Rapid use of resources as a basis of the <Emphasis Type="Italic">Heracleum sosnowskyi</Emphasis> invasive syndrome

The functional properties of the invasive Heracleum sosnowskyi were compared with those of the native Heracleum sibiricum. The leaf and root traits, as well as those of a whole plant and of the mycorrhiza formation, were studied. H. sosnowskyi can fulfill the high-productivity potential only under the optimum availability of resources (especially water), while it is poorly adapted to maintain a high physiological activity under the unfavorable soil conditions. An important component of the H. sosnowskyi invasiveness is an ability to utilize rapidly resources in both the above- and underground areas at the optimum conditions.     

The Impacts of Above- and Belowground Plant Input on Soil Microbiota: Invasive <Emphasis Type="Italic">Spartina alterniflora</Emphasis> Versus Native <Emphasis Type="Italic">Phragmites australis</Emphasis>

Invasive plants affect soil food webs through various resource inputs including shoot litter, root litter and living root input. The net impact of invasive plants on soil biota has been recognized; however, the relative contributions of different resource input pathways have not been quantified. Through a 2 × 2 × 2 factorial field experiment, a pair of invasive and native plant species (Spartina alterniflora vs. Phragmites australis) was compared to determine the relative impacts of their living roots or shoots and root litter on soil microbial and nematode communities. Living root identity affected bacteria-to-fungi PLFA ratios, abundance of total nematodes, plant-feeding nematodes and omnivorous nematodes. Specifically, the plant-feeding nematodes were 627% less abundant when living roots of invasive S. alterniflora were present than those of native P. australis. Likewise, shoot and root biomass (within soil at 0–10 cm depth) of S. alterniflora was, respectively, 300 and 100% greater than those of P. australis. These findings support the enemy release hypothesis of plant invasion. Root litter identity affected other components of soil microbiota (that is, bacterial-feeding nematodes), which were 34% more abundant in the presence of root litter of P. australis than S. alterniflora. Overall, more variation associated with nematode community structure and function was explained by differences in living roots than root or shoot litter for this pair of plant species sharing a common habitat but contrasting invasion degrees. We conclude that belowground resource input is an important mechanism used by invasive plants to affect ecosystem structure and function.     

Diversity of root-endophytic <Emphasis Type="Italic">Trichoderma</Emphasis> from Malaysian Borneo

Trichoderma species form endophytic associations with plant roots and may provide a range of benefits to their hosts. However, few studies have systematically examined the diversity of Trichoderma species associated with plant roots in tropical regions. During the evaluation of Trichoderma isolates for use as biocontrol agents, root samples were collected from more than 58 genera in 35 plant families from a range of habitats in Malaysian Borneo. Trichoderma species were isolated from surface-sterilised roots and identified following analysis of partial translation elongation factor-1α (tef1) sequences. Species present included Trichoderma afroharzianum, Trichoderma asperelloides, Trichoderma asperellum, Trichoderma guizhouense, Trichoderma reesei, Trichoderma strigosum and Trichoderma virens. Trichoderma asperellum/T. asperelloides, Trichoderma harzianum s.l. and T. virens were the most frequently isolated taxa. tef1 sequence data supported the recognition of undescribed species related to the T. harzianum complex. The results suggest that tropical plants may be a useful source of novel root-associated Trichoderma for biotechnological applications.     

Fast seedling root growth leads to competitive superiority of invasive plants

The competitive superiority of invasive plants plays a key role in the process of plant invasions, enabling invasive plants to overcome the resistance of local plant communities. Fast aboveground growth and high densities lead to the competitive superiority of invasive species in the competition for light. However, little is understood of the role belowground root competition may play in invasion. We conducted an experiment to test the effect of root growth on the performance of an invasive shrub Cassia alata, a naturalized, non-invasive shrub Corchorus capsularis, and a native shrub Desmodium reticulatum. We compared seedling growth of the three species and their competitive ability in situ. The roots of the C. alata seedlings grew much faster than those of C. capsularis and D. reticulatum during the entire growth period although C. alata had shorter shoots than D. reticulatum. Furthermore, C. alata showed an apparent competition advantage compared to the other two species as evidenced by less biomass reduction in intraspecific competition and higher competitive effects in interspecific competition. Our study reveals that fast seedling root growth may be important in explaining the competitive advantages of invasive plants. Future studies should pay more attention to the belowground traits of invasive plants, the trade-off between shoot and root growth, and the role of root competition in affecting the population dynamics of invasive plants and the structures of invaded communities.     

