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

Background and Aims

Plants are expected to maximize their net photosynthetic gains and efficiently use available resources, but the fundamental principles governing trade-offs in suites of traits related to resource-use optimization remain uncertain. This study investigated whether Acer saccharum (sugar maple) saplings could maximize their net photosynthetic gains through a combination of crown structure and foliar characteristics that let all leaves maximize their photosynthetic light-use efficiency (ɛ).

Methods

A functional–structural model, LIGNUM, was used to simulate individuals of different leaf area index (LAI_ind) together with a genetic algorithm to find distributions of leaf angle (L_A) and leaf photosynthetic capacity (A_max) that maximized net carbon gain at the whole-plant level. Saplings grown in either the open or in a forest gap were simulated with A_max either unconstrained or constrained to an upper value consistent with reported values for A_max in A. saccharum.

Key Results

It was found that total net photosynthetic gain was highest when whole-plant PPFD absorption and leaf ɛ were simultaneously maximized. Maximization of ɛ required simultaneous adjustments in L_A and A_max along gradients of PPFD in the plants. When A_max was constrained to a maximum, plants growing in the open maximized their PPFD absorption but not ɛ because PPFD incident on leaves was higher than the PPFD at which ɛ_max was attainable. Average leaf ɛ in constrained plants nonetheless improved with increasing LAI_ind because of an increase in self-shading.

Conclusions

It is concluded that there are selective pressures for plants to simultaneously maximize both PPFD absorption at the scale of the whole individual and ɛ at the scale of leaves, which requires a highly integrated response between L_A, A_max and LAI_ind. The results also suggest that to maximize ɛ plants have evolved mechanisms that co-ordinate the L_A and A_max of individual leaves with PPFD availability.     

Stem hydraulic traits and leaf water-stress tolerance are co-ordinated with the leaf phenology of angiosperm trees in an Asian tropical dry karst forest

Background and Aims

The co-occurring of evergreen and deciduous angiosperm trees in Asian tropical dry forests on karst substrates suggests the existence of different water-use strategies among species. In this study it is hypothesized that the co-occurring evergreen and deciduous trees differ in stem hydraulic traits and leaf water relationships, and there will be correlated evolution in drought tolerance between leaves and stems.

Methods

A comparison was made of stem hydraulic conductivity, vulnerability curves, wood anatomy, leaf life span, leaf pressure–volume characteristics and photosynthetic capacity of six evergreen and six deciduous tree species co-occurring in a tropical dry karst forest in south-west China. The correlated evolution of leaf and stem traits was examined using both traditional and phylogenetic independent contrasts correlations.

Key Results

It was found that the deciduous trees had higher stem hydraulic efficiency, greater hydraulically weighted vessel diameter (D_h) and higher mass-based photosynthetic rate (A_m); while the evergreen species had greater xylem-cavitation resistance, lower leaf turgor-loss point water potential (π₀) and higher bulk modulus of elasticity. There were evolutionary correlations between leaf life span and stem hydraulic efficiency, A_m, and dry season π₀. Xylem-cavitation resistance was evolutionarily correlated with stem hydraulic efficiency, D_h, as well as dry season π₀. Both wood density and leaf density were closely correlated with leaf water-stress tolerance and A_m.

Conclusions

The results reveal the clear distinctions in stem hydraulic traits and leaf water-stress tolerance between the co-occurring evergreen and deciduous angiosperm trees in an Asian dry karst forest. A novel pattern was demonstrated linking leaf longevity with stem hydraulic efficiency and leaf water-stress tolerance. The results show the correlated evolution in drought tolerance between stems and leaves.Key words: Tropical dry forest, karst, leaf habit, hydraulic conductivity, cavitation resistance, leaf water-stress tolerance, wood density, leaf density, phylogenetic independent contrasts     

Leaf age dependent changes in within-canopy variation in leaf functional traits: a meta-analysis

