Recent emergence of the wheat Lr34 multi-pathogen resistance: insights from haplotype analysis in wheat, rice, sorghum and Aegilops tauschii

Spontaneous sequence changes and the selection of beneficial mutations are driving forces of gene diversification and key factors of evolution. In highly dynamic co-evolutionary processes such as plant-pathogen interactions, the plant's ability to rapidly adapt to newly emerging pathogens is paramount. The hexaploid wheat gene Lr34, which encodes an ATP-binding cassette (ABC) transporter, confers durable field resistance against four fungal diseases. Despite its extensive use in breeding and agriculture, no increase in virulence towards Lr34 has been described over the last century. The wheat genepool contains two predominant Lr34 alleles of which only one confers disease resistance. The two alleles, located on chromosome 7DS, differ by only two exon-polymorphisms. Putatively functional homoeologs and orthologs of Lr34 are found on the B-genome of wheat and in rice and sorghum, but not in maize, barley and Brachypodium. In this study we present a detailed haplotype analysis of homoeologous and orthologous Lr34 genes in genetically and geographically diverse selections of wheat, rice and sorghum accessions. We found that the resistant Lr34 haplotype is unique to the wheat D-genome and is not found in the B-genome of wheat or in rice and sorghum. Furthermore, we only found the susceptible Lr34 allele in a set of 252 Ae. tauschii genotypes, the progenitor of the wheat D-genome. These data provide compelling evidence that the Lr34 multi-pathogen resistance is the result of recent gene diversification occurring after the formation of hexaploid wheat about 8,000 years ago.

Developmental and growth controls of tillering and water-soluble carbohydrate accumulation in contrasting wheat (Triticum aestivum L.) genotypes: can we dissect them?

In wheat, tillering and water-soluble carbohydrates (WSCs) in the stem are potential traits for adaptation to different environments and are of interest as targets for selective breeding. This study investigated the observation that a high stem WSC concentration (WSCc) is often related to low tillering. The proposition tested was that stem WSC accumulation is plant density dependent and could be an emergent property of tillering, whether driven by genotype or by environment. A small subset of recombinant inbred lines (RILs) contrasting for tillering was grown at different plant densities or on different sowing dates in multiple field experiments. Both tillering and WSCc were highly influenced by the environment, with a smaller, distinct genotypic component; the genotypeenvironment range covered 350750 stems m(2) and 25210mg g(1) WSCc. Stem WSCc was inversely related to stem number m(2), but genotypic rankings for stem WSCc persisted when RILs were compared at similar stem density. Low tilleringhigh WSCc RILs had similar leaf area index, larger individual leaves, and stems with larger internode cross-section and wall area when compared with high tilleringlow WSCc RILs. The maximum number of stems per plant was positively associated with growth and relative growth rate per plant, tillering rate and duration, and also, in some treatments, with leaf appearance rate and final leaf number. A common threshold of the red:far red ratio (0.390.44; standard error of the difference0.055) coincided with the maximum stem number per plant across genotypes and plant densities, and could be effectively used in crop simulation modelling as a ocut-off' rule for tillering. The relationship between tillering, WSCc, and their component traits, as well as the possible implications for crop simulation and breeding, is discussed.

Current and emerging screening methods to identify post-head-emergence frost adaptation in wheat and barley

Cereal crops can suffer substantial damage if frosts occur at heading. Identification of post-head-emergence frost (PHEF) resistance in cereals poses a number of unique and difficult challenges. Many decades of research have failed to identify genotypes with PHEF resistance that could offer economically significant benefit to growers. Research and breeding gains have been limited by the available screening systems. Using traditional frost screening systems, genotypes that escape frost injury in trials due to spatial temperature differences and/or small differences in phenology can be misidentified as resistant. We believe that by improving techniques to minimize frost escapes, such ofalse-positive' results can be confidently identified and eliminated. Artificial freezing chambers or manipulated natural frost treatments offer many potential advantages but are not yet at the stage where they can be reliably used for frost screening in breeding programmes. Here we describe the development of a novel photoperiod gradient method (PGM) that facilitates screening of genotypes of different phenology under natural field frosts at matched developmental stages. By identifying frost escapes and increasing the efficiency of field screening, the PGM ensures that research effort can be focused on finding genotypes with improved PHEF resistance. To maximize the likelihood of identifying PHEF resistance, we propose that the PGM form part of an integrated strategy to (i) source germplasm;(ii) facilitate high throughput screening; and (iii) permit detailed validation. PGM may also be useful in other studies where either a range of developmental stages and/or synchronized development are desired.

