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Concept: Heterotroph


This work critically reviews modeling concepts for standard activated sludge wastewater treatment processes (e.g., hydrolysis, growth and decay of organisms, etc.) for some of the most commonly used models. Based on a short overview on the theoretical biochemistry knowledge this review should help model users to better understand (i) the model concepts used; (ii) the differences between models, and (iii) the limits of the models. The seven analyzed models are: (1) ASM1; (2) ASM2d; (3) ASM3; (4) ASM3 + BioP; (5) ASM2d + TUD; (6) Barker & Dold model; and (7) UCTPHO+. Nine standard processes are distinguished and discussed in the present work: hydrolysis; fermentation; ordinary heterotrophic organisms (OHO) growth; autotrophic nitrifying organisms (ANO) growth; OHO & ANO decay; poly-hydroxyalkanoates (PHA) storage; polyphosphate (polyP) storage; phosphorus accumulating organisms PAO) growth; and PAO decay. For a structured comparison, a new schematic representation of these processes is proposed. Each process is represented as a reaction with consumed components on the left of the figure and produced components on the right. Standardized icons, based on shapes and color codes, enable the representation of the stoichiometric modeling concepts and kinetics. This representation allows highlighting the conceptual differences of the models, and the level of simplification between the concepts and the theoretical knowledge. The model selection depending on their theoretical limitations and the main research needs to increase the model quality are finally discussed. Biotechnol. Bioeng. 2013; 110: 24-46. © 2012 Wiley Periodicals, Inc.

Concepts: Bacteria, Organism, Truth, Sewage treatment, Model, Heterotroph, Activated sludge, Autotroph


The presence of the synthetic estrogen 17α-ethinylestradiol (EE2) in the environment is of increasing concern due to the endocrine disruption of aquatic organisms. Incomplete removal from wastewater (WW) is one of the main sources of EE2 in aquatic ecosystems, thus improving processes like biological WW treatment/activated sludge (AS) is becoming significantly important. There are opposing results regarding EE2 biodegradability by AS; one discrepancy is the efficacy of heterotrophic bacteria. This research demonstrated the ability of heterotrophs commonly present in AS (B. subtilis, P. aeruginosa, P. putida, R. equi, R. erythropolis, R. rhodochrous, R. zopfii) to remove EE2. R. rhodochrous was the most successful with no detectable EE2 after 48 h; the other bacteria achieved 21%-61% EE2 removal. No additive or synergistic effects were observed due to the combination of the bacterial cultures with maximum EE2 removals of 43% after 300 h.

Concepts: DNA, Bacteria, Organism, Natural environment, Ecosystem, Bioremediation, Heterotroph, Removal


Factors controlling the community composition of marine heterotrophic prokaryotes include organic-C, mineral nutrients, predation, and viral lysis. Two mesocosm experiments, performed at an Arctic location and bottom-up manipulated with organic-C, had very different results in community composition for both prokaryotes and viruses. Previously, we showed how a simple mathematical model could reproduce food web level dynamics observed in these mesocosms, demonstrating strong top-down control through the predator chain from copepods via ciliates and heterotrophic nanoflagellates. Here, we use a steady-state analysis to connect ciliate biomass to bacterial carbon demand. This gives a coupling of top-down and bottom-up factors whereby low initial densities of ciliates are associated with mineral nutrient-limited heterotrophic prokaryotes that do not respond to external supply of labile organic-C. In contrast, high initial densities of ciliates give carbon-limited growth and high responsiveness to organic-C. The differences observed in ciliate abundance, and in prokaryote abundance and community composition in the two experiments were in accordance with these predictions. Responsiveness in the viral community followed a pattern similar to that of prokaryotes. Our study provides a unique link between the structure of the predator chain in the microbial food web and viral abundance and diversity.

