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Aims Cracks and biopores in compacted soil such as plough pans could aid deep rooting, mitigating constraints to seasonal upland use of paddy fields for rice production. This research investigated how soil macropores through a simulated plough pan affects root growth of contrasting deep and shallow rooting rice genotypes. Methods Deep rooting Black Gora and shallow rooting IR64 rice varieties were grown in packed cores of unsaturated soil in a controlled greenhouse. Simulated biopores and cracks (macropores) were inserted through the plough pan to form treatments with no macropores, biopores, cracks, and combined cracks and biopores. Different root parameters such as root length density (RLD), root volume, root diameter, number of root tips and branches were measured. The number of roots was calculated manually, including the number of roots growing through macropores in the plough pan layer. Results Plough pans with macropores had 25–32% more roots than with no macropores. RLD was 55% greater in the plough pan layer if cracks were present compared to biopores. Conversely, RLD was 26% less in subsoil if the plough pan had cracks compared to biopores. Different root parameters were greatly influenced by the presence of macropores in the plough pan, and deep-rooted Black Gora produced 81% greater RLD, 30% more root numbers and 103% more branching than the shallow rooted rice genotype IR64 within the plough pan layer. Conclusions Macropores greatly improve rice root growth through plough pans for a deep rooting but not a shallow rooting rice variety. Whereas cracks produce a greater number of roots in the plough pan, biopores result in greater root branching and root numbers deeper in subsoil.

Alternate wetting and drying (AWD) technique has been developed and evaluated on rice (Oryza sativa) systems in several countries worldwide for increasing water use efficiency and reducing negative effects of permanent flooding, like the increase in methane emissions and arsenic availability in soil. In this study, a paddy field experiment was carried out for two years to evaluate the application in Northern Italy rice area of a moderate AWD, i.e. only implemented during the vegetative phase of the crop and ponded water maintained thereafter, compared with Continuous Flooding (CF) system. The adaptability of 12 European commercial rice cultivars to AWD was investigated in terms of crop phenology, morphological traits, root production, nitrogen (N) uptake, yield, milled rice yield and microelement concentration in grains. Results showed substantial (40.7 %) water saving probably favoured by the presence of a shallow water table. In these pedoclimatic conditions, very limited effects of a mild AWD on crop status and final productivity were recorded and the commercial cultivars did not display significant different adaptabilities to the water stress. Moreover, AWD decreased arsenic (As) concentration in grain but increased grain Cadmium (Cd) being the degree of such a response dependent upon the variety, suggesting that the genotype plays an important role in this aspect of adaptation to AWD. info:eu-repo/semantics/acceptedVersion

Paddy soils in Bangladesh experience extensive irrigation with groundwater and surface water, both having variable geochemical constituents. The soils also have topological variations across the landscape. To understand the geochemical variability in the soils as affected by the different sources of irrigation water and the topographical variability, cultivation zones of paddy soils irrigated with both groundwater (n = 904) and surface water (n = 281) across Bangladesh were sampled and analyzed for a suit of seventeen geochemical elements. This study also assessed the extent and distribution of arsenic and the other geochemical elements in the paddy soils (n = 1209) as well as in a set of neighboring non-paddy soils (n = 235) within the different inundation land types (highland, medium highland-1, medium highland-2, medium lowland, lowland and very lowland) of Bangladesh. The mean concentrations of aluminum (26,000 mg/kg), cobalt (13 mg/kg), copper (32 mg/kg), iron (28,250 mg/kg), lead (18 mg/kg), magnesium (8050 mg/kg), molybdenum (1.02 mg/kg), nickel (41 mg/kg), potassium (4870 mg/kg), sodium (750 mg/kg) and zinc (70 mg/kg) in the surface water-irrigated paddy soils were found to be significantly (0.001 ≥ p ≤ 0.05) higher compared to the concentrations in the soils irrigated with groundwater (23,400; 12; 28; 25,650; 17; 7000; 0.96; 36; 4350; 600; and 62 mg/kg, respectively). Therefore, surface water used for paddy irrigation could increase the inputs of a number of toxic elements in the paddy soils having potential risk of crop contamination. Arsenic in the paddy and non-paddy soils varied significantly (F = 24.74, p F = 3.42, p

The AWD is an irrigation technology for rice cultivation, consisting in implementing alternate draining and flooded periods over the growing season, that delivers multiple environmental benefits, such as reduced water consumption, CH4 emissions and arsenic (As) grain content, but can be offset by yield losses. The trade-offs between the agronomic and environmental effects of AWD are crop context-dependent and they also vary among the different versions of AWD studied. Therefore, the implementation of a safe AWD needs to be preceded by studies conducted within a specific rice cropping system. A two-year field experiment was conducted to assess the effect of AWD on grain yield, As and heavy metal content in grains, and greenhouse gas emissions in nine representative European rice cultivars grown in a Mediterranean growing area. The experiment was performed in a split-plot design with four replications. The study revealed a significant cultivar effect on the agronomic response to AWD. Among the studied cultivars, one of them performed as tolerant to AWD while a group formed by four cultivars showed slight non-significant yield decline. AWD significantly reduced CH4 emissions and the global warming potential by 90% being such a large mitigation capacity explained by the negligible N2O emissions found in both water treatments. Finally, the implementation of AWD significantly reduced by ca. 40% As grain concentration but increased cadmium content, though the levels remained below the recommended thresholds. Further, AWD increased key nutritional elements like cupper, selenium, and zinc. In conclusion, this study confirms that AWD can be safely implemented in Mediterranean rice cultivation conditions with limited or null yield impact while obtaining the associated environmental benefits of this practice info:eu-repo/semantics/acceptedVersion

As the global population grows, demand on food production will also rise. For rice, one limiting factor effecting production could be availability of fresh water, hence adoption of techniques that decrease water usage while maintaining or increasing crop yield are needed. Alternative wetting and drying (AWD) is one of these techniques. AWD is a method by which the level of water within a rice field cycles between being flooded and nonflooded during the growth period of the rice crop. The degree to which AWD affects cultivars differently has not been adequately addressed to date. In this study, 22 rice cultivars, mostly landraces of the aus subpopulation, plus some popular improved indica cultivars from Bangladesh, were tested for their response to AWD across three different field sites in Bangladesh. Grain and shoot elemental concentrations were determined at harvest. Overall, AWD slightly increased grain mass and harvest index compared to plants grown under continually flooded (CF) conditions. Plants grown under AWD had decreased concentrations of nitrogen in their straw compared to plants grown under CF. The concentration of elements in the grain were also affected when plants were grown under AWD compared to CF: Nickel, copper, cadmium and iron increased, but sodium, potassium, calcium, cobalt, phosphorus, molybdenum and arsenic decreased in the grains of plants grown under AWD. However, there was some variation in these patterns across different sites. Analysis of variance revealed no significant cultivar × treatment interaction, or site × cultivar × treatment interaction, for any of the plant mass traits. Of the elements analyzed, only grain cadmium concentrations were significantly affected by treatment × cultivar interactions. These data suggest that there is no genetic adaptation amongst the cultivars screened for response to AWD, except for grain cadmium concentration and imply that breeding specifically for AWD is not needed.

