Rice Leaf Sheath Research Articles

Secretome analysis of the rice bacteriumXanthomonas oryzae(Xoo) using in vitro and in planta systems

Xanthomonas oryzae pv. oryzae (Xoo) causes bacterial blight disease in rice, and that severely affects yield loss (upto 50%) of total rice production. Here, we report a proteomics investigation of Xoo (compatible race K3)-secreted proteins, isolated from its in vitro culture and in planta infected rice leaves. 2DE coupled with MALDI-TOF-MS and/or nLC-ESI-MS/MS approaches identified 139 protein spots (out of 153 differential spots), encoding 109 unique proteins. Identified proteins belonged to multiple biological and molecular functions. Metabolic and nutrient uptake proteins were common up to both in vitro and in planta secretomes. However, pathogenicity, protease/peptidase, and host defense-related proteins were highly or specifically expressed during in planta infection. A good correlation was observed between protein and transcript abundances for nine proteins secreted in planta as per semiquantitative RT-PCR analysis. Transgenic rice leaf sheath (carrying PBZ1 promoter::GFP cell death reporter), when used to express a few of the identified secretory proteins, showed a direct activation of cell death signaling, suggesting their involvement in pathogenicity related with secretion effectors. This work furthers our understanding of rice bacterial blight disease, and serves as a resource for possible translation in generating disease resistant rice plants for improved seed yield.

Stimulation of phenolic metabolism by silicon contributes to rice resistance to sheath blight

AbstractSheath blight caused by Rhizoctonia solani is a major disease of rice worldwide. Silicon (Si) can enhance rice resistance to sheath blight, but the relation with phenolic metabolism is poorly known. Two rice cultivars with different levels of resistance to R. solani (resistant Teqing and susceptible Ningjing 1) were studied to determine the effects of Si on disease intensity (rated from 0 to 9) and the involvement of phenolic compounds in disease resistance. Variation in the concentrations of phenolics (including total soluble phenolics, flavonoids, and lignin) and in the activities of defense‐related enzymes polyphenoloxidase (PPO) and phenylalanine ammonia‐lyase (PAL) in rice leaf sheaths was investigated. The results show that Si application reduced sheath‐blight disease ratings of Ningjing 1 and Teqing by 2.96 and 0.65, respectively. In uninoculated plants, Si application alone had no significant effects on the concentrations of phenolic compounds or on the activities of PPO and PAL. In inoculated plants, Si application increased phenolics concentrations and PPO and PAL activities only in the susceptible cultivar Ningjing 1. We conclude that Si‐induced enhancement of phenolic metabolism contributed to the improved resistance of rice to sheath blight in the sensitive cultivar.

Large-Scale Gene Disruption in Magnaporthe oryzae Identifies MC69, a Secreted Protein Required for Infection by Monocot and Dicot Fungal Pathogens

To search for virulence effector genes of the rice blast fungus, Magnaporthe oryzae, we carried out a large-scale targeted disruption of genes for 78 putative secreted proteins that are expressed during the early stages of infection of M. oryzae. Disruption of the majority of genes did not affect growth, conidiation, or pathogenicity of M. oryzae. One exception was the gene MC69. The mc69 mutant showed a severe reduction in blast symptoms on rice and barley, indicating the importance of MC69 for pathogenicity of M. oryzae. The mc69 mutant did not exhibit changes in saprophytic growth and conidiation. Microscopic analysis of infection behavior in the mc69 mutant revealed that MC69 is dispensable for appressorium formation. However, mc69 mutant failed to develop invasive hyphae after appressorium formation in rice leaf sheath, indicating a critical role of MC69 in interaction with host plants. MC69 encodes a hypothetical 54 amino acids protein with a signal peptide. Live-cell imaging suggested that fluorescently labeled MC69 was not translocated into rice cytoplasm. Site-directed mutagenesis of two conserved cysteine residues (Cys36 and Cys46) in the mature MC69 impaired function of MC69 without affecting its secretion, suggesting the importance of the disulfide bond in MC69 pathogenicity function. Furthermore, deletion of the MC69 orthologous gene reduced pathogenicity of the cucumber anthracnose fungus Colletotrichum orbiculare on both cucumber and Nicotiana benthamiana leaves. We conclude that MC69 is a secreted pathogenicity protein commonly required for infection of two different plant pathogenic fungi, M. oryzae and C. orbiculare pathogenic on monocot and dicot plants, respectively.

