Optimality of nitrogen distribution among leaves in plant canopies

The vertical gradient of the leaf nitrogen content in a plant canopy is one of the determinants of vegetation productivity. The ecological significance of the nitrogen distribution in plant canopies has been discussed in relation to its optimality; nitrogen distribution in actual plant canopies is close to but always less steep than the optimal distribution that maximizes canopy photosynthesis. In this paper, I review the optimality of nitrogen distribution within canopies focusing on recent advancements. Although the optimal nitrogen distribution has been believed to be proportional to the light gradient in the canopy, this rule holds only when diffuse light is considered; the optimal distribution is steeper when the direct light is considered. A recent meta-analysis has shown that the nitrogen gradient is similar between herbaceous and tree canopies when it is expressed as the function of the light gradient. Various hypotheses have been proposed to explain why nitrogen distribution is suboptimal. However, hypotheses explain patterns observed in some specific stands but not in others; there seems to be no general hypothesis that can explain the nitrogen distributions under different conditions. Therefore, how the nitrogen distribution in canopies is determined remains open for future studies; its understanding should contribute to the correct prediction and improvement of plant productivity under changing environments.     

Is shadier better? The effect of agroforestry management on small mammal diversity

Agroforestry systems play fundamental roles for wildlife conservation, but are prone to disturbances from management practices aiming at increasing local productivity. This work investigates the small mammal assemblages present in cacao agroforests, which differ in shade tree density. We tested the prediction that higher tree density increases shade level, analyzed how some environmental variables important for small mammals (vegetation complexity, tree basal area, and invertebrate biomass) vary across the shade level gradient, and how the assemblages respond to these variations. We also tested the effect of the environmental variables on the abundance of the three most common species: Rhipidomys mastacalis, Hylaeamys seuanezi, and Marmosa murina. We captured 651 individuals belonging to 18 species. A positive relationship was observed between an abundance of non-forest specialists and tree basal area, while species diversity within this group showed positive association with vegetation complexity. Assemblage structure (described by a matrix of species abundance per site) was not affected by our environmental variables, but R. mastacalis was more abundant in sites with lower vegetation complexity. Higher shade levels in cacao agroforest tended to occur in sites with greater tree basal area, which was not a good predictor of small mammal diversity. This suggests that environmental management to reduce shade with the purpose of increasing cacao productivity is not necessarily negative for small mammal conservation. Species diversity was favored by structurally complex systems, a possible response to greater niche diversity.     

Phytochrome control of the growth and development of Rumex obtusifolius under simulated canopy light environments*

Abstract Studies on the growth and development of Rumex obtusifolius seedlings under simulated shade conditions show that both light quantity and quality contribute to the observed responses. In the shade situation the plants have lower dry matter, lower leaf area and lower net assimilation rates. Petiole elongation occurs under shade conditions only after transfer of the plants from non-shade environments. The effects of light quality are related to phytochrome photoequilibria set up by the increased relative photon flux in the far-red found under vegetation canopies.     

Cooling streams with riparian trees: Thermal regime depends on total solar radiation penetrating the canopy

Riparian planting is often recommended for stream restoration, notably to mitigate solar heating of stream waters. However, plant leaves shade photosynthetically active radiation (PAR, 400–700 nm) more efficiently than the near-infrared (NIR, 700–3000 nm) wavelengths that comprise about half of incident solar radiation and so contribute equally to water heating. Total solar radiation (NIR + PAR) exposure at the reach scale is needed to avoid bias when predicting the thermal response of streams. In this article, we alert stream ecologists and riparian restorers to the water heating contribution of NIR and provide a means to account for both NIR and PAR in total solar radiation penetrating riparian canopies. We used spectral pyranometers to simultaneously measure total solar radiation and its NIR component under different tree canopies over a wide range of shade conditions as indexed by PAR exposure. Measurements were made during full overcast (diffuse lighting) conditions so as to ‘sample’ transmission of radiation through the complete canopy. The NIR proportion of sub-canopy solar radiation increased steadily with increasing shade and was appreciably greater under willow than pine canopies, while NZ native broadleaf rainforest had intermediate (rather variable) NIR content. Our trend lines for different tree canopies permit total sub-canopy solar radiation to be estimated from PAR exposure for unbiased modelling of stream thermal regimes.     

