Quantifying foliar trait variation and covariation in sun and shade leaves using leaf spectroscopy in eastern North America

Characterizing foliar trait variation in sun and shade leaves can provide insights into inter-and intra-species resource use strategies and plant response to environmental change.However,datasets with records of multiple foliar traits from the same individual and including shade leaves are sparse,which limits our ability to investigate trait-trait,trait-environment relationships and trait coordination in both sun and shade leaves.We presented a comprehensive dataset of 15 foliar traits from sun and shade leaves sampled with leaf spectroscopy,including 424 individuals of 110 plant species from 19 sites across eastern North America.We investigated trait variation,covariation,scaling relationships with leaf mass,and the effects of environment,canopy position,and taxonomy on trait expression.Generally,sun leaves had higher leaf mass per area,nonstructural carbohydrates and total phenolics,lower mass-based chlorophyll a+b,carotenoids,phosphorus,and potassium,but exhibited species-specific characteristics.Covariation between sun and shade leaf traits,and trait-environment relationships were overall consistent across species.The main dimensions of foliar trait variation in seed plants were revealed including leaf economics traits,photosynthetic pigments,defense,and structural traits.Taxonomy and canopy position collectively explained most of the foliar trait variation.This study highlights the importance of including intra-individual and intra-specific trait variation to improve our understanding of ecosystem functions.Our findings have implications for efficient field sampling,and trait mapping with remote sensing.

Extremely thin but very robust:Surprising cryptogam trait combinations at the end of the leaf economics spectrum

Leaf economics spectrum(LES)describes the fundamental trade-offs between leaf structural,chemical,and physiological investments.Generally,structurally robust thick leaves with high leaf dry mass per unit area(LMA)exhibit lower photosynthetic capacity per dry mass(Amass).Paradoxically,“soft and thinleaved”mosses and spikemosses have very low Amass,but due to minute-size foliage elements,their LMA and its components,leaf thickness(LT)and density(LD),have not been systematically estimated.Here,we characterized LES and associated traits in cryptogams in unprecedented details,covering five evolutionarily different lineages.We found that mosses and spikemosses had the lowest LMA and LT values ever measured for terrestrial plants.Across a broad range of species from different lineages,Amass and LD were negatively correlated.In contrast,Amass was only related to LMA when LMA was greater than 14 g cm^(-2).In fact,low Amass reflected high LD and cell wall thickness in the studied cryptogams.We conclude that evolutionarily old plant lineages attained poorly differentiated,ultrathin mesophyll by increasing LD.Across plant lineages,LD,not LMA,is the trait that represents the trade-off between leaf robustness and physiology in the LES.

Dominance of rock exposure and soil depth in leaf trait networks outweighs soil quality in karst limestone and dolomite habitats

Leaf trait networks(LTNs)visualize the intricate linkages reflecting plant trait-functional coordination.Typical karst vegetation,developed from lithological dolomite and limestone,generally exhibits differential communities,possibly due to habitat rock exposure,soil depth,and soil physicochemical properties variations,leading to a shift from plant trait variation to functional linkages.However,how soil and habitat quality affect the differentiation of leaf trait networks remains unclear.LTNs were constructed for typical dolomite and limestone habitats by analyzing twenty-one woody plant leaf traits across fifty-six forest subplots in karst mountains.The differences between dolomite and limestone LTNs were compared using network parameters.The network association of soil and habitat quality was analyzed using redundancy analysis(RDA),Mantle's test,and a random forest model.The limestone LTN exhibited significantly higher edge density with lower diameter and average path length when compared to the dolomite LTN.It indicates LTN differentiation,with the limestone network displaying a more compact architecture and higher connectivity than the dolomite network.The specific leaf phosphorus and leaf nitrogen contents of dolomite LTN,as well as the leaf mass and leaf carbon contents of limestone LTN,significantly contributed to network degree and closeness,serving as crucial node traits regulating LTN connectedness.Additionally,both habitat LTNs significantly correlated with soil nitrogen and phosphorus,stoichiometric ratios,pH,and organic carbon,as well as soil depth and rock exposure rates,with soil depth and rock exposure showing greater relative importance.Soil depth and rock exposure dominate trait network differentiation,with the limestone habitat exhibiting a more compact network architecture than the dolomite habitat.

