Phyllosphere nitrogen relations: reciprocal transfer of nitrogen between epiphyllous liverworts and host plants in the understorey of a lowland tropical wet forest in Costa Rica

Epiphyllous bryophytes on tropical rainforest plants acquire nutrients from throughfall and free-living N2-fixing organisms, but may also depend directly on host leaf leachates. By contrast, after drying events bryophytes lose significant quantities of nutrients through leaching that can be taken up by host leaves. To assess a potential nutritional interdependency, nitrogen fluxes between epiphyllous liverworts and their host leaves (Carludovica drudei, Costus laevis, Dieffenbachia concinna, Pentagonia wendlandii) were quantified by in situ15N-labelling techniques in a lowland rainforest, Piedras Blancas National Park, Costa Rica. Depending on host species, epiphyllous bryophytes met between 1 and 57% of their N demand from host leaf leachates. Externally supplied 15N was taken up both by epiphylls and host leaves, but N from epiphyll leachates accounted for < 2.5% of host leaf N after 14 d. Long-term observations (180 d) demonstrated the highly dynamic nature of phyllosphere N of the investigated tropical rainforest understorey and an intermittent sink capacity of epiphyllous bryophytes.     

Functional differences between woodland savannas and seasonally dry forests from south‐eastern Brazil: Evidence from 15N natural abundance studies

Nitrogen availability and N‐cycling dynamics across ecosystems play a critical role in plant functioning and species distribution. Measurements of ¹⁵N natural abundance provides a way to assess ecosystem N dynamics, and the range of nitrogen stable isotope values (δ¹⁵N) for plants in an ecosystem can indicate divergent strategies for N uptake. We tested the hypotheses that the N‐rich seasonally dry forest would have higher soil and leaf δ¹⁵N and a smaller range of leaf δ¹⁵N values compared to the N‐poor cerradão (savanna woodland). We measured N concentration and δ¹⁵N in two soil depths and leaves of 27 woody species in cerradão and 26 in seasonally dry forest. As expected, total soil N concentration decreased while soil δ¹⁵N value increased with soil depth. Regardless of soil depth, seasonally dry forest soils had higher δ¹⁵N and total N concentration compared to cerradão soils. Foliar δ¹⁵N values varied from ?6.4‰ to 5.9‰ in cerradão and from ?2.3‰ to 8.4‰ in seasonally dry forest plants. Phylogenetically independent contrasts analysis and comparisons of δ¹⁵N mean values of the most abundant species and species co‐occurring in both sites confirmed the hypothesis of higher δ¹⁵N for seasonally dry forest in comparison to cerradão. These results corroborate the expectation of higher soil and leaf δ¹⁵N values in sites with higher soil N availability. However, except for the most abundant species, no across‐site leaf–soil (δ¹⁵N leaf –δ¹⁵N soil) differences (Δδ¹⁵N) were found suggesting that differences in leaf δ¹⁵N between cerradão and seasonally dry forest are driven by differences in soil δ¹⁵N. Variation of leaf δ¹⁵N was large in both sites and only slightly higher in cerradão, suggesting high diversity of N use strategies for both cerradão and seasonally dry forest communities.     

Environmental and species‐specific controls on δ13C and δ15N in dominant woody plants from central‐western Argentinian drylands

