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

Background and Aims

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

Methods

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

Key Results

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

Conclusions

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

Establishment of

We compared establishment of Douglas fir (Pseudotsuga menziesii) and Corsican pine (Pinus nigra) seedlings in kanuka (Kunzea ericoides) and manuka (Leptospermum scoparium) shrubland to test the hypothesis that Douglas fir, because of its greater shade tolerance, is better able to establish in woody communities than pine species. Seed of the conifer species was sown under a range of canopy covers at six sites, the cover being low-statured vegetation in openings between stands, stand edges, and moderate and dense canopies. After three growing seasons, survival of Corsican pine seedlings was greatest in the open and declined progressively as canopy cover increased. This contrasted with Douglas fir, where survival was greatest at the canopy edge. Survival of Douglas fir seedlings significantly exceeded that of Corscican pine seedlings under dense canopy positions. Seedling numbers of both species declined significantly with increasing leaf area index of manuka, but not kanuka stands, where seedling numbers were lower. Leaf area index of manuka stands accounted for substantially greater variation in number and survival of Corsican pine than Douglas fir seedlings. It is concluded that Douglas fir is better able to establish in shaded environments in woody communities than Corsican pine; however, further monitoring is required to confirm the long-term survival of both species under the moderate and dense canopy positions in this trial.     

Photosynthesis and Photoprotection in Overwintering Plants

Abstract: Seasonal differences in the capacity of photosynthetic electron transport, leaf pigment composition, xanthophyll cycle characteristics and chlorophyll fluorescence emission were investigated in two biennial mesophytes ( Malva neglecta and Verbascum thapsus ) that grow in full sunlight, and in leaves/needles of sun and shade populations of several broad-leafed evergreens and conifers (Vinca minor, Euonymus kiautschovicus, Mahonia repens, Pseudotsuga menziesii [Douglas fir], and Pinus ponderosa). Both mesophytic species maintained or upregulated photosynthetic capacity in the winter and exhibited no upregulation of photoprotection. In contrast, photosynthetic capacity was downregulated in sun leaves/needles of V. minor, Douglas fir, and Ponderosa pine, and even in shade needles of Douglas fir. Interestingly, photosynthetic capacity was upregulated during the winter in shade leaves/needles of V. minor, Ponderosa pine and Euonymus kiautschovicus. Nocturnal retention of zeaxanthin and antheraxanthin, and their sustained engagement in a state primed for energy dissipation, were observed largely in the leaves/needles of sun-exposed evergreen species during winter. Factors that may contribute to these differing responses to winter stress, including chloroplast redox state, the relative levels of source and sink activity at the whole plant level, and apoplastic versus symplastic phloem loading, are discussed.     

A leaf gas exchange model that accounts for intra-canopy variability by considering leaf nitrogen content and local acclimation to radiation in grapevine (Vitis vinifera L.)

Understanding the distribution of gas exchange within a plant is a prerequisite for scaling up from leaves to canopies. We evaluated whether leaf traits were reliable predictors of the effects of leaf ageing and leaf irradiance on leaf photosynthetic capacity (V(cmax) , J(max) ) in field-grown vines (Vitis vinifera L). Simultaneously, we measured gas exchange, leaf mass per area (LMA) and nitrogen content (N(m) ) of leaves at different positions within the canopy and at different phenological stages. Daily mean leaf irradiance cumulated over 10 d (PPFD(10) ) was obtained by 3D modelling of the canopy structure. N(m) decreased over the season in parallel to leaf ageing while LMA was mainly affected by leaf position. PPFD(10) explained 66, 28 and 73% of the variation of LMA, N(m) and nitrogen content per area (N(a) ), respectively. Nitrogen content per unit area (N(a) = LMA × N(m) ) was the best predictor of the intra-canopy variability of leaf photosynthetic capacity. Finally, we developed a classical photosynthesis-stomatal conductance submodel and by introducing N(a) as an input, the model accurately simulated the daily pattern of gas exchange for leaves at different positions in the canopy and at different phenological stages during the season.     

