were different for each plant. Leaves were steeply declined, and oriented towards the largest canopy gap at each site. Steep leaf angles reduced interception of direct and diffuse PAR when compared to interception by plant with a hypothetical horizontal leaf arrangement. When compared to a hypothetical arrangement with steep leaf declination and a uniform azimuth distribution, the observed leaf arrangement increased the efficiency of interception of diffuse PAR, but had a variable effect on the interception of direct PAR. Results indicate that the developing leaves of juvenile P. crassifolius orient towards the strongest sources of diffuse light, regardless of their value as a source of direct light. By maximising diffuse light interception while reducing direct light interception, leaf orientation may be a partial determinant of the types of habitats exploited by this species. This study emphasises the importance of considering diffuse light interception for plants growing in partially shaded environments.  相似文献   

11.
The transmission of direct, diffuse and global solar radiation in and around canopy gaps occurring in an uneven-aged, evergreen Nothofagus betuloides forest during the growing season (October 2006–March 2007) was estimated by means of hemispherical photographs. The transmission of solar radiation into the forest was affected not only by a high level of horizontal and vertical heterogeneity of the forest canopy, but also by low angles of the sun’s path. The below-canopy direct solar radiation appeared to be variable in space and time. On average, the highest amount of transmitted direct solar radiation was estimated below the undisturbed canopy at the southeast of the gap centre. The transmitted diffuse and global solar radiation above the forest floor exhibited lower variability and, on average, both were higher at the centre of the canopy gaps. Canopy structure and stand parameters were also measured to explain the variation in the below-canopy solar radiation in the forest. The model that best fit the transmitted below-canopy direct solar radiation was a growth model, using plant area index with an ellipsoidal angle distribution as the independent variable (R 2 = 0.263). Both diffuse and global solar radiation were very sensitive to canopy openness, and for both cases a quadratic model provided the best fit for these data (R 2 = 0.963 and 0.833, respectively). As much as 75% and 73% of the variation in the diffuse and global solar radiation, respectively, were explained by a combination of stand parameters, namely basal area, crown projection, crown volume, stem volume, and average equivalent crown radius.  相似文献   

12.
Muller O  Hikosaka K  Hirose T 《Oecologia》2005,143(4):501-508
In a temperate climate, evergreen species in the understory are exposed to large changes in photosynthetic photon flux density (PPFD) and temperature over the year. We determined the photosynthetic traits of leaves of an evergreen understory shrub Aucuba japonica at three sites at monthly intervals: understorys of a deciduous forest; an evergreen forest; and a gap in a mixed forest. This set up enabled us to separate the effects of seasonal change in PPFD and temperature on photosynthetic acclimation under natural conditions. The effects of PPFD and temperature were analysed by simple and multiple regression analyses. The amounts of light utilisation components (LU), represented by nitrogen and rubisco contents per area, were higher in winter, when temperature was low and PPFD was high. The LU relative to the amount of light harvesting components (LH), represented by chlorophyll a/b and rubisco/chlorophyll ratios, and the inverse of chlorophyll/nitrogen ratio were also higher in winter. We quantified the effects of PPFD and temperature on the LU and LH components. Across sites PPFD had stronger effects than air temperature, while within a site temperature had stronger effects on photosynthetic acclimation. We concluded that the photosynthetic apparatus is strongly affected by the prevailing PPFD at the time of leaf development. Within a given light regime, however, plants acclimated by increasing LU relative to LH primarily in response to temperature and to a lesser extent to PPFD.  相似文献   

13.
Yamasaki M  Kikuzawa K 《Oecologia》2003,137(2):226-232
This study investigated spatio-temporal variation in the leaf area consumed by insect herbivores within a canopy of Fagus crenata, with reference to the light conditions of leaf clusters. There was no clear relationship between photosynthetic photon flux density (PPFD) and consumed leaf area (CLA) in May, immediately after leaf flush, but CLA decreased with an increase in PPFD after June. Leaf mass per area, carbon concentration, C/N ratio, concentration of total phenolics, and condensed tannin concentration were higher in leaves under high light intensity than those of leaves under low light. On the other hand, the nitrogen concentration of leaves decreased as light availability increased. Consequently, within-tree variation in light availability affects the consumption of leaves by insect herbivores through temporal changes in leaf characteristics.  相似文献   

