盐胁迫环境下植物促生菌的作用机制研究进展

盐胁迫是限制干旱和半干旱地区作物生产的主要非生物胁迫之一,严重影响作物的生长发育,植物促生菌(Plant growth-promoting bacteria,PGPB)可有效减轻植物的盐胁迫损伤,合理施用PGPB是盐胁迫下促进作物生长的重要途径。本文从盐胁迫环境下PGPB在调节植物激素内稳态、促进养分吸收和诱导植物产生系统耐受性等方面的作用阐述了PGPB提高植物耐盐性、减轻植物胁迫损伤的作用机制。讨论了能够在植物根际稳定定殖并在盐生环境下稳定保持PGP活性的功能菌株对未来农业的可持续发展的重要意义,同时,对该研究方向的重难点和未来的发展趋势作出展望。     

Enhanced Cd extraction of oilseed rape (Brassica napus) by plant growth-promoting bacteria isolated from Cd hyperaccumulator Sedum alfredii Hance

Four plant growth-promoting bacteria (PGPB) were used as study materials, among them two heavy metal-tolerant rhizosphere strains SrN1 (Arthrobacter sp.) and SrN9 (Bacillus altitudinis) were isolated from rhizosphere soil, while two endophytic strains SaN1 (Bacillus megaterium) and SaMR12 (Sphingomonas) were identified from roots of the cadmium (Cd)/zinc (Zn) hyperaccumulator Sedum alfredii Hance. A pot experiment was carried out to investigate the effects of these PGPB on plant growth and Cd accumulation of oilseed rape (Brassica napus) plants grown on aged Cd-spiked soil. The results showed that the four PGPB significantly boosted oilseed rape shoot biomass production, improved soil and plant analyzer development (SPAD) value, enhanced Cd uptake of plant and Cd translocation to the leaves. By fluorescent in situ hybridization (FISH) and green fluorescent protein (GFP), we demonstrated the studied S. alfredii endophytic bacterium SaMR12 were able to colonize successfully in the B. napus roots. However, all four PGPB could increase seed Cd accumulation. Due to its potential to enhance Cd uptake by the plant and to restrict Cd accumulation in the seeds, SaMR12 was selected as the most promising microbial partner of B. napus when setting up a plant–microbe fortified remediation system.     

Effect of plant growth promoting bacterium; Pseudomonas putida UW4 inoculation on phytoremediation efficacy of monoculture and mixed culture of selected plant species for PAH and lead spiked soils

Plant growth promoting bacteria (PGPB) enhanced phytoremediation (PEP) is an attractive remedial strategy for the remediation of polycyclic aromatic hydrocarbon (PAH) and heavy metal (HM) contaminated sites. The effect of PGPB; Pseudomonas putida UW4 inoculation on the phytoremediation efficiency of Medicago sativa, Festuca arundinacea, Lolium perenne, and mixed plants (L. perenne and F. arundinacea) was assessed. This involved two contaminant treatments; “PAH” (phenanthrene; 300?mg·kg^?1, fluoranthene; 200?mg·kg^?1, and benzo[a]pyrene; 5?mg·kg^?1) and “PAH?+?HM” (‘PAH’ treatments +100?mg of Pb/kg). PGPB inoculation significantly enhanced root biomass yield of F. arundinacea in PAH treatment, and the mixed plant shoot biomass and L. perenne root biomass yields of the PAH?+?HM treatment. PGPB significantly enhanced dissipation of phenanthrene and fluoranthene for M. sativa-PAH?+?PGPB treatment and fluoranthene for F. arundinacea-PAH?+?HM?+?PGPB treatment. In others, PGPB inoculation either had no impact or inhibited PAH dissipation. PAH dissipation for the single and mixed plant treatments with PGPB inoculation were not different. The efficiency of PEP is dependent on different factors such as PGPB inoculum biomass, plant species, plant–microbe specificity and type of contaminants. Exploiting PEP technology would require proper understanding of plant tolerance and growth promoting mechanisms, and rhizosphere activities.     