Responses of leaf structure and photosynthetic properties to intra-canopy light gradients: a common garden test with four broadleaf deciduous angiosperm and seven evergreen conifer tree species

Spectra of leaf traits in northern temperate forest canopies reflect major differences in leaf longevity between evergreen conifers and deciduous broadleaf angiosperms, as well as plastic modifications caused by within-crown shading. We investigated (1) whether long-lived conifer leaves exhibit similar intra-canopy plasticity as short-lived broadleaves, and (2) whether global interspecific relationships between photosynthesis, nitrogen, and leaf structure identified for sun leaves adequately describe leaves differentiated in response to light gradients. We studied structural and photosynthetic properties of intra-tree sun and shade foliage in adult trees of seven conifer and four broadleaf angiosperm species in a common garden in Poland. Shade leaves exhibited lower leaf mass-per-area (LMA) than sun leaves; however, the relative difference was smaller in conifers than in broadleaves. In broadleaves, LMA was correlated with lamina thickness and tissue density, while in conifers, it was correlated with thickness but not density. In broadleaves, but not in conifers, reduction of lamina thickness was correlated with a thinner palisade layer. The more conservative adjustment of conifer leaves could result from a combination of phylogenetic constraints, contrasting leaf anatomies and shoot geometries, but also from functional requirements of long-lived foliage. Mass-based nitrogen concentration (N(mass)) was similar between sun and shade leaves, and was lower in conifers than in deciduous broadleaved species. Given this, the smaller LMA in shade corresponded with a lower area-based N concentration (N(area)). In evergreen conifers, LMA and N(area) were less powerful predictors of area-based photosynthetic rate (A (max(area))) in comparison with deciduous broadleaved angiosperms. Multiple regression for sun and shade leaves showed that, in each group, A (max(mass)) was related to N(mass) but not to LMA, whereas LMA became a significant codeterminant of A (max(mass)) in analysis combining both groups. Thus, a fundamental mass-based relationship between photosynthesis, nitrogen, and leaf structure reported previously also exists in a dataset combining within-crown and across-functional type variation.     

Photosynthetic Response to Irradiance in Valeriana jatamansi Jones,a Threatened Understorey Medicinal Herb of Western Himalaya

Net photosynthetic rate (P _N) of Valeriana jatamansi plants, grown under nylon net shade or under different tree canopies, was saturated with photons at 1 000 mol m^–2 s^–1 photosynthetic photon-flux-density (PPFD), whereas open-grown plants were able to photosynthesise even at higher PPFD, e.g. of 2 000 mol m^–2 s^–1. Plants grown under net shade had higher total chlorophyll (Chl) content per unit area of leaf surface. However, Chl a/b ratio was maximal in open-grown plants, but remained unchanged in plants grown in nylon net shade and under different tree canopies. Sun-grown plants had thicker leaves (higher leaf mass per leaf area unit), higher wax content, and higher P _N than shade grown plants. Thus V. jatamansi is able to acclimate to high PPFD and therefore this Himalayan species may be cultivated in open habitat to meet the ever-increasing industrial demand.     

