Soil warming accelerates decomposition of fine woody debris

Aims

There is evidence that increased N inputs to boreal forests, via atmospheric deposition or intentional fertilization, may impact negatively on ectomycorrhizal (ECM) fungi leading to a reduced flux of plant-derived carbon (C) back to the atmosphere via ECM. Our aim was to investigate the impact of N fertilization of a Pinus sylvestris (L.) forest stand on the return of recently photoassimilated C via the ECM component of soil respiration.

Methods

We used an in situ, large-scale, ¹³C-CO₂ isotopic pulse labelling approach and monitored the ¹³C label return using soil gas efflux chambers placed over three different types of soil collar to distinguish between heterotrophic (R_H), autotrophic (R_A; partitioned further into contributions from ECM hyphae and total R_A) and total (R_S) soil respiration.

Results

The impact of N fertilization was to significantly reduce R_A, particularly respiration via extramatrical ECM hyphae. ECM hyphal flux in control plots showed substantial spatial variability, resulting in mean flux estimates exceeding estimates of total R_A, while ECM contributions to R_A in N treated plots were estimated at around 30%.

Conclusion

Significant impacts on soil C cycling may be caused by reduced plant C allocation to ECM fungi in response to increased N inputs to boreal forests; ecosystem models so far lack this detail.     

Influence of elevation on canopy transpiration of temperate deciduous forests in a complex mountainous terrain of South Korea

Background and aims

Variations in microclimate and soil characteristics on mountain slopes influence forest structure and function. Precipitation, incoming solar radiation and relative humidity change along a mountain slope. Equally, soil depth and the amount of stored soil moisture vary. The objective of this study was to examine the impacts of these factors on forest water use in mountainous terrains.

Methods

Transpiration of four temperate deciduous forest stands located at different elevations in South Korea was monitored with a sap flow technique throughout the growing season in 2010. The study sites were located on the north slope at 450 m (450 N), 650 m (650 N), and 950 m (950 N). To examine the effect of aspect, an additional site with a southern aspect was studied at 650 m (650S). All the sites were dominated by Quercus species, with leaf area index (L) ranging between 5 ? 6 m² m^?2.

Results

Rainfall increased, while air temperature (T _A ) and daytime vapor pressure deficit (D) decreased with increasing elevation. We did not observe any gradients in solar radiation (R _S ), soil moisture and sap flux density of the individual trees (J _st ) with an elevational gradient. Sapwood area (A _S ), i.e., hydro-active xylem area, and daily maximum tree water use (max TWU) increased non-linearly with increasing diameter at breast height (DBH). Neither A _S nor max TWU varied among tree species or along the elevation. The total annual canopy transpiration (E _C ) was 175, 115, 110, and 90 mm for 450 N, 650 N, 650S, and 950 N, respectively. E _C declined with increasing elevation as a result of decreasing length of the growing season, D, and T _A along the elevation. Significantly (P < 0.001) higher stomatal sensitivity to changes in D was found at the 950 N, leading to lower annual E _C and lower water use efficiency (WUE) at this elevation.

Conclusions

We conclude that differences in E _C exist along the mountain slope studied, corresponding to changing T _A , D, length of the growing season, and stomatal sensitivity to D, which should be considered when establishing forest catchment water balances.     

Optimizing nitrogen economy under drought: increased leaf nitrogen is an acclimation to water stress in willow (Salix spp.)

Background and Aims

The major objective was to identify plant traits functionally important for optimization of shoot growth and nitrogen (N) economy under drought. Although increased leaf N content (area basis) has been observed in dry environments and theory predicts increased leaf N to be an acclimation to drought, experimental evidence for the prediction is rare.

Methods

A pedigree of 200 full-sibling hybrid willows was pot-grown in a glasshouse in three replicate blocks and exposed to two water regimes for 3 weeks. Drought conditions were simulated as repeated periods of water shortage. The total leaf mass and area, leaf area efficiency (shoot growth per unit leaf area, E_A), area-based leaf N content (N_A), total leaf N pool (N_L) and leaf N efficiency (shoot growth per unit leaf N, E_N) were assessed.

