Mesophyll Diffusion Conductance Research Articles

Functional responses of two Mediterranean pine species in an ozone Free-Air Controlled Exposure (FACE) experiment.

Effects of the phytotoxic and widespread ozone (O3) pollution may be species specific, but knowledge on Mediterranean conifer responses to long-term realistic exposure is still limited. We examined responses regarding to photosynthesis, needle biochemical stress markers and carbon and nitrogen (N) isotopes of two Mediterranean pine species (Pinus halepensis Mill. and Pinus pinea L.). Seedlings were grown in a Free-Air Controlled Exposure experiment with three levels of O3 (ambient air, AA [38.7p.p.b. as daily average]; 1.5×AA and 2.0×AA) during the growing season (May-October 2019). In P. halepensis, O3 caused a significant decrease in the photosynthetic rate, which was mainly due to a reduction of both stomatal and mesophyll diffusion conductance to CO2. Isotopic analyses indicated a cumulative or memory effect of O3 exposure on this species, as the negative effects were highlighted only in the late growing season in association with a reduced biochemical defense capacity. On the other hand, there was no clear effect of O3 on photosynthesis in P. pinea. However, this species showed enhanced N allocation to leaves to compensate for reduced photosynthetic N- use efficiency. We conclude that functional responses to O3 are different between the two species determining that P. halepensis with thin needles was relatively sensitive to O3, while P. pinea with thicker needles was more resistant due to a potentially low O3 load per unit mass of mesophyll cells, which may affect species-specific resilience in O3-polluted Mediterranean pine forests.

Vertical profile of photosynthetic CO<sub>2</sub> response within rice canopy

Leaf-specific Farquhar-von Caemmerer-Berry (FvCB) model was fitted to characterize the vertical profile of photosynthetic CO2 response within rice canopy. Leaf-position-specific and canopy average FvCB models were fitted to study a suitable leaf representing photosynthetic parameters at the canopy scale. The results showed that leaf photosynthesis was limited by Rubisco activity or ribulose-1,5-bisphosphate regeneration under field conditions. The maximum rate of carboxylation, maximum rate of electron transport, rate of triose phosphates utilization, and light respiration rate in the FvCB model reached the highest values for the top second leaf and then decreased, while the mesophyll diffusion conductance kept decreased in downward leaves. The integrated photosynthetic CO2-response curves for the top fourth and fifth leaves were appropriate for estimating parameters in the FvCB model at the canopy scale.

Mesophyll conductance exerts a significant limitation on photosynthesis during light induction.

Past studies have established mesophyll diffusion conductance to CO2 (gm ) as a variable and significant limitation to plant photosynthesis under steady-state conditions. However, the role of gm in influencing photosynthesis (A) during the transient period of light induction is largely unknown. We combined gas exchange measurements with laser-enabled carbon isotope discrimination measurements to assess gm during photosynthetic induction, using Arabidopsis as the measurement species. Our measurements revealed three key findings: (1) we found that the rate at which gm approached steady state during induction was not necessarily faster than the induction rate of the carboxylation process, contradictory to what has been suggested in previous studies; (2) gm displayed a strong and consistent coordination with A under both induction and steady-state settings, hinting that the mechanism driving gm -A coupling does not require physiological stability as a prerequisite; and (3) photosynthetic limitation analysis of our data revealed that when integrated over the entire induction period, the relative limitation of A imposed by gm can be as high as > 35%. The present study provides the first demonstration of the important role of gm in limiting CO2 assimilation during photosynthetic induction, thereby pointing to a need for more research attention to be devoted to gm in future induction studies.

A process-based coupled model of stomatal conductance-photosynthesis-transpiration during leaf ontogeny for water-saving irrigated rice.

