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This paper reviews the interactions between water and nitrogen from physiological, agronomic, economic, breeding and modelling perspectives. Our primary focus is wheat; we consider forage crops, sorghum and legumes where relevant aspects of water–nitrogen interactions have been advanced.
From a physiological perspective, we ask: How does nitrogen deficit influence the water economy of the crop? How does water deficit influence the nitrogen economy of the crop? How do combined water and nitrogen deficit affect crop growth and yield? We emphasise synergies, and the nitrogen-driven trade-off between the efficiency in the use of water and nitrogen. The concept of nitrogen–water co-limitation is discussed briefly.
From agronomic and economic perspectives, the need to match supply of nitrogen and water is recognised, but this remains a challenge in dryland systems with uncertain rainfall. Under-fertilisation commonly causes gaps between actual and water-limited potential yield. We discuss risk aversion and the role of seasonal rainfall forecasts to manage risk.
From a breeding perspective, we ask how selection for yield has changed crop traits relating to water and nitrogen. Changes in nitrogen traits are more common and profound than changes in water-related traits. Comparison of shifts in the wheat phenotype in Australia, UK, Argentina and Italy suggests that improving yield per unit nitrogen uptake is straightforward; it requires selection for yield and allowing grain protein concentration to drift unchecked. A more interesting proposition is to increase nitrogen uptake to match yield gains and conserve protein in grain. Increased stomatal conductance is a conspicuous response to selection for yield which partially conflicts with the perception that reduced conductance at high vapour pressure deficit is required to increase water- use efficiency; but high stomatal conductance at high vapour pressure deficit may be adaptive for thermal stress.
From a modelling perspective, water and nitrogen are linked in multiple ways. In crops where water limits growth, reduced biomass reduces nitrogen demand. Reciprocally, nitrogen limitation during crop expansion reduces leaf area index and increases the soil evaporation : transpiration ratio. Water–nitrogen interactions are also captured in the water-driven uptake of nitrogen by mass flow and diffusion and in the water-driven processes of nitrogen in soil (e.g. mineralisation).
The paper concludes with suggestions for future research on water-nitrogen interactions.
A survey of Barley yellow dwarf virus (BYDV) incidence in cereal crops in the Czech Republic over 4 years showed, on average, 13.3% BYDV-positive, randomly tested wheat and barley samples. The cultivated wheat and barley cultivars had different levels of susceptibility to BYDV infection. Field trials were performed with different barley and wheat breeding lines and cultivars, and resistance traits were evaluated after artificial inculcation by the viruliferous aphid vector Rhopalosiphum padi L. with BYDV-PAV. Our results showed high variability of visual symptom score (VSS) and reduction in grain weight per spike (GWS-R) in trials within the tested lines and cultivars. The barley line (WBON 96-123) and cultivars (Wysor, Travira) that contained RYd2 differed significantly from other cultivars in VSS. Line WBON 96-123 and cvv. Wysor and Yatzi showed the lowest GWS-R. Wheat line PSR 3628 and cvv. Altigo, Elan, Sparta, Aladin and Hewit showed significant difference from other cultivars in VSS. PSR 3628, Sparta, and Elan showed the lowest GWS-R. Similar results were obtained from BYDV titre analysis by RT-qPCR corresponding to the VSS and GWS-R traits. A low virus titre corresponded to low VSS and GWS-R. Hence, our results suggest that laborious and time-consuming GWS-R analysis could be replaced in some cases by qPCR-based BYDV titre analysis and, together with VSS evaluation, could enhance the efficiency of resistance assessment.
