Foliar nutrient allocation patterns in Banksia attenuata and Banksia sessilis differing in growth rate and adaptation to low-phosphorus habitats

Background and AimsPhosphorus (P) and nitrogen (N) are essential nutrients that frequently limit primary productivity in terrestrial ecosystems. Efficient use of these nutrients is important for plants growing in nutrient-poor environments. Plants generally reduce foliar P concentration in response to low soil P availability. We aimed to assess ecophysiological mechanisms and adaptive strategies for efficient use of P in Banksia attenuata (Proteaceae), naturally occurring on deep sand, and B. sessilis, occurring on shallow sand over laterite or limestone, by comparing the allocation of P among foliar P fractions.MethodsWe carried out pot experiments with slow-growing B. attenuata, which resprouts after fire, and faster growing opportunistic B. sessilis, which is killed by fire, on substrates with different P availability using a randomized complete block design. We measured leaf P and N concentrations, photosynthesis, leaf mass per area, relative growth rate and P allocated to major biochemical fractions in B. attenuata and B. sessilis.Key ResultsThe two species had similarly low foliar total P concentrations, but distinct patterns of P allocation to P-containing fractions. The foliar total N concentration of B. sessilis was greater than that of B. attenuata on all substrates. The foliar total P and N concentrations in both species decreased with decreasing P availability. The relative growth rate of both species was positively correlated with concentrations of both foliar nucleic acid P and total N, but there was no correlation with other P fractions. Faster growing B. sessilis allocated more P to nucleic acids than B. attenuata did, but other fractions were similar.ConclusionsThe nutrient allocation patterns in faster growing opportunistic B. sessilis and slower growing B. attenuata revealed different strategies in response to soil P availability which matched their contrasting growth strategy.     

Relationship between the Relative Limitation and Resorption Efficiency of Nitrogen vs Phosphorus in Woody Plants

Most previous studies have ascribed variations in the resorption of a certain plant nutrient to its corresponding environmental availability or level in tissues, regardless of the other nutrients’ status. However, given that plant growth relies on both sufficient and balanced nutrient supply, the nutrient resorption process should not only be related to the absolute nutrient status, but also be regulated by the relative limitation of the nutrient. Here, based on a global woody-plants dataset from literature, we test the hypothesis that plants resorb proportionately more nitrogen (or phosphorus) when they are nitrogen (or phosphorus) limited, or similar proportions of nitrogen (N) and phosphorus (P) when co-limited by both nutrients (the relative resorption hypothesis). Using the N:P ratio in green foliage as an indicator of nutrient limitation, we found an inverse relationship between the difference in the proportionate resorption of N vs P and this foliar N:P ratio, consistent across species, growth-forms, and vegetation-types globally. Moreover, according to the relative resorption hypothesis, communities with higher/lower foliar N:P (more likely P/N limited) tend to produce litter with disproportionately higher/lower N:P, causing a worsening status of P/N availability; this positive feedback may somehow be counteracted by several negative-feedback mechanisms. Compared to N, P generally shows higher variability in resorption efficiency (proportion resorbed), and higher resorption sensitivity to nutrient availability, implying that the resorption of P seems more important for plant nutrient conservation and N:P stoichiometry. Our findings elucidate the nutrient limitation effects on resorption efficiency in woody plants at the global scale, and thus can improve the understanding of nutrient resorption process in plants. This study also suggests the importance of the foliar N:P ratio as a key parameter for biogeochemical modeling, and the relative resorption hypothesis used to deduce the critical (optimal) N:P ratio for a specific plant community.     