Metabolic profile and antioxidant responses during drought stress recovery in sugarcane treated with humic acids and endophytic diazotrophic bacteria

Water deficit is the major yield‐limiting factor for sugarcane crop production that can be enhanced by inoculating with plant growth promoting bacteria (PGPB) combined with humic substances. The aim of this work was to examine changes to the metabolic profile and antioxidant enzyme activity of sugarcane treated with PGPB and humic acid (HA) after drought and then rehydration. The drought was imposed by withholding irrigation for 21 days thereby measuring enzyme activity, metabolic profile and photosynthetic rate 1 week after rehydratation. Growth of plants treated with HA, PGPB and with both treatments combined (PGPB + HA) was higher than control plants. The antioxidant enzymes superoxide dismutase (SOD), catalase (CAT) and ascorbate peroxidase (APX) activities remained higher after rehydration only in plants treated with HA. Plants treated with HA and PGPB + HA exhibited increased transpiration, stomatal conductance and net photosynthesis than plants treated with PGPB. The PGPB‐treated plants exhibited drought resistance that resembled ‘delayed stress onset’, which is a new term for preserving water in the plants tissues. Water preservation in plants treated with PGPB was corroborated by higher relative water content (RWC) than control plants at the end of the drought period. Plants treated with HA + PGPB exhibited the highest water potential after rehydration and high RWC. Osmotic adjustment in the other treatments (control, HA and PGPB) was indicated by a new pattern of metabolic response after rehydration, including generally enhanced carbohydrates and proteins and specific changes induced by HA‐enhancing aromatic compounds, whereas PGPB exhibited enhanced fatty acids and other aliphatic H species. Humic acids assist with drought stress recovery by inducing antioxidant enzyme activity whereas PGPB induced preservation of leaf water potential and RWC by closing stomata efficiently, resulting in plant water preservation.     

Are drought-resistance promoting bacteria cross-compatible with different plant models?

The association between plant and plant growth promoting bacteria (PGPB) contributes to the successful thriving of plants in extreme environments featured by water shortage. We have recently shown that, with respect to the non-cultivated desert soil, the rhizosphere of pepper plants cultivated under desert farming hosts PGPB communities that are endowed with a large portfolio of PGP traits. Pepper plants exposed to bacterial isolates from plants cultivated under desert farming exhibited a higher tolerance to water shortage, compared with untreated control. This promotion was mediated by a larger root system (up to 40%), stimulated by the bacteria, that enhanced plant ability to uptake water from dry soil. We provide initial evidence that the nature of the interaction can have a limited level of specificity and that PGPB isolates may determine resistance to water stress in plants others than the one of the original isolation. It is apparent that, in relation to plant resistance to water stress, a feature of primary evolutionary importance for all plants, a cross-compatibility between PGPB and different plant models exists at least on a short-term.     

Improvement of Maize Yield by Foliar Application of Azospirillum brasilense Az39

Journal of Plant Growth Regulation - Azospirillum brasilense is a plant growth promoting bacteria (PGPB) widely used as an inoculant in diverse species, including maize (Zea mays L.) to improve...     

Associative bacteria influence maize (Zea mays L.) growth,physiology and root anatomy under different nitrogen levels

