Source water, phenology and growth of two tropical dry forest tree species growing on shallow karst soils

Seasonally dry tropical forests are dominated by deciduous and evergreen tree species with a wide range of leaf phenology. We hypothesized that Piscidia piscipula is able to extend leaf senescence until later in the dry season due to deeper and more reliable water sources than Gymnopodium floribundum, which loses leaves earlier in the dry season. Physiological performance was assessed as timing of leaf production and loss, growth, leaf water potential, depth of water uptake determined by stable isotopes, and leaf stable isotopic composition of carbon (δ¹³C) and oxygen (δ¹⁸O). P. piscipula took water primarily from shallow sources, whereas G. floribundum took water from shallow and deep sources. The greatest variation in water sources occurred during the onset of the dry season, when G. floribundum was shedding old leaves and growing new leaves, but P. piscipula maintained its leaves from the previous wet season. P. piscipula showed greater relative growth rate, greater leaf expansion rates, and more negative predawn and midday water potentials than G. floribundum. P. piscipula also exhibited greater leaf organic δ¹³C and lower δ¹⁸O values, indicating that the decrease in photosynthetic carbon isotope discrimination was associated with greater stomatal conductance and greater photosynthesis. Our results indicate that the contrasting early and late dry season leaf loss phenology of these two species is not simply determined by rooting depth, but rather a more complicated suite of characteristics based on opportunistic use of dynamic water sources, maximizing carbon gain, and maintenance of water potential during the dry season.     

Carbon assimilation,nitrogen, and photochemical efficiency of different Himalayan tree species along an altitudinal gradient

In the area of Jumla region in Western Nepal, measurements of saturated leaf net photosynthetic rate (P_sat), nitrogen content, leaf fluorescence, carbon isotopic composition, and water status were performed on woody coniferous (Pinus wallichiana, Picea smithiana, Abies spectabilis, Juniperus wallichiana, Taxus baccata), evergreen (Quercus semecarpifolia, Rhododendron campanulatum), and deciduous broadleaved species (Betula utilis, Populus ciliata, Sorbus cuspidata) spreading from 2 400 m up to the treeline at 4 200 m a.s.l. With the exception of J. wallichiana, P_sat values were lower in coniferous than broadleaved species. Q. semecarpifolia, that in this area grows above the coniferous belt between 3 000 and 4 000 m, showed the highest P_sat at saturating irradiance and the highest leaf N content. This N content was higher and P_sat lower than those of evergreen oak species of tempe forests at middle and low altitudes. For all species, P_sat and N content were linearly correlated, but instantaneous nitrogen use efficiency was lower than values measured in lowland and temperate plant communities. The values of carbon isotopic composition, estimated by δ₁₃C, showed the same range reported for temperate tree species. The ranking of δ₁₃C values for the different tree types was conifers < evergreen broadleaved<deciduous, suggesting tighter stomatal closure and higher water use efficiency for the evergreen types, confirming trends found elsewhere. No relevant differences of δ₁₃C were found along the altitudinal gradient. Quantum yield of photochemistry at saturating irradiance, measured by leaf fluorescence (δF/F_m’), was highest in J. wallichiana and lowest in T. baccata. Overall, photochemical efficiency was more strongly related to species than to altitude. Interestingly, changes of .δF/F_m’ along the altitudinal gradient correlated well with the reported altitudinal distribution of the species.     

Changes of Leaf Morphological, Anatomical Structure and Carbon Isotope Ratio with the Height of the Wangtian Tree （Parashorea chinensis） in Xishuangbanna, China

Leaf morphological and anatomical structure and carbon isotope ratio （δ^13C） change with increasing tree height. To determine how tree height affects leaf characteristics, we measured the leaf area, specific leaf mass （ratio of leaf mass to leaf area [LMA]）, thickness of the total leaf, cuticle, epidermis, palisade and sponge mesophyll, stomata traits and δ^13C at different heights of Parashorea chinensis with methods of light and scanning electron microscopy （SEM） and isotope-ratio mass spectrometry. The correlation and stepwise regression between tree height and leaf structure traits were carried out with SPSS software. The results showed that leaf structures and δ^13C differed significantly along the tree height gradient. The leaf area, thickness of sponge mesophyll and size of stomata decreased with increasing height, whereas the thickness of lamina, palisade mesophyll, epidermis, and cuticle, ratios of palisade to spongy thickness, density of stomata and vascular bundles, LMA and δ^13C increased with tree height. Tree height showed a significant relationship with all leaf indices and the most significant relationship was with epidermis thickness, leaf area, cuticle thickness, δ^13C. The δ^13C value showed a significantly positive relationship with LMA （R = 0.934）. Our results supported the hypothesis that the leaf structures exhibited more xeromorphic characteristics with the increasing gradient of tree height.     

