Water use efficiency as a function of leaf age and position within the cotton canopy

The gas exchange properties of whole plant canopies are an integral part of crop productivity and have attracted much attention in recent years. However, insufficient information exists on the coordination of transpiration and CO₂ uptake for individual leaves during the growing season. Single-leaf determinations of net photosynthesis (Pn), transpiration (E) and water use efficiency (WUE) for field-grown cotton (Gossypium hirsutum L.) leaves were recorded during a 2-year field study. Measurements were made at 3 to 4 day intervals on the main-stem and first three sympodial leaves at main-stem node 10 from their unfolding through senescence. Results indicated that all gas exchange parameters changed with individual main-stem and sympodial leaf age. Values of Pn, E and WUE followed a rise and fall pattern with maximum rates achieved at a leaf age of 18 to 20 days. While no significant position effects were observed for Pn, main-stem and sympodial leaves did differ in E and WUE particularly as leaves aged beyond 40 days. For a given leaf age, the main-stem leaf had a significantly lower WUE than the three sympodial leaves. WUE's for the main-stem and three sympodial leaves between the ages of 41 to 50 days were 0.85, 1.30, 1.36 and 1.95 μmol CO₂ mmol⁻¹ H₂O, respectively. The mechanisms which mediated leaf positional differences for WUE were not strictly related to changes in stomatal conductance (g_s·H₂O) since decreases in g_s·H₂O with leaf age were similar for the four leaves. However, significantly different radiant environments with distance along the fruiting branch did indicate the possible involvement of mutual leaf shading in determining WUE. The significance of these findings are presented in relation to light competition within the plant canopy during development.     

A pump/leak/buffer model for plant nitrate uptake

A simple simulation model is described to account for the rates at which plants take up nitrate and reduce it to protein. It is based on the pump and leak principle, with the pump working at a constant rate per unit sap volume provided that there is an adequate concentration of nitrate at the root surface. The rate of leakage is assumed to be proportional to the concentration difference between the inside and the outside of the plant. Nitrogen is removed from the plant nitrate pool (the buffer) at a constant fraction of the photosynthesis rate. When applied to data for the diurnal variation in nitrate uptake by ryegrass, the model predicts an uptake pattern similar to that actually observed, with a time lag of about 5 hours between photosynthesis and uptake.     

Photoinhibition at chilling temperature

The effects of moderate light at chilling temperature on the photosynthesis of unhardened (acclimated to +18° C) and hardened (cold-acclimated) spinach (Spinacea oleracea L.) leaves were studied by means of fluorescence-induction measurements at 20° C and 77K and by determination of quantum yield of O₂ evolution. Exposure to 550 mol photons·m^-2·s^-1 at +4° C induced a strong photoinhibition in the unhardened leaves within a few hours. Photoinhibition manifested by a decline in quantum yield was characterized by an increase in initial fluorescence (F _o) and a decrease in variable fluorescence (F _v) and in the ratio of variable to maximum fluorescence (F _V/F _M), both at 77K and 20° C. The decline in quantum yield was more closely related to the decrease in the F _V/F _M ratio measured at 20° C, as compared with F _V/F _M at 77K. Quenching of the variable fluorescence of photosystem II was accompanied by a decline in photosystem-I fluorescence at 77K, indicating increased thermal de-excitation of pigments as the main consequence of the light treatment. All these changes detected in fluorescence parameters as well as in the quantum yield of O₂ evolution were fully reversible within 1–3 h at a higher temperature in low light. The fast recovery led us to the view that this photoinhibition represents a regulatory mechanism protecting the photosynthetic apparatus from the adverse effects of excess light by increasing thermal energy dissipation. Long-term cold acclimation probably enforces other protective mechanisms, as the hardened leaves were insensitive to the same light treatment that induced strong inhibition of photosynthesis in unhardened leaves.Abbreviations F ₀ initial fluorescence - F _M maximum fluorescence - F _V variable fluorescence (F _M-F ₀ - PFD photon flux density - PS photosystem     