Unveiling the adaptation strategies of woody plants in remnant forest patches to spatiotemporal urban expansion through leaf trait networks

Background:With the expansion of urban areas,the remnants of forested areas play a crucial role in preserving biodiversity in urban environments.This study aimed to explore the impact of spatiotemporal urban expansion on the networks of leaf traits in woody plants within remnant forest patches,thereby enhancing our understanding of plant adaptive strategies and contributing to the conservation of urban biodiversity.Methods:Our study examined woody plants within 120 sample plots across 15 remnant forest patches in Guiyang,China.We constructed leaf trait networks (LTNs) based on 26 anatomical,structural,and compositional leaf traits and assessed the effects of the spatiotemporal dynamics of urban expansion on these LTNs.Results and conclusions:Our results indicate that shrubs within these patches have greater average path lengths and diameters than trees.With increasing urban expansion intensity,we observed a rise in the edge density of the LTN-shrubs.Additionally,modularity within the networks of shrubs decreased as road density and urban expansion intensity increased,and increases in the average path length and average clustering coefficient for shrubs were observed with a rise in the composite terrain complexity index.Notably,patches subjected to‘leapfrog’expansion exhibited greater average patch length and diameter than those experiencing edge growth.Stomatal traits were found to have high degree centrality within these networks,signifying their substantial contribution to multiple functions.In urban remnant forests,shrubs bolster their resilience to variable environmental pressures by augmenting the complexity of their leaf trait networks.

Geographical differences of leaf traits of the endangered plant Litsea coreana Levl.var.sinensis and its relationship with climate

Seventeen morphological and anatomical characteristics of the leaves were selected from five natural populations to explore the variation in leaf traits of Lits ea core ana var.sinensis and the effects of geographical environment on these variations.Nested analysis of variance,multiple comparisons,principal component analysis(PCA),and correlation analysis were conducted to explore the variations within and between populations and their correlation with geographical and climatic factors.Significant differences in the 17 leaf traits were observed within and among populations.On average,the relative contribution of within population variation to total variation was 24.8%,which was lower than among population variation(54.6%).The average differentiation coefficient of the traits was 65.8%,and the average coefficient of variation 11.8%,ranging from6.7%for main vein thickness to 21.4%for petiole length.The PC A results showed that morphological characteristics were divided into two categories,and the level of variation was greater than that of leaf anatomy.Most of the leaf traits were significantly correlated with geography and climate and showed a gradual variation with longitude,latitude,and altitude.In areas with high temperatures,less rainfall,and strong seasonal rainfall,the leaves are larger,longer and thicker.This study shows that variations in leaf traits of L.coreana var.sinensis mainly come from variations among populations.The level of trait differentiation among populations is high and the level of variation within populations low.These findings help further understand leaf morphological characteristics of this species and can provide a valuable reference for the protection and sustainable utilization of this natural resource.

High-throughput phenotyping of plant leaf morphological, physiological,and biochemical traits on multiple scales using optical sensing

Acquisition of plant phenotypic information facilitates plant breeding, sheds light on gene action, and can be applied to optimize the quality of agricultural and forestry products. Because leaves often show the fastest responses to external environmental stimuli, leaf phenotypic traits are indicators of plant growth,health, and stress levels. Combination of new imaging sensors, image processing, and data analytics permits measurement over the full life span of plants at high temporal resolution and at several organizational levels from organs to individual plants to field populations of plants. We review the optical sensors and associated data analytics used for measuring morphological, physiological, and biochemical traits of plant leaves on multiple scales. We summarize the characteristics, advantages and limitations of optical sensing and data-processing methods applied in various plant phenotyping scenarios. Finally, we discuss the future prospects of plant leaf phenotyping research. This review aims to help researchers choose appropriate optical sensors and data processing methods to acquire plant leaf phenotypes rapidly,accurately, and cost-effectively.