High activities and mRNA expression of pyrophosphate-fructose-6-phosphate-phosphotransferase and 6-phosphofructokinase are induced as a response to Rhizoctonia solani infection in rice leaf sheaths

Rice sheath blight disease caused by Rhizoctonia solani Kuhn results in significant yield and quality losses in rice growing areas worldwide. The glycolytic pathway is important in the resistance response to R. solani infection in rice. This study examined one of the regulatory steps in this pathway catalyzed by pyrophosphate-fructose-6-phosphate-phosphotransferase (PFP) and 6-phosphofructokinase (PFK). PFP and PFK activity in R. solani-infected rice plants increased. The mRNA expression of PFP/PFK isozymes showed that PFK 1, 2, 4 and 5 in the resistant line at 1 dpi were high as compared to the gradual increase observed in the expression of all PFP isozymes. Also PFK 1, PFK 3, PFK 4, PFK 5, PFP 2 and PFP 5 were adaptive to sheath blight disease infection and linked to defence response while, the expressions of PFK 2, PFP 1, PFP 3 and PFP 4 although adaptive, were not specific to R. solani infection. These observations provide evidence that (a) both PFP and PFK have isozymes that play an adaptive role after R. solani infection but while those of PFK are expressed at higher levels within a short time after infection those of PFP are expressed gradually, (b) the adaptive activation of PFP in R. solani-infected rice plants is correlated with the paired expression of its α- and β-subunits as shown by PFP 2 and PFP 5, and (c) the expression of some α-subunits is not specific to R. solani infection as shown by PFP 1, PFP 3 and PFP 4.

Chemical composition, angiotensin‐converting enzyme‐inhibitory activity and antioxidant activities of few‐flower wild rice (Zizania latifolia Turcz.)

The chemical compositions of the stem and leaf sheath of few-flower wild rice were analysed. In addition, their extracts were evaluated for diphenylpicrylhydrazyl (DPPH) free radical-scavenging activity, ferric-reducing antioxidant power and angiotensin-converting enzyme (ACE)-inhibitory activity, since these are important properties of sources of nutraceuticals or functional foods. The stems contained more ascorbic acid (0.06 g kg(-1) fresh weight), protein (28.18 g kg(-1) dry weight (DW)), reducing sugars (308.54 g kg(-1) DW), water-soluble pectin (20.63 g kg(-1) DW), Na(2) CO(3) -soluble pectin (44.14 g kg(-1) DW), K (8 g kg(-1) dry matter (DM), S (6 g kg(-1) DM) and P (5 g kg(-1) DM) but less starch, total dietary fibre, Si, Na and Ca than the leaf sheaths. The DPPH free radical-scavenging IC(50) values of the stem and leaf sheath extracts were 19.28 and 21.22 mg mL(-1) respectively. In addition, the ACE-inhibitory IC(50) value of the stem extracts was 38.54 mg mL(-1). Both the stem and leaf sheath extracts exhibited good antioxidant properties, while good ACE-inhibitory activity was detected only in the phosphate buffer solution extracts of the stem. Few-flower wild rice could be processed into formula feeds for fish, poultry, etc. or functional foods for persons with high blood pressure.

Exploring the applicability of field measurements of mid-infrared reflectance using a portable band-pass filter type radiometer to assess paddy rice yields

There are few case studies concerning field radiometric measurements in the mid-infrared (MIR) wavelengths for crop plants. This article describes radiometric experiments measuring reflectance in the MIR bands (3.425 ± 0.1 and 3.9 ± 0.1 μm) to evaluate the number of panicles (NP, m−2) of ripening paddy rice using a portable band-pass filter type spectroradiometer. First, laboratory experiments were conducted using containers filled with rice grains of various 1000-grain weights and fresh and dry slashed rice culms plus leaf sheaths, leaf blades and panicles. The intensity of light from the sun or electric bulbs reflected from the grains or plant organs was measured in the MIR, visible (VIS), near-infrared (NIR) and short-wave infrared (SIR) bands. Heavier grains and dense panicles reflected less in the MIR bands than lighter grains and the same occurred in the SIR bands. Field measurements were conducted for several plots of ripening rice where the panicles were partly plucked beforehand. The trial showed that the instrumentation and the method for measuring MIR band reflectance factors were usable for investigating the possibility of MIR measurements for NP.

