Characterization Free-living (FL) and Particle-attached (PA) bacterial communities of a canyon river reservoir on Yungui Plateau, China

doi:10.21203/rs.3.rs-1535823/v1

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Characterization Free-living (FL) and Particle-attached (PA) bacterial communities of a canyon river reservoir on Yungui Plateau, China

https://doi.org/10.21203/rs.3.rs-1535823/v1

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Bacteria play an important role in nutrient recycling and energy conversion in any aquatic ecosystem. Revealing the composition of free-living (FL) and particle-attached (PA) bacterial communities provide insights into their roles for biogeochemical processes and algal bloom dynamics. This study revealed the structure of FL and PA bacterial communities in a canyon reservoir in the Yungui-Plateau region, southern China. Firstly, we compared the difference using 3.0 µm and 5.0 µm filter for PA and FL bacterial communities (defined as PA3, FL3 and PA5, FL5), in order to choose the better sampling protocol. Secondly, we identified the differences between FL and PA in respect of their composition, diversity as well as function. Results showed that (1) dominant taxa did not change between FL3 and FL5, PA3 and PA5, but the relative abundance varied. Community composition and predicted gene functions showed significant differences between PA3 and PA5, while insignificant difference was noticed between FL3 and FL5; (2) FL was dominated by Actinobacteria and Proteobacteria, while PA was dominated by Planctomycetes and Proteobacteria; (3) there was insignificant difference on α-diversity between PA3 and PA3, but PA5 community exhibited greater dissimilarity (β-diversity) than PA3; (4) differences were detected between FL and PA community, with PA community exhibiting greater diversity and metabolic versatility than FL; (5) nitrogen metabolism related genes were abundant, implying an important role of both FL and PA in aquatic nitrogen cycling and hence in the microbial loop in this reservoir. This study provided evidence for using 5.0 µm filter to separate PA bacterial community, and also presented fundamental information on FL and PA bacterioplankton in Wujiangdu reservoir.

River reservoir

bacterioplankton

community composition

bacterial diversity

gene functions

environmental drivers

Bacteria, are a major component of the microbial loop, and play an important role in biogeochemical processes. They mineralize dissolved organic matter, such as algal exudate and transfer carbon and energy to higher trophic levels, and play an essential role in the microbial food web (Azam et al., 1983). Furthermore, bacterioplankton can play different roles in the initiation, development and termination of a broad range of algal blooms (Paver et al., 2013; Buchan et al., 2014). Thereby, the bacterioplankton community can, to some extent, indicate trophic status and certain organic or heavy metal pollution, mirroring the state of water quality (Pinto et al., 2021). Therefore, it was proposed that bacteria act as sentinels of environmental change due to their sensitivity to changing environmental conditions including trophic status and anthropogenic pollution (Harnisz, 2013; Savio et al., 2015). Based on their life style, there are two bacterial fractions, i.e. free-living (FL), with a size range between 0.22 to 3.0 µm or 5 µm (Crespo et al., 2013; Grossart, 2010), and particle-attached (PA), which can be separated from FL by filtration on 3.0 µm/ 5 µm filters. FL refers to bacteria floating freely in the water column, while the PA fraction comprises bacteria attached to organic particles and living organisms. Several studies revealed that there are significant phylogenetic differences between FL and PA fractions, whereby some bacteria can rapidly exchange between the size fractions (Tang et al., 2015; Bizic-Ionescu et al., 2014). The first step to study FL and PA assemblages is to collect the most representative PA samples. Generally, the prevalent method is filtration and centrifugation, earlier study has proposed that centrifugation was proper in eutrophic and turbulent lakes, but it could enrich some taxa disproportionately due to various sinking velocity (Xie et al., 2020). Filtration with 3.0 µm (ImLiu et al., 2019; Wang et al., 2020) or 5.0 µm (Allgaier and Grossart, 2006; Parveen et al., 2011; Zhao et al., 2017; Hu et al., 2020) pore size filter is the most frequently used. Different pore sizes may affect the structure of community. Moreover, the methods applied for collection may be trophic-dependent because nutrients are associated with the composition and size of particles in the water column (Tang et al., 2017).