An Analysis of Growth of Oil Palm Seedlings in Full Daylight and in Shade

Growth of seedling oil palms (Elaeis guineensis), in full daylightand under three levels of shade, was studied using growth analysistechniques. In full day-light, net assimilation rates (E_A) betweeno.15 and 0.32 g./dm.²/week were obtained associated with lowleaf-area ratios (F) giving relative growth-rates (R_W) rangingfromI I.8 to 3.2 per cent, per day. There were no indicationsof seasonal differences within the small range of values found. The plants take about 90 days to adapt to shade conditions becausethe mean plastochron is 24 days, and shading effects are beststudied on plants grown since germination under the shade treatments.Very different pictures of response to shade were obtained usingplants grown initially in full light followed by 90 days' shadebefore sampling compared with plants grown under shade sincegermination. In the latter, except at the lowest light levelused, I I.I per cent, of full day-light, there was very littleeffect of light on E_A or R_w, although the F values decreasedas light increased. Extrapolation of the F values to the extinctionpoint gave values similar to those obtained in another experimenton the effect of a number of shade levels on F. The physiological and ecological implications of these findings,particularly the low growth-rates and shade tolerance, are discussed.     

Uncoupling the effects of shade and food resources of vegetation on Mediterranean ants: an experimental approach at the community level

Vegetation is one of the main factors affecting the composition and structure of ant communities. The effect of vegetation is both by offering food resources and by modifying the proportion of ground cover exposed to shade conditions. However, it is difficult in field studies to uncouple the effects of food resources and shade on animal communities. The goal of this study was to analyze experimentally the effects of vegetation through food and shade supply on the structure and composition of a Mediterranean ant community. We have crossed these two factors in a factorial design where we have eliminated vegetation in experimental treatments and we have simulated the effects of vegetation by manipulating separately the supply of food resources and shade. The expected decrease in ant abundance and richness in plots without food resources but with shade was only partially confirmed by the results, because there was a decrease in the number of ant species but not a significant decrease in overall ant abundance in plots without food resources. We did not confirm the second hypothesis that the decrease in shade while maintaining food resources resulted in an increase of heat-tolerant, subordinate species and, consequently, ant richness. Species composition differed in the different experimental treatments. Stress-tolerant species were abundant in areas without shade and without food resources. Heat-tolerant species were mainly present in plots without shade but with food resources. Finally, species with high requirements of shade and/or food resources were associated to control plots with both shade and food resources from vegetation. Overall, this study provides an experiment that uncouples experimentally shade and food resources and dissects their effects on the whole ant community, and shows that the effects of the two factors are independent and affect different components of ant community structure and composition.     

The nitrogen budget of a pine forest under free air CO2 enrichment

Elevated concentrations of atmospheric CO₂ increase plant biomass, net primary production (NPP) and plant demand for nitrogen (N). The demand for N set by rapid plant growth under elevated CO₂ could be met by increasing soil N availability or by greater efficiency of N uptake. Alternatively, plants could increase their nitrogen-use efficiency (NUE), thereby maintaining high rates of growth and NPP in the face of nutrient limitation. We quantified dry matter and N budgets for a young pine forest exposed to 4 years of elevated CO₂ using free-air CO₂ enrichment technology. We addressed three questions: Does elevated CO₂ increase forest NPP and the demand for N by vegetation? Is demand for N met by greater uptake from soils, a shift in the distribution of N between plants, microbes, and soils, or increases in NUE under elevated CO₂? Will soil N availability constrain the NPP response of this forest as CO₂ fumigation continues? A step-function increase in atmospheric CO₂ significantly increased NPP during the first 4 years of this study. Significant increases in NUE under elevated CO₂ modulated the average annual requirement for N by vegetation in the first and third growing seasons under elevated CO₂; the average stimulation of NPP in these years was 21% whereas the average annual stimulation of the N requirement was only 6%. In the second and fourth growing seasons, increases in NPP increased the annual requirement for N by 27-33%. Increases in the annual requirement for N were largely met by increases in N uptake from soils. Retranslocation of nutrients prior to senescence played only a minor role in supplying the additional N required by trees growing under elevated CO₂. NPP was highly correlated with between-plot variation in the annual rate of net N mineralization and CO₂ treatment. This demonstrates that NPP is co-limited by C availability, as CO₂ from the atmosphere, and N availability from soils. There is no evidence that soil N mineralization rates have increased under elevated CO₂. The correlation between NPP and N mineralization rates and the increase in the annual requirement for N in certain years imply that soil N availability may control the long-term productivity response of this ecosystem to elevated CO₂. Although we have no evidence suggesting that NPP is declining in response to >4 years of CO₂ fumigation, if the annual requirement of N continues to be stimulated by elevated CO₂, we predict that the productivity response of this forest ecosystem will decline over time.     

