Contrasting leaf characteristics of trees and lianas in secondary and mature forests in southwestern China

We compared variation in sun-canopy leaf anatomy, morphology and photosynthetic rates of coexisting woody species (trees and lianas) in an 8-year-old secondary forest (SF) and mature forest (MF) in the wet season in Xishuangbanna, SW China. Variability of leaf traits of 66 species within growth-form groups in each forest was quantified using coefficients of variation (CV). For the mean values, the woody species in the SF had significantly higher leaf thickness and stomatal density, but lower nonmesophyll/mesophyll ratios than those in the MF. The average leaf area and leaf mass area (LMA) in the studied woody species did not change greatly during the successional process, but differed significantly between the growth forms, with trees having higher values than lianas. The light-saturated photosynthetic rate per unit leaf area (A _a) of the woody species in the SF ranged from 11.2 to 34.5 μmol m⁻² s⁻¹, similarly to pioneer tree species from literature data in southeast Asia. The A _a and photosynthetic nitrogen-use efficiency (PNUE) were significantly higher than those in the MF; whereas A _a in the MF ranged between 9 to 21 μmol m⁻² s⁻¹, with similar values between lianas and trees. For all woody species in both SF and MF, there were no significant differences in the average values of the CV of all measured variables for both lianas and trees. However, considerable variation in leaf anatomy, morphology, and photosynthetic rates within both growth forms and forests existed, as well as substantial variation in leaf size and stomatal density. We concluded that the tropical woody species formed a heterogeneous functional group in terms of leaf morphology and physiology in both secondary and mature forests.     

Short-term coherency between gross primary production and community respiration in an algal-dominated reef flat

Rates of net community carbon production (mmol C m⁻² h⁻¹) were measured continuously in an algal-dominated reef flat community on the Kaneohe Bay barrier reef, Hawaii, for 12 days at the end of October 2006. The weather became increasingly cloudy during the last 5 days of measurements, resulting in a sevenfold decline in daily incident light (28–4 Ein m⁻² d⁻¹). In response, gross primary production (P) for the reef flat community also decreased sevenfold, varying linearly with light (r ² = 0.92, n = 12). Community respiration (R) decreased fivefold over this same period and was highly correlated with changes in P (r ² = 0.84, n = 12). We reason that this short-term coherence between P and R indicates that most of the carbon fixed during this period was rapidly metabolized via plant respiration. We further conclude that the dominance of autotrophic respiration under general conditions of nutrient-limited growth can explain much of the balance between P and R that is commonly observed in shallow reef communities.     

Effect of light intensity on partitioning of photosynthetic electron transport to photorespiration in four subtropical forest plants

Photosynthetic rate (P_n) and the partitioning of noncyclic photosynthetic electron transport to photorespiration (J_O) in seedlings of four subtropical woody plants growing at three light intensities were studied in the summer time by measurements of chlorophyll fluorescence and CO₂ exchange. ExceptSchima superba, an upper canopy tree species, the tree speciesCastanopsis fissa and two understory shrubsPsychotria rubra, Ardisia quinquegona had the highestP _n at 36% of sunlight intensity. The total photosynthetic electron transport rate (J_F) and the ratio ofJ _O/J_F were elevated in leaves under full sunlight.J _O/J_F ratio reached 0.5–0.6 and coincided with the increasing of oxygenation rate of Rubisco (V_O), the activity of glycolate oxidase and photorespiration rate at full sunlight. It is suggested that an increasing partitioning proportion of photosynthetic electron transport to photorespiration might be one of the protective regulation mechanisms in forest plant under strong summer light and high temperature conditions.     