Spatial variation in mean annual precipitation is the principal driver of plant water and nitrogen status in drylands. The natural abundance of carbon stable isotopes (δ¹³C) in photosynthetic tissues of C3 plants is an indicator of time‐integrated behaviour of stomatal conductance; while that of nitrogen stable isotopes (δ¹⁵N) is an indicator of the main source of plant N (soil N vs. atmospheric N₂). Previous studies in drylands have documented that plant δ¹³C and δ¹⁵N values increase with decreasing mean annual precipitation due to reductions in stomatal conductance, and soil enriched in ¹⁵N, respectively. However, evidence for this comes from studies focused on stable isotopes measurements integrated at the plant community level or on dominant plants at the site level, but little effort has been made to study C and N isotope variations within a species growing along rainfall gradients. We analysed plant δ¹³C, δ¹⁵N and C/N values of three woody species having different phenological leaf traits (deciduous, perennial and aphyllous) along a regional mean annual precipitation gradient from the central‐western Argentinian drylands. Noticeably, plant δ¹³C and δ¹⁵N values in the three woody species did not increase towards sites with low precipitation or at the start of the growing season (drier period), as we expected. These results suggest that environmental factors other than mean annual precipitation may be affecting plant δ¹³C and δ¹⁵N. The short‐term environmental conditions may interact with species‐specific plant traits related to water and nitrogen use strategies and override the predictive influence of the mean annual precipitation on plant δ¹³C and δ¹⁵N widely reported in drylands.     

Temporal dynamics of bio-available Si fluxes in a temperate forested catchment (Meerdaal forest, Belgium)

Stable isotopes of nitrogen (N) in plants are increasingly used to evaluate ecosystem N cycling patterns. A basic assumption in this research is that plant δ¹⁵N reflects the δ¹⁵N of the N source. Recent evidence suggests that plants may fractionate on uptake, transport, or transformation of N. If the dominant source of plant N is via roots, a difference in δ¹⁵N by tissue type would suggest fractionation on transport and assimilation of N. In order to evaluate differences between species and plant parts, we measured δ¹⁵N in root, stem, and leaf tissues of individual sugar maple (Acer saccharum; SM) and American beech (Fagus grandifolia; BE) plants ranging in age from germinants to mature trees at the Hubbard Brook Experimental Forest, New Hampshire (USA). For SM, root δ¹⁵N > stem δ¹⁵N > leaf δ¹⁵N; for BE seedlings, root δ¹⁵N > stem δ¹⁵N and root δ¹⁵N > leaf δ¹⁵N. These differences suggest that fractionation occurs during plant transport and assimilation of N. Beech δ¹⁵N (root, stem, and leaf) was consistently higher than SM δ¹⁵N for 1–7 year-old seedlings. At one site, we found no differences with age in foliar δ¹⁵N (range: 4.1–4.8 ‰) for seedlings, saplings, and trees which suggests that it may be possible to compare foliar δ¹⁵N of plants of different ages at some sites. However, at another site, foliar and root δ¹⁵N were higher for trees than 1–2 year-old seedlings. This study suggests that physiological differences in N assimilation and transport processes that differ by species likely control plant δ¹⁵N.     

15N natural abundance of plant and soil components of a Banksia woodland ecosystem in relation to nitrate utilization, life form, mycorrhizal status and N2-fixing abilities of component species

Studies of the variation in δ¹⁵N values for plants from a fire-prone Banksia woodland in South West Australia showed that pioneer herbaceous, non-mycorrhizal species which were active in nitrate reduction and storage, had the highest values (1.81%c). A detailed study of one such species Ptilotus polystachus demonstrated a close correspondence between the δ¹⁵N values of soil nitrate, xylem nitrate and leaf total nitrogen, suggesting an exclusive reliance on nitrate ions as nitrogen source. These pioneer species also showed a preponderance of the chloroplastic isoform of glutamine synthetase while woody species generally had higher activity associated with the cytosolic isoform. The group comprising monocotyledonous hemicryptophytes and geophytes contained species with slightly positive δ¹⁵N values and moderately active in nitrate reduction and storage. Nitrogen-fixing species had the lowest δ¹⁵N values (–0.36‰), irrespective of their apparent utilisation of nitrate. However, woody resprouter species which had low levels of nitrate reduction and storage had δ¹⁵N values which fell within the range of values obtained for the miscellaneous assemblage of N₂-fixing species. Consequently, ¹⁵N abundance values failed to distinguish N₂ fixing from non-fixing woody species, and therefore, could not be used in the ecosystem to determine the dependence of putative nitrogen fixing species on N₂ fixation. The study demonstrated complex patterns of nitrogen utilization in the ecosystem in which exploitation of different nitrogen resources related to plant life form and the physiological attributes of nitrogen assimilation by component species.     