Evaluation of <Emphasis Type="Italic">Nastus fausti</Emphasis> Reitter (Coleoptera: Curculionidae: Entiminae: Nastini) for biological control of invasive giant hogweeds (<Emphasis Type="Italic">Heracleum</Emphasis> spp.)

The weevil Nastus fausti Reitter (Coleoptera, Curculionidae) was evaluated for its potential in the biological control of invasive giant hogweeds (Heracleum spp.). Quantitative sampling suggested that at a high population density (more that 3–4 mature larvae per plant) damage by N. fausti larvae could have some negative impact on the above-ground part of the plant. However, no-choice laboratory tests showed that N. fausti females were able to feed on a number of Apiaceae genera, including such important cultivated crops as carrot, parsnip, and celeriac. Feeding on these plants did not cause any significant decrease in female survival or fecundity. Moreover, at least part of N. fausti larvae may feed and develop on roots of these plants, and the rate of their growth and development does not differ significantly from that in larvae fed on roots of H. mantegazzianum. N. fausti adult and larval feeding on Angelica purpurascens, representative of related genus of the same tribe, was recorded under natural conditions, too. In combination, these data suggest that N. fausti is an oligophagous species connected with plants from at least several genera of Apiaceae and thus it cannot be considered a potential agent for biological control of invasive Heracleum species.     

Contribution of different arbuscular mycorrhizal fungal inoculum to <Emphasis Type="Italic">Elymus nutans</Emphasis> under nitrogen addition

Arbuscular mycorrhizal (AM) fungi are known to promote plant growth and nutrient uptake, but their role in nitrogen (N) uptake still remains unclear. Therefore, a pot experiment was set up to evaluate the impacts of N addition and AM inoculation (Diversispora eburnea, Claroideoglomus etunicatum, Paraglomus occultum, and their mixture) on AM root colonization, plant biomass, N and P nutrition in Elymus nutans. Our results showed that AM root colonization was unaffected by N addition but was significantly affected by different AM fungal species. D. eburnea and C. etunicatum showed significant higher root colonization than P. occultum. The E. nutans exhibited the highest biomass when inoculated with D. eburnea and significantly higher than non-mycorrhizal (the control) regardless of N addition. Under N addition treatment, D. eburnea significantly enhanced P content of roots, N content of shoots and roots, while AM mixture significantly enhanced shoot P content compared with non-mycorrhizal. However, N and P content in shoots and roots did not significantly vary among treatments when no N was added. In addition, inoculation with C. etunicatum and P. occultum showed no significant effect on plant biomass, N and P content regardless of N addition. In conclusion, this study revealed that the plant response to N addition depends on AM fungal species and also confirmed that significant functional diversity exists among AM fungal species.     

Effects of leaf litter on inter-specific competitive ability of the invasive plant <Emphasis Type="Italic">Wedelia trilobata</Emphasis>

Allelochemicals released by invasive plants contribute to the successful invasion of new habitats. However, the relationship between allelopathic effects and competitive ability of invasive plants has not been characterized. We quantified the neighbor effects of Wedelia trilobata (family: Asteraceae) and the allelopathic effects of its leaf litter on two Asteraceae competitor species (invasive Eupatorium catarium and non-invasive Lactuca sativa) and on its own ramet growth. The seed germination rate and seedling biomass of the two competitor species decreased following treatment with W. trilobata leaf extracts. When co-cultured with W. trilobata, the total biomass of the two competitor species significantly decreased regardless of whether leaf extracts were present. Under low plant density co-culture conditions, W. trilobata leaf extracts enhanced the inhibitory effects on E. catarium. In contrast, W. trilobata leaf extracts promoted the growth of W. trilobata adventitious roots, resulting in increased competitive ability. Therefore, W. trilobata growth was promoted by its own allelochemicals in leaf extracts, whereas the growth of the invasive and non-invasive competitors was inhibited by the same chemicals. These responses facilitated the invasion by W. trilobata. Our study demonstrates that leaf litter of invasive plants may inhibit the growth of neighboring species to enhance the competitive ability of the invasive plants during the early stages of invasion.     

Formation of phenolic compounds in the roots of <Emphasis Type="Italic">Hedysarum theinum</Emphasis> cultured in vitro

A steadily growing culture of genetically transformed roots of a valuable medicinal Altaic plant Hedysarum theinum Krasnob. was established using a Ri-plasmid (pRi) T-DNA of A4 wild strain of Agrobacterium rhizogenes. The composition of secondary substances accumulated in the in vitro roots and the roots of H. theinum intact seedlings was investigated. Isoflavonoids were found to represent by their main low-molecular metabolites. Preparative HPLC made it possible to isolate four substances from the methanolic extract of H. theinum cultured roots. Using ¹H- and ¹³C-NMR-spectrometry, these substances were identified as formononetin, ononin (formononetin glycoside), malonyl ononin, and texasin glucoside. The qualitative composition of secondary metabolites of the genetically transformed roots and the roots of H. theinum seedlings was essentially the same, except that malonyl ononin was not found in the latter. The technique of producing artificial seeds on the basis of H. theinum roots cultured in vitro was tested, and the possibility of their use as a rhizogenic inoculum was substantiated. The culture of H. theinum roots is considered as a potential source of ecologically pure raw material for medicinal preparations, and the artificial seeds with root inoculum are a promising vehicle for propagation and conservation of this valuable plant.     