Photosynthesis and resource distribution through plant canopies

Plant canopies are characterized by dramatic gradients of light between canopy top and bottom, and interactions between light, temperature and water vapour deficits. This review summarizes current knowledge of potentials and limitations of acclimation of foliage photosynthetic capacity (A(max)) and light-harvesting efficiency to complex environmental gradients within the canopies. Acclimation of A(max) to high light availability involves accumulation of rate-limiting photosynthetic proteins per unit leaf area as the result of increases in leaf thickness in broad-leaved species and volume: total area ratio and mesophyll thickness in species with complex geometry of leaf cross-section. Enhancement of light-harvesting efficiency in low light occurs through increased chlorophyll production per unit dry mass, greater leaf area per unit dry mass investment in leaves and shoot architectural modifications that improve leaf exposure and reduce within-shoot shading. All these acclimation responses vary among species, resulting in species-specific use efficiencies of low and high light. In fast-growing canopies and in evergreen species, where foliage developed and acclimated to a certain light environment becomes shaded by newly developing foliage, leaf senescence, age-dependent changes in cell wall characteristics and limited foliage re-acclimation capacity can constrain adjustment of older leaves to modified light availabilities. The review further demonstrates that leaves in different canopy positions respond differently to dynamic fluctuations in light availability and to multiple environmental stresses. Foliage acclimated to high irradiance respond more plastically to rapid changes in leaf light environment, and is more resistant to co-occurring heat and water stress. However, in higher light, co-occurring stresses can more strongly curb the efficiency of foliage photosynthetic machinery through reductions in internal diffusion conductance to CO(2). This review demonstrates strong foliage potential for acclimation to within-canopy environmental gradients, but also highlights complex constraints on acclimation and foliage functioning resulting from light x foliage age interactions, multiple environmental stresses, dynamic light fluctuations and species-specific leaf and shoot structural constraints.     

Scaling CO2-photosynthesis relationships from the leaf to the canopy

Responses of individual leaves to short-term changes in CO₂ partial pressure have been relatively well studied. Whole-plant and plant community responses to elevated CO₂ are less well understood and scaling up from leaves to canopies will be complicated if feedbacks at the small scale differ from feedbacks at the large scale. Mathematical models of leaf, canopy, and ecosystem processes are important tools in the study of effects on plants and ecosystems of global environmental change, and in particular increasing atmospheric CO₂, and might be used to scale from leaves to canopies. Models are also important in assessing effects of the biosphere on the atmosphere. Presently, multilayer and big leaf models of canopy photosynthesis and energy exchange exist. Big leaf models — which are advocated here as being applicable to the evaluation of impacts of global change on the biosphere — simplify much of the underlying leaf-level physics, physiology, and biochemistry, yet can retain the important features of plant-environment interactions with respect to leaf CO₂ exchange processes and are able to make useful, quantitative predictions of canopy and community responses to environmental change. The basis of some big leaf models of photosynthesis, including a new model described herein, is that photosynthetic capacity and activity are scaled vertically within a canopy (by plants themselves) to match approximately the vertical profile of PPFD. The new big leaf model combines physically based models of leaf and canopy level transport processes with a biochemically based model of CO₂ assimilation. Predictions made by the model are consistent with canopy CO₂ exchange measurements, although a need exists for further testing of this and other canopy physiology models with independent measurements of canopy mass and energy exchange at the time scale of 1 h or less.Abbreviations LAI leaf area index - NIR near infrared (700–3000 nm) radiation - PAR photosynthetically active (400–700 nm) radiation - PI photosynthetic irradiance (400–700 nm) - PPFD photosynthetic photon flux area density (400–700 nm) - PS I Photosystem I - PS II Photosystem II - Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase - RuP₂ ribulose-1,5-bisphosphate     

Photosynthetic acclimation of the liana Stigmaphyllon lindenianum to light changes in a tropical dry forest canopy