Within-canopy variation in leaf structural and photosynthetic characteristics is a major means by which whole canopy photosynthesis is maximized at given total canopy nitrogen. As key acclimatory modifications, leaf nitrogen content (N _A) and photosynthetic capacity (A _A) per unit area increase with increasing light availability in the canopy and these increases are associated with increases in leaf dry mass per unit area (M _A) and/or nitrogen content per dry mass and/or allocation. However, leaf functional characteristics change with increasing leaf age during leaf development and aging, but the importance of these alterations for within-canopy trait gradients is unknown. I conducted a meta-analysis based on 71 canopies that were sampled at different time periods or, in evergreens, included measurements for different-aged leaves to understand how within-canopy variations in leaf traits (trait plasticity) depend on leaf age. The analysis demonstrated that in evergreen woody species, M _A and N _A plasticity decreased with increasing leaf age, but the change in A _A plasticity was less suggesting a certain re-acclimation of A _A to altered light. In deciduous woody species, M _A and N _A gradients in flush-type species increased during leaf development and were almost invariable through the rest of the season, while in continuously leaf-forming species, the trait gradients increased constantly with increasing leaf age. In forbs, N _A plasticity increased, while in grasses, N _A plasticity decreased with increasing leaf age, reflecting life form differences in age-dependent changes in light availability and in nitrogen resorption for growth of generative organs. Although more work is needed to improve the coverage of age-dependent plasticity changes in some plant life forms, I argue that the age-dependent variation in trait plasticity uncovered in this study is large enough to warrant incorporation in simulations of canopy photosynthesis through the growing period.     

Contrasting correlation networks between leaf structure,nitrogen and chlorophyll in herbaceous and woody canopies

We studied acclimation patterns in leaf dry mass per area (M_A), nitrogen (N_A) and chlorophyll (ζ_A) content per area, and chlorophyll to nitrogen ratio (ζ/N) along vertical light gradients in natural temperate mixed herbaceous canopy and deciduous tree canopy. In the deciduous tree canopy, all leaves are formed at approximately the same time, and the light gradient during the rest of the growing season reflects the differences in light availability during leaf development, whereas in the herbaceous canopy, leaf production continues during most of the growing season and major changes in light conditions occur after leaf maturation. M_A and N_A increased strongly with increasing current light availability (I_D) in the tree canopy. In the herbaceous canopy, M_A and N_A were generally unrelated to I_D. Depending on species, the correlation between chlorophyll content per leaf area (ζ_A) and I_D was positive, negative, or non-significant. Path analyses revealed two opposite effects of I_D on the amount of leaf chlorophyll. In the tree canopy, increasing I_D enhanced ζ_A through changes in M_A and N_A, whereas the direct effect of light was negative in both canopies. The overall correlation network between foliage structural and chemical traits and the relationships with I_D were significantly stronger in the tree canopy, suggesting limited re-acclimation potential in the mixed herbaceous canopy. Within-species acclimation patterns reflected the patterns within the main functional types. These data demonstrate that the relationships of current light availability vs. leaf dry mass per area, leaf nitrogen and chlorophyll contents, and chlorophyll to nitrogen ratio differ among multi-species herbaceous canopies and deciduous tree canopies due to contrasting canopy development.     

What is the most prominent factor limiting photosynthesis in different layers of a greenhouse cucumber canopy?

Background and Aims

Maximizing photosynthesis at the canopy level is important for enhancing crop yield, and this requires insights into the limiting factors of photosynthesis. Using greenhouse cucumber (Cucumis sativus) as an example, this study provides a novel approach to quantify different components of photosynthetic limitations at the leaf level and to upscale these limitations to different canopy layers and the whole plant.

Methods

A static virtual three-dimensional canopy structure was constructed using digitized plant data in GroIMP. Light interception of the leaves was simulated by a ray-tracer and used to compute leaf photosynthesis. Different components of photosynthetic limitations, namely stomatal (S_L), mesophyll (M_L), biochemical (B_L) and light (L_L) limitations, were calculated by a quantitative limitation analysis of photosynthesis under different light regimes.

