Natural variation of the root morphological response to nitrate supply in Arabidopsis thaliana

Nitrogen fertilization increases crop yield but excessive nitrate use can be a major environmental problem due to soil leaching or greenhouse gas emission. Root traits have been seldom considered as selection criteria to improve Nitrogen Use Efficiency of crops, due to the difficulty of measuring root traits under field conditions. Nonetheless, learning about mechanisms of lateral root (LR) growth stimulation or repression by nitrate availability could help to redesign root system architecture (RSA), a strategy aimed at developing plants with a dense and profound root system and with higher N uptake efficiency. Here, we explored the genetic diversity provided by natural populations of the model species Arabidopsis thaliana to identify potentially adaptive differences in biomass production and root morphology in response to nitrate availability. A core collection of 24 accessions that maximizes the genetic diversity within the species and Col-0 (the reference accession) were grown vertically on agar medium at moderate (N+) nitrate level for 6 days and then transferred to the same condition or to low (N?) nitrate concentration for 7 days. There was a major nutritional effect on the shoot biomass and root to shoot biomass ratio. The variation of the root biomass and RSA traits (primary root length, LRs number, LR mean length, total LRs length and LR densities) was primarily genetically determined. Differences in RSA traits between accessions were somewhat more pronounced at N?. Some accessions produced almost no visible LRs (Pyl-1, N13) at N?, while other produced up to a dozen (Kn-0). Taken together our data illustrate that natural variation exists within Arabidopsis for the studied traits. The identification of RSA ideotypes in the N response will facilitate further analysis of quantitative traits for root morphology.     

Evolving technologies for growing,imaging and analyzing 3D root system architecture of crop plants

A plant's ability to maintain or improve its yield under limiting conditions,such as nutrient de ficiency or drought,can be strongly in fluenced by root system architecture(RSA),the three-dimensional distribution of the different root types in the soil. The ability to image,track and quantify these root system attributes in a dynamic fashion is a useful tool in assessing desirable genetic and physiological root traits. Recent advances in imaging technology and phenotyping software have resulted in substantive progress in describing and quantifying RSA. We have designed a hydroponic growth system which retains the three-dimensional RSA of the plant root system,while allowing for aeration,solution replenishment and the imposition of nutrient treatments,as well as high-quality imaging of the root system. The simplicity and flexibility of the system allows for modi fications tailored to the RSA of different crop species and improved throughput. This paper details the recent improvements and innovations in our root growth and imaging system which allows for greater image sensitivity(detection of fine roots and other root details),higher ef ficiency,and a broad array of growing conditions for plants that more closely mimic those found under field conditions.     

Dwarf alleles differentially affect barley root traits influencing nitrogen acquisition under low nutrient supply

Sustainable food production depends critically on the development of crop genotypes that exhibit high yield under reduced nutrient inputs. Rooting traits have been widely advocated as being able to influence optimal plant performance, while breeding-based improvements in yield of spring barley suggest that this species is a good model crop. To date, however, molecular genetics knowledge has not delivered realistic plant ideotypes, while agronomic trials have been unable to identify superior traits. This study explores an intermediate experimental system in which root traits and their effect on plant performance can be quantified. As a test case, four modern semi-dwarf barley varieties, which possess either the ari-e.GP or the sdw1 dwarf allele, were compared with the long-stemmed old variety Kenia under two levels of nutrient supply. The two semi-dwarf types differed from Kenia, exhibiting smaller stem mass and total plant nitrogen (N), and improved partitioning of mass and N to grain. Amongst the semi-dwarfs, the two ari-e.GP genotypes performed better than the two sdw1 genotypes under standard and reduced nutrient supply, particularly in root mass, root investment efficiency, N acquisition, and remobilization of N and mass to grain. However, lack of between-genotype variation in yield and N use efficiency indicated limited potential for exploiting genetic variation in existing varieties to improve barley performance under reduced nutrient inputs. Experimental approaches to test the expression of desirable root and shoot traits are scrutinized, and the potential evaluated for developing a spring barley ideotype for low nutrient conditions.     