MoSfl1 Is Important for Virulence and Heat Tolerance in Magnaporthe oryzae

The formation of appressoria, specialized plant penetration structures of Magnaporthe oryzae, is regulated by the MST11-MST7-PMK1 MAP kinase cascade. One of its downstream transcription factor, MST12, is important for penetration and invasive growth but dispensable for appressorium formation. To identify additional downstream targets that are regulated by Pmk1, in this study we performed phosphorylation assays with a protein microarray composed of 573 M. oryzae transcription factor (TF) genes. Three of the TF genes phosphorylated by Pmk1 in vitro were further analyzed by coimmunoprecipitation assays. One of them, MoSFL1, was found to interact with Pmk1 in vivo. Like other Sfl1 orthologs, the MoSfl1 protein has the HSF-like domain. When expressed in yeast, MoSFL1 functionally complemented the flocculation defects of the sfl1 mutant. In M. oryzae, deletion of MoSFl1 resulted in a significant reduction in virulence on rice and barley seedlings. Consistent with this observation, the Mosfl1 mutant was defective in invasive growth in penetration assays with rice leaf sheaths. In comparison with that of vegetative hyphae, the expression level of MoSFL1 was increased in appressoria and infected rice leaves. The Mosfl1 mutant also had increased sensitivity to elevated temperatures. In CM cultures of the Mosfl1 and pmk1 mutants grown at 30°C, the production of aerial hyphae and melanization were reduced but their growth rate was not altered. When assayed by qRT-PCR, the transcription levels of the MoHSP30 and MoHSP98 genes were reduced 10- and 3-fold, respectively, in the Mosfl1 mutant. SFL1 orthologs are conserved in filamentous ascomycetes but none of them have been functionally characterized in non-Saccharomycetales fungi. MoSfl1 has one putative MAPK docking site and three putative MAPK phosphorylation sites. Therefore, it may be functionally related to Pmk1 in the regulation of invasive growth and stress responses in M. oryzae.

Relationships between exogenous-toxin quantity and increased biomass of transgenic Bt crops under elevated carbon dioxide

Field-OTC experiments were conducted with the goals of ascertaining if increased biomass in Bt transgenic cotton and rice grown under elevated CO 2 results in diminished exogenous-Bt toxin, and assessing the effectiveness of Bt transgenes against lepidopteran pests. Bt cotton responded differently, in terms of Bt-toxin quantity, than Bt rice, and both indicated differences among developmental stages. Dramatic biomass increase significantly diluted Bt-toxin content in 45-DAS (“days after seedling”) petioles and shoots and 90-DAS Bt cotton squares, and in the 50-DAS tissues and 100-DAS leaf sheaths of Bt rice. Moreover, the dilution effect was partially responsible for decreased Bt-toxin in these tissues, but not responsible for significant decreases in Bt-toxin in 90-DAS Bt cotton leaves and bolls. Furthermore, elevated CO 2 significantly affected the fitness and performance of Chilo suppressalis, and the susceptible and resistant colonies of Helicoverpa armigera, although adversely affected Bt-gene expression for the transgenic cotton and rice.

High activities and mRNA expression of pyrophosphate-fructose-6-phosphate-phosphotransferase and 6-phosphofructokinase are induced as a response to Rhizoctonia solani infection in rice leaf sheaths

High activities and mRNA expression of pyrophosphate-fructose-6-phosphate-phosphotransferase and 6-phosphofructokinase are induced as a response to Rhizoctonia solani infection in rice leaf sheaths

