Plant species richness and diversity of the serpentine areas on the Witwatersrand

Soil chemistry can play an important role in determining plant diversity. Serpentine soils are usually toxic to many plant taxa, which limits plant diversity compared to that on adjacent non-serpentine soils. The usually high concentrations of toxic metals in serpentine soils are considered to be the edaphic factors that cause low diversity and high endemism. This paper aimed primarily to determine whether there is a relationship between serpentine soil chemistry and species richness on the Witwatersrand and to compare species richness of the serpentine areas with that of adjacent non-serpentine areas as well as with the species richness of the serpentine areas in the Barberton Greenstone Belt. The alpha- and beta-diversity of the Witwatersrand serpentine and non-serpentine areas was also investigated. A secondary aim of this study was to determine which of the non-serpentine taxa were more common on the serpentine than off the serpentine, which taxa were more common off the serpentine than on the serpentine and which taxa were equally common on and off serpentine soils. There was no significant difference in alpha-diversity between the serpentine and the adjacent non-serpentine areas, but beta-diversity is higher between serpentine plots than between non-serpentine plots. Although soil factors do affect species richness and diversity of plants on the Witwatersrand to a limited extent, the concentrations of soil chemicals in serpentine soils are not sufficiently different from those in non-serpentine soils to significantly influence the species richness and diversity of the serpentine soils. The high, but similar, diversity on serpentine and non-serpentine soils on the Witwatersrand indicates that soil factors do not play a significant role in determining diversity on potentially toxic soils in the area.     

Responses of two populations of an Iranian nickel-hyperaccumulating serpentine plant,Alyssum inflatum Nyar., to substrate Ca/Mg quotient and nickel

We selected two geographically close serpentine and non-serpentine populations of a Ni-hyperaccumulating plant (Alyssum inflatum) to investigate the influence of two common factors of serpentine soils: high Ni concentrations and low Ca/Mg quotients. Soils and plants were sampled from serpentine and non-serpentine substrates, and concentrations of Ca, Mg and Ni were measured. A hydroponic culture was used to compare growth and elemental composition responses of serpentine and non-serpentine plants to different Ca/Mg quotients and Ni concentrations in the nutrient solution. The Ca/Mg quotient for non-serpentine soils was 15 times higher than for serpentine soils, but there was no difference in the Ca/Mg quotient of plants from the two populations. In hydroponic culture, plants from both populations were able to survive at high Ca/Mg quotients. This result suggests that serpentine plants of A. inflatum do not necessarily need a substrate with a low Ca/Mg quotient for survival. Decreases in the Ca/Mg quotient in hydroponics decreased growth. The magnitude of this decrease was significantly greater in non-serpentine plants, suggesting a greater resistance of serpentine plants to low Ca/Mg quotients. Total Ni concentration in serpentine soils was 13 times higher than in non-serpentine soils, but ammonium nitrate-extractable concentrations of Ni in both soil types were similar. Ni concentrations in non-serpentine plants from their natural habitat were significantly lower than in serpentine plants, but there was no significant difference in Ni accumulation by plants of the two populations in hydroponic culture. However, increased concentrations of Ni in the hydroponic medium caused similar decreases in growth of both populations, indicating that Ni tolerance of the two populations was similar.     

Nutrient resorption from senescing leaves in two Stipa species native to central Argentina

Abstract Nutrient resorption from senescing leaves enables plants to conserve and reuse nutrients. As such, it could be expected that plant species adapted to infertile soils have a higher nutrient resorption efficiency (percentage reduction of nutrients between green and senesced leaves) and/or higher nutrient resorption proficiency (absolute reduction of nutrients in senesced leaves) than those adapted to fertile soils. Our objective was to compare nitrogen (N) and phosphorous (P) resorption of two congener grasses that successfully occupy uplands of relatively low fertility (Stipa gynerioides) or lowlands of relatively high fertility (Stipa brachychaeta) in natural grasslands of central Argentina. The two Stipa species did not differ in N and P resorption efficiency, but S. gynerioides had a higher N and P resorption proficiency than S. brachychaeta. As a consequence, leaf‐level N and P use efficiency were higher in the species adapted to low fertility conditions than in the species adapted to high fertility conditions. The higher nutrient resorption proficiency of S. gynerioides was also associated with relatively low leaf‐litter decomposition and nutrient release rates found in a previous study.     

