The Planctomycetes, Verrucomicrobia, Chlamydiae and sister
phyla comprise a superphylum with biotechnological and
medical relevance
Michael Wagner and Matthias Horn
In the rRNA-based tree of life four bacterial phyla, comprising
the Planctomycetes, Verrucomicrobia, Chlamydiae and
Lentisphaerae, form together with the candidate phyla
Poribacteria and OP3 a monophyletic group referred to as the
PVC superphylum. This assemblage contains organisms that
possess dramatically different lifestyles and which colonize
sharply contrasting habitats. Some members of this group are
among the most successful human pathogens, others are
abundant soil microbes, and others still are of major
importance for the marine nitrogen cycle and hold much
promise for sustainable wastewater treatment. Recent
comparative genomic and metagenomic analyses of a few
representatives of this group revealed many unusual features
and generated unexpected hypotheses regarding their
physiology, some of which have already been confirmed
experimentally. Furthermore, the availability of these genome
sequences offered new insights into the evolutionary history of
this peculiar group of microbes with major medical, ecological
and biotechnological relevance.
Addresses
Department of Microbial Ecology, University of Vienna, 1090 Vienna,
Austria
Corresponding author: Wagner, Michael (wagner@microbial-
ecology.net)
Current Opinion in Biotechnology 2006, 17:241249
This review comes from a themed issue on
Environmental biotechnology
Edited by David A Stahl and Michael Wagner
Available online 15th May 2006
0958-1669/$ see front matter
# 2006 Elsevier Ltd. All rights reserved.
DOI 10.1016/j.copbio.2006.05.005
Introduction
The most recent version of the bacterial tree of life
consists of at least 50 phyla [1]. Owing to difficulties in
the reconstruction of ancient relationships among bacter-
ial phyla, however, their branching order in this tree
remains largely unresolved. An interesting exception to
this mistiness is the PVC superphylum an assemblage
of four bacterial phyla together with the Poribacteria and
OP3 candidate phyla (which contain no cultured rela-
tives), which is consistently recovered as a monophyletic
group in 16S rRNA trees with different treeing methods
www.sciencedirect.com
(Figure 1). At first glance, this grouping is unexpected
because no obvious similarities between the different
phyla, which would reflect their shared evolutionary
history, are apparent. Furthermore, bootstrap support
for this superphylum is not very high (Figure 1) and there
are a few reports in which some of its members do not
cluster together [24]. However, owing to limited data at
the time of their publication, these reports did not include
all postulated phyla of the PVC superphylum, which
might have resulted in biased tree topologies. Further-
more, a closer inspection of the available literature,
including all recently published genome sequences of
members of the PVC superphylum, reveals several unex-
pected similarities which lend additional support to the
monophyly of this group. A better understanding of the
evolution and biology of the bacteria of the PVC super-
phylum is of particular importance, because it encom-
passes organisms of major medical or environmental
relevance. Furthermore, this grouping includes the
recently discovered anaerobic ammonia oxidizers, which
are increasingly exploited for the cost-effective removal
of nitrogen compounds in sewage treatment.
The first part of this review summarizes separately for
each phylum selected research highlights published since
January 2004 (some earlier publications are also cited if
required to provide context), while the second part aims
to provide a synthesis of these findings in an evolutionary
context.
Planctomycetes
Of all phyla grouped in the PVC superphylum, the
Planctomycetes are most frequently represented in the
current 16S rRNA Ribosomal Database Project (RDP)
database (release 9.38) and make up 0.8% of all depos-
ited sequences. This relatively low number has to be
interpreted with caution, because the 16S rRNA genes of
Planctomycetes do have mismatches to some PCR primers
that are widely used in environmental diversity surveys
[5]. Although this feature might lead to underrepresenta-
tion of the Planctomycetes in environmental 16S rRNA
libraries, the past few years have nonetheless seen a flood
of papers reporting the detection of Planctomycetes in
aquatic and terrestrial ecosystems. For example, the
Planctomycetes was the most numerous bacterial group
in an acidic Sphagnum peat bog detectable by fluorescence
in situ hybridization [6] and they were also found in
hot springs [7]. As for the other phyla within the PVC
superphylum, some Planctomycetes live together with
Current Opinion in Biotechnology 2006, 17:241249
242 Environmental biotechnology

Figure 1

The Planctomycetes/Verrucomicrobia/Chlamydiae (PVC) superphylum. Relationships between the phyla Planctomycetes, Verrucomicrobia,
Chlamydiae, Lentisphaerae and OP3, and their major subgroups as inferred from comparative 16S rRNA sequence analysis using maximum
likelihood (AxML, PhyML), maximum parsimony (DNAPARS), and neighbour joining for tree reconstructions. The monophyly of all five phyla was
supported by all treeing methods applied. A consensus tree based on the obtained maximum likelihood tree is shown. Only the position of the
Poribacteria remained unresolved, and the position of the Verrucomicrobia subdivision 5 was not monophyletic with the Verrucomicrobia with
all treeing methods (dotted lines). Verrucomicrobia subdivision 6, as well as other lineages represented by single or only a few environmental
sequences, have not been included in the analysis. Bootstrap values calculated with PhyML (500 resamplings, using a reduced dataset),
DNAPARS (100 resamplings) and estimated with the respective tool implemented in ARB are indicated for each phylum and for major internal
nodes. A selection of representatives of seven bacterial phyla was used as outgroup, and a filter considering only those positions that are
conserved in at least 50% of all Bacteria was applied in all calculations. Bar, 10% estimated evolutionary distance. Note that a consistent tree
topology (without the Poribacteria sequences which were not available at that time) has recently been published by [89].

eukaryotes. For example, members of the genus Blasto-
pirellula [8] were isolated from the giant tiger prawn [9]
and 16S rRNA sequences of Planctomycetes were also
detected in a termite gut [10] and in sponges [11].
The aerobic marine organism Rhodopirellula baltica is the
first planctomycete for which a complete genome
sequence (7.15 MB) was determined [12]. Surprisingly,
despite its large size this genome contains a relatively
small pool of transcriptional regulators [13] but carries 110
genes encoding sulfatases. It is believed that these
enzymes are important for the metabolism of sulfated
carbohydrates, such as chondroitin, by R. baltica. These
enantioselective sulfatases [14] might also become bio-
technologically important for the quantitative production
of single stereoisomers from racemates, as they were
shown to possess a stereocomplementary retaining activ-
ity [15]. Recently, a two-dimensional reference map of
soluble proteins of R. baltica was established and 558
genes were shown to be expressed by protein spot iden-
tification [16]. The reference map was also used to
reconstruct the central catabolic routes of R. baltica [17]

Current Opinion in Biotechnology 2006, 17:241249 www.sciencedirect.com


and to investigate the growth phase dependent regulation
of protein expression in this organism [18].
