Root responses to nitrogen pulse frequency under different nitrogen amounts

Responses of morphology and biomass allocation of roots to frequency of nitrogen (N) pulse potentially influence the fitness of plants, but such responses may be determined by root size. We grew 12 plant species of three functional groups (grasses, forbs, and legumes) under two N pulse frequencies (high vs. low supply frequency) and two N amounts (high vs. low supply amount). Compared to low-amount N supply, high-amount N supply stimulated biomass accumulation and root growth by either increasing the thickness and length of roots or decreasing the root mass fraction. Compared to low-frequency N supply, high-frequency N supply improved biomass accumulation and root growth in forbs or grasses, but not in legumes. Furthermore, the magnitude of the response to N frequency was significantly negatively correlated with root size at the species scale, but this was only true when the N amount was high. We conclude that root responses to N frequency are related to plant functional types, and non-legume species is more sensitive to N frequency than legume species. Our results also suggest that root size is a determinant of root responses to N frequency when N supply amount is high.     

不同氮磷添加浓度对豆科3种树木幼苗生长及生物量分配的影响

选取海南尖峰岭热带山地雨林中不同功能类群的豆科树木幼苗（代表低氮需求的长脐红豆Ormosia balansae Drake、中氮需求的荔枝叶红豆Ormosia semicastrata Hance f. litchifolia How和高氮需求的猴耳环Archidendron clypearia（Jack）I.C.Nielsen）为对象,设置5个浓度梯度的氮（N）添加和2个浓度梯度的磷（P）添加养分控制实验,研究苗木的生长表现。结果显示：（1）氮添加条件下,3个树种幼苗的苗高、总叶面积、根长、根表面积、生物量5个指标对中高浓度氮添加的敏感性大小均为长脐红豆 > 猴耳环 > 荔枝叶红豆;其中,叶总面积对氮肥浓度变化的响应最敏感,长脐红豆的根长、根表面积以及猴耳环根长的响应敏感性次之。（2）长脐红豆和猴耳环幼苗的根冠比受氮肥添加浓度的影响不显著;荔枝叶红豆幼苗的根冠比则随氮肥添加浓度的升高而增大,这种适应策略反映出荔枝叶红豆幼苗对添加中高浓度氮肥有较强的耐受能力。（3）磷添加条件下,长脐红豆和猴耳环幼苗的生长速率为低磷>高磷,表明这2个树种在幼苗阶段为低氮、低磷需求;荔枝叶红豆在低氮处理下的生长速率为高磷>低磷,表明该树种幼苗阶段为低氮、高磷需求。     

Understanding the Influences of Spatial Patterns on N Availability Within the Brazilian Amazon Forest

Nitrogen variations at different spatial scales and integrated across functional groups were addressed for lowland tropical forests in the Brazilian Amazon as follows: (1) how does N availability vary across the region over different spatial scales (regional × landscape scale); (2) how are these variations in N availability integrated across plant functional groups (legume × non-legume trees). Leaf N, P, and Ca concentrations as well the leaf N isotope ratios (δ¹⁵N) from a large set of legume and non-legume tree species were measured. Legumes had higher foliar N/Ca ratios than non-legumes, consistent with the high energetic costs in plant growth associated with higher foliar P/Ca ratios found in legumes than in non-legumes. At the regional level, foliar δ¹⁵N decreased with increasing rainfall. At the landscape level, N availability was higher in the forests on clayey soils on the plateau than in forests on sandier soils. The isotope as well as the non-isotope data relationships here documented, explain to a large extent the variation in δ¹⁵N signatures across gradients of rainfall and soil. Although at the regional level, the precipitation regime is a major determinant of differences in N availability, at the landscape level, under the same precipitation regime, soil type seems to be a major factor influencing the availability of N in the Brazilian Amazon forest.     

Coupled soil-availability and tree-limitation nutritional shifts induced by N deposition: insights from N to P relationships in Abies pinsapo forests

Background and aims

Interacting effects of atmospheric N deposition on the degree to which tree demand for other nutrients is met by soil supply has seldom been explored in Mediterranean-type ecosystems. We hypothesized that patterns for the relative availability of N and P in soils will be matched by variations in process rates related to soil organic P cycling and by shifts from N to P limitation of tree growth.