Process-based coupled model of stomatal conductance-photosynthesis-transpiration was developed to estimate simultaneously stomatal conductance gsw, photosynthetic rate Pn, and transpiration rate Tr during leaf ontogeny. The modified Jarvis model was constructed by superposing the influence of leaf age LA on gsw in traditional Jarvis model. And the modified Farquhar model was constructed by incorporating the relationships of the LA with parameters in Farquhar model into traditional Farquhar model. The average and leaf-age-based coupled models were constructed, respectively, by combining traditional Farquhar and Penman-Monteith models with traditional Jarvis, and combining modified Farquhar and Penman-Monteith models with modified Jarvis. The results showed that the gsw, the maximum rate of carboxylation, maximum rate of electron transport, rate of triose phosphates utilization, and mitochondrial respiration rate varied in a positive skew pattern, while the mesophyll diffusion conductance decreased linearly with increase in LA. The average coupled model underestimated gsw, Pn, and Tr for young leaves and overestimated gsw, Pn, and Tr for old leaves. And the leaf-age-based coupled model generally perfected well in estimating gsw, Pn, and Tr for all leaves during leaf ontogeny. The study will provide basic information for either modeling leaf gsw, Pn, and Tr continuously, or upscaling them from leaf to canopy scale by considering the variation of LA within canopy.

Chloride as a macronutrient increases water-use efficiency by anatomically driven reduced stomatal conductance and increased mesophyll diffusion to CO2.

Chloride (Cl- ) has been recently described as a beneficial macronutrient, playing specific roles in promoting plant growth and water-use efficiency (WUE). However, it is still unclear how Cl- could be beneficial, especially in comparison with nitrate (NO3- ), an essential source of nitrogen that shares with Cl- similar physical and osmotic properties, as well as common transport mechanisms. In tobacco plants, macronutrient levels of Cl- specifically reduce stomatal conductance (gs ) without a concomitant reduction in the net photosynthesis rate (AN ). As stomata-mediated water loss through transpiration is inherent in the need of C3 plants to capture CO2 , simultaneous increase in photosynthesis and WUE is of great relevance to achieve a sustainable increase in C3 crop productivity. Our results showed that Cl- -mediated stimulation of larger leaf cells leads to a reduction in stomatal density, which in turn reduces gs and water consumption. Conversely, Cl- improves mesophyll diffusion conductance to CO2 (gm ) and photosynthetic performance due to a higher surface area of chloroplasts exposed to the intercellular airspace of mesophyll cells, possibly as a consequence of the stimulation of chloroplast biogenesis. A key finding of this study is the simultaneous improvement of AN and WUE due to macronutrient Cl- nutrition. This work identifies relevant and specific functions in which Cl- participates as a beneficial macronutrient for higher plants, uncovering a sustainable approach to improve crop yield.

Structural controls on photosynthetic capacity through juvenile‐to‐adult transition and needle ageing in Mediterranean pines

Abstract Needle photosynthetic potentials strongly vary among primary (juvenile) and secondary (adult) needles (heteroblasty) in Pinus species, but there is limited understanding of the underlying structural, diffusional and chemical controls. We studied differences in needle photosynthetic characteristics among current‐year juvenile and adult needles and among different‐aged adult needles in Mediterranean pines Pinus halepensis Mill., P. pinea L. and P. nigra J. F. Arnold subsp. salzmannii (Dunal) Franco, hypothesizing that needle anatomical modifications upon juvenile‐to‐adult transition lead to reduced photosynthetic capacity due to greater limitation of photosynthesis by mesophyll conductance and due to an increase in the share of support tissues at the expense of photosynthetic tissues. We also hypothesized that such alterations occur with needle ageing, but to a lower degree. Photosynthetic capacity per dry mass was 2.4‐ to 2.7‐fold higher in juvenile needles, and this was associated with 3.4‐ to 3.7‐fold greater mesophyll diffusion conductance, 2‐ to 2.5‐fold greater maximum carboxylase activity of Rubisco (Vcmax) and 2.2‐ to 3‐fold greater capacity for photosynthetic electron transport (Jmax). The latter differences were driven by modifications in mesophyll volume fraction and changes in the share of nitrogen between structural and photosynthetic functions. Analogous changes in photosynthetic characteristics occurred with needle ageing, but their extent was less. These results indicate that conifer foliage photosynthetic machinery undergoes a profound change from a fast return strategy in juveniles to slow return stress‐resistant strategy in adults and that this strategy shift is driven by modifications in foliage biomass investments in support and photosynthetic functions as well as by varying mesophyll diffusional controls on photosynthesis. Changes in needle morphophysiotype during tree and needle ageing need consideration in predicting changes in tree photosynthetic potentials through tree ontogeny and during and among growing seasons. A plain language summary is available for this article.

