Flood events overrule fertiliser effects on biomass production and species richness in riverine grasslands

Question: Do severe winter flood events lift the nutrient limitation of biomass production in a river floodplain? How does this affect plant species richness? How long do the effects last? Location: Floodplain grassland on calcareous sandy loam near river Rhine in The Netherlands. Methods: Plots were fertilised with four treatments (control, N, P, N+P) for 21 years; plant species composition, vegetation biomass and tissue nutrient concentrations were determined every year between 1985 and 2005. Results: Fertilisation with N generally increased biomass production and reduced species richness, but these effects varied over time. During the first four years of the experiment, biomass production appeared to be co‐limited by N and P, while N fertilisation dramatically reduced plant species richness; these effects became weaker subsequently. Following two extreme winter floods in 1993–94 and 1994–95 and a drought in spring 1996, the effects of fertilisation disappeared between 1998 and 2001 and then appeared again. Flooding caused an overall reduction in species richness (from c. 24 to 15 species m^‐2) and an increase in biomass production, which were only partly reversed after ten years. Conclusions: Long time series are necessary to understand vegetation dynamics and nutrient limitation in river floodplains, since they are influenced by occasional flood and drought events, whose effects may persist for more than ten years. A future increase in flooding frequency might be detrimental to species richness in floodplain grasslands.     

Nutrient limitation and botanical diversity in wetlands: can fertilisation raise species richness?

The 'resource balance hypothesis' proposes that the species richness of grassland vegetation is potentially highest when the N:P ratio of plant tissues is 10–15 (co-limitation), so that species richness could be raised by fertilisation with N or P at sites with lower or higher N:P ratios, respectively. Here we use data from field surveys in Swiss, Dutch and American fens or wet grasslands to analyse what changes in N:P ratios might produce noticeable changes in species richness. Plant species numbers, above-ground biomass, tissue N and P concentrations and soil pH were recorded in plots of 0.06–4 m². In each data set, plots with intermediate tissue N:P ratios (6–20) were on average most species-rich, but N:P ratios explained only 5–37% of the variation in species richness. Moreover, these effects were partially confounded with those of vegetation biomass and/or soil pH. The unique effects of N:P ratios (excluding those shared with biomass and pH) explained 11–17% of variation in species richness. The relationship between species richness and N:P ratios was asymmetric: plots with high N:P ratios were more species-poor than those with low N:P ratios. This was paralleled by a smaller species pool size at high N:P ratios (estimated from species numbers in multiple records), suggesting that fewer species are adapted to P-limited conditions than to N-limited conditions. According to these data, species richness in wetlands may possibly be raised by P-fertilisation when the initial N:P ratio of the vegetation is well above 20, but this option is not recommended for nature conservation as it might promote common species at the expense of rare ones.     

Global changes in soil stocks of carbon,nitrogen, phosphorus,and sulphur as influenced by long‐term agricultural production

Quantifying changes in stocks of C, N, P, and S in agricultural soils is important not only for managing these soils sustainably as required to feed a growing human population, but for C and N, they are also important for understanding fluxes of greenhouse gases from the soil environment. In a global meta‐analysis, 102 studies were examined to investigate changes in soil stocks of organic C, total N, total P, and total S associated with long‐term land‐use changes. Conversion of native vegetation to cropping resulted in substantial losses of C (?1.6 kg m^?2, ?43%), N (?0.15 kg m^?2, ?42%), P (?0.029 kg m^?2, ?27%), and S (?0.015 kg m^?2, ?33%). The subsequent conversion of conventional cropping systems to no‐till, organic agriculture, or organic amendment systems subsequently increased stocks, but the magnitude of this increase (average of +0.47 kg m^?2 for C and +0.051 kg m^?2 for N) was small relative to the initial decrease. We also examined the conversion of native vegetation to pasture, with changes in C (?11%), N (+4.1%), and P (+25%) generally being modest relative to changes caused by conversion to cropping. The C:N ratio remained relatively constant irrespective of changes in land use, whilst in contrast, the C:S ratio decreased by 21% in soils converted to cropping – this suggesting that biochemical mineralization is of importance for S. The data presented here will assist in the assessment of different agricultural production systems on soil stocks of C, N, P, and S – this information assisting not only in quantifying the effects of existing agricultural production on these stocks, but also allowing for informed decision‐making regarding the potential effects of future land‐use changes.     

