Nitrogen-fixing and non-nitrogen-fixing legume plants differ in leaf nutrient concentrations and relationships between photosynthetic and hydraulic traits.

Legumes account for a significant proportion of plants in the terrestrial ecosystems. Nitrogen (N)-fixing capability of certain legumes is a pivotal trait that contributes to their ecological dominance. Yet, the functional traits and trait relationships between N-fixer and non-N-fixer legumes are poorly understood. Here, we investigated 27 functional traits associated with morphology, nutrients, hydraulic conductance and photosynthesis in 42 woody legumes (19 N-fixers and 23 non-N-fixers) in a common garden. Our results showed that N-fixers had higher specific leaf area, photosynthetic phosphorus (P)-use efficiency, leaf N, and iron concentrations on both area and mass basis, N/P ratio, and carbon (C) to P ratio, but lower wood density, area-based maximum photosynthetic rate (Aa), photosynthetic N-use efficiency, leaf mass- and area-based P and molybdenum and area-based boron concentrations, and C/N ratio, compared with non-N-fixers. The mass-based maximum photosynthetic rate (Am), stomatal conductance (gs), intrinsic water-use efficiency (WUEi), mass- and area-based leaf potassium and mass-based boron concentrations, leaf hydraulic conductance (Kleaf), and whole-shoot hydraulic conductance (Kshoot) showed no difference between N-fixers and non-N-fixers. Significant positive associations between all hydraulic and photosynthetic trait pairs were found in N-fixers, but only one pair (Kshoot-Aa) in non-N-fixers, suggesting that hydraulic conductance plays a more important role in mediating photosynthetic capacity in N-fixers compared with non-N-fixers. Higher mass-based leaf N was linked to lower time-integrated gs and higher WUEi among non-N-fixer legumes or all legumes pooled after phylogeny was considered. Moreover, mass-based P concentration was positively related to Am and gs in N-fixers, but not in non-N-fixers, indicating that the photosynthetic capacity and stomatal conductance in N-fixers were more dependent on leaf P status than in non-N-fixers. These findings expand our understanding of the trait-based ecology within and across N-fixer and non-N-fixer legumes in tropics.

Effects of lianas on forest biogeochemistry during their lives and afterlives.

Climate change and other anthropogenic disturbances are increasing liana abundance and biomass in many tropical and subtropical forests. While the effects of living lianas on species diversity, ecosystem carbon, and nutrient dynamics are receiving increasing attention, the role of dead lianas in forest ecosystems has been little studied and is poorly understood. Trees and lianas coexist as the major woody components of forests worldwide, but they have very different ecological strategies, with lianas relying on trees for mechanical support. Consequently, trees and lianas have evolved highly divergent stem, leaf, and root traits. Here we show that this trait divergence is likely to persist after death, into the afterlives of these organs, leading to divergent effects on forest biogeochemistry. We introduce a conceptual framework combining horizontal, vertical, and time dimensions for the effects of liana proliferation and liana tissue decomposition on ecosystem carbon and nutrient cycling. We propose a series of empirical studies comparing traits between lianas and trees to answer questions concerning the influence of trait afterlives on the decomposability of liana and tree organs. Such studies will increase our understanding of the contribution of lianas to terrestrial biogeochemical cycling, and help predict the effects of their increasing abundance.

Linkages among stem xylem transport, biomechanics, and storage in lianas and trees across three contrasting environments.

