Interspecific variation in xylem vulnerability to cavitation among tropical tree and shrub species

In tropical moist forests, seasonal drought limits plant survival, productivity and diversity. Drought-tolerance mechanisms of tropical species should reflect the maximum seasonal water deficits experienced in a particular habitat. We investigated stem xylem vulnerability to cavitation in nine tropical species with different life histories and habitat associations. Stem xylem vulnerability was scored as the xylem water potential causing 50 and 75% loss of hydraulic conductivity (P50 and P75, respectively). Four shade-tolerant shrubs ranged from moderately resistant (P50=-1.9 MPa for Ouratea lucens Kunth. Engl.) to highly resistant to cavitation (P50=-4.1 MPa for Psychotria horizontalis Sw.), with shallow-rooted species being the most resistant. Among the tree species, those characteristic of waterlogged soils, Carapa guianensis Aubl., Prioria copaifera Griseb. and Ficus citrifolia Mill., were the most vulnerable to cavitation (P50=-0.8 to -1.6 MPa). The wet-season, deciduous tree, Cordia alliodora (Ruiz and Pav.) Oken., had resistant xylem (P50=-3.2 MPa), whereas the dry-season, deciduous tree, Bursera simaruba (L.) Sarg. was among the most vulnerable to cavitation (P50=-0.8 MPa) of the species studied. For eight out of the nine study species, previously reported minimum seasonal leaf water potentials measured in the field during periods of drought correlated with our P50 and P75 values. Rooting depth, deciduousness, soil type and growth habit might also contribute to desiccation tolerance. Our results support the functional dependence of drought tolerance on xylem resistance to cavitation.     

Interspecific variability of stem photosynthesis among tree species

The photosynthetic characteristics of current-year stems of six deciduous tree species, two evergreen tree species and ginkgo (Ginkgo biloba L.) were compared. Gas exchange, chlorophyll concentration, nitrogen concentration and maximum quantum yield of PSII were measured in stems in summer and winter. A light-induced decrease in stem CO(2) efflux was observed in all species. The apparent gross photosynthetic rate in saturating light ranged from 0.72 micromol m(-2 )s(-1) (ginkgo, in winter) to 3.73 micromol m(-2) s(-1) (Alnus glutinosa (L.) Gaertn., in summer). Despite this variability, a unique correlation (slope = 0.75), based on our results and those reported in the literature, was found between gross photosynthetic rate and dark respiration rate. Mass-based gross photosynthetic rate decreased with stem mass per area and correlated to chlorophyll concentration and nitrogen concentration, both in summer and winter. The radial distribution of stem chlorophyll differed among species, but all species except ginkgo had chlorophyll as deep as the pith. In summer, the maximum quantum yield of stem PSII (estimated from the ratio of variable to maximal fluorescence; F (v)/F (m)) of all species was near the optimal value found for leaves. By contrast, the values were highly variable in winter, suggesting large differences in sensitivity to low-temperature photoinhibition. The winter values of Fv/Fm were only 31-60% of summer values for the deciduous species, whereas the evergreen conifer species maintained high F (v)/F (m) in winter. The results highlight the interspecific variability of gross photosynthesis in the stem and its correlation with structural traits like those found for leaves. The structural correlations suggest that the selection of photosynthetic traits has operated under similar constraints in stems and leaves.     

Responses of leaf respiration to temperature and leaf characteristics in three deciduous tree species vary with site water availability