Research about bacterioplankton in river reservoirs on Yungui-Plateau are scarce. Zhang et al. (2020) investigated the relationship between phytoplankton and bacterioplankton in Dianchi Lake. Wu et al. (2019) revealed the vertical bacterial profile in Lake Lugu. Both studies were conducted in Yunnan Province. Wu River is the largest southern tributary of Yangtze River and also the largest river in Guizhou Province. Wujiangdu reservoir was the first major dam established in the middle and upper reaches of the Wu River. There are many streams supplying water to the impoundment, such as Xifeng River and Pianyan River. Due to the karstification characteristics in this region, Wu River water is abundant with carbonated weathering (Han and Liu, 2004). Yue et al. (2021) investigated the vertical distribution of bacterial community in Wujiangdu reservoir and emphasized the role of stratification on the vertical pattern of bacterial community structure. Cascade dams along Wu River could affect overall microbial community via altering hydrological features. However, their community structure and function, classified as FL and PA fraction, in such plateau reservoirs are still poorly understood. Information about the bacterial distribution, assembly, diversity and functioning in this reservoir could help us understand the mechanisms underlying microbial food web and phytoplankton succession in such plateau freshwater ecosystems. Therefore, it is necessary to unveil the bacterial community structure in reservoirs of Yungui Plateau.

We collect samples from 14 sites along Wujiangdu reservoir, a typical canyon river reservoir. Based on 16S rRNA marker gene, we firstly compared the results of PA bacterioplankton by using 3.0 µm and 5.0 µm filter, and simultaneously investigate both FL and PA bacterial community structure, diversity and gene function. We hypothesized that (1) different pore size filters would result in different PA bacterioplankton community; (2) FL and PA bacterioplankton differ in composition, diversity and functions in Wujiangdu reservoir.

2.1 Site description and sampling

Wujiangdu Reaservoir, built in 1979, is located at the lower basin of Wu River, and at the conjunction of Xifeng Town and Zunyi city, Guizhou Province. It has a watershed of 27790 km², a mean depth of 154 m, and volume of 23 x 10⁸ m³. The average water residence time is 53 days. This reservoir has a history of cage aquaculture activities since 1999.

Water samples were collected from 14 locations (HS, YL, YLr, KC, JK, JKr, DTD, TL, TLr, XT, XF, XFr, PY, and PYr) in Wujiangdu Reservoir on Aug 19th, 2021 (Fig. 1). These locations were chosen due to their position at the confluence of small rivers and the main river. Water Temperature, Dissolved Oxygen (DO), pH and Conductivity (Cond) were measured on site via a portable YSI probe (HANNA HI98194), Secchi depth (SD) was measured by a Secchi disk at each station. Based on the temperature profile, there was weak thermal stratification. Thus, we regard that the surface samples could represent the epilimnion. For chemical analyses, 1L of surface water (0.5 m) was taken using a water sampler. And for microbial analyses, an additional 2 liter was filled in a sterile PE bottle which was kept at in site temperature for transport to the lab.

2.2 Water chemistry analysis

Total phosphorus (TP), phosphate (PO₄³⁻), total nitrogen (TN), nitrate (NO₃⁻), nitrite (NO₂⁻), silicate (SiO₂), total suspended solids (TTS), and Chlorophyll a (Chl a) were measured according to standard methods (A.P.H.A., 2012). Based on these measurements, trophic status index (TSI) was calculated according to the equations below (Carlson, 1991; Matthews et al., 2002; An & Park, 2003):

TSI_SD=60-14.42ln (SD, m)

TSI_TN=54.45 + 14.43ln (TN, mg/L)

TSI_TP=14.42ln (TP, µg/L) + 4.15

TSI_CHL=9.81ln (CHL, µg/L) + 30.6

2.3 Bacterial community analysis

For bacterial DNA, two types of polycarbonate filter (3.0 µm and 5.0 µm) were applied to collect PA bacteria (PA3 and PA5). 400 mL water were filtrated on a 47 mm diameter filter firstly, and then filtered through a 0.22 µm filter to collect FL fraction (FL3 and FL 5). All filters were stored at -80 ℃ for further analyses. The ALFA-seq Advanced water Kit was used for DNA extraction according to the manufacturer’s instruction. DNA concentration and purity were measured using the NanoDrop One. V4-V5 region of 16S rRNA gene were amplified used the primer 806R/515F. Based on NEBNext Ultra™ DNA library Prep kit for Illumina standard protocol, a library was established before sequencing on an IlluminaHiseq2500 platform and 250bp paired-end reads were generated. Quality filtering on the paired-end raw reads were performed according to the Trimmomatic (V0.33) quality-controlled process. Paired-end clean reads were merged using FLASH (V1.2.11). Then16S rRNA gene sequences were analyzed by usearch software (V8.0.1517). sequences with ≥ 97% similarity were assigned to the same OTU. For each representative sequence, the GreenGene database was used based on RDP classifier algorithm and the assign_taxonomy.py script in Qiime to annotate taxonomic information. Rarefaction curve and rank abundance curve were seen in Fig. S1.

