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D C Coleman

Publications and source records attributed to D C Coleman.

At least 19 recordsLinked to original sources

Comparison of Candida dubliniensis and C. albicans based on polar lipid composition.

AIMS: To test the hypothesis that strains of Candida dubliniensis and C. albicans can be differentiated on the basis of polar lipid profiles. METHODS AND RESULTS: Five isolates of C. dubliniensis and six isolates of C. albicans were tested by growth at 45 degrees C, production of chlamydospores on cornmeal agar, colonial colour on CHROMagar Candida medium and assimilation of DL-lactate, alpha-methyl-D-glucoside and xylose. Polar lipids were then extracted from freeze-dried cultures and analysed using fast atom bombardment mass spectrometry. Isolates were grouped by single linkage clustering based on correlation coefficients for strain pairs calculated with carboxylate and phospholipid molecular species distributions. The most intense carboxylate and phospholipid molecular species anions were of m/z 281 (C(18 : 1)) and m/z 515 (PA 23 : 2). Phosphatidylethanolamine and phosphatidylglycerol were the predominant phospholipid families in C. dubliniensis, compared with phosphatidic acid in C. albicans isolates. All of the C. dubliniensis isolates grouped together in one cluster, whereas all of the C. albicans isolates grouped in a separate cluster. CONCLUSIONS: Fast atom bombardment mass spectrometry can differentiate the two species based on analysis of polar lipid distributions. SIGNIFICANCE AND IMPACT OF THE STUDY: These findings demonstrate that C. dubliniensis and C. albicans have distinct polar lipid profiles.

Anions↗

Effective control of dental chair unit waterline biofilm and marked reduction of bacterial contamination of output water using two peroxide-based disinfectants.

Bacterial biofilm in dental unit waterlines (DUWs) is a widespread problem, and poses a potentially significant risk of infection to dental staff and patients, particularly those who are medically compromised or immunocompromised. The purpose of the present study was to investigate the level of bacterial contamination of dental chair unit output water in the Dublin Dental Hospital, and to investigate the efficacy of two hydrogen peroxide-based disinfectants in reducing bacterial loads to < or =200 cfu/mL as recommended by the American Dental Association. The chemical quality of dental chair unit input and output water was well within the limits recommended for potable water. Water supplied to the units yielded an average aerobic heterotrophic bacterial cell density of 184 cfu/mL. However, the corresponding density in output water was considerably higher; the average cell density in water from the three-in-one air/water syringes and cup fillers in 12 chairs was 8200 and 4300 cfu/mL, respectively. Dental unit water obtained from 18 separate reservoir-supplied units in general practices in the Dublin area yielded an average of 66000 cfu/mL. The bacterial species found were predominantly environmental organisms, which were also present at low levels in the input water. Some of the species identified (e.g., Burkholderia cepacia and Pseudomonas fluorescens) are known opportunistic pathogens. The capacity of two disinfectants, Sterilex Ultra and Sanosil, to reduce bacterial contamination to safe levels was compared. In a controlled study, once weekly overnight (15 h) disinfection using either agent reduced the bacterial density to below the American Dental Association recommended level of 200 cfu/mL. However, once disinfection ceased the bacterial loads increased to unacceptably high levels within three weeks. Electron microscopic analysis showed that both disinfectants markedly reduced biofilm in the DUWs, but the biofilm rapidly became extensive again when once weekly disinfection ceased. While both disinfectants were equally effective in lowering the bacterial counts to acceptable levels, Sterilex Ultra was associated with clogging of DUWs in some dental chair units after repeated usage, suggesting that Sanosil is a more suitable agent for routine use.

Biofilms↗

Candida dubliniensis candidaemia in an HIV-positive patient in Ireland.

Candida dubliniensis was first identified in Dublin in 1995 in oral isolates recovered from human immunodeficiency virus (HIV)-infected individuals. Although C. dubliniensis has been primarily recovered from the oral cavities of HIV-infected individuals, the number of reports describing its isolation from HIV-negative individuals, including cases of candidaemia, is growing. To date there has only been one report of C. dubliniensis candidaemia in an HIV-infected patient, in this case from the USA. In the present study, 2 Candida isolates recovered from blood samples were presumptively identified as C. dubliniensis on the basis of their dark green coloration on CHROMagar Candida medium and lack of growth at 45 degrees C. This identification was confirmed by carbohydrate assimilation profile analysis and by polymerase chain reaction (PCR) analysis with C. dubliniensis-specific PCR primers. Both isolates were susceptible to fluconazole. The isolates were found retrospectively to be from a single HIV-infected patient who was receiving broad-spectrum antibacterials at the time of isolation of C. dubliniensis from blood. This study represents the first documented case of C. dubliniensis bloodstream infection in Ireland and is only the second case of C. dubliniensis bloodstream infection identified in an HIV-infected individual anywhere in the world.