PREMISE: Stem xylem transports water and nutrients, mechanically supports aboveground tissues, and stores water and nonstructural carbohydrates. These three functions are associated with three types of cells-vessel, fiber, and parenchyma, respectively. METHODS: We measured stem theoretical hydraulic conductivity (Kt), modulus of elasticity (MOE), tissue water content, starch, soluble sugars, cellulose, and xylem anatomical traits in 15 liana and 16 tree species across three contrasting sites in Southwest China. RESULTS: Lianas had higher hydraulic efficiency and tissue water content, but lower MOE and cellulose than trees. Storage traits (starch and soluble sugars) did not significantly differ between lianas and trees, and trait variation was explained mainly by site, highlighting how environment shapes plant storage strategies. Kt was significantly positively correlated with vessel diameter and vessel area fraction in lianas and all species combined. The MOE was significantly positively correlated with fiber area fraction, wood density, and cellulose in lianas and across all species. The tissue water content was significantly associated with parenchyma area fraction in lianas. Support function was strongly linked with transport and storage functions in lianas. In trees, transport and support functions were not correlated, while storage function was tightly linked with transport and support functions. CONCLUSIONS: These findings enhance our understanding of the relationship between stem xylem structure and function in lianas and trees, providing valuable insights into how plants adapt to environmental changes and the distinct ecological strategies employed by lianas and by trees to balance the demands of hydraulic transport, mechanical support, and storage.

Convergent relationships between flower economics and hydraulic traits across aquatic and terrestrial herbaceous plants.

Maintaining open flowers is critical for successful pollination and depends on long-term water and carbon balance. Yet the relationship between how flower hydraulic traits are coordinated in different habitats is poorly understood. Here, we hypothesize that the coordination and trade-offs between floral hydraulics and economics traits are independent of environmental conditions. To test this hypothesis, we investigated a total of 27 flower economics and hydraulic traits in six aquatic and six terrestrial herbaceous species grown in a tropical botanical garden. We found that although there were a few significant differences, most flower hydraulics and economics traits did not differ significantly between aquatic and terrestrial herbaceous plants. Both flower mass per area and floral longevity were significantly positively correlated with the time required for drying full-hydrated flowers to 70% relative water content. Flower dry matter content was strongly and positively related to drought tolerance of the flowers as indicated by flower water potential at the turgor loss point. In addition, there was a trade-off between hydraulic efficiency and the construction cost of a flower across species. Our results show that flowers of aquatic and terrestrial plants follow the same economics spectrum pattern. These results suggest a convergent flower economics design across terrestrial and aquatic plants, providing new insights into the mechanisms by which floral organs adapt to aquatic and terrestrial habitats.

Hydraulic traits and photosynthesis are coordinated with trunk sapwood capacitance in tropical tree species.

Water stored in trunk sapwood is vital for the canopy to maintain its physiological function under high transpiration demands. Little is known regarding the anatomical properties that contribute to the hydraulic capacitance of tree trunks and whether trunk capacitance is correlated with the hydraulic and gas exchange traits of canopy branches. We examined sapwood capacitance, xylem anatomical characteristics of tree trunks, embolism resistance, the minimal xylem water potential of canopy branches, leaf photosynthesis and stomatal conductance in 22 species from a tropical seasonal rainforest and savanna. The results showed that the mean trunk sapwood capacitance did not differ between the two biomes. Capacitance was closely related to the fiber lumen fraction and fiber wall reinforcement and not to the axial and ray parenchyma fractions. Additionally, it was positively correlated with the theoretical hydraulic conductivity of a trunk and the specific hydraulic conductivity of branches, and showed a trade-off with branch embolism resistance. Species with a high trunk sapwood capacitance maintained less negative canopy water potentials in the dry season, but higher leaf photosynthetic rates and stomatal conductance in the wet season. This study provides a functional link among trunk sapwood capacitance, xylem anatomy, canopy hydraulics and photosynthesis in tropical trees.

Can leaf drought tolerance predict species abundance and its changes in tropical-subtropical forests?

Climate change has resulted in an increase in drought severity in the species-rich tropical and subtropical forests of southern China. Exploring the spatiotemporal relationship between drought-tolerance trait and tree abundance provides a means to elucidate the impact of droughts on community assembly and dynamics. In this study, we measured the leaf turgor loss point (πtlp) for 399 tree species from three tropical forest plots and three subtropical forest plots. The plot area was 1 ha and tree abundance was calculated as total basal area per hectare according to the nearest community census data. The first aim of this study was to explore πtlp abundance relationships in the six plots across a range of precipitation seasonality. Additionally, three of the six plots (two tropical forests and one subtropical forest) had consecutive community censuses data (12-22 years) and the mortality ratios and abundance year slope of tree species were analyzed. The second aim was to examine whether πtlp is a predictor of tree mortality and abundance changes. Our results showed that tree species with lower (more negative) πtlp were more abundant in the tropical forests with relative high seasonality. However, πtlp was not related to tree abundance in the subtropical forests with low seasonality. Moreover, πtlp was not a good predictor of tree mortality and abundance changes in both humid and dry forests. This study reveals the restricted role of πtlp in predicting the response of forests to increasing droughts under climate change.