Genetic Structure of Cochliobolus sativus Populations Sampled from Roots and Leaves of Barley and Wheat in North Dakota

Common root rot (CRR) and spot blotch, caused by Cochliobolus sativus (Ito and Kurib.) Drechsl. ex Dast., are important diseases of barley (Hordeum vulgare L.) and wheat (Triticum aestivum L.) worldwide. However, the population biology of C. sativus is still poorly understood. In this study, the genetic structure of three C. sativus populations, consisting of isolates sampled respectively from barley leaves (BL), barley roots (BR) and wheat roots (WR) in North Dakota, was analysed with amplified fragment length polymorphism (AFLP) markers. A total of 127 AFLP loci were generated among 208 C. sativus isolates analysed with three primer combinations. Gene diversity (H = 0.277-0.335) were high in all three populations. Genetic variation among C. sativus individuals within population accounted for 74%, whereas 26% of the genetic variation was explained among populations. Genetic differentiation was high (empty set PT = 0.261, corrected G ''(st)= 0.39), whereas gene flow (Nm) ranged from 1.27 to 1.56 among the three populations analysed. The multilocus linkage disequilibrium (LD) ((r) over bard = 0.0760.117) was moderate in C. sativus populations. Cluster analyses indicate that C. sativus populations differentiated according to the hosts (barley and wheat) and tissues (root and leaf) although generalists also exist in North Dakota. Crop breeding may benefit from combining genes for resistance against both specialists and generalists of C. sativus.

Diploid and tetraploid progenitors of wheat are valuable sources of resistance to the root lesion nematode Pratylenchus thornei

The root lesion nematode Pratylenchus thornei is widely distributed in Australian wheat (Triticum aestivum) producing regions and can reduce yield by more than 50%, costing the industry AU$50 M/year. Genetic resistance is the most effective form of management but no commercial cultivars are resistant (R) and the best parental lines are only moderately R. The wild relatives of wheat have evolved in P. thornei-infested soil for millennia and may have superior levels of resistance that can be transferred to commercial wheats. To evaluate this hypothesis, a collection of 251 accessions of wheat and related species was tested for resistance to P. thornei under controlled conditions in glasshouse pot experiments over two consecutive years. Diploid accessions were more R than tetraploid accessions which proved more R than hexaploid accessions. Of the diploid accessions, 11 (52%) Aegilops speltoides (S-[B]-genome), 10 (43%) Triticum monococcum (A (m) -genome) and 5 (24%) Triticum urartu (A (u) -genome) accessions were R. One tetraploid accession (Triticum dicoccoides) was R. This establishes for the first time that P. thornei resistance is located on the A-genome and confirms resistance on the B-genome. Since previous research has shown that the moderate levels of P. thornei resistance in hexaploid wheat are dose-dependent, additive and located on the B and D-genomes, it would seem efficient to target A-genome resistance for introduction to hexaploid lines through direct crossing, using durum wheat as a bridging species and/or through the development of amphiploids. This would allow resistances from each genome to be combined to generate a higher level of resistance than is currently available in hexaploid wheat.