Adult↗

A polymeric system for the intra-oral delivery of an anti-fungal agent.

Oral candidal infections are often persistent and intractable and thus the aim of this study was to develop a polymeric sustained release device to improve the topical treatment of these infections. A self curing system based on poly(ethyl methacrylate) and tetrahydrofurfuryl methacrylate (PEM/THFM) was used with chlorhexidine diacetate (CX) added at levels between 0 and 12% w/w. Water uptake by the device was assessed gravimetrically and CX release measured by UV spectrometry. Anti candidal activity was established by culturing azole sensitive and resistant strains of Candida albicans in the presence of the polymeric delivery device with and without CX. Candidal growth was measured by turbidimetry or surviving colony-forming unit (CFU) formation. There was an initial high release of CX over 24 h followed by a slow diffusion up to 7 days. CX inhibited candidal growth and survival markedly in vitro, with the test samples showing less than 0.5 x 10(-7) CFU/ml compared to controls (3-4 x 10(-7) CFU/ml). These results indicate the potential of a chlorhexidine containing PEM/THFM polymeric system in the treatment of persistent candidal infections.

Administration, Oral↗

MDR1-mediated drug resistance in Candida dubliniensis.

Candida dubliniensis is a recently described opportunistic fungal pathogen that is closely related to Candida albicans. Candida dubliniensis readily develops resistance to the azole antifungal agent fluconazole, both in vitro and in infected patients, and this resistance is usually associated with upregulation of the CdMDR1 gene, encoding a multidrug efflux pump of the major facilitator superfamily. To determine the role of CdMDR1 in drug resistance in C. dubliniensis, we constructed an mdr1 null mutant from the fluconazole-resistant clinical isolate CM2, which overexpressed the CdMDR1 gene. Sequential deletion of both CdMDR1 alleles was performed by the MPA(R)-flipping method, which is based on the repeated use of a dominant mycophenolic acid resistance marker for selection of integrative transformants and its subsequent deletion from the genome by FLP-mediated, site-specific recombination. In comparison with its parental strain, the mdr1 mutant showed decreased resistance to fluconazole but not to the related drug ketoconazole. In addition, we found that CdMDR1 confers resistance to the structurally unrelated drugs 4-nitroquinoline-N-oxide, cerulenin, and brefeldin A, since the enhanced resistance to these compounds of the parent strain CM2 compared with the matched susceptible isolate CM1 was abolished in the mdr1 mutant. In contrast, CdMDR1 inactivation did not cause increased susceptibility to amorolfine, terbinafine, fluphenazine, and benomyl, although overexpression of CdMDR1 in a hypersusceptible Saccharomyces cerevisiae strain had previously been shown to confer resistance to these compounds. The effect of CdMDR1 inactivation was identical to that seen in two similarly constructed C. albicans mdr1 mutants. Therefore, despite species-specific differences in the amino acid sequences of the Mdr1 proteins, overexpression of CaMDR1 and CdMDR1 in clinical C. albicans and C. dubliniensis strains seems to confer the same drug resistance profile in both species.

Antifungal Agents↗

Isogenic strain construction and gene targeting in Candida dubliniensis.

Candida dubliniensis is a recently described opportunistic fungal pathogen that is closely related to Candida albicans but differs from it with respect to epidemiology, certain virulence characteristics, and the ability to develop fluconazole resistance in vitro. A comparison of C. albicans and C. dubliniensis at the molecular level should therefore provide clues about the mechanisms used by these two species to adapt to their human host. In contrast to C. albicans, no auxotrophic C. dubliniensis strains are available for genetic manipulations. Therefore, we constructed homozygous ura3 mutants from a C. dubliniensis wild-type isolate by targeted gene deletion. The two URA3 alleles were sequentially inactivated using the MPA(R)-flipping strategy, which is based on the selection of integrative transformants carrying a mycophenolic acid resistance marker that is subsequently deleted again by site-specific, FLP-mediated recombination. The URA3 gene from C. albicans (CaURA3) was then used as a selection marker for targeted integration of a fusion between the C. dubliniensis MDR1 (CdMDR1) promoter and a C. albicans-adapted GFP reporter gene. Uridine-prototrophic transformants were obtained with high frequency, and all transformants of two independent ura3-negative parent strains had correctly integrated the reporter gene fusion into the CdMDR1 locus, demonstrating that the CaURA3 gene can be used for efficient and specific targeting of recombinant DNA into the C. dubliniensis genome. Transformants carrying the reporter gene fusion did not exhibit detectable fluorescence during growth in yeast extract-peptone-dextrose medium in vitro, suggesting that CdMDR1 is not significantly expressed under these conditions. Fluconazole had no effect on MDR1 expression, but the addition of the drug benomyl strongly activated the reporter gene fusion in a dose-dependent fashion, demonstrating that the CdMDR1 gene, which encodes an efflux pump mediating resistance to toxic compounds, is induced by the presence of certain drugs.

