Deposition of particulate matter of different size fractions on leaf surfaces and in waxes of urban forest species

Particulate matter (PM) is an air contaminant in urban and industrial areas that often exceeds limit values, creating serious problems due to its harmful effects on health. Planting trees and shrubs as air filters is a way to improve air quality in these areas. However,further knowledge on species effectiveness in air purification is essential This study compared four species of tree (Acer campestre L, Fraxinus excelsior L, Platanus x hispanica Mill. ex Muenchh. 'Acerifolia', Tilia cordata Mill.), three species of shrub (Forsythia x intermedia Zabel, Physocarpus opulifolius (L.) Maxim., Spiraea japonica L.), and one climber species (Hedera helix L) that are commonly cultivated along streets in Poland to capture fine, coarse and larger particles from air. Separate gravimetric analyses were performed to quantify PM deposited on surfaces and trapped in waxes. Significant differences were found between the plant species tested. The distribution of different particle size fractions differed between and within species and also between leaf surfaces and in waxes.     

广州市常见行道树种叶片表面形态与滞尘能力

以广州市常见的18种行道树为对象,通过扫描电镜观察比较了行道树的叶表面形态结构、应用接触角测定仪测定了绿化树种叶片的接触角对滞尘能力的影响.结果表明:不同树种的滞尘量差异显著,18种植物叶片雨后第26天的最大滞尘量在0.066-1.831 g/m2,物种间相差达27倍以上.叶表面具有网状结构,气孔密度较大(20＜气孔密度＜60个)且气孔开口较大(如芒果)容易滞留粉尘;叶表面平滑具有蜡质层,气孔排列整齐,无明显起伏(如红花羊蹄甲、桃花心木、大叶紫薇、鹅掌藤),滞尘能力较弱.植物叶片接触角与滞尘量呈负相关(r=-0.614),接触角＜90°的表现为亲水性.易润湿的植物叶片雨后第26天最大滞尘量在1.0-1.831 g/m2,叶片表面的形态结构凹凸不平,具有钩状或脊状褶皱、突起等且20＜气孔密度＜60范围内,测得的接触角较小(芒果、重阳木、高山榕),使得粉尘与植物叶片接触面积较大,粉尘不易从叶面脱落,滞尘能力较强.而接触角较大的盆架树、麻楝、大叶紫薇、鹅掌藤和红花羊蹄甲的滞尘量均＜1.0g/m2,其特殊的表面结构和疏水的蜡质使颗粒物不易吸附在植物叶片上,因此滞尘能力较弱.由此可见,植物叶表面蜡质含量和气孔密度及其叶片接触角的大小是影响植物叶片滞尘能力的主要因素,在进行城市绿化时,适当考虑选择叶表面形态有利于滞尘的绿化树种,将可提高城市植被的环境效应.     

Accumulation of particulate matter,heavy metals,and polycyclic aromatic hydrocarbons on the leaves of Tilia cordata Mill. in five Polish cities with different levels of air pollution

Urbanized areas are struggling with the problem of air pollution and as the number of people living in cities is increasing, the situation is likely to deteriorate. One of the most harmful pollutants is particulate matter (PM). Increased levels of PM in the atmosphere are likely to have a negative impact on human health. Phytoremediation technology could be a solution. It involves plants acting as bio-filters by accumulating particles on, and in the leaves, thus removing the particles from the atmosphere. This study investigates the accumulation of PM including heavy metals (HMs) and polycyclic aromatic hydrocarbons (PAHs), on the foliage of small-leaved lime (Tilia cordata Mill.) in five Polish cities. There were significantly different PM amounts found in the trees between the cities which related to the different quantities of PM in the atmosphere at these cities. Significant differences were found between cities for the amounts of the different particulate size fractions, and the HMs and PAHs in leaves. Strong winds reduced the amount of PM on leaves, especially the smallest fractions, but no relationship with precipitation was found. The results suggest that T. cordata improves the air quality in cities and can be used as an effective bioindicator for PM air pollution.     

