Distinct Responses of Leaf Traits to Environment and Phylogeny Between Herbaceous and Woody Angiosperm Species in China

Leaf traits play key roles in plant resource acquisition and ecosystem processes; however, whether the effects of environment and phylogeny on leaf traits differ between herbaceous and woody species remains unclear. To address this, in this study, we collected data for five key leaf traits from 1,819 angiosperm species across 530 sites in China. The leaf traits included specific leaf area, leaf dry matter content, leaf area, leaf N concentration, and leaf P concentration, all of which are closely related to trade-offs between resource uptake and leaf construction. We quantified the relative contributions of environment variables and phylogeny to leaf trait variation for all species, as well as for herbaceous and woody species separately. We found that environmental factors explained most of the variation (44.4–65.5%) in leaf traits (compared with 3.9–23.3% for phylogeny). Climate variability and seasonality variables, in particular, mean temperature of the warmest and coldest seasons of a year (MTWM/MTWQ and MTCM/MTCQ) and mean precipitation in the wettest and driest seasons of a year (MPWM/MPWQ and MPDM/MPDQ), were more important drivers of leaf trait variation than mean annual temperature (MAT) and mean annual precipitation (MAP). Furthermore, the responses of leaf traits to environment variables and phylogeny differed between herbaceous and woody species. Our study demonstrated the different effects of environment variables and phylogeny on leaf traits among different plant growth forms, which is expected to advance the understanding of plant adaptive strategies and trait evolution under different environmental conditions.

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Article Variation and Tradeoff of Leaf Traits of Karst Shrub in Southwest China

10.21203/rs.3.rs-38026/v1 ◽

2020 ◽

Author(s):

zhigang zou ◽

Fuping Zeng ◽

Zhaoxia Zeng ◽

Jianxiong Liao ◽

Hu Du ◽

...

Keyword(s):

Leaf Area ◽

Southwest China ◽

Scientific Information ◽

Leaf Traits ◽

Karst Region ◽

Research Station ◽

Dry Matter Content ◽

Leaf Trait ◽

Environmental Selection ◽

Trade Offs

Abstract Long-term droughts were found to have guided the environmental selection ofshrub plant characteristics in a karst region of China, as the plants were found to have developed a set of leaf trait combinations,includinga small specific leaf area (SLA), leaf area (LA), and large leaf dry matter content (LDMC), that are known to be suitable for drought environments.Leaf traits of plants are not only the intuitive and operable taxonomic traits in plant taxonomy, but also reflect the responses and adaptations of plants to their habitats. This is helpful when trying to understand the role of environmental screening and when filtering plant functional traits. The objective of this investigation was to determine the leaf trait variations, adaptations, and patterns in the shrubs from a karst region in China.Weinvestigated 11 leaf traits to quantify the variations in their trade-offs and the trait–habitat /species relationships for the shrubs at theHuanjiang karst ecosystem observation and research station, China, using multivariate analyses.There were significant intraspecific and interspecific changes in the leaf traits ofthe shrub plants, and there were differences among the traits. Except for carbonmass, nitrogenarea, and phosphorousarea, the interspecific variations of the leaf traits were generally higher than the interspecific variation. The correlation between the leaf traits in the karst shrubs was also significant. Species differences had a higher explanatory degree for the leaf traits than topography or soil nutrients. The findings of this study will enhance our understanding of the variations in leaf traits in the karst shrubregions and the adaptative strategies of the plants in degraded habitats.Furthermore, these results may provide scientific information to help guide vegetation recovery programs in the karst region of southwest China.

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Responses of leaf traits to climatic gradients: adaptive variation versus compositional shifts

Biogeosciences ◽

10.5194/bg-12-5339-2015 ◽

2015 ◽

Vol 12 (18) ◽

pp. 5339-5352 ◽

Cited By ~ 28

Author(s):

T.-T. Meng ◽

H. Wang ◽

S. P. Harrison ◽

I. C. Prentice ◽

J. Ni ◽

...