14.
M. Méthy  P. Alpert  J. Roy 《Oecologia》1990,84(2):265-271
Summary Plant canopy shade reduces photosynthetic photon flux density (PPFD) and ratio of red to far-red light (z). Both effects can cause plants to increase potential for light acquisition through vertical growth and leaf area expansion. Clonal plants such as Eichhornia crassipes might alternatively increase light interception via horizontal growth of stolons or rhizomes and placement of new ramets in less shaded microsites. Effect of simulated canopy shade and component effects of PPFD and z were tested by filtering or adding light uniformly, to a whole group of connected ramets, or locally, to individual ramets within a group. In uniform treatments, low PPFD reduced total growth but low z did not. Low PPFD and low z independently reduced stolon and ramet production and caused etiolation of petioles; effect of low PPFD plus low z on ramet production was greater than that of either factor alone. Lateral clonal growth thus did not seem to be a response to uniform shading; instead, uniformly low PPFD or low z increased partitioning to established ramets. Low z changed partitioning without changing total growth. In local treatments, reduction of growth of individual ramets due to low PPFD and inhibition of new ramet production attributable to spectral composition of light were mitigated when connected ramets were unshaded; plants may respond differently to patchy than to uniform shade.  相似文献   

15.
Abstract This paper demonstrates the changes in structure, floristics and forest floor light regimes across the boundary between open forest and rainforest at Kirrama in northeastern Queensland. Hemispherical photographs of the canopy were used to estimate spatial and temporal variations in potential (clear-sky) direct and diffuse photosynthetic photon flux density (PPFD = 400–700 nm) across the boundary. Five vegetation zones were identified across the boundary: (A) open forest, (B) tall open forest, (C) tall open forest with a rainforest under-storey, (D) young rainforest, and (E) mature rainforest. During the summer (September–March) direct PPFD declined semi-exponentially across the boundary, while in the winter (April–August), the decline was more linear. However, the decline in diffuse PPFD across the boundary was linear throughout the year. Mean per cent grass cover was positively correlated and mean per cent shrub cover negatively correlated with annual average total PPFD across the boundary. Mean percentages of diffuse radiation relative to that above the forest (diffuse site factors) ranged from 9.8% in the mature rainforest to 66.4% in the open forest, while mean percentages of direct radiation relative to that above the forest (direct site factors) ranged from 2.9–38.3% at the same sites, respectively. Daily PPFD in the rainforest understorey is probably insufficient for the shade intolerant (pioneer) species which prefer the more open environments on the rainforest margin where light intensities and soil temperatures are higher. Towards the rainforest end of zone C, light conditions appear to be similar to those experienced within small treefall gaps in rainforests. Typically, such light conditions are preferred by shade tolerant (primary) tree species that are unable to grow and reproduce successfully in the adjacent shaded understorey and to compete with the fast growing pioneer tree species in the more open end of zone C.  相似文献   

16.
Distribution of photosynthetic photon flux density (PPFD) in a culture vessel was estimated using a sensor film for measuring the integrated solar radiation. A plantlet model whose leaves were constructed from sensor film was used to simulate a potato plantlet. The plantlet model was put into a culture vessel and, after exposure to culture conditions for 20 days, PPFD was estimated for each individual model leaf based on the degree of fading of the sensor film. The method was able to demonstrate the light intensity distribution patterns inside a small test tube under downward lighting conditions. This technique allows for the estimation of light intensity distribution patterns inside a small culture vessel, which was previously difficult to measure by conventional methods.  相似文献   

