Wood density is related to aboveground biomass and productivity along a successional gradient in upper Andean tropical forests.

Wood density (WD) is a key functional trait related to ecological strategies and ecosystem carbon dynamics. Despite its importance, there is a considerable lack of information on WD in tropical Andean forests, particularly regarding its relationship with forest succession and ecosystem carbon cycling. Here, we quantified WD in 86 upper Andean tree and shrub species in central Colombia, with the aim of determining how WD changes with forest succession and how it is related to productivity. We hypothesized that WD will increase with succession because early successional forests will be colonized by acquisitive species, which typically have low WD, while the shaded understory of older forests should favor higher WD. We measured WD in 481 individuals from 27 shrub and 59 tree species, and quantified aboveground biomass (AGB), canopy height, net primary production (NPP) and species composition and abundance in 14, 400-m2, permanent plots. Mean WD was 0.513 ± 0.114 (g/cm3), with a range between 0.068 and 0.718 (g/cm3). Shrubs had, on average, higher WD (0.552 ± 0.095 g/cm3) than trees (0.488 ± 0.104 g/cm3). Community weighted mean WD (CWMwd) decreased with succession (measured as mean canopy height, AGB, and basal area); CWMwd also decreased with aboveground NPP and stem growth. In contrast, the percentage of NPP attributed to litter and the percent of shrubs in plots increased with CWMwd. Thus, our hypothesis was not supported because early successional forests had higher CWMwd than late successional forests. This was related to a high proportion of shrubs (with high WD) early in succession, which could be a consequence of: 1) a low seed availability of trees due to intense land use in the landscape and/or 2) harsh abiotic conditions early in succession that filter out trees. Forest with high CWMwd had a high %NPP attributed to litter because they were dominated by shrubs, which gain little biomass in their trunks. Our findings highlight the links between WD, succession and carbon cycling (biomass and productivity) in this biodiversity hotspot. Thus, WD is an important trait that can be used to understand upper Andean forest recovery and improve forest restoration and management practices.

Diverging functional strategies but high sensitivity to an extreme drought in tropical dry forests.

Extreme drought events have negative effects on forest diversity and functioning. At the species level, however, these effects are still unclear, as species vary in their response to drought through specific functional trait combinations. We used long-term demographic records of 21,821 trees and extensive databases of traits to understand the responses of 338 tropical dry forests tree species to ENSO2015 , the driest event in decades in Northern South America. Functional differences between species were related to the hydraulic safety-efficiency trade-off, but unexpectedly, dominant species were characterised by high investment in leaf and wood tissues regardless of their leaf phenological habit. Despite broad functional trait combinations, tree mortality was more widespread in the functional space than tree growth, where less adapted species showed more negative net biomass balances. Our results suggest that if dry conditions increase in this ecosystem, ecological functionality and biomass gain would be reduced.

Effect of temporally heterogeneous light on photosynthetic light use efficiency, plant acclimation and growth in Abatia parviflora.

Individual leaves have a unique instantaneous photosynthetic photon flux density (PPFD) at which net photosynthetic light use efficiency (ϵL, the ratio between net photosynthesis and PPFD) is maximised (PPFDϵmax). When PPFD is above or below PPFDϵmax, efficiency declines. Thus, we hypothesised that heterogeneous PPFD conditions should increase the amount of time leaves photosynthesise at a PPFD different than PPFDϵmax and result in reduced growth. To date, this prediction has not been rigorously tested. Here, we exposed seedlings of Abatia parviflora Ruiz & Pav to light regimes of equal total daily irradiance but with three different daily time courses of PPFD: constant PPFD (No_H), low heterogeneity (Low_H) and high heterogeneity (High_H). Mean ϵL, leaf daily photosynthesis and plant growth were all significantly higher in No_H and Low_H plants than in High_H plants, supporting our hypothesis. In addition, mean ϵL was positively related to final plant biomass. Unexpectedly, High_H plants had more etiolated stems and more horizontal leaves than No_H and Low_H plants, possibly due to exposure to low PPFD in the morning and afternoon. In conclusion, PPFD heterogeneity had an important effect on average ϵL, photosynthesis and growth, but also on allocation and plant morphology.

Norway maple displays greater seasonal growth and phenotypic plasticity to light than native sugar maple.

