Peter E.L. van der Putten

Using combined measurements of gas exchange and chlorophyll fluorescence to estimate parameters of a biochemical C3 photosynthesis model: a critical appraisal and a new integrated approach applied to leaves in a wheat (Triticum aestivum) canopy

We appraised the literature and described an approach to estimate the parameters of the Farquhar, von Caemmerer and Berry model using measured CO(2) assimilation rate (A) and photosystem II (PSII) electron transport efficiency (Phi(2)). The approach uses curve fitting to data of A and Phi(2) at various levels of incident irradiance (I(inc)), intercellular CO(2) (C(i)) and O(2). Estimated parameters include day respiration (R(d)), conversion efficiency of I(inc) into linear electron transport of PSII under limiting light [kappa(2(LL))], electron transport capacity (J(max)), curvature factor (theta) for the non-rectangular hyperbolic response of electron flux to I(inc), ribulose 1.5-bisphosphate carboxylase/oxygenase (Rubisco) CO(2)/O(2) specificity (S(c/o)), Rubisco carboxylation capacity (V(cmax)), rate of triose phosphate utilization (T(p)) and mesophyll conductance (g(m)). The method is used to analyse combined gas exchange and chlorophyll fluorescence measurements on leaves of various ages and positions in wheat plants grown at two nitrogen levels. Estimated S(c/o) (25 degrees C) was 3.13 mbar microbar(-1); R(d) was lower than respiration in the dark; J(max) was lower and theta was higher at 2% than at 21% O(2); kappa(2(LL)), V(cmax), J(max) and T(p) correlated to leaf nitrogen content; and g(m) decreased with increasing C(i) and with decreasing I(inc). Based on the parameter estimates, we surmised that there was some alternative electron transport.

Using a biochemical C4 photosynthesis model and combined gas exchange and chlorophyll fluorescence measurements to estimate bundle-sheath conductance of maize leaves differing in age and nitrogen content

Bundle-sheath conductance (g(bs) ) affects CO(2) leakiness, and, therefore, the efficiency of the CO(2) -concentrating mechanism (CCM) in C(4) photosynthesis. Whether and how g(bs) varies with leaf age and nitrogen status is virtually unknown. We used a C(4) -photosynthesis model to estimate g(bs) , based on combined measurements of gas exchange and chlorophyll fluorescence on fully expanded leaves of three different ages of maize (Zea mays L.) plants grown under two contrasting nitrogen levels. Nitrogen was replenished weekly to maintain leaf nitrogen content (LNC) at a similar level across the three leaf ages. The estimated g(bs) values on leaf-area basis ranged from 1.4 to 10.3 mmol m(-2) s(-1) and were affected more by LNC than by leaf age, although g(bs) tended to decrease as leaves became older. When converted to resistance (r(bs) = 1/g(bs)), r(bs) decreased monotonically with LNC. The correlation was presumably associated with nitrogen effects on leaf anatomy such as on wall thickness of bundle-sheath cells. Despite higher g(bs), meaning less efficient CCM, the calculated loss due to photorespiration was still low for high-nitrogen leaves. Under the condition of ambient CO(2) and saturating irradiance, photorespiratory loss accounted for 3-5% of fixed carbon for the high-nitrogen, versus 1-2% for the low-nitrogen, leaves.

Early competition shapes maize whole-plant development in mixed stands

Mixed cropping is practised widely in developing countries and is gaining increasing interest for sustainable agriculture in developed countries. Plants in intercrops grow differently from plants in single crops, due to interspecific plant interactions, but adaptive plant morphological responses to competition in mixed stands have not been studied in detail. Here the maize (Zea mays) response to mixed cultivation with wheat (Triticum aestivum) is described. Evidence is provided that early responses of maize to the modified light environment in mixed stands propagate throughout maize development, resulting in different phenotypes compared with pure stands. Photosynthetically active radiation (PAR), red:far-red ratio (R:FR), leaf development, and final organ sizes of maize grown in three cultivation systems were compared: pure maize, an intercrop with a small distance (25cm) between maize and wheat plants, and an intercop with a large distance (44cm) between the maize and the wheat. Compared with maize in pure stands, maize in the mixed stands had lower leaf and collar appearance rates, increased blade and sheath lengths at low ranks and smaller sizes at high ranks, increased blade elongation duration, and decreased R:FR and PAR at the plant base during early development. Effects were strongest in the treatment with a short distance between wheat and maize strips. The data suggest a feedback between leaf initiation and leaf emergence at the plant level and coordination between blade and sheath growth at the phytomer level. A conceptual model, based on coordination rules, is proposed to explain the development of the maize plant in pure and mixed stands.