Microbial community profiles in intercellular fluid of rice

Plants harbor microorganisms that are thought to stimulate plant defense systems or promote plant growth. Individual species in these intercellular microbial communities are often not sufficiently abundant to be easily described, although some endophytic microorganisms amenable to culture have been characterized. To better understand the microbial population of plants, we collected intercellular fluid (IF) from leaf blades and sheaths of rice and subsequently isolated DNA from the IF. Denaturing gradient gel electrophoresis (DGGE) analysis of 16S and 18S rDNA fragments amplified from IF DNA by PCR indicated that these band patterns were distinguishable from those of a leaf surface-wash fluid (SF). Analysis of a set of rDNA fragments amplified from IF DNA of rice with different genotypes, paddies or growth stages for the primary survey of overall microbial community in the IF suggested that this approach is suitable for analyzing microbial diversity in the IF from various plant samples. Actually, comparative analysis of amplified rDNA fragments of rice and other five plant species indicated that the microbial diversity in IF is likely to vary substantially among plant species. We can also use sequence analysis of 16S rDNA fragments amplified from rice IF DNA to identify species including unculturable bacteria and proteobacteria and Xanthomonas and 18S rDNA fragments to identify Tilletiaria anomala, Tilletia iowensis, Ustilago maydis and unculturable eukaryotes. Thus, IF DNA analysis seems to be a good tool to further study the microbial ecology of plants.

Relationship Between Lodging Resistance and Chemical Contents in Culms and Sheaths of Japonica Rice During Grain Filling

To understand the relationship between lodging resistance and chemical component contents in culms and leaf sheaths of rice, the physical strength and maximum bearing capacity of culm, and the contents and amounts of potassium (K), silicon (Si) and soluble sugars in culms and leaf sheaths were investigated using four japonica rice varieties with different lodging resistance characteristics during grain filling. There were significant differences in the total amounts of K, Si and soluble sugars in culms and leaf sheaths among the tested rice varieties. The difference in the total amount of Si was greater than that of K or soluble sugars. The physical strength and maximum bearing capacity of culm continuously decreased from heading to ripening, with a rapid decrease at the dough stage. However, the contents and total amounts of K and Si in culms and the Si content in leaf sheaths gradually increased and an accumulation of K and Si in culms was exhibited, whereas the content and total amount of K and the total amount of Si in leaf sheaths gradually decreased and an exportation of K and Si in leaf sheaths was presented. The physical strength was positively and significantly correlated with the total amounts of K and Si in culms during grain filling except that at the heading stage, the total amount of soluble sugars in culms at the heading and milky stages, the total amounts of Si and soluble sugars in leaf sheaths at the heading stage, the total amount of K in leaf sheaths at the heading and milky stages, and the maximum bearing capacity during grain filling. It is suggested that the lodging resistance of japonica rice would be improved by increasing the amount of soluble sugars in plants at the early filling stage, and enhancing the amount of Si in plants during grain filling through topdressing Si fertilizer at the early filling stage.

Transcriptional analysis of molecular interactions between Pseudomonas fluorescens strain TDK1, Oryza sativa and Cnaphalocrocis medinalis

AbstractRice leaffolder Cnaphalocrocis medinalis is a devastating insect pest posing major threat to rice cultivation. Plant growth‐promoting rhizobacteria Pseudomonas fluorescens strain TDK1 was found to induce high degree of resistance in rice plants against leaffolder insect through Induced Systemic Resistance in our previous studies. Thus the present study was aimed to understand the molecular mechanisms involved in P. fluorescens TDK1‐mediated host plant resistance against leaffolder infestation on rice. Transcript profiling was done through DD‐RT‐PCR method in leaffolder‐infested rice plants that were treated with and without Pf TDK1. A total of 3500 transcripts were displayed, of which 165 transcripts exhibited differential expression during rice–Pf TDK1–leaffolder interactions. The differential transcripts were 58 in Pf TDK1‐treated + leaffolder‐challenged, 51 in leaffolderinfested and 56 in Pf TDK1‐treated rice leaf sheath tissues. Annotation and mapping of differentially expressed genes onto metabolic pathways revealed the interaction of leaffolder and Pf TDK1 in rice plant at the molecular level. Furthermore, Pf TDK1 induced over‐expression of genes found to regulate the diverse metabolic processes by the selective integration of defensive signals in rice plants.

EGFP-Rhm51 foci enable the visualization and enumeration of DNA double-strand breaks in Magnaporthe oryzae

To detect and count DNA double-strand breaks (DSBs) during the life cycle of Magnaporthe oryzae, we constructed an expression vector for EGFP-Rhm51 fusion protein and introduced it into strain Ina86-137. EGFP-Rhm51 foci were detected, and the number of foci in mycelia treated with mitomycin C, which induces DSBs, was higher than in untreated samples, indicating that the foci enabled the visualization of DSBs during the life cycle. EGFP-Rhm51 foci were observed at all stages of the asexual cycle including when invasive hyphae formed in an intact rice leaf sheath, demonstrating that M. oryzae undergoes DSBs during vegetative and parasitic growth.