Influence of stand density on soil CO2 efflux for a Pinus densiflora forest in Korea

We investigated the influence of stand density [938 tree ha⁻¹ for high stand density (HD), 600 tree ha⁻¹ for medium stand density (MD), and 375 tree ha⁻¹ for low stand density (LD)] on soil CO₂ efflux (R _S) in a 70-year-old natural Pinus densiflora S. et Z. forest in central Korea. Concurrent with R _S measurements, we measured litterfall, total belowground carbon allocation (TBCA), leaf area index (LAI), soil temperature (ST), soil water content (SWC), and soil nitrogen (N) concentration over a 2-year period. The R _S (t C ha⁻¹ year⁻¹) and leaf litterfall (t C ha⁻¹ year⁻¹) values varied with stand density: 6.21 and 2.03 for HD, 7.45 and 2.37 for MD, and 6.96 and 2.23 for LD, respectively. In addition, R _S was correlated with ST (R ² = 0.77–0.80, P < 0.001) and SWC (R ² = 0.31–0.35, P < 0.001). It appeared that stand density influenced R _S via changes in leaf litterfall, LAI and SWC. Leaf litterfall (R ² = 0.71), TBCA (R ² = 0.64–0.87), and total soil N contents in 2007 (R ² = 0.94) explained a significant amount of the variance in R _S (P < 0.01). The current study showed that stand density is one of the key factors influencing R _S due to the changing biophysical and environmental factors in P. densiflora.     

Modeling nitrous oxide emission from rivers: a global assessment

Estimates of global riverine nitrous oxide (N₂O) emissions contain great uncertainty. We conducted a meta‐analysis incorporating 169 observations from published literature to estimate global riverine N₂O emission rates and emission factors. Riverine N₂O flux was significantly correlated with NH₄, NO₃ and DIN (NH₄ + NO₃) concentrations, loads and yields. The emission factors EF(a) (i.e., the ratio of N₂O emission rate and DIN load) and EF(b) (i.e., the ratio of N₂O and DIN concentrations) values were comparable and showed negative correlations with nitrogen concentration, load and yield and water discharge, but positive correlations with the dissolved organic carbon : DIN ratio. After individually evaluating 82 potential regression models based on EF(a) or EF(b) for global, temperate zone and subtropical zone datasets, a power function of DIN yield multiplied by watershed area was determined to provide the best fit between modeled and observed riverine N₂O emission rates (EF(a): R² = 0.92 for both global and climatic zone models, n = 70; EF(b): R² = 0.91 for global model and R² = 0.90 for climatic zone models, n = 70). Using recent estimates of DIN loads for 6400 rivers, models estimated global riverine N₂O emission rates of 29.6–35.3 (mean = 32.2) Gg N₂O–N yr⁻¹ and emission factors of 0.16–0.19% (mean = 0.17%). Global riverine N₂O emission rates are forecasted to increase by 35%, 25%, 18% and 3% in 2050 compared to the 2000s under the Millennium Ecosystem Assessment's Global Orchestration, Order from Strength, Technogarden, and Adapting Mosaic scenarios, respectively. Previous studies may overestimate global riverine N₂O emission rates (300–2100 Gg N₂O–N yr⁻¹) because they ignore declining emission factor values with increasing nitrogen levels and channel size, as well as neglect differences in emission factors corresponding to different nitrogen forms. Riverine N₂O emission estimates will be further enhanced through refining emission factor estimates, extending measurements longitudinally along entire river networks and improving estimates of global riverine nitrogen loads.     

Effects of root diameter and root nitrogen concentration on in situ root respiration among different seasons and tree species

Knowledge of root respiration is a prerequisite for a better understanding of ecosystem carbon budget and carbon allocation. However, there are not many relevant data in the literature on direct measurements of in situ root respiration by root chamber method. Furthermore, few studies have been focused on the effects of root diameter (D _r) and root nitrogen concentration (N _r) on in situ root respiration among different seasons and tree species. To address these goals, we used a simplified root-chamber system to measure in situ root respiration rates of Acacia crassicarpa and Eucalyptus urophylla in subtropical plantations of south China. We found that the species and season variation in root respiration were affected by D _r and N _r. Also, the root respiration per unit dry mass (R _r, nmol CO₂ g⁻¹ s⁻¹) and root respiration per unit N (R _n, nmol CO₂ g N⁻¹ s⁻¹) were affected by D _r and N _r. The R _r, R _n, N _r and soil temperature sensitivity (Q ₁₀) of R _r for the two species significantly decreased with an increase of D _r. The R _r of the two species showed significant an inter-seasonal and diurnal pattern, and this trend decreased with increasing D _r. Both the R _r and Q ₁₀ of the two species increased with increasing N _r. The D _r and N _r explained 54 and 52% of the observed variation in R _r for A. crassicarpa, and 65 and 70% for E. urophylla. The R _r, N _r, and Q ₁₀ of A. crassicarpa were significantly higher than those of E. urophylla. Our results indicated that root respiration was dependent on D _r and N _r, and this dependence varied with season and plant species.     