Keyword(s):

Leaf Area ◽

Dry Matter ◽

Linear Models ◽

Environmental Gradients ◽

Empirical Studies ◽

Matter Content ◽

Dry Matter Content ◽

Adaptive Variation ◽

Trait Variation ◽

Leaf Trait

Abstract. Dynamic global vegetation models (DGVMs) typically rely on plant functional types (PFTs), which are assigned distinct environmental tolerances and replace one another progressively along environmental gradients. Fixed values of traits are assigned to each PFT; modelled trait variation along gradients is thus driven by PFT replacement. But empirical studies have revealed "universal" scaling relationships (quantitative trait variations with climate that are similar within and between species, PFTs and communities); and continuous, adaptive trait variation has been proposed to replace PFTs as the basis for next-generation DGVMs. Here we analyse quantitative leaf-trait variation on long temperature and moisture gradients in China with a view to understanding the relative importance of PFT replacement vs. continuous adaptive variation within PFTs. Leaf area (LA), specific leaf area (SLA), leaf dry matter content (LDMC) and nitrogen content of dry matter were measured on all species at 80 sites ranging from temperate to tropical climates and from dense forests to deserts. Chlorophyll fluorescence traits and carbon, phosphorus and potassium contents were measured at 47 sites. Generalized linear models were used to relate log-transformed trait values to growing-season temperature and moisture indices, with or without PFT identity as a predictor, and to test for differences in trait responses among PFTs. Continuous trait variation was found to be ubiquitous. Responses to moisture availability were generally similar within and between PFTs, but biophysical traits (LA, SLA and LDMC) of forbs and grasses responded differently from woody plants. SLA and LDMC responses to temperature were dominated by the prevalence of evergreen PFTs with thick, dense leaves at the warm end of the gradient. Nutrient (N, P and K) responses to climate gradients were generally similar within all PFTs. Area-based nutrients generally declined with moisture; Narea and Karea declined with temperature, but Parea increased with temperature. Although the adaptive nature of many of these trait-climate relationships is understood qualitatively, a key challenge for modelling is to predict them quantitatively. Models must take into account that community-level responses to climatic gradients can be influenced by shifts in PFT composition, such as the replacement of deciduous by evergreen trees, which may run either parallel or counter to trait variation within PFTs. The importance of PFT shifts varies among traits, being important for biophysical traits but less so for physiological and chemical traits. Finally, models should take account of the diversity of trait values that is found in all sites and PFTs, representing the "pool" of variation that is locally available for the natural adaptation of ecosystem function to environmental change.

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Responses of leaf traits to climatic gradients: adaptive variation vs. compositional shifts

Biogeosciences Discussions ◽

10.5194/bgd-12-7093-2015 ◽

2015 ◽

Vol 12 (9) ◽

pp. 7093-7124 ◽

Cited By ~ 1

Author(s):

T.-T. Meng ◽

H. Wang ◽

S. P. Harrison ◽

I. C. Prentice ◽

J. Ni ◽

...

Keyword(s):