Selection of common bean (Phaseolus vulgaris L.) for N2 fixation at different levels of available phosphorus under field and environmentally-controlled conditions

Phosphorus is one of several factors which affect N₂ fixation and along with N, is a principal yield-limiting nutrient in many regions of the world. Since the legume plant is an essential partner in symbiotic N₂ fixation, knowledge of host genotype variability for this process at different levels of P availability will be useful when breeding bean cultivars for enhanced N₂ fixation. The objective of this study was to obtain common bean (Phaseolus vulgaris L.) lines able with enhanced ability to support biological N₂ fixation under different levels of available phosphorus.Experiments were conducted in a growth room using a sand-alumina system to provide different levels of available P and in the field on a low-N soil. In the growth room studies, P availability strongly affected plant growth and traits related to N₂ fixation. No significant interaction was detected for P levels × bean lines, indicating that bean lines performed similarly at both high and low P levels.Total shoot N was used as a direct and indirect measure of N₂-fixation potential under growth room (N-free media) and field (low-N soil) conditions, respectively. Based on this criterion, two of the 41 and 54 inbred backcross lines of a segregating population evaluated in the growth room and the field, respectively, contained greater shoot N content than the recurrent parent and N shoot contents similar to the donor parent. Variability of N₂ fixation under low available P was observed, and high N₂ fixing and high yielding progeny lines were detected.     

Variability of traits associated with phosphorus efficiency in wild and cultivated genotypes of common bean

Genetic variation in plant growth under limited phosphorus (P) supply is necessary to obtain more productive cultivars on low P-available soils. Two pot experiments were conducted to evaluate the variability of some traits associated with efficiency of P absorption and utilization in wild and cultivated genotypes of common bean (Phaseolus vulgaris L.) under biological N2 fixation. At two P levels (20 and 80 mg P kg-1 soil, P1 and P2, respectively), 20 wild and 6 cultivated genotypes were grown in Experiment 1, and 4 wild and 27 cultivated genotypes were grown in Experiment 2. Plants were harvested at flowering, but in Experiment 1 wild accessions that did not flower were harvested at the beginning of leaf senescence. In Experiment 1, part of the genotypic variability of wild accessions was attributed to a less homogeneous ontogenetic stage at harvest, whereas in Experiment 2 some variation in biomass production was due to distinct phenologies of cultivated genotypes. Wild lines did not seem more tolerant to low P conditions, but the genotypic variation observed suggests these materials as a source of genetic diversity. Part of the variation in the root area and root efficiency ratio (total P content:root area) was compensatory, resulting in narrow genotypic differences in the total P content. The total P content and root efficiency ratio presented a wider amplitude of variation at P2 than at P1, and P uptake was more influenced by P supply than root production. Since the genotype × P level interaction was not significant for shoot biomass and shoot P concentration in Experiment 2, P utilization efficiency may be a useful selection criterion for cultivars between limited and adequate P supply. Within the sample of genetic diversity evaluated herein, there was large genotypic variability for traits related to P efficiency among wild and cultivated genotypes of common bean.     

From lab to field, new approaches to phenotyping root system architecture

Plant root system architecture (RSA) is plastic and dynamic, allowing plants to respond to their environment in order to optimize acquisition of important soil resources. A number of RSA traits are known to be correlated with improved crop performance. There is increasing recognition that future gains in productivity, especially under low input conditions, can be achieved through optimization of RSA. However, realization of this goal has been hampered by low resolution and low throughput approaches for characterizing RSA. To overcome these limitations, new methods are being developed to facilitate high throughput and high content RSA phenotyping. Here we summarize laboratory and field approaches for phenotyping RSA, drawing particular attention to recent advances in plant imaging and analysis. Improvements in phenotyping will facilitate the genetic analysis of RSA and aid in the identification of the genetic loci underlying useful agronomic traits.     