Zinc allocation and re-allocation in rice

Aims: Agronomy and breeding actively search for options to enhance cereal grain Zn density. Quantifying internal (re-)allocation of Zn as affected by soil and crop management or genotype is crucial. We present experiments supporting the development of a conceptual model of whole plant Zn allocation and re-allocation in rice. Methods: Two solution culture experiments using 70Zn applications at different times during crop development and an experiment on within-grain distribution of Zn are reported. In addition, results from two earlier published experiments are re-analyzed and re-interpreted. Results: A budget analysis showed that plant zinc accumulation during grain filling was larger than zinc allocation to the grains. Isotope data showed that zinc taken up during grain filling was only partly transported directly to the grains and partly allocated to the leaves. Zinc taken up during grain filling and allocated to the leaves replaced zinc re-allocated from leaves to grains. Within the grains, no major transport barrier was observed between vascular tissue and endosperm. At low tissue Zn concentrations, rice plants maintained concentrations of about 20 mg Zn kg−1 dry matter in leaf blades and reproductive tissues, but let Zn concentrations in stems, sheath, and roots drop below this level. When plant zinc concentrations increased, Zn levels in leaf blades and reproductive tissues only showed a moderate increase while Zn levels in stems, roots, and sheaths increased much more and in that order. Conclusions: In rice, the major barrier to enhanced zinc allocation towards grains is between stem and reproductive tissues. Enhancing root to shoot transfer will not contribute proportionally to grain zinc enhancement.

Temperature response of bundle-sheath conductance in maize leaves

Highlight Bundle-sheath conductance, g bs , is commonly assumed to be independent of temperature. We report that temperature response of maize g bs followed a peaked or non-peaked Arrhenius equation, triggering further investigations on this response.

Modelling the relationship between CO2 assimilation and leaf anatomical properties in tomato leaves

The CO2 concentration near Rubisco and, therefore, the rate of CO2 assimilation, is influenced by both leaf anatomical factors and biochemical processes. Leaf anatomical structures act as physical barriers for CO2 transport. Biochemical processes add or remove CO2 along its diffusion pathway through mesophyll. We combined a model that quantifies the diffusive resistance for CO2 using anatomical properties, a model that partitions this resistance and an extended version of the Farquhar-von Caemmerer-Berry model. We parametrized the model by gas exchange, chlorophyll fluorescence and leaf anatomical measurements from three tomato cultivars. There was generally a good agreement between the predicted and measured light and CO2 response curves. We did a sensitivity analysis to assess how the rate of CO2 assimilation responds to changes in various leaf anatomical properties. Next, we conducted a similar analysis for assumed diffusive properties and curvature factors. Some variables (diffusion pathway length in stroma, diffusion coefficient of the stroma, curvature factors) substantially affected the predicted CO2 assimilation. We recommend more research on the measurements of these variables and on the development of 2-D and 3-D gas diffusion models, since these do not require the diffusion pathway length in the stroma as predefined parameter.

Intercropping with wheat lowers nutrient uptake and biomass accumulation of maize, but increases photosynthetic rate of the ear leaf

Maize and wheat are globally important food crops. The two species can be grown as an intercrop, with substantial land sparing in the order of 20 %, as expressed by a land equivalent ratio of ~1.2. Here, we study nutrient uptake and the photosynthesis rate of intercropped maize and show that nutrient uptake is constrained by competition with wheat, while the photosynthesis rate is not decreased, but—surprisingly—increased. Ecophysiological mechanisms potentially underlying the unexpected high photosynthesis rate in intercropped maize are discussed.

Calibration matters : On the procedure of using the chlorophyll fluorescence method to estimate mesophyll conductance

Estimates of mesophyll conductance (gm), when calculated from chlorophyll fluorescence, are uncertain, especially when the photosystem II (PSII) operating efficiency is measured from the traditional single saturation pulse methodology. The multiphase flash method has recently been recommended to replace the single saturation pulse method, allowing a more reliable estimation of gm. Also, many researchers still directly use the PSII operating efficiency to derive linear electron transport rate J (that is required to estimate gm), without appropriate calibration using measurements under non-photorespiratory conditions. Here we demonstrate for tomato and rice that (i) using the multiphase flash method did not yield realistic estimates of gm if no calibration was conducted; and (ii) using the single saturation pulse method still gave reasonable estimates of gm when calibration based on the non-photorespiratory measurements was properly conducted. Therefore, conducting calibration based on data under non-photorespiratory conditions was indispensable for a reliable estimation of gm, regardless whether the multiphase flash or the single saturation pulse method was used for measuring the PSII operating efficiency. Other issues related to the procedure of using the chlorophyll fluorescence method to estimate gm were discussed.