Effects of nitrogen supply on xylem cytokinin delivery, transpiration and leaf expansion of pea genotypes differing in xylem-cytokinin concentration

The rms2 and rms4 pea ( Pisum sativum L.) branching mutants have higher and lower xylem-cytokinin concentration, respectively, relative to wild type (WT) plants. These genotypes were grown at two levels of nitrogen (N) supply for 18 - 20 d to determine whether or not xylem-cytokinin concentration (X-CK) or delivery altered the transpiration and leaf growth responses to N deprivation. Xylem sap was collected by pressurising de-topped root systems. As sap-flow rate increased, X-CK declined in WT and rms2, but did not change in rms4. When grown at 5.0 mM N, X-CKs of rms2 and rms4 were 36% higher and 6-fold lower, respectively, than WT at sap-flow rates equivalent to whole-plant transpiration. Photoperiod cytokinin (CK) delivery rates ( the product of transpiration and X-CK) decreased more than 6-fold in rms4. Growth of plants at 0.5 mM N had negligible (< 10%) effects on transpiration rates expressed on a leaf area basis in WT and rms4, but decreased transpiration rates of rms2. The low-N treatment decreased leaf expansion by 20 - 25% and expanding leaflet N concentration by 15%. These changes were similar in all genotypes. At sap-flow rates equivalent to whole-plant transpiration, the low N treatment decreased X-CK in rms2 but had no discernible effect in WT and rms4. Since the low N treatment decreased transpiration of all genotypes, photoperiod CK delivery rates also decreased in all genotypes. The similar leaf growth response of all genotypes to N deprivation despite differences in both absolute and relative X-CKs and deliveries suggests that shoot N status is more important in regulating leaf expansion than xylem-supplied cytokinins. The decreased X-CK and transpiration rate of rms2 following N deprivation suggests that changes in xylem-supplied CKs may modify water use.

Genetic variation for carbon isotope discrimination in sunflower: Association with transpiration efficiency and evidence for cytoplasmic inheritance

Plants accumulate isotopes of carbon at different rates because of discrimination against C-13 relative to C-12. In plants that fix carbon by the C-3 pathway, the amount of discrimination correlates negatively with transpiration efficiency (TE) where TE is the amount of dry matter accumulated per unit water transpired. Therefore, carbon isotope discrimination (Delta) has become a useful tool for selecting genotypes with improved TE and performance in dry environments. Surveys of 161 sunflower (Helianthus spp.) genotypes of diverse origin revealed a large and unprecedented range of genetic variation for Delta (19.5-23.8parts per thousand). A strong negative genetic correlation (r(g)) between TE and Delta (r(g) = -0.87, P < 0.001) was observed in glasshouse studies. Gas exchange measurements of field grown plants indicated that Delta was strongly correlated with stomatal conductance to water vapor (g), (r(g) 0.64, P < 0.01), and the ratio of net assimilation rate (A) to g, (r(g) = 0.86, P < 0.001), an instantaneous measure of TE. Genotype CMSHA89MAX1 had the lowest TE (and highest Delta) of all genotypes tested in these studies and low yields in hybrid combination. Backcrossing studies showed that the TE of this genotype was due to an adverse effect of the MAX1 cytoplasm, which was inherited from the diploid perennial H. maximiliani Schrader. Overall, these studies suggested that there is an excellent opportunity for breeders to develop sunflower germplasm with improved TE. This can be achieved, in part, by avoiding cytoplasms such as the MAX1 cytoplasm.

