Rhizosphere Soil Bacterial Community Structure Research Articles

Sesame bacterial wilt significantly alters rhizosphere soil bacterial community structure, function, and metabolites in continuous cropping systems

Bacterial wilt is the leading disease of sesame and alters the bacterial community composition, function, and metabolism of sesame rhizosphere soil. However, its pattern of change is unclear. Here, the purpose of this study was to investigate how these communities respond to three differing severities of bacterial wilt in mature continuously cropped sesame plants by metagenomic and metabolomic techniques, namely, absence (WH), moderate (WD5), and severe (WD9) wilt. The results indicated that bacterial wilt could significantly change the bacterial community structure in the rhizosphere soil of continuously cropped sesame plants. The biomarker species with significant differences will also change with increasing disease severity. In particular, the gene expression levels of Ralstonia solanacearum in the WD9 and WD5 treatments increased by 25.29% and 33.61%, respectively, compared to those in the WH treatment (4.35 log10 copies g-1). The occurrence of bacterial wilt significantly altered the functions of the bacterial community in rhizosphere soil. KEEG and CAZy functional annotations revealed that the number of significantly different functions in WH was greater than that in WD5 and WD9. Bacterial wilt significantly affected the relative content of metabolites, especially acids, in the rhizosphere soil, and compared with those in the rhizosphere soil from WH, 10 acids (including S-adenosylmethionine, N-acetylleucine, and desaminotyrosine, etc.) in the rhizosphere soil from WD5 or WD9 significantly increased. In comparison, the changes in the other 10 acids (including hypotaurine, erucic acid, and 6-hydroxynicotinic acid, etc.) were reversed. The occurrence of bacterial wilt also significantly inhibited metabolic pathways such as ABC transporter and amino acid biosynthesis pathways in rhizosphere soil and had a significant impact on two key enzymes (1.1.1.11 and 2.6.1.44). In conclusion, sesame bacterial wilt significantly alters the rhizosphere soil bacterial community structure, function, and metabolites. This study enhances the understanding of sesame bacterial wilt mechanisms and lays the groundwork for future prevention and control strategies against this disease.

Response of bacterial community characteristics in the rhizosphere soil of Stellera chamaejasme L. to its expansion on the Qinghai‐Tibet Plateau

AbstractQinghai‐Tibet Plateau is facing a serious environmental and ecological problem of Meadow degradation. Toxic weed invasion is a typical characteristic of grassland degradation. Soil microbial community composition is sensitive to environmental changes; however, the effects of poisonous weed expansion on soil bacterial communities are unclear. Here, we investigated the effects of Stellera chamaejasme L. expansion on the rhizosphere soil bacterial community structure and function using high‐throughput sequencing. The results showed that expansion of Stellera chamaejasme L. changed soil nitrogen(e.g., total nitrogen [TN, −39.02%], available nitrogen [AN, −32.95%]) and other soil nutrients. Redundancy analysis (RDA) and Variance partitioning analysis (VPA) showed that soil nutrients changed, leading to significant changes in the bacterial community structure. The expansion of Stellera chamaejasme L significantly reduced its rhizosphere bacterial alpha diversity, and the beta diversity had significant differences (p < 0.05). Principal coordinates analysis (PCoA) and analysis of similarity (ANOSIM) indicated that the expansion caused significant variations in the rhizosphere bacterial community (R = 0.7037, p < 0.01). The linear discriminant analysis (LDA) effect size (LEfSe) analysis identified 23 biomarkers, most of which were Proteobacteria, indicating that bacteria involved in soil nutrient cycling were better able to survive in the alpine grassland. The Biolog EcoPlate method was used to determine the soil microbial metabolic capacity in different S. chamaejasme expansions. The result showed that heavy expansion had higher carbon source usage ability and microbial diversity index values. Furthermore, it was also found that heavy expansion improved the usage rate of amino acid carbon sources. Tax4Fun prediction analysis further indicated that carbohydrate metabolism, amino acid metabolism, and membrane transport were central metabolic pathways of rhizosphere soil bacteria. Our study found that Stellera chamaejasme L. changed rhizosphere soil nutrient and bacterial community structure during expansion and helped it tolerate harsh conditions by enriching bacterial communities actively involved in carbon and nitrogen metabolism and promoting plant growth. These findings provided evidence to propose effective restoration measures for poisonous grassland degradation.