水稻冠层结构变化对二向反射率的影响

通过对植被二向反射特性的研究,可反演植被冠层结构信息,如叶面积指数、平均叶角、株高、覆盖率等。在大田晚稻（秀水63）移栽后第26、35、41、49、62、67和86d,对水稻冠层结构及二向反射率进行实测,并分析了二向反射率随冠层结构变化规律,结果表明,植被冠层二向反射率对入射角与观测角的敏感性随着植被冠层结构的变化而变化,蕴涵着丰富的植被结构信息。     

Effects of kelp canopies on bleaching and photosynthetic activity of encrusting coralline algae

Canopy-forming algae often coexist with an understorey of encrusting coralline algae that bleach following the loss of canopies. We tested the hypothesis that canopy loss causes a reduction in photosynthetic activity of encrusting coralline algae concomitant with their bleaching. When canopies were experimentally removed, corallines bleached and their photosynthetic activity was rapidly reduced to half their activity observed under canopies. This result prompted us to test, and subsequently accept, the hypothesis that exposure of understorey corallines to enhanced light intensity per se (simulation of canopy loss) acts as a mechanism that causes bleaching and reduced photosynthetic activity. Despite bleaching, encrusting corallines maintained reduced levels of photosynthetic activity, and this may explain why, under certain conditions, bleached corallines can persist in the absence of canopy-forming algae. Nevertheless, our data provide evidence that the positive association between canopy-forming algae and encrusting coralline algae is maintained because of shade provided by the canopy.     

Contrasting shade avoidance responses in two perennial grasses: a field investigation in simulated sparse and dense canopies

We designed an experiment with potted plants grown outdoors to investigate the expression of shade avoidance in simulated sparse and dense canopies by two perennial grasses known to express contrasting responses to low red:far-red ratios (R:FR). Plants were grown in canopy microenvironments designed to lower the R:FR by reflection of horizontally propagated FR from neighbors and by direct attenuation of R by filters located above plants. Two specific hypotheses were tested: (1) Paspalum dilatatum will express greater shade avoidance than Schizachyrium scoparium to low R:FR in both sparse and dense canopies, and (2) low R:FR will produce greater expressions of shade avoidance in sparse than in dense canopies in both species. P. dilatatum was more responsive to low R:FR than S. scoparium in both the sparse and dense canopies and lower ramet number plant^–1 was the only common shade avoidance response between species in sparse canopies. P. dilatatum also showed significant reductions in juvenile ramet initiation, juvenile ramet mass, total shoot mass, and shoot:root ratios in sparse canopies, but only juvenile ramet initiation was reduced in dense canopies. The suppression of juvenile ramet initiation in the dense canopy was at least partially modulated by the vertically propagated R:FR because a similar reduction in PFD and horizontally propagated R:FR showed 42% greater juvenile ramet initiation in the respective control. S. scoparium only showed a significant reduction in ramet number plant^–1 and a significant increase in blade length in sparse canopies, but no significant responses occurred in dense canopies. Consequently, neither hypothesis was rejected. Variable shade avoidance responses between species and canopy densities indicate that both interspecific variation and various proportions of vertically and horizontally propagated low R:FR can influence the expression of shade avoidance responses of perennial grasses in field settings.     

