Cultivated alien plants with high invasion potential are more likely to be traded online in China.

Biological invasions have become a worldwide problem, and measures to efficiently prevent and control invasions are still in development. Like many other parts of the world, China is undergoing a dramatic increase in plant invasions. Most of the currently 933 established (i.e., naturalized) plant species, of which 214 are categorized as invasive, have been introduced into China for cultivation. It is likely that many of those species are still being traded, particularly online, by plant nurseries. However, studies assessing whether naturalized and invasive species are currently being traded more or less than nonnaturalized aliens are rare. We extracted online-trade information for 13,718 cultivated alien plant taxa on 1688.com, the largest website for domestic B2B in China. We analyzed how the presence in online-nursery catalogs, the number of online nurseries that offerred the species for sale, and the product type (i.e., seeds, live plants and vegetative organs) differed among nonnaturalized, naturalized noninvasive, and invasive species. Compared to nonnaturalized taxa, naturalized noninvasive and invasive taxa were 3.7-5.2 times more likely to be available for purchase. Naturalized noninvasive and invasive taxa were more frequently offered as seeds by online nurseries, whereas nonnaturalized taxa were more frequently offered as live plants. Based on these findings, we propose that, to reduce the further spread of invasive and potentially invasive plants, implementation of plant-trade regulations and a monitoring system of the online horticultural supply chain will be essential.

Dose dependent effect of nitrogen on the phyto extractability of Cd in metal contaminated soil using Wedelia trilobata.

Cadmium (Cd) is one of the toxic heavy metal that negatively affect plant growth and compromise food safety for human consumption. Nitrogen (N) is an essential macronutrient for plant growth and development. It may enhance Cd tolerance of invasive plant species by maintaining biochemical and physiological characteristics during phytoextraction of Cd. A comparative study was conducted to evaluate the phenotypical and physiological responses of invasive W. trilobata and native W. chinensis under low Cd (10 µM) and high Cd (80 µM) stress, along with different N levels (i.e., normal 91.05 mg kg-1 and low 0.9105 mg kg-1). Under low-N and Cd stress, the growth of leaves, stem and roots in W. trilobata was significantly increased by 35-23%, 25-28%, and 35-35%, respectively, compared to W. chinensis. Wedelia trilobata exhibited heightened antioxidant activities of catalase and peroxidase were significantly increased under Cd stress to alleviate oxidative stress. Similarly, flavonoid content was significantly increased by 40-50% in W. trilobata to promote Cd tolerance via activation of the secondary metabolites. An adverse effect of Cd in the leaves of W. chinensis was further verified by a novel hyperspectral imaging technology in the form of normalized differential vegetation index (NDVI) and photochemical reflectance index (PRI) compared to W. trilobata. Additionally, W. trilobata increased the Cd tolerance by regulating Cd accumulation in the shoots and roots, bolstering its potential for phytoextraction potential. This study demonstrated that W. trilobata positively responds to Cd with enhanced growth and antioxidant capabilities, providing a new platform for phytoremediation in agricultural lands to protect the environment from heavy metals pollution.

Pathogen resistance in Sphagneticola trilobata (Singapore daisy): molecular associations and differentially expressed genes in response to disease from a widespread fungus.

