By:John A. Fuerst, Ph.D. (School the Chemistry and also Molecular Biosciences, The college of Queensland)©2010ubraintv-jp.com Education
Citation:Fuerst ,J.A.(2010)Beyond Prokaryotes and Eukaryotes : Planctomycetes and Cell Organization.ubraintv-jp.com Education3(9):44
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Planctomycetes challenge our concept of the bacter cell and of a prokaryote together a cell framework type, and also our ideas around origins of the eukaryotic bio nucleus.

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To what level do all living cells have actually the very same structure? carry out shared functions only extend to a bounding membrane and DNA, RNA, and proteins as simple component components of your molecular biological engines? If so, why is this the case? come answer such questions we first need come know just how many varieties of cell structure there are. The planctomycete bacteria have challenged our assumptions about these basic questions.


What different kinds of cell do species in various kingdoms of life possess? every the same type or numerous different species or just a few? All modern living cell we recognize of are bounded by a membrane the surrounds the cell"s cytoplasm and a genome that DNA. This uses to members of every three great Domains that life, the Bacteria, the Archaea, and also the Eucarya (Woese, Kandler, & Wheelis 1990), an initial separated native each other on the basis of ribosome RNA order (Woese & Fox 1977). The cell of Bacteria and also Archaea, with a simple cell architecture, differ basically in cell structure from those the Eucarya, which have actually more complex structure, with inner compartments characterized by membranes. This two varieties have been termed prokaryotes (cells through naked DNA there is no a membrane envelope) and also eukaryotes (cells v a membrane-bounded nucleus). Standard scale (meaning ‘true nuclei" wherein "karyote" is derived from the ancient Greek for "kernel") encompass fungi, protozoa, eukaryotic algae, and of food plants and animals consisting of ourselves. Eukaryotic cells have actually a cell nucleus containing chromosomal DNA bounded by membranes. The cell types in Bacteria and Archaea are taken into consideration as "prokaryotes." The usage of the term prokaryote was very first introduced as a result of electron microscopic lense studies reflecting a shared basic cell structure amongst bacteria (Stanier & Van niel 1962). It has been topic to controversy because two distinct Domains the life, Bacteria and also Archaea, are grouped as prokaryotes (Pace 2006; Whitman 2009, martin & Koonin 2006; Cavalier-Smith 2007; Dolan & Margulis 2007; Glansdorff, Xu & Labedan 2008) . Interestingly, "prokaryote" means "before the nucleus," and refers to a cell framework where the chromosomal DNA is not enveloped in any internal membrane but is free in the cell cytoplasm.

It is a useful rule the thumb, for example for a scientist making use of a light microscope in a hospital microbiology lab, the eukaryotes such as the yeast Candida are bigger than prokaryotes through an bespeak of ten. However, cell size does not specify eukaryotes, because eukaryotes and prokaryotes overlap in size. The naval alga Ostreococcus manages the internal complexity of chloroplasts, mitochondria, and also nucleus in a cell around 1 μm long and 0.7 μm wide, similar to plenty of Bacteria and also Archaea (Courties, 1994). In contrast, one of the largest well-known bacteria — Epulopiscium fishelsoni uncovered in fish intestines — can be up to 600 μm long and 80 μm wide, simply visible come the nude eye (Angert, Clements & speed 1993). The greatest bacterium of every is a large sulfur-depositing bacter Thiomargarita namibiensis, frequently at the very least 100 μm broad and as much as 750 μm vast (Schulz 1999). Genome sizes of prokaryotes and eukaryotes likewise overlap: just 2.9 Mb (million basic pairs) lengthy in the eukaryotic bio parasite Encephalitozoon cuniculi but 9.1 Mb in gliding bacterium Myxococcus xanthus.