Key Results

In the water-stress treatment, shoot biomass growth was N limited in the genotypes with low N_L, but increasingly limited by other factors in the genotypes with greatest N_L. The N_A was increased by drought, and drought-induced shift in N_A varied between genotypes (significant G × E). Judged from the E_A–N_A relationship, optimal N_A was 16 % higher in the water-stress compared with the well-watered treatment. Biomass allocation to leaves and shoots varied between treatments, but the treatment response of the leaf : shoot ratio was similar across all genotypes.

Conclusions

It is concluded that N-uptake efficiency and leaf N efficiency are important traits to improve growth under drought. Increased leaf N content (area basis) is an acclimation to optimize N economy under drought. The leaf N content is an interesting trait for breeding of willow bioenergy crops in a climate change future. In contrast, leaf biomass allocation is a less interesting breeding target to improve yield under drought.     

Root phenotyping by root tip detection and classification through statistical learning

Background and aims

Variations in the water and soil background in the signal path can cause variations in canopy spectral reflectance, which leads to uncertainty in estimating the canopy nitrogen (N) status. The primary objective of this study was to explore the optimum vegetation indices that were highly correlated with canopy leaf N concentration (LNC) but less influenced by the canopy leaf area index (LAI) and vegetation coverage (VC) in rice.

Methods

A systematic analysis of the quantitative relationships between various hyperspectral vegetation indices and LNC, VC and LAI was conducted based on 4-year rice field experiments using different rice varieties, N rates and planting densities. New spectral indices were derived to estimate LNC in rice under variable vegetation coverage.

Results

Although the newly developed simple green ratio indices, SR (R₅₅₃, R₅₃₇) and SR (R₅₄₅, R₅₃₈), and the three-band index (R₆₀₅-R₅₂₁-R₆₈₂)/(R₆₀₅+R₅₂₁+R₆₈₂) correlated well with the LNC. Only SR (R₅₅₃, R₅₃₇) was less influenced by VC/LAI and showed a stable performance in both the independent calibration and validation datasets. For the published indices tested in the present study, NDVIg-b and ND (R₅₀₃, R₄₈₃) showed a good predictive ability for the LNC. However, both of these indices and other published indices were found to be significantly dominated by the VC/LAI.

Conclusion

SR (R₅₅₃, R₅₃₇) was the best index to reliably estimate the LNC in rice under various cultivation conditions, and is recommended for this use. However, other spectral indices need to be examined to determine if they influenced by factors such as VC/LAI. Such studies will improve the applicability of these indices to different types of rice cultivars and production systems.     

Within-canopy variation in photosynthetic capacity,SLA and foliar N in temperate broad-leaved trees with contrasting shade tolerance

Key message

The relative shade tolerance of T. cordata , F. sylvatica , and C. betulus in mature stands is based on different species-specific carbon and nitrogen allocation patterns.

Abstract

The leaf morphology and photosynthetic capacity of trees are remarkably plastic in response to intra-canopy light gradients. While most studies examined seedlings, it is not well understood how plasticity differs in mature trees among species with contrasting shade tolerance. We studied light-saturated net photosynthesis (A _max), maximum carboxylation rate (V _cmax), electron transport capacity (J _max) and leaf dark respiration (R _d) along natural light gradients in the canopies of 26 adult trees of five broad-leaved tree species in a mixed temperate old-growth forest (Fraxinus excelsior, Acer pseudoplatanus, Carpinus betulus, Tilia cordata and Fagus sylvatica), representing a sequence from moderately light-demanding to highly shade-tolerant species. We searched for species differences in the dependence of photosynthetic capacity on relative irradiance (RI), specific leaf area (SLA) and nitrogen per leaf area (N _a ). The three shade-tolerant species (C. betulus, T. cordata, F. sylvatica) differed from the two more light-demanding species by the formation of shade leaves with particularly high SLA but relatively low N _a and consequently lower area-based A _max, and a generally higher leaf morphological and functional plasticity across the canopy. Sun leaf morphology and physiology were more similar among the two groups. The three shade-tolerant species differed in their shade acclimation strategies which are primarily determined by the species’ plasticity in SLA. Under low light, T. cordata and F. sylvatica increased SLA, mass-based foliar N and leaf size, while C. betulus increased solely SLA exhibiting only low intra-crown plasticity in leaf morphology and N allocation patterns. This study with mature trees adds to our understanding of tree species differences in shade acclimation strategies under the natural conditions of a mixed old-growth forest.     