Methods

We examined N/P relationships in Mediterranean-fir (Abies pinsapo) forests from two nearby regions differing in N deposition levels.

Results

N pools and transformation rates and the contribution of organic fractions to the labile P pool in soils showed increasing trends toward the pollution source. Phosphomonoesterase activity (PME) in bulk soils, root PME per unit biomass (but not per unit soil volume) and biomass accumulation in P-fertilized root-in-growth cores incubated in situ were also the highest at the sites receiving elevated N deposition, indicating P limitation. In contrast, forest stands in the region farther from the pollutant source were N-limited (preferential root growth in N-rich soil microsites) and showed lower PME activities and higher total fine root biomass.

Conclusions

In the forests under elevated N deposition, higher values for an overall indicator of soil N status matched with indications of an accelerated soil organic P subcycle and P-limitation of tree growth.     

Interspecific differences in the responses of root phosphatase activities and morphology to nitrogen and phosphorus fertilization in Bornean tropical rain forests

Soil organic phosphorus (P) compounds can be the main P source for plants in P‐limited tropical rainforests. Phosphorus occurs in diverse chemical forms, including monoester P, diester P, and phytate, which require enzymatic hydrolysis by phosphatase into inorganic P before assimilation by plants. The interactions between plant interspecific differences in organic P acquisition strategies via phosphatase activities with root morphological traits would lead to P resource partitioning, but they have not been rigorously evaluated. We measured the activities of three classes of phosphatases (phosphomonoesterase, PME; phosphodiesterase, PDE; and phytase, PhT), specific root length (SRL), root diameter, and root tissue density in mature tree species with different mycorrhizal associations (ectomycorrhizal [ECM] or arbuscular mycorrhizal [AM]) and different successional status (climax or pioneer species) in Sabah, Malaysia. We studied nitrogen (N)‐ and P‐fertilized plots to evaluate the acquisition strategies for organic P under P‐limited conditions 7 years after fertilization was initiated. P fertilization reduced the PME activity in all studied species and reduced PhT and PDE activities more in climax species than in the two pioneer species, irrespective of the mycorrhizal type. PDE activity increased in some climax species after N fertilization, suggesting that these species allocate excess N to the synthesis of PDE. Moreover, PME and PhT activities, but not PDE activity, correlated positively with SRL. We suggest that climax species tend to be more strongly dependent on recalcitrant organic P (i.e., phytate and/or diester P) than pioneer species, regardless of the mycorrhizal type. We also suggest that trees in which root PME or PhT activity is enhanced can increase their SRL to acquire P efficiently. Resource partitioning of soil organic P would occur among species through differences in their phosphatase activities, which plays potentially ecologically important role in reducing the competition among coexisting tree species in lowland tropical rainforests.     

How do nitrogen and phosphorus deficiencies affect strigolactone production and exudation?

Plants exude strigolactones (SLs) to attract symbiotic arbuscular mycorrhizal fungi in the rhizosphere. Previous studies have demonstrated that phosphorus (P) deficiency, but not nitrogen (N) deficiency, significantly promotes SL exudation in red clover, while in sorghum not only P deficiency but also N deficiency enhances SL exudation. There are differences between plant species in SL exudation under P- and N-deficient conditions, which may possibly be related to differences between legumes and non-legumes. To investigate this possibility in detail, the effects of N and P deficiencies on SL exudation were examined in Fabaceae (alfalfa and Chinese milk vetch), Asteraceae (marigold and lettuce), Solanaceae (tomato), and Poaceae (wheat) plants. In alfalfa as expected, and unexpectedly in tomato, only P deficiency promoted SL exudation. In contrast, in Chinese milk vetch, a leguminous plant, and in the other non-leguminous plants examined, N deficiency as well as P deficiency enhanced SL exudation. Distinct reductions in shoot P levels were observed in plants grown under N deficiency, except for tomato, in which shoot P level was increased by N starvation, suggesting that the P status of the shoot regulates SL exudation. There seems to be a correlation between shoot P levels and SL exudation across the species/families investigated.     