Concepts: Archaea, Bacteria, Organism, Microbiology, Ecology, Food chain, Heterotroph, Plankton


We previously reported that Synechococcus elongatus PCC 7942, engineered with the sucrose transporter CscB, can export up to 85% of its photosynthetically-fixed carbon as sucrose and shows considerable promise as an alternative carbohydrate source. One approach to effectively utilize this cyanobacterium is to generate synthetic, light-driven consortia in which sucrose-metabolizing heterotrophs catalyze the conversion of the low-value carbohydrate into higher-value compounds in co-culture. Here, we report an improved synthetic photoautotroph/chemoheterotroph consortial design in which sucrose secreted by S. elongatus CscB directly supports the bacterium Halomonas boliviensis, a natural producer of the bioplastic precursor, PHB. We show that alginate encapsulation of S. elongatus CscB enhances sucrose-export rates ~2-fold within 66h, to ~290mg sucrose L(-1) d(-1) OD750(-1) and enhances the co-culture stability. Consortial H. boliviensis accumulate up to 31% of their dry-weight as PHB, reaching productivities up to 28.3mg PHB L(-1) d(-1). This light-driven, alginate-partitioned co-culture platform achieves PHB productivities that match or exceed those of traditionally engineered cyanobacterial monocultures. Importantly, S. elongatus CscB/H. boliviensis co-cultures were continuously productive for over 5 months and resisted invasive microbial species without the application of antibiotics or other chemical selection agents.

Concepts: Cyanobacteria, Photosynthesis, Archaea, Bacteria, Metabolism, Organism, Microorganism, Heterotroph


The nitrogen (N) cycle consists of complex microbe-mediated transformations driven by a variety of factors, including diversity and concentrations of N compounds. In this study, we examined taxonomic diversity and N substrate utilization by heterotrophic bacteria isolated from streams under complex and simple N-enrichment conditions.

Concepts: Bacteria, Evolution, Metabolism, Enzyme, Organism, Nitrogen, Domain, Heterotroph


There is increasing awareness that many terrestrial and aquatic organisms are not strictly heterotrophic or autotrophic but rather mixotrophic. Mixotrophy is an intermediate nutritional strategy, merging autotrophy and heterotrophy to acquire organic carbon and/or other elements, mainly N, P or Fe. We show that both terrestrial and aquatic mixotrophs fall into three categories, namely necrotrophic (where autotrophs prey on other organisms), biotrophic (where heterotrophs gain autotrophy by symbiosis) and absorbotrophic (where autotrophs take up environmental organic molecules). Here we discuss their physiological and ecological relevance since mixotrophy is found in virtually every ecosystem and occurs across the whole eukaryotic phylogeny, suggesting an evolutionary pressure towards mixotrophy. Ecosystem dynamics tend to separate light from non-carbon nutrients (N and P resources): the biological pump and water stratification in aquatic ecosystems deplete non-carbon nutrients from the photic zone, while terrestrial plant successions create a canopy layer with light but devoid of non-carbon soil nutrients. In both aquatic and terrestrial environments organisms face a grand écart (dancer’s splits, i.e., the need to reconcile two opposing needs) between optimal conditions for photosynthesis vs. gain of non-carbon elements. We suggest that mixotrophy allows adaptation of organisms to such ubiquist environmental gradients, ultimately explaining why mixotrophic strategies are widespread.

Concepts: Bacteria, Evolution, Organism, Plant, Ecosystem, Heterotroph, Autotroph, Primary nutritional groups


Microbial community diversity determines the function of each chamber of multi-stage moving bed biofilm reactor (MBBR) systems. How the microbial community data can be further used to serve wastewater treatment process modelling and optimization has been rarely studied. In this study, a MBBR system was set up to investigate the microbial community diversity of biofilm in each functional chamber. The compositions of microbial community of biofilm from different chambers of MBBR were quantified by high-throughput sequencing. Significantly higher proportion of autotrophs were found in the second aerobic chamber (15.4%), while 4.3% autotrophs were found in the first aerobic chamber. Autotrophs in anoxic chamber were negligible. Moreover, ratios of active heterotrophic biomass and autotrophic biomass (XH/XA) were obtained by performing respiration tests. By setting heterotroph/autotroph ratios obtained from sequencing analysis equal to XH/XA, a novel approach for kinetic model parameters estimation was developed. This work not only investigated microbial community of MBBR system, but also it provided an approach to make further use of molecular microbiology analysis results.

