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G D Farquhar

Publications and source records attributed to G D Farquhar.

At least 19 recordsLinked to original sources

The influence of protein-protein interactions on the organization of proteins within thylakoid membranes.

The influence of attractive protein-protein interactions on the organization of photosynthetic proteins within the thylakoid membrane was investigated. Protein-protein interactions were simulated using Monte Carlo techniques and the influence of different interaction energies was examined. It was found that weak interactions led to protein clusters whereas strong interactions led to ramified chains. An optimum curve for the relationship between interaction energy and the number of contact sites emerged. With increasing particle densities the effect decreased. In a mixture of interacting and noninteracting particles the distance between the noninteracting particles was increased and there seemed to be much more free space around them. In thylakoids, this could lead to a more homogeneous distribution of the noninteracting but rate-limiting cytochrome bf complexes. Due to the increased free space between cytochrome bf, obstruction of binding sites--occurring unavoidably in a random distribution--may be drastically reduced. Furthermore, protein-protein interactions in thylakoids may lead to a decrease in plastoquinone diffusion.

Binding Sites↗

Breeding for high water-use efficiency.

There is a pressing need to improve the water-use efficiency of rain-fed and irrigated crop production. Breeding crop varieties with higher water-use efficiency is seen as providing part of the solution. Three key processes can be exploited in breeding for high water-use efficiency: (i) moving more of the available water through the crop rather than it being wasted as evaporation from the soil surface or drainage beyond the root zone or being left behind in the root zone at harvest; (ii) acquiring more carbon (biomass) in exchange for the water transpired by the crop, i.e. improving crop transpiration efficiency; (iii) partitioning more of the achieved biomass into the harvested product. The relative importance of any one of these processes will vary depending on how water availability varies during the crop cycle. However, these three processes are not independent. Targeting specific traits to improve one process may have detrimental effects on the other two, but there may also be positive interactions. Progress in breeding for improved water-use efficiency of rain-fed wheat is reviewed to illustrate the nature of some of these interactions and to highlight opportunities that may be exploited in other crops as well as potential pitfalls. For C3 species, measuring carbon isotope discrimination provides a powerful means of improving water-use efficiency of leaf gas exchange, but experience has shown that improvements in leaf-level water-use efficiency may not always translate into higher crop water-use efficiency or yield. In fact, the reverse has frequently been observed. Reasons for this are explored in some detail. Crop simulation modelling can be used to assess the likely impact on water-use efficiency and yield of changing the expression of traits of interest. Results of such simulations indicate that greater progress may be achieved by pyramiding traits so that potential negative effects of individual traits are neutralized. DNA-based selection techniques may assist in such a strategy.

Agriculture↗

Dependence of plastoquinol diffusion on the shape, size, and density of integral thylakoid proteins.

The diffusion of plastoquinol in the chloroplast thylakoid membrane is modelled using Monte Carlo techniques. The integral proteins are seen as obstacles to diffusion, and features of percolation theory emerge. Thus, the diffusion coefficient diminishes with increasing distance and there is a critical threshold of protein concentration, above which the long-range diffusion coefficient is zero. The area occupied by proteins in vivo is assessed and appears to be around this threshold, as determined from calculations assuming randomly distributed noninteracting proteins. Slight changes in the protein arrangement lead to pronounced changes in diffusion behaviour under such conditions. Mobility of the proteins increases the protein occupancy threshold, while boundary lipids impermeable to PQ diffusion decrease it. Further, the obstruction of plastoquinone/plastoquinol binding sites in a random arrangement is evaluated.

Binding Sites↗

The effect of exogenous abscisic acid on stomatal development, stomatal mechanics, and leaf gas exchange in Tradescantia virginiana.

Gas exchange parameters and stomatal physical properties were measured in Tradescantia virginiana plants grown under well-watered conditions and treated daily with either distilled water (control) or 3.0 mM abscisic acid (ABA). Photosynthetic capacity (CO(2) assimilation rate for any given leaf intercellular CO(2) concentration [c(i)]) and relative stomatal sensitivity to leaf-to-air vapor-pressure difference were unaffected by the ABA treatment. However, at an ambient CO(2) concentration (c(a)) of 350 micromol mol(-1), ABA-treated plants operated with significantly lower c(i). ABA-treated plants had significantly smaller stomata and higher stomatal density in their lower epidermis. Stomatal aperture versus guard cell pressure (P(g)) characteristics measured with a cell pressure probe showed that although the form of the relationship was similar in control and ABA-treated plants, stomata of ABA-treated plants exhibited more complete closure at P(g) = 0 MPa and less than half the aperture of stomata in control plants at any given P(g). Scaling from stomatal aperture versus P(g) to stomatal conductance versus P(g) showed that plants grown under ABA treatment would have had significantly lower maximum stomatal conductance and would have operated with lower stomatal conductance for any given guard cell turgor. This is consistent with the observation of lower c(i)/c(a) in ABA-treated plants with a c(a) of 350 micromol mol(-1). It is proposed that the ABA-induced changes in stomatal mechanics and stomatal conductance versus P(g) characteristics constitute an improvement in water-use efficiency that may be invoked under prolonged drought conditions.