Key Results

In the virtual cucumber canopy, B_L and L_L were the most prominent factors limiting whole-plant photosynthesis. Diffusional limitations (S_L + M_L) contributed <15 % to total limitation. Photosynthesis in the lower canopy was more limited by the biochemical capacity, and the upper canopy was more sensitive to light than other canopy parts. Although leaves in the upper canopy received more light, their photosynthesis was more light restricted than in the leaves of the lower canopy, especially when the light condition above the canopy was poor. An increase in whole-plant photosynthesis under diffuse light did not result from an improvement of light use efficiency but from an increase in light interception. Diffuse light increased the photosynthesis of leaves that were directly shaded by other leaves in the canopy by up to 55 %.

Conclusions

Based on the results, maintaining biochemical capacity of the middle–lower canopy and increasing the leaf area of the upper canopy would be promising strategies to improve canopy photosynthesis in a high-wire cucumber cropping system. Further analyses using the approach described in this study can be expected to provide insights into the influences of horticultural practices on canopy photosynthesis and the design of optimal crop canopies.     

Optimizing nitrogen economy under drought: increased leaf nitrogen is an acclimation to water stress in willow (Salix spp.)

Background and Aims

The major objective was to identify plant traits functionally important for optimization of shoot growth and nitrogen (N) economy under drought. Although increased leaf N content (area basis) has been observed in dry environments and theory predicts increased leaf N to be an acclimation to drought, experimental evidence for the prediction is rare.

Methods

A pedigree of 200 full-sibling hybrid willows was pot-grown in a glasshouse in three replicate blocks and exposed to two water regimes for 3 weeks. Drought conditions were simulated as repeated periods of water shortage. The total leaf mass and area, leaf area efficiency (shoot growth per unit leaf area, E_A), area-based leaf N content (N_A), total leaf N pool (N_L) and leaf N efficiency (shoot growth per unit leaf N, E_N) were assessed.

Key Results

In the water-stress treatment, shoot biomass growth was N limited in the genotypes with low N_L, but increasingly limited by other factors in the genotypes with greatest N_L. The N_A was increased by drought, and drought-induced shift in N_A varied between genotypes (significant G × E). Judged from the E_A–N_A relationship, optimal N_A was 16 % higher in the water-stress compared with the well-watered treatment. Biomass allocation to leaves and shoots varied between treatments, but the treatment response of the leaf : shoot ratio was similar across all genotypes.

Conclusions

It is concluded that N-uptake efficiency and leaf N efficiency are important traits to improve growth under drought. Increased leaf N content (area basis) is an acclimation to optimize N economy under drought. The leaf N content is an interesting trait for breeding of willow bioenergy crops in a climate change future. In contrast, leaf biomass allocation is a less interesting breeding target to improve yield under drought.     

A model of canopy photosynthesis incorporating protein distribution through the canopy and its acclimation to light, temperature and CO2

Background and Aims

The distribution of photosynthetic enzymes, or nitrogen, through the canopy affects canopy photosynthesis, as well as plant quality and nitrogen demand. Most canopy photosynthesis models assume an exponential distribution of nitrogen, or protein, through the canopy, although this is rarely consistent with experimental observation. Previous optimization schemes to derive the nitrogen distribution through the canopy generally focus on the distribution of a fixed amount of total nitrogen, which fails to account for the variation in both the actual quantity of nitrogen in response to environmental conditions and the interaction of photosynthesis and respiration at similar levels of complexity.

Model

A model of canopy photosynthesis is presented for C₃ and C₄ canopies that considers a balanced approach between photosynthesis and respiration as well as plant carbon partitioning. Protein distribution is related to irradiance in the canopy by a flexible equation for which the exponential distribution is a special case. The model is designed to be simple to parameterize for crop, pasture and ecosystem studies. The amount and distribution of protein that maximizes canopy net photosynthesis is calculated.

Key Results

The optimum protein distribution is not exponential, but is quite linear near the top of the canopy, which is consistent with experimental observations. The overall concentration within the canopy is dependent on environmental conditions, including the distribution of direct and diffuse components of irradiance.

Conclusions

The widely used exponential distribution of nitrogen or protein through the canopy is generally inappropriate. The model derives the optimum distribution with characteristics that are consistent with observation, so overcoming limitations of using the exponential distribution. Although canopies may not always operate at an optimum, optimization analysis provides valuable insight into plant acclimation to environmental conditions. Protein distribution has implications for the prediction of carbon assimilation, plant quality and nitrogen demand.     