Concepts: Bacteria, Organism, Microbiology, Biotechnology, Sewage treatment, Heterotroph, Autotroph


Phosphorus (P) limitation has been demonstrated for micro-polluted surface water denitrification treatment in previous study. In this paper, a lab-scale comparative study of autotrophic denitrification (ADN) and heterotrophic denitrification (HDN) in phosphorus-limited surface water was investigated, aiming to find out the optimal nitrogen/phosphorus (N/P) ratio and the mechanism of the effect of P limitation on ADN and HDN. Furthermore, the optimal denitrification process was applied to the West Lake denitrification project, aiming to improve the water quality of the West Lake from worse than grade V to grade IV (GB3838-2006). The lab-scale study showed that the lack of P indeed inhibited HDN more greatly than ADN. The optimal N/P ratio for ADN and HDN was 25 and a 0.15 mg PO43–P L-1of microbial available phosphorus (MAP) was observed. P additions could greatly enhance the resistance of ADN and HDN to hydraulic loading shock. Besides, The P addition could effectively stimulate the HDN performance via enriching the heterotrophic denitrifiers and the denitrifying phosphate-accumulating organisms (DNPAOs). Additionally, HDN was more effective and cost-effective than ADN for treating P-limited surface water. The study of the full-scale HDBF (heterotrophic denitrification biofilter) indicated that the denitrification performance was periodically impacted by P limitation, particularly at low water temperatures.

Concepts: Bacteria, Organism, Water, Hydrology, Water quality, Heterotroph, Denitrification, Autotroph


A rich eukaryotic planktonic community exists in high-mountain lakes despite the diluted, oligotrophic and cold, harsh prevailing conditions. Attempts of an overarching appraisal have been traditionally hampered by observational limitations of small, colorless, and soft eukaryotes. We aimed to uncover the regional eukaryotic biodiversity of a mountain lakes district to obtain general conclusions on diversity patterns, dominance, geographic diversification, and food-web players common to oligotrophic worldwide distributed freshwater systems. An unprecedented survey of 227 high-altitude lakes comprising large environmental gradients was carried out using Illumina massive tag sequencing of the 18S rRNA gene. We observed a large Chrysophyceae dominance in richness, abundance and novelty, and unveiled an unexpected richness in heterotrophic phagotrophs and parasites. In particular, Cercozoa and Chytridiomycota showed diversity features similar to the dominant autotrophic groups. The prominent beta-dispersion shown by parasites suggests highly specific interactions and a relevant role in food webs. Interestingly, the freshwater Pyrenean metacommunity contained more diverse specific populations than its closest marine oligotrophic equivalent, with consistently higher beta-diversity. The relevance of unseen groups opens new perspectives for the better understanding of planktonic food webs. Mountain lakes, with remarkable environmental idiosyncrasies, may be suitable environments for the genetic diversification of microscopic eukaryotic life forms.

Concepts: Bacteria, Organism, Life, Ribosomal RNA, Fungus, Natural environment, Phytoplankton, Heterotroph


Heterotrophic plants provide excellent opportunities to study the effects of altered selective regimes on genome evolution. Plastid genome (plastome) studies in heterotrophic plants are often based on one or a few highly divergent species or sequences as representatives of an entire lineage, thus missing important evolutionary-transitory events. Here, we present the first infraspecific analysis of plastome evolution in any heterotrophic plant. By combining genome skimming and targeted sequence capture, we address hypotheses on the degree and rate of plastome degradation in a complex of leafless orchids (Corallorhiza striata) across its geographic range. Plastomes provide strong support for relationships and evidence of reciprocal monophyly between C. involuta and the endangered C. bentleyi. Plastome degradation is extensive, occurring rapidly over a few million years, with evidence of differing rates of genomic change among the two principal clades of the complex. Genome skimming and targeted sequence capture differ widely in coverage depth overall, with depth in targeted sequence capture datasets varying immensely across the plastome as a function of GC content. These findings will help to fill a knowledge gap in models of heterotrophic plastid genome evolution, and have implications for future studies in heterotrophs.

Concepts: DNA, Genetics, Organism, Genome, Plastid, Species, Heterotroph, Orchidaceae