Understanding the molecular associations underlying pathogen resistance in invasive plant species is likely to provide useful insights into the effective control of alien plants, thereby facilitating the conservation of native biodiversity. In the current study, we investigated pathogen resistance in an invasive clonal plant, Sphagneticola trilobata, at the molecular level. Sphagneticola trilobata (i.e., Singapore daisy) is a noxious weed that affects both terrestrial and aquatic ecosystems, and is less affected by pathogens in the wild than co-occurring native species. We used Illumina sequencing to investigate the transcriptome of S. trilobata following infection by a globally distributed generalist pathogen (Rhizoctonia solani). RNA was extracted from leaves of inoculated and un-inoculated control plants, and a draft transcriptome of S. trilobata was generated to examine the molecular response of this species following infection. We obtained a total of 49,961,014 (94.3%) clean reads for control (un-inoculated plants) and 54,182,844 (94.5%) for the infected treatment (inoculated with R. solani). Our analyses facilitated the discovery of 117,768 de novo assembled contigs and 78,916 unigenes. Of these, we identified 3506 differentially expressed genes and 60 hormones associated with pathogen resistance. Numerous genes, including candidate genes, were associated with plant-pathogen interactions and stress response in S. trilobata. Many recognitions, signaling, and defense genes were differentially regulated between treatments, which were confirmed by qRT-PCR. Overall, our findings improve our understanding of the genes and molecular associations involved in plant defense of a rapidly spreading invasive clonal weed, and serve as a valuable resource for further work on mechanism of disease resistance and managing invasive plants.

Allelopathy confers an invasive Wedelia higher resistance to generalist herbivore and pathogen enemies over its native congener.

The Novel Defense Hypothesis predicts that introduced plants may possess novel allelochemicals which act as a defense against native generalist enemies. Here, we aim to test if the chemicals involved in allelopathy in the invasive plant Wedelia trilobata can contribute to higher resistance against generalist herbivore and pathogen enemies by comparing with its native congener W. chinensis in controlled laboratory conditions. The allelopathic effects of the leaf extract from W. trilobata on the generalist enemies were also assessed. We showed that the larvae of two moth species preferred W. chinensis over W. trilobata. The growth rate of larvae feeding on W. trilobata leaves was significantly lower than those feeding on W. chinensis leaves. When detached leaves were inoculated with phytopathogens, the infected leaf area of W. trilobata was significantly smaller than that of W. chinensis. In addition, the leaf extract of W. trilobata also effectively inhibited the growth of the larvae and the mycelial growth of the phytopathogens. Our results indicate that the defenses of invasive W. trilobata against generalist herbivore and pathogen enemies are stronger than that of its native congener, which may be attributed to the allelopathic effects. This study provides novel insights that can comprehensively link the Novel Defense, Behavioral Constraint and Enemy Release hypotheses. These combined hypotheses would explain how invasive plants escape from their natural specialist enemies, where their allelopathic chemicals may deter herbivorous insects and inhibit pathogen infection.

Gene Expression Profiling Reveals Enhanced Defense Responses in an Invasive Weed Compared to Its Native Congener During Pathogenesis.

Invasive plants are a huge burden on the environment, and modify local ecosystems by affecting the indigenous biodiversity. Invasive plants are generally less affected by pathogens, although the underlying molecular mechanisms responsible for their enhanced resistance are unknown. We investigated expression profiles of three defense hormones (salicylic acid, jasmonic acid, and ethylene) and their associated genes in the invasive weed, Alternanthera philoxeroides, and its native congener, A. sessilis, after inoculation with Rhizoctonia solani. Pathogenicity tests showed significantly slower disease progression in A. philoxeroides compared to A. sessilis. Expression analyses revealed jasmonic acid (JA) and ethylene (ET) expressions were differentially regulated between A. philoxeroides and A. sessilis, with the former having prominent antagonistic cross-talk between salicylic acid (SA) and JA, and the latter showing weak or no cross-talk during disease development. We also found that JA levels decreased and SA levels increased during disease development in A. philoxeroides. Variations in hormonal gene expression between the invasive and native species (including interspecific differences in the strength of antagonistic cross-talk) were identified during R. solani pathogenesis. Thus, plant hormones and their cross-talk signaling may improve the resistance of invasive A. philoxeroides to pathogens, which has implications for other invasive species during the invasion process.

Different Responses of an Invasive Clonal Plant Wedelia trilobata and its Native Congener to Gibberellin: Implications for Biological Invasion.