Abscisic Acid↗

Variation in branchlet delta13C in relation to branchlet nitrogen concentration and growth in 8-year-old hoop pine families (Araucaria cunninghamii) in subtropical Australia.

Carbon isotope composition (delta13C) of branchlet tissue at nine canopy positions, and nitrogen concentration (N(mass)) at four canopy positions, were assessed in 8-year-old hoop pine (Araucaria cunninghamii Ait. ex D. Don) trees from 23 half-sib families, grown in six blocks of a progeny test in southeastern Queensland, Australia. There was considerable variation among sampling positions, families and blocks in both delta13C and N(mass). The delta13C was positively related to N(mass) only for samples from the upper outer crown (P < 0.005). Phenotypic correlations existed between tree growth and canopy delta13C. Branchlet delta13C of the inner and lower outer crown was positively related (P < 0.037) to tree height, but delta13C in branchlets of the upper outer crown was not related to tree height, or was related negatively (P < 0.045). There were significant differences in delta13C between hoop pine families for six canopy positions (upper canopy positions as well as lower canopy positions on the northern side), with heritabilities greater than 0.40. The significance of these findings is discussed in relation to water and light competition within the tree canopy of hoop pine.

Carbon Isotopes↗

Variation in the oxygen isotope ratio of phloem sap sucrose from castor bean. Evidence in support of the Péclet effect.

Theory suggests that the level of enrichment of (18)O above source water in plant organic material (Delta) may provide an integrative indicator of control of water loss. However, there are still gaps in our understanding of the processes affecting Delta. One such gap is the observed discrepancy between modeled enrichment of water at the sites of evaporation within the leaf and measured enrichment of the leaf water as a whole (Delta(L)). Farquhar and Lloyd (1993) suggested that this may be caused by a Péclet effect. It is also unclear whether organic material formed in the leaf reflects enrichment of water at the sites of evaporation within the leaf or Delta(L). To investigate this question castor bean (Ricinus communis L.) leaves, still attached to the plant, were sealed into a controlled-environment gas exchange chamber and subjected to a step change in leaf-to-air vapor pressure difference. Sucrose was collected from a cut on the petiole of the leaf in the chamber under equilibrium conditions and every hour for 6 h after the change in leaf-to-air vapor pressure difference. Oxygen isotope composition of sucrose in the phloem sap (Delta(suc)) reflected modeled Delta(L). A model is presented describing Delta(suc) at isotopic steady state, and accounts for 96% of variation in measured Delta(suc). The data strongly support the Péclet effect theory.

Ricinus communis↗

Effects of elevated [CO(2)] and nitrogen nutrition on cytokinins in the xylem sap and leaves of cotton.

We measured the level of xylem-derived cytokinins (CKs) entering a cotton leaf, and the CK levels in the same leaf, thus enabling xylem sap and foliar CKs to be compared concurrently. Although zeatin was the dominant CK in xylem sap, zeatin, dihydrozeatin, and N(6)-(2-isopentenyl) adenine were present in approximately equimolar levels in leaves. Elevated [CO(2)] (EC) has an effect on the levels of cytokinins in sap and leaf tissues. This effect was modulated by the two levels of root nitrogen nutrition (2 and 12 mM nitrate). Growth enhancement (70%) in EC over plants in ambient [CO(2)] (AC) was observed for both nitrogen nutrition treatments. Low-nitrogen leaves growing in EC exhibited photosynthetic acclimation, whereas there was no sign of photosynthetic acclimation in high-nitrogen grown leaves. Under these prevailing conditions, xylem sap and leaf tissues were obtained for CK analysis. Higher nitrogen nutrition increased the delivery per unit leaf area of CKs to the leaf at AC. EC caused a greater increase in CK delivery to the leaf at low nitrogen conditions (106%) than at high nitrogen conditions (17%). EC induced a significant increase in CK content in low-nitrogen leaves, whereas CK content in leaf tissues was similar for high-nitrogen leaves growing in AC and EC.

Carbon Dioxide↗

Carbon isotope discrimination varies genetically in c(4) species.