As the world’s population increases, demands on staple crops like rice (Oryza sativa L.) will also increase, requiring additional fresh water supplies for irrigation of rice fields. Safe alternate wetting and drying (AWD) is a water management technique that is being adopted across a number of countries to reduce the water input for rice cultivation. The impact of AWD on plant growth, yield and grain quality is not well understood. A field trial of AWD was conducted at Mymensingh, Bangladesh over two boro (dry) seasons using eight field plots, four under AWD and four continuously flooded (CF). This manuscript describes the results of check cultivar BRRI dhan28 which was replicated in 35–40 rows per plot giving a total of 140–160 replicates per treatment. A study on the soil solution concentration of many elements indicated that manganese, iron, zinc, and arsenic were different under AWD conditions compared to CF on a number of sampling time points, but did not show a pattern related to the AWD treatment. A survey of soil strength using a penetrometer detected a small, but statistically significant, hardening of the surface soil of the AWD plots. At harvest the shoot and grain mass was significantly greater for the plants grown under AWD (9.0-9.4% and 12.0-15.4%, respectively) with the plants grown under AWD having a greater number of productive tillers. Physiological examination in the first year showed that although AWD decreased (∼21%) leaf elongation rate (LER) of recently transplanted seedlings during the first drying cycle, subsequent drying cycles did not affect LER, while tillering was slightly increased by AWD and there was evidence of higher leaf abscisic acid (ABA) in AWD plants. In the second year analysis of six phytohormones revealed that AWD increased plant foliar isopentenyladenine (iP) concentrations by 37% while leaf trans-zeatin concentrations decreased (36%) compared to CF plants. The elemental composition of the shoots and grains was also examined. In both years AWD decreased grain concentration of sulphur (by 4% and 15%), calcium (by 6% and 9%), iron (by 11% and 16%), and arsenic (by 14% and 26%), while it increased the grain concentration of manganese (by 19% and 28%), copper (by 81% and 37%), and cadmium (by 28% and 67%). These results indicate that plants grown under safe AWD conditions at this site have an increased grain mass compared to plants grown under CF, and this may be partly due to a high number of productive tillers. AWD decreases the concentration of arsenic in the grains in this site, but it elevates the concentration of cadmium.

Made available in DSpace on 2020-12-10T19:59:24Z (GMT). No. of bitstreams: 0 Previous issue date: 2020-02-01 Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) FAPEG-Goias Research Foundation FAPEMA-Maranhao Research Foundation United Kingdom, Biotechnology and Biological Sciences Research Council In weathered tropical soil, low nutrient use efficiency can lead to agricultural systems becoming unsustainable. Therefore, tropical agriculture is highly dependent on ecosystem services, such as nutrient recycling and carbon sequestration, to enhance soil fertility, increase nutrient uptake, and facilitate sustainable production of agricultural goods. This research aimed to find the balance between sustainability and profitability of tropical agriculture by evaluating the changes in soil caused by the ecosystem services provided by the biomass of leguminous trees (Gliricidia) and assessing how these changes (associated with potassium) can affect nitrogen-use efficiency and maize yield. An experiment was conducted testing the impact of Glircidia biomass addition vs. bare soil, with or without addition of both nitrogen and/or potassium. Changes in soil organic matter, (SOM) base cations sum, soil resistance, N uptake, N-use efficiency, and maize yield were evaluated. Gliricidia biomass, when used with N and K, contributed to increasing SOM by 5.0 g/kg and the sum of base cations by 1458. 65 kg/ha in the 0-30 cm layer. Moreover, grain yield was increased by approximately 70% in the treatments with Gliricidia when compared to treatments without biomass where yield was very low. In bare soil, the additional yield of 1.5 tons/ha would not be enough to convince farmers to change slash and burn to conventional bare soil systems. Our results showed that leguminous trees, such as Gliricidia, might contribute to ensuring sustainable agricultural intensification in humid tropical soils with low natural fertility by providing ecosystem services such as biomass production, carbon sequestration, base cation recycling, and increased N acquisition. These findings might be an important strategy to replace the common slash-and-burn-system and preserve the rainforest against the traditional shifting cultivation system. In contrast, the conventional system with bare soil showed that the addition of nitrogen was unfeasible, mainly in conditions of high rainfall precipitation. In these circumstances, the use of potassium may increase nitrogen-use efficiency only when biomass is not used. Sao Paulo State Univ, Coll Agr Sci, Dept Crop Sci, BR-18610307 Botucatu, SP, Brazil Univ Estadual Maranhao, Postgrad Program Agroecol, BR-65000000 Sao Luis, Maranhao, Brazil Fed Inst Educ Sci & Technol Piaui, BR-64860000 Urucui, Piaui, Brazil Univ Fed Maranhao, Dept Biol, BR-65080805 Sao Luis, Maranhao, Brazil Univ Aberdeen, Inst Biol & Environm Sci, Aberdeen AB24 3UU, Scotland Univ Nottingham, Sch Biosci, Sutton Bonington Campus, Loughborough LE12 5RD, Leics, England Sao Paulo State Univ, Coll Agr Sci, Dept Crop Sci, BR-18610307 Botucatu, SP, Brazil FAPESP: 2015/50305-8 FAPEG-Goias Research Foundation: 201510267001479 FAPEMA-Maranhao Research Foundation: RCUK-02771/16 United Kingdom, Biotechnology and Biological Sciences Research Council: BB/N013201/1

Background Rice is a global staple crop, being the main calorific component of many people living subsistence livelihoods. Rice can accumulate toxic elements such as arsenic, with the crop water management strongly affecting uptake. This study utilises the Bengal and Assam Aus Panel to conduct genome wide association (GWA) mapping for arsenic in shoots and grains of rice grown over 2 years under continually flooded (CF) and alternate wetting and drying (AWD). The aim was to assess genotype by water management interaction, identify quantitative trait loci (QTL) for arsenic accumulation, and propose candidate genes for lowering grain arsenic. Results AWD significantly reduced grain arsenic across all cultivars on average by 15.7 and 15.1% in year 1 and 2 respectively and shoot arsenic by 27.0% compared to the plants grown under CF. There was a weak cultivar by treatment interaction for grain for arsenic. All traits were strongly influenced by cultivar. GWA mapping identified a large number of 74 individual QTLs for arsenic, with six QTLs showing stability across years and/or water treatments. Three of the loci (one on chromosome 3, one on chromosome 4, and one on chromosome 5) were investigated in detail using an approach of clustering cultivars that had similar haplotypes for the QTL regions and then looking at the phenotypic values across the clusters. Two of the identified QTLs co-localised with known genes involved in arsenic accumulation, including Lsi2 which has not previously been reported to underlie a grain arsenic QTL. Conclusions This study has identified a number of novel QTLs for arsenic accumulation, as well as cultivars that consistently accumulate less arsenic over multiple field traits. The use of a haplotype clustering approach after GWA mapping has allowed for the effect, in terms of arsenic accumulation, to be determined for cultivars that share similar genomic sequence. Allocating nine high yielding Bangladeshi cultivars to these clusters has identified the potential of utilising these QTLs in breeding programmes; for example, incorporation of the QTL on chromosome 5 should decrease grain arsenic in elite high yielding Bangladeshi cultivars by 10% in all high yielding cultivars studied. Electronic supplementary material The online version of this article (10.1186/s12284-019-0307-9) contains supplementary material, which is available to authorized users.

Water saving techniques, such as alternate wetting and drying (AWD), are becoming a necessity in modern rice farming because of climate change mitigation and growing water use scarcity. Reducing water can vastly reduce methane (CH4) emissions; however, this net climate benefit may be offset by enhanced carbon dioxide (CO2) emissions from soil. The main aims of this study were: to determine the effects of AWD on yield and ecosystem C dynamics, and to establish the underlying mechanistic basis for observed trends in net ecosystem C gain or loss in an Italian rice paddy. We investigated the effects of conventional water management (i.e. conventionally flooded paddy; CF) and AWD on biomass accumulation (aboveground, belowground, grain), key ecosystem C fluxes (net ecosystem exchange (NEE), net primary productivity (NPP), gross primary productivity (GPP), ecosystem respiration (ER), autotrophic respiration (RA), heterotrophic respiration (RH)), and soil organic matter (SOM) decay for four common commercial European rice cultivars. The most significant finding was that neither treatment nor cultivar affected NEE, GPP, ER or SOM decomposition. RA was the dominant contributor to ER for both CF and AWD treatments. Cultivar and treatment affected the total biomass of the rice plants; specifically, with greater root production in CF compared to AWD. Importantly, there was no effect of treatment on the overall yield for any cultivar. Possibly, the wetting-drying cycles may have been insufficient to allow substantial soil C metabolism or there was a lack of labile substrate in the soil. These results imply that AWD systems may not be at risk of enhancing soil C loss, making it a viable solution for climate change mitigation and water conservation. Although more studies are needed, the initial outlook for AWD in Europe is positive; with no net loss of soil C from SOM decomposition, whilst also maintaining yield., Graphical abstract Unlabelled Image, Highlights • Alternate wetting and drying does not enhance carbon loss in European rice paddies. • Aerobic conditions do not increase soil organic matter decomposition. • Periods of drought do not affect rice yields in four European rice cultivars. • Alternate wetting and drying reduces rice root production.