Potential yield and water-use effficiency benefits in sorghum from limited maximum transpiration rate

Limitations on maximum transpiration rates, which are commonly observed as midday stomatal closure, have been observed even under well-watered conditions. Such limitations may be caused by restricted hydraulic conductance in the plant or by limited supply of water to the plant from uptake by the roots. This behaviour would have the consequences of limiting photosynthetic rate, increasing transpiration efficiency, and conserving soil water. A key question is whether the conservation of water will be rewarded by sustained growth during seed fill and increased grain yield. This simulation analysis was undertaken to examine consequences on sorghum yield over several years when maximum transpiration rate was imposed in a model. Yields were simulated at four locations in the sorghum-growing area of Australia for 115 seasons at each location. Mean yield was increased slightly ( 5 - 7%) by setting maximum transpiration rate at 0.4 mm h(-1). However, the yield increase was mainly in the dry, low-yielding years in which growers may be more economically vulnerable. In years with yield less than similar to 450 g m(-2), the maximum transpiration rate trait resulted in yield increases of 9 - 13%. At higher yield levels, decreased yields were simulated. The yield responses to restricted maximum transpiration rate were associated with an increase in efficiency of water use. This arose because transpiration was reduced at times of the day when atmospheric demand was greatest. Depending on the risk attitude of growers, incorporation of a maximum transpiration rate trait in sorghum cultivars could be desirable to increase yields in dry years and improve water use efficiency and crop yield stability.

Transpiration efficiency in a segregating population of sunflower: Inheritance, correlation with other traits and association with hybrid grain yield

Plants incorporate isotopes of carbon into their tissue at different rates because of discrimination against 13C relative to 12C during photosynthesis. This difference in discrimination has been negatively correlated with transpiration efficiency (TE) in many C3 species and so, carbon isotope discrimination (Δ) of leaf tissues has been proposed as a potential tool for selecting genotypes with improved performance under water limited conditions. The relationship between Δ and TE in sunflower has been described previously using diverse genotypes, but this relationship has not been investigated with material selected from a segregating population. In this study, the TE of twenty recombinant inbred lines from a population (HAR4 x SA52) segregating for Δ was evaluated in a rainout shelter experiment. A strong negative genetic correlation between TE and Δ was observed (rg = -0.58), confirming previous studies of sunflower with unrelated lines. In addition, TE was strongly correlated to plant height at the final harvest (rg = 0.64) and TDW (rg = 0.58), and moderately correlated to SLW (rg = 0.46) and SPAD (rg = 0.21) but not leaf number (rg = 0.02). Estimates of narrow sense heritability of TE and Δ were very high (0.82 and 0.77, respectively) suggesting that selection for these traits could occur in early generations of segregating populations. Grain yield evaluations under field conditions of hybrids contrasting for Δ showed that low Δ (high TE) hybrids had a yield advantage between 22-35% in dry environments where the yield was less than 2t/ha. While this level of yield advantage may not be realized in commercial breeding programs, computer simulations suggest that 10-15% yield improvements may be possible. Low Δ material selected from the population HAR4 x SA52 has been distributed to private seed companies for further evaluation.

Selection of species and provenances for low-rainfall areas: physiological responses of Eucalyptus cloeziana and Eucalyptus argophloia to seasonal conditions in subtropical Queensland

Responses of stomatal conductance (g(s)) and net photosynthesis (A) to changes in soil water availability, photosynthetic photon flux density (Q), air temperature (1) and leaf-to-air vapour pressure deficit (D) were investigated in 4-year-old trees of a dry inland provenance of Eucalyptus argophloia Blakely, and two dry inland provenances (Coominglah and Hungry Hills) and a humid coastal provenance (Wolvi) of Eucalyptus cloeziana F. Muell. between April 2001 and April 2002 in southeast Queensland, Australia. There were minimal differences in A, g, and water relations variables among the coastal and inland provenances of E. cloeziana but large differences between E. argophloia and E. cloeziana. E. argophloia and to a lesser extent the Hungry Hills (inland) provenance of E. cloeziana maintained relatively higher pre-dawn water potential (psi(pd)) during the dry season suggesting possible access to water at depth. Simple phenomenological models of stomatal conductance as a function of Q, T and D explained 60% of variation in gs in E. cloeziana and more than 75% in E. argophloia, when seasonal effect was incorporated in the model. A Ball-Berry model for net photosynthesis explained between 70 and 80% of observed variation in A in both species. These results have implications in matching the dry and humid provenances of E. cloeziana and E. argophloia to suitable sites in subtropical environments. (C) 2004 Elsevier B.V. All rights reserved.

