Development of drought-resistant cultivars using physiomorphological traits in rice

Drought is a major problem for rice grown under rainfed lowland and upland conditions, but progress in breeding to improve drought resistance has been slow. This paper describes patterns of water-stress development in rice fields, reviews genetic variation in physio-morphological traits for drought resistance in rice, and suggests how knowledge of stress physiology can contribute to plant breeding programmes that aim to increase yield in water-limiting environments. To provide a basis for integrating physiological research with plant-breeding objectives we define drought resistance in terms of relative yield of genotypes. Therefore, a drought-resistant genotype will be one which has a higher grain yield than others when all genotypes are exposed to the same level of water stress.A major reason for the slow progress in breeding for drought resistance in rice is the complexity of the drought environment, which often results in the lack of clear identification of the target environment(s). There is a need to identify the relative importance of the three common drought types; early-season drought which often causes delay in transplanting, mild intermittent stress which can have a severe cumulative effect, and late stress which affects particularly late-maturing genotypes. In addition, in rainfed lowland rice, flooded and non-flooded soil conditions may alternate during the growing season, and affect nutrient availability or cause toxicity.Several drought-resistance mechanisms, and putative traits which contribute to them, have been identified for rice; important among these being drought escape via appropriate phenology, root characteristics, specific dehydration avoidance and tolerance mechanisms, and drought recovery. Some of these mechanisms/traits have been shown to confer drought resistance and others show potential to do so in rice. The most important is the appropriate phenology which matches crop growth and development with the water environment. A deep root system, with high root length density at depth is useful in extracting water thoroughly in upland conditions, but does not appear to offer much scope for improving drought resistance in rainfed lowland rice where the development of a hard pan may prevent deep root penetration. Under water-limiting environments, genotypes which maintain the highest leaf water potential generally grow best, but it is not known if genotypic variation in leaf water potential is solely caused by root factors. Osmotic adjustment is promising, because it can potentially counteract the effects of a rapid decline in leaf water potential and there is large genetic variation for this trait. There is genotypic variation in expression of green leaf retention which appears to be a useful character for prolonged droughts, but it is affected by plant size which complicates its use as a selection criterion for drought resistance.There is a general lack of drought related research for rice in rainfed lowland conditions. This needs to be rectified, particularly considering their importance relative to upland conditions in Asian countries. We suggest that focussing physiological-genetic research efforts onto clearly defined, major target environments should provide a basis for increasing the relevance of stress physiology and the efficiency of breeding programmes for development of drought-resistant genotypes.     

Genotypic variations in the plasticity of nodal root penetration through the hardpan during soil moisture fluctuations among four rice varieties

Rainfed lowland rice fields are characterized by soil moisture fluctuations (SMF) and the presence of hardpan that impedes deep rooting and thus limits water extraction from deep soil layer during the periods of drought. In this study, we used rootboxes with three layers; shallow layer, artificial hardpan, and deep and wet layer below the hardpan, to evaluate differences in the plasticity of nodal roots elongation through the hardpan and promote root branching below the hardpan in response to SMF among four rice varieties; Sasanishiki, Habataki, Nipponbare, and Kasalath. Experiments were conducted during the summer and autumn seasons. Plasticity was computed as the difference in root traits within each variety between the SMF and continuously well-watered treatments. In both experiments, Habataki consistently tended to exhibit higher root plasticity than the other three varieties by increasing number of nodal roots that penetrated the hardpan during rewatering period in SMF, when the soil moisture increased and penetration resistance decreased. This root plasticity then contributed to greater water use at the deeper soil during the subsequent drought period and overall shoot dry matter production. Habataki had significantly higher δ¹³C value in roots at deep layer than roots at the shallow and hardpan layers under SMF, which may indicate that these were relatively newly grown roots as a consequence of root plasticity. This study also indicates that CSSLs derived from Sasanishiki and Habataki varieties may be suitable for the analysis of QTLs associated with root plasticity expression in rainfed lowland with hardpan and experiencing SMF.     