Conifer Presence May Negatively Affect Sugar Maple’s Ability to Migrate into the Boreal Forest Through Reduced Foliar Nutritional Status

The discipline of ecology suffers from a lack of knowledge of non-climatic factors (for example, plant–soil, plant–plant and plant–insect interactions) to predict tree species range shifts under climate change. The next generation of simulation models of forest response to climate change must build upon local observations of species interactions and growth along climatic gradients. We examined whether sugar maple (Acer saccharum) seedlings were disadvantaged with respect to soil nutrient uptake under coniferous canopies, as this species would need to migrate northward into conifer-dominated forests in response to climate change. An experimental design was applied to 3 sites, forming the largest possible latitudinal/climatic gradient for sugar maple in Quebec (Canada) and isolating the effect of conifer presence on its seedling’s nutritional status. We tested whether: (1) both soil and climate and (2) presence of conifers affected foliar nutrient levels of sugar maple seedlings. Climate and soil (through pH) strongly affected nutrient availability for sugar maple seedlings and predicted 63.7% of their foliar nutrient variability. When controlling for site effects, we found a significant negative effect of conifers on foliar Ca and Mg levels of maple seedlings, which can adversely affect their overall health and vigour. When considering projected modifications of the forest environment due to climate change, we suggest that northward migration of sugar maple will be negatively affected by the presence of conifers through reduced foliar nutrition.     

Influence of Precipitation on Soil and Foliar Nutrients Across Nine Costa Rican Forests

We explored patterns of soil and foliar nutrients across nine mature forest sites in Costa Rica, where mean annual precipitation (MAP) ranged between 3500 and 5500 mm, altitude ranged between 200 and 1200 m, and species composition varied among sites. Our objective was to investigate the relationship between rainfall and plant or soil nutrient characteristics to better understand the potential long‐term effects that alterations in MAP could have on the nutrient dynamics of wet forest plant communities. Indicators of soil N availability (net mineralization and nitrification) decreased with MAP but were not related to foliar N. Soil and foliar P, by contrast, were not correlated with MAP but were positively correlated with each other. Thus, across our gradient foliar P was a better predictor of soil nutrient availability than foliar N. There were wide differences in foliar nutrient concentrations and N:P ratios among species within sites. At each site, legumes had higher mean percent N than nonlegumes, resulting in higher N:P ratios for legumes. Taken together, these data suggest that, at least in these forests, a climate‐driven decrease in MAP could cause an increase in net N mineralization and nitrification for the wetter sites. However, this may not affect productivity at the community level because of low P availability, complex feedbacks between soil and foliar nutrients, and interactions with other biological and environmental factors such as elevation.     

Controls of Bedrock Geochemistry on Soil and Plant Nutrients in Southeastern Utah

The cold deserts of the Colorado Plateau contain numerous geologically and geochemically distinct sedimentary bedrock types. In the area near Canyonlands National Park in Southeastern Utah, geochemical variation in geologic substrates is related to the depositional environment with higher concentrations of Fe, Al, P, K, and Mg in sediments deposited in alluvial or marine environments and lower concentrations in bedrock derived from eolian sand dunes. Availability of soil nutrients to vegetation is also controlled by the formation of secondary minerals, particularly for P and Ca availability, which, in some geologic settings, appears closely related to variation of CaCO₃ and Ca-phosphates in soils. However, the results of this study also indicate that P content is related to bedrock and soil Fe and Al content suggesting that the deposition history of the bedrock and the presence of P-bearing Fe and Al minerals, is important to contemporary P cycling in this region. The relation between bedrock type and exchangeable Mg and K is less clear-cut, despite large variation in bedrock concentrations of these elements. We examined soil nutrient concentrations and foliar nutrient concentration of grasses, shrubs, conifers, and forbs in four geochemically distinct field sites. All four of the functional plant groups had similar proportional responses to variation in soil nutrient availability despite large absolute differences in foliar nutrient concentrations and stoichiometry across species. Foliar P concentration (normalized to N) in particular showed relatively small variation across different geochemical settings despite large variation in soil P availability in these study sites. The limited foliar variation in bedrock-derived nutrients suggests that the dominant plant species in this dryland setting have a remarkably strong capacity to maintain foliar chemistry ratios despite large underlying differences in soil nutrient availability.     

