Aerenchyma Formation in the Seminal Roots of Japanese Wheat Cultivars in Relation to Growth under Waterlogged Conditions

Abstract

Morphological adaptation of roots is critical for plants to survive under waterlogging. In this study, we evaluated the capacity of wheat to form aerenchyma in seminal roots in combination with the growth angle of the roots. We used five Japanese cultivars from the waterlogging-prone Kanto-Kyushu region in Japan, and a non-Japanese cultivar, Bobwhite for comparison. Seedlings in pot culture were waterlogged at a 3-cm depth for 7 days. The first adverse effect of waterlogging on plant growth was a significant reduction of root dry mass. The reduction rate varied with the cultivar, and it was 19.2% in cv. Shiroganekomugi and 40.0% in cv. Norin 61. Root aerenchyma was initially observed on the 2nd day of waterlogging and developed until the 7th day, in all 6 cultivars. Quantitative analysis of the aerenchyma development revealed no significant difference in radial distribution among the cultivars, whereas a slight difference was found in the axial distribution. As a consequence, the heavier root weight of Shiroganekomugi was not related to either the radial or axial developing capacity of aerenchyma but might be due to the effect of its shallow root angle in the soil. These results suggest that the capacity to form aerenchyma in the seminal root is not sufficient for expression of waterlogging tolerance in the Japanese wheat cultivars.     

Roles of Root Aerenchyma Development and Its Associated QTL in Dry Matter Production under Transient Moisture Stress in Rice

Abstract

Enhanced aerenchyma development in rice under transient drought-to-waterlogged (TD-W) stress promotes root system development by promoting lateral root production. This study analyzed the quantitative trait loci (QTLs) associated with the plasticity in aerenchyma development under TD-W stress. A mapping population of 60 F₂ genotypes of chromosome segment substituted lines (CSSL) derived from CSSL47 and Nipponbare crosses were grown in rootboxes and evaluated for shoot and root growth, and aerenchyma development (expressed as root porosity). The TD-W stress was imposed starting with water saturated soil condition at sowing and then to progressive drought from 0 to 21 days after sowing (DAS) prior to exposure to sudden waterlogging for another 17 days (21 to 38 DAS). We performed simple and composite interval mapping to identify QTLs for aerenchyma development. QTL associated with aerenchyma development was mapped on the short-arm of chromosome 12 and designated as qAER-12. The effect of qAER-12 on the plasticity in aerenchyma development under TD-W was significantly associated with the increase in lateral root elongation and branching. This resulted in greater root system development as expressed in total root length and consequently contributed to higher dry matter production. This qAER-12 is probably the first reported QTL associated with aerenchyma development in rice under TD-W and is a useful trait for the improvement of the adaptive capability under fluctuating soil moisture conditions.     

Secondary Aerenchyma Formation and its Relation to Nitrogen Fixation in Root Nodules of Soybean Plants (Glycine max) Grown under Flooded Conditions

Abstract

Soybean (Glycine max (L.) Merr.) is considered to be susceptible to flooding, a major agronomic problem in the world, and nitrogenase activity rapidly declines due to oxygen deficiency in root nodules. We investigated nodule acclimation to flooding at the morphological level using a soybean cultivar possessing the ability to form secondary aerenchyma. After 1 week of treatment, lenticels were formed on the surface of the root nodules, and secondary aerenchyma were observed through the lenticels under both irrigated and flooded conditions. As the plant grew, the nodule epidermis came off, and well-developed secondary aerenchyma covered the nodule surface. The secondary aerenchyma originated from the secondary meristem (phellogen) girdling the sclerenchyma, and the degree of development was greater in flooded nodules than in irrigated ones. Although root nodulation and total nitrogenase activity (TNA) decreased under flooded conditions, there were no differences in shoot N concentration, specific nitrogenase activity (SNA) and relative ureide-N in the xylem bleeding sap between plants in the irrigated and flooded conditions. Under flooded conditions, however, when the entry of oxygen into the secondary aerenchyma formed in the hypocotyl was inhibited by vaseline treatment (pasting on the surface of the hypocotyl), the shoot N concentration, TNA, SNA, the ureide-N concentration and the relative ureide-N in the sap declined remarkably. These results suggested that secondary aerenchyma formation in soybean plants is a morphological acclimation response to flooding stress, and that one of the functions is to supply atmospheric oxygen to root nodules, which consequently enables nodule activity to be maintained.     