Comparison of Root System Development in Two Rice Cultivars During Stress Recovery from Drought and the Plant Traits for Drought Resistance

Summary

A greenhouse experiment was conducted to compare root system development of two upland rice cultivars, IRAT 13 and Senshou, during recovery from drought stress and to identify the plant traits that confer drought resistance. From 62 days after sowing (62 DAS), drought stress was given for 6 d followed by rewatering for 14 d. Root length density (RLD) and root diameter (thickness) were measured at the end of the stress and rewatering periods. Control plants were well-watered throughout the study. Gultivar IRAT 13 had thicker roots and higher relative RLD (ratio of RLD in drought-stressed plants to that in control plants) than under drought stress, and significantly higher root growth recovery after rewatering cultivar Senshou. Related plant traits such as evapotranspiration (ET), leaf and stem dry weights and weight of senescent leaves (dead leaves) in IRAT 13 were significantly more favorable for drought resistance compared to Senshou.     

栽培稻抗旱性研究的现状与策略

 水资源短缺正成为制约我国农业发展的重要因素。培育抗旱的栽培稻品种并实现水稻旱作，不但可在很大程度上节约水资源，而且有利于增产稳产，节约能源和减少环境污染。抗旱性包括逃旱性、避旱性、耐旱性和复原抗旱性。形态生理学的研究揭示出大量的与栽培稻抗旱性有关的形态特征和生理特性，如根系和叶片性状、生育期、渗透调节、脱落酸含量与栽培稻抗旱性密切相关，且已利用分子标记对上述性状进行了基因定位（QTL）研究。旱稻品种改良也已取得重大进展。在进行抗旱品种改良的基础上，通过引进相应的栽培技术，节水种植，实现水稻旱作，并达到稳产与增产的目的，是抗旱性研究的战略目标。在增产、稳产和优质的前提下，以培育耐旱性极强的水稻（或旱稻）为中心，建立有代表性的抗旱性研究基地, 进一步加强稻属抗旱基因资源的发掘和创新、抗旱生理学和遗传学的研究、利用现代生物技术实现不同物种间抗旱基因的转移、建立节水种植栽培技术新体系是目前抗旱性研究的主要内容。     

Yield response of rice (Oryza sativa L.) genotypes to drought under rainfed lowland: 3. Plant factors contributing to drought resistance

A series of experiments were conducted in drought-prone northeast Thailand to examine the magnitude of yield responses of diverse genotypes to drought stress environments and to identify traits that may confer drought resistance to rainfed lowland rice. One hundred and twenty eight genotypes were grown under non-stress and four different types of drought stress conditions.Under severe drought conditions, the maintenance of PWP of genotypes played a significant role in determining final grain yield. Because of their smaller plant size (lower total dry matter at anthesis) genotypes that extracted less soil water during the early stages of the drought period, tended to maintain higher PWP and had a higher fertile panicle percentage, filled grain percentage and final grain yield than other genotypes. PWP was correlated with delay in flowering (r=−0.387) indicating that the latter could be used as a measure of water potential under stress. Genotypes with well-developed root systems extracted water too rapidly and experienced severe water stress at flowering. RPR which showed smaller coefficient of variation was more useful than root mass density in identifying genotypes with large root system.Under less severe and prolonged drought conditions, genotypes that could achieve higher plant dry matter at anthesis were desirable. They had less delay in flowering, higher grain yield and higher drought response index, indicating the importance of ability to grow during the prolonged stress period.Other shoot characters (osmotic potential, leaf temperature, leaf rolling, leaf death) had little effect on grain yield under different drought conditions. This was associated with a lack of genetic variation and difficulty in estimating trait values precisely.Under mild stress conditions (yield loss less than 50%), there was no significant relationship between the measured drought characters and grain yield. Under these mild drought conditions, yield is determined more by yield potential and phenotype than by drought resistant mechanisms per se.     

Variation in root system architecture and drought response in rice (Oryza sativa): Phenotyping of the OryzaSNP panel in rainfed lowland fields

Root growth at soil depths below 30 cm may provide access to critical soil water reserves during drought in rainfed lowland rice. In this study, the OryzaSNP panel, a set of 20 lines representing genetic diversity in rice used for the discovery of DNA sequence polymorphisms, was evaluated for root characteristics in the field over three seasons varying in drought severity. Root length density (RLD) at a depth of 30–45 cm varied up to 74–92% among genotypes under drought stress (2008–2009 dry seasons), ranging from 0.024 to 0.23 cm cm⁻³ in 2008 and from 0.19 to 0.81 cm cm⁻³ in 2009. Real-time monitoring of soil moisture profiles revealed significant differences among genotypes, and these differences were correlated with RLD at those soil depths. Among the lines evaluated, the Aus isozyme group, particularly the genotype Dular, showed greater drought resistance associated with deep root growth and the highest drought response index (less reduction in yield by drought stress). Since the set of genotypes used in this study has been completely sequenced for SNP markers, the phenotypic information on root growth and drought avoidance responses presented here could be used in initial analysis of the genetic basis of dehydration avoidance traits and in facilitating improvement in drought resistance in rice.     

