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Earthworms Enhance Crop Resistance to Insects Under Microplastic Stress by Mobilizing Physical and Chemical Defenses
Abstract
To assess the ecological risk of microplastics (MPs) in agricultural systems, it is critical to simultaneously focus on MP-mediated single-organism response and different trophic-level organism interaction. Herein, we placed earthworms in soils contaminated with different concentrations (0.02% and 0.2% w/w) of polyethylene (PE) and polypropylene (PP) MPs to investigate the effect of earthworms on tomato against Helicoverpa armigera (H. armigera) under MPs stress. We found that earthworms alleviated the inhibitory effects of MPs stress on tomato growth and disrupted H. armigera growth. Compared to individual MPs exposure, earthworm incorporation significantly increased the silicon and lignin content in herbivore-damaged tomato leaves by 19.1% and 57.6%, respectively. Metabolites involved in chemical defense (chlorogenic acid) and phytohormones (jasmonic acid) were also activated by earthworm incorporation. Furthermore, earthworms effectively reduced oxidative damage induced by H. armigera via promoting antioxidant metabolism. Overall, our results suggest that utilizing earthworms to regulate above-and below-ground interactions could be a promising strategy for promoting green agriculture.
Microplastics (MPs) contamination presents a significant global environmental challenge, with its potential to influence soil carbon (C) dynamics being a crucial aspect for understanding soil C changes and global C cycling. This meta-analysis synthesizes data from 110 peer-reviewed publications to elucidate the directional, magnitude, and driving effects of MPs exposure on soil C dynamics globally. We evaluated the impacts of MPs characteristics (including type, biodegradability, size, and concentration), soil properties (initial pH and soil organic C [SOC]), and experimental conditions (such as duration and plant presence) on various soil C components. Key findings included the significant promotion of SOC, dissolved organic C, microbial biomass C, and root biomass following MPs addition to soils, while the net photosynthetic rate was reduced. No significant effects were observed on soil respiration and shoot biomass. The study highlights that the MPs concentration, along with other MPs properties and soil attributes, critically influences soil C responses. Our results demonstrate that both the nature of MPs and the soil environment interact to shape the effects on soil C cycling, providing comprehensive insights and guiding strategies for mitigating the environmental impact of MPs.
In terrestrial ecosystems, plant leaves provide the largest biological habitat for highly diverse microbial communities, known as the phyllosphere microbiota. However, the underlying mechanisms of host-driven assembly of these ubiquitous communities remain largely elusive. Here, we conduct a large-scale and in-depth assessment of the rice phyllosphere microbiome aimed at identifying specific host-microbe links. A genome-wide association study reveals a strong association between the plant genotype and members of four bacterial orders, Pseudomonadales, Burkholderiales, Enterobacterales and Xanthomonadales. Some of the associations are specific to a distinct host genomic locus, pathway or even gene. The compound 4-hydroxycinnamic acid (4-HCA) is identified as the main driver for enrichment of bacteria belonging to Pseudomonadales. 4-HCA can be synthesized by the host plant’s OsPAL02 from the phenylpropanoid biosynthesis pathway. A knockout mutant of OsPAL02 results in reduced Pseudomonadales abundance, dysbiosis of the phyllosphere microbiota and consequently higher susceptibility of rice plants to disease. Our study provides a direct link between a specific plant metabolite and rice phyllosphere homeostasis opening possibilities for new breeding strategies.
Strigolactones (SLs) are carotenoid‐derived phytohormones that regulate plant growth and development. While root‐secreted SLs are well‐known to facilitate plant symbiosis with beneficial microbes, the role of SLs in plant interactions with pathogenic microbes remains largely unexplored.
Using genetic and biochemical approaches, we demonstrate a negative role of SLs in rice (Oryza sativa) defense against the blast fungus Pyricularia oryzae (syn. Magnaporthe oryzae).
We found that SL biosynthesis and perception mutants, and wild‐type (WT) plants after chemical inhibition of SLs, were less susceptible to P. oryzae. Strigolactone deficiency also resulted in a higher accumulation of jasmonates, soluble sugars and flavonoid phytoalexins in rice leaves. Likewise, in response to P. oryzae infection, SL signaling was downregulated, while jasmonate and sugar content increased markedly. The jar1 mutant unable to synthesize jasmonoyl‐l‐isoleucine, and the coi1‐18 RNAi line perturbed in jasmonate signaling, both accumulated lower levels of sugars. However, when WT seedlings were sprayed with glucose or sucrose, jasmonate accumulation increased, suggesting a reciprocal positive interplay between jasmonates and sugars. Finally, we showed that functional jasmonate signaling is necessary for SL deficiency to induce rice defense against P. oryzae.
We conclude that a reduction in rice SL content reduces P. oryzae susceptibility by activating jasmonate and sugar signaling pathways, and flavonoid phytoalexin accumulation.
Crop breeding for mechanized harvesting has driven modern agriculture. In tomato, machine harvesting for industrial processing varieties became the norm in the 1970s. However, fresh-market varieties whose fruits are suitable for mechanical harvesting are difficult to breed because of associated reduction in flavour and nutritional qualities. Here we report the cloning and functional characterization of fs8.1, which controls the elongated fruit shape and crush resistance of machine-harvestable processing tomatoes. FS8.1 encodes a non-canonical GT-2 factor that activates the expression of cell-cycle inhibitor genes through the formation of a transcriptional module with the canonical GT-2 factor SlGT-16. The fs8.1 mutation results in a lower inhibitory effect on the cell proliferation of the ovary wall, leading to elongated fruits with enhanced compression resistance. Our study provides a potential route for introducing the beneficial allele into fresh-market tomatoes without reducing quality, thereby facilitating mechanical harvesting.
