Does local soil factor drive functional leaf trait variation? A test on Neilingding Island, South China.

Leaf traits were affected by soil factors and displayed varietal differences in forest. However, few examples have been reported on the Island ecosystems. We comprehensively investigated 9 leaf traits (leaf length, leaf width, leaf area, SLA, leaf fresh weight, leaf C content, leaf N content, leaf K content, leaf C:N ratio) of 54 main subtropical woody species and soil parameters (soil pH, total C content, total N content, total K content, available N content, available P content, available K content and soil moisture) in Neilingding Island, Shenzhen, southern China. Intra-and interspecific variation of leaf traits were measured and their correlations with soil parameters were explored. The interspecific variations of leaf C:N ratio, leaf N content and leaf fresh weight were higher than their intraspecific variations. The intraspecific variation of leaf K content was larger than that of interspecific one, accounting for 80.69% of the total variance. Positive correlations were found among intraspecific coefficients of variations in leaf morphological traits. The correlation analysis between the variation of intraspecific traits and the variation of soil parameters showed that changes in soil factors affected leaf morphology and stoichiometry. The interaction between soil moisture and soil available P content was the key factor on intraspecific variations of leaf traits including leaf area, leaf fresh weight, leaf C and leaf K content. We concluded that leaf traits of plants in the island were tightly related to soil parameters. Soil parameters, especially soil moisture and available P content, affected plant leaf morphology and stoichiometry at the local scale.

Response of leaf functional traits to soil nutrients in the wet and dry seasons in a subtropical forest on an island.

Introduction: Island ecosystems often have a disproportionate number of endemic species and unique and fragile functional characteristics. However, few examples of this type of ecosystem have been reported. Methods: We conducted a comprehensive field study on Neilingding Island, southern China. The leaf samples of 79 subtropical forest tree species were obtained and their functional traits were studied in the dry and wet seasons to explain the relationships between plant functional traits and soil nutrients. Results: We found a greater availability of soil moisture content (SMC) and nutrients in the wet season than in the dry season. The values of wet season soil available phosphorus (5.97 mg·kg-1), SMC (17.67%), and soil available potassium (SAK, 266.96 mg·kg-1) were significantly higher than those of the dry season. The leaf dry matter content, specific leaf weight, leaf density, leaf total carbon, leaf total nitrogen, leaf total calcium, and the N/P and C/P ratios of leaves were all significantly higher in the dry season than in the wet season, being 18.06%, 12.90%, 12.00%, 0.17%, 3.41%, 9.02%, 26.80%, and 24.14% higher, respectively. In contrast, the leaf area (51.01 cm2), specific leaf area (152.76 cm2·g-1), leaf water content (0.59%), leaf total nitrogen (1.31%), leaf total phosphorus (0.14%), and leaf total magnesium (0.33%) were much lower in the dry season than in the wet one. There were significant pairwise correlations between leaf functional traits, but the number and strength of correlations were significantly different in the dry and wet seasons. The SAK, soil total phosphorus (STP), and pH impacted plant leaf functional traits in the dry season, whereas in the wet season, they were affected by SAK, STP, pH, and NO3- (nitrate). Discussion: Both soil nutrients and water availability varied seasonally and could cause variation in a number of leaf traits.

Plant species identity and mycorrhizal type explain the root-associated fungal pathogen community assembly of seedlings based on functional traits in a subtropical forest.