Nitrogen use efficiency in selected rice (Oryza sativa L.) genotypes under different water regimes and nitrogen levels

Water and nutrient availability are two major constraints in most rice-based rainfed shallow lowland systems of Asia. Both stresses interact and contribute to the low productivity and widespread poverty in this environment. The objective of this study was to improve the understanding of interaction between the two factors and to identify varietal characteristics beneficial for productivity in a water- and nutrient-limited rice environment. For this purpose, we screened 19 rice genotypes adapted to different rice environments under two water and two nutrient treatments during the wet season of 2004 and 2005 in southern Luzon, Philippines. Across all genotypes tested and in comparison with the irrigated control, rainfed conditions reduced grain yield of the treatment without N application by 69% in 2004 and by 59% in 2005. The mean nitrogen fertilizer response was highest in the dryer season of 2004 and the rainfed treatment, indicating that water stress had no effect on fertilizer response. Nitrogen application reduced the relative yield loss to 49% of the irrigated treatment in 2004 and to 52% of the irrigated treatment in 2005. Internal efficiency of N (IEN) and recovery efficiency of applied N (REN) were significantly different between genotypes, but were not affected by water availability (REN) or by water and nutrient availability (IEN). In contrast, grain yield and total N uptake were affected by cultivar, N and water availability. Therefore, germplasm for rainfed environments should be screened under conditions of limited and good nitrogen and water supplies. The four best cultivars, CT6510-24-1-2, IR55423-01, IR72, and IR57514-PMI5-B-1-2, performed well across all treatments and both years. Except for IR72, they were all characterized by medium height, medium duration, high early vigor, and a moderate level of drought tolerance. This combination of characteristics seems to enable the optimal use of limited water and nutrient resources occurring in many shallow rainfed lowlands. We also concluded that moderate drought stress does not necessarily affect the response to moderate N rates, provided that drought does not induce high spikelet sterility and that fertilizer N is properly managed.     

Matching the Expression of Root Plasticity with Soil Moisture Availability Maximizes Production of Rice Plants Grown in an Experimental Sloping Bed having Soil Moisture Gradients

Abstract

Soil moisture distributions in rainfed lowland rice environments are largely determined by the position in the toposequence. In this study, we developed an experimental sloping bed that can simulate the soil hydrological conditions in sloping rainfed lowland rice environments to examine if the expression of promoted root system development in relation to soil moisture availability along the soil profile may maximize water uptake and dry matter production under drought. The gradient of available water along both the surface soil layer and the vertical soil profile was successfully created by manipulating ground water levels in the experimental sloping bed indicating the practical effectiveness of this experimental system. Then, two contrasting genotypes, IRAT109 (upland rice adapted japonica) and KDML105 (lowland adapted indica) were grown for plasticity evaluation. Dry matter production was maintained even at a higher position in the toposequence in IRAT109, but decreased in KDML105. Such maintenance of dry matter production in IRAT109 was attributed to its greater ability to increase root length density in a deeper soil layer, where more soil moisture is available. In contrast, KDML105 maintained root length density in the upper soil layer, and could not utilize the soil moisture available in the deeper soil layer. These results imply that the genotype that expressed root plasticity with root system developing in the soil portion where more soil moisture was available showed greater dry matter production than the genotype that showed root plasticity in the soil layer where soil moisture was less available.     

