Partial root-zone drying irrigation improves intrinsic water-use efficiency and maintains high photosynthesis by uncoupling stomatal and mesophyll conductance in cotton leaves.

Partial root-zone drying irrigation (PRD) can improve water-use efficiency (WUE) without reductions in photosynthesis; however, the mechanism by which this is attained is unclear. To amend that, PRD conditions were simulated by polyethylene glycol 6000 in a root-splitting system and the effects of PRD on cotton growth were studied. Results showed that PRD decreased stomatal conductance (gs) but increased mesophyll conductance (gm). Due to the contrasting effects on gs and gm, net photosynthetic rate (AN) remained unaffected, while the enhanced gm/gs ratio facilitated a larger intrinsic WUE. Further analyses indicated that PRD-induced reduction of gs was related to decreased stomatal size and stomatal pore area in adaxial and abaxial surface which was ascribed to lower pore length and width. PRD-induced variation of gm was ascribed to the reduced liquid-phase resistance, due to increases in chloroplast area facing to intercellular airspaces and the ratio of chloroplast surface area to total mesophyll cell area exposed to intercellular airspaces, as well as to decreases in the distance between cell wall and chloroplast, and between adjacent chloroplasts. The above results demonstrate that PRD, through alterations to stomatal and mesophyll structures, decoupled gs and gm responses, which ultimately increased intrinsic WUE and maintained AN.

Effect of light-induced changes in leaf anatomy on intercellular and cellular components of mesophyll resistance for CO2 in Fagus sylvatica.

Mesophyll resistance for CO2 diffusion (rm) is one of the main limitations for photosynthesis and plant growth. Breeding new varieties with lower rm requires knowledge of its distinct components. We tested new method for estimating the relative drawdowns of CO2 concentration (c) across hypostomatous leaves of Fagus sylvatica. This technique yields values of the ratio of the internal CO2 concentrations at the adaxial and abaxial leaf side, cd/cb, the drawdown in the intercellular air space (IAS), and intracellular drawdown between IAS and chloroplast stroma, cc/cbd. The method is based on carbon isotope composition of leaf dry matter and epicuticular wax isolated from upper and lower leaf sides. We investigated leaves from tree-canopy profile to analyse the effects of light and leaf anatomy on the drawdowns and partitioning of rm into its inter- (rIAS) and intracellular (rliq) components. Validity of the new method was tested by independent measurements of rm using conventional isotopic and gas exchange techniques. 73% of investigated leaves had adaxial epicuticular wax enriched in 13C compared to abaxial wax (by 0.50‰ on average), yielding 0.98 and 0.70 for average of cd/cb and cc/cbd, respectively. The rIAS to rliq proportion were 5.5:94.5% in sun-exposed and 14.8:85.2% in shaded leaves. cc dropped to less than half of the atmospheric value in the sunlit and to about two-thirds of it in shaded leaves. This method shows that rIAS is minor but not negligible part of rm and reflects leaf anatomy traits, i.e. leaf mass per area and thickness.

Plasticity of mesophyll cell density and cell wall thickness and composition play a pivotal role in regulating plant growth and photosynthesis under shading in rapeseed.

BACKGROUND AND AIMS: Plasticity of leaf growth and photosynthesis is an important strategy of plants to adapt to shading stress; however, their strategy of leaf development to achieve a simultaneous increase in leaf area and photosynthesis under shading remains unknown. METHODS: In the present study, a pot experiment was conducted using three rapeseed genotypes of Huayouza 50 (HYZ50), Zhongshuang 11 (ZS11) and Huayouza 62 (HYZ62), and the responses of plant growth, leaf morphoanatomical traits, cell wall composition and photosynthesis to shading were investigated. KEY RESULTS: Shading significantly increased leaf area per plant (LAplant) in all genotypes, but the increase in HYZ62 was greater than that in HYZ50 and ZS11. The greater increment of LAplant in HYZ62 was related to the larger decrease in leaf mass per area (LMA) and leaf density (LD), which were in turn related to less densely packed mesophyll cells and thinner cell walls (Tcw). Moreover, shading significantly increased photosynthesis in HYZ62 but significantly decreased it in HYZ50. The enhanced photosynthesis in HYZ62 was related to increased mesophyll conductance (gm) due primarily to thinner cell walls. CONCLUSIONS: The data presented indicate that the different plasticity of mesophyll cell density, cell wall thickness and cell wall composition in response to shading can dramatically affect leaf growth and photosynthesis.