Concentric-tube airlift bioreactors

Mass transfer coefficients were measured in three concentric-tube airlift reactors of different scales (RIMP, V _L =0.07 m³;RIS?1,V _L =2.50 m³;RIS?2, V _L =5.20 m³). The effects of top and bottom clearance and flow resistances at downcorner entrance were studied in water-air system. Experimental results show that h _s ,h _B and A _d /A _R ratio affect K _L a values as a result of their influence on gas holdup and liquid velocity. The gas-liquid mass-transfer coefficients for all the geometric variables were successfully correlated as Sherwood number with Froude and Galilei numbers, the bottom spatial ratio (B=h _B /D _R ), the top spatial ratio , the gas separation ratio and the downcomer flow resistance ratio (R=A _d /A _R ). The proposed empirical model satisfactorily fitted the experimental data obtained in large airlift reactors and some data presented in literature.     

Decoupling litter barrier and soil moisture influences on the establishment of an invasive grass

Background and aims

Through recruitment, plants establish in novel environments. Recruitment also is the stage where plants undergo the highest mortality. We investigate the recruitment niche for Microstegium vimineum, an annual grass from East Asia spreading throughout eastern North American forests.

Methods

Current observational and greenhouse research indicates that M. vimineum recruitment may be inhibited by leaf litter and promoted by soil moisture; we use field studies to experimentally test how these factors influence M. vimineum germination, seedling survival and reproduction. Specifically, we introduce M. vimineum seeds into forest microhabitats with experimentally varied levels of soil moisture and leaf litter.

Results

Soil moisture increases M. vimineum germination regardless of leaf litter thickness and ameliorates seedling mortality in deep leaf litter. Seed production per m² increases with watering, reflecting higher germination and survival, whereas per capita seed production increases with leaf litter thickness, reflecting density-dependent limits on seed production.

Conclusions

The interactive effects of varied levels of soil moisture and leaf litter thickness on key M. vimineum life history stages highlight the need to consider multiple drivers, such as rainfall and local forest disturbance, when assessing how soil properties influence the establishment of invasive plants.     

Precipitation variability does not affect soil respiration and nitrogen dynamics in the understorey of a Mediterranean oak woodland

Background and aims

Future climate scenarios for the Mediterranean imply increasing precipitation variability. This study presents a large-scale water manipulation experiment simulating changes in precipitation variability, aiming at a better understanding of the effects of rainfall patterns on soil C and N cycling and understorey productivity in a Mediterranean oak woodland.

Methods

We used rain-out shelters to achieve (1) a normal dry period (7 days), and (2) a dry period increased three-fold (21 days), without altering total annual precipitation inputs.

Results

The temporal patterns of soil respiration (R _s) and soil inorganic N were not affected by treatment. However, water infiltration and N leaching increased with large infrequent watering events. R _s and soil NH₄ ⁺-N correlated with soil temperature, with soil NO₃ ^?-N being influenced by leaching.

Conclusions

The lack of significant treatment effects on either R _s or soil inorganic N can be explained by (1) minor differences in plant productivity between the treatments, suggesting equal plant N demand, and (2) the absence of moisture dependence of R _s and soil NH₄ ⁺-N. Increased N leaching with large infrequent precipitation events may have longer-term consequences for ecosystem functioning. Our results contribute to an improved understanding of possible climate change effects on key ecosystem processes in Mediterranean ecosystems.     

Conservation of functional traits leads to shrub expansion across a chronosequence of shrub thicket development

Key message

Robust physiology of Myrica cerifera across a chronosequence (i.e., space for time substitution) of shrub thicket age classes contributes to rapid cover expansion observed in the last 50 years.