Vessel dimorphism and wood traits in lianas and trees among three contrasting environments.

PREMISE: Determining how xylem vessel diameters vary among plants and across environments gives insights into different water-use strategies among species and ultimately their distributions. Here, we tested the vessel dimorphism hypothesis that the simultaneous occurrence of many narrow and a few wide vessels gives lianas an advantage over trees in seasonally dry environments. METHODS: We measured the diameters of 13,958 vessels from 15 liana species and 10,430 vessels from 16 tree species in a tropical seasonal rainforest, savanna, and subtropical evergreen broadleaved forest. We compared differences in mean and hydraulically weighted vessel diameter (MVD and Dh ), vessel density (VD), theoretical hydraulic conductivity (Kt ), vessel area fraction (VAF), and wood density (WD) between lianas and trees and among three sites. RESULTS: Nine liana species and four tree species had dimorphic vessels. From the tropical seasonal rainforest to the savanna, liana MVD, Dh and Kt decreased, and VD and WD increased, while only tree WD increased. From the tropical seasonal rainforest to the subtropical forest, six wood traits remained unchanged for lianas, while tree MVD, Dh and Kt decreased and VD increased. Trait space for lianas and trees were more similar in the savanna and more divergent in the subtropical forest compared to the tropical seasonal rainforest. CONCLUSIONS: These results suggest that lianas tend to possess greater vessel dimorphism, which may explain how lianas grow well during seasonal drought, influencing their unique distribution across tropical rainfall gradients.

Physiological response and photosynthetic recovery to an extreme drought: Evidence from plants in a dry-hot valley savanna of Southwest China.

The frequency of extreme drought events has been rising worldwide, but due to its unpredictability, how plants will respond remains poorly understood. Here, we aimed to characterize how the hydraulics and photosynthesis of savanna plants respond to extreme drought, and tested whether they can subsequently recover photosynthesis after drought. There was an extreme drought in 2019 in Southwest (SW) China. We investigated photosynthetic gas exchange, leaf-, stem-, and whole-shoot hydraulic conductance of 18 plant species with diverse leaf habits (deciduous, semi-deciduous and evergreen) and growth forms (tree and shrub) from a dry-hot valley savanna in SW China for three rainy seasons from 2019 to 2021. We also compared photosynthetic gas exchange to those of a regular year (2014). We found that leaf stomatal and hydraulic conductance and maximum photosynthetic rate were significantly lower during the drought in 2019 than in the wetter years. In 2019, all studied plants maintained stomatal conductance at their minimum level observed, which could be related to high vapor pressure deficits (VPD, >2 kPa). However, no significant difference in stem and shoot hydraulic conductance was detected across years. The reductions in leaf hydraulic conductance and stomatal regulation under extreme drought might help keep the stem hydraulic function. Stomatal conductance and photosynthesis after drought (2020 and 2021) showed comparable or even higher values compared to that of 2014, suggesting high recovery of photosynthetic gas exchange. In addition, the response of hydraulic and photosynthetic traits to extreme drought was convergent across leaf habits and growth forms. Our results will help better understand the physiological mechanism underlying the response of savanna ecosystems to climate change.

You are what you eat: nutrient and water relations between mistletoes and hosts.

