Mark S. Wilson, Ph.D.

Genetic and Biochemical Characterization of PAH-degrading Microorganisms from Marine and Estuarine Environments

Studies of naphthalene degradation are significant because naphthalene is a common pollutant that serves as a chemical model for the degradation of polycyclic aromatic hydrocarbons (PAHs), which are often carcinogenic. Genetic and biochemical studies of PAH degradation expand our abilities to effectively manage and treat polluted environments and to engineer novel technologies for pollution abatement. Additionally, these studies provide insights into the behavior and evolution of conjugative plasmids, the lateral transfer of genetic information among bacteria, and the diversity and function of oxygenase genes and enzymes.

In all pure cultures of naphthalene-mineralizing bacteria that have been examined, the aerobic metabolism of naphthalene is initiated by a multicomponent enzyme system called naphthalene dioxygenase (NDO). The ability to degrade naphthalene has been demonstrated for a wide range of bacterial genera, and these organisms have been shown to possess a number of distinct NDOs that have widely varying degrees of nucleotide and amino acid sequence similarities.  Frequently, the genes for PAH degradation are located on large conjugative plasmids that range from 60-500 kbp in size. These plasmids are transmitted between a variety of bacteria in polluted environments, thus disseminating the ability to derive carbon and energy from the chemical pollutants.

The vast majority of studies characterizing PAH degradation have examined bacteria residing in soils and freshwater habitats. The few studies focusing on marine and estuarine environments have indicated that distinct bacterial genera, using novel NDO and other gene sequences, are responsible for attenuation of PAHs in these sites. This study examines the genetic basis and molecular diversity of PAH degradation in Humboldt Bay in northern CA, with a focus on the diversity of genes, plasmids and organisms responsible for naphthalene catabolism.

Objectives and Approaches

Objective 1: Isolate and characterize a range of microorganisms from Humboldt Bay that are capable of naphthalene catabolism. Naphthalene-degrading microorganisms will be isolated by plating dilutions of PAH-impacted sediments onto an artificial seawater-based medium supplemented with nitrogen and phosphorous and solidified with purified noble agar.  Naphthalene will be provided in vapor phase. In preliminary studies done last year, 2 undergraduates working with me isolated 50 PAH-degrading bacteria from sediments at the base of a creosote-soaked piling in Humboldt Bay.  The students characterized the morphology, Gram Stain reactions, and biochemical characteristics of the isolates. They cloned and sequenced 16S rRNA genes from 12 of the isolates and demonstrated that they belonged to 4 different taxonomic groups, including Gram positive Rhodococci and Gram negative alpha and gamma proteobacteria. These 50 isolates are stored as frozen stocks. Starting in fall of 2004, two other undergraduates have begun to isolate naphthalene-degrading bacteria from an area of Humboldt Bay that was heavily impacted by a 5000-gallon fuel oil spill in 1997 and again by a 2000-gallon oil spill in 1999.

Objective 2: Characterize the diversity of NDO genes that are present in the Humboldt Bay isolates. DNA will be extracted from naphthalene-degrading isolates and the presence of characterized NDO genes will be examined by attempting to PCR-amplify previously described NDO gene sequences using a suite of primer sets. Some of these primers will be designed by students as part of this project.  PCR products will be cloned, sequenced and compared. DNA from strains not producing a PCR product will be examined in Southern hybridizations using labeled probes produced from strains that are positive for amplification.  We will attempt to clone novel NDO genes from strains that don’t produce a PCR product or hybridize with NDO probes. These cloning attempts will use an NDO-indicative indole-to-indigo color change reaction.  This reaction allows us to directly screen colonies resulting from ligations of size-selected genomic DNA samples partially digested with restriction enzymes. Starting in fall of 2004, two additional undergraduates (not those involved in culturing efforts) have begun these experiments with a subset of the isolates from the creosote site.

Objective 3: Characterize the large conjugative plasmids that are present in the Humboldt Bay isolates. We will identify the presence and size of large plasmids in site isolates using Pulsed-Field Gel Electrophoresis (PFGE).  Transfer of PFGE-separated DNA to nitrocellulose membranes will allow us to assess the plasmid or chromosomal location of NDO genes, and potentially whether multiple copies of NDO genes exist in site organisms. Restriction Fragment Length Polymorphisms of plasmid DNA will be used to assess the relatedness of different plasmids. Future studies will use labeled plasmid DNA as probes to examine plasmid diversity via Southern hybridization.