We measured responses of leaf respiration to temperature and leaf characteristics in three deciduous tree species (Quercus rubra L., Quercus prinus L. and Acer rubrum L.) at two sites differing in water availability within a single catchment in the Black Rock Forest, New York. The response of respiration to temperature differed significantly among the species. Acer rubrum displayed the smallest increase in respiration with increasing temperature. Corresponding Q(10) values ranged from 1.5 in A. rubrum to 2.1 in Q. prinus. Dark respiration at ambient air temperatures, expressed on a leaf area basis (Rarea), did not differ significantly between species, but it was significantly lower (P < 0.01) in trees at the wetter (lower) site than at the drier (upper) site (Q. rubra: 0.8 versus 1.1 micromol m(-2) s(-1); Q. prinus: 0.95 versus 1.2 micromol m(-2) s(-1)). In contrast, when expressed on a leaf mass basis (R(mass)), respiration rates were significantly higher (P < 0.01) in A. rubrum (12.5-14.6 micromol CO(2) kg(-1) s(-1)) than in Q. rubra (8.6-9.9 micromol CO(2) kg(-1) s(-1)) and Q. prinus (9.2-10.6 micromol CO(2) kg(-1) s(-1)) at both the lower and upper sites. Respiration on a nitrogen basis (R(N)) displayed a similar response to R(mass). The consistency in R(mass) and R(N) between sites indicates a strong coupling between factors influencing respiration and those affecting leaf characteristics. Finally, the relationships between dark respiration and A(max) differed between sites. Trees at the upper site had higher rates of leaf respiration and lower A(max) than trees at the lower site. This shift in the balance of carbon gain and loss clearly limits carbon acquisition by trees at sites of low water availability, particularly in the case of A. rubrum.     

Leaf litterfall concentrations and fluxes of elements in deciduous tree species

The influence of tree species and soil properties on leaf litterfall concentrations and fluxes of elements were studied in three mixed deciduous forest stands. Leaf litterfall fluxes of sixteen elements were measured during autumn defoliation in 100 to 150 yr old individuals of Fagus sylvatica L., Quercus robur L., Carpinus betulus L., Tilia cordata Mill, and Acer platanoides L. Compared to throughfall and precipitation, leaf litterfall dominated the flux of P, Ca, Mn, N and Mg (in decreasing order) to the soil below the tree canopies. For K and especially S and Na, however, the importance of leaf litterfall input to the total fluxes was small. Fluxes and especially concentrations of Ca, Mg, K, P and partly N were positively related to base saturation of the soil, while Mn was negatively related due to a higher mobilization rate and plant uptake of this element on acid soils. Litterfall fluxes for C. betulus were strongly positively influenced by base saturation and fluxes were usually higher than for F. sylvatica and Q. robur at the richest site. Quercus robur had often the lowest fluxes, especially of Ca and Mg. Tilia cordata had the highest leaf litter concentrations in twelve of the sixteen elements, and the highest fluxes particularly of N and K. Acer platanoides had the highest or among the highest concentrations and fluxes of Ca, Zn and Mn. The concentration of Al in C. betulus leaf litterfall, was about five times higher than in all other tree species at all sites.     

Evaluation of leaf display of evergreen broadleaved tree species and deciduous tree species in warm temperate conifer plantations

We investigated the sapling leaf display in the shade among trees of various leaf lifespans co-occurring under the canopy of a warm-temperate conifer plantation. We measured leaf-area ratio (aLAR) and morphological traits of saplings of evergreen broadleaved tree species and a deciduous tree species. Although we found large interspecific and intraspecific differences in aLAR even among saplings of similar size in the homogeneous light environment, we did not find a consistent trend in aLAR with leaf lifespan among the species. While deciduous trees annually produced a large leaf area, some evergreen broadleaved trees retained their leaves across years and had aLAR values as high as those of deciduous trees. Among leaf-level, shoot-level, and individual-level morphological traits, aLAR was positively correlated with current-year shoots mass per aboveground biomass in deciduous trees, and with the area of old leaves per aboveground mass in evergreen broadleaved trees. Thus, tree-to-tree variation in the degrees of annual shoot production and the accumulation of old leaves were responsible for the interspecific and intraspecific variations in aLAR.     

