Root cortical aerenchyma enhances the growth of maize on soils with suboptimal availability of nitrogen, phosphorus, and potassium

Root cortical aerenchyma (RCA) is induced by hypoxia, drought, and several nutrient deficiencies. Previous research showed that RCA formation reduces the respiration and nutrient content of root tissue. We used SimRoot, a functional-structural model, to provide quantitative support for the hypothesis that RCA formation is a useful adaptation to suboptimal availability of phosphorus, nitrogen, and potassium by reducing the metabolic costs of soil exploration in maize (Zea mays). RCA increased the growth of simulated 40-d-old maize plants up to 55%, 54%, or 72% on low nitrogen, phosphorus, or potassium soil, respectively, and reduced critical fertility levels by 13%, 12%, or 7%, respectively. The greater utility of RCA on low-potassium soils is associated with the fact that root growth in potassium-deficient plants was more carbon limited than in phosphorus- and nitrogen-deficient plants. In contrast to potassium-deficient plants, phosphorus- and nitrogen-deficient plants allocate more carbon to the root system as the deficiency develops. The utility of RCA also depended on other root phenes and environmental factors. On low-phosphorus soils (7.5 μM), the utility of RCA was 2.9 times greater in plants with increased lateral branching density than in plants with normal branching. On low-nitrate soils, the utility of RCA formation was 56% greater in coarser soils with high nitrate leaching. Large genetic variation in RCA formation and the utility of RCA for a range of stresses position RCA as an interesting crop-breeding target for enhanced soil resource acquisition.     

Architectural Tradeoffs between Adventitious and Basal Roots for Phosphorus Acquisition

Adventitious rooting contributes to efficient phosphorus acquisition by enhancing topsoil foraging. However, metabolic investment in adventitious roots may retard the development of other root classes such as basal roots, which are also important for phosphorus acquisition. In this study we quantitatively assessed the potential effects of adventitious rooting on basal root growth and whole plant phosphorus acquisition in young bean plants. The geometric simulation model SimRoot was used to dynamically model root systems with varying architecture and C availability growing for 21 days at 3 planting depths in 3 soil types with contrasting nutrient mobility. Simulated root architectures, tradeoffs between adventitious and basal root growth, and phosphorus acquisition were validated with empirical measurements. Phosphorus acquisition and phosphorus acquisition efficiency (defined as mol phosphorus acquired per mol C allocated to roots) were estimated for plants growing in soil in which phosphorus availability was uniform with depth or was greatest in the topsoil, as occurs in most natural soils. Phosphorus acquisition and acquisition efficiency increased with increasing allocation to adventitious roots in stratified soil, due to increased phosphorus depletion of surface soil. In uniform soil, increased adventitious rooting decreased phosphorus acquisition by reducing the growth of lateral roots arising from the tap root and basal roots. The benefit of adventitious roots for phosphorus acquisition was dependent on the specific respiration rate of adventitious roots as well as on whether overall C allocation to root growth was increased, as occurs in plants under phosphorus stress, or was lower, as observed in unstressed plants. In stratified soil, adventitious rooting reduced the growth of tap and basal lateral roots, yet phosphorus acquisition increased by up to 10% when total C allocation to roots was high and adventitious root respiration was similar to that in basal roots. With C allocation to roots decreased by 38%, adventitious roots still increased phosphorus acquisition by 5%. Allocation to adventitious roots enhanced phosphorus acquisition and efficiency as long as the specific respiration of adventitious roots was similar to that of basal roots and less than twice that of tap roots. When adventitious roots were assigned greater specific respiration rates, increased adventitious rooting reduced phosphorus acquisition and efficiency by diverting carbohydrate from other root types. Varying the phosphorus diffusion coefficient to reflect varying mobilities in different soil types had little effect on the value of adventitious rooting for phosphorus acquisition. Adventitious roots benefited plants regardless of basal root growth angle. Seed planting depth only affected phosphorus uptake and efficiency when seed was planted below the high phosphorus surface stratum. Our results confirm the importance of root respiration in nutrient foraging strategies, and demonstrate functional tradeoffs among distinct components of the root system. These results will be useful in developing ideotypes for more nutrient efficient crops.     

