Endophytic Bacteria Promote the Growth of Suaeda glauca in Saline-Alkali Stress:Regulation of Osmotic Pressure and Antioxidative Defense System

Endophytic bacteria are promising bacterial fertilizers to improve plant growth under adverse environment.For ecological remediation of coastal wetlands,it was necessary to investigate the effect and interaction of endophytes on halophytes under saline-alkali stress.In this study,an endophytic bacterium strain HK1 isolated from halophytes was selected to infect Suaeda glauca under pH(7 and 8)and salinity gradient(150,300 and 450mmolL^(-1)).Strain HK1 was identified as Pantoea ananatis and it had ability to fix nitrogen,dissolve inorganic phosphorus and produce indole-3-aceticacid(IAA).The results showed that strain HK1 could promote the growth of S.glauca seedings when the salinity was less than 300mmolL^(-1),in view of longer shoot length and heavier fresh weight.The infected plants could produce more proline to decrease the permeability of cells,which content increased by 26.2%–61.1%compared to the non-infected group.Moreover,the oxidative stress of infected plants was relieved with the malondialdehyde(MDA)content decreased by 16.8%–32.9%,and the peroxidase(POD)activity and catalase(CAT)activity increased by 100%–500%and 6.2%–71.4%,respectively.Statistical analysis revealed that increasing proline content and enhancing CAT and POD activities were the main pathways to alleviate saline-alkali stress by strain HK1 infection,and the latter might be more important.This study illustrated that endophytic bacteria could promote the growth of halophytes by regulation of osmotic substances and strengthening antioxidant activities.This finding would be helpful for the bioremediation of coastal soil.

Utilizing the γ-Irradiated Sodium Alginate as a Plant Growth Promoter for Enhancing the Growth,Physiological Activities,and Alkaloids Production in Catharanthus roseus L.

Sodium alginate is a polysaccharide that is largely obtained from the brown algae （Sargassum sp.）.It has been used as a wonderful growth promoting substance in its depolymerized form for various plants.The aim of this study was to find out the effects of various concentrations of γ-irradiated sodium alginate （ISA）,viz.,deionized water （control,T0）,20 （T1）,40 （T2）,60 （T3）,80 （T4）,and 100 ppm （T5） on the agricultural performance of Catharanthus roseus L.（Rosea） in terms of growth attributes,photosynthesis,physiological activities,and alkaloid production.The present work revealed that ISA applied as leaf-sprays at concentrations from 20 to 100 ppm might improve growth,photosynthesis,physiological activities,and alkaloid production in C.roseus L.significantly.Of the various ISA concentrations,80 ppm proved to be the best one compared to other concentrations applied.

Transcriptomic insights into growth promotion effect of Trichoderma afroharzianum TM2-4 microbial agent on tomato plants

Plant growth promoting fungi are receiving increased attention as valuable beneficial microorganisms in crop cultivation due to their capacity to produce bioactive substances,promote plant growth and enhance immune defense functions.In this study,a novel Trichoderma isolate,designated as TM2-4,was screened from healthy tomato rhizosphere soil and identified as Trichoderma afroharzianum.Culture filtrate of the isolate TM2-4 displayed obvious bioactive substance production and an evident effect in promoting tomato seed germination,with hypocotyl length,radical length and vigor index increased by 28.7,19.4 and 62.1%,respectively,after a 100-fold dilution treatment.To assess the promotion effect and related mechanism of isolate TM2-4,the plant biological indexes and gene expression profiles of tomato plants treated with or without T.afroharzianum TM2-4 microbial agent were investigated by greenhouse pot experiment and RNA sequencing.The results demonstrated that T.afroharzianum TM2-4 significantly promoted tomato plant growth in terms of plant height,dry weight,number of leaves per plant and root activity,through efficient colonization in the rhizosphere and root system of the plants.Transcriptome analyses identified a total of 984 differentially expressed genes in T.afroharzianum microbial agent inoculated tomato roots,which were mainly engaged in the biological process of phytohormone homeostasis,antioxidant activity,as well as metabolic pathways including phenylpropanoid biosynthesis and glutathione metabolism.These findings provide useful information for understanding the mechanism of isolate TM2-4 for tomato plant growth promotion,which would facilitate further development of T.afroharzianum TM2-4 microbial agent for use in vegetable crop production.