Interspecific variation in nighttime transpiration and stomatal conductance in a mixed New England deciduous forest

Transpiration is generally assumed to be insignificant at night when stomata close in response to the lack of photosynthetically active radiation. However, there is increasing evidence that the stomata of some species remain open at night, which would allow for nighttime transpiration if there were a sufficient environmental driving force. We examined nighttime water use in co-occurring species in a mixed deciduous stand at Harvard Forest, MA, using whole-tree and leaf-level measurements. Diurnal whole-tree water use was monitored continuously with Granier-style sap flux sensors in paper birch (Betula papyrifera Marsh.), red oak (Quercus rubra L.) and red maple (Acer rubrum L.). An analysis was conducted in which nighttime water flux could be partitioned between refilling of internal water stores and transpiration. Substantial nighttime sap flux was observed in all species and much of this flux was attributed to the refilling of depleted water stores. However, in paper birch, nighttime sap flux frequently exceeded recharge estimates. Over 10% of the total daily sap flux during the growing season was due to transpiration at night in paper birch. Nighttime sap flux was over 8% of the total daily flux in red oak and 2% in red maple; however, this flux was mainly associated with recharge. On nights with elevated vapor pressure deficit, sap flux continued through the night in paper birch, whereas it reached zero during the night in red oak and red maple. Measurements of leaf-level gas exchange on a night with elevated vapor pressure deficit showed stomatal conductance dropping by only 25% in paper birch, while approaching zero in red oak and red maple. The study highlighted differences in ecophysiological controls on sap flux exerted by co-occurring species. Paper birch is a fast-growing, shade-intolerant species with an earlier successional status than red oak and red maple. Risking water loss through nighttime transpiration may provide paper birch with an ecological advantage by enabling the species to maximize photosynthesis and support rapid growth. Nighttime transpiration may also be a mechanism for delivering oxygen to respiring cells in the deep sapwood of paper birch.     

Direct effect of elevated CO(2) on nocturnal in situ leaf respiration in nine temperate deciduous tree species is small

Direct (i.e., short-term) effects of elevated CO(2) on nocturnal in situ leaf respiration rate were measured in nine deciduous tree species (seven genera) in 20 3.5-4.0-h experiments. During the experiments, CO(2) concentration was alternated between 400 and 800 ppm (approximately 40 and 80 Pa of CO(2)). Data analysis accounted for effects on respiration rate of the normal decline in temperature with time after sunset. The median response to a 40-Pa increase in CO(2) was a 1.5% decrease in respiration rate, with responses ranging from a 5.6% inhibition to a 0.4% stimulation. Direct effects of elevated CO(2) on respiration were similar among the species. Thus, the response of nocturnal leaf respiration rate to a short-term CO(2) increase was small, and of little practical importance to the accuracy of measurements of respiration involving similar changes in CO(2) concentration during measurement. These direct respiratory responses of leaves to elevated CO(2) would translate into only slight, if any, effects on the carbon balance of temperate deciduous forests in a future atmosphere containing as much as 80 Pa CO(2).     

Sodium and chloride distribution in salt-stressed Prunus salicina, a deciduous tree species

Measurements were made over four growing seasons of the Na(+) and Cl(-) content of leaves and woody tissues (twigs, branches, trunk and roots) of mature, fruit-bearing Prunus salicina Lindl. (on Marianna 2624 rootstock) trees irrigated during the growing season with water containing 3, 14 or 28 mM salt (2/1 molar ratio of NaCl and CaCl(2)). At the beginning of the study, the trees were 19 years old. Woody tissues of trees irrigated with water containing 14 or 28 mM salt accumulated Na(+) and Cl(-). Leaves of trees irrigated with water containing 14 or 28 mM salt accumulated Cl(-), but not Na(+), unless they had visible symptoms of salt injury. X-Ray microanalysis of leaf mesophyll cells indicated some ability of the cells to sequester Cl(-) in the vacuole. The data demonstrate a capacity for ion compartmentation among tissues and cell organelles in mature Prunus salicina, which may explain the ability of the species to survive low levels of salinity for several years in the field.     

