Abstract: Stay-green sorghum plants exhibit greener leaves and stems during the grain-filling period under water-limited conditions compared with their senescent counterparts, resulting in increased grain yield, grain mass, and lodging resistance. Stay-green has been mapped to a number of key chromosomal regions, including Stg1, Stg2, Stg3, and Stg4, but the functions of these individual quantitative trait loci (QTLs) remain unclear. The objective of this study was to show how positive effects of Stg QTLs on grain yield under drought can be explained as emergent consequences of their effects on temporal and spatial water-use patterns that result from changes in leaf-area dynamics. A set of four Stg near-isogenic lines (NILs) and their recurrent parent were grown in a range of field and semicontrolled experiments in southeast Queensland, Australia. These studies showed that the four Stg QTLs regulate canopy size by: (1) reducing tillering via increased size of lower leaves, (2) constraining the size of the upper leaves; and (3) in some cases, decreasing the number of leaves per culm. In addition, they variously affect leaf anatomy and root growth. The multiple pathways by which Stg QTLs modulate canopy development can result in considerable developmental plasticity. The reduction in canopy size associated with Stg QTLs reduced pre-flowering water demand, thereby increasing water availability during grain filling and, ultimately, grain yield. The generic physiological mechanisms underlying the stay-green trait suggest that similar Stg QTLs could enhance post-anthesis drought adaptation in other major cereals such as maize, wheat, and rice.

Abstract: Digital repeat photography is becoming widely used for near-surface remote sensing of vegetation. Canopy greenness, which has been used extensively for phenological applications, can be readily quantified from camera images. Important questions remain, however, as to whether the observed changes in canopy greenness are directly related to changes in leaf-level traits, changes in canopy structure, or some combination thereof. We investigated relationships between canopy greenness and various metrics of canopy structure and function, using five years (2008-2012) of automated digital imagery, ground observations of phenological transitions, leaf area index (LAI) measurements, and eddy covariance estimates of gross ecosystem photosynthesis from the Harvard Forest, a temperate deciduous forest in the northeastern United States. Additionally, we sampled canopy sunlit leaves on a weekly basis throughout the growing season of 2011. We measured physiological and morphological traits including leaf size, mass (wet/dry), nitrogen content, chlorophyll fluorescence, and spectral reflectance and characterized individual leaf color with flatbed scanner imagery. Our results show that observed spring and autumn phenological transition dates are well captured by information extracted from digital repeat photography. However, spring development of both LAI and the measured physiological and morphological traits are shown to lag behind spring increases in canopy greenness, which rises very quickly to its maximum value before leaves are even half their final size. Based on the hypothesis that changes in canopy greenness represent the aggregate effect of changes in both leaf-level properties (specifically, leaf color) and changes in canopy structure (specifically, LAI), we developed a two end-member mixing model. With just a single free parameter, the model was able to reproduce the observed seasonal trajectory of canopy greenness. This analysis shows that canopy greenness is relatively insensitive to changes in LAI at high LAI levels, which we further demonstrate by assessing the impact of an ice storm on both LAI and canopy greenness. Our study provides new insights into the mechanisms driving seasonal changes in canopy greenness retrieved from digital camera imagery. The nonlinear relationship between canopy greenness and canopy LAI has important implications both for phenological research applications and for assessing responses of vegetation to disturbances.

Abstract: Increasing crop productivity while simultaneously reducing the environmental footprint of crop production is considered a major challenge for the coming decades. Even short episodes of heat stress can reduce crop yield considerably causing low resource use efficiency. Studies on the impact of heat stress on crop yields over larger regions generally rely on temperatures measured by standard weather stations at 2 m height. Canopy temperatures measured in this study in field plots of rye were up to 7 °C higher than air temperature measured at typical weather station height with the differences in temperatures controlled by soil moisture contents. Relationships between heat stress and grain number derived from controlled environment studies were only confirmed under field conditions when canopy temperature was used to calculate stress thermal time. By using hourly mean temperatures measured by 78 weather stations located across Germany for the period 1994–2009 it is estimated, that mean yield declines in wheat due to heat stress during flowering were 0.7% when temperatures are measured at 2 m height, but yield declines increase to 22% for temperatures measured at the ground. These results suggest that canopy temperature should be simulated or estimated to reduce uncertainty in assessing heat stress impacts on crop yield.

