The scaling of elemental stoichiometry and growth rate over the course of bamboo ontogeny.

Stoichiometric rules may explain the allometric scaling among biological traits and body size, a fundamental law of nature. However, testing the scaling of elemental stoichiometry and growth to size over the course of plant ontogeny is challenging. Here, we used a fast-growing bamboo species to examine how the concentrations and contents of carbon (C), nitrogen (N) and phosphorus (P), relative growth rate (G), and nutrient productivity scale with whole-plant mass (M) at the culm elongation and maturation stages. The whole-plant C content vs M and N content vs P content scaled isometrically, and the N or P content vs M scaled as a general 3/4 power function across both growth stages. The scaling exponents of G vs M and N (and P) productivity in newly grown mass vs M relationships across the whole growth stages decreased as a -1 power function. These findings reveal the previously undocumented generality of stoichiometric allometries over the course of plant ontogeny and provide new insights for understanding the origin of ubiquitous quarter-power scaling laws in the biosphere.

Weak transgenerational effects of ancestral nitrogen and phosphorus availabilities on offspring phenotypes in Arabidopsis thaliana.

Nutrient availability significantly regulates plant growth and metabolic functions, but whether and how the long-term exposure of ancestral plants to contrasting nutrient environments influences offspring phenotypic performance (i.e., transgenerational plasticity) remain poorly addressed. Here we conducted experimental manipulations using Arabidopsis thaliana with the ancestral plants grown in different nitrogen (N) and phosphorus (P) availabilities over eleven consecutive generations, and then examined the offspring phenotypic performance under the interactive effects of current and ancestral nutrient environments. We found that current rather than ancestral nutrient environments dominantly explained the variations in offspring plant traits (i.e., flowering time, aboveground biomass and biomass allocation fractions), suggesting the relatively weak transgenerational effects of ancestral N and P availabilities on offspring phenotypes. In contrast, increasing N and P availabilities in the offspring generation remarkably shortened the flowering time, increased the aboveground biomass, and altered biomass allocation fractions differentially among organs. Despite the overall weak transgenerational phenotypic plasticity, under the low nutrient environment, the offspring of ancestral plants from the low nutrient environment had a significantly higher fruit mass fraction than those from the suitable nutrient environment. Taken together, our findings suggest that A. thaliana exhibits a much stronger within- than trans-generational trait plasticity under contrasting nutrient availabilities, and may provide important insights into the understanding of plant adaptation and evolutionary processes under changing nutrient environments.

Decoupling of N and P aggravated upward along food chains in an urban river ecosystem.

Anthropogenic input of nutrient has profoundly influenced water quality and aquatic organisms, however, large and unbalanced nitrogen (N) and phosphorus (P) inputs (decoupling) can lead to a range of ecological health problems such as eutrophication. Whether and how the decoupling varies along the aquatic food chain remains poorly addressed. Here we chose an urban river ecosystem in the cosmopolis region of Beijing, with reclaimed water as the entire replenishment water source over 20 years, to demonstrate the decoupling pattern of N vs P across trophic levels. Results showed that organism C, N and P concentration increased, but N:P ratio decreased upward along the food chains, suggesting that this decoupling of N and P increased as trophic level ascends. Compared with natural freshwater ecosystem, the decoupling of N and P was aggravated in the reclaimed water river. Moreover, the homeostasis of N and P were higher at higher relative to lower trophic levels, and higher in macro-food chain relative to planktonic food chain. This study, for the first time, revealed the increasing decoupling of N vs P upward along the major food chains in an urban aquatic ecosystem, and could improve the understanding of nutrient cycling at the food chain level under human disturbance, and provide useful information for ecological restoration and eutrophication control of urban wetlands replenished with reclaimed water.

Scaling the leaf nutrient resorption efficiency: Nitrogen vs phosphorus in global plants.