Stripe (yellow) rust, caused by Puccinia striiformis Westend. f. sp. tritici Erikss., is one of the most damaging diseases in wheat and is especially damaging for winter and facultative wheat. The objective of this study was to understand stripe rust resistance in 100 wheat and facultative wheat entries from the International Winter Wheat Improvement Program by conducting experiments in a greenhouse and in four field environments in Washington State, USA, and by genotyping molecular markers linked to Yr genes. Percentages of entries resistant to the rust races at the seedling stage were: PST-17, 44%; PST-37, 32%; PST-43, 45%; PST-45, 49%; PST-116, 18%; PST-100, 17%; and PST-127, 8%. Molecular markers were positive for genes Yr9, Yr17, and Yr18 and negative for Yr5, Yr10, and Yr15. Yr18 was present in 44 entries (44%). By using the highly virulent races PST-127 and PST-100 under controlled conditions, 16 entries were shown to have high-temperature adult-plant (HTAP) resistance and resistant–moderately resistant field reactions at all four field sites. Resistant entries, especially those with HTAP resistance, were also identified in the field experiments.
Light absorption and light extinction of wheat (Triticum aestivum L.) are among the most important parameters affecting wheat growth and yield production. However, these properties are affected by plant and environmental factors. Despite wheat being an important food crop, there is not much information on these light parameters in wheat. Accordingly, light parameters were investigated in wheat plants in the present study in three field experiments under two distinct climatic conditions, a warm arid and semi-arid climate and a cool climate. The aims of the study were to determine how light absorption and light extinction of wheat are affected by: (1) planting date and plant genotype under arid and semi-arid conditions (Experiment I); and (2) N chemical fertilisation and plant genotype under arid and semi-arid conditions (Experiment II) and cool temperate conditions (Experiment III). Light absorption by the canopy was determined using a lightmeter and coefficients of extinction were calculated. Analyses of variance indicated significant effects of experimental treatments on light properties, wheat growth and yield production. There was a high rate of variability in light absorption, with a maximum of 59.27%, and light extinction coefficients were in the range 0.45–0.66. The experimental treatments resulted in high variability in the leaf area index (2.08–7.49), wheat biological yield (7831.1–22 515.96 kg ha–1), grain yield (2481.3–9273.57 kg ha–1) and harvest index (32.86–53.90%). The interaction between planting date and plant genotypes indicated that the responses of different wheat genotypes to planting date were highly variable, significantly affecting light absorption and light extinction by wheat. It is possible to make the optimum use of solar light and produce the highest rate of yield if the most efficient genotype (Line 14-C81, Pishvaz and Pishtaz) is planted on the right planting date (15 November) using the optimum rate of N chemical fertilisation (50, 100, 150 kg ha–1).
Drought stress is the most important production constraint in maize (Zea mays L.), especially in rainfed agriculture. To improve productivity of rainfed maize, the development of hybrids with tolerance to drought stress is an important objective in maize breeding programs. The present study was undertaken to identify maize hybrids that perform better under drought-stress and drought-free conditions by using various selection indices. These selection indices were calculated on the basis of yield (t ha–1) performance of hybrids measured under drought stress and optimum environments. A set of 38 cultivars was evaluated at 10 environments (representing five each of drought stress and optimum growing conditions). The average reduction in grain yield due to drought stress was 52%. Effects of genotype, environment and their interaction were significant sources of variation in determining grain yield, respectively explaining 5.0–7.4%, 55.0–60.2% and 12.0–15.0% of total variation in yield under drought-stress and drought-free conditions. Of eight selection indices considered for study, three indices such as harmonic mean, geometric mean, and stress tolerance index were identified as suitable for selection of genotypes capable of performing well both under drought-stress and drought-free environments. Drought response index and drought resistance index were found useful in identifying hybrids that performed better under drought stress. Stress susceptibility index was negatively correlated with yield measured under drought stress. Stress susceptibility index could be used as selection index but only in combination with yield performance data under water-deficit conditions in order to identify drought-tolerant hybrids with reasonable productivity. Test weight, shelling percentage, days to maturity, and ear girth were found to be useful traits for improving yield performance across diverse environments. Cultivation of identified drought-tolerant hybrids would be useful to enhance maize productivity in drought-stress environments.