Utilizing Chromosome Segment Substitution Lines (CSSLs) for Evaluation of Root Responses to Transient Moisture Stresses in Rice

Abstract

Drought and waterlogging that occur sequentially under field conditions are important abiotic stresses affecting plant growth and development. The ability to maintain the root system development during the contrasting moisture stresses may be one of the key traits for plant adaptation. This study aimed to identify the key root traits that contributed to the above ability by comparatively examining the effects of the two moisture stresses in succession on root system development. The chromosome segment substitution lines (CSSLs) from the crosses between the japonica rice cultivar Nipponbare and indica rice cultivar Kasalath were used for precise comparison of root system development. The rice seedlings were grown by hydroponics under a continuously well-aerated condition for 14 days (non-stressed), a drought condition for 7 days followed by an oxygen (O₂)-deficient (stagnant) condition for 7 days (drought-to-stagnant, D-S), or a stagnant condition for 7 days followed by drought condition for 7 days (stagnant-to-drought, S-D). CSSL43 and 47 did not show any significant differences in growth from Nipponbare under the non-stressed condition, but exhibited greater lateral root production under the stresses. Lateral root production was most closely related to faster seminal root elongation mediated by higher aerenchyma formation in the D-S condition, and to more branching of lateral roots on the seminal root axis in the S-D condition. The D-S condition severely affected lateral root production due to reduced seminal root elongation and aerenchyma formation. These results confirmed the fact that those root traits previously identified using different cultivars greatly contribute to plant adaptation. Oxygen deficiency preceded by drought (D-S) was more stressful to roots than drought preceded by O₂ deficiency (S-D), because drought reduced root aerenchyma formation during the subsequent stagnant condition.     

Effect of pre- and post-heading waterlogging on growth and grain yield of four millets

Seeds of Panicum miliaceum, Panicum sumatrense, Setaria glauca, and Setaria italica were raised in polyvinylchloride tubes filled with soil to determine interspecific differences in waterlogging tolerance and the effect of pre- and post-heading waterlogging on growth and grain yield. Four treatments were conducted including control (no-waterlogging stress during growth). Pre-heading waterlogging treatment was initiated 17 days after sowing to heading (TC). Post-heading waterlogging treatment was initiated heading till harvest (CT). Waterlogging treatment was initiated 17 days after sowing to harvesting (TT). The grain yield of P. miliaceum, S. glauca, and S. italica decreased 16, 18, and 4%, while that of P. sumatrense increased 210% under TT treatment and this showed P. sumatrense had most waterlogging tolerance. The grain yield was more affected under TC treatment in S. italica and P. miliaceum. However, there was not significant differences the grain yield between TC and CT treatment in P. sumatrense and S. glauca. Total dry weight, total root dry weight, number of crown root, and the proportion of lysigenous aerenchyma of P. sumatrense were significantly higher than those of other millets at harvesting. Plant growth rate, total root dry weight, number of crown root, and the proportion of lysigenous aerenchyma of P. sumatrense were significantly higher than those of other millets at heading. These results suggest that P. sumatrense exhibits waterlogging tolerance by enhancing root growth characterized by a high proportion of lysigenous aerenchyma in the crown root.     

Alleviation of waterlogging damage in winter rape by application of uniconazole: Effects on morphological characteristics,hormones and photosynthesis

Oilseed rape seedlings (Brassica napus L.) treated with uniconazole were transplanted at the five-leaf stage into specially designed experimental containers, and then exposed to waterlogging for 3 weeks. Pre-treatment of rape seedlings with uniconazole significantly increased seedling height, shoot width, number of green leaves and leaf area per plant, and consequently the shoot, root, and total dry weight after waterlogging. Following waterlogging stress, the uniconazole treated plants had a significantly improved growth including plant height, length and width of leaves, number of green leaves, leaf area per plant, and canopy width at the stem elongation stage, and plant height, stem width, and root, shoot and total dry weight at the flowering stage. At harvest, uniconazole treatment increased the number of primary and secondary branches, seeds per pod, and number of effective pods in branches and in terminal raceme after waterlogging treatment. The uniconazole-induced increase in the number of pods per plant and number of seeds per pod were the two yield components responsible for the significantly greater seed and oil yields obtained from the uniconazole plus waterlogging treated plants, over either the control or waterlogged plants. Uniconazole also reduced waterlogging-induced rise in the erucic acid content of the seeds. The modification of GA₃, zeatin, ABA and ethylene levels due to pre-treatment of rape seedlings with uniconazole might have helped to delay the chlorosis and senescence induced by waterlogging. Uniconazole treatment also increased the leaf photosynthetic rates of waterlogged plants, in part, due to the changes in leaf conductance and hormone levels which ultimately affected various physiological processes.     