Effects of nutrients and arbuscular mycorrhizal colonization on the growth of Salix gracilistyla seedlings in a nutrient-poor fluvial bar

A field survey and a pot culture experiment were conducted to examine the effects of nutrients (N and P) and arbuscular mycorrhizal (AM) fungi on the growth of Salix gracilistyla, a pioneer plant in riparian habitats. The plants growing in the field were colonized by AM and/or ectomycorrhizal fungi. However, the direct effect of AM colonization on seedling growth was not detected in the pot culture experiment. In contrast, N application significantly promoted plant growth, suggesting that the growth of S. gracilistyla seedlings is largely limited by the availability of N in the field.     

Tree physiological responses to above-ground herbivory directly modify below-ground processes of soil carbon and nitrogen cycling

Above‐ground herbivory is ubiquitous in terrestrial ecosystems, yet its impacts on below‐ground processes and consequences for plants remain ambiguous. To examine whether physiological responses of individual trees may potentially modify soil nutrient availability, we subjected Fagus sylvatica L. (European beech) and Abies alba Mill. (silver fir) to simulated foliar herbivory over two growing seasons. Above‐ground herbivory enhanced N mineralization and inorganic N availability in the soil. The total input of C from the plant roots to the soil is not known; however, carbon sequestration in the soil, measured using stable isotopic techniques, was unaffected by herbivory. Fagus responded to herbivory by producing larger leaves, with increased photosynthetic capacity and N content, which largely compensated for the loss of biomass; Abies exhibited no such response. We conclude that despite large interspecific differences in the growth response, tree physiological responses to foliar herbivory are capable of directly modifying soil biological processes.     

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.     

Coupled Nitrogen and Calcium Cycles in Forests of the Oregon Coast Range

Nitrogen (N) is a critical limiting nutrient that regulates plant productivity and the cycling of other essential elements in forests. We measured foliar and soil nutrients in 22 young Douglas-fir stands in the Oregon Coast Range to examine patterns of nutrient availability across a gradient of N-poor to N-rich soils. N in surface mineral soil ranged from 0.15 to 1.05% N, and was positively related to a doubling of foliar N across sites. Foliar N in half of the sites exceeded 1.4% N, which is considered above the threshold of N-limitation in coastal Oregon Douglas-fir. Available nitrate increased five-fold across this gradient, whereas exchangeable magnesium (Mg) and calcium (Ca) in soils declined, suggesting that nitrate leaching influences base cation availability more than soil parent material across our sites. Natural abundance strontium isotopes (⁸⁷Sr/⁸⁶Sr) of a single site indicated that 97% of available base cations can originate from atmospheric inputs of marine aerosols, with negligible contributions from weathering. Low annual inputs of Ca relative to Douglas-fir growth requirements may explain why foliar Ca concentrations are highly sensitive to variations in soil Ca across our sites. Natural abundance calcium isotopes (δ⁴⁴Ca) in exchangeable and acid leachable pools of surface soil measured at a single site showed 1 per mil depletion relative to deep soil, suggesting strong Ca recycling to meet tree demands. Overall, the biogeochemical response of these Douglas-fir forests to gradients in soil N is similar to changes associated with chronic N deposition in more polluted temperate regions, and raises the possibility that Ca may be deficient on excessively N-rich sites. We conclude that wide gradients in soil N can drive non-linear changes in base-cation biogeochemistry, particularly as forests cross a threshold from N-limitation to N-saturation. The most acute changes may occur in forests where base cations are derived principally from atmospheric inputs.     

Leaf litter variation influences invasion dynamics in the invasive wetland grass Phalaris arundinacea