Nitrogen and water addition reduce leaf longevity of steppe species

Background and aims

Changes in supplies of resources will modify plant functional traits. However, few experimental studies have addressed the effects of nitrogen and water variations, either singly or in combination, on functional traits.

Methods

A 2-year field experiment was conducted to test the effects of nitrogen and water addition on leaf longevity and other functional traits of the two dominant (Agropyron cristatum and Stipa krylovii) and three most common species (Cleistogenes squarrosa, Melilotoides ruthenica and Potentilla tanacetifolia) in a temperate steppe in northern China.

Key Results

Additional nitrogen and water increased leaf nitrogen content and net photosynthetic rate, and changed other measured functional traits. Leaf longevity decreased significantly with both nitrogen addition (–6 days in 2007 and –5·4 days in 2008; both P < 0·001) and watering (–13 days in 2007 and –9·9 days in 2008; both P < 0·001), and significant differences in leaf longevity were also found among species. Nitrogen and water interacted to affect leaf longevity and other functional traits. Soil water content explained approx. 70 % of the shifts in leaf longevity. Biomass at both species and community level increased under water and nitrogen addition because of the increase in leaf biomass production per individual plant.

Conclusions

The results suggest that additional nitrogen and water supplies reduce plant leaf longevity. Soil water availability might play a fundamental role in determining leaf longevity and other leaf functional traits, and its effects can be modified by soil nitrogen availability in semi-arid areas. The different responses of species to resource alterations may cause different global change ramifications under future climate change scenarios.     

Optimal use of leaf nitrogen explains seasonal changes in leaf nitrogen content of an understorey evergreen shrub

Background and Aims

Understorey evergreen species commonly have a higher leaf nitrogen content in winter than in summer. Tested here is a hypothesis that such changes in leaf nitrogen content maximize nitrogen-use efficiency, defined as the daily carbon gain per unit nitrogen, under given temperature and irradiance levels.

Methods

The evergreen shrub Aucuba japonica growing naturally at three sites with different irradiance regimes in Japan was studied. Leaf photosynthetic characteristics, Rubisco and leaf nitrogen with measurements of temperature and irradiance monthly at each site were determined. Daily carbon gain was determined as a function of leaf nitrogen content to calculate the optimal leaf nitrogen content that maximized daily nitrogen-use efficiency.

Key Results

As is known, the optimal leaf nitrogen content increased with increasing irradiance. The optimal leaf nitrogen content also increased with decreasing temperature because the photosynthetic capacity per Rubisco decreased. Across sites and months, the optimal leaf nitrogen content was close to the actual leaf nitrogen content and explained the variation in actual leaf nitrogen by 64 %. Sensitivity analysis showed that the effect of temperature on optimal nitrogen content was similar in magnitude to that of irradiance.

Conclusions

Understorey evergreen species regulate leaf nitrogen content so as to maximize nitrogen-use efficiency in daily carbon gain under changing irradiance and temperature conditions.     

Feeding enhances photosynthetic efficiency in the carnivorous pitcher plant Nepenthes talangensis

Background and Aims

Cost–benefit models predict that carnivory can increase the rate of photosynthesis (A_N) by leaves of carnivorous plants as a result of increased nitrogen absorption from prey. However, the cost of carnivory includes decreased A_N and increased respiration rates (R_D) of trapping organs. The principal aim of the present study was to assess the costs and benefits of carnivory in the pitcher plant Nepenthes talangensis, leaves of which are composed of a lamina and a pitcher trap, in response to feeding with beetle larvae.

Methods

Pitchers of Nepenthes grown at 200 µmol m⁻² s⁻¹ photosynthetically active radiation (PAR) were fed with insect larvae for 2 months, and the effects on the photosynthetic processes were then assessed by simultaneous measurements of gas exchange and chlorophyll fluorescence of laminae and pitchers, which were correlated with nitrogen, carbon and total chlorophyll concentrations.