Deep rooting and global change facilitate spread of invasive grass

Abiotic global change factors, such as rising atmospheric CO₂, and biotic factors, such as exotic plant invasion, interact to alter the function of terrestrial ecosystems. An invasive lineage of the common reed, Phragmites australis, was introduced to North America over a century ago, but the belowground mechanisms underlying Phragmites invasion and persistence in natural systems remain poorly studied. For instance, Phragmites has a nitrogen (N) demand higher than native plant communities in many of the ecosystems it invades, but the source of the additional N is not clear. We exposed introduced Phragmites and native plant assemblages, containing Spartina patens and Schoenoplectus americanus, to factorial treatments of CO₂ (ambient or +300 ppm), N (0 or 25 g m^?2 year^?1), and hydroperiod (4 levels), and focused our analysis on changes in root productivity as a function of depth and evaluated the effects of introduced Phragmites on soil organic matter mineralization. We report that non-native invasive Phragmites exhibited a deeper rooting profile than native marsh species under all experimental treatments, and also enhanced soil organic matter decomposition. Moreover, exposure to elevated atmospheric CO₂ induced a sharp increase in deep root production in the invasive plant. We propose that niche separation accomplished through deeper rooting profiles circumvents nutrient competition where native species have relatively shallow root depth distributions; deep roots provide access to nutrient-rich porewater; and deep roots further increase nutrient availability by enhancing soil organic matter decomposition. We expect that rising CO₂ will magnify these effects in deep-rooting invasive plants that compete using a tree-like strategy against native herbaceous plants, promoting establishment and invasion through niche separation.     

The Role of MPK6 as Mediator of Ethylene/Jasmonic Acid Signaling in <Emphasis Type="Italic">Serendipita indica</Emphasis>-Colonized Arabidopsis Roots

Serendipita indica is an axenically cultivable fungus, which colonizes a broad range of plant species including the model plant Arabidopsis thaliana. Root colonization by this endophyte leads to enhanced plant fitness and performance and promotes resistance against different biotic and abiotic stresses. The involvement of MPK6 in this mutualistic interaction had been previously shown with an mpk6 A. thaliana mutant, which failed to respond to S. indica colonization. Here, we demonstrate that mpk6 roots are significantly less colonized by S. indica compared to wild-type roots and the foliar application of plant hormones, ethylene, or jasmonic acid, restores the colonization rate at least to the wild-type level. Further, hormone-treated mpk6 plants show typical S. indica-induced growth promotion effects. Moreover, expression levels of several genes related to plant defense and hormone signaling are significantly changed at different colonization phases. Our results demonstrate that the successful root colonization by S. indica depends on efficient suppression of plant immune responses. In A. thaliana, this process relies on intact hormone signaling in which MPK6 seems to play a pivotal role.     

Effects of the density of the invasive macrophyte <Emphasis Type="Italic">Hydrilla verticillata</Emphasis> and root competition on growth of one native macrophyte in different sediment fertilities

The occurrence of non-native species at high densities may generate competition for resources and possibly exclude native species in various environments. We evaluated the effects of increased densities of the non-native invasive macrophyte Hydrilla verticillata on the growth of the native species Egeria najas in different sediment types and with only root interactions or root?+?shoot interactions. We tested the hypothesis that the effect of the invasive on the native species is density dependent and that it is greater when competition for light and nutrients occurs (root?+?shoot interactions). The results of these experiments demonstrated that increased density of the invasive species H. verticillata significantly decreased the growth of the native species independent of sediment type (sand or mud sediments). When plants competed for water and sediment resources (root?+?shoot interactions), the native species was more impacted by the invasive than when they competed only for water resources (only shoots interacting). Our results show that E. najas is probably unable to colonize sites highly colonized by hydrilla, and this applies to both sand and mud sediments. This outcome suggests that H. verticillata is a threat for E. najas and likely other native submerged species in South America.     

