Stimulation of root hair elongation in Arabidopsis thaliana by low phosphorus availability

Low phosphorus availability stimulates root hair elongation in many plants, which may have adaptive significance in soil phosphorus acquisition. We investigated the effect of low phosphorus on the elongation of Arabidopsis thaliana root hairs. Arabidopsis thaliana plants were grown in plant culture containing high (1000 mmol m^?3) or low (1 mmol m^?3) phosphorus concentrations, and root hair elongation was analysed by image analysis. After 15d of growth, low-phosphorus plants developed root hairs averaging 0.9 mm in length while high-phosphorus plants of the same age developed root hairs averaging 0.3 mm in length. Increased root hair length in low-phosphorus plants was a result of both increased growth duration and increased growth rate. Root hair length decreased logarithmically in response to increasing phosphorus concentration. Local changes in phosphorus availability influenced root hair growth regardless of the phosphorus status of the plant. Low phosphorus stimulated root hair elongation in the hairless axr2 mutant, exogenously applied IAA stimulated root hair elongation in wild-type high-phosphorus plants and the auxin antagonist CM PA inhibited root hair elongation in low-phosphorus plants. These results indicate that auxin may be involved in the low-phosphorus response in root hairs.     

The efficiency of Arabidopsis thaliana (Brassicaceae) root hairs in phosphorus acquisition

Arabidopsis thaliana root hairs grow longer and denser in response to low-phosphorus availability. In addition, plants with the root hair response acquire more phosphorus than mutants that have root hairs that do not respond to phosphorus limiting conditions. The purpose of this experiment was to determine the efficiency of root hairs in phosphorus acquisition at high- and low-phosphorus availability. Root hair growth, root growth, root respiration, plant phosphorus uptake, and plant phosphorus content of 3-wk-old wild-type Arabidopsis (WS) were compared to two root hair mutants (rhd6 and rhd2) under high (54 mmol/m) and low (0.4 mmol/m) phosphorus availability. A cost-benefit analysis was constructed from the measurements to determine root hair efficiency. Under high-phosphorus availability, root hairs did not have an effect on any of the parameters measured. Under low-phosphorus availability, wild-type Arabidopsis had greater total root surface area, shoot biomass, phosphorus per root length, and specific phosphorus uptake. The cost-benefit analysis shows that under low phosphorus, wild-type roots acquire more phosphorus for every unit of carbon respired or unit of phosphorus invested into the roots than the mutants. We conclude that the response of root hairs to low-phosphorus availability is an efficient strategy for phosphorus acquisition.     

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.     

在控制条件下云南松幼苗根系对低磷胁迫的响应

磷是控制生命过程的重要元素,植物在生长过程中需要大量的磷,低磷常导致一些植物发生适应性变化。云南松(PinusyunnanensisFranch.)以云南高原为起源和分布中心,其对低磷土壤环境表现出了很强的适应能力,广泛分布并正常生长于贫瘠的低磷红壤上,研究云南松对低磷环境的适应机制,对人类探索高效利用有限的磷素资源的方法具有现实意义。本实验通过对不同磷处理水平下培养的云南松幼苗根系生物量和根冠比等的研究,分析了云南松幼苗根系对低磷胁迫的响应。实验所用云南松种子采集自云南省通海县秀山森林公园内的健壮云南松林。结果表明:当磷浓度下降到0.5mmol/L时,云南松幼苗主根长度开始随磷浓度的降低而增加,根冠比随磷浓度的降低而增大,而侧根发生数没有随磷浓度的降低而显示出显著的增减规律,根系生物量也没有随磷浓度的降低而呈现出有规律的增减,根系生物量始终保持在一定的水平。进一步的分析表明:低磷胁迫下,云南松幼苗保证了物质分配对根的优先地位,以维持其根的生物量在一定水平,进而维持整个生命;云南松幼苗主要是靠主根长度的增加而不是靠侧根数量的增加来适应低磷环境。     

The formation,morphology and anatomy of cluster root of Lupinus albus L. as dependent on soil type and phosphorus supply

