Plant Resource-use Strategies Research Articles

Mycorrhizal inoculation alters rhizosphere fungal community and root morphology to improve drought tolerance of resource-acquisitive but not resource-conservative Quercus species

The mutualistic association between mycorrhizal fungi and plants is well known to improve plant drought resistance. However, the influence of external inoculants on the interactions between soil microbial communities and plant functional traits and how these interactions improve plant drought tolerance under drought stress remain unclear. We conducted a pot inoculation experiment to assess the effect of two inoculated fungal strains (Clitopolius hobsonii NL-19 and C. sp. HSL-YX-7-A) on morphological traits and rhizosphere fungal communities of eight Quercus species seedlings. Plant and soil fungal traits were measured 2 months after inoculation and then underwent a short-term (33 days) drought rewetting treatment. The biomass of all tree species reduced significantly under drought stress followed by rewetting (drought treatment) compared with that in the control treatment (well-watered); however, inoculated mycorrhizal fungi (drought + mycorrhizal inoculation treatment) increased the biomass of five Quercus species (growth-promoting species (GPS) exhibiting a significant increase in biomass under mycorrhizal inoculation treatment than under the uninoculated treatment during drought stress, characterized by a resource-use acquisitive strategy) and had no effect on the other three Quercus species (non-growth-promoting species (NGPS) exhibiting no significant difference in biomass between mycorrhizal inoculation and uninoculated treatments during drought stress, characterized by a resource-use conservative strategy). The leaf traits of the two species groups (GPS and NGPS) varied little among the three treatments. The values of most root traits (e.g., specific root length and root length) of GPS increased significantly under drought and drought + mycorrhizal inoculation treatments compared with those in the control, whereas relatively minor variations were observed in NGPS among the three treatments. The fungal community composition and functional groups (primary trophic modes) in the drought treatment were altered for NGPS but not for GPS; additionally, they changed in the drought + mycorrhizal inoculation treatment for GPS but not for NGPS when compared with those in the drought treatment. Furthermore, the drought tolerance of GPS was improved directly by fungal functional groups (i.e., Pathotroph-Saprotroph- Symbiotrophs proliferation) and indirectly by root traits (e.g., increased specific root length), whereas NGPS may adapt to drought stress through other pathways (e.g., physiological and biochemical regulation). Overall, drought and mycorrhizal inoculation affected the root traits and fungal community structure of Quercus seedlings, which rely on plant resource-use strategies. Our results provide new insights into the relationship between plant resource-use strategies and soil microbes for improving plant performance under drought stress.

Enhanced rock weathering increased soil phosphorus availability and altered root phosphorus-acquisition strategies.

Enhanced rock weathering (ERW) has been proposed as a measure to enhance the carbon (C)-sequestration potential and fertility of soils. The effects of this practice on the soil phosphorus (P) pools and the general mechanisms affecting microbial P cycling, as well as plant P uptake are not well understood. Here, the impact of ERW on soil P availability and microbial P cycling functional groups and root P-acquisition traits were explored through a 2-year wollastonite field addition experiment in a tropical rubber plantation. The results show that ERW significantly increased soil microbial carbon-use efficiency and total P concentrations and indirectly increased soil P availability by enhancing organic P mobilization and mineralization of rhizosheath carboxylates and phosphatase, respectively. Also, ERW stimulated the activities of P-solubilizing (gcd, ppa and ppx) and mineralizing enzymes (phoADN and phnAPHLFXIM), thus contributing to the inorganic P solubilization and organic P mineralization. Accompanying the increase in soil P availability, the P-acquisition strategy of the rubber fine roots changed from do-it-yourself acquisition by roots to dependence on mycorrhizal collaboration and the release of root exudates. In addition, the direct effects of ERW on root P-acquisition traits (such as root diameter, specific root length, and mycorrhizal colonization rate) may also be related to changes in the pattern of belowground carbon investments in plants. Our study provides a new insight that ERW increases carbon-sequestration potential and P availability in tropical forests and profoundly affects belowground plant resource-use strategies.

Hemp cultivation opportunities for marginal lands development.