Carbon, nitrogen, and hydrogen isotope ratios in creekside trees in western Kansas

Three species of creekside trees were monitored weekly during the 2007 and 2008 growing seasons. The 2007 growing season was wet early, but became drier as the season progressed. In contrast, the 2008 growing season was dry early, but became wetter as the season progressed. Creekside trees were measured to determine effects of changing water regimes on leaf-level processes. Lonicera tatarica plants were compared to Morus alba and Celtis occidentalis trees. Leaves were monitored for changes in stomatal conductance, transpiration, δ¹³C, δ¹⁵N, δD, leaf temperature, and heat losses via latent, sensible, and radiative pathways. δD of creek water was more similar to ground water than to rain water, but the creek was partially influenced by summer rains. δD of bulk leaf material was significantly higher in individuals of C. occidentalis compared to the other species, consistent with source water coming from shallower soil layers. Despite decreasing water levels, none of these tree species showed signs of water stress. There were no significant differences between species in stomatal conductance or transpiration. Leaf δ¹³C was significantly lower in individuals of L. tatarica compared to the other species. Differences in δ¹³C were attributed to a lower carboxylation capacity, consistent with lower leaf nitrogen content in L. tatarica plants. Leaf δ¹⁵N was significantly lower in individuals of L. tatarica compared to the other species, consistent with uptake of a different N source. Two of the three sites appeared to be affected by inorganic N from fertilizer run-off. Evidence is presented that these species acquired water and nitrogen from different sources, resulting from differences in root uptake patterns.     

Shifts in bryophyte carbon isotope ratio across an elevation?��?soil age matrix on Mauna Loa, Hawaii: do bryophytes behave like vascular plants?

The carbon isotope ratio (δ(13)C) of vascular plant leaf tissue is determined by isotope discrimination, primarily mediated by stomatal and mesophyll diffusion resistances and by photosynthetic rate. These effects lead to predictable trends in leaf δ(13)C across natural gradients of elevation, irradiance and nutrient supply. Less is known about shifts in δ(13)C for bryophytes at landscape scale, as bryophytes lack stomata in the dominant gametophyte phase, and thus lack active control over CO(2) diffusion. Twelve bryophyte species were sampled across a matrix of elevation and soil ages on Mauna Loa, Hawaii Island. We tested hypotheses based on previous findings for vascular plants, which tend to have less negative δ(13)C at higher elevations or irradiances, and for leaves with higher leaf mass per area (LMA). Across the matrix, bryophytes spanned the range of δ(13)C values typical of C(3) vascular plants. Bryophytes were remarkably similar to vascular plants in exhibiting less negative δ(13)C with increasing elevation, and with lower overstory cover; additionally δ(13)C was related to bryophyte canopy projected mass per area, a trait analogous to LMA in vascular plants, also correlated negatively with overstory cover. The similarity of responses of δ(13)C in bryophytes and vascular plants to environmental factors, despite differing morphologies and diffusion pathways, points to a strong direct role of photosynthetic rate in determining δ(13)C variation at the landscape scale.     

Stable nitrogen isotope patterns of trees and soils altered by long-term nitrogen and phosphorus addition to a lowland tropical rainforest