The Effects of Coexisting Elements (Zn and Ni) on Cd Accumulation and Rhizosphere Bacterial Community in the Soil-Tomato System

The increasing cadmium (Cd) levels in agricultural soils have become a worldwide concern for food crop security. Cd accumulation in the soil-plant system is closely related to other coexisting factors. In this study, the effects of different levels of Zn or Ni on Cd accumulation in tomato plants and on the rhizosphere soil bacterial community structure were analyzed by coupling pot experiments with high-throughput sequencing. The results demonstrated that tomato plants (Lycopersicon esculentum) in Zn-Cd and Ni-Cd co-contaminated soils exhibited lower relative growth rates. Co-contamination at low levels tended to reduce the bioaccumulation of heavy metals in the roots of plants, whereas increased contaminant concentrations produced the opposite effect. In the presence of 200 mg/kg Zn or 20 mg/kg Ni, the biomass of plant roots increased by 4.95–23.16% and the Cd content of the plant roots decreased by 17.36–68.93% due to the antagonistic effects between Cd and Zn/Ni. In addition, the richness and diversity of the bacterial community were significantly altered under HMs co-contamination, and the number of special bacteria was positively correlated with the level of heavy metals in the rhizosphere soil. The relative abundance of Proteobacteria increased and that of Actinobacteria decreased in soils with low levels of heavy metals. This may improve the tolerance of plant roots to heavy metals and reduce the accumulation of Cd in plant roots. These findings highlight the important role of coexisting elements in the inhibition of Cd accumulation in tomatoes and offer important information for the production of safe crops.

Effects of Irrigation Using Activated Brackish Water on the Bacterial Community Structure of Rhizosphere Soil

The technology for the activation of irrigation brackish water was successful at improving the soil environment and crop yields. However, very few studies have examined how activated brackish water irrigation affects the rhizosphere bacterial communities and network patterns. By combining 16S rRNA high-throughput sequencing, we evaluated the diversity and composition of the rhizosphere bacterial community after being subjected to different types of irrigation with activated brackish water. We also analyzed the correlation and co-occurrence networks among the bacterial diversity, composition, and rhizosphere soil properties. The results showed that compared with brackish water irrigation, the salt content of activated irrigation water significantly decreased by 9.35%, 9.83%, and 12.95%, respectively. Irrigation with different types of activated brackish water had no significant (p > 0.05) effect on the diversity of the rhizosphere bacterial community, but it significantly changed its community composition, which was primarily dominated by soil nutrient indicators. The soil total nitrogen (TN) showed a significant (p < 0.01) negative correlation with the Chao1 index. Additionally, the changes in bacterial communities under different types of activated brackish irrigation water mainly occurred at the genus level. We showed that the rhizosphere soil that had been irrigated with oxygenated brackish water and magnetized brackish water better supported the reproduction of some soil-borne pathogens. Magnetization-oxygenation coupling treatment could significantly reduce the colonization of soil-borne pathogens of the rhizosphere soil, while also favoring the function of functional bacteria involved in soil nutrient transformation. This study highlights the main factors affecting the rhizosphere soil bacterial community structure by activated brackish water irrigation, while also providing new technical support for brackish water irrigation.

Root exudates and rhizosphere soil bacterial relationships of Nitraria tangutorum are linked to k-strategists bacterial community under salt stress.

When plants are subjected to various biotic and abiotic stresses, the root system responds actively by secreting different types and amounts of bioactive compounds, while affects the structure of rhizosphere soil bacterial community. Therefore, understanding plant–soil-microbial interactions, especially the strength of microbial interactions, mediated by root exudates is essential. A short-term experiment was conducted under drought and salt stress to investigate the interaction between root exudates and Nitraria tangutorum rhizosphere bacterial communities. We found that drought and salt stress increased rhizosphere soil pH (9.32 and 20.6%) and electrical conductivity (1.38 and 11 times), respectively, while decreased organic matter (27.48 and 31.38%), total carbon (34.55 and 29.95%), and total phosphorus (20 and 28.57%) content of N. tangutorum rhizosphere soil. Organic acids, growth hormones, and sugars were the main differential metabolites of N. tangutorum under drought and salt stress. Salt stress further changed the N. tangutorum rhizosphere soil bacterial community structure, markedly decreasing the relative abundance of Bacteroidota as r-strategist while increasing that of Alphaproteobacteria as k-strategists. The co-occurrence network analysis showed that drought and salt stress reduced the connectivity and complexity of the rhizosphere bacterial network. Soil physicochemical properties and root exudates in combination with salt stress affect bacterial strategies and interactions. Our study revealed the mechanism of plant–soil-microbial interactions under the influence of root exudates and provided new insights into the responses of bacterial communities to stressful environments.