Leaf Area ◽

Dry Matter ◽

Linear Models ◽

Environmental Gradients ◽

Empirical Studies ◽

Matter Content ◽

Dry Matter Content ◽

Adaptive Variation ◽

Trait Variation ◽

Leaf Trait

Abstract. Dynamic global vegetation models (DGVMs) typically rely on plant functional types (PFTs), which are assigned distinct environmental tolerances and replace one another progressively along environmental gradients. Fixed values of traits are assigned to each PFT; modelled trait variation along gradients is thus driven by PFT replacement. But empirical studies have revealed "universal" scaling relationships (quantitative trait variations with climate that are similar within and between species, PFTs and communities); and continuous, adaptive trait variation has been proposed to replace PFTs as the basis for next-generation DGVMs. Here we analyse quantitative leaf-trait variation on long temperature and moisture gradients in China with a view to understanding the relative importance of PFT replacement vs. continuous adaptive variation within PFTs. Leaf area (LA), specific leaf area (SLA), leaf dry matter content (LDMC) and nitrogen content of dry matter were measured on all species at 80 sites ranging from temperate to tropical climates and from dense forests to deserts. Chlorophyll fluorescence traits and carbon, phosphorus and potassium contents were measured at 47 sites. Generalized linear models were used to relate log-transformed trait values to growing-season temperature and moisture indices, with or without PFT identity as a predictor, and to test for differences in trait responses among PFTs. Continuous trait variation was found to be ubiquitous. Responses to moisture availability were generally similar within and between PFTs, but biophysical traits (LA, SLA and LDMC) of forbs and grasses responded differently from woody plants. SLA and LDMC responses to temperature were dominated by the prevalence of evergreen PFTs with thick, dense leaves at the warm end of the gradient. Nutrient (N, P and K) responses to climate gradients were generally similar within all PFTs. Area-based nutrients generally declined with moisture; Narea and Karea declined with temperature, but Parea increased with temperature. Although the adaptive nature of many of these trait–climate relationships is understood qualitatively, a key challenge for modelling is to predict them quantitatively. Models must also take into account that community-level responses to climatic gradients can be influenced by shifts in PFT composition, such as the replacement of deciduous by evergreen trees, which may run either parallel or counter to trait variation within PFTs. The importance of PFT shifts varies among traits, being important for biophysical traits but less so for physiological and chemical traits.

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Within-species leaf trait variation and ecological flexibility in resprouting tropical trees

Journal of Tropical Ecology ◽

10.1017/s0266467412000429 ◽

2012 ◽

Vol 28 (5) ◽

pp. 527-530 ◽

Cited By ~ 1

Author(s):

Carl F. Salk

Keyword(s):

Leaf Area ◽

Tree Species ◽

Environmental Changes ◽

Functional Trait ◽

Leaf Mass Per Area ◽

Trait Variation ◽

Leaf Functional Traits ◽

Leaf Trait ◽

Neotropical Tree ◽

Ecological Flexibility

Plants have an inherent flexibility to respond to different environmental conditions. One axis of plant ecophysiological strategy is seen in the spectrum of leaf functional traits. Flexibility in these traits would be suggestive of plants’ phenotypic plasticity in response to environmental changes. This research seeks to identify differences between leaves of sprout and non-sprout shoots of a broad ecological range of neotropical tree species. Using a functional-trait approach, this study assesses a large pool of species for within-species physiological flexibility. Leaf mass per area (LMA) and leaf area were measured for plants of sprout and non-sprout origin for 26 tree species grown in a reforestation plantation in Panama. Sprouts had a consistently lower LMA than non-sprouts, but there was no consistent pattern for leaf area. These trends show that sprouts are more like pioneer species than conspecific saplings, a finding in general agreement with fast sprout growth seen in previous studies. Further, later-successional (high LMA) species showed a greater reduction of LMA in sprouts. These results show that tropical tree species adjust physiologically to changing ecological roles and suggest that certain species may be more resilient than realized to changing climate and disturbance patterns.

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Interspecific covariation between stomatal density and other functional leaf traits in a local flora

Botany ◽

10.1139/b09-103 ◽

2010 ◽

Vol 88 (1) ◽

pp. 30-38 ◽

Cited By ~ 18

Author(s):

Jessy Loranger ◽

Bill Shipley

Keyword(s):

Chlorophyll Content ◽

Stomatal Density ◽

Leaf Traits ◽

Functional Ecology ◽

Leaf Trait ◽

Leaf Chlorophyll ◽

Sun And Shade Leaves ◽

Angiosperm Species ◽

Sun And Shade ◽

Shade Leaves

Despite the importance of stomata in leaf functioning, and despite the recent interest in interspecific leaf trait covariation in functional ecology, little is known about how stomatal density relates to other leaf traits in a broad interspecific context. This is especially important because stomatal density has been widely used to deduce temporal variation in atmospheric CO2 concentrations [CO2atm] from fossilized or herbarium leaves. We therefore measured stomatal density, specific leaf area (SLA) and its components, leaf thickness, and leaf chlorophyll content in both sun and shade leaves of 169 individuals from 52 angiosperm species in southwestern Quebec. Using mixed models, we show that stomatal density decreases allometrically with increasing SLA and chlorophyll content, and increases allometrically with increasing lamina thickness. The sun–shade contrast changes the intercepts, but not the slopes, of these relationships. It is important to take into consideration these relations when correlating stomatal density with [CO2], to avoid spurious interpretations.