Temperature Variation under Continuous Light Restores Tomato Leaf Photosynthesis and Maintains the Diurnal Pattern in Stomatal Conductance.

The response of tomato plants (Solanum lycopersicum L. cv. Aromata) to continuous light (CL) in relation to photosynthesis, abscisic acid (ABA) and reactive oxygen species (ROS) was investigated to improve the understanding of the development and/or alleviation of CL-induced leaf injury in constant and diurnal temperature fluctuations with similar daily light integral and daily mean temperature. The plants were grown in three photoperiodic treatments for 15 days; One treatment with a 16/8 h light/dark period and a light/dark temperature of 27/17°C (Control), two CL treatments with 24 h photoperiods, one with a constant temperature of 24°C (CLCT) and the other one with variable temperature of 27/17°C for 16/8 ho, respectively (CLVT). A diurnal pattern of stomatal conductance (gs) and [ABA] was observed in the plants grown in the control and CLVT conditions, while the plants in CLCT conditions experienced a significant decrease in stomatal conductance aligned with an increase in ABA. The net photosynthesis (A) was significantly reduced in CLCT, aligned with a significant decrease in the maximum rate of Rubisco carboxylation (Vcmax), the maximum rate of electron transport (Jmax) and mesophyll diffusion conductance to CO2 (gm) in comparison to the control and CLVT. An increased production of H2O2 and O2•- linked with increased activities of antioxidative enzymes was seen in both CL treatments, but despite of this, leaf injuries were only observed in the CLCT treatment. The results suggest that the diurnal temperature fluctuations alleviated the CL injury symptoms, probably because the diurnal cycles of cellular mechanisms were maintained. The ROS were shown not to be directly involved in CL-induced leaf injury, since both ROS production and scavenging was highest in CLVT without leaf chlorotic symptoms.

Two tropical conifers show strong growth and water-use efficiency responses to altered CO2 concentration.

Conifers dominated wet lowland tropical forests 100million years ago (MYA). With a few exceptions in the Podocarpaceae and Araucariaceae, conifers are now absent from this biome. This shift to angiosperm dominance also coincided with a large decline in atmospheric CO2 concentration (ca). We compared growth and physiological performance of two lowland tropical angiosperms and conifers at ca levels representing pre-industrial (280 ppm), ambient (400 ppm) and Eocene (800 ppm) conditions to explore how differences in ca affect the growth and water-use efficiency (WUE) of seedlings from these groups. Two conifers (Araucaria heterophylla and Podocarpus guatemalensis) and two angiosperm trees (Tabebuia rosea and Chrysophyllum cainito) were grown in climate-controlled glasshouses in Panama. Growth, photosynthetic rates, nutrient uptake, and nutrient use and water-use efficiencies were measured. Podocarpus seedlings showed a stronger (66 %) increase in relative growth rate with increasing ca relative to Araucaria (19 %) and the angiosperms (no growth enhancement). The response of Podocarpus is consistent with expectations for species with conservative growth traits and low mesophyll diffusion conductance. While previous work has shown limited stomatal response of conifers to ca, we found that the two conifers had significantly greater increases in leaf and whole-plant WUE than the angiosperms, reflecting increased photosynthetic rate and reduced stomatal conductance. Foliar nitrogen isotope ratios (δ15N) and soil nitrate concentrations indicated a preference in Podocarpus for ammonium over nitrate, which may impact nitrogen uptake relative to nitrate assimilators under high ca SIGNIFICANCE: Podocarps colonized tropical forests after angiosperms achieved dominance and are now restricted to infertile soils. Although limited to a single species, our data suggest that higher ca may have been favourable for podocarp colonization of tropical South America 60MYA, while plasticity in photosynthetic capacity and WUE may help account for their continued persistence under large changes in ca since the Eocene.

Seasonal variations in photosynthesis, intrinsic water-use efficiency and stable isotope composition of poplar leaves in a short-rotation plantation.