The effect of phosphorus supply on the formation, morphology and anatomy of cluster roots of Lupinus albus L. cv Ultra grown in a loam and two sandy soils was examined relative to its effect on total root length, shoot weight and the phosphorus concentration of the shoots. The loam soil was most conducive to the formation of cluster roots. Cluster roots growing in the sandy soils developed to a lesser extent on plants of an equivalent phosphorus status, suggesting that some biotic or abiotic factors independent of phosphorus supply were also operating. The presence of mature cluster rootlets on a length of lateral root increased the root surface area by 14–22 times of an equal length of lateral roots not bearing cluster rootlets. The application of phosphorus decreased cluster-root length, whereas total root length showed a steady increase. There was an inverse relationship between cluster-root production and phosphorus concentration in shoots ranging from 2 to 8.5 mg g^–1 with the critical phosphorus level for maximum shoot growth being around 2.5 mg g^–1. Cluster roots formed in solution culture were not well developed in comparison with those grown in the loam soil or nutrient solution with added loam soil. The organisation of the cluster rootlet was similar to that of the lateral roots. Mature rootlets lacked an apical meristem and a vascular cambium with a reduced root cap and cortical tissue.     

The influence of the phosphorus and nitrogen nutrition of pea plants on the growth of their progeny

Summary In glasshouse experiments with peas grown in nutrient cultures, seed from phosphorus-deficient plants contained much lower phosphorus concentrations than seed from plants not deficient in this element, but differences in the supply of mineral nitrogen to the plants (which were nodulated) had little effect on the chemical composition of the seed.Lower haulm weights and pea yields were obtained from sowings of low-phosphorus as compared with high-phosphorus seed when the plants were grown in cultures deficient in phosphorus. In cultures not deficient in phosphorus, there were no such differences between the performance of the seed.In field experiments carried out on a fertile soil, low-phosphorus seeds gave 20 to 25 per cent lower yields of both haulms and peas than were obtained from sowings of high-phosphorus seed.It appeared that the differences in the performance of the seeds weer related to differences in the concentration of phosphorus they contained rather than to any other variation in their composition or size.     

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     

The effect of local application of fertilizer on the content of cytokinins in the xylem sap of maize

Cytokinin content in xylem sap was higher in plants grown under local supply of fertilizers as compared to those grown under homogenous distribution of nutrients in soil. The separate assay of cytokinins in xylem exudate from split root system showed that roots, which were in contact with fertilizer mainly contributed to cytokinins transported from roots to shoots. This revised version was published online in July 2006 with corrections to the Cover Date.     

Phosphate Availability Alters Lateral Root Anatomy and Root Architecture of Fraxinus mandshurica Rupr. Seedlings

Plants have evolved some mechanisms to maximize the efficiency of phosphorus acquisition.Changes in root architecture are one such mechanism. When Fraxinus mandshurica Rupr. seedlings were grown under conditions of low phosphorus availability, the length of cells in the meristem zone of the lateral roots was longer, but the length of cells in the elongation and mature zones of the lateral roots was shorter,compared with seedlings grown under conditions of high phosphorus availability. The elongation rates of primary roots increased as phosphorus availability increased, but the elongation rates of the branched zones of the primary roots decreased. The number of lateral root primordia and the length of the lateral roots decreased as phosphorus availability increased. The topological index (altitude slope) decreased as phosphorus availability increased, suggesting that root architecture tended to be herringbone-like when seedlings were grown under conditions of low phosphate availability. Herringbone-like root systems exploit nutrients more efficiently, but they have higher construction costs than root systems with a branching pattern.     

Phosphorus sources and availability modify growth and distribution of root clusters and nodules of native Australian legumes

A variety of native Western Australian legumes produced root clusters in sand culture confirming field and published observations. In general, these legumes grew equally well when supplied with organic or inorganic sources of phosphorus. The nitrogen content of shoots and roots varied little among treatments for all species, however, phosphorus content was always greater in plants supplied with inositol‐P. The plasticity of root growth in response to localized placement of organic and inorganic sources of phosphorus was demonstrated using a simple ‘split root’ technique. Total root dry weight was, on average, more than doubled in P‐amended sand when compared with non‐amended sand. Root clusters tended to be produced in areas of relatively high phosphorus concentration and nodules in areas of low phosphorus concentration. Levels of phosphorus in lateral roots grown in P‐amended sand were significantly different from lateral roots grown in the corresponding non‐amended sand. Growth increases averaging 70% for white sand to over 100% for yellow sand indicated a large degree of ‘plasticity’ in roots under conditions of heterogeneous supply of phosphorus. Spatially exclusive development of organs for the acquisition of nutrients is discussed in relation to requirements for carbon in organ production and maintenance.     