Differences in Bacterial Community Structure in Rhizosphere Soil of Three Caragana Species and Its Driving Factors in a Common Garden Experiment

The soil bacterial diversity and community structures in rhizosphere soil of Caragana microphylla, Caragana liouana, and Caragana roborovskyi in a common garden experiment were measured using the high-throughput sequencing technique, with the aim of investigating the factors driving the variation in the bacterial community structure. The results indicated that 42 phyla, 55 classes, 123 orders, 244 families, and 558 genera were obtained from the rhizosphere soil. The dominant phyla in all sample sites were Proteobacteria, Cyanobacteria, Actinobacteria, Bacteroidetes, Firmicutes, and Acidobacteria (relative abundance>1%). At the genus level, Phenylobacterium, Ensifer, and Chitinophaga were dominant. Two-way analysis of variance showed that species had a significant effect on the Shannon index and Simpson index of rhizosphere soil bacteria of the three Caragana species, whereas the Chao1 index, Shannon index, and Simpson index were significantly affected by the interaction of provenances and species. There was a significant difference among the three species in the composition of bacterial communities, and the cluster analysis indicated that the composition of the soil bacterial community significantly differed among provenances in C. liouana and C. roborovskyi. Based on the redundancy analysis, mean annual precipitation and altitude were the dominant factors influencing the rhizosphere soil bacterial community structure. Overall, the present results indicated that there were intraspecific and interspecific differences in the diversity and community structures of rhizosphere soil bacteria, and the bacterial community structure was mainly affected by the provenance climate. These results provide a theoretical basis for understanding the adaptation strategies and ecological restoration of the three Caragana species.

Microplastic particles alter wheat rhizosphere soil microbial community composition and function

The wide existence of microplastics (MPs) in the terrestrial systems is proved by –many studies, and their presence could potentially change the soil chem-physical properties and processes. Various types of microplastics may have different behaviors, inducing distinct effects on the soil ecosystems. However, the knowledge of microplastic impacts on rhizosphere soil bacterial community structure is limited. In our study, three types of microplastics, i.e., polyethylene (PE), polyvinylchloride (PVC) and polystyrene (PS), with the same particle size (200 µm) and concentration (2%) were used to investigate their influences on the rhizosphere soil bacterial communities. Results revealed that the alpha diversities (richness, evenness and diversity) of microbiota in the rhizosphere soil were variously decreased by the microplastics, especially the PE MPs. The relative abundance of some various phyla and genera related to pollution degradation was miscellaneously increased, indicating that the MPs with different characterizations may have miscellaneous biodegradation pathways. Moreover, the PICRUSt2 analysis demonstrated that PS decreased most functional category levels and led to a decrease of bacterial genus number, however, PE and PVC improved metabolic pathways and xenobiotics biodegradation and metabolism. Our findings offer important knowledge of how the microplastics with different characterizations influence rhizosphere soil bacterial communities and their related function.

Effects of Cr Stress on Rhizosphere Soil Bacterial Community Structure and Diversity of Iris Pseudacorus

Effects of Cr Stress on Rhizosphere Soil Bacterial Community Structure and Diversity of Iris Pseudacorus

Rhizosphere bacteria community and functions under typical natural halophyte communities in North China salinized areas.