Abstract: * Stay-green is an integrated drought adaptation trait characterized by a distinct green leaf phenotype during grain filling under terminal drought. We used sorghum (Sorghum bicolor), a repository of drought adaptation mechanisms, to elucidate the physiological and genetic mechanisms underpinning stay-green. * Near-isogenic sorghum lines (cv RTx7000) were characterized in a series of field and managed-environment trials (seven experiments and 14 environments) to determine the influence of four individual stay-green (Stg1–4) quantitative trait loci (QTLs) on canopy development, water use and grain yield under post-anthesis drought. * The Stg QTL decreased tillering and the size of upper leaves, which reduced canopy size at anthesis. This reduction in transpirational leaf area conserved soil water before anthesis for use during grain filling. Increased water uptake during grain filling of Stg near-isogenic lines (NILs) relative to RTx7000 resulted in higher post-anthesis biomass production, grain number and yield. Importantly, there was no consistent yield penalty associated with the Stg QTL in the irrigated control. * These results establish a link between the role of the Stg QTL in modifying canopy development and the subsequent impact on crop water use patterns and grain yield under terminal drought.

Abstract: Patterns of tropical forest functional diversity express processes of ecological assembly at multiple geographic scales and aid in predicting ecological responses to environmental change. Tree canopy chemistry underpins forest functional diversity, but the interactive role of phylogeny and environment in determining the chemical traits of tropical trees is poorly known. Collecting and analyzing foliage in 2,420 canopy tree species across 19 forests in the western Amazon, we discovered (i) systematic, community-scale shifts in average canopy chemical traits along gradients of elevation and soil fertility; (ii) strong phylogenetic partitioning of structural and defense chemicals within communities independent of variation in environmental conditions; and (iii) strong environmental control on foliar phosphorus and calcium, the two rock-derived elements limiting CO2 uptake in tropical forests. These findings indicate that the chemical diversity of western Amazonian forests occurs in a regionally nested mosaic driven by long-term chemical trait adjustment of communities to large-scale environmental filters, particularly soils and climate, and is supported by phylogenetic divergence of traits essential to foliar survival under varying environmental conditions. Geographically nested patterns of forest canopy chemical traits will play a role in determining the response and functional rearrangement of western Amazonian ecosystems to changing land use and climate.

Abstract: Leaf life span is an important plant trait associated with interspecific variation in leaf, organismal, and ecosystem processes. We hypothesized that intraspecific variation in gymnosperm needle traits with latitude reflects both selection and acclimation for traits adaptive to the associated temperature and moisture gradient. This hypothesis was supported, because across 127 sites along a 2,160-km gradient in North America individuals of Picea glauca, Picea mariana, Pinus banksiana, and Abies balsamea had longer needle life span and lower tissue nitrogen concentration with decreasing mean annual temperature. Similar patterns were noted for Pinus sylvestris across a north–south gradient in Europe. These differences highlight needle longevity as an adaptive feature important to ecological success of boreal conifers across broad climatic ranges. Additionally, differences in leaf life span directly affect annual foliage turnover rate, which along with needle physiology partially regulates carbon cycling through effects on gross primary production and net canopy carbon export. However, most, if not all, global land surface models parameterize needle longevity of boreal evergreen forests as if it were a constant. We incorporated temperature-dependent needle longevity and %nitrogen, and biomass allocation, into a land surface model, Community Atmosphere Biosphere Land Exchange, to assess their impacts on carbon cycling processes. Incorporating realistic parameterization of these variables improved predictions of canopy leaf area index and gross primary production compared with observations from flux sites. Finally, increasingly low foliage turnover and biomass fraction toward the cold far north indicate that a surprisingly small fraction of new biomass is allocated to foliage under such conditions.

Abstract: Plant phenology, a sensitive indicator of climate change, influences vegetation-atmosphere interactions by changing the carbon and water cycles from local to global scales. Camera-based phenological observations of the color changes of the vegetation canopy throughout the growing season have become popular in recent years. However, the linkages between camera phenological metrics and leaf biochemical, biophysical, and spectral properties are elusive. We measured key leaf properties including chlorophyll concentration and leaf reflectance on a weekly basis from June to November 2011 in a white oak forest on the island of Martha's Vineyard, Massachusetts, USA. Concurrently, we used a digital camera to automatically acquire daily pictures of the tree canopies. We found that there was a mismatch between the camera-based phenological metric for the canopy greenness (green chromatic coordinate, gcc) and the total chlorophyll and carotenoids concentration and leaf mass per area during late spring/early summer. The seasonal peak of gcc is approximately 20 days earlier than the peak of the total chlorophyll concentration. During the fall, both canopy and leaf redness were significantly correlated with the vegetation index for anthocyanin concentration, opening a new window to quantify vegetation senescence remotely. Satellite- and camera-based vegetation indices agreed well, suggesting that camera-based observations can be used as the ground validation for satellites. Using the high-temporal resolution dataset of leaf biochemical, biophysical, and spectral properties, our results show the strengths and potential uncertainties to use canopy color as the proxy of ecosystem functioning.