Unexpectedly, R. baltica and several other planctomy-
cetes encode most (or all, in the case of Gemmata obscur-
iglobus) of the 16 proteins that are involved in
methanopterin- and methanofuran-mediated C1 transfers
and which are shared between archaeal methanogens and
bacterial methylotrophs [12,19,20]. Furthermore, pro-
teome analysis of R. baltica, which cannot grow on com-
mon C1 compounds, demonstrated expression of some of
these genes [19]. Moreover, the functionality of some of
these genes has been confirmed by their expression in the
heterologous host Methylobacterium extorquens [20]. These
interesting results complicate our perception of the evo-
lutionary history of these C1 tranfer genes [19,20,21] and
raise the question of what function these genes fulfil in
Planctomycetes. Recently, it has been demonstrated that
Gemmata sp. strain Wa1-1 has a novel and functional
methylene tetrahydromethanopterin dehydrogenase
but, as for R. baltica, does not contain serine-cycle genes.
This observation suggests that the methanopterin-linked
C1 transfer reactions might be involved in formaldehyde
detoxification [22].
Besides R. baltica, the so-called anammox bacteria, a deep
branching monophyletic group of the Planctomycetes [23]
capable of anaerobically oxidizing ammonium, have been
intensively studied over the past few years (for recent
reviews see [24,25]). Anammox metabolism takes place in
the anammoxosome [26], the membrane of which is
composed of lipids with peculiar staircase-like ladderane
hydrocarbon chains that comprise three or four linearly
concatenated cyclobutane structures [27,28,29]. These
anammox bacteria have yet to be obtained in pure culture,
but can be enriched (e.g. in sequencing batch and gas lift
reactors). Furthermore, molecular [30] tools and biomar-
kers are available to detect and identify them in natural or
man-made systems [31,32]. Using these approaches, ana-
mmox bacteria have been found in many marine and
freshwater ecosystems and were estimated to contribute
up to 50% of oceanic nitrogen loss [33]. Furthermore, the
anammox process is currently implemented in water
treatment for the low-cost removal of ammonia from
high-strength waste streams and is an impressive example
of the value of basic microbiological research for civil
engineering and society in general (e.g. [3436]).
In 2006 the almost-complete genome of Candidatus
Kuenenia stuttgartiensis, which was assembled in the
framework of a six year project using an environmental
genomics approach from a reactor community genome,
was published [37]. On the basis of this genome
sequence, refined models for anammox catabolism and
CO2 fixation were inferred. Furthermore, the genome
suggests that anammox organisms are much more versa-
tile than previously thought and could, for example, use
www.sciencedirect.com
The PVC superphylum Wagner and Horn 243
organic acids to convert nitrate and nitrite into dinitrogen
gas. This hypothesis was also confirmed experimentally
[37,38] and, in the meantime, a new anammox organism
that co-oxidizes propionate and ammonium has been
identified [30].
Chlamydiae
In contrast to all other members of the PVC superphylum,
the Chlamydiae have been studied extensively for dec-
ades. Chlamydiae are obligate intracellular bacteria char-
acterized by an unique developmental cycle and, as such,
are well-known pathogens of animals and humans. They
cause several severe diseases such as trachoma, genital
tract infections and pneumonia, with millions of infec-
tions reported each year. It was thus surprising that, in
addition to the medically important chlamydiae (grouped
into the family Chlamydiaceae), a large diversity of pre-
viously unrecognized chlamydiae was recently discovered
in the environment. As pathogenic chlamydiae have been
well covered in recent reviews (e.g. [3941]), we focus
here on environmental chlamydiae. These chlamydiae
haven been found associated with amoebae, arthropods,
crustaceans, fish and as contaminants of a human cell
culture ([42,43,44] and earlier work reviewed in [45]).
Like the Chlamydiaceae, they replicate within eukaryotic
host cells and show a biphasic developmental cycle.
Owing to their obligate intracellular lifestyle and the lack
of a genetic system for chlamydiae, genome sequencing
has been of major importance for the recent progress in
chlamydia research [40,46]. In addition to the ten Chla-
mydiaceae genomes available to date, the first genome
sequence of an environmental chlamydia strain, the
amoeba symbiont Protochlamydia amoebophila UWE25
(Parachlamydiaceae), was recently determined [47]. Com-
parison of the genomes of classical and environmental
Chlamydiae, which diverged about 700 million years
ago, suggested that their last common ancestor already
lived intracellularly and that chlamydiae have been asso-
ciated with eukaryotic host cells since primordial times.
Most remarkably, several virulence-associated proteins
of the modern Chlamydiaceae, like the type three secre-
tion system or the chlamydial protease-like activity
factor [48], originate from the chlamydial ancestor and
were most likely developed during its interaction with
ancient unicellular eukaryotes. Furthermore, the high
number of proteins found in the P. amoebophila genome
showing greatest similarity with plant (plastid) proteins
(5%) suggests that the chlamydial ancestors might also
have been associated with plants at some time during
evolution [47]. Further evidence for this hypothesis
comes from the analysis of nucleotide transport proteins,
a class of proteins that until recently seemed to be
restricted to chlamydiae, rickettsiae and plants (plastids)
and whose function seemed to be limited to the
exchange of bacterial/plastidic ADP with ATP from
the host cell.
Current Opinion in Biotechnology 2006, 17:241249
244 Environmental biotechnology
Phylogenetic analysis of these integral membrane pro-
teins suggested that they were invented in the chlamydial
evolutionary lineage and were subsequently passed to
rickettsiae and plants or cyanobacteria by lateral gene
transfer [4951]. Biochemical analysis of all five nucleo-
tide transporter isoforms encoded in the P. amoebophila
genome revealed that they also catalyse the import of
NAD+ and nucleotides other than ATP from the host
cytosol, and revealed a tight coupling of symbiont and
host metabolism by means of these proteins [52,53].
There is some evidence that the chlamydial symbionts
might also be able to infect humans [54,55] and nucleo-
tide transporters might thus represent a promising target
for future antichlamydia therapy.
Verrucomicrobia
Verrucomicrobia are widely distributed and in the period
reviewed here 16S rRNA sequences of members of this
phylum were retrieved from various soils (e.g. see
[56,57]), confirming their importance in these systems.
It has been estimated that Verrucomicrobia comprise upto
10% of the total bacteria in soil and contribute up to 9.8%
of the bacterial 16S rRNA in these systems [58]. Although
most members of this phylum remain uncultivated, Jans-
sens group recently isolated representatives of subdivi-
sions 2 [58,59] and 3 [59,60] and showed that for
Verrucomicrobia isolation from soil liquid serial dilution
is inferior to solid media [61].
Members of the Verrucomicrobia phylum were also
recently detected in bogs [6,62] and in a mesocosm
simulating the flooding of an unplanted paddy soil [63].