Retranslocation of foliar nutrients in evergreen tree species planted in a Mediterranean environment

Internal nutrient recycling through retranslocation (resorption) is important for meeting the nutrient demands of new tissue production in trees. We conducted a comparative study of nutrient retranslocation from leaves of five tree species from three genera grown in plantation forests for commercial or environmental purposes in southern Australia--Acacia mearnsii De Wild., Eucalyptus globulus Labill., E. fraxinoides H. Deane & Maiden, E. grandis W. Hill ex Maiden and Pinus radiata D. Don. Significant amounts of nitrogen, phosphorus and potassium were retranslocated during three phases of leaf life. In the first phase, retranslocation occurred from young leaves beginning 6 months after leaf initiation, even when leaves were physiologically most active. In the second phase, retranslocation occurred from mature green leaves during their second year, and in the third phase, retranslocation occurred during senescence before leaf fall. Nutrient retranslocation occurred mainly in response to new shoot production. The pattern of retranslocation was remarkably similar in the leaves of all study species (and in the phyllodes of Casuarina glauca Sieber ex Spreng.), despite their diverse genetics, leaf forms and growth rates. There was no net retranslocation of calcium in any of the species. The amounts of nutrients at the start of each pre-retranslocation phase had a strong positive relationship with the amounts subsequently retranslocated, and all species fitted a common relationship. The percentage reduction in concentration or content (retranslocation efficiency) at a particular growth phase is subject to many variables, even within a species, and is therefore not a meaningful measure of interspecific variation. It is proposed that the pattern of retranslocation and its governing factors are similar among species in the absence of interspecies competition for growth and crown structure which occurs in mixed species stands.     

Allometric differences between current-year shoots and large branches of deciduous broad-leaved tree species

Current-year shoots are mostly made of primary tissues, whereas first-order branches comprise mainly secondary tissues. Differences in tissue composition of these units reflect differences in functional design. We compared the allometry of current-year shoots and first-order branches in eight deciduous broad-leaved tree species and examined the functional differences underlying the design of current-year shoots and first-order branches. Allometric relationships of first-order branches tended to be compatible with predictions of the pipe model and elastic similarity model. That is, allometric constants of the relationships between leaf mass and stem diameter at the branch base and between stem diameter and stem mass were 2.0 and 0.33-0.38, respectively, indicating that the functional regulation of stem form of first-order branches can be predicted by the two models. However, allometric relationships of current-year shoots were not compatible with the predictions of the pipe and elastic similarity models. Thus, the allometric constant of the relationship between leaf mass and stem diameter at the base of current-year shoots was larger than 2.0, and the allometric constant of the relationship between stem length and stem diameter of shoots was larger than 1.0 in all species examined. However, current-year shoots had an allometric constant of leaf mass against stem length that was less than 1.0, suggesting a functional demand on shoot design to reduce self-shading. Also, allometric constants of stem length against stem diameter at the shoot base were larger in monopodial species than in sympodial species, whereas allometric constants of leaf mass per shoot against stem length were smaller in monopodial species than in sympodial species. We propose that the allometries of current-year shoots reflect their function as disposable units for temporary leaf arrangement.     

Spatial distributions of tropical tree species in northern Vietnam under environmentally variable site conditions

Ecological interactions of species and thus their spatial pat- terns may differ between homogeneous and heterogeneous forests. To account for this, techniques of point pattern analysis were implemented on mapped locations of tree individuals from two 1-ha tropicalforest plots in Vietnam. We analyzed the effect of environmental heterogeneity on tree distributions; spatial distribution patterns of dominant species; inter-specific associations; and conspecific associations between life stages. Our analyses showed that： environmental conditions were homo- geneous at plot 1 but heterogeneous at plot 2; in both plots, all six domi- nant species were aggregated at various scales up to 30 m, and tree spe- cies were aggregated at larger scales in the homogeneous site than in the heterogeneous site; attraction between pairs of species was remarkably higher at the homogeneous site while negative associations were more frequent in the heterogeneous site; some species, H. kurzii, T. ilicifolia （homogeneous plot） and D. sylvatica, S. wightianum （heterogeneous plot） showed a lack of early life-stage individuals near conspecific adults. Moreover, additional clustering of young individuals was independent from conspecific adults, except D. sylvatica in both sites. These findings are consistent with the Janzen-ConneU hypothesis. Overall, habitat het- erogeneity influences spatial patterns and inter-specific associations of the tree species and evidences of self-thinning are shown in most species.     