ATP Binding Cassette Transporter, Subfamily B, Mem↗

Differentiation of Candida dubliniensis from Candida albicans on staib agar and caffeic acid-ferric citrate agar.

The methods currently available for the identification of the pathogenic yeast Candida dubliniensis all have disadvantages in that they are time-consuming, expensive, and/or, in some cases, unreliable. In a recent study (P. Staib and J. Morschhäuser, Mycoses 42:521-524; 1999) of 14 C. dubliniensis and 11 C. albicans isolates, it was suggested that the ability of C. dubliniensis to produce rough colonies and chlamydospores (chlamydoconidia) on Staib agar (SA) provided a simple means of differentiating it from its close relative C. albicans. In the present investigation, we examined the colony morphology and chlamydospore production of 130 C. dubliniensis and 166 C. albicans isolates on SA and on the related defined medium caffeic acid-ferric citrate agar (CAF). All of the C. dubliniensis and C. albicans isolates produced chlamydospores on the control medium, i.e., rice-agar-Tween agar. However, while none of the C. albicans isolates produced chlamydospores on either SA or CAF, 85.4 and 83.8% of the C. dubliniensis isolates produced chlamydospores on SA and CAF, respectively. All of the C. albicans isolates grew as smooth, shiny colonies on SA after 48 to 72 h of incubation at 30 degrees C, while 97.7% of the C. dubliniensis isolates grew as rough colonies, many (65%) with a hyphal fringe. In contrast, 87.4% of the C. albicans and 93.8% of the C. dubliniensis isolates yielded rough colonies on CAF. Although the results of this study confirm that SA is a good medium for distinguishing between C. dubliniensis and C. albicans, we believe that discrimination between these two species is best achieved on the basis of colony morphology rather than chlamydospore production.

Agar↗

Serological differentiation of experimentally induced Candida dubliniensis and Candida albicans infections.

Using a rabbit model of systemic infection, we show that it is possible to differentiate infections caused by Candida dubliniensis and other Candida species by detecting the antibody response mounted by the infected animals. These results confirm our previous observation in a patient with C. dubliniensis candidemia and suggest that detection of C. dubliniensis-specific antibodies is useful in the diagnosis of invasive candidiasis caused by this yeast.

Animals↗

Isolation of C. dubliniensis from insulin-using diabetes mellitus patients.

The identification of the novel candidal species, C. dubliniensis, from oral swab studies of HIV-seropositive and -seronegative individuals has led to speculation that such a species may also reside in the oral cavity of other patient groups. In this study involvement of the newly described species, C. dubliniensis, was investigated in oral carriage and disease in 414 insulin-using diabetes mellitus patients. Seventy-seven percent of the diabetic patients carried candidal species in the oral cavity. C. albicans was the most commonly identified candidal species. This was followed by C. dubliniensis, which was isolated on 64 occasions. Colonisation with multiple candidal species was common, and C. dubliniensis was present in both carriage and disease states. Seven patients without signs of oral disease had C. dubliniensis isolated as the sole candidal species, while the same species was associated with various forms of oral candidosis in six patients.

Candida↗

A simple and rapid technique for the detection of Epstein-Barr virus DNA in HIV-associated oral hairy leukoplakia biopsies.