© 2018, The Author(s). Background and aims: Alternate wetting and drying (AWD) saves water in paddy rice production but could influence soil physical conditions and root growth. This study investigated the interaction between contrasting rice genotypes, soil structure and mechanical impedance influenced by hydraulic stresses typical of AWD. Methods: Contrasting rice genotypes, IR64 and deeper-rooting Black Gora were grown in various soil conditions for 2 weeks. For the AWD treatments the soil was either maintained in a puddled state, equilibrated to −5kPa (WET), or dried to −50kPa and then rewetted at the water potential of −5kPa (DRY-WET). There was an additional manipulated macropore structure treatment, i.e. the soil was broken into aggregates, packed into cores and equilibrated to −5kPa (REPACKED). A flooded treatment (puddled soil remained flooded until harvest) was set as a control (FLOODED). Soil bulk density, penetration resistance and X-ray Computed Tomography (CT) derived macropore structure were measured. Total root length, root surface area, root volume, average diameter, and tip number were determined by WinRhizo. Results: AWD induced formation of macropores and slightly increased soil mechanical impedance. The total root length of the AWD and REPACKED treatments were 1.7–2.2 and 3.5–4.2 times greater than that of the FLOODED treatment. There was no significant difference between WET and DRY-WET treatments. The differences between genotypes were minimal. Conclusions: AWD influenced soil physical properties and some root characteristics of rice seedlings, but drying soil initially to −50kPa versus −5kPa had no impact. Macropores formed intentionally from repacking caused a large change in root characteristics.

In a large scale survey of rice grains from markets (13 countries) and fields (6 countries), a total of 1578 rice grain samples were analysed for lead. From the market collected samples, only 0.6% of the samples exceeded the Chinese and EU limit of 0.2 μg g − 1 lead in rice (when excluding samples collected from known contaminated/mine impacted regions). When evaluating the rice grain samples against the Food and Drug Administration's (FDA) provisional total tolerable intake (PTTI) values for children and pregnant women, it was found that only people consuming large quantities of rice were at risk of exceeding the PTTI from rice alone. Furthermore, 6 field experiments were conducted to evaluate the proportion of the variation in lead concentration in rice grains due to genetics. A total of 4 of the 6 field experiments had significant differences between genotypes, but when the genotypes common across all six field sites were assessed, only 4% of the variation was explained by genotype, with 9.5% and 11% of the variation explained by the environment and genotype by environment interaction respectively. Further work is needed to identify the sources of lead contamination in rice, with detailed information obtained on the locations and environments where the rice is sampled, so that specific risk assessments can be performed.

Root research requires high throughput phenotyping methods that provide meaningful information on root depth if the full potential of the genomic revolution is to be translated into strategies that maximise the capture of water deep in soils by crops. A very simple, low cost method of assessing root depth of seedlings using a layer of herbicide (TRIK or diuron) buried 25 or 30 cm deep in soil-filled boxes of varying size is described that is suitable for screening hundreds or thousands of rice accessions in controlled environment conditions. Variation in cultivar sensitivity to the herbicide when injected into pots was detected but considered small in relation to the variation detected when the herbicide was buried. Using 32 rice cultivars previously characterised for root traits in rhizotron and hydroponic systems, 80% of variation in herbicide score at 35 days was explained by cultivar and herbicide score correlated strongly with rooting depth traits. Using 139 genotypes of the Bala × Azucena mapping population, heritability for herbicide symptoms reached 55% and quantitative trait loci were detected which match those previously reported in this population. In repeated experiments using different soils, the method did not always perform to its maximum potential (in terms of speed of symptom development or discrimination between cultivars). This was not due to degradation or reduced bio-availability of the herbicide in the soil but is believed to be due to the soil water content and water release characteristics as it relates to plant water use. Therefore, when using this technique, thorough preliminary experiments to determine the best water application regime for the particular combination of soil and environmental conditions are required. The method should be applicable to seedling stage screening of rice and other crops.

Background The development of relatively cheap and high throughput methods of genotyping and phenotyping plants offers the opportunity to explore local germplasm more thoroughly than before and should accelerate the identification of sources of genetic variation suitable for breeding. In this study, 135 Sri Lankan accessions, mostly identified as landraces, for which data was available at the International Rice Research Institute on drought scores were genotyped using a 384 SNP array and assessed for root depth using a newly developed buried herbicide method. Roots of 36 accessions were assessed using hydroponics and 12 using soil-filled rhizotrons to establish if variation in herbicide score could be attributed to root traits. Results Population structure based on the SNPs using STRUCTURE revealed six groups, being tropical japonica, aus and four indica subpopulations. Three of these indica subpopulations do not seem to be represented in the Rice Diversity Panel I (RDP1) of 372 global rice accessions and appear to represent genetic diversity so far poorly studied by the global scientific community. The herbicide score was highly discriminatory between landraces and correlated very strongly with hydroponic and rhizotron root traits. The mean herbicide score strongly differentiated between landraces according to the province and the latitude from which they were collected. It also differed between subpopulations, being high in indica 2 and tropical japonica and low in indica 1 and aus. Drought scores suggest that indica 2 is more drought resistant than the other groups. Correlations indicate that those landraces with high herbicide scores are more drought resistant in the vegetative stage. The landrace Niyan Wee, whose name in Sinhalese means “drought rice” belongs to the indica 2 subgroup, has high herbicide scores and deep roots. Conclusions Niyan Wee and other cultivars within the indica 2 subgroup should be a valuable source of breeding for drought resistance at least partly because of their superior root traits, not normally associated with the indica rice cultivars. Electronic supplementary material The online version of this article (doi:10.1186/s12284-016-0092-7) contains supplementary material, which is available to authorized users.

Elements in grain crops such as iron, zinc and selenium are essential in the human diet, whereas elements such as arsenic are potentially toxic to humans. This study aims to identify quantitative trait loci (QTLs) for trace elements in rice grain. A field experiment was conducted in an arsenic enriched field site in Qiyang, China using the Bala x Azucena mapping population grown under standard field conditions. Grains were subjected to elemental analysis by inductively coupled plasma mass spectroscopy. QTLs were detected for the elemental composition within the rice grains, including for iron and selenium, which have previously been detected in this population grown at another location, indicating the stability of these QTLs. A correlation was observed between flowering time and a number of the element concentrations in grains, which was also revealed as co-localisation between flowering time QTLs and grain element QTLs. Unravelling the environmental conditions that influence the grain ionome appears to be complex, but from the results in this study one of the major factors which controls the accumulation of elements within the grain is flowering time.

• Inorganic arsenic (As(i) ) in rice (Oryza sativa) grains is a possible threat to human health, with risk being strongly linked to total dietary rice consumption and consumed rice As(i) content. This study aimed to identify the range and stability of genetic variation in grain arsenic (As) in rice. • Six field trials were conducted (one each in Bangladesh and China, two in Arkansas, USA over 2 yr, and two in Texas, USA comparing flooded and nonflood treatments) on a large number of common rice cultivars (c. 300) representing genetic diversity among international rice cultivars. • Within each field there was a 3-34 fold range in grain As concentration which varied between rice subpopulations. Importantly, As(i) correlated strongly with total As among a subset of 40 cultivars harvested in Bangladesh and China. • Genetic variation at all field sites was a large determining factor for grain As concentration, indicating that cultivars low in grain As could be developed through breeding. The temperate japonicas exhibited lower grain As compared with other subpopulations. Effects for year, location and flooding management were also statistically significant, suggesting that breeding strategies must take into account environmental factors.