The Basic Leucine Zipper Transcription Factor Moatf1 Mediates Oxidative Stress Responses and Is Necessary for Full Virulence of the Rice Blast Fungus Magnaporthe oryzae

Magnaporthe oryzae is the causal agent of rice blast disease, leading to enormous losses of rice production. Here, we characterized a basic leucine zipper (bZIP) transcription factor, Moatf1, in M. oryzae, a homolog of Schizosaccharomyces pombe ATF/CREB that regulates the oxidative stress response. Moatf1 deletion caused retarded vegetative growth of mycelia, and the Moatf1 mutant exhibited higher sensitivity to hydrogen peroxide (H(2)O(2)) than did the wild-type strain. The mutant showed severely reduced activity of extracellular enzymes and transcription level of laccases and peroxidases and exhibited significantly reduced virulence on rice cultivar CO-39. On rice leaf sheath, most of the infectious hyphae of the mutant became swollen and displayed restricted growth in primary infected cells. Defense response was strongly activated in plants infected by the mutant. Diamino benzidine staining revealed an accumulation of H(2)O(2) around Moatf1 mutant appressoria and rice cells with Moatf1 hyphae that was absent in the wild type. Inhibition of the plant NADPH oxidase by diphenyleneiodonium prevented host-derived H(2)O(2) accumulation and restored infectious hyphal growth of the mutant in rice cells. Thus, we conclude that Moatf1 is necessary for full virulence of M. oryzae by regulating the transcription of laccases and peroxidases to impair reactive oxygen species-mediated plant defense.

Identification of gibberellin acid-responsive proteins in rice leaf sheath using proteomics

The phytohormone gibberellin acid (GA) controls many aspects of plant development. In this study, we identified proteins that are differentially expressed between the rice (Oryza sativa L.) GA-deficient cultivar, Aijiaonante, and its parental line, Nante. Proteins were extracted from rice leaf sheath and examined by 2DGE. Among more than 1200 protein spots reproducibly detected on each gel, 29 were found to be highly up-regulated by GAs in Nante, and 6 were down-regulated by GAs in Aijiaonante. These 35 proteins were identified by MALDI-TOF MS and were classified into three groups based on their putative function in metabolism, stress/defense processes and signal transduction. These data suggest that metabolic pathways are the main target of regulation by GAs during rice development. Our results provide new information about the involvement of GAs in rice development.

Characterization of β-tubulin 4 regulated by gibberellins in rice leaf sheath

Tubulins are basic components of microtubules and are encoded by a multigene family in eukaryotes. The expression of OsTUB4, one of eight β-tubulin isotypes identified in the rice genome, was characterized. OsTUB4 was expressed in root primodia and the shoot apical meristem in basal parts of the leaf sheath in rice seedlings. OsTUB4 transcript abundance in leaf sheath increased by treatment with gibberellic acid (GA3) in a dose- and time-dependent manner. OsTUB4 transcript levels in gibberellin (GA)-deficient mutants were less than those of wild type rice. An OsTUB4 promoter::GUS assay also confirmed the responsiveness of OsTUB4 to exogenous GA3, suggesting that OsTUB4 expression was regulated by GA and may be involved in GA-regulated leaf sheath growth. In addition, OsTUB4 could interact with different specific proteins in vitro as assayed by a yeast two-hybrid system, indicating that OsTUB4 may have diverse functions through interaction with different proteins.

The Supernatant of a Conidial Suspension of Magnaporthe oryzae Contains a Factor that Promotes the Infection of Rice Plants

AbstractMany factors produced by the pathogen Magnaporthe oryzae enhance its ability to infect rice. We found a novel infection‐promoting activity in the supernatant of a conidia suspension (SCS) of M. oryzae. The addition of SCS promoted the invasion of excised rice leaf sheaths by infectious hyphae. The activity was heat‐stable and was found in SCSs from five virulent and three avirulent isolates of M. oryzae on the rice cv. Nipponbare (Pia). The effect was exclusively detected in compatible interactions. The infection of rice plants by non‐rice blast fungi was not enhanced by SCS. These results suggest that SCS includes a heat‐stable factor(s) that promotes M. oryzae infection during compatible interactions.