A method of generating nucleic acid probes by polymerase chain reaction (PCR) for the detection of Epstein-Barr virus (EBV)-DNA by in situ hybridization in oral hairy leukoplakia (OHL) lesions is described. This method has the advantage over older methods of being cheaper, quicker and retaining sensitivity and specificity. Purified PCR products of Epstein-Barr virus DNA of 110 bp and 328 bp were labelled with biotin by nick translation or random primer labelling and were compared in in situ hybridization experiments with probes prepared by incorporation of biotin-labelled nucleotides in the PCR reaction mixture, with EBV viral DNA as a template. These probes were applied to 18 OHL tongue biopsies known to be positive for EBV-DNA, using a commercially available biotin-labelled BamHI "V" fragment EBV-DNA probe. To determine the specificity of the probes, we applied them to 20 normal tongue tissue samples and to 12 biopsies taken from keratotic tongue lesions from patients without risk factors for HIV infection and known to be negative for EBV-DNA. Clear positive signals for EBV-DNA were detected in all 18 cases of OHL biopsies using the amplimer of 328 bp labelled by PCR and random primer labelling. However, nick translation labelling was less efficient and sensitive. All control specimens were negative for EBV-DNA.

AIDS-Related Opportunistic Infections↗

Emerging pathogens.

The ever increasing numbers of immunosuppressed individuals has led to a significant increase in the incidence of opportunistic infections, particularly those caused by fungi. The epidemiology of infections caused by the common fungal pathogens such as Candida albicans, Cryptococcus neoformans and Aspergillus fumigatus has been well documented. However, in addition to these, a number of species which have previously been unrecognized (e.g., C. dubliniensis) or have previously been assumed to be non-pathogenic (e.g., Saccharomyces cerevisiae, Scedosporium spp. and Fusarium spp.) have emerged as agents of human disease. Since these species have only been identified recently as human pathogens, their role in disease is poorly understood. In most cases, identification of these species is problematic and therefore their epidemiology has yet to be elucidated adequately. In addition, several of these species fail to respond to conventional antifungal therapies. In this article, we describe the emergence of two separate yeast species (C. dubliniensis and S. cerevisiae) and two separate groups of moulds (Scedosporium prolificans and Fusarium spp.), as human pathogens. It is apparent from what we already know, that much work has yet to be performed before we have a clear understanding of how these species cause disease and most importantly how they can be controlled.

Candida↗

Recovery of Candida dubliniensis from non-human immunodeficiency virus-infected patients in Israel.

Candida dubliniensis is a recently discovered yeast species principally associated with carriage and disease in the oral cavities of human immunodeficiency virus (HIV)-infected individuals. To date the majority of isolates of this species have been identified in Europe and North America. In this study, five Candida isolates recovered from separate HIV-negative hospitalized patients in Jerusalem, Israel, were presumptively identified as C. dubliniensis on the basis of their dark green coloration when grown on CHROMagar Candida medium. Their identification was confirmed by a variety of techniques, including carbohydrate assimilation profiles, absence of growth at 45 degrees C, positive reaction with C. dubliniensis-specific antibodies as determined by indirect immunofluorescence analysis, and positive amplification with C. dubliniensis-specific PCR primers. All five strains were shown to be susceptible to a range of antifungal agents, including fluconazole. One of the five isolates was recovered from urine specimens, while the remaining four were recovered from upper respiratory tract and oral samples. While none of the patients was HIV positive, all were receiving broad-spectrum antibacterials at the time isolates of C. dubliniensis were obtained from clinical specimens. This study describes the first isolates of C. dubliniensis from the Middle East and confirms that this yeast can be associated with carriage and infection in the absence of HIV infection.

Adult↗

Lack of a relationship between Lewis antigen expression and cagA, CagA, vacA and VacA status of Irish Helicobacter pylori isolates.

The cagA gene, vacA gene, CagA (cytotoxin-associated gene A product) and VacA (vacuolating cytotoxin) status of a collection of Helicobacter pylori isolates from the geographically distinct Irish population was determined, the potential association of these traits with Lewis (Le) antigen expression was assessed, and the relationship between these bacterial properties and the pathology associated with H. pylori infection was evaluated. Of the 57 isolates, a higher proportion from ulcer than from non-ulcer patients expressed VacA (71% vs. 53%). H. pylori isolates which were cagA-positive were no more significantly associated with peptic ulcers than non-ulcer disease (71% vs. 67%, P = 0.775), nor were CagA-positive isolates (57% vs. 50%, P = 0.783), but 80% of the isolates from duodenal ulcer patients were cagA-positive. Thirty-seven of the 57 isolates were tested for Le antigen expression. No statistically significant relationship (P > 0.05) was found between the occurrence and level of expression of Le(x) or Le(y) and cagA, vacA, or VacA status. This lack of an association in the Irish H. pylori isolates contrasts with that previously reported for predominantly North American isolates, and may be attributable to the adaptation of H. pylori strains with differing attributes to different human populations.