Plant performance under drought stress is not solely defined by an inadequate water supply but by an interaction among many factors, including climatic, edaphic, and biological factors. An important interacting factor affecting root growth, and therefore the ability of a plant to access and take up water, is the soil physical environment. Soil penetration resistance can restrict, or even halt, root system growth. For rice, a soil penetration resistance of 1.4 MPa is sufficient to inhibit root system expansion. This review describes the effects of the soil physical environment on root growth and its interaction with drought stress. A large variation in soil penetration resistance exists among rainfed rice-growing areas of South and Southeast Asia and within experimental stations used for managed-drought field phenotyping. This variability may influence genotypic performance across experimental sites/countries and the response of crop genotypes to drought stress. A case study is presented in which differences in the soil physical environment may partially elucidate differences in experimental results between two field studies conducted at different locations. These results highlight the need for increased knowledge of environmental interactions to allow the outputs of genomics to increase drought tolerance at the field level.

Irrigation with arsenic contaminated groundwater in the Bengal Delta may lead to As accumulation in the soil and rice grain. The dynamics of As concentration and speciation in paddy fields during dry season (boro) rice cultivation were investigated at 4 sites in Bangladesh and West Bengal, India. Three sites which were irrigated with high As groundwater had elevated As concentrations in the soils, showing a significant gradient from the irrigation inlet across the field. Arsenic concentration and speciation in soil pore water varied temporally and spatially; higher As concentrations were associated with an increasing percentage of arsenite, indicating a reductive mobilization. Concentrations of As in rice grain varied by 2-7 fold within individual fields and were poorly related with the soil As concentration. A field site employing alternating flooded-dry irrigation produced the lowest range of grain As concentration, suggesting a lower soil As availability caused by periodic aerobic conditions.

It has previously been shown that across different arsenic (As) soil environments, a decrease in grain selenium (Se), zinc (Zn), and nickel (Ni) concentrations is associated with an increase in grain As. In this study we aim to determine if there is a genetic element for this observation or if it is driven by the soil As environment. To determine the genetic and environmental effect on grain element composition, multielement analysis using ICP-MS was performed on rice grain from a range of rice cultivars grown in 4 different field sites (2 in Bangladesh and 2 in West Bengal). At all four sites a negative correlation was observed between grain As and grain Ni, while at three of the four sites a negative correlation was observed between grain As and grain Se and grain copper (Cu). For manganese, Ni, Cu, and Se there was also a significant genetic interaction with grain arsenic indicating some cultivars are more strongly affected by arsenic than others.

Genetic analysis of heat tolerance will help breeders produce rice ( Oryza sativa L.) variet-ies adapted to future climates. An F 6 population of 181 recombinant of Aberdeen, Aberdeen, AB24 3UU, UK. J. Cairns, current address: inbred lines of Bala (toler-ant) × Azucena (susceptible) was screened for heat tolerance at anthesis by measuring spikelet fertility at 30°C (control) and 38°C (high temper-ature) in experiments conducted in the Philip-pines and the United Kingdom. The parents varied significantlyfor absolute spikelet fertility under control (79–87%) and at high tempera-ture (2.9–47.1%), and for relative spikelet fertility (high temperature/control) at high temperature (3.7–54.9%). There was no correlation between spikelet fertility in control and high-temperature conditions and no common quantitative trait loci (QTLs) were identified.Two QTLs for spike-let fertility under control conditions were identi-fiedon chromosomes 2 and 4. Eight QTLs for spikelet fertility under high-temperature condi-tions were identifiedon chromosomes 1, 2, 3, 8, 10, and 11. The most significantheat-responsive QTL, contributed by Bala and explaining up to 18% of the phenotypic variation, was identifiedon chromosome 1 (38.35 mega base pairs on the rice physical genome map). This QTL was also found to influenceplant height, explain-ing 36.6% of the phenotypic variation. A com-parison with other studies of abiotic (drought, cold, salinity) stresses showed QTLs at similar positions on chromosomes 1, 3, 8, and 10, sug-gesting common underlying stress-responsive regions of the genome.S.V.K. Jagadish, T.R. Wheeler, and P.Q. Craufurd, Plant Environment Lab., Univ. of Reading, Cutbush Lane, Shinfield, Reading, RG2 9AF, UK; J. Cairns and R. Lafitte, Crop and Environmental Sciences Divi-sion, International Rice Research Institute, DAPO Box 7777, Metro Manila, Philippines; A.H. Price, Dep. of Plant and Soil Science, Univ. CIMMYT, Km. 45, Carretera Mexico-Veracruz, El Batan, Texcoco, Edo. de Mexico, CP 56130, Mexico. Received 18 Sept. 2009. *Cor-responding author (p.craufurd@cgiar.org).

Arsenic (As) accumulation in rice grains is a risk to human health. The mechanism of transfer of As from the shoot into the grain during grain filling is unknown at present. In this study As speciation in the shoot and grains at maturity were examined, and the relationships between phosphorus (P) and As, and silicon (Si) and As were established in a wide range of cultivars grown in As contaminated field trials in Bangladesh and China. No correlations were observed between shoot and grain speciation, with the inorganic form comprising 93.0-97.0% of As in the shoot and 63.0-83.7% in the grains. The percentage of dimethylarsinic acid (DMA) was between 1.4 and 6.6% in the shoot and 14.6 and 37.0% in the grains; however, the concentrations were comparable, ranging from 0.07 to 0.26 mg kg(-1) in the shoots and 0.03 to 0.25 mg kg(-1) in the grains. A positive correlation was observed between shoot As and shoot Si, however, no correlation was observed between shoot Si and grain As. A significant negative correlation was observed between shoot P and grain As concentrations. These results suggest that the translocation of As into the grain from the shoots is potentially using P rather than Si transport mechanisms. The findings also indicate that inorganic As and DMA translocation to the grain differ considerably.

Sixteen bacterial strains isolated from the roots and rhizosphere of rice plants growing in saline and non-saline soils from the Shorkot area of Pakistan were tested for their ability to promote plant growth and reduce the incidence of rice blast disease. When applied to the soil, many of the isolated rhizobacterial strains increased seedling growth and/or suppressed rice blast disease in greenhouse-grown plants of the cultivars Super Basmati and Azucena, but each cultivar responded to different subsets of the bacteria. In the cv Super Basmati, increased blast resistance was correlated with the production of siderophores by the rhizobacteria. Several strains inhibited the growth of the causative agent of rice blast disease, the fungal pathogen Magnaporthe grisea, in an in vitro dual culture assay. Direct bioantagonism was correlated with disease resistance in Super Basmati, but not in Azucena, and direct antagonism as a cause for the reduced disease incidence is also unlikely since no epiphytic colonisation of leaves was detected. Rhizosphere colonisation by the bacteria in plants grown in sterile sand was correlated with disease resistance in Super Basmati, but not in Azucena. As well as the differences in strains that protected each cv against blast disease, we also found that there were differences in the ability of some strains to protect plants against blast depending on soil type. Hence, there are complex interactions between rhizobacteria and rice plants with respect to biocontrol of rice blast disease, dependent upon both rice cv and soil type. The identity of strains that promoted high levels of disease protection, including three that performed well across all plant cultivars and growth conditions, was determined by 16S rRNA gene sequencing.