Soil salinity is a serious environmental issue in arid China. Halophytes show extreme salt tolerance and are grow in saline-alkaline environments. There rhizosphere have complex bacterial communities, which mediate a variety of interactions between plants and soil. High-throughput sequencing was used to investigated rhizosphere bacterial community changes under the typical halophyte species in arid China. Three typical halophytes were Leymus chinensis (LC), Puccinellia tenuiflora (PT), Suaeda glauca (SG). The dominant phyla were Proteobacteria, Actinobacteria, Chloroflexi, Gemmatimonadetes, Acidobacteria and Bacteroidetes, Suaeda glauca rhizosphere has stronger enrichment of Nitrospirae and Cyanobacteria. The Ace, Chao and Shannon indices were significantly higher in soils under LC and SG (P<0.05). Functional predictions, based on 16S rRNA gene by PICRUSt, indicated that Energy metabolism, Amino acid metabolism, Carbohydrate metabolism and Fatty acid metabolism are dominant bacterial functions in three halophytes rhizosphere soil. Carbon metabolism, Oxidative phosphorylation, Methane metabolism, Sulfur metabolism and Nitrogen metabolism in SG were significantly higher than that in LC and PT. Regression analysis revealed that rhizosphere soil bacterial community structure is influenced by soil organic matter (SOM) and soil water content (SWC), while soil bacterial community diversity is affected by soil pH. This study contributes to our understanding of the distribution characteristics and metabolic functions under different halophyte rhizosphere bacterial communities, and will provide references for the use of rhizosphere bacteria to regulate the growth of halophytes and ecological restoration of saline soil.

Fresh Compost Tea Application Does Not Change Rhizosphere Soil Bacterial Community Structure, and Has No Effects on Soybean Growth or Yield.

Soil bacteria drive key ecosystem functions, including nutrient mobilization, soil aggregation and crop bioprotection against pathogens. Bacterial diversity is thus considered a key component of soil health. Conventional agriculture reduces bacterial diversity in many ways. Compost tea has been suggested as a bioinoculant that may restore bacterial community diversity and promote crop performance under conventional agriculture. Here, we conducted a field experiment to test this hypothesis in a soybean-maize rotation. Compost tea application had no influence on bacterial diversity or community structure. Plant growth and yield were also unresponsive to compost tea application. Combined, our results suggest that our compost tea bacteria did not thrive in the soil, and that the positive impacts of compost tea applications reported elsewhere may be caused by different microbial groups (e.g., fungi, protists and nematodes) or by abiotic effects on soil (e.g., contribution of nutrients and dissolved organic matter). Further investigations are needed to elucidate the mechanisms through which compost tea influences crop performance.

Variation in physicochemical properties and bacterial community structure in rhizosphere soil of Coptis chinensis tow cropping modes

The natural forest and artificial shed are the main cropping modes of Coptis chinensis. This study is aimed to reveal the rhizosphere soil bacterial community structure difference between under tow C. chinensis cropping modes-natural forest and artificial shed, and to assist us to completely understand soil quality condition,and provide theoretical guidance for soil improvement and C. chinensis planting. The rhizosphere soil samples of 1-5-year-old C. chinensis under tow cropping modes-natural forest and artificial shed were collected. Illumina high-throughput sequencing technology was used to analyze the alpha diversity, community composition, community structure of soil bacteria under the tow cropping modes,and the effects of soil nutriment indices on soil bacterial community structure. Through the analysis of species number, Shannon, Chao1 index and ACE index of bacterial community, it was found that the bacterial diversity of 1-year-old C. chinensis soil under natural forest cropping mode was significantly lower than that under artificial shed cropping mode, and the diversity of bacterial communities in soil of 2-5-years old C. chinensis were not significant different between two cropping modes. A total of 53 phyla,60 classes,140 orders and 266 families were detected in the rhizosphere soil of C. chinensis under the cropping modes of natural forest, respectively. The rhizosphere soil of C. chinensis under the cropping modes of artificial shed included 54 phyla,65 classes,140 orders and 264 families, respectively. Under the two cropping modes, the top 10 dominant species of bacterial community abundance are the same, they are Proteobacteria, Acidobacteria, Actinobacteria,Bacteroidetes, Planctomycetes, Chloroflexi, Verrucomicrobia, Gemmatimonadetes, Firmicutes and Cyanobacteria, but there are differences in the abundance sequence. The top 10 dominant species of bacterial community abundance accounted for 74.36% to 74.30% of the total bacteria, and 3.15% to 3.92% of the bacteria are unclassified. The results of Metastat analysis showed that the abundance of Gemmatimonadetes in the rhizosphere soil of C. chinensis under the cropping modes the artificial shed was significantly higher than that under the natural forest cropping mode(P&lt;0.05). MRPP analysis of community structure differences showed that under tow cropping modes, there were significant differences in the bacterial community structure of 1-4-year-old soil bacteria, among which the difference between 1-year-old soil samples was the largest. With the increase of cropping years, the difference gradually decreases, and there is no significant difference in the bacterial community structure between 5-year-old soil samples. RDA analysis and correlation analysis of bacterial community structure and soil physical and chemical properties showed that the order of environmental factors on the rhizosphere soil bacteria of Coptis chinensis was: pH&gt;available P&gt; total P&gt; total K&gt;bulk density&gt;total N&gt;available N&gt;organic matter. The results are helpful to understand the soil health of C. chinensis and provide scientific basis and theoretical guidance for soil improvement and C. chinensis planting.