High litter mass is hypothesized to produce an invader-directed invasion by changing ecosystem properties such as nutrient cycling rates and light availability. An invasive plant species that stimulates litter accumulation may induce a positive feedback when it benefits from high litter conditions. Phalaris arundinacea is an invasive wetland grass that may induce positive litter feedback, as it produces abundant litter that varies in quality due to a wide range of foliar C:N content. In this study we investigated the range of growth responses within native and invasive genotypes of Phalaris that varied in initial foliar C:N levels (high C:N content was present in the invasive genotypes) when grown under varying litter mass. Overwintering with high litter reduced establishing tiller survivorship and the presence of litter delayed tiller emergence by 2 weeks. Overall, genotypes exhibited high trait plasticity in response to litter. Our results indicate that high litter mass can stimulate Phalaris growth, specifically for the genotypes with high initial C:N foliar tissue. Additionally, genotypes with initially high C:N ratios exhibited plastic responses consistent with a Master-of-some strategy indicating that their performance under high litter may depend upon the nutrient conditions under which they are grown. This study provides evidence for conditions that may lead to a positive feedback in Phalaris’ introduced range. Future studies should investigate how changing litter quantity alters nutrient cycling and competitor growth.     

Nutrient Limitation of Native and Invasive N2-Fixing Plants in Northwest Prairies

Nutrient rich conditions often promote plant invasions, yet additions of non-nitrogen (N) nutrients may provide a novel approach for conserving native symbiotic N-fixing plants in otherwise N-limited ecosystems. Lupinus oreganus is a threatened N-fixing plant endemic to prairies in western Oregon and southwest Washington (USA). We tested the effect of non-N fertilizers on the growth, reproduction, tissue N content, and stable isotope δ¹⁵N composition of Lupinus at three sites that differed in soil phosphorus (P) and N availability. We also examined changes in other Fabaceae (primarily Vicia sativa and V. hirsuta) and cover of all plant species. Variation in background soil P and N availability shaped patterns of nutrient limitation across sites. Where soil P and N were low, P additions increased Lupinus tissue N and altered foliar δ¹⁵N, suggesting P limitation of N fixation. Where soil P was low but N was high, P addition stimulated growth and reproduction in Lupinus. At a third site, with higher soil P, only micro- and macronutrient fertilization without N and P increased Lupinus growth and tissue N. Lupinus foliar δ¹⁵N averaged −0.010‰ across all treatments and varied little with tissue N, suggesting consistent use of fixed N. In contrast, foliar δ¹⁵N of Vicia spp. shifted towards 0‰ as tissue N increased, suggesting that conditions fostering N fixation may benefit these exotic species. Fertilization increased cover, N fixation, and tissue N of non-target, exotic Fabaceae, but overall plant community structure shifted at only one site, and only after the dominant Lupinus was excluded from analyses. Our finding that non-N fertilization increased the performance of Lupinus with few community effects suggests a potential strategy to aid populations of threatened legume species. The increase in exotic Fabaceae species that occurred with fertilization further suggests that monitoring and adaptive management should accompany any large scale applications.     

Growth,nutrient acquisition and ectomycorrhizae of <Emphasis Type="Italic">Eucalyptus regnans</Emphasis> F. Muell. seedlings in fertilized or diluted air-dried and undried forest soil

Seedlings of Eucalyptus regnans (mountain ash) grow poorly in undried forest soil, where they develop purple coloration in the foliage, but their growth is markedly improved when forest soil has been air dried. Whether this growth promotion is purely due to improved nutrient status of the soil, as a result of air drying, was investigated. In several pot experiments, E. regnans seedlings were grown (i) in air-dried and undried forest soil with addition of different levels of complete fertiliser, (ii) in air-dried or undried soil diluted to different extents with sand, or (iii) in undried soil mixed with different amounts of air-dried soil. Seedling dry weight, P content and incidence of ectomycorrhizal root tips were determined.In all experiments, the dry weights of seedlings were 3–6 times greater in 100% air-dried soil than in 100% undried soil. Fertiliser application resulted in a significant increase in dry weight of seedlings in both air-dried and undried soil, but the dry weights in air-dried soil were always significantly greater than those in undried soil at the same level of fertiliser application. Even at the highest level of fertiliser application, the growth difference between seedlings in air-dried and undried soil remained. When air-dried soil was diluted with sand, there was a significant reduction in seedling dry weight only when soil was diluted to 20% or less (air-dried soil:total mix). Conversly, when air-dried soil was mixed with undried soil, there was a proportional decrease in seedling dry weight with increasing amounts of undried soil. In all experiments, the dominant ectomycorrhizal morphotypes in 100% air-dried soil were different from those in undried soil. Fertilisation and dilution of air-dried and undried soil did not result in a reduction in the overall incidence of ectomycorrhizal root tips, although the frequency of occurrence of different ectomycorrhizal morphotypes was affected.It is concluded that the growth difference between seedlings in air-dried and undried forest soils is not due solely to differences in the direct availability of nutrients in the soils, and different ectomycorrhizae may indirectly affect nutrient availability to the plant.     