模拟氮沉降对低磷胁迫下马尾松生长和磷效率的影响

近年来大气N沉降日趋严重,导致森林土壤有效N含量增加,N/P发生改变,将会影响低P胁迫下林木的生长发育和P效率.本文以马尾松家系为研究对象,设置模拟N沉降与同质低P(介质表层与深层均缺P)、异质低P(介质表层P丰富、深层缺P)耦合的盆栽试验,研究N沉降对马尾松生长性状以及P吸收和利用效率的影响.结果表明： 同质低P下,模拟N沉降对马尾松生长性状和P效率影响较小,但存在显著的N×家系互作效应,家系40×44和71×20在N沉降后生物量增加,家系36×29和73×23生物量降低;异质低P下,模拟N沉降显著增加了马尾松苗高、生物量以及P吸收效率等,其原因是促进了根系生长和表层土壤中根系分布比例的增加.不同P环境下模拟N沉降对马尾松生长的影响,还与植株N/P有关.同质低P环境下,马尾松植株N/P为13.8,植株对N敏感性低,酸性磷酸酶活性增加,但未改善马尾松生长状况.在异质低P环境中,植株N/P为9.7,模拟N沉降显著增加了苗木生物量和P吸收效率,但未显著改变酸性磷酸酶活性.     

Phosphorus and Nitrogen Interactions in Field-Grown Soybean as Related to Genetic Attributes of Root Morphological and Nodular Traits

Two field experiments with different soybean (Glycine max L.) materials were conducted to investigate the interactions between phosphorus (P) and nitrogen (N) as related to the genetic attributes of root morphological and nodular traits. In experiment one, 13 cultivated soybean varieties were grown in a field with relatively low soil P and N availability. P application with 160 kg P/hm2 as triple superphosphate produced a significant simultaneous increase in the content of both P and N in shoot, demonstrating positive P and N interactions. The addition of P also increased root dry weight, root nodule number, nodule mass, nodule size, and nodulation index, but decreased root length and root surface area, indicating that P may affect N nutrition in soybean through a number of root morphological and nodular traits. Interestingly,like P content, N content appeared to be more correlated with root morphological traits (root weight, root length, and root surface area) than with root nodular traits (nodule number, nodule size, nodule mass, and nodulation index) at both P levels, implying that N taken up by the roots may contribute more to the plant N status than biological N2 fixation under the present experimental conditions. In experiment two, 57 soybean lines of a recombinant inbred line (RIL) population derived from a cross between a cultivated variety and a wild genotype were grown on another field site with moderately sufficient P and N levels to further characterize the genetic attributes of root morphological and nodular traits and their relationships with P and N interactions. The results indicated that all morphological and nodular traits measured continually segregated in the RIL population with a normal distribution of the phenotypic values, indicating that these traits are possibly controlled by quantitative trait loci (QTLs). Genetic analysis revealed that all these root traits had relatively low heritabilities (h2b=74.12, 70.65, 73.76, 56.34, 52.59, and 52.24 for root weight, root length,root surface area, nodule number, nodule mass, and nodule size, respectively), suggesting that root morphology and nodule formation are influenced greatly by environmental factors. Correlation analysis of the RILs showed that shoot N content was significantly correlated with P content, confirming positive P×N interactions. Similar to experiment one, shoot N content was only significantly correlated with root morpho logical traits, but not with root nodular traits, again denoting the fact that the N status in soybean could be attributed more to N uptake from the soil than to biological N2 fixation under the present experimental conditions.     

Phenotype of Arabidopsis thaliana semi-dwarfs with deep roots and high growth rates under water-limiting conditions is independent of the GA5 loss-of-function alleles

Background and Aims The occurrence of Arabidopsis thaliana semi-dwarf accessions carrying inactive alleles at the gibberellin (GA) biosynthesis GA5 locus has raised the question whether there are pleiotropic effects on other traits at the root level, such as rooting depth. In addition, it is unknown whether semi-dwarfism in arabidopsis confers a growth advantage under water-limiting conditions compared with wild-type plants. The aim of this research was therefore to investigate whether semi-dwarfism has a pleiotropic effect in the root system and also whether semi-dwarfs might be more tolerant of water-limiting conditions.Methods The root systems of different arabidopsis semi-dwarfs and GA biosynthesis mutants were phenotyped in vitro using the GROWSCREEN-ROOT image-based software. Semi-dwarfs were phenotyped together with tall, near-related accessions. In addition, root phenotypes were investigated in soil-filled rhizotrons. Rosette growth trajectories were analysed with the GROWSCREEN-FLUORO setup based on non-invasive imaging.Key Results Mutations in the early steps of the GA biosynthesis pathway led to a reduction in shoot as well as root size. Depending on the genetic background, mutations at the GA5 locus yielded phenotypes characterized by decreased root length in comparison with related wild-type ones. The semi-dwarf accession Pak-3 showed the deepest root system both in vitro and in soil cultivation experiments; this comparatively deep root system, however, was independent of the ga5 loss-of-function allele, as shown by co-segregation analysis. When the accessions were grown under water-limiting conditions, semi-dwarf accessions with high growth rates were identified.Conclusions The observed diversity in root system growth and architecture occurs independently of semi-dwarf phenotypes, and is probably linked to a genetic background effect. The results show that there are no clear advantages of semi-dwarfism at low water availability in arabidopsis.     