In prokaryote cell company there is a nucleoid containing genomic DNA but it is not surrounded by membranes such together what specifies the eukaryotic bio nucleus (Martin & Koonin 2006). Eukaryotes such together fungi, protozoa, plants, and also animals, and also thus we humans, have cells with complicated structure (Figure 1) with interior membranes and membrane-bounded organelles. These include the chromosome-containing nucleus surrounded by a atom envelope consisting of 2 membranes. Many eukaryotes additionally have more membrane-bounded organelles, mitochondria and chloroplasts. Some eukaryotes possess neither mitochondria or chloroplasts; these incorporate parasitic protozoa causing human disease such as Giardia, ancestors the which are thought to have actually lost your mitochondria however retained remnant mitochondrial genes or even in some cases remnants of mitochondria referred to as mitosomes or hydrogenosomes (Embley 2006). Mitochondria and also chloroplasts represent old symbiotic events in i m sorry bacteria were got by early on eukaryotes or your ancestors; molecular evidence for their relationships to bacteria is really strong.


A eukaryotic cell (left) has membrane-enclosed DNA, which creates a structure referred to as the cell nucleus (located at center of the eukaryotic cell; keep in mind the violet DNA fastened in the pink nucleus). A typical eukaryotic cell also has added membrane-bound oribelles of differing shapes and also sizes. In contrast, a prokaryotes cell (right) does not have actually membrane-bound DNA and also lacks various other membrane-bound organelles together well.

Would you suppose some features of eukaryotic bio molecular biology to be characteristic that this kind of cell? Yes, for example transcription and translation are not coupled in eukaryotes however are separated; protein-coding genes come as exons interrupted through introns which have to be spliced out; DNA is compacted via proteins referred to as histones; and also there is a facility endomembrane system. That system contains membrane vesicle-dependent motion of products into and out that the cell-endocytosis and exocytosis. Cell department in eukaryotes is a visibly complicated multi-stage procedure called mitosis entailing microtubules written of cytoskeletal proteins.