In the latter experiment, the late appearance of Verruco-
microbia after flooding suggested that they were adapted
to low substrate concentrations and can be regarded as K-
strategists. 16S rRNA sequences affiliated with the Ver-
rucomicrobia were also obtained from aquatic systems,
including drinking water [64], lake mesocosms [65,66],
lakes [67,68], marine sediments [69], and even from hot
springs [7]. Furthermore, Verrucomicrobia thrive in man-
made ecosystems such as acid rock drainage [70] and
leachate from a municipal solid waste landfill [71] and in
various reactor systems [72,73].
Interestingly, several Verrucomicrobia live in association
with eukaryotes, a feature that might explain why the yet
unpublished genome of Verrucomicrobium spinosum carries
genes for the non-flagellar type III secretion system [74],
a protein secretion apparatus known to mediate the
interaction between bacteria and eukaryotic cells. For
example, Verrucomicrobia were isolated from the gut of the
sea cucumber Stichopus japonicus [75] and were also
detected and cultivated from termite guts [10,76] and
the human intestine [77,78], where they contribute to
mucine degradation. Furthermore, Verrucomicrobia live as
obligate endosymbionts in ectoparasitic nematodes of the
genus Xiphinema, which feed on fruit crops. Nematodes
Current Opinion in Biotechnology 2006, 17:241249
and bacteria have co-evolved for about 150 million years
and symbiont transmission seems to be strictly vertical via
the female host [79]. Other Verrucomicrobia live as ecto-
symbionts on hypotrich ciliates of the genus Euplotidium
and defend their host against predation in a unique way
by being ejected [80]. Unexpectedly, microtubulin-like
structures have been observed in these bacteria [81], an
observation that is consistent with the recent detection of
the btubA and btubB genes in the genome of the verru-
comicrobium Prosthecobacter dejongeii [82]. P. dejongeii
encodes proteins with a higher sequence homology and
greater structural similarity to eukaryotic tubulin than to
its their bacterial homologue FtsZ [83]. BtubA and BtubB
assemble after heterologous expression in vitro to form
long protofilament bundles, although all attempts to
visualize these structures in Prosthecobacter by electron
microscopy have so far been unsuccessful [84].
Poribacteria, Lentisphaerae, the OP3
phylum and other environmental lineages
Poribacteria were detected in 2004 in marine demos-
ponges and, to date, remain unknown from any other
environmental niche [85]. No pure culture of Poribacteria
is available but, according to specific fluorescence in situ
hybridization analysis, they can occur in high numbers in
these sponges. Furthermore, indirect evidence exists that
Poribacteria, like Planctomycetes, possess a compartmented
cytoplasm [85]. Recently, a 39 kb genomic fragment from
an uncultured poribacterium was published [86]. This
fragment contained the 16S rRNA gene, but this gene was
found to be unlinked to other rRNA genes, a feature that
was previously observed in some but not all Planctomycetes
[12,87,88]. Furthermore, this genome fragment carried
two other genes with high sequence homology to R.
baltica, lending additional weight to the inclusion of
the Poribacteria in the PVC superphylum.
The Lentisphaerae phylum was proposed in 2004 based on
the isolation of two phylogenetically distinct marine
strains producing transparent exopolymers from coastal
seawater by dilution to extinction in low-nutrient sea-
water media. Dot-blot hybridization demonstrated that
these strains make up less than 1% of the bacterial
community in the Pacific and Atlantic Ocean [89].
The Lentisphaerae phylum also encompasses the former
candidate phylum VadinBE97 and harbours several
organisms that live in association with eukaryotes includ-
ing cows, termites and the Pompeii worm, Alvinella
pompejana, which thrives at hydrothermal vents in the
Pacific. Interestingly, the Lentisphaerae phylum also con-
tains Victivallis vadensis, a strictly anaerobic cellobiose-
degrading isolate from human faeces [90].
The OP3 candidate phylum contains no cultured bacteria
and was proposed in 1998 on the basis of several 16S
rRNA gene sequences retrieved from Obsidian Pool in
the Yellowstone National Park [91]. In addition to this
www.sciencedirect.com

candidate phylum, which belongs to the PVC superphy-
lum, the candidate phylum WWE2 from a municipal
anaerobic sludge digester [73] has also been reported
to be associated with members of the superphylum. With
our dataset and treeing methods, however, we were
unable to confirm this association.
Evolutionary history of the PVC superphylum
As discussed above, many members of the superphylum
live in close association with eukaryotes and, for at least
two phyla (Chlamydia and Poribacteria), no free-living
members have been described. Therefore, it is tempting
to speculate that the last common ancestor of the super-
phylum already possessed genes that could easily be
modified to successfully colonize or invade eukaryotic
hosts. First hints for possible candidates genes were
obtained by extracting from all published genomes of
Planctomycetes and Chlamydiae a list of proteins that are
(almost) exclusively found in these phyla [37]. Surpris-
ingly, this analysis revealed that nucleotide transporter
genes are not only present in all sequenced Chlamydiae,
but also in the genome of the planctomycete R. baltica
(but not in Kuenenia stuttgartiensis). Consequently, nucleo-
tide transporters might represent an ancient trait of this
superphylum and an example for pre-adaptation of its last
common ancestor to a symbiotic lifestyle. Possibly this
ancestor, like the Planctomycetes and the Poribacteria
[92], also possessed intracellular membrane compart-
mentation and used nucleotide transporters for nucleo-
tide or NAD+ transport between the compartments. Some
evolving groups like the Chlamydiae then lost the intra-
cellular compartments, but managed to relocate the
nucleotide transporters to their cytoplasmic membrane
to tap the nucleotide pool of their eukaryotic host cells. If
this scenario is correct, we would expect the transporter of
R. baltica to be functional and located in an intracellular
membrane and other members of the discussed phyla
(e.g. the endosymbiotic Verrucomicrobia) to make use of
such nucleotide transporters.
In any case, one can add the nucleotide transporters (and
the other superphylum signature proteins found by Strous
and coworkers [37]) to the rapidly growing list of recog-
nized shared features of the superphylum members, thus
lending rRNA-independent support to the common
ancestry of this group. For example, all available genomes
of Planctomycetes (including the anammox bacterium
K. stuttgartiensis and the unpublished genome of Gemmata
sp. Wa-1), Chlamydiae and the unpublished genome of the
verrucomicrobium Prosthecobacter dejongeii lack the bac-
terial tubulin homologue FtsZ, which is highly conserved
among prokaryotes and found in almost all bacterial
groups and the Euryarchaeota [93]. As FtsZ is essential
for the division of most prokaryotic cells, it will be
interesting to examine whether one of the above-men-
tioned signature proteins replaces its function within
this superphylum. Furthermore, the Chlamydiae and
www.sciencedirect.com
The PVC superphylum Wagner and Horn 245
Planctomycetes lack significant amounts of peptidoglycan
in their cell wall, although genome studies revealed that
they descended from an ancestor that obviously possessed
a classical Gram-negative cell wall [37,47,94]. This is
consistent with the observation that some members of the
superphylum, like the Verrucomicrobia, possess a typical
peptidoglycan cell wall. Furthermore, many members of
the superphylum have a complex life cycle and, as men-
tioned above, the Poribacteria and the Planctomycetes both
have intracellular compartments. In addition, the verruco-
microbium P. dejongeii and the planctomycete Gemmata sp.