城市绿地植物群落特征对亚微米颗粒物的影响

植物群落是构成城市绿地的基本单元之一,也是绿地发挥空气颗粒物调节功能的基础。亚微米级颗粒物,即PM1颗粒物对大气能见度、环境质量与人体健康等均存在严重的负面危害。为探究城市绿地植物群落对亚微米颗粒物水平的调节机制,本研究基于实地测量,针对不同类型植物群落内PM1颗粒物水平的差异,以及群落冠层结构对PM1浓度的影响开展定量研究。结果表明,植物群落对亚微米颗粒物具有一定的减滞效果。8种常见群落类型中,针阔-乔草型群落减滞PM1颗粒物的能力最优,阔叶-乔灌草群落次之,针叶-灌草型群落最弱,但不同类型植物群落内部的PM1浓度差异并不显著。植物群落的冠层结构同PM1颗粒物间存在明显的非线性响应关系,部分冠层指标存在关键的拐点阈值。当冠层郁闭度大于75%,疏透度小于55%左右时,植物群落减滞PM1颗粒物的效果最佳。     

北京城区行道树国槐叶面尘分布及重金属污染特征

叶面尘分布范围广、累积时间长,能够较好地指示近地面大气颗粒物污染特征和累积状况。为揭示城市道路交通的大气颗粒物污染特征,研究了北京城区不同类型道路两侧行道树国槐(Sophora japonica)的叶面滞尘量,分析了叶面尘主要重金属的浓度和污染特征,采用相关分析和主成分分析法探讨了颗粒物重金属的主要来源。结果表明:国槐叶面滞尘量在快速路、主干路、次干路、支路上的比值为100∶84∶75∶75,受车流量影响较大。行道树国槐的平均叶面滞尘量为0.68 g/m2,庭院树国槐(远离交通)叶面滞尘量是0.51 g/m2,两者差异显著。行道树叶面尘重金属元素Cu、Zn、Pb在叶面尘中的浓度达到土壤背景值的6倍,空间分布较均匀,主要来自交通排放,庭院树叶面尘重金属污染也受到交通排放的重要影响。     

C(4) photosynthesis in tree form euphorbia species from hawaiian rainforest sites

The ¹³C ¹²C isotope ratios and the leaf anatomy of 18 species and varieties of Euphorbia native to the Hawaian Islands indicated that all possess C₄ photosynthesis. These species range from small prostrate coastal strand shrubs to shrubs and trees in rainforest and bog habitats. The results show that C₄ photosynthesis occurs in plants from a much wider range of habitats and life-forms than has been previously reported.     

Variation in woody plant mortality and dieback from severe drought among soils, plant groups, and species within a northern Arizona ecotone

Vegetation change from drought-induced mortality can alter ecosystem community structure, biodiversity, and services. Although drought-induced mortality of woody plants has increased globally with recent warming, influences of soil type, tree and shrub groups, and species are poorly understood. Following the severe 2002 drought in northern Arizona, we surveyed woody plant mortality and canopy dieback of live trees and shrubs at the forest–woodland ecotone on soils derived from three soil parent materials (cinder, flow basalt, sedimentary) that differed in texture and rockiness. Our first of three major findings was that soil parent material had little effect on mortality of both trees and shrubs, yet canopy dieback of trees was influenced by parent material; dieback was highest on the cinder for pinyon pine (Pinus edulis) and one-seed juniper (Juniperus monosperma). Ponderosa pine (Pinus ponderosa) dieback was not sensitive to parent material. Second, shrubs had similar mortality, but greater canopy dieback, than trees. Third, pinyon and ponderosa pines had greater mortality than juniper, yet juniper had greater dieback, reflecting different hydraulic characteristics among these tree species. Our results show that impacts of severe drought on woody plants differed among tree species and tree and shrub groups, and such impacts were widespread over different soils in the southwestern U.S. Increasing frequency of severe drought with climate warming will likely cause similar mortality to trees and shrubs over major soil types at the forest–woodland ecotone in this region, but due to greater mortality of other tree species, tree cover will shift from a mixture of species to dominance by junipers and shrubs. Surviving junipers and shrubs will also likely have diminished leaf area due to canopy dieback.     