Modest enhancement of nitrogen conservation via retranslocation in response to gradients in N supply and leaf N status

Plant nutrient resorption, a ubiquitous mechanism of nutrient conservation, has often been proposed to be more pronounced in infertile than fertile habitats, and in species common to infertile compared to fertile habitats, because of the presumed advantage when nutrients are scarce. However, previous studies provide weak and inconsistent empirical support for these hypotheses, although few have examined intraspecific variation across well-quantified resource gradients. This study addresses intraspecific patterns of nutrient resorption for eight species across two N availability gradients on similar soils in an N-limited oak savanna ecosystem: a long-term fire frequency gradient with a negatively correlated N fertility gradient and a long-term N fertilization gradient. We hypothesized that both resorption proficiency (the minimum nutrient level retained in a senesced leaf) and efficiency (the proportional change in leaf nutrient concentration) would decrease with increasing soil N availability and plant N status. For the seven non-N fixers, either resorption proficiency or efficiency decreased modestly in treatments with higher N availability. In contrast, the legume Amorpha canescens Pursh had higher N levels in green and senesced leaves, and resorbed N much more weakly than the non-fixers, and did not respond in terms of proficiency or efficiency to soil N availability. Across all species and sites in each N fertility gradient, a scaling analysis showed greater resorption efficiency in plants with lower N concentrations. Our data suggest that species can have modest resorption responses reflective of soil nutrient availability and differences in resorption related to their N economy that represent mechanisms of nutrient conservation in nutrient-limited soils.     

Resorption proficiency and efficiency of leaf nutrients in woody plants in eastern China

Aims (i) To explore variations in nutrient resorption of woody plants and their relationship with nutrient limitation and (ii) to identify the factors that control these variations in forests of eastern China.Methods We measured nitrogen (N) and phosphorus (P) concentrations in both green and senesced leaves of 172 woody species at 10 forest sites across eastern China. We compared the nutrient resorption proficiency (NuRP) and efficiency (NuRE) of N and P in plant leaves for different functional groups; we further investigated the latitudinal and altitudinal variations in NuRP and NuRE and the impacts of climate, soil and plant types on leaf nutrient resorptions.Important findings On average, the leaf N resorption proficiency (NRP) and P resorption proficiency (PRP) of woody plants in eastern China were 11.1mg g ? 1 and 0.65 mg g ? 1, respectively; and the corresponding N resorption efficiency (NRE) and P resorption efficiency (PRE) were 49.1% and 51.0%, respectively. Angiosperms have higher NRP (are less proficient) values and lower NRE and PRE values than gymnosperms, but there are no significant differences in NRP, PRP and PRE values between species with different leaf habits (evergreen vs. deciduous angiosperms). Trees have higher NRE and PRE than shrubs. Significant geographical patterns of plant nutrient resorption exist in forests of eastern China. In general, NRP and PRE decrease and PRP and NRE increase with increasing latitude/altitude for all woody species and for the different plant groups. Plant functional groups show more controls than environmental factors (climate and soil) on the N resorption traits (NRP and NRE), while site-related variables present more controls than plant types on PRP and PRE. NRP increases and PRP and NRE decrease significantly with increasing temperature and precipitation for the overall plants and for most groups, except that significant PRE–climate relationship holds for only evergreen angiosperms. Leaf nutrient resorption did not show consistent responses in relation to soil total N and P stoichiometry, probably because the resorption process is regulated by the relative costs of drawing nutrients from soil versus from senescing leaves. These results support our hypothesis that plants growing in P-limited habitats (low latitudes/altitudes or areas with high precipitation/temperature) should have lower PRP and higher PRE, compared with their counterparts in relatively N-limited places (high latitudes/altitudes or areas with low precipitation/temperature). Our findings can improve the understanding of variations in N and P resorption and their responses to global change, and thus facilitate to incorporate these nutrient resorption processes into future biogeochemical models.     