Numerical design of RN<Stack><Subscript><Emphasis Type="Italic">n</Emphasis></Subscript><Superscript>ν</Superscript></Stack> symmetry-based RF pulse schemes for recoupling and decoupling of nuclear spin interactions at high MAS frequencies

An approach for the efficient implementation of RN _n ^ν symmetry-based pulse schemes that are often employed for recoupling and decoupling of nuclear spin interactions in biological solid state NMR investigations is demonstrated at high magic-angle spinning frequencies. RF pulse sequences belonging to the RN _n ^ν symmetry involve the repeated application of the pulse sandwich {R _ϕ R _−ϕ}, corresponding to a propagator U _RF = exp(−i4ϕI _z), where ϕ = πν/N and R is typically a pulse that rotates the nuclear spins through 180° about the x-axis. In this study, broadband, phase-modulated 180° pulses of constant amplitude were employed as the initial ‘R’ element and the phase-modulation profile of this ‘R’ element was numerically optimised for generating RN _n ^ν symmetry-based pulse schemes with satisfactory magnetisation transfer characteristics. At representative MAS frequencies, RF pulse sequences were implemented for achieving ¹³C–¹³C double-quantum dipolar recoupling and through bond scalar coupling mediated chemical shift correlation and evaluated via numerical simulations and experimental measurements. The results from these investigations are presented here. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The resprouting response of co‐occurring temperate woody plant and grass species to elevated [CO2]: An insight into woody plant encroachment of grasslands

The phenomenon of woody plant thickening in grasslands has been observed globally and is likely to have widespread ecological consequences. It has been proposed that woody plant thickening is driven in part by rising atmospheric [CO₂] enhancing the resprouting ability of woody plants relative to grasses so they respond more strongly after disturbances such as herbivory and fire. The aim of this study was to examine the CO₂ effect on the resprouting ability of 16 co‐occurring temperate woody plant and grass species (eight species from each growth form). Plants were grown in a controlled glasshouse experiment under ambient (400 ppm) and elevated [CO₂] (600 ppm) for 14 weeks after which their resprouting ability was measured. Root non‐structural carbohydrate (NSC_mass) and nitrogen (N_mass) storage was used as proxies to measure the resprouting ability of woody plants while for the grasses it was measured directly. We found that both the woody plants (22% on average; P = 0.003) and grasses (20% on average; P = 0.003) produced more biomass under elevated [CO₂]. Despite the woody plants not allocating additional carbon to belowground storage under elevated [CO₂], they had significantly greater root NSC_mass (23% on average; P = 0.007) due to increased root biomass production (8% on average; P = 0.007). In contrast, root N_mass of the woody plants did not differ between CO₂ treatments (P = 0.373). Surprisingly, the resprouting ability of the grasses did not significantly differ between the CO₂ treatments (P = 0.067). These results provide evidence that the differing resprouting response of woody plants and grasses under elevated [CO₂] may be contributing to woody plant encroachment of grasslands worldwide.     

Relationship between net photosynthesis and leaf respiration in Mediterranean evergreen species

The relationship between net photosynthetic (P _N) and leaf respiration (R) rates of Quercus ilex, Phillyrea latifolia, Myrtus communis, Arbutus unedo, and Cistus incanus was monitored in the period February 2006 to February 2007. The species investigated had low R and P _N during winter, increasing from March to May, when mean air temperature reached 19.2 °C. During the favourable period, C. incanus and A. unedo had a higher mean P _N (16.4±2.4 μmol m⁻² s⁻¹) than P. latifolia, Q. ilex, and M. communis (10.0±1.3 μmol m⁻² s⁻¹). The highest R (1.89±0.30 μmol m⁻² s⁻¹, mean of the species), associated to a significant P _N decrease (62 % of the maximum, mean value of the species), was measured in July (mean R/P _N ratio 0.447±0.091). Q₁₀, indicating the respiration sensitivity to short-term temperature increase, was in the range 1.49 to 2.21. Global change might modify R/P _N determining differences in dry matter accumulation among the species, and Q. ilex and P. latifolia might be the most favoured species by their ability to maintain sufficiently higher P _N and lower R during stress periods.     