Theoretical evidence for the functional benefit of root cortical aerenchyma in soils with low phosphorus availability

Background and Aims

The formation of root cortical aerenchyma (RCA) reduces root respiration and nutrient content by converting living tissue to air volume. It was hypothesized that RCA increases soil resource acquisition by reducing the metabolic and phosphorus cost of soil exploration.

Methods

To test the quantitative logic of the hypothesis, SimRoot, a functional–structural plant model with emphasis on root architecture and nutrient acquisition, was employed. Sensitivity analyses for the effects of RCA on the initial 40 d of growth of maize (Zea mays) and common bean (Phaseolus vulgaris) were conducted in soils with varying degrees of phosphorus availability. With reference to future climates, the benefit of having RCA in high CO₂ environments was simulated.

Key Results

The model shows that RCA may increase the growth of plants faced with suboptimal phosphorus availability up to 70 % for maize and 14 % for bean after 40 d of growth. Maximum increases were obtained at low phosphorus availability (3 µm). Remobilization of phosphorus from dying cells had a larger effect on plant growth than reduced root respiration. The benefit of both these functions was additive and increased over time. Larger benefits may be expected for mature plants. Sensitivity analysis for light-use efficiency showed that the benefit of having RCA is relatively stable, suggesting that elevated CO₂ in future climates will not significantly effect the benefits of having RCA.

Conclusions

The results support the hypothesis that RCA is an adaptive trait for phosphorus acquisition by remobilizing phosphorus from the root cortex and reducing the metabolic costs of soil exploration. The benefit of having RCA in low-phosphorus soils is larger for maize than for bean, as maize is more sensitive to low phosphorus availability while it has a more ‘expensive’ root system. Genetic variation in RCA may be useful for breeding phosphorus-efficient crop cultivars, which is important for improving global food security.     

Rapid decline in nitrate uptake and respiration with age in fine lateral roots of grape: implications for root efficiency and competitive effectiveness

Changes in function as an individual root ages has important implications for understanding resource acquisition, competitive ability and optimal lifespan. Both nitrate uptake and respiration rates of differently aged fine roots of grape (Vitis rupestris x V. riparia cv. 3309 C) were measured. The resulting data were then used to simulate nitrate uptake efficiency and nutrient depletion as a function of root age. Both nitrate uptake and root respiration declined remarkably quickly with increasing root age. The decline in both N uptake and root respiration corresponded with a strong decline in root N concentration, suggesting translocation of nitrogen out of the roots. For simulations where no nutrient depletion occurs at the root surface, daily uptake efficiency was maximal at root birth and lifetime nitrate uptake efficiency slowly increased as the roots aged. Simulations of growth of roots into unoccupied soil using a solute transport model indicated the advantage of high uptake capacity in new roots under competitive conditions where nitrate availability is very transitory.     

Rhizoeconomics: Carbon costs of phosphorus acquisition

Plants display a wide array of physiological adaptations to low soil phosphorus availability. Here we discuss metabolic and ecological costs associated with these strategies, focusing on the carbon costs of root traits related to phosphorus acquisition in crop plants. We propose that such costs are an important component of adaptation to low phosphorus soils. In common bean, genotypes with superior low phosphorus adaptation express traits that reduce the respiratory burden of root growth, including greater allocation to metabolically inexpensive root classes, such as adventitious roots, and greater formation of cortical aerenchyma, which reduces specific root respiration. Root hair formation increases phosphorus acquisition at minimal carbon cost, but may have other unknown ecological costs. Mycorrhizas and root exudates enhance phosphorus acquisition in some taxa, but at significant carbon cost. Root architectural patterns that enhance topsoil foraging enhance phosphorus acquisition but appear to incur tradeoffs for water acquisition and spatial competition. A better understanding of the metabolic and ecological costs associated with phosphorus acquisition strategies is needed for an intelligent deployment of such traits in crop improvement programs.     