Understanding rice plant resistance to the Brown Planthopper (Nilaparvata lugens): A proteomic approach

Engineering and breeding resistant plant varieties are the most effective and environmentally friendly ways to control agricultural pests and improve crop performance. However, the mechanism of plant resistance to pests is poorly understood. Here we used a quantitative mass-spectrometry-based proteomic approach for comparative analysis of expression profiles of proteins in rice leaf sheaths in responses to infestation by the brown planthopper (Nilaparvata lugens Stål, BPH), which is a serious rice crop pest. Proteins involved in multiple pathways showed significant changes in expression in response to BPH feeding, including jasmonic acid synthesis proteins, oxidative stress response proteins, beta-glucanases, protein; kinases, clathrin protein, glycine cleavage system protein, photosynthesis proteins and aquaporins. The corresponding genes of eight important proteins were further analyzed by quantitative RT-PCR. Proteomic and transcript responses that were related to wounding, oxidative and pathogen stress overlapped considerably between BPH-resistant (carrying the resistance gene BPH15) and susceptible rice lines. In contrast, proteins and genes related to callose metabolism remained unchanged and glycine cleavage system protein was up-regulated in the BPH-resistant lines, indicating that they have an efficient and specific defense mechanism. Our results provide new information about the interaction between rice and the BPH.

The QTL Analysis of RuBisCO in Flag Leaves and Non-Structural Carbohydrates in Leaf Sheaths of Rice Using Chromosome Segment Substitution Lines and Backcross Progeny F2 Populations

In rice (Oryza sativa L.), the maintenance of high photosynthetic rate of flag leaves and the carbon remobilization from leaf sheaths after heading is a critical physiological component affecting the yield. To clarify the genetic basis of RuBisCO content of the flag leaf, a major determinant of photosynthetic rate, and non-structural carbohydrate (NSC) concentration in the third leaf sheath at heading, we carried out quantitative trait loci (QTL) analysis with 39 Koshihikari/Kasalath chromosome segment substitution lines (CSSLs) and backcross progeny F2 population derived from target CSSL holding the QTL/Koshihikari in the field. QTLs for RuBisCO content and NSC concentration at heading were detected between R2447-C1286 and R2447-R716 on chromosome 10, respectively, by comparing Koshihikari with four CSSLs for chromosome 10 (SL-229, -230, -231 and -232). The progeny QTL for RuBisCO content and for NSC concentration at heading qRCH-10 and qNSCLSH-10-1, respectively, were detected at similar marker intervals between RM8201 and RM5708. In addition, QTLs for RuBisCO content at 14 d after heading, qRCAH-10-1 and qRCAH-10-2, were detected in regions different from that of qRCH-10. No QTL for NSC concentration at 14 d after heading was detected between RM8201 and R716, the region analyzed in this study. The QTLs qRCH-10 and qRCAH-10-1 for RuBisCO content would have additive effects. These QTLs for RuBisCO content and NSC concentration newly found using CSSLs and their backcross progeny F2 population should be useful for better understanding the genetic basis of source and temporary-sink functions in rice and for genetic improvement of Koshihikari in terms of their functions.

Understanding the molecular basis of plant growth promotional effect of Pseudomonas fluorescens on rice through protein profiling.

BackgroundPlant Growth Promoting Rhizobacteria (PGPR), Pseudomonas fluorescens strain KH-1 was found to exhibit plant growth promotional activity in rice under both in-vitro and in-vivo conditions. But the mechanism underlying such promotional activity of P. fluorescens is not yet understood clearly. In this study, efforts were made to elucidate the molecular responses of rice plants to P. fluorescens treatment through protein profiling. Two-dimensional polyacrylamide gel electrophoresis strategy was adopted to identify the PGPR responsive proteins and the differentially expressed proteins were analyzed by mass spectrometry.ResultsPriming of P. fluorescens, 23 different proteins found to be differentially expressed in rice leaf sheaths and MS analysis revealed the differential expression of some important proteins namely putative p23 co-chaperone, Thioredoxin h- rice, Ribulose-bisphosphate carboxylase large chain precursor, Nucleotide diPhosphate kinase, Proteosome sub unit protein and putative glutathione S-transferase protein.ConclusionFunctional analyses of the differential proteins were reported to be directly or indirectly involved in growth promotion in plants. Thus, this study confirms the primary role of PGPR strain KH-1 in rice plant growth promotion.