Inheritance of resistance to root-lesion nematodes (Pratylenchus thornei and P. neglectus) in five doubled-haploid populations of wheat

Nematode species Pratylenchus thornei and P. neglectus are the two most important root-lesion nematodes affecting wheat (Triticum aestivum L.) and other grain crops in Australia. For practical plant breeding, it will be valuable to know the mode of inheritance of resistance and whether the same set of genes confer resistance to both species. We evaluated reactions to P. thornei and P. neglectus of glasshouse-inoculated plants of five doubled-haploid populations derived from five resistant synthetic hexpaloid wheat lines, each crossed to the susceptible Australian wheat cultivar Janz. For each cross we determined genetic variance, heritability and minimum number of effective resistance genes for each nematode species. Distributions of nematode numbers for both species were continuous for all doubled-haploid populations. Heritabilities were high and the resistances were controlled by 4-7 genes. There was no genetic correlation between resistance to P. thornei and to P. neglectus in four of the populations and a significant but low correlation in one. Therefore, resistances to P. thornei and to P. neglectus are probably inherited quantitatively and independently in four of these synthetic hexaploid wheat populations, with the possibility of at least one genetic factor contributing to resistance to both species in one of the populations. Parents with the greatest level of resistance will be the best to use as donor parents to adapted cultivars, and selection of resistance to both species in early generations will be optimal to carry resistance through successive cycles of inbreeding to produce resistant cultivars for release.

The importance of grasslands for animal production and other functions: a review on management and methodological progress in the tropics

The global importance of grasslands is indicated by their extent; they comprise some 26% of total land area and 80% of agriculturally productive land. The majority of grasslands are located in tropical developing countries where they are particularly important to the livelihoods of some one billion poor peoples. Grasslands clearly provide the feed base for grazing livestock and thus numerous high-quality foods, but such livestock also provide products such as fertilizer, transport, traction, fibre and leather. In addition, grasslands provide important services and roles including as water catchments, biodiversity reserves, for cultural and recreational needs, and potentially a carbon sink to alleviate greenhouse gas emissions. Inevitably, such functions may conflict with management for production of livestock products. Much of the increasing global demand for meat and milk, particularly from developing countries, will have to be supplied from grassland ecosystems, and this will provide difficult challenges. Increased production of meat and milk generally requires increased intake of metabolizable energy, and thus increased voluntary intake and/or digestibility of diets selected by grazing animals. These will require more widespread and effective application of improved management. Strategies to improve productivity include fertilizer application, grazing management, greater use of crop by-products, legumes and supplements and manipulation of stocking rate and herbage allowance. However, it is often difficult to predict the efficiency and cost-effectiveness of such strategies, particularly in tropical developing country production systems. Evaluation and on-going adjustment of grazing systems require appropriate and reliable assessment criteria, but these are often lacking. A number of emerging technologies may contribute to timely low-cost acquisition of quantitative information to better understand the soil-pasture-animal interactions and animal management in grassland systems. Development of remote imaging of vegetation, global positioning technology, improved diet markers, near IR spectroscopy and modelling provide improved tools for knowledge-based decisions on the productivity constraints of grazing animals. Individual electronic identification of animals offers opportunities for precision management on an individual animal basis for improved productivity. Improved outcomes in the form of livestock products, services and/or other outcomes from grasslands should be possible, but clearly a diversity of solutions are needed for the vast range of environments and social circumstances of global grasslands.

Genetic Structure of Cochliobolus sativus Populations Sampled from Roots and Leaves of Barley and Wheat in North Dakota