Photosynthetic carbon assimilation and transpirational water loss play an important role in the yield and the carbon sequestration potential of bioenergy-devoted cultures of fast-growing trees. For six poplar (Populus) genotypes in a short-rotation plantation, we observed significant seasonal and genotypic variation in photosynthetic parameters, intrinsic water-use efficiency (WUEi) and leaf stable isotope composition (δ13C and δ18O). The poplars maintained high photosynthetic rates (between 17.8 and 26.9 μmol m−2 s−1 depending on genotypes) until late in the season, in line with their fast-growth habit. Seasonal fluctuations were mainly explained by variations in soil water availability and by stomatal limitation upon photosynthesis. Stomatal rather than biochemical limitation was confirmed by the constant intrinsic photosynthetic capacity (Vcmax) during the growing season, closely related to leaf nitrogen (N) content. Intrinsic water-use efficiency scaled negatively with carbon isotope discrimination (Δ13Cbl) and positively with the ratio between mesophyll diffusion conductance (gm) and stomatal conductance. The WUEi – Δ13Cbl relationship was partly influenced by gm. There was a trade-off between WUEi and photosynthetic N-use efficiency, but only when soil water availability was limiting. Our results suggest that seasonal fluctuations in relation to soil water availability should be accounted for in future modelling studies assessing the carbon sequestration potential and the water-use efficiency of woody energy crops.

Isoprenoid emissions, photosynthesis and mesophyll diffusion conductance in response to blue light

The effects of blue light (BL) on leaf gas exchange of Populus×canadensis, a strong isoprene emitter, and Quercus ilex and Citrus reticulata, two monoterpene emitters with respectively small and large storage pools for monoterpenes, were studied. Leaves were initially exposed to a saturating photosynthetic photon flux density (PPFD) of white light (WL), which was then progressively reduced to perform WL-response curves. Leaves acclimated to saturating WL were then quickly exposed to equivalent BL levels to perform BL-response curves. Blue light did not significantly affect photosynthetic parameters in the light-limited portion of the PPFD-response curves in both P.×canadensis and Q. ilex. Whereas photosynthesis (A), stomatal conductance (gs), and mesophyll conductance (gm) were significantly decreased at high PPFDs of BL. A was similarly inhibited by BL in C. reticulata, but there was no significant effect of light quality on gs. Overall these results show that the negative effect of BL on photosynthesis is widespread in tree species with different leaf characteristics, and that this involves coordinated reductions in gs and gm. BL negatively affected isoprene emission and, to a lesser extent monoterpene emissions, in concert with photosynthetic inhibition. Interesting, both isoprene and monoterpene emissions were shown to be inversely dependent upon intercellular [CO2]. These results indicate that a change in light spectral quality, which can vary during the day, between days and within seasons, can alter photosynthesis and isoprenoid emissions, depending on the PPFD intensity. Such effects should be strongly considered in photosynthesis and volatile isoprenoid emission models.

Importance of leaf anatomy in determining mesophyll diffusion conductance to CO2 across species: quantitative limitations and scaling up by models

Foliage photosynthetic and structural traits were studied in 15 species with a wide range of foliage anatomies to gain insight into the importance of key anatomical traits in the limitation of diffusion of CO2 from substomatal cavities to chloroplasts. The relative importance of different anatomical traits in constraining CO2 diffusion was evaluated using a quantitative model. Mesophyll conductance (g m) was most strongly correlated with chloroplast exposed surface to leaf area ratio (S c/S) and cell wall thickness (T cw), but, depending on foliage structure, the overall importance of g m in constraining photosynthesis and the importance of different anatomical traits in the restriction of CO2 diffusion varied. In species with mesophytic leaves, membrane permeabilities and cytosol and stromal conductance dominated the variation in g m. However, in species with sclerophytic leaves, g m was mostly limited by T cw. These results demonstrate the major role of anatomy in constraining mesophyll diffusion conductance and, consequently, in determining the variability in photosynthetic capacity among species.