Plants face attackers aboveground and belowground. Insect root herbivores can lead to severe crop losses, yet the underlying transcriptomic responses have rarely been studied.
We studied the dynamics of the transcriptomic response of Brussels sprouts (Brassica oleracea var. gemmifera) primary roots to feeding damage by cabbage root fly larvae (Delia radicum), alone or in combination with aboveground herbivory by cabbage aphids (Brevicoryne brassicae) or diamondback moth caterpillars (Plutella xylostella). This was supplemented with analyses of phytohormones and the main classes of secondary metabolites; aromatic, indole and aliphatic glucosinolates.
Root herbivory leads to major transcriptomic rearrangement that is modulated by aboveground feeding caterpillars, but not aphids, through priming soon after root feeding starts. The root herbivore downregulates aliphatic glucosinolates. Knocking out aliphatic glucosinolate biosynthesis with CRISPR‐Cas9 results in enhanced performance of the specialist root herbivore, indicating that the herbivore downregulates an effective defence.
This study advances our understanding of how plants cope with root herbivory and highlights several novel aspects of insect–plant interactions for future research. Further, our findings may help breeders develop a sustainable solution to a devastating root pest.
Plants have evolved plastic defence strategies to deal with the uncertainty of when, by which species and in which order attack by herbivores will take place1–3. However, the responses to current herbivore attack may come with a cost of compromising resistance to other, later arriving herbivores. Due to antagonistic cross-talk between physiological regulation of plant resistance to phloem-feeding and leaf-chewing herbivores4–8, the feeding guild of the initial herbivore is considered to be the primary factor determining whether resistance to subsequent attack is compromised. We show that, by investigating 90 pairwise insect–herbivore interactions among ten different herbivore species, resistance of the annual plant Brassica nigra to a later arriving herbivore species is not explained by feeding guild of the initial attacker. Instead, the prevalence of herbivore species that arrive on induced plants as approximated by three years of season-long insect community assessments in the field explained cross-resistance. Plants maintained resistance to prevalent herbivores in common patterns of herbivore arrival and compromises in resistance especially occurred for rare patterns of herbivore attack. We conclude that plants tailor induced defence strategies to deal with common patterns of sequential herbivore attack and anticipate arrival of the most prevalent herbivores. Plants have developed defences against herbivores, but with multiple threats at different times, those defences may be compromised. This experiment finds that plants adapt over time to anticipate prevalent herbivores and common patterns of attack.
Plastic pollution accumulating in an area of the environment is considered “poorly reversible” if natural mineralization processes occurring there are slow and engineered remediation solutions are improbable. Should negative outcomes in these areas arise as a consequence of plastic pollution, they will be practically irreversible. Potential impacts from poorly reversible plastic pollution include changes to carbon and nutrient cycles; habitat changes within soils, sediments, and aquatic ecosystems; co-occurring biological impacts on endangered or keystone species; ecotoxicity; and related societal impacts. The rational response to the global threat posed by accumulating and poorly reversible plastic pollution is to rapidly reduce plastic emissions through reductions in consumption of virgin plastic materials, along with internationally coordinated strategies for waste management.
Lignin is a complex aromatic biopolymer that strengthens and waterproofs plant secondary cell walls, enabling mechanical stability in trees and long-distance water transport in xylem. Lignin removal is a key step in paper production and biomass conversion to biofuels, motivating efforts to re-engineer lignin biosynthesis. However, the physical nature of lignin’s interactions with wall polysaccharides is not well understood. Here we show that lignin self-aggregates to form highly hydrophobic and dynamically unique nanodomains, with extensive surface contacts to xylan. Solid-state NMR spectroscopy of intact maize stems, supported by dynamic nuclear polarization, reveals that lignin has abundant electrostatic interactions with the polar motifs of xylan. Lignin preferentially binds xylans with 3-fold or distorted 2-fold helical screw conformations, indicative of xylans not closely associated with cellulose. These findings advance our knowledge of the molecular-level organization of lignocellulosic biomass, providing the structural foundation for optimization of post-harvest processing for biofuels and biomaterials.
Worldwide, microplastics (MPs) have been commonly recognized as a threat to soil ecosystems. Terrestrial soils are widely contaminated by MPs due to the application of sewage sludge and wastewater, plastic mulching, littering, the input of tire wear from roads and atmospheric deposition. Within the last decade, an increasing number of individual studies focused on item counts and masses of MPs in different global soil environments.
