Diallel analysis for mineral element absorption in tropical adapted soybeans [Glycine max (L.) Merrill]

The Brazilian tropical adapted soybeans contains, in addition to superior morphological characters, genetic factors for tolerance to cultivation in acidic, mineral-stressed soils. However, the selection process for these hindrances has been empirical, and information on the genetics of mineral element uptake by the plant is necessary. The objective of this investigation was to identify the mode of inheritance for the absorption of phosphorus, potassium, calcium, magnesium, iron, aluminium, manganese, zinc and copper in a 9 × 9 diallel cross. General combining ability (GCA) was higher than specific combining ability (SCA), with the exception of copper, manganese and zinc, indicating predominantly additive effects. The ratios of GCA/SCA varied between 3.4 (calcium) and 8.5 (magnesium). The regression of covariance (W_r) on variance (V_r) showed that the additive-dominance model explained the genetic differences in this germ plasm. However, the detection of overdominance could be related to possible heterozygosity in the parental varieties for mineral absorption. Broad-sense heritability values were higher than narrow sense heritability values for aluminium, iron, potassium, calcium and magnesium, being in the range of 67.9–86.9% and 42.0–56.6%, respectively. This is an indication that soybeans can be further improved to efficient utilisation of nutrients and to tolerate toxic factors in the soil.     

等渗Ca(NO3)2和NaCl胁迫对黄瓜幼苗生长及渗透调节物质含量的影响

采用营养液培养方法,以耐盐性较弱的‘津春2号’黄瓜品种为试材,研究了等渗Ca(NO3)2和NaCl胁迫对黄瓜幼苗生长、根系电解质渗透率、根系活力、Na+和K+含量及渗透调节物质含量的影响。结果显示:(1)在84mmol.L-1 NaCl和56mmol.L-1 Ca(NO3)2等渗胁迫下,黄瓜幼苗鲜重和干重均显著下降,且NaCl处理下降的幅度大于等渗Ca(NO3)2处理。(2)NaCl主要通过对黄瓜根系的伤害来抑制植株生长,表现为根系活力下降、根系质膜透性增大、Na+大量积累、K+含量显著下降、Na+/K+明显上升,最终导致根冠比下降;而Ca(NO3)2处理对根系质膜透性、K+含量、Na+/K+的影响均小于NaCl胁迫,且根系活力和根冠比上升,但Ca(NO3)2胁迫后叶片含水量和渗透调节能力均小于NaCl胁迫。(3)NaCl胁迫条件下,黄瓜幼苗内渗透调节物质以可溶性糖为主,而Ca(NO3)2胁迫以可溶性蛋白为主。研究表明,NaCl胁迫对黄瓜幼苗的伤害大于等渗Ca(NO3)2,NaCl主要通过破坏根系质膜结构影响植株生长,而Ca(NO3)2主要通过引起地上部生理干旱来影响植株生长。     

Primary and Lateral Root Development of Dark- and Light-Grown Cotton Seedlings under Salinity Stress*

Primary root growth dynamics and lateral root development of dark- and light grown cotton seedlings (Gossypium hirsutum L., cv. Acala SJ-2) were studied under control and salinity stress conditions. The seedlings were grown by two methods: A) in paper-lined, vermiculite-filled beakers with the plants growing between the paper and the glass wall (Gladish and Rost, 1993), and B) in hydroponics after germination and initial growth in germination paper rolls saturated with the treatment solutions (Kent and Läuchli, 1985). After germination, daily primary root elongation rate gradually incrased to a maximum, then gradually declined to close to zero for dark-grown seedlings, or to sustained rates of about 10 mm per day for light-grown control plants. Salinity stress delayed primary root growth and reduced peak elongation rates, without changing the general primary root growth pattern. These results suggest that salinity changed the time-scale, but did not modify the normal developmental sequence. Lateral root growth was more inhibited by salinity than primary root growth. In addition, elongation of lateral roots was more inhibited by salinity than their initiation and emergence. Light exposure of the shoot favored both sustained primary root growth from 7 days after planting, and lateral root emergence and growth. Salinity effects were more severe on seedlings germinated and grown in hydroponics (method B) than on vermiculite-grown plants (method A). These results emphasize the importance of growing conditions for the NaCl-induced effects on cotton root development. In addition, the differential effects of salinity on primary and lateral roots became evident, pointing to diverse control mechanisms for the development of these root types.     