Abstract

Many studies have documented the causes of woody expansion into grasslands, but few address unique morphological and physiological traits that facilitate expansion. Myrica cerifera, an evergreen N-fixer, is the dominant shrub on many barrier islands of the southeastern United States. Cover of Myrica cerifera has expanded by ~400 % on Hog Island, Virginia, in the past 50 years. Accretion of the northern end of the island has resulted in a chronosequence (i.e., space for time substitution) of both soil age and shrub thicket development. We investigated functional traits and physiological parameters related to light capture, processing and water balance of M. cerifera across shrub thickets of four age classes from ~10 to ~50 years. We hypothesized that light processing capabilities and hydraulic capacity would be reduced with thicket age. Spatial variation in morphology (i.e., leaf thickness, leaf area) and structure (i.e., leaf angle) related to light capture was observed. Yet, little or no differences were detected in stomatal density, photosynthetic pigments, electron transport rate (ETR) and hydraulic conductivity across sites. Previous research has shown declines in leaf N content, productivity and leaf litter production across the chronosequence. In contrast, we observed that physiology remains consistent despite considerable differences in thicket age and development. Myrica cerifera maintains high photosynthetic and hydraulic efficiency, factors which enable expansion and maintenance of shrub thickets in mesic coastal environments.     

The effects of the moss layer on the decomposition of intercepted vascular plant litter across a post-fire boreal forest chronosequence

Aims

Feather mosses form a thick ground layer in boreal forests that can intercept incoming litter fall. This interception may influence the decomposition of incoming litter but this has been little explored. We investigated how the moss layer influences decomposition of intercepted litter along a 362-year fire driven forest chronosequence in northern Sweden across which soil fertility declines.

Methods

We placed leaf litter from three plant species into plots in which mosses and dwarf shrubs were either experimentally removed or left intact, at each of ten stands across the chronosequence. After one year we measured litter mass loss, and litter nitrogen and phosphorous.

Results

Litter decomposed consistently faster, and had higher nitrogen and phosphorus, in the presence of mosses and at greater depth in the moss layer. Despite an increase in moss depth across the chronosequence we did not find consistent increases in effects of moss removal on litter decomposition or on litter N or P.

Conclusions

Our findings identify a clear role of the moss layer in boreal forests in promoting the decomposition of intercepted leaf litter, and highlight that this role is relatively consistent across chronosequence stages that vary greatly in productivity and moss depth.     

Seedling growth and soil nutrient availability in exotic and native tree species: implications for afforestation in southern China

Background and aims

The relationship between tree species and soil nutrient availability is critical for evaluating plantation succession and promoting forest restoration. This study was conducted to evaluate the impact of exotic and native tress species on soil nutrient availability.

Methods

Four exotic species (Eucalyptus urophylla, E. tereticornis, Acaia auriculaeformis, A. mangium) and four native species (Castanopsis fissa, Schima superba, C. hystrix, Michelia macclurei) were planted and grown for one-year. Soil solution (DOC, DON, NH₄?N, NO₃?N) was sampled and analyzed during the study. After the experiment, soil properties were determined, and plant tissues were analyzed.

Results

DOC levels were greater in soils with trees planted than controls without trees. Compared to native species, exotic species had much faster growth rates and greatly reduced DON and NO₃?N concentrations. Exotic species always had less P concentrations in leaves and stems than native species. Furthermore, N-fixing A. auriculaeformis led to greater soil available P compared to other species.

Conclusions

Based on these findings, we provide some recommendations for afforestation practice. This study highlights that a better understanding of the pros and cons of exotic species would be beneficial to advance afforestation in China and the world.     

N and P colimitation of N2-fixing and N-supplied fynbos legumes from the Cape Floristic Region

Background and aims

Legume species in the fynbos vegetation of the Cape Floristic Region, that fix N₂ in soils with low P, may have evolved for enhanced acquisition and efficient use of P. It was hypothesized that N₂-fixing and combined-N supplied (N-supplied) A. linearis, P. calyptrata and C. genistoides are adapted to low P and would be relatively unresponsive to increased P of 100 μM.

