Root development and the expression of aquaporin genes in rice seedlings under osmotic stress

We conducted hydroponic culture experiments to characterize root traits in a rice cultivar ‘Puluik Arang’ that has been identified in a previous study as a cultivar that is adaptable to unflooded conditions. Root morphological traits and the expression of 11 aquaporin genes in rice seedlings (cv. Puluik Arang and cv. Akitakomachi) subjected to osmotic stress by polyethylene glycol (PEG) treatments (10 and 20%) were analysed. ‘Puluik Arang’ exhibited significantly greater water uptake under 10% PEG treatment than ‘Akitakomachi’. Lateral root development was maintained in ‘Puluik Arang’ under PEG treatments. The expression of some aquaporin genes, particularly OsTIP2;1, was higher in ‘Puluik Arang’ than in ‘Akitakomachi’. Immunocytochemical analysis showed that the OsTIP2;1 protein mainly accumulated in endodermal cells. The results suggest that better lateral root development and the function of aquaporins could contribute to water uptake in ‘Puluik Arang’ under osmotic stress.     

Growth Responses of Drought Resistant Rice Cultivars to Soil Compaction under Irrigated and Succeeding Non-irrigated Conditions during the Vegetative Stage

Abstract

We investigated whether drought resistant rice cultivars exhibit higher dry-matter production under wet and dry compacted soil conditions in the vegetative stage and determined the dominant factors governing resistance to soil compaction. Three rice cultivars, a drought-sensitive Nipponbare, and drought-resistant Senshou and Dular, were grown in pots at four soil bulk densities (SBD) ranging from 1300 to 1600 dry soil kg m^?3. Root and shoot dry matter productions was slightly smaller in Nipponbare over the 29 days after sowing under irrigated conditions than in the other cultivars at all SBDs. Senshou and Dular also maintained a higher dry matter production, both in relative and absolute values, than Nipponbare under the condition of withheld irrigation from days 29 — 39 after sowing. The higher stomatal conductance and leaf water potential of these two cultivars were supported by a larger root system which was mostly accompanied by lower top-root ratios in the irrigated and compacted soils. The higher plant growth rate under the non-irrigated condition might have been a result of both the higher water absorption rate and water use efficiency, which in turn were supported by the larger root biomass. We conclude that the ability of rice to rapidly develop a root system in the early vegetative phase under compacted soils facilitates plant production under subsequent soil desiccated conditions.     

Genotypic Variation in Biomass Production at the Early Vegetative Stage among Rice Cultivars Subjected to Deficient Soil Moisture Regimes and Its Association with Water Uptake Capacity

References

Genetic improvement in water uptake ability and/or water use efficiency (WUE) of rice cultivars is one option to enhance productivity under water-limited conditions. We examined the genotypic variation in biomass production among 70 rice cultivars (69 cultivars of NIAS global rice core collection and Azucena) under different soil moisture conditions, and to identify whether water uptake ability or WUE is responsible for the variation, if any. Two-week-old seedlings were transplanted into pots and grown for three weeks in an environmentally-regulated growth chamber under three soil moisture regimes: flooded (?0.02 MPa soil water potential) and two unflooded (?0.10 and ?0.52 MPa) conditions. Substantial genotypic variations in total dry weight (TDW) were observed under all three regimes. Among all the cultivars tested, TDW was significantly correlated with water uptake ability, but not with WUE. However, several cultivars exhibited comparably higher WUE while showing superior biomass production under the ?0.52 MPa regime. The amount of water uptake was significantly correlated with root dry weight among cultivars regardless of moisture regimes, while substantial genotypic difference in the amount of water uptake per unit root dry weight was observed. These results indicate that a marked genotypic difference exists in biomass production at the early vegetative growth under water-deficient conditions, and that this difference appears to be ascribed primarily to greater water uptake capacity, and additionally to higher WUE in drought-tolerant cultivars.     