Tropical plant canopies show abrupt changes in light conditions across small differences in spatial and temporal scales. Given the canopy light heterogeneity, plants in this stratum should express a high degree of plasticity, both in space (allocation to plant modules as a function of opportunity for resource access) and time (photosynthetic adjustment to temporal changes in the local environment). Using a construction crane for canopy access, we studied light acclimation of the liana Stigmaphyllon lindenianum to sun and shade environments in a tropical dry forest in Panama during the wet season. Measured branches were randomly distributed in one of four light sequences: high- to low-light branches started the experiment under sun and were transferred to shade during the second part of the experiment; low- to high-light branches (LH) were exposed to the opposite sequence of light treatments; and high-light and low-light controls , which were exposed only to sun and shade environments, respectively, throughout the experiment. Shade branches were set inside enclosures wrapped in 63% greenhouse shade cloth. After 2 months, we transferred experimental branches to opposite light conditions by relocating the enclosures. Leaf mortality was considerably higher under shade, both before and after the transfer. LH branches reversed the pattern of mortality by increasing new leaf production after the transfer. Rates of photosynthesis at light saturation, light compensation points, and dark respiration rates of transferred branches matched those of controls for the new light treatment, indicating rapid photochemical acclimation. The post-expansion acclimation of sun and shade foliage occurred with little modification of leaf structure. High photosynthetic plasticity was reflected in an almost immediate ability to respond to significant changes in light. This response did not depend on the initial light environment, but was determined by exposure to new light conditions. Stigmaphyllon responded rapidly to light changes through the functional adjustment of already expanded foliage and an increase in leaf production in places with high opportunity for carbon gain. Received: 24 April 1998 / Accepted: 11 May 1999     

Orientation behavior of the predator Laricobius nigrinus (Coleoptera: Derodontidae) to hemlock woolly adelgid and host tree odors in a multi-chambered olfactometer

We studied the adult ambulatory response of the predator, Laricobius nigrinus Fender (Coleoptera: Derodontidae), to odors from its prey, Adelges tsugae Annand, the hemlock woolly adelgid, and foliage of hemlock woolly adelgid, host hemlocks (Tsuga spp.), and other conifers. Both the predator and hemlock woolly adelgid are apparently native to western North America, but the predator is being released in the eastern United States, which has different hemlock species, for biological control of a lineage of hemlock woolly adelgid inadvertently introduced from Japan. L. nigrinus responded to odors from hemlock woolly adelgid host trees, but not to odors from hemlock woolly adelgid. L. nigrinus collected from hemlock woolly adelgid-infested western hemlock were more strongly attracted to odors from western hemlock [Tsuga heterophylla (Rafinesque) Sargent] than eastern hemlock [Tsuga canadensis (L.) Carrière] in most trials. Odors from western white pine (Pinus monticola Douglas ex D. Don) and white spruce [Picea glauca (Moench) Voss] were as attractive as western hemlock odors whereas odors from Douglas-fir [Pseudotsuga menziesii variety menziesii (Mirbel)] and ponderosa pine (Pinus ponderosa Douglas ex Lawson) were avoided. L. nigrinus reared on hemlock woolly adelgid-infested eastern hemlock in the laboratory were lethargic and were not attracted to either eastern or western hemlock odors. Predators collected in the field and tested monthly from December to March responded similarly each month, except February, when they flew rather than walked in the olfactometer, suggesting a period of dispersal or mate finding at that time of year. The implications of these results for programs to release L. nigrinus in the eastern United States for control of hemlock woolly adelgid are discussed.     

A review of light interception in plant stands from leaf to canopy in different plant functional types and in species with varying shade tolerance