Heritable bacterial endosymbionts in native and invasive populations of <Emphasis Type="Italic">Harmonia axyridis</Emphasis>

Harmonia axyridis Pallas (1773) (Coleoptera: Coccinellidae) is the well-studied system of invasive insect species. Native and invasive parts of the area of H. axyridis are isolated geographically. We studied the species composition and the distribution of bacterial symbionts Spiroplasma and Rickettsia in seven localities of the native area and six localities of the invasive area of H. axyridis. Rickettsia was detected in H. axyridis populations for the first time. We found that the proportion of beetles infected with Rickettsia in native and invasive populations of H. axyridis is about 0.03. Spiroplasma was found only in native populations of H. axyridis. The proportion of infected individuals with Spiroplasma in native populations of H. axyridis is about 0.08. All studied native populations of H. axyridis are infected with Spiroplasma, while all invasive populations are not. We discuss the possible influence of Spiroplasma and Rickettsia in the formation of invasive populations of H. axyridis.     

Involvement of reactive oxygen species and Ca<Superscript>2+</Superscript> in the differential responses to low-boron in rapeseed genotypes

Background and aims

Invasive weeds may exert negative impact on other plant species and soil processes. The observed negative impact of an invasive weed species may be driven by allelopathy or nutrient availability.

Methodology

Sorghum halepense is one of the worst invasive weeds in crop fields. We quantified the species richness in the S. halepense-invaded communities and communities not yet invaded by the weed. Sorghum soil and no-sorghum soil were analysed for total phenolics, microbial activity, available nitrogen (N) and organic carbon. Manipulative experiments were carried out to understand the interference potential of S. halepense. Soil was amended with root or shoot leachate of S. halepense, and its impact on plant growth and soil properties was studied.

Results

Out of four S. halepense-sites, lower plant species richness was observed in one site compared to uninvaded sites. S. halepense-invaded soil had higher levels of total phenolics and lower levels of available N. Higher inhibition in the root growth of Brassica juncea or Bidens pilosa was observed in root leachate-amended soil than shoot leachate-amended soil. Shoot leachate-amended soil had higher levels of total phenolics and available N than root leachate-amended soils. Significant reduction in the available N was observed in soil amended with root leachate. Significant decline in the total phenolics over a period of time was observed in soil amended with root leachate or shoot leachate of S. halepense. Higher CO₂ release was observed 24 h after amending soil with root leachate or shoot leachate of S. halepense.

Conclusions

Sorghum halepense interference potential in its soil is likely due to lower levels of available N. Greater reduction in root dry weight of assay species in root leachate amended soil compared to shoot leachate amended soil was likely due to lower levels of available N in root leachate-amended soil. Relative interference potential of both root and shoot leachates or extracts should be evaluated in allelopathy bioassays and further experiments should be designed to distinguish the role of allelochemicals and nutrient availability in plant growth inhibition.

     

Nitrogen addition drives convergence of leaf litter decomposition rates between <Emphasis Type="Italic">Flaveria bidentis</Emphasis> and native plant

Evidence is growing that invasive species can change decomposition rates and associated nutrient cycling within an ecosystem by changing the quality of the litter entering a system. However, the relative contribution of their distinct litter types to carbon turnover is less understood, especially in the context of enhanced N deposition. The objective of this study was to investigate the whole-plant responses of an invasive plant Flaveria bidentis in litter decay to simulated N eutrophication. A 1-year study was conducted to assess if N enhancement influenced decomposition and nutrient dynamics of litters from foliage, fine roots and twigs of F. bidentis compared to co-occurring native species Setaria viridis. N fertilization significantly decreased the decomposition rate of the foliage of the invasive F. bidentis by more than 25% relative to the water control, but had relatively minor effects on decomposition of its twigs and fine root litter or leaf litter from the native species. Collectively, decomposition rates of foliar litters of the invasive and native species become convergent over time in the presence of N addition. Moreover, net N loss was predominately influenced by litter species, followed by the litter type, while N addition had little effect on net N loss. Our study showed that the variation in litter decomposition was much greater between litter types of the invasive F. bidentis than between different plant species under the N addition and that the litter of invasive species with higher inherent decomposability did not always decompose more rapidly than the litter of native species in response to predicted N deposition enhancement.     