Key Results

A_N and maximum (F_v/F_m) and effective quantum yield of photosystem II (Φ_PSII) were greater in the fed than unfed laminae but not in the fed compared with unfed pitchers. Respiration rate was not significantly affected in fed compared with unfed plants. The unfed plants had greater non-photochemical quenching (NPQ) of chlorophyll fluorescence. Higher NPQ in unfed lamina did not compensate for their lower Φ_PSII, resulting in lower photochemical quenching (QP) and thus higher excitation pressure on PSII. Biomass and nitrogen and chlorophyll concentration also increased as a result of feeding. The cost of carnivory was shown by lower A_N and Φ_PSII in pitchers than in laminae, but R_D depended on whether it was expressed on a dry weight or a surface area basis. Correlation between nitrogen and A_N in the pitchers was not found. Cost–benefit analysis showed a large beneficial effect on photosynthesis from feeding as light intensity increased from 200 to 1000 µmol m⁻² s⁻¹ PAR after which it did not increase further. All fed plants began to flower.

Conclusion

Feeding pitchers with insect larvae increases A_N of leaf laminae, due to higher nutrient acquisition, with strong correlation with nitrogen concentration, but A_N of pitchers does not increase, despite increased nitrogen concentration in their tissue. Increased A_N improves growth and reproduction and is likely to increase the competitive advantage of carnivorous over non-carnivorous plants in nutrient-poor habitats.Key words: carnivorous plants, chlorophyll fluorescence, Nepenthes talangensis, nitrogen, pitcher plant, photosynthetic rate, photosystem II, respiration rate     

Leaf investment and light partitioning among leaves of different genotypes of the clonal plant Potentilla reptans in a dense stand after 5 years of competition

Background and Aims

While within-species competition for light is generally found to be asymmetric – larger plants absorbing more than proportional amounts of light – between-species competition tends to be more symmetric. Here, the light capture was analysed in a 5-year-old competition experiment that started with ten genotypes of the clonal plant Potentilla reptans. The following hypotheses were tested: (a) if different genotypes would do better in different layers of the canopy, thereby promoting coexistence, and (b) if leaves and genotypes with higher total mass captured more than proportional amounts of light, possibly explaining the observed dominance of the abundant genotypes.

Methods

In eight plots, 100 leaves were harvested at various depths in the canopy and their genotype determined to test for differences in leaf biomass allocation, leaf characteristics and the resulting light capture, calculated through a canopy model using the actual vertical light and leaf area profiles. Light capture was related to biomass to determine whether light competition between genotypes was asymmetric.

Key Results

All genotypes could reach the top of the canopy. The genotypes differed in morphology, but did not differ significantly in light capture per unit mass (Φ_mass) for leaves with the laminae placed at the same light levels. Light capture did increase disproportionately with leaf mass for all genotypes. However, the more abundant genotypes did not capture disproportionately more light relative to their mass than less-abundant genotypes.

Conclusions

Vertical niche differentiation in light acquisition does not appear to be a factor that could promote coexistence between these genotypes. Contrary to what is generally assumed, light competition among genetic individuals of the same species was size-symmetric, even if taller individual leaves did capture disproportionately more light. The observed shifts in genotype frequency cannot therefore be explained by asymmetric competition for light.Key words: Potentilla reptans, light, competition, symmetric, clonal, genotype, investment, petiole, canopy, allocation     

Exploring the spatial distribution of light interception and photosynthesis of canopies by means of a functional-structural plant model

Background and Aims

At present most process-based models and the majority of three-dimensional models include simplifications of plant architecture that can compromise the accuracy of light interception simulations and, accordingly, canopy photosynthesis. The aim of this paper is to analyse canopy heterogeneity of an explicitly described tomato canopy in relation to temporal dynamics of horizontal and vertical light distribution and photosynthesis under direct- and diffuse-light conditions.

Methods

Detailed measurements of canopy architecture, light interception and leaf photosynthesis were carried out on a tomato crop. These data were used for the development and calibration of a functional–structural tomato model. The model consisted of an architectural static virtual plant coupled with a nested radiosity model for light calculations and a leaf photosynthesis module. Different scenarios of horizontal and vertical distribution of light interception, incident light and photosynthesis were investigated under diffuse and direct light conditions.