Rice near-isogenic-lines (NILs) contrasting for grain yield under lowland drought stress

The development of near-isogenic-lines (NILs) is a very important tool for both genetic and physiological dissection of drought resistance in rice. Two pairs of NILs differing for grain yield under drought stress were isolated and characterized for yield, yield related traits, and several physiological traits in a range of contrasting environments. In replicated field trials both NIL pairs differed significantly for grain yield under drought stress but showed similar yield potential, phenology, and yield component traits under non-stress conditions. A polymorphism analysis study with 491 SSRs revealed that both NIL pairs are at least 96% genetically similar. These NILs show that small genetic differences can cause large difference in grain yield under drought stress in rice. In both pairs the drought-tolerant NILs showed a significantly higher assimilation rate at later stages both under stress and non-stress conditions. They also had a higher transpiration rate under non-stress condition. The most tolerant NIL (IR77298-14-1-2-B-10) had significantly higher transpiration rate and stomatal conductance in severe stress conditions. In one pair the tolerant NIL had constitutively deeper roots than the susceptible NIL. In the second pair, which had higher mean root length than the first pair, the tolerant NIL had more roots, greater root thickness, and greater root dry weight than the susceptible NIL. Deeper root length may allow tolerant NILs to extract more water at deeper soil layers. It is concluded that enhanced rooting depth is an important strategy for dehydration avoidance and rice adaptation to drought stress, but root architecture might not be the only mechanism causing the significant yield increase we observed in lowland drought stress environments. To further dissect the drought avoidance mechanisms in rice, analysis of root hydraulic properties may be necessary.     

Functional roles of root plasticity and its contribution to water uptake and dry matter production of CSSLs with the genetic background of KDML105 under soil moisture fluctuation

Soil moisture fluctuation (SMF) stress due to erratic rainfall in rainfed lowland (RFL) rice ecosystems negatively affect production. Under such condition, root plasticity is one of the key traits that play important roles for plant adaptation. This study aimed to evaluate root plasticity expression and its functional roles in water uptake, dry matter production and yield under SMF using three chromosome segment substitution lines (CSSLs) with major genetic background of KDML105 and a common substituted segment in chromosome 8. The CSSLs showed greater shoot dry matter production than KDML105 under SMF, which was attributed to the maintenance of stomatal conductance resulting in higher grain yield. The root system development based on total root length of the CSSLs were significantly higher than that of KDML105 due to the promoted production of nodal and lateral roots. These results implied that the common substituted segments in chromosome 8 of the 3 CSSLs may be responsible for the expression of their root plasticity under SMF and contributed to the increase in water uptake and consequently dry matter production and yield. These CSSLs could be used as a good source of genetic material for drought resistance breeding programs targeting rainfed lowland condition with fluctuating soil moisture environments and for further genetic studies to elucidate mechanisms underlying root plasticity.     

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.     

水稻生理特性与抗旱性的相关分析及QTL定位

利用籼稻品种IR64和粳稻品种Azucena杂交产生的包含110个加倍单倍体株系的群体,在干旱胁迫和正常水分条件下,连续在2004年和2005年于抽穗期分别测定了叶片水势、相对含水量、叶绿素含量(SPAD值)、游离脯氨酸含量、气孔导度和蒸腾速率,并于成熟期取样,计算抗旱系数。与正常水分状况下相比,干旱胁迫条件下叶片的游离脯氨酸含量的增加达极显著水平,干旱胁迫条件下叶片的相对含水量、水势、叶绿素含量和气孔导度的降低均达显著或极显著水平。相关分析表明,在干旱胁迫条件下,叶片相对含水量、叶片水势与抗旱系数呈显著或极显著正相关。 利用175个RFLP标记构建的遗传连锁图谱分析了与抗旱性相关的叶片生理指标,共检测到与抗旱性相关的6个生理指标的7个加性QTL,31对上位性QTL,其中有2个主效QTL、9对上位性QTL存在环境互作效应。在两种水分条件下检测到的QTL结果有较大差异,说明干旱胁迫对控制与抗旱性相关的叶片生理性状基因的表达有显著的影响。在6个抗旱相关生理指标中,检测到的控制叶片气孔导度和水势的QTL较多,有3个加性QTL和8对上位性QTL控制气孔导度,有8对上位性QTL控制水势。     