Diversity index was calculated in R, including OTU reads number, Chao1 and Abundance-based coverage estimator (ACE), Simpson index, Shannon index, and Evenness. Chao1 and ACE are used to estimate richness. Shannon and Simspon index provide more inference about the community composition than simple species richness or evenness. Analysis of variance (ANOVA) tests were used to test for significant differences in diversity between FL and PA fractions. Community dissimilarity was calculated using Bray-Curtis dissimilarity. Non-parametric multidimensional scaling (NMDS) was performed to display the distance of bacterial communities. Permutational multivariate analysis of variance (PerMANOVA) was used to test whether there was significant difference on community composition between groups (Anderson, 2008). PERMDISP, a common test completed in conjunction with PerMANOVA, a multivariate analog of Levene’s test for homogeneity of multivariate variances, was performed to test the null hypothesis of ‘no difference in dispersion between groups’ (Anderson and Walsh, 2013). Distance decay pattern was tested using Mantel tests, which calculates geographic distance based on sampling coordinates, and Euclidean distance using environmental parameters. Redundancy analysis (RDA) was used to summarize the relationship between bacterial community and measured environmental factors. PerMANOVA, PERMDISP, NMDS and RDA were performed using the ‘vegan’ package in R (Oksanen et al., 2020).

OTU data was used for Phylogenetic Investigation of Communities by Reconstruction of Unobserved States (PICRUSTs2) and for gene function prediction based on clusters of orthologs groups (COG) database and Kyoto Encyclopedia of Genes and Genomes (KEGG) pathways (Douglas et al, 2020).

3.1 Water physio-chemical parameters

Water transparency, indicated by SD, ranged from 2.6 m to 5.3 m. Nutrient concentrations, including total nitrogen (TN), total phosphorus (TP), phosphate (PO₄³⁻), nitrate (NO₃⁻) and total suspended solid (TSS) and Chlorophyll a (Chl a), exhibited spatial heterogeneity in this reservoir with various degrees. TN ranged from 2.31 mg/L to 3.25 mg/L, and TP from 0.01 mg/L to 0.02 mg/L. Nitrate (NO₃⁻) was the major component of dissolved nitrogen, ranging from 1.72 mg/L to 2.55 mg/L, followed by NO₂⁻ (0.008 mg/L- 0.035 mg/L) and NH₄⁺ (0.008 mg/L-0.110 mg/L) (nitrogen and phosphorus concentration seen in Fig. S2). According to Trophic Status Index (TSI), Wujiangdu reservoir was mesotrophic to eutrophic (Table 1).

Table 1

Summary of Trophic Status Index (TSI) in Wujiagndu Reservoir
TSI	Mean	Min	Max
TSI_SD	46.99	40.37	51.52
TSI_TN	68.68	66.55	71.47
TSI_TP	45.05	43.31	47.19
TSI_CHL	50.54	43.09	56.62

3.3 FL and PA bacterioplankton community in surface water

In the surface waters of Wujiangdu reservoir, the detected bacterial OTUs belonged to 37 phyla and 73 class. Within FL or PA, the dominant taxa were the same, but with varying relative abundance. Dominant phyla in FL include Actinobacteria and Proteobacteria, whereas dominant phyla in PA were Planctomycetes and Proteobacteria. On class level, Actinobacteria and Acidimicrobiia were dominant taxa in FL. Planctomycetes predominant in PA, followed by α-Proteobacteria. On order level, Frankiales and Microtrichales (Actinobacteria) were dominant in FL, whereas Pirellulales (Planctomycetes) predominated in PA. On family level, the dominant group in FL fraction were Sporichthyaceae and Illumatobacteraceae (Actinobacteria) and Burkholderiaceae (Proteobacteria). Whereas Pirellulaceae (Planctomycetes) dominated the PA bacterial communities (Fig. 2). ANOVA tests showed that there were insignificant differences on the relative abundance of dominant taxa between FL3 and FL5 fraction at phylum, class, order and family level (p > 0.05). While significant differences were noticed between PA3 and PA5 fraction on relative abundance (p < 0.05). Beside, significant different was detected between PA 3 and PA5 (p < 0.05), and in-significant difference was noticed between FL 3 and FL5 by PerMANOVA test (p > 0.05).