Effects of Dark Septate Endophytes Strain A024 on Damping-off Biocontrol, Plant Growth and the Rhizosphere Soil Enviroment of Pinus sylvestris var. mongolica Annual Seedlings.

Dark septate endophytes (DSEs) exert a vital role in promoting plant growth, improving mineral absorption, biological disease control, and enhancing plant stress resistance. The effects of dark septate endophyte strain, Phialocephala bamuru A024 on damping-off biocontrol, plant development, nutrients within the rhizosphere soil, as well as bacterial communities in the annual seedlings of P. sylvestris var. Mongolica were studied. According to our findings, following P. bamuru A024 inoculation, the damping-off disease morbidity decreased significantly compared with control, some physiological indices such as β-1,3-glucanase, chitinase enzyme activity as well as a soluble protein and proline content in P. sylvestris var. mongolica were elevated under R. solani stress. After inoculation with P. bamuru A024, the biomass in seedlings, nutrients in soil, root structure index, together with activities of soil enzymes were remarkably up-regulated relative to control (p < 0.05). As suggested by the results of high-throughput sequencing, the microbial structure in the rhizosphere soil of the P. sylvestris var. mongolica showed significant differences (p < 0.05) after P. bamuru A024 inoculation compared to control treatment and the rhizosphere soil bacterial community structure after DSE A024 inoculation was positively correlated to the main soil nutrition indices.

Effects of salt and drought stresses on rhizosphere soil bacterial community structure and peanut yield

A pot experiment with Huayu 25 as experimental material was conducted, with treatments of drought and salt stresses. The effects of drought and salt stresses at the flowering stage on the plant morphology, pod yield, and soil bacterial community structure in the rhizosphere were examined. The results showed that Proteobacteria, Actinobacteria, Saccharibacteria, Chloroflexi, Cyanobacteria, and Acidobacteria were the dominant phyla in the rhizosphere soil of peanut. Compared with that under normal conditions, the relative abundance of Proteobacteria and Actinobacteria dramatically decreased, while that of Cyanobacteria evidently increased in drought-treated and salt-treated soil. Moreover, the variation of Cyanobacteria abundance caused by combined drought and salt stresses was stronger than that caused by single drought or salt stress. Functional meta-genomic profiling indicated that a series of sequences related to signaling transduction, defense mechanism and post-translational modification, protein turnover, chaperones were enriched in rhizosphere soil under stressed conditions, which might have implications for plant survival and stress tolerance. Drought and salt stress affectedpeanut growth and reduced pod yield. Results from this study would present reference on the future improvement of stress tolerance of peanuts via modifying soil microbial community.

Effects of shade stress on turfgrasses morphophysiology and rhizosphere soil bacterial communities