青藏高原高寒草甸植被和土壤对短期禁牧的响应

为探讨青藏高原高寒草甸对短期禁牧的响应,设置冬季自由放牧和短期(2年)禁牧的对比试验。采用随机样方法调查植被群落盖度,分析地上和地下生物量、根冠比、植被地上和地下部分以及表层(0-10cm)土壤全碳、全氮和全磷含量、生态化学计量以及营养元素的关联性。研究结果显示:1)短期禁牧显著改变高寒草甸植被盖度、地上生物量、根冠比、植被全磷含量和N∶P,以及土壤全磷含量。2)相关性分析表明,禁牧后土壤全碳含量与植被地上全碳含量呈显著相关性,自由放牧后土壤全碳和全氮含量分别与植被地下部分全碳和全氮含量呈显著相关性。结果表明,不同的草原管理措施(禁牧、放牧)会改变高寒草甸植被与土壤养分分配及其平衡关系,同时,植被与表层土壤主要养分含量之间的关联性仅存在于部分植物器官与部分营养元素之间。     

Calcium and iron as key drivers of brown moss composition through differential effects on phosphorus availability

Brown moss-dominated rich fens are characterized by minerotrophic conditions, in which calcium (Ca) and iron (Fe) concentrations show large variations. We examined the relative importance of Ca and Fe in relation to the occurrence of three typical brown moss species: Scorpidium scorpioides, Scorpidium cossonii, and Hamatocaulis vernicosus. Peat chemistry was examined in 24 stands of brown moss-dominated rich fens: 12 in the Netherlands and 12 in central Sweden. Ca and Fe turned out to be important drivers of brown moss composition. Fens dominated by Scorpidium scorpioides or Scorpidium cossonii were characterized by high pore water Ca-concentrations and total soil Ca-contents, but low P-availability. In these Ca-rich, but Fe-poor fens, foliar N:P ratios of vascular vegetation exceeded 20?g?g^?1, indicating phosphorus (P)-limitation due to Ca-P precipitation or low P-sorption capacity due to low Fe-levels. In contrast, fens dominated by Hamatocaulis vernicosus were characterized by high pore water Fe-concentrations and total soil Fe-contents, but also relatively high P-availability. N:P ratios in these fens were below 13.5?g?g^?1, indicating potential nitrogen (N)-limitation. We conclude that the relative roles of Ca and Fe, as related to the geohydrological conditions present, strongly determine the brown moss composition in rich fens through their differential effects on plant P-availability.     

Plant functional traits along environmental gradients in seasonally dry and fire‐prone ecosystem

Questions: How does the abundance and richness of plant assemblages with different functional (regeneration and nutrient acquisition) traits vary with fire regime, moisture availability and substrate fertility? What is the role of different functional traits in maintaining plant diversity under changing environmental conditions in seasonally dry and fire‐prone environments? Location: Southwest Western Australia. Methods: Plant species abundance and soil nutrients were determined at 16 forest sites with variable fire histories across an aridity gradient. All plant species were classified based on their functional traits as (1) perennial or annual, (2) ectomycorrhizal, arbuscular mycorrhizal, ericoid mycorrhizal, orchid mycorrhizal, proteoid or other non‐mycorrhizal, (3) resprouters or seeder, and (4) nitrogen fixer or non‐fixer. We used a multivariate (fourth‐corner) technique to simultaneously test the significance and direction of the relationship between each of these traits and fire frequency, fire interval length, aridity, and soil N, P and C fractions. Results: The functional response of the vegetation to fire regime was minor and restricted to annual species, which comprised only ～4% of taxa. Proteoid and ectomycorrhizal species dominated over species with arbuscular and orchid mycorrhizal roots, N‐fixers dominated over non‐fixers, and seeders dominated over resprouters when N fertility was low but organic labile P was high. Further, proteoid and ectomycorrhizal species richness increased with aridity, while arbuscular mycorrhizal species richness decreased. Conclusions: While the functional composition of southwest Australian vegetation is largely insensitive to changes in fire regime, nutrient acquisition and, to a lesser extent, regeneration traits provide mechanisms for the vegetation community to adjust to changes in resource availability. Thus, diversity responses to environmental change in seasonally dry and fire‐prone ecosystems are likely to be primarily mediated by the composition of nutrient acquisition traits in the vegetation community.     