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.     

Adaptive Changes and Genotypic Variation for Root Architecture of Common Bean in Response to Phosphorus Deficiency

Root architectural responses to phosphorus (p) availability may be an important trait for P acquisition efficiency. In the present study, The authors examined the effects of P availability on root architectural responses of different common bean genotypes. Five common bean (Phaseolus vulgaris L.) genotypes representing different origins and ecotypic races were compared both in a specially designed paper pouch system and a stratified P buffer sand culture system with computer image analysis. The results showed that root architecture was regulated by P availability. P deficiency led to form a shallower root system, as indicated by increased relative distribution of basal root length in the upper layers and decreased the growth angle of basal roots. There was significant genetic variation in root architecture in response to P deficiency both in the paper pouch system and the stratified sand culture system. Under low P conditions some genotypes were more gravitropically sensitive to low P availability, resulting in producing a shallower root system and enhanced root exploration into the surface soil, where soil available P is more concentrated. G19833 and DOR364, which were most contrasting in P efficiency, were also very different in root architectural response to P availability. The results from this study suggest that P availability regulates root architecture and P deficiency leads to shallower root architecture in beans. The genetic potential of root architecture provides the possibility of selecting this trait for improving P acquisition efficiency in common bean.     

Phosphate Availability Alters Lateral Root Anatomy and Root Architecture of Fraxinus mandshurica Rupr. Seedlings

Plants have evolved some mechanisms to maximize the efficiency of phosphorus acquisition. Changes in root architecture are one such mechanism. When Fraxinus mandshurica Rupr. seedlings were grown under conditions of low phosphorus availability, the length of cells in the meristem zone of the lateral roots was longer, but the length of cells in the elongation and mature zones of the lateral roots was shorter,compared with seedlings grown under conditions of high phosphorus availability. The elongation rates of primary roots increased as phosphorus availability increased, but the elongation rates of the branched zones of the primary roots decreased. The number of lateral root primordia and the length of the lateral roots decreased as phosphorus availability increased. The topological index (altitude slope) decreased as phosphorus availability increased, suggesting that root architecture tended to be herringbone-like when seedlings were grown under conditions of low phosphate availability. Herringbone-like root systems exploit nutrients more efficiently, but they have higher construction costs than root systems with a branching pattem.     

Long-term warming effects on root morphology, root mass distribution, and microbial activity in two dry tundra plant communities in northern Sweden

Effects of warming on root morphology, root mass distribution and microbial activity were studied in organic and mineral soil layers in two alpine ecosystems over>10 yr, using open-top chambers, in Swedish Lapland. Root mass was estimated using soil cores. Washed roots were scanned and sorted into four diameter classes, for which variables including root mass (g dry matter (g DM) m(-2)), root length density (RLD; cm cm(-3) soil), specific root length (SRL; m g DM(-1)), specific root area (SRA; m2 kg DM(-1)), and number of root tips m(-2) were determined. Nitrification (NEA) and denitrification enzyme activity (DEA) in the top 10 cm of soil were measured. Soil warming shifted the rooting zone towards the upper soil organic layer in both plant communities. In the dry heath, warming increased SRL and SRA of the finest roots in both soil layers, whereas the dry meadow was unaffected. Neither NEA nor DEA exhibited differences attributable to warming. Tundra plants may respond to climate change by altering their root morphology and mass while microbial activity may be unaffected. This suggests that carbon may be incorporated in tundra soils partly as a result of increases in the mass of the finer roots if temperatures rise.     