Developmental changes in mesophyll diffusion conductance and photosynthetic capacity under different light and water availabilities in Populus tremula: how structure constrains function

Finite mesophyll diffusion conductance (g(m) ) significantly constrains net assimilation rate (A(n) ), but g(m) variations and variation sources in response to environmental stresses during leaf development are imperfectly known. The combined effects of light and water limitations on g(m) and diffusion limitations of photosynthesis were studied in saplings of Populus tremula L. An one-dimensional diffusion model was used to gain insight into the importance of key anatomical traits in determining g(m) . Leaf development was associated with increases in dry mass per unit area, thickness, density, exposed mesophyll (S(mes) /S) and chloroplast (S(c) /S) to leaf area ratio, internal air space (f(ias) ), cell wall thickness and chloroplast dimensions. Development of S(mes) /S and S(c) /S was delayed under low light. Reduction in light availability was associated with lower S(c) /S, but with larger f(ias) and chloroplast thickness. Water stress reduced S(c) /S and increased cell wall thickness under high light. In all treatments, g(m) and A(n) increased and CO(2) drawdown because of g(m) , C(i) -C(c) , decreased with increasing leaf age. Low light and drought resulted in reduced g(m) and A(n) and increased C(i) -C(c) . These results emphasize the importance of g(m) and its components in determining A(n) variations during leaf development and in response to stress.

Effects of potassium supply on limitations of photosynthesis by mesophyll diffusion conductance in Carya cathayensis

Potassium (K) influences the photosynthesis process in a number of ways; however, the mechanisms underlying the photosynthetic response to differences in K supply are not well understood. Concurrent measurements of gas exchange and chlorophyll fluorescence were made to investigate the effect of K nutrition on photosynthetic efficiency and mesophyll conductance (g(m)) in hickory seedlings (Carya cathayensis Sarg.) in a greenhouse. The results show that leaf K concentrations < 0.7-0.8% appeared to limit the leaf net CO2 assimilation rate (A), and that the relative limitation of photosynthesis due to g(m) and stomatal conductance (g(s)) decreased with increasing supplies of K. However, a sensitivity analysis indicated that A was most sensitive to the maximum carboxylation rate of Rubisco (V(c,max)) and the maximum rate of electron transport (J(max)). These results indicate that the photosynthetic rate is primarily limited by the biochemical processes of photosynthesis (V(c,max) and J(max)), rather than by g(m) and g(s) in K-deficient plants. Additionally, g(m) was closely correlated with g(s) and the leaf dry mass per unit area (M(A)) in hickory seedlings, which indicates that decreased g(m) and g(s) may be a consequence of leaf anatomical adaptation.

Role of mesophyll diffusion conductance in constraining potential photosynthetic productivity in the field

Limited mesophyll diffusion conductance to CO(2) (g(m)) can significantly constrain plant photosynthesis, but the extent of g(m)-limitation is still imperfectly known. As g(m) scales positively with foliage photosynthetic capacity (A), the CO(2) drawdown from substomatal cavities (C(i)) to chloroplasts (C(C), C(i)-C(C)=A/g(m)) rather than g(m) alone characterizes the mesophyll diffusion limitations of photosynthesis. The dependencies of g(m) on A, foliage structure (leaf dry mass per unit area, M(A)), and the resulting drawdowns across a dataset of 81 species of contrasting foliage structure and photosynthetic potentials measured under non-stressed conditions were analysed to describe the structure-driven potential photosynthetic limitations due to g(m). Further the effects of key environmental stress factors and leaf and plant developmental alterations on g(m) and CO(2) drawdown were evaluated and the implications of varying g(m) on foliage photosynthesis in the field were simulated. The meta-analysis demonstrated that g(m) of non-stressed leaves was negatively correlated with M(A), and despite the positive relationship between g(m) and A, the CO(2) drawdown was larger in leaves with more robust structure. The correlations were stronger with mass-based g(m) and A, probably reflecting the circumstance that mesophyll diffusion is a complex three-dimensional process that scales better with mesophyll volume-weighted than with leaf area-weighted traits. The analysis of key environmental stress effects on g(m) and CO(2) drawdowns demonstrated that the effect of individual stresses on CO(2) drawdowns varies depending on the stress effects on foliage structure and assimilation rates. Leaf diffusion limitations are larger in non-senescent older leaves and also in senescent leaves, again reflecting more robust leaf structure and/or non-co-ordinated alterations in leaf photosynthesis and g(m). According to simulation analyses, in plants with a larger part of the overall diffusion conductance from the ambient atmosphere to the chloroplasts in the mesophyll, photosynthesis is less sensitive to changes in stomatal conductance. Accordingly, in harsher environments that support vegetation with tougher long-living stress-tolerant leaves with lower g(m), reductions in stomatal conductance that are common during stress periods are expected to alter photosynthesis less than in species where a larger part of the total diffusion limitation is determined by stomata. While structural robustness improves plant performance under environmental stress, low g(m) and inherently large CO(2) drawdown in robust leaves limits the photosynthesis of these plants more severely under favourable conditions when stomatal conductance is high. The differences in overall responsiveness to environmental modifications of plants with varying g(m) need consideration in current large-scale ecosystem productivity models.