• Despite the great diversity of plant growth‐promoting bacteria (PGPB ) with potential to partially replace the use of N fertilisers in agriculture, few PGPB have been explored for the production of commercial inoculants, reinforcing the importance of identifying positive plant–bacteria interactions. Aiming to better understand the influence of PGPB inoculation in plant development, two PGPB species with distant phylogenetic relationship were inoculated in maize.
• Maize seeds were inoculated with Bacillus sp. or Azospirillum brasilense . After germination, the plants were subjected to two N treatments: full (N+) and limiting (N?) N supply. Then, anatomical, biometric and physiological analyses were performed.
• Both PGPB species modified the anatomical pattern of roots, as verified by the higher metaxylem vessel element (MVE ) number. Bacillus sp. also increased the MVE area in maize roots. Under N+ conditions, both PGPB decreased leaf protein content and led to development of shorter roots; however, Bacillus sp. increased root and shoot dry weight, whereas A. brasilense increased photosynthesis rate and leaf nitrate content. In plants subjected to N limitation (N?), photosynthesis rate and photosystem II efficiency increased in maize inoculated with Bacillus sp., whilst A. brasilense contained higher ammonium, amino acids and total soluble sugars in leaves, compared to the control.
• Plant developmental and metabolical patterns were switched by the inoculation, regardless of the inoculant bacterium used, producing similar as well as distinct modifications to the parameters studied. These results indicate that even non‐diazotrophic inoculant strains can improve the plant N status as result of the morpho‐anatomical and physiological modifications produced by the PGPB .

     

Mechanisms of action of plant growth promoting bacteria

The idea of eliminating the use of fertilizers which are sometimes environmentally unsafe is slowly becoming a reality because of the emergence of microorganisms that can serve the same purpose or even do better. Depletion of soil nutrients through leaching into the waterways and causing contamination are some of the negative effects of these chemical fertilizers that prompted the need for suitable alternatives. This brings us to the idea of using microbes that can be developed for use as biological fertilizers (biofertilizers). They are environmentally friendly as they are natural living organisms. They increase crop yield and production and, in addition, in developing countries, they are less expensive compared to chemical fertilizers. These biofertilizers are typically called plant growth-promoting bacteria (PGPB). In addition to PGPB, some fungi have also been demonstrated to promote plant growth. Apart from improving crop yields, some biofertilizers also control various plant pathogens. The objective of worldwide sustainable agriculture is much more likely to be achieved through the widespread use of biofertilizers rather than chemically synthesized fertilizers. However, to realize this objective it is essential that the many mechanisms employed by PGPB first be thoroughly understood thereby allowing workers to fully harness the potentials of these microbes. The present state of our knowledge regarding the fundamental mechanisms employed by PGPB is discussed herein.     

Genetics of phosphate solubilization and its potential applications for improving plant growth-promoting bacteria

Plant growth-promoting bacteria (PGPB) are soil and rhizosphere bacteria that can benefit plant growth by different mechanisms. The ability of some microorganisms to convert insoluble phosphorus (P) to an accessible form, like orthophosphate, is an important trait in a PGPB for increasing plant yields. In this mini-review, the isolation and characterization of genes involved in mineralization of organic P sources (by the action of enzymes acid phosphatases and phytases), as well as mineral phosphate solubilization, is reviewed. Preliminary results achieved in the engineering of bacterial strains for improving capacity for phosphate solubilization are presented, and application of this knowledge to improving agricultural inoculants is discussed.     

Mixed planting with a leguminous plant outperforms bacteria in promoting growth of a metal remediating plant through histidine synthesis

The effectiveness of plant growth promoting bacteria (PGPB) in improving metal phytoremediation is still limited by stunted plant growth under high soil metal concentrations. Meanwhile, mixed planting with leguminous plants is known to improve yield in nutrient deficient soils but the use of a metal tolerant legume to enhance metal tolerance of a phytoremediator has not been explored. We compared the use of Pseudomonas brassicacearum, Rhizobium leguminosarum, and the metal tolerant leguminous plant Vicia sativa to promote the growth of Brassica juncea in soil contaminated with 400 mg Zn kg^–1, and used synchrotron based microfocus X-ray absorption spectroscopy to probe Zn speciation in plant roots. B. juncea grew better when planted with V. sativa than when inoculated with PGPB. By combining PGPB with mixed planting, B. juncea recovered full growth while also achieving soil remediation efficiency of >75%, the maximum ever demonstrated for B. juncea. μXANES analysis of V. sativa suggested possible root exudation of the Zn chelates histidine and cysteine were responsible for reducing Zn toxicity. We propose the exploration of a legume-assisted-phytoremediation system as a more effective alternative to PGPB for Zn bioremediation.     