Phenotypic, Stress Tolerance and Plant Growth Promoting Characteristics of Rhizobial Isolates from Selected Wild Legumes of Semiarid Region, Tirupati, India

Rhizobia are vital for nitrogen input, fertility of soil and legume plant growth. Knowledge on rhizobial diversity from arid and semiarid areas is important for dry land agriculture in the context of climatic change and for economic utilization. This study provides morphological, biochemical, stress tolerance and plant growth promoting characteristics of fifteen rhizobial isolates from the nodules of same number of wild legumes and one isolate from cultivated Arachis hypogea from semi-arid region, Tirupati. The bacterial isolates were confirmed as rhizobia based on colony morphology and biochemical tests. Based on the colour change of YMA-BTB medium, eight isolates were identified as slow growers and six were fast growers. The isolates differed in growth pattern, colony morphology, antibiotic resistance at higher concentrations and uniformity in utilization of carbon and nitrogen sources. The isolates are tolerant to NaCl up to one percent, displayed normal growth at temperatures 28℃ - 30℃, at neutral pH and poor growth at pH 5and 9. The isolates varied in the production of EPS and IAA, positive for phosphate solubilization and siderophore formation. This functional diversity displayed by the isolates can be utilised for the legume crop production by cross inoculation.

Evaluation of Plant Growth Promoting Ability of Bacillus amyloliquefaciens Bc2 and Trichoderma harzianum TR In Vivo

Bacillus amyloliquefaciens Bc2 and Trichderma harzianum TR were used to evaluate their growth-promoting activity on cultivated strawberries, under laboratory and field conditions, and we have noticed that the percentage of achene germination is important for ones treated with TR (=97%) followed by those treated with Bc2 strain (=90%) and the control (=84%). Inoculations on field showed that on untreated soil with insecticide, TR is effective and allows the development of plants and extends the duration of flowering and fruiting. On treated soil, Bc2 clearly promotes the growth and development of strawberry seedlings and its role as plant growth promoting microorganisms has been proved.

Plant growth-promoting characteristics of root fungal endophytes isolated from a traditional Cordillera rice landrace

This study was conducted to isolate and characterize the plant growth-promoting potential of fungal endophytes from the roots of Diket red,a traditional rice plant from the Cordillera,Northern Luzon.Eighteen morphospecies of filamentous endophytes were isolated of which twelve isolates were successfully identified to the species level.These isolates were identified as Aspergillus versicolor,Aspergillus sp.,Chaetosphaeria sp.,Cladosporium cladosporioides,Hypocrea lixii,Microascus murinus,and Trichoderma harzianum.The identified twelve isolates were selected to screen in vitro for their plant growth-promoting characteristics,and evaluated in vivo for their beneficial effects on seedling vigor and early seedling growth.Isolate PPL14 produced the highest IAA at 55.5μg ml^(-1)and M.murinus PPL10 produced the highest amount of IAA at 3.73μg IAA mg^(-1)dry mycelia wt.The seedling vigor assay and in vivo plant growth promotion bioassay indicated overall positive effects of culture filtrate(CF)application of the endophyte isolates.Rice seeds and seedlings grown in aseptic condition and treated with endophyte CFs displayed significantly enhanced levels of germination,seedling vigor,shoot,root,and total plant growth,and biomass compared to non-treated control.Other plant growth-promoting characteristics were also studied including phosphate solubilization,siderophore production,ammonia production,and catalase activity.This study supports the potential use of fungal endophytes as bio-inoculants for plant growth promotion and enhancement of nutrient assimilation of agriculturally important crops.

The rhizosphere microbial complex in plant health:A review of interaction dynamics

Climate change,urbanization,and population increase limit food availability.To sustain human existence,there is the need to increase food and agricultural production to mitigate the impact of these factors.Scientists have been working for years on ways to increase food production.From plant breeding techniques to soil science,scientists have made tremendous progress.The rhizobiome has been proven to be important to crop production,and the impact of the rhizobiome on plant health cannot be overemphasized.Being rich in diverse complex microbial interactions,the rhizosphere has become a major force in recent plant growth promotion studies.The upsurge in next-generation sequencing applications with the various“omics”technologies is helping to unearth information relating to rhizosphere impact on plant growth.Explaining the complex interactions between and across microbial species present in the rhizosphere is important to further enhance our understanding of their mechanistic and mutualistic functions.Knowledge from this can be used in rhizosphere biome engineering for improved plant growth and yield in the face of the various biotic and abiotic challenges.