Root plasticity as the key root trait for adaptation to various intensities of drought stress in rice

Roots play an important role in rice adaptation to drought conditions. This study aimed to identify the key root traits that contribute to plant adaptation to drought stress. We used chromosome segment substitution lines (CSSLs) derived from Nipponbare and Kasalath crosses, which were grown in the field and hydroponics. In field experiments, the plants were grown under soil moisture gradients with line source sprinkler system up to around heading. Among the 54 CSSLs, only CSSL50 consistently showed significantly higher shoot dry matter production than its parent Nipponbare as the drought intensified for 3?years while most of the CSSLs reduced dry matter production to similar extents with Nipponbare under the same conditions. CSSL50 showed significantly greater total root length through promoted lateral root branching and elongation than Nipponbare, especially under mild stress conditions (15?30% w/w of soil moisture contents), which is considered as phenotypic plasticity. Such plastic root development was the key trait that effectively contributed to plant dry matter production through increased total root length and thus water uptake. However, there was no relationship between root plasticity and plant growth under the stress conditions induced by polyethylene glycol in hydroponics.     

In vitro evaluation of osmotic stress tolerance using a novel root recovery assay

We established a novel in vitro method, termed the root recovery assay, to evaluate the survival under osmotic stress of lettuce (Lactuca sativa L.) seedlings. Under salinity and drought stress, combination of the root-bending assay and root recovery assay showed the same trends in dry weight and survival rate as a hydroponic culture. Both in vitro assays and hydroponics ranked the three lettuce cultivars in the same order of drought tolerance. The root-bending assay evaluated the plant’s growth and the root recovery assay indicated the plant’s survival. In addition, the combined assay required less space and approximately half the time period compared with the hydroponic culture. These results suggested that application of the root-bending and root recovery assay should be a rapid and space-saving method with which to evaluate the osmotic stress tolerance of lettuce from both growth and survival standpoints. Hideki Maruyama and Ryohei Koyama contributed equally to this work.     

盐分胁迫对啤酒大麦幼苗生长、离子平衡和根际pH变化的影响

盐分胁迫不仅影响植物的生长,而且会影响植物根际微域环境。根际pH的改变对土壤养分的有效性和微生物群落组成的变化有重要影响。为了探究啤酒大麦幼苗对不同类型盐分胁迫的生理生态响应机制和根际pH变化影响的生理机制,采用水培法,通过不同类型盐分(对照、混合Na盐、混合Cl盐和NaCl)胁迫处理啤酒大麦幼苗,对其生长、离子平衡和根际pH变化进行了研究。结果表明,1)在3种不同类型盐分胁迫下,啤酒大麦幼苗地上部干重、含水量均有所降低,而根冠比增加,尤其在NaCl胁迫下啤酒大麦幼苗地上部干重较对照显著降低了17.88%,而根干重和根冠比则分别增加了19.12%和43.86%。不同类型盐分胁迫抑制了啤酒大麦幼苗根长的生长,尤其在混合Na盐胁迫下根长降低明显(P0.05),但促进了根表面积和根体积的增加,尤其在混合Cl盐胁迫下,根表面积和根体积分别增加了41.76%和84.38%。2)不同类型盐分胁迫下啤酒大麦幼苗地上部离子平衡发生改变,在混合Na盐和NaCl胁迫下啤酒大麦幼苗主要吸收Na~+,地上部K~+/Na~+、Ca~(2+)/Na~+和Mg~(2+)/Na~+显著降低;混合Cl盐和NaCl胁迫下则过量吸收Cl~-,抑制了H_2PO_4~-、NO_3~-和SO_4~(2-)的吸收。3)在混合Na盐、混合Cl盐和NaCl盐分胁迫下,啤酒大麦幼苗对阴离子的吸收总量高于对阳离子的吸收总量,离子平衡计算结果表明根际呈碱化现象,与原位显色结果一致,且在混合Cl盐胁迫下根际碱化程度最大。     

LaALMT1 mediates malate release from phosphorus-deficient white lupin root tips and metal root to shoot translocation

Under phosphorus (P) deficiency, Lupinus albus (white lupin) releases large amounts of organic acid anions from specialized root structures, so-called cluster or proteoid roots, to mobilize and acquire sparingly soluble phosphates from a restricted soil volume. The molecular mechanisms underlying this release and its regulation are, however, poorly understood. Here, we identified a gene belonging to the aluminium (Al)-activated malate transporter (ALMT) family that specifically contributes to malate, but not citrate release. This gene, LaALMT1, was most prominently expressed in the root apices under P deficiency, including those of cluster roots and was also detected in the root stele. Contrary to several ALMT homologs in other species, the expression was not stimulated, but moderately repressed by Al. Aluminium-independent malate currents were recorded from the plasma membrane localized LaALMT1 expressed in Xenopus oocytes. In composite lupins with transgenic roots, LaALMT1 was efficiently mutated by CRISPR-Cas9, leading to diminished malate efflux and lower xylem sap malate concentrations. When grown in an alkaline P-deficient soil, mutant shoot phosphate concentrations were similar, but iron and potassium concentrations were diminished in old leaves, suggesting a role for ALMT1 in metal root to shoot translocation, a function that was also supported by growth in hydroponics.     

