The key role of cell walls in setting mesophyll conductance to CO2 (gm) and, consequently, photosynthesis, is reviewed. First, the theoretical properties of cell walls that can affect gm are presented. Then, we focus on cell wall thickness (Tcw) reviewing empirical evidence showing that Tcw varies strongly among species and phylogenetic groups in a way that correlates with gm and photosynthesis i.e. the thicker the mesophyll cell walls, the lower gm and photosynthesis. Potential interplays of gm, Tcw, dehydration tolerance and hydraulic properties of leaves are also discussed. Dynamic variations of Tcw in response to the environment and their implications in the regulation of photosynthesis are discussed, and recent evidence suggesting an influence of cell wall composition on gm are presented. We then propose a hypothetical mechanism for the influence of cell walls on photosynthesis, combining the effects of thickness and composition, particularly pectins. Finally, we discuss the prospects for using biotechnology for enhancing photosynthesis by means of altering cell wall-related genes.

Under drought conditions, leaf photosynthesis is limited by the supply of CO2 . Drought induces production of abscisic acid (ABA), and ABA decreases stomatal conductance (gs ). Previous papers reported that the drought stress also causes the decrease in mesophyll conductance (gm ). However, the relationships between ABA content and gm are unclear. We investigated the responses of gm to the leaf ABA content [(ABA)L ] using an ABA-deficient mutant, aba1, and the wild type (WT) of Nicotiana plumbaginifolia. We also measured leaf water potential (ΨL ) because leaf hydraulics may be related to gm . Under drought conditions, gm decreased with the increase in (ABA)L in WT, whereas both (ABA)L and gm were unchanged by the drought treatment in aba1. Exogenously applied ABA decreased gm in both WT and aba1 in a dose-dependent manner. ΨL in WT was decreased by the drought treatment to -0.7 MPa, whereas ΨL in aba1 was around -0.8 MPa even under the well-watered conditions and unchanged by the drought treatment. From these results, we conclude that the increase in (ABA)L is crucial for the decrease in gm under drought conditions. We discuss possible relationships between the decrease in gm and changes in the leaf hydraulics.

Summary β-Cyclocitral (β-CC) is a volatile compound deriving from 1 O 2 oxidation of β-carotene in plant leaves. β-CC elicits a retrograde signal, modulating 1 O 2 -responsive genes and enhancing tolerance to photooxidative stress. Here, we show that β-CC is converted into water-soluble β-cyclocitric acid (β-CCA) in leaves. This metabolite is a signal that enhances plant tolerance to drought by a mechanism different from known responses such as stomatal closure, osmotic potential adjustment, and jasmonate signaling. This action of β-CCA is a conserved mechanism, being observed in various plant species, and it does not fully overlap with the β-CC-dependent signaling, indicating that β-CCA induces only a branch of β-CC signaling. Overexpressing SCARECROW-LIKE14 (SCL14, a regulator of xenobiotic detoxification) increased drought tolerance and potentiated the protective effect of β-CCA, showing the involvement of the SCL14-dependent detoxification in the phenomenon. β-CCA is a bioactive apocarotenoid that could potentially be used to protect crop plants against drought.

NRT1.1 is a putative nitrate sensor and is involved in many nitrate-dependent responses. On the other hand, a nitrate-independent function of NRT1.1 has been implied, but the clear-cut evidence is unknown. We found that NRT1.1 mutants showed enhanced tolerance to concentrated ammonium as sole N source in Arabidopsis thaliana. This unique phenotype was not observed in mutants of NLP7, which has been suggested to play a role in the nitrate-dependent signaling pathway. Our real-time PCR analysis, and evidence from a literature survey revealed that several genes relevant to the aliphatic glucosinolate-biosynthetic pathway were regulated via a nitrate-independent signal from NRT1.1. When taken together, the present study strongly suggests the existence of a nitrate-independent function of NRT1.1.