Nutrient availability effects on vesicular-arbuscular mycorrhizal bell pepper (Capsicum annuum) seedlings and transplants

Pepper (Capsicum annuum) seeds were sown in nutrient-poor sand or nutrient-rich peat/vermiculite amended or not amended with Glomus macrocarpum. The vesicular-arbuscular mycorrhizal (VAM) seedlings were irrigated with three levels of nutrient solution, and transplanted into four levels of P-amended soil, each of which was irrigated with two levels of nutrient solution minus P. Mycorrhizal seedlings in sand were responsive to increasing nutrient levels; in nutrient-rich peat the seedlings did not respond to additional fertilisation. The greatest seedling development accompanied by good fungus colonisation was in nutrient-poor medium irrigated with the highest nutrient solution tested (18 mM N, 1.2 mM P, and 7 mM K). Non-VAM plants almost ceased growing between the weeks 4 and 5, whereas VAM plants increased in weight by 41–188%. After transplanting, sand-grown seedlings benefited from VAM when 300 mg P/kg or more was added to the soil but peat-grown plants did not. Fruit development was delayed in all non-VAM plants compared with VAM ones.     

Effects of vesicular-arbuscular mycorrhizal inoculation at different soil P availabilities on growth and nutrient uptake of in vitro propagated coffee (Coffea arabica L.) plants

 In a pot experiment, the growth and the nutrient status of in vitro propagated coffee (Coffea arabica L.) microcuttings were investigated for 5 months following vesicular-arbuscular mycorrhizal (VAM) inoculation with either Acaulospora melleae or Glomus clarum at four soil P availabilities. Control plants remained P-deficient even at the highest soil P availability while mycorrhizal plants were P-sufficient at all soil P availabilities. Growth of control plants was only improved at the highest soil P availability. In P-deficient soil, neither of the two VAM species improved plant growth. Plant growth increased by 50% following inoculation with either A. melleae or G. clarum when P availability went from deficient to low. No further plant growth improvement was induced by either VAM species at intermediate and high soil P levels. Nevertheless, growth of plants inoculated with G. clarum was still significantly greater than that of non-mycorrhizal plants at the highest soil P availability. Root colonization by G. clarum increased with increasing soil P availability while root colonization by A. mellea decreased with soil P level increasing above low P availability. Soil P availability also affected Zn nutrition through its influence on VAM symbiosis. With increasing soil P availability, foliar Zn status increased with G. clarum or decreased with A. mellea in parallel to root colonization by VAM. This study demonstrates the beneficial effects of VAM inoculation on in vitro propagated Arabica coffee microcuttings, as shown previously for seedlings. This study also demonstrates differences in tolerance to soil P availability between VAM species, most likely resulting from their differing abilities to enhance coffee foliar P status. Accepted: 14 November 1996     

Tight Coupling Between Shoot Level Foliar N and P,Leaf Area,and Shoot Growth in Arctic Dwarf Shrubs Under Simulated Climate Change