A mapping population of 114 lines from Bala × Azucena was grown under drought stress at two field sites with contrasting soil physical properties. Drought was imposed between 35 and 65 days after sowing (DAS) and root density at 35 cm depth was measured 70 DAS. Leaf rolling, leaf drying and relative water content were recorded as indicators of drought avoidance. Root density correlated with indicators of drought avoidance. Two significant and two putative quantitative trait loci (QTLs) for root density and 28 QTLs for drought avoidance were identified. Most QTLs did not agree between sites. There was also reasonable agreement between leaf-drying QTLs and previously reported root-growth QTLs detected under controlled conditions (in contrast to a previous screen on soil with a higher penetration resistance). These data also reveal QTL × environment interaction, which will need to be understood more clearly if progress towards breeding for drought resistance via alterations of root morphology is to be achieved.

The concentration of arsenic (As) in rice grains has been identified as a risk to human health. The high proportion of inorganic species of As (As(i)) is of particular concern as it is a nonthreshold, class 1 human carcinogen. To be able to breed rice with low grain As, an understanding of genetic variation and the effect of different environments on genetic variation is needed. In this study, 13 cultivars grown at two field sites each in Bangladesh, India, and China are evaluated for grain As. There was a significant site, genotype, and site by genotype interaction for total grain As. Correlations were observed only between sites in Bangladesh and India, not between countries or within the Chinese sites. For seven cultivars the As was speciated which revealed significant effects of site, genotype, and site by genotype interaction for percentage As(i). Breeding low grain As cultivars that will have consistently low grain As and low As(i), over multiple environments using traditional breeding approaches may be difficult, although CT9993-5-10-1-M, Lemont, Azucena, and Te-qing in general had low grain As across the field sites.

Research into the composition of cereal grains is motivated by increased interest in food quality. Here multi-element analysis is conducted on leaves and grain of the Bala x Azucena rice mapping population grown in the field. Quantitative trait loci (QTLs) for the concentration of 17 elements were detected, revealing 36 QTLs for leaves and 41 for grains. Epistasis was detected for most elements. There was very little correlation between leaf and grain element concentrations. For selenium, lead, phosphorus and magnesium QTLs were detected in the same location for both tissues. In general, there were no major QTL clusters, suggesting separate regulation of each element. QTLs for grain iron, zinc, molybdenum and selenium are potential targets for marker assisted selection to improve seed nutritional quality. An epistatic interaction for grain arsenic also looks promising to decrease the concentration of this carcinogenic element.

The identification of markers linked to genes contributing to drought resistance promises opportunities to breed high yielding rice varieties for drought prone areas. Several studies using different mapping populations have previously identified quantitative trait loci (QTLs) for traits theoretically related to drought resistance. A mapping population of 176 F 6 recombinant inbred lines (RILs) derived from two upland rice varieties with contrasting aboveground drought avoidance traits (Bala and Azucena) with a linkage map of 157 markers was used to map QTLs for aboveground leaf morphological and physiological traits related to drought avoidance. Plants were grown for 6 weeks under controlled environmental conditions with three replications. Leaves were excised and placed on a balance. The rate of leaf rolling and water loss was recorded, after which leaf area, dry weight and specific leaf area were characterized. A simple method of estimating time to stomatal closure was employed. A total of 13 QTLs were detected for leaf morphological traits, three for initial transpiration and four for the proportion of water loss required to reach a specific advanced state of leaf rolling. No QTLs were detected for time of stomatal closure or speed of leaf rolling, nor for either water loss or transpiration at stomatal closure despite clear parental differences and moderate heritabilities in most of these traits. The co-location of QTLs for traits measured here and for drought avoidance previously reported from field experiments on chromosome 1, 3 and 5 link the genetics of drought resistance to leaf dimensions and physiology. However, a physiological explanation for a QTL for drought avoidance on chromosome 7 remains elusive.

Modifying plant root systems is considered a means of crop improvement targeted to low-resource environments, particularly low nutrient and drought-prone agriculture. The identification of quantitative trait loci (QTLs) for root traits has stimulated marker-assisted breeding to this end, but different QTLs have been detected in different populations of the same species, and importantly, in the same population when grown in different experimental environments. The presence of QTL x environment interaction is implicated, and this must be characterised if the utility of the target QTLs is to be realised. Previous attempts to do this suffer from a lack of control over replicate environments and inadequate statistical rigour. The Bala x Azucena mapping population was grown in two replicate experiments of four treatment environments, a control, a low light, a low soil nitrogen and a low soil water treatment. After a 4 weeks growth, maximum root length, maximum root thickness, root mass below 50 cm, total plant dry mass, % root mass and shoot length were measured. A summary of the overall results is presented in an accompanying paper. Here, QTL analysis by composite interval mapping is presented. A total of 145 QTLs were detected, mapping to 37 discrete loci on all chromosomes. Superficial evidence of QTL x E (great difference in LOD score) was tested by single-marker analysis which confirmed QTL x E for five loci representing only five individual trait-loci interactions. Some loci appeared to be stable across environments. Some QTLs were clearly more or less active under low light, low nitrogen or drought. A few notable loci on chromosomes 1, 2, 3, 5, 7 and 9 are briefly discussed. Also discussed are some remaining statistical shortcomings that will be addressed in another companion paper.

Altering root system architecture is considered a method of improving crop water and soil nutrient capture. The analysis of quantitative trait loci (QTLs) for root traits has revealed inconsistency in the same population evaluated in different environments. It must be clarified if this is due to genotype x environment interaction or considerations of statistics if the value of QTLs for marker-assisted breeding is to be estimated. A modified split-plot design was used where a main plot corresponded to a separate experiment. The main plot factor had four treatments (environments), which were completely randomized among eight trials, so that each treatment was replicated twice. The sub-plot factor consisted of 168 recombinant inbreed lines of the Bala x Azucena rice mapping population, randomly allocated to the seven soil-filled boxes. The aim of the trial was to quantify QTL x environment interaction. The treatments were chosen to alter partitioning to roots; consisting of a control treatment (high-soil nitrogen, high light and high-water content) and further treatments where light, soil nitrogen or soil water was reduced singly. After 4 weeks growth, maximum root length (MRL), maximum root thickness, root mass below 50 cm, total plant dry mass (%), root mass and shoot length were measured. The treatments affected plant growth as predicted; low nitrogen and drought increased relative root partitioning, low-light decreased it. The parental varieties Bala and Azucena differed significantly for all traits. Broad-sense heritability of most traits was high (57-86%). Variation due to treatment was the most important influence on the variance, while genotype was next. Genotype x environment interaction was detected for all traits except MRL, although the proportion of variation due to this interaction was generally small. It is concluded that genotype x environment interaction is present but less important than genotypic variation. A companion paper presents QTL x environment analysis of data.

Drought stress is a major constraint for rice (Oryza sativa L.) production and yield stability in rainfed ecosystems. Identifying genomic regions (QTLs) contributing in drought resistance will help to develop rice cultivars suitable for water-limiting environments through marker-assisted breeding. QTLs linked to physio-morphological and plant production traits under drought stress in the field were mapped by evaluating 177 F6 recombinant inbred (RI) lines of Bala × Azucena under rainfed conditions in the target environment (TE). The rice lines were subjected to severe drought stress during reproductive phase due to a natural rainfall failure event. The RI lines showed significant variation in physio-morphological and plant production traits under stress. A total of 24 QTLs were identified for various traits under stress, which individually explained 4.6 to 22.3% phenotypic variation. Composite interval mapping detected three markers viz., RM3894, RG409 and G1073 on chromosomes 3 and 8 linked to grain yield under drought stress in TE, respectively explaining 22..3, 17.1 and 10.9% of phenotypic variation. QTLs for leaf drying, days to 50% flowering and number of productive tillers under drought stress co-located at certain of these regions. Further, QTLs for several root traits overlapped with QTLs for grain yield under stress in these RI lines, indicating the pleiotropic effects of root trait QTLs on rice performance under stress. Correlation coefficients between potential root traits determined in another study and plant production under stress in this study were not significant in these RI lines. Consistent QTLs for drought resistance traits and yield under drought stress in TE were detected and might be useful for rainfed rice improvement.