Close linkages between leaf functional traits and soil and leaf C:N:P stoichiometry under altered precipitation in a desert steppe in northwestern China

Leaf functional traits are important for characterizing plant nutrient strategies. The C:N:P stoichiometric balance in soils and plants, which could indicate types of nutrient limitation, is altered under changing precipitation patterns. However, whether such alterations affect leaf functional traits remains unclear. We conducted a three-year simulated precipitation experiment in a desert steppe in northwestern China to determine changes in leaf photosynthetic traits and nutrient conservation traits in five plant species and tested the relationships of these traits with soil and leaf C:N:P stoichiometry. The five species showed few changes in their leaf traits under drought conditions, but they adjusted these traits (especially P traits) under extremely wet conditions (50% increase in precipitation). Improved leaf photosynthetic N and P use, lowered leaf P uptake, and enhanced leaf N resorption might help Lespedeza potaninii to rely less on soil nutrients in extremely wet environments than other species do. Leaf photosynthetic traits were regulated primarily by soil and leaf C:N:P stoichiometry. Leaf nutrient conservation traits were controlled by both leaf C:N:P stoichiometry and soil properties (i.e., enzyme activity and microbial biomass), a condition especially true for P traits. The results suggest that precipitation-induced alteration in the C:N:P stoichiometric balance might have important influences on plant nutrient use strategies and even on the nutrient cycling of desert steppes.

     

Root and leaf attributes accounting for the performance of fast- and slow-growing grasses at different nutrient supply

Despite their difference in potential growth rate, the slow-growing Brachypodium pinnatum and the fast-growing Dactylis glomerata co-occur in many nutrient-poor calcareous grasslands. They are known to respond differently to increasing levels of N and P. An experiment was designed to measure which characteristics are affected by nutrient supply and contribute to the ecological performance of these species. Nutrient acquisition and root and shoot traits of these grasses were studied in a garden experiment with nine nutrient treatments in a factorial design of 3 N and 3 P levels each. D. glomerata was superior to B. pinnatum in nutrient acquisition and growth in all treatments. B. pinnatum was especially poor in P acquisition. Both species responded to increasing N supply and to a lesser extent to increasing P supply by decreasing their root length and increasing their leaf area per total plant weight. D. glomerata showed a higher plasticity. In most treatments, the root length ratio (RLR) and the leaf area ratio (LAR) were higher for D. glomerata. A factorization of these parameters into components expressing biomass allocation, form (root fineness or leaf thickness) and density (dry matter content) shows that the low density of the biomass of D. glomerata was the main cause for the higher RLR and LAR. The biomass allocation to the roots showed a considerable plasticity but did not differ between the species. B. pinnatum had the highest leaf weight ratio. Root fineness was highly plastic in D. glomerata, the difference with B. pinnatum being mainly due to the thick roots of D. glomerata at high nutrient supply. The leaf area/leaf fresh weight ratio did not show any plasticity and was slightly higher for B. pinnatum. It is concluded, that the low density of the biomass of D. glomerata is the pivotal trait responsible for its faster growth at all nutrient levels. It enables simultaneously a good nutrient acquisition capacity by the roots as well as a superior carbon acquisition by the leaves. The high biomass density of B. pinnatum will then result in a lower nutrient requirement due to a slower turnover, which in the long term is advantageous under nutrient-poor conditions.     