Soil water content and nitrogen differentially correlate with multidimensional leaf traits of two temperate broadleaf species

The variation and correlation of leaf economics and vein traits are crucial for predicting plant ecological strategies under different environmental changes.However,correlations between these two suites of traits and abiotic factors such as soil water and nitrogen content remain ambiguous.We measured leaf economics and vein traits as well as soil water and nitrogen content for two different shade-tolerant species(Betula platyphylla and Acer mono)in four mixed broadleaved-Korean pine(Pinus koraiensis)forests along a latitudinal gradient in Northeast China.We found that leaf economics traits and vein traits were decoupled in shade-intolerant species,Betula platphylla,but significantly coupled in a shadetolerant species,A.mono.We found stronger correlations among leaf traits in the shade tolerant species than in the shade intolerant species.Furthermore,leaf economic traits were positively correlated with the soil water gradient for both species,whereas vein traits were positively correlated with soil water gradient for the shade intolerant species but negatively correlated in the shade tolerant species.Although economic traits were positively correlated with soil nitrogen gradient in shade intolerant species but not correlated in shade tolerant species,vein traits were negatively correlated with soil nitrogen gradient in shade tolerant species but not correlated in shade intolerant species.Our study provides evidence for distinct correlations between leaf economics and vein traits and local abiotic factors of species differing in light demands.We recommend that the ecological significance of shade tolerance be considered for species when evaluating ecosystem functions and predicting plant responses to environmental changes.

Responses of maize germination,root morphology and leaf trait to characteristics of lead pollution:a case study

On base of the content of Pb in the soil under different land use patterns in Lanping Lead-zinc mining area,Yunnan in southwest China,the root morphology and leaf traits of maize in different concentration Pb(20,40,60,80,100,150,200,500,1000,2000,3000 mg/L)were analyzed.The results showed that maize germination rate,germination vigor and growth index decreased with the increase of Pb concentration.The root length,surface area of maize increased by 0.21%-81.58%,8.99%-73.43%,1.50%-77.37%,respectively,under 20-500 mg/L Pb concentration.However,these parameters under 1000-3000 mg/L Pb concentration decreased by 37.86%-553.54%,44.99%-766.16%,55.99%-92.81%,respectively,and these lowest value appeared in 3000 mg/L Pb treatment.The root volume of maize increased by 4.57%-89.25%in 20-80 mg/L Pb concentration,and it decreased with the increase of Pb concentration when the Pb concentration was higher than 80 mg/L and decreased by 94.13%in 3000 mg/L Pb.The root surface area and length of 0.50-1.00 diameter class were higher than those of other diameter classes,and these value of maize under 500 mg/L Pb were higher than those of other concentrations.The length and perimeter of maize leaves with the highest value of 220.36 and 962.68 mm,respectively appeared in 60 mg/L Pb treatment.The leaf width and area of maize with the highest value of 15.68 mm and 2448.31 mm^(2),respectively,appeared in 40 mg/L Pb treatment,which indicated that the leaf traits of maize were promoted by low concentration Pb and inhibited by high concentration Pb.

Metabolic profiles and morphological characteristics of leaf tips among different sweet potato(Ipomoea batatas Lam.)varieties

Sweet potato leaf tips have high nutritional value,and exploring the differences in the metabolic profiles of leaf tips among different sweet potato varieties can provide information to improve their qualities.In this study,a UPLC-Q-Exactive Orbitrap/MS-based untargeted metabolomics method was used to evaluate the metabolites in leaf tips of 32 sweet potato varieties.Three varieties with distinct overall metabolic profiles(A01,A02,and A03),two varieties with distinct profiles of phenolic acids(A20 and A18),and three varieties with distinct profiles of flavonoids(A05,A12,and A16)were identified.In addition,a total of 163 and 29 differentially expressed metabolites correlated with the color and leaf shape of sweet potato leaf tips,respectively,were identified through morphological characterization.Group comparison analysis of the phenotypic traits and a metabolite-phenotypic trait correlation analysis indicated that the color differences of sweet potato leaf tips were markedly associated with flavonoids.Also,the level of polyphenols was correlated with the leaf shape of sweet potato leaf tips,with lobed leaf types having higher levels of polyphenols than the entire leaf types.The findings on the metabolic profiles and differentially expressed metabolites associated with the morphology of sweet potato leaf tips can provide useful information for breeding sweet potato varieties with higher nutritional value.