大青沟自然保护区主要森林群落优势种的叶性状

植物功能性状是近年来生态学研究的热点。其中叶功能性状与植株生物量和植物对资源的获得、利用及利用效率的关系最为密切。大青沟森林植物群落分布于科尔沁沙地,生境条件非常特殊,在沙沟里存在着一片茂密的森林,与周围浩瀚无垠的沙坨景观形成极为鲜明的对照。从沟底到沟顶,虽然海拔高度仅相差六、七十米,但由于距离沟底水源不同,土壤条件差异大,形成了不同的森林植物群落类型。大果榆群落、蒙古栎群落和水曲柳群落分别分布在大青沟自然保护区的沟顶、沟中和沟底。为了对大青沟自然保护区,不同环境梯度下的森林植物群落叶片功能性状进行研究,以大青沟自然保护区大果榆、蒙古栎、水曲柳3种主要森林群落为研究对象,分别测定不同群落优势种的叶厚度、比叶面积、叶干物质含量、叶大小和叶干重等5项叶功能性状,研究叶功能性状之间的关系,并对不同生长型、不同群落叶功能性状进行比较。相关分析结果表明,叶厚度与比叶面积呈极显著负相关,与叶大小和叶干重呈极显著正相关;比叶面积与叶干物质含量、叶干重呈极显著负相关,与叶大小呈显著正相关;叶干物质含量与叶大小呈极显著负相关,与叶干重呈极显著正相关;叶大小与叶干重呈极显著正相关。不同生长型植物叶片性状的分析表明,草本植物的叶干物质含量比乔木和灌木低,而其比叶面积高于乔木和灌木;对不同群落叶功能性状进行比较发现,大果榆群落和蒙古栎群落乔木、灌木叶厚度与干物质含量显著高于水曲柳群落的叶厚度与干物质含量,二者比叶面积显著低于水曲柳群落。大果榆群落和蒙古栎群落从叶功能性状的角度,它们具有较高的叶干物质含量和较低的比叶面积,体现出适应干旱生境叶片的特征。水曲柳群落呈现出低叶干物质含量、高比叶面积的特征,体现出适应湿润、土壤水分较好生境的特征。不同群落通过调节自身的物种组成,形成不同的功能性状组合来适应环境。     

Leaf litter of <Emphasis Type="Italic">Kageneckia angustifolia</Emphasis> D. Don (Rosaceae) inhibits seed germination in sclerophyllous montane woodlands of central Chile

Leaf litter accumulation can have either positive, negative or neutral effects on seed germination and seedling recruitment. In montane woodlands of the Mediterranean zone of central Chile, large amounts of leaf litter accumulate beneath the crowns of the summer semi-deciduous tree Kageneckia angustifolia and no regeneration of this or other plant species has been observed beneath this tree throughout the year. In a sample plot of 5000 m² we selected ten K. angustifolia trees and measured (1) leaf litter accumulation beneath and outside canopy; (2) the effects of time elapsed since burial on viability of K. angustifolia seeds with and without a leaf litter cover; (3) field seed germination with presence or absence of leaf litter and (4) the possible chemical effects of K. angustifolia leaf litter leachates on seed germination of its own seeds and of other two co-occurring native shrubs species (Guindilia trinervis and Solanum ligustrinum). Our results show that a considerable accumulation of leaf litter occurred beneath K. angustifolia, and litter negatively affected seed viability and germination of this species in the field. Under laboratory conditions, K. angustifolia leaf litter leachates inhibited seed germination of its own seeds and of the two native shrub species. Chemical effects are likely involved in the negative effects of leaf litter on the recruitment of K. angustifolia in the montane sclerophyllous woodland of central Chile.     

Effect of epicuticular wax crystals on the localization of artificially deposited sub-micron carbon-based aerosols on needles of <Emphasis Type="Italic">Cryptomeria japonica</Emphasis>

Elucidation of the mechanism of adsorption of particles suspended in the gas-phase (aerosol) to the outer surfaces of leaves provides useful information for understanding the mechanisms of the effect of aerosol particles on the growth and physiological functions of trees. In the present study, we examined the localization of artificially deposited sub-micron-sized carbon-based particles on the surfaces of needles of Cryptomeria japonica, a typical Japanese coniferous tree species, by field-emission scanning electron microscopy. The clusters (aggregates) of carbon-based particles were deposited on the needle surface regions where epicuticular wax crystals were sparsely distributed. By contrast, no clusters of the particles were found on the needle surface regions with dense distribution of epicuticular wax crystals. Number of clusters of carbon-based particles per unit area showed statistically significant differences between regions with sparse epicuticular wax crystals and those with dense epicuticular wax crystals. These results suggest that epicuticular wax crystals affect distribution of carbon-based particles on needles. Therefore, densely distributed epicuticular wax crystals might prevent the deposition of sub-micron-sized carbon-based particles on the surfaces of needles of Cryptomeria japonica to retain the function of stomata.     