Abstract: The ecophysiology of tropical montane cloud forest (TMCF) trees is influenced by crown-level microclimate factors including regular mist/fog water inputs, and large variations in evaporative demand, which in turn can significantly impact water balance. We investigated the effect of such microclimatic factors on canopy ecophysiology and branch-level water balance in the dry season of a seasonal TMCF in Veracruz, Mexico, by quantifying both water inputs (via foliar uptake, FU) and outputs (day- and night-time transpiration, NT). Measurements of sap flow, stomatal conductance, leaf water potential and pressure-volume relations were obtained in Quercus lanceifolia, a canopy-dominant tree species. Our results indicate that FU occurred 34% of the time and led to the recovery of 9% (24 ± 9.1 L) of all the dry-season water transpired from individual branches. Capacity for FU was independently verified for seven additional common tree species. NT accounted for approximately 17% (46 L) of dry-season water loss. There was a strong correlation between FU and the duration of leaf wetness events (fog and/or rain), as well as between NT and the night-time vapour pressure deficit. Our results show the clear importance of fog and NT for the canopy water relations of Q. lanceifolia.

Abstract: Forest gaps, openings in the canopy caused by death of one or more trees, have a profound effect on forest regeneration and drive the forest growth cycle. It is therefore necessary to understand the effects of forest gaps on regeneration for modern forest management. In order to provide a quantitative assessment of the effects of forest gaps on regeneration of woody plants, we conducted this review of gap effects on woody plant regeneration on the basis of 527 observations from 42 individual papers, and reported the results of these data in a meta-analysis. Overall, densities of regenerated woody plants were significantly greater (359%) in forest gaps than on the closed-canopy forest floor. The regeneration density in gaps of plantation forests was significantly greater (P < 0.05) than that of natural forest because the regeneration in gaps of plantation forests was improved by both gap effects and experimental measures. Similarly, in comparison to natural gaps, regeneration was better enhanced in artificial gaps. Regeneration density exhibited a significantly positive correlation with gap size, but a negative correlation with gap age because the gap size decreased with increasing gap age. Shade tolerance of woody plants affected regeneration density in gaps and understory. Average regeneration density of shade-tolerant species exhibited a significantly positive response to gaps but densities remained lower in total than those of intermediate and shade-intolerant species. Gap effects on regeneration decreased in response to increasing temperature and precipitation because of the limiting effects of lower temperature and moisture on woody plant regeneration. In summary, forest gaps enhance woody plant regeneration, and the effects of gaps varied by forest type, gap characteristics, environmental factors and plant traits. The results of this meta-analysis are useful for better understanding the effects and roles of gaps on forest regeneration and forest management.

Abstract: Much of our understanding of natural forest dynamics in the temperate region of Europe is based on observational studies in old-growth remnants that have emphasized small-scale gap dynamics and equilibrium stand structure and composition. Relatively little attention has been given to the role of infrequent disturbance events in forest dynamics. In this study, we analyzed dendroecological data from four stands and three windthrow patches in an old-growth landscape in the Dinaric Mountains of Bosnia and Herzegovina to examine disturbance history, tree life history traits, and compositional dynamics. Over all stands, most decades during the past 340 years experienced less than 10% canopy loss, yet each stand showed evidence of periodic intermediate-severity disturbances that removed > 40% of the canopy, some of which were synchronized over the study area landscape. Analysis of radial growth patterns indicated several life history differences among the dominant canopy trees; beech was markedly older than fir, while growth patterns of dead and dying trees suggested that fir was able to tolerate longer periods of suppressed growth in shade. Maple had the fastest radial growth and accessed the canopy primarily through rapid early growth in canopy gaps, whereas most beech and fir experienced a period of suppressed growth prior to canopy accession. Peaks in disturbance were roughly linked to increased recruitment, but mainly of shade-tolerant beech and fir; less tolerant species (i.e., maple, ash, and elm) recruited successfully on some of the windthown sites where advance regeneration of beech and fir was less abundant. The results challenge the traditional notions of stability in temperate old-growth forests of Europe and highlight the nonequilibrial nature of canopy composition due to unique histories of disturbance and tree life history differences. These findings provide valuable information for developing natural disturbance-based silvicultural systems, as well as insight into maintaining less shade-tolerant, but valuable broadleaved trees in temperate forests of Europe.