Community structure differed between FL and PA, with taxa showing various relative abundance. For community structure, PerMANOVA suggested that there was significant difference between FL and PA bacterial community (both 3.0 and 5.0) (p = 0.001). SIMPER analysis showed that significant different phyla between FL and PA were Actinobacteria and Planctomycetes.

3.3 Bacterial community diversity

Reads number, Chao1, ACE, Shannon index and Simpson index, and Evenness index were calculated. It showed that all richness (reads number, Chao1 and ACE) and diversity indexes (Simpson index, Shannon index and evenness) were greater in PA5 than PA3. But no such pattern was observed for FL3 and FL5 (Fig. 3). ANOVA tests suggested that there was insignificant difference on diversity indexes between FL3 and FL5, whereas significant difference was detected on Shannon index between PA3 and PA5 (p = 0.04). Besides, richness showed higher values in PA than in FL. Whereas diversity indexes exhibited various patterns in PA and FL, with FL (both FL3 and FL5) greater than PA3, but similar with PA5. Significant differences on diversity indices were detected between FL3 and PA3 diversity, and FL5 and PA5.

NMDS was performed to identify the β-diversity of both FL and PA bacterioplankton community using Bray-Curtis dissimilarity index. FL5 had greater dissimilarity than FL3, and PA5 was more dissimilar than PA3 (Fig. 4a). PERMDISP test suggested that there was significant difference on β-diversity between PA3 and PA5 (p = 0.02) (Fig. 4b).

3.4 Functional analysis of bacterial communities

Dominant gene functions clustered on metabolism based on KEGG pathways for both FL and PA bacterial community, primarily including carbohydrate, amino acid, cofactor and vitamins metabolism. An overview on the relevant diseases is given in Fig. 5a and b. In addition, sub-dominant functions were associated with genetic information processing, such as replication and repair, folding, sorting and degradation, translation, transcription, transport and catabolism. Other less-dominant categories of KEGG pathways were related to cellular processes and environmental information processes. It was worthy noticing that several OTUs associated with human diseases were detected in this study, such as neurodegenerative disease, parasitic and bacterial infectious diseases, immune systems and cancer. Significant differences were noticed on gene function between PA3 and PA5 (p < 0.01).

Wujiangdu reservoir exhibited a relatively high level of nitrogen, as indicated by TN, nitrate, nitrite and ammonium concentration. Genes associated with nitrogen metabolism, clustered as KO, were observed based on KEGG pathway analysis. There were 47 nitrogen-relevant KO noticed in this reservoir. Among them, the predominant gene is ko01915 (glnA, GLUL), coding for glutamine synthetase (GS).

3.5 Bacterioplankton community with environmental factors

Mantel tests suggested that environmental matrix had a strong relationship with the FL bacterial communities (p < 0.01), while weak relationship with PA bacterial communities (p = 0.02). in other words, bacterial communities became more dissimilar along with the environmental conditions. Moreover, both FL and PA bacterioplankton communities exhibited significant relationship with geographic separation (p = 0.001) (Fig. 6). Therefore, distance decay pattern occurs in this reservoir for FL and PA bacteria (Table 2).

Table 2

Results of Mantel tests (Env: environmental factors; Geo: Geographical factor, R: correlation coefficient; p: probability value)
	FL3	FL5	PA3	PA5
Env-R	0.65	0.45	0.44	0.24
Env-p	0.001	0.001	0.001	0.02
Geo-R	0.61	0.56	0.61	0.60
Geo-p	0.001	0.002	0.001	0.001

RDA showed that FL and PA bacterioplankton, dominated by varying relative abundance of different phyla, were affected by different environmental factors (Fig. 7a). FL fraction was present in the right panel on the biplot, along the NH₄⁺, Chl a, TP, TN and NO₃⁻ gradients, whereas PA fraction, distributed on the left side, was associated with pH, TSS, and SD. Correlation analysis results suggested that dominant phyla (top 10) exhibiting different correlation relationships with environmental factors (Fig. 7b and c). In addition, Cyanobacteria showed positively correlation with Planctomycetes in FL, whereas negatively correlation in PA. Proteobacteria negatively correlated with Cyanobacteria in both FL and PA.