Potential of Biostimulants Based on PGPR Rhizobacteria Native to Benin’s Soils on the Growth and Yield of Maize (<i>Zea mays </i>L.) under Greenhouse Conditions

The application of biostimulants in agriculture represents an environmentally friendly alternative while increasing agricultural production. The aims of the study were to develop solid biostimulants based on five rhizobacteria native to Benin’s soils and to evaluate their efficacy on the growth and biomass yield of maize under greenhouse conditions on ferrallitic and ferruginous soils. Clay and peat were used as a conservation binder for the preparation of the biostimulants. These binders were used alone or combined in the different formulations with maize flour and sucrose. 10 g of biostimulants were applied at sowing in pots containing five kilograms of sterilised soil. The experimental design was a completely randomised block of 24 treatments with three replicates. The results obtained showed significant improvements (P < 0.001) in height (49.49%), stem diameter (32.7%), leaf area (66.10%), above-ground biomass (97.12%) and below-ground biomass (53.98%) on ferrallitic soil with the application of the clay + Pseudomonas putida biostimulant compared to the control. On the other hand, the use of the peat biostimulant + Pseudomonas syringae was more beneficial for plant growth on ferruginous soil. The height, stem diameter, leaf area, above-ground biomass and below-ground biomass of the plants under the influence of this biostimulant were improved by 83.06%, 44.57%, 102.94%, 86.84% and 42.68%, respectively, compared to the control. Therefore, these results confirm that Rhizobacteria express their potential through biostimulants formulated on maize. The formulated biostimulants can later be used by producers to improve crop productivity for sustainable agriculture.

Increased ammonification,nitrogenase,soil respiration and microbial biomass N in the rhizosphere of rice plants inoculated with rhizobacteria

Azospirillum brasilense and Pseudomonas fluorescens are well-known plant growth promoting rhizobacteria.However,the effects of A.brasilense and P.fluorescens on the N cycles in the paddy field and rice plant growth are little known.This study investigated whether and how A.brasilense and P.fluorescens contribute to the N transformations and N supply capacities in the rhizosphere,and clarified the effects of A.brasilense and P.fluorescens on the N application rate in rice cultivation.Inoculations with A.brasilense and P.fluorescens coupled with N application rate trials were conducted in the paddy field in 2016 and 2017.The inoculations of rice seedlings included four treatments:sterile saline solution(M_(0)),A.brasilense(M_(b)),P.fluorescens(M_(p)),and co-inoculation with a mixture of A.brasilense and P.fluorescens(M_(bp)).The N application rate included four levels:0 kg N ha^(–1)(N_(0)),90 kg N ha^(–1)(N_(90)),180 kg N ha^(–1)(N_(180)),and 270 kg N ha^(–1)(N_(270)).The results indicated that the M_(bp) and M_(p) treatments significantly enhanced the ammonification activities in the rhizosphere compared with the M_(0) treatment,especially for higher N applications,while the Mbp and M_(b) treatments greatly enhanced the nitrogenase activities in the rhizosphere compared with the M_(0) treatments,especially for lower N applications.Azospirillum brasilense and P.fluorescens did not participate in the nitrification processes or the denitrification processes in the soil.The soil respiration rate and microbial biomass N were greatly affected by the interactions between the rhizobacteria inoculations and the N fertilizer applications.In the M_(bp) treatment,N supply capacities and rice grain yields showed no significant differences among the N_(90),N_(180),and N_(270) applications.The N application rate in the study region can be reduced to 90 kg N ha^(–1) for rice seedlings co-inoculated with a mixture of A.brasilense and P.fluorescens.

Research Progress in Biological Control of Soft Rot of Amorphophallus konjac

In this paper,the main control methods of soft rot of Amorphophallus konjac are reviewed,with a focus on the current research status of using plant growth promoting rhizobacteria for biological control of soft rot of A.konjac,and future research directions are looked forward to.