WA-1 share four eukaryotic signature proteins, which were
most probably laterally acquired (see below) [95].
The Planctomycetes and Verrucomicrobia possess several
phenotypic and molecular features that were considered
to be typical for eukaryotes; for example, some Verruco-
microbia carry genes encoding tubulin-like proteins. In
Planctomycetes the chromosomal DNA is surrounded by at
least one intracytoplasmic membrane and in the case of
Gemmata obscuriglobus the bacterial chromosome is even
enveloped in two membranes, analogous to the eukar-
yotic nuclear body which was previously considered to
represent a unique structure of eukaryotes [92].
Furthermore Gemmata produces primitive sterols [96].
Consequently, some researchers hypothesised that an
ancestor of the Planctomycetes and Verrucomicrobia might
have been a possible candidate as progenitor of the
eukaryotic lineage (for a review on the current state of
the discussion see [92]). However, the recent finding
that only a low number of eukaryotic signature proteins
can be detected in P. dejongeii, Gemmata sp. WA-1 and
K. stuttgartiesis [37,95] refutes this hypothesis. Consis-
tently, supertrees based on concatenated amino acid
sequences of phylogenetic marker molecules obtained
from genome studies showed that Planctomycetes and
Chlamydiae are phylogenetically related [37,97,98] and
that this group is not at the root of the bacterial tree [37],
as was previously suggested [99]. The presence of eukar-
yotic genes in many members of the superphylum is
thus most likely explained by lateral gene transfer and
there is accumulating data in support of this hypothesis
[83,95,96,100]. In this context, it is important to keep in
mind the (possibly ancient) association of many super-
phylum members with eukaryotes that might have pro-
moted such interdomain gene transfer events. Recently,
it has even been speculated that verrucomicrobial ecto-
symbionts with microtubulin structures similar to those
described by Petroni and colleagues [80] already colo-
nized the mid-ancestor of eukaryotic cells and, via an
endosymbiotic intermediate stage, evolved into the fla-
gellae of todays eukaryotes [101].
Conclusions
In conclusion, our understanding of the evolutionary
history and biology of the Planctomycetes, the Chlamydiae,
the Verrucomicrobia and their sister phyla has largely been
Current Opinion in Biotechnology 2006, 17:241249
246 Environmental biotechnology
propelled by genome and community genome analysis.
The currently ongoing genome sequencing projects of
Blastopirellula marina, Planctomyces maris, Gemmata obscur-
iglobus, Gemmata sp. strain Wa1-1 (Planctomycetes), Prosthe-
cobacter dejongeii, Akkermansia muciniphila, Chthoniobacter
flavus, Opitutus terrae, Verrucomicrobium sp. strain 2D6,
Verrucomicrobium spinosum (Verrucomicrobia), Simkania nege-
vensis Z (Chlamydiae), Lentisphaera araneosa, and Victivallis
vadensis (Lentisphaerae) (http://www.genomesonline.org)
can thus be expected to considerably extend and deepen
this knowledge and to shed more light onto one of the most
enigmatic branches of the bacterial tree of life.
Update
In April 2006 an interesting paper appeared that reports
on the diversity and distribution of PVC superphylum
members in the suboxic zone of the Black Sea [102]. This
study also includes a phylogenetic tree which clusters
Planctomycetes, Chlamydiae, Verrucomicrobia, Lentisphaerae
and candidate phylum OP3, but does not display the
Poribacteria.
Acknowledgements
The authors would like to express their gratitude to Mike Taylor for his
critical comments on the manuscript. We would also like to thank Mike
Jetten and Marc Strous (University of Nijmegen, Netherlands), Jean
Weissenbach and Denis LePaslier (Genoscope, France), as well as
Thomas Rattei and Werner Mewes (GSF, Germany) for great
collaboration on this subject. Work in the authors laboratory is supported
by the Austrian Science Fund (FWF) and the Federal Ministry for
Education, Science and Culture (bm:bwk).
References and recommended reading
Papers of particular interest, published within the annual period of
review, have been highlighted as:
 of special interest
 of outstanding interest
1. Schloss PD, Handelsman J: Status of the microbial census.
Microbiol Mol Biol Rev 2004, 68:686-691.
2. Ciccarelli FD, Doerks T, von Mering C, Creevey CJ, Snel B, Bork P:
Toward automatic reconstruction of a highly resolved tree of
life. Science 2006, 311:1283-1287.
3. Ward NL, Rainey FA, Hedlund BP, Staley JT, Ludwig W,
Stackebrandt E: Comparative phylogenetic analyses of
members of the order Planctomycetales and the division
Verrucomicrobia: 23S rRNA gene sequence analysis supports
the 16S rRNA gene sequence-derived phylogeny. Int J Syst Evol
Microbiol 2000, 50:1965-1972.
4. Jenkins C, Fuerst JA: Phylogenetic analysis of evolutionary
relationships of the planctomycete division of the domain
bacteria based on amino acid sequences of elongation factor
Tu. J Mol Evol 2001, 52:405-418.
5. Vergin KL, Urbach E, Stein JL, DeLong EF, Lanoil BD,
Giovannoni SJ: Screening of a fosmid library of marine
environmental genomic DNA fragments reveals four clones
related to members of the order Planctomycetales.
Appl Environ Microbiol 1998, 64:3075-3078.
6. Dedysh SN, Pankratov TA, Belova SE, Kulichevskaya IS,
Liesack W: Phylogenetic analysis and in situ identification
of bacteria community composition in an acidic Sphagnum
peat bog. Appl Environ Microbiol 2006, 72:2110-2117.
7. Kanokratana P, Chanapan S, Pootanakit K, Eurwilaichitr L:
Diversity and abundance of Bacteria and Archaea in the
Current Opinion in Biotechnology 2006, 17:241249
Bor Khlueng hot spring in Thailand. J Basic Microbiol 2004,
44:430-444.