Oomycota species associated with deciduous and coniferous seedlings in forest tree nurseries of Western Turkey

Occurrence and pathogenicity of Oomycota species causing root rot were investigated in 10 forest tree nurseries in western Turkey. Soil samples (129 in total) taken from the rhizosphere of symptomatic seedlings were baited for Oomycota using young leaves of Quercus suber, Rhododendron simsii and R. ponticum. Oomycota (178 isolates) were obtained by culturing on selective media, and identified using morphological methods followed by PCR and sequencing of the ITS rDNA and cox1 regions. Phytophthora aff. cactorum, P. citricola sensu lato, P. crassamura, P. syringae, Pythium aphanidermatum, Py. intermedium, Py. irregulare, Py. ultimum and Phytopythium vexans were common amongst the isolates. The highest diversity of Oomycota was found in the forest nurseries at Adapazar?‐Hendek and ?zmir‐Torbal?. Pathogenicity tests showed that the isolates caused lesions on a range of host plants. The importance of these nursery infections in transferring potentially damaging oomycete species to Turkish forests is discussed.     

Foliage dark respiration in Abies amabilis (Dougl.) Forbes: variation within the canopy

Dark respiration of foliage was measured in a 30-year-old stand of Abies amabilis in western Washington from June to November. Both laboratory and field measurements were used to study the effect of environmental and tree variables on respiration. Foliage respiration rates were most strongly influenced by needle temperature. After accounting for leaf temperature differences, foliage respiration decreased with depth in the canopy for all age classes of foliage. Respiration differences attributed to location within the canopy were greatest early in the growing season, but were still significant in November. Younger foliage respired more than older foliage in the upper canopy, but not in the lower canopy. Respiration differences due to foliage age were highly significant in the early growing season, but were not detectable by mid-October.     

Component and whole-system respiration fluxes in northern deciduous forests

We measured component and whole-system respiration fluxes in northern hardwood (Acer saccharum Marsh., Tilia americana L., Fraxinus pennsylvanica Marsh.) and aspen (Populus tremuloides Michx.) forest stands in Price County, northern Wisconsin from 1999 through 2002. Measurements of soil, leaf and stem respiration, stem biomass, leaf area and biomass, and vertical profiles of leaf area were combined with biometric measurements to create site-specific respiration models and to estimate component and whole-system respiration fluxes. Hourly estimates of component respiration were based on site measurements of air, soil and stem temperature, leaf mass, sapwood volume and species composition. We also measured whole-system respiration from an above-canopy eddy flux tower. Measured soil respiration rates varied significantly among sites, but not consistently among dominant species (P < 0.05 and P > 0.1). Annual soil respiration ranged from 8.09 to 11.94 Mg C ha(-1) year(-1). Soil respiration varied linearly with temperature (P < 0.05), but not with soil water content (P > 0.1). Stem respiration rates per unit volume and per unit area differed significantly among species (P < 0.05). Stem respiration per unit volume of sapwood was highest in F. pennsylvanica (up to 300 micro mol m(3) s(-1)) and lowest in T. americana (22 micro mol m(3) s(-1)) when measured at peak summer temperatures (27 to 29 degrees C). In northern hardwood stands, south-side stem temperatures were higher and more variable than north-side temperatures during leaf-off periods, but were not different statistically during leaf-on periods. Cumulative annual stem respiration varied by year and species (P < 0.05) and averaged 1.59 Mg C ha(-1) year(-1). Leaf respiration rates varied significantly among species (P < 0.05). Respiration rates per unit leaf mass measured at 30 degrees C were highest for P. tremuloides (38.8 nmol g(-1) s(-1)), lowest for Ulmus rubra Muhlenb. (13.1 nmol g(-1) s(-1)) and intermediate and similar (30.2 nmol g(-1) s(-1)) for T. americana, F. pennsylvanica and Q. rubra. During the growing season, component respiration estimates were dominated by soil respiration, followed by leaf and then stem respiration. Summed component respiration averaged 11.86 Mg C ha(-1) year(-1). We found strong covariance between whole-ecosystem and summed component respiration measurements, but absolute rates and annual sums differed greatly.     