Root plasticity under fluctuating soil moisture stress exhibited by backcross inbred line of a rice variety,Nipponbare carrying introgressed segments from KDML105 and detection of the associated QTLs

In rainfed lowland rice ecosystem, rice plants are often exposed to alternating recurrences of waterlogging and drought due to erratic rainfall. Such soil moisture fluctuation (SMF) which is completely different from simple or progressive drought could be stressful for plant growth, thereby causing reduction in yield. Root plasticity is one of the key traits that play important roles for plant adaptation under such conditions. This study aimed to evaluate root plasticity expression and its functional roles in dry matter production and yield under SMF using Nipponbare, KDML 105 and three backcross inbred lines (BILs) and to identify QTL(s) associated with root traits in response to SMF at two growth stages using Nipponbare/KDML105 F₂ plants. A BIL, G3-3 showed higher shoot dry matter production and yield than Nipponbare due to its greater ability to maintain stomatal conductance concomitant with greater root system development caused by promoted production of nodal and lateral roots under SMF. QTLs were identified for total nodal root length, total lateral root length, total root length, number of nodal roots, and branching index under SMF at vegetative and reproductive stages. The QTLs detected at vegetative and reproductive stages were different. We discuss here that relationship between root system of G3-3 and the detected QTLs. Therefore, G3-3 and the identified QTLs could be useful genetic materials in breeding program for improving the adaptation of rice plants in target rainfed lowland areas.     

优化栽培模式对水稻根-冠生长特性、水氮利用效率和产量的影响

【目的】探明优化栽培模式对水稻根冠发育以及产量与肥水利用效率的影响。【方法】以甬优1540(三系籼/粳杂交稻)为材料,设置3个处理：0N(空白)栽培模式、当地农户习惯栽培模式(对照)以及优化栽培模式。【结果】优化栽培处理两年的平均产量为11.5 t/hm²,与对照差异不显著;但其氮肥偏生产力、产谷利用率以及水分利用率较对照显著提高。与对照相比,优化栽培处理改善了水稻根系形态与生理特征,降低了根系生物量与根-冠比,提高了深根比与比根长,增加了齐穗期与灌浆中期根系活跃吸收表面积,提高了灌浆中后期根系氧化力与根系伤流液中玉米素(Z)+玉米素核苷(ZR)的浓度。此外,与对照相比,优化栽培处理显著提高了灌浆中后期剑叶净光合速率、叶片中Z+ZR含量以及籽粒中蔗糖-淀粉代谢途径关键酶活性。【结论】优化与集成现有栽培技术,可以改善水稻根系形态与生理特征,提高地上部生理活性,进而实现肥水利用效率的提高。     

QTL mapping rolling,stomatal conductance and dimension traits of excised leaves in the Bala × Azucena recombinant inbred population of rice

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 F₆ 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.     

氮和磷对不同基因型小麦水分状况的影响

为了阐明不同水分条件下氮、磷对不同基因型小麦的水分生理指标的作用及其变化趋势，试验逸用6个不同基因型的小麦进化材料，研究了拔节期各基因型小麦的叶片保水力、叶水势、叶片膜透性与气孔导度。结果表明，当土壤水分充足时，施肥会提高叶片保水力、叶水势，降低叶片膜透性，增加二倍体小走的气孔导度，而降低六倍体小走的气孔导度。在水分胁迫下，施肥会提高叶片保水力，降低叶水势、叶片膜透性和气孔导度；而且随小麦染色体倍性从2n→4n→6n的进化方向，小麦叶片保水力、叶水势、叶片膜透性均先降低，然后又升高。     