Introduction: As a crucial factor in determining ecosystem functioning, interaction between plants and soil-borne fungal pathogens deserves considerable attention. However, little attention has been paid into the determinants of root-associated fungal pathogens in subtropical seedlings, especially the influence of different mycorrhizal plants. Methods: Using high-throughput sequencing techniques, we analyzed the root-associated fungal pathogen community for 19 subtropical forest species, including 10 ectomycorrhizal plants and 9 arbuscular mycorrhizal plants. We identified the roles of different factors in determining the root-associated fungal pathogen community. Further, we identified the community assembly process at species and mycorrhizal level and managed to reveal the drivers underlying the community assembly. Results: We found that plant species identity, plant habitat, and plant mycorrhizal type accounted for the variations in fungal pathogen community composition, with species identity and mycorrhizal type showing dominant effects. The relative importance of different community assembly processes, mainly, homogeneous selection and drift, varied with plant species identity. Interestingly, functional traits associated with acquisitive resource-use strategy tended to promote the relative importance of homogeneous selection, while traits associated with conservative resource-use strategy showed converse effect. Drift showed the opposite relationships with functional traits compared with homogeneous selection. Notably, the relative importance of different community assembly processes was not structured by plant phylogeny. Drift was stronger in the pathogen community for ectomycorrhizal plants with more conservative traits, suggesting the predominant role of stochastic gain and loss in the community assembly. Discussion: Our work demonstrates the determinants of root-associated fungal pathogens, addressing the important roles of plant species identity and plant mycorrhizal type. Furthermore, we explored the community assembly mechanisms of root-associated pathogens and stressed the determinant roles of functional traits, especially leaf phosphorus content (LP), root nitrogen content (RN) and root tissue density (RTD), at species and mycorrhizal type levels, offering new perspectives on the microbial dynamics underlying ecosystem functioning.

Host Recognition and Specific Infection of Endomelanconiopsis endophytica during Early Infection.

Coevolution between the pathogen and host plant drives pathogenic effector diversity. However, the molecular mechanism behind host-specific pathogenesis remains to be explored. Here, we present a 43 Mb whole-genome sequence of Endomelanconiopsis endophytica strain LS29, a host-specific pathogen of the common subtropical tree Castanopsis fissa. We described its genome annotations and identified its effector candidates. By performing temporal transcriptome sequencing of E. endophytica on C. fissa during early infection, we found that E. endophytica repressed other microbes in order to attack the tissue of the host by producing antibiotics earlier than 24 h post-inoculation (hpi). Simultaneously, a variety of effectors were secreted to recognize the host plant, but most of them showed a significantly opposing expression regulation trend after 24 hpi, indicating that 24 hpi represents a key time point between host recognition and specific infection. Furthermore, a comparison of isoenzymes showed that only a few effectors were identified as specific effectors, which were involved in hydrolyzing the compounds of the plant cell wall and releasing fatty acids during the early infection of C. fissa. Our results determined host recognition timing and identified a specific catalog of effectors, which are crucial for revealing the molecular mechanism of host-specific pathogenesis.

Negative impacts of sea-level rise on soil microbial involvement in carbon metabolism.

Sea-level rise has been threatening the terrestrial ecosystem functioning of coastal islands, of which the most important component is carbon (C) cycling. However, metagenomic and metabolomic evidence documenting salt intrusion effects on molecular biological processes of C cycling are still lacking. Here, we investigated microbial communities, metagenomic taxonomy and function, and metabolomic profiles in the marine-terrestrial transition zone of low- and high-tide, and low- and high-land areas based on distances of 0 m, 50 m, 100 m, and 200 m, respectively, to the water-land junction of Neilingding Island. Our results showed that soil salinity (EC) was the dominant driver controlling bacterial abundance and community composition and metagenomic taxonomy and function. The metabolomic profiling at the low-tide site was significantly different from that of other sites. The low-tide site had greater abundance of Proteobacteria and Bacteroidetes (1.6-3.7 fold), especially Gammaproteobacteria, but lower abundance (62-83%) of Acidobacteria and Chloroflexi, compared with other three sites. The metagenomic functional genes related to carbohydrate metabolism decreased at the low-tide site by 15.2%, including the metabolism of aminosugars, di- and oligo-saccharides, glycoside hydrolases, and monosaccharides, leading to significant decreases in 21 soil metabolites, such as monosaccharide (l-gulose), disaccharide (sucrose and turanose), and oligosaccharides (stachyose and maltotetraose). Our study demonstrates that elevated salinity due to sea-level rise may suppress C-cycling genes and their metabolites, therefore having negative impacts on microbial metabolism of organic matter.

Asexual reproduction and vegetative growth of Bionectria ochroleuca in response to temperature and photoperiod.