Changes in the efficiency of light interception and in the costs for light harvesting along the light gradients from the top of the plant canopy to the bottom are the major means by which efficient light harvesting is achieved in ecosystems. In the current review analysis, leaf, shoot and canopy level determinants of plant light harvesting, the light-driven plasticity in key traits altering light harvesting, and variations among different plant functional types and between species of different shade tolerance are analyzed. In addition, plant age- and size-dependent alterations in light harvesting efficiency are also examined. At the leaf level, the variations in light harvesting are driven by alterations in leaf chlorophyll content modifies the fraction of incident light harvested by given leaf area, and in leaf dry mass per unit area (M _A) that determines the amount of leaf area formed with certain fraction of plant biomass in the leaves. In needle-leaved species with complex foliage cross-section, the degree of foliage surface exposure also depends on the leaf total-to-projected surface area ratio. At the shoot scale, foliage inclination angle distribution and foliage spatial aggregation are the major determinants of light harvesting, while at the canopy scale, branching frequency, foliage distribution and biomass allocation to leaves (F _L) modify light harvesting significantly. F _L decreases with increasing plant size from herbs to shrubs to trees due to progressively larger support costs in plant functional types with greater stature. Among trees, F _L and stand leaf area index scale positively with foliage longevity. Plant traits altering light harvesting have a large potential to adjust to light availability. Chlorophyll per mass increases, while M _A, foliage inclination from the horizontal and degree of spatial aggregation decrease with decreasing light availability. In addition, branching frequency decreases and canopies become flatter in lower light. All these plastic modifications greatly enhance light harvesting in low light. Species with greater shade tolerance typically form a more extensive canopy by having lower M _A in deciduous species and enhanced leaf longevity in evergreens. In addition, young plants of shade tolerators commonly have less strongly aggregated foliage and flatter canopies, while in adult plants partly exposed to high light, higher shade tolerance of foliage allows the shade tolerators to maintain more leaf layers, resulting in extended crowns. Within a given plant functional type, increases in plant age and size result in increases in M _A, reductions in F _L and increases in foliage aggregation, thereby reducing plant leaf area index and the efficiency of light harvesting. Such dynamic modifications in plant light harvesting play a key role in stand development and productivity. Overall, the current review analysis demonstrates that a suite of chemical and architectural traits at various scales and their plasticity drive plant light harvesting efficiency. Enhanced light harvesting can be achieved by various combinations of traits, and these suites of traits vary during plant ontogeny.     

Inter- and intraspecific relationships between shade tolerance and shade avoidance in temperate trees

Patterns of physiological and architectural adaptation and acclimation to decreasing light availability were investigated along a light gradient for saplings of 12 common species of temperate deciduous trees in southeastern Ontario, Canada. Physiological adaptation and acclimation (shade tolerance physiology) were quantified at the leaf level by measuring leaf mass per unit area (LMA), dark respiration per unit leaf nitrogen, chlorophyll per unit leaf nitrogen and the chlorophyll a:b ratio for the newest fully expanded leaf on the leader. Architectural adaptation and acclimation (shade avoidance) were quantified by measuring branching intensity and side shoot:main shoot length ratios for the most recent three years of growth on the leader and selected side branches. Within species, increases in LMA, chlorophyll a:b ratio and respiration per unit nitrogen and decreases in chlorophyll per unit nitrogen indicated that shade tolerance physiology generally increased with decreasing canopy openness. Increases in the branching intensity and side shoot:main shoot ratios of the leader and side branches indicated that shade avoidance also increased with decreasing canopy openness for the majority of species; however, in some species, stem bending under deep shade resulted in lateral growth. Interspecific variation in shade tolerance physiology was minimal when species were compared under equal amounts of canopy openness. In contrast, interspecific variation in shade avoidance variables was relatively high under equal canopy openness, with saplings of shade tolerant canopy species exhibiting higher shade avoidance than saplings of shade tolerant understory species.     

Seasonal and diurnal leaf movements of Rhododendron maximum L. in contrasting irradiance environments

Summary Leaf orientation (azimuth and angle) and leaf curling were measured seasonally and diurnally on Rhododendron maximum L. under an evergreen and a deciduous canopy. The microclimatic conditions under the evergreen canopy (mixed pine and hemlock) were characterized by lower irradiance but similar temperature, and vapor pressure deficit (vpd) to that under the deciduous canopy (mixed oak and maple). Under both canopies irradiance was more intense during winter months.On a seasonal basis leaf angle was closer to horizontal under the evergreen canopy but there was no difference between leaf curling in the two sites. Stomatal conductance was higher under the deciduous canopy but stomata were closed in the winter (following canopy abscission) under the evergreen and deciduous canopies even during warm winter days. Leaf water potentials were lower in the winter and Rhododendron maximum had higher leaf water potentials under the evergreen canopy.Significant association between mean leaf angle and curling index were found above a mean leaf angle of 70°. Leaf curling was highly associated with leaf temperature where 0° C was a critical value stimulating leaf curling. Leaf angle was linearly related to leaf temperatures above 0° C although this relationship was different under the two canopy types as a result of differing irradiance or differing water potential.     