Growth and Yield of Six Rice Cultivars under Three Water-saving Cultivations

Abstract

We evaluated the genotypic differences in growth, grain yield, and water productivity of six rice (Oryza sativa L.) cultivars from different agricultural ecotypes under four cultivation conditions: continuously flooded paddy (CF), alternate wetting and drying system (AWD) in paddy field, and aerobic rice systems in which irrigation water was applied when soil moisture tension at 15 cm depth reached ?15 kPa (A15) and ?30 kPa (A30). In three of the sixcultivars, we also measured bleeding rate and predawn leaf water potential (LWP) to determine root activity and plant water status. Soil water potential (SWP) in the root zone averaged ?1.3 kPa at 15 cm in AWD, -5.5 and -6.6 kPa at 15 and 35 cm, respectively, in A15, and ?9.1 and ?7.6 kPa at 15 and 35 cm, respectively, in A30. The improved lowland cultivar, Nipponbare gave the highest yield in CF and AWD. The improved upland cultivar, UPLRi-7, and the traditional upland cultivar, Sensho gave the highest yield in A15 and A30, respectively. The yields of traditional upland cultivars,Sensho and Beodien in A30 were not lower than the yields in CF. However, the yields of the improved lowland cultivars, Koshihikari and Nipponbare, were markedly lower in A15 and A30. Total water input was 2145 mm in CF, 1706 mm in AWD, 804 mm in A15, and 627 mm in A30. The water productivity of upland rice cultivars in aerobic plots was 2.2 to 3.6 times higher than that in CF, while those of lowland cultivars in aerobic plots were lower than those in CF. The bleeding rate of Koshihikari was lower in A15 and A30 than in CF and AWD, and its LWP was significantly lower in A15 and A30 than in CF and AWD, but Sensho and Beodien showed no differences among the four cultivation conditions. We conclude that aerobic rice systems are promising technologies for farmers who lack access to enough water to grow flooded lowland rice. However, lowland cultivars showed severe growth and yield reductions under aerobic soil conditions. This might result from poor root systems and poor root function, which limits water absorption and thus decreases LWP. More research on the morphological and physiological traits under aerobic rice systems is needed.     

Physiological and Molecular Analysis of Applied Nitrogen in Rice Genotypes

Ten genotypes of rice (Oryza sativa L.) were grown for 30 d in complete nutrient solution with 1 mmol/L (N-insufficient),4 mmol/L (N-moderate) and 10 mmol/L (N-high) nitrogen levels,and nitrogen efficiency (NE) was analyzed.Growth performance,measured in terms of fresh weight,dry weight and lengths of root and shoot,was higher in N-efficient than in N-inefficient rice genotypes at low N level.Of these 10 genotypes,Suraksha was identified as the most N-efficient,while Vivek Dhan the most N-inefficient.To find out the physiological basis of this difference,the nitrate uptake rate of root and the activities of nitrate assimilatory enzymes in leaves of N-efficient and N-inefficient rice genotypes were studied.Uptake experiments revealed the presence of two separate nitrate transporter systems mediating high-and low-affinity nitrate uptake.Interestingly,the nitrate uptake by the roots of Suraksha is mediated by both high-and low-affinity nitrate transporter systems,while that of Vivek Dhan by only low-affinity nitrate transporter system.Study of the activities and expression levels of nitrate assimilatory enzymes in N-efficient and N-inefficient rice genotypes showed that nitrate reductase (NR) and glutamine synthetase (GS) play important roles in N assimilation under low-nitrogen conditions.     