Serpentine and non-serpentine Silene dioica plants do not differ in nickel tolerance

Serpentine soils are hostile to plant life. They are dry, contain high concentrations of nickel and have an unfavorable calcium/magnesium ratio. The dioecious plant Silene dioica (L.) Clairv. (Caryophyllaceae) is the most common herb on serpentine soils in the Swedish mountains. It also commonly grows on non-serpentine soils in the subalpine and coastal area. I have compared the germination frequency, plant establishment and growth of serpentine and subalpine non-serpentine populations in serpentine soil under greenhouse conditions. Further more I have studied the specific effect of nickel on root and shoot growth of serpentine and non-serpentine plants from the subalpine and coastal area in solutions with different concentrations of nickel. Plants from serpentine and non-serpentine populations grew well and in a similar fashion in serpentine soil. Moreover, S. dioica plants, irrespective of original habitat, tolerated enhanced concentrations of nickel when grown in solutions. An analysis of metal content in serpentine plants from natural populations shows that S. dioica has a higher nickel concentration in the roots than in the shoots. The growth studies show that S. dioica is constitutively adapted to serpentine, and that all populations have the genetic and ecological tolerance to grow on serpentine.     

Nutrient resorption in wetland macrophytes: comparison across several regions of different nutrient status

This study explored patterns of nutrient resorption in wetland macrophytes to test the prediction that plants from regions with a strong nutrient limitation will show higher resorption of the limiting nutrient. Nitrogen and phosphorus resorption was assessed in macrophytes from marshes of different nutrient status in tropical and temperate regions, and expressed as resorption efficiency (NRE, PRE) and proficiency (NRP, PRP). Macrophytes were grouped into three categories: Typha, graminoids and broadleaved plants. Nitrogen was less limiting than P, consequently N availability varied less than P availability, NRP and NRE were lower, and N resorption was mostly incomplete. NRP was determined more by growth form than by local conditions. The large range of soil P concentrations allowed an exploration of relationships between P availability and resorption along a wide gradient. P-limited macrophytes (N : P > 16) had significantly higher PRP and PRE. Resorption proficiency was found to be a more sensitive indicator of changes in nutrient availability than resorption efficiency. The results confirmed that resorption in wetland macrophytes depends on nutrient availability, and is higher at nutrient-limited sites. A particularly strong relationship was found between resorption indicators and P limitation expressed either as live tissue N : P or soil P.     

Congeneric Serpentine and Nonserpentine Shrubs Differ More in Leaf Ca:Mg than in Tolerance of Low N, Low P, or Heavy Metals

Serpentine soils limit plant growth by NPK deficiencies, low Ca availability, excess Mg, and high heavy metal levels. In this study, three congeneric serpentine and nonserpentine evergreen shrub species pairs were grown in metalliferous serpentine soil with or without NPKCa fertilizer to test which soil factors most limit biomass production and mineral nutrition responses. Fertilization increased biomass production and allocation to leaves while decreasing allocation to roots in both serpentine and nonserpentine species. Simultaneous increases in biomass and leaf N:P ratios in fertilized plants of all six species suggest that N is more limiting than P in this serpentine soil. Neither N nor P concentrations, however, nor root to shoot translocation of these nutrients, differed significantly between serpentine and nonserpentine congeners. All six species growing in unfertilized serpentine soil translocated proportionately more P to leaves compared to fertilized plants, thus maintaining foliar P. Leaf Ca:Mg molar ratios of the nonserpentine species were generally equal to that of the soil. The serpentine species, however, maintained significantly higher leaf Ca:Mg than both their nonserpentine counterparts and the soil. Elevated leaf Ca:Mg in the serpentine species was achieved by selective Ca transport and/or Mg exclusion operating at the root-to-shoot translocation level, as root Ca and Mg concentrations did not differ between serpentine and nonserpentine congeners. All six species avoided shoot toxicity of heavy metals by root sequestration. The comparative data on nutrient deficiencies, leaf Ca:Mg, and heavy metal sequestration suggest that the ability to maintain high leaf Ca:Mg is a key evolutionary change needed for survival on serpentine soil and represents the physiological feature distinguishing the serpentine shrub species from their nonserpentine congeners. The results also suggest that high leaf Ca:Mg is achieved in these serpentine species by selective translocation of Ca and/or inhibited transport of Mg from roots, rather than by uptake/exclusion at root surfaces.     