The importance of nitrogen-fixation for an invader of a coastal California grassland

Whether a novel trait of an invader directly contributes to increased establishment of that invader is a relatively unstudied question in plant ecology. Nitrogen (N)-fixing shrubs comprise a significant subset of grassland invaders worldwide, which suggests the potential importance of the novel trait of N-fixation in the invasion process. We indirectly tested the importance of N-fixation in the invasion of Genista monspessulana (French broom) in a California grassland by alleviating N and phosphorus (P) limitation to the grassland matrix. Grassland productivity was co-limited by N and P; N alone did not release the resident vegetation, and did not affect Genista performance. Genista was strongly limited by P: seedlings had more nodules, greater leaf N concentration, and higher growth and survival with P additions. When N was added with P, however, growth of the resident vegetation was 50?C70% greater than with N or P alone, accompanied by decreases in Genista performance. This suggests that the advantage conferred to Genista by N-fixation was dampened when the resident vegetation was released from nutrient limitation.     

中国主要湿地植被氮和磷生态化学计量学特征

研究湿地植物氮(N)和磷(P)的生态化学计量学特征对揭示植物与生境的耦合关系具有重要意义。通过收集中国52个采样区湿地植物不同器官和全株样本的N和P含量, 对其进行分类和统计分析, 探讨植物器官、生长期、植物类型、湿地类型和气候带对湿地植物N和P生态化学计量学特征的影响。结果表明: 1)湿地植物各器官N、P和N:P的几何平均值均表现为叶片(N, 16.07 mg·g^-1; P, 1.85 mg·g^-1; N:P, 8.67) >地上部分(N, 13.54 mg·g^-1; P, 1.72 mg·g^-1; N:P, 7.96) >茎(N, 7.86 mg·g^-1; P, 1.71 mg·g^-1; N:P, 4.58); 2)叶片N含量随时间变化呈现“三峰”型变化, 峰值分别出现在5月、7月和9月; 茎的N含量随时间变化表现为“双峰”型, 峰值出现在5月和9月; 成熟期之前, 植物叶片的N:P与N趋同波动, N:P主要受N含量控制; 衰老期N:P受P含量控制。3)湿地类型是影响植物叶片N和P生态化学计量特征的关键因素, N和P含量最高值出现在河流, 最低值出现在沼泽湿地, N:P的变化趋势大致与之相反。4)植物叶片N、P和N:P的几何平均值都表现为热带>温带>亚热带, 但总体差异不显著(p > 0.05)。5)中国大部分湿地植物叶片N:P < 14, 表现为N限制。     

Plant responses to nutrient addition and predictive ability of vegetation N:P ratio in an austral fen

1. Previous studies of the N:P ratio in wetland plants have been carried out in northern hemisphere wetlands where atmospheric nitrogen deposition is higher. There is little research on foliar N:P ratio as a potential indicator of nutrient limitation in vegetation communities in southern hemisphere wetlands. This study aimed to redress this knowledge gap and answer the following questions: how well does the plant tissue nitrogen to phosphorus (N:P) ratio predict wetland plant community nutrient limitation, as indicated by vegetation standing stocks and below-ground biomass, in southern hemisphere fens? Secondly, what are the impacts of realistic upper levels of farm nutrient run-off on natural montane fen vegetation?
2. Low (35 kg ha⁻¹ year⁻¹) and high (70 kg ha⁻¹ year⁻¹) levels of nitrate-N or ammonium-N with and without P (20 kg ha⁻¹ year⁻¹) were added to 81 vegetation plots over a period of 2.75 years. Species composition, plant nutrient status, and above-ground live vegetation standing stocks were assessed after 3 years, and below-ground biomass after 2 years.
3. Plant tissue analysis suggested the community was N limited or N and P co-limited; we found greater standing stocks of vegetation in plots treated with 70 kg ha⁻¹ year⁻¹ ammonium-N, indicating N limitation. No difference between other treatments was found in above-ground standing stocks or below-ground biomass. Plant species cover increased in both high N treatments, consistent with N limitation. These changes in plant species cover were accompanied by significant decreases in species richness in both high N treatments. Native species dominated the vegetation and this was unaffected by nutrient addition (90% cover).
4. This is one of the first studies to test and find support for the N:P ratio in southern hemisphere wetlands. Observed declines in species richness after N fertilisation in an N-limited fen suggests increased N may pose risks to austral wetlands. Responses by plant communities (changes in composition, biomass) to lower levels of nutrient addition may require longer periods of fertilisation to be apparent in slow growing ecosystems.