Agricultural diversification and high-quality products deriving from sustainable crops such as hemp can represent a solution to revitalize marginal areas and reverse land abandonment. This study aimed at comparing four different hemp cultivars (Carmagnola Selezionata, "CS"; Futura 75, "FUT"; Felina 32, "FEL"; Secuieni Jubileu, "JUB") to provide information to select the best suited cultivar for cultivation in mountain marginal areas and for specific end-use applications. Hemp cultivars were cultivated in a single experimental field to compare their ecological and agronomic behavior (duration of life cycle phases, plant size and biomass allocation, and plant resource-use strategies). Through metabolomic analysis of both vegetative and reproductive parts of the plants we tested the presence of substances of nutraceutical interest and traced seed nutritional profile. The four cultivars had different ecological and agronomic behavior, and nutritional profile. We found several compounds with potential pharmaceutical and nutraceutical values in all parts of the plant (leaves, inflorescences, and stems). JUB resulted the most suitable for seed production while CS showed the highest content of bioactive compounds in flowers and leaves. FUT, showed the best suitability for multi-purpose cultivation, while FEL seemed to be not appropriate for the cultivation in mountain area. The multi-disciplinary approach we adopted was effective in distinguish across hemp cultivars and provided information to farmers for the selection of the best hemp cultivar to select. Hemp had a high potential for cultivation in marginal lands, demonstrating to be an economic resource due to its multi-purpose use and to the possibility to generate high-added values products. Our results could serve as a stimulus for the reintroduction of this culture in the study area and in other similar environments.

Liming shift above- and belowground functional traits of Chinese fir from conservative to acquisitive

Plant functional traits can serve as indicators of plant resource-use strategies in response to environmental changes. Plants growing in acidified soil exhibit conservative traits and limit growth, but there is still little known about changes in above- and belowground functional traits and their coordination under alleviation of soil acidification. Here, we conducted a pot experiment on raising soil pH by liming in Chinese fir (Cunninghamia lanceolata) saplings in southern China. The experiment involved in two degrees of acidified soil (Mild, pH = 4.3; Severe, pH = 3.6) and three liming treatments: Control (CK, 0), low liming treatment (L, 1000 kg CaO hm−2) and high liming treatment (H, 4000 kg CaO hm−2). We measured 22 above- and belowground functional traits of Chinese fir including allocation, morphological, photosynthetic, and chemical traits to assess their response to liming. The results show that liming enhanced aboveground mass fraction and total leaf area (TLA), and reduced construction cost (lower C concentration). However, specific leaf area (SLA), nutrient concentrations of N and P, and photosynthetic rate (Pn) remained unchanged. These results suggest that aboveground components exhibited acquisitive strategies in response to liming with greater TLA and lower construction cost. Moreover, liming reduced root mass fraction (RMF), increased specific root length (SRL), specific root area (SRA), and root N and Ca nutrient concentrations. These results suggest that belowground components also exhibited acquisitive strategies through constructing longer and thinner roots with greater metabolic rates. Our findings show that liming shifts above- and belowground resource-use strategies of Chinese fir from conservative to acquisitive. These changes in functional traits support the ability of Chinese fir to grow rapidly in response to alleviation of soil acidification. Our research not only contributes to a deeper understanding of plant adaptation mechanisms in the face of environment changes but also provides valuable insights for improving carbon sequestration of plantations in subtropical China.

Size-dependent and environment-mediated shifts in leaf traits of a deciduous tree species in a subtropical forest.

AimsUnderstanding the joint effects of plant development and environment on shifts of intraspecific leaf traits will advance the understandings of the causes of intraspecific trait variation. We address this question by focusing on a widespread species Clausena dunniana in a subtropical broad‐leaved forest.MethodsWe sampled 262 individuals of C. dunniana at two major topographic habitat types, the slope and hilltop, within the karst forests in Maolan Nature Reserve in southwestern China. We measured individual plant level leaf traits (i.e., specific leaf area (SLA), leaf area, leaf dry‐matter content (LDMC), and leaf thickness) that are associated with plant resource‐use strategies. We adopted a linear mixed‐effects model in which the plant size (i.e., the first principal component of plant basal diameter and plant height) and environmental factors (i.e., topographic habitat, canopy height, and rock‐bareness) were used as independent variables, to estimate their influences on the shifts of leaf traits.Key ResultsWe found that (1) plant size and the environmental factors independently drove the intraspecific leaf trait shifts of C. dunniana, of which plant size explained less variances than environmental factors. (2) With increasing plant size, C. dunniana individuals had increasingly smaller SLA but larger sized leaves. (3) The most influential environmental factor was topographic habitat; it drove the shifts of all the four traits examined. Clausena dunniana individuals on hilltops had leaf traits representing more conservative resource‐use strategies (e.g., smaller SLA, higher LDMC) than individuals on slopes. On top of that, local‐scale environmental factors further modified leaf trait shifts.ConclusionsPlant size and environment independently shaped the variations in intraspecific leaf traits of C. dunniana in the subtropical karst forest of Maolan. Compared with plant size, the environment played a more critical role in shaping intraspecific leaf trait variations, and potentially also the underlying individual‐level plant resource‐use strategies.