Root respiration and its relation to nutrient contents in soil and root and EVI among 8 ecosystems, northern China

Root respiration is a critical and uncertain component of ecosystem carbon budgets. We assessed whether variation in root respiration at a reference temperature were associated with that of root and soil nutrient contents and enhanced vegetation index (EVI) among 8 ecosystems, including three forests, two shrublands, two meadow grasslands, and one meadow in a forest-steppe ecotone in northern China. Mass specific root respiration was positively related to root nitrogen content when data from all the different classes of root diameter were pooled. Area specific root respiration increased with soil available nitrogen through increasing fine root biomass. Beside the effect of soil or root nutrient contents, mass specific respiration of fine roots followed EVI patterns, suggesting that root respiration was strongly dependent on photosynthesis. A simple respiration–EVI relationship may provide a convenient way to estimate root respiration and has potential applications in quantitative spatial extrapolation.     

夜间增温对冬小麦根系生长和土壤养分有效性的影响

气候变暖存在明显的昼夜不对称性,夜间气温升高幅度显著高于白天.本研究采用夜间被动式增温系统,于2009-2010年在我国冬小麦主产区（石家庄、徐州、许昌和镇江）进行全生育期田间增温试验,研究了土壤pH值、速效养分和抽穗期冬小麦根系对夜间增温的响应.结果表明： 与不增温对照相比,夜间增温显著降低了土壤pH值和速效养分含量,并在一定程度上提高了根系干质量和根冠比.冬小麦整个生育期,夜间增温分别使石家庄、徐州、许昌和镇江试验点土壤pH值平均降低0.4%、0.4%、0.7%和0.9%,碱解氮含量平均降低8.1%、8.1%、7.1%和6.0%,速效磷含量平均降低15.7%、12.1%、19.6%和25.8%;速效钾含量平均降低11.5%、7.6%、7.6%和10.1%.增温处理下,石家庄、徐州和镇江试验点抽穗期冬小麦根系干质量分别平均增加31.5%、27.0%和14.5%;石家庄、许昌和镇江试验点抽穗期冬小麦根冠比分别平均提高23.8%、13.7%和9.7%.夜间增温可能通过改变土壤化学特性影响土壤养分供应和冬小麦生长     

树木根系碳分配格局及其影响因子

根系作为树木提供养分和水分的“源”和消耗C的“汇”,在陆地生态系统C平衡研究中具有重要的理论意义。尽管20多年来的研究已经认识到根系消耗净初级生产力占总净初级生产力较大的比例,但是,根系（尤其是细根）消耗C的机理以及C分配的去向一直没有研究清楚。主要原因是细根消耗光合产物的生理生态过程相当复杂,准确估计各个组分消耗的C具有很大的不确定性,常常受树种和环境空间和时间异质性、以及研究方法的限制。综述了分配到地下的C主要去向,即细根生产和周转、呼吸及养分吸收与同化、分泌有机物、土壤植食动物,及有关林木地下碳分配机理的几种假说,分析了地下碳分配估计中存在的不确定性。目的是在全球变化C循环研究中对生态系统地下部分根系消耗的C以及分配格局引起重视。     

Heterogeneous nutrient supply promotes maize growth and phosphorus acquisition: additive and compensatory effects of lateral roots and root hairs

Background and AimsRoot proliferation is a response to a heterogeneous nutrient distribution. However, the growth of root hairs in response to heterogeneous nutrients and the relationship between root hairs and lateral roots remain unclear. This study aims to understand the effects of heterogeneous nutrients on root hair growth and the trade-off between root hairs and lateral roots in phosphorus (P) acquisition.MethodsNear-isogenic maize lines, the B73 wild type (WT) and the rth3 root hairless mutant, were grown in rhizoboxes with uniform or localized supply of 40 (low) or 140 (high) mg P kg⁻¹ soil.ResultsBoth WT and rth3 had nearly two-fold greater shoot biomass and P content under local than uniform treatment at low P. Significant root proliferation was observed in both WT and rth3 in the nutrient patch, with the WT accompanied by an obvious increase (from 0.7 to 1.2 mm) in root hair length. The root response ratio of rth3 was greater than that of WT at low P, but could not completely compensate for the loss of root hairs. This suggests that plants enhanced P acquisition through complementarity between lateral roots and root hairs, and thus regulated nutrient foraging and shoot growth. The disappearance of WT and rth3 root response differences at high P indicated that the P application reduced the dependence of the plants on specific root traits to obtain nutrients.ConclusionsIn addition to root proliferation, the root response to a nutrient-rich patch was also accompanied by root hair elongation. The genotypes without root hairs increased their investment in lateral roots in a nutrient-rich patch to compensate for the absence of root hairs, suggesting that plants enhanced nutrient acquisition by regulating the trade-off of complementary root traits.     