Norway maple (Acer platanoides L), which is among the most invasive tree species in forests of eastern North America, is associated with reduced regeneration of the related native species, sugar maple (Acer saccharum Marsh) and other native flora. To identify traits conferring an advantage to Norway maple, we grew both species through an entire growing season under simulated light regimes mimicking a closed forest understorey vs. a canopy disturbance (gap). Dynamic shade-houses providing a succession of high-intensity direct-light events between longer periods of low, diffuse light were used to simulate the light regimes. We assessed seedling height growth three times in the season, as well as stem diameter, maximum photosynthetic capacity, biomass allocation above- and below-ground, seasonal phenology and phenotypic plasticity. Given the north European provenance of Norway maple, we also investigated the possibility that its growth in North America might be increased by delayed fall senescence. We found that Norway maple had significantly greater photosynthetic capacity in both light regimes and grew larger in stem diameter than sugar maple. The differences in below- and above-ground biomass, stem diameter, height and maximum photosynthesis were especially important in the simulated gap where Norway maple continued extension growth during the late fall. In the gap regime sugar maple had a significantly higher root : shoot ratio that could confer an advantage in the deepest shade of closed understorey and under water stress or browsing pressure. Norway maple is especially invasive following canopy disturbance where the opposite (low root : shoot ratio) could confer a competitive advantage. Considering the effects of global change in extending the potential growing season, we anticipate that the invasiveness of Norway maple will increase in the future.

Contributions of leaf photosynthetic capacity, leaf angle and self-shading to the maximization of net photosynthesis in Acer saccharum: a modelling assessment.

BACKGROUND AND AIMS: Plants are expected to maximize their net photosynthetic gains and efficiently use available resources, but the fundamental principles governing trade-offs in suites of traits related to resource-use optimization remain uncertain. This study investigated whether Acer saccharum (sugar maple) saplings could maximize their net photosynthetic gains through a combination of crown structure and foliar characteristics that let all leaves maximize their photosynthetic light-use efficiency (ε). METHODS: A functional-structural model, LIGNUM, was used to simulate individuals of different leaf area index (LAI(ind)) together with a genetic algorithm to find distributions of leaf angle (L(A)) and leaf photosynthetic capacity (A(max)) that maximized net carbon gain at the whole-plant level. Saplings grown in either the open or in a forest gap were simulated with A(max) either unconstrained or constrained to an upper value consistent with reported values for A(max) in A. saccharum. KEY RESULTS: It was found that total net photosynthetic gain was highest when whole-plant PPFD absorption and leaf ε were simultaneously maximized. Maximization of ε required simultaneous adjustments in L(A) and A(max) along gradients of PPFD in the plants. When A(max) was constrained to a maximum, plants growing in the open maximized their PPFD absorption but not ε because PPFD incident on leaves was higher than the PPFD at which ε(max) was attainable. Average leaf ε in constrained plants nonetheless improved with increasing LAI(ind) because of an increase in self-shading. CONCLUSIONS: It is concluded that there are selective pressures for plants to simultaneously maximize both PPFD absorption at the scale of the whole individual and ε at the scale of leaves, which requires a highly integrated response between L(A), A(max) and LAI(ind). The results also suggest that to maximize ε plants have evolved mechanisms that co-ordinate the L(A) and A(max) of individual leaves with PPFD availability.

Relationships among precipitation regime, nutrient availability, and carbon turnover in tropical rain forests.

The effect of high precipitation regime in tropical forests is poorly known despite indications of its potentially negative effects on nutrient availability and carbon (C) cycling. Our goal was to determine if there was an effect of high rainfall on nitrogen (N) and phosphorous (P) availability and indexes of C cycling in lowland tropical rain forests exposed to a broad range of mean annual precipitation (MAP). We predicted that C turnover time would increase with MAP while the availability of N and P would decrease. We studied seven Neotropical lowland forests covering a MAP range between 2,700 and 9,500 mm. We used radiocarbon (∆(14)C) from the atmosphere and respired from soil organic matter to estimate residence time of C in plants and soils. We also used C, N, and P concentrations and the stable isotope ratio of N (δ(15)N) in live and dead plant tissues and in soils as proxies for nutrient availability. Negative δ(15)N values indicated that the wettest forests had N cycles that did not exhibit isotope-fractionating losses and were potentially N-limited. Element ratios (N:P and C:P) in senescent leaves, litter, and live roots showed that P resorption increased considerably with MAP, which points towards increasing P-limitation under high MAP regimes. Soil C content increased with MAP but C turnover time only showed a weak relationship with MAP, probably due to variations in soil parent material and age along the MAP gradient. In contrast, comparing C turnover directly to nutrient availability showed strong relationships between C turnover time, N availability (δ(15)N), and P availability (N:P) in senescent leaves and litter. Thus, an effect of MAP on carbon cycling appeared to be indirectly mediated by nutrient availability. Our results suggest that soil nutrient availability plays a central role in the dynamic of C cycling in tropical rain forests.

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 (phi) and curvature (theta)]. 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.