Responses to Mg/Ca balance in an Iranian serpentine endemic plant, Cleome heratensis (Capparaceae) and a related non-serpentine species, C. foliolosa

A soil Ca/Mg quotient greater than unity is generally considered necessary for normal plant growth but some serpentine plants are adapted to much lower Ca/Mg quotients, resulting from a major cation imbalance in their substrata. In order to investigate the growth and tolerance responses of serpentine and non-serpentine species to varied Ca/Mg quotients, controlled nutrient solution experiments were performed using an a newly reported Iranian endemic serpentine plant, Cleome heratensis Bunge et Bien. Ex Boiss. and a related non-serpentine species Cleome foliolosa DC. and a Eurasian Ni-hyperaccumulating species Alyssum murale Waldst. and Kit. Seedlings were grown in modified Hoagland’s solutions with varying Ca and Mg concentrations (0.2–2.5 and 0.5–10 mM, respectively) in a fully factorial randomised block design. The yields of the two serpentine plants increased significantly as Mg concentrations in the nutrient solution were increased from 0.5 to 4 mM but decreased in the 10 mM Mg treatment. For C. foliolosa yields decreased significantly from 0.5 to 10 mM Mg, indicating the sensitivity of this non-serpentine plant, and the relative tolerance of the serpentine plants to extremely high levels of Mg. Shoot and root Mg and Ca concentrations in C. heratensis and A. murale were higher than those in C. foliolosa in the low and moderate Mg treatments, supporting the view that many serpentine plants have a relatively high requirement for Mg. Maximum Mg concentrations were found in the roots of C. heratensis. Yields of C. heratensis and A. murale did not change significantly as Ca levels in nutrient solution increased from 0.2 to 2.5 mM Ca, However the yield of C. foliolosa increased significantly from 0.2 to 1.5 mM Ca, indicating sensitivity in this non-serpentine plant and tolerance of the two serpentine plants to low levels of Ca correlated with tissue Ca concentrations, probably because of a greater ability for Ca uptake at low-Ca availability. Calcium deficiency in the low-Ca treatments could be a reason for reduced yield in the non-serpentine plants.     

Rapid development of phosphorus limitation in temperate rainforest along the Franz Josef soil chronosequence