Actinobacteria-enhanced plant growth, nutrient acquisition, and crop protection:Advances in soil, plant, and microbial multifactorial interactions

Agricultural areas of land are deteriorating every day owing to population increase, rapid urbanization, and industrialization. To feed today’s huge populations, increased crop production is required from smaller areas, which warrants the continuous application of high doses of inorganic fertilizers to agricultural land. These cause damage to soil health and, therefore, nutrient imbalance conditions in arable soils. Under these conditions, the benefits of microbial inoculants (such as Actinobacteria) as replacements for harmful chemicals and promoting ecofriendly sustainable farming practices have been made clear through recent technological advances. There are multifunctional traits involved in the production of different types of bioactive compounds responsible for plant growth promotion, and the biocontrol of phytopathogens has reduced the use of chemical fertilizers and pesticides. There are some well-known groups of nitrogen-fixing Actinobacteria, such as Frankia, which undergo mutualism with plants and offer enhanced symbiotic trade-offs.In addition to nitrogen fixation, increasing availability of major plant nutrients in soil due to the solubilization of immobilized forms of phosphorus and potassium compounds, production of phytohormones, such as indole-3-acetic acid, indole-3-pyruvic acid, gibberellins, and cytokinins, improving organic matter decomposition by releasing cellulases, xylanase, glucanases, lipases, and proteases, and suppression of soil-borne pathogens by the production of siderophores, ammonia, hydrogen cyanide, and chitinase are important features of Actinobacteria useful for combating biotic and abiotic stresses in plants.The positive influence of Actinobacteria on soil fertility and plant health has motivated us to compile this review of important findings associated with sustaining plant productivity in the long run.

Genomic analyses of metal resistance genes in three plant growth promoting bacteria of legume plants in Northwest mine tailings, China

To better understand the diversity of metal resistance genetic determinant from microbes that survived at metal tailings in northwest of China, a highly elevated level of heavy metal containing region, genomic analyses was conducted using genome sequence of three native metal-resistant plant growth promoting bacteria（PGPB）. It shows that： Mesorhizobium amorphae CCNWGS0123 contains metal transporters from P-type ATPase, CDF（Cation Diffusion Facilitator）, Hup E/Ure J and CHR（chromate ion transporter） family involved in copper, zinc, nickel as well as chromate resistance and homeostasis. Meanwhile, the putative Cop A/Cue O system is expected to mediate copper resistance in Sinorhizobium meliloti CCNWSX0020 while Znt A transporter, assisted with putative Czc D, determines zinc tolerance in Agrobacterium tumefaciens CCNWGS0286. The greenhouse experiment provides the consistent evidence of the plant growth promoting effects of these microbes on their hosts by nitrogen fixation and/or indoleacetic acid（IAA） secretion,indicating a potential in-site phytoremediation usage in the mining tailing regions of China.

Applications of beneficial plant growth promoting rhizobacteria and mycorrhizae in rhizosphere and plant growth:A review

Because of climate change and the highly growing world population,it becomes a huge challenge to feed the whole population.To overcome this challenge and increase the crop yield,a large number of fertilizers are applied but these have many side effects.Instead of these,scientists have discovered beneficial rhizobacteria,which are environmentally friendly and may increase crop yield and plant growth.The microbial population of the rhizosphere shows a pivotal role in plant development by inducing its physiology.Plant depends upon the valuable interactions among the roots and microbes for the growth,nutrients availability,growth promotion,disease suppression and other important roles for plants.Recently numerous secrets of microbes in the rhizosphere have been revealed due to huge development in molecular and microscopic technologies.This review illustrated and discussed the current knowledge on the development,maintenance,interactions of rhizobacterial populations and various proposed mechanisms normally used by PGPR in the rhizosphere that encouraging the plant growth and alleviating the stress conditions.In addition,this research reviewed the role of single and combination of PGPR,mycorrhizal fungi in plant development and modulation of the stress as well as factors affecting the microbiome in the rhizosphere.

Improvement of Sporulation Conditions of a New Strain of Bacillus amyloliquefaciens in Liquid Fermentation

The desirable active ingredients for the development of bioproducts based in Bacillus sp. for the control of soil pathogens are the spores because these structures exhibit more resistance and stability to conditions present during the fermentation, formulation, and storage processes. To improve the sporulation of a native strain of Bacillus amyloliquefaciens (Bs006) using liquid fermentation, some modifications in the concentrations of the components in a previously standardized culture media were made. Subsequently, five sporulation inducers (iron nitrate, mixture of salts, peroxide hydrogen, temperature, and initial cell concentration) were evaluated. The treatment with a mixture of salts in combination with an initial cell concentration of 1 × 108 cells/ml was selected because a final spore concentration of 1.05 × 1010 spores/ml after 66 hours with a fully substrate consumption and sporulation efficiency of 88.7% was obtained. To demonstrate the biological activity of B. amyloliquefaciens Bs006 in Cape gooseberry seedlings, a greenhouse bioassay was conducted, and statistical differences in plant growth-promoting parameters compared with previous media were not found. Additionally, the proposed modified media (coded as JM) presented a benefit-cost ratio 2.65 times higher compared with the baseline media.