C3 photosynthesis is often limited by CO2 diffusivity or stomatal (gs) and mesophyll (gm) conductances. To characterize effects of stomatal closure induced by either high CO2 or abscisic acid (ABA) application on gm, we examined gs and gm in the wild type (Col‐0) and ost1 and slac1‐2 mutants of Arabidopsis thaliana grown at 390 or 780 μmol mol−1 CO2. Stomata of these mutants were reported to be insensitive to both high CO2 and ABA. When the ambient CO2 increased instantaneously, gm decreased in all these plants, whereas gs in ost1 and slac1‐2 was unchanged. Therefore, the decrease in gm in response to high CO2 occurred irrespective of the responses of gs. gm was mainly determined by the instantaneous CO2 concentration during the measurement and not markedly by the CO2 concentration during the growth. Exogenous application of ABA to Col‐0 caused the decrease in the intercellular CO2 concentration (Ci). With the decrease in Ci, gm did not increase but decreased, indicating that the response of gm to CO2 and that to ABA are differently regulated and that ABA content in the leaves plays an important role in the regulation of gm.

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Decreases in photosynthetic rate, stomatal conductance (gs), and mesophyll conductance (gm) are often observed under elevated CO2 conditions. However, which anatomical and/or physiological factors contribute to the decrease in gm is not fully understood. Arabidopsis thaliana wild-type and carbon-metabolism mutants (gwd1, pgm1, and cfbp1) with different accumulation patterns of non-structural carbohydrates were grown at ambient (400 ppm) and elevated (800 ppm) CO2. Anatomical and physiological traits of leaves were measured to investigate factors causing the changes in gm and in the mesophyll resistance (expressed as the reciprocal of mesophyll conductance per unit chloroplast surface area facing to intercellular space, Sc/gm). When grown at elevated CO2, all the lines showed increases in cell wall mass, cell wall thickness, and starch content, but not in leaf thickness. gm measured at 800 ppm CO2 was significantly lower than at 400 ppm CO2 in all the lines. Changes in Sc/gm were associated with thicker cell walls rather than with excess starch content. The results indicate that the changes in gm and Sc/gm that occur in response to elevated CO2 are independent of non-structural carbohydrates, and the cell wall represents a greater limitation factor for gm than starch.

Background Plants invest photosynthates in construction and maintenance of their structures and functions. Such investments are considered costs. These costs are recovered by CO2 assimilation rate (A) in the leaves and thus A is regarded as the immediate, short-term benefit. In photosynthesising leaves, CO2 diffusion from the air to the carboxylation site is hindered by several structural and biochemical barriers. CO2 diffusion from the intercellular air space to the chloroplast stroma is obstructed by the mesophyll resistance. The inverses is the mesophyll conductance (gm). Whether various plants realize an optimal gm, and how much investments are needed for relevant gm, remain unsolved. Scope This review examines relationships among leaf construction costs (CC), leaf maintenance costs (MC) and gm in various plants under diverse growth conditions. Through a literature survey, we demonstrate a strong linear relationship between leaf mass per area (LMA) and leaf CC. The overall correlation of CC versus gm across plant phylogenetic groups is weak, but significant trends are evident within specific groups and/or environments. Investment in CC is necessary for increase in LMA and mesophyll cell surface area (Smes). This allows the leaf to accommodate more chloroplasts, thus increasing A. However, increases in LMA and/or Smes often accompany other changes, such as cell wall thickening, which diminishes gm. Such factors that make the correlations of CC-gm elusive are identified. Conclusions For evaluation of contribution of gm to recover CC, leaf life-span is the key factor. The estimation of MC in relation to gm, especially in terms of costs required to regulate aquaporins, could be essential for efficient control of gm over the short-term. Over the long-term, costs are mainly reflected in CC, while benefits also include ultimate fitness attributes in terms of integrated carbon gain over the life of a leaf, plant survival and reproductive output.