Foliar nitrogen (N) isotope ratios (δ¹⁵N) are used as a proxy for N-cycling processes, including the “openness” of the N cycle and the use of distinct N sources, but there is little experimental support for such proxies in lowland tropical forest. To address this, we examined the δ¹⁵N values of soluble soil N and canopy foliage of four tree species after 13 years of factorial N and P addition to a mature lowland rainforest. We hypothesized that N addition would lead to ¹⁵N-enriched soil N forms due to fractionating losses, whereas P addition would reduce N losses as the plants and microbes adjusted their stoichiometric demands. Chronic N addition increased the concentration and δ¹⁵N value of soil nitrate and δ¹⁵N in live and senesced leaves in two of four tree species, but did not affect ammonium or dissolved organic N. Phosphorus addition significantly increased foliar δ¹⁵N in one tree species and elicited significant N × P interactions in two others due to a reduction in foliar δ¹⁵N enrichment under N and P co-addition. Isotope mixing models indicated that three of four tree species increased their use of nitrate relative to ammonium following N addition, supporting the expectation that tropical trees use the most available form of mineral N. Previous observations that anthropogenic N deposition in this tropical region have led to increasing foliar δ¹⁵N values over decadal time-scales is now mechanistically linked to greater usage of ¹⁵N-enriched nitrate.     

Caribbean octocorals record changing carbon and nitrogen sources from 1862 to 2005

During the last century, the global biogeochemical cycles of carbon (C) and nitrogen (N) have been drastically altered by human activities. A century of land‐clearing and biomass burning, followed by fossil fuel combustion have increased the concentration of atmospheric CO₂ by approximately 20%, and since the mid‐1900s, the use of agricultural fertilizers has been the primary driver of an approximate 90% increase in bioavailable N. Geochemical records obtained through stable isotope analysis of terrestrial and marine biota effectively illustrate rising anthropogenic C inputs. However, there are fewer records of anthropogenic N, despite the enormous magnitude of change and the known negative effects of N on ecosystem health. We used stable isotope values from independent octocorals (gorgonians) sampled across the Western Atlantic over the last 143 years to document human perturbations of the marine C and N pools. Here, we demonstrate that in sea plumes δ¹³C values and in both sea plumes and sea fans δ¹⁵N values declined significantly from 1862 to 2005. Sea plume δ ¹³C values were negatively correlated with increasing atmospheric CO₂ concentrations and corroborate known rates of change resulting from global fossil fuel combustion, known as the Suess effect. We suggest that widespread input of agricultural fertilizers to near‐shore coastal waters is the dominant driver for the decreasing δ ¹⁵N trend, though multiple anthropogenic sources are likely affecting this trend. Given the interest in using δ ¹⁵N as an indicator for N pollution in aquatic systems, we highlight the risk of underestimating contributions of pollutants as a result of source mixing as demonstrated by a simple isotope‐mixing model. We conclude that signals of major human‐induced perturbations of the C and N pools are detectable in specimens collected over wide geographic scales, and that archived materials are invaluable for establishing baselines against which we can assess environmental change.     

Evaluating the consequences of salmon nutrients for riparian organisms: Linking condition metrics to stable isotopes

Stable isotope ratios (δ¹³C and δ¹⁵N) have been used extensively to trace nutrients from Pacific salmon, but salmon transfer more than carbon and nitrogen to stream ecosystems, such as phosphorus, minerals, proteins, and lipids. To examine the importance of these nutrients, metrics other than isotopes need to be considered, particularly when so few studies have made direct links between these nutrients and how they affect riparian organisms. Our study specifically examined δ¹³C and δ¹⁵N of riparian organisms from salmon and non‐salmon streams in Idaho, USA, at different distances from the streams, and examined whether the quality of riparian plants and the body condition of invertebrates varied with access to these nutrients. Overall, quality and condition metrics did not mirror stable isotope patterns. Most notably, all riparian organisms exhibited elevated δ¹⁵N in salmon streams, but also with proximity to both stream types suggesting that both salmon and landscape factors may affect δ¹⁵N. The amount of nitrogen incorporated from Pacific salmon was low for all organisms (<20%) and did not correlate with measures of quality or condition, probably due to elevated δ¹⁵N at salmon streams reflecting historical salmon runs instead of current contributions. Salmon runs in these Idaho streams have been declining, and associated riparian ecosystems have probably seen about a 90% reduction in salmon‐derived nitrogen since the 1950s. In addition, our results support those of other studies that have cautioned that inferences from natural abundance isotope data, particularly in conjunction with mixing models for salmon‐derived nutrient percentage estimates, may be confounded by biogeochemical transformations of nitrogen, physiological processes, and even historical legacies of nitrogen sources. Critically, studies should move beyond simply describing isotopic patterns to focusing on the consequences of salmon‐derived nutrients by quantifying the condition and fitness of organisms putatively using those resources.     