Cell wall thickness has phylogenetically consistent effects on the photosynthetic nitrogen-use efficiency of terrestrial plants.

Leaf photosynthetic nitrogen-use efficiency (PNUE) diversified significantly among C3 species. To date, the morpho-physiological mechanisms and interrelationships shaping PNUE on an evolutionary time scale remain unclear. In this study, we assembled a comprehensive matrix of leaf morpho-anatomical and physiological traits for 679 C3 species, ranging from bryophytes to angiosperms, to comprehend the complexity of interrelationships underpinning PNUE variations. We discovered that leaf mass per area (LMA), mesophyll cell wall thickness (Tcwm ), Rubisco N allocation fraction (PR ), and mesophyll conductance (gm ) together explained 83% of PNUE variations, with PR and gm accounting for 65% of those variations. However, the PR effects were species-dependent on gm , meaning the contribution of PR on PNUE was substantially significant in high-gm species compared to low-gm species. Standard major axis (SMA) and path analyses revealed a weak correlation between PNUE and LMA (r2 = 0.1), while the SMA correlation for PNUE-Tcwm was robust (r2 = 0.61). PR was inversely related to Tcwm , paralleling the relationship between gm and Tcwm , resulting in the internal CO2 drawdown being only weakly proportional to Tcwm . The coordination of PR and gm in relation to Tcwm constrains PNUE during the course of evolution.

Stomatal, mesophyll and biochemical limitations to photosynthesis and their relationship with leaf structure over an elevation gradient in two conifers.

Photosynthetic responses across complex elevational gradients provides insight into fundamental processes driving responses of plant growth and net primary production to environmental change. Gas exchange of needles and twig water potential were measured in two widespread coniferous tree species, Pinus contorta and Picea engelmannii, over an 800-m elevation gradient in southeastern Wyoming, USA. We hypothesized that limitations to photosynthesis imposed by mesophyll conductance (gm) would be greatest at the highest elevation sites due to higher leaf mass per area (LMA) and that estimations of maximum rate of carboxylation (Vcmax) without including gm would obscure elevational patterns of photosynthetic capacity. We found that gm decreased with elevation for P. contorta and remained constant for P. engelmannii, but in general, limitation to photosynthesis by gm was small. Indeed, estimations of Vcmax when including gm were equivalent to those estimated without including gm and no correlation was found between gm and LMA nor between gm and leaf N. Stomatal conductance (gs) and biochemical demand for CO2 were by far the most limiting processes to photosynthesis at all sites along the elevation gradient. Photosynthetic capacity (A) and gs were influenced strongly by differences in soil water availability across the elevation transect, while gm was less responsive to water availability. Based on our analysis, variation in gm plays only a minor role in driving patterns of photosynthesis in P. contorta and P. engelmannii across complex elevational gradients in dry, continental environments of the Rocky Mountains and accurate modeling of photosynthesis, growth and net primary production in these forests may not require detailed estimation of this trait value.

Variations in leaf anatomical characteristics drive the decrease of mesophyll conductance in poplar under elevated ozone.

Decline in mesophyll conductance (gm ) plays a key role in limiting photosynthesis in plants exposed to elevated ozone (O3 ). Leaf anatomical traits are known to influence gm , but the potential effects of O3 -induced changes in leaf anatomy on gm have not yet been clarified. Here, two poplar clones were exposed to elevated O3 . The effects of O3 on the photosynthetic capacity and anatomical characteristics were assessed to investigate the leaf anatomical properties that potentially affect gm . We also conducted global meta-analysis to explore the general response patterns of gm and leaf anatomy to O3 exposure. We found that the O3 -induced reduction in gm was critical in limiting leaf photosynthesis. Changes in liquid-phase conductance rather than gas-phase conductance drive the decline in gm under elevated O3, and this effect was associated with thicker cell walls and smaller chloroplast sizes. The effects of O3 on palisade and spongy mesophyll cell traits and their contributions to gm were highly genotype-dependent. Our results suggest that, while anatomical adjustments under elevated O3 may contribute to defense against O3 stress, they also cause declines in gm and photosynthesis. These results provide the first evidence of anatomical constraints on gm under elevated O3 .

Interspecific variation in the temperature response of mesophyll conductance is related to leaf anatomy.