Growth and reproduction are two essential life-history traits for fungi. Understanding life-history strategies provides insight into the environmental adaption of species. Here, we investigated the colonial morphology, vegetative growth, and asexual reproduction of the ascomycete fungus Bionectria ochroleuca in response to a variety of environmental conditions. We demonstrated that the increased temperature from 15 to 25°C induced mycelial growth and conidiation in B. ochroleuca. We also found that the optimal temperatures for mycelial growth and conidial formation in this fungus species were 25 and 30°C, respectively. However, as the temperature increased from 25 to 30°C, mycelial growth was suppressed, but the total number of conidia was significantly increased. The shift in light-dark cycles dramatically changed the morphological features of the colonies and affected both vegetative growth and asexual reproduction. Under incubation environments of alternating light and dark (16:8 and 8:16 light:dark cycles), conidiophores and conidia in the colonies formed dense-sparse rings and displayed synchronous wave structures. When the light duration was prolonged in the sequence of 0, 8, 16, and 24 hr per day, mycelial growth was suppressed, but conidiation was promoted. Together, our results indicate that temperature and light period may trigger a trade-off between vegetative growth and asexual reproduction in B. ochroleuca.

Compatible Mycorrhizal Types Contribute to a Better Design for Mixed Eucalyptus Plantations.

Mixed-species forest plantation is a sound option to facilitate ecological restoration, plant diversity and ecosystem functions. Compatible species combinations are conducive to reconstruct plant communities that can persist at a low cost without further management and even develop into natural forest communities. However, our understanding of how the compatibility of mycorrhizal types mediates species coexistence is still limited, especially in a novel agroforestry system. Here, we assessed the effects of mycorrhizal association type on the survival and growth of native woody species in mixed-species Eucalyptus plantations. To uncover how mycorrhizal type regulates plant-soil feedbacks, we first conducted a pot experiments by treating distinct mycorrhizal plants with soil microbes from their own or other mycorrhizal types. We then compared the growth response of arbuscular mycorrhizal plants and ectomycorrhizal plants to different soil microbial compositions associated with Eucalyptus plants. We found that the type of mycorrhizal association had a significant impact on the survival and growth of native tree species in the Eucalyptus plantations. The strength and direction of the plant-soil feedbacks of focal tree species depended on mycorrhizal type. Non-mycorrhizal plants had consistent negative feedbacks with the highest survival in the Eucalyptus plantations, whereas nitrogen-fixing plants had consistent positive feedbacks and the lowest survival. Arbuscular mycorrhizal and ectomycorrhizal plants performed varied feedback responses to soil microbes from distinct mycorrhizal plant species. Non-mycorrhizal plants grew better with Eucalyptus soil microbes while nitrogen-fixing plants grew worse with their own conspecific soil microbes. Different soil microbial compositions of Eucalyptus consistently increased the aboveground growth of arbuscular mycorrhizal plants, but the non-mycorrhizal microbial composition of the Eucalyptus soil resulted in greater belowground growth of ectomycorrhizal plants. Overall, Eucalyptus plants induced an unfavorable soil community, impeding coexistence with other mycorrhizal plants. Our study provides consistent observational and experimental evidence that mycorrhizal-mediated plant-microbial feedback on species coexistence among woody species. These findings are with important implications to optimize the species combinations for better design of mixed forest plantations.

Soil fungal networks moderate density-dependent survival and growth of seedlings.

Pathogenic and mutualistic fungi have contrasting effects on seedling establishment, but it remains unclear whether density-dependent survival and growth are regulated by access to different types of mycorrhizal fungal networks supported by neighbouring adult trees. Here, we conducted an extensive field survey to test how mycorrhizal and pathogenic fungal colonization of arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) seedlings in a subtropical forest respond to density of neighbouring adult trees. In addition, we undertook a hyphal exclusion experiment to explicitly test the role of soil fungal networks in driving density-dependent effects on seedling growth and survival. Conspecific adult density was a strong predictor for the relative abundance of putative pathogens, which was greater in roots of AM than of ECM seedlings, while mycorrhizal fungal abundance and colonization were not consistently affected by conspecific adult density. Both ECM and AM fungal networks counteracted conspecific density-dependent mortality, but ECM fungi were more effective at weakening the negative effects of high seedling density than AM fungi. Our findings reveal a critical role of common fungal networks in mitigating negative density-dependent effects of pathogenic fungi on seedling establishment, which provides mechanistic insights into how soil fungal diversity shapes plant community structure in subtropical forests.