Mistletoes play important roles in biogeochemical cycles. Although many studies have compared nutrient concentrations between mistletoes and their hosts, no general patterns have been found and the nutrient uptake mechanisms in mistletoes have not been fully resolved. To address the water and nutrient relations in mistletoes compared with their hosts, we measured 11 nutrient elements, two isotope ratios and two leaf morphological traits for 11 mistletoe and 104 host species from four sites across a large environmental gradient in southwest China. Mistletoes had significantly higher phosphorus, potassium, and boron concentrations, nitrogen isotope ratio, and lower carbon isotope ratio (δ13 C) indicative of lower water-use efficiency than hosts, but other elements were similar to those in hosts. Sites explained most of the variation in the multidimensional trait space. With increasing host nitrogen concentration, both mistletoe δ13 C and the difference between mistletoe and host δ13 C increased, providing evidence to support the 'nitrogen parasitism hypothesis'. Host nutrient concentrations were the best predictors for that of the mistletoe nutrient elements in most cases. Our results highlight the important roles of environmental conditions and host nutrient status in determining mistletoe nutrient pools, which together explain their trophic interactions with hosts in subtropical and tropical ecosystems.

A critical thermal transition driving spring phenology of Northern Hemisphere conifers.

Despite growing interest in predicting plant phenological shifts, advanced spring phenology by global climate change remains debated. Evidence documenting either small or large advancement of spring phenology to rising temperature over the spatio-temporal scales implies a potential existence of a thermal threshold in the responses of forests to global warming. We collected a unique data set of xylem cell-wall-thickening onset dates in 20 coniferous species covering a broad mean annual temperature (MAT) gradient (-3.05 to 22.9°C) across the Northern Hemisphere (latitudes 23°-66° N). Along the MAT gradient, we identified a threshold temperature (using segmented regression) of 4.9 ± 1.1°C, above which the response of xylem phenology to rising temperatures significantly decline. This threshold separates the Northern Hemisphere conifers into cold and warm thermal niches, with MAT and spring forcing being the primary drivers for the onset dates (estimated by linear and Bayesian mixed-effect models), respectively. The identified thermal threshold should be integrated into the Earth-System-Models for a better understanding of spring phenology in response to global warming and an improved prediction of global climate-carbon feedbacks.

Convergence in phosphorus constraints to photosynthesis in forests around the world.

Tropical forests take up more carbon (C) from the atmosphere per annum by photosynthesis than any other type of vegetation. Phosphorus (P) limitations to C uptake are paramount for tropical and subtropical forests around the globe. Yet the generality of photosynthesis-P relationships underlying these limitations are in question, and hence are not represented well in terrestrial biosphere models. Here we demonstrate the dependence of photosynthesis and underlying processes on both leaf N and P concentrations. The regulation of photosynthetic capacity by P was similar across four continents. Implementing P constraints in the ORCHIDEE-CNP model, gross photosynthesis was reduced by 36% across the tropics and subtropics relative to traditional N constraints and unlimiting leaf P. Our results provide a quantitative relationship for the P dependence for photosynthesis for the front-end of global terrestrial C models that is consistent with canopy leaf measurements.

The complete chloroplast genome of Bauhinia racemosa Lam. (Fabaceae): a versatile tropical medicinal plant.

Bauhinia racemosa Lam. (1783), a versatile medicinal plant, belongs to the family Fabaceae (subfamily Cercidoideae). In this study, we analyzed the complete chloroplast genome to facilitate its use in genetic research. The complete chloroplast genome of B. racemosa was found to be 155,501 bp long, including two inverted repeat (IR) regions of 25,446 bp, which are separated by a small single-copy (SSC) region of 18,295 bp and a large single-copy (LSC) region of 86,314 bp. The overall GC content is 36.4%. The genome of B. racemosa contains 129 genes, including 83 protein-coding genes, 37 tRNAs, 8 rRNAs, and 1 pseudogene (rps19). Phylogenetic analysis suggests that B. racemosa forms a monophyletic clade with the other four Bauhinia species (B. brachycarpa, B. purpurea, B. blakeana and B. variegata var. variegata).

Sources and consequences of mismatch between leaf disc and whole-leaf leaf mass per area (LMA).