Importance of mesophyll diffusion conductance in estimation of plant photosynthesis in the field

Mesophyll diffusion conductance to CO(2) (g(m)) is an important leaf characteristic determining the drawdown of CO(2) from substomatal cavities (C(i)) to chloroplasts (C(C)). Finite g(m) results in modifications in the shape of the net assimilation (A) versus C(i) response curves, with the final outcome of reduced maximal carboxylase activity of Rubisco (V(cmax)), and a greater ratio of the capacity for photosynthetic electron transport to V(cmax) (J(max)/V(cmax)) and alterations in mitochondrial respiration rate (R(d)) when estimated from A/C(i) responses without considering g(m). The influence of different Farquhar et al. model parameterizations on daily photosynthesis under non-stressed (C(i) kept constant throughout the day) and stressed conditions (mid-day reduction in C(i)) was compared. The model was parameterized on the basis of A/C(C) curves and A/C(i) curves using both the conventional fitting procedure (V(cmax) and R(d) fitted separately to the linear part of the response curve and J(max) to the saturating part) and a procedure that fitted all parameters simultaneously. The analyses demonstrated that A/C(i) parameterizations overestimated daily assimilation by 6-8% for high g(m) values, while they underestimated if by up to 70% for low g(m) values. Qualitative differences between the A/C(i) and A/C(C) parameterizations were observed under stressed conditions, when underestimated V(cmax) and overestimated R(d) of A/C(i) parameterizations resulted in excessive mid-day depression of photosynthesis. Comparison with measured diurnal assimilation rates in the Mediterranean sclerophyll species Quercus ilex under drought further supported this bias of A/C(i) parameterizations. While A/C(i) parameterization predicted negative carbon balance at mid-day, actual measurements and simulations with the A/C(C) approach yielded positive carbon gain under these conditions. In addition, overall variation captured by the best A/C(i) parameterization was poor compared with the A/C(C) approach. This analysis strongly suggests that for correct parameterization of daily time-courses of photosynthesis under realistic field conditions, g(m) must be included in photosynthesis models.

Leaf mesophyll diffusion conductance in 35 Australian sclerophylls covering a broad range of foliage structural and physiological variation

Foliage structure, chemistry, photosynthetic potentials (V(cmax) and J(max)), and mesophyll diffusion conductance (g(m)) were quantified for 35 broad-leaved species from four sites with contrasting rainfall and soil fertility in eastern Australia. The aim of the study was to estimate the extent to which g(m) and related leaf properties limited photosynthesis (A), focusing on highly sclerophyllous species typical of the 'slow-return' end of the leaf economics spectrum. Leaf dry mass per unit area (M(A)) varied approximately 5-fold, leaf life span (L(L)) and N (N(M)) and P (P(M)) contents per dry mass approximately 8-fold, and various characteristics of foliage photosynthetic machinery 6- to 12-fold across the data set. As is characteristic of the 'leaf economics spectrum', more robust leaves with greater M(A) and longevity were associated with lower nutrient contents and lower foliage photosynthetic potentials. g(m) was positively correlated with V(cmax) and J(max), and these correlations were stronger on a mass basis. Only g(m)/mass was negatively associated with M(A). CO(2) drawdown from substomatal cavities to chloroplasts (C(i)-C(C)) characterizing mesophyll CO(2) diffusion limitations was larger in leaves with greater M(A), lower g(m)/mass, and lower photosynthetic potentials. Relative limitation of A due to finite mesophyll diffusion conductance, i.e. 1-A(infinite g(m))/A(actual g(m)), was always >0.2 and up to 0.5 in leaves with most robust leaf structure, demonstrating the profound effect of finite g(m) on realized photosynthesis rates. Data from different sites were overlapping in bivariate relationships, and the variability of average values between the sites was less than among the species within the sites. Nevertheless, photosynthesis was more strongly limited by g(m) in low rain/high nutrient and high rain/low nutrient sites that supported vegetation with more sclerophyllous foliage. These data collectively highlight a strong relationship between leaf structure and g(m), and demonstrate that realized photosynthesis rates are strongly limited by g(m) in this highly sclerophyllous flora.