The impact of epicuticular wax on gas-exchange and photoinhibition in Leucadendron lanigerum (Proteaceae)

This study investigated the seasonal modification of wax deposition, and the impact of epicuticular wax on gas-exchange as well as photoinhibition in Leucadendron lanigerum, a species from the Proteaceae family with wax-covered leaf surfaces and the stomata also partially occluded by wax. The results of this study demonstrated that the deposition of epicuticular wax in L. lanigerum is dependent on the age of the leaf as well as the season, and generation and regeneration of wax occur mostly in spring while transformation and also degeneration of wax crystals occur in winter. Epicuticular waxes decreased cuticular water loss, but had little impact on leaf reflectance. The temperature of leaves without wax was lower than that of wax-covered leaves, indicating that the rate of transpiration impacted more on leaf temperature than reflectance of light in the PAR range in L. lanigerum. The wax coverage at the entrance of stomata in L. lanigerum increased resistance to gas diffusion and as a consequence decreased stomatal conductance, transpiration and photosynthesis. Also, the results indicated that epicuticular waxes do help prevent photodamage in L. lanigerum, and so this property could benefit plants living in arid environments with high solar radiation.     

Variability in water use efficiency at the leaf level among Mediterranean plants with different growth forms

Assessing natural variability of leaf water use efficiency in plants adapted to extreme conditions of the Mediterranean climate represents an important step in the evaluation of the usefulness of some plant ecophysiological traits under water stress. Eleven Mediterranean species naturally inhabiting the Balearic Islands and corresponding to different growth forms (herbs, semi-deciduous shrubs, woody evergreen shrubs and woody evergreen semi-shrubs) were subject to progressive soil water depletion. Leaf intrinsic water use efficiency was measured by gas exchange at four different degrees of water stress. Under well watered conditions, differences in leaf intrinsic water use efficiency (A _N/g _s) among growth forms were limited to woody evergreen semi-shrubs, which presented the highest values. Under water stress conditions, differences became more evident, with a trend for an increase in A _N/g _s from woody evergreen shrubs, through semi-deciduous shrubs and herbaceous to woody evergreen semi-shrubs. The observed variation in A _N/g _s correlated with several physiological (leaf water potential, soil to leaf hydraulic conductance and stomatal conductance) and morphological (stomatal density) parameters, displaying a general relationship for all growth forms. This suggests that the capacity for withstanding water limitation is adaptive for all Mediterranean species. However, when A _N/g _s was related to leaf mass area, this relationship was not generally applicable, and depended on growth forms, suggesting that different growth forms display specific morphological adjustments in response to water shortage.     

Developmental Changes in Composition and Morphology of Cuticular Waxes on Leaves and Spikes of Glossy and Glaucous Wheat (Triticum aestivum L.)

The glossy varieties (A14 and Jing 2001) and glaucous varieties (Fanmai 5 and Shanken 99) of wheat (Triticum aestivum L.) were selected for evaluation of developmental changes in the composition and morphology of cuticular waxes on leaves and spikes. The results provide us with two different wax development patterns between leaf and spike. The general accumulation trend of the total wax load on leaf and spike surfaces is first to increase and then decrease during the development growth period, but these changes were caused by different compound classes between leaf and spike. Developmental changes of leaf waxes were mainly the result of variations in composition of alcohols and alkanes. In addition, diketones were the third important contributor to the leaf wax changes in the glaucous varieties. Alkanes and diketones were the two major compound classes that caused the developmental changes of spike waxes. For leaf waxes, β- and OH-β-diketones were first detected in flag leaves from 200-day-old plants, and the amounts of β- and OH-β-diketones were significantly higher in glaucous varieties compared with glossy varieties. In spike waxes, β-diketone existed in all varieties, but OH-β-diketone was detectable only in the glaucous varieties. Unexpectedly, the glaucous variety Fanmai 5 yielded large amounts of OH-β-diketone. There was a significant shift in the chain length distribution of alkanes between early stage leaf and flag leaf. Unlike C₂₈ alcohol being the dominant chain length in leaf waxes, the dominant alcohol chain length of spikes was C₂₄ or C₂₆ depending on varieties. Epicuticular wax crystals on wheat leaf and glume were comprised of platelets and tubules, and the crystal morphology changed constantly throughout plant growth, especially the abaxial leaf crystals. Moreover, our results suggested that platelets and tubules on glume surfaces could be formed rapidly within a few days.     