Fruit photosynthesis

Abstract. In addition to photosynthesis as in the leaf, fruit possess a system which refixes CO₂ from the mitochondrial respiration of predominantly imported carbon. This pathway produces malate by the action of phosphoenolpyruvate carboxylase, PEPC, (E.C. 4.1.1.31) and appears to be regulated primarily by the cytosolic concentration of HCO₃/CO₂ and malate. Malate is stored in the vacuole as malic acid, constituting a major carbon pool and a potential substrate for respiration. The PEPC in apple fruit proves to be an efficient form of the enzyme with low Michaelis constants, i.e. Km = 0.09 mol m^-3 PEP and 0.2 mol m^–3 HCO₃, and large Ki= 110 mol m^-3 HCO³. In fleshy fruit, chlorophyll and chloroplasts are unevenly distributed; they resemble the C₃ sun-type and arc concentrated in the perivascular tissue, with smaller chloroplasts, fewer grana per chloroplast and a larger degree of vacuolation than commonly found in a leaf of the same species. Fruit photosynthesis often compensates for respiratory CO₂ loss in the light. However, due to respiration in the dark, CO₂ loss is in excess of photosynthetic gain in the light, such that a continual loss of CO₂ was observed in the diurnal cycle and which is maintained throughout fruit development. The rate of CO₂ exchange decreases on a fresh weight or surface basis, but increases with fruit ontogeny on a per fruit basis, causing accumulation of several percent CO₂ in the internal cavity. Stomata are present in the outer epidermis of those fruits examined, but with a 10-to 100-fold lesser frequency than in the abaxial epidermis of leaf of the same species. The number of Stomata is set at anthesis and remained constant, while the stomatal frequency decreases as the fruit surface expands. Stomata are as sensitive as in leaves in the early stages of fruit development, but often are transformed into lenticels during fruit ontogeny, thereby decreasing the permeability of the outer epidermis. The discrepancy between the CO₂-concentrating mechanism provided by PEPC analogous to C₄/CAM Photosynthesis and the kinetics of fruit PEPC, characteristic of C₃/non-autotrophic tissue, suggests the definition of a new type of ‘fruit photosynthesis’ rather than its categorization within an existing type.     

Expression of C4-like photosynthesis in several species of Flaveria

Abstract Photosynthetic metabolism was investigated in leaves of five species of Flaveria (Asteraceac), all previously considered to be C₄ plants. Leaves were exposed to ¹⁴CO₂ for different intervals up to 16s. Extrapolation of ¹⁴C-product curves to zero time indicated that only F. trinervia and F.bidentis assimilated atmospheric CO₂ exclusively through phosphoenolpyruvate carboxylase. The proportion of direct fixation of ¹⁴CO₂ by ribulose-I, 5-bisphosphate carboxylase/oxygenase (Rubisco) ranged from 5 to 10% in leaves of F. australasica. F. palmeri and F. vaginata. Protoplasts of leaf mesophyll and bundle sheath cells were utilized to examine the intercellular compartmentation of principal photosynthetic enzymes. Leaves of F. australasica, F. palmeri and F. vaginata contained 5 to 7% of the leaf's Rubisco activity in the mesophyll cells, while leaves of F. trinervia and F. bidentis contained at most 0.2 to 0.8% of such activity in their mesophyll cells. Thus, F. trinervia and F. bidentis have the complete C₄ syndrome, while F. australasica, F. palmeri and F. vaginata are less advanced, C₄-like species.     

Measurement of the extent of electron transfer to the bacteriopheophytin in the M-subunit in reaction centers of Rhodopseudomonas viridis