     

The effects of groundwater discharge,mowing and eutrophication on fen vegetation evaluated over half a century

Questions: Were continued groundwater discharge and mowing regimes sufficient for vegetation preservation from 1944 to 1993? Which has a stronger effect on vegetation development; groundwater discharge or mowing? What is the role of surface water eutrophication as driver of vegetation change? Location: Het Hol, The Netherlands (ca. 92 ha, 52°13′N, 5°05′E). Methods: Hydrology was simulated for the late 1940s, early 1960s and 1987. Vegetation maps (1944, 1960, 1975 and 1993) were compared for biotope cover. Vegetation recordings in 1944 and 1987 were compared. Surface water quality was compared between 1950 and 1987. Which sites were mown was reconstructed from an interview. Effects of periodic mowing and groundwater discharge on vegetation development were tested for correlation. Results: Biotope diversity reduced significantly through decrease of semi‐aquatic and tall‐herb biotopes, and expansion of forest. The quagfen terrestrialization sere nearly disappeared from 1987 recordings, while the reed sere did well concerning abundance and species richness. Several typical (rich) fen species disappeared from recordings, while new species were mostly field margin species. Periodic mowing and discharge combined are correlated with increasing species numbers. The P‐concentration in surface water increased while N‐concentration decreased. Conclusions: Preservation of the reed sere was successful, whereas preservation of the quagfen sere was not. Periodic mowing and discharge stimulate species richness, discharge more so than periodic mowing. But slight eutrophication likely induced a shift from P‐limitation to N‐limitation, which stimulated the reed sere at the expense of the quagfen sere.     

Nitrogen inputs accelerate phosphorus cycling rates across a wide variety of terrestrial ecosystems

? Biologically essential elements--especially nitrogen (N) and phosphorus (P)--constrain plant growth and microbial functioning; however, human activities are drastically altering the magnitude and pattern of such nutrient limitations on land. Here we examine interactions between N and P cycles of P mineralizing enzyme activities (phosphatase enzymes) across a wide variety of terrestrial biomes. ? We synthesized results from 34 separate studies and used meta-analysis to evaluate phosphatase activity with N, P, or N×P fertilization. ? Our results show that N fertilization enhances phosphatase activity, from the tropics to the extra-tropics, both on plant roots and in bulk soils. By contrast, P fertilization strongly suppresses rates of phosphatase activity. ? These results imply that phosphatase enzymes are strongly responsive to changes in local nutrient cycle conditions. We also show that plant phosphatases respond more strongly to fertilization than soil phosphatases. The tight coupling between N and P provides a mechanism for recent observations of N and P co-limitation on land. Moreover, our results suggest that terrestrial plants and microbes can allocate excess N to phosphatase enzymes, thus delaying the onset of single P limitation to plant productivity as can occur via human modifications to the global N cycle.     

Isotopic evidence of partial mycoheterotrophy in the Gentianaceae: Bartonia virginica and Obolaria virginica as case studies