Trait-based responses to cessation of nutrient enrichment in a tundra plant community

Plant communities worldwide show varied responses to nutrient enrichment—including shifts in species identity, decreased diversity, and changes in functional trait composition—but the factors determining community recovery after the cessation of nutrient addition remain uncertain. We manipulated nutrient levels in a tundra community for 6 years of nutrient addition followed by 8 years of recovery. We examined how community recovery was mediated by traits related to plant resource-use strategy and plant ability to modify their environment. Overall, we observed persistent effects of fertilization on plant communities. We found that plants with fast-growing traits, including higher specific leaf area, taller stature and lower foliar C:N, were more likely to show a persistent increase in fertilized plots than control plots, maintaining significantly higher cover in fertilized plots 8 years after cessation of fertilization. Additionally, although graminoids responded most strongly to the initial fertilization treatment, forb species were more vulnerable to fertilization effects in the long-term, showing persistent decline and no recovery in 8 years. Finally, these persistent fertilization effects were accompanied by modified environmental conditions, including persistent increases in litter depth and soil phosphorous and lower soil C:N. Our results demonstrate the potential for lasting effects of nutrient enrichment in nutrient-limited systems and identify species traits related to rapid growth and nutrient-use efficiency as the main predictors of the persistence of nutrient enrichment effects. These findings highlight the usefulness of trait-based approach for understanding the persistent feedbacks of nutrient enrichment, plant dynamics, and niche construction via litter and nutrient build-up.

Root traits mediate functional guilds of soil nematodes in an ex-arable field

One of the greatest challenges in soil ecology is to disentangle the plant-mediated bottom-up factors regulating soil biodiversity and community composition. The soil food web, fundamentally driving nearly all ecosystem functions, is controlled by the quantity and quality of root-mediated resources. Here, a trait-based approach was adopted to explore the divergence of soil nematode functional guilds and their connections to root traits belonging to distinct plant resource-use strategies. Root traits and rhizosphere nematode functional guilds were measured on four plant species in an ex-arable field. Results showed that plant species exhibiting acquisitive strategies promoted nematode abundance in contrast to species with conservative strategies. Further, the results also supported that plant resource-use strategies could regulate nematode life strategies in a bottom-up manner. However, lower proportions of opportunist nematodes in rhizosphere were found in acquisitive plants rather than conservative plants, mainly attributed to the stronger top-down regulation as a dominant control within the former than the later ones. Structure equation modeling revealed that root length density could primarily modulate nematode abundance and functional guilds mainly through changes in nitrogen availability of rhizosphere, as indicated by mineral N and enzymatic stoichiometry. Overall, our findings extend the conceptual framework based on a trait-centred view spanning plants to the soil food web, and this knowledge is critical to understand the mechanisms of ecosystem process.

Relationships between root morphology, root exudate compounds and rhizosphere microbial community in durum wheat

A multidisciplinary approach was adopted to investigate multiple relationships between root morphology, rhizosphere soil exudates and soil microbial community in eight durum wheat [Triticum turgidum ssp. durum (Desf.)] cultivars previously characterized in terms of their ability to interact with mycorrhizal fungi. Six plants from each cultivar were grown to the five-leaf stage in a growth chamber. Root system architecture was characterized by root scans, root exudates were identified and quantified by gas chromatography–mass spectrometry (GC–MS), and the rhizosphere bacterial, archaeal and fungal community structure was assessed using terminal-restriction fragment length polymorphism (T-RFLP). Wheat cultivars were very different in terms of the morphological root traits. Cappelli, Grecale, Ofanto and PR22D89 cultivars showed the most developed root system. Root exudates included sugars, amino acids, polyalcohols and organic acids, with carbohydrates accounting for 87.3% of the whole data set. All the compounds showed significant differences among cultivars, with Grecale and Claudio showing the highest exudation ability of metabolites. T-RFLP profiles showed a greater relative abundance of bacterial taxa (55.8% on average) than those of archaea and fungi (35.6% and 8.6%, on average, respectively) in all cultivars, and responded positively to root exudation. Principal component analysis identified root dry weight, surface area and volume, carbohydrate exudation, bacterial and archaeal Shannon–Weaver diversity indices and the fungal evenness index as the traits that mainly contributed to the cultivar variability. A larger root system did not always promote a higher diversity of the rhizosphere microbial biomass. This then suggests complex interactions between traits, probably due to the differences in the quality and/or composition of the root exudates. Based on our results, we propose that durum wheat cultivars could be classified according to the different plant resource-use strategies employed, such as conservative (i.e. Cappelli, with a large root system but low exudation ability) and explorative/competitive (i.e. Grecale and Claudio, with a high exudation ability) cultivars. The results are discussed with reference to the complex relationships between the plant and the rhizospheric microorganisms in order to determine the best combinations that would enhance root efficiency.