Root Growth and Water Extraction Response of Doubled-Haploid Rice Lines to Drought and Rewatering during the Vegetative Stage

Abstract

Doubled haploid lines (DHLs) of rice (Oryza sativa L.) were used to examine responses to drought and rewatering in controlled rainfed lowland conditions, in order to determine whether confounding by unrelated traits would be less than has been reported previously for contrasting cultivars that differ in genetic background. IR62266 and four DHLs derived from the cross between IR62266 and CT9993 (DHL-32, -51, -54 and -79) were grown in pot experiments in the greenhouse during the 2000 dry and wet seasons at IRRI, Los Baños, Philippines. There were two water regimes (well-watered and drought). Estimated water extraction obtained by time domain reflectometry (TDR) was similar to cumulative transpiration estimated from pot weighing for each genotype. Genotypic variation was observed in root traits and water extraction, with extraction slower in DHL-32 and faster in DHL-79, especially in deeper soil layers. An upper bound relationship between water extraction from a soil layer and root length density (RLD) in that layer was readily apparent over DHLs and soil depths, suggesting a critical value of RLD for water extraction of 0.30 cm cm^?3 in these conditions. Because soils in the field would not be as homogenous as the puddled soils used in these greenhouse experiments, this critical RLD for water extraction from a soil layer is a reference for ideal conditions, and requires careful validation in the field. Use of DHLs permitted comparisons with reduced confounding by genetic background, with consequent improvements in precision.     

Frequency of occurrence of various drought types and its impact on performance of photoperiod-sensitive and insensitive rice genotypes in rainfed lowland conditions in Cambodia

Lowland rice production in the Mekong region is generally low because crops are cultivated under rainfed conditions and often exposed to drought. To examine how field water availability affects productivity of different genotypes in rainfed lowland rice, the field experiments were carried out for six years at eight locations in Cambodia. We classified 34 genotypes used in the experiments into photoperiod-insensitive [short (IS) and medium (IM) maturity] and sensitive [medium (SM) and long-duration (SL) maturity] genotype groups. Mean days-to-flower from sowing was 87 in IS, 112 in IM, 112 in SM and 132 in SL and mean grain yield was 2.0, 2.8, 2.5 and 2.4 t ha⁻¹, respectively. Drought environment was quantified for each experiment by determining whether free water level was observed to be less than the soil surface during three growth stages: GS1 (maximum tillering), GS2 (panicle development) and GS3 (grain filling). The drought frequency estimated from 44 field experiments was 18% in all IM, SM and SL at GS1, 23%, 25% and 32% in IM, SM and SL, respectively at GS2, and 43%, 45% and 57% at GS3. Thus, the drought occurred more frequently after flowering, particularly in the SL group. Based on the results, the IM genotypes may be grown to escape from drought during the GS2 and GS3 periods by sowing early (June) in the drought environment. When sown late (August), the SM genotypes are exposed to less drought risk because they flower earlier than the IM genotypes. Compared with the SM genotypes, the SL genotypes are highly exposed to water stress during the GS3 period, resulting in yield reduction. SM and IM had similar occurrence of drought environment, but yield reduction due to drought was less in SM than in IM. Thus, photoperiod-sensitive cultivars with medium maturity are preferred in drought-prone lowland fields, particularly when sowing is delayed. In the favourable water environment, the SM genotypes can be better than the IM genotypes when sown early in the season, while the IM genotypes performs better than the SM genotypes with late sowing. This suggests that for lowland fields without drought photoperiod-insensitive cultivars are recommended and photoperiod-sensitive cultivars can be preferred if sown early.     

甜菜碱对玉米幼苗抗旱性的诱导效应

以旱玉5号和太单32为材料,在20%PEG模拟干旱条件下,研究外源甜菜碱对玉米幼苗抗旱性的诱导效应。结果表明,水分胁迫显著降低了玉米幼苗的生物量和叶片相对含水量,提高了幼苗的根冠比和渗透调节物质、抗氧化酶活性、ABA含量。喷施0.5mmol/L的甜菜碱溶液能缓解玉米幼苗的水分胁迫,提高幼苗的生物量、叶片相对含水量、根冠比及渗透调节物质、抗氧化酶活性、ABA含量。     

Breeding for drought tolerance: Direct selection for yield, response to selection and use of drought-tolerant donors in upland and lowland-adapted populations