8. Schlesner H, Rensmann C, Tindall BJ, Gade D, Rabus R, Pfeiffer S,
Hirsch P: Taxonomic heterogeneity within the
Planctomycetales as derived by DNA-DNA hybridization,
description of Rhodopirellula baltica gen. nov., sp. nov.,
transfer of Pirellula marina to the genus Blastopirellula gen.
nov. as Blastopirellula marina comb. nov. and emended
description of the genus Pirellula. Int J Syst Evol Microbiol 2004,
54:1567-1580.
9. Fuerst JA, Gwilliam HG, Lindsay M, Lichanska A, Belcher C,
Vickers JE, Hugenholtz P: Isolation and molecular identification
of planctomycete bacteria from postlarvae of the giant
tiger prawn, Penaeus monodon. Appl Environ Microbiol 1997,
63:254-262.
10. Shinzato N, Muramatsu M, Matsui T, Watanabe Y: Molecular
phylogenetic diversity of the bacterial community in the gut
of the termite Coptotermes formosanus. Biosci Biotechnol
Biochem 2005, 69:1145-1155.
11. Hentschel U, Hopke J, Horn M, Friedrich AB, Wagner M, Hacker J,
Moore BS: Molecular evidence for a uniform microbial
community in sponges from different oceans. Appl Environ
Microbiol 2002, 68:4431-4440.
12. Glockner FO, Kube M, Bauer M, Teeling H, Lombardot T,
 Ludwig W, Gade D, Beck A, Borzym K, Heitmann K et al.:
Complete genome sequence of the marine planctomycete
Pirellula sp. strain 1. Proc Natl Acad Sci USA 2003,
100:8298-8303.
This paper reports the first genome sequence of a planctomycete,
Pirellula sp. strain 1, which is today classified as Rhodopirellula baltica.
The R. baltica genome, among the largest bacterial genomes known to
date, encodes more than 100 sulfatases used for the degradation of
sulfated glycopolymers produced by algae.
13. Lombardot T, Bauer M, Teeling H, Amann R, Glockner FO:
The transcriptional regulator pool of the marine bacterium
Rhodopirellula baltica SH 1T as revealed by whole genome
comparisons. FEMS Microbiol Lett 2005, 242:137-145.
14. Wallner SR, Bauer M, Wurdemann C, Wecker P, Glockner FO,
Faber K: Highly enantioselective sec-alkyl sulfatase activity
of the marine planctomycete Rhodopirellula baltica shows
retention of configuration. Angew Chem Int Ed Engl 2005,
44:6381-6384.
15. Gadler P, Glueck SM, Kroutil W, Nestl BM, Larissegger-Schnell B,
Ueberbacher BT, Wallner SR, Faber K: Biocatalytic approaches
for the quantitative production of single stereoisomers from
racemates. Biochem Soc Trans 2006, 34:296-300.
16. Gade D, Theiss D, Lange D, Mirgorodskaya E, Lombardot T,
Glockner FO, Kube M, Reinhardt R, Amann R, Lehrach H et al.:
Towards the proteome of the marine bacterium
Rhodopirellula baltica: mapping the soluble proteins.
Proteomics 2005, 5:3654-3671.
17. Gade D, Gobom J, Rabus R: Proteomic analysis of
carbohydrate catabolism and regulation in the marine
bacterium Rhodopirellula baltica. Proteomics 2005,
5:3672-3683.
18. Gade D, Stuhrmann T, Reinhardt R, Rabus R: Growth phase
dependent regulation of protein composition in Rhodopirellula
baltica. Environ Microbiol 2005, 7:1074-1084.
19. Bauer M, Lombardot T, Teeling H, Ward NL, Amann RI,
Glockner FO: Archaea-like genes for C1-transfer enzymes
in Planctomycetes: phylogenetic implications of their
unexpected presence in this phylum. J Mol Evol 2004,
59:571-586.
20. Chistoserdova L, Jenkins C, Kalyuzhnaya MG, Marx CJ,
 Lapidus A, Vorholt JA, Staley JT, Lidstrom ME: The enigmatic
planctomycetes may hold a key to the origins of
methanogenesis and methylotrophy. Mol Biol Evol 2004,
21:1234-1241.
A provocative paper which nicely demonstrated by heterologous com-
plementation experiments that some of the C1-transfer genes which were
unexpectedly found to be present in Planctomycetes are indeed func-
tional.
www.sciencedirect.com

21. Kalyuzhnaya MG, Korotkova N, Crowther G, Marx CJ,
Lidstrom ME, Chistoserdova L: Analysis of gene islands involved
in methanopterin-linked C1 transfer reactions reveals new
functions and provides evolutionary insights. J Bacteriol 2005,
187:4607-4614.
22. Vorholt JA, Kalyuzhnaya MG, Hagemeier CH, Lidstrom ME,
Chistoserdova L: MtdC, a novel class of methylene
tetrahydromethanopterin dehydrogenases. J Bacteriol 2005,
187:6069-6074.
23. Schmid M, Walsh K, Webb R, Rijpstra WI, van de Pas-Schoonen K,
Verbruggen MJ, Hill T, Moffett B, Fuerst J, Schouten S et al.:
Candidatus Scalindua brodae, sp. nov., Candidatus
Scalindua wagneri, sp. nov., two new species of anaerobic
ammonium oxidizing bacteria. Syst Appl Microbiol 2003,
26:529-538.
24. Jetten MS, Cirpus I, Kartal B, van Niftrik L, van de Pas-
Schoonen KT, Sliekers O, Haaijer S, van der Star W, Schmid M,
van de Vossenberg J et al.: 1994-2004: 10 years of research on
the anaerobic oxidation of ammonium. Biochem Soc Trans
2005, 33:119-123.
25. Op den Camp HJ, Kartal B, Guven D, van Niftrik LA, Haaijer SC,
van der Star WR, van de Pas-Schoonen KT, Cabezas A, Ying Z,
Schmid MC et al.: Global impact and application of the
anaerobic ammonium-oxidizing (anammox) bacteria. Biochem
Soc Trans 2006, 34:174-178.
26. van Niftrik LA, Fuerst JA, Sinninghe Damste JS, Kuenen JG,
Jetten MS, Strous M: The anammoxosome: an intracytoplasmic
compartment in anammox bacteria. FEMS Microbiol Lett 2004,
233:7-13.
27. Sinninghe Damste JS, Strous M, Rijpstra WI, Hopmans EC,
 Geenevasen JA, van Duin AC, van Niftrik LA, Jetten MS:
Linearly concatenated cyclobutane lipids form a dense
bacterial membrane. Nature 2002, 419:708-712.
An excellent paper which shows that so-called ladderanes (linearly fused
cyclobutane rings) represent the dominant membrane lipids in the ana-
mmoxosome of anammox bacteria. This finding is particularly important
because anammox bacteria use these ladderanes to build a very dense
anammoxosome membrane and thus protect themselves from toxic
intermediates like hydrazine. Furthermore, it is noteworthy that ladder-
anes, which are promising compounds for optoelectronics, have never
been observed before in nature.