The aim of this study was to examine how shifts in soil nutrient availability along a soil chronosequence affected temperate rainforest vegetation. Soil nutrient availability, woody plant diversity, composition and structure, and woody species leaf and litter nutrient concentrations were quantified along the sequence through ecosystem progression and retrogression. In this super-wet, high leaching environment, the chronosequence exhibited rapid soil development and decline within 120,000 years. There were strong gradients of soil pH, N, P and C, and these had a profound effect on vegetation. N:P_leaf increased along the chronosequence as vegetation shifted from being N- to P- limited. However, high N:P_leaf ratios, which indicate P-limitation, were obtained on soils with both high and low soil P availability. This was because the high N-inputs from an N-fixing shrub caused vegetation to be P-limited in spite of high soil P availability. Woody species nutrient resorption increased with site age, as availability of N and P declined. Soil P declined 8-fold along the sequence and P resorption proficiency decreased from 0.07 to 0.01%, correspondingly. N resorption proficiency decreased from 1.54 to 0.26%, corresponding to shifts in mineralisable N. Woody plant species richness, vegetation cover and tree height increased through ecosystem progression and then declined. During retrogression, the forest became shorter, more open and less diverse, and there were compositional shifts towards stress-tolerant species. Conifers (of the Podocarpaceae) were the only group to increase in richness along the sequence. Conifers maintained a lower N:P_leaf than other groups, suggesting superior acquisition of P on poor soils. In conclusion, there was evidence that P limitation and retrogressive forests developed on old soils, but N limitation on very young soils was not apparent because of inputs from an abundant N-fixing shrub.Electronic Supplementary Material Supplementary material is available in the online version of this article at .     

Leaf nutrient dynamics and nutrient resorption: a comparison between larch plantations and adjacent secondary forests in Northeast China

Aims Conversion of secondary forests to pure larch plantations is a common management practice driven by the increasing demand for timber production in Northeast China, resulting in a reduction in soil nutrient availability after a certain number of years following conversion. Nutrient resorption prior to leaf senescence was related to soil fertility, an important nutrient conservation strategy for plants, being especially significant in nutrient-poor habitats. However, the seasonal dynamics of leaf nutrients and nutrient resorption in response to secondary forest conversion to larch plantations is not well understood.Methods A comparative experiment between larch plantations (Larix spp.) and adjacent secondary forests (dominant tree species including Quercus mongolica, Acer mono, Juglans mandshurica and Fraxinus rhynchophylla) was conducted. We examined the variations in leaf nutrient (macronutrients: N, P, K, Ca and Mg; micronutrients: Cu and Zn) concentrations of these tree species during the growing season from May to October in 2013. Nutrient resorption efficiency and proficiency were compared between Larix spp. and the broadleaved species in the secondary forests.Important findings Results show that the seasonal variation of nutrient concentrations in leaves generally exhibited two trends, one was a downward trend for N, P, K, Cu and Zn, and another was an upward trend for Ca and Mg. The variations in foliar nutrient concentrations were mainly controlled by the developmental stage of leaves rather than by tree species. Resorption of the observed seven elements varied among the five tree species during leaf senescence. Nutrient resorption efficiency varied 6–75% of N, P, K, Mg, Cu and Zn, while Ca was not retranslocated in the senescing leaves of all species, and Mg was not retranslocated in Larix spp. Generally, Larix spp. tended to be more efficient and proficient (higher than 6–30% and 2–271% of nutrient resorption efficiency and resorption proficiency, respectively) in resorbing nutrients than the broadleaved species in the secondary forests, indicating that larch plantations had higher leaf nutrient resorption and thus nutrient use efficiency. Compared with Larix spp., more nutrients would remain in the leaf litter of the secondary forests, indicating an advantage of secondary forests in sustaining soil fertility. In contrast, the larch plantation would reuse internal nutrients rather than lose nutrients with litter fall and thus produce a positive feedback to soil nutrient availability. In summary, our results suggest that conversion from secondary forests to pure larch plantations would alter nutrient cycling through a plant-mediated pathway.     

Mutualism and adaptive divergence: co-invasion of a heterogeneous grassland by an exotic legume-rhizobium symbiosis