We reviewed these studies to achieve a representative picture of common degrees of contamination. The majority of the prospected agricultural and horticultural sites were exposed to sewage sludge and mulching film application and showed concentrations of <13000 items kg-1 dry soil and 4.5 mgkg-1 dry soil. Microplastic concentrations in soils in the vicinity of municipal areas were thereby 10 times larger compared to rural sites. The measurement of masses was generally underrepresented compared to item numbers, and mass data were often generated from microscopic analyses by using shape-to-mass models instead of direct measurements. Extreme values, such as on industrial sites, exceed the common concentrations by 2 to 4 orders of magnitude, which might be attributed not only to the land use but also to the applied methods of measurement. Campaigns that focus on other entry pathways like composts, road dust runoff and littering or land uses like grassland, forest, fallow and wilderness as well as industrial sites and landfills were underrepresented or nonexistent. Background loads, such as atmospheric deposition, were often not excluded from the measurements and, thus, the studies might overestimate the contribution of the analyzed entry pathway to the total load. Other studies focused on light density MP, e.g., from mulching films, and therefore underestimated the amount of soil MP.
Despite these limitations, the data give an impression of the spectrum of global MP concentrations and are a good basis for experiments examining the effects of MPs on exposed soils. Based on the collected data, we identified problems of past studies and recommend that future experimentation take into account standardized methods of extraction and quantification, a proper characterization of the sampling sites and their history as well as the exploration of as yet underrepresented entry pathways and land uses.
In plants, pathogen attack can induce an immune response known as systemic acquired resistance that protects against a broad spectrum of pathogens. In the search for safer agrochemicals, silica nanoparticles (SiO2 NPs; food additive E551) have recently been proposed as a new tool. However, initial results are controversial, and the molecular mechanisms of SiO2 NP-induced disease resistance are unknown. Here we show that SiO2 NPs, as well as soluble Si(OH)4, can induce systemic acquired resistance in a dose-dependent manner, which involves the defence hormone salicylic acid. Nanoparticle uptake and action occurred exclusively through the stomata (leaf pores facilitating gas exchange) and involved extracellular adsorption in the air spaces in the spongy mesophyll of the leaf. In contrast to the treatment with SiO2 NPs, the induction of systemic acquired resistance by Si(OH)4 was problematic since high Si(OH)4 concentrations caused stress. We conclude that SiO2 NPs have the potential to serve as an inexpensive, highly efficient, safe and sustainable alternative for plant disease protection.
Trichomes are specialized epidermal cells that contribute to plant resistance against herbivores. Their formation is controlled precisely by multiple genetic and environmental signals. Previous studies have shown that microRNA319 (miR319) and gibberellin (GA) signaling are involved in trichome development in Arabidopsis, but little is known about their interaction between these factors.
Here we reported that the miR319a/TEOSINTE BRANCHED/CYCLOIDEA/PCF (TCP) module participates in trichome initiation synergistically with GA signaling in Populus tomentosa. We demonstrated that overexpression of miR319a decreased transcription levels of its targeted TCPs and significantly elevated leaf trichome density in transgenic poplar, resulting in decreasing insect herbivory. Conversely, repressing miR319a by short tandem target mimics (STTM) elevated TCP expression levels and decreased trichome density in transgenic plants.
The trichome phenotype of 35S:miR319a plants could be abolished by introducing a miR319a‐resistant form of TCP19. Furthermore, the miR319a‐targeted TCP19 interacted directly with REPRESSOR OF ga1‐3 (RGA), a downstream repressor of GA signaling. TCP19 and RGA synergistically inhibited the GLABROUS1 (GL1)‐induced expression of trichome marker gene GLABRA2 (GL2), thereby repressing leaf trichome initiation.
Our results provide an insight into the molecular mechanism by which miR319/TCP19 module and GA signaling coordinated regulating trichome initiation in P. tomentosa.
Despite the increasing prevalence of atmospheric nanoplastics (NPs), there remains limited research on their phytotoxicity, foliar absorption, and translocation in plants. In this study, we aimed to fill this knowledge gap by investigating the physiological effects of tomato leaves exposed to differently charged NPs and foliar absorption and translocation of NPs. We found that positively charged NPs caused more pronounced physiological effects, including growth inhibition, increased antioxidant enzyme activity, and altered gene expression and metabolite composition and even significantly changed the structure and composition of the phyllosphere microbial community. Also, differently charged NPs exhibited differential foliar absorption and translocation, with the positively charged NPs penetrating more into the leaves and dispersing uniformly within the mesophyll cells. Additionally, NPs absorbed by the leaves were able to translocate to the roots. These findings provide important insights into the interactions between atmospheric NPs and crop plants and demonstrate that NPs' accumulation in crops could negatively impact agricultural production and food safety.
Microplastic pollution can have detrimental effects on soil environments and inhibit crop growth. Earthworms, known as soil engineers, promote crop growth, but their role and impact on the amelioration of microplastic-polluted soil is not yet clear. In this study, we investigated the impact and pathways of earthworm activity on microplastic-contaminated soil by introducing varying densities (without earthworm:0, low-density: 1, medium-density: 2, high-density: 5 ind column-1) of earthworms (epi-endogeic) into soil contaminated with two types of microplastics: polyethylene and polyvinyl chloride. Our results showed that earthworms all survived in soil polluted with two types of microplastics. Meanwhile, earthworm activity increased nutrient content and enzyme activity by 0.2-36.1% and 2.9-34.3%, respectively, and significantly increased soil microbial biomass and community diversity index. Earthworm activity also decreased antioxidant enzyme activity and promoted maize plant growth, including agronomic traits such as plant height, biomass, root length, and root surface area. Furthermore, the nutrient content of maize organs increased by 1.1-29.7%. Partial least squares models confirmed that earthworm activity alleviated the stress effect of microplastic pollution on plant growth by improving soil structure, fertility, and microbial abundance and diversity. The greatest effect on maize growth was observed with the improvement of soil physical-chemical properties. Our results suggest that medium densities of earthworms have the greatest soil improvement effect and provide an important basis for bioremediation of farmland contaminated by microplastics and promoting green and efficient development in agriculture.