Evolution of plant water relations research in Australia

Ecophysiological research in Australia has focussed, at different times, on the fundamental similarities in function between all plant species, and on the peculiarity of Australian species with respect to their survival in stressful environments. Early work on plant water relations emphasised the differences between species, and indicated that diverse structural and functional attributes occurred in species from the same water-limited environment. Most recent research has emphasised processes that optimise rates of carbon dioxide exchange, but the understanding of functioning in plants with different morphological arrangements is incomplete. Variation in functions between individual plants and geographic populations in wild species has been examined to a lesser extent. The great variety within and between populations of wild plant species warrants further study for both understanding and more effective management of this biological resource.

Trait physiology and crop modelling as a framework to link phenotypic complexity to underlying genetic systems

New tools derived from advances in molecular biology have not been widely adopted in plant breeding for complex traits because of the inability to connect information at gene level to the phenotype in a manner that is useful for selection. In this study, we explored whether physiological dissection and integrative modelling of complex traits could link phenotype complexity to underlying genetic systems in a way that enhanced the power of molecular breeding strategies. A crop and breeding system simulation study on sorghum, which involved variation in 4 key adaptive traits-phenology, osmotic adjustment, transpiration efficiency, stay-green-and a broad range of production environments in north-eastern Australia, was used. The full matrix of simulated phenotypes, which consisted of 547 location-season combinations and 4235 genotypic expression states, was analysed for genetic and environmental effects. The analysis was conducted in stages assuming gradually increased understanding of gene-to-phenotype relationships, which would arise from physiological dissection and modelling. It was found that environmental characterisation and physiological knowledge helped to explain and unravel gene and environment context dependencies in the data. Based on the analyses of gene effects, a range of marker-assisted selection breeding strategies was simulated. It was shown that the inclusion of knowledge resulting from trait physiology and modelling generated an enhanced rate of yield advance over cycles of selection. This occurred because the knowledge associated with component trait physiology and extrapolation to the target population of environments by modelling removed confounding effects associated with environment and gene context dependencies for the markers used. Developing and implementing this gene-to-phenotype capability in crop improvement requires enhanced attention to phenotyping, ecophysiological modelling, and validation studies to test the stability of candidate genetic regions.

Estimating percolation and lateral water flow on sloping land in rainfed lowland rice ecosystem

Quantifying water losses in paddy fields assists estimation of water availability in rainfed lowland rice ecosystem. Little information is available on water balance in different toposequence positions of sloped rainfed lowland. Therefore, the aim of this work was to quantify percolation and the lateral water flow with special reference to the toposequential variation. Data used for the analysis was collected in Laos and northeast Thailand. Percolation and water tables were measured on a daily basis using a steel cylindrical tube with a lid and perforated PVC tubes, respectively. Percolation rate was determined using linear regression analysis of cumulative percolation. Assuming that the total amount of evaporation and transpiration was equivalent to potential evapotranspiration, the lateral water flow was estimated using the water balance equation. Separate perched water and groundwater tables were observed in paddy fields on coarse-textured soils. The percolation rate varied between 0 and 3 mm/day across locations, and the maximum water loss by lateral movement was more than 20 mm/day. Our results are in agreement with the previously reported findings, and the methodology of estimating water balance components appears reasonably acceptable. With regard to the toposequential variation, the higher the position in the topoesquence, the greater potential for water loss because of higher percolation and lateral flow rates.