Rhizobial Association with Non-Legumes: Mechanisms and Applications

It has been known for more than a century that rhizobia can promote the growth of legumes through the formation of nitrogen-fixing nodules, but the interaction of rhizobia with non-legumes has been neglected as an experimental system. During the last couple of decades, work on rhizobial interaction with non-legumes has been done progressively and it has been demonstrated that rhizobia can associate with roots of non-legumes also, without forming true nodules, and can promote their growth by using one or more of the direct or indirect mechanisms of actions. Phytohormone production, secretion of other chemicals like lipo-chito-oligosaccharides (LCOs) and lumichrome, solubilization of precipitated phosphorus and mineralization of organic P, improvement in uptake of plant nutrients by altering root morphology, production of siderophores to meet the iron requirements of the plant under iron-stressed conditions and lowering of ethylene level through ACC deaminase enzyme, are some examples of the rhizobial mechanisms with direct positive effects on non-leguminous plant growth. Indirectly, rhizobia improve the growth of non-legumes through biocontrol of pathogens via antibiosis, parasitism or competition with pathogens for nutrients and space, by inducing systemic resistance in the host plant and through increasing root adhering soil by releasing exopolysaccarides which regulate the water movement and facilitate the root growth. However, no influence or even inhibitory effects of rhizobial inoculation on non-legumes has also been demonstrated in some cases. Plant growth promoting mechanisms of rhizobia and its practical application in non-legumes are the major focus of this review.     

Consumption rates and food preferences of slugs in a calcareous grassland under current and future CO<Subscript>2</Subscript> conditions

This study explored consumption of a generalist herbivore feeding on leaf tissue of various plant species of a calcareous grassland, and tested whether consumption levels and preferences changed when plants were exposed to 5 years of in situ CO₂ enrichment. The first part of this experiment tested whether the consumption patterns of slugs (Deroceras reticulatum) observed in single-species feeding tests were altered when slugs were given a choice of food sources. Overall consumption increased 270% when slugs were given a choice, and they preferred having a choice of food sources more than they preferred having any one food source. Surprisingly, slugs consumed fewer legumes and grasses and more non-leguminous forbs when given a choice. In the second part of this experiment, feeding behaviors of slugs in response to elevated CO₂ were investigated by feeding them leaves of two legumes, one grass, and a non-leguminous forb (Trifolium medium, Lotus corniculatus, Bromus erectus, and Sanguisorba minor, respectively) in two or four species combinations. In the leguminous species mix, the non-leguminous species mix, and the combined mix (legumes and non-legumes), neither overall consumption by herbivores nor species preference was significantly altered by long-term CO₂ enrichment. In the combined species mix, slugs preferred legumes to non-legumes (P=0.012) and exhibited a weak functional group preference shift from non-legumes to legumes (P=0.089) in response to CO₂ enrichment. This is the first time such a shift has been observed, and provides evidence that there may be multiple herbivore responses to rising atmospheric CO₂ concentrations. Numerous single-species feeding tests using insects have shown that consumption by herbivores may increase when herbivores are fed plants grown in enriched CO₂ atmospheres. This study clearly demonstrates the limited applicability of non-choice feeding trials to generalist herbivores in species-rich communities. Received: 19 August 1999 / Accepted: 20 April 2000     

Differences in rhizosphere carbon-partitioning among plant species of different families

Interspecific variations in carbon (C) allocation and partitioning in the rhizosphere were investigated on 12 Mediterranean species belonging to different family groups (grasses, legumes, non-legume forbs) and having different life cycles. Plants grown individually in artificial soil, in a greenhouse and inoculated with rhizosphere microflora were labelled with ¹⁴CO₂ for 3 h at the vegetative stage. Rhizosphere respiration was measured during 6 days after which labelled C partitioning between shoots, roots, soil, root washing solution and respiration was estimated. The percentage of assimilated ¹⁴C allocated below ground differed significantly between species (41 – 76%) but no significant difference was found between grasses, legumes and non-legume forbs. When expressed as percentage of below-ground ¹⁴C, rhizosphere respiration was significantly smaller for non-legume forbs (42%) than for grasses (46%) and legumes (51%). Consequently more ¹⁴C was incorporated into root biomass in the former. Half-life of ¹⁴CO₂ evolution through respiration ranged from 23 h in legumes to 27 h for non-legume forbs and 37 h for grasses. This suggested differences in microbial activities due to quantities and quality of root exuded C. Rhizosphere respiration was positively correlated with the amount of ¹⁴C in the solution used to wash the roots on one hand, and root N concentration on the other hand. This led to a functional hierarchy between plant family groups of the overall rhizosphere activity. It went from non-legume forbs being the less active (except Crepis sancta)in terms of respiration and exudation, to grasses and then legumes, the most active but also the richest in nitrogen.     