The effects of density dependence and habitat preference on species coexistence and relative abundance.

In plant communities, some mechanisms maintain differences in species' abundances, while other mechanisms promote coexistence. Asymmetry in conspecific negative density dependence (CNDD) and/or habitat preference is hypothesized to shape relative species abundance, whereas community compensatory trends (CCTs) induced by community-level CNDD and heterospecific facilitation are hypothesized to promote coexistence. We use survey data from three 1-ha permanent dynamic plots in a subtropical forest over the course of a decade to find out which of these processes are important and at which life-history stages (the seedling, sapling, and juvenile stages) they exert their effects. CNDD was not related to abundance in any of the life-history stages. Suitable habitats positively influenced plant abundance at all tested life stages, but especially so for juveniles. Community-level CNDD of seedling neighbors was detected at the seedling stage, while heterospecific facilitation was detected across all tested life-history stages. A CCT in seedling survival was detected, but there was no evidence for such trends across the other life-history stages. Altogether, our results suggest that habitat specificity increases the rarity of species, whereas a CCT at the seedling stage, which is likely to be induced by CNDD and heterospecific facilitation, enables such species to maintain their populations.

Light and competition alter leaf stoichiometry of introduced species and native mangrove species.

Ecological stoichiometry is the study of the balance of ecosystem energy and nutrient cycling, especially carbon (C), nitrogen (N), and phosphorus (P). C, N, and P are the key elements for plant growth and metabolism. Systematic research on leaf stoichiometry in mangrove forest ecosystems is still lacking. To understand the leaf stoichiometry of introduced species and native species in mangrove forests, we selected four species (one introduced species, Sonneratia apetala, and three native species, Avicennia marina, Aegiceras corniculatum, and Kandelia obovate) and measured leaf C, N, and P contents under different light conditions. The results showed that there were significant negative scaling relationships of leaf C versus N and C versus P but positive scaling relationships of leaf N versus P in the four mangrove species. Light and competition had significant effects on leaf stoichiometry, especially under the full light condition. S. apetala influenced leaf elements in a mixture with native species. Interspecific competition reduced leaf N and P contents in A. corniculatum and K. obovate but increased leaf N and P contents in A. marina. Leaf N and P contents of the four species showed similar responses to both intraspecific and interspecific competition. The ratio of leaf C:N:P (108:11:1) in the mangrove forests was lower than that in other ecosystems, and species with a higher growth rate had a higher leaf P content and lower N:P ratio, supporting the growth rate hypothesis. Leaf N:P was 11.04, indicating that there was N limitation in the mangrove forests. This systematic research of leaf stoichiometry of mangrove forests improves our understanding of mangrove growth and nutrient use strategies in response to different environmental stresses.

Neighboring trees regulate the root-associated pathogenic fungi on the host plant in a subtropical forest.

Root-associated fungi and host-specific pathogens are major determinants of species coexistence in forests. Phylogenetically related neighboring trees can strongly affect the fungal community structure of the host plant, which, in turn, will affect the ecological processes. Unfortunately, our understanding of the factors influencing fungal community composition in forests is still limited. In particular, investigation of the relationship between the phytopathogenic fungal community and neighboring trees is incomplete. In the current study, we tested the host specificity of members of the root-associated fungal community collected from seven tree species and determined the influence of neighboring trees and habitat variation on the composition of the phytopathogenic fungal community of the focal plant in a subtropical evergreen forest. Using high-throughput sequencing data with respect to the internal transcribed spacer (ITS) region, we characterized the community composition of the root-associated fungi and found significant differences with respect to fungal groups among the seven tree species. The density of conspecific neighboring trees had a significantly positive influence on the relative abundance of phytopathogens, especially host-specific pathogens, while the heterospecific neighbor density had a significant negative impact on the species richness of host-specific pathogens, as well as phytopathogens. Our work provides evidence that the root-associated phytopathogenic fungi of a host plant depend greatly on the tree neighbors of the host plant.

Soil fungal networks maintain local dominance of ectomycorrhizal trees.