Root traits across environmental gradients in Mediterranean woody communities: are they aligned along the root economics spectrum?

Root traits play a critical role in plant resource-use strategies and ecosystem functioning, but there is great controversy regarding their identity and functionality in different dimensions of belowground functional variation. Here, we explored the level of covariation among a suite of key root traits (i.e. specific root length, root dry matter content, diameter and density) as well as between them and two aboveground traits related with plant function (leaf nutrient concentration and specific leaf area). We also evaluated whether these patterns of trait covariation were consistent at different spatial scales and organisational levels. We collected fine root (< 2 mm) samples of 534 individuals (of 80 woody species) along a wide regional range of environmental conditions in southern Spain. In general, strong correlations among most of the root morphological traits were found, supporting the existence of a ‘root economics spectrum’, as well as between root traits and the aboveground traits. However, root diameter was not completely aligned along this ecological axis, supporting the idea of a multidimensional spectrum of root traits. The main syndrome of root trait covariation was consistent at the different spatial scales and organisational levels. Soil nutrients and water availability were the main drivers of root trait variation. Our results indicate that root trait variation is primarily aligned along a leading dimension related to resource economics. However, the distinct pattern of root diameter may indicate a multidimensionality of belowground traits that needs to be explored in greater depth.

Phylogeny is a powerful tool for predicting plant biomass responses to nitrogen enrichment.

Increasing rates of anthropogenic nitrogen (N) enrichment to soils often lead to the dominance of nitrophilic plant species and reduce plant diversity in natural ecosystems. Yet, we lack a framework to predict which species will be winners or losers in soil N enrichment scenarios, a framework that current literature suggests should integrate plant phylogeny, functional tradeoffs, and nutrient co-limitation. Using a controlled fertilization experiment, we quantified biomass responses to N enrichment for 23 forest tree species within the genus Eucalyptus that are native to Tasmania, Australia. Based on previous work with these species' responses to global change factors and theory on the evolution of plant resource-use strategies, we hypothesized that (1) growth responses to N enrichment are phylogenetically structured, (2)species with more resource-acquisitive functional traits have greater growth responses to N enrichment, and (3) phosphorus (P) limits growth responses to N enrichment differentially across species, wherein P enrichment increases growth responses to N enrichment more in some species than others. We built a hierarchical Bayesian model estimating effects of functional traits (specific leaf area, specific stem density, and specific root length) and P fertilization on species' biomass responses to N, which we then compared between lineages to determine whether phylogeny explains variation in responses to N. In concordance with literature on N limitation, a majority of species responded strongly and positively to N enrichment. Mean responses ranged three-fold, from 6.21 (E.pulchella) to 16.87 (E.delegatensis) percent increases in biomass per g N·m-2 ·yr-1 added. We identified a strong difference in responses to N between two phylogenetic lineages in the Eucalyptus subgenus Symphyomyrtus, suggesting that shared ancestry explains variation in N limitation. However, our model indicated that after controlling for phylogenetic non-independence, eucalypt responses to N were not associated with functional traits (although post-hoc analyses show a phylogenetic pattern in specific root length similar to that of responses to N), nor were responses differentially limited by P. Overall, our model results suggest that phylogeny is a powerful predictor of winners and losers in anthropogenic N enrichment scenarios in Tasmanian eucalypts, which may have implications for other species.

Community aggregated traits disclose functional responses to seasonal resource fluctuations and spatial heterogeneity

Aim To investigate changes in community-level functional responses to rainfall seasonality in Neotropical grasslands through the analysis of community aggregated traits. Location Semi-natural grasslands in the Rio Grande de Matagalpa watershed, Nicaragua. Methods We measured 14 functional response traits that are indicators of plant resource-use strategies, across 32 herbaceous and four woody species in eight permanent plots: leaf size, specific leaf area, leaf dry matter content, leaf lifespan, foliar concentrations of P, N, Ca, K and Mg, plant height, lateral spread by clonal growth, root depth, start of flowering period and length of the period from flowering to seed shed. We calculated the community-weighted means (CWM) with trait values weighted by species cover for five different paddocks 11 times (between 2 Jul 2007 and 26 May 2008), and grouped them into four periods according to rainfall seasonality: early rainy season, late rainy season, early dry season and late dry season. Results Community aggregated values of specific leaf area, leaf dry matter content, leaf lifespan, foliar concentrations of P, N, Ca, K and Mg, length of the period from flowering to seed shed, and to a lesser extent lateral spread by clonal growth responded to temporal variations in rainfall, and corresponded to a considerable extent to expectations based on plant resource-use strategies along resource supply gradients. Community aggregated values of specific leaf area and foliar nutrient concentration were higher in the rainy season than in the dry season, while we observed an opposite trend with leaf dry matter content and leaf lifespan. Conclusions Semi-natural grasslands of central Nicaragua experience shifts in plant trait dominance that correspond with changes in resource supply given by rainfall seasonality, indicating that the stabilizing function of net primary productivity found in earlier studies can in part be a consequence of temporal differentiation in functional responses.