Species interactions play a critical role in biological invasions. For example, exotic plant and microbe mutualists can facilitate each other's spread as they co-invade novel ranges. Environmental context may influence the effect of mutualisms on invasions in heterogeneous environments, however these effects are poorly understood. We examined the mutualism between the legume, Medicago polymorpha, and the rhizobium, Ensifer medicae, which have both invaded California grasslands. Many of these invaded grasslands are composed of a patchwork of harsh serpentine and relatively benign non-serpentine soils. We grew legume genotypes collected from serpentine or non-serpentine soil in both types of soil in combination with rhizobium genotypes from serpentine or non-serpentine soils and in the absence of rhizobia. Legumes invested more strongly in the mutualism in the home soil type and trends in fitness suggested that this ecotypic divergence was adaptive. Serpentine legumes had greater allocation to symbiotic root nodules in serpentine soil than did non-serpentine legumes and non-serpentine legumes had greater allocation to nodules in non-serpentine soil than did serpentine legumes. Therefore, this invasive legume has undergone the rapid evolution of divergence for soil-specific investment in the mutualism. Contrary to theoretical expectations, the mutualism was less beneficial for legumes grown on the stressful serpentine soil than on the non-serpentine soil, possibly due to the inhibitory effects of serpentine on the benefits derived from the interaction. The soil-specific ability to allocate to a robust microbial mutualism may be a critical, and previously overlooked, adaptation for plants adapting to heterogeneous environments during invasion.     

Phosphorus conservation during post‐fire regeneration in a Chilean temperate rainforest

Nitrogen and phosphorus are the main elements limiting net primary production in terrestrial ecosystems. When growing in nutrient‐poor soils, plants develop physiological mechanisms to conserve nutrients, such as reabsorbing elements from senescing foliage (i.e. nutrient retranslocation). We investigated the changes in soil N and P in post‐fire succession in temperate rainforests of southern Chile. In this area, forest recovery often leads to spatially scattered, discrete regeneration with patches varying in age, area, species richness and tree cover, representing different degrees of recovery from disturbance. We hypothesized that soil nutrient concentrations should differ among tree regenerating patches depending on the progress of forest regeneration and that nutrient resorption should increase over time as colonizing trees respond to limited soil nutrients. To evaluate these hypotheses, we sampled 40 regeneration patches in an area of 5 ha, spanning a broad range of vegetation complexity, and collected soil, tree foliage and litter samples to determine N and P concentrations. Nutrient concentrations in leaf litter were interpreted as nutrient resorption proficiency. We found that soil P was negatively correlated with all the indicators of successional progress, whereas total soil N was independent of the successional progress. Foliar N and P were unrelated to soil nutrient concentrations; however, litter N was negatively related to soil N, and litter P was positively related with soil P. Finally, foliar N:P ratios ranged from 16 to 25, which suggests that P limitation can hamper post‐fire regeneration. We provide evidence that after human‐induced fires, succession in temperate forests of Chile can become nutrient limited and that high nutrient retranslocation is a key nutrient conservation strategy for regenerating tree communities.     

Combined effects of nitrogen addition and litter manipulation on nutrient resorption of Leymus chinensis in a semi‐arid grassland of northern China

Plant growth in semi‐arid ecosystems is usually severely limited by soil nutrient availability. Alleviation of these resource stresses by fertiliser application and aboveground litter input may affect plant internal nutrient cycling in such regions. We conducted a 4‐year field experiment to investigate the effects of nitrogen (N) addition (10 g N·m^?2·year^?1) and plant litter manipulation on nutrient resorption of Leymus chinensis, the dominant native grass in a semi‐arid grassland in northern China. Although N addition had no clear effects on N and phosphorus (P) resorption efficiencies in leaves and culms, N fertilisation generally decreased leaf N resorption proficiency by 54%, culm N resorption proficiency by 65%. Moreover, N fertilisation increased leaf P resorption proficiency by 13%, culm P resorption proficiency by 20%. Under ambient or enriched N conditions, litter addition reduced N and P resorption proficiencies in both leaves and culms. The response of P resorption proficiency to litter manipulation was more sensitive than N resorption proficiency: P resorption proficiency in leaves and culms decreased strongly with increasing litter amount under both ambient and enriched N conditions. In contrast, N resorption proficiency was not significantly affected by litter addition, except for leaf N resorption proficiency under ambient N conditions. Furthermore, although litter addition caused a general decrease of leaf and culm nutrient resorption efficiencies under both ambient and enriched N conditions, litter addition effects on nutrient resorption efficiency were much weaker than the effects of litter addition on nutrient resorption proficiency. Taken together, our results show that leaf and non‐leaf organs of L. chinensis respond consistently to altered soil N availability. Our study confirms the strong effects of N addition on plant nutrient resorption processes and the potential role of aboveground litter, the most important natural fertiliser in terrestrial ecosystems, in influencing plant internal nutrient cycling.     