The mechanisms regulating community composition and local dominance of trees in species-rich forests are poorly resolved, but the importance of interactions with soil microbes is increasingly acknowledged. Here, we show that tree seedlings that interact via root-associated fungal hyphae with soils beneath neighbouring adult trees grow faster and have greater survival than seedlings that are isolated from external fungal mycelia, but these effects are observed for species possessing ectomycorrhizas (ECM) and not arbuscular mycorrhizal (AM) fungi. Moreover, survival of naturally-regenerating AM seedlings over ten years is negatively related to the density of surrounding conspecific plants, while survival of ECM tree seedlings displays positive density dependence over this interval, and AM seedling roots contain greater abundance of pathogenic fungi than roots of ECM seedlings. Our findings show that neighbourhood interactions mediated by beneficial and pathogenic soil fungi regulate plant demography and community structure in hyperdiverse forests.

Linking Aboveground Traits to Root Traits and Local Environment: Implications of the Plant Economics Spectrum.

The plant economics spectrum proposes that ecological traits are functionally coordinated and adapt along environmental gradients. However, empirical evidence is mixed about whether aboveground and root traits are consistently linked and which environmental factors drive functional responses. Here we measure the strength of relationships between aboveground and root traits, and examine whether community-weighted mean trait values are adapted along gradients of light and soil fertility, based on the seedling censuses of 57 species in a subtropical forest. We found that aboveground traits were good predictors of root traits; specific leaf area, dry matter, nitrogen and phosphorus content were strongly correlated with root tissue density and specific root length. Traits showed patterns of adaptation along the gradients of soil fertility and light; species with fast resource-acquisitive strategies were more strongly associated with high soil phosphorus, potassium, openness, and with low nitrogen, organic matter conditions. This demonstrates the potential to estimate belowground traits from known aboveground traits in seedling communities, and suggests that soil fertility is one of the main factors driving functional responses. Our results extend our understanding of how ecological strategies shape potential responses of plant communities to environmental change.

Soil microbes drive phylogenetic diversity-productivity relationships in a subtropical forest.

The relationship between plant diversity and productivity and the mechanisms underpinning that relationship remain poorly resolved in species-rich forests. We combined extensive field observations and experimental manipulations in a subtropical forest to test how species richness (SR) and phylogenetic diversity (PD) interact with putative root-associated pathogens and how these interactions mediate diversity-productivity relationships. We show that (i) both SR and PD were positively correlated with biomass for both adult trees and seedlings across multiple spatial scales, but productivity was best predicted by PD; (ii) significant positive relationships between PD and productivity were observed in nonsterile soil only; and (iii) root fungal diversity was positively correlated with plant PD and SR, while the relative abundance of putative pathogens was negatively related to plant PD. Our findings highlight the key role of soil pathogenic fungi in tree diversity-productivity relationships and suggest that increasing PD may counteract negative effects of plant-soil feedback.

Effect of root exudates of Eucalyptus urophylla and Acacia mearnsii on soil microbes under simulated warming climate conditions.

BACKGROUND: Recent studies demonstrated that warming and elevated carbon dioxide (CO2) indirectly affect the soil microbial community structure via plant root exudates. However, there is no direct evidence for how the root exudates affect soil microbes and how the compositions of root exudates respond to climate change. RESULTS: The results showed that warming directly decreased biomass of soil-borne bacteria and fungi for Acacia mearnsii De Willd but it did not impact soil microbial community for Eucalyptus urophylla S.T. Blake. In contrast, elevated CO2 had strong direct effect on increasing soil microbial biomass for both plant species. However, plant roots could significantly increase the secretion of antibacterial chemicals (most probable organic acids), which inhibited the growth of bacteria and fungi in elevated CO2 environment. This inhibitory effect neutralized the facilitation from increasing CO2 concentration on microbial growth. CONCLUSIONS: We concluded that climate change can directly affect microorganisms, and indirectly affect the soil microbial community structure by changes in composition and content of plant root exudates.

Effects of neighborhood trait composition on tree survival differ between drought and postdrought periods.