The population genetic structure of a common tropical damselfish on the Great Barrier Reef and eastern Papua New Guinea

Understanding patterns of connectivity in marine species is vital for the management and conservation of marine biodiversity. Here, the population genetic structure of a common and abundant tropical reef damselfish, Pomacentrus amboinensis, is reported. Using nine polymorphic microsatellite loci, the genetic structure at both small (i.e., around Lizard Island, Great Barrier Reef [GBR]) and large (GBR and Papua New Guinea [PNG]) spatial and temporal scales (2–1,600 km; 28 days– 6 years; n = 1,119) was analyzed. Temporal analyses found no evidence of genetic differentiation within or between Lizard Island recruitment pulses (R _ST = −0.001, P = 0.788), or corresponding established adult populations separated by 6 years of sampling (R _ST = 0.003, P = 0.116). The spatial analysis revealed that P. amboinensis populations are largely panmictic on the GBR and eastern PNG (R _ST = 0.001, P = 0.913), the only genetic discontinuity being between Kimbe Bay to the north of PNG and all populations south of PNG (R _ST = 0.077, P < 0.0001). Despite assumed high levels of self-recruitment based on previous tagging studies (15–60%), data presented here indicate that enough recruits are dispersing to impede the evolution of genetic structure over distances as great as 1,600 kms in this species. Data therefore indicate that the temporal genetic stability recorded here is maintained by high levels of gene flow.     

Photosynthesis,respiration, and carbon allocation of two cool-season perennial grasses in response to surface soil drying

The study was conducted to investigate carbon metabolic responses to surface soil drying for cool-season grasses. Kentucky bluegrass (Poa pratensis L.) and tall fescue (Festuca arundinaceae Schreb.) were grown in a greenhouse in split tubes consisting of two sections. Plants were subjected to three soil moisture regimes: (1) well-watered control; (2) drying of upper 20-cm soil (upper drying); and (3) drying of whole 40-cm soil profile (full drying). Upper drying for 30 d had no dramatic effects on leaf water potential (Ψ_leaf) and canopy photosynthetic rate (P_n) in either grass species compared to the well-watered control, but it reduced canopy respiration rate (R_canopy) and root respiration rate in the top 20 cm of soil (R_top). For both species in the lower 20 cm of wet soil, root respiration rates (R_bottom) were similar to the control levels, and carbon allocation to roots increased with the upper soil drying, particularly for tall fescue. The proportion of roots decreased in the 0-20 cm drying soil, but increased in the lower 20 cm wet soil for both grass species; the increase was greater for tall fescue. The Ψ_leaf, P_n, R_canopy, R_top, R_bottom, and carbon allocation to roots in both soil layers were all significantly higher for upper dried plants than for fully dried plants of both grass species. The reductions in R_canopy and R_top in surface drying soil and increases in root respiration and carbon allocation to roots in lower wet soil could help these grasses cope with surface-soil drought stress. This revised version was published online in June 2006 with corrections to the Cover Date.     

Leaf phosphorus influences the photosynthesis–nitrogen relation: a cross-biome analysis of 314 species