Carbon-isotope discrimination (Delta) is used to distinguish between different photosynthetic pathways. It has also been shown that variation in Delta occurs among varieties of C(3) species, but not as yet, in C(4) species. We now report that Delta also varies among genotypes of sorghum (Sorghum bicolor Moench), a C(4) species. The discrimination in leaves of field-grown plants of 12 diverse genotypes of sorghum was measured and compared with their grain yields. Discrimination varied significantly among genotypes, and there was a significant negative correlation between grain yield and Delta. The variation in Delta may be caused by genetic differences in either leakiness of the bundle-sheath cells or by differences in the ratio of assimilation rate to stomatal conductance. At the leaf level, the former should be related to light-use efficiency of carbon fixation and the latter should be related to transpiration efficiency. Both could relate to the yield of the crop.

Journal Article↗

Short-term changes in leaf carbon isotope discrimination in salt- and water-stressed c(4) grasses.

Online carbon isotope discrimination (Delta) and leaf gas exchange measurements were made with control and salt-stressed Zea mays and Andropogon glomeratus, two NADP-ME type C(4) grasses. Linear relationships between Delta and p(i)/p(a) (the ratio of intercellular to atmospheric CO(2) partial pressure) were found for control plants which agreed well with theoretical models describing carbon isotope discrimination in C(4) plants. These data provided estimates of phi, the proportion of CO(2) fixed by phosphoenolpyruvate carboxylase which leaks out of the bundle sheath and the component of fractionation due to diffusion in air. Salt-stressed plants had wider variation in Delta for the same or less range in p(i)/p(a). Additional work indicated Delta changed independently of p(i)/p(a) in both water- and salt-stressed plants, suggesting a possible diurnal change in phi as plant water status changed linked to a decrease in the activity of the C(3) photosynthetic pathway relative to C(4) pathway activity. The possible effect of stress-induced changes in phi on organic matter delta(13) C of C(4) plants is apt to be most apparent in chronically stressed environments.

Journal Article↗

Effects of soil strength on the relation of water-use efficiency and growth to carbon isotope discrimination in wheat seedlings.

The ratio of carbon accumulation to transpiration, W, of wheat (Triticum aestivum L.) seedlings increased with increasing soil strength, measured as soil penetrometer resistance, and this was already apparent at the two leaf stage. The ratio was negatively correlated with carbon isotope discrimination, in accord with theory. This means that decrease in intercellular partial pressure of CO(2) accounted for an important part of the increase in W with increasing soil strength. Despite a lower CO(2) concentration in the leaves at high soil strength, assimilation rate per unit leaf area was enhanced. Greater ribulose 1,5-bisphosphate carboxylase activity confirmed that photosynthetic capacity was actually increased. This pattern of opposite variation of assimilation rate and of stomatal conductance is unusual. The ratio of plant carbon mass to leaf area increased markedly with increasing soil strength, mainly because of a greater investment of carbon into roots than into shoots. A strong negative correlation was found between this ratio and carbon isotope discrimination. For a given increase in discrimination, decrease in carbon mass per leaf area was proportionally larger than decrease in assimilation rate, so that relative growth rate was positively correlated to carbon isotope discrimination.

Journal Article↗

Gradients of Intercellular CO(2) Levels Across the Leaf Mesophyll.

Most current photosynthesis models, and interpretations of many wholeleaf CO(2) gas exchange measurements, are based on the often unstated assumption that the partial pressure of CO(2) is nearly uniform throughout the airspaces of the leaf mesophyll. Here we present measurements of CO(2) gradients across amphistomatous leaves allowed to assimilate CO(2) through only one surface, thus simulating hypostomatous leaves. We studied five species: Eucalyptus pauciflora Sieb. ex Spreng., Brassica chinensis L., Gossypium hirsutum L., Phaseolus vulgaris L., and Spinacia oleracea L. For Eucalyptus, maximum CO(2) pressure differences across the leaf mesophyll were 73 and 160 microbar when the pressures outside the lower leaf surface were 310 and 590 microbar, respectively. Using an approximate theoretical calculation, we infer that if the CO(2) had been supplied equally at both surfaces then the respective mean intercellular CO(2) pressures would have been roughly 12 and 27 microbar less than the pressures in the substomatal cavities in these cases. For ambient CO(2) pressures near 320 microbar, the average and minimum pressure differences across the mesophyll were 45 and 13 microbar. The corresponding mean intercellular CO(2) pressures would then be roughly 8 and 2 microbar less than those in the substomatal cavities. Pressure differences were generally smaller for the four agricultural species than for Eucalyptus, but they were nevertheless larger than previously reported values.