The invasive clonal plant Wedelia trilobata contains higher levels of ent-kaurane diterpenes, which are precursors of gibberellins (GAs), and higher rates of clonal growth than its native congener W. chinensis in invaded habitats. We hypothesized that the higher levels of endogenous GAs facilitate greater ramet growth in W. trilobata compared with W. chinensis. We quantified endogenous levels of GA1+3 in the two species and compared their growth responses to the changes of endogenous and exogenous GA3 by using short-term and long-term hydroponics experiments. After a period of homogeneous cultivation, levels of endogenous GA1+3 were higher in W. trilobata than in W. chinensis. The reduction of endogenous GAs repressed the emergence of adventitious roots and the growth of W. trilobata in the initial cultivation stage, and inhibited its shoot elongation and biomass. Levels of endogenous GA1+3 were positively correlated with the length of shoots and adventitious roots of W. trilobata. Adventitious roots of W. trilobata also emerged earlier and grew faster when treated with exogenous GA3. In contrast, exogenous GA3 treatment inhibited the length of adventitious roots in W. chinensis, and levels of endogenous GA1+3 did not correlate with shoot or adventitious root length. Our study suggests that GAs accelerate the rapid clonal growth of W. trilobata, more than that of its native congener W. chinensis, illustrating the relationship between plant hormones and the clonal growth of invasive plants. These findings are important for understanding the mechanisms associated with the invasiveness of clonal plants and their potential management.

Deforestation and fragmentation of natural forests in the upper Changhua watershed, Hainan, China: implications for biodiversity conservation.

Hainan, the largest tropical island in China, belongs to the Indo-Burma biodiversity hotspot. The Changhua watershed is a center of endemism for plants and birds and the cradle of Hainan's main rivers. However, this area has experienced recent and ongoing deforestation and habitat fragmentation. To quantify habitat loss and fragmentation of natural forests, as well as the land-cover changes in the Changhua watershed, we analyzed Landsat images obtained in 1988, 1995, and 2005. Land-cover dynamics analysis showed that natural forests increased in area (97,909 to 104,023 ha) from 1988 to 1995 but decreased rapidly to 76,306 ha over the next decade. Rubber plantations increased steadily throughout the study period while pulp plantations rapidly expanded after 1995. Similar patterns of land cover change were observed in protected areas, indicating a lack of enforcement. Natural forests conversion to rubber and pulp plantations has a general negative effect on biodiversity, primarily through habitat fragmentation. The fragmentation analysis showed that natural forests area was reduced and patch number increased, while patch size and connectivity decreased. These land-cover changes threatened local biodiversity, especially island endemic species. Both natural forests losses and fragmentation should be stopped by strict enforcement to prevent further damage. Preserving the remaining natural forests and enforcing the status of protected areas should be a management priority to maximize the watershed's biodiversity conservation value.

Light limitation and litter of an invasive clonal plant, Wedelia trilobata, inhibit its seedling recruitment.

BACKGROUND AND AIMS: Invasive clonal plants have two reproduction patterns, namely sexual and vegetative propagation. However, seedling recruitment of invasive clonal plants can decline as the invasion process proceeds. For example, although the invasive clonal Wedelia trilobata (Asteraceae) produces numerous seeds, few seedlings emerge under its dense population canopy in the field. In this study it is hypothesized that light limitation and the presence of a thick layer of its own litter may be the primary factors causing the failure of seedling recruitment for this invasive weed in the field. METHODS: A field survey was conducted to determine the allocation of resources to sexual reproduction and seedling recruitment in W. trilobata. Seed germination was also determined in the field. Effects of light and W. trilobata leaf extracts on seed germination and seedling growth were tested in the laboratory. KEY RESULTS: Wedelia trilobata blooms profusely and produces copious viable seeds in the field. However, seedlings of W. trilobata were not detected under mother ramets and few emerged seedlings were found in the bare ground near to populations. In laboratory experiments, low light significantly inhibited seed germination. Leaf extracts also decreased seed germination and inhibited seedling growth, and significant interactions were found between low light and leaf extracts on seed germination. However, seeds were found to germinate in an invaded field after removal of the W. trilobata plant canopy. CONCLUSIONS: The results indicate that lack of light and the presence of its own litter might be two major factors responsible for the low numbers of W. trilobata seedlings found in the field. New populations will establish from seeds once the limiting factors are eliminated, and seeds can be the agents of long-distance dispersal; therefore, prevention of seed production remains an important component in controlling the spread of this invasive clonal plant.