Carbon and nitrogen stable isotope ratios of macroinvertebrates in the littoral zone of Lake Biwa as indicators of anthropogenic activities in the watershed

Carbon and nitrogen stable isotope ratios (δ¹³C and δ¹⁵N) of macroinvertebrates inhabiting littoral zones of lakes can serve as useful indicators of material loading from the watershed. We collected snails (Semisulcospira spp.) and bivalves (Unio douglasiae biwae Kobelt) from 29 littoral sites in Lake Biwa near the mouths of river tributaries with various human population density (HPD) and land-use patterns. The δ¹³C and δ¹⁵N signatures were determined for three potential food sources: particulate organic matter in the pelagic zone (PPOM), riverine particulate organic matter from tributaries (RPOM) and epilithic organic matter in the littoral zone (EOM). The stable isotope mixing model revealed that snails relied mainly on EOM, and bivalves on PPOM and RPOM. Multiple regression analysis showed that intersite variation in δ¹⁵N for snails was best explained by HPD, while variation in δ¹⁵N of EOM and nitrate was explained to a lesser extent by HPD. Comparison with isotope signatures of their food sources and riverine nutrients revealed that snails assimilated anthropogenic nitrogen from wastewater in the watershed. Our results also showed that the δ¹³C value of bivalves was marginally related to the fraction of paddy fields in the watersheds. In conclusion, the isotope signatures of macroinvertebrates inhabiting the littoral zone can be useful indicators of anthropogenic impacts from the watershed.     

Variation in leaf and soil δ<Superscript>15</Superscript>N in diverse tree species in a lowland dipterocarp rainforest,Malaysia

Key message

Large variations in leaf δ ¹⁵ N in Bornean tropical rainforest trees may indicate that various tropical species have species-specific strategy for nitrogen uptake under low soil nutrient conditions, including root symbiotic microorganisms such as ectomycorrhiza.

Abstract

Lowland tropical rainforests in Southeast Asia are characterized by high species diversity despite limited soil nutrient conditions. The plant nitrogen isotope ratio (δ¹⁵N) reflects plant uptake of soil nitrogen. We analyzed δ¹⁵N values and nitrogen content (N %) in leaves and roots of 108 woody species with different types of symbiotic microorganisms, of different life forms (emergent, canopy, sub-canopy, understory, and canopy gap species), and from different families in a Bornean lowland dipterocarp forest to gain more insight into the diversity of nitrogen uptake strategy in the rhizosphere. Leaf δ¹⁵N values in the species studied varied largely from ?7.2 to 5.0 ‰, which is comparable to the values of known Asian trees including temperate, sub-tropical, and tropical mountain forests. Leaf δ¹⁵N also varied significantly among both life forms and families, though the phylogenetically independent contrast (PIC) relationships were not statistically significant among life form, family, and symbiotic types. Some families showed specific leaf δ¹⁵N values; Dipterocarpaceae, the dominant family in the canopy layer with symbiotic ectomycorrhiza in Southeast Asia, had small intraspecific variation and higher leaf δ¹⁵N values (0.03 ‰) compared with species exhibiting arbuscular mycorrhiza, whereas several families such as Burseraceae, Euphorbiaceae, and Myrtaceae showed large interspecific variation in leaf δ¹⁵N (e.g., from ?7.2 to 5.0 ‰ in Euphorbiaceae). These variations suggest that tropical species may have family- or species-specific strategy, such as root symbiotic microorganisms, for nitrogen uptake under low-nutrient conditions in tropical rainforests in Southeast Asia.