Future studies will examine the transmission of conjugative plasmids among site isolates and in situ in Humboldt Bay. We will use long-term non-selective culturing to produce plasmid-cured derivatives of a diverse set of naphthalene degraders. Rifampicin-resistant, plasmid-cured derivative strains will be obtained using rifampicin gradient plates.  The host range and transmission characteristics of the plasmids will be studied by using these derivative strains. These derivative strains can also be used to trap and recover conjugative plasmids being transferred in situ in Humboldt Bay. I intend to use these results to determine the relative significance and distribution of the NDO analogs in situ. Additionally, I intend to seek funding for the sequencing and comparison of several of the distinct conjugative plasmids involved in PAH degradation that are discovered during the course of the projects outlined above.

                  Greater understanding of these processes can assist with the development of new technologies and management strategies for use in contaminated field sites. For example, we expect to identify rapidly-transmissible and widely-transmissible plasmids involved in natural attenuation of PAHs in Humboldt Bay.  Comparison of baseline and post-treatment levels of the genes and plasmids identified in these studies can be used to monitor the efficacy of remediation efforts. Additionally, engineering efforts may be brought to bear to increase the rate at which naturally-occurring plasmids are disseminated, and particular plasmids may be engineered for specific applications or enhanced activities. The results will also help us to understand how these plasmids evolve, and the diversity of plasmids and genes present in these environments.

                  A major goal of this project is to engage several undergraduates (4-6 per semester) in independent research projects that train them in techniques important in diverse biotechnological fields.  These techniques include microbiological isolation and culturing techniques, DNA extractions, PCR, PCR primer design, pulsed field gel electrophoresis, molecular cloning, DNA sequencing, RFLP analysis, and Southern hybridizations.  Additionally, these students will be introduced to the fields of biodegradation and bioremediation.

The following abstract is for a poster presentation on this work at the 106th General Meeting of the American Society for Microbiology, Orlando FL May 21-25.

Isolation and Genetic Characterization of Naphthalene-Degrading Bacteria from Humboldt Bay, CA

Donnie L. Carter*, Amber L. Orloff*, Erika L. Kraft* and Mark S. Wilson

The ability of microorganisms to catabolize PAH is frequently the consequence of acquiring large, low-copy-number conjugative plasmids that carry genes for degradative pathways.  This study seeks to understand the diversity of microorganisms, genes and conjugative plasmids responsible for the biodegradation of naphthalene and other PAHs in marine and estuarine systems, with a focus on Humboldt Bay, CA. PAH-degrading bacteria were isolated from creosote-contaminated salt marsh sediments by liquid enrichment with naphthalene followed by plating and single colony isolation on a synthetic seawater medium exposed to naphthalene vapors. Of >50 isolates obtained, six isolates were identified for further study because 16S rRNA sequences, microscopy and biochemical tests indicated that they represented taxonomically diverse groups.  Pulsed field gel electrophoresis (PFGE) was used to identify the size and presence of large plasmids in the isolates. PCR and Southern hybridization were used to identify the presence and location (chromosome or plasmid) of previously characterized dioxygenase genes. Isolates were identified as Rhodococcus opacus, Thalassolituus oleivorans, Chromohalobacter salinarum and Sphingomonas sp. PFGE indicated that the two Sphingomonas isolates each had 2 circular plasmids (~ 220 and 270 kbp), and T. oleivorans had 3 (possibly linear) plasmids (~ 170, 220 and 250 kbp). No plasmids were detected in the Chromohalobacter isolate, and problems with lysing the Gram positive R. opacus isolates have so far led to inconclusive results with those strains. PCR and Southern hybridization indicated that diverse alleles of dioxygenases were present in these strains, that some strains had multiple dioxygenase alleles, and that some of these genes were located chromosomally. Taxonomically diverse strains of PAH-degrading bacteria have been isolated from creosote-contaminated sediments of Humboldt Bay. Characterization of these organisms indicates that diverse large plasmids and diverse genetic systems for PAH degradation are present in microorganisms living in the estuary.

Life in Boiling Acid

 

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