Prospecting microbial biofilms as climate smart strategies for improving plant and soil health: A review

Microbes and their products play key roles in complementing chemical fertilizers and plant protection chemicals by eliciting defence mechanisms in crop plants,thereby providing immunity and resistance against diverse stresses.Among the different environmental technologies used to mitigate climate change,several microbiological interventions are promising,among which biofilms,both naturally-existing and laboratory-developed,and their application have gained momentum recently.Microbial biofilms are aggregations of microbial cells in a self-generated polymeric matrix,which are produced by several genera of bacteria,yeasts,cyanobacteria,and fungi as a mode of reproductive fitness,thereby advantageous in successful proliferation,even in extreme environments.The propensity of microbial biofilms to grow in diverse niches and adapt to biotic and abiotic stresses illustrates their immense potential as biofertilizers and disease suppression options in various crops across diverse ecologies.In the inhospitable habitats of deserts and denudated land,biofilms play an important role in preventing soil erosion and sustaining vegetation,microflora,and fauna.Biofilms represent a mode of growth for several microbes with plant growth-promoting and biocontrol potential,which are important in seed establishment and the facilitation of exchanging nutrients and metabolites from the environment.In this review,we discuss the prospects of microbial biofilms as a green option in agriculture in general and,more specifically,their potential in mitigating climate change.

Utilization of lignocellulosic plant residues for compost formation and its role in improving soil fertility

Globally,urbanization and a steady increase in population generate a huge amount of wastes,which leads to a series of economic,social,and environmental changes,mainly in developing countries.There is an utmost need for efficient management strategies for the beneficial utilization of these wastes into useful products.Among these strategies,composting is gaining attention due to its benefits of solid waste management,such as proper sterilization,and economical and effective bioconversion of lignocellulosic wastes to valuable products.Composting is an effective and sustainable approach for the management of various lignocellulosic wastes.This process comprises a series of effective waste treatment steps to ensure sustainable agriculture.Different composting methods have been explored for solid waste management.Furthermore,the influence of various factors relevant to composting has been elucidated.Microbes play a significant role in enhancing the degradation of lignocellulosic residues by secreting different hydrolytic enzymes.Compost has been utilized for increasing soil properties and improving plant growth.

Role of beneficial microbial gene pool in mitigating salt/nutrient stress of plants in saline soils through underground phytostimulating signalling molecules

Soil salinity diminishes soil health and reduces crop yield,which is becoming a major global concern.Salinity stress is one of the primary stresses,leading to several other secondary stresses that restrict plant growth and soil fertility.The major secondary stresses induced in plants under saline-alkaline conditions include osmotic stress,nutrient limitation,and ionic stress,all of which negatively impact overall plant growth.Under stressed conditions,certain beneficial soil microflora are known to have evolved phytostimulating mechanisms,such as the synthesis of osmoprotectants,siderophores,1-aminocyclopropane-1-carboxylic acid(ACC)deaminase activity,phosphate solubilization,and hormone production,which enhance plant growth and development while mitigating nutrient stress.Beneficial soil-borne bacterial species such as Bacillus,Pseudomonas,and Klebsiella and fungal strains such as Trichoderma,Aspergillus,Penicillium,Alternaria,and Fusarium also aid in reducing salinity stress.Phosphate-solubilizing microorganisms also assist in nutrient acquisition via both enzymatic and non-enzymatic processes.In the case of enzymatic processes,they produce different enzymes such as alkaline phosphatases and phytases,whereas non-enzymatic processes produce organic acids such as gluconic,citric,malic,and oxalic acids.The native halotolerant/halophilic soil microbial gene pool with multifunctional traits and stress-induced gene expression can be developed as suitable bio-inoculants to enhance stress tolerance and optimize plant growth in saline soils.