Identification of QTLs for root characteristics in maize grown in hydroponics and analysis of their overlap with QTLs for grain yield in the field at two water regimes

We investigated the overlap among quantitative trait loci (QTLs) in maize for seminal root traits measured in hydroponics with QTLs for grain yield under well-watered (GY-WW) and water-stressed (GY-WS) field conditions as well as for a drought tolerance index (DTI) computed as GY-WS/GY-WW. In hydroponics, 11, 7, 9, and 10 QTLs were identified for primary root length (R1L), primary root diameter (R1D), primary root weight (R1W), and for the weight of the adventitious seminal roots (R2W), respectively. In the field, 7, 8, and 9 QTLs were identified for GY-WW, GY-WS, and DTI, respectively. Despite the weak correlation of root traits in hydroponics with GY-WW, GY-WS, and DTI, a noticeable overlap between the corresponding QTLs was observed. QTLs for R2W most frequently and consistently overlapped with QTLs for GY-WW, GY-WS, and/or DTI. At four QTL regions, an increase in R2W was positively associated with GY-WW, GY-WS, and/or DTI. A 10 cM interval on chromosome 1 between PGAMCTA205 and php20644 showed the strongest effect on R1L, R1D, R2W, GY-WW, GY-WS, and DTI. These results indicate the feasibility of using hydroponics in maize to identify QTL regions controlling root traits at an early growth stage and also influencing GY in the field. A comparative analysis of the QTL regions herein identified with those described in previous studies investigating root traits in different maize populations revealed a number of QTLs in common.     

Interaction of aluminium and drought stress on root growth and crop yield on acid soils

Background

Aluminium (Al) toxicity and drought stress are two major constraints for crop production in the world, particularly in the tropics. The variation in rainfall distribution and longer dry spells in much of the tropics during the main growing period of crops are becoming increasingly important yield-limiting factors with the global climate change. As a result, crop genotypes that are tolerant of both drought and Al toxicity need to be developed.

Scope

The present review mainly focuses on the interaction of Al and drought on root development, crop growth and yield on acid soils. It summarizes evidence from our own studies and other published/related work, and provides novel insights into the breeding for the adaptation to these combined abiotic stresses. The primary symptom of Al phytotoxicity is the inhibition of root growth. The impeded root system will restrict the roots for exploring the acid subsoil to absorb water and nutrients which is particularly important under condition of low soil moisture in the surface soil under drought. Whereas drought primarily affects shoot growth, effects of phytotoxic Al on shoot growth are mostly secondary effects that are induced by Al affecting root growth and function, while under drought stress root growth may even be promoted. Much progress has recently been made in the understanding of the physiology and molecular biology of the interaction between Al toxicity and drought stress in common bean (Phaseolus vulgaris L.) in hydroponics and in an Al-toxic soil.

Conclusions

Crops growing on acid soils yield less than their potential because of the poorly developed root system that limits nutrient and water uptake. Breeding for drought resistance must be combined with Al resistance, to assure that drought resistance is expressed adequately in crops grown on soils with acid Al-toxic subsoils.     

High‐throughput two‐dimensional root system phenotyping platform facilitates genetic analysis of root growth and development

High‐throughput phenotyping of root systems requires a combination of specialized techniques and adaptable plant growth, root imaging and software tools. A custom phenotyping platform was designed to capture images of whole root systems, and novel software tools were developed to process and analyse these images. The platform and its components are adaptable to a wide range root phenotyping studies using diverse growth systems (hydroponics, paper pouches, gel and soil) involving several plant species, including, but not limited to, rice, maize, sorghum, tomato and Arabidopsis. The RootReader2D software tool is free and publicly available and was designed with both user‐guided and automated features that increase flexibility and enhance efficiency when measuring root growth traits from specific roots or entire root systems during large‐scale phenotyping studies. To demonstrate the unique capabilities and high‐throughput capacity of this phenotyping platform for studying root systems, genome‐wide association studies on rice (Oryza sativa) and maize (Zea mays) root growth were performed and root traits related to aluminium (Al) tolerance were analysed on the parents of the maize nested association mapping (NAM) population.     