     

Nitrogen stable isotopes indicate differences in nitrogen cycling between two contrasting Jamaican montane forests

Background and aims

The aim of this study is to enhance our knowledge of nitrogen (N) cycling and N acquisition in tropical montane forests through analysis of stable N isotopes (δ¹⁵N).

Methods

Leaves from eight common tree species, leaf litter, soils from three depths and roots were sampled from two contrasting montane forest types in Jamaica (mull ridge and mor ridge) and were analysed for δ¹⁵N.

Results

All foliar δ¹⁵N values were negative and varied among the tree species but were significantly more negative in the mor ridge forest (by about 2 ‰). δ¹⁵N of soils and roots were also more negative in mor ridge forests by about 3 ‰. Foliar δ¹⁵N values were closer to that of soil ammonium than soil nitrate suggesting that trees in these forests may have a preference for ammonium; this may explain the high losses of nitrate from similar tropical montane forests. There was no correlation between the rankings of foliar δ¹⁵N in the two forest types suggesting a changing uptake ratio of different N forms between forest types.

Conclusions

These results indicate that N is found at low concentrations in this ecosystem and that there is a tighter N cycle in the mor ridge forest, confirmed by reduced nitrogen availability and lower rates of nitrification. Overall, soil or root δ¹⁵N values are more useful in assessing ecosystem N cycling patterns as different tree species showed differences in foliar δ¹⁵N between the two forest types.     

Nitrogen isotope signature variability in plant species from open peatland

We report natural abundance of ¹⁵N content in a suite of plants and lichens from a sloping open peatland in coastal northern British Columbia, Canada. A wide range of δ¹⁵N values varying between −11 and +3‰ were observed in the studied species. Herbs, mosses, and shrubs generally grouped individually along the δ¹⁵N gradient from enriched to more depleted values. This grouping probably reflects similar nitrogen acquiring strategies within the group, but some species with different strategies showed deviated δ¹⁵N values from the cluster. Hummock-lawn gradient, which is illustrated as depth to water table, explains the variation in δ¹⁵N signature of cryptogams: the deeper the groundwater table, the more depleted is the ¹⁵N. This trend might be explained by differences in nitrogen sources and/or differences in internal fractionation during nitrogen translocation processes.     

A δ<Superscript>15</Superscript>N assessment of nitrogen deposition for the endangered epiphytic orchid <Emphasis Type="Italic">Laelia speciosa</Emphasis> from a city and an oak forest in Mexico

Atmospheric nitrogen deposition poses a major threat to global biodiversity. Tropical epiphytic plants are especially at risk given their reliance on atmospheric sources of nutrients. The leaf, pseudobulb, and root carbon and nitrogen content, C:N ratio, as well as the nitrogen isotopic composition were studied for individuals of Laelia speciosa from a city and from an oak forest in Mexico. The nitrogen content of leaves was similar between the city and the oak forest, reaching 1.3 ± 0.2 % (dry mass). The δ¹⁵N of leaves, pseudobulbs, and roots reached 5.6 ± 0.2 ‰ in the city, values found in sites exposed to industrial and vehicular activities. The δ¹⁵N for plant from the oak forest amounted to –3.1 ± 0.3 ‰, which is similar to values measured from sites with low industrial activities. Some orchids such as Laelia speciosa produce a single pseudobulb per year, i.e., a water and nutrient storage organ, so the interannual nitrogen deposition was studied by considering the ten most recent pseudobulbs for plants from either site formed between 2003 and 2012. The C:N ratio of the ten most recent pseudobulbs from the oak forest, as well as that of the pseudobulbs formed before 2010 for plants in the city were indistinguishable from each other, averaging 132.4 ± 6.5, while it was lower for the two most recent pseudobulbs in the city. The δ¹⁵N values of pseudobulbs from the oak forest averaged ?4.4 ± 0.1 ‰ for the entire series. The δ¹⁵N ranged from 0.1 ± 1.6 ‰ for the oldest pseudobulb to 4.7 ± 0.2 ‰ for the pseudobulb formed in the city from 2008 onwards. Isotopic analysis and the C:N ratio for L. speciosa revealed that rates of nitrogen deposition were higher in the city than in the forest. The δ¹⁵N values of series of pseudobulbs showed that it is possible to track nitrogen deposition over multiple years.     