Nutrient resorption from senescent leaves is one essential plant nutrient strategy. Allocation of nitrogen (N) and phosphorus (P) reflects the influences of evolution and ecological processes on plant functional traits, and thus is related to functional types and environmental factors. However, we know little about the pattern among plant functional types (PFTs) and the driving factors of the allometric relationship of N resorption efficiency (NRE) against P resorption efficiency (PRE) in plant leaves (NRE ~ PREb; b, scaling exponent). We compiled N and P resorption data from the literature, including 2541 records, 894 plant species, and 488 sites worldwide, and then explored the allometric relationships between NRE and PRE across different PFTs and environmental factors (i.e. climate and soil nutrients). The scaling exponent for overall species was 0.88, suggesting that plants generally re-absorb P from senesced leaves at a higher rate than N. Among diverse PFTs, the scaling exponents of broadleaved (0.91), deciduous (0.92), non-leguminous (0.88), and woody plants (0.90) were higher than those of coniferous (0.81), evergreen (0.89), leguminous (0.74), and herbaceous plants (0.76), respectively. The scaling exponents increased with increasing latitude and soil nutrient (N and P) availability, and decreased with increasing mean annual temperature. Our results suggest that terrestrial plants utilize P relative to N more effectively through resorbing a higher proportion of P than N from senescent leaves. However, the differential resorption efficiency between N and P may vary among diverse plant types, and displayed a biogeographic pattern at global scale through the plant-environment interactions. These findings can broaden our understanding of the nutrient recycling processes within plants, and help in better prediction of nutrient balance in response to global changes.

[Nitrogen and phosphorus stoichiometry of aquatic macrophytes in Kaidu River of Xinjiang and the potential influencing factors].

Nitrogen (N) and phosphorus (P) are two common mineral elements constraining plant growth. Nutrients in aquatic macrophytes are mainly absorbed from water and sediments, and thus elemental composition in plant tissues can reflect the nutritional status in freshwater ecosystems. Kaidu River is an important river flowing through the alpine meadows, deserts, and desert oases in Xinjiang. Herein, samples of aquatic macrophytes, water, and sediments across the Kaidu River were collected. Foliar stoichiometric characteristics of N and P in plants were examined among life forms and phylogeny. The differences and correlations among the stoichiometric characteristics of plants, water bodies, and sediments in the upstream and downstream and across different land use types were elaborated. Results showed that the means of leaf N and P concentrations and N/P mass ratio were 24.9 mg·g-1, 2.49 mg·g-1, and 12.6, respectively. There were significant differences in the leaf N and P stoichiometry among various life forms. Specifically, leaf N and P concentrations in submerged species were significantly higher than that in floating-leaved species and emergent species. The N/P of floating-leaved species was 19.2 and significantly higher than that in other two life-forms, indicating that their growth might be limited by P. Leaf N and P concentrations were higher in the upstream than in the downstream. Nitrogen contents in water and sediment were the higher in the upstream of the river. Compared with the river reaches in farming areas, the higher N and P concentrations of aquatic macrophytes in the grassland regions might be related to the higher contents of organic matter in grassland soils and of animal slurries from flocks and herds, suggesting that grazing exerted larger impact on the stoichiometric characteristics of the Kaidu River ecosystem. Our findings highlighted that overgrazing might accelerate the deterioration of water quality in the upstream of the Kaidu River, disturb the balance of N and P in the aquatic ecosystem, and potentially influence the biogeochemical cycling.

Family-level leaf nitrogen and phosphorus stoichiometry of global terrestrial plants.