Although mesophyll conductance (gm ) is known to be sensitive to temperature (T), the mechanisms underlying the temperature response of gm are not fully understood. In particular, it has yet to be established whether interspecific variation in gm -T relationships is associated with mesophyll anatomy and vein traits. In the present study, we measured the short-term response of gm in eight crop species, and leaf water potential (Ψleaf ) in five crop species over a temperature range of 15-35°C. The considered structural parameters are surface areas of mesophyll cells and chloroplasts facing intercellular airspaces per unit leaf area (Sm and Sc ), cell wall thickness (Tcw ), and vein length per area (VLA). We detected large interspecific variations in the temperature responses of gm and Ψleaf . The activation energy for gm (Ea,gm ) was found to be positively correlated with Sc , although it showed no correlation with Tcw . In contrast, VLA was positively correlated with the slope of the linear model of Ψleaf -T (a), whereas Ea,gm was marginally correlated with VLA and a. A two-component model was subsequently used to model gm -T relationships, and the mechanisms underlying the temperature response of gm are discussed. The data presented here indicate that leaf anatomy is a major determinant of the interspecific variation in gm -T relationships.

Structural and functional leaf diversity lead to variability in photosynthetic capacity across a range of Juglans regia genotypes.

Similar to other cropping systems, few walnut cultivars are used as scion in commercial production. Germplasm collections can be used to diversify cultivar options and hold potential for improving crop productivity, disease resistance and stress tolerance. In this study, we explored the anatomical and biochemical bases of photosynthetic capacity and response to water stress in 11 Juglans regia accessions in the U.S. department of agriculture, agricultural research service (USDA-ARS) National Clonal Germplasm. Net assimilation rate (An ) differed significantly among accessions and was greater in lower latitudes coincident with higher stomatal and mesophyll conductances, leaf thickness, mesophyll porosity, gas-phase diffusion, leaf nitrogen and lower leaf mass and stomatal density. High CO2 -saturated assimilation rates led to increases in An under diffusional and biochemical limitations. Greater An was found in lower-latitude accessions native to climates with more frost-free days, greater precipitation seasonality and lower temperature seasonality. As expected, water stress consistently impaired photosynthesis with the highest % reductions in lower-latitude accessions (A3, A5 and A9), which had the highest An under well-watered conditions. However, An for A3 and A5 remained among the highest under dehydration. J. regia accessions, which have leaf structural traits and biochemistry that enhance photosynthesis, could be used as commercial scions or breeding parents to enhance productivity.

Cell-level anatomy explains leaf age-dependent declines in mesophyll conductance and photosynthetic capacity in the evergreen Mediterranean oak Quercus ilex subsp. rotundifolia.

Leaves of Mediterranean evergreen tree species experience a reduction in net CO2 assimilation (AN) and mesophyll conductance to CO2 (gm) during aging and senescence, which would be influenced by changes in leaf anatomical traits at cell level. Anatomical modifications can be accompanied by the dismantling of photosynthetic apparatus associated to leaf senescence, manifested through changes at the biochemical level (i.e., lower nitrogen investment in photosynthetic machinery). However, the role of changes in leaf anatomy at cell level and nitrogen content in gm and AN decline experienced by old non-senescent leaves of evergreen trees with long leaf lifespan is far from being elucidated. We evaluated age-dependent changes in morphological, anatomical, chemical and photosynthetic traits in Quercus ilex subsp. rotundifolia Lam., an evergreen oak with high leaf longevity. All photosynthetic traits decreased with increasing leaf age. The relative change in cell wall thickness (Tcw) was less than in chloroplast surface area exposed to intercellular air space (Sc/S), and Sc/S was a key anatomical trait explaining variations in gm and AN among different age classes. The reduction of Sc/S was related to ultrastructural changes in chloroplasts associated to leaf aging, with a concomitant reduction in cytoplasmic nitrogen. Changes in leaf anatomy and biochemistry were responsible for the age-dependent modifications in gm and AN. These findings revealed a gradual physiological deterioration related to the dismantling of the photosynthetic apparatus in older leaves of Q. ilex subsp. rotundifolia.

The response of mesophyll conductance to ozone-induced oxidative stress is genotype-dependent in poplar.