Nitrogen isotope composition of tomato (Lycopersicon esculentum Mill. cv. T-5) grown under ammonium or nitrate nutrition

Studies that quantify plant δ¹⁵N often assume that fractionation during nitrogen uptake and intra-plant variation in δ¹⁵N are minimal. We tested both assumptions by growing tomato (Lycopersicon esculetum Mill. cv. T-5) at NH₄⁺ or NO^?₃ concentrations typical of those found in the soil. Fractionation did not occur with uptake; whole-plant δ¹⁵N was not significantly different from source δ¹⁵ N for plants grown on either nitrogen form. No intra-plant variation in δ¹⁵N was observed for plants grown with NH⁺₄. In contrast. δ¹⁵N of leaves was as much as 5.8% greater than that of roots for plants grown with NO^?₃. The contrasting patterns of intra-plant variation are probably caused by different assimilation patterns. NH⁺₄ is assimilated immediately in the root, so organic nitrogen in the shoot and root is the product of a single assimilation event. NO^?₃ assimilation can occur in shoots and roots. Fractionation during assimilation caused the δ¹⁵N of NO^?₃ to become enriched relative to organic nitrogen; the δ¹⁵N of NO^?₃ was 11.1 and 12.9% greater than the δ¹⁵N of organic nitrogen in leaves and roots, respectively. Leaf δ¹⁵N may therefore be greater than that of roots because the NO^?₃ available for assimilation in leaves originates from a NO^?₃ pool that was previously exposed to nitrate assimilation in the root.     

The “isohydric trap”: A proposed feedback between water shortage,stomatal regulation,and nutrient acquisition drives differential growth and survival of European pines under climatic dryness

Climatic dryness imposes limitations on vascular plant growth by reducing stomatal conductance, thereby decreasing CO₂ uptake and transpiration. Given that transpiration‐driven water flow is required for nutrient uptake, climatic stress‐induced nutrient deficit could be a key mechanism for decreased plant performance under prolonged drought. We propose the existence of an “isohydric trap,” a dryness‐induced detrimental feedback leading to nutrient deficit and stoichiometry imbalance in strict isohydric species. We tested this framework in a common garden experiment with 840 individuals of four ecologically contrasting European pines (Pinus halepensis, P. nigra, P. sylvestris, and P. uncinata) at a site with high temperature and low soil water availability. We measured growth, survival, photochemical efficiency, stem water potentials, leaf isotopic composition (δ¹³C, δ¹⁸O), and nutrient concentrations (C, N, P, K, Zn, Cu). After 2 years, the Mediterranean species Pinus halepensis showed lower δ¹⁸O and higher δ¹³C values than the other species, indicating higher time‐integrated transpiration and water‐use efficiency (WUE), along with lower predawn and midday water potentials, higher photochemical efficiency, higher leaf P, and K concentrations, more balanced N:P and N:K ratios, and much greater dry‐biomass (up to 63‐fold) and survival (100%). Conversely, the more mesic mountain pine species showed higher leaf δ¹⁸O and lower δ¹³C, indicating lower transpiration and WUE, higher water potentials, severe P and K deficiencies and N:P and N:K imbalances, and poorer photochemical efficiency, growth, and survival. These results support our hypothesis that vascular plant species with tight stomatal regulation of transpiration can become trapped in a feedback cycle of nutrient deficit and imbalance that exacerbates the detrimental impacts of climatic dryness on performance. This overlooked feedback mechanism may hamper the ability of isohydric species to respond to ongoing global change, by aggravating the interactive impacts of stoichiometric imbalance and water stress caused by anthropogenic N deposition and hotter droughts, respectively.     