Root morphological characteristics are known to be important in the drought resistance of some rice (Oryza sativa L.) varieties. The identification of quantitative trait loci (QTLs) associated with root morphology and other drought resistance-related traits should help breeders produce more drought resistant varieties. Stability in the expression of root growth QTL across rooting environments is critical for their use in breeding programs. A greenhouse experiment in which a mapping population of 140 recombinant inbred lines and the parental varieties Bala and Azucena were grown in glass-sided soil chambers and evaluated for root growth and water uptake was conducted. In each of 2 years, two treatments were used; an early water-deficit (WD0) in which seeds were sown into wet soil but received no more water, and a late water-deficit (WD49) in which the plants were watered for 49 days and then received no water for a week. The major differences between treatments and years in dry matter partitioning and root growth traits are reported elsewhere. Here, the identification of QTLs for root growth traits by composite interval mapping is described. At LOD>3.2, there were six QTLs for the weight of roots below 90 cm and maximum root length, 11 for root to shoot ratio, 12 for the number of roots past 100 cm, and 14 for root thickness. A total of 24 regions were identified as containing QTLs (these regions often contained several QTLs identified for different root traits). Some were revealed only in individual experiments and/or for individual traits, while others were common to different traits or experiments. Seven QTLs, on chromosomes 1, 2, 4, 7, 9 (two QTLs) and 11, where considered particularly noteworthy. The complex results are discussed in the context of previously reported QTLs for root growth in other populations, the interaction between QTL with the environment and the value of QTLs for breeding.

Root morphological characteristics are known to be important in the drought resistance of some rice ( Oryza sativa L.) varieties. As part of a research programme aimed at using molecular marker technology for the improvement of drought resistance in rice, it is necessary to identify quantitative trait loci (QTLs) associated with root morphology and other drought resistance-related traits. Stability across environments is critically important for the identification of root growth QTLs that are valuable in breeding. Here, a mapping population of 140 recombinant inbred lines and the parental varieties Bala and Azucena were screened for root growth in thin glass-sided soil-filled chambers. In each of 2 years, two treatments were used: an early water deficit (WD0) in which seeds were sown into wet soil but received no more water, and a late water deficit (WD49) in which the plants were watered for 49 days and water was withheld for a final 7 days. Plant height, visible root growth, soil moisture at four depths and water use were monitored weekly. After 56 days, shoot mass, relative water content, root mass in four depth sections and adventitious root thickness at two depths were measured. In the WD0 treatment, shoot growth slowed markedly, while maximum root length was increased, indicating major changes in partitioning. This change in partitioning was also revealed as major differences in root mass (total and at depth) and root to shoot ratio between treatments. There were also large differences in partitioning between similar treatments in different years, which may in part reflect plant responses to soil water and nutrient status. In both treatments, water was extracted first from the upper sections of the rooting zone and then from progressively deeper layers. Effective water extraction required a root length density of at least 0.4 cm cm −3 in both treatments, and in the WD49 treatment water extraction at 100 cm depth was also related to shoot size and stomatal conductance. Azucena had thicker roots and more roots at depth compared to Bala, which slowed shoot growth sooner and became less water-stressed than Azucena. Azucena has root traits that potentially contribute to drought resistance, while Bala has a number of shoot-related mechanisms that make it adapted to drought-prone environments. The data presented here have been used to identify QTLs for root growth traits in a companion paper.

Localizing genes that contribute to drought avoidance in a quantitative way should enable the exploitation of these genes in breeding through marker-assisted selection, and may lead to the discovery of gene identity and function. Between 110 and 176 F6 recombinant inbred lines from a mapping population derived from a cross of upland rice varieties Bala and Azucena have been evaluated for indicators of drought avoidance in sites in the Philippines and West Africa over two dry seasons. A molecular map with 102 RFLP, 34 AFLP and six microsatellite markers has been used to map (by composite interval mapping) quantitative trait loci (QTLs) for the visual scores of leaf rolling and leaf drying and leaf relative water content. QTLs were mapped for each site and across sites. A total of 17 regions were identified which contained QTLs with a LOD score greater than 3.2. For leaf rolling, Bala was the parent contributing the majority of positive alleles whilst for the other traits, Bala and Azucena contributed more evenly. Six of the 17 regions influenced more than one trait, explaining the phenotypic correlations between traits that were observed. Three QTLs appeared to be specific to the Philippines experiments. One QTL had opposing effects in the Philippines and West Africa. QTLs for relative water content were detected on chromosome 8, congruent with an osmotic adjustment QTL identified in another population. Only three of the QTLs identified here have not been reliably identified in the two other populations that have been screened for drought avoidance. By using several populations assessed for drought avoidance in different sites, the distribution and utility of QTLs for drought avoidance in rice is being elucidated.

The use of molecular markers in the mapping of traits of agronomic importance holds great promise for speeding the development of improved plant varieties and increasing our understanding of the physiological or molecular mechanisms behind biological phenomena. The technique is now being applied to drought resistance in rice (Oryza sativa L.). Drought is important because a considerable proportion of the world rice area is not irrigated and can be prone to water deficit. A large number of people, particular some of the poorest rice farmers, stand to gain if new varieties which combine high yield and drought resistance can be developed. Rice should be particularly useful for the molecular genetic analysis of drought resistance because of its growing role as a model monocot species and the diversity of drought resistance mechanisms which are found in the germplasm. We briefly review the traits which might be considered important in improving drought resistance in rice, before explaining the molecular mapping approach. We review progress at locating quantitative trait loci (QTLs) for individual mechanisms of drought resistance in controlled environment conditions. This includes a detailed comparison of reported QTLs related to root morphological characters. The search for QTLs associated with field performance under drought stress is analysed and the problems associated with understanding the genetic control of a complex physical and physiological phenomenon under conditions of substantial environmental variation are highlighted. We emphasise that the use of near isogenic lines in overcoming some of the problems offers considerable promise for the future.

summary Improving the drought resistance of high yielding rice (Oryza sativa L.) varieties for upland areas is a goal of rice breeders since upland rice relies exclusively on rainfall for water and is generally low yielding. Identifying quantitative trait loci (QTLs) which confer drought resistance promises to speed this goal. We have previously described the use of 82 restriction fragment length polymorphism (RFLP) markers in the genetic mapping of root growth characteristics that are potentially involved in drought resistance in a population of 178 F2 plants derived from a cross of two drought resistant upland varieties, Azucena and Bala. Here we report the characterization and QTL mapping of three other mechanisms of drought resistance, leaf rolling, stomatal behaviour and early flowering, in this population. When the youngest fully expanded leaf was excised from 12 rice varieties,‘IR20’and‘Bala’ were identified as slow at rolling. Although ‘Azucena’ rolled its leaves much faster than ‘Bala’ upon excision, stomatal resistance rose more rapidly in‘Bala’, resulting in a lower rate of water loss. By assessing leaf rolling and stomatal resistance in the F2 population, it was possible to identify QTLs for these traits. A QTL for slow leaf rolling from ‘Bala’ was found on chromosome 1. Two QTLs associated with resting stomatal conductance were found on chromosomes 3 and 12. QTLs for the rate of stomatal closure were found on chromosomes 3 and 7. In addition, it was possible to map QTLs associated with the days to heading which were found on chromosomes 3, 8 and 10. These results identify markers of potential value to breeders and are discussed in the context of previously reported data for heading date QTLs in rice and for stomatal conductance QTLs in maize.

Root growth is an important component of the adaptation of rice to drought-prone environments. A hydroponic screen was used to study root growth of 28 rice varieties. Both maximum root length and adventitious root thickness varied widely between varieties. In general, japonica varieties had larger root systems than indica varieties. Two F2 populations involving the thick- and long-rooted upland japonica variety ‘Azucena’ and two poor-rooting varieties, namely the upland indica‘Bala’ and the Italian japonica‘Maratelli’, were made and screened in hydroponics. Generation means analysis revealed significant additive and dominance main effects for the root length traits with a prevalence of dominance gene effects in both crosses. The dominance×dominance type of non-allelic interactions were important for maximum root length from day 7 to day 28, root volume, root thickness and root cell length in the cross ‘Bala’בAzucena’. The heritability (broad-sense) estimates varied from low to high for the traits and displayed differences between populations. This suggested that recombinant lines with improved root traits can be developed from the two crosses with selection methods that involve some form of progeny evaluation. In a companion paper, we report the mapping of quantitative trait loci (QTLs) for root growth traits in the ‘Bala’בAzucena’ population using restriction fragment length polymorphisms (RFLPs).