The CO2 diffusion conductance within the leaf mesophyll (gm) is considered a major limiting factor of photosynthesis. However, the effects of the major secondary air pollutant ozone (O3) on gm have been poorly investigated. Eight genotypes of the economically important tree species Populus × canadensis Moench were exposed to 120 ppb O3 for 21 d. gm showed a genotype-dependent response to O3-induced oxidative stress and was a major limiting factor of net assimilation rate (Anet), ahead of stomatal conductance to CO2 (gsc) and of the maximum carboxylation capacity of the Rubisco enzyme (Vcmax) in half of the tested genotypes. Increased leaf dry mass per area (LMA) and decreased chlorophyll content were linked to the observed gm decrease, but this relationship did not entirely explain the different genotypic gm responses. Moreover, the oxidative stress defence metabolites ascorbate and glutathione were not related to O3 tolerance of gm. However, malondialdehyde probably mitigated the observed gm decrease in some genotypes due to its oxidative stress signalling function. The large variation of gm suggests different regulation mechanisms amongst poplar genotypes under oxidative stress.

Structural organization of the spongy mesophyll.

Many plant leaves have two layers of photosynthetic tissue: the palisade and spongy mesophyll. Whereas palisade mesophyll consists of tightly packed columnar cells, the structure of spongy mesophyll is not well characterized and often treated as a random assemblage of irregularly shaped cells. Using micro-computed tomography imaging, topological analysis, and a comparative physiological framework, we examined the structure of the spongy mesophyll in 40 species from 30 genera with laminar leaves and reticulate venation. A spectrum of spongy mesophyll diversity encompassed two dominant phenotypes: first, an ordered, honeycomblike tissue structure that emerged from the spatial coordination of multilobed cells, conforming to the physical principles of Euler's law; and second, a less-ordered, isotropic network of cells. Phenotypic variation was associated with transitions in cell size, cell packing density, mesophyll surface-area-to-volume ratio, vein density, and maximum photosynthetic rate. These results show that simple principles may govern the organization and scaling of the spongy mesophyll in many plants and demonstrate the presence of structural patterns associated with leaf function. This improved understanding of mesophyll anatomy provides new opportunities for spatially explicit analyses of leaf development, physiology, and biomechanics.

New insight into pectin effects on mesophyll conductance in four species of deciduous Forest.

The amount of pectin in the cell wall is considered a critical element regulating cell wall porosity. Thus, it is likely to influence the diffusional processes particularly that from the substomatal cavities to the carboxylation sites in the chloroplast stroma (i.e. mesophyll conductance, gm). We analyzed in four deciduous species (Acer campestre, Crateaugus monogyna, Corylus avellana, Quercus robur) the correlation between pectin and gm in two phase of leaves development (i.e. mature green leaves and senescent leaves). Results showed that in A. campestre, C. monogyna and Q. robur to higher value of pectin in mature green leaves corresponded the higher gm value. Taking into account this result, we can assume that a higher amount of pectins is associated to an increased cell wall hydrophilicity and elasticity, thus increasing gm because CO2 molecules cross the wall dissolved in water. An opposite behavior was observed in C. avellana.

Expression of a CO2-permeable aquaporin enhances mesophyll conductance in the C4 species Setaria viridis.

A fundamental limitation of photosynthetic carbon fixation is the availability of CO2. In C4 plants, primary carboxylation occurs in mesophyll cytosol, and little is known about the role of CO2 diffusion in facilitating C4 photosynthesis. We have examined the expression, localization, and functional role of selected plasma membrane intrinsic aquaporins (PIPs) from Setaria italica (foxtail millet) and discovered that SiPIP2;7 is CO2-permeable. When ectopically expressed in mesophyll cells of Setaria viridis (green foxtail), SiPIP2;7 was localized to the plasma membrane and caused no marked changes in leaf biochemistry. Gas exchange and C18O16O discrimination measurements revealed that targeted expression of SiPIP2;7 enhanced the conductance to CO2 diffusion from the intercellular airspace to the mesophyll cytosol. Our results demonstrate that mesophyll conductance limits C4 photosynthesis at low pCO2 and that SiPIP2;7 is a functional CO2 permeable aquaporin that can improve CO2 diffusion at the airspace/mesophyll interface and enhance C4 photosynthesis.

Coordination of leaf hydraulic, anatomical, and economical traits in tomato seedlings acclimation to long-term drought.