Journal Article↗

Correlation between the Carbon Isotope Discrimination in Leaf Starch and Sugars of C(3) Plants and the Ratio of Intercellular and Atmospheric Partial Pressures of Carbon Dioxide.

Carbon isotope discrimination (Delta) was analyzed in leaf starch and soluble sugars, which represent most of the recently fixed carbon. Plants of three C(3) species (Populus nigra L. x P. deltoides Marsh., Gossypium hirsutum L. and Phaseolus vulgaris L.) were kept in the dark for 24 hours to decrease contents of starch and sugar in leaves. Then gas exchange measurements were made with constant conditions for 8 hours, and subsequently starch and soluble sugars were extracted for analysis of carbon isotope composition. The ratio of intercellular, p(i), and atmospheric, p(a), partial pressures of CO(2), was calculated from gas exchange measurements, integrated over time and weighted by assimilation rate, for comparison with the carbon isotope ratios in soluble sugars and starch. Carbon isotope discrimination in soluble sugars correlated strongly (r = 0.93) with p(i)/p(a) in all species, as did Delta in leaf starch (r = 0.84). Starch was found to contain significantly more (13)C than soluble sugar, and possible explanations are discussed. The strong correlation found between Delta and p(i)/p(a) suggests that carbon isotope analysis in leaf starch and soluble sugars may be used for monitoring, indirectly, the average of p(i)/p(a) weighted by CO(2) assimilation rate, over a day. Because p(i)/p(a) has a negative correlation with transpiration efficiency (mol CO(2)/mol H(2)O) of isolated plants, Delta in starch and sugars may be used to predict differences in this efficiency. This new method may be useful in ecophysiological studies and in selection for improved transpiration efficiency in breeding programs for C(3) species.

Journal Article↗

Models relating subcellular effects of temperature to whole plant responses.

The analysis of the temperature dependence of the rate of photosynthetic carbon assimilation by leaves of C3 plants has involved subdivision into a diffusional (mainly stomatal) component and a biochemical one. The latter has been further analysed in terms of Rubisco activity and of the capacity for regeneration of RubP. Rubisco activity has been modelled in terms of the number of active (carbamylated) enzyme sites and the kinetics of these sites. RubP regeneration capacity has been modelled in terms of photosynthetic electron transport and photophosphorylation capacity and the rate of supply of inorganic phosphate. The short-term (minutes to hours) responses of assimilation rate to changes in temperature of Eucalyptus pauciflora leaves, in the range 15 to 35 degrees C, are examined using this framework. It is argued that the above approach has some useful features, even if these are mainly pedagogical. Problems with the simplified framework are also discussed, including heterogeneity of the light intensity and CO2 concentration within the leaf. The latter is discussed in terms of the resistances to diffusion within the leaf and observations of non-uniformity ('patches') of stomatal opening. The effects of temperature, couched in the above terms, appear to be mainly those on the supply of carbon as a substrate for growth. Yet the model (or description of carbon assimilation) can accommodate certain effects of demand for carbon (sink limitation). These occur, notably, via the supply of inorganic phosphate, but potentially also via Rubisco activation and other mechanisms. A striking example of effects of reduced temperature on assimilation and growth being caused by changed demand for carbon, is given for the tropical crop, Arachis hypogaea. This example prompts an appraisal of the effects of temperature on components of growth other than photosynthesis, for example, of leaf production rate.

Carbon↗

Investigation of the CO(2) Dependence of Quantum Yield and Respiration in Eucalyptus pauciflora.

In leaves of C(3) plants, the rate of nonphotorespiratory respiration appears to be higher in darkness than in the light. This change from a high to a low rate of carbon loss with increasing photon flux density leads to an increase in the apparent quantum yield of photosynthetic CO(2) assimilation at low photon flux densities (Kok effect). The mechanism of this suppression of nonphotorespiratory respiration is not understood, but biochemical evidence and the observation that a Kok effect is often not observed under low O(2), has led to the suggestion that photorespiration might be involved in some way. This hypothesis was tested with snowgum (Eucalyptus pauciflora Sieb. ex Spreng.) using gas exchange methods. The test was based on the assumption that if photorespiration were involved, then it would be expected that the intercellular partial pressure of CO(2) would also have an influence on the Kok effect. Under normal atmospheric levels of CO(2) and O(2), a Kok effect was found. Changing the intercellular partial pressure of CO(2), however, did not affect the estimate of nonphotorespiratory respiraton, and it was concluded that its decrease with increasing photon flux density did not involve photorespiration. Concurrent measurements showed that the quantum yield of net assimilation of CO(2) increased with increasing intercellular partial pressure of CO(2), and this increase agreed closely with predictions based on recent models of photosynthesis.