Leaf nitrogen (N) and phosphorus (P) concentrations are critical for photosynthesis, growth, reproduction and other ecological processes of plants. Previous studies on large-scale biogeographic patterns of leaf N and P stoichiometric relationships were mostly conducted using data pooled across taxa, while family/genus-level analyses are rarely reported. Here, we examined global patterns of family-specific leaf N and P stoichiometry using a global data set of 12,716 paired leaf N and P records which includes 204 families, 1,305 genera, and 3,420 species. After determining the minimum size of samples (i.e., 35 records), we analyzed leaf N and P concentrations, N:P ratios and N∼P scaling relationships of plants for 62 families with 11,440 records. The numeric values of leaf N and P stoichiometry varied significantly across families and showed diverse trends along gradients of mean annual temperature (MAT) and mean annual precipitation (MAP). The leaf N and P concentrations and N:P ratios of 62 families ranged from 6.11 to 30.30 mg g-1, 0.27 to 2.17 mg g-1, and 10.20 to 35.40, respectively. Approximately 1/3-1/2 of the families (22-35 of 62) showed a decrease in leaf N and P concentrations and N:P ratios with increasing MAT or MAP, while the remainder either did not show a significant trend or presented the opposite pattern. Family-specific leaf N∼P scaling exponents did not converge to a certain empirical value, with a range of 0.307-0.991 for 54 out of 62 families which indicated a significant N∼P scaling relationship. Our results for the first time revealed large variation in the family-level leaf N and P stoichiometry of global terrestrial plants and that the stoichiometric relationships for at least one-third of the families were not consistent with the global trends reported previously. The numeric values of the family-specific leaf N and P stoichiometry documented in the current study provide critical synthetic parameters for biogeographic modeling and for further studies on the physiological and ecological mechanisms underlying the nutrient use strategies of plants from different phylogenetic taxa.

Effects of nitrogen and phosphorus supply on stoichiometry of six elements in leaves of Arabidopsis thaliana.

BACKGROUND AND AIMS: Plant elemental composition is of fundamental importance for plant growth and metabolic functions. However, knowledge of how multi-elemental stoichiometry responds to varying nitrogen (N) and phosphorus (P) availabilities remains limited. METHODS: We conducted experimental manipulations with nine repeat experiments to investigate the effects of N and P supply on the concentrations and variability of six elements, carbon (C), N, P, potassium (K), calcium (Ca) and magnesium (Mg), in leaves of Arabidopsis thaliana. KEY RESULTS: N supply increased the concentrations of N, K and Mg, decreased the concentration of P, but exerted little influence on the concentrations of C and Ca in green leaves. P supply increased the concentrations of P and Ca, decreased the concentration of C, initially increased and then decreased the concentration of K, but showed little influence on the concentrations of N and Mg in green leaves. Multivariate patterns among the concentrations of these six elements in green leaves was influenced by the type of nutrient supply (i.e. N or P). Elemental variability decreased with increasing elemental concentrations in green leaves at the intraspecific level, supporting the Stability of Limiting Elements Hypothesis that was originally proposed from a meta-analysis of pooled data across species or communities. Compared with green leaves, the senesced leaves showed greater variability in C, N, P, K and Mg concentrations but lower variability in Ca concentration. CONCLUSIONS: N and P supplies exerted differential influences on the concentrations of C, N, P, K, Ca and Mg in green leaves. The specific C content should be considered when assessing C cycling under global nutrient changes. Stage-dependent patterns of leaf stoichiometric homeostasis differed among elements with various chemical characteristics. These findings can help to improve our understanding of plant eco-physiological responses and acclimation under global nutrient changes from the stoichiometric perspective of multiple elements.

An assessment on the uncertainty of the nitrogen to phosphorus ratio as a threshold for nutrient limitation in plants.

BACKGROUND AND AIMS: The nitrogen (N) to phosphorus (P) ratio (N:P) has been widely used as a threshold for identifying nutrient limitations in terrestrial plants; however, the associated reliability has not been well assessed. METHODS: The uncertainty of nutrient limitations caused by the N:P threshold was evaluated using two approaches: fertilization experiments synthesized across multiple ecosystems; and random sampling simulation of the impacts of different nutrient sufficiencies and deficiencies. KEY RESULTS: The fertilization experiment data indicated that the types of nutrient limitation determined via N:P thresholds were partly inconsistent with the growth responses observed under N and P additions, i.e. under N:P thresholds of 14 and 16 (or 10 and 20), 32.5 % (or 16.2 %) of the data were inconsistent between these two. The random sampling simulation suggested that N:P thresholds may indicate N (or P) limitations when leaf N (or P) content is sufficient, whereas these thresholds may not indicate N (or P) limitations when leaf N (or P) content is deficient. The error risks calculated from the sampling simulation presented large fluctuations at small sample sizes and decreased as the thresholds of nutrient content sufficiency (or deficiency) increased (or decreased). The N:P thresholds of 10 and 20 showed lower error risks than the thresholds of 14 and 16. CONCLUSIONS: These findings highlight that canonical N:P thresholds have the potential to introduce a large uncertainty when used to detect plant nutrient limitations, suggesting that the error risks should be cautioned in future studies.