BACKGROUND: Leaf hydraulic and economics traits are critical for balancing plant water and CO2 exchange, and their relationship has been widely studied. Leaf anatomical traits determine the efficiency of CO2 diffusion within mesophyll structure. However, it remains unclear whether leaf anatomical traits are associated with leaf hydraulic and economics traits acclimation to long-term drought. RESULTS: To address this knowledge gap, eight hydraulic traits, including stomatal and venation structures, four economics traits, including leaf dry mass per area (LMA) and the ratio between palisade and spongy mesophyll thickness (PT/ST), and four anatomical traits related to CO2 diffusion were measured in tomato seedlings under the long-term drought conditions. Redundancy analysis indicated that the long-term drought decreased stomatal conductance (gs) mainly due to a synchronized reduction in hydraulic structure such as leaf hydraulic conductance (Kleaf) and major vein width. Simultaneously, stomatal aperture on the adaxial surface and minor vein density (VDminor) also contributed a lot to this reduction. The decreases in mesophyll thickness (Tmes) and chlorophyll surface area exposed to leaf intercellular air spaces (Sc/S) were primarily responsible for the decline of mesophyll conductance (gm) thereby affecting photosynthesis. Drought increased leaf density (LD) thus limited CO2 diffusion. In addition, LMA may not be important in regulating gm in tomato under drought. Principal component analysis revealed that main anatomical traits such as Tmes and Sc/S were positively correlated to Kleaf, VDminor and leaf thickness (LT), while negatively associated with PT/ST. CONCLUSIONS: These findings indicated that leaf anatomy plays an important role in maintaining the balance between water supply and CO2 diffusion responses to drought. There was a strong coordination between leaf hydraulic, anatomical, and economical traits in tomato seedlings acclimation to long-term drought.

TGD5 is required for normal morphogenesis of non-mesophyll plastids, but not mesophyll chloroplasts, in Arabidopsis.

Stromules are dynamic membrane-bound tubular structures that emanate from plastids. Stromule formation is triggered in response to various stresses and during plant development, suggesting that stromules may have physiological and developmental roles in these processes. Despite the possible biological importance of stromules and their prevalence in green plants, their exact roles and formation mechanisms remain unclear. To explore these issues, we obtained Arabidopsis thaliana mutants with excess stromule formation in the leaf epidermis by microscopy-based screening. Here, we characterized one of these mutants, stromule biogenesis altered 1 (suba1). suba1 forms plastids with severely altered morphology in a variety of non-mesophyll tissues, such as leaf epidermis, hypocotyl epidermis, floral tissues, and pollen grains, but apparently normal leaf mesophyll chloroplasts. The suba1 mutation causes impaired chloroplast pigmentation and altered chloroplast ultrastructure in stomatal guard cells, as well as the aberrant accumulation of lipid droplets and their autophagic engulfment by the vacuole. The causal defective gene in suba1 is TRIGALACTOSYLDIACYLGLYCEROL5 (TGD5), which encodes a protein putatively involved in the endoplasmic reticulum (ER)-to-plastid lipid trafficking required for the ER pathway of thylakoid lipid assembly. These findings suggest that a non-mesophyll-specific mechanism maintains plastid morphology. The distinct mechanisms maintaining plastid morphology in mesophyll versus non-mesophyll plastids might be attributable, at least in part, to the differential contributions of the plastidial and ER pathways of lipid metabolism between mesophyll and non-mesophyll plastids.

Mitochondrial movement during its association with chloroplasts in Arabidopsis thaliana.

Plant mitochondria move dynamically inside cells and this movement is classified into two types: directional movement, in which mitochondria travel long distances, and wiggling, in which mitochondria travel short distances. However, the underlying mechanisms and roles of both types of mitochondrial movement, especially wiggling, remain to be determined. Here, we used confocal laser-scanning microscopy to quantitatively characterize mitochondrial movement (rate and trajectory) in Arabidopsis thaliana mesophyll cells. Directional movement leading to long-distance migration occurred at high speed with a low angle-change rate, whereas wiggling leading to short-distance migration occurred at low speed with a high angle-change rate. The mean square displacement (MSD) analysis could separate these two movements. Directional movement was dependent on filamentous actin (F-actin), whereas mitochondrial wiggling was not, but slightly influenced by F-actin. In mesophyll cells, mitochondria could migrate by wiggling, and most of these mitochondria associated with chloroplasts. Thus, mitochondria migrate via F-actin-independent wiggling under the influence of F-actin during their association with chloroplasts in Arabidopsis.

Stomatal, mesophyll conductance, and biochemical limitations to photosynthesis during induction.