Soil microplastic pollution can have negative effects on organisms, including plants, but the underlying mechanisms are not fully understood. We tested whether structural or chemical properties of a microplastic cause its effects on plant above- and belowground growth and whether these effects can be influenced by earthworms. We conducted a factorial experiment in a greenhouse with seven common Central European grassland species. Microplastic granules of the synthetic rubber ethylene propylene diene monomer (EPDM),1 a frequently used infill material of artificial turfs, and cork granules with a comparable size and shape to the EPDM granules were used to test for structural effects of granules in general. To test for chemical effects, EPDM-infused fertilizer was used, which should have contained any leached water-soluble chemical components of EPDM. Two Lumbricus terrestris individuals were added to half of the pots, to test whether these earthworms modify effects of EPDM on plant growth. EPDM granules had a clear negative effect on plant growth, but since cork granules had a negative effect of similar magnitude, with an average decrease in biomass of 37 % in presence of granules, this is likely due to the structural properties of granules (i.e., size and shape). For some belowground plant traits, EPDM had a stronger effect than cork, which shows that there must be other factors playing into the effects of EPDM on plant growth. The EPDM-infused fertilizer did not have any significant effect on plant growth by itself, but it had in interaction with other treatments. Earthworms had an overall positive effect on plant growth and mitigated most of the negative effects of EPDM. Our study shows that EPDM microplastic can have negative effects on plant growth, and that these might be more related to its structural than to its chemical properties.
Tetracycline pollution in soil irreversibly damages the biosafety of plants by inhibiting the mitochondrial function. Some traditional Chinese medicine (TCM) plants, such as Salvia miltiorrhiza Bunge, have a strong tolerance to mitochondrial damage. We comprehensively compared the doxycycline (DOX) tolerances of two ecotypes of S. miltiorrhiza in the Sichuan and Shandong provinces and found that the Sichuan ecotype had a lower yield reduction, more stable accumulation of medicinal ingredients, higher mitochondrial integrity, and a more robust antioxidant system. The synergetic response networks under DOX pollution of both ecotypes were constructed using RNA sequencing and ultrahigh-performance liquid chromatography-tandem mass spectrometry. The differentiation of the downstream pathways of aromatic amino acids (AAAs) produced variations in the DOX tolerance of S. miltiorrhiza in different regions. The Sichuan ecotype maintained redox homeostasis and xylem development by activating salvianolic acid and indole biosynthesis, while the Shandong ecotype balanced chemical and mechanical defenses by regulating the flavonoid biosynthesis. Rosmarinic acid, a downstream AAA molecule, maintains the mitochondrial homeostasis of plant seedlings under DOX pollution by targeting the ABCG28 transporter. We also highlight the significance of downstream AAA small molecules in guiding the development of bio-based environmental pollution remediation agents.
In recent years, the prevalence and accumulation of microplastics (MPs) in the soil environment have raised worldwide concerns for their resistance to degradation and long-term ecological impacts. However, compared with aquatic ecosystems, there is still a lack of critical reviews on the ecological effects and effective remediation methods of MPs in soil. In this review, the distribution characteristics, sources and transportation, identification techniques of MPs are summarized, followed by an elaboration of the interactions between MPs contamination and soil ecosystems. Moreover, we generalized the effects and degradation mechanisms of different organisms on the degradation of MPs. There are a variety of microorganisms in the environment that can degrade MPs, but the enzymes that really work are yet to be explored. It may be a breakthrough in the field of biodegradation to transform the stability and performance of enzymes based on computer protein design. This review will guide researchers to further explore feasible approaches and formulate advanced remediation strategies for MPs contamination in the future.
Nanosilicon applications have been shown to increase plant defenses against both abiotic and biotic stresses. Silicon quantum nanodots (Si NDs), a form of nanosilicon, possess excellent biological and physiochemical properties (e.g., minimal size, high water solubility, stability, and biocompatibility), potentially making them more efficient in regulating plant responses to stress than other forms of silicon. However, to date, we still lack mechanistic evidence for how soil-applied Si NDs alter the regulation of plant physical and chemical defenses against insect herbivores. To address this gap, we compared the effect of fluorescent amine-functionalized Si NDs (5 nm) and the conventional fertilizer sodium silicate on maize (Zea mays L.) physical and chemical defenses against the oriental armyworm (Mythimna separata, Walker) caterpillars. We found that 50 mg/kg Si NDs and sodium silicate additions inhibited the growth of caterpillars the most (35.7% and 22.8%, respectively) as compared to other application doses (0, 10, and 150 mg/kg). Both Si NDs and silicate addition activated biosynthesis genes responsible for chemical (benzoxazinoids) and physical (lignin) defense production. Moreover, Si NDs upregulated the gene expression of antioxidant enzymes (SOD, CAT, and POD) and promoted the antioxidant metabolism (flavonoids) in maize leaves under M. separata attack. Finally, we show that, under field conditions, Si ND addition increased maize cob weight (28.7%), cob grain weight (40.8%), and 100-grain weight (26.5%) as compared to the control, and more so than the conventional silicon fertilizer. Altogether, our findings highlight the potential for Si NDs to be used as an effective and ecofriendly crop protection strategy in agroecosystems.