Leaf carbon isotope ratio and water use efficiency of urban roadside trees in summer in Kyoto city

The effect of summer climate on leaf carbon isotope composition (δ¹³C) of the major roadside tree species Prunus × yedoensis (P. yedoensis) was investigated in Kyoto city, Japan, to explore the implications for alterations in urban environments. Temperature and the vapor pressure deficit were higher at sites of higher traffic volumes, possibly affected by a heat island effect. The leaf δ¹³C of P. yedoensis trees was affected strongly by leaf carbon isotope discrimination (Δ), with much less effect of δ¹³C on atmospheric CO₂. Leaf Δ values in the summer were smaller at sites of higher traffic volumes with high atmospheric temperatures, suggesting a higher long-term water use efficiency (WUE) at these sites. Gas exchange measurements of P. yedoensis leaves indeed suggested a higher intrinsic WUE at sites of higher traffic volumes with high atmospheric temperatures. These results suggest that leaf Δ is related to the response of WUE to summer climates, and that leaf δ¹³C in urban areas is a useful tracer for understanding the influences of urban environments on plant photosynthetic processes.     

Individual tree context and contrast dictate tree physiological features and arthropod biodiversity patterns across multiple trophic levels

1. The present study tested the hypothesis that tree context (natural, semi-natural or planted) and contrast (height of surrounding vegetation) affect tree physiological characters (leaf size, nutrient content, and stress-related factors), and also alter the arthropod biodiversity patterns either directly or indirectly. 2. Arthropods were collected from tree canopies using chemical fogging from the ecologically important South African native tree Podocarpus elongatus. 3. Low contrast trees had significantly larger leaves than those in high contrast environments and harboured an overall higher richness, abundance, and biomass of arthropods, although this was guild-dependent. 4. Trees in natural contexts had less foliar δ¹³C, suffered less from artificial nitrogen enrichment, and harboured significantly higher herbivore and predator arthropod richness and unique assemblages, compared with planted trees. 5. Semi-natural trees supported natural levels of arthropod richness, although these were mostly generalist species. 6. Tree context and contrast can therefore dictate associated biota at multiple trophic levels and native trees may fail to maintain natural biodiversity in transformed landscapes.     

Plasticity of photosynthetic performance of the Indian tree Butea monosperma TAUB. at three sites with different microclimates

The Fabaceae tree Butea monosperma (TAUB.; syn. Erythrina monosperma (LAM.)) is widely distributed in Central and West-India. We studied it at three sites, i.e. at two locations with contrasting exposure (NE and SW, respectively) in a small mountain range with poor soil on highly drained rocky slopes and at a third location in a plane with deeper soils and better water supply. The two mountain range sites differed in the light climate where the NE-slope obtained more day-integrated irradiance. Chlorophyll fluorescence was measured with a portable fluorometer and leaf samples for stable isotope analyses (δ¹³C, δ¹⁵N, δ¹⁸O) were collected. No differences were seen in carbon and nitrogen contents of leaves at the three sites. N and O isotope signatures of the leaves were similar at the two rocky hill slope sites. More positive values for both signatures were obtained in the leaves in the plane. For all sites saturation of ETR was only achieved well above a PPFD of 1,000 μmol m^?2 s^?1 indicating that the leaves were sun-type leaves. The photosynthetic performance of Butea at the plane was very similar to that at the SW-slope of the mountain range and higher ETRs were obtained at the NE-slope. Ecophysiological flexibility allows Butea to perform well in a variety of habitats and yet gives it particular fitness at specific sites. The best performance was observed in the highly insolated steep rocky hill site (NE-slope) underlining the suitability of the tree for reforestation.     