Key Results

Simulated light interception showed a good correspondence to the measured values. Explicitly described leaf angles resulted in higher light interception in the middle of the plant canopy compared with fixed and ellipsoidal leaf-angle distribution models, although the total light interception remained the same. The fraction of light intercepted at a north–south orientation of rows differed from east–west orientation by 10 % on winter and 23 % on summer days. The horizontal distribution of photosynthesis differed significantly between the top, middle and lower canopy layer. Taking into account the vertical variation of leaf photosynthetic parameters in the canopy, led to approx. 8 % increase on simulated canopy photosynthesis.

Conclusions

Leaf angles of heterogeneous canopies should be explicitly described as they have a big impact both on light distribution and photosynthesis. Especially, the vertical variation of photosynthesis in canopy is such that the experimental approach of photosynthesis measurements for model parameterization should be revised.     

Characterization of three-dimensional spatial aggregation and association patterns of brown rot symptoms within intensively mapped sour cherry trees

Background and Aims

Characterization of spatial patterns of plant disease can provide insights into important epidemiological processes such as sources of inoculum, mechanisms of dissemination, and reproductive strategies of the pathogen population. Whilst two-dimensional patterns of disease (among plants within fields) have been studied extensively, there is limited information on three-dimensional patterns within individual plant canopies. Reported here are the detailed mapping of different symptom types of brown rot (caused by Monilinia laxa) in individual sour cherry tree (Prunus cerasus) canopies, and the application of spatial statistics to the resulting data points to determine patterns of symptom aggregation and association.

Methods

A magnetic digitizer was utilized to create detailed three-dimensional maps of three symptom types (blossom blight, shoot blight and twig canker) in eight sour cherry tree canopies during the green fruit stage of development. The resulting point patterns were analysed for aggregation (within a given symptom type) and pairwise association (between symptom types) using a three-dimensional extension of nearest-neighbour analysis.

Key Results

Symptoms of M. laxa infection were generally aggregated within the canopy volume, but there was no consistent pattern for one symptom type to be more or less aggregated than the other. Analysis of spatial association among symptom types indicated that previous year''s twig cankers may play an important role in influencing the spatial pattern of current year''s symptoms. This observation provides quantitative support for the epidemiological role of twig cankers as sources of primary inoculum within the tree.

Conclusions

Presented here is a new approach to quantify spatial patterns of plant disease in complex fruit tree canopies using point pattern analysis. This work provides a framework for quantitative analysis of three-dimensional spatial patterns within the finite tree canopy, applicable to many fields of research.     

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.     

Simplification of a light-based model for estimating final internode length in greenhouse cucumber canopies

Background and Aims

Light quantity and quality affect internode lengths in cucumber (Cucumis sativus), whereby leaf area and the optical properties of the leaves mainly control light quality within a cucumber plant community. This modelling study aimed at providing a simple, non-destructive method to predict final internode lengths (FILs) using light quantity and leaf area data.

Methods

Several simplifications of a light quantity and quality sensitive model for estimating FILs in cucumber have been tested. The direct simplifications substitute the term for the red : far-red (R : FR) ratios, by a term for (a) the leaf area index (LAI, m² m⁻²) or (b) partial LAI, the cumulative leaf area per m² ground, where leaf area per m² ground is accumulated from the top of each plant until a number, n, of leaves per plant is reached. The indirect simplifications estimate the input R : FR ratio based on partial leaf area and plant density.

Key Results

In all models, simulated FILs were in line with the measured FILs over various canopy architectures and light conditions, but the prediction quality varied. The indirect simplification based on leaf area of ten leaves revealed the best fit with measured data. Its prediction quality was even higher than of the original model.

Conclusions

This study showed that for vertically trained cucumber plants, leaf area data can substitute local light quality data for estimating FIL data. In unstressed canopies, leaf area over the upper ten ranks seems to represent the feedback of the growing architecture on internode elongation with respect to light quality. This highlights the role of this domain of leaves as the primary source for the specific R : FR signal controlling the final length of an internode and could therefore guide future research on up-scaling local processes to the crop level.     