28. Boumann HA, Hopmans EC, van de Leemput I, Op den Camp HJ,
van de Vossenberg J, Strous M, Jetten MS, Sinninghe Damste JS,
Schouten S: Ladderane phospholipids in anammox bacteria
comprise phosphocholine and phosphoethanolamine
headgroups. FEMS Microbiol Lett 2006, 258:297-304.
29. Sinninghe Damste JS, Rijpstra WI, Geenevasen JA, Strous M,
Jetten MS: Structural identification of ladderane and other
membrane lipids of planctomycetes capable of anaerobic
ammonium oxidation (anammox). FEBS J 2005, 272:4270-4283.
30. Kartal B, Rattray J, van Niftrik LA, van de Vossenberg J,
Schmid MC, Webb RI, Schouten S, Fuerst JA, Damste JS,
Jetten MS et al.: Candidatus Anammoxoglobus propionicus a
new propionate oxidizing species of anaerobic ammonium
oxidizing bacteria. Syst Appl Microbiol 2006.
31. Schmid MC, Maas B, Dapena A, van de Pas-Schoonen K,
van de Vossenberg J, Kartal B, van Niftrik L, Schmidt I,
Cirpus I, Kuenen JG et al.: Biomarkers for in situ detection
of anaerobic ammonium-oxidizing (anammox) bacteria.
Appl Environ Microbiol 2005, 71:1677-1684.
32. Jetten M, Schmid M, van de Pas-Schoonen K, Sinninghe
Damste J, Strous M: Anammox organisms: enrichment,
cultivation, and environmental analysis. Methods Enzymol
2005, 397:34-57.
33. Kuypers MM, Lavik G, Woebken D, Schmid M, Fuchs BM,
Amann R, Jorgensen BB, Jetten MS: Massive nitrogen loss from
the Benguela upwelling system through anaerobic ammonium
oxidation. Proc Natl Acad Sci USA 2005, 102:6478-6483.
34. Guven D, van de Pas-Schoonen K, Schmid MC, Strous M,
Jetten MS, Sozen S, Orhon D, Schmidt I: Implementation of the
anammox process for improved nitrogen removal. J Environ
Sci Health A Tox Hazard Subst Environ Eng 2004, 39:1729-1738.
www.sciencedirect.com
The PVC superphylum Wagner and Horn 247
35. Third KA, Paxman J, Schmid M, Strous M, Jetten MS,
Cord-Ruwisch R: Treatment of nitrogen-rich wastewater
using partial nitrification and anammox in the CANON
process. Water Sci Technol 2005, 52:47-54.
36. Fux C, Siegrist H: Nitrogen removal from sludge digester liquids
by nitrification/denitrification or partial nitritation/anammox:
environmental and economical considerations. Water Sci
Technol 2004, 50:19-26.
37. Strous M, Pelletier E, Mangenot S, Rattei T, Lehner A, Taylor MW,
 Horn M, Daims H, Bartol-Mavel D, Wincker P et al.: Deciphering
the evolution and metabolism of an anammox bacterium from
a community genome. Nature 2006, 440:790-794.
This manuscript shows the power of environmental genomics by the
successful assembly of an annamox genome from a bioreactor commu-
nity consisting of at least 29 different bacteria. Although no template
genome of a closely related organism was available to assist the assem-
bly process, more than 98% of the genome could be reconstructed and
was used to develop new hypotheses on the metabolism and evolution of
this uncultured organism.
38. Guven D, Dapena A, Kartal B, Schmid MC, Maas B,
van de Pas-Schoonen K, Sozen S, Mendez R, Op den Camp HJ,
Jetten MS et al.: Propionate oxidation by and methanol
inhibition of anaerobic ammonium-oxidizing bacteria.
Appl Environ Microbiol 2005, 71:1066-1071.
39. Abdelrahman YM, Belland RJ: The chlamydial developmental
cycle. FEMS Microbiol Rev 2005, 29:949-959.
40. Subtil A, Dautry-Varsat A: Chlamydia: five years A.G. (after
genome). Curr Opin Microbiol 2004, 7:85-92.
41. Belland R, Ojcius DM, Byrne GI: Chlamydia. Nat Rev Microbiol
2004, 2:530-531.
42. Collingro A, Toenshoff ER, Taylor MW, Fritsche TR, Wagner M,
Horn M: Candidatus Protochlamydia amoebophila, an
endosymbiont of Acanthamoeba spp. Int J Syst Evol Microbiol
2005, 55:1863-1866.
43. Draghi A II, Popov VL, Kahl MM, Stanton JB, Brown CC,
 Tsongalis GJ, West AB, Frasca S Jr: Characterization of
Candidatus Piscichlamydia salmonis (Order Chlamydiales), a
chlamydia-like bacterium associated with epitheliocystis in
farmed Atlantic salmon (Salmo salar). J Clin Microbiol 2004,
42:5286-5297.
Candidatus Piscichlamydia salmonis represents a novel family within the
phylum Chlamydiae and is the deepest branching chlamydia-like organ-
ism identified so far. A more detailed analysis of these bacteria, which
cause epitheliocystis in salmon but which have so far not been obtained in
cell culture, should reveal novel insights into the evolution of chlamydiae
and might allow further conclusions on their last common ancestor.
44. Kostanjsek R, Strus J, Drobne D, Avgustin G: Candidatus
Rhabdochlamydia porcellionis, an intracellular bacterium
from the hepatopancreas of the terrestrial isopod Porcellio
scaber (Crustacea: Isopoda). Int J Syst Evol Microbiol 2004,
54:543-549.
45. Corsaro D, Valassina M, Venditti D: Increasing diversity within
Chlamydiae. Crit Rev Microbiol 2003, 29:37-78.
46. Vandahl BB, Birkelund S, Christiansen G: Genome and proteome
analysis of Chlamydia. Proteomics 2004, 4:2831-2842.
47. Horn M, Collingro A, Schmitz-Esser S, Beier CL, Purkhold U,
Fartmann B, Brandt P, Nyakatura GJ, Droege M, Frishman D et al.:
Illuminating the evolutionary history of chlamydiae.
Science 2004, 304:728-730.
48. Dong F, Zhong Y, Arulanandam B, Zhong G: Production
of a proteolytically active protein, chlamydial protease/
proteasome-like activity factor, by five different Chlamydia
species. Infect Immun 2005, 73:1868-1872.
49. Linka N, Hurka H, Lang BF, Burger G, Winkler HH, Stamme C,
Urbany C, Seil I, Kusch J, Neuhaus HE: Phylogenetic
relationships of non-mitochondrial nucleotide transport
proteins in bacteria and eukaryotes. Gene 2003, 306:27-35.