Seedling growth of five tropical dry forest tree species in relation to light and nitrogen gradients

Aims Increasing anthropogenic nitrogen (N) deposition has been claimed to induce changes in species composition and community dynamics. A greenhouse experiment was conducted to examine the effect of increased N availability on growth and functional attributes of seedlings of five tree species with different life history characteristics under varying irradiances. The following questions have been addressed: (i) how do the pioneer and non-pioneer species respond in absolute growth and relative growth rate (RGR) to the interaction of light and nitrogen? (ii) how does the interaction between irradiance and nitrogen availability modulate growth attributes (i.e. functional attributes)? (iii) is there any variation in growth responses between leguminous and non-leguminous species along the light and nitrogen gradients?Methods Seedlings of five tree species (Acacia catechu, Bridelia retusa, Dalbergia sissoo, Lagerstroemia parviflora and Terminalia arjuna) were subjected to twelve combinations of irradiance and N levels. Various growth traits, including height (HT), basal area (BA), whole plant dry biomass (M D), leaf mass per unit area (LMA), leaf area ratio (LAR), net assimilation rate (NAR), RGR, biomass fractions, root-to-shoot ratio (R:S) and leaf nitrogen content, were studied to analyse intra- and inter-specific responses to interacting light and N gradients.Important findings Significant interactions for irradiance and N availability for majority of growth attributes indicates that growth and biomass allocation of seedlings were more responsive to N availability under high irradiance. However, species responded differentially to N addition and they did not follow successional status. Slow growers (B. retusa, a shade-tolerant species and L. parviflora, a light demander) exhibited greater response to N enrichment than the fast growers (A. catechu, D. sissoo and T. arjuna). However, N-mediated increment in growth traits was greater in non-legumes (B. retusa, L. parviflora and T. arjuna) compared with that of legumes (A. catechu and D. sissoo). Allocation of biomass to root was strongly suppressed at the highest N supply across species; however, at high irradiance and high N availability, a greater suppression in R:S ratio was observed for B. retusa. NAR was a stronger determinant of RGR relative to LAR, suggesting its prominent role in increased RGR along increasing irradiances. Overall, a higher growth response of slow-growing species to elevated N levels, particularly the non-pioneers (B. retusa and L. parviflora) suggests that future N deposition may lead to perturbations in competition hierarchies and species composition, ultimately affecting community dynamics in nutrient-poor tropical dry forests.     

Growth and nodulation of tropical food legumes in dilute solution culture

Twenty-two tropical food legumes were grown in dilute nutrient solution with or without rhizobium inoculation and supplied with either low or adequate amounts of inorganic N. Growth of legumes supplied with adequate inorganic N was generally satisfactory. However, solution phosphorus (P) concentration (15μM) was excessive for black gram, while the initial solution manganese concentration (1.8μM) was excessive for green gram. Growth responses to inoculation with rhizobium at low inorganic N supply were obtained in only 9 of the 22 legumes studied, and shoot dry matter yields were ≤ 51% of those obtained with adequate N supply. Poor growth by inoculated plants with a low N supply was attributed to failure of the inoculated strain of Bradyrhizobium to infect roots (lima bean and Mexican yam bean), to low nodule numbers (green gram, black gram and navy bean), or to excessive uptake of P (black gram, adzuki bean, pigeonpea, winged bean and cowpea cv. Vita 4) and/or manganese (green gram and black gram). High solution temperatures may have limited N fixation by some of the legumes, particularly chickpea.     