? Premise of the study: An estimated 10% of plant species have evolved to steal C from their symbiotic fungal partners (mycoheterotrophy), and while physiological evidence for full and partial mycoheterotrophy is well developed in the Orchidaceae and Ericaceae, it is lacking for the majority of other mycoheterotrophic taxa. The family Gentianaceae not only contains several lineages of achlorophyllous mycoheterotrophs, but also contains species that are putative partially mycoheterotrophic. The North American genera Bartonia and Obolaria (Gentianaceae) are green but have leaves reduced to scales or foliose bracts and so have ambiguous mycoheterotrophic status. ? Methods: We investigated the natural abundance (13)C and (15)N profiles of both genera along with total N and chlorophyll content and investigated mycorrhizal infection using light microscopy. ? Key results: The shoots of B. virginica were significantly more enriched in (15)N than the surrounding vegetation but not in (13)C. In contrast, the shoots of O. virginica are not enriched in (15)N compared to the surrounding vegetation but were significantly enriched in (13)C. Total N concentrations were significantly higher than the surrounding vegetation in B. virginica, while the collaroid roots of both species were infected by arbuscular mycorrhizal fungi. ? Conclusions: This microscopic evidence coupled with the natural abundance stable isotope profiles strongly suggests that both species are partially mycoheterotrophic. However, differences in the root-shoot stable isotopic patterns relative to surrounding vegetation between B. virginica and O. virginica are suggestive of the utilization of different physiological pathways or extent of commitment to mycoheterotrophic C gain.     

Can tissue element concentration patterns at the individual species level indicate the factors underlying vegetation gradients in wetlands?

Question: Do tissue element concentrations at the individual species level vary along major vegetation gradients in wetlands, and can they indicate environmental conditions? Location: West Carpathians. Methods: Total plant species composition was recorded in plots distributed along a poor to rich gradient within spring fens and along the gradient from fens to wet meadows. Eriophorum angustifolium (Cyperaceae) and three broadleaf dicotyledonous herb species were collected from the vegetation plots. Tissue N, P, K, Ca and Fe concentrations, N:P and N:K ratios of the species were determined. Each variable was correlated with the sample scores along the first two axes of the DCA ordination, which represented the two main vegetation gradients. Results: K and Ca concentrations in a particular species correlated well with the vegetation gradients, thus indicating changes in the element availability to the species. The trends were sometimes contradictory to known patterns at the community level, but the differences could be ecologically interpreted. Contrary to Ca and K, patterns in N, P and Fe concentration appeared to be more species‐specific. E. angustifolium had a lower K and Ca concentration than the broadleaf herbs. Conclusions: Compared to community‐level measurements, element concentrations in individual species correlated less with observed vegetation gradients. Trends found at the species level may indicate changes in ecological conditions affecting the species, although they need not correspond with trends found at the community level. We conclude that the species‐level approach cannot substitute, but can advance, the community‐level approach in searching for mechanisms underlying vegetation gradients within wetlands.     

Groundwater drawdown drives ecophysiological adjustments of woody vegetation in a semi‐arid coastal ecosystem

Predicted droughts and anthropogenic water use will increase groundwater lowering rates and intensify groundwater limitation, particularly for Mediterranean semi‐arid ecosystems. These hydrological changes may be expected to elicit differential functional responses of vegetation either belowground or aboveground. Yet, our ability to predict the impacts of groundwater changes on these ecosystems is still poor. Thus, we sought to better understand the impact of falling water table on the physiology of woody vegetation. We specifically ask (a) how is woody vegetation ecophysiological performance affected by water table depth during the dry season? and (b) does the vegetation response to increasing depth to groundwater differ among water‐use functional types? We examined a suite of physiological parameters and water‐uptake depths of the dominant, functionally distinct woody vegetation along a water‐table depth gradient in a Mediterranean semi‐arid coastal ecosystem that is currently experiencing anthropogenic groundwater extraction pressure. We found that groundwater drawdown did negatively affect the ecophysiological performance of the woody vegetation. Across all studied environmental factors, depth to groundwater was the most important driver of ecophysiological adjustments. Plant functional types, independent of groundwater dependence, showed consistent declines in water content and generally reduced C and N acquisition with increasing depths to groundwater. Functional types showed distinct operating physiological ranges, but common physiological sensitivity to greater water table depth. Thus, although differences in water‐source use exist, a physiological convergence appeared to happen among different functional types. These results strongly suggest that hydrological drought has an important impact on fundamental physiological processes, constraining the performance of woody vegetation under semi‐arid conditions. By disentangling the functional responses and vulnerability of woody vegetation to groundwater limitation, our study establishes the basis for predicting the physiological responses of woody vegetation in semi‐arid coastal ecosystems to groundwater drawdown.     