High-molecular-weight secretory proteins and low-molecular-weight exudates (carboxylates, phenols, free amino acids and sugars) released from roots of soybean (Glycine max (L.) Merr.) differentially influence genotypic phosphorus (P) acquisition efficiency (PAE). We hypothesised that genotypes with higher root exudation potential would exhibit enhanced P acquisition, and screened 116 diverse soybean genotypes by labelling shoots with 14CO2. A root exudation index (REI) derived from total 14C in the root exudate at sufficient (250 μm) and low (4 μm) P levels was used to classify genotypes for PAE. Genotypes with REI >2.25 exhibited significantly higher exudation at low than at sufficient P, which in turn increased PAE. Under low P availability, efficient genotypes exude a greater quantity of organic compounds into the rhizosphere. This increases P availability to meet the crop requirement, enabling the crop to produce consistent biomass and seed yield with reduced fertiliser addition. Such maintenance of growth and yield potential by mining the inherent soil P is a favourable trait in genotypes, reducing dependence on P fertilisers. Measuring REI at seedling stage to select P-efficient plants accelerates the screening process by accommodating large numbers of genotypes.
Colletotrichum trifolii, the causal agent of southern anthracnose, is a major fungal disease in red clover (Trifolium pratense) and lucerne (alfalfa, Medicago sativa), with varietal differences reported. However, results on the reactions of other fodder legumes grown in Central Europe are limited. In this study, a greenhouse test for resistance evaluated the response of T. repens, T. alexandrinum, T. incarnatum, T. resupinatum, Lotus corniculatus, M. lupulina, Onobrychis viciifolia, Vicia sativa, and V. villosa against C. trifolii compared with M. sativa. Conditions of the test were not suitable for T. alexandrinum and the two Vicia species; however, T. repens, T. resupinatum, and L. corniculatus proved highly resistant against the fungus. The other species showed mean plant survival rates of 61.5–84.7%.
Foraging strategies in arbuscular mycorrhizal fungi (AMF) for heterogeneously distributed resources in the soil remain to be explored. We used nylon-mesh bags of 30 μm to simulate patches of different phosphorus (P) supply levels (Expt 1) and P forms (organic v. inorganic, Expts 1 and 2). In Expt 1, host maize (Zea mays) was unfertilised; in each pot, five P-enriched bags were supplied with either Na-phytate or KH2PO4 at P rates of 0 (P0), 50 (P50), 100 (P100), 150 (P150) and 200 (P200) mg P kg–1. In Expt 2, maize plants were supplied with 20 (P20) or 50 (P50) mg P kg–1, and five P-enriched bags were supplied with different P forms (Na-phytate, lecithin, RNA, KH2PO4) and a nil-P control. Three fungal species (Funneliformis mosseae, Rhizophagus irregularis, and Glomus etunicatum) were compared in Expt 1, and the first two species in Expt 2.
In Expt 1, the hyphal-length density (HLD) of G. etunicatum was not significantly different among different P levels when supplied with KH2PO4, whereas the HLD of R. irregularis tended to increase at higher P supply (above P50) in the Na-phytate treatment. The HLD of F. mosseae increased at P150 when supplied with KH2PO4, and increased at P100 and P150 in the Na-phytate treatment relative to P0. APase activity levels were more related to P supply level, in particular with F. mosseae inoculation and uninoculated control, showing that P200 significantly reduced APase relative to P0. In Expt 2, greater hyphal growth of both fungal species tended to occur with KH2PO4. At P20, the HLD of R. irregularis in treatments with KH2PO4 and lecithin, and of F. mosseae with KH2PO4, were higher than in P0. At P50, the HLD of F. mosseae was higher than of R. irregularis; but P form had no significant influence on HLD of F. mosseae, whereas the HLD of R. irregularis in the P-amended treatment (except with Na-phytate) was higher than in P0. APase activity did not differ significantly between the two fungal species. Highest APase activity generally occurred with lecithin, with no significant difference among the other P forms. Our results indicate that the response of AMF to P-enriched patches is complex, and both the form and amount of P supplied should be considered. Variations between AMF in the proliferation of hyphae to heterogeneous nutrient patches might be a mechanism by which these species can maintain diversity in intensive agricultural ecosystems.
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