High-throughput imaging and analysis of root system architecture in Brachypodium distachyon under differential nutrient availability

Nitrogen (N) and phosphorus (P) deficiency are primary constraints for plant productivity, and root system architecture (RSA) plays a vital role in the acquisition of these nutrients. The genetic determinants of RSA are poorly understood, primarily owing to the complexity of crop genomes and the lack of sufficient RSA phenotyping methods. The objective of this study was to characterize the RSA of two Brachypodium distachyon accessions under different nutrient availability. To do so, we used a high-throughput plant growth and imaging platform, and developed software that quantified 19 different RSA traits. We found significant differences in RSA between two Brachypodium accessions grown on nutrient-rich, low-N and low-P conditions. More specifically, one accession maintained axile root growth under low N, while the other accession maintained lateral root growth under low P. These traits resemble the RSA of crops adapted to low-N and -P conditions, respectively. Furthermore, we found that a number of these traits were highly heritable. This work lays the foundation for future identification of important genetic components of RSA traits under nutrient limitation using a mapping population derived from these two accessions.     

Genetic and environmental differences in the mineral composition of ryegrass herbage

Summary The herbage of seven ryegrass clones, growing in the field as spaced clones in a randomised layout, was analysed for twelve mineral constituents. Significant differences in mineral composition of the clones were obtained for all minerals except potassium; nitrate, sulphate, and sodium showed the largest differences. The clones showed significant heritabilities for all the differences in mineral content except potassium; the extent to which these were due to additive genetic effects was not determined. In the case of nitrate, differences between clones persisted for twelve months.Interrelationships between differences in mineral composition, root cation-exchange capacities and herbage growth differences were examined. Total correlations were first calculated and then separated into genetic and environmental correlations, a number of which were significant. The most notable were genetic correlations between root cation-exchange capacities and five mineral constituents (iron, aluminium, titanium, nitrate, and acid-soluble phosphorus). No genetic correlations were found between growth and mineral differences.     

Varied tolerance to NaCl salinity is related to biochemical changes in two contrasting lettuce genotypes

Salt stress perturbs a multitude of physiological processes such as photosynthesis and growth. To understand the biochemical changes associated with physiological and cellular adaptations to salinity, two lettuce varieties (Verte and Romaine) were grown in a hydroponics culture system supplemented with 0, 100 or 200 mM NaCl. Verte displayed better growth under 100 mM NaCl compared to Romaine, but both genotypes registered relatively similar reductions in growth under 200 mM NaCl treatment. Both varieties showed differences in net photosynthetic activity in the absence of salt and 8 days after salt treatment. These differences diminished subsequently under prolonged salt stress (14 days). Verte showed enhanced leaf proline and restricted total cations especially Na⁺, lesser malondialdehyde (MDA) formation and lignification in the roots under 100 mM NaCl salinity. Membrane damage estimated by electrolyte leakage increased with elevated salt concentrations in roots of both varieties, but Verte had significantly lower electrolyte leakage relative to Romaine under 100 mM NaCl. Moreover, Verte also accumulated greater levels of carotenoids under increasing NaCl concentrations compared to Romaine. Taken together, these findings suggest that the greater tolerance of Verte to 100 mM NaCl is related to the more restricted accumulation of total cations and toxic Na⁺ in the roots and enhanced levels of antioxidative metabolites in root and leaf tissue.     

Transpiration does not control growth and nutrient supply in the amphibious plant Mentha aquatica

Mentha aquatica L. was grown at different nutrient availabilities in water and in air at 60% RH. The plants were kept at 600 mmol m^?3 free CO₂ dissolved in water (40 times air equilibrium) and at 30 mmol m^?3 CO₂ in air to ensure CO₂ saturation of growth in both environments. We quantified the transpiration-independent water transport from root to shoot in submerged plants relative to the transpiration stream in emergent plants and tested the importance of transpiration in sustaining nutrient flux and shoot growth. The acropetal water flow was substantial in submerged Mentha aquatica, reaching 14% of the transpiration stream in emergent plants. The transpiration-independent mass flow of water from the roots, measured by means of tritiated water, was diverted to leaves and adventitious shoots in active growth. The plants grew well and at the same rates in water and air, but nutrient fluxes to the shoot were greater in plants grown in air than in those that were submerged when they were rooted in fertile sediments. Restricted O₂ supply to the roots of submerged plants may account for the smaller nutrient concentrations, though these exceeded the levels required to saturate growth. In hydroponics, the root medium was aerated and circulated between submerged and emergent plants to minimize differences in medium chemistry, and here the two growth forms behaved similarly and could fully exploit nutrient enrichment. It is concluded that the lack of transpiration from leaf surfaces in a vapour-saturated atmosphere, or under water, is not likely to constrain the transfer of nutrients from root to shoot in herbaceous plants. Nutrient deficiency under these environmental conditions is more likely to derive from restricted development and function of the roots in waterlogged anoxic soils or from low porewater concentrations of nutrients.     