Impacts of eriophyoid gall mites on arctic willow in a rapidly changing Arctic

Arctic plants and herbivores are subject to ongoing climatic changes that are more rapid and extreme than elsewhere on the planet, and thus it is pivotal to understand the arctic plant-herbivore interactions in a global change context. We examined how infestation by an eriophyoid gall mite affects the circumpolar shrub Salix arctica, and how the effects vary across vegetation types. Specifically, we compared multiple leaf characteristics (leaf area, biomass, nutrient levels, δ¹⁵N and δ¹³C, and stress and performance of the photosynthetic apparatus) of infested leaves to those of un-infested leaves. Furthermore, we examined how altered environmental conditions, here experimentally manipulated levels of temperature, water and nutrients, shading, and UV-B radiation, affect the prevalence, density, and intensity of gall mite infestation and its impacts on S. arctica. Infested leaves were smaller in area and biomass and had lower nitrogen and carbon pools. However, their carbon concentration was higher, possibly because the galls acted as carbon sinks. The smaller photosynthetic area and lower nutrient content caused increased stress on the photosynthetic apparatus in infested leaves. The remaining leaf tissue responded with a higher photosynthetic performance, although there were indications of a general reduction in photosynthesis. Female leaves were more affected than male leaves. The experimental manipulations of environmental conditions did not affect the gall prevalence, density, or intensity on S. arctica leaves. Rather, plants responded positively to the treatments, reducing the effects of the galls to in-significance. This suggests a higher tolerance and defense against gall mites under future climate conditions.     

Species-specific differences in temporal and spatial variation in δ13C of plant carbon pools and dark-respired CO2 under changing environmental conditions

Stable carbon isotope signatures are often used as tracers for environmentally driven changes in photosynthetic δ¹³C discrimination. However, carbon isotope signatures downstream from carboxylation by Rubisco are altered within metabolic pathways, transport and respiratory processes, leading to differences in δ¹³C between carbon pools along the plant axis and in respired CO₂. Little is known about the within-plant variation in δ¹³C under different environmental conditions or between species. We analyzed spatial, diurnal, and environmental variations in δ¹³C of water soluble organic matter (δ¹³C_WSOM) of leaves, phloem and roots, as well as dark-respired δ¹³CO₂ (δ¹³C_res) in leaves and roots. We selected distinct light environments (forest understory and an open area), seasons (Mediterranean spring and summer drought) and three functionally distinct understory species (two native shrubs—Halimium halimifolium and Rosmarinus officinalis—and a woody invader—Acacia longifolia). Spatial patterns in δ¹³C_WSOM along the plant vertical axis and between respired δ¹³CO₂ and its putative substrate were clearly species specific and the most δ¹³C-enriched and depleted values were found in δ¹³C of leaf dark-respired CO₂ and phloem sugars, ~?15 and ~?33 ‰, respectively. Comparisons between study sites and seasons revealed that spatial and diurnal patterns were influenced by environmental conditions. Within a species, phloem δ¹³C_WSOM and δ¹³C_res varied by up to 4 ‰ between seasons and sites. Thus, careful characterization of the magnitude and environmental dependence of apparent post-carboxylation fractionation is needed when using δ¹³C signatures to trace changes in photosynthetic discrimination.     

Optimal photosynthetic use of light by tropical tree crowns achieved by adjustment of individual leaf angles and nitrogen content

Background and Aims

Theory for optimal allocation of foliar nitrogen (ONA) predicts that both nitrogen concentration and photosynthetic capacity will scale linearly with gradients of insolation within plant canopies. ONA is expected to allow plants to efficiently use both light and nitrogen. However, empirical data generally do not exhibit perfect ONA, and light-use optimization per se is little explored. The aim was to examine to what degree partitioning of nitrogen or light is optimized in the crowns of three tropical canopy tree species.

Methods

Instantaneous photosynthetic photon flux density (PPFD) incident on the adaxial surface of individual leaves was measured along vertical PPFD gradients in tree canopies at a frequency of 0·5 Hz over 9–17 d, and summed to obtain the average daily integral of PPFD for each leaf to characterize its insolation regime. Also measured were leaf N per area (N_area), leaf mass per area (LMA), the cosine of leaf inclination and the parameters of the photosynthetic light response curve [photosynthetic capacity (A_max), dark respiration (R_d), apparent quantum yield (ϕ) and curvature (θ)]. The instantaneous PPFD measurements and light response curves were used to estimate leaf daily photosynthesis (A_daily) for each leaf.