Mechanical stimulation in Brachypodium distachyon: Implications for fitness,productivity, and cell wall properties

Mechanical stimulation, including exposure to wind, is a common environmental variable for plants. However, knowledge about the morphogenetic response of the grasses (Poaceae) to mechanical stimulation and impact on relevant agronomic traits is very limited. Two natural accessions of Brachypodium distachyon were exposed to wind and mechanical treatments. We surveyed a wide range of stem-related traits to determine the effect of the two treatments on plant growth, development, and stem biomass properties. Both treatments induced significant quantitative changes across multiple scales, from the whole plant down to cellular level. The two treatments resulted in shorter stems, reduced biomass, increased tissue rigidity, delayed flowering, and reduced seed yield in both accessions. Among changes in cell wall-related features, a substantial increase in lignin content and pectin methylesterase activity was most notable. Mechanical stimulation also reduced the enzymatic sugar release from the cell wall, thus increasing biomass recalcitrance. Notably, treatments had a distinct and opposite effect on vascular bundle area in the two accessions, suggesting genetic variation in modulating these responses to mechanical stimulation. Our findings highlight that exposure of grasses to mechanical stimulation is a relevant environmental factor affecting multiple traits important for their utilization in food, feed, and bioenergy applications.     

Resource allocation to growth or luxury consumption drives mycorrhizal responses

Highly variable phenotypic responses in mycorrhizal plants challenge our functional understanding of plant‐fungal mutualisms. Using non‐invasive high‐throughput phenotyping, we observed that arbuscular mycorrhizal (AM) fungi relieved phosphorus (P) limitation and enhanced growth of Brachypodium distachyon under P‐limited conditions, while photosynthetic limitation under low nitrogen (N) was exacerbated by the fungus. However, these responses were strongly dependent on host genotype: only the faster growing genotype (Bd3‐1) utilised P transferred from the fungus to achieve improved growth under P‐limited conditions. Under low N, the slower growing genotype (Bd21) had a carbon and N surplus that was linked to a less negative growth response compared with the faster growing genotype. These responses were linked to the regulation of N : P stoichiometry, couples resource allocation to growth or luxury consumption in diverse plant lineages. Our results attest strongly to a mechanism in plants by which plant genotype‐specific resource economics drive phenotypic outcomes during AM symbioses.     

硝态氮（NO3^—）对水稻侧根生长及其氮吸收的影响

侧根是植物吸收利用土壤养分的重要器官 ,其生长发育受内部遗传因子和外部环境矿质养分的影响。通过琼脂分层培养发现 :局部供应NO-3 可以诱导水稻 (OryzasativaL .)主根或不定根上侧根的生长。为研究旱种条件下NO-3 对水稻侧根发育及其N吸收的影响 ,设置了 3个蛭石培养实验 :分根处理、全株缺N、全株供N处理。分根处理 (一半根系供应 3mmol/LKNO3,另一半根系供应 3mmol/LKCl)结果表明 :局部供应NO-3 能够促进水稻侧根生长。而在全株处理下 ,N饥饿诱导了侧根的伸长。水稻根系对NO-3 的这两种反应都存在着显著的基因型差异。同时对地上部N浓度、可溶性总糖含量及N含量分析表明 ,这些生理指标在分根处理与全株加N处理中的差异均不显著 ,表明分根处理也能基本满足植株正常生长对N的需求。在分根处理中 ,水稻的N含量与分根处理中供N一侧的平均侧根长度存在显著正相关 ,这表明在养分不均一的介质中 ,侧根长度对水稻N素吸收具有十分重要的作用。而在N素充足的条件下 ,两者之间的相关性并不显著 ,这暗示在养分充足的环境下 ,侧根长度可能并不是决定根系吸收N素的主要因素     

Genetic dissection of seminal root architecture in elite durum wheat germplasm

Although root architecture has been shown to play an important role in crop performance, particularly under drought conditions, no information is available on the genetic control of root traits in durum wheat, a crop largely grown in rainfed areas with low rainfall. In our study, a panel of 57 elite durum wheat accessions were evaluated under controlled conditions for root and shoot traits at the seedling stage. Significant genetic variability was detected for all the root and shoot traits that were investigated. Correlation analysis suggested that root and shoot features were only partially controlled by common sets of genes. The high linkage disequilibrium (up to 5 cM) present in the germplasm collection herein considered allowed us to use simple sequence repeat‐based association mapping to identify chromosome regions with significant effects on the investigated traits. In total, 15 chromosome regions showed significant effects on one or more root architectural features. A number of these regions also influenced shoot traits and, in some cases, plant height measured in field conditions. Major effects were detected on chromosome arms 2AL (at Xgwm294), 7AL (at Xcfa2257 and Xgwm332) and 7BL (at Xgwm577 and Xcfa2040). The accessions with the most remarkable differences in root features will provide a valuable opportunity to assemble durum wheat mapping populations well suited for ascertaining the effects of root architecture on water use efficiency and grain yield.     

