Iron concentrations and EBPR performance

Interestingly, only eight out of a total twenty-eight genera showed negative correlation with Fe, while

nineteen and eighteen genera displayed negative correlations with specific PO4-P release and specific

PO4-P uptake, respectively. This suggests that only nine/ten genera in this sample were responsible for

improving EBPR performance. Of these nine/ten genera responsible for EBPR Alkalispirillum, Dechloromonas, Micropruina, Ochrobactrum, Simplicispira, and Tetrasphaera measured the strongest positive correlations.

Figure 5-7a and Figure 5-7b show clearly Tetrasphaera abundance increases as Fe decreases. Tetrasphaera is able to withstand acidic conditions and was found to have higher relative abundance at pH 4 than pH 10 (Yuan et al., 2015; Weissbrodt et al., 2013). This is supported by the strong negative correlation with pH in this research (r=-0.62). Tetrasphaera has a high metabolic versatility and can utilise glucose, it can be found in industrial wastewaters and septic tanks, suggesting Tetrasphaera is prolific and resistant to contaminants (Kristiansen et al., 2013; Muszynski et al., 2015). Tetrasphaera was present in twenty-two of thirty-two samples in total, showing that it is the most prolific genus and resistant to various levels of Fe dosing and EBPR performance. Ju et al. (2014) suggest that Tetrasphaera are present at higher abundances in winter months, but in this study correlations with temperature and rainfall were weak (r=-0.06 and r=-0.15, respectively). Tetrasphaera measured strong positive correlation with Aestuariimicrobium (r=0.51) and strong negative correlation with Lactobacillus (r=- 0.46).

Figure 5-7a and Figure 5-7b display that Terrimonas has a positive correlation with Fe concentrations and negative correlation with EBPR performance. This is in contrast to Tetrasphaera, but similar to fifteen other genera detected. From forty-eight parameters measured in correlation analysis Terrimonas and Tetrasphaera displayed opposite behaviour in thirty-two parameters, i.e. showing positive correlations when the other was negative and vice versa. This opposite behaviour of the genera were

uptake reaction rate, release reaction rate, and total P. Terrimonas and Tetrasphaera each also showed strong and opposite correlations with Ca with r=0.53 and r=-0.52, respectively. Terrimonas was the dominant genus in aerobic, anoxic and anaerobic sludges (Zhang et al., 2015). Little is known of the environmental significance of Terrimonas (Zhang et al., 2011), but this analysis may help illuminate the behaviour of Terrimonas in EBPR.

Ochrobactrum displayed its highest abundances at medium Fe and medium EBPR performance as shown in Figure 5-7c and Figure 5-7d. Ochrobactrum are obligately aerobic, their optimum growth temperature ranges from 20-37°C (Holt et al., 1994). Little information on Ochrobactrum could be found, therefore the results presented here hope to add to the knowledge of this genus. Ochrobactrum showed the strongest negative correlation with alkalinity (r=-0.52) than the other genera. A very strong negative correlation with diversity (r=-0.81) was measured showing that Ochrobactrum may outcompete with other genera in the EBPR system. Ochrobactrum measured negative correlations with every genera (twenty-eight in total) except Clostridium XI (r=0.25) and Lactobacillus (r=0.65). Ochrobactrum and Lactobacillus conveyed similar behaviours showing opposite correlations with only five parameters

measured: Al, Ca, dry solids %, release reaction rate and specific PO4-P uptake. Overall, Ochrobactrum

and Lactobacillus measured very similar correlation values differing most by 0.23 in Al and uptake reaction rate correlations.

Dechloromonas in Figure 5-7c and Figure 5-7d showed opposite behaviour of Ochrobactrum, a weak correlation of r=-0.14 exists between the two genera. Dechloromonas are facultatively anaerobic bacteria and grow in a range of environments (optimum pH 7.2 and 30°C) and thus are environmentally dominant (Nor et al., 2011; Wolterink et al., 2005). Dechloromonas are not involved in P removal and no evidence of polyP accumulation was seen in samples taken after phosphate addition (Ahn et al., 2007).