The ecophysiological linkage of leaf phosphorus (P) to photosynthetic capacity (A _max) and to the A _max–nitrogen relation remains poorly understood. To address this issue we compiled published and unpublished field data for mass-based A _max, nitrogen (N) and P (n = 517 observations) from 314 species at 42 sites in 14 countries. Data were from four biomes: arctic, cold temperate, subtropical (including Mediterranean), and tropical. We asked whether plants with low P levels have low A _max, a shallower slope of the A _max–N relationship, and whether these patterns have a geographic signature. On average, leaf P was substantially lower in the two warmer than in the two colder biomes, with the reverse true for N:P ratios. The evidence indicates that the response of A _max to leaf N is constrained by low leaf P. Using a full factorial model for all data, A _max was related to leaf N, but not to leaf P on its own, with a significant leaf N ×  leaf P interaction indicating that the response of A _max to N increased with increasing leaf P. This was also found in analyses using one value per species per site, or by comparing only angiosperms or only woody plants. Additionally, the slope of the A _max–N relationship was higher in the colder arctic and temperate than warmer tropical and subtropical biomes. Sorting data into low, medium, and high leaf P groupings also showed that the A _max–N slope increases with leaf P. These analyses support claims that in P-limited ecosystems the A _max–N relationship may be constrained by low P, and are consistent with laboratory studies that show P-deficient plants have limited ribulose-1,5-bisphosphate regeneration, a likely mechanism for the P influence upon the A _max–N relation. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Adaptability to elevated temperature and nitrogen addition is greater in a high-elevation population than in a low-elevation population of Hippophae rhamnoides

Hippophae rhamnoides L. is a broadleaf deciduous woody shrub occurring in southwest China, where it has been widely used in ecological restoration. In this study we investigated growth and physiological responses of 2-year-old healthy seedlings to elevated temperature, nitrogen (N) addition and their combination in two contrasting populations from high and low elevations. In closed-top chamber experiments, two populations were subjected to two temperature conditions (ambient temperature and temperature elevated by 2.2 ± 0.2°C) and two N levels (0 and 25 g N m⁻² a⁻¹). Compared with the control, increases in total leaf area (TLA), total chlorophyll content (TC), light-saturated photosynthetic rate (P _max), guaiacol peroxidase activity (POD), catalase activity (CAT) and carbon isotope composition (δ¹³C) were greater in the high-elevation population than in the low-elevation population under elevated temperature. On the other hand, decreases in root and shoot biomass ratio (RS), TC, P _max, light saturation point (L _SP), light compensation point (L _CP), superoxide dismutase (SOD), POD, CAT and δ¹³C were lower in the high-elevation population than in the low-elevation population under N addition. Moreover, the combination of elevated temperature and N addition decreased RS, P _max, apparent quantum efficiency (Φ), SOD, POD and δ¹³C significantly more in the low-elevation population than in the high-elevation population. These results demonstrated that there are different adaptive strategies among H. rhamnoides populations, the high-elevation population exhibiting higher adaptability to elevated temperature and N addition than the low-elevation population.     

Autotrophic and heterotrophic respiration in needle fir and Quercus-dominated stands in a cool-temperate forest, central Korea

To investigate annual variation in soil respiration (R _S) and its components [autotrophic (R _A) and heterotrophic (R _H)] in relation to seasonal changes in soil temperature (ST) and soil water content (SWC) in an Abies holophylla stand (stand A) and a Quercus-dominated stand (stand Q), we set up trenched plots and measured R _S, ST and SWC for 2 years. The mean annual rate of R _S was 436 mg CO₂ m⁻² h⁻¹, ranging from 76 to 1,170 mg CO₂ m⁻² h⁻¹, in stand A and 376 mg CO₂ m⁻² h⁻¹, ranging from 82 to 1,133 mg CO₂ m⁻² h⁻¹, in stand Q. A significant relationship between R _S and its components and ST was observed over the 2 years in both stands, whereas a significant correlation between R _A and SWC was detected only in stand Q. On average over the 2 years, R _A accounted for approximately 34% (range 17–67%) and 31% (15–82%) of the variation in R _S in stands A and Q, respectively. Our results suggested that vegetation type did not significantly affect the annual mean contributions of R _A or R _H, but did affect the pattern of seasonal change in the contribution of R _A to R _S.     