热岛效应下亚热带城市植被叶气孔权衡特征及其与叶功能性状的关系

叶片气孔不仅是植物平衡光合-蒸腾关系的重要门户,也是影响大气碳循环与水循环的关键结构。分析热岛效应下福州市乔木、灌木、草本3种生活型和常绿、落叶2种叶习性植物的气孔性状间的差异及其与其他叶功能性状间的权衡关系有助于探究不同类型植物在热环境下的适应策略。以福州市区的自然和半自然植被为研究对象,测定441个植物样本的气孔特征、化学计量特征和形态特征,结果表明：（1）3种生活型、2种叶习性植物的气孔长度（SL）、气孔密度（SD）差异显著（P<0.05）,潜在气孔导度指数（PCI）不存在显著差异（P>0.05）。草本的SL高于灌木和乔木,乔木的SD最高,灌木次之,草本最低;落叶植物的SL高于常绿植物,SD低于常绿植物。（2）SL与SD间的权衡关系稳定存在于3种生活型和2种叶习性植物中,且随着不同生活型和落叶习性植物的生态策略而呈现各异的权衡特征,即当SL一定时,乔木的SD最大,灌木的SD最小,常绿植物的SD大于落叶植物。（3）气孔性状和叶片形态、化学计量特征紧密联系,SL与比叶面积（SLA）正相关（P<0.01）,与叶面积（LA）负相关（P<0.01）;SD与叶氮含量（LNC）、叶磷含量（LPC）、SLA负相关（P<0.01）,与LA正相关（P<0.01）;PCI与LNC、SLA负相关（P<0.01）,与叶厚度（LT）正相关（P<0.05）。（4）复杂的环境是气孔性状变异的重要驱动因素,SL、PCI均与年均温（MAT）负相关（P<0.05）。     

Hydrogen isotopic compositions of n-alkanes from terrestrial plants correlate with their ecological life forms

Stable hydrogen isotopic compositions (δD) of compound-specific biomarkers, such as n-alkanes from plant leaf waxes, can be used as a proxy for paleoclimatic change. However, the relationship between hydrogen isotopes of plant leaf wax and plant ecological life forms is not well understood. Here, we report the δD of n-alkanes from 34 modern terrestrial plants, including twenty-one C₃ plants and thirteen C₄ plants from northwestern China, determined using gas chromatography/thermal conversion/isotope ratio mass spectrometry. Our data show that the stable hydrogen isotopes are poorly correlated with the plant photosynthetic pathway (C₃ vs. C₄) and that they do not give clear regional precipitation signals. Together with a comparative analysis of published δD values from plant leaf waxes in other regions, we believe that the stable hydrogen isotope of plant leaf waxes is more closely related to ecological life forms of these terrestrial plants (i.e. tree, shrub, and grass). In general, the grasses have more negative δD values than the co-occurring trees and shrubs. Our findings suggest that the δD values of sedimentary leaf waxes from higher plants may record changes of a plant ecosystem under the influence of environmental alteration and imply that reconstruction of the paleoclimate using δD values from plant n-alkanes should be based upon specific plant taxa, and comparison should be made among plants with similar ecological life forms.     

南京市14种绿化树种对空气中重金属的累积能力

分析了南京化工厂（污染区）和江苏省林科院（对照区）不同绿化树种叶片重金属元素（Pb、Cd和Cu）的含量,揭示了14种绿化树种对3种重金属污染物的累积能力。结果表明,城市绿化树种对大气重金属污染物具有一定的吸收净化能力,并依重金属和树种的不同具有明显差异;同种植物对不同重金属的累积量也有很大差别,其中对Cu的累积量最高;采样区植物叶片重金属含量明显高于对照区;对Pb累积量高的树种有：杨树、广玉兰、女贞和紫叶李; Cd累积量高的树种为杨树;Cu累积量高的树种为构树;累积3种重金属综合能力最强的树种有杨树、构树、雪松、广玉兰、悬铃木、栾树。     

Duvatrienediol,alkanes, and fatty acids in cuticular wax of tobacco leaves of various physiological maturity

4,8,13-Duvatriene-1,3-diol found in the leaf wax of Nicotiana tabacum was of highest concentration in the wax from young leaves, and quantitatively decreased in importance with leaf age. Tobacco plants in flower had less duvatrienediol than those that were less mature; however, the total wax content did not change with leaf age. Air drying leaves reduced the duvatrienediol concentration. Changes of total fatty acids and n-alkanes with leaf age and air drying were also examined.     