Anatomical responses of leaf and stem of Arabidopsis thaliana to nitrogen and phosphorus addition.

Nitrogen (N) and phosphorus (P) availabilities play crucial roles in plant morphogenesis and physiological processes, but how plant anatomical traits respond to the N and P supply is not well elucidated. We evaluated the effects of N and P supply on multiple leaf and stem anatomical traits of Arabidopsis thaliana. The addition of N increased the stem diameter, cortex thickness, rosette radius, midrib thickness, and size of leaf and stem vasculature significantly. Abaxial stomatal length (LSL) increased while adaxial epidermal cell density decreased significantly with increasing N supply. P addition did not affect stem size and leaf epidermal traits, but enhanced the thickness of stem xylem. The nutrient limiting status did not affect most traits except for LSL. The anatomical traits measured varied a lot in the extent of response to N and P addition, despite relatively stronger response to N addition overall. Cortex thickness, rosette radius, stomatal density and epidermal cell density exhibited relatively high plasticity to both nutrients, while stomatal length and stomatal index were relatively stable. Thus, these results suggested that the anatomical traits of shoot vasculature of A. thaliana were enhanced by both nutrients but more affected by N addition, satisfying the plant growth and nutrient requirements. Our findings may help shed light on plant adaptation to nutrient availability changes under the ongoing anthropogenic impacts, but the generality across numerous plant species still warrants further researches.

Stage-dependent stoichiometric homeostasis and responses of nutrient resorption in Amaranthus mangostanus to nitrogen and phosphorus addition.

Stoichiometric homeostasis is the ability of plants remaining their element composition relatively stable regardless of changes in nutrient availability, via various physiological mechanisms. Nutrient resorption is one of such key mechanisms, but whether and how nitrogen and phosphorus homeostasis and resorption in plants would change with growth-stages under variable nutrient supply was unclear. A nitrogen (N) and phosphorus (P) fertilizer addition experiment was conducted to evaluate the dynamics of N and P homeostasis and resorption efficiency during different growth-stages of Amaranthus mangostanus in a greenhouse. The homeostasis regulation coefficient of green-leaf P varied significantly, while that of green-leaf N maintained relatively stable across growth stages. Moreover, homeostasis regulation coefficient of N was higher at seedling stage but lower at flowering stage than that of P at corresponding stages, suggesting that the growth of A. mangostanus may switch from being more N- to P-limited from vegetative to reproductive stage. N resorption efficiency (NRE) was higher and P resorption efficiency (PRE) was lower at flowering than seed-filling stage. The nutrient dynamics displayed here suggested contrasting nutrient homeostasis and resorption responses of plants to environmental nutrient availability across growth stages. These findings can improve the understanding of nutrition maintenance mechanism of plants during their growth.

Phosphorus accumulates faster than nitrogen globally in freshwater ecosystems under anthropogenic impacts.

Combined effects of cumulative nutrient inputs and biogeochemical processes that occur in freshwater under anthropogenic eutrophication could lead to myriad shifts in nitrogen (N):phosphorus (P) stoichiometry in global freshwater ecosystems, but this is not yet well-assessed. Here we evaluated the characteristics of N and P stoichiometries in bodies of freshwater and their herbaceous macrophytes across human-impact levels, regions and periods. Freshwater and its macrophytes had higher N and P concentrations and lower N : P ratios in heavily than lightly human-impacted environments, further evidenced by spatiotemporal comparisons across eutrophication gradients. N and P concentrations in freshwater ecosystems were positively correlated and N : P was negatively correlated with population density in China. These results indicate a faster accumulation of P than N in human-impacted freshwater ecosystems, which could have large effects on the trophic webs and biogeochemical cycles of estuaries and coastal areas by freshwater loadings, and reinforce the importance of rehabilitating these ecosystems.

Nutrient allocation strategies of woody plants: an approach from the scaling of nitrogen and phosphorus between twig stems and leaves.