We have measured the extent of flash-induced electron transfer from the bacteriochlorophyll dimer, P, to the bacteriopheophytin in the M-subunit, H_M, in reaction centers of Rhodopseudomonas viridis. This has been done by measuring the transient states produced by excitation of reaction centers trapped in the PH_L ^–H_M state at 90 K. Under these conditions the normal forward electron transfer to the bacteriopheophytin in the L-subunit, H_L, is blocked and the yield of transient P⁺H_M ^– can be estimated with respect to the lifetime of P^*. Under these conditions flash induced absorbance decreases of the bacteriochlorophyll dimer 990 nm band suggest that a transient P⁺ state is formed with a quantum yield of 0.09±0.06 compared to that formed during normal photochemistry. These transient measurements provide an upper limited on the yield of a transient P⁺ H_M ^– state. An estimate of 0.09 as the yield of the P⁺ H_M ^– state is consistent with all current observations. This estimate and the lifetime of P^* suggest that the electron transfer rate from P^* to H_M, k_M, is about 5 × 10⁹ sec^–1 (_M = 200ps). These measurements suggest that the a branching ratio k_L/k_M is on the order of 200. The large value of the branching ratio is remarkable in view of the structural symmetry of the reaction center. This measurement should be useful for electron transfer calculations based upon the reaction center structure.     

Expression of a C3 plant Rubisco SSU gene in regenerated C4 Flaveria plants

We report the successful transformation, via Agrobacterium tumefaciens infection, and regeneration of two species of the genus Flaveria: F. brownii and F. palmeri. We document the expression of a C₃ plant gene, an abundantly expressed ribulose 1,5-bisphosphate carboxylase/oxygenase small subunit gene isolated from petunia, in these C₄ plants. The organ-specific expression of this petunia gene in Flaveria brownii is qualitatively identical to its endogenous pattern of expression.     

Influence of temperature and nitrogen concentration on photosynthesis ofDunaliella viridis Teodoresco

The photosynthetic behaviour ofDunaliella viridis has been studied under a combination of three variables: irradiance (0–900 mol m^–2 s^–1), temperature (15, 23, 31, 38, 42 °C) and nitrogen concentration (0.05, 0.5, 1.5, 5, 10 mM NO ₃ ^- ) at a salinity of 2 M NaCl.The highest rates of photosynthesis have been found at 31 °C and a nitrate concentration of 10 mM. There exists a synergistic effect between temperature and nitrogen availability on the photosynthesis ofD. viridis; under nitrogen deficiency oxygen evolution is low, even null at high temperature. The interaction between these two variables of control occurs in a multiplicative way. There is also a general increase in photosynthetic pigments following the increase in nitrogen concentration in the culture medium. The normalization of net photosynthesis data in relation to chlorophylla shows that nitrogen concentration makes an indirect control of the photosynthetic rate ofD. viridis through the variation of pigment concentration.     

Seasonal patterns of water relations and enzyme activity of the facultative CAM plant Portulacaria afra (L.) Jacq.

Abstract. Portulacaria afra (L.) Jacq. is a perennial facultative CAM species showing a seasonal shift from C₃ to CAM photosynthesis. The shift to CAM during the summer occurs despite continued irrigation of the plants. The authors examined the hypothesis that the seasonal shift to CAM occurred because of low transient water potentials. They measured changes in whole leaf water, osmotic and pressure potentials over the course of the shift. They also studied changes in enzyme activity to ascertain if PEP carboxylase and PEP carboxykinase were induced during the seasonal shift to CAM. Water potentials were high, from -0.1 to -0.5 MPa, predawn and midday, when the C₃ pathway of photosynthesis was utilized. Osmotic potentials were constant, from -0.7 to - 0.8 MPa, indicating very little change in turgor. P. afra shifted to CAM indicated by large diurnal acid fluctuations (300 400 meq m⁻²) despite C₃-like predawn water potentials. Midday water potentials usually decreased 0.2-0.7 MPa, while the osmotic potential remained unchanged or decreased slightly. Thus, a midday loss of turgor was associated with the use of the CAM pathway. The results support the hypothesis that the induction of CAM occurred due to low transient water potentials and may be partially mediated through the loss of turgor. The shift to CAM is only a partial induction with PEP carboxykinase showing high activity all year round while PEP carboxylase increases three-to five-fold over C₃ levels. Relatively high levels of CAM enzyme activity enables the utilization of the CAM pathway in the winter and spring in response to high daytime temperatures and increased evaporative demand. These results would lead to an increase in water use efficiency during such periods when compared to other inducible CAM species.