Waterlogging and fire impacts on nitrogen availability and utilization in a subtropical wet heathland (wallum)

Protein, amino acids and ammonium were the main forms of soluble soil nitrogen in the soil solution of a subtropical heathland (wallum). After fire, soil ammonium and nitrate increased 90- and 60-fold, respectively. Despite this increase in nitrate availability after fire, wallum species exhibited uniformly low nitrate reductase activities and low leaf and xylem nitrate. During waterlogging soil amino acids increased, particularly γ-aminobutyric acid (GABA) which accounted for over 50% of amino nitrogen. Non-mycorrhizal wallum species were significantly (P < 0.05) ¹⁵N-enriched (0.3–4.3‰) compared to species with mycorrhizal associations (ericoid-type, ecto-, va-mycorrhizal) which were strongly depleted in ¹⁵N (-6.3 to -1.8‰). Lignotubers and roots had δ¹⁵N signatures similar to that of the leaves of respective species. The exceptions were fine roots of ecto-, ecto/va-, and ericoid type mycorrhizal species which were enriched in ¹⁵N (0.1–2.4‰). The 5¹⁵N signatures of δ¹⁵N_{total soil N} and δ¹⁵N_{soil NH4+} were in the range 3.7–4.5‰, whereas δ¹⁵N_{soil NO3?} was significantly (P < 0.05) more enriched in ¹⁵N (9.2–9.8‰). It is proposed that there is discrimination against ¹⁵N during transfer of nitrogen from fungal to plant partner. Roots of selected species incorporated nitrogen sources in the order of preference: ammonium > glycine > nitrate. The exception were proteoid roots of Hakea (Proteaceae) which incorporated equal amounts of glycine and ammonium.     

Do grazers alter nitrogen dynamics on grazing lawns in a South African savannah?

The presence of grazers on grazing lawns in East Africa and North America often alters nitrogen cycling and availability. Grazing lawns can be defined as areas where grasses are kept in a short, actively growing, palatable state by the action of grazers. Our aim was to test whether lawns have enhanced leaf nitrogen (N) concentrations, total soil N and δ¹⁵N when compared to tall grass areas in a South African savannah. Previous studies have used ecosystem δ¹⁵N as a proxy of N availability, and enriched δ¹⁵N values have been suggested to indicate higher N availability or higher N transformation rates. Across all sites, foliar N concentrations (but not soil N) were higher when compared to tall grass areas, and evidence of enriched foliar and soil δ¹⁵N values was found on the lawns. These results suggest that grazers may be involved in altering the rates of N transformations directly on grazing lawns. Regardless of whether these N transformations included increased net N mineralization, higher N concentrations in above‐ground foliage attract grazers back to the lawns, encouraging their maintenance.     

Reconstructing terrestrial nutrient cycling using stable nitrogen isotopes in wood

Although recent anthropogenic effects on the global nitrogen (N) cycle have been significant, the consequences of increased anthropogenic N on terrestrial ecosystems are unclear. Studies of the impact of increased reactive N on forest ecosystems—impacts on hydrologic and gaseous loss pathways, retention capacity, and even net primary productivity—have been particularly limited by a lack of long-term baseline biogeochemical data. Stable nitrogen isotope analysis (ratio of ¹⁵N to ¹⁴N, termed δ¹⁵N) of wood chronologies offers the potential to address changes in ecosystem N cycling on millennial timescales and across broad geographic regions. Currently, nearly 50 studies have been published utilizing wood δ¹⁵N records; however, there are significant differences in study design and data interpretation. Here, we identify four categories of wood δ¹⁵N studies, summarize the common themes and primary findings of each category, identify gaps in the spatial and temporal scope of current wood δ¹⁵N chronologies, and synthesize methodological frameworks for future research by presenting eight suggestions for common methodological approaches and enhanced integration across studies. Wood δ¹⁵N records have the potential to provide valuable information for interpreting modern biogeochemical cycling. This review serves to advance the utility of this technique for long-term biogeochemical reconstructions.