QTL analysis of ear leaf traits in maize(Zea mays L.)under different planting densities

Modem maize varieties have become more productive than ever, owing largely to increased tolerance of high plant density. However, the genetics of ear leaf traits under different densities remains poorly understood. In this study, Zhongdan 909 recombinant inbred lines(RILs) derived from a cross between Z58 and HD568 were genotyped for 3072 single-nucleotide polymorphisms(SNPs), and phenotyped for leaf length(LL), leaf width(LW), and leaf angle(LA) of the uppermost ear leaf under three planting densities(52,500,67,500, and 82,500 plants ha-1, respectively). A genetic map was then constructed using1358 high-quality SNPs. The total length of the linkage map was 1985.2 cM and the average interval between adjacent markers 1.46 cM. With increasing density, LL and LW decreased from 63.68 to 63.02 cm and from 8.56 to 8.21 cm, respectively, while LA increased from19.42° to 19.66°. All three traits had high heritabilities, of 0.75, 0.78, and 0.84, respectively.Using inclusive composite interval mapping, 23, 25, and 17 quantitative trait loci(QTL) were detected for LL, LW, and LA, respectively. Of these, 35 were simultaneously detected under two or three plant densities, while 30 were detected under only one. Sixty-five individual QTL explained 2.41% to 16.53% of phenotypic variation, while eight accounted for >10%.These findings will help us understand the genetic basis of leaf traits in maize as well as the response of maize to increased plant density.

Differences in leaf functional traits between exotic and native Compositae plant species

This study aims to determine the differences in leaf functional traits and phenotypic plasticity of leaf functional traits between exotic and native Compositae plant species. Leaf width of exotic plants was significantly lower than that of native species. Leaf length, specific leaf area(SLA), single-leaf wet and dry weights, leaf moisture, and leaf thickness of exotic plants were also lower than those of native species but not significantly. The leaf shape index of exotic plants was higher than that of native species but not significantly. This implies that the relatively low leaf construction cost for exotic plants may play an important role in the success of their invasions. The higher leaf shape index and lower leaf width of exotic plants can enhance the efficiency of resource capture(especially sunlight capture) via adjustments to leaf shape and size, thereby increasing the survival of exotic plants. The plasticity indices of single-leaf wet weight and leaf thickness of exotic plants were significantly lower than those of native species. The lower phenotypic plasticity of single-leaf wet weight and leaf thickness of exotic plants may be the result of a cost to plasticity. That is, if the plasticity is too high, the fitness of plant species might be reduced sharply under unfavorable environments. Thus, lower plasticity of leaf functional traits may compensate for the negative impact of adverse environments and stabilize leaf construction costs for exotic plants. Moreover, reduced phenotypic plasticity might be one of the key competitive strategies by which exotic plants successfully invade new habitats. Overall, exotic plants did not always exhibit higher values of leaf functional traits or increased phenotypic plasticity of leaf functional traits compared with native species.

Identification of quantitative trait loci for the dead leaf rate and the seedling dead rate under alkaline stress in rice