Common root rot (CRR) and spot blotch, caused by Cochliobolus sativus (Ito and Kurib.) Drechsl. ex Dast., are important diseases of barley (Hordeum vulgare L.) and wheat (Triticum aestivum L.) worldwide. However, the population biology of C. sativus is still poorly understood. In this study, the genetic structure of three C. sativus populations, consisting of isolates sampled respectively from barley leaves (BL), barley roots (BR) and wheat roots (WR) in North Dakota, was analysed with amplified fragment length polymorphism (AFLP) markers. A total of 127 AFLP loci were generated among 208 C. sativus isolates analysed with three primer combinations. Gene diversity (H = 0.277-0.335) were high in all three populations. Genetic variation among C. sativus individuals within population accounted for 74%, whereas 26% of the genetic variation was explained among populations. Genetic differentiation was high (empty set PT = 0.261, corrected G ''(st)= 0.39), whereas gene flow (Nm) ranged from 1.27 to 1.56 among the three populations analysed. The multilocus linkage disequilibrium (LD) ((r) over bard = 0.0760.117) was moderate in C. sativus populations. Cluster analyses indicate that C. sativus populations differentiated according to the hosts (barley and wheat) and tissues (root and leaf) although generalists also exist in North Dakota. Crop breeding may benefit from combining genes for resistance against both specialists and generalists of C. sativus.

Rapid phenotyping for adult-plant resistance to stripe rust in wheat

Stripe or yellow rust (YR) is a significant problem in wheat crops worldwide. The deployment of adult-plant resistance (APR) genes in wheat cultivars is considered a sustainable management strategy, as these genes confer partial resistance that is usually non-race specific. Screening for APR typically involves assessment of adult plants in the field, where expression may be influenced by environmental factors. We report a high-throughput screening method for YR APR that can be used to assess fixed lines or segregating populations grown under controlled environmental conditions (CEC). Inoculation of 3-week-old wheat plants from lines with known APR responses to YR, when grown under constant light and temperature, provided disease responses typical of adult plants. Two F-2 populations ('H45' x 'ST93' and 'Wyalkatchem' x 'ST93') segregating for APR were assessed under both CEC and field conditions. These populations showed similar variation in disease response and lines assessed in both environments attained similar rankings. Phenotypic screening using CEC and continuous light provides an opportunity to accelerate the development of new wheat cultivars with durable resistance.

More fertile florets and grains per spike can be achieved at higher temperature in wheat lines with high spike biomass and sugar content at booting

An understanding of processes regulating wheat floret and grain number at higher temperatures is required to better exploit genetic variation. In this study we tested the hypothesis that at higher temperatures, a reduction in floret fertility is associated with a decrease in soluble sugars and this response is exacerbated in genotypes low in water soluble carbohydrates (WSC). Four recombinant inbred lines contrasting for stem WSC were grown at 20/10 degrees C and 11 h photoperiod until terminal spikelet, and then continued in a factorial combination of 20/10 degrees C or 28/14 degrees C with 11 h or 16 h photoperiod until anthesis. Across environments, High WSC lines had more grains per spike associated with more florets per spike. The number of fertile florets was associated with spike biomass at booting and, by extension, with glucose amount, both higher in High WSC lines. At booting, High WSC lines had higher fixed C-13 and higher levels of expression of genes involved in photosynthesis and sucrose transport and lower in sucrose degradation compared with Low WSC lines. At higher temperature, the intrinsic rate of floret development rate before booting was slower in High WSC lines. Grain set declined with the intrinsic rate of floret development before booting, with an advantage for High WSC lines at 28/14 degrees C and 16 h. Genotypic and environmental action on floret fertility and grain set was summarised in a model.

Improving the prediction of odour impacts from meat chicken farms with detailed ventilation and odour data

Odour from meat chicken (broiler) farms is an environmental issue affecting the sustainable development of the chicken meat industry but is a normal part of broiler production. Odour plumes exhausted from broiler sheds interact with the environment, where dispersion and dilution of the odours varies constantly, especially diurnally. The potential for odour impacts is greatest when odour emission rates are high and/or when atmospheric dispersion and dilution of odour plumes is limited (i.e. during stable conditions). We continuously monitored ventilation rate, on-site weather conditions, atmospheric stability, and estimated odour concentration with an artificial olfaction system. Detailed inspection of odour emission rates at critical times, i.e. dawn, dusk and night time, revealed that maximum daily and batch odour emission rates are not necessarily the cause of odour impacts. Periods of lower odour emission rates on each day are more likely to correspond with odour impacts. Odour emission rates need to be measured at the times when odour impacts are most likely to occur, which is likely to be at night. Additionally, high resolution ventilation rate data should be sought after to improve odour emission models, especially at critical times of the day. Consultants, regulators and researchers need to give more thought to odour emission rates from meat chicken farms to improved prediction and management of odour impacts.