Changes in antioxidant gene expression and induction of oxidative stress in pea (<Emphasis Type="Italic">Pisum sativum</Emphasis> L.) under Al stress

Aluminium toxicity has been recognized as a primary growth-limiting factor in acid soil, resulting in a decrease in plant growth and production. In this experiment we have studied the induction of oxidative stress and changes in antioxidant gene expression in pea (var. ALASKA) under aluminium (Al) stress. We have found that Al treatment affected the growth of pea plant and induced oxidative stress with a change in antioxidant gene expression profile. While the expression of glutathione-s-transferase (GST) and catalase (CAT) was more in root, cytosolic Ascorbate peroxidase (cAPX) expression increased in shoot under aluminium stress. Copper- Zinc Superoxide dismutase (Cu-Zn SOD) gene expression was higher after 24 h but decreased after 48 h along with elevated expression of manganese superoxide dismutase (MnSOD) and iron-superoxide dismutase (FeSOD) at higher aluminium contentrations after 24 and 48 h. Aluminium stress elevated hydrogen peroxide (H₂O₂) level and affected the growth. The proline content did not change significantly, whereas glutathione content increased with a decreased ascorbate content under Al stress. The present study indicates that aluminium treatment affected the antioxidant gene expression and induced oxidative stress in pea plant.     

Mapping QTLs for root traits under different nitrate levels at the seedling stage in maize (Zea mays L.)

Nitrogen (N) loss is a worldwide problem in crop production. Apart from reasonable N fertilizer application, breeding N efficient cultivars provides an alternative way. Root architecture is an important factor determining N acquisition. However, little is known about the molecular genetic basis for root growth in relation to N supply. In the present study, an F₈ maize (Zea may L.) recombinant inbred (RI) population consisting of 94 lines was used to identify the QTLs for root traits under different nitrate levels. The lateral root length (LRL), axial root length (ARL), maximal axial root length (MARL), axial root number (ARN) and average axial root length (AARL) were evaluated under low N (LN) and high N (HN) conditions in a hydroponics system. A total of 17 QTLs were detected among which 14 loci are located on the same chromosome region as published QTLs for root traits. A major QTL on chromosome 1 (between bnlg1025 and umc2029) for the AARL under LN could explain 43.7% of the phenotypic variation. This QTL co-localizes with previously reported QTLs that associate with root traits, grain yield, and N uptake. Our results indicate that longer axial roots are important for efficient N acquisition and the major QTL for AARL may be used as a marker in breeding N efficient maize genotypes.     

Microbial ACC-Deaminase: Prospects and Applications for Inducing Salt Tolerance in Plants

Salinity is one of the most important stresses that hamper agricultural productivity in nearly every part of the world. Enhanced biosynthesis of ethylene in plants under salinity stress is well established. Higher ethylene concentration inhibits root growth and ultimately affects the overall plant growth. Overcoming this ethylene-induced root inhibition is a prerequisite for successful crop production. Recent studies have shown that ethylene level in plants is regulated by a key enzyme 1-aminocyclopropane-1-carboxylicacid (ACC)-deaminase. This enzyme is present in plant growth-promoting bacteria (PGPR) and lowers the ethylene level by metabolizing its precursor ACC into α-ketobutyrate and ammonia (NH₃). Inoculation of plants under salinity stress with PGPR having ACC-deaminase activity mitigates the inhibitory effects of salinity on root growth by lowering the ethylene concentration in the plant. This in turn results in prolific root growth, which is beneficial for the uptake of nutrients and maintenance of growth under stressful environment. The present review critically discusses the effects of salinity stress on plant growth with special reference to ethylene production and the effects of rhizobacteria containing ACC-deaminase on crop improvement under salinity stress. It also discusses how much progress has been made in producing transgenic lines of different crops over-expressing the gene encoding ACC-deaminase and how far such transformed lines can tolerate salinity stress.