Methods

18 legume species were evaluated for their nodulation response to low P availability. The N X P interaction was then examined in A. linearis, P. calyptrata and C. genistoides reliant on either N₂-fixation or 300 μM N (NH₄NO₃), and receiving 0.1, 1.0, 10 and 100 μM P (NaH₂PO₄).

Results

In the species selection experiment, A. linearis, P. calyptrata and C. genistoides, with the greatest nodule fresh weight (FW) and nodule FW to root FW ratio, were the most prolific nodulating species. In the N X P experiment, with low P supply, the biomass of N₂-fixing P. calyptrata and C. genistoides was consistently greater than that of N-supplied plants. In contrast, with high P supply of 100 μM P, all N-supplied plants accumulated more biomass than the corresponding N₂-fixing plants. High P-use efficiency, poor down-regulation of P uptake and P storage was evident in A. linearis and P. calyptrata.

Conclusion

The growth response to P and the significant N X P interactions indicate that N₂-fixing and N-supplied plants were not adapted to low P, but rather colimited by both N and P.     

Photosynthetic characteristics and nitrogen allocation in the black locust (<Emphasis Type="Italic">Robinia pseudoacacia</Emphasis> L.) grown in a FACE system

Key message

The black locust is adapted to elevated [CO ₂ ] through changes in nitrogen allocation characteristics in leaves.

Abstract

The black locust (Robinia pseudoacacia L.) is an invasive woody legume within Japan. This prolific species has a high photosynthetic rate and growth rate, and undergoes symbiosis with N₂-fixing micro-organisms. To determine the effect of elevated CO₂ concentration [CO₂] on its photosynthetic characteristics, we studied the chlorophyll (Chl) and leaf nitrogen (N) content, and the leaf structure and N allocation patterns in the leaves and acetylene reduction activity after four growing seasons, in R. pseudoacacia. Our specimens were grown at ambient [CO₂] (370 μmol mol^?1) and at elevated [CO₂] (500 μmol mol^?1), using a free air CO₂ enrichment (FACE) system. Net photosynthetic rate at growth [CO₂] (A _growth) and acetylene reduction activity were significantly higher, but maximum carboxylation rate of RuBisCo (V _cmax), maximum rate of electron transport driving RUBP regeneration (J _max), net photosynthetic rate under enhanced CO₂ concentration and light saturation (A _max), the N concentration in leaf, and in leaf mass per unit area (LMA) and ribulose-1,5-bisphosphate carboxylase oxygenase (RuBisCo) content were significantly lower grown at elevated [CO₂] than at ambient [CO₂]. We also found that RuBisCo/N were less at elevated [CO₂], whereas Chl/N increased significantly. Allocation characteristics from N in leaves to photosynthetic proteins, N_L (Light-harvesting complex: LHC, photosystem I and II: PSI and PSII) and other proteins also changed. When R. pseudoacacia was grown at elevated [CO₂], the N allocation to RuBisCo (N_R) decreased to a greater extent but N_L and N remaining increased relative to specimens grown at ambient [CO₂]. We suggest that N remobilization from RuBisCo is more efficient than from proteins of electron transport (N_E), and from N_L. These physiological responses of the black locust are significant as being an adaptation strategy to global environmental changes.

     

Roots affect the response of heterotrophic soil respiration to temperature in tussock grass microcosms

Aims and Background

While the temperature response of soil respiration (R_S) has been well studied, the partitioning of heterotrophic respiration (R_H) by soil microbes from autotrophic respiration (R_A) by roots, known to have distinct temperature sensitivities, has been problematic. Further complexity stems from the presence of roots affecting R_H, the rhizosphere priming effect. In this study the short-term temperature responses of R_A and R_H in relation to rhizosphere priming are investigated.

Methods

Temperature responses of R_A, R_H and rhizosphere priming were assessed in microcosms of Poa cita using a natural abundance δ¹³C discrimination approach.

Results

The temperature response of R_S was found to be regulated primarily by R_A, which accounted for 70 % of total soil respiration. Heterotrophic respiration was less sensitive to temperature in the presence of plant roots, resulting in negative priming effects with increasing temperature.