Strong Invasive Mechanism of Wedelia trilobata via Growth and Physiological Traits under Nitrogen Stress Condition.

Nitrogen (N) is one of the most crucial elements for plant growth. However, a deficiency of N affects plant growth and development. Wedelia trilobata is a notorious invasive plant species that exhibits superior tolerance to adapt to environmental stresses. Yet, research on the growth and antioxidant defensive system of invasive Wedelia under low N stress, which could contribute to understanding invasion mechanisms, is still limited. Therefore, this study aims to investigate and compare the tolerance capability of invasive and native Wedelia under low and normal N conditions. Native and invasive Wedelia species were grown in normal and low-N conditions using a hydroponic nutrient solution for 8 weeks to assess the photosynthetic parameters, antioxidant activity, and localization of reactive oxygen species (ROS). The growth and biomass of W. trilobata were significantly (p < 0.05) higher than W. chinensis under low N. The leaves of W. trilobata resulted in a significant increase in chlorophyll a, chlorophyll b, and total chlorophyll content by 40.2, 56.2, and 46%, respectively, compared with W. chinensis. W. trilobata significantly enhanced antioxidant defense systems through catalase, peroxidase, and superoxide dismutase by 18.6%, 20%, and 36.3%, respectively, providing a positive response to oxidative stress caused by low N. The PCA analysis showed that W. trilobata was 95.3% correlated with physiological traits by Dim1 (79.1%) and Dim2 (16.3%). This study provides positive feedback on W. trilobata with respect to its comprehensive invasion mechanism to improve agricultural systems via eco-friendly approaches in N deficit conditions, thereby contributing to the reclamation of barren land.

Plant growth promoting rhizobacteria and biochar production from Parthenium hysterophorus enhance seed germination and productivity in barley under drought stress.

Drought stress can significantly affect plant growth and development. Biochar (BC) and plant growth-promoting rhizobacteria (PGPR) have been found to increase plant fertility and development under drought conditions. The single effects of BC and PGPR in different plant species have been widely reported under abiotic stress. However, there have been relatively few studies on the positive role of PGPR, BC, and their combination in barley (Hordeum vulgare L.). Therefore, the current study investigated the effects of BC from Parthenium hysterophorus, drought tolerant PGPR (Serratia odorifera), and the combination of BC + PGPR on the growth, physiology, and biochemical traits of barley plants under drought stress for two weeks. A total of 15 pots were used under five treatments. Each pot of 4 kg soil comprised the control (T0, 90% water), drought stress alone (T1, 30% water), 35 mL PGPR/kg soil (T2, 30% water), 2.5%/kg soil BC (T3, 30% water), and a combination of BC and PGPR (T4, 30% water). Combined PGPR and BC strongly mitigated the negative effects of drought by improving the shoot length (37.03%), fresh biomass (52%), dry biomass (62.5%), and seed germination (40%) compared to the control. The PGPR + BC amendment treatment enhanced physiological traits, such as chlorophyll a (27.9%), chlorophyll b (35.3%), and total chlorophyll (31.1%), compared to the control. Similarly, the synergistic role of PGPR and BC significantly (p< 0.05) enhanced the antioxidant enzyme activity including peroxidase (POD), catalase (CAT), and superoxide dismutase (SOD) to alleviate the toxicity of ROS. The physicochemical properties (N, K, P, and EL) of the soils were also enhanced by (85%, 33%, 52%, and 58%) respectively, under the BC + PGPR treatment compared to the control and drought stress alone. The findings of this study have suggested that the addition of BC, PGPR, and a combination of both will improve the soil fertility, productivity, and antioxidant defense systems of barley under drought stress. Therefore, BC from the invasive plant P. hysterophorus and PGPR can be applied to water-deficient areas to improve barley crop production.