Physiology of the Potato: New Insights into Root System and Repercussions for Crop Management

Potato roots are concentrated mostly in the plow layer up to 30 cm in soil depth. Some roots extend up to 100 cm depth and the total root length throughout the soil profile reaches about 10–20 km m^?2 area. There are large differences in root mass (dry weight and length) in the plow layer between cultivars, breeding lines and wild relatives. The differences are generally stable across different environmental conditions, such as locations with different soil types, fertilizer rates and planting densities. Under favourable environmental conditions without severe shortage of water and nutrients, root mass differences between genotypes are related to maturity class: late genotypes continue root growth longer, and attain larger root mass and deeper rooting than early genotypes. Differences in root mass become clear at the start of flowering, much earlier than differences in shoot mass. Root mass is negatively correlated with early tuber bulking. However, root mass generally shows positive correlations with shoot mass and final tuber yield. Differences in root mass also exist amongst genotypes of the same maturity class. Using root mass in the plow layer and tuber yield as selection criteria, Konyu cultivars were bred in Japan. They showed significantly less reduction of leaf conductance and photosynthesis, leaf area and tuber yield than commercial cultivars under dry soil conditions. To assist breeding for root characters, new methods have been developed to assess the ability of roots to penetrate into hard soil layers using pots with paraffin-vaseline discs and the ability to absorb under low water potential in vitro. Physiological research on root characteristics contributed in the past, and will continue to do so in the future, to the development of new cultivars with high drought tolerance and to the improvement of irrigation practice.     

盐胁迫对不同水稻品种光合特性和生理生化特性的影响

 用2个耐盐水稻品种和2个盐敏感型水稻品种为材料,研究盐胁迫对水稻植株生物量积累、光合特性等生理特性的影响。结果表明,在盐胁迫条件下,耐盐水稻品种和盐敏感型水稻品种地上和根系的干鲜质量均呈下降趋势,其中,以盐敏感型水稻品种的地上部鲜质量与根系干质量的下降最为显著。盐胁迫条件下,水稻叶片的净光合速率（Pn）、气孔导度（Gs）、蒸腾作用（Tr）和表观叶肉导度（AMC）均呈不同程度的下降趋势。其中,耐盐水稻品种Pn、Gs、Tr和Pn/Ci的下降均低于盐敏感型水稻品种。同时,耐盐品种水分利用效率（WUE）也高于盐敏感型品种。盐胁迫条件下,耐盐水稻品种和盐敏感型水稻品种的胞间二氧化碳浓度（Ci）变化并不明显,气孔限制百分率（Ls）均较低,品种间差异也不显著,而表观叶肉导度显著下降,由此推测盐胁迫条件下Pn的下降并非因为气孔的限制,而与RuBPCase活性的下降有关。盐胁迫条件下水稻叶片和根系的渗透性调节物质脯氨酸、总氨基酸和可溶性糖含量上升,其中以可溶性糖含量上升最为显著（P＜0.01）。盐胁迫下叶片保护酶SOD、POD和CAT活性增强,膜透性增强,丙二醛含量增加,同时,根系活力下降。     

梯度干旱胁迫对水稻叶片光合和水分状况的影响

采用温室营养液培养方式,通过添加0%、10%、20%、30%PEG6000模拟干旱胁迫,对水稻幼苗叶片的光合作用和水分状况进行比较分析。结果表明:1)在干旱胁迫下,水稻叶片的光合速率、气孔导度、叶肉导度、总导度和叶绿体内CO2浓度等都显著降低;2)在干旱胁迫条件下,限制光合作用的非气孔限制值并没有显著提高,而气孔限制值则大幅提高;与正常水分条件相比,扬稻6号和汕优63在30%PEG干旱胁迫下气孔限制值分别提高了42%和81%;3)光合速率与气孔导度、叶肉导度、总导度及叶绿体内CO2浓度呈正相关;4)在重度干旱胁迫下(20%和30%),叶片水势和含水量都显著下降,并且叶片水势与气孔导度、叶肉导度和总导度呈正相关。因此,气孔关闭导致的叶绿体内CO2浓度降低是限制光合作用的最主要因素,同时叶片水势的降低增加了叶片内CO2传输的阻力。     