Root cortical aerenchyma inhibits radial nutrient transport in maize (Zea mays)

Background and Aims

Formation of root cortical aerenchyma (RCA) can be induced by nutrient deficiency. In species adapted to aerobic soil conditions, this response is adaptive by reducing root maintenance requirements, thereby permitting greater soil exploration. One trade-off of RCA formation may be reduced radial transport of nutrients due to reduction in living cortical tissue. To test this hypothesis, radial nutrient transport in intact roots of maize (Zea mays) was investigated in two radiolabelling experiments employing genotypes with contrasting RCA.

Methods

In the first experiment, time-course dynamics of phosphate loading into the xylem were measured from excised nodal roots that varied in RCA formation. In the second experiment, uptake of phosphate, calcium and sulphate was measured in seminal roots of intact young plants in which variation in RCA was induced by treatments altering ethylene action or genetic differences.

Key Results

In each of three paired genotype comparisons, the rate of phosphate exudation of high-RCA genotypes was significantly less than that of low-RCA genotypes. In the second experiment, radial nutrient transport of phosphate and calcium was negatively correlated with the extent of RCA for some genotypes.

Conclusions

The results support the hypothesis that RCA can reduce radial transport of some nutrients in some genotypes, which could be an important trade-off of this trait.     

Modeling root loss reveals impacts on nutrient uptake and crop development

Despite the widespread prevalence of root loss in plants, its effects on crop productivity are not fully understood. While root loss reduces the capacity of plants to take up water and nutrients from the soil, it may provide benefits by decreasing the resources required to maintain the root system. Here, we simulated a range of root phenotypes in different soils and root loss scenarios for barley (Hordeum vulgare), common bean (Phaseolus vulgaris), and maize (Zea mays) using and extending the open-source, functional–structural root/soil simulation model OpenSimRoot. The model enabled us to quantify the impact of root loss on shoot dry weight in these scenarios and identify in which scenarios root loss is beneficial, detrimental, or has no effect. The simulations showed that root loss is detrimental for phosphorus uptake in all tested scenarios, whereas nitrogen uptake was relatively insensitive to root loss unless main root axes were lost. Loss of axial roots reduced shoot dry weight for all phenotypes in all species and soils, whereas lateral root loss had a smaller impact. In barley and maize plants with high lateral branching density that were not phosphorus-stressed, loss of lateral roots increased shoot dry weight. The fact that shoot dry weight increased due to root loss in these scenarios indicates that plants overproduce roots for some environments, such as those found in high-input agriculture. We conclude that a better understanding of the effects of root loss on plant development is an essential part of optimizing root system phenotypes for maximizing yield.

Root loss has a major impact on crop development and nutrient uptake; modeling reveals that the magnitude of the effect depends on species, root phenotype, and soil conditions.     

Root cortical aerenchyma improves the drought tolerance of maize (Zea mays L.)

Root cortical aerenchyma (RCA) reduces root respiration in maize by converting living cortical tissue to air volume. We hypothesized that RCA increases drought tolerance by reducing root metabolic costs, permitting greater root growth and water acquisition from drying soil. To test this hypothesis, recombinant inbred lines with high and low RCA were observed under water stress in the field and in soil mesocosms in a greenhouse. In the field, lines with high RCA had 30% more shoot biomass at flowering compared with lines with low RCA under water stress. Root length density in deep soil was significantly greater in the high RCA lines compared with the low RCA lines. Mid‐day leaf relative water content in the high RCA lines was 10% greater than in the low RCA lines under water stress. The high RCA lines averaged eight times the yield of the low RCA lines under water stress. In mesocosms, high RCA lines had less seminal root respiration, deeper rooting, and greater shoot biomass compared with low RCA lines under water stress. These results support the hypothesis that RCA is beneficial for drought tolerance in maize by reducing the metabolic cost of soil exploration.     