Invasive Plant Alternanthera philoxeroides Benefits More Competition Advantage from Rhizosphere Bacteria Regardless of the Host Source.

The rhizosphere plays a vital role in the exchange of materials in the soil-plant ecosystem, and rhizosphere microorganisms are crucial for plant growth and development. In this study, we isolated two strains of Pantoea rhizosphere bacteria separately from invasive Alternanthera philoxeroides and native A. sessilis. We conducted a control experiment to test the effects of these bacteria on the growth and competition of the two plant species using sterile seedlings. Our findings showed that the rhizobacteria strain isolated from A. sessilis significantly promoted the growth of invasive A. philoxeroides in monoculture compared to native A. sessilis. Both strains significantly enhanced the growth and competitiveness of invasive A. philoxeroides under competition conditions, regardless of their host source. Our study suggests that rhizosphere bacteria, including those from different host sources, can contribute to the invasion of A. philoxeroides by significantly enhancing its competitiveness.

A Green Approach Used for Heavy Metals 'Phytoremediation' Via Invasive Plant Species to Mitigate Environmental Pollution: A Review.

Heavy metals (HMs) normally occur in nature and are rapidly released into ecosystems by anthropogenic activities, leading to a series of threats to plant productivity as well as human health. Phytoremediation is a clean, eco-friendly, and cost-effective method for reducing soil toxicity, particularly in weedy plants (invasive plant species (IPS)). This method provides a favorable tool for HM hyperaccumulation using invasive plants. Improving the phytoremediation strategy requires a profound knowledge of HM uptake and translocation as well as the development of resistance or tolerance to HMs. This review describes a comprehensive mechanism of uptake and translocation of HMs and their subsequent detoxification with the IPS via phytoremediation. Additionally, the improvement of phytoremediation through advanced biotechnological strategies, including genetic engineering, nanoparticles, microorganisms, CRISPR-Cas9, and protein basis, is discussed. In summary, this appraisal will provide a new platform for the uptake, translocation, and detoxification of HMs via the phytoremediation process of the IPS.

Rubber and pulp plantations represent a double threat to Hainan's natural tropical forests.

Hainan, the largest tropical island in China, belongs to the Indo-Burma biodiversity hotspot and harbors large areas of tropical forests, particularly in the uplands. The Changhua watershed is the cradle of Hainan's main river and a center of endemism for plants and birds. The watershed contains great habitat diversity and is an important conservation area. We analyzed the impact of rubber and pulp plantations on the distribution and area of tropical forest in the watershed, using remote sensing analysis of Landsat images from 1988, 1995 and 2005. From 1988 to 1995, natural forest increased in area (979-1040 sq km) but decreased rapidly (763 sq km) over the next decade. Rubber plantations increased steadily through the study period while pulp plantations appeared after 1995 but occupied 152 sq km by 2005. Rubber and pulp plantations displace different types of natural forest and do not replace one another. Because pulp is not as profitable as rubber and existing pulp processing capacity greatly exceeds local supply, considerable pressure exists on remaining upland forests. We recommend for future management that these plantation forests be reclassified as 'industrial', making a clear policy distinction between natural and industrial forestry. Additionally, the local government should work to enforce existing laws preventing forest conversion on marginal and protected areas.

Both Adaptability and Endophytic Bacteria Are Linked to the Functional Traits in the Invasive Clonal Plant Wedelia trilobata.