低磷胁迫下磷高效基因型大麦的根系形态特征

在根袋土培盆栽条件下,以磷高效基因型DH110+、DH147和低效基因型DH49大麦为试验材料,利用根系分析系统分析不同施磷(P₂O₅)水平(极低磷25 mg·kg^-1、低磷50 mg·kg^-1和正常磷75 mg·kg^-1)下,磷高效基因型大麦的根系形态特征及其与植株磷素吸收的关系.结果表明： 低磷胁迫显著降低大麦生物量和磷吸收量,其中磷高效基因型的生物量和磷吸收量在各施磷水平下分别为低效基因型的1.24～1.70和1.18～1.83倍;大麦的总根长、总根表面积、平均根系直径、不定根长及其根表面积、侧根长及其根表面积均随施磷水平的降低而显著降低,其中磷高效基因型大麦在各施磷水平下的总根长、总根表面积、比根长、侧根长及根表面积分别为低效基因型的1.46～2.06、1.12～1.51、1.35～1.72、1.69～2.42和1.40～1.78倍,而平均根系直径为低效基因型的70.6%～90.2%;主成分分析表明,平均根系直径、比根表面积和比根长受基因型差异的影响较为明显,是区分两类磷效率基因型大麦根系形态差异的主要指标;偏最小二乘回归分析表明,各施磷水平下,总根长、总根表面积对大麦植株磷素吸收贡献均较大,随施磷水平降低,不定根长、不定根表面积对大麦植株磷素吸收的贡献明显降低,而平均根系直径、比根长、侧根长及其根表面积的贡献明显增加.磷高效基因型大麦可通过维持侧根的生长、根细度和比根长的增加来适应低磷胁迫.     

Morphological and physiological responses of rice (Oryza sativa) to limited phosphorus supply in aerated and stagnant solution culture

BACKGROUND AND AIMS: Rain-fed lowland rice commonly encounters stresses from fluctuating water regimes and nutrient deficiency. Roots have to acquire both oxygen and nutrients under adverse conditions while also acclimating to changes in soil-water regime. This study assessed responses of rice roots to low phosphorus supply in aerated and stagnant nutrient solution. METHODS: Rice (Oryza sativa 'Amaroo') was grown in aerated solution with high P (200 micro m) for 14 d, then transferred to high or low (1.6 micro m) P supply in aerated or stagnant solution for up to 8 d. KEY RESULTS: After only 1 d in stagnant conditions, root radial oxygen loss (ROL) had decreased by 90 % in subapical zones, whereas near the tip ROL was maintained. After 4 d in stagnant conditions, maximum root length was 11 % less, and after 8 d, shoot growth was 25 % less, compared with plants in aerated solution. The plants in stagnant solution had up to 19 % more adventitious roots, 24 % greater root porosity and 26 % higher root/shoot ratio. Rice in low P supply had fewer tillers in both stagnant and aerated conditions. After 1-2 d in stagnant solution, relative P uptake declined, especially at low P supply. Aerated roots at low P supply maintained relative P uptake for 4 d, after which uptake decreased to the same levels as in stagnant solution. CONCLUSIONS: Roots responded rapidly to oxygen deficiency with decreased ROL in subapical zones within 1-2 d, indicating induction of a barrier to ROL, and these changes in ROL occurred at least 2 d before any changes in root morphology, porosity or anatomy were evident. Relative P uptake also decreased under oxygen deficiency, showing that a sudden decline in root-zone oxygen adversely affects P nutrition of rice.     

Earthworm effect on root morphology in a split root system

Plants respond to their environment through adaptations such as root proliferation in nutrient-rich patches. Through their burrows and casts production in soil, earthworms create heterogeneity which could lead to local root adaptations or systemic effects. To investigate the effect of earthworms on root system morphology and determine whether earthworm effect is local or systemic, we set up two independent split root experiments with rice or barley, (i) without earthworm (CC), (ii) with earthworms in both compartments (EE), and (iii) with earthworms in one single compartment (CE). Earthworms had an effect on belowground plant biomass. The relative length of thick roots decreased with an increasing abundance of earthworms. Some root diameter classes responded to earthworm number in a linear or curvilinear way, making simple conclusions difficult. We found no difference in root biomass or morphology between the two compartments of the split root system in the CE treatment, but a positive effect of earthworm biomass on root biomass, volume, surface area, and length at the whole plant level. Results supported a systemic effect dependent on earthworm abundance. Modification of nutrient mineralization, soil physical structure, and/or the concentration of signal molecules could all be responsible for this systemic effect.     