Agro-ecosystem contamination with microplastics (MPs) is of great concern. However, limited research has been conducted on the agricultural soil of tropical regions. This paper investigated MPs in the agro-ecosystem of Hainan Island, China, as well as their relationships with plastic mulching, farming practices, and social and environmental factors. The concentration of MPs in the study area ranged from 2800 to 82500 particles/kg with a mean concentration of 15461.52 particles/kg. MPs with sizes between 20 and 200 μm had the highest abundance of 57.57%, fragment (58.16%) was the most predominant shape, while black (77.76%) was the most abundant MP colour. Polyethylene (PE) (71.04%) and polypropylene (PP) (19.83%) were the main types of polymers. The mean abundance of MPs was significantly positively correlated (p < 0.01) with all sizes, temperature, and shapes except fibre, while weakly positively correlated with the population (p = 0.21), GDP (p =0.33), and annual precipitation (p = 0.66). In conclusion, plastic mulching contributed to significant contamination of soil MPs in the study area, while environmental and social factors promoted soil MPs fragmentation.
The current study results indicate serious contamination with MPs, which poses a concern regarding ecological and environmental safety.
To promote the sustainable and safe application of nanotechnology employing engineered nanoparticles (NPs) in agroecosystems, it is crucial to pay more attention to the NP-mediated biological response process and environmental impact assessment simultaneously. Herein, 50 mg kg-1 La2O3 NPs were added to soils without and with earthworms for cherry radish growth for 50 days to investigate the response changes of metabolites in radish above- and below-ground organs and rhizosphere bacterial communities. We found that La2O3 NP exposure, especially with earthworms, notably increased the La bioavailability and uptake by taproots and eventually increased radish leaf sucrose content and plant biomass. The La2O3 NP exposure significantly altered metabolite profiles in taproot flesh and peel tissues, and particularly La2O3 NP exposure combined with earthworms was more conducive to La2O3 NPs to promote radish taproot peel to synthesize more secondary antioxidant metabolites. Moreover, compared with the control, the La2O3 NP exposure resulted in weaker and fewer correlations between rhizosphere bacteria and taproot metabolites, but this was recovered somewhat after the inoculation of earthworms. Altogether, our results provide novel insights into the soil-fauna-driven biological and biochemical impact of La2O3 NP exposure on edible root crops and the long-term environmental risks to the rhizosphere microbiota in agroecosystems.
Plants have evolved a series of defensive mechanisms against pathogens and herbivores, but the defense response always leads to decreases in growth or reproduction, which has serious implications for agricultural production. Growth and defense are negatively regulated not only through metabolic consumption but also through the antagonism of different phytohormones, such as jasmonic acid (JA) and salicylic acid (SA). Meanwhile, plants can limit the expression of defensive metabolites to reduce the costs of defense by producing constitutive defenses such as glandular trichomes or latex and accumulating specific metabolites, determining the activation of plant defense or the maintenance of plant growth. Interestingly, plant defense pathways might be prepared in advance which may be transmitted to descendants. Plants can also use external organisms to protect themselves, thus minimizing the costs of defense. In addition, plant relatives exhibit cooperation to deal with pathogens and herbivores, which is also a way to regulate growth and defense.
Microplastics (MPs), especially biodegradable MPs (BMPs) have attracted increasing attention in recent years. However, the effects of MPs with different biodegradability on the soil-plant systems are not well explored. In this study, the effects of polyethylene MPs (PEMPs) and polylactic acid MPs (PLAMPs) on physio-biochemical performance and metabolomic profile of soybean (Glycine max), as well as the bacterial communities in soil were investigated. Our results showed that PEMPs had no noticeable toxicity on the plant growth, while 0.1% PLAMPs significantly decreased the root length by 27.53% when compared with the control. The peroxidase (POD) activity was reduced and catalase (CAT) activity was increased by MPs in plant leaves. The metabolomics study suggested that the significantly affected metabolic pathway is amino acid metabolism. Additionally, Shannon and Simpson indices of rhizosphere soil were changed only under 0.1% PLAMPs. The key bacteria involved in the dinitrogen fixation were also altered. This study provides a novel insight into the potential effects of MPs with different biodegradability on soil-plant systems and highlights that BMPs might have stronger negative effects for terrestrial ecosystem, which needs to be further explored in future research.