Allocation of limited nutrients, such as nitrogen (N) and phosphorus (P), among plant organs reflects the influences of evolutionary and ecological processes on functional traits of plants, and thus is related to functional groups and environmental conditions. In this study, we tested this hypothesis by exploring the stoichiometric scaling of N and P concentrations between twig stems and leaves of 335 woody species from 12 forest sites across eastern China. Scaling exponents of twig stem N (or P) to leaf N (or P) varied among functional groups. With increasing latitude, these scaling exponents significantly decreased from >1 at low latitude to <1 at high latitude across the study area. These results suggested that, as plant nutrient concentration increased, plants at low latitudes showed a faster increase in twig stem nutrient concentration, whereas plants at high latitudes presented a faster increase in leaf nutrient concentration. Such shifts in nutrient allocation strategy from low to high latitudes may be controlled by temperature. Overall, our findings provide a new approach to explore plant nutrient allocation strategies by analysing the stoichiometric scaling of nutrients among organs, which could broaden our understanding of the interactions between plants and their environments.

Reproductive organ and young tissues show constrained elemental composition in Arabidopsis thaliana.

BACKGROUND AND AIMS: The identification of stoichiometric homeostasis is crucial for understanding plant adaptive strategies under a changing environment. However, current knowledge of plant stoichiometric homeostasis has mainly been obtained from mature leaves, with little from other organs across different developmental stages. METHODS: We conducted a greenhouse nitrogen (N) and phosphorus (P) addition experiment to evaluate the strength of stoichiometric homeostasis across different organs and developmental stages of Arabidopsis thaliana. KEY RESULTS: Homeostatic regulation coefficients (H) for N (HN), P (HP) and N : P ratio (HNP) were highest in reproductive tissue, followed by stem and leaf at the same stage. All H parameters in the same organ decreased significantly over the developmental stages. Leaf HN, HP and HNP were highest at stage 1, followed by stages 2 and 3. Both stem and silique at stage 2 relative to stage 3 had higher HN, HP and HNP. These results suggested that reproductive tissue relative to other organs and young tissue relative to old tissue showed more constrained elemental composition in response to nutrient availabilities, and such trends were also evidenced by stoichiometric scaling relationships. CONCLUSIONS: Our findings highlight that stoichiometric homeostasis is tightly related to the ontogenesis of plant tissue. These results could have a strong implication for diagnosing relative availabilities of N and P in ecosystems, suggesting that the N and P stoichiometry of old tissues might be stronger indicators of nutrient status for plants, but further study is needed to test the generality across species with more distinguishable functional traits.

Biogeographic patterns of structural traits and C:N:P stoichiometry of tree twigs in China's forests.

There have been a number of studies on biogeographic patterns of plant leaf functional traits; however, the variations in traits of other plant organs such as twigs are rarely investigated. In this study, we sampled current-year twigs of 335 tree species from 12 forest sites across a latitudinal span of 32 degrees in China, and measured twig specific density (TSD), twig dry matter content (TDMC), and carbon (C), nitrogen (N) and phosphorous (P) contents, to explore the latitudinal and environmental patterns of these twig traits. The overall mean of TSD and TDMC was 0.37 g cm(-3) and 41%, respectively; mean twig C, N and P was 472 mg g(-1), 9.8 mg g-1 and 1.15 mg g(-1), respectively, and mean N:P mass ratio was 10.6. TSD was positively correlated with TDMC which was positively associated with twig C but negatively with twig N and P. There were no significant differences in TSD between conifer, deciduous-broadleaf and evergreen-broadleaf plants, but evergreen-broadleaf plants had the lowest and conifers the highest TDMC. Conifer twigs were lowest in C, N, P and N:P, whereas deciduous-plant twigs were highest in N and P and evergreen-plant twigs were highest in C and N:P. As latitude increased or temperature/precipitation dropped, TDMC and P increased, but N:P ratio decreased. Our results also showed that the patterns of twig P and N:P stoichiometry were consistent with those reported for leaves, but no significant trends in twig N were observed along the gradient of latitude, climate and soils. This study provides the first large-scale patterns of the twig traits and will improve our understanding of the biogeochemistry of carbon and other key nutrients in forest ecosystems.