Root Development,Water Uptake,and Shoot Dry Matter Production under Water Deficit Conditions in Two CSSLs of Rice: Functional Roles of Root Plasticity

Abstract

Root traits that can contribute to drought resistance have not been clearly indentified. We examined the role of root system development in enhancing water uptake and contribution to dry matter production by using the root box-pinboard method, with which quantitative assessment of root system development and the water uptake of root are possible. Chromosome segment substitution lines CSSL45 and CSSL50, and the recurrent parent Nipponbare were grown under continuously waterlogged conditions (control), and various intensities of water deficit in root boxes. There was no significant difference among the genotypes in shoot growth and root development, while CSSL45 and CSSL50 showed greater shoot dry weight than Nipponbare under water deficit conditions. This was due to their abilities to promote root system development as compared with Nipponbare, which facilitated greater water extraction than Nipponbare, especially under the mild water deficit condition of 20–25% w/w soil moisture contents. Furthermore, the increased root length density did not exceed the estimated critical value for water uptake, which indicates that plastic root system development was functionally effective and efficient for the enhancement of water uptake under mild water deficit conditions.     

Genotypic Variations in Responses of Lateral Root Development to Transient Moisture Stresses in Rice Cultivars

Abstract

Soil water regimes under field conditions inevitably tend to fluctuate ranging from drought to waterlogging. Genotypes that adapt better to such changing hydrologic conditions are assumed to have the ability to maintain root system development under such conditions. This study aimed to evaluate the responses of root system development based on lateral root production to transient moisture stresses, and the contribution of the elongation of seminal and nodal root axes and their lateral, root branching, and aerenchyma development in the seminal root axis, to root system development. The seedlings of two aerobic genotypes (UPLRi7 and NSICRc9) and one irrigated-lowland genotype (PSBRc82), and two parental genotypes (Nipponbare and Kasalath) of chromosome segment substitution lines (CSSLs) were grown by hydroponics. The seedlings were exposed to a drought condition by adding polyethylene glycol to the solution for 7 days and then to an O₂-deficient stagnant condition for 7 days (drought-to-stagnant condition), or to reverse successive conditions (stagnant-to-drought condition). Under both conditions, the aerobic genotypes showed greater ability to produce lateral roots than the irrigated-lowland genotype. Under the transient stagnant-to-drought condition, the root traits that contributed to greater lateral root production in the aerobic genotypes were faster seminal root elongation that was closely associated with branching of lateral roots, and greater nodal root production. Under transient drought to stagnant condition; these were faster seminal root elongation mediated by higher aerenchyma formation, and greater nodal root production. Kasalath showed much greater ability to produce lateral roots under both transient moisture stress conditions than Nipponbare. This indicates the potential utility of the CSSLs for precise identification of desirable root traits with less genetic confounding.     

淹水胁迫下小麦根通气组织形成的PCD特征及活性氧作用初探

为了解淹水条件下小麦根通气组织形成的细胞学特点和活性氧(Reactive oxygen species, ROS)在通气组织形成中的作用,以高度耐湿的华麦8号幼苗为材料进行淹水处理,在显微水平上系统地观察了通气组织的形成过程,在分子水平上检测了细胞核DNA的断裂情况,并对ROS的动态变化进行了荧光观察.结果表明:(1)通气组织最先起源于根皮层中部,然后逐渐扩展,到淹水8 d时基本形成,淹水60 d时更为发达;(2)淹水1.5～48 h根皮层出现大量细胞核DNA断裂,且细胞核DNA降解为180～200 bp的片段,证明小麦根通气组织的形成过程是根皮层细胞发生细胞程序化死亡(Programmed cell death, PCD)的过程,而且淹水1.5～48 h是根皮层细胞死亡高峰期;(3)ROS在细胞核DNA发生断裂前开始积累,在通气组织形成中呈现动态变化.上述结果表明, 淹水胁迫下小麦根通气组织形成是一个皮层细胞PCD的过程,并且ROS可能参与介导了PCD的进程.另外,淹水胁迫在前期抑制了次生根的产生,随后又促进了次生根的产生.     