Root symbionts： Powerful drivers of plant above- and belowground indirect defenses

Soil microbial mutualists of plants, including mycorrhizal ftmgi, non- mycorrhizal fungi and plant growth promoting rhizobacteria, have been typically characterized for increasing nutrient acquisition and plant growth. More recently, soil microbes have also been shown to increase direct plant defense against above- and below- ground herbivores. Plants, however, do not only rely on direct defenses when attacked, but they can also recruit pest antagonists such as predators and parasitoids, both above and belowground, mainly via the release of volatile organic compounds （i.e., indirect defenses）. In this review, we illustrate the main features and effects of soil microbial mutualists of plants on plant indirect defenses and discuss possible applications within the framework of sustainable crop protection against root- and shoot-feeding arthropod pests. We indicate the main knowledge gaps and the future challenges to be addressed in the study and application of these rnultifaceted interactions.

Benefits of a root fungal endophyte on physiological processes and growth of the vulnerable legume tree Prosopis chilensis(Fabaceae)

Aims Desertification is a major concern in arid and semi-arid regions globally.Understanding interactions between vulnerable plant spe-cies and associated microbial symbionts may have important appli-cations for conservation and restoration strategies in affected areas.Methods In this study,we evaluated the root-associated fungal endophyte community in Prosopis chilensis,a vulnerable and threatened arid-adapted tree of northern Chile.Host benefits in terms of physiologi-cal performance and plant growth were also assessed.Endophytic fungi were isolated from asymptomatic roots by the culture method for molecular identification of the 18S rRNA gene.The dominant fungal endophyte(Penicillium sp.)in the community was later used in inoculation experiments to assess its effect on maximum quan-tum efficiency of photosystem II(_(PSII)),(Fv/Fm)actual _(PSII) efficiency(Φ_(PSII)),and non-photochemical quenching(NPQ).Total sugars,starch content,malonaldehyde(MDA),nitrogen content(%)and growth traits were also measured.Important Findings WhereasΦ_(PSII) increased significantly in endophyte-inoculated plants,NPQ was found to decrease.No effect of endophyte inocula-tion on sugars and MDA was detected,but starch content,leaf nitro-gen content,number of leaves and shoot biomass were found to increase.Results revealed that inoculation of endophytic Penicillium fungal isolate can provide significant physiological benefits to the host plant P.chilensis.Its presence resulted in greater _(PSII) efficiency and higher leaf nitrogen and carbohydrate content,enhancing host plant growth.These findings highlight the importance of consider-ing the fungal endophyte community of this vulnerable species as an important tool to the design of further revegetation and conserva-tion programs.

Mutualistic fungus Piriformospora indica modulates cadmium phytoremediation properties of host plant via concerted action of enzymatic and non-enzymatic biochemicals

Soils and ecosystems contaminated with cadmium (Cd) threaten human health and adversely affect morphological,physiological,and biochemical parameters of plants.The symbiotic association of endophytic fungi with their host plants is the best strategy to improve various plant characteristics and remediate soils polluted with heavy metal(loid)s (HMs).Being a well-known plant growth-promoting fungus,Piriformospora indica confers resistance against a number of abiotic stresses,including HM stress.This pot experiment explored the potential and ameliorative effects of P.indica on Artemisia annua L.plants treated with different concentrations (0,40,80,and 120 mg kg-1) of Cd.Inoculation with P.indica significantly increased plant performance,especially by enhancing chlorophyll concentration and water potential and by decreasing electrolytic leakage,when compared with un-inoculated plants,despite the high Cd levels.Similarly,P.indica enhanced antioxidant enzyme activities,thereby reducing the drastic effects of Cd in inoculated plants.In addition,P.indica accumulated Cd in the roots of colonized plants,as revealed by atomic absorption spectroscopy,and restricted Cd translocation to aerial parts.Furthermore,P.indica showed in vitro resistance (up to a certain level) to Cd stress;however,fungus growth was inhibited at very high Cd concentrations,proving it an excellent candidate for use as a potential phytoremediator in fields affected by Cd contamination.The transcriptional analysis showed that the signaling genes and artemisinin and flavonoid biosynthetic pathway genes were significantly upregulated in P.indica-co-cultivated plants when compared with un-inoculated plants,suggesting a fine collaboration between primary and secondary metabolisms to modulate resistance capacity and to enhance the phytoremediation capability of A.annua against Cd toxicity.