Trait physiology and crop modelling to link phenotypic complexity to underlying genetic systems

New tools derived from advances in molecular biology have not been widely adopted in plant breeding because of the inability to connect information at gene level to the phenotype in a manner that is useful for selection. We explore whether a crop growth and development modelling framework can link phenotype complexity to underlying genetic systems in a way that strengthens molecular breeding strategies. We use gene-to-phenotype simulation studies on sorghum to consider the value to marker-assisted selection of intrinsically stable QTLs that might be generated by physiological dissection of complex traits. The consequences on grain yield of genetic variation in four key adaptive traits – phenology, osmotic adjustment, transpiration efficiency, and staygreen – were simulated for a diverse set of environments by placing the known extent of genetic variation in the context of the physiological determinants framework of a crop growth and development model. It was assumed that the three to five genes associated with each trait, had two alleles per locus acting in an additive manner. The effects on average simulated yield, generated by differing combinations of positive alleles for the traits incorporated, varied with environment type. The full matrix of simulated phenotypes, which consisted of 547 location-season combinations and 4235 genotypic expression states, was analysed for genetic and environmental effects. The analysis was conducted in stages with gradually increased understanding of gene-to-phenotype relationships, which would arise from physiological dissection and modelling. It was found that environmental characterisation and physiological knowledge helped to explain and unravel gene and environment context dependencies. We simulated a marker-assisted selection (MAS) breeding strategy based on the analyses of gene effects. When marker scores were allocated based on the contribution of gene effects to yield in a single environment, there was a wide divergence in rate of yield gain over all environments with breeding cycle depending on the environment chosen for the QTL analysis. It was suggested that knowledge resulting from trait physiology and modelling would overcome this dependency by identifying stable QTLs. The improved predictive power would increase the utility of the QTLs in MAS. Developing and implementing this gene-to-phenotype capability in crop improvement requires enhanced attention to phenotyping, ecophysiological modelling, and validation studies to test the stability of candidate QTLs.

The water balance on sloping land in rainfed lowland rice ecosystem

Quantifying water losses in paddy fields assists estimation of water availability in rainfed lowland rice ecosystem. There is, however, no definite method for determining the water losses, and little information is available on water balance in different toposequence positions of a sloped rainfed lowland. Therefore, the aim of this work was to quantify percolation and the lateral water flow with special reference to the toposequential variation. Data used for the analysis was collected in Laos and northeast Thailand. Percolation and water tables were measured on a daily basis. The percolator is a steel cylindrical tube with a lid to prevent water loss from evapotranspiration. The water table meter is a short PVC tube for determination of perched water table and a long PVC tube for groundwater table, and the side is perforated with 5-mm diameter holes at 20-mm distance. Percolation rate was determined using linear regression analysis of cumulative percolation. Assuming that the total amount of evaporation and transpiration was equivalent to potential evapotranspiration, the lateral water flow was estimated using the water balance equation. Our results are in agreement with the previously reported findings, and the methodology of estimating water balance components appears reasonably acceptable. With regard to the toposequential variation, the higher position in the topoesquence, the greater potential of the water losses because of higher percolation and lateral flow rates.

Cutting management of multipurpose tree legumes: effects on green herbage production, leaf retention and water-use-efficiency during the dry season in Timor, Indonesia

Edible herbage production and water-use-efficiency of three tree legumes (Leucaena leucocephala cv. Tarramba, L. pallida x L. leucocephala (KX2) and Gliricidia sepium), cut at different times of the year (February, April, June and uncut) were compared in a semi-arid area of Timor Island, Indonesia. Cutting in the early and mid dry-season (April and June) resulted in higher total leaf production (P< 0.05) and water-use-efficiency (P< 0.05), than cutting late in the wet-season (February) or being left uncut. For the leucaena treatments removing leaf in the early to mid dry-season reduced transpiration, saving soil water for subsequent regrowth as evidenced by the higher relative water contents of leaves from these treatments. This cutting strategy can be applied to local farming conditions to increase the supply of feed for livestock during the dry season.