Leaf internal diffusion conductance limits photosynthesis more strongly in older leaves of Mediterranean evergreen broad‐leaved species

ABSTRACTLeaf age‐dependent changes in structure, nitrogen content, internal mesophyll diffusion conductance (gm), the capacity for photosynthetic electron transport (Jmax) and the maximum carboxylase activity of Rubisco (Vcmax) were investigated in mature non‐senescent leaves of Laurus nobilis L., Olea europea L. and Quercus ilex L. to test the hypothesis that the relative significance of biochemical and diffusion limitations of photosynthesis changes with leaf age. The leaf life‐span was up to 3 years in L. nobilis and O. europea and 6 years in Q. ilex. Increases in leaf age resulted in enhanced leaf dry mass per unit area (MA), larger leaf dry to fresh mass ratio, and lower nitrogen contents per dry mass (NM) in all species, and lower nitrogen contents per area (NA) in L. nobilis and Q. ilex. Older leaves had lower gm, Jmax and Vcmax. Due to the age‐dependent increase in MA, mass‐based gm, Jmax and Vcmax declined more strongly (7‐ to 10‐fold) with age than area‐based (5‐ to 7‐fold) characteristics. Diffusion conductance was positively associated with foliage photosynthetic potentials. However, this correlation was curvilinear, leading to lower ratio of chloroplastic to internal CO2 concentration (Cc/Ci) and larger drawdown of CO2 from leaf internal air space to chloroplasts (ΔC) in older leaves with lower gm. Overall the age‐dependent decreases in photosynthetic potentials were associated with decreases in NM and in the fraction of N in photosynthetic proteins, whereas decreases in gm were associated with increases in MA and the fraction of cell walls. These age‐dependent modifications altered the functional scaling of foliage photosynthetic potentials with MA, NM, and NA. The species primarily differed in the rate of age‐dependent modifications in foliage structural and functional characteristics, but also in the degree of age‐dependent changes in various variables. Stomatal openness was weakly associated with leaf age, but due to species differences in stomatal openness, the distribution of total diffusion limitation between stomata and mesophyll varied among species. These data collectively demonstrate that in Mediterranean evergreens, structural limitations of photosynthesis strongly interact with biochemical limitations. Age‐dependent changes in gm and photosynthetic capacities do not occur in a co‐ordinated manner in these species such that mesophyll diffusion constraints curb photosynthesis more in older than in younger leaves.

Theoretical Considerations when Estimating the Mesophyll Conductance to CO2 Flux by Analysis of the Response of Photosynthesis to CO2

The conductance for CO(2) diffusion in the mesophyll of leaves can limit photosynthesis. We have studied two methods for determining the mesophyll conductance to CO(2) diffusion in leaves. We generated an ideal set of photosynthesis rates over a range of partial pressures of CO(2) in the stroma and studied the effect of altering the mesophyll diffusion conductance on the measured response of photosynthesis to intercellular CO(2) partial pressure. We used the ideal data set to test the sensitivity of the two methods to small errors in the parameters used to determine mesophyll conductance. The two methods were also used to determine mesophyll conductance of several leaves using measured rather than ideal data sets. It is concluded that both methods can be used to determine mesophyll conductance and each method has particular strengths. We believe both methods will prove useful in the future.