Differences in interactions of aboveground and belowground herbivores on the invasive plant <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis> and native host <Emphasis Type="Italic">A. sessilis</Emphasis>

Plant invasions may result in novel plant-herbivore interactions. However, we know little about whether and how invasive plants can mediate native above- and belowground herbivore interactions. In this study, we conducted greenhouse experiments to examine the interaction between a native defoliating beetle, Cassida piperata, and a native root-knot nematode, Meloidogyne incognita, on the invasive alligator weed, Alternanthera philoxeroides. We also included their native host A. sessilis in the experiments to examine whether the patterns of above- and belowground herbivore interaction vary with host plants (invasive vs. native). We analyzed total carbon and nitrogen in leaves and roots attacked by M. incognita and C. piperata. M. incognita slightly negatively affected feeding by C. piperata on A. philoxeroides, and the leaf area damaged decreased as the number of M. incognita increased. M. incognita had a negative impact on total leaf nitrogen, but had no impact on total leaf carbon. M. incognita egg production on A. philoxeroides roots decreased as the amount of damage caused by C. piperata increased. Herbivory by C. piperata did not affect total root carbon or nitrogen. M. incognita and C. piperata did not affect each other on the native plant A. sessilis. These results suggest that invasive plants can mediate native above- and belowground herbivore interactions. The knowledge of how invasive plants affect those interactions is crucial for better understanding the impacts of biological invasions on native above- and belowground organisms.     

<Emphasis Type="Italic">Piriformospora indica</Emphasis>-mediated salinity tolerance in <Emphasis Type="Italic">Aloe vera</Emphasis> plantlets

Piriformospora indica, a root endophytic fungus, has been reported to promote growth of many plants under normal condition and allow the plants to survive under stress conditions. However, its impact on an important medicinal plant Aloe vera L. has not been well studied. Therefore, this study was undertaken to investigate the effect of P. indica on salinity stress tolerance of A. vera plant. P. indica inoculated and non-inoculated A. vera plantlets were subjected to four levels of salinity treatment- 0, 100, 200 and 300 mM NaCl. The salinity stress decreased the ability of the fungus to colonize roots of A. vera but the interaction of A. vera with P. indica resulted in an overall increase in plant biomass and greater shoot and root length as well as number of shoots and roots. The photosynthetic pigment (Chl a, Chl b and total Chl) and gel content were significantly higher for the fungus inoculated A. vera plantlets, at respective salinity concentrations. Furthermore, the inoculated plantlets had higher phenol, flavonoid, flavonol, aloin contents and radical scavenging activity at all salinity concentrations. The higher phenolic and flavonoid content may help the plants ameliorate oxidative stress resulting from high salinity.     

<Emphasis Type="Italic">Sedum</Emphasis> root foraging in layered green roof substrates

Background and aims

Layered profiles of designed soils may provide long-term benefits for green roofs, provided the vegetation can exploit resources in the different layers. We aimed to quantify Sedum root foraging for water and nutrients in designed soils of different texture and layering.

Methods

In a controlled pot experiment we quantified the root foraging ability of the species Sedum album (L.) and S. rupestre (L.) in response to substrate structure (fine, coarse, layered or mixed), vertical fertiliser placement (top or bottom half of pot) and watering (5, 10 or 20 mm week^?1).

Results

Water availability was the main driver of plant growth, followed by substrate structure, while fertiliser placement only had marginal effects on plant growth. Root foraging ability was low to moderate, as also reflected in the low proportion of biomass allocated to roots (5–13%). Increased watering reduced the proportion of root length and root biomass in deeper layers.

Conclusions

Both S. album and S. rupestre had a low ability to exploit water and nutrients by precise root foraging in substrates of different texture and layering. Allocation of biomass to roots was low and showed limited flexibility even under water-deficient conditions.

     

An invasive plant provides refuge to native plant species in an intensely grazed ecosystem

Invasion by exotic plant species and herbivory can individually alter native plant species diversity, but their interactive effects in structuring native plant communities remain little studied. Many exotic plant species escape from their co-evolved specialized herbivores in their native range (in accordance with the enemy release hypothesis). When these invasive plants are relatively unpalatable, they may act as nurse plants by reducing herbivore damage on co-occurring native plants, thereby structuring native plant communities. However, the potential for unpalatable invasive plants to structure native plant communities has been little investigated. Here, we tested whether presence of an unpalatable exotic invader Opuntia ficus-indica was associated with the structure of native plant communities in an ecosystem with a long history of grazing by ungulate herbivores. Along 17 transects (each 1000 m long), we conducted a native vegetation survey in paired invaded and uninvaded plots. Plots that harboured O. ficus-indica had higher native plant species richness and Shannon–Wiener diversity H′ than uninvaded plots. However, mean species evenness J was similar between invaded and uninvaded plots. There was no significant correlation between native plant diversity and percentage plot cover by O. ficus-indica. Presence of O. ficus-indica was associated with a compositional change in native community assemblages between paired invaded and uninvaded plots. Although these results are only correlative, they suggest that unpalatable exotic plants may play an important ecological role as refugia for maintenance of native plant diversity in intensely grazed ecosystems.