The quantitative trait loci （QTLs） for the dead leaf rate （DLR） and the dead seedling rate （DSR） at the different rice growing periods after transplanting under alkaline stress were identified using an F2：3 population, which included 200 individuals and lines derived from a cross between two japonica rice cultivars Gaochan 106 and Changbai 9 with microsatellite markers. The DLR detected at 20 days to 62 days after transplanting under alkaline stress showed continuous normal or near normal distributions in F3 lines, which was the quantitative trait controlled by multiple genes. The DSR showed a continuous distribution with 3 or 4 peaks and was the quantitative trait controlled by main and multiple genes when rice was grown for 62 days after transplanting under alkaline stress. Thirteen QTLs associated with DLR were detected at 20 days to 62 days after transplanting under alkaline stress. Among these, qDLR9-2 located in RM5786-RM160 on chromosome 9 was detected at 34 days, 41 days, 48 days, 55 days, and 62 days, respectively; qDLR4 located in RM3524-RM3866 on chromosome 4 was detected at 34 days, 41 days, and 48 days, respectively; qDLR7-1 located in RM3859-RM320 on chromosome 7 was detected at 20 days and 27 days; and qDLR6-2 in RM1340-RM5957 on chromosome 6 was detected at 55 days and 62 days, respectively. The alleles of both qDLR9-2 and qDLR4 were derived from alkaline sensitive parent ＂Gaochanl06＂. The alleles of both qDLR7-1 and qDLR6-2 were from alkaline tolerant parent Changbai 9. These gene actions showed dominance and over dominance primarily. Six QTLs associated with DSR were detected at 62 days after transplanting under alkaline stress. Among these, qDSR6-2 and qDSR8 were located in RM1340-RM5957 on chromosome 6 and in RM3752-RM404 on chromosome 8, respectively, which were associated with DSR and accounted for 20.32% and 18.86% of the observed phenotypic variation, respectively; qDSR11-2 and qDSR11-3 were located in RM536-RM479 and RM2596-RM286 on chromosome 11, respectively, which were associated with DSR explaining 25.85% and 15.41% of the observed phenotypic variation, respectively. The marker flanking distances of these QTLs were quite far except that of qDSR6-2, which should be researched further.

Differences in leaf functional traits between red and green leaves of two evergreen shrubs Photinia×fraseri and Osmanthus fragrans

Leaf functional traits are adaptations that enable plants to live under different environmental conditions. This study aims to evaluate the differences in leaf functional traits between red and green leaves of two evergreen shrubs Photinia × fraseri and Osrnanthus fragrans. Specific areas of red leaves are higher than that of green leaves in both species. Thus, the material investment per unit area and per lamina of red leaves is significantly lower than that of green leaves, implying an utmost effort of red leaves to increase light capture and use efficiency because of their low leafchlorophyll concentration. The higher petiole length of green leaves compared with that of red leaves indicates that adult green leaves may have large fractional biomass allocation to support the lamina structures in capturing light with maximum efficiency and obtaining a high growth rate. The high range of the phenotypic plasticity of leaf size, leaf thickness, single-leaf wet and dry weights, and leaf moisture of green leaves may be beneficial in achieving efficient control of water loss and nutrient deprivation. The high range of phenotypic plasticity of leaf chlorophyll concentration of red leaves may be advantageous in increasing resource （especially light） capture anduse efficiency because this leaf type is juvenile in the growth stage and has low leaf-chlorophyll concentration.

Seasonal variations of leaf traits and drought adaptation strategies of four common woody species in South Texas, USA

Understanding physiological responses and drought adaptation strategies of woody plant leaf traits in sub-humid to semi-arid regions is of vital importance to understand the interplay between ecological processes and plant resource-allocation strategies of different tree species.Seasonal variations of leaf morphological traits,stoichiometric traits and their relationships of two drought tolerant woody species,live oak(Quercus virginiana)and honey mesquite(Prosopis glandulosa)and two less drought tolerant species,sugarberry(Celtis laevigata)and white ash(Fraxinus americana)were analyzed in a sub-humid to semi-arid area of south Texas,USA.Our findings demonstrate that for the two drought tolerant species,the leguminous P.glandulosa had the highest specific leaf area,leaf N,P,and lowest leaf area and dry mass,indicating that P.glandulosa adapts to an arid habitat by decreasing leaf area,thus reducing water loss,reflecting a resource acquisition strategy.While the evergreen species Q.virginiana exhibited higher leaf dry mass,leaf dry matter content,C content,C:N,C:P and N:P ratios,adapts to an arid habitat through increased leaf thickness and thus reduced water loss,reflecting a resource conservation strategy in south Texas.For the two less drought tolerant deciduous species,the variations of leaf traits in C.laevigata and F.americana varied between Q.virginiana and P.glandulosa,reflecting a trade-off between rapid plant growth and nutrient maintenance in a semi-arid environment.