Limitations on photosynthesis of competing individuals in stands and the consequences for canopy structure

The canopy structure of a stand of vegetation is determined by the growth patterns of the individual plants within the stand and the competitive interactions among them. We analyzed the carbon gain of individuals in two dense monospecific stands of Xanthium canadense and evaluated the consequences for intra-specific competition and whole-stand canopy structure. The stands differed in productivity, and this was associated with differences in nitrogen availability. Canopy structure, aboveground mass, and nitrogen contents per unit leaf area (N_area) were determined for individuals, and leaf photosynthesis was measured as a function of N_area. These data were used to calculate the daily carbon gain of individuals. Within stands, photosynthesis per unit aboveground mass (P_mass) of individual plants increased with plant height, despite the lower leaf area ratios of taller plants. The differences in P_mass between the tallest most dominant and shortest most subordinate plants were greater in the high-nitrogen than in the low-nitrogen stand. This indicated that competition was asymmetric and that this asymmetry increased with nitrogen availability. In the high-nitrogen stand, taller plants had a higher P_mass than shorter ones, because they captured more light per unit mass and because they had higher photosynthesis per unit of absorbed light. Conversely, in the low-nitrogen stand, the differences in P_mass between plants of different heights resulted only from differences in their light capture per unit mass. Sensitivity analyses revealed that an increase in N_area, keeping leaf area of plants constant, increased whole-plant carbon gain for the taller more dominant plants but reduced carbon gain in the shorter more subordinate ones, which implies that the N_area values of shorter plants were greater than the optimal values for maximum photosynthesis. On the other hand, the carbon gain of all individual plants, keeping their total canopy N constant, was positively related to an increase in their individual leaf area. At the same time, however, increasing the leaf area for all plants simultaneously reduced the carbon gain of the whole stand. This result shows that the optimal leaf area index (LAI), which maximizes photosynthesis of a stand, is not evolutionarily stable because at this LAI, any individual can increase its carbon gain by increasing its leaf area.     

Changes in the composition of rainwater upon passage through the canopies of trees and of ground vegetation in a Dutch oak-birch forest

The composition of canopy throughfall water in an oak-birch woodland, heavily affected by atmospheric deposition of N and S, changed markedly upon contact with the above-ground parts of the ground vegetation, which consisted mainly of bracken. The fluxes of nitrate and H⁺ decreased, simultaneously with an increase in the flux of bicarbonate, indicative of above-ground uptake of nitrate by the ground vegetation. This above-ground assimilation takes place in spite of abundant availability of inorganic nitrogen in the root zone of the ground vegetation. Fluxes of phosphate were somewhat lower, and those of ammonium somewhat higher in throughfall of the ground vegetation than in that of the tree layer. Although those differences were not statistically significant, they do suggest assimilation of some P and extra net dry deposition of atmospheric ammonia below the tree canopies.     

Optimal nitrogen distribution within a leaf canopy under direct and diffuse light

Nitrogen distribution within a leaf canopy is an important determinant of canopy carbon gain. Previous theoretical studies have predicted that canopy photosynthesis is maximized when the amount of photosynthetic nitrogen is proportionally allocated to the absorbed light. However, most of such studies used a simple Beer's law for light extinction to calculate optimal distribution, and it is not known whether this holds true when direct and diffuse light are considered together. Here, using an analytical solution and model simulations, optimal nitrogen distribution is shown to be very different between models using Beer's law and direct–diffuse light. The presented results demonstrate that optimal nitrogen distribution under direct–diffuse light is steeper than that under diffuse light only. The whole‐canopy carbon gain is considerably increased by optimizing nitrogen distribution compared with that in actual canopies in which nitrogen distribution is not optimized. This suggests that optimization of nitrogen distribution can be an effective target trait for improving plant productivity.     