Water relations and photosynthetic capacity of two species of Calotropis in a tropical semi-arid ecosystem

Background and Aims

Calotropis procera and Calotropis gigantea, originally from warm parts of Africa and Asia, are now pan-tropical and in ecological terms considered an indicator of overgrazed, disturbed lands; they grow successfully in dry areas. Variations in water relations, morphology and photosynthesis of the two species growing in the same habitat were studied to assess possible mechanisms of tolerance to drought and how these relate to their ecophysiological success. Also the hypothesis that their photosynthetic rate (A) under drought would be affected by stomatal and non-stomatal limitations was tested.

Methods

Water relations, gas exchange, water use efficiency (WUE), fluorescence parameters, pubescence and specific leaf area (SLA) of Calotropis procera and C. gigantea plants growing in the field were evaluated during the wet (WS) and dry (DS) seasons.

Results

The xylem water potential (ψ) was similar in both species during the WS and DS; drought caused a 28 % decrease of ψ. In C. procera, A, stomatal conductance (g_s) and carboxylation efficiency (CE) were higher in the WS with half the values of those during the DS, this species being more affected by drought than C. gigantea. A high δ¹³C of C. gigantea (–26·2 ‰) in the WS indicated a higher integrated WUE, in agreement with its lower g_s. Leaves of C. gigantea were more pubescent than C. procera. Relative stomatal and non-stomatal limitation of A increased with drought in both species; no changes in maximum quantum yield of photosystem II (PSII; F_v/F_m) were observed. The decrease in the relative quantum yield of PSII (φ_PSII) and in the photochemical quenching coefficient (q_P) was more pronounced in C. procera than in C. gigantea.

Conclusions

The photosynthetic capacity of C. procera was higher than that of C. gigantea. During the DS, A was regulated by stomatal and non-stomatal factors in a coordinated manner and drought did not cause chronic photoinhibition. A higher density of trichomes and leaf angle in C. gigantea may contribute to the maintenance of A and confer more efficient protection of photochemical activity in the DS. Ecophysiological traits such as high photosynthetic rate throughout the year even during the DS, and high WUE, highly pubescent leaves and low SLA observed in both species contribute to the establishment and growth of Calotropis in dry conditions.     

Quantitative characterization of clumping in Scots pine crowns

Background and Aims

Proper characterization of the clumped structure of forests is needed for calculation of the absorbed radiation and photosynthetic production by a canopy. This study examined the dependency of crown-level clumping on tree size and growth conditions in Scots pine (Pinus sylvestris), and determined the ability of statistical canopy radiation models to quantify the degree of self-shading within crowns as a result of the clumping effect.

Methods

Twelve 3-D Scots pine trees were generated using an application of the LIGNUM model, and the crown-level clumping as quantified by the crown silhouette to total needle area ratio (STAR_crown) was calculated. The results were compared with those produced by the stochastic approach of modelling tree crowns as geometric shapes filled with a random medium.

Key Results

Crown clumping was independent of tree height, needle area and growth conditions. The results supported the capability of the stochastic approach in characterizing clumping in crowns given that the outer shell of the tree crown is well represented.

Conclusions

Variation in the whole-stand clumping index is induced by differences in the spatial pattern of trees as a function of, for example, stand age rather than by changes in the degree of self-shading within individual crowns as they grow bigger.     

Effects of soil water and nitrogen on growth and photosynthetic response of Manchurian ash (Fraxinus mandshurica) seedlings in northeastern China

Background

Soil water and nitrogen (N) are considered to be the main environmental factors limiting plant growth and photosynthetic capacity. However, less is known about the interactive effects of soil water and N on tree growth and photosynthetic response in the temperate ecosystem.

Methods/Principal Findings

We applied N and water, alone and in combination, and investigated the combined effect of different water and N regimes on growth and photosynthetic responses of Fraxinus mandshurica seedlings. The seedlings were exposed to three water regimes including natural precipitation (CK), higher precipitation (HW) (CK +30%) and lower precipitation (LW) (CK −30%), and both with and without N addition for two growing seasons. We demonstrated that water and N supply led to a significant increase in the growth and biomass production of the seedlings. LW treatment significantly decreased biomass production and leaf N content, but they showed marked increases in N addition. N addition could enhance the photosynthetic capability under HW and CK conditions. Leaf chlorophyll content and the initial activity of Rubisco were dramatically increased by N addition regardless of soil water condition. The positive relationships were found between photosynthetic capacity, leaf N content, and SLA in response to water and N supply in the seedling. Rubisco expression was up-regulated by N addition with decreasing soil water content. Immunofluorescent staining showed that the labeling for Rubisco was relatively low in leaves of the seedlings under LW condition. The accumulation of Rubisco was increased in leaf tissues of LW by N addition.