The crop management practice of alternate wetting and drying (AWD) is being promoted by IRRI and the national research and extension program in Bangladesh and other parts of the world as a water-saving irrigation practice that reduces the environmental impact of dry season rice production through decreased water usage, and potentially increases yield. Evidence is growing that AWD will dramatically reduce the concentration of arsenic in harvested rice grains conferring a third major advantage over permanently flooded dry season rice production. AWD may also increase the concentration of essential dietary micronutrients in the grain. However, three crucial aspects of AWD irrigation require further investigation. First, why is yield generally altered in AWD? Second, is AWD sustainable economically (viability of farmers' livelihoods) and environmentally (aquifer water table heights) over long-term use? Third, are current cultivars optimized for this irrigation system? This paper describes a multidisciplinary research project that could be conceived which would answer these questions by combining advanced soil biogeochemistry with crop physiology, genomics, and systems biology. The description attempts to show how the breakthroughs in next generation sequencing could be exploited to better utilize local collections of germplasm and identify the molecular mechanisms underlying biological adaptation to the environment within the context of soil chemistry and plant physiology.

Cereal grains are the dominant source of cadmium in the human diet, with rice being to the fore. Here we explore the effect of geographic, genetic, and processing (milling) factors on rice grain cadmium and rice consumption rates that lead to dietary variance in cadmium intake. From a survey of 12 countries on four continents, cadmium levels in rice grain were the highest in Bangladesh and Sri Lanka, with both these countries also having high per capita rice intakes. For Bangladesh and Sri Lanka, there was high weekly intake of cadmium from rice, leading to intakes deemed unsafe by international and national regulators. While genetic variance, and to a lesser extent milling, provide strategies for reducing cadmium in rice, caution has to be used, as there is environmental regulation as well as genetic regulation of cadmium accumulation within rice grains. For countries that import rice, grain cadmium can be controlled by where that rice is sourced, but for countries with subsistence rice economies that have high levels of cadmium in rice grain, agronomic and breeding strategies are required to lower grain cadmium.

Interest in belowground plant growth is increasing, especially in relation to arguments that shallow-rooted cultivars are efficient at exploiting soil phosphorus while deep-rooted ones will access water at depth. However, methods for assessing roots in large numbers of plants are diverse and direct comparisons of methods are rare. Three methods for measuring root growth traits were evaluated for utility in discriminating rice cultivars: soil-filled rhizotrons, hydroponics and soil-filled pots whose bottom was sealed with a non-woven fabric (a potential method for assessing root penetration ability). A set of 38 rice genotypes including the OryzaSNP set of 20 cultivars, additional parents of mapping populations and products of marker-assisted selection for root QTLs were assessed. A novel method of image analysis for assessing rooting angles from rhizotron photographs was employed. The non-woven fabric was the easiest yet least discriminatory method, while the rhizotron was highly discriminatory and allowed the most traits to be measured but required more than three times the labour of the other methods. The hydroponics was both easy and discriminatory, allowed temporal measurements, but is most likely to suffer from artefacts. Image analysis of rhizotrons compared favourably to manual methods for discriminating between cultivars. Previous observations that cultivars from the indica subpopulation have shallower rooting angles than aus or japonica cultivars were confirmed in the rhizotrons, and indica and temperate japonicas had lower maximum root lengths in rhizotrons and hydroponics. It is concluded that rhizotrons are the preferred method for root screening, particularly since root angles can be assessed.

Arsenic accumulation in rice grain has been identified as a major problem in some regions of Asia. A study was conducted to investigate the effect of increased organic matter in the soil on the release of arsenic into soil pore water and accumulation of arsenic species within rice grain. It was observed that high concentrations of soil arsenic and organic matter caused a reduction in plant growth and delayed flowering time. Total grain arsenic accumulation was higher in the plants grown in high soil arsenic in combination with high organic matter, with an increase in the percentage of organic arsenic species observed. The results indicate that the application of organic matter should be done with caution in paddy soils which have high soil arsenic, as this may lead to an increase in accumulation of arsenic within rice grains. Results also confirm that flooding conditions substantially increase grain arsenic.

Altering root morphology of rice (Oryza sativa L.) cultivars could improve yields in drought-prone upland ecosystems. Marker-assisted backcross breeding was used to introgress four QTLs for root traits into an upland rice cultivar. The QTLs had previously been identified under experimental conditions in a different genetic background. The introgressed lines and the recurrent parent were grown for 6 years by resource-poor farmers in upland sites in Eastern India and yields recorded. In combination the QTLs significantly increased yield by 1 t ha(-1) under relatively favourable field conditions. In less favourable trials, the QTL effects were not detected due to greater heterogeneity in soil-water availability in very low yielding environments and consequent yield variability. Root studies under controlled conditions showed that lines with the introgressions had longer roots throughout tillering than the recurrent parent (14 cm longer 2 weeks after sowing). Therefore, both improved roots and increased yield can be attributed to the introgression of QTLs. This is the first demonstration that marker-assisted backcross breeding (MABC) to introgress multiple root QTLs identified under controlled conditions is an effective strategy to improve farmers' yields of upland rice. The strategy was used to breed a novel upland rice cultivar that has been released in India as Birsa Vikas Dhan 111.

Thermal imaging using infrared (IR) is now an established technology for the study of stomatal responses and for phenotyping plants for differences in stomatal behaviour. This paper outlines the potential applications of IR sensing in drought phenotyping, with particular emphasis on a description of the problems with extrapolation of the technique from the study of single leaves in controlled environments to the study of plant canopies is field plots, with examples taken from studies on grapevine (Vitis vinifera L.) and rice (Oryza sativa L.). Particular problems include the sensitivity of leaf temperature (and potentially the temperature of reference surfaces) to both temporal and spatial variation in absorbed radiation, with leaf temperature varying by as much as 15°C between full sun and deep shade. Examples of application of the approach to phenotyping in the field and the steps in data analysis are outlined, demonstrating that clear genotypic variation may be detected despite substantial variation in soil moisture status or incident radiation by the use of appropriate normalisation techniques.

For the world's population, rice consumption is a major source of inorganic arsenic (As), a nonthreshold class 1 carcinogen. Reducing the amount of total and inorganic As within the rice grain would reduce the exposure risk. In this study, grain As was measured in 76 cultivars consisting of Bangladeshi landraces, improved Bangladesh Rice Research Institute (BRRI) cultivars, and parents of permanent mapping populations grown in two field sites in Bangladesh, Faridpur and Sonargaon, irrigated with As-contaminated tubewell water. Grain As ranged from 0.16 to 0.74 mg kg(-1) at Faridpur and from 0.07 to 0.28 mg kg(-1) at Sonargaon. Highly significant cultivar differences were detected and a significant correlation (r = 0.802) in the grain As between the two field sites was observed, indicating stable genetic differences in As accumulation. The cultivars with the highest concentration of grain As were the Bangladeshi landraces. Landraces with red bran had significantly more grain As than the cultivars with brown bran. The percent of inorganic As decreased linearly with increasing total As, but genetic variation within this trend was identified. A number of local cultivars with low grain As were identified. Some tropical japonica cultivars with low grain As have the potential to be used in breeding programs and genetic studies aiming to identify genes which decrease grain As.