50. Greub G, Raoult D: History of the ADP/ATP-translocase-
encoding gene, a parasitism gene transferred from a
Chlamydiales ancestor to plants 1 billion years ago.
[corrected]. Appl Environ Microbiol 2003, 69:5530-5535.
Current Opinion in Biotechnology 2006, 17:241249
248 Environmental biotechnology
51. Schmitz-Esser S, Linka N, Collingro A, Beier CL, Neuhaus HE,
Wagner M, Horn M: ATP/ADP translocases: a common feature
of obligate intracellular amoebal symbionts related to
chlamydiae and rickettsiae. J Bacteriol 2004, 186:683-691.
52. Haferkamp I, Schmitz-Esser S, Linka N, Urbany C, Collingro A,
Wagner M, Horn M, Neuhaus HE: A candidate NAD+ transporter
in an intracellular bacterial symbiont related to chlamydiae.
Nature 2004, 432:622-625.
53. Haferkamp I, Schmitz-Esser S, Wagner M, Neigel N, Horn M,
Neuhaus HE: Tapping the nucleotide pool of the host: novel
nucleotide carrier proteins of Protochlamydia amoebophila.
Mol Microbiol 2006, 60:1534-1545.
54. Greub G, Mege J-L, Raoult D: Parachlamydia acanthamoeba
enters and multiplies within human macrophages and induces
their apoptosis. [corrected]. Infect Immun 2003, 71:5979-5985.
55. Collingro A, Poppert S, Heinz E, Schmitz-Esser S, Essig A,
Schweikert M, Wagner M, Horn M: Recovery of an
environmental chlamydia strain from activated sludge by
co-cultivation with Acanthamoeba sp. Microbiology
2005:301-309.
56. He J, Xu Z, Hughes J: Molecular bacterial diversity of a forest
soil under residue management regimes in subtropical
Australia. FEMS Microbiol Ecol 2006, 55:38-47.
57. Hackl E, Zechmeister-Boltenstern S, Bodrossy L, Sessitsch A:
Comparison of diversities and compositions of bacterial
populations inhabiting natural forest soils. Appl Environ
Microbiol 2004, 70:5057-5065.
58. Sangwan P, Chen X, Hugenholtz P, Janssen PH: Chthoniobacter
flavus gen. nov., sp. nov., the first pure-culture representative
of subdivision two, Spartobacteria classis nov., of the phylum
Verrucomicrobia. Appl Environ Microbiol 2004, 70:5875-5881.
59. Sangwan P, Kovac S, Davis KE, Sait M, Janssen PH: Detection
and cultivation of soil verrucomicrobia. Appl Environ Microbiol
2005, 71:8402-8410.
60. Joseph SJ, Hugenholtz P, Sangwan P, Osborne CA, Janssen PH:
Laboratory cultivation of widespread and previously
uncultured soil bacteria. Appl Environ Microbiol 2003,
69:7210-7215.
61. Schoenborn L, Yates PS, Grinton BE, Hugenholtz P, Janssen PH:
Liquid serial dilution is inferior to solid media for isolation of
cultures representative of the phylum-level diversity of soil
bacteria. Appl Environ Microbiol 2004, 70:4363-4366.
62. Juottonen H, Galand PE, Tuittila ES, Laine J, Fritze H, Yrjala K:
Methanogen communities and Bacteria along an
ecohydrological gradient in a northern raised bog complex.
Environ Microbiol 2005, 7:1547-1557.
63. Noll M, Matthies D, Frenzel P, Derakshani M, Liesack W:
Succession of bacterial community structure and diversity in a
paddy soil oxygen gradient. Environ Microbiol 2005, 7:382-395.
64. Martiny AC, Albrechtsen HJ, Arvin E, Molin S: Identification of
bacteria in biofilm and bulk water samples from a
nonchlorinated model drinking water distribution system:
detection of a large nitrite-oxidizing population associated
with Nitrospira spp. Appl Environ Microbiol 2005, 71:8611-8617.
65. Haukka K, Heikkinen E, Kairesalo T, Karjalainen H, Sivonen K:
Effect of humic material on the bacterioplankton community
composition in boreal lakes and mesocosms. Environ Microbiol
2005, 7:620-630.
66. Haukka K, Kolmonen E, Hyder R, Hietala J, Vakkilainen K,
Kairesalo T, Haario H, Sivonen K: Effect of nutrient loading
on bacterioplankton community composition in lake
mesocosms. Microb Ecol 2006, 51:137-146.
67. Lindstrom ES, Kamst-Van Agterveld MP, Zwart G: Distribution of
typical freshwater bacterial groups is associated with pH,
temperature, and lake water retention time. Appl Environ
Microbiol 2005, 71:8201-8206.
68. Eiler A, Bertilsson S: Composition of freshwater bacterial
communities associated with cyanobacterial blooms in four
Swedish lakes. Environ Microbiol 2004, 6:1228-1243.
Current Opinion in Biotechnology 2006, 17:241249
69. Polymenakou PN, Bertilsson S, Tselepides A, Stephanou EG:
Bacterial community composition in different sediments
from the Eastern Mediterranean Sea: a comparison of four
16S ribosomal DNA clone libraries. Microb Ecol 2005,
50:447-462.
70. Okabayashi A, Wakai S, Kanao T, Sugio T, Kamimura K: Diversity
of 16S ribosomal DNA-defined bacterial population in acid
rock drainage from Japanese pyrite mine. J Biosci Bioeng 2005,
100:644-652.
71. Huang LN, Zhu S, Zhou H, Qu LH: Molecular phylogenetic
diversity of bacteria associated with the leachate of a closed
municipal solid waste landfill. FEMS Microbiol Lett 2005,
242:297-303.
72. Kimura N, Shinozaki Y, Lee TH, Yonezawa Y: The microbial
community in a 2,4-dinitrophenol-digesting reactor as
revealed by 16S rDNA gene analysis. J Biosci Bioeng 2003,
96:70-75.
73. Chouari R, Le Paslier D, Dauga C, Daegelen P, Weissenbach J,
Sghir A: Novel major bacterial candidate division within a
municipal anaerobic sludge digester. Appl Environ Microbiol
2005, 71:2145-2153.
74. Pallen MJ, Beatson SA, Bailey CM: Bioinformatics, genomics
and evolution of non-flagellar type-III secretion systems:
a Darwinian perspective. FEMS Microbiol Rev 2005,
29:201-229.
75. Sakai T, Ishizuka K, Kato I: Isolation and characterization of a
fucoidan-degrading marine bacterium. Mar Biotechnol 2003,
5:409-416.
76. Stevenson BS, Eichorst SA, Wertz JT, Schmidt TM, Breznak JA:
New strategies for cultivation and detection of previously
uncultured microbes. Appl Environ Microbiol 2004,
70:4748-4755.