Conclusions/Significance

Our study has presented better understanding of the interactions between soil water and N on the growth and photosynthetic response in F. mandschurica seedlings, which may provide novel insights on the potential responses of the forest ecosystem to climate change associated with increasing N deposition.     

Interspecific prediction of photosynthetic light response curves using specific leaf mass and leaf nitrogen content: effects of differences in soil fertility and growth irradiance

Background and Aims

Previous work has shown that the entire photosynthetic light response curve, based on both Mitscherlich and Michaelis–Menten functions, could be predicted in an interspecific context through allometric relations linking the parameters of these functions to two static leaf traits: leaf nitrogen (N) content and leaf mass per area (LMA). This paper describes to what extent these allometric relations are robust to changes in soil fertility and the growth irradiance of the plants.

Methods

Plants of 25 herbaceous species were grown under controlled conditions in factorial combinations of low/high soil fertility and low/high growth irradiance. Net photosynthetic rates per unit dry mass were measured at light intensities ranging from 0 to 700 µmol m⁻² s⁻¹ photosynthetically active radiation (PAR).

Key Results

The differing growth environments induced large changes in N, LMA and in each of the parameter estimates of the Mitscherlich and Michaelis–Menten functions. However, the differing growth environments induced only small (although significant) changes in the allometric relationships linking N and LMA to the parameters of the two functions. As a result, 88 % (Mitcherlich) and 89 % (Michaelis–Menten) of the observed net photosynthetic rates over the full range of light intensities (0–700 µmol m⁻² s⁻¹ PAR) and across all four growth environments could be predicted using only N and LMA using the same allometric relations.

Conclusions

These results suggest the possibility of predicting net photosynthetic rates in nature across species over the full range of light intensities using readily available data.     

Within-canopy variation in photosynthetic capacity,SLA and foliar N in temperate broad-leaved trees with contrasting shade tolerance

Key message

The relative shade tolerance of T. cordata , F. sylvatica , and C. betulus in mature stands is based on different species-specific carbon and nitrogen allocation patterns.

Abstract

The leaf morphology and photosynthetic capacity of trees are remarkably plastic in response to intra-canopy light gradients. While most studies examined seedlings, it is not well understood how plasticity differs in mature trees among species with contrasting shade tolerance. We studied light-saturated net photosynthesis (A _max), maximum carboxylation rate (V _cmax), electron transport capacity (J _max) and leaf dark respiration (R _d) along natural light gradients in the canopies of 26 adult trees of five broad-leaved tree species in a mixed temperate old-growth forest (Fraxinus excelsior, Acer pseudoplatanus, Carpinus betulus, Tilia cordata and Fagus sylvatica), representing a sequence from moderately light-demanding to highly shade-tolerant species. We searched for species differences in the dependence of photosynthetic capacity on relative irradiance (RI), specific leaf area (SLA) and nitrogen per leaf area (N _a ). The three shade-tolerant species (C. betulus, T. cordata, F. sylvatica) differed from the two more light-demanding species by the formation of shade leaves with particularly high SLA but relatively low N _a and consequently lower area-based A _max, and a generally higher leaf morphological and functional plasticity across the canopy. Sun leaf morphology and physiology were more similar among the two groups. The three shade-tolerant species differed in their shade acclimation strategies which are primarily determined by the species’ plasticity in SLA. Under low light, T. cordata and F. sylvatica increased SLA, mass-based foliar N and leaf size, while C. betulus increased solely SLA exhibiting only low intra-crown plasticity in leaf morphology and N allocation patterns. This study with mature trees adds to our understanding of tree species differences in shade acclimation strategies under the natural conditions of a mixed old-growth forest.