Schizosaccharomyces pombe

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Schizosaccharomyces pombe
Scientific classification
Kingdom: Fungi
Phylum: Ascomycota
Subphylum: Taphrinomycotina
Class: Schizosaccharomycetes
Order: Schizosaccharomycetales
Family: Schizosaccharomycetaceae
Genus: Schizosaccharomyces
Species: S. pombe
Binomial name
Schizosaccharomyces pombe
Lindner (1893)

Schizosaccharomyces pombe, also called "fission yeast", is a species of yeast. It is used as a model organism in molecular and cell biology. It is a unicellular eukaryote, whose cells are rod-shaped. Cells typically measure 3 to 4 micrometres in diameter and 7 to 14 micrometres in length. Its genome, which is approximately 14.1 million base pairs, is estimated to contain 4,970 protein-coding genes and at least 450 non-coding RNAs.[1]

These cells maintain their shape by growing exclusively through the cell tips and divide by medial fission to produce two daughter cells of equal sizes, which makes them a powerful tool in cell cycle research.

Fission yeast was isolated in 1893 by Paul Lindner from East African millet beer. The species name is derived from the Swahili word for beer (Pombe). It was first developed as an experimental model in the 1950