Extensive use of plastic film mulch has resulted in accumulation of a large amount of residual plastic, which will eventually fragment into microplastics in agricultural soils. However, it is unclear how microplastics from plastic mulch film affect crops. To address this issue, rice plants exposed to microplastics derived from poly(butyleneadipate-co-terephthalate) (PBAT)-based biodegradable mulch film (BM) and polyethylene (PE) mulch film (PM) were investigated for plant growth, physio-biochemical processes, and gene expressions. Both types of microplastics significantly reduced the height and dry weight of rice plant. Oxidative stress was induced by microplastics in rice shoot and root, with levels of ROS relatively higher under treatment PM than that under treatment BM. Transcriptomic data showed that more genes were down-regulated by treatment PM than that by treatment BM. Genes encoding ammonium and nitrate transporters were down-regulated by both types of microplastics in rice roots at vegetative stage, whereas up-regulated at reproductive stage, as compared to their respective treatment with no microplastics (CK). Similar results regarding phenylpropanoid biosynthesis pathway and lignin content were also observed in rice roots. Net photosynthetic rate and SPAD value were significantly inhibited by treatments BM and PM in rice shoot, and the expression of genes involved in light reaction was reduced at vegetative stage, whereas there were no differences of them at reproductive stage, as compared to their respective treatment CK. Our study suggests that microplastics from BM and PM both affect the growth of rice plants via nitrogen metabolism and photosynthesis. The negative effects imposed by both types of microplastics on rice plant can be mitigated with the growth of plants, and the negative effects of microplastics from PE mulch film on rice plant are relatively stronger than that from the PBAT-based biodegradable film.
Silicon (Si) acquisition by plants are stimulated by root-associated rhizosphere processes which are usually mediated by nanomaterials and soil fauna. In this study, 10 mg kg-1 silica nanomaterials (SiO2 NMs)...
Adaptive responses of plants are important not only for local processes in populations and communities but also for global processes in the biosphere through the primary production of ecosystems. In recent years, the concept of environmental hormesis has been increasingly used to explain the adaptive responses of living organisms, including plants, to low doses of natural factors, both abiotic and biotic, as well as various anthropogenic impacts. However, the issues of whether plant hormesis is similar/different when it is induced by mild stressors having different specific effects and what is the contribution of hormetic stimulation of non-specific and specific adaptive mechanisms in plant resilience to strong stressors (i.e., preconditioning) remains unclear. This paper analyses hormetic stimulation of non-specific and specific adaptive mechanisms in plants and its significance for preconditioning, the phenomenon of the hormetic trade-off for these mechanisms, and the position of hormetic stimulation of non-specific and specific adaptive mechanisms in the system of plant adaptations to environmental challenges. The analysis has shown that both non-specific and specific adaptive mechanisms of plants can be stimulated hormetically by mild stressors and are important for plant preconditioning. Due to limited plant resources, non-specific and specific adaptive mechanisms have hormetic trades-offs 1 (hormesis accompanied by the deterioration of some plant traits) and 2 (hormesis of some plant traits with the invariability of others). At the same time, hormetic trade-off 2 is observed much more often than hormetic trade-off 1, at least, this was demonstrated here for non-specific adaptive responses of plants. The hormetic stimulation of non-specific and specific adaptive mechanisms is part of the inducible adaptation of plants caused by stress factors and is an adaptation to random (unpredictable) changes in the environment.
Pollution by microplastics in natural and cultivated soils is an emerging issue of global interest. Microplastics may interfere with the soil structure, microorganism and invertebrate biodiversity, and related biological processes. Surprisingly, there is a poor understanding of how microplastics influence the interactions between members of different trophic levels. We investigated whether a high level of contamination (5% w/w concentration) with high-density polyethylene (HDPE) microplastics (< 0.25 mm size) in soil interferes with the attraction of an insect herbivore towards a plant-soil system under short-term laboratory conditions. As a model system, we used the black fungus gnat Bradysia difformis (Diptera: Sciaridae), which is an important crop pest, and lentil (Lens culinaris), a host plant. Through one-way olfactometer bioassays, we recorded the female responses to plant-soil systems subjected to different combinations of HDPE exposures and watering regimes. Female residence time (i.e., the time spent in the olfactometer sector close to the stimulus) was higher when testing high-watered (75.3% ± 2.99) than low-watered, microplastic-free plant-soil systems (58.6% ± 3.88). However, residence time was lower towards systems maintained 7 days with 5% HDPE (53.9% ± 4.79) compared to HDPE-free (76.5% ± 2.64) systems. Notably, the attraction of females towards HDPE-free systems compared to 5% HDPE was higher only under high watering regime. Attractiveness of 5% HDPE plant-soil system was restored when cues from fungal colonies were added (66.1% ± 4.33). Here we demonstrated for the first time that microplastics alter the attraction of female fungus gnats towards a plant-soil system. Our results depict an extreme scenario with high HDPE (5%), which is occasionally found in soils close to industrial areas. Therefore it is still too early to derive strong conclusions for typical agricultural settings. However, if microplastic pollution increases in agricultural soils, possible interferences in the established soil-plant-herbivore communication mechanisms should be carefully considered.
Microplastics (MPs) are drawing increasing attention from the international community due to their potential threats to the ecosystem and human health. Although their occurrence and spatial distribution have been extensively studied in recent years, the relationship between their abundance and sizes remains unclear. Moreover, the underlying mechanisms dominating their size distribution have rarely been explored. In the present study, we developed a novel conditional fragmentation model to describe MP size distribution in the soil environment. It is proposed that the distribution of MPs is not a coincidence but controlled by conditional aging. The applicability of this model was tested using data collected from different land use settings in Beijing, China. A distinct downsizing phenomenon from fibers, films, and fragments to granules is observed. Undisturbed land use types accumulated larger sized MPs with higher stability, while human interference accelerated the fragmentation of MPs. Both morphological analysis and time-of-flight secondary ion mass spectroscopy (TOF-SIMS) observations provided direct evidence for the conditional fragmentation process. Furthermore, the model has proven to be suitable for describing the size distribution of MPs from various sources (including atmospheric deposition, transportation, and agriculture) and aging processes (such as mechanical abrasion, chemical oxidation, and photochemical transformation). It is proposed that this model can be used for various purposes in MP-related studies, especially source identification, transport modeling, and risk assessment.