The dynamics of leaf photosynthesis in fluctuating light affects carbon gain by plants. Mesophyll conductance (gm) limits CO2 assimilation rate (A) under the steady state, but the extent of this limitation under non-steady-state conditions is unknown. In the present study, we aimed to characterize the dynamics of gm and the limitations to A imposed by gas diffusional and biochemical processes under fluctuating light. The induction responses of A, stomatal conductance (gs), gm, and the maximum rate of RuBP carboxylation (Vcmax) or electron transport (J) were investigated in Arabidopsis (Arabidopsis thaliana (L.)) and tobacco (Nicotiana tabacum L.). We first characterized gm induction after a change from darkness to light. Each limitation to A imposed by gm, gs and Vcmax or J was significant during induction, indicating that gas diffusional and biochemical processes limit photosynthesis. Initially, gs imposed the greatest limitation to A, showing the slowest response under high light after long and short periods of darkness, assuming RuBP-carboxylation limitation. However, if RuBP-regeneration limitation was assumed, then J imposed the greatest limitation. gm did not vary much following short interruptions to light. The limitation to A imposed by gm was the smallest of all the limitations for most of the induction phase. This suggests that altering induction kinetics of mesophyll conductance would have little impact on A following a change in light. To enhance the carbon gain by plants under naturally dynamic light environments, attention should therefore be focused on faster stomatal opening or activation of electron transport.

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.

Coleataenia prionitis, a C4-like species in the Poaceae.

C4-like plants represent the penultimate stage of evolution from C3 to C4 plants. Although Coleataenia prionitis (formerly Panicum prionitis) has been described as a C4 plant, its leaf anatomy and gas exchange traits suggest that it may be a C4-like plant. Here, we reexamined the leaf structure and biochemical and physiological traits of photosynthesis in this grass. The large vascular bundles were surrounded by two layers of bundle sheath (BS): a colorless outer BS and a chloroplast-rich inner BS. Small vascular bundles, which generally had a single BS layer with various vascular structures, also occurred throughout the mesophyll together with BS cells not associated with vascular tissue. The mesophyll cells did not show a radial arrangement typical of Kranz anatomy. These features suggest that the leaf anatomy of C. prionitis is on the evolutionary pathway to a complete C4 Kranz type. Phosphoenolpyruvate carboxylase (PEPC) and pyruvate, Pi dikinase occurred in the mesophyll and outer BS. Glycine decarboxylase was confined to the inner BS. Ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) accumulated in the mesophyll and both BSs. C. prionitis had biochemical traits of NADP-malic enzyme type, whereas its gas exchange traits were close to those of C4-like intermediate plants rather than C4 plants. A gas exchange study with a PEPC inhibitor suggested that Rubisco in the mesophyll could fix atmospheric CO2. These data demonstrate that C. prionitis is not a true C4 plant but should be considered as a C4-like plant.

Anatomically induced changes in rice leaf mesophyll conductance explain the variation in photosynthetic nitrogen use efficiency under contrasting nitrogen supply.

BACKGROUND: The ratio of CO2 mesophyll conductance (gm) to Ribulose-1, 5-bisphosphate carboxylase/oxygenase (Rubisco) content has been suggested to positively affect photosynthetic nitrogen use efficiency (PNUE). The anatomical basis of gm has been quantified, but information on the relationship between cell-level anatomies and PNUE is less advanced. Here, hydroponic experiments were conducted in rice plants supplied with ammonium (NH4+) and nitrate (NO3-) under three N levels (low, 0.71 mM; intermediate, 2.86 mM; high, 7.14 mM) to investigate the gas exchange parameters, leaf anatomical structure and PNUE. RESULTS: The results showed a lower PNUE in plants supplied with high nitrogen and NH4+, which was positively correlated with the gm/Rubisco ratio. A one-dimensional within-leaf model revealed that the resistance to CO2 diffusion in the liquid phase (rliq) dominated the overall mesophyll resistance (rm), in which CO2 transfer resistance in the cell wall, cytoplasm and stroma were significantly affected by nitrogen supply. The chloroplast surface area exposed to intercellular space (Sc) per Rubisco rather than the gm/Sc ratio was positively correlated with PNUE and was thus considered a key component influencing PNUE. CONCLUSION: In conclusion, our study emphasized that Sc was the most important anatomical trait in coordinating gm and PNUE with contrasting N supply.