Traffic and industrial activities around Riyadh cause the accumulation of heavy metals in legumes: A case study

The objective of this study was to analyse the effect of the continuously increasing anthropogenic activities around Riyadh, Saudi Arabia on the accumulation of heavy metals in leguminous crops. This study determined whether four legume crops, Pisum sativum L., Vicia faba L., Glycine max and Vigna sinensis, could accumulate the heavy metals Cu, Mn, Pb and Zn in their leaves, pods and grains during the summer when grown under conditions with ambient air pollution from heavy traffic and industrial activities in Riyadh, Saudi Arabia. The effect of the air pollution was examined by quantifying the protein and trace element Cu, Mn, Pb and Zn concentrations in the leaves, pods and grains of the four plant species. Analysis of the results indicated that air pollution significantly increased the heavy metal concentrations in the leaves, pods and grains. Toxic concentrations of the heavy metals were found in the plants grown at L3, L4 and L5. In conclusion, the air pollution increases as the traffic, industrial activities and population density increase.     

Leaf growth dynamics in four plant species of the Patagonian Monte, Argentina

Studying plant responses to environmental variables is an elemental key to understand the functioning of arid ecosystems. We selected four dominant species of the two main life forms. The species selected were two evergreen shrubs: Larrea divaricata and Chuquiraga avellanedae and two perennial grasses: Nassella tenuis and Pappostipa speciosa. We registered leaf/shoot growth, leaf production and environmental variables (precipitation, air temperature, and volumetric soil water content at two depths) during summer-autumn and winter-spring periods. Multiple regressions were used to test the predictive power of the environmental variables. During the summer-autumn period, the strongest predictors of leaf/shoot growth and leaf production were the soil water content of the upper layer and air temperature while during the winter-spring period, the strongest predictor was air temperature. In conclusion, we found that the leaf/shoot growth and leaf production were associated with current environmental conditions, specially to soil water content and air temperature.     

Comparative ecophysiological responses to drought of two shrub and four tree species from karst habitats of southwestern China

Drought stress is one of the most important factors in limiting the survival and growth of plants in the harsh karst habitats of southwestern China, especially at the seedling establishment stage. The ecophysiological response to drought stress of native plants with different growth forms is useful for re-vegetation programs. Two shrub and four tree species were studied, including Pyracantha fortuneana (evergreen shrub), Rosa cymosa (deciduous shrub), Cinnamomum bodinieri (evergreen tree), and other three deciduous trees, Broussonetia papyrifera, Platycarya longipes, and Pteroceltis tatarinowii. The seedlings were randomly assigned to four drought treatments, i.e., well-watered, mild drought stress, moderate drought stress, and severe drought stress. Leaf water relations, gas exchange, chlorophyll fluorescence, and growth of the seedlings were investigated. Under severe drought stress, the two shrubs with low leaf area ratio (LAR) maintained higher water status, higher photosynthetic capacity, and larger percent biomass increase than the most of the trees. The two shrubs also had lower specific leaf area, greater intrinsic water use efficiency, and thermal dissipation than the trees. This suggested that the two shrubs had high tolerance to severe drought and were suitable for re-vegetation in harsh habitats. The evergreen C. bodinieri exhibited higher leaf mass ratio (LMR) and LAR than the deciduous species under mild and moderate stress. However, the low maximum quantum efficiency of PSII photochemistry (F _v/F _m) and net assimilation rate, and the sharp decreases of water potential, LMR, LAR, and biomass under severe stress indicated C. bodinieri’s weak tolerance to severe drought. In response to drought stress, the three deciduous trees revealed sharp reductions of biomass due to the large drought-induced decreases of gas exchange, LAR, and LMR. Under drought conditions, the deciduous trees minimized water loss by stomatal closure and by reducing transpiration leaf area and light harvesting through shedding leaves. This suggested that the three deciduous trees were more sensitive to water availability than the shrubs and used avoidance strategies against drought stress. However, the better growth performance of the deciduous trees than that of the shrubs under favorable conditions suggested that deciduous trees could be suitable for habitats with mild and temporary drought stress.