Key Results

Leaf N_area and A_max changed as a hyperbolic asymptotic function of the PPFD regime, not the linear relationship predicted by ONA. Despite this suboptimal nitrogen partitioning among leaves, A_daily did increase linearly with PPFD regime through co-ordinated adjustments in both leaf angle and physiology along canopy gradients in insolation, exhibiting a strong convergence among the three species.

Conclusions

The results suggest that canopy tree leaves in this tropical forest optimize photosynthetic use of PPFD rather than N per se. Tropical tree canopies then can be considered simple ‘big-leaves’ in which all constituent ‘small leaves’ use PPFD with the same photosynthetic efficiency.Key words: Optimal resource allocation, nitrogen, photosynthetic capacity, leaf mass per area, tropical trees, radiation use efficiency, scaling, leaf angle, canopy architecture, big leaf model     

Stable carbon isotope ratio of tree leaves, boles and fine litter in a tropical forest in Rondônia, Brazil

Leaves of 208 trees were collected for isotopic analysis together with wood from 36 tree boles and 18 samples of fine litter from a terra-firme forest located at Samuel Ecological Reserve, Rondônia State, in the southwestern Amazon region. The range of δ¹³C values in leaves was from ?28 to ?36‰, with an average (±1 SD) of ?32.1?±?1.5‰, which was more negative than the δ¹³C values of bole samples (?28.4?±?2.0‰) and fine litter (?28.7?±?2.0‰). These values are within the range found for tropical and subtropical forests. Pooling the δ¹³C values for leaf samples from trees of the same height gave averages which were positively correlated with plant height at a highly significant level, with a slope of 0.06 and an intercept of ?33.3‰ and a correlation coefficient r ²=0.70 (P<0.001).     

Species differences in carbon isotope ratios, specific leaf area and nitrogen concentrations in leaves of Eucalyptus growing in a common garden compared with along an aridity gradient

Leaves produced in 2004 of 422 species of Eucalyptus whose natural habitat is southern Australia were sampled at the Currency Creek Arboretum in South Australia where the annual (mainly winter) rainfall is about 400 mm. Tree height, leaf area, leaf dry weight, leaf nitrogen (N) concentration and leaf carbon isotope ratio (δ¹³C) were measured and the specific leaf area (SLA) calculated. Among the 422 species, the SLA varied from 1.5 to 8.8 m² kg^?1 and N concentration varied from 0.6 to 2.1%, much greater than in 64 species collected along an aridity transect from southwestern Western Australia to central Australia in 2003. Also, the range of leaf δ¹³C values was similar in the common garden to that across the aridity transect. For the 45 species present in both studies, the SLA and leaf N concentration in the common garden were similar to those measured in leaves along the aridity transect, indicating that these characteristics are inherent in the species and vary little with environment. The variation in leaf δ¹³C in the common garden was just as great as along the transect, but the values measured in the one location were poorly correlated with those along the transect. This was not expected, as the variation in δ¹³C at one common site in South Australia was anticipated to be less than along the aridity gradient where annual rainfall varied from 250 to 1200 mm. Path analysis on the 45 species common to both studies indicated that rainfall did not have a direct effect on δ¹³C, but the differences in δ¹³C resulted from indirect effects of rainfall on SLA and N concentration. δ¹³C was negatively correlated with SLA but positively correlated with N. Thus, both effects may compensate for each other so that no significant relationship between δ¹³C and rainfall was observable. However, there is a large degree of variation of δ¹³C at any level of rainfall. The origin and ecological implications of this observation are discussed.     