Response to CO2 enrichment of understory vegetation in the shade of forests

Responses of forest ecosystems to increased atmospheric CO₂ concentration have been studied in few free‐air CO₂ enrichment (FACE) experiments during last two decades. Most studies focused principally on the overstory trees with little attention given to understory vegetation. Despite its small contribution to total productivity of an ecosystem, understory vegetation plays an important role in predicting successional dynamics and future plant community composition. Thus, the response of understory vegetation in Pinus taeda plantation at the Duke Forest FACE site after 15–17 years of exposure to elevated CO₂, 6–13 of which with nitrogen (N) amendment, was examined. Aboveground biomass and density of the understory decreased across all treatments with increasing overstory leaf area index (LAI). However, the CO₂ and N treatments had no effect on aboveground biomass, tree density, community composition, and the fraction of shade‐tolerant species. The increases of overstory LAI (~28%) under elevated CO₂ resulted in a reduction of light available to the understory (~18%) sufficient to nullify the expected growth‐enhancing effect of elevated CO₂ on understory vegetation.     

Does foliage on the same branch compete for the same water? Experiments on Douglas-fir trees

Do branchlets within a branch have autonomous water supplies, or do they share a common water supply system? We hypothesized that if branchlets shared a common water supply, then stomatal conductance (g_s) on sunlit foliage would increase with reduced transpiration of competing foliage on the branch. We reduced transpiration of other foliage on the branch through bagging and shading, and we monitored the gas-exchange responses of the remaining sunlit foliage on the branch relative to control branches for several age classes of Douglas-fir trees (aged ~10 years, 20 years, and 450 years old). Contrary to our hypothesis, we found no increases in g_s in either young or old trees following transient reductions in the amount of transpiring leaf area. The diurnal change in water potential, mid-day stomatal closure and associated photosynthetic decline occurred at the same time and were of the same magnitude on both treated and untreated branches, with the exception of photosynthesis in one 450-year-old tree. Hydraulic conductance measurements of branch junctions indicate that xylem within branches is only partially interconnected which would reduce the effectiveness of shading as a means of increasing water supply to the remaining sunlit foliage. The lack of a response implies that when a branch is in partial shade, the remaining sunlit foliage has no advantage with respect to water status over foliage on a branch completely in the sun.     

Drivers of native species regeneration in the process of restoring natural forests from mono‐specific,even‐aged tree plantations: a quantitative review

A substantial proportion of the existing tree plantations has been established following clearing of native forests. This form of conversion has become widely unaccepted, and there are increasing demands to reverse it through ecological restoration. Yet, there is a lack of integrated knowledge on how best to restore. Here, we reviewed 68 studies to identify the main factors determining establishment success of regeneration of native woody species when restoring natural forests from plantation forests using active and passive approaches, beneath existing canopies, and following their removal. According to the evidence collected, herbivory, within‐gap position, soil properties, and ground cover type and structure had limited influence on regeneration, showing significant effects in less than 26% of cases in which their influence was tested. In contrast, spatial landscape configuration, overstorey structure, ground vegetation structure, overstorey composition, and climate and geomorphology had significant effects in 67, 47, 47, 52, and 63% of cases, respectively. Regeneration diversity and abundance increased with proximity to natural vegetation remnants and seed sources. Lower canopy and understorey stocking levels positively influenced regeneration, as did interventions to reduce them. Canopy cover reduction proved especially effective in warmer regions, in stands of broadleaved species, younger ages (<30 years), higher densities (>1,000 trees/ha), and taller canopies (>20 m). Restoration of native forests can be optimized by adopting interventions that prove most effective, and prioritizing more responsive stand types. However, the specific stand attributes and environmental factors described should be further studied to understand the mechanisms underlying their influence on regeneration.     

Field evidences for the positive effects of aerosols on tree growth

Theoretical and eddy covariance studies demonstrate that aerosol‐loading stimulates canopy photosynthesis, but field evidence for the aerosol effect on tree growth is limited. Here, we measured in situ daily stem growth rates of aspen trees under a wide range of aerosol‐loading in China. The results showed that daily stem growth rates were positively correlated with aerosol‐loading, even at exceptionally high aerosol levels. Using structural equation modeling analysis, we showed that variations in stem growth rates can be largely attributed to two environmental variables covarying with aerosol loading: diffuse fraction of radiation and vapor pressure deficit (VPD). Furthermore, we found that these two factors influence stem growth by influencing photosynthesis from different parts of canopy. Using field observations and a mechanistic photosynthesis model, we demonstrate that photosynthetic rates of both sun and shade leaves increased under high aerosol‐loading conditions but for different reasons. For sun leaves, the photosynthetic increase was primarily attributed to the concurrent lower VPD; for shade leaves, the positive aerosol effect was tightly connected with increased diffuse light. Overall, our study provides the first field evidence of increased tree growth under high aerosol loading. We highlight the importance of understanding biophysical mechanisms of aerosol‐meteorology interactions, and incorporating the different pathways of aerosol effects into earth system models to improve the prediction of large‐scale aerosol impacts, and the associated vegetation‐mediated climate feedbacks.     