海岸草本植物细根、叶片功能性状及其与土壤因子的关系

植物根、叶是对环境敏感性最高的器官,探究根叶功能性状之间的相互关系以及对环境因子的响应,有助于揭示植物对资源的利用状况及其对环境的适生策略。为探讨沿海植物的适应策略,该文以平潭岛砂质海岸草本植物为研究对象,由海及陆设置了3个距离梯度,选取6个叶功能性状和5个细根功能性状指标,分析海岸植物叶片与细根功能性状及其对土壤因子的响应。结果表明:(1)根、叶功能性状变异系数幅度在潮间带最小,在距高海潮线30～60 m的梯度上最大。单叶面积、叶磷含量、根平均直径、根组织密度、根磷含量随着由海到陆的距离增加呈上升趋势; 叶干物质含量、叶组织密度、比根长、比根面积呈下降趋势。(2)植物通过性状组合,在生长与防御间进行资源分配的权衡,表现在叶性状间、根性状间以及根-叶性状间具不同程度的相关性。其中,在地上-地下对应性状中,叶厚与根平均直径、叶磷含量与根磷含量呈极显著正相关; 而比叶面积与比根面积、比根长,叶组织密度与根组织密度均未表现出显著的相关性。(3)土壤因子对海岸植物功能性状变化的解释度为52.05%,其中土壤含盐量的影响最大,其次是土壤含水量、电导率、pH值。总体而言,在恶劣的海岸环境下,由海向陆土壤含盐量、电导率、含水量及pH值逐渐下降,整体为低磷高盐碱,植物表现为不同的生存策略:距海近的植物采取“叶片资源保守型、根系资源获取型”策略; 距海远的植物则采取“叶片资源获取型、根系资源保守型”。该研究结果为了解海岸草本植物对环境梯度变化的响应机制和适应性提供了一定参考价值,同时也利于通过分析土壤等环境特性按梯度筛选栽种适宜的物种,促进海岸植物的恢复和保护。     

Improving crop nutrient efficiency through root architecture modifications

Improving crop nutrient ef ficiency becomes an essential consideration for environmentally friendly and sustainable agriculture. Plant growth and development is dependent on 17 essential nutrient elements,among them,nitrogen(N) and phosphorus(P) are the two most important mineral nutrients. Hence it is not surprising that low N and/or low P availability in soils severely constrains crop growth and productivity,and thereby have become high priority targets for improving nutrient ef ficiency in crops. Root exploration largely determines the ability of plants to acquire mineral nutrients from soils. Therefore,root architecture,the 3-dimensional con figuration of the plant's root system in the soil,is of great importance for improving crop nutrient ef ficiency. Furthermore,the symbiotic associations between host plants and arbuscular mycorrhiza fungi/rhizobial bacteria,are additional important strategies to enhance nutrient acquisition. In this review,we summarize the recent advances in the current understanding of crop species control of root architecture alterations in response to nutrient availability and root/microbe symbioses,through gene or QTL regulation,which results in enhanced nutrient acquisition.     

Root hair length and rhizosheath mass depend on soil porosity,strength and water content in barley genotypes