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Leaf nitrogen from first principles: field evidence for adaptive variation with climate

10.5194/bg-2016-89 ◽

2016 ◽

Author(s):

Ning Dong ◽

Iain Colin Prentice ◽

Bradley J. Evans ◽

Stefan Caddy-Retalic ◽

Andrew J. Lowe ◽

...

Keyword(s):

Environmental Gradients ◽

Species Turnover ◽

Leaf Nitrogen ◽

Functional Ecology ◽

Leaf Mass Per Area ◽

Mean Annual Temperature ◽

Rubisco Activity ◽

Trait Variation ◽

Leaf Trait ◽

Field Evidence

Abstract. Nitrogen content per unit leaf area (Narea) is a key variable in plant functional ecology and biogeochemistry. Narea comprises a structural component, which scales with leaf mass per area (LMA), and a metabolic component, which scales with Rubisco capacity. The co-ordination hypothesis, as implemented in LPJ and related global vegetation models, predicts that Rubisco capacity should be directly proportional to irradiance but should decrease with ci:ca and temperature because the amount of Rubisco required to achieve a given assimilation rate declines with both. We tested these predictions using LMA, leaf δ13C and leaf N measurements on complete species assemblages sampled at sites on a North-South transect from tropical to temperate Australia. Partial effects of mean canopy irradiance, mean annual temperature and ci:ca (from δ13C) on Narea were all significant and their directions and magnitudes were in line with predictions. Over 80 % of the variance in community-mean (ln) Narea was accounted for by these predictors plus LMA. Moreover, Narea could be decomposed into two components, one proportional to LMA (slightly steeper in N-fixers), the other to predicted Rubisco activity. Trait gradient analysis revealed ci:ca to be perfectly plastic, while species turnover contributed about half the variation in LMA and Narea. Interest has surged in methods to predict continuous leaf-trait variation from environmental factors, in order to improve ecosystem models. Our results indicate that Narea has a useful degree of predictability, from a combination of LMA and ci:ca – themselves in part environmentally determined – with Rubisco activity, as predicted from local growing conditions. This is consistent with a 'plant-centred' approach to modelling, emphasizing the adaptive regulation of traits. Models that account for biodiversity will also need to partition community-level trait variation into components due to phenotypic plasticity and/or genotypic differentiation within species, versus progressive species replacement, along environmental gradients. Our analysis suggests that variation in Narea is about evenly split between these two modes.

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Reliability of leaf functional traits after delayed measurements

Australian Journal of Botany ◽

10.1071/bt19166 ◽

2020 ◽

Vol 68 (2) ◽

pp. 100

Author(s):

Dinesh Thakur ◽

Lakhbeer Singh ◽

Amit Chawla

Keyword(s):

Leaf Area ◽

Leaf Traits ◽

Western Himalaya ◽

Matter Content ◽

Dry Matter Content ◽

Leaf Dry Matter Content ◽

Leaf Functional Traits ◽

Temporary Storage ◽

Delayed Measurements ◽

Stable Leaf

In this study, the effect of temporary storage (at 4°C) on measurement of leaf traits was tested. We collected leaf samples from 25 species, which represented different functional types in the high altitude vegetation of western Himalaya, to measure leaf area (LA), leaf rehydration, specific leaf area (SLA) and leaf dry matter content (LDMC). Repeated trait measurements were performed for up to 7 days. We found that in all the species, LA increased in initial 24 h of rehydration and thereafter remained stable. Leaf rehydration was found to be sensitive to delayed measurements and changed significantly for up to 7 days. For SLA and LDMC, the effect of storage time was significant only for a few species. On the basis of our findings, we recommend that, for samples stored in dark at 4°C, LA, SLA and LDMC can reliably be estimated after a delay of up to 7 days. Further, these key leaf traits should be estimated only after 24 h of rehydration. Also, trait measurements after prolonged rehydration of leaves should be avoided. Outcomes of this study will be beneficial when a large number of samples are collected from locations far away from laboratory and temporary storage is necessitated before trait measurements.