Increased N affects P uptake of eight grassland species: the role of root surface phosphatase activity

Increased N deposition may change species composition in grassland communities by shifting them to P limitation. Interspecific differences in P uptake traits might be a crucial yet poorly understood factor in determining the N effects. To test the effects of increased N supply (relative to P), we conducted two greenhouse fertilization experiments with eight species from two functional groups (grasses, herbs), including those common in P and N limited grasslands. We investigated plant growth and P uptake from two P sources, orthophosphate and not‐readily available P (bound‐P), under different N supply levels. Furthermore, to test if the N effects on P uptake was due to N availability alone or altered N:P ratio, we examined several uptake traits (root‐surface phosphatase activity, specific root length (SRL), root mass ratio (RMR)) under varying N:P supply ratios. Only a few species (M. caerulea, A. capillaris, S. pratensis) could take up a similar amount of P from bound‐P to that from orthophosphate. These species had neither higher SRL, RMR, phosphatase activity per unit root (Pase_root), nor higher total phosphatase activity (Pase_tot: Pase_root times root mass), but higher relative phosphatase activity (Pase_rel: Pase_tot divided by biomass) than other species. The species common from P‐limited grasslands had high Pase_rel. P uptake from bound‐P was positively correlated with Pase_tot for grasses. High N supply stimulated phosphatase activity but decreased RMR and SRL, resulting in no increase in P uptake from bound‐P. Pase_root was influenced by N:P supply ratio, rather than by only N or P level, whereas SRL and RMR was not dominantly influenced by N:P ratio. We conclude that increased N stimulates phosphatase activity via N:P stoichiometry effects, which potentially increases plant P uptake in a species‐specific way. N deposition, therefore, may alter plant community structure not only by enhancing productivity, but also by favouring species with traits that enable them to persist better under P limited conditions.     

Nitrogen:phosphorous supply ratio and allometry in five alpine plant species

In terrestrial ecosystems, atmospheric nitrogen (N) deposition has greatly increased N availability relative to other elements, particularly phosphorus (P). Alterations in the availability of N relative to P can affect plant growth rate and functional traits, as well as resource allocation to above‐ versus belowground biomass (M_A and M_B). Biomass allocation among individual plants is broadly size‐dependent, and this can often be described as an allometric relationship between M_A and M_B, as represented by the equation , or log M_A = logα + βlog M_B. Here, we investigated whether the scaling exponent or regression slope may be affected by the N:P supply ratio. We hypothesized that the regression slope between M_A and M_B should be steeper under a high N:P supply ratio due to P limitation, and shallower under a low N:P supply ratio due to N limitation. To test these hypotheses, we experimentally altered the levels of N, P, and the N:P supply ratio (from 1.7:1 to 135:1) provided to five alpine species representing two functional groups (grasses and composite forbs) under greenhouse conditions; we then measured the effects of these treatments on plant morphology and tissue content (SLA, leaf area, and leaf and root N/P concentrations) and on the scaling relationship between M_A and M_B. Unbalanced N:P supply ratios generally negatively affected plant biomass, leaf area, and tissue nutrient concentration in both grasses and composite forbs. High N:P ratios increased tissue N:P ratios in both functional groups, but more in the two composite forbs than in the grasses. The positive regression slopes between log M_A and log M_B exhibited by plants raised under a N:P supply ratio of 135:1 were significantly steeper than those observed under the N:P ratio of 1.7:1 and 15:1. Synthesis: Plant biomass allocation is highly plastic in response to variation in the N:P supply ratio. Studies of resource allocation of individual plants should focus on the effects of nutrient ratios as well as the availability of individual elements. The two forb species were more sensitive than grasses to unbalanced N:P supplies. To evaluate the adaptive significance of this plasticity, the effects of unbalanced N:P supply ratio on individual lifetime fitness must be measured.     