During the last 10 years, a large number of quantitative trait loci (QTLs) controlling rice root morphological parameters have been detected in several mapping populations by teams interested in improving drought resistance in rice. Compiling these data could be extremely helpful in identifying candidate genes by positioning consensus QTLs with more precision through meta-QTL analysis. We extracted information from 24 published papers on QTLs controlling 29 root parameters including root number, maximum root length, root thickness, root/shoot ratio, and root penetration index. A web-accessible database of 675 root QTLs detected in 12 populations was constructed. This database includes also all QTLs for drought resistance traits in rice published between 1995 and 2007. The physical position on the pseudo-chromosomes of the markers flanking each QTL was determined. An overview of the number of root QTLs in 5-Mb segments covering the whole genome revealed the existence of “hot spots,” The 32 trait × chromosome combinations comprising six or more QTLs were subjected to a meta-QTL analysis using the software package MetaQTL. The method enabled us both to determine the likely number of true QTLs in these areas using an Akaike information criterion and to estimate their position. The meta-QTL confidence intervals were notably reduced and, for the smallest ones, encompassed only a few genes.

Summary • Striga is a parasitic weed attacking mainly maize, sorghum, millet and cowpea. Studying the interaction between rice and Striga is valuable since rice is a model monocot. In this paper, the susceptibility of different rice cultivars to S. hermonthica was tested and quantitative trait loci (QTL) for Striga tolerance mapped on the Bala × Azucena F6 population. • Seven rice cultivars were grown with and without S. hermonthica for 14 wk. For the mapping experiment, 115 recombinant inbred lines (RILs), along with Azucena and Bala, were grown with and without Striga for 11 wk. • Rice cultivars tested had different susceptibilities to Striga, ranging from highly susceptible to completely resistant. Azucena and Bala differed in the speed of Striga emergence and the impact on host growth. A genomic region between positions 139 and 166 cM on chromosome 1 was identified containing strong QTL (LOD = 4.9–15.7) for all traits measured. • This indicates that genes for Striga tolerance exist in rice germplasm and the mapped QTL can be further studied to promote understanding of the nature of resistance/tolerance and breeding for Striga-resistant crop plants.

Summary • The root-knot nematode Meloidogyne graminicola is an obligate biotrophic parasite and a major pest of rice (Oryza sativa) for which resistant varieties are not currently available. Quantitative trait loci (QTLs) for partial resistance to M. graminicola were identified using a mapping population based on two rice varieties, Bala × Azucena. • Experiments were carried out to investigate the interactions between M. graminicola and these two varieties in terms of nematode establishment, reproduction and effect on rice yield. Nematode establishment was also assessed in the mapping population. • Meloidogyne graminicola consistently caused more galling and had higher reproductive success in Azucena than in Bala. M. graminicola did not significantly reduce yield in Bala, but caused a yield reduction of almost half in Azucena, suggesting that the partial resistance to nematode establishment was related to nematode tolerance. A total of six significant or putative QTLs for nematode tolerance were detected. • For two of the QTLs detected, Azucena was the donor of the tolerance alleles, suggesting it may be possible to breed plants with greater tolerance than Bala.

The rapid progress in rice genotyping must be matched by advances in phenotyping. A better understanding of genetic variation in rice for drought response, root traits, and practical methods for studying them are needed. In this study, the OryzaSNP set (20 diverse genotypes that have been genotyped for SNP markers) was phenotyped in a range of field and container studies to study the diversity of rice root growth and response to drought. Of the root traits measured across more than 20 root experiments, root dry weight showed the most stable genotypic performance across studies. The environment (E) component had the strongest effect on yield and root traits. We identified genomic regions correlated with root dry weight, percent deep roots, maximum root depth, and grain yield based on a correlation analysis with the phenotypes and aus, indica, or japonica introgression regions using the SNP data. Two genomic regions were identified as hot spots in which root traits and grain yield were co-located; on chromosome 1 (39.7-40.7 Mb) and on chromosome 8 (20.3-21.9 Mb). Across experiments, the soil type/ growth medium showed more correlations with plant growth than the container dimensions. Although the correlations among studies and genetic co-location of root traits from a range of study systems points to their potential utility to represent responses in field studies, the best correlations were observed when the two setups had some similar properties. Due to the co-location of the identified genomic regions (from introgression block analysis) with QTL for a number of previously reported root and drought traits, these regions are good candidates for detailed characterization to contribute to understanding rice improvement for response to drought. This study also highlights the utility of characterizing a small set of 20 genotypes for root growth, drought response, and related genomic regions.

Rice is the one of the oldest crop cereals in Asia and has been grown since ancient times. In the present study, a rice diversity panel was exposed to drought and drought recovery was scored to identify QTLs and candidate genes related to drought resistance. There are no reports of QTL mapping using Genome wide association mapping for drought recovery has been published. Only one significant association on chromosome 2 for drought recovery with physical position at 24559374 bp was found. positional candidate genes underneath QTL was examined bioinformatically and through the literature revealing several interesting genes which may offer potential for developing drought resistant rice cultivars.

Two upland rice varieties, Azucena and Bala, were screened for root growth under droughted and irrigated treatments in two field sites at the West Africa Rice Development Association (WARDA) experimental farm, Cote d’Ivoire, during the dry season of 1999/2000. The sites were chosen to represent contrasting soil profile penetration resistance (PR) characteristics on upland sites, although both were relatively impeding. The number of nodal root axes per unit area passing through horizontal transects (root density) was counted at 35, 56, 77 and 98 days after sowing (DAS) at 10 cm depth intervals. Azucena consistently maintained a greater root density than Bala and a greater proportion of Azucena roots grew to 30 cm depth (22.7% vs. 8.4% at 77 DAS). There was little detectable effect of water regime on root distribution but evidence of lower root numbers at depths below 20 cm in the higher PR site was revealed. A site by variety by soil depth interaction suggests that Azucena roots are more strongly affected by very high PR than those of Bala. PR between 0–30 cm depth increased greatly with decreasing soil water content during the drought as the soil dried. This increase is likely to have prevented or greatly impaired further nodal root growth within this layer. At 40 cm depth, PR was high (3–4 MPa) but did not increase during the drought. At this depth root growth rate was likely to be greatly reduced despite the availability of water. These results demonstrate that varietal differences in root morphology characterised in the laboratory can be also detected in impeding field soils as differences in the density of roots at depth. Relatively poor root growth in these fields in the absence of drought was probably due to the high mechanical impedance and/or the physiological stress of the plants in the dry season. Our results indicate that high mechanical impedance was a more fundamental constraint on root growth than soil water availability during the drought. Thus, varietal differences in root penetration ability might be very important for drought avoidance in soils of this type.

A population of recombinant inbred rice lines from a cross between the upland japonica cultivar Azucena and the upland indica cultivar Bala was evaluated in a series of upland field experiments. Water stress was imposed during the reproductive stage by managed irrigation during the dry season, while control treatments were maintained in aerobic, well-irrigated conditions. Water deficit resulted in a yield reduction of 17 to 50%. The genetic correlation between stress and control yields was quite high when stress was mild, and the heritability of yield was similar in stress and control treatments across both years of this study. Genetic correlations between secondary traits such as leaf rolling and drying and yield under stress varied from high (leaf drying) to insignificant (leaf rolling). Lines with superior yield tended to have fewer panicles and larger grain size than the high-yielding parent, Bala, even though the panicle number was positively correlated with yield and the thousand-grain weight was not associated with yield for the population as a whole. Analysis of quantitative trait loci (QTLs) for yield and yield components allowed the identification of 31 regions associated with growth or yield components. Superior alleles came from either parent. Several of the regions identified had also been reported for root mass at depth or maximum root length in this population in other studies made under controlled environments, and for leaf drying (LD) in field studies. However, the direction of the effect of QTLs was not consistent, which indicates that there was not necessarily a causal relationship between these secondary traits and performance. We conclude that mapping populations can provide novel insights on the actual relationships between yield components and secondary traits in stress and control environments and can allow identification of significant QTLs for yield components under drought stress.