Pirellula marina", "true", "Figure 2", "A cryosubstituted whole-cell thin section of Pirellula marina strain ACM 3344 (= ATCC 49060, DSM 3645) is shown false-colored to improve contrast. Paryphoplasm (from the Greek paryphe, a border woven along a robe) is the peripheral cytoplasm surrounding the membrane-bounded pirellulosome, and lacks ribosomes. Riboplasm, within the pirellulosome has ribosomes.", "true", "All legal rights reserved.", "600", "403", "http://www.sgm.ac.uk/");">
Pirellula marina", "true", "Figure 2", "A cryosubstituted whole-cell thin ar of Pirellula marina stress, overload ACM 3344 (= ATCC 49060, DSM 3645) is displayed false-colored to enhance contrast. Paryphoplasm (from the Greek paryphe, a border woven along a robe) is the peripheral cytoplasm bordering the membrane-bounded pirellulosome, and also lacks ribosomes. Riboplasm, within the pirellulosome consists of ribosomes.", "true", "All civil liberties reserved.", "600", "403", "http://www.sgm.ac.uk/");">Figure 2:Pirellula marina
Pirellula marina", "true", "Figure 2", "A cryosubstituted whole-cell thin section of Pirellula marina strain ACM 3344 (= ATCC 49060, DSM 3645) is displayed false-colored to boost contrast. Paryphoplasm (from the Greek paryphe, a border woven follow me a robe) is the peripheral cytoplasm surrounding the membrane-bounded pirellulosome, and also lacks ribosomes. Riboplasm, inside the pirellulosome includes ribosomes.", "true", "All rights reserved.", "600", "403", "http://www.sgm.ac.uk/");">A cryosubstituted whole-cell thin section of Pirellula marina stress, overload ACM 3344 (= ATCC 49060, DSM 3645) is presented false-colored to enhance contrast. Paryphoplasm (from the Greek paryphe, a border woven along a robe) is the peripheral cytoplasm surrounding the membrane-bounded pirellulosome, and also lacks ribosomes. Riboplasm, inside the pirellulosome contains ribosomes.
© 1997 society for basic Microbiology Lindsay, M. R., Webb, R. I. & Fuerst, J. A. Pirellulosomes: A new form of membrane-bounded cell compartment in planctomycete bacteria of the genus Pirellula. Microbiology-uk 143 739–748 (1997). All legal rights reserved.
Pirellula marina", "Figure 2", "A cryosubstituted whole-cell thin section of Pirellula marina strain ACM 3344 (= ATCC 49060, DSM 3645) is presented false-colored to boost contrast. Paryphoplasm (from the Greek paryphe, a border woven along a robe) is the peripheral cytoplasm bordering the membrane-bounded pirellulosome, and lacks ribosomes. Riboplasm, within the pirellulosome has ribosomes.", "600","http://www.sgm.ac.uk/", "The cell has an oval shape outlined by 2 lines representing barriers: the outermost great is labeling the \"cell wall\" and also the inner obstacle is labeled the \"cell membrane.\" Two-thirds that the cabinet is easy shaded. This lightly-shaded region is an organelle labeled the \"pirellulosome\" and its shape is reinforced by a barrier labeled the \"intracytoplasmic membrane.\" at the center of the pirellulosome, a dense, darkly-shaded circular an ar is labeled the \"nucleoid.\" The lightly-shaded an are surrounding the newly closed is labeling the \"riboplasm,\" and is peppered by small black dots, which are ribosomes. The staying volume that the cell the is not attributed come the pirellulosome includes no ribosomes and also is labeling the \"paryphoplasm.\"")" class="inlineLinks"> Figure detail
What are planctomycetes and also why room they so different? Members the the phylum Planctomycetes are certainly Bacteria ~ above the communication of gene sequence trees such as those native ribosomal RNA, however they have actually several considerably distinctive functions separating them from other bacteria (Fuerst 1995). They happen in many marine and freshwater aquatic habitats and in soils, and also in too much habitats such together the ultradry Mars-like soil of the Atacama desert (Schlesner, 1994) and living stromatolite microbial mats in hypersaline marine shallows (Papineau et al 2005). They are so various from various other bacteria that, when very first detected through microscopy that lake water, castle were puzzled with fungi due to the fact that some have stalks conveniently mistaken for cellular filaments choose those of fungi. Microbiologists have discovered planctomycetes in the setting by culturing making use of media include antibiotics, since planctomycetes are naturally resistant come penicillins, or via clone libraries created from DNA adhered to by PCR and sequencing of 16S rRNA genes. One group, anammox planctomycetes, has a unique metabolism and also generate energy via oxidation that ammonia without oxygen (the "anammox" process-that is, anaerobic ammonia oxidation). This procedure is crucial for worldwide nitrogen cycling, is supplied for ecological cleanup of nitrogen in wastewater (Kuenen 2008; Strous, M. Et al) , and also may in future significantly reduce the energy expenses of sewer cleanup (Kartal, Kuenen & valve Loosdrecht 2010). A scientist native a biotechnology agency discovered the procedure when trying to account for unusual disappearances that ammonia in a wastewater remediation plant. Various other microbiologists sequenced gene from copy DNA from bioreactors and also examined bioreactor cells by electron microscopy, showing that planctomycetes were responsible for anammox. All planctomycetes own the remarkable attribute of lacking peptidoglycan, a cell wall polymer existing in many bacteria yet not archaea, and also they every reproduce through budding rather than by binary fission properties of E. Coli. Their absence of peptidoglycan provides them resistant come β-lactam antibiotics consisting of penicillins, the basis of your selective culture from blended communities. Among their most remarkable attributes is the their cells are compartmentalized — split into separate internal compartments — by membranes. To uncover these membranes, scientists supplied special "cryo" methods for prepare sectioned cells because that examination in electron microscopes, since rapid freezing better preserves these fragile internal structures. All planctomycete types examined have cells whereby a major intracytoplasmic membrane the end an external ribosome-free an ar termed the paryphoplasm native a significant inner compartment containing every the cell DNA condensed right into a fibrillar nucleoid. This major inner compartment has actually been termed the pirellulosome, after planctomycete genus Pirellula (Lindsay, Webb & Fuerst 1997) (Figure 2). Cell of every planctomycete species possess a pirellulosome, a paryphoplasm, and a membrane separating them, for this reason in all species nucleoid DNA is surrounding by at least one internal membrane. In the situation of one species, Gemmata obscuriglobus, one even more startling structure occurs, where the nucleoid within the pirellulosome is surrounded by one envelope consists of two closely apposed membranes (Figure 3). This outcomes in a nucleus-like structure, the atom body, at least superficially similar to the of the eukaryotic nucleus, which additionally has a nuclear envelope consist of of two membranes. The envelope appears to entirely surround the nucleoid-containing nuclear body. Microbiologists identified this from examining serial part of cells by electron microscopic lense — pictures of a collection of sections from the same cell can be provided to derive computer reconstructions in one analogous means to just how CAT scans that the human body are offered in medicine (Lindsay 2001).