Effects of inoculation of plant-growth promoting bacteria on Ni uptake by Indian mustard

In this study, among a collection of Ni resistant bacterial strains isolated from serpentine soil, two plant growth promoting bacteria (PGPB), Ps29C and Bm4C were selected based on their ability to utilize ACC as the sole N source and promote seedling growth in roll towel assay. The Ni resistant PGPB, Ps29C and Bm4C were characterized as Pseudomonas sp. and Bacillus megaterium, respectively, on the basis of their 16s rDNA sequences. Assessment of the parameters of plant growth promotion revealed the intrinsic ability of the strains for the production of IAA, siderophore and solubilization of insoluble phosphate. Further, the plant growth promoting activity of Ps29C and Bm4C on the Indian mustard (Brassica juncea) were assessed with different concentrations of Ni in soil. Inoculation of Ps29C or Bm4C promoted plant growth and protected the plant from Ni toxicity. However, the maximum growth was observed in the plants inoculated with strain Bm4C. Inoculation with Ps29C or Bm4C had little influence on the accumulation of Ni in root and shoot system, but produced a much larger aboveground biomass. The present observations showed that the strains Ps29C and Bm4C protect the plants against the inhibitory effects of nickel, probably due to the production of IAA, siderophore and solubilization of phosphate. The above results provided a new insight into the phytoremediation of Ni contaminated soil.     

Effects of plant growth promoting bacteria and mycorrhizal on Capsicum annuum L. var. aviculare ([Dierbach] D’Arcy and Eshbaugh) germination under stressing abiotic conditions

Capsicum annuum var. aviculare to Tarahumara and Papago Indians and farmers of Sonora desert is a promising biological and commercial value as a natural resource from arid and semiarid coastal zones. Traditionally, apply synthetic fertilizers to compensate for soil nitrogen deficiency. However, indiscriminate use of these fertilizers might increase salinity. The inoculation by plant growth promoting bacteria (PGPB) and arbuscular mycorrhizal fungi (AMF) represents an alternative as potential bio fertilizer resources for salty areas. Seeds ecotypes from four areas of Sonora desert (Mazocahui, Baviacora, Arizpe, La Tortuga), in order to inoculate them with one species of PGPB and AMF. Two germination tests were carried out to study the effect of salinity, temperature regime (night/day) and inoculation with PGPB and AMF growth factors measured on germination (percentage and rate), plant height, root length, and produced biomass (fresh and dry matter). The results indicated that from four studied ecotypes, Mazocahui was the most outstanding of all, showing the highest germination under saline and non-saline conditions. However, the PGPB and AMF influenced the others variables evaluated. This study is the first step to obtain an ideal ecotype of C. a. var. aviculare, which grows in the northwest of México and promoting this type of microorganisms as an efficient and reliable biological product. Studies of the association of PGPB and AMF with the C. a. var. aviculare-Mazocahui ecotype are recommended to determine the extent to which these observations can be reproduced under field conditions.     

Plant Growth-Promoting Bacteria Facilitate the Growth of Barley and Oats in Salt-Impacted Soil: Implications for Phytoremediation of Saline Soils

Plant growth-promoting bacteria (PGPB) strains that contain the enzyme 1-amino- cyclopropane-1-carboxylate (ACC) deaminase can lower stress ethylene levels and improve plant growth. In this study, ACC deaminase-producing bacteria were isolated from a salt-impacted (～50 dS/m) farm field, and their ability to promote plant growth of barley and oats in saline soil was investigated in pouch assays (1% NaCl), greenhouse trials (9.4 dS/m), and field trials (6–24 dS/m). A mix of previously isolated PGPB strains UW3 (Pseudomonas sp.) and UW4 (P. sp.) was also tested for comparison. Rhizobacterial isolate CMH3 (P. corrugata) and UW3+UW4 partially alleviated plant salt stress in growth pouch assays. In greenhouse trials, CMH3 enhanced root biomass of barley and oats by 200% and 50%, respectively. UW3+UW4, CMH3 and isolate CMH2 also enhanced barley and oat shoot growth by 100%–150%. In field tests, shoot biomass of oats tripled when treated with UW3+UW4 and doubled with CHM3 compared with that of untreated plants. PGPB treatment did not affect salt uptake on a per mass basis; higher plant biomass led to greater salt uptake, resulting in decreased soil salinity. This study demonstrates a method for improving plant growth in marginal saline soils. Associated implications for salt remediation are discussed.     