Functional groups' performances as influenced by nitrogen,phosphorus and nodule inhibition of legumes

Aims Nitrogen (N) and phosphorus (P) constitute essential elements for plant growth and their availability influence species diversity in herbaceous plant communities. Legumes exhibit relatively high abundance in N-limited soils. Moreover, the legumes' N:P ratios are much higher than those of the other plant species grown in the same site, probably because they are able to fix atmospheric N 2. The objective of this study was to determine how the relative proportion in N and P availability and the restriction of legumes to fix atmospheric N 2 affect: (i) the primary productivity of plant species, (ii) species composition and (iii) N and P concentrations of species.Methods In an outdoor experiment, mixtures containing grasses, legumes and non-legume forbs were established in 32 containers under four soil treatments (control, N addition, P addition and disinfected soil), in a completely randomized design with eight replicates. Plant growth was examined when N and P were limited in the control soil:sand mixture, in a pot experiment sown with Plantago lanceolata .Important findings The pot experiment indicated that both N and P were limiting for the growth of P. lanceolata. Soil treatments affected primary productivity and species composition. Legumes had a relatively high abundance in the control and their growth was favoured, especially that of Medicago sativa, by P addition. Grasses' growth was increased by the addition of N. Inhibition of rhizobia resulted in poor growth of legumes and concomitant higher growth of grasses, in comparison to the control. The N:P ratios of non-legume species differed between treatments and were always higher in the legume species, even in the disinfected soil. The latter provides evidence that the high N concentrations found in legumes are a physiological characteristic of this specific group of plants.     

Metallophytes—a view from the rhizosphere

Abundance of Fabaceae declines in representation through post-fire-succession in fynbos vegetation of the Cape Floristic Region (CFR). This reduction in legume occurrence coincides with a known decline in post-fire soil P availability. It was hypothesized that the disappearance of legume species during post-fire succession is due to an inability to acquire P effectively from sparingly soluble sources. P-acquisition strategies and response to P supply were compared between legume (Aspalathus, Cyclopia, Indigofera, Podalyria) and non-legume (Elegia, Leucadendron, Protea) genera when supplied with 1 or 10 mg P kg^?1 dry sand. Each genus consisted of a seeder (non-persistent) and resprouter (persistent) species. Non-legumes showed a greater investment in below-ground biomass, more root clusters, with higher concentrations of carboxylates exuded by cluster roots and carboxylates that were better suited to the mobilization of sparingly soluble P compared to legumes. The growth response to increased P supply was 53% higher in legumes than in non-legumes. The lack of a growth response to an elevated P supply in the non-legumes was attributed to N-limitation. Legume resprouters had a higher investment in cluster-root biomass and a lower capacity to down-regulate P-uptake than the seeders. Therefore the inability to acquire sufficient P from low concentration and sparingly soluble soil P-sources may contribute to the lack of indigenous legume persistence in fynbos vegetation of the CFR.     

Growth rate, root development and nutrient uptake of 55 plant species from the Great Plains Grasslands, USA

Empirical and theoretical studies have highlighted that plantcompetition and species diversity are substantially affected by interactionsamong plant growth and nutrient uptake rates, root lateral spread, rootplasticity, and small scale soil nutrient heterogeneity. This study wasdesignedto (a) experimentally estimate parameters regarding rootscaling patterns, root biomass allocation, growth rates, nutrient productivity,and root nutrient influx rates of 55 plant species common to Great Plainsgrasslands; and (b) determine if grasses and forbs can beclassified into statistically distinct groups based on these characteristics.Wefound that: (1) In all species root lateral spread, rootlength, and root surface area had significant allometric scaling relationshipswith root biomass, but that the relationships were unaffected by Navailability.(2) Reductions in the supply of N increased the root:shootratio in 62% of the species. (3) The frequency distributionandmean values of maximum relative growth rates were very similar for grasses andforbs/shrubs, but mid successional grasses had a higher relative growth ratethan late successional ones. (4) In 78% of the species tested,N productivity was increased by reductions in the N supply.(5)When subjected to a high N supply, the N and P productivity of grasses was, onaverage, higher than that of forbs/shrubs, and the N and P productivity ofC₄ grasses was, on average, higher than that of C₃grasses. No differences were found under a low N supply. (6)Nodifferences on the average maximum N and P influx rates per unit of rootsurfacearea were found between grasses and forbs or between C₃ andC₄ grasses, but both were correlated with maximum relative growthrate. (7) The set of parameters we measured were able toseparate grasses and forbs/shrubs into statistically distinct groups that tendto follow in broad terms the coarse vs. fine scaleforaging strategies hypothesis.     