The role of the interactions between endophytes and host plants is unclear in invasive plants from different geographical latitudes. In this study, we aimed to explore the relationship between endophytic microbes and the functional traits of the invasive plant Wedelia trilobata. We explored the relationship between endophytes and the clonal growth traits of the invasive clonal plant Wedelia trilobata from different geographical latitudes using high-throughput sequencing technology and a common garden-planting experiment. We found that: (1) Different W. trilobata populations had similar endophytic fungi but different endophytic bacteria. However, no latitudinal variation pattern of the overall microbial community was found; (2) plant clonal growth performance (i.e., spacer length) was significantly correlated with endophytic bacterial diversity but not fungal diversity; and (3) the latitudinal variation pattern of the plant clonal growth performance of W. trilobata populations was found in pre-cultivated (i.e., wild) individuals but disappeared in post-cultivated W. trilobata. Our results suggest both environmental adaptability and the endophytic bacterial community are linked to the functional traits of the invasive clonal plant W. trilobata, and these functional traits tend to increase its invasiveness, which may enhance its invasion success.

Cadmium hyperaccumulation as an inexpensive metal armor against disease in Crofton weed.

Invasive plants readily invade metal-contaminated areas. The hyperaccumulation of toxic heavy metals is not an uncommon feature among plant species. Although several hypotheses were proposed to explain this phenomenon, it is currently unclear how hyperaccumulation may benefit plants. The invasive Crofton weed (Ageratina adenophora) is a known hyperaccumulator of chromium and lead. We previously found that the species can also hyperaccumulate cadmium. The role of phytoaccumulation in defense to pathogen attack is unclear. We inoculated A. adenophora plants with a common generalist pathogen (Rhizoctonia solani) to test its resistance under cadmium treatment. We found evidence that cadmium hyperaccumulation reduced pathogen infection in A. adenophora. Our findings indicate elemental defense is highly cost efficient for hyperaccumulators inhabiting metal-contaminated sites, where plants were only modestly affected by cadmium. The reduction in pathogen damage conferred by cadmium was relatively high, particularly under lower cadmium levels. However, the benefits at higher levels may be capped. Elemental defense may be a key mechanism for plant invasion into polluted sites, especially in regions with widespread industrial activity. Our study highlights the importance of testing different metal concentrations when testing plant resistance and the importance of considering enemy attack when selecting plants for phytoremediation.

Different Growth Promoting Effects of Endophytic Bacteria on Invasive and Native Clonal Plants.

The role of the interactions between endophytes and alien plants has been unclear yet in plant invasion. We used a completely germ-free culture system to quantify the plant growth-promoting (PGP) effects of endophytic bacteria Bacillus sp. on aseptic seedlings of Wedelia trilobata and of its native clonal congener W. chinensis. The endophytic bacteria did not affect the growth of W. chinensis, but they significantly promoted the growth of W. trilobata. With the PGP effects of endophytic bacteria, relative change ratios of the clonal traits and the ramets' growth traits of W. trilobata were significantly greater than those of W. chinensis. Our results indicate that the growth-promoting effects of endophytes may differ between invasive and native clonal plants, and the endophytes of invasive plant may be host-specific to facilitate plant invasion.

Curvilinear effects of invasive plants on plant diversity: plant community invaded by Sphagneticola trilobata.

The effects of invasive plants on the species diversity of plant communities are controversial, showing either a positive or negative linear relationship. Based on community data collected from forty 5 m×5 m plots invaded by Sphagneticola trilobata in eight cities across Hainan Island, China, we found S. trilobata decreased plant community diversity once its cover was beyond 10%. We demonstrated that the effects of invasive/native plants on the plant diversity of communities invaded by S. trilobata were curvilinear. These effects, which showed peaks under different degrees of vegetation cover, appeared not only for S. trilobata and all invasive plants, but also for all native plants. Invasive plants primarily had negative effects on plant diversity when they became abundant at a much lower cover level (less than 35%), compared with the native plants (over 60%). Thus, it is necessary to distinguish a range for assessing the effects of plants, especially invasive plants. Our results also confirmed that the invasion intensity of invasive alien plants increased with the intensity of local economic development. We highlight and further discuss the critical importance of curvilinear effects of biological invasion to provide ideas regarding the conservation of local biodiversity and the management of invasive plants.