System. App!. Microbiol. 18,549-559 (1995)
© Gu;;tav Fischer Verlag· Stuttgart· .lena· New York
Phylogenetic Analysis of Seven New Isolates of Ammonia-Oxidizing
Bacteria Based on 16S rRNA Gene Sequences
JANNE B. UTAKER1, LARS BAKKEN
2
, QING QIAO JIANG
2
, and INGOLF F. NES
1
1 Laboratory of Microbial Gen'e Technology, Department of Biotechnology, Agricultural University of Norway, As, Norway
Section of Microbiology, Department of Biotechnology, Agricultural University of Norway, As, Norway
Received June 23, 1995
Summary
In this report 165 rRNA gene sequences from seven new isolates of the genus Nitrosospira were deter-
mined, and used in a phylogenetic analysis. The homology values for these 16S rDNA sequences ranged
from 97.4% to complete homology. In general, the three closest related genera of ammonia-oxidizers -
Nitrosospira, Nitrosoiobus and Nitrosovibrio - have 165 rDNA sequences with homology values from
96.9%, giving low resolution when the taxa are positioned in a phylogenetic tree. Thus, while these
ammonia-oxidizers possess distinct morphological differences; these differences seem not to be reflected in
the 165 ribosomal DNA sequences. This suggests that the morphological traits have evolved recently, and
that 16S ribosomal RNA analysis alone is not sufficient to determine the evolutionary relationship be-
tween them. The results of this phylogenetic analysis confirm the previously proposed emendation of the
genus classification within these bacteria.
Key words: 165 rRNA - Phylogeny - Ammonia-oxidizing bacteria - PCR
Introduction
The ammonia-oxidizing bacteria constitute a group of
obligate chemoautotrophs which use carbon dioxide as
the sole source of carbon, and obtain energy from oxida-
tion of ammonia to nitrite (Watson et aI., 1989). Five
genera of ammonia-oxidizers have been characterized;
Nitrosomonas (Winogradsky, 1892), Nitrosococcus
(Winogradsky, 1892). Nitrosospira (Winogradsky and
Winogradsky, 1933), Nitrosolobus (Watson et aI., 1971)
and Nitrosovibrio (Harms et aI., 1976), being classified as
nitrifying bacteria and belonging to the Nitrobacteraceae
family. The nitrifiers are found in most aerobic environ-
ments where organic matter is mineralized, and have been
isolated from soil, freshwater, brackish water, seawater,
and sewage disposal system (Watson et aI., 1989).
The earliest classification of ammonia-oxidizers was
based on morphological characters, and the progress re-
garding comparative studies on physiology, genetics and
phylogeny of this group has been slow (MacDonald,
1986). One obvious reason for this is the fact that isolation
and maintenance of ammonium-oxidizing bacteria is both
difficult and time consuming. The difficulties are con-
nected to elimination of heterotrophic organisms during
isolation, since the ammonia-oxidizers have very low
growth rates (8-24 hrs generation time) and
Harms, 1985), and a low cell yield per mol substrate
Uiang, 1995). Hence, a culture seldom grows to visible
turbidity unless excessive amounts of substrate is added
Uiang, 1995). High ammonium concentrations are not re-
commended, however, due to the potential toxicity of am-
monium and of the nitrite formed during growth (Schmidt
and Belser, 1982). The ability to make stock cultures of
ammonia-oxidizers is also limited, because of low survival
both after preserveration by freeze drying and by freezing
at -80°C Uiang, 1995). Because of the low availability of
ammonium-oxidizers, molecular biology techniques are
well suited for studies on these bacteria.
The ammonia-oxidizers have been found to have very
similar physiology (Koops and Moiler, 1992), and the
DNA base composition (mol% G+C) among the genera
of ammonia-oxidizers varies within the limited range of 45
to 56% GC (Koops and Harms, 1985). A study on lipid
profiles (Blumer et aI., 1969), has also concluded that