A cryosubstituted whole-cell thin ar of Gemmata obscuriglobus handle via high-pressure freezing showing a nuclear human body consisting of nuclear envelope membranes around a an ar containing condensed nucleoid and ribosomes, several of which have the right to be checked out lining the inside membrane the the nuclear body envelope. Ribosomes are additionally present in the cytoplasm outside of the atom body, however a ribosome-free paryphoplasm is current at the outer component of the cell. Tiny red arrows indicate regions of the inside nuclear body envelope whereby electron-dense ribosome-like particles room lined up along this membrane. Bar 500 nm.
© 2010 ubraintv-jp.com education and learning Courtesy the Kuo-Chang Lee, activities of J. A. Fuerst. All civil liberties reserved.
Gemmata obscuriglobus", "Figure 3", "A cryosubstituted whole-cell thin ar of Gemmata obscuriglobus handle via high-pressure freezing reflecting a nuclear body consisting of nuclear envelope membranes around a region containing condensed nucleoid and also ribosomes, few of which deserve to be checked out lining the within membrane that the nuclear body envelope. Ribosome are also present in the cytoplasm exterior of the atom body, but a ribosome-free paryphoplasm is existing at the outer part of the cell. Little red arrows suggest regions that the inside nuclear body envelope whereby electron-dense ribosome-like particles room lined up along this membrane. Bar 500 nm. ", "650","http://www.ubraintv-jp.com/ubraintv-jp.com_education", "There is a space between the outermost border of the cell and also an interior barrier. The interior obstacle is labeling the \"intracytoplasmic membrane\" and the room is labeling \"paryphoplasm.\" The inside of the cell contains multiple membrane layers, labeling \"nuclear envelope membranes.\" 4 red arrows point to small, black dots lining the inside of the atom membranes. In the space enclosed through the nuclear membrane, a circular, darkly-shaded main point is labeling \"condensed new closed DNA.\" The membrane-enclosed an ar inside the cell and the interior room surrounding it room both labeled \"riboplasm.\"")" class="inlineLinks"> Figure information
Candidatus Kuenenia stuttgartiensis", "true", "Figure 4", "Transmission electron micrograph the a Candidatus Kuenenia stuttgartiensis cell. Together a member of the Planctomycetes, Candidatus K. Stuttgartiensis consists of subcellular compartments, consisting of the anammoxosome, where power conservation take away place. The sample was highpressure frozen, freeze substituted and Epon embedded. The riboplasm is the equivalent of the ribosome-containing cytoplasm in many other bacteria. The range bar represents 200 nm.", "true", "All legal rights reserved.", "605", "503", "");">
Candidatus Kuenenia stuttgartiensis", "true", "Figure 4", "Transmission electron micrograph that a Candidatus Kuenenia stuttgartiensis cell. As a member the the Planctomycetes, Candidatus K. Stuttgartiensis contains subcellular compartments, consisting of the anammoxosome, where power conservation take away place. The sample was highpressure frozen, freeze substituted and also Epon embedded. The riboplasm is the indistinguishable of the ribosome-containing cytoplasm in many other bacteria. The scale bar to represent 200 nm.", "true", "All legal rights reserved.", "605", "503", "");">Figure 4
Candidatus Kuenenia stuttgartiensis", "Figure 4", "Transmission electron micrograph of a Candidatus Kuenenia stuttgartiensis cell. As a member of the Planctomycetes, Candidatus K. Stuttgartiensis has subcellular compartments, including the anammoxosome, where energy conservation take away place. The sample was highpressure frozen, freeze substituted and Epon embedded. The riboplasm is the indistinguishable of the ribosome-containing cytoplasm in many other bacteria. The scale bar to represent 200 nm.", "605","", "The plantomycete has a one shape, and also is outlined by 2 barriers: the outermost obstacle is labeling the \"cell wall\" and the inner obstacle is labeling the \"cytoplasmic membrane.\" A range bar represents 200 nanometers, and the plantomycete is around three time the width of the scale bar, or 600 nanometers. The room inside the cell"s inner is labeled \"riboplasm\" which is separated native the cell"s numerous outer membrane great by a thin line labeled \"intracytoplasmic membrane.\" The room between the riboplasm and also the cytoplasmic membrane is labeling \"paryphoplasm.\" A dark region of the cell, barely distinguishable indigenous the bordering riboplasm, is labeling \"nucleoid.\" beside the nucleoid, approximately two-thirds the the cell"s interior volume is labeling \"anammoxosome,\" and also is easy shaded. The anammoxosome is outlined by one \"anammoxosome membrane.\"")" class="inlineLinks"> Figure information
Membranes the the nuclear human body envelope space formed throughout budding (Lee, Webb & Fuerst 2009). In the anammox bacteria discussed above, there is also an additional but this time single-membrane-bounded organelle, the anammoxosome (Figure 4), comprise at least one enzyme essential for their unique anammox ammonia-oxidizing physiology in the lack of oxygen (van Niftrik et al 2004). An additional interesting function of planctomycete cells is your condensed nucleoid, a fixed of urgently chromosomal DNA fibrils clearly shows in sectioned cell via electron microscopy, contrasting v the nucleoids of other bacteria which often tend to be spread throughout the cytoplasm. In the case of Gemmata obscuriglobus this has actually been attached to the high resistance of cell in this types to UV and also gamma radiation (Lieber et al 2009). The planctomycetes are hence Bacteria that kind a significant exception to simple prokaryote structure. Space there possibly other exceptions to be discovered? Perhaps; Ignicoccus hospitalis, an Archaean life at 90°C may pose similar problems because that the prokaryote idea (Kuper, U. Et al 2010).