Lichen responses to nitrogen and phosphorus additions can be explained by the different symbiont responses

? Responses to simulated nitrogen (N) deposition with or without added phosphorus (P) were investigated for three contrasting lichen species - the N-sensitive Alectoria sarmentosa, the more N-tolerant Platismatia glauca and the N(2) -fixing Lobaria pulmonaria- in a field experiment. ? To examine whether nutrient limitation differed between the photobiont and the mycobiont within the lichen, the biomass responses of the respective bionts were estimated. ? The lichenized algal cells were generally N-limited, because N-stimulated algal growth in all three species. The mycobiont was P-limited in one species (A. sarmentosa), but the growth response of the mycobionts was complex, as fungal growth is also dependent on a reliable carbon export from the photobiont, which may have been the reason for the decrease of the mycobiont with N addition in P. glauca. ? Our findings showed that P availability was an important factor when studying effects of N deposition, as P supply can both mitigate and intensify the negative effects of N on epiphytic lichens.     

Biomass production of the last remaining fen with Saxifraga hirculus in Switzerland is controlled by nitrogen availability

Olde Venterink H. and Vittoz P. 2008. Biomass production of the last remaining fen with Saxifraga hirculus in Switzerland is controlled by nitrogen availability. Bot. Helv. 118: 165 – 174. For conservation management of endangered plants it is important to know which nutrient(s) control growth of the vegetation, because maintenance of low nitrogen (N), phosphorus (P) or potassium (K) availability requires different management measures. The aim of this study was to determine the type of nutrient limitation for the vegetation in the last remaining site with Saxifraga hirculus in Switzerland, using nutrient ratios in the aboveground vegetation as an indicator. We made vegetation relevees, collected biomass of the vascular plants, and took soil samples in three plots at this site. The biomass was very low (152–231 g m ^-2), and all three plots were clearly N-limited with N:P ratios of 7– 8. Soil extractable N concentrations were generally low, and P and K concentrations were moderate to high, which was consistent with the indicated N limitation. Hence conservation management first of all needs to prevent N-enrichment, and needs to avoid increased mineralization rates through drainage, or the accumulation of N in the system from atmospheric deposition. Therefore N output seems required through for instance grazing or mowing. The current grazing management seems to function well, since total aboveground biomass is very low and S. hirculus has a high abundance in this last remnant. Submitted 5 June 2008; Accepted 14 October 2008 Subject editor: Sonja Wipf     

Responses of wetland graminoids to the relative supply of nitrogen and phosphorus

The biomass production of wetland vegetation can be limited by nitrogen or phosphorus. Some species are most abundant in N-limited vegetation, and others in P-limited vegetation, possibly because growth-related traits of these species respond differently to N versus P supply. Two growth experiments were carried out to examine how various morphological and physiological traits respond to the relative supply of N and P, and whether species from sites with contrasting nutrient availability respond differently. In experiment 1, four Carex species were grown in nutrient solutions at five N:P supply ratios (1.7, 5, 15, 45, 135) combined with two levels of supply (geometric means of N and P supply). In experiment 2, two Carex and two grass species were grown in sand at the same .ve N:P supply ratios combined with three levels of supply and two light intensities (45% or 5% daylight). After 12-13 weeks of growth, plant biomass, allocation, leaf area, tissue nutrient concentrations and rates and nutrient uptake depended signi.cantly on the N:P supply ratio, but the type and strength of the responses differed among these traits. The P concentration and the N:P ratio of shoots and roots as well as the rates of N and P uptake were mainly determined by the N:P supply ratio; they showed little or no dependence on the supply level and relatively small interspeci.c variation. By contrast, the N concentration, root mass ratio, leaf dry matter content and speci.c leaf area were only weakly related to the N:P supply ratio; they mainly depended on plant species and light, and partly on overall nutrient supply. Plant biomass was determined by all factors together. Within a level of light and nutrient supply, biomass was generally maximal (i.e. co-limited by N and P) at a N:P supply ratio of 15 or 45. All species responded in a similar way to the N:P supply ratio. In particular, the grass species Phalaris arundinacea and Molinia caerulea showed no differences in response that could clearly explain why P. arundinacea tends to invade P-rich (N-limited) sites, and M. caerulea P-limited sites. This may be due to the short duration of the experiments, which investigated growth and nutrient acquisition but not nutrient con­servation.     