However, in this study Dechloromonas showed a moderate correlation with specific PO4-P

release/uptake with r=0.34 and r=0.31. Dechloromonas abundance peaked on day 349 when it was the most abundant genus at 35.7%. Day 349 also marks the significant increase in microbial community diversity (Figure 3a). Dechloromonas showed strong positive correlations with Alkalispirillum (r=0.57) and Simplicispira (r=0.78). Dechloromonas and Simplicispira differed in correlation response in six of

forty-eight parameters, namely, Dongia, Elioraea, Terrimonas, Fe concentration, dry solids %, and NH4.

5.6 Conclusion

Simultaneous chemical removal of P can support the EBPR process, but overdosing can have a deleterious effect on microorganisms responsible for biological P removal. Chemical removal of P

The observed inverse relationship between chemical dosing and EBPR performance in this system, supports the hypothesis of this research.

Proteobacteria and Actinobacteria measured positive correlations with specific PO4-P release and

specific PO4-P uptake showing each plays a major role in EBPR. Bacteroidetes, Firmicutes, Acidobacteria,

and Actinobacteria were reported as playing an important role in nutrient removal processes (Ma et al., 2015). However, Bacteroidetes and Firmicutes were absent from highly performing EBPR samples in this study. During the sampling regime, Firmicutes and Actinobacteria appeared to substitute positions with each other due to these bacteria containing a family of proteins functionally equivalent to each other (Ravagnani et al., 2005).

In terms of genera Simplicispira, Tetrasphaera, and Micropruina were present in highest performing

specific PO4-P release and specific PO4-P uptake samples. Each of the genera measured strong

correlations with specific PO4-P release and specific PO4-P uptake and therefore play an important role

in EBPR. Micropruina measured weaker correlation with specific PO4-P release but stronger correlation

with specific PO4-P uptake indicating in which aspect of EBPR they play a larger role. Tetrasphaera were

the third most dominant genus across the sampling regime while Micropruina being the least dominant. This analysis shows that the dominance of microorganisms does not reflect its importance in the EBPR process.

Most literature suggests that increased diversity provides improved process stability and reduces risk of

EBPR failure. However, no strong relationship was found between diversity and specific PO4-P release

and specific PO4-P uptake. A dramatic decrease in diversity was measured followed by a recovery of

diversity and mutual improvement of EBPR performance. The recovery of diversity may be due to the mutation and adaptation of new dominant bacteria strains (Li & Jin, 2009). Lactobacilli were present at low diversities due to their ability to outcompete other microorganisms and their fast growth rates (Gois et al., 2013). However, as diversity increased Lactobacillus was replaced by Ochrobactrum with which it has a strong positive correlational relationship. A very strong negative correlation with diversity was measured showing that Ochrobactrum outcompetes genera in the EBPR system. Ochrobactrum measured negative correlations with every genera (twenty-eight in total) except Clostridium XI and Lactobacillus.

Heavy metals can cause PAO infection and death through intra-cellular phage induction (Motlagh et al., 2015). However, presence of Fe increased the diversity of microorganisms, which does not agree with the hypothesis of this research. A positive correlation was measured between Fe concentration and diversity, and Fe concentration and genus count. A strong positive correlation was measured between Bacteroidetes and Fe concentration due to the fact that Bacteroidetes are very resistant versatile microorganisms (Wan et al., 2013; Naumoff & Dedysh, 2012). In terms of genera, Terrimonas measured

the highest correlation with Fe concentration, but little is known of the environmental significance of Terrimonas. Terrimonas measured high abundance at elevated Fe concentrations and diversity, displaying it is a resilient and competitive genus. Aestuariimicrobium and Dokdonella are also considered resistant to Fe, because they were present in all five highest Fe samples.

Eight out of a total twenty-eight genera showed negative correlation with Fe, while nineteen and

eighteen genera displayed negative correlations with specific PO4-P release and specific PO4-P uptake,

respectively. This suggests that only nine/ten genera in this sample were responsible for improving EBPR performance. Tetrasphaera were present in twenty-two of thirty-two samples in total, showing that it is the most prolific genus and resistant to various levels of Fe dosing and EBPR performance. Terrimonas and Tetrasphaera displayed opposite behaviour in thirty-two parameters, i.e. showing positive correlations when the other was negative and vice versa. Little is known of the environmental significance of Terrimonas, but this analysis may help illuminate the behaviour of Terrimonas in EBPR.

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