Nutrient availability limits productivity of arctic ecosystems, and this constraint means that the amount of nitrogen (N) in plant canopies is an exceptionally strong predictor of vegetation productivity. However, climate change is predicted to increase nutrient availability leading to increases in carbon sequestration and shifts in community structure to more productive species. Despite tight coupling of productivity with canopy nutrients at the vegetation scale, it remains unknown how species/shoot level foliar nutrients couple to growth, or how climate change may influence foliar nutrients–productivity relationships to drive changes in ecosystem carbon gain and community structure. We investigated the influence of climate change on arctic plant growth relationships to shoot level foliar N and phosphorus (P) in three dominant subarctic dwarf shrubs using an 18-year warming and nutrient addition experiment. We found a tight coupling between total leaf N and P per shoot, leaf area and shoot extension. Furthermore, a steeper shoot length-leaf N relationship in deciduous species (Vaccinium myrtillus and Vaccinium uliginosum) under warming manipulations suggests a greater capacity for nitrogen to stimulate growth under warmer conditions in these species. This mechanism may help drive the considerable increases in deciduous shrub cover observed already in some arctic regions. Overall, our work provides the first evidence at the shoot level of tight coupling between foliar N and P, leaf area and growth i.e. consistent across species, and provides mechanistic insight into how interspecific differences in alleviation of nutrient limitation will alter community structure and primary productivity in a warmer Arctic.     

Response of conifer seedlings to nitrate and ammonium sources of nitrogen

Summary Differences in growth responses of Douglas fir, western hemlock, Sitka spruce, and white spruce to nitrate and ammonium N sources were examined in sand culture and artificial soil culture. Effects of the two forms of N on growth, needle area, and N uptake of three Douglas fir halb-sib progenies were examined in a second sand culture. Response of Douglas fir to the two forms of N was followed over two years in nursery soil of different pH levels. In sand culture 1 mean seedling dry weight of all species, except hemlock, was greatest when ammonium N and nitrate N were provided in equal amounts. In all species, except Sitka spruce, ammonium alone resulted in greater growth than nitrate alone. Use of ammonium N resulted in greater growth of all species, than was obtained with nitrate N, at pH values in the region 5.4 and 7.5 in artificial soil culture. Only Douglas fir showed substantial differences due to N source below pH 5. Growth of all species was greater at pH 5.4 than at 7.5 in each N source treatment. Growth of Douglas fir seedlings was greatest with ammonium N and least with nitrate N in sand culture 2. Supply of nitrate and ammonium in equal proportions resulted in intermediate growth. Leaf area/plant weight ratio was unaffected by N source. Analysis of nutrient solutions showed appreciable nitrification of ammonium N during the 7 days between solution changes. In the three greenhouse experiments, with little exception, increase in proportion of ammonium in N supply resulted in increase of seedling tissue N concentration. This effect was more pronounced in roots than shoots. Total N uptake by ammonium fed seedlings was about double the N uptake of nitrate fed seedlings in sand culture 2. Nursery grown Douglas fir seedlings showed greater growth response to ammonium sulphate than to calcium nitrate, and this appeared due entirely to form of N supply in the first year. A similar response in the second year was partly due to greater soil acidification by ammonium sulphate. Compared with calcium nitrate, ammonium sulphate increased N concentration of one-year old shoots, but this difference was not detected by foliar analysis of two-year old seedlings.     

Plant availability of nutrients recovered as solids from human urine tested in climate chamber on Triticum aestivum L

Recovered nutrients by freezing-thawing from human urine in combination with struvite precipitation and nitrogen adsorption on zeolite and activated carbon have been tested in pot trials with wheat, Triticum aestivum L., in a climate chamber during 21 days. A simple test design using sand as substrate was chosen to give a first, general evaluation of the nutrient (P and N) availability from these sources. Dry weight, plant growth morphology, total-P and total-N were analysed. The tests show a slow-release of nutrients (P and N) from struvite and from N-adsorbents. The nitrogen in all treatments was in the deficiency range for optimum yield for wheat. Higher pH than usual for soil tests contributed to the difficulties in plant uptake, especially in the pots with only struvite (with highest MgO addition) as nutrient source.     