Canopy structure,light microclimate and leaf gas exchange of Quercus coccifera L. in a Portuguese macchia: measurements in different canopy layers and simulations with a canopy model

Summary The structural characteristics of a diverse array of Quercus coccifera canopies were assessed and related to measured and computed light attenuation, proportion of sunlit foliage, foliage temperatures, and photosynthesis and diffusive conductance behavior in different canopy layers. A canopy model incorporating all components of shortwave and longwave radiation, and the energy balance, conductance, and CO₂ and H₂O exchanges of all leaf layers was developed and compared with measurements of microclimate and gas exchange in canopies in four seasons of the year. In the denser canopies with a leaf area index (LAI) greater than 5, there is little sunlit foliage and the diffuse radiation (400–700 nm) is attenuated to 5% or less of the global radiation (400–700 nm) incident on the top of the canopy. Foliage of this species is nonrandomly distributed with respect to azimuth angle, and within each canopy layer, foliage azimuth and inclination angles are correlated. A detailed version of the model which computed radiation interception and photosynthetic light harvesting according to these nonrandom distributions indicated little difference in whole-canopy gas exchange from calculations of the normal model, which assumes random azimuth orientation. The contributions of different leaf layers to canopy gas exchange are not only a function of the canopy microclimate, but also the degree to which leaves in the lower layers of the canopy exhibit more shade-leaf characteristics, such as low photosynthetic and respiratory capacity and maximal conductance. On cloudless days, the majority of the foliage in a canopy of 5.4 LAI is shaded —70%–90% depending on the time of year. Yet, the shaded foliage under these conditions is calculated to contribute only about one-third of the canopy carbon gain. This contribution is about the same as that of the upper 13% of the canopy foliage. Computed annual whole-canopy carbon gain and water use are, respectively, 60% and 100% greater for a canopy of 5 LAI than for one of 2 LAI. Canopy water-use efficiency is correspondingly less for the canopy of 5 LAI than for that of 2 LAI, but most of this difference is apparent during the cool months of the year, when moisture is more abundant.     

Coupled response of stomatal and mesophyll conductance to light enhances photosynthesis of shade leaves under sunflecks

Light gradients within tree canopies play a major role in the distribution of plant resources that define the photosynthetic capacity of sun and shade leaves. However, the biochemical and diffusional constraints on gas exchange in sun and shade leaves in response to light remain poorly quantified, but critical for predicting canopy carbon and water exchange. To investigate the CO₂ diffusion pathway of sun and shade leaves, leaf gas exchange was coupled with concurrent measurements of carbon isotope discrimination to measure net leaf photosynthesis (A_n), stomatal conductance (g_s) and mesophyll conductance (g_m) in Eucalyptus tereticornis trees grown in climate controlled whole‐tree chambers. Compared to sun leaves, shade leaves had lower A_n, g_m, leaf nitrogen and photosynthetic capacity (A_max) but g_s was similar. When light intensity was temporarily increased for shade leaves to match that of sun leaves, both g_s and g_m increased, and A_n increased to values greater than sun leaves. We show that dynamic physiological responses of shade leaves to altered light environments have implications for up‐scaling leaf level measurements and predicting whole canopy carbon gain. Despite exhibiting reduced photosynthetic capacity, the rapid up‐regulation of g_m with increased light enables shade leaves to respond quickly to sunflecks.     