Effects of Ethephon on Aerenchyma Formation in Rice Roots

The effects of ethephon on the constitutive aerenchyma formation in roots were studied with a rice variety Yangdao 6 as material. The number of air spaces formed by disintegrated cells in mediopellis increased significantly with the rising ethephon concentrations, whereas superoxide dismutase (SOD) activity showed downward trends. Compared with the control, the expression levels of xyloglucan endotransglycosylase (XET) gene were markedly higher both at the apical 10 mm and distal parts of roots in 100 mg/L ethephon treated plants. The accumulation of XET was supposed to be associated with the aerenchyma development. Furthermore, earlier cortical cell death was observed under the ethephon treatments, and most of nuclei of cells at 4 mm from the root apex disintegrated with many Golgi apparatus, mitochondria and membrane- bound vesicles around the cell wall.     

乙烯利诱导水稻根内通气组织形成的研究

 以扬稻6号（Oryza sativa L.）幼苗为材料,研究乙烯利对根内组成型通气组织形成的影响。结果表明,随营养液中乙烯利浓度的增高,中皮层部位细胞解体形成通气空腔数增多,超氧化物歧化酶的活性呈下降趋势;以正常生长的水稻根系为对照,100 mg/L乙烯利处理后根尖10 mm和距根尖10 mm以上部位的木葡聚糖转葡糖苷酶（XET）基因表达量均显著高于对照相应部位,XET基因的积累与通气组织形成具有一致性;对处理与对照根尖10 mm部位超微结构比较发现,乙烯利处理后根系皮层细胞发生死亡解体的时间早,在距离根尖4 mm处多数细胞内核已解体消亡,细胞壁周围可见较多的高尔基体、线粒体和膜包裹的空泡等。     

Interactive Effects of Elevated Atmospheric CO2 and Waterlogging on Vegetative Growth of Soybean (Glycine max (L.) Merr.)

Abstract

Waterlogging is a major predicted agricultural problem for crop production in some areas under current climate change, but no studies are available on the interactive effects of waterlogging and elevated atmospheric CO₂ concentration ([CO₂]). We hypothesized that elevated [CO₂] could alleviate the damage caused by waterlogging, and tested the hypothesis using vegetative growth of soybean (Glycine max) in 10 experiments (different sowing time and different soil type) conducted at Morioka and Tsukuba for three years. The 2-week-old plants grown under elevated and ambient [CO₂] were exposed to waterlogging for 2 weeks. Total dry weight at the end of the treatment was higher under elevated [CO₂] than under ambient [CO₂], and it was significantly reduced by waterlogging under both levels of [CO₂], without significant [CO₂]×waterlogging interactions, at both locations. The negative effects of the waterlogging were greater in root dry weight than in top dry weight, and the root exudation per unit root dry weight was also reduced by waterlogging, without a [CO₂] ×waterlogging interaction. Therefore, the hypothesis of a [CO₂]×waterlogging interaction can be rejected, and provide an important basis for predicting future damage caused by waterlogging under elevated [CO₂] conditions.     

Effects of the Combined Application of Ethephon and Gibberellin on Growth of Rice (Oryza sativa L.) Seedlings

Abstract

Improvement of early seedling growth, such as seedling emergence and vigor is one of the most important agronomic traits in direct seeding rice cultivation. The effects of two plant growth regulators (PGRs), gibberellic acid (GA₃) and ethephon (ET), on seedling growth under flooded soil conditions at different temperatures and water depths were investigated. The PGRs were applied during the seed soaking process. A single treatment with GA₃ or ET increased seedling growth. However, combined application of GA₃ and ET was more effective than that of GA₃ or ET alone in many cases at both growing temperatures (15 and 20ºC). The growth of different organs in the rice seedlings, such as the coleoptiles, first leaves, and second leaves was also increased by PGR treatment. The nitrogen concentration of the shoot and the ratio of shoot dry weight to shoot length did not differ significantly among the treatments. The results of our study show that rice seedling growth in direct seeding cultivation may be improved by treatment with GA₃ and ET in combination.     