QTLs for water absorption and flour yield identified in the doubled haploid wheat population Lang/QT8766

Two key quality traits in milling wheat are flour yield (FY) and water absorption (WA). Ideally, breeders would prefer to use markers to select promising lines rather than time consuming rheology tests. In this study, we measured FY and WA on a wheat mapping population (Lang/QT8766) of 162 individuals grown in two replicated field experiments at three locations over 2 years. We also carried out near infrared reflectance spectroscopy (NIRS) predictions on the grain for these traits to see if NIRS phenotypic data could provide useful mapping results when compared to the reference phenotypic data. Several common QTLs were identified for FY and WA by both sets of data. The QTL on chromosome 4D was a consistently recurring QTL region for both traits. The QTL on chromosome 2A was positively linked to protein content which was supported by genetic correlation data. The results also indicated it was possible to obtain useful phenotypic data for mapping FY and WA using NIRS data. This would save time and costs as NIRS is quicker and cheaper than current rheology methods.

Improved chemical control of Conyza bonariensis in wheat limits problems in the following fallow

Conyza bonariensis is a major weed infesting zero-tilled cropping systems in subtropical Australia, particularly in wheat and winter fallows. Uncontrolled C.bonariensis survives to become a problem weed in the following crops or fallows. As no herbicide has been registered for C.bonariensis in wheat, the effectiveness of 11 herbicides, currently registered for other broad-leaved weeds in wheat, was evaluated in two pot and two field experiments. As previous research showed that the age of C.bonariensis, and to a lesser extent, the soil moisture at spraying affected herbicide efficacy, these factors also were investigated. The efficacy of the majority of herbicide treatments was reduced when large rosettes (5-15cm diameter) were treated, compared with small rosettes (<5cm diameter). However, for the majority of herbicide treatments, the soil moisture did not affect the herbicide efficacy in the pot experiments. In the field, a delay in herbicide treatment of 2 weeks reduced the herbicide efficacy consistently across herbicide treatments, which was related to weed age but not to soil moisture differences. Across all the experiments, four herbicides controlled C.bonariensis in wheat consistently (83-100%): 2,4-D; aminopyralid + fluroxypyr; picloram + MCPA + metsulfuron; and picloram + high rates of 2,4-D. Thus, this problem weed can be effectively and consistently controlled in wheat, particularly when small rosettes are treated, and therefore C.bonariensis will have a less adverse impact on the following fallow or crop.

Assessing the sustainability of wheat-based cropping systems using simulation modelling: sustainability

Concepts of agricultural sustainability and possible roles of simulation modelling for characterising sustainability were explored by conducting, and reflecting on, a sustainability assessment of rain-fed wheat-based systems in the Middle East and North Africa region. We designed a goal-oriented, model-based framework using the cropping systems model Agricultural Production Systems sIMulator (APSIM). For the assessment, valid (rather than true or false) sustainability goals and indicators were identified for the target system. System-specific vagueness was depicted in sustainability polygons-a system property derived from highly quantitative data-and denoted using descriptive quantifiers. Diagnostic evaluations of alternative tillage practices demonstrated the utility of the framework to quantify key bio-physical and chemical constraints to sustainability. Here, we argue that sustainability is a vague, emergent system property of often wicked complexity that arises out of more fundamental elements and processes. A 'wicked concept of sustainability' acknowledges the breadth of the human experience of sustainability, which cannot be internalised in a model. To achieve socially desirable sustainability goals, our model-based approach can inform reflective evaluation processes that connect with the needs and values of agricultural decision-makers. Hence, it can help to frame meaningful discussions, from which actions might emerge.