Drought is the most important constraint reducing rice yield in rainfed areas. Earlier efforts to improve rice yield under drought mainly focused on improving secondary traits because the broad-sense heritability (H) of grain yield under drought stress was assumed to be low, however gains in yield by selecting for secondary traits have not been clearly demonstrated in rice. In present study, the effectiveness of direct selection for grain yield was assessed under lowland reproductive stage stress at Raipur in eastern India and under upland reproductive stage drought stress at IRRI. The selection under severe stress (in both upland and lowland trials) resulted in greater gains under similar stress levels (yield reduction of 65% or greater under stress) in evaluation experiments than did selection under non-stress conditions, with no yield reduction under non-stress conditions. We observed similar H of grain yield under stress and non-stress conditions, indicating direct selection for yield under drought will be effective under both lowland and upland drought stresses. None of the secondary traits (panicle exsertion, harvest index, leaf rolling, leaf drying) included in our study showed a higher estimate for H than grain yield under stress. Secondary traits as well as indirect selection for grain yield under non-stress situation were predicted to be less effective in improving yield under drought in both lowland and upland ecosystem than direct selection for grain yield under the respective stress situations. The low, but positive values observed for genetic correlation (r_G) between yield under stress and non-stress indicated that it is possible to combine drought tolerance with high-yield potential but low values also indicated that selection for grain yield needs to be carried under stress environments. The study also indicated that under lowland drought stress, the use of highly drought-tolerant donors, as parents in crosses to high yielding but susceptible varieties resulted in a much higher frequency of genotypes combining high-yield potential with tolerance than did crosses among elite lines with high-yield potential but poor tolerance. Breeding strategies that use drought-tolerant donors and that combine screening for yield under managed drought stress with screening for yield potential are likely to result in the development of improved cultivars for drought-prone rainfed rice producing areas.     

A water balance model for characterization of length of growing period and water stress development for rainfed lowland rice

There is large year-to-year variation in rice production across the Mekong region (Laos, Cambodia and Thailand) due to uncertainty in the timing of the onset of the wet season and drought stress that may develop at any time during the growth of rainfed lowland rice. Unique to the nature of lowland water balance is a large component of deep percolation water loss, which depends on soil texture. The objectives of this study were to develop a soil water balance model for calculating the amount of water held in field storage (i.e. in soil and, if there is standing water, above the soil surface) and to apply it to determine the length of growing period (LGP) and water stress development in relation to soil type and rainfall pattern for the rice ecosystem. The water balance is computed separately for above-ground plus topsoil layer and subsoil layer. Components of the water balance are the existing amount of stored water, rainfall, evapotranspiration, deep percolation, and runoff. The deep percolation rate was determined from clay content in each soil layer. The model runs with daily or weekly weather data to estimate the soil water level for the growing period in the wet season. The model was validated with data collected from top, middle and bottom of rainfed lowland fields in Savannakhet province, Laos. The best correlation between the observed and simulated water level was obtained (r² = 0.41) for middle fields. The simulation results showed that LGP varied greatly from year to year, particularly in locations with sandy soils, due mostly to variation in monthly rainfall occurring at the early part of the growing season (April), but also to some extent by variation at the end of growing season (October). Soil texture on the other hand is shown to have a large influence on the end of the rice growing period and hence LGP, and also water stress development during growth. Sandy soils with clay content less than 7% that are prevalent in the province are shown to cause frequent water stress and early finish in rainfed lowland rice. The model accordingly provides reasonable outputs that can provide a geographical dimension of soil hydrological patterns for various rice growing environments, and also identify the spatial pattern of drought stress that is likely to occur. Model outputs can be used to provide guidelines for practical advice to the rice farmers and researchers for determination of appropriate crop management strategies (e.g. time of planting, varieties), and policy makers for investment decisions on inputs (e.g. fertilizer price) aimed at increasing rice productivity in this Mekong region.     

植物生长物质冠菌素诱导旱稻、水稻幼苗抗旱性的效应及生理机制

以旱稻 297 和水稻越富幼苗为材料,在 20% PEG 模拟干旱条件下,研究了植物生长物质冠菌素处理对叶片水分状况、质膜透性、渗透调节物质（脯氨酸、可溶性糖和可溶性蛋白）及内源激素（ABA、IAA 和GA3）含量的影响。干旱胁迫下,冠菌素处理可以维持旱稻 297（0.01 μmol/L）和越富（0.1 μmol/L）较高的叶片相对含水量,促进幼苗叶片中脯氨酸、可溶性糖和可溶性蛋白的积累,降低质膜透性,维持细胞质膜的完整性;同时,冠菌素（0.01 和0.1 μmol/L）处理明显促进旱稻 297 和越富幼苗叶片中ABA的积累,并改变了 IAA 和GA3 的浓度及比例。冠菌素处理能改善旱稻和水稻幼苗耐干旱胁迫的能力,最适浓度分别为0.01 μmol/L和0.1 μmol/L。     