PREMISE: Leaf mass per area (LMA), which is an important functional trait in leaf economic spectrum and plant growth analysis, is measured from leaf discs or whole leaves. Differences between the measurement methods may lead to large differences in the estimates of LMA values. METHODS: We examined to what extent estimates of LMA based on whole leaves match those based on discs using 334 woody species from a wide range of biomes (tropics, subtropics, savanna, and temperate), whether the relationship varied by leaf morphology (tissue density, leaf area, leaf thickness), punch size (0.6- and 1.0-cm diameter), and whether the extent of intraspecifc variation for each species matches. RESULTS: Disc-based estimates of species mean LMA matched the whole-leaf estimates well, and whole-leaf LMA tended to be 9.69% higher than leaf-disc LMA. The ratio of whole-leaf LMA to leaf-disc LMA was higher for species with higher leaf tissue density and larger leaves, and variance in the ratio was greater for species with lower leaf tissue density and thinner leaves. Estimates based on small leaf discs also inflated the ratio. The extent of the intraspecific variation only weakly matched between whole-leaf and disc-based estimates (R2 = 0.08). CONCLUSIONS: Our results suggest that simple conversion between whole-leaf and leaf-disc LMA is difficult for species obtained with a small leaf punch, but it should be possible for species obtained with a large+ leaf punch. Accurately representing leaf traits will likely require careful selection between leaf-disc and whole-leaf traits depending on the objectives. Quantifying intraspecific variation using leaf discs should be also considered with caution.

Leaf trait association in relation to herbivore defense, drought resistance, and economics in a tropical invasive plant.

PREMISE: Exploring how functional traits vary and covary is important to understand plant responses to environmental change. However, we have limited understanding of the ways multiple functional traits vary and covary within invasive species. METHODS: We measured 12 leaf traits of an invasive plant Chromolaena odorata, associated with plant or leaf economics, herbivore defense, and drought resistance on 10 introduced populations from Asia and 12 native populations from South and Central America, selected across a broad range of climatic conditions, and grown in a common garden. RESULTS: Species' range and climatic conditions influenced leaf traits, but trait variation across climate space differed between the introduced and native ranges. Traits that confer defense against herbivores and drought resistance were associated with economic strategy, but the patterns differed by range. Plants from introduced populations that were at the fast-return end of the spectrum (high photosynthetic capacity) had high physical defense traits (high trichome density), whereas plants from native populations that were at the fast-return end of the spectrum had high drought escape traits (early leaf senescence and high percentage of withered shoots). CONCLUSIONS: Our results indicate that invasive plants can rapidly adapt to novel environmental conditions. Chromolaena odorata showed multiple different functional trait covariation patterns and clines in the native and introduced ranges. Our results emphasize that interaction between multiple traits or functions should be considered when investigating the adaptive evolution of invasive plants.

Higher water and nutrient use efficiencies in savanna than in rainforest lianas result in no difference in photosynthesis.

Differences in traits between lianas and trees in tropical forests have been studied extensively; however, few have compared the ecological strategies of lianas from different habitats. Here, we measured 25 leaf and stem traits concerning leaf anatomy, morphology, physiology and stem hydraulics for 17 liana species from a tropical seasonal rainforest and for 19 liana species from a valley savanna in south-west China. We found that savanna lianas had higher vessel density, wood density and lower hydraulically weighted vessel diameter and theoretical hydraulic conductivity than tropical seasonal rainforest lianas. Compared with tropical seasonal rainforest lianas, savanna lianas also showed higher leaf dry matter content, carbon isotope composition (δ13C), photosynthetic water use efficiency, ratio of nitrogen to phosphorus, photosynthetic phosphorus use efficiency and lower leaf size, stomatal conductance and nitrogen, phosphorus and potassium concentrations. Interestingly, no differences in light-saturated photosynthetic rate were found between savanna and tropical seasonal rainforest lianas either on mass or area basis. This is probably due to the higher water and nutrient use efficiencies of savanna lianas. A principal component analysis revealed that savanna and tropical seasonal rainforest lianas were significantly separated along the first axis, which was strongly associated with acquisitive or conservative resource use strategy. Leaf and stem functional traits were coordinated across lianas, but the coordination or trade-off was stronger in the savanna than in the tropical seasonal rainforest. In conclusion, a relatively conservative (slow) strategy concerning water and nutrient use may benefit the savanna lianas, while higher nutrient and water use efficiencies allow them to maintain similar photosynthesis as tropical seasonal rainforest species. Our results clearly showed divergences in functional traits between lianas from savanna and tropical seasonal rainforest, suggesting that enhanced water and nutrient use efficiencies might contribute to the distribution of lianas in savanna ecosystems.