Conclusions

The results emphasize the importance of roots in regulating the temperature response of R_S, and a framework is presented for further investigation into temperature effects on heterotrophic respiration and rhizosphere priming, which could be applied to other soil and vegetation types to improve models of soil carbon turnover.     

Progressive and retrogressive ecosystem development coincide with soil bacterial community change in a dune system under lowland temperate rainforest in New Zealand

Background

It is established that plant communities show patterns of change linked to progressive and retrogressive stages of ecosystem development. It is not known, however, whether bacterial communities also show similar patterns of change associated with long-term ecosystem development.

Methods

We studied soil bacterial communities along a 6,500 year dune chronosequence under lowland temperate rain forest at Haast, New Zealand. Pyrosequencing of 16S rRNA genes was used to observe structural change in bacterial communities during the process of pedogenesis and ecosystem development.

Results

Bacterial communities showed patterns of change during pedogenesis, with the largest change during the first several hundred years after dune stabilization. The most abundant bacterial taxa were Alphaproteobacteria, Actinobacteria and Acidobacteria. These include taxa most closely related to nitrogen-fixing bacteria, and suggest heterotrophic nitrogen input may be important throughout the chronosequence. Changes in bacterial community structure were related to changes in several soil properties, including total phosphorus, C:N ratio, and pH. The Bacteroidetes, Actinobacteria, Cyanobacteria, Firmicutes, and Betaproteobacteria all showed a general decline in abundance as pedogenesis proceeded, while Acidobacteria, Alphaproteobacteria, and Plantctomycetes tended to increase as soils aged.

Conclusions

There were trends in the dynamics of bacterial community composition and structure in soil during ecosystem development. Bacterial communities changed in ways that appear to be consistent with a model of ecosystem progression and retrogression, perhaps indicating fundamental processes underpin patterns of below and above-ground community change during ecosystem development.     

A unifying concept for the dependence of whole-crop N : P ratio on biomass: theory and experiment

Background and Aims

Numerous estimates have been made of the concentrations of N and P required for good growth of crop species but they have not been defined by any unifying model. The aim of the present study was to develop such a model for the dependence of the N : P ratio on crop mass, to test its validity and to use it to identify elements of similarity between different crop species and wild plants.

Methods

A model was derived between plant N : P ratio (R_w) and its dry biomass per unit area (W) during growth with near optimum nutrition by considering that plants consist of growth-related tissue and storage-related tissue with N : P ratios R_g and R_s, respectively. Testing and calibration against experimental data on different crop species led to a simple equation between R_w and W which was tested against independent experimental data.

Key Results

The validity of the model and equation was supported by 365 measurements of R_w in 38 field experiments on crops. R_g and R_s remained approximately constant throughout growth, with average values of 11·8 and 5·8 by mass. The model also approximately predicted the relationships between leaf N and P concentrations in 124 advisory estimates on immature tissues and in 385 wild species from published global surveys.

Conclusions

The N : P ratio of the biomass of very different crops, during growth with near optimum levels of nutrients, is defined entirely in terms of crop biomass, an average N : P ratio of the storage/structure-related tissue of the crop and an average N : P ratio of the growth-related tissue. The latter is similar to that found in leaves of many wild plant species, and even micro-organisms and terrestrial and freshwater autotrophs.Key words: Biomass, growth rate, model, N : P ratio, plant allometry, plant mass, stoichiometry     

Selenium accumulation in durum wheat and spring canola as a function of amending soils with selenite, selenate and or sulphate

Aims

A comparison was performed between plant species to determine if extractable, rather than total soil Se, is more effective at predicting plant Se accumulation over a full growing season.

Methods

Durum wheat (Triticum turgidum L.) and spring canola (Brassica napus L.) were sown in potted soil amended with 0, 0.1, 1.0, or 5.0 mg kg^?1 Se as SeO₄ ^2? or SeO₃ ^2?. In addition, SeO₄ ^2?-amended soils were amended with 0 or 50 mg kg^?1 S as SO₄ ^2?. Soils were analyzed for extractable and total concentration of Se ([Se]). Twice during the growing season plants were harvested and tissue [Se] was determined.