11种乡土阔叶树在广州南沙的早期生长表现

于2008—2011年连续监测了广州南沙黄山鲁森林公园11种乡土阔叶树种的高生长、胸径生长及冠幅生长,及与邻近白楸次生林的根区土肥力比较研究,探讨这些树种的生长特点及改良土壤的潜力。结果表明,千果榄仁和白锥生长不适,死亡率较高,其他树种大都生长良好。其中,楝叶吴茱萸、西南桦、米老排和山杜英等表现出较快的树高、胸径和冠幅生长。与邻近的白楸次生林相比,3年生试验林林木根区土的有机质较低;不同树种的根区土肥力比较结果表明,火力楠和米老排的根区土肥力较高,生长快速的西南桦和楝叶吴茱萸次之,生长缓慢的白锥的根区土肥力较低,反映了不同树种改良土壤的潜力有较大差异。初步研究结果表明,火力楠和米老排等树种有利于土壤肥力的改良,西南桦和楝叶吴茱萸等速生树种有利于快速提高植被的覆盖率。     

帽儿山地区四种树种树干呼吸的比较研究

树干呼吸是森林生态系统碳平衡的重要组成部分,它每年消耗碳同化总量(NPP) 的11%～33 %.本文采用Li-6400-09土壤呼吸气室于2010 年5 月测定了帽儿山地区4个主要树种(白桦、落叶松、山杨、红松)在不同高度、不同方位的树干呼吸速率,同时监测了树干表面温度.研究结果表明:树干呼吸速率随树干表面温度的升高而...     

Patterns of root respiration rates and morphological traits in 13 tree species in a tropical forest

The root systems of forest trees are composed of different diameters and heterogeneous physiological traits. However, the pattern of root respiration rates from finer and coarser roots across various tropical species remains unknown. To clarify how respiration is related to the morphological traits of roots, we evaluated specific root respiration and its relationships to mean root diameter (D) of various diameter and root tissue density (RTD; root mass per unit root volume; gcm(-3)) and specific root length (SRL; root length per unit root mass; mg(-1)) of the fine roots among and within 14 trees of 13 species from a primary tropical rainforest in the Pasoh Forest Reserve in Peninsular Malaysia. Coarse root (2-269mm) respiration rates increased with decreasing D, resulting in significant relationships between root respiration and diameter across species. A model based on a radial gradient of respiration rates of coarse roots simulated the exponential decrease in respiration with diameter. The respiration rate of fine roots (<2mm) was much higher and more variable than those of larger diameter roots. For fine roots, the mean respiration rates for each species increased with decreasing D. The respiration rates of fine roots declined markedly with increasing RTD and increased with increasing SRL, which explained a significant portion of the variation in the respiration among the 14 trees from 13 species examined. Our results indicate that coarse root respiration in tree species follows a basic relationship with D across species and that most of the variation in fine root respiration among species is explained by D, RTD and SRL. We found that the relationship between root respiration and morphological traits provides a quantitative basis for separating fine roots from coarse roots and that the pattern holds across different species.     

Quantifying stomatal and non-stomatal limitations to carbon assimilation resulting from leaf aging and drought in mature deciduous tree species

Gas exchange techniques were used to investigate light-saturated carbon assimilation and its stomatal and non-stomatal limitations over two seasons in mature trees of five species in a closed deciduous forest. Stomatal and non-stomatal contributions to decreases in assimilation resulting from leaf age and drought were quantified relative to the maximum rates obtained early in the season at optimal soil water contents. Although carbon assimilation, stomatal conductance and photosynthetic capacity (V(cmax)) decreased with leaf age, decreases in V(cmax) accounted for about 75% of the leaf-age related reduction in light-saturated assimilation rates, with a secondary role for stomatal conductance (around 25%). However, when considered independently from leaf age, the drought response was dominated by stomatal limitations, accounting for about 75% of the total limitation. Some of the analytical difficulties associated with computing limitation partitioning are discussed, including path dependence, patchy stomatal closure and diffusion in the mesophyll. Although these considerations may introduce errors in our estimates, our analysis establishes some reasonable boundaries on relative limitations and shows differences between drought and non-drought years. Estimating seasonal limitations under natural conditions, as shown in this study, provides a useful basis for comparing limitation processes between years and species.