Why are planctomycetes important for understanding development of eukaryotes? There are two significant types that hypothesis about how eukaryotes and their nuclei may have actually originated (Poole & penny 2007). Among these involves fusion between cells from domain names Archaea and Bacteria: one of the cells involved in the fusion becomes the eukaryote nucleus (Martin & Muller 1998; boy name 2005; boy name et al. 2001). There room several different versions the this hypothesis (Martin & Koonin 2006). The other kind of hypothesis involves "autogenous" development of internal membranes from just one non-eukaryote cell form via an infolding the cytoplasmic membrane to form an endomembrane system and eventually to type a cell core (Cavalier-Smith 2002; Cavalier-Smith 2009). Nutrient salvation may have selected because that this (de Duve 2007).

The existence of planctomycetes v their membrane compartments support an autogenous (self generating) design for development of membrane-bounded nuclei (Glansdorff, Xu & Labedan 2008; Fuerst 2005). Us don"t yet understand to what extent compartments of planctomycetes deserve to be explained by molecular mechanisms comparable to those the eukaryotes. Also if it turns out that there space no straight links between planctomycetes and also eukaryotes (i.e. If convergent evolution can describe their similarities), the planctomycete cell arrangement implies that combination models for beginnings of eukaryote-like cabinet compartments might not be necessary to describe how eukaryotes obtained their nucleus.


Planctomycete cell structure differs from the of all other known prokaryotes. We space not certain yet whether the cell plan of a planctomycete favor Gemmata resembles that of standard scale at a deeper molecular level. Together a eukaryote-like cabinet plan developing in a member that the Bacteria says that it may be worth trying to find such depth similarities.