硝态氮(NO-3)对水稻侧根生长及其氮吸收的影响

侧根是植物吸收利用土壤养分的重要器官,其生长发育受内部遗传因子和外部环境矿质养分的影响.通过琼脂分层培养发现:局部供应NO-3可以诱导水稻( Oryza sativa L.)主根或不定根上侧根的生长.为研究旱种条件下NO-3对水稻侧根发育及其N吸收的影响,设置了3个蛭石培养实验:分根处理、全株缺N、全株供N处理.分根处理(一半根系供应3 mmol/L KNO3,另一半根系供应3 mmol/L KCl)结果表明:局部供应NO-3 能够促进水稻侧根生长.而在全株处理下,N饥饿诱导了侧根的伸长.水稻根系对NO-3的这两种反应都存在着显著的基因型差异.同时对地上部N浓度、可溶性总糖含量及N含量分析表明,这些生理指标在分根处理与全株加N处理中的差异均不显著,表明分根处理也能基本满足植株正常生长对N的需求.在分根处理中,水稻的N含量与分根处理中供N一侧的平均侧根长度存在显著正相关,这表明在养分不均一的介质中,侧根长度对水稻N素吸收具有十分重要的作用.而在N素充足的条件下,两者之间的相关性并不显著,这暗示在养分充足的环境下,侧根长度可能并不是决定根系吸收N素的主要因素.     

纯化腐植酸对低氮胁迫下黄瓜幼苗生长及养分吸收的影响

以黄瓜‘新泰密刺’、‘津优1号’为供试品种,研究沙培条件下不同浓度(0、50、100、150、200 mg·L^-1)纯化腐植酸(PHA)浇灌对低氮胁迫(1 mmol·L^-1 NO₃^-)下黄瓜幼苗生长及养分吸收的影响.结果表明: 在较低N供应条件下,沙培浇灌PHA可显著增加黄瓜幼苗的根总长、根表面积及根尖数,增大根体积,促进黄瓜幼苗株高和茎粗生长,增大叶面积;显著提高黄瓜幼苗叶片中脯氨酸及可溶性糖含量;促进N素以及P、K、Ca、Mg、Fe元素的吸收.由参试两黄瓜品种对低氮胁迫下PHA处理响应效果来看,不同品种的某些性状对PHA处理浓度的敏感程度稍有差异,综合结果显示,施用100～150 mg·L^-1PHA可显著促进两品种幼苗生长及养分吸收.     

Exploring the natural variation for seedling traits and their link with seed dimensions in tomato

The success of germination, growth and final yield of every crop depends to a large extent on the quality of the seeds used to grow the crop. Seed quality is defined as the viability and vigor attribute of a seed that enables the emergence and establishment of normal seedlings under a wide range of environments. We attempt to dissect the mechanisms involved in the acquisition of seed quality, through a combined approach of physiology and genetics. To achieve this goal we explored the genetic variation found in a RIL population of Solanum lycopersicum (cv. Moneymaker) x Solanum pimpinellifolium through extensive phenotyping of seed and seedling traits under both normal and nutrient stress conditions and root system architecture (RSA) traits under optimal conditions. We have identified 62 major QTLs on 21 different positions for seed, seedling and RSA traits in this population. We identified QTLs that were common across both conditions, as well as specific to stress conditions. Most of the QTLs identified for seedling traits co-located with seed size and seed weight QTLs and the positive alleles were mostly contributed by the S. lycopersicum parent. Co-location of QTLs for different traits might suggest that the same locus has pleiotropic effects on multiple traits due to a common mechanistic basis. We show that seed weight has a strong effect on seedling vigor and these results are of great importance for the isolation of the corresponding genes and elucidation of the underlying mechanisms.