渍涝对不同耐性花生品种根系形态特征及解剖结构的影响

为解析渍涝对花生根系形态特征的影响，以耐性品种湘花2008与敏感品种中花4号根系为研究材料，在自制根架系统内进行盆栽培养，于幼苗期分别进行0、3、6、9、12 d渍涝胁迫处理，通过根系的鲜（干）重以及显微和超微结构研究不同耐性花生品种根系形态特征的差异。结果表明，渍涝胁迫下中花4号根系受影响较大，随着淹水天数延长，根系颜色加深，臭味加重，而湘花2008仅根系中、上部根色变黄，臭味较轻，湘花2008根系鲜重耐渍系数高于中花4号。湘花2008出现了通气组织，且随着淹水时间增加，通气组织数量增加、横截面积加大；主根中段木质部基本保持正常，导管大体呈射线状分布。而中花4号未形成明显的通气组织，木质部和导管呈不规则分布。湘花2008主根细胞内的细胞核、线粒体虽受到影响，但仍能维持细胞活动，保持根系生长状态。中花4号细胞器受渍涝胁迫较敏感，加速衰老死亡。综上认为，渍涝胁迫下湘花2008根色和根味受影响较小，主根出现了通气组织，主根中段木质部与细胞中的细胞核、线粒体基本正常。而中花4号根色和根味受影响较重，未形成明显的通气组织，主根中段木质部和导管呈不规则分布，主根中段细胞中的细胞质溶解，线粒体异常。本研究可为解析花生耐渍涝机理提供依据。     

淹水胁迫对4种暖季型草坪草光合特性的影响

为了筛选耐淹护坡植物,本研究以双穗雀稗（Paspalum paspaloides）、伪针茅（Pseudoraphis spinescens）、海滨雀稗（P. vaginatum）和假俭草（Eremochloa ophiuroides）4种暖季型草坪草为材料,研究不同淹水时间（0、6、12、18、24、30 d）处理对上述材料生长、叶绿素含量和叶片光合参数的影响。结果表明：淹水胁迫对双穗雀稗和伪针茅地上与根系生长未产生显著影响,但显著抑制了海滨雀稗和假俭草地上和根系生长;假俭草和海滨雀稗的光合色素含量呈明显的下降趋势,而双穗雀稗和伪针茅的光合色素含量保持相对稳定,到淹水后期其含量反而有所升高。此外,海滨雀稗和假俭草的净光合速率（P_n）随淹水时间的延长显著下降,而双穗雀稗和伪针茅的光合作用保持相对稳定,在淹水胁迫30 d时反而有所提高;淹水胁迫对伪针茅和双穗雀稗的气孔导度（G_s）、蒸腾速率（T_r）没有显著影响,而在淹水胁迫30 d时,海滨雀稗和假俭草的G_s分别较对照降低了25.7%和37.7%。结果表明,双穗雀稗和伪针茅在淹水胁迫下能够保持稳定的光合作用,生长未受到明显的影响,具有很强的耐淹性,是河湖库区优良的固土护坡植物。     

不同耐铝性大麦品种对铝锰胁迫的生理响应

为进一步明确大麦品种对铝锰胁迫的生理效应,研究了不同胁迫处理对大麦生长、叶片功能和氧化胁迫指标的影响。结果表明,铝毒害导致大麦根长和根系干物质量降低,锰毒害显著降低植株株高和地上部干物质量,铝锰共存时锰加重了大麦铝毒害。铝锰胁迫降低了功能叶叶绿素含量和光合能力,铝锰复合处理下叶片功能指标下降幅度高于单独铝或锰处理。胁迫处理下叶片功能指标的下降幅度以叶绿素a含量和叶片蒸腾速率最明显。胁迫处理对铝敏感品种的生长和叶片功能毒害强于耐铝品种。铝锰胁迫引起大麦功能叶丙二醛（MDA）含量增加,过氧化物酶（POD）和超氧化物歧化酶（SOD）活性增强,铝敏感品种增幅更大,复合处理时两品种叶片MDA积累量及铝敏感品种POD和SOD活性增幅显著高于单独铝或锰处理。     