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Seasonal variation in foliar properties in Mediterranean Pine forests of different post-fire age

10.5194/egusphere-egu21-1064 ◽

2021 ◽

Author(s):

Christodoulos I Sazeides ◽

Nikolaos M Fyllas ◽

Anastasia Christopoulou

Keyword(s):

Leaf Area ◽

Intraspecific Variation ◽

Light Response ◽

Leaf Traits ◽

Seasonal Patterns ◽

Dry Matter Content ◽

Sapwood Area ◽

Pinus Brutia ◽

Foliar Traits ◽

Trait Variability

<p>Foliar properties play a crucial role in local and global biochemical cycles. Systematic variation in key leaf traits has been reported both between and within species. Intraspecific variation in leaf traits is controlled by micro-environmental conditions and follows seasonal patterns. In this study we examine the seasonal patterns of six foliar traits including leaf area (LA), leaf thickness (Lth), leaf mass per area (LMA), leaf dry matter content (LDMC), leaf area to sapwood area ratio (LA/SA) and branch wood density (WD) in addition to the associated parameters of the Michaelis-Menten light response curve (i.e. light saturated net photosynthetic rate (Asat), half saturation coefficient (Km) and dark respiration rate (Rd)). We measured on a monthly basis the foliar traits and developed light response curves in four Pinus brutia dominated stands along a post-fire chronosequence (15, 40, 70 and 90 years) from sunlit branches. Significant differences in the interannual trait variability were found between stands for LDMC, WD and Asat, with the highest variability identified in the younger plot. LA/SA, Rd and Km also showed strong interannual variability although not statistically different between plots. A mixed effect model analysis revealed high intraclass correlation coefficients for Km and Asat suggesting that net photosynthesis is following systematic seasonal patterns. Overall LA was higher and LDMC was lower in the oldest plot and WD was higher in the denser (40 years) plot. Interestingly gas exchange parameters did not show differences in their overall mean values. Across plots, Asat was strongly positively related to Km, and LMA was positively related to LDMC and Lth. LDMC was also positively related with Asat and negatively with Lth. A principal component analysis (PCA) revealed two major dimensions of intraspecific trait variability within our plots. The first PCA axis was positively related to Asat, Km, LDMC and LMA suggesting that regardless of the stand age needles are placed along a fast-slow carbon gain dimension with denser needles illustrating faster area-based photosynthesis. The second PCA axis was positively related to LA and Lth suggesting that bigger needles are also thicker. A subsequent permutational multivariate analysis of variance revealed that the centroids and the dispersion of the trait syndromes differed between stands, with the youngest plot illustrating higher trait dispersion and the oldest plot characterized by bigger and thicker needles. Thus, in older stands were competition for light is higher, needles are deployed to be bigger and thicker to optimize light capture, while in younger stands they are optimized along a leaf density - photosynthetic capacity spectrum depending on (more heterogeneous) microenvironmental conditions. Our findings illustrate that intraspecific variation can be attributed to either seasonal (abiotic) light availability or to (biotic) heterogeneity related to stand structure, and have important implications for local scale forest dynamics models.</p><p>&#171;This research is co-financed by Greece and the European Union (European Social Fund- ESF) through the Operational Programme &#171;Human Resources Development, Education and Lifelong Learning 2014-2020&#187; in the context of the project &#8220;Carbon fluxes across a post-fire chronosequence in Pinus brutia Ten forests.&#8221; (MIS 5049513)&#187;.</p>

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Effects of grazing on leaf traits and ecosystem functioning in Inner Mongolia grasslands: scaling from species to community

Biogeosciences Discussions ◽

10.5194/bgd-6-9945-2009 ◽

2009 ◽

Vol 6 (5) ◽

pp. 9945-9975 ◽

Cited By ~ 4

Author(s):

S. X. Zheng ◽

H. Y. Ren ◽

Z. C. Lan ◽

W. H. Li ◽

Y. F. Bai

Keyword(s):