Selecting plants with improved root hair growth is a key strategy for improving phosphorus-uptake efficiency in agriculture. While significant inter- and intra-specific variation is reported for root hair length, it is not known whether these phenotypic differences are exhibited under conditions that are known to affect root hair elongation. This work investigates the effect of soil strength, soil water content (SWC) and soil particle size (SPS) on the root hair length of different root hair genotypes of barley. The root hair and rhizosheath development of five root hair genotypes of barley (Hordeum vulgare L.) was compared in soils with penetrometer resistances ranging from 0.03 to 4.45 MPa (dry bulk densities 1.2–1.7 g cm^?3). A “short” (SRH) and “long” root hair (LRH) genotype was selected to further investigate whether differentiation of these genotypes was related to SWC or SPS when grown in washed graded sand. In low-strength soil (<1.43 MPa), root hairs of the LRH genotype were on average 25 % longer than that of the SRH genotype. In high-strength soil, root hair length of the LRH genotype was shorter than that in low-strength soil and did not differ from that of the SRH genotype. Root hairs were shorter in wetter soils or soils with smaller particles, and again SRH and LRH did not differ in hair length. Longer root hairs were generally, but not always, associated with larger rhizosheaths, suggesting that mucilage adhesion was also important. The root hair growth of barley was found to be highly responsive to soil properties and this impacted on the expression of phenotypic differences in root hair length. While root hairs are an important trait for phosphorus acquisition in dense soils, the results highlight the importance of selecting multiple and potentially robust root traits to improve resource acquisition in agricultural systems.     

A multiple ion-uptake phenotyping platform reveals shared mechanisms affecting nutrient uptake by roots

Nutrient uptake is critical for crop growth and is determined by root foraging in soil. Growth and branching of roots lead to effective root placement to acquire nutrients, but relatively little is known about absorption of nutrients at the root surface from the soil solution. This knowledge gap could be alleviated by understanding sources of genetic variation for short-term nutrient uptake on a root length basis. A modular platform called RhizoFlux was developed for high-throughput phenotyping of multiple ion-uptake rates in maize (Zea mays L.). Using this system, uptake rates were characterized for the crop macronutrients nitrate, ammonium, potassium, phosphate, and sulfate among the Nested Association Mapping (NAM) population founder lines. The data revealed substantial genetic variation for multiple ion-uptake rates in maize. Interestingly, specific nutrient uptake rates (nutrient uptake rate per length of root) were found to be both heritable and distinct from total uptake and plant size. The specific uptake rates of each nutrient were positively correlated with one another and with specific root respiration (root respiration rate per length of root), indicating that uptake is governed by shared mechanisms. We selected maize lines with high and low specific uptake rates and performed an RNA-seq analysis, which identified key regulatory components involved in nutrient uptake. The high-throughput multiple ion-uptake kinetics pipeline will help further our understanding of nutrient uptake, parameterize holistic plant models, and identify breeding targets for crops with more efficient nutrient acquisition.

A platform for quantifying root uptake rates of multiple, simultaneous nutrients reveals these rates are correlated among nutrients, are heritable, and may have a common genetic basis.     

Root hairs confer a competitive advantage under low phosphorus availability

Root hairs are presumably important in the acquisition of immobile soil resources such as phosphorus. The density and length of root hairs vary substantially within and between species, and are highly regulated by soil phosphorus availability, which suggests that at high nutrient availability, root hairs may have a neutral or negative impact on fitness. We used a root-hairless mutant of the small herbaceous dicot Arabidopsis thaliana to assess the effect of root hairs on plant competition under contrasting phosphorus regimes. Wildtype plants were grown with hairless plants in a replacement series design at high (60 m phosphate in soil solution) and low (1 m phosphate in soil solution) phosphorus availability. At high phosphorus availability, wildtype and mutant plants were equal in growth, phosphorus acquisition, fecundity and relative crowding coefficient (RCC). At low phosphorus availability, hairless plants accumulated less biomass and phosphorus, and produced less seed when planted with wildtype plants. Wildtype plants were unaffected by the presence of hairless plants in mixed genotype plantings. Wildtype plants had RCC values greater than one while hairless plants had RCC values less than one. We conclude that root hairs increase the competitiveness of plants under low phosphorus availability but do not reduce growth or competitiveness under high phosphorus availability.     