Silicon, in its nanoscale form, has shown plant-promoting and insecticidal properties. To date, however, we lack mechanistic evidence for how nanoscale silicon influences the regulation of plant chemical defenses against herbivore attacks. To address this gap, we compared the effect of Si nanodots and sodium silicate, a conventional silicate fertilizer, on maize (Zea mays L.) chemical defenses against the oriental armyworm (Mythimna separata, Walker) caterpillars. We found that Si NDs and sodium silicate additions, at the dose of 50 mg/L, significantly inhibited the growth of caterpillars by 53.5% and 34.2%, respectively. This increased plant resistance was associated with a 44.2% increase in the production of chlorogenic acid, as well as the expression of PAL, C4H, 4CL, C3H and HCT, core genes involved in the biosynthesis of chlorogenic acid, by 1.7, 2.4, 1.9, 1.8 and 4.5 folds, respectively. Particularly, in the presence of M. separata, physiological changes in maize plants treated with 50 mg/L Si NDs, including changes in shoot biomass, leaf nutrients (e.g., K, P, Si), and chemical defense compounds (e.g., chlorogenic acid, total phenolics), were higher than those of plants added with equivalent concentrations of conventional silicate fertilizer. Taken together, our findings indicate that Si, in nanoscale form, could replace synthetic pesticides, and be implemented for a more effective and ecologically-sound management of insect pests in maize crop farming.
Tomato (Solanum lycopersicum L.) is a natural host for the Helicoverpa-Chloridea (Lepidoptera: Noctuidae: Heliothinae) pest complex. The species Helicoverpa armigera (Hübner) was responsible for significant yield losses in several crops after its detection in Brazil. The morphology of its larval stage resembles common Heliothinae species, making pest control decisions difficult. The overall lack of studies on the Heliothinae associated with tomatoes in Brazil and the establishment of H. armigera in the country plus their recent outbreaks supported our investigation about the relative importance of the insects from the Helicoverpa-Chloridea complex in this vegetable crop. A nationwide survey was carried out across fresh-market and processing tomato fields. Molecular analyses targeting a segment of the mitochondrial cytochrome oxidase subunit I (mtCOI) gene and their sequence analyses indicated the presence of a pest complex, comprising the introduced species H. armigera and the indigenous species, Helicoverpa zea (Boddie), and Chloridea virescens (Fabricius). The Harm 1 haplotype of H. armigera was identified as the predominant Heliothinae pest infesting fresh-market tomatoes. The New World species Chloridea subflexa (Guenée) as well as the exotic Solanaceae-specific species Helicoverpa assulta (Guenée) were not found in our survey. Additional larvae surveys in processing tomato fields during 2013/2014 in Central Brazil also indicated H. armigera as the most abundant Heliothinae species (95%) together with H. zea (4.75%) and C. virescens (0.25%). The occurrence of distinct Helicoverpa species (which are potentially capable of interbreeding) indicates that novel crop management strategies will be necessary in order to minimize damages caused by this pest complex in tomatoes.
Silicon improves plant resistance to a wide range of biotic and abiotic stresses. However, available silicon as a
plant nutrient in soil is scarce. A continuous supply of available silicon is vital for plants. Earthworms are
important detritivores in many terrestrial ecosystems. Therefore, we hypothesized that earthworms harbor silicate
solubilizing bacteria in their guts and play a key role in silicon weathering and dynamics. Silicate solubilizing
bacteria were isolated from the gut of geophagous endogeic earthworm (Pheretima guillelmi) and their
effects on silicon weathering and availability to maize (Zea mays) were examined. Significantly more silicate
solubilizing bacteria were observed in the earthworms gut than those in the surrounding soil in all sampling
locations. The 16S rRNA analysis showed a higher silicate solubilizing bacteria diversity with 20 bacterial strains
isolated from the gut walls, belonging to the genera of Aeromonas, Bacillus, Cellvibrio, Ensifer, Flavobacterium,
Microbacterium, Paracoccus, Pseudomonas, Rhizobium, and Streptomyces. Of these bacteria, only three species were
found in the surrounding soil. Inoculation with the three fastest-growing strains in liquid culture increased
soluble silicon released from the feldspar and quartz powder in the media. Inoculation with the 3C1 strain in the
potting soil significantly increased soluble silicon content in the soil, enhanced silicon uptake and accumulation
in maize plants and promoted seedling growth. Taken together, these results not only highlight the importance
of earthworm gut microbes in soil silicon weathering but also indicate a new approach in searching for silicate
solubilizing bacteria to increase soil silicon availability in agriculture.