17.
In a deciduous broad-leaved forest, we investigated the seasonality and heterogeneity of understory light conditions in relation to the leaf phenology of overstory trees. Fisheye photographs were taken from spring to autumn to estimate direct and indirect light conditions above the understory. Spatial variation in daily direct photosynthetic photon flux density (PPFD) was highest in late May, when the early and the intermediate-flushing trees had finished flushing and the late-flushing trees had just started flushing. After whole canopy closure, spatial variation in direct PPFD became low. Thus, asynchronous overstory leaf flushing in spring resulted in spatial heterogeneity in understory light conditions. These results suggest that the leaf phenological patterns of overstory trees are an important factor in the formation of the understory community.  相似文献   

18.
We investigated the morphological and physiological acclimation of leaves grown within a canopy of Japanese oak tree (Quercus mongolica var. crispula) in terms of the susceptibility to photoinhibition under various growth light conditions. The maximum rates of photosynthesis (P(max) ) and electron transport (ETR(max) ) were higher in mature leaves grown under stronger light with higher area-based leaf nitrogen (N) content closely associated with higher leaf mass per area. The net photosynthetic (P(n) ) and electron transport (ETR) rates corresponding to the daily peak photosynthetic photon flux density (PPFD(max) ) during leaf maturation were almost comparable to P(max) and ETR(max) , respectively. Conversely, P(n) and ETR at the daily average PPFD (PPFD(avg) ) were substantially low in shade-grown leaves when compared with P(max) and ETR(max) . The susceptibility to photoinhibition at PPFD(max) , i.e. at sunflecks for the shade-grown leaves, was assessed by the rate of excess energy production. Although sun leaves showed higher rates of electron transport and thermal energy dissipation than shade leaves under PPFD(max) conditions, the rate of excess energy production was almost constant across shade to sun leaves. The shade leaves of the Japanese oak grown within a crown were suggested to adjust their N investment to maintain higher photosynthetic capacities compared with those required to maximize the net carbon gain, which may facilitate the dissipation of the excessive light energy of sunflecks to circumvent photoinhibition in cooperation with thermal energy dissipation.  相似文献   

19.
Light environments in two lowland New Zealand podocarp rainforests are described using data from quantum sensors. Mean daily total photosynthetic photon flux density (PPFD) in the forest understorey varies from 2.6-5.2% incident PPFD in summer and 1.0-2.5% in autumn, and in gaps from 5.0-16.6% in summer and 6.3-8.3% in autumn. Pronounced differences in understorey PPFD occur between clear and overcast days. Overcast days tend to have a lower proportion of 2-minute periods with very low mean PPFD than clear days. In summer, 37.7-91.7% of PPFD occurs as sunflecks, but these only occur for 4.0-27.9% of the time. Most sunflecks are short duration (42.2-72.7% less than 4 minutes in summer) but some are very long (>32 minutes). Overall, light environments are similar to those found in other forests, both in New Zealand and elsewhere. However, canopy structure has a pronounced influence on spatial distribution of light environments within a forest, and differences in the size and frequency of canopy disturbances are a contributing factor to the floristic differences between the two forests studied.  相似文献   

20.
Spatial heterogeneity in light availability for tree seedlings under the canopy of a temperate pine forest was studied. Six-day measurements at 10-s intervals revealed a great variety in the temporal patterns of photosynthetic photon flux density (PPFD) and histograms among observation days and microsites; mean daily total PPFD relative to full sun varied from 1.5% to 10.4% depending on the microsites. The occurrence and duration of PPFD above 80 μmol m−2 s−1, which might reflect sunfleck activity, varied greatly among the microsites. However, several simple empirical relationships were found between some parameters characterizing microsite light availability and sunfleck activity; the diffuse site factor was correlated well with other parameters, including daily total PPFD, daily totals and daily summed durations of high PPFD above any examined threshold level, and its contribution to daily total PPFD. Diffuse site factors which were obtained for 700 microsites within an area of 28 m2 on three different occasions during the growing season showed high correlations within the microsite. Based on the regressed relationship between the relative growth rate of current-year seedlings ofQuercus serrata and the microsite diffuse site factor and the results of area-survey measurement of the diffuse site factor, an estimation was made of the abundance of potential ‘safe-sites’ for seedling growth of the species; the ‘safe sites’ were estimated to cover 40% and 0% of the total area of the sunny and shady sites of the forest, respectively.  相似文献   

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