An amazing recent breakthrough suggesting the planctomycetes and also their cells might possess deeper resemblances with eukaryotes is the exploration of an endocytosis-like procedure for protein absorb by cell of Gemmata obscuriglobus entailing membrane infolding and vesicle formation (Lonhienne et al 2010). Like the membrane-bounded nuclear human body in this species, together a procedure is unknown in various other Bacteria or Archaea. This is constant with the nutrient-acquisition-driven model for endomembrane advancement referred to above as a possible an initial step top top the roadway to eukaryotes. Can planctomycetes have actually a whole collection of further attributes previously well-known only in eukaryotes, consisting of eukaryote-homologous genes and proteins? cabinet biologists and also microbiologists don"t understand yet, but are excited by this prospect which may lead to new insights right into the evolutionary relationships between simpler and more complicated kinds the cells.


Cell biologists and microbiologists have in the previous thought that just two major types of structure cell organization exist, prokaryotes — wherein the chromosomal DNA is naked and also exposed in ~ the cabinet cytoplasm — and eukaryotic — where chromosomal DNA is surrounding by a membranous envelope. Every members of domain names Bacteria and also Archaea have actually been believed to possess prokaryotes organization. The planctomycete bacteria however complicate this simple scheme, due to the fact that their cells space compartmentalized by inner membranes including those neighboring the chromosomal DNA. Space they a third form of cell company that originated separately of other known types, or space they key to understanding how eukaryotes and their facility internal framework evolved? In either situation they are significant for expertise the development of cells and the varied ways in which cells have the right to potentially evolve.


Angert, E. R., Clements, K. D. & Pace, N. R. The largest bacterium. Ubraintv-jp.com 362, 239–241 (1993) doi:10.1038/362239a0.

Cavalier-Smith, T. Ide of a bacterium still precious in prokaryote debate. Ubraintv-jp.com 446, 257 (2007) doi:10.1038/446257c.

Cavalier-Smith, T. The phagotrophic origin of eukaryotes and phylogenetic classification of Protozoa. Int J Syst Evol Microbiol 52, 297–354 (2002).

Cavalier-Smith, T. Predation and eukaryote cell origins: a coevolutionary perspective. Int J Biochem cabinet Biol 41, 307–322 (2009) doi:10.1016/j.biocel.2008.10.002.

Courties, C. Et al. Smallest eukaryotic organism. Ubraintv-jp.com 370, 255 (1994).

de Duve, C. The beginning of eukaryotes: a reappraisal. Nat Rev Microbiol 8, 395–403 (2007) doi:10.1038/nrg2071.

Dolan, M. F. & Margulis, L. Advances in biology expose truth around prokaryotes. Ubraintv-jp.com 445, 21 (2007) doi:10.1038/445021b.

Embley, T. M. Multiple an additional origins that the anaerobic way of living in eukaryotes. Philos infectious diseases world fashion R Soc lonely B Biol Sci 361, 1055–1067 (2006) doi:10.1098/rstb.2006.1844.

Fuerst, J. A. & Webb, R. I. Membrane-bounded nucleoid in the eubacterium Gemmata obscuriglobus. Proc Natl Acad Sci U S A 88, 8184–8188 (1991).

Fuerst, J. A. The planctomycetes: emerging models because that microbial ecology, evolution and cell biology. Microbiology 141 ( Pt 7), 1493–1506 (1995).

Kartal, B., Kuenen, J. G. & van Loosdrecht, M. C. Engineering. Sewage treatment v anammox. Scientific research 328, 702–703 (2010) doi:10.1126/science.1185941.

Kuenen, J. G. Anammox bacteria: from discovery to application. Nat Rev Microbiol 6, 320–326 (2008) doi:10.1038/nrmicro1857.

Lindsay, M. R. Et al. Cell compartmentalisation in planctomycetes: novel varieties of structure organisation because that the bacterial cell. Arch Microbiol 175, 413–429 (2001).

Lonhienne, T. G. Et al. Endocytosis-like protein absorb in the bacterium Gemmata obscuriglobus. Proc Natl Acad Sci USA (2010) doi:10.1073/pnas.1001085107.