Plant growth-promoting bacteria and nitrate availability: impacts on root development and nitrate uptake

Plant growth-promoting bacteria (PGPB) and NO-3 availability both affect NO-3 uptake and root architecture. The presence of external NO-3 induces the expression of NO-3 transporter genes and elicits lateral root elongation in the part of the root system exposed to the NO-3 supply. By contrast, an increase in NO-3 supply leads to a higher plant N status (low N demand), which represses both the NO-3 transporters and lateral root development. The effects of PGPB on NO-3 uptake and root development are similar to those of low NO-3 availability (concomitant stimulation of NO-3 uptake rate and lateral root development). The mechanisms responsible for the localized and long-distance regulation of NO-3 uptake and root development by NO-3 availability are beginning to be elucidated. By contrast, the signalling and transduction pathways elicited by the rhizobacteria remain totally unknown. This review will compare the effects of NO-3 availability and PGPB on root morphogenesis and NO-3 uptake, in order to determine whether interactions exist between the NO-3-dependent and the PGPB-dependent regulatory pathways.     

ACC deaminase from plant growth-promoting bacteria affects crown gall development

In addition to the well-known roles of indoleacetic acid and cytokinin in crown gall formation, the plant hormone ethylene also plays an important role in this process. Many plant growth-promoting bacteria (PGPB) encode the enzyme 1-aminocyclopropane-1-carboxylate (ACC) deaminase, which can degrade ACC, the immediate precursor of ethylene in plants, to alpha-ketobutyrate and ammonia and thereby lower plant ethylene levels. To study the effect of ACC deaminase on crown gall development, an ACC deaminase gene from the PGPB Pseudomonas putida UW4 was introduced into Agrobacterium tumefaciens C58, so that the effect of ACC deaminase activity on tumour formation in tomato and castor bean plants could be assessed. Plants were also coinoculated with A. tumefaciens C58 and P. putida UW4 or P. putida UW4-acdS- (an ACC deaminase minus mutant strain). In both types of experiments, it was observed that the presence of ACC deaminase generally inhibited tumour development on both tomato and castor bean plants.     

Comparative genomics reveals the acquisition of mobile genetic elements by the plant growth-promoting Pantoea eucrina OB49 in polluted environments

Heavy metal-tolerant plant growth-promoting bacteria (PGPB) have gained popularity in bioremediation in recent years. A genome-assisted study of a heavy metal-tolerant PGPB Pantoea eucrina OB49 isolated from the rhizosphere of wheat grown on a heavy metal-contaminated site is presented. Comparative pan-genome analysis indicated that OB49 acquired heavy metal resistance genes through horizontal gene transfer. On contigs S10 and S12, OB49 has two arsRBCH operons that give arsenic resistance. On the S12 contig, an arsRBCH operon was discovered in conjunction with the merRTPCADE operon, which provides mercury resistance. P. eucrina OB49 may be involved in an ecological alternative for heavy metal remediation and growth promotion of wheat grown in metal-polluted soils. Our results suggested the detection of mobile genetic elements that harbour the ars operon and the fluoride resistance genes adjacent to the mer operon.     