Journal Article↗

Leaf Conductance in Relation to Rate of CO(2) Assimilation: I. Influence of Nitrogen Nutrition, Phosphorus Nutrition, Photon Flux Density, and Ambient Partial Pressure of CO(2) during Ontogeny.

Plants of Zea mays were grown with different concentrations of nitrate (0.6, 4, 12, and 24 millimolar) and phosphate (0.04, 0.13, 0.53, and 1.33 millimolar) supplied to the roots, photon flux densities (0.12, 0.5, and 2 millimoles per square meter per second), and ambient partial pressures of CO(2) (305 and 610 microbars). Differences in mineral nutrition and irradiance led to a large variation in rate of CO(2) assimilation per unit leaf area (A, 11 to 58 micromoles per square meter per second) when measured under standard conditions. The variation was shown, with the plants that had received different amounts of nitrate, to be related to variations in the nitrogen and chlorophyll contents, and phosphoenolpyruvate and ribulose-1,5-bisphosphate carboxylase activities per unit leaf area. Irrespective of growth treatment, A and leaf conductance to CO(2) transfer (g), measured under standard conditions were in almost constant proportion, implying that intercellular partial pressure of CO(2) (p(i)), was almost constant at 95 microbars. The same proportionality was maintained as A and g increased in an initially nitrogen-deficient plant that had been supplied with abundant nitrate. It was shown that p(i) measured at a given ambient partial pressure was not affected by the ambient partial pressure at which the plants had been grown, although it was different when measured at different ambient partial pressures. This suggests that the close coupling between A and g in these experiments is not associated with sensitivity of stomata to change in p(i).Similar, though less comprehensive, experiments were done with Gossypium hirsutum, and yielded similar conclusions, except that the proportionality between A and g at normal ambient partial pressure of CO(2) implied P(i) approximately 200 microbars.

Journal Article↗

Leaf Conductance in Relation to Rate of CO(2) Assimilation: II. Effects of Short-Term Exposures to Different Photon Flux Densities.

When photon flux density incident on attached leaves of Zea mays L. was varied from the equivalent of 0.12 of full sunlight to full sunlight, leaf conductance to CO(2) transfer, g, changed in proportion to the change in rate of CO(2), assimilation, A, with the result that intercellular partial pressure of CO(2) remained almost constant. The proportionality was the same as that previously found in g and A measured at one photon flux density in plants of Zea mays L. grown at different levels of mineral nutrition, light intensities, and ambient partial pressures of CO(2). In shade-grown Phaseolus vulgaris L. plants, A as photon flux density was increased from about 0.12 up to about 0.5 full sunlight, the proportionality being almost the same in plants grown at low and at high light intensity.When photon flux density incident on the adaxial and abaxial surfaces of the isolateral leaves of Eucalyptus pauciflora Sieb. ex Spreng was varied, g and A also varied proportionally. The leaf conductance in a particular surface was affected by the photon flux density at the opposite surface to a greater extent than was expected on the basis of transmittance. The results indicated that stomata may, in some way, be sensitive to the photon flux absorbed within the leaf as a whole.

Journal Article↗

Leaf Conductance in Relation to Rate of CO(2) Assimilation: III. Influences of Water Stress and Photoinhibition.

Rates of CO(2) assimilation and leaf conductances to CO(2) transfer were measured in plants of Zea mays during a period of 14 days in which the plants were not rewatered, and leaf water potential decreased from -0.5 to -8.0 bar. At any given ambient partial pressure of CO(2), water stress reduced rate of assimilation and leaf conductance similarly, so that intercellular partial pressure of CO(2) remained almost constant. At normal ambient partial pressure of CO(2), the intercellular partial pressure of CO(2) was estimated to be 95 microbars. This is the same as had been estimated in plants of Zea mays grown with various levels of nitrogen supply, phosphate supply and irradiance, and in plants of Zea mays examined at different irradiances.After leaves of Phaseolus vulgaris L. and Eucalyptus pauciflora Sieb. ex Spreng had been exposed to high irradiance in an atmosphere of CO(2)-free N(2) with 10 millibars O(2), rates of assimilation and leaf conductances measured in standard conditions had decreased in similar proportions, so that intercellular partial pressure of CO(2) remained almost unchanged. As the conductance of each epidermis that had not been directly irradiated had declined as much as that in the opposite, irradiated surface it was hypothesized that conductance may have been influenced by photoinhibition within the mesophyll tissue.

Journal Article↗