Abstract: Spanning 15% of the global ice-free terrestrial surface, agricultural lands provide an immense and near-term opportunity to address climate change, food, and water security challenges. Through the computationally informed breeding of canopy structural traits away from those of modern cultivars, we show that solutions exist that increase productivity and water use efficiency, while increasing land-surface reflectivity to offset greenhouse gas warming. Plants have evolved to maximize capture of radiation in the upper leaves, thus shading competitors. While important for survival in the wild, this is suboptimal in monoculture crop fields for maximizing productivity and other biogeophysical services. Crop progenitors evolved over the last 25 million years in an atmosphere with less than half the [CO2] projected for 2050. By altering leaf photosynthetic rates, rising [CO2] and temperature may also alter the optimal canopy form. Here using soybean, the world's most important protein crop, as an example we show by applying optimization routines to a micrometeorological leaf canopy model linked to a steady-state model of photosynthesis, that significant gains in production, water use, and reflectivity are possible with no additional demand on resources. By modifying total canopy leaf area, its vertical profile and angular distribution, and shortwave radiation reflectivity, all traits available in most major crop germplasm collections, increases in productivity (7%) are possible with no change in water use or albedo. Alternatively, improvements in water use (13%) or albedo (34%) can likewise be made with no loss of productivity, under Corn Belt climate conditions.

Abstract: Fruits from three cultivars of apple (Malus × domestica Borkh.)-'McIntosh', 'Gala' and 'Mutsu'-were harvested from the exterior and interior of the tree canopy. Peel and flesh tissues were sampled separately to determine how the position of the fruit on the tree might affect the levels of the primary and secondary metabolites in the fruit. Fruit from the outer-canopy had a higher fresh weight and a higher soluble solids content compared with inner-canopy fruit. Both the flesh and peel of the outer-canopy fruit had higher concentrations of soluble sugars and sugar alcohols, but lower starch concentrations than the inner-canopy fruit. Canopy position did not significantly affect malic acid concentrations, except in the peel of 'McIntosh' and the flesh of 'Mutsu'. Although levels of ascorbic and succinic acids were higher in the peel of the outer-canopy fruit, the responses of other organic acids to canopy position depended on tissue type and cultivar. Except for histidine, lysine, threonine and glycine, most amino acids accumulated at higher concentrations in the inner-canopy fruit. By contrast, levels of phenolic compounds from both the peel and flesh were significantly higher in the outer-canopy fruit. The significant effects of location within the canopy on both primary metabolites and secondary metabolites demonstrate the importance of light exposure on apple fruit quality.

Abstract: Precise knowledge of fuel conditions is important for predicting fire hazards and simulating fire growth and intensity across the landscape. We present a methodology to retrieve and map forest canopy fuel and other forest structural parameters using small-footprint full-waveform airborne light detection and ranging (LiDAR) data. Full-waveform LiDAR sensors register the complete returned backscattered signal through time and can describe physical properties of the intercepted objects. This study was undertaken in a mixed forest dominated by Douglas-fir, occasionally mixed with other conifers, in north-west Oregon (United States). We extracted two sets of LiDAR metrics using pulse detection and waveform modelling and then constructed several predictive models using forward stepwise multiple linear regression. The resulting models explained ~80% of the variability for many of the canopy fuel and forest structure parameters: aboveground biomass (R2=0.84), quadratic mean diameter (R2=0.82), canopy height (R2=0.79), canopy base height (R2=0.78) and canopy fuel load (R2=0.79). The lowest performing models included basal area (R2=0.76), stand volume (R2=0.73), canopy bulk density (R2=0.67) and stand density index (R2=0.66). Our results indicate that full-waveform LiDAR systems show promise in systematically characterising the structure and canopy fuel loads of forests, which may enable accurate fire behaviour forecasting that in turn supports the development of prevention and planning policies.

Abstract: Global climate warming is one of the key forces driving plant community shifts, such as range shifts of temperate species into boreal forests. As plant community shifts are slow to observe, ecotones, boundaries between two ecosystems, are target areas for providing early evidence of ecological responses to warming. The role of soil fauna is poorly explored in ecotones, although their positive and negative effects on plant species can influence plant community structure. We studied nematode communities in response to experimental warming (ambient, +1.7, +3.4 °C) in soils of closed and open canopy forest in the temperate-boreal ecotone of Minnesota, USA and calculated various established nematode indices. We estimated species-specific coverage of understory herbaceous and shrub plant species from the same experimental plots and tested if changes in the nematode community are associated with plant cover and composition. Individual nematode trophic groups did not differ among warming treatments, but the ratio between microbial-feeding and plant-feeding nematodes increased significantly and consistently with warming in both closed and open canopy areas and at both experimental field sites. The increase in this ratio was positively correlated with total cover of understory plant species, perhaps due to increased predation pressure on soil microorganisms causing higher nutrient availability for plants. Multivariate analyses revealed that temperature treatment, canopy conditions and nematode density consistently shaped understory plant communities across experimental sites. Our findings suggest that warming-induced changes in nematode community structure are associated with shifts in plant community composition and productivity in the temperate-boreal forest ecotones.

Abstract: 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.