Effect of nitrogen,phosphorus and potassium availability on emergence,nodulation and growth of Trifolium medium L. in alkaline soil

Little is known about the effects of nutrient availability on the growth of Trifolium medium in alkaline soil. In 2010, a pot experiment (10 N, P and K fertiliser treatments) with seeding of T. medium into alkaline soil was performed and emergence of seedlings, survival, aboveground and belowground organs were studied. The positive effects of increased nutrient availability on seedling emergence ranged from 5% in the control to 17% in the high P treatment. The lowest mortality was in treatments with P and K supply and the highest in treatments with N supply, due to the sensitivity of young plants to high N availability. The highest values of most measured aboveground plant traits were recorded in treatments with simultaneous application of N, P and K. There were highly positive effects of P supply alone or in combination with N and K on the development of belowground organs. Taproot length ranged from 11.5 in high N to 40.2 cm in P treatment. There was a negative effect of N application on nodulation, especially in N treatments, where growth of T. medium was limited by insufficient P supply. The number of nodules per plant ranged from 0.8 to 4.5 in the high N and P treatments. As demonstrated in this study, T. medium is a potentially suitable legume for alkaline soils. It requires a relatively high P and K supply as well as moderate mineral N supply to achieve its maximum growth potential.     

Nitrogen, phosphorus and potassium nutritional status of semiarid steppe grassland in Inner Mongolia

In grazed semiarid steppe ecosystems, much attention has been paid to aspects of growth limitation by water. So far, potential limitation of primary production by plant nutrients was rarely considered. This knowledge is essential for identification of sustainable land-use practices in these large and important ecosystems on the background of over-exploitation and climate change. In the present study plant nutrient concentrations and ratios were investigated with factorial additions of water and N fertilizer at two sites with contrasting soil nutrient availability. Combined analysis of nutrient concentrations, contents, biomass production, and plant N:P ratios consistently confirmed primary growth limitation by water and a strong N limitation when sufficient amounts of water were supplied. P limitation only occurred at the site with low P availability when in addition to the natural supply, water and N fertilizer were given. According to reported thresholds of N:K and K:P ratios, K was not limiting in any plot. The observed nutritional patterns in the plant community were related to the dynamics of species composition and their specific nutrient status. Stipa grandis had the highest N:P ratio whereas Artemisia frigida showed lowest N:P. These nutrient characteristics were related to growth strategies of dominant species. Accordingly, the relative biomass contribution of S. grandis and A. frigida strongly affected the nutrient status of the plant community. Plant N:P ratios indicate the relative limitation by N or P in the semiarid grasslands under sufficient water supply, but other methods of nutritional diagnosis should be used when plant N:P ratios remain below critical values.     

Imaging spectroscopy reveals the effects of topography and logging on the leaf chemistry of tropical forest canopy trees

Logging, pervasive across the lowland tropics, affects millions of hectares of forest, yet its influence on nutrient cycling remains poorly understood. One hypothesis is that logging influences phosphorus (P) cycling, because this scarce nutrient is removed in extracted timber and eroded soil, leading to shifts in ecosystem functioning and community composition. However, testing this is challenging because P varies within landscapes as a function of geology, topography and climate. Superimposed upon these trends are compositional changes in logged forests, with species with more acquisitive traits, characterized by higher foliar P concentrations, more dominant. It is difficult to resolve these patterns using traditional field approaches alone. Here, we use airborne light detection and ranging‐guided hyperspectral imagery to map foliar nutrient (i.e. P, nitrogen [N]) concentrations, calibrated using field measured traits, over 400 km² of northeastern Borneo, including a landscape‐level disturbance gradient spanning old‐growth to repeatedly logged forests. The maps reveal that canopy foliar P and N concentrations decrease with elevation. These relationships were not identified using traditional field measurements of leaf and soil nutrients. After controlling for topography, canopy foliar nutrient concentrations were lower in logged forest than in old‐growth areas, reflecting decreased nutrient availability. However, foliar nutrient concentrations and specific leaf area were greatest in relatively short patches in logged areas, reflecting a shift in composition to pioneer species with acquisitive traits. N:P ratio increased in logged forest, suggesting reduced soil P availability through disturbance. Through the first landscape scale assessment of how functional leaf traits change in response to logging, we find that differences from old‐growth forest become more pronounced as logged forests increase in stature over time, suggesting exacerbated phosphorus limitation as forests recover.