Detrital nutrient content and leaf species differentially affect growth and nutritional regulation of detritivores

Resource nutrient content and identity are common bottom–up controls on organismal growth and nutritional regulation. One framework to study these factors, ecological stoichiometry theory, predicts that elevated resource nitrogen (N) and phosphorus (P) contents enhance organism growth by alleviating constraints on N and P acquisition. However, the regulatory mechanisms underlying this response – including whether responses depend on resource identity – remain poorly understood. In this study, we tested roles of detrital N and P contents and identity (leaf species) in constraining growth of aquatic invertebrate detritivores. We synthesized results from seven detritivore species fed wide nutrient gradients of oak and maple detritus in the laboratory. Across detritivore taxa, we used a meta‐analytic approach quantifying effects of detrital leaf species and N and P contents on growth, consumption, and N‐ and P‐specific assimilation and growth efficiencies. Detritivore growth rates increased on higher‐N and P detritus and on oak compared to maple detritus. Notably, the mechanisms of improved growth differed between the responses to detrital nutrients versus leaf species, with the former driven by greater consumption rates despite lower assimilation efficiencies on higher‐nutrient detritus, and the latter driven by improved N and P assimilation and N growth efficiencies on oak detritus. These findings suggest animal nutrient acquisition changes flexibly in response to resource changes, altering the fate of detrital N and P throughout regulation. We affirm resource identity and nutrients as important bottom–up controls, but suggest these factors act through separate pathways to affect organism growth and thereby change detrital ecosystems under anthropogenic forest compositional change and nutrient enrichment.     

Reduced root cortical burden improves growth and grain yield under low phosphorus availability in maize

Root phenes and phene states that reduce the metabolic cost of soil exploration may improve plant growth under low phosphorus availability. We tested the hypothesis that under low phosphorus, reduced living cortical area (LCA) would increase soil exploration, phosphorus capture, biomass, and grain yield. Maize genotypes contrasting in LCA were grown in the field and in greenhouse mesocosms under optimal and suboptimal phosphorus regimes. Percent LCA in nodal roots ranged from 25% to 67%. Plants with 0.2 mm² less LCA under low phosphorus had 75% less root segment respiration, 54% less root phosphorus content, rooted 20 cm deeper, allocated up to four times more roots between 60 and 120 cm depth, had between 20% and 150% more biomass, 35–40% greater leaf phosphorus content, and 60% greater grain yield compared with plants with high LCA. Low‐LCA plants had up to 55% less arbuscular mycorrhizal colonization in axial roots, but this decrease was not correlated with biomass or phosphorus content. The LCA components cortical cell file number and cortical cell size were important for biomass and phosphorus content under low phosphorus. These results are consistent with the hypothesis that root phenes that decrease the metabolic cost of soil exploration are adaptive under phosphorus stress.     

Trait‐abundance relation in response to nutrient addition in a Tibetan alpine meadow: The importance of species trade‐off in resource conservation and acquisition

In competition‐dominated communities, traits promoting resource conservation and competitive ability are expected to have an important influence on species relative abundance (SRA). Yet, few studies have tested the trait‐abundance relations in the line of species trade‐off in resource conservation versus acquisition, indicating by multiple traits coordination. We measured SRA and key functional traits involving leaf economic spectrum (SLA, specific leaf area; LDMC, leaf dry matter content; LCC, leaf carbon concentration; LNC, leaf nitrogen concentration; LPC, leaf phosphorus concentration; Hs, mature height) for ten common species in all plots subjected to addition of nitrogen fertilizer (N), phosphorus fertilizer (P), or both of them (NP) in a Tibetan alpine meadow. We test whether SRA is positively related with traits promoting plant resource conservation, while negatively correlated with traits promoting plant growth and resource acquisition. We found that species were primarily differentiated along a trade‐off axis involving traits promoting nutrient acquisition and fast growth (e.g., LPC and SLA) versus traits promoting resource conservation and competition ability (e.g., large LDMC). We further found that SRA was positively correlated with plant height, LDMC, and LCC, but negatively associated with SLA and leaf nutrient concentration irrespective of fertilization. A stronger positive height‐SRA was found in NP‐fertilized plots than in other plots, while negative correlations between SRA and SLA and LPC were found in N or P fertilized plots. The results indicate that species trade‐off in nutrient acquisition and resource conservation was a key driver of SRA in competition‐dominated communities following fertilization, with the linkage between SRA and traits depending on plant competition for specific soil nutrient and/or light availability. The results highlight the importance of competitive exclusion in plant community assembly following fertilization and suggest that abundant species in local communities become dominated at expense of growth while infrequent species hold an advantage in fast growth and dispersals to neighbor meta‐communities.