Cavitation resistance of peduncle, petiole and stem is correlated with bordered pit dimensions in Magnolia grandiflora.

Variation in resistance of xylem to embolism among flowers, leaves, and stems strongly influences the survival and reproduction of plants. However, little is known about the vulnerability to xylem embolism under drought stress and their relationships to the anatomical traits of pits among reproductive and vegetative organs. In this study, we investigated the variation in xylem vulnerability to embolism in peduncles, petioles, and stems in a woody plant, Magnolia grandiflora. We analyzed the relationships between water potentials that induced 50% embolism (P50) in peduncles, petioles, and stems and the conduit pit traits hypothesized to influence cavitation resistance. We found that peduncles were more vulnerable to cavitation than petioles and stems, supporting the hypothesis of hydraulic vulnerability segmentation that leaves and stems are prioritized over flowers during drought stress. Moreover, P50 was significantly correlated with variation in the dimensions of inter-vessel pit apertures among peduncles, petioles and stems. These findings highlight that measuring xylem vulnerability to embolism in reproductive organs is essential for understanding the effect of drought on plant reproductive success and mortality under drought stress.

Traits, strategies, and niches of liana species in a tropical seasonal rainforest.

Plant functional traits and strategies hold the promise to explain species distribution, but few studies have linked multiple traits to multiple niche dimensions (i.e., light, water, and nutrients). Here, we analyzed for 29 liana species in a Chinese tropical seasonal rainforest how: (1) trait associations and trade-offs lead to different plant strategies; and (2) how these traits shape species' niche dimensions. Eighteen functional traits related to light, water, and nutrient use were measured and species niche dimensions were quantified using species distribution in a 20-ha plot combined with data on canopy gaps, topographic water availability, and soil nutrients. We found a tissue toughness spectrum ranging from soft to hard tissues along which species also varied from acquisitive to conservative water use, and a resource acquisition spectrum ranging from low to high light capture and nutrient use. Intriguingly, each spectrum partly reflected the conservative-acquisitive paradigm, but at the same time, the tissue toughness and the resource acquisition spectrum were uncoupled. Resource niche dimensions were better predicted by individual traits than by multivariate plant strategies. This suggests that trait components that underlie multivariate strategy axes, rather than the plant strategies themselves determine species distributions. Different traits were important for different niche dimensions. In conclusion, plant functional traits and strategies can indeed explain species distributions, but not in a simple and straight forward way. Although the identification of global plant strategies has significantly advanced the field, this research shows that global, multivariate generalizations are difficult to translate to local conditions, as different components of these strategies are important under different local conditions.

Divergence of stem biomechanics and hydraulics between Bauhinia lianas and trees.

Liana abundance and biomass are increasing in neotropical and Asian tropical seasonal forests over the past decades. Stem mechanical properties and hydraulic traits influence the growth and survival of plants, yet the differences in stem mechanical and hydraulic performance between congeneric lianas and trees remain poorly understood. Here, we measured 11 stem mechanical and hydraulic traits for 10 liana species and 10 tree species from Bauhinia grown in a tropical common garden. Our results showed that Bauhinia lianas possessed lower stem mechanical strength as indicated by both modulus of elasticity and modulus of rupture, and higher stem potential hydraulic conductivity than congeneric trees. Such divergence was mainly attributed to the differentiation in liana and tree life forms. Whether the phylogenetic effect was considered or not, mechanical strength was positively correlated with wood density, vessel conduit wall reinforcement and sapwood content across species. Results of principle component analysis showed that traits related to mechanical safety and hydraulic efficiency were loaded in the opposite direction, suggesting a trade-off between biomechanics and hydraulics. Our results provide evidence for obvious differentiation in mechanical demand and hydraulic efficiency between congeneric lianas and trees.