Growth and photosynthetic responses in Jatropha curcas L. seedlings of different provenances to watering regimes

Seedlings from four provenances of Jatropha curcas were subjected to 80, 50, and 30% of soil field capacity in potted experiments in order to study their responses to water availability. Our results showed that with the decline of soil water availability, plant growth, biomass accumulation, net photosynthetic rate, stomatal conductance (g_s), and transpiration rate (E) decreased, whereas leaf carbon isotope composition (δ¹³C), leaf pigment contents, and stomatal limitation value increased, while maximal quantum yield of PSII photochemistry was not affected. Our findings proved that stomatal limitation to photosynthesis dominated in J. curcas under low water availability. The increase of δ¹³C should be attributed to the decrease in g_s and E under the lowest water supply. J. curcas could adapt to low water availability by adjusting its plant size, stomata closure, reduction of E, increasing δ¹³C, and leaf pigment contents. Moreover, effects of provenance and the interaction with the watering regime were detected in growth and many physiological parameters. The provenance from xeric habitats showed stronger plasticity in the plant size than that from other provenances under drought. The variations may be used as criteria for variety/provenance selection and improvement of J. curcas performance.     

Foliar and litter needle carbon and oxygen isotope compositions relate to tree growth of an exotic pine plantation under different residue management regimes in subtropical Australia

Background and aim

Significant differences in tree growth were observed in an exotic pine plantation under different harvest residue management regimes at ages 2–10 years. However, the variations in tree growth between residue management treatments could not be explained by soil and foliar nutrient analyses, except by potassium (K) concentration. Therefore, this study determined the carbon isotope composition (δ¹³C) and oxygen isotope composition (δ¹⁸O) of current and archived foliar samples from the exotic pine plantation to establish relationships with foliar K concentration and tree growth indices as a means to determine changes in stomatal conductance (g_s) and photosynthetic rate (A_max) or water use efficiency (WUE), and therefore understand the variations in tree growth across treatments.

Methods

The harvest residue treatments were: (1) residue removal, RR₀; (2) single level residue retention, RR₁; and (3) double level residue retention, RR₂. Foliar δ¹³C and δ¹⁸O were determined for samples at ages 2, 4, 6 and 10 years, and the atmospheric ¹³C discrimination (Δ¹³C), intercellular CO₂ concentration (C_i) and WUE were determined from the δ¹³C data. Litter needle δ¹³C and δ¹⁸O were also determined over 15 months between ages 9 and 10 years. These parameters or variables where correlated to each other as well as to the periodic mean annual increment of basal area (PAIB) and the periodic mean annual increment of tree diameter at breast height (PAID) across the treatments and over time. Foliar δ¹³C and δ¹⁸O were also related to published data of foliar K concentrations of the same trees.

Results

Significant variations of foliar δ¹³C, and therefore WUE and Δ¹³C, across treatments were only observed at ages 4 and 10 years old, and foliar δ¹⁸O at age 4 years old only. The results showed increasing foliar δ¹³C, δ¹⁸O and WUE, and decreasing Δ¹³C and Ci, from RR₀ to RR₂ treatments. However, while the WUE was positively related to the PAID and PAIB at age 4 years, it was negatively related to PAID and PAIB at age 10 years old. Litter needle δ¹³C, indicative of WUE, was also negatively related to the PAID at age 10 years old. . At age 4 years, foliar δ¹³C and δ¹⁸O were positively related with a steep slope of 7.70 ‰ across treatments, and that both isotopes were positively related to foliar K concentrations. Similarly, δ¹⁸O was negatively related to the Δ¹³C. No significant relationship can be determined between foliar δ¹³C, or Δ¹³C, and δ¹⁸O at age 10 years old. In addition, WUE was increasing (p?<?0.001) and Δ¹³C and C_i decreasing (p?<?0.001) with decreasing PAID over time.

Conclusions

The variations at age 4 years in foliar δ¹³C or Δ¹³C and δ¹⁸O and increasing WUE with increasing growth rate suggest growth induced water-stress with increasing residue-loading rate as a result of the nutritional effect of the harvest residues on tree growth. At age 10 years, the negative relationships between WUE and PAID indicate nutrient limitation has an over-riding effect on δ¹³C variations rather than g_s. This was due to the lack of a significant relationship between foliar Δ¹³C and δ¹⁸O at this age, as well as over time.     