Interrelationship of Bradyrhizobium sp. and plant growth-promoting bacteria in cowpea: Survival and symbiotic performance

The objective of this study was to evaluate the survival of cowpea during bacterial colonization and evaluate the interrelationship of the Bradyrhizobium sp. and plant growth-promoting bacteria (PGPB) as a potential method for optimizing symbiotic performance and cowpea development. Two experiments using the model legume cowpea cv. “IPA 206” were conducted. In the first experiment, cowpea seeds were disinfected, germinated and transferred to sterilized Gibson tubes containing a nitrogen-free nutritive solution. The experimental design was randomized blocks with 24 treatments [Bradyrhizobium sp. (BR 3267); 22 PGPB; absolute control (AC)] with three replicates. In the second experiment, seeds were disinfected, inoculated according to their specific treatment and grown in Leonard jars containing washed and autoclaved sand. The experimental design was randomized blocks with 24 treatments [BR 3267; 22 BR 3267 + PGPB; AC] with three replicates. Scanning electron microscopy demonstrated satisfactory colonization of the roots of inoculated plants. Additionally, synergism between BR 3267 and PGPB in cowpeas was observed, particularly in the BR 3267 + Paenibacillus graminis (MC 04.21) and BR 3267 + P. durus (C 04.50), which showed greater symbiotic performance and promotion of cowpea development.     

Plant growth-promoting bacteria as inoculants in agricultural soils

Plant-microbe interactions in the rhizosphere are the determinants of plant health, productivity and soil fertility. Plant growth-promoting bacteria (PGPB) are bacteria that can enhance plant growth and protect plants from disease and abiotic stresses through a wide variety of mechanisms; those that establish close associations with plants, such as the endophytes, could be more successful in plant growth promotion. Several important bacterial characteristics, such as biological nitrogen fixation, phosphate solubilization, ACC deaminase activity, and production of siderophores and phytohormones, can be assessed as plant growth promotion (PGP) traits. Bacterial inoculants can contribute to increase agronomic efficiency by reducing production costs and environmental pollution, once the use of chemical fertilizers can be reduced or eliminated if the inoculants are efficient. For bacterial inoculants to obtain success in improving plant growth and productivity, several processes involved can influence the efficiency of inoculation, as for example the exudation by plant roots, the bacterial colonization in the roots, and soil health. This review presents an overview of the importance of soil-plant-microbe interactions to the development of efficient inoculants, once PGPB are extensively studied microorganisms, representing a very diverse group of easily accessible beneficial bacteria.     

Physiological responses and antioxidant enzyme changes in <Emphasis Type="Italic">Sulla coronaria</Emphasis> inoculated by cadmium resistant bacteria

Plant growth promoting bacteria (PGPB) may help to reduce the toxicity of heavy metals on plants growing in polluted soils. In this work, Sulla coronaria inoculated with four Cd resistant bacteria (two Pseudomonas spp. and two Rhizobium sullae) were cultivated in hydroponic conditions treated by Cd; long time treatment 50 µM CdCl₂ for 30 days and short time treatment; 100 µM CdCl₂ for 7 days. Results showed that inoculation with Cd resistant PGPB enhanced plant biomass, thus shoot and root dry weights of control plants were enhanced by 148 and 35% respectively after 7 days. Co-inoculation of plants treated with 50 and 100 µM Cd increased plant biomasses as compared to Cd-treated and uninoculated plants. Cadmium treatment induced lipid peroxidation in plant tissues measured through MDA content in short 7 days 100 µM treatment. Antioxidant enzyme studies showed that inoculation of control plants enhanced APX, SOD and CAT activities after 30 days in shoots and SOD, APX, SOD, GPOX in roots. Application of 50 µM CdCl₂ stimulated all enzymes in shoots and decreased SOD and CAT activities in roots. Moreover, 100 µM of CdCl₂ increased SOD, APX, CAT and GPOX activities in shoots and increased significantly CAT activity in roots. Metal accumulation depended on Cd concentration, plant organ and time of treatment. Furthermore, the inoculation enhanced Cd uptake in roots by 20% in all treatments. The cultivation of this symbiosis in Cd contaminated soil or in heavy metal hydroponically treated medium, showed that inoculation improved plant biomass and increased Cd uptake especially in roots. Therefore, the present study established that co-inoculation of S. coronaria by a specific consortium of heavy metal resistant PGPB formed a symbiotic system useful for soil phytostabilization.