A mathematical and empirical model of the performance of a membrane-coupled anaerobic fermentor

A mathematical model was developed to describe the performance of a membrane-coupled anaerobic fermentor (MCAF)-based process. In our experimental results, higher volatile fatty acid (VFA) recovery ratios were obtained at greater filtration ratios. The VFA recovery ratio peaked at an HRT of 12 h and a membrane filtration ratio of 0.95 at a constant SRT. Based on our simulation, the HRT and filtration ratio should be maintained at less than 1 day and above 0.9, respectively, to exceed an organic materials recovery ratio of 35% at a constant SRT of 10 days. Our empirical model, which predicts the effluent VFA concentration (C _o ), described the performance of the MCAF adequately. The model demonstrated that the outlet VFA concentration was a function of three independent parameters—HLR, input organic concentration (C _i ), and membrane filtration ratio (ϕ). Multiple regression analyses were conducted using 50 measurements of the MCAF, yielding the following relationship: C _o  = 0.278ϕ^1.13 C _i ^1.93 HLR^0.11. The correlation coefficient (R ²) was 0.90. The simulation results were consistent with the observed data; therefore, due to its simplicity, this model predicts the effluent VFA concentration of an MCAF adequately.     

Photosynthesis and dark respiration of leaves of terrestrial carnivorous plants

Using CO₂ gasometry, net photosynthetic (P _N) and dark respiration rates (R _D) were measured in leaves or traps of 12 terrestrial carnivorous plant species usually grown in the shade. Generally, mean maximum P _N (60 nmol CO₂ g⁻¹(DM) s⁻¹ or 2.7 μmol m⁻² s⁻¹) was low in comparison with that of vascular non-carnivorous plants but was slightly higher than that reported elsewhere for carnivorous plants. After light saturation, the facultatively heliophytic plants behaved as shade-adapted plants. Mean R _D in leaves and traps of all species reached about 50% of maximum P _N and represents the high photosynthetic (metabolic) cost of carnivory.     

A proposal for universal formulas for estimating leaf water status of herbaceous and woody plants based on spectral reflectance properties

The present study deals with the relationships between water status parameters of plant leaves and reflectances (R_λ) at characteristic wavelengths, between 522 and 2450 nm, as well as reflectance ratios, R_λ/R₁₄₃₀, R_λ/R₁₆₅₀, R_λ/R₁₈₅₀, R_λ/R₁₉₂₀, and R_λ/R₁₉₅₀, based on the air-drying experimental results of soybean (Glycine max Merr.), maize (Zea mays L.), tuliptree (Liriodendron tulipifera L.) and viburnum (Viburnum awabuki K. Koch.) plants. The water status parameters include leaf water content per unit leaf area (LWC), specific leaf water content (SWC), leaf moisture percentage of fresh weight (LMP), relative leaf water content (RWC) and relative leaf moisture percentage on fresh weight basis (RMP). Effective spectral reflectances and reflectance ratios for estimating the LWC, SWC, LMP, RWC and RMP were identified. With these spectral indices, approaches to estimating LWC, RWC and RMP were discussed. Eventually, an attempt on universal formulas was made for estimating the leaf moisture conditions of both herbaceous and woody plants as mentioned above. Moreover, applicability of these formulas was checked with the field experimental results of soybean and maize grown under water and nutrient stresses. This revised version was published online in June 2006 with corrections to the Cover Date.     

Ecosystem respiration depends strongly on photosynthesis in a temperate heath

We measured net ecosystem CO₂ flux (F _n) and ecosystem respiration (R _E), and estimated gross ecosystem photosynthesis (P _g) by difference, for two years in a temperate heath ecosystem using a chamber method. The exchange rates of carbon were high and of similar magnitude as for productive forest ecosystems with a net ecosystem carbon gain during the second year of 293 ± 11 g C m⁻² year⁻¹ showing that the carbon sink strength of heather-dominated ecosystems may be considerable when C. vulgaris is in the building phase of its life cycle. The estimated gross ecosystem photosynthesis and ecosystem respiration from October to March was 22% and 30% of annual flux, respectively, suggesting that both cold-season carbon gain and loss were important in the annual carbon cycle of the ecosystem. Model fit of R _E of a classic, first-order exponential equation related to temperature (second year; R ² = 0.65) was improved when the P _g rate was incorporated into the model (second year; R ² = 0.79), suggesting that daytime R _E increased with increasing photosynthesis. Furthermore, the temperature sensitivity of R _E decreased from apparent Q ₁₀ values of 3.3 to 3.9 by the classic equation to a more realistic Q ₁₀ of 2.5 by the modified model. The model introduces R _photo, which describes the part of respiration being tightly coupled to the photosynthetic rate. It makes up 5% of the assimilated carbon dioxide flux at 0°C and 35% at 20°C implying a high sensitivity of respiration to photosynthesis during summer. The simple model provides an easily applied, non-intrusive tool for investigating seasonal trends in the relationship between ecosystem carbon sequestration and respiration.     