Antigens, Bacterial↗

Rapid identification of Candida dubliniensis with commercial yeast identification systems.

Candida dubliniensis is a newly described species that is closely related phylogenetically to Candida albicans and that is commonly associated with oral candidiasis in human immunodeficiency virus-positive patients. Several recent studies have attempted to elucidate phenotypic and genotypic characteristics of use in separating the two species. However, results obtained with simple phenotypic tests were too variable and tests that provided more definitive data were too complex for routine use in the clinical laboratory setting. The objective of this study was to determine if reproducible identification of C. dubliniensis could be obtained with commercial identification kits. The substrate reactivity profiles of 80 C. dubliniensis isolates were obtained by using the API 20C AUX, ID 32 C, RapID Yeast Plus, VITEK YBC, and VITEK 2 ID-YST systems. The percentages of C. dubliniensis isolates capable of assimilating or hydrolyzing each substrate were compared with the percentages from the C. albicans profiles in each kit's database, and the results were expressed as percent C. dubliniensis and percent C. albicans. Any substrate that showed >50% difference in reactivity was considered useful in differentiating the species. In addition, assimilation of methyl-alpha-D-glucoside (MDG), D-trehalose (TRE), and D-xylose (XYL) by the same isolates was investigated by the traditional procedure of Wickerham and Burton (L. J. Wickerham and K. A. Burton, J. Bacteriol. 56:363-371, 1948). At 48 h (the time recommended by the manufacturer for its new database), we found that the assimilation of four carbohydrates in the API 20C AUX system could be used to distinguish the species, i.e., glycerol (GLY; 88 and 14%), XYL (0 and 88%), MDG (0 and 85%), and TRE (15 and 97%). Similarly, results with the ID 32 C system at 48 h showed that XYL (0 and 98%), MDG (0 and 98%), lactate (LAT; 0 and 96%), and TRE (30 and 96%) could be used to separate the two species. Phosphatase (PHS; 9 and 76%) and alpha-D-glucosidase (23 and 94%) proved to be the most useful for separation of the species in the RapID Yeast Plus system. While at 24 h the profiles obtained with the VITEK YBC system showed that MDG (10 and 95%), XYL (0 and 95%), and GLY (26 and 80%) could be used to separate the two species, at 48 h only XYL (6 and 95%) could be used to separate the two species. The most useful substrates in the VITEK 2 ID-YST system were TRE (1 and 89%), MDG (1 and 99%), LAT (4 and 98%), and PHS (83 and 1%). While the latter kit was not yet commercially available at the time of the study, it would appear to be the most valuable for the identification of C. dubliniensis. Although assimilation of MDG, TRE, and XYL proved to be the most useful for species differentiation by the majority of commercial systems, the results with these carbohydrates by the Wickerham and Burton procedure were essentially the same for both species, albeit following protracted incubation. Thus, it is the rapidity of the assimilation achieved with the commercial systems that allows the differentiation of C. dubliniensis from C. albicans.

AIDS-Related Opportunistic Infections↗

Identification of Candida dubliniensis based on temperature and utilization of xylose and alpha-methyl-D-glucoside as determined with the API 20C AUX and vitek YBC systems.

To have a better understanding of the role of Candida dubliniensis in clinical infections, it is essential that microbiology laboratories can identify this species rapidly and accurately in clinical specimens. C. dubliniensis has been reported to lack the ability to utilize xylose (XYL) and alpha-methyl-D-glucoside (MDG) and to grow poorly or not at all at 45 degrees C, whereas Candida albicans isolates utilize XYL and MDG and usually grow well at 45 degrees C. We tested 66 isolates of C. dubliniensis and 100 isolates of C. albicans with both the API 20C AUX and Vitek YBC systems to evaluate the ability of the XYL and MDG tests contained within each of these systems to distinguish between the two species. The ability to grow at 45 degrees C was also examined. None of the C. dubliniensis isolates grew at 45 degrees C, and 23 of 100 C. albicans isolates (23%) exhibited poor or no growth at 45 degrees C. The XYL and MDG tests contained within the API 20C AUX system were both negative for all 66 C. dubliniensis isolates and were positive for 98 (XYL) and 56 (MDG) of the 100 C. albicans isolates. With the Vitek system, 64 of 66 C. dubliniensis isolates (97.0%) were XYL negative and 63 (95.0%) were MDG negative. Conversely, 96 of 100 C. albicans isolates (96.0%) were XYL positive and 100 (100.0%) were MDG positive with the Vitek system. Clinical microbiology laboratories could use lack of growth at 45 degrees C and a negative XYL test with either the API 20C AUX or Vitek yeast identification system to provide a presumptive identification of C. dubliniensis. A negative MDG test result with either system would also be helpful but may misclassify C. albicans as C. dubliniensis, especially when the API 20C AUX system is used.