Leaf Area ◽

River Basin ◽

Ecosystem Functioning ◽

Leaf Traits ◽

Northern China ◽

Leaf Biomass ◽

Area Index ◽

Semiarid Grassland ◽

Leaf Trait ◽

Leaf Mass

Abstract. More attention has focused on using some easily measured plant functional traits to predict grazing influence on plant growth and ecosystem functioning. However, there has been much controversy on leaf traits response to grazing, thus more research should be conducted at the species level. Here we investigated the leaf area, leaf mass and specific leaf area (SLA) of 263 species in eight grassland communities along a soil moisture gradient in the Xilin River Basin, a semiarid grassland of northern China, to explore the grazing effects on ecosystem functioning. Results demonstrated that grazing decreased the leaf area and leaf mass in more than 56% of species in the Xilin River Basin, however, responses of SLA to grazing varied widely between species. Grazing increased SLA in 38.4% of species, decreased SLA in 31.3% of species and had no effect on 30.3% of species. Annuals and biennials generally developed high SLA as grazing tolerance traits, while perennial graminoids developed low SLA as grazing avoidance traits. Considering the water ecotypes, the SLA-increased and SLA-unchanged species were dominated by hygrophytes and mesophytes, while the SLA-decreased species were dominated by xerophytes. At the community level, grazing decreased the mean leaf area index (LAI) of six communities by 16.9%, leaf biomass by 35.2% and standing aboveground biomass (SAB) by 35.0% in the Xilin River Basin, indicating that overgrazing greatly decreased the ecosystem functioning in the semi-arid grassland of northern China. Soil properties, especially fielding holding capacity and soil organic carbon and total nitrogen could mediate the negative grazing impacts. The results suggest SLA is a better leaf trait to reveal plant adaptability to grazing. Our findings have practical implications for range management and productivity maintenance in the semiarid grassland, and it is feasible to take some measures such as ameliorating soil water and nutrient availabilities to prevent grassland degradation.

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Scaling of petiole anatomies, mechanics and vasculatures with leaf size in the widespread Neotropical pioneer tree species Cecropia obtusa Trécul (Urticaceae)

Tree Physiology ◽

10.1093/treephys/tpz136 ◽

2020 ◽

Vol 40 (2) ◽

pp. 245-258 ◽

Cited By ~ 1

Author(s):

Sébastien Levionnois ◽

Sabrina Coste ◽

Eric Nicolini ◽

Clément Stahl ◽

Hélène Morel ◽

...

Keyword(s):

Leaf Area ◽

Leaf Size ◽

Leaf Traits ◽

Hydraulic Efficiency ◽

Cross Sectional ◽

Leaf Trait ◽

Key Factor ◽

Leaf Economic Spectrum ◽

Geometrical Effect ◽

Trait Variability

Abstract Although the leaf economic spectrum has deepened our understanding of leaf trait variability, little is known about how leaf traits scale with leaf area. This uncertainty has resulted in the assumption that leaf traits should vary by keeping the same pace of variation with increases in leaf area across the leaf size range. We evaluated the scaling of morphological, tissue-surface and vascular traits with overall leaf area, and the functional significance of such scaling. We examined 1,271 leaves for morphological traits, and 124 leaves for anatomical and hydraulic traits, from 38 trees of Cecropia obtusa Trécul (Urticaceae) in French Guiana. Cecropia is a Neotropical genus of pioneer trees that can exhibit large laminas (0.4 m2 for C. obtusa), with leaf size ranging by two orders of magnitude. We measured (i) tissue fractions within petioles and their second moment of area, (ii) theoretical xylem hydraulic efficiency of petioles and (iii) the extent of leaf vessel widening within the hydraulic path. We found that different scaling of morphological trait variability allows for optimisation of lamina display among larger leaves, especially the positive allometric relationship between lamina area and petiole cross-sectional area. Increasing the fraction of pith is a key factor that increases the geometrical effect of supportive tissues on mechanical rigidity and thereby increases carbon-use efficiency. We found that increasing xylem hydraulic efficiency with vessel size results in lower leaf lamina area: xylem ratios, which also results in potential carbon savings for large leaves. We found that the vessel widening is consistent with hydraulic optimisation models. Leaf size variability modifies scaling of leaf traits in this large-leaved species.

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