Light and VPD gradients drive foliar nitrogen partitioning and photosynthesis in the canopy of European beech and silver fir

While foliar photosynthetic relationships with light, nitrogen, and water availability have been well described, environmental factors driving vertical gradients of foliar traits within forest canopies are still not well understood. We, therefore, examined how light availability and vapour pressure deficit (VPD) co-determine vertical gradients (between 12 and 42 m and in the understorey) of foliar photosynthetic capacity (Amax), 13C fractionation (∆), specific leaf area (SLA), chlorophyll (Chl), and nitrogen (N) concentrations in canopies of Fagus sylvatica and Abies alba growing in a mixed forest in Switzerland in spring and summer 2017. Both species showed lower Chl/N and lower SLA with higher light availability and VPD at the top canopy. Despite these biochemical and morphological acclimations, Amax during summer remained relatively constant and the photosynthetic N-use efficiency (PNUE) decreased with higher light availability for both species, suggesting suboptimal N allocation within the canopy. ∆ of both species were lower at the canopy top compared to the bottom, indicating high water-use efficiency (WUE). VPD gradients strongly co-determined the vertical distribution of Chl, N, and PNUE in F. sylvatica, suggesting stomatal limitation of photosynthesis in the top canopy, whereas these traits were only related to light availability in A. alba. Lower PNUE in F. sylvatica with higher WUE clearly indicated a trade-off in water vs. N use, limiting foliar acclimation to high light and VPD at the top canopy. Species-specific trade-offs in foliar acclimation to environmental canopy gradients may thus be considered for scaling photosynthesis from leaf to canopy to landscape levels.     

Decomposition and nitrogen release from the foliage litter of fir (Abies sachalinensis) and oak (Quercus crispula) under different forest canopies in Hokkaido, Japan

When two tree species co-occur, decomposition and nitrogen (N) release from the foliage litter depend on two factors: the forest floor conditions under each canopy type and the species composition of the litter. We conducted an experiment using fir and oak to answer several questions regarding decomposition beneath canopies of the two species and the effects of litter species composition on decomposition. We compared the rates of decomposition and N release from three different litters (fir needle, oak leaf, and a mixture of the two) in 1-mm-mesh litterbags on the forest floor under three different canopies (a 40-year-old fir plantation, large oak trees, and mixed fir and oak trees) in Hokkaido, Japan, over a 2-year period. Beneath each of these canopy types, the litter decomposition rate and percentage of N remaining in the litterbags containing a mixture of fir and oak litter were not significantly different from the expected values calculated for litterbags containing litter from a single tree species. Oak leaf litter decomposed significantly faster than fir needle litter beneath each canopy type. The litter decomposition rate was significantly higher beneath the fir canopy than under the oak canopy, and was intermediate under the mixed canopy of fir and oak. No net N release, that is, a decrease in the total N compared to the original amount, was detected from fir litter under each canopy type or from oak leaf litter beneath the oak canopy. N increased over the original amount in the fir litter beneath the oak canopy and the mixed canopy of fir and oak, but N was released from the oak litter under the fir canopy and the mixed canopy of fir and oak. These results suggest that oak leaf litter blown onto fir forest floor enhances nutrient cycling, and this might be a positive effect of a mixed stand of conifer and broad-leaved trees.     

Distribution of leaf characteristics in relation to orientation within the canopy of woody species

Over the last few decades considerable effort has been devoted to research of leaf adaptations to environmental conditions. Many studies have reported strong differences in leaf mass per unit area (LMA) within a single tree depending on the photosynthetic photon flux density (PPFD) incident on different locations in the crown. There are fewer studies, however, of the effects of differences in the timing of light incidence during the day on different crown orientations.Leaves from isolated trees of Quercus suber and Quercus ilex in a cold Mediterranean climate were sampled to analyze differences in LMA and other leaf traits among different crown orientations. Gas-exchange rates, leaf water potentials, leaf temperatures and PPFD incident on leaf surfaces in different crown orientations were also measured throughout one entire summer day for each species.Mean daily PPFD values were similar for the leaves from the eastern and western sides of the canopy. On the western side, PPFD reached maximum values during the afternoon. Maximum leaf temperatures were approximately 10–20% higher on the west side, whereas minimum leaf water potentials were between 10 and 24% higher on the east side. Maximum transpiration rates were approximately 22% greater on the west, because of the greater leaf-to-air vapor pressure deficits (LAVPD). Mean individual leaf area was around 10% larger on the east than on the west side of the trees. In contrast, there were no significant differences in LMA between east and west sides of the crown.Contrary to our expectations, more severe water stress on the west side did not result in increases in LMA, although it was associated with lower individual leaf area. We conclude that increases in LMA measured by other authors along gradients of water stress would be due to differences in light intensity between dry and humid sites.     