4.1 Comparison of PA3 and PA5 bacterioplankton composition, diversity and gene functions

Actinobacteria, small and slow-growing (Allgaier et al., 2007), are ubiquitous in lakes. they are ultramicrobacteria, able to decompose high molecular weight compounds, such as cellulose, lignin, are generally free-living, open-water defense specialists (Newton et al., 2011; Koblizek, 2015). This group represents a unique pelagic freshwater lineage, and they are uniform in the metabolism and ecological requirements.

Proteobacteria appeared to be comprised more from PA (27.15%) than from FL (23.4%) bacterioplankton, with alpha-Proteobacteria and gamma-Proteobacteria as the most dominant subphylum of Proteobacteria in both FL and PA. Proteobacteria with these two patterns implying their adaptation strategy to the environment (Dang and Lovell, 2016). Alpha-Proteobacteria are phototrophic bacteria with preference for low-nutrient conditions and resistant to predation (Imhoff, 2006, Newton et al., 2011). Whereas gamma -Proteobacteria prefer nutrient-rich environment, with the capacity to rapidly exploit the extra nutrient available.

Planctomycetes, predominant in PA community, normally present in low abundance in lakes and can be abundant in nutrient-enriched waters (Liu et al., 2014; Newton et al., 2011). They usually have large cell sizes due to their cellular structures and budding cell division. This group prefer to be attached than free-living (Allgaier and Grossart, 2006). Earlier studies suggested observed its high abundance associated with diatom (Morris et al., 2006) and cyanobacterial blooms (Woodhouse et al., 2016; Guedes et al., 2018). Ruber et al. (2017) reported high Planctomycetes abundance in a lake dominated by Synechococcus, and Salmaso et al. (2017) observed that Planctomyces was the most abundant genus within Planctomycetes in a bacterioplankton with Synechoccus accounted for 30% of Cyanobacteria. Consistent with other studies, we observed that Synechococcus was relatively high in this reservoir. This observvation might imply the co-occurrence pattern of Plantomycetes and Synechococcus in this reservoir. But their functional roles in association with phytoplankton, particularly cyanobacterial bloom remain unclear and need to be further researched. One possible mechanism could be their degradation capacity of sulfated polysaccharides produced by cyanobacteria (Cai et al. 2013).

Richness and diversity were greater in PA5 than PA3. Besides, greater dispersion of PA5 was observed than PA3, which can be interpreted as higher dissimilarity of PA5 community structure. For comparison between FL and PA, the richness was higher in PA than FL, but diversity was lower in PA3 than FL, and PA5 was similar with FL. This indicated that more abundance but less diverse bacteria were collected on 3.0 µm filter. The lower diversity of PA3 suggested that filtration through 5.0 µm filter was better than 3.0 µm filter for collecting PA bacteria in respect of diversity. Therefore, filtration using 5.0 µm was better at separating PA bacterioplankton in this reservoir when diversity was the priority for research.

Significant differences were also observed for predicted gene functions between PA3 and PA5. In this reservoir, the observed abundant metabolism function suggested that bacterioplankton, both FL and PA, involved in the decomposition and recycling of nutrients and other compounds. Furthermore, the predominant nitrogen-relevant gene was coding for glutamine synthetase. This enzyme catalyzes the ATP-dependent conversion of glutamate and ammonia to glutamine. It uses the bioavailable nitrogen in the form of ammonium into cellular metabolism. The abundant nitrogen metabolism gene, as well as relatively high level of nitrogen observed in Wujiangdu reservoir implies the productivity of bacterioplankton in the recycling of nitrogen.

4.2 Comparison of filter with different pore size for PA collection

Different porosity may be one of the reasons for low diversity in PA samples on 3.0 µm filters than 5.0 µm filters in Wujiangdu reservoir. This might be associated with the filter clogging and cell retention. During filtration process, clogging could lead to a progressive increase in the size range and diversity of the retained particles (Padilla et al., 2015). FL bacteria and bacteria attached to smaller particles could be detained on filters due to clogging as the increase of filtration time. Therefore, when the filtration time using 3.0 µm filters increases, smaller cells less than 3.0 µm and potentially free-living cells could also be retained. This explains that the relative abundances of some taxa were greater in PA3 than PA5. Besides, it would accumulate more dominant taxa, even though rare species could be collected using 3.0 µm. Thus, the diversity of PA3 was lower than PA5.