Is distribution of hydraulic constraints within tree crowns reflected in photosynthetic water-use efficiency? An example of <Emphasis Type="Italic">Betula pendula</Emphasis>

Spatial and daily variation in photosynthetic water-use efficiency was examined in leaves of Betula pendula Roth with respect to distribution of hydraulic conductance within the crown, morphological properties of stomata, and water availability. Intrinsic water-use efficiency (A _n/g _s) was determined from gas-exchange measurements performed both in situ in a natural forest stand and on detached shoots under laboratory conditions. In intact foliage, sun leaves demonstrated significantly higher (P < 0.001) A _n/g _s than shade leaves, as photosynthesis in the lower canopy was chronically limited by low light availability. However, this difference reversed in the mid-day period under sufficient irradiance (I > 800 μmol m⁻² s⁻¹): A _n/g _s averaged 28.8 and 24.0 μmol mol⁻¹ (P < 0.01) for shade and sun leaves, respectively. This last finding coincided with the data obtained in laboratory conditions: under equivalent leaf water supply and light, A _n/g _s in shade foliage was greater (P < 0.001) than in sun foliage across a wide range of irradiance. Thus, shade foliage of B. pendula is characterized by inherently higher A _n/g _s than sun foliage, associated with more conservative stomatal behavior, and lower soil-to-leaf (K _T) and leaf hydraulic conductances. Under unlimited light conditions, a within-crown trade-off between A _n/g _s and K _T becomes apparent. Differences in stomatal conductance between the detached shoots from sunlit and shaded canopy layers were largely attributable to the variation in stomatal morphology; significant relationships were established with characteristics combining stomatal size and density (relative stomatal surface, stomatal pore area index). Stomatal morphology is very likely involved in long-term adjustment of photosynthetic WUE.     

Relationships of photosynthetic capacity to PSII efficiency and to photochemical reflectance index of Pinus taiwanensis through different seasons at high and low elevations of sub-tropical Taiwan

We investigated the relationships of photosynthetic capacity (P _nsat, near light-saturated net photosynthetic rate measured at 1,200 μmol m⁻² s⁻¹ PPFD) to photosystem II efficiency (F _v/F _m) and to photochemical reflectance index [PRI = (R ₅₃₁ − R ₅₇₀)/(R ₅₃₁ + R ₅₇₀)] of Pinus taiwanensis Hay. needles at high (2,600 m a.s.l) and low-elevation (800 m a.s.l) sites through different seasons. Results indicate that at high-elevation site, P _nsat, F _v/F _m and PRI (both measured at predawn) paralleled in general with the air temperature. On the coolest measuring day with the minimum air temperature dropping to −2°C, P _nsat could decrease to ca. 15% of its highest value, which was measured in autumn. At low-elevation site, with the minimum air temperature of 10–12°C in cooler season and almost no seasonal variation of F _v/F _m, P _nsat dropped to ca. 65% of its highest value and PRI decreased ca. 0.02 in winter. Even though seasonal variation of P _nsat was affected by many factors, it was still closely related to PRI based on statistical analyses using data from both sites, through different seasons. On the contrary, seasonal variation of F _v/F _m of P. taiwanensis needles was influenced mainly by low temperature at high elevation. Therefore, the correlation of P _nsat − F _v/F _m was lower than that of P _nsat − PRI when data combined from both high- and low-elevation sites were analyzed. It is concluded that predawn PRI could be used as an indicator to estimate the seasonal potential of photosynthetic capacity of P. taiwanensis grown at low- and high-elevations of sub-tropical Taiwan.