Importance of internal hydraulic redistribution for prolonging the lifespan of roots in dry soil

Redistribution of water within plants could mitigate drought stress of roots in zones of low soil moisture. Plant internal redistribution of water from regions of high soil moisture to roots in dry soil occurs during periods of low evaporative demand. Using minirhizotrons, we observed similar lifespans of roots in wet and dry soil for the grapevine 'Merlot' (Vitis vinifera) on the rootstock 101-14 Millardet de Gramanet (Vitis riparia x Vitis rupestris) in a Napa County, California vineyard. We hypothesized that hydraulic redistribution would prevent an appreciable reduction in root water potential and would contribute to prolonged root survivorship in dry soil zones. In a greenhouse study that tested this hypothesis, grapevine root systems were divided using split pots and were grown for 6 months. With thermocouple psychrometers, we measured water potentials of roots of the same plant in both wet and dry soil under three treatments: control (C), 24 h light + supplemental water (LW) and 24 h light only (L). Similar to the field results, roots in the dry side of split pots had similar survivorship as roots in the wet side of the split pots (P = 0.136) in the C treatment. In contrast, reduced root survivorship was directly associated with plants in which hydraulic redistribution was experimentally reduced by 24 h light. Dry-side roots of plants in the LW treatment lived half as long as the roots in the wet soil despite being provided with supplemental water (P < 0.0004). Additionally, pre-dawn water potentials of roots in dry soil under 24 h of illumination (L and LW) exhibited values nearly twice as negative as those of C plants (P = 0.034). Estimates of root membrane integrity using electrolyte leakage were consistent with patterns of root survivorship. Plants in which nocturnal hydraulic redistribution was reduced exhibited more than twice the amount of electrolyte leakage in dry roots compared to those in wet soil of the same plant. Our study demonstrates that besides a number of ecological advantages to protecting tissues against desiccation, internal hydraulic redistribution of water is a mechanism consistent with extended root survivorship in dry soils.     

Growth and morphology of eucalypt seedling-roots, in relation to soil strength arising from compaction

Information on the response of root growth and morphology to soil strength is useful for testing suitability of existing and new tillage methods and/or for selecting plants suitable for a specific site with or without tillage. Although there is extensive published information on the root growth-soil strength relationships for annual agricultural plants, such information is scarce for woody, perennial tree species. The purpose of this study is to examine growth and morphology of the root systems of 17-day-old eucalypt seedlings with respect to variation in soil strength. Soil strength in this study was varied by compaction of a well-aggregated clay soil to bulk densities of 0.7–1.0 Mg m^-3 whilst maintaining adequate water availability and aeration for plant growth. Lengths and tip-diameters of primary and lateral roots were measured on the excavated root systems of seedlings.With increase in bulk density and also soil strength (expressed as penetrometer resistance), total length of primary and lateral roots decreased. There were 71 and 31% reduction in the lengths of primary and lateral roots respectively with an increase in penetrometer resistance from 0.4 to 4.2 MPa. This indicated primary roots to be more sensitive to high soil strength than the lateral roots. Average length of lateral roots and diameters of both primary and lateral root tips increased with an increase in soil strength as well. There was greater abundance of lateral roots (no. of lateral roots per unit length of primary root) and root hairs with increased soil strength. The observed root behaviour to variable soil strength is discussed in the context of compensatory growth of roots and overall growth of plants.     

Optimizing reproductive phenology in a two-resource world: a dynamic allocation model of plant growth predicts later reproduction in phosphorus-limited plants

Background and Aims

Timing of reproduction is a key life-history trait that is regulated by resource availability. Delayed reproduction in soils with low phosphorus availability is common among annuals, in contrast to the accelerated reproduction typical of other low-nutrient environments. It is hypothesized that this anomalous response arises from the high marginal value of additional allocation to root growth caused by the low mobility of phosphorus in soils.

Methods

To better understand the benefits and costs of such delayed reproduction, a two-resource dynamic allocation model of plant growth and reproduction is presented. The model incorporates growth, respiration, and carbon and phosphorus acquisition of both root and shoot tissue, and considers the reallocation of resources from senescent leaves. The model is parameterized with data from Arabidopsis and the optimal reproductive phenology is explored in a range of environments.

Key Results

The model predicts delayed reproduction in low-phosphorus environments. Reproductive timing in low-phosphorus environments is quite sensitive to phosphorus mobility, but is less sensitive to the temporal distribution of mortality risks. In low-phosphorus environments, the relative metabolic cost of roots was greater, and reproductive allocation reduced, compared with high-phosphorus conditions. The model suggests that delayed reproduction in response to low phosphorus availability may be reduced in plants adapted to environments where phosphorus mobility is greater.

Conclusions

Delayed reproduction in low-phosphorus soils can be a beneficial response allowing for increased acquisition and utilization of phosphorus. This finding has implications both for efforts to breed crops for low-phosphorus soils, and for efforts to understand how climate change may impact plant growth and productivity in low-phosphorus environments.