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.     

Relationship between acid phosphatase activity and P concentration in organs of Cyclopia and Aspalathus species,and a non-legume of the Cape Floristic Region

Aims The role of tissue acid phosphatase (APase) activity of legumes and non-legumes in their P nutrition and adaptation to low-P soils is not well understood. To better understand this, a relationship between APase activity and P concentration in leaves, stems, roots and nodules of legumes,CyclopiaandAspalathusand a non-legume,Leucadendron strictum, all native to the P-poor soils of the Cape fynbos biome, was assessed.     

Linking leaf and root trait syndromes among 39 grassland and savannah species

Here, we tested hypothesized relationships among leaf and fine root traits of grass, forb, legume, and woody plant species of a savannah community. CO2 exchange rates, structural traits, chemistry, and longevity were measured in tissues of 39 species grown in long-term monocultures. Across species, respiration rates of leaves and fine roots exhibited a common regression relationship with tissue nitrogen (N) concentration, although legumes had lower rates at comparable N concentrations. Respiration rates and N concentration declined with increasing longevity of leaves and roots. Species rankings of leaf and fine-root N and longevity were correlated, but not specific leaf area and specific root length. The C3 and C4 grasses had lower N concentrations than forbs and legumes, but higher photosynthesis rates across a similar range of leaf N. Despite contrasting photosynthetic pathways and N2-fixing ability among these species, concordance in above- and below-ground traits was evident in comparable rankings in leaf and root longevity, N and respiration rates, which is evidence of a common leaf and root trait syndrome linking traits to effects on plant and ecosystem processes.     

The effect of nitrogen nutrition on cluster root formation and proton extrusion by Lupinus albus

Nitrogen nutrition can influence cluster root formation in many wild species, but the effect of N form on cluster root formation and root exudation by white lupin is not known. In a solution culture study, we examined the effect of N nutrition (ammonium, nitrate, both or N2 fixation) on cluster root formation and H+ extrusion by white lupin plants under deficient and adequate P supply. The number of cluster roots increased greatly when plants were supplied with I microM P compared with 50 microM P, the increase being 7.8-fold for plants treated with (NH4)2SO4, 3-fold for plants treated with KNO3 and NH4NO3, and 2-4-fold for N2-fixing plants. Under P deficiency. NH4+-N supply resulted in production of a greater number and biomass of cluster roots than other N sources. Dry weight of cluster roots was 30 % higher than that of non-cluster roots in P-deficient plants treated with (NH4)2SO4 and NH4NO3. In plants treated with sufficient P (50 microM), the weight of non-cluster roots was approx. 90 % greater than that of cluster roots. Both total (micromol per plant h(-1)) and specific (micromol g(-1) root d. wt h(-1)) H+ extrusions were greatest from roots of plants supplied with (NH4)2SO4, followed by those supplied with NH4NO3 and N2 fixation, whereas plants receiving KNO3 had negative net H+ extrusion between the third and fifth week of growth (indicating uptake of protons or release of OH- ions). The rate of proton extrusion by NH4+-N-fed plants was similar under P-deficient and P-sufficient conditions. In contrast, proton exudation by N2-fixing plants and KNO3-treated plants was ten-fold greater under P deficiency than under P sufficiency. In comparison with P deficiency, plants treated with 50 microM P had a significantly higher concentration of P in roots, shoots and youngest expanded leaves (YEL). Compared with the N2 fixation and KNO3 treatments, total N concentration was highest in roots, shoots and YEL of plants supplied with (NH4)2SO4 and NH4NO3, regardless of P supply. Under P deficiency, K concentrations in roots decreased at all N supplies, especially in plants treated with (NH4)2SO4 and NH4NO3, which coincided with the greatest H+ extrusion at these P and N supplies. In conclusion, NH4-N nutrition stimulated cluster root formation and H+ extrusion by roots of P-deficient white lupin.