BackgroundThe shade represents one of the major environmental limitations for turfgrass growth. Shade influences plant growth and alters plant metabolism, yet little is known about how shade affects the structure of rhizosphere soil microbial communities and the role of soil microorganisms in plant shade responses. In this study, a glasshouse experiment was conducted to examine the impact of shade on the growth and photosynthetic capacity of two contrasting shade-tolerant turfgrasses, shade-tolerant dwarf lilyturf (Ophiopogon japonicus, OJ) and shade-intolerant perennial turf-type ryegrass (Lolium perenne, LP). We also examined soil-plant feedback effects on shade tolerance in the two turfgrass genotypes. The composition of the soil bacterial community was assayed using high-throughput sequencing.ResultsOJ maintained higher photosynthetic capacity and root growth than LP under shade stress, thus OJ was found to be more shade-tolerant than LP. Shade-intolerant LP responded better to both shade and soil microbes than shade-tolerant OJ. The shade and live soil decreased LP growth, but increased biomass allocation to shoots in the live soil. The plant shade response index of LP is higher in live soil than sterile soil, driven by weakened soil-plant feedback under shade stress. In contrast, there was no difference in these values for OJ under similar shade and soil treatments. Shade stress had little impact on the diversity of the OJ and the LP bacterial communities, but instead impacted their composition. The OJ soil bacterial communities were mostly composed of Proteobacteria and Acidobacteria. Further pairwise fitting analysis showed that a positive correlation of shade-tolerance in two turfgrasses and their bacterial community compositions. Several soil properties (NO3−-N, NH4+-N, AK) showed a tight coupling with several major bacterial communities under shade stress. Moreover, OJ shared core bacterial taxa known to promote plant growth and confer tolerance to shade stress, which suggests common principles underpinning OJ-microbe interactions.ConclusionSoil microorganisms mediate plant responses to shade stress via plant-soil feedback and shade-induced change in the rhizosphere soil bacterial community structure for OJ and LP plants. These findings emphasize the importance of understanding plant-soil interactions and their role in the mechanisms underlying shade tolerance in shade-tolerant turfgrasses.

Influence of salt stress on the rhizosphere soil bacterial community structure and growth performance of groundnut (Arachis hypogaea L.).

Soil salinity is regarded as severe environmental stress that can change the composition of rhizosphere soil bacterial community and import a plethora of harms to crop plants. However, relatively little is known about the relationship between salt stress and root microbial communities in groundnuts. The goal of this study was to assess the effect of salt stress on groundnut growth performance and rhizosphere microbial community structure. Statistical analysis exhibited that salt stress indeed affected groundnut growth and pod yield. Further taxonomic analysis showed that the bacterial community predominantly consisted of phyla Proteobacteria, Actinobacteria, Saccharibacteria, Chloroflexi, Acidobacteria, and Cyanobacteria. Among these bacteria, numbers of Cyanobacteria and Acidobacteria mainly increased, while that of Actinobacteria and Chloroflexi decreased after salt treatment via taxonomic and qPCR analysis. Moreover, Sphingomonas and Microcoleus as the predominant genera in salt-treated rhizosphere soils might enhance salt tolerance as plant growth-promoting rhizobacteria. Metagenomic profiling showed that series of sequences related to signaling transduction, posttranslational modification, and chaperones were enriched in the salt-treated soils, which may have implications for plant survival and salt tolerance. These data will help us better understand the symbiotic relationship between the dominant microbial community and groundnuts and form the foundation for further improvement of salt tolerance of groundnuts via modification of soil microbial community.

Effects of Sphaeropsis Blight on Rhizosphere Soil Bacterial Community Structure and Soil Physicochemical Properties of Pinus sylvestris var. mongolica in Zhanggutai, China

Pinus sylvestris var. mongolica is an important tree species for ecological construction and environmental restoration owing to its rapid growth rate and excellent stress resistance. Pinus sylvestris var. mongolica sphaeropsis blight is a widespread disease caused by Sphaeropsis sapinea. This study was focused on non-infected (CK) and infected (SS) Pinus sylvestris var. mongolica plants in Zhanggutai area, Liaoning Province, China. Illumina high-throughput sequencing based on the templates of sequencing-by-synthesis working with reversible terminators is a widely used approach. In the present study, systematic differences in relationships among rhizosphere soil physicochemical properties, bacterial community structure, diverse bacterial genera, and alpha diversity indices between the two categories were evaluated. The current findings are as follows: (1) Shannon’s index of SS soil was significantly higher than CK, and it was significantly lower in May than July and September (p &lt; 0.05). (2) Non-metric multidimensional scaling (NMDS) showed a difference in bacterial community structure during May (spring), July (summer), and September. (3) At the phylum level, no significant difference was found in the bacterial genera between CK and SS soil for three seasons; however, at the genus level, there were about 19 different bacterial genera. The correlation studies between 19 different bacterial genera and environmental factors and α-diversity indicated that bacterial genera of non-infected and infected Pinus sylvestris var. mongolica were distributed differently. The bacterial genera with CK were positively correlated with soil physicochemical properties, while a negative correlation was found for SS. In conclusion, the differences in nutrient and microbial community structure in the rhizosphere soil of Pinus sylvestris var. mongolica are the main causes of shoot blight disease.