Nutrient resorption in tropical savanna forests and woodlands of central Brazil

Nutrient limitation in Brazilian savanna (known as cerrado) presumably causes trees to maximize nutrient resorption from senesced leaves to reduce their dependence on nutrient availability. To assess patterns between nutrient resorption and soil fertility, we measured community-level nitrogen (N), phosphorus (P), and potassium (K) concentrations in mature and senesced leaves and soil fertility in the upper 50 cm soil layer in structurally diverse cerrado ecosystems in the Cuiaba Basin (CB) and Pantanal (PAN) of Mato Grosso, Brazil. Foliar nutrient concentration data were used to estimate resorption efficiency and proficiency, and correlation was used to determine whether resorption efficiency and proficiency varied across soil fertility gradients. We found that N and P resorption proficiency (NRP and PRP, respectively) and P resorption efficiency (PRE) increased significantly as total soil N (NRP) and extractable P (PRP and PRE) declined. In contrast, K resorption efficiency (KRE) declined as soil sand content and bulk density increased, which was likely due to a reduction in soil water-holding capacity. Leaf N/P ratios indicate potential N limitation and/or N + P co-limitation for ecosystems in the PAN and P limitation and/or N + P co-limitation for ecosystems in the CB, while trends in leaf N/K ratios indicate possible K or K + P co-limitation for the CB only. Our results illustrate that cerrado forests and woodlands have highly variable nutrient resorption capacities that vary predictably across soil fertility or textural gradients and indicate that cerrado communities have flexible nutrient resorption that can reduce their dependence on soil nutrient availability.     

Genomic analysis of differentiation between soil types reveals candidate genes for local adaptation in Arabidopsis lyrata

Serpentine soil, which is naturally high in heavy metal content and has low calcium to magnesium ratios, comprises a difficult environment for most plants. An impressive number of species are endemic to serpentine, and a wide range of non-endemic plant taxa have been shown to be locally adapted to these soils. Locating genomic polymorphisms which are differentiated between serpentine and non-serpentine populations would provide candidate loci for serpentine adaptation. We have used the Arabidopsis thaliana tiling array, which has 2.85 million probes throughout the genome, to measure genetic differentiation between populations of Arabidopsis lyrata growing on granitic soils and those growing on serpentinic soils. The significant overrepresentation of genes involved in ion transport and other functions provides a starting point for investigating the molecular basis of adaptation to soil ion content, water retention, and other ecologically and economically important variables. One gene in particular, calcium-exchanger 7, appears to be an excellent candidate gene for adaptation to low CaratioMg ratio in A. lyrata.     

Nickel hyperaccumulation by Thlaspi montanum var. montanum (Brassicaceae): a constitutive trait

Adaptations to particular stresses may occur only in populations experiencing those stresses or may be widespread within a species. Nickel hyperaccumulation is viewed as an adaptation to high-Ni (serpentine) soils, but few studies have determined if hyperaccumulation ability is restricted to populations from high-Ni soils or if it is a constitutive trait found in populations on both high- and low-Ni soils. We compared mineral element concentrations of Thlaspi montanum var. montanum plants grown on normal and high-Ni greenhouse soils to address this question. Seed sources were from four populations (two serpentine, two non-serpentine) in Oregon and northern California, USA. Plants from all populations were able to hyperaccumulate Ni, showing Ni hyperaccumulation to be a constitutive trait in this species. Populations differed in their ability to extract some elements (e.g., Ca, Mg, P) from greenhouse soils. We noted a negative correlation between tissue concentrations of Ni and Zn. We suggest that the ability to hyperaccumulate Ni has adaptive value to populations growing on non- serpentine soil. This adaptive value may be a consequence of metal-based plant defense against herbivores/pathogens, metal- based interference against neighboring plant species, or an efficient nutrient scavenging system. We suggest that the Ni hyperaccumulation ability of T. montanum var. montanum may be an inadvertent consequence of an efficient nutrient (possibly Zn or Ca) uptake system.     