Candida↗

In vitro susceptibilities of Candida dubliniensis isolates tested against the new triazole and echinocandin antifungal agents.

Candida dubliniensis is a newly recognized fungal pathogen causing mucosal disease in AIDS patients. Although preliminary studies indicate that most strains of C. dubliniensis are susceptible to established antifungal agents, fluconazole-resistant strains have been detected. Furthermore, fluconazole-resistant strains are easily derived in vitro, and these strains exhibit increased expression of multidrug resistance transporters, especially MDR1. Because of the potential for the development of resistant strains of C. dubliniensis, it is prudent to explore the in vitro activities of several of the newer triazole and echinocandin antifungals against isolates of C. dubliniensis. In this study we tested 71 isolates of C. dubliniensis against the triazoles BMS-207147, Sch 56592, and voriconazole and a representative of the echinocandin class of antifungal agents, MK-0991. We compared the activities of these agents with those of the established antifungal agents fluconazole, itraconazole, amphotericin B, and 5-fluorocytosine (5FC) by using National Committee for Clinical Laboratory Standards microdilution reference methods. Our findings indicate that the vast majority of clinical isolates of C. dubliniensis are highly susceptible to both new and established antifungal agents. Strains with decreased susceptibilities to fluconazole remained susceptible to the investigational agents as well as to amphotericin B and 5FC. The increased potencies of the new triazole and echinocandin antifungal agents may provide effective therapeutic options for the treatment of infections due to C. dubliniensis.

AIDS-Related Opportunistic Infections↗

Rapid PCR test for discriminating between Candida albicans and Candida dubliniensis isolates using primers derived from the pH-regulated PHR1 and PHR2 genes of C. albicans.

The development of a satisfactory means to reliably distinguish between the two closely related species Candida albicans and Candida dubliniensis in the clinical mycology laboratory has proved difficult because these two species are phenotypically so similar. In this study, we have detected homologues of the pH-regulated C. albicans PHR1 and PHR2 genes in C. dubliniensis. Restriction fragment length polymorphism analysis suggests that there are significant sequence differences between the genes of the two species. In order to exploit this apparent difference, oligonucleotide primers based on the coding sequence of the C. albicans PHR1 structural gene were designed and used in PCR experiments. Use of these primers with C. albicans template DNA from 17 strains yielded a predicted 1.6-kb product, while C. dubliniensis template DNA from 19 strains yielded no product. We therefore propose that PCR using these primers is a rapid and reliable means of distinguishing the two germ tube- and chlamydospore-producing species C. albicans and C. dubliniensis.

Apoenzymes↗

Prokaryotes: the unseen majority.

The number of prokaryotes and the total amount of their cellular carbon on earth are estimated to be 4-6 x 10(30) cells and 350-550 Pg of C (1 Pg = 10(15) g), respectively. Thus, the total amount of prokaryotic carbon is 60-100% of the estimated total carbon in plants, and inclusion of prokaryotic carbon in global models will almost double estimates of the amount of carbon stored in living organisms. In addition, the earth's prokaryotes contain 85-130 Pg of N and 9-14 Pg of P, or about 10-fold more of these nutrients than do plants, and represent the largest pool of these nutrients in living organisms. Most of the earth's prokaryotes occur in the open ocean, in soil, and in oceanic and terrestrial subsurfaces, where the numbers of cells are 1.2 x 10(29), 2.6 x 10(29), 3.5 x 10(30), and 0. 25-2.5 x 10(30), respectively. The numbers of heterotrophic prokaryotes in the upper 200 m of the open ocean, the ocean below 200 m, and soil are consistent with average turnover times of 6-25 days, 0.8 yr, and 2.5 yr, respectively. Although subject to a great deal of uncertainty, the estimate for the average turnover time of prokaryotes in the subsurface is on the order of 1-2 x 10(3) yr. The cellular production rate for all prokaryotes on earth is estimated at 1.7 x 10(30) cells/yr and is highest in the open ocean. The large population size and rapid growth of prokaryotes provides an enormous capacity for genetic diversity.

Prokaryotic Cells↗