Although direct tree demographic responses to drought are widely recognized, studies of drought-mediated changes in tree interactions are rare. We hypothesize that drought exacerbates soil-water limitation and intensifies competition for water, but reduces light limitation and competition for light. We predict that competition would be stronger for trees (1) consuming more water or more susceptible to water deficits during drought and (2) intercepting more light or more susceptible to shade during postdrought periods. We tested these predictions in a 50-ha tropical forest plot by quantifying the effects of neighborhood mean trait values on tree survival during versus after a severe drought. We used wood density (WD) and leaf mass per area (LMA) as proxies for water and light use strategies, respectively. Tree survival was lower, canopy loss was greater, and sapling recruitment was greater during the drought relative to postdrought census intervals. This suggests that drought pushed water deficits to lethal extremes and increased understory light availability. Relationships between survival and neighborhood WD were independent of drought, which is inconsistent with our first prediction. In contrast, relationships between survival and neighborhood LMA differed strongly with drought. Survival time was unaffected by neighborhood LMA during drought, but was longer for trees of all sizes in low-LMA neighborhoods in the postdrought census interval, consistent with the prediction of reduced competition for light during drought. Our results suggest that severe drought might increase light availability and reduce competition for light in moist tropical forests.

Habitat effects on intra-species variation in functional morphology: Evidence from freshwater fish.

Biotic-environment interactions have long been considered an important factor in functional phenotype differentiation in organisms. The differentiation processes determining functional phenotypes can reveal important mechanisms yielding differences in specific functions of animal traits in the ecosystem. In the present study, we examined functional morphological variations in relation to increasing geographic altitude. Six fish species were examined for how environment factors affect intra-specific functional morphology in the subtropical Pearl River in southern China. Functional morphology traits revealed variable effects due to geographic elevation, although spatial autocorrelation existed among the species tested. The results showed that high-elevation individuals had a more narrow-bodied morphology, with more flexible maneuvrability when swimming, and more evenly distributed musculature than low-elevation individuals. Low-elevation individuals preyed upon larger food sources than high-elevation individuals in some species. Fish functional morphology was strongly affected by regional environmental factors (such as elevation and water temperature) and physical characteristics of local rivers (such as flow velocity, river fractals, and coefficients of fluvial facies). In addition, the effects of the regional factors were stronger than those of the local factors in the Pearl River. Furthermore, it was found that morphological traits associated with locomotion were primarily effected by the river's physical characteristics. While morphological traits associated with food acquisition were primarily affected by water chemical factors (such as DO, water clarity, NH 4-N concentration, and TDS). These results demonstrated that habitat has an influence on the biological morphology of fish species, which further affects the functioning of the organism within the ecosystem.

Development of polymorphic EST-SSR markers in Itea chinensis (Iteaceae) and cross-amplification in related species.

PREMISE OF THE STUDY: We isolated and characterized 16 expressed sequence tag-simple sequence repeat (EST-SSR) markers in Itea chinensis (Iteaceae), a common evergreen broadleaf tree, for future studies on the genetic diversity and spatial genetic structure of the species. METHODS AND RESULTS: Based on transcriptome data of I. chinensis, a total of 36 primer pairs were initially designed and tested. Of these, 16 were successfully amplified and showed clear polymorphism. For these markers, the number of alleles per locus varied from two to 15. The observed and expected heterozygosity ranged from 0 to 0.600 and 0.072 to 0.554, respectively. Furthermore, all loci were successfully cross-amplified in two congeneric species, I. oblonga and I. yangchunensis. CONCLUSIONS: The EST-SSR markers described here can be used to study the genetic diversity and phylogeographic patterns of I. chinensis and other related species in Itea.

Abundance-dependent effects of neighbourhood dissimilarity and growth rank reversal in a neotropical forest.

Why tropical forests harbour an exceptional number of species with striking differences in abundances remains an open question. We propose a theoretical framework to address this question in which rare species may have different extirpation risks depending on species ranks in tree growth and sensitivities to neighbourhood interactions. To evaluate the framework, we studied tree growth and its responses to neighbourhood dissimilarity (ND) in traits and phylogeny for 146 species in a neotropical forest. We found that tree growth was positively related to ND, and common species were more strongly affected by ND than rare species, which may help delay dominance of common species. Rare species grew more slowly at the community-wide average ND than common species. But rare species grew faster when common species tended to dominate locally, which may help reduce extirpation risk of rare species. Our study highlights that tree growth rank among species depends on their responses to neighbourhood interactions, which can be important in fostering diversity maintenance in tropical forests.