Photosynthetic Recovery of a Perennial Grass Leymus chinensis after Different Periods of Soil Drought

Abstract

The effects of rewatering after different periods of soil drought stress on the photosynthetic capacity of Leymus chinensis in pots were investigated. The plants were subjected to short-term (10-d), moderate-term (20-day) and long-term (30-d) drought each followed by rewatering. Control plants were well watered during the experimental periods. The long-term water stress without rewatering decreased the chlorophyll content, Chl a/b ratio, carbonic anhydrase (CA) activity, net photosynthetic rate (A), and leaf area compared with the control. Rewatering increased the ratio of Chl a/b, CA activity and A, but decreased the leaf area and ion leakage from the cut leaf pieces. The long-term water stress without rewatering reduced the maximal efficiency of PSII (F_v/F_m), the actual quantum yield (Φ_p), and photochemical quenching (q_p), but these values were increased by rewatering to more than the control level, though non-photochemical quenching (q_N) was decreased as compared with the control. This implied that long-term drought aggravated PSII, but rewatering improved it. The net CO₂-exchange rate showed similar diurnal changes in all treatments, but the rate in the morning was lower in long-term drought (before rewatering) than in the other treatments. These results suggest the photosynthesis of Leymus chinensis may be well adapted to episodical soil drought.     

Increasing Production of Rainfed Lowland Rice in Drought Prone Environments

Summary

Drought is a major production constraint of rainfed lowland rice grown in Thailand and Laos. Adverse soil conditions also reduce yield. In an attempt to increase rainfed lowland rice production in these countries, a major collaborative international project was conducted during a 6-year period in the region. The objectives of the project were to quantify production constraints, determine genotypic variation in yield, and identify an effective breeding strategy. A rice simulation model was developed also and used to investigate the potential impact of strategies for genetic improvement and agronomic management.

Four major physical or biological constraints to higher production levels of rainfed lowland rice were identified, (1) the lack of standing water at the appropriate time of transplanting, (2) severe water stress that often develops at the end of the growing season, (3) low yield potential of the present cultivars, particularly in Thailand, and (4) adverse soil conditions including low pH and low soil fertility. The results of the field experiments and simulation modelling exercises showed that the influence of these constraints can be reduced and yield increased by several methods : in particular, choice of appropriate cultivars and time of sowing to match crop phenology with water availability, application of appropriate fertilizer, adoption of high yielding cultivars, adoption of direct seeding in place of the traditional transplanting system, and reduction of percolation water loss from the paddies.

A technology package currendy being investigated for the rainfed lowland rice is direct seeding early in the season, using cultivars that flower by the end of the rainy season, with application of organic or chemical fertilizer. The appropriate cultivars are early flowering and short-intermediate statured, possess high yield potential and ability to maintain favourable plant water status at flowering, and have the ability to establish well and compete against weed under direct seeding.     

Yield constraints of rainfed lowland rice in Central Java, Indonesia

The low and unstable yields of rainfed lowland rice in Central Java can be attributed to drought, nutrient stress, pest infestation or a combination of these factors. Field experiments were conducted in six crop seasons from 1997 to 2000 at Jakenan Experiment Station to quantify the yield loss due to these factors. Experimental treatments—two water supply levels (well-watered, rainfed) in the main plots and five fertilizer levels (0-22-90, 120-0-90, 120-22-0, 120-22-90, 144-27-108 kg NPK ha⁻¹) in the subplots—were laid out in a split-plot design with four replications. Crop, soil, and water parameters were recorded and pest infestations were assessed.

In all seasons, rice yield was significantly influenced by fertilizer treatments. Average yield reduction due to N omission was 42%, to K omission 33–36%, and to P omission 3–4%. Water by nutrient interactions did not affect rice yield and biomass production. In two of the three dry seasons, an average of 20% of the panicles were damaged by pests and estimated yield loss from pests was 56–59% in well-watered and well-fertilized treatments. In one out of six seasons, yields under rainfed conditions were 20–23% lower than under well-watered conditions. Drought, N and K deficiencies, and pest infestation are the major determinants for high yields in rainfed environments in Jakenan. Supplying adequate nutrient and good pest control are at least as important as drought management for increasing crop productivity of rainfed rice-growing areas in Central Java. The relative importance of drought, nutrient and pest management may vary in other rainfed areas. Yield constraints analysis should be systematically carried out to identify appropriate management strategies.     