不同耐铝大麦品种对铝锰互作胁迫的生理响应研究

铝（Al）和锰（Mn）是限制酸性土壤作物生长的两大主要因素。以耐铝性差异较大的两个大麦品种为材料,研究了不同处理对大麦生长、叶片功能和氧化胁迫指标的影响。结果表明,与对照相比,铝毒害导致大麦根长下降,根系干物质量降低,锰毒害显著降低植株株高和地上部干物质量,铝锰共存时锰加重大麦铝毒害。铝锰胁迫降低功能叶叶绿素含量和光合能力,复合处理下叶片功能指标下降幅度高于单独铝或锰处理。胁迫处理下叶片功能指标的下降幅度以叶绿素a含量和叶片蒸腾速率最明显。胁迫处理对铝敏感品种的生长和叶片功能毒害强于耐铝品种。铝锰胁迫引起大麦功能叶丙二醛（MDA）含量增加,过氧化物酶（POD）和 超氧化物歧化酶（SOD）活性增强,铝敏感品种增幅更大,复合处理时两品种叶片 MDA 积累量及铝敏感品种 POD 和 SOD 活性增幅显著高于单独铝或锰处理。     

【Short Report】Alternative Experimental Method Using a FRP Pot for Evaluating Wet Damage in Soybean and Morning Glory Grown underExcess Soil Water Conditions

Abstract

A fiber reinforced plastic (FRP) pot, equipped with a slanted pipe through which water is supplied, has been used for assessment of rice herbicides. This pot may also be useful for controlling water table in upland conditions. In this experiment, early growths of soybean and morning glory grown under waterlogging and excess soil water conditions were evaluated. The waterlogging condition was attained by adjusting the top of the slanted pipe to soil surface level, and excess soil water condition (0.2 m³ m^?3) by adjusting to 15 cm below the soil surface. The water content of the soil did not fluctuate during the treatment. Shoot growths of soybean and morning glory were poorer under waterlogging than excess soil water condition. The present experimental procedures using a FRP pot could be available for evaluation of wet damage of field crops grown under ill-drained field.     

Mechanisms for coping with submergence and waterlogging in rice

Rice (Oryza sativa L.), unlike other cereals, can grow well in paddy fields and is highly tolerant of excess water stress, from either submergence (in which part or all of the plant is under water) or waterlogging (in which excess water in soil limits gas diffusion). Rice handles submergence stress by internal aeration and growth controls. A quiescence strategy based on Submergence-1A (SUB1A) or an escape strategy based on SNORKEL1 (SK1) and SNORKEL2 (SK2) is used for the growth controls. On the other hand, rice handles waterlogging stress by forming lysigenous aerenchyma and a barrier to radial O₂ loss (ROL) in roots in order to supply O₂ to the root tip. In this article, we summarize recent advances in understanding the mechanisms of responding to excess water stresses (i.e., submergence and waterlogging) in rice and other gramineous plants.     

Foliar N/P ratio and nutrient limitation to vegetation growth on Keerqin sandy grassland of North‐east China

To examine whether the critical leaf N/P ratios (of 14, 16) are valid to test nutrient limitation in the context of semi‐arid sandy grasslands, an experiment was conducted on a Keerqin sandy grassland in North‐east China to investigate the responses of plant biomass and nutrient concentrations to fertilization. Plant biomass production and leaf nutrient concentrations were measured after five consecutive years of fertilization with N (20 g N m^?2 year^?1) and/or P (10 g P₂O₅ m^?2 year^?1). Nitrogen fertilization increased the shoot biomass by twofold and consequently the shoot/root ratio, whereas P fertilization had little effect on either shoot biomass or shoot/root ratio. Leaf N/P ratio varied among species with an average of 5·6 in the control, while the mean leaf N/P ratio (7·5) under the N fertilization treatment remained below the threshold of 14. Our results suggest that the critical N/P ratio (14, 16) is not applicable as a test for nutrient limitations in the context of semi‐arid, sandy grassland.