Significance of mesophyll conductance for photosynthetic capacity and water-use efficiency in response to alkaline stress in Populus cathayana seedlings

Cuttings of Populus cathayana were exposed to three different alkaline regimes (0, 75, and 150 mM Na₂CO₃) in a semicontrolled environment. The net photosynthesis rate (P _N), mesophyll conductance (g _m), the relative limitations posed by stomatal conductance (L _s) and by mesophyll conductance (L _m), photosynthetic nitrogen-use efficiency (PNUE), carbon isotope composition (δ¹³C), as well as specific leaf area (SLA) were measured. P _N decreased due to alkaline stress by an average of 25% and g _m decreased by an average of 57%. Alkaline stress caused an increase of L _m but not L _s, with average L _s of 26%, and L _m average of 38% under stress conditions. Our results suggested reduced assimilation rate under alkaline stress through decreased mesophyll conductance in P. cathayana. Moreover, alkaline stress increased significantly δ¹³C and it drew down CO₂ concentration from the substomatal cavities to the sites of carboxylation (C _i-C _c), but decreased PNUE. Furthermore, a relationship was found between PNUE and C _i-C _c. Meanwhile, no correlation was found between δ¹³C and C _i/C _a, but a strong correlation was proved between δ¹³C and C _c/C _a, indicating that mesophyll conductance was also influencing the ¹³C/¹²C ratio of leaf under alkaline stress.     

Leaf characteristics and diurnal variation of chlorophyll fluorescence in leaves of the ‘bana’ vegetation of the amazon region

In six dominant species of the Amazonian ‘Bana’ vegetation, leaf blade characteristics, pigment composition, and chlorophyll (Chl) fluorescence parameters were measured in young and mature leaves under field conditions. Leaf δ¹³C was comparable in the six species, which suggested that both expanding and expanded leaves contained organic matter fixed under similar intercellular and ambient CO₂ concentration (C _i/C _a). High leaf C/N and negative δ¹⁵N values found in this habitat were consistent with the extreme soil N-deficiency. Analysis of Chl and carotenoids showed that expanding leaves had an incomplete development of photosynthetic antenna when compared to adult leaves. Dynamic inactivation of photosystem 2 (PS2) at midday was observed at both leaf ages as F_v/F_m decreased compared to predawn values. Adult leaves reached overnight F_v/F_m ratios typical of healthy leaves. Overnight recovery of F_v/F_m in expanding leaves was incomplete. F₀ remained unchanged from midday to predawn and F_v tended to increase from midday to predawn. The recovery from midday depression observed in adult leaves suggested an acclimatory down-regulation associated with photo-protection and non-damage of PS2.     

Needle δ13C and mobile carbohydrates in Pinus koraiensis in relation to decreased temperature and increased moisture along an elevational gradient in NE China

A tree’s crown interacts with atmospheric variables such as CO₂, temperature, and humidity. Physioecology of leaves/needles (e.g. δ¹³C, mobile carbohydrates, and nitrogen) is, therefore, strongly affected by microclimate in and surrounding a tree crown. To understand the physiological responses of leaves to changes in air temperature and moisture, we measured δ¹³C, soluble sugars, starch, and total nitrogen (N) concentrations in current year and 1-yr-old needles of Pinus koraiensis trees, and compared the growing season air temperature and relative humidity within and outside P. koraiensis crowns along an elevational gradient from 760 to 1,420 m a.s.l. on Changbai Mountain, NE China. Our results indicated that needle N and mobile carbohydrates concentrations, as well as needle δ¹³C values changed continuously with increasing elevation, corresponding to a continuous decrease in air temperature and an increase in relative humidity. Needle carbon and nitrogen status is highly significantly negatively correlated with temperature, but positively correlated with relative humidity. These results indicate that increases in air temperature in combination with decreases in relative humidity may result in lower levels of N and mobile carbohydrates in P. koraiensis trees, suggesting that future climate changes such as global warming and changes in precipitation patterns will directly influence the N and carbon physiology at P. koraiensis individual level, and indirectly affect the competitive ability, species composition, productivity and functioning at the stand and ecosystem level in NE China. Due to the relatively limited range of the transect (760–1,420 m) studied, further research is needed to explain whether the present results are applicable to scales across large elevational gradients.     