Interactive Effects Between Reindeer and Habitat Fertility Drive Soil Nutrient Availabilities in Arctic Tundra

Herbivores impact nutrient availability and cycling, and the net effect of herbivory on soil nutrients is generally assumed to be positive in nutrient-rich environments and negative in nutrient-poor ones. This is, however, far from a uniform pattern, and there is a recognized need to investigate any interactive effects of herbivory and habitat fertility (i.e., plant C/N ratios) on soil nutrient availabilities. We determined long-term effects of reindeer on soil extractable nitrogen (N) and phosphorus (P) and their net mineralization rates along a fertility gradient of plant carbon (C) to N and P ratios in arctic tundra. Our results showed that reindeer had a positive effect on soil N in the more nutrient-poor sites and a negative effect on soil P in the more nutrient-rich sites, which contrasts from the general consensus. The increase in N availability was linked to a decrease in plant and litter C/N ratios, suggesting that a shift in vegetation composition toward more graminoids favors higher N cycling. Soil P availability was not as closely linked to the vegetation and is likely regulated more by herbivore-induced changes in soil physical and chemical properties. The changes in soil extractable N and P resulted in higher soil N/P ratios, suggesting that reindeer could drive the vegetation toward P-limitation. This research highlights the importance of including both the elements N and P and conducting studies along environmental gradients in order to better understand the interactive effects of herbivory and habitat fertility on nutrient cycling and primary production.     

Mycorrhizal colonisation and P-supplement effects on N uptake and N assimilation in perennial ryegrass under well-watered and drought-stressed conditions

To compare the effect of arbuscular mycorrhiza (AM) and P-supplement on N uptake and N assimilation under well-watered or drought-stressed conditions, Glomus intraradices-colonised, P-supplemented non-mycorrhizal (P) and non-mycorrhizal (control) plants of Lolium perenne were exposed to 12?days of water treatment. Leaf water potential (Ψ (w)), photosynthetic ability, and N and P nutritional status were measured at the beginning (day 0) and end (day 12) of water treatment. N absorption, amino acid and protein synthesis were quantified using the isotopic tracer (15)N at day?12. Under well-watered conditions, growth response and physiological parameters were similar in AM and P plants, as compared to controls. Drought (10% water) significantly decreased these parameters in all three treatments. As compared to control plants, the negative impact of water deficit on the Ψ (w), photosynthesis, biomass, and N and P content was highly alleviated in AM plants, while only slightly improved or remained the same level in P plants. The effect of AM symbiosis on N absorption and N assimilation was greater than that of the P supplement under well-watered and drought-stressed conditions, and this effect was highly enhanced under drought-stressed conditions. At terminal drought stress on day?12, the effect of AM colonisation on de novo synthesis of amino acids and proteins was 4.4- and 4.8-fold higher than that of the P supplement. These results indicate that the AM symbiosis plays an integrative role in N nutrition by alleviating the negative impacts of drought on N or P uptake and N assimilation, whereas the efficiency of a direct P supplement is very limited under drought-stressed conditions.     

Growth in epiphytic bromeliads: response to the relative supply of phosphorus and nitrogen

Insufficient nitrogen (N) and phosphorus (P) frequently limit primary production. Although most nutrient studies on vascular epiphytes have focused on N uptake, circumstantial evidence suggests that P rather than N is the most limiting element for growth in this plant group. We directly tested this by subjecting a total of 162 small individuals of three bromeliad species ( Guzmania monostachia , Tillandsia elongata , Werauhia sanguinolenta ) to three N and three P levels using a full-factorial experimental design, and determined relative growth rates (RGR) and nutrient acquisition over a period of 11 weeks. Both N and P supply had a significant effect on RGR, but only tissue P concentrations were correlated with growth. Uptake rates of N and P, in contrast, were not correlated with RGR. Increased nutrient supply led to an up to sevenfold increase in tissue P concentration compared to natural conditions, while concentrations of N hardly changed or even decreased. All treatment combinations, even at the lowest experimental P supply, led to decreased N:P ratios. We conclude that P is at least as limiting as N for vegetative function under natural conditions in these epiphytic bromeliads. This conclusion is in line with the general notion of the prevalence of P limitation for the functioning of terrestrial vegetation in the tropics.