Results

Plants exposed to SeO₃ ^2? accumulated the least Se. Fitted predictive models for whole plant accumulation based on extractable soil [Se] were similar to models based on total [Se] in soil (R²?=?0.73 or 0.74, respectively) and selenium speciation and soil [S] were important soil parameters to consider. As well, soil S amendments limited Se toxicity.

Conclusions

Soil quality guidelines (SQGs) based on extractable Se should be considered for risk assessment, particularly when Se speciation is unknown. Predictive models to estimate plant Se uptake should include soil S, a modifier of Se accumulation.     

Contrasting correlation networks between leaf structure,nitrogen and chlorophyll in herbaceous and woody canopies

We studied acclimation patterns in leaf dry mass per area (M_A), nitrogen (N_A) and chlorophyll (ζ_A) content per area, and chlorophyll to nitrogen ratio (ζ/N) along vertical light gradients in natural temperate mixed herbaceous canopy and deciduous tree canopy. In the deciduous tree canopy, all leaves are formed at approximately the same time, and the light gradient during the rest of the growing season reflects the differences in light availability during leaf development, whereas in the herbaceous canopy, leaf production continues during most of the growing season and major changes in light conditions occur after leaf maturation. M_A and N_A increased strongly with increasing current light availability (I_D) in the tree canopy. In the herbaceous canopy, M_A and N_A were generally unrelated to I_D. Depending on species, the correlation between chlorophyll content per leaf area (ζ_A) and I_D was positive, negative, or non-significant. Path analyses revealed two opposite effects of I_D on the amount of leaf chlorophyll. In the tree canopy, increasing I_D enhanced ζ_A through changes in M_A and N_A, whereas the direct effect of light was negative in both canopies. The overall correlation network between foliage structural and chemical traits and the relationships with I_D were significantly stronger in the tree canopy, suggesting limited re-acclimation potential in the mixed herbaceous canopy. Within-species acclimation patterns reflected the patterns within the main functional types. These data demonstrate that the relationships of current light availability vs. leaf dry mass per area, leaf nitrogen and chlorophyll contents, and chlorophyll to nitrogen ratio differ among multi-species herbaceous canopies and deciduous tree canopies due to contrasting canopy development.     

Nitrogen:phosphorous supply ratio and allometry in five alpine plant species

In terrestrial ecosystems, atmospheric nitrogen (N) deposition has greatly increased N availability relative to other elements, particularly phosphorus (P). Alterations in the availability of N relative to P can affect plant growth rate and functional traits, as well as resource allocation to above‐ versus belowground biomass (M_A and M_B). Biomass allocation among individual plants is broadly size‐dependent, and this can often be described as an allometric relationship between M_A and M_B, as represented by the equation , or log M_A = logα + βlog M_B. Here, we investigated whether the scaling exponent or regression slope may be affected by the N:P supply ratio. We hypothesized that the regression slope between M_A and M_B should be steeper under a high N:P supply ratio due to P limitation, and shallower under a low N:P supply ratio due to N limitation. To test these hypotheses, we experimentally altered the levels of N, P, and the N:P supply ratio (from 1.7:1 to 135:1) provided to five alpine species representing two functional groups (grasses and composite forbs) under greenhouse conditions; we then measured the effects of these treatments on plant morphology and tissue content (SLA, leaf area, and leaf and root N/P concentrations) and on the scaling relationship between M_A and M_B. Unbalanced N:P supply ratios generally negatively affected plant biomass, leaf area, and tissue nutrient concentration in both grasses and composite forbs. High N:P ratios increased tissue N:P ratios in both functional groups, but more in the two composite forbs than in the grasses. The positive regression slopes between log M_A and log M_B exhibited by plants raised under a N:P supply ratio of 135:1 were significantly steeper than those observed under the N:P ratio of 1.7:1 and 15:1. Synthesis: Plant biomass allocation is highly plastic in response to variation in the N:P supply ratio. Studies of resource allocation of individual plants should focus on the effects of nutrient ratios as well as the availability of individual elements. The two forb species were more sensitive than grasses to unbalanced N:P supplies. To evaluate the adaptive significance of this plasticity, the effects of unbalanced N:P supply ratio on individual lifetime fitness must be measured.