Role of early vigor in adaptation of rice to water-saving aerobic culture: Effects of nitrogen utilization and leaf growth

Early vigor and rapid canopy development are important characteristics in aerobic rice culture, where they are highly susceptible to soil water deficits. To elucidate the response of rice's vegetative growth to water management regimes, we evaluated the leaf growth and the concomitant nitrogen (N) utilization of nine cultivars grown in flooded and aerobic culture in 2 years. In aerobic culture, the soil water potential at a depth of 20 cm frequently reached −60 kPa in 2007, but remained above −30 kPa in 2008. The average leaf area index (LAI) in the middle of the vegetative growth stage, N uptake and leaf N content per unit leaf area (specific leaf N; SLN) in aerobic culture were comparable to those in flooded culture. However, there was a significant cultivar × water regime interaction in LAI: cultivars with higher LAI during the vegetative growth stage achieved higher yield in aerobic rice culture. IR72 and Takanari (high-yielding cultivars of flood-irrigated rice) showed poor leaf growth as well as lower N uptake and higher SLN in aerobic culture compared with flooded culture. Our results show that early vigor is closely associated with yield stability to the soil moisture fluctuations in aerobic rice culture, even if weeds are properly controlled. Greater N uptake from aerobic soil and better balancing between the N demand for leaf growth and the N supply to the leaves under fluctuating soil moisture would be, at least in part, relevant to a rice cultivar's adaptation to aerobic conditions.     

Root growth dynamics and stomatal behaviour of rice (Oryza sativa L.) grown under aerobic and flooded conditions

Aerobic rice culture is a new technology designed to reduce water use, but the vulnerability of rice to aerobic condition has limited its development. The objective of this study was to characterize the root growth and stomatal behaviour of four rice cultivars grown in flooded and aerobic culture for 2 years. In aerobic culture, where the soil water potential at 20-cm depth averaged between −15 and −30 kPa, total root biomass was significantly lower than in flooded culture for the whole growth period, owing to a reduction in root biomass in the surface layer. Dry-matter partitioning to roots decreased, but the ratio of deep root biomass to total root biomass tended to be higher in aerobic culture than in flooded culture. The low root-to-shoot ratio and poor root growth in the surface layer in aerobic culture are attributable to the considerable reduction in adventitious root number. As a result, the varietal difference in total root biomass was due largely to individual root growth in aerobic culture. Stomatal closure was distinct at the vegetative stage in aerobic culture, even when the soil water potential was near field capacity, partly because of the poor rooting vigour. When the soil water potential at 20-cm depth was below −50 kPa, the stomatal behaviour reflected the root growth in the subsurface layer. These results suggest the role of vigorous root growth in soil water uptake and hence, in maintaining transpiration in aerobic rice culture.     

不同土壤肥力下旱后复水对冬小麦光合特性及水分利用效率的影响

为给冬小麦节水高产栽培提供理论依据,采用盆栽试验,研究了不同土壤肥力下返青后干旱及拔节期复水对冬小麦光合特性和水分利用效率的影响。结果表明,同一土壤水分条件下,光合速率、蒸腾速率、气孔导度、胞间CO2浓度均随土壤肥力的降低而降低;同一土壤肥力下,光合速率、蒸腾速率、气孔导度、胞间CO2浓度均随干旱程度的增加而降低。干旱胁迫复水24 h后,光合速率、蒸腾速率、气孔导度、胞间CO2浓度均出现了补偿或超补偿效应,但中度干旱恢复度大于重度干旱。光合速率以HA处理（高肥力、最大田间持水量的85%）最高,单叶水分利用效率在干旱胁迫下和复水24 h后均以HB处理（高肥力、最大田间持水量的55%）最高。以上结果说明,冬小麦返青期中度干旱条件下,高肥力土壤有利于复水后光合系统恢复和单叶水分利用效率提高,而重度干旱下贫瘠土壤应减少基肥的施入量。