77. Wang M, Ahrne S, Jeppsson B, Molin G: Comparison of bacterial
diversity along the human intestinal tract by direct cloning and
sequencing of 16S rRNA genes. FEMS Microbiol Ecol 2005,
54:219-231.
78. Derrien M, Vaughan EE, Plugge CM, de Vos WM: Akkermansia
muciniphila gen. nov., sp. nov., a human intestinal
mucin-degrading bacterium. Int J Syst Evol Microbiol 2004,
54:1469-1476.
79. Vandekerckhove TT, Coomans A, Cornelis K, Baert P, Gillis M:
Use of the Verrucomicrobia-specific probe EUB338-III and
fluorescent in situ hybridization for detection of Candidatus
Xiphinematobacter cells in nematode hosts. Appl Environ
Microbiol 2002, 68:3121-3125.
80. Petroni G, Spring S, Schleifer KH, Verni F, Rosati G: Defensive
extrusive ectosymbionts of Euplotidium (Ciliophora) that
contain microtubule-like structures are bacteria related to
Verrucomicrobia. Proc Natl Acad Sci USA 2000, 97:1813-1817.
81. Bermudes D, Hinkle G, Margulis L: Do prokaryotes contain
microtubules? Microbiol Rev 1994, 58:387-400.
82. Jenkins C, Samudrala R, Anderson I, Hedlund BP, Petroni G,
Michailova N, Pinel N, Overbeek R, Rosati G, Staley JT:
Genes for the cytoskeletal protein tubulin in the bacterial
genus Prosthecobacter. Proc Natl Acad Sci USA 2002,
99:17049-17054.
83. Schlieper D, Oliva MA, Andreu JM, Lowe J: Structure of bacterial
tubulin BtubA/B: evidence for horizontal gene transfer.
Proc Natl Acad Sci USA 2005, 102:9170-9175.
84. Sontag CA, Staley JT, Erickson HP: In vitro assembly and
GTP hydrolysis by bacterial tubulins BtubA and BtubB.
J Cell Biol 2005, 169:233-238.
85. Fieseler L, Horn M, Wagner M, Hentschel U: Discovery of the
novel candidate phylum Poribacteria in marine sponges.
Appl Environ Microbiol 2004, 70:3724-3732.
86. Fieseler L, Quaiser A, Schleper C, Hentschel U: Analysis of the
first genome fragment from the marine sponge-associated,
novel candidate phylum Poribacteria by environmental
genomics. Environ Microbiol 2006, 8:612-624.
www.sciencedirect.com

87. Schmid M, Schmitz-Esser S, Jetten M, Wagner M: 16S-23S rDNA
intergenic spacer and 23S rDNA of anaerobic ammonium-
oxidizing bacteria: implications for phylogeny and in situ
detection. Environ Microbiol 2001, 3:450-459.
88. Liesack W, Stackebrandt E: Evidence for unlinked rrn operons
in the planctomycete Pirellula marina. J Bacteriol 1989,
171:5025-5030.
89. Cho JC, Vergin KL, Morris RM, Giovannoni SJ: Lentisphaera
 araneosa gen. nov., sp. nov, a transparent exopolymer
producing marine bacterium, and the description of a
novel bacterial phylum, Lentisphaerae. Environ Microbiol 2004,
6:611-621.
The continuation of an impressive success story which shows again that
many bacteria previously considered to be unculturable can be isolated
using very low nutrient concentrations.
90. Zoetendal EG, Plugge CM, Akkermans AD, de Vos WM: Victivallis
 vadensis gen. nov., sp. nov., a sugar-fermenting anaerobe
from human faeces. Int J Syst Evol Microbiol 2003, 53:211-215.
A very good paper that reports the first isolation of a bacterium belonging
to the Lentisphaera phylum by using a smart enrichment technique.
91. Hugenholtz P, Pitulle C, Hershberger KL, Pace NR: Novel division
level bacterial diversity in a Yellowstone hot spring. J Bacteriol
1998, 180:366-376.
92. Fuerst JA: Intracellular compartmentation in planctomycetes.
 Annu Rev Microbiol 2005, 59:299-328.
A detailed and well structured review on the structural cell plan of
Planctomycetes. A must for all people interested in the field.
93. Margolin W: FtsZ and the division of prokaryotic cells and
organelles. Nat Rev Mol Cell Biol 2005, 6:862-871.
94. McCoy AJ, Maurelli AT: Building the invisible wall: updating the
 chlamydial peptidoglycan anomaly. Trends Microbiol 2006,
14:70-77.
To date, peptidogycan has not been detected in chlamydiae, although all
chlamydiae encode an almost complete peptidoglycan biosynthesis
www.sciencedirect.com
The PVC superphylum Wagner and Horn 249
pathway. This is an excellent review summarizing our current knowledge
on potential roles of peptidoglycan in chlamydiae.
95. Staley JT, Bouzek H, Jenkins C: Eukaryotic signature proteins
of Prosthecobacter dejongeii and Gemmata sp. Wa-1 as
revealed by in silico analysis. FEMS Microbiol Lett 2005,
243:9-14.
96. Pearson A, Budin M, Brocks JJ: Phylogenetic and biochemical
evidence for sterol synthesis in the bacterium Gemmata
obscuriglobus. Proc Natl Acad Sci USA 2003, 100:15352-15357.
97. Teeling H, Lombardot T, Bauer M, Ludwig W, Glockner FO:
Evaluation of the phylogenetic position of the planctomycete
Rhodopirellula baltica SH 1 by means of concatenated
ribosomal protein sequences, DNA-directed RNA polymerase
subunit sequences and whole genome trees. Int J Syst Evol
Microbiol 2004, 54:791-801.
98. Bern M, Goldberg D: Automatic selection of representative
proteins for bacterial phylogeny. BMC Evol Biol 2005, 5:34.
99. Brochier C, Philippe H: Phylogeny: a non-hyperthermophilic
ancestor for bacteria. Nature 2002, 417:244.
100. Budd A, Blandin S, Levashina EA, Gibson TJ: Bacterial a2-
 macroglobulins: colonization factors acquired by horizontal
gene transfer from the metazoan genome? Genome Biol 2004,
5:R38.
A very interesting paper which suggests that several bacteria that live in
tight association with eukaryotes have acquired a2-macroglobulin from
their hosts and might use this to block host antimicrobial defence.
101. Li JY, Wu CF: New symbiotic hypothesis on the origin of
eukaryotic flagella. Naturwissenschaften 2005, 92:305-309.
102. Kirkpatrick J, Oakley B, Fuchsman C, Srinivasan S, Staley JT,
Murray JW: Diversity and distribution of Planctomycetes and
related bacteria in the suboxic zone of the Black sea. Appl
Environ Microbiol 2006, 72:3079-3083.
Current Opinion in Biotechnology 2006, 17:241249