Growth and water relations of simulated swards of Lolium perenne L. selection lines with contrasting leaf osmotic potentials

Two selection lines of Lolium perenne , which had different leaf lamina osmotic potential (Ψ_s) when droughted in controlled environments, were grown as vegetative simulated swards in containers of soil 1·0 × 1·3 × 0·9 m deep in the glasshouse. Water was withheld during July, August and September.
Under irrigation the low-ψ_s line had leaf lamina and leaf base ψ^s 0·06 MPa lower than the high-ψ_s line. Under drought, the two lines did not differ significantly in lamina ψ_s but leaf base ψ_s was 0·16 MPa lower in the low line than in the high line. Upon rewatering, most of the osmotic adjustment in the leaf bases was lost within 2 d, but that in the laminae was more stable.
Herbage growth under irrigation was similar in the two lines until September, when it was greater in the highψ_s line. Drought reduced herbage growth, and also regrowth rates following rewatering, but the two lines did not differ in their sensitivity to drought.
Selection for low ψ^s also increased leaf extension rates, and reduced tillering and leaf water conductance of irrigated plants.
The results are discussed in relation to selecting for improved drought resistance on the basis of physiological characters.     

Morpho-Physiological Response of Oryza glaberrima to Gradual Soil Drying

Soil drought occurrence during dry season has been the main constraint, besides prolonged flooding during rainy season, in increasing cropping intensity and rice productivity in tropical riparian wetland. Use of drought tolerant rice genotype might be a suitable option for overcoming such problem. This study focused on the effects of gradual soil drying during early vegetative growth stage on morphological and physiological traits of five Oryza glaberrima genotypes, namely RAM12, RAM14, RAM59, RAM97 and RAM101, and two Oryza sativa subsp japonica genotypes, i.e. Koshihikari and Minamihatamochi. The plants were subjected to 6 d of gradual soil drying condition from 15 days after transplanting (DAT) to 20 DAT, and were allowed to recover until 22 DAT. Gradual soil drying reduced plant growth as indicated by dry mass accumulation. Drought reduced stomatal conductance and increased leaf rolling score of all the genotypes. All the genotypes showed comparable response on stomatal conductance, but O. glaberrima genotypes performed higher in leaf rolling recovery. Meanwhile, O. sativa genotypes decreased total leaf area and specific leaf area, but increased specific leaf weight in order to avoid further damages due to drought stress. Drought tolerance mechanisms in RAM101, RAM12, RAM59 and RAM14 were associated with leaf morpho-physiological responses, root traits and dry biomass accumulation.     

Root signalling and osmotic adjustment during intermittent soil drying sustain grain yield of field grown wheat

A field study was conducted to investigate the effect of intermittent soil drying on resulting non-hydraulic and hydraulic root signals, leaf gas exchange, leaf growth, day of heading, leaf osmotic adjustment and yield of wheat grown in sand and loam soils in lysimeters. A 40-day-drought treatment was imposed when the flag leaf started to emerge and was terminated close to maturity. Soil water content and soil water potential of various soil layers were measured using the neutron moderation method and tensiometers, respectively. Soil drying in the top soil layers induced increase in both xylem and bulk-leaf abscisic acid (ABA) content and reduced the stomatal conductance and leaf growth even before a measurable change in leaf water potential could be detected in droughted plants when compared with fully watered plants. Further, heading and flowering occurred 4 days earlier in the droughted than in the well-watered plants before any loss in leaf water potential had occurred as compared with the fully watered plants. When more severe drought reduced the leaf water status, further accumulation of leaf ABA occurred and transpiration decreased in addition to gradual osmotic adjustment and senescence of older leaves. The osmotic adjustment sustained leaf turgor pressure during soil drying. At severe drought, the osmotic adjustment at full turgor in the flag leaves was 0.85 MPa. In sand, the kernel dry weight increased and as a result similar grain yield was obtained in both the treatments. In loam which had more water available than sand, no significant reduction in the final yield was induced by the drought. It is concluded that (1) non-hydraulic root signals caused early drought adaptation at mild water stress by reducing leaf growth and stomatal conductance and hastening of heading and flowering; (2) osmotic adjustment sustained turgor maintenance and hence the yield-forming processes during moderate and severe water stress.