Lee, K. C., Webb, R. I. & Fuerst, J. A. The cell cycle that the planctomycete Gemmata obscuriglobus v respect to cell compartmentalization. BMC cabinet Biol 10, 4 (2009) doi:10.1186/1471-2121-10-4.

Lieber, A., Leis, A., Kushmaro, A., Minsky, A. & Medalia, O. Chromatin organization and also radio resistance in the bacterium Gemmata obscuriglobus. J Bacteriol, 191 1439–1445 (2009) doi:10.1128/jb.01513-08.

Lindsay, M. R., Webb, R. I. & Fuerst, J. A. Pirellulosomes: A new form of membrane-bounded cell compartment in planctomycete bacteria of the genus Pirellula. Microbiol-Uk 143, 739–748 (1997).

Martin, W. Archaebacteria (Archaea) and the beginning of the eukaryotic bio nucleus. Curr Opin Microbiol 8, 630–637 (2005) doi:10.1016/j.mib.2005.10.004.

Martin, W. Et al. An introduction of endosymbiotic models for the origins of eukaryotes, their ATP-producing organelles (mitochondria and hydrogenosomes), and also their heterotrophic lifestyle. Biol Chem 382, 1521–1539 (2001) doi:10.1515/bc.2001.187.

Martin, W. & Koonin, E. V. A positive an interpretation of prokaryotes. Ubraintv-jp.com 442, 868 (2006) doi:10.1038/442868c.

Martin, W. & Koonin, E. V. Introns and also the origin of nucleus-cytosol compartmentalization. Ubraintv-jp.com 440, 41–45 (2006) doi:10.1038/ubraintv-jp.com04531.

Martin, W. & Muller, M. The hydrogen theory for the first eukaryote. Ubraintv-jp.com 392, 37–41 (1998) doi:10.1038/32096.

Pace, N. R. Time for a change. Ubraintv-jp.com 441, 289 (2006) doi:10.1038/441289a.

Pace, N. R. Troubles with "procaryote". J Bacteriol 191, 2008–2010; discussion 2011 (2009) doi:10.1128/jb.01224-08

Papineau, D. Et. Al Composition and also structure the microbial neighborhoods from stromatolites the Hamelin pool in Shark Bay, western Australia. Appl Environ Microbiol 71, 4822–4832 (2005) doi:10.1128/aem.71.8.4822-4832.2005

Poole, A. M. & Penny, D. Analyzing hypotheses for the origin of eukaryotes. Bioessays 29, 74–84 (2007) doi:10.1002/bies.20516.

Schlesner, H. The advance of media an ideal for the microorganisms morphologically resembling Planctomyces spp., Pirellula spp., and other Planctomycetales from miscellaneous aquatic habitats making use of dilute media. Systematic and Applied Microbiology 17, 135–145 (1994).

Schulz, H. N. Et al. Dense populations of a gigantic sulfur bacterium in Namibian shelf sediments. Scientific research 284, 493–495 (1999).

Stanier, R. Y. & valve Niel, C. B. The concept of a bacterium. Arch Mikrobiol 42, 17–35 (1962).

Strous, M. Et al. Lacking lithotroph established as brand-new planctomycete. Ubraintv-jp.com 400, 446–449 (1999) doi:10.1038/22749.

van Niftrik, L. A. Et al. The anammoxosome: an intracytoplasmic compartment in anammox bacteria. FEMS Microbiol Lett 233, 7–13 (2004).

Whitman, W. B. The modern-day concept the the procaryote. J Bacteriol 191, 2000–2005; conversation 2006–2007 (2009) doi:10.1128/jb.00962-08.

Woese, C. R., Kandler, O. & Wheelis, M. L. In the direction of a natural system of organisms: proposal because that the domains Archaea, Bacteria, and also Eucarya. Proc Natl Acad Sci U S A 87, 4576–4579 (1990).

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Woese, C. R. & Fox, G. E. Phylogenetic framework of the prokaryotic domain: the main kingdoms. Proc Natl Acad Sci U S A 74, 5088–5090 (1977).