Monoterpene emission from coniferous trees in response to elevated CO2 concentration and climate warming

It was hypothesized that high CO₂ availability would increase monoterpene emission to the atmosphere. This hypothesis was based on resource allocation theory which predicts increased production of plant secondary compounds when carbon is in excess of that required for growth. Monoterpene emission rates were measured from needles of (a) Ponderosa pine grown at different CO₂ concentrations and soil nitrogen levels, and (b) Douglas fir grown at different CO₂ concentrations. Ponderosa pine grown at 700 μmol mol^–1 CO₂ exhibited increased photosynthetic rates and needle starch to nitrogen (N) ratios when compared to trees grown at 350 μmol mol^–1 CO₂. Nitrogen availability had no consistent effect on photosynthesis. Douglas fir grown at 550 μmol mol^–1 CO₂ exhibited increased photosynthetic rates as compared to growth at 350 μmol mol^–1 CO₂ in old, but not young needles, and there was no influence on the starch/N ratio. In neither species was there a significant effect of elevated growth CO₂ on needle monoterpene concentration or emission rate. The influence of climate warming and leaf area index (LAI) on monoterpene emission were also investigated. Douglas fir grown at elevated CO₂ plus a 4 °C increase in growth temperature exhibited no change in needle monoterpene concentration, despite a predicted 50% increase in emission rate. At elevated CO₂ concentration the LAI increased in Ponderosa pine, but not Douglas fir. The combination of increased LAI and climate warming are predicted to cause an 80% increase in monoterpene emissions from Ponderosa pine forests and a 50% increase in emissions from Douglas fir forests. This study demonstrates that although growth at elevated CO₂ may not affect the rate of monoterpene emission per unit biomass, the effect of elevated CO₂ on LAI, and the effect of climate warming on monoterpene biosynthesis and volatilization, could increase canopy monoterpene emission rate.     

Acclimation of plants to light gradients in leaf canopies: evidence for a possible role for cytokinins transported in the transpiration stream.

The mechanism of response of plants to vertical light intensity gradients in leaf canopies was investigated. Since shaded leaves transpire less than leaves in high light, it was hypothesized that cytokinins (CKs) carried by mass transport in the transpiration stream would be distributed over the leaf area of partially shaded plants parallel to the gradient in light intensity. It was also hypothesized that this causes the distribution of leaf growth, leaf N and photosynthetic capacity, and possibly chloroplast acclimation as observed in plants growing in leaf canopies. In a field experiment, the distribution of Ca, N and CKs in a bean leaf canopy of a dense and an open stand supported the concept of a role for CKs in the response of N allocation to the light gradient when a decreasing sensitivity for CKs with increasing leaf age is assumed. Both shading of one leaf of the pair of primary bean leaves and independent reduction of its transpiration rate in a growth cabinet experiment caused lower dry mass, N and Ca per unit leaf area in comparison to the opposite not treated leaf. Shading caused a parallel reduction in CK concentration, which supports the hypothesis, but independent reduction of transpiration rate failed to do the same. Application of benzylaminopurine (BA) counteracted the reduction caused by shade of leaf N, photosynthetic capacity and leaf area growth. The experiments show an important role for the transpiration stream in the response of plants to light gradients. Evidence is presented here that CKs carried in the transpiration stream may be important mediators for the acclimation of plants to leaf canopy density.