Foliar nutrient resorption patterns of four functional plants along a precipitation gradient on the Tibetan Changtang Plateau

Nutrient resorption from senesced leaves as a nutrient conservation strategy is important for plants to adapt to nutrient deficiency, particularly in alpine and arid environment. However, the leaf nutrient resorption patterns of different functional plants across environmental gradient remain unclear. In this study, we conducted a transect survey of 12 communities to address foliar nitrogen (N) and phosphorus (P) resorption strategies of four functional groups along an eastward increasing precipitation gradient in northern Tibetan Changtang Plateau. Soil nutrient availability, leaf nutrient concentration, and N:P ratio in green leaves ([N:P]_g) were linearly correlated with precipitation. Nitrogen resorption efficiency decreased, whereas phosphorus resorption efficiency except for sedge increased with increasing precipitation, indicating a greater nutrient conservation in nutrient‐poor environment. The surveyed alpine plants except for legume had obviously higher N and P resorption efficiencies than the world mean levels. Legumes had higher N concentrations in green and senesced leaves, but lowest resorption efficiency than nonlegumes. Sedge species had much lower P concentration in senesced leaves but highest P resorption efficiency, suggesting highly competitive P conservation. Leaf nutrient resorption efficiencies of N and P were largely controlled by soil and plant nutrient, and indirectly regulated by precipitation. Nutrient resorption efficiencies were more determined by soil nutrient availability, while resorption proficiencies were more controlled by leaf nutrient and N:P of green leaves. Overall, our results suggest strong internal nutrient cycling through foliar nutrient resorption in the alpine nutrient‐poor ecosystems on the Plateau. The patterns of soil nutrient availability and resorption also imply a transit from more N limitation in the west to a more P limitation in the east Changtang. Our findings offer insights into understanding nutrient conservation strategy in the precipitation and its derived soil nutrient availability gradient.     

Soil characteristics and nitrogen resorption in Stipa krylovii native to northern China

Nitrogen (N) resorption from senescing tissues enables plants to conserve and reuse this important nutrient. As such, it is expected that plant species adapted to infertile soils could have a higher N-resorption efficiency (percentage reduction of nitrogen between green and senescing tissues) and/or higher N-resorption proficiency (absolute reduction of nitrogen in senescing tissues) than those adapted to fertile soils. To test this hypothesis, we investigated the relationships among soil characteristics (total N, nitrate-N, ammonium-N, pH and moisture) and N resorption in Stipa kryloviiRoshev., a species occurred widely in natural grasslands of northern China. N contents in green and senescing tissues were 6.7±0.1 and 3.3±0.1 mg g^–1, respectively. The mean value of N-resorption efficiency was found to be 72.1%. The N-resorption efficiency in S. kryloviiwas independent of soil characteristics. The N-resorption proficiency in S. kryloviiwas dependent on soil nitrate- and ammonium-N, but it was relatively independent of soil total N. The N-resorption proficiency was negatively correlated with soil pH and moisture. There was a positive correlation between N concentration in green tissues and resorption efficiency. However, N-resorption efficiency was not correlated significantly with N concentration in senescing tissues. These results indicate that the intraspecific variation in N resorption of Stipa kryloviiRoshev. is associated with soil regimes and that higher N resorption on N-poor soils is an adaptive strategy for S. kryloviito maximize N use under conditions of limited N supply.