The impact of diffuse sunlight on canopy light-use efficiency, gross photosynthetic product and net ecosystem exchange in three forest biomes

For three forest canopies (a sparse, boreal needleleaf; a temperate broadleaf; and a dense, tropical, broadleaf stand) light‐use efficiency (LUE) is found to be 6–33% higher when sky radiance is dominated by diffuse rather than direct sunlight. This enhancement is much less than that reported previously for both crops (110%; Choudbury, 2001 ) and moderately dense temperate woodland (50–180%). We use the land‐surface scheme JULES to interpret the observed canopy response. Once sunflecks and leaf orientation are incorporated explicitly into the scheme, our simulations reproduce convincingly the overall level of canopy gross photosynthetic product (GPP), its enhancement with respect to diffuse sunlight and the mean 15% reduction in productivity observed during the afternoon due to stomatal closure. The LUE enhancement under diffuse sunlight can be explained by sharing of the canopy radiation‐load, which is reduced under direct sky radiance. Once sunflecks are accounted for the advantage of implementing more sophisticated calculations of stomatal conductance (e.g. Ball–Berry and SPA submodels) is less obvious even for afternoon assimilation. Empirical relations are developed between observed carbon flux and the environmental variables total downwelling shortwave radiation (SW), canopy temperature (T) and the fraction of diffuse sky radiance (fDIF). These relations allow us to gauge the impact of increased/reduced insolation on GPP and net ecosystem exchange (NEE). Overall the three stands appear to be fairly stable within global trends and typical interannual variability (SW changing by <15%). Greatest sensitivity is exhibited by the boreal site, Zotino, where NEE falls by 9±4% for a 15% reduction in SW.     

A simulation of the importance of length of growing season and canopy functional properties on the seasonal gross primary production of temperate alpine meadows

Background and Aims: Along snowmelt gradients, the canopies of temperate alpine meadowsdiffer strongly in their structural and biochemical properties.Here, a study is made of the effects of these canopy dissimilaritiescombined with the snow-induced changes in length of growingseason on seasonal gross primary production (GPP). Methods: Leaf area index (LAI) and community-aggregated values of leafangle and leaf nitrogen content were estimated for seven alpineplant canopies distributed along a marked snowmelt gradient,and these were used as input variables in a sun–shadecanopy bulk-photosynthesis model. The model was validated forplant communities of early and late snowmelt sites by measuringthe instantaneous CO₂ fluxes with a canopy closed-chamber technique.A sensitivity analysis was conducted to estimate the relativeimpact of canopy properties and environmental factors on thedaily and seasonal GPP. Key Results: Carbon uptake was primarily related to the LAI and total canopynitrogen content, but not to the leaf angle. For a given levelof photosynthetically active radiation, CO₂ assimilation washigher under overcast conditions. Sensitivity analysis revealedthat increase of the length of the growing season had a highereffect on the seasonal GPP than a similar increase of any otherfactor. It was also found that the observed greater nitrogencontent and larger LAI of canopies in late-snowmelt sites largelycompensated for the negative impact of the reduced growing season. Conclusions: The results emphasize the primary importance of snow-inducedchanges in length of growing season on carbon uptake in alpinetemperate meadows. It was also demonstrated how using leaf-traitvalues of the dominants is a useful approach for modelling ecosystemcarbon-cycle-related processes, particularly when continuousmeasurements of CO₂ fluxes are technically difficult. The studythus represents an important step in addressing the challengeof using a plant functional-trait approach for biogeochemicalmodelling.