Spatial distribution of leaf water-use efficiency and carbon isotope discrimination within an isolated tree crown

The spatial variations in the stable carbon isotope composition (δ¹³C) of air and leaves (total matter and soluble sugars) were quantified within the crown of a well‐watered, 20‐year‐old walnut tree growing in a low‐density orchard. The observed leaf carbon isotope discrimination (Δ) was compared with that computed by a three‐dimensional model simulating the intracanopy distribution of irradiance, transpiration and photosynthesis (previously parameterized and tested for the same tree canopy) coupled to a biophysically based model of carbon isotope discrimination. The importance of discrimination associated with CO₂ gradients encountered from the substomatal sites to the carboxylation sites was evaluated. We also assessed by simulation the effect of current irradiance on leaf gas exchange and the effect of long‐term acclimation of photosynthetic capacity and stomatal and internal conductances to light regime on intracanopy gradients in Δ. The main conclusions of this study are: (i) leaf Δ can exhibit important variations (5 and 8‰ in total leaf material and soluble sugars, respectively) along light gradients within the foliage of an isolated tree; (ii) internal conductance must be taken into account to adequately predict leaf Δ, and (iii) the spatial variations in Δ and water‐use efficiency resulted from the short‐term response of leaf gas exchange to variations in local irradiance and, to a much lesser extent, from the long‐term acclimation of leaf characteristics to the local light regime.     

Responses of four succession tree species in low subtropics to enhanced UV-B radiation in the field

The tested tree species included pioneer species Acacia mangium, early succession stage species Schima superba, mesophyte intermediate-succession species Machilus chinensis, and shade-tolerant plant or late-succession species Cryptocarya concinna which occur in the lower subtropical forest community. A comparison with the current ambient level of UV-B radiation (UV-B) showed the leaf net photosynthetic rate (P _N), transpiration rate (E), and stomatal conductance (g _s) of the four species ranged from significantly decreased to no significant change. Additionally, the thickness of palisade and mesophyll in leaves of four tree species were decreased sharply by enhanced UV-B. The thickness of spongy parenchyma in leaves was also decreased except for M. chinensis. UV-B increased the leaf width of A. mangium but its leaf length, leaf thickness, and dry mass per unit area were not affected. Significantly increased stomata width was observed in A. mangium leaf epidermis in response to UV-B. Significantly decreased stomata width and significantly increased stomata density of leaf abaxial epidermis in M. chinensis were also observed. The stomata density of abaxial epidermis of C. concinna was remarkably increased by enhanced UV-B. The height and branch biomass of A. mangium and the height of S. superba were reduced visibly by enhanced UV-B. The four plant species could be classified into three groups of UV-B sensitiveness by hierarchical cluster analysis. A. mangium was sensitive to enhanced UV-B, while C. concinna showed more tolerance.     

Growth and stable isotope signals associated with drought-related mortality in saplings of two coexisting pine species

Drought-induced events of massive tree mortality appear to be increasing worldwide. Species-specific vulnerability to drought mortality may alter patterns of species diversity and affect future forest composition. We have explored the consequences of the extreme drought of 2005, which caused high sapling mortality (approx. 50 %) among 10-year-old saplings of two coexisting pine species in the Mediterranean mountains of Sierra Nevada (Spain): boreo-alpine Pinus sylvestris and Mediterranean P. nigra. Sapling height growth, leaf δ¹³C and δ¹⁸O, and foliar nitrogen concentration in the four most recent leaf cohorts were measured in dead and surviving saplings. The foliar isotopic composition of dead saplings (which reflects time-integrated leaf gas-exchange until mortality) displayed sharp increases in both δ¹³C and δ¹⁸O during the extreme drought of 2005, suggesting an important role of stomatal conductance (g_s) reduction and diffusional limitations to photosynthesis in mortality. While P. nigra showed decreased growth in 2005 compared to the previous wetter year, P. sylvestris maintained similar growth levels in both years. Decreased growth, coupled with a sharper increase in foliar δ¹⁸O during extreme drought in dead saplings, indicate a more conservative water use strategy for P. nigra. The different physiological behavior of the two pine species in response to drought (further supported by data from surviving saplings) may have influenced 2005 mortality rates, which contributed to 2.4-fold greater survival for P. nigra over the lifespan of the saplings. This species-specific vulnerability to extreme drought could lead to changes in dominance and distribution of pine species in Mediterranean mountain forests.