Tea trichomes contain special flavor-determining metabolites, however, little is known about how and why tea trichomes produce them. Integrated metabolite and transcriptome profiling on tea trichomes in comparison with leaves showed that trichomes contribute to tea plant defense and tea flavor and nutritional quality. These unicellular, non-glandular, and unbranched tea trichomes produce a wide array of tea characteristic metabolites, such as UV-protective flavonoids, insect-toxic caffeine, herbivore-defensive volatiles, and theanine, as evidenced by expression of whole sets of genes involved in different metabolic pathways. Both dry and fresh trichomes contain several volatiles and flavonols that were not found or at much low levels in trichome-removed leaves, including benzoic acid derivatives, lipid oxidation derivatives, and monoterpene derivatives. Trichomes also specifically expressed many diseases signaling genes, and various anti-herbivore or anti-abiotic peptides. Trichomes are one of the domestication traits in tea plants. Tea trichomes contribute to tea plant defenses and tea flavors.
Potential adverse effects of nanoplastics (NPs) on marine organisms have received increased attention in recent years. In contrast, few data are available on terrestrial plants, especially on the mechanisms for transport of NPs in plants and phytotoxicity (at both phenotypic and molecular levels) of plants induced by NPs. To address this knowledge gap, we conducted a microcosm study in which hydroponically-cultured rice (Oryza sativa L.) seedlings were exposed to polystyrene (PS)-NPs at 0, 10, 50, and 100 mg L⁻¹ for 16 d and examined for morphological and physiological phenotypes and transcriptomics. Laser confocal scanning micrographs confirmed PS-NPs were uptaken by rice roots, greatly benefitted from the transport activity of aquaporin in rice roots. The significant enhancement (p < 0.05) of antioxidant enzyme activities reflected the oxidative stress response of rice roots upon exposure to PS-NPs. Treatment by PS-NPs decreased root length and increased lateral root numbers. Carbon metabolism was activated (e.g., increased carbon and soluble sugar contents) whereas jasmonic acid and lignin biosynthesis were inhibited. The present study demonstrated the likelihood for transport of PS-NPs in rice roots and induced phytotoxicity by PS-NPs, which should inspire further investigations into the potential human health risks from rice consumption.
Widespread application of sewage sludge produced from wastewater treatment plants for agricultural use has been regarded as a primary source of microplastics (MPs) into soils. However, little is known regarding MPs in sludge-based fertilizers and their relevant fate in soils as being applied in agriculture. We comprehensively investigated the abundance, polymer size, type and morphology of MPs in dewatered sludge, sewage sludge composts, sludge-based fertilizer-amended soils and earthworms by stereoscopy and micro-Fourier-transform infrared (µ-FTIR) spectrometry methods. Results clearly showed that the quantity of MPs in soils exhibited a close correlation with application rate of sludge-based fertilizers. The total abundances of MPs were 545.9 and 87.6 items/kg in soils after annual amendment with 30 (field A) and 15 t/ha (field B) of sludge composts, which is significantly higher than that without compost application (field C, 5.0 items/kg). Correspondingly, MPs were found in earthworms with low quantities of 1.8 and 0.4 items/individual in the field A and field B, respectively, while no MP was detected in field C. We speculate that sludge composts may act as a vehicle of MPs into soils, and then enter soil biota and in turn influence the spread of MPs in the environment.
Environmental contamination by microplastics is now considered an emerging threat to biodiversity and ecosystem functioning. Soil ecosystems, particularly agricultural land, have been recognized as a major sink of microplastics, but the impacts of microplastics on soil ecosystems (e.g., above and below ground) remain largely unknown. In this study, different types of microplastics [biodegradable polylactic acid (PLA)], conventional high-density polyethylene (HDPE), and microplastic clothing fibers were added to soil containing the endogeic Aporrectodea rosea (rosy-tipped earthworm) and planted with Lolium perenne (perennial ryegrass) to assess the biophysical soil response in a mesocosm experiment. When exposed to fibers or PLA microplastics, fewer seeds germinated. There was also a reduction in shoot height with PLA. The biomass of A. rosea exposed to HDPE was significantly reduced compared to control samples. Furthermore, with HDPE present there was a decrease in soil pH. The size distribution of water-stable soil aggregates was altered when microplastics were present, suggesting potential alterations of soil stability. This study provides evidence that microplastics manufactured of HDPE and PLA, and synthetic fibers can affect the development of L. perenne, health of A. rosea and basic, but crucial soil properties, with potential further impacts on soil ecosystem functioning.
Microplastics can affect biophysical properties of the soil. However, little is known about the cascade of events on fundamental levels of terrestrial ecosystems, i.e. starting with the changes in soil abiotic properties and propagating across the various components of soil-plant interactions, including soil microbial communities and plant traits. We investigated here the effects of six different microplastics (polyester fibers, polyamide beads, and four fragment types- polyethylene, polyester terephthalate, polypropylene, and polystyrene) on a broad suite of proxies for soil health and performance of spring onion (Allium fistulosum). Significant changes were observed in plant biomass, tissue elemental composition, root traits, and soil microbial activities. These plant and soil responses to microplastic exposure were used to propose a causal model for the mechanism of the effects. Impacts were dependent on particle type, i.e. microplastics with a shape similar to other natural soil particles elicited smaller differences from control. Changes in soil structure and water dynamics may explain the observed results, in which polyester fibers and polyamide beads triggered the most pronounced impacts on plant traits and function. The findings reported here imply that the pervasive microplastic contamination in soil may have consequences for plant performance, and thus for agroecosystems and terrestrial biodiversity.