Plant resource-use strategies: the importance of phenotypic plasticity in response to a productivity gradient for two subalpine species.

Functional traits are indicators of plant interactions with their environment and the resource-use strategies of species can be defined through some key functional traits. The importance of genetic variability and phenotypic plasticity in trait variations in response to a common environmental change was investigated in two subalpine species. Two species with contrasted resource-use strategies, Dactylis glomerata and Festuca paniculata, were grown along a productivity gradient in a greenhouse experiment. Functional traits of different genotypes were measured to estimate the relative roles of phenotypic plasticity and genetic variability, and to compare their levels of phenotypic plasticity. Trait variability in the field for the two species is more likely to be the result of phenotypic plasticity rather than of genetic differentiation between populations. The exploitative species D. glomerata expressed an overall higher level of phenotypic plasticity compared with the conservative species F. paniculata. In addition to different amplitudes of phenotypic plasticity, the two species differed in their pattern of response for three functional traits relevant to resource use (specific leaf area, leaf dry matter content and leaf nitrogen content). Functional trait variability was mainly the result of phenotypic plasticity, with the exploitative species showing greater variability. In addition to average trait values, two species with different resource-use strategies differed in their plastic responses to productivity.

Specific Leaf Area and Leaf Dry Matter Content of Plants Growing in Sand Dunes

We investigated the variations in specific leaf area (SLA) and leaf dry matter content (LDMC) of 20 species (10 annuals and 10 perennials) that have different distributional patterns in the Kerqin Sandy Land in northern China. The main purpose of our study was to determine if SLA and/or LDMC could be used as indicators of plant resource-use strategy in sand dune environments. The selected species were mainly distributed in three types of sand dunes: mobile sand dune, semi-fixed sand dune, and fixed sand dune. The three differed in their soil nutrient content and vegetation productivity. Both SLA and LDMC varied substantially among species in the different dunes. Maximum SLA (28.2 m^2 kg^(-1)) was observed in the annual grass Setaria viridis in a fixed sand dune, and the lowest SLA (8.6 m^2 kg^(-1)) was found in the perennial grass Pennisetum centrasiatum in a semi-fixed sand dune. LDMC varied from 0.10 g g^(-1) for the annual forb Chenopodium glaucum in a fixed sand dune to 0.41 g g^(-1) for the perennial grass Phragmitis australis in a fixed sand dune. For species common to two or three sites, their SLA and LDMC were significantly different between sites (P＜0.05), except for two species for SLA and six species for LDMC. SLA and LDMC were negatively correlated both in the three types of sand dunes and for each functional type (annual versus perennial species). However, SLA and LDMC were significantly correlated only for the species in fixed sand dune, perennial species as well as all species. The results of our study showed that the SLA of annual plant species was generally higher than that of perennial species and that LDMC was significantly different between annual compared to perennial plants (P＜0.05).

Leaf traits as indicators of resource‐use strategy in floras with succulent species

Summary • Associations between specific leaf area (SLA), leaf water content (LWC) and leaf thickness (LT) in 77 species were analysed to identify which of these traits gave a better indicator value of general plant resource-use strategy within the flora of central-western Argentina, in which succulent species are common. • When all species were considered together, SLA and LWC were not significantly correlated. All high-SLA tender-leafed species showed high LWC. Low SLA, however, was associated both with low LWC (sclerophyllous species) and with high LWC (succulents). When succulents were excluded, the association between SLA and LWC was significant and positive. A similar trend was found for a mixed set of nonsucculent species from other floras of the world. • In the Argentine data set, SLA and LT, but not LWC, were significantly correlated with species’ scores along a multivariate axis of plant resource-use strategy. • Because of its clearer ecological interpretation and its applicability across different floras, SLA appears to be the best candidate for inclusion in large comparative databases.