Meanwhile, PA bacteria could also enter into FL due to detachment from particles (Padilla et al., 2015). Clogging and detachment would increase the similarity between FL and PA fraction. Additionally, the lifestyle of FL and PA was not absolutely defined, many taxa possess an alternative lifestyle (Grossart, 2010). In our results, the higher diversity in PA5 than PA3 suggested that using 5.0 µm filter was better choice for PA separation.

Particles in deep lakes are generally rich in organic detrital matter which is densely colonized by bacteria, while particles in turbulent estuaries tend to be smaller with significant content of inorganic matter (Simon et al., 2002; Zhang et al., 2016). Wujaingdu reservoir is a typical riverine reservoir, with the water residence time about 0.14 year (Wang et al., 2019), and the particulate organic matter was mainly from phytoplankton-derived particulate organic carbon (Shi et al., 2018). The choice of pore size to separate PA from FL fractions is dependent on the size of the particles present in the aquatic ecosystem. Therefore, heterotrophic bacteria were mainly colonized on large organic particles in Wujiangdu reservoir. Our results suggested that using 5.0 um filters is the better choice to separate PA bacteria. (1) There are significant differences between PA3 and PA5 on community composition; (2) diversity of PA5 is greater than PA3; (3) bacterial community of PA5 was significantly different from FL community, indicating a distinct discrimination of FL and PA.

4.3 The relationship between environmental factors and bacterioplankton in Wujiangdu reservoir

Distance decay pattern refers that communities become increasingly different as the distance increases (Nekola and White, 1999). The underlying mechanism for this phenomenon is that environmental conditions are more similar with shorter distance. Therefore, species adapted to such habitats would also appear and colonize in similar habitats. Another reason is dispersal limitation, meaning that individuals tend to arrive at nearby sites. For microbes, however, it is generally regarded as no dispersal limitation, but the environment selects (Becking, 1934). In this reservoir, distance decay pattern was observed for both FL and PA, implying that both spatial and environmental factors affect the bacterioplankton community. For PA5 community, the influence of environment was relatively weak, indicating other environmental factors might play more important role. For PA bacterioplankton, the particle was crucial for determining the specific species attached. Thus, the composition of PA is dependent on the composition of particle in this reservoir.

This study proposed that 5.0 µm pore size filter was better to collect PA bacterioplankton because it could retain more diverse PA community. Moreover, it investigated both FL and PA bacterioplankton community structure in this typical Plateau reservoir in Yungui Plateau. FL and PA bacterioplankton exhibited different composition, diversity, and gene functions. The mechanisms underlying the community assemble and the observed human disease relevant gene need to be further researched in the next step.

Funding

This study was funded by National Scientific Funding of China (No. 32060270), the Science and Technology Foundation of Guizhou Province (No. Qian ke he ji chu [2020]1Y072), and the Doctor Scientific Research Foundation of Guizhou Normal University for 2017 (No. 11904/0519087).

Competing interests

The authors have no relevant financial or non-financial interests to disclose.

Author contribution

All authors contributed to the study conception and design. Material preparation, data collection and analysis were performed by Chen Chen and Xu Tao. The first draft of the manuscript was written by Yang Yang and all authors commented on previous versions of the manuscript. All authors read and approved the final manuscript.

Ethical approval - Not applicable

Consent to participate – Not applicable

Consent to publish – Not applicable

Availability of data and materials

Readers interested in this study may contact the corresponding author for dataset access.

Acknowledgements

We thank Professor Hans-Peter Grossart from Leibniz Institute for Freshwater Ecology and Inland Fisheries for commenting on the manuscript.

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Characterization Free-living (FL) and Particle-attached (PA) bacterial communities of a canyon river reservoir on Yungui Plateau, China

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Abstract

Figures

1. Introduction

2. Materials And Methods

2.1 Site description and sampling

2.2 Water chemistry analysis

2.3 Bacterial community analysis

3. Results

3.1 Water physio-chemical parameters

3.3 FL and PA bacterioplankton community in surface water

3.3 Bacterial community diversity

3.4 Functional analysis of bacterial communities

3.5 Bacterioplankton community with environmental factors

4. Discussions

4.1 Comparison of PA3 and PA5 bacterioplankton composition, diversity and gene functions

4.2 Comparison of filter with different pore size for PA collection

4.3 The relationship between environmental factors and bacterioplankton in Wujiangdu reservoir

5. Conclusions

Declarations

References

Supplementary Files

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