Leaf Traits and Antioxidant Defense for Drought Tolerance During Early Growth Stage in Some Popular Traditional Rice Landraces from Koraput, India

Three popular traditional rice landraces, namely Kalajeera, Machakanta and Haladichudi, from Koraput, India were used to analyse the leaf traits and antioxidant defence for drought tolerance. When rice plants were exposed to different levels of drought stress by varying concentrations of polyethylene glycol （PEG） 6000, seed germination and growth parameters were significantly declined in all the rice landraces compared to the control. Drought stress also altered the leaf phenotypic traits based on chlorophyll fluorescence parameters and chlorophyll index, with more significant differences in susceptible variety IR64 than in traditional landraces. Furthermore, activities of antioxidative enzymes and proline and protein contents overtly increased under drought stress. The traditional rice landraces showed higher relative ratios for different parameters compared to the susceptible variety IR64. Taken together, the traditional landraces had superior leaf physiological efficiency compared to the susceptible and tolerant check varieties under drought stress.

Higher leaf area through leaf width and lower leaf angle were the primary morphological traits for yield advantage of japonica/indica hybrids

The yield potential of japonica/indica hybrids(JIH)has been achieved over 13.5 t ha–1 in large-scale rice fields,and some physiological traits for yield advantage of JIH over japonica inbred rice(JI)and indica hybrid rice(IH)were also identified.To date,little attention has been paid to morphological traits for yield advantage of JIH over JI and IH.For this reason,three JIH,three JI,and three IH were field-grown at East China(Ningbo,Zhejiang Province)in 2015 and 2016.Compared with JI and IH,JIH had 14.3 and 20.8%higher grain yield,respectively,attributed to its more spikelets per panicle and relatively high percentage of filled grains.The advantage in spikelets per panicle of JIH over JI and IH was shown in number of grains on the upper,middle,and lower branches.Compared with JI and IH,JIH had higher leaf area through leaf width and lower leaf angle of upper three leaves,higher leaf area index and leaf area per tiller at heading and maturity stages,higher stem weight per tiller and K and Si concentrations of stem at maturity,higher dry matter weight in leaf,stem,and panicle at heading and maturity stages,and higher biomass accumulation after heading and lower biomass translocation from stem during ripening.Leaf width of upper three leaves were correlated positively,while leaf angle of upper three leaves were correlated negatively with biomass accumulation after heading,stem weight per tiller,and per unit length.Our results indicated that the grain yield advantage of JIH was ascribed mainly to the more spikelets per panicle and relatively high percentage of filled grains.Higher leaf area through leaf width and more erect leaves were associated with improved biomass accumulation and stem weighing during ripening,and were the primary morphological traits underlying higher grain yield of JIH.

Elevation gradient distribution of indices of tree population in a montane forest:The role of leaf traits and the environment

Background:To disentangle the controls on species distribution in the context of climate change is a central element in proposed strategies to maintain species diversity.However,previous studies have focused mainly on the roles of abiotic factors(e.g.,climate and soil properties),with much less attention given to the roles of biotic factors such as functional traits.Here,we measured eight leaf traits for 240 individual trees of 53 species and analyzed the variation in traits and population composition indices and their relationships with soil properties,climate factors,and leaf traits.Results:The tree density,frequency and species importance values of the overall species and saplings significantly increased with increasing elevation,while the same indices(except for species frequency)of adults did not significantly change.The largest percentage of variation of species importance value(greater than 50%)was explained by climate,but leaf traits played a critical role in driving elevation distribution patterns of both saplings and adults;the abundance of saplings significantly increased with elevation,with increased leaf carbon contents,while the abundance of adults did not change in accordance with a nutrient conservation strategy associated with the leaf economic spectrum.Conclusions:Our results suggest that the elevation gradient distribution of woody plant species is dependent on tree size and that local atmospheric humidity and leaf traits cause considerable variation in species distribution along subtropical mountain elevations.We provide evidence of which leaf traits play a key role in the elevation gradient distribution of different sizes of woody tree species.

QTL analysis of flag leaf in barley (Hordeum vulgare L.) for morphological traits and chlorophyll content

To understand genetic patterns of the morphological and physiological traits in flag leaf of barley, a double haploid （DH） population derived from the parents Yerong and Franklin was used to determine quantitative trait loci （QTL） controlling length, width, length/width, and chlorophyll content of flag leaves. A total of 9 QTLs showing significantly additive effect were detected in 8 intervals on 5 chromosomes. The variation of individual QTL ranged from 1.9% to 20.2%. For chlorophyll content expressed as SPAD value, 4 QTLs were identified on chromosomes 2H, 3H and 6H; for leaf length and width, 2 QTLs located on chromosomes 5H and 7H, and 2 QTLs located on chromosome 5H were detected; and for length/width, I QTL was detected on chromosome 7H. The identification of these QTLs associated with the properties of flag leaf is useful for barley improvement in breeding programs.

Changes in leaf vein traits among vein types of alpine grassland plants on the Tibetan Plateau

Vein traits influence photosynthesis and drought resistance and are sensitive to climate change.It is unclear whether vein traits,similar to other leaf traits,have obvious regional characteristics and covariance with other leaf traits,especially in Tibetan Plateau grasslands.We collected 66 species from 37 sites in late July of 2012 and early August of 2013 to investigate leaf vein traits and their relationships with other leaf traits in comparison with the available global database data and to elucidate vein investment of plants with different vein types.The average vein length per area(VLA)of plants in the Tibetan Plateau was within the range of the global dataset,and the relationships among vein traits and other leaf traits of alpine plants were consistent with the global models.The VLAs of parallel-veined grasses and pinnateveined forbs were significantly lower and higher than the global mean value,respectively.The leaf mass perarea and total nitrogen content of parallel-veined grasses were significantly lower and higher,respectively,than the global mean values;the opposite was observed in pinnate-veined forbs.The parallel-veined grasses exhibited almost a four-fold higher vein biomass investment(i.e.,vein mass per leaf mass)than pinnate-veined forbs in the same region.The average VLA and its relationships with other leaf traits of the alpine grassland plants on the Tibetan Plateau had no regional characteristics,suggesting a convergence in plant leaf functioning.Plants with different leaf types differ in their adaptation strategies to plateau environments,and this may relate to biomass investment into leaf veins.Our study fills the gap with regard to vein density in alpine grassland species and provides a new perspective for understanding plant physiology and ecology by calculating and comparing the proportion of vein investment among different vein types.

Variations in leaf functional traits and physiological characteristics of Abies georgei var.smithii along the altitude gradient in the Southeastern Tibetan Plateau

Variations in leaf functional traits of Abies georgei var. smithii at 3700, 3900, 4100, 4300, and 4390 m altitude were investigated in 15 typical plots in the Southeastern Tibetan Plateau. In each plot, three seedlings were selected, of which functional leaves in current-year sunny branches were chosen for the measurement of morphological, photosynthetic, and physiological and biochemical characteristics, and their variations were analyzed. Results showed that significant variations existed among the leaf functional traits of A. georgei var. smithii along the altitudinal gradient, as well as their physiological adaption indicators. Leaf area decreased, while the mass per area and thickness of leaf increased at an altitude above 4,100 m. The maxima of pigment, total nitrogen concentration, net photosynthesis rate during light-saturated, and when water use efficiency appeared at 4100 m altitude. In addition, A. georgei var. smithii seedlings regulated the activities of superoxide dismutase and ascorbate peroxidase to resist abiotic stress under 4100 m altitude. Meanwhile, malondialdehyde concentration and the dark respiration rate rapidly increased, which indicates that A. georgei var. smithii seedlingssuffered from heavy abiotic stress from 4100 m to 4390 m altitude. Basing on variations in leaf functional traits along the altitude gradient, we inferred that 4100 m altitude was the suitable region for A. georgei var. smithii growth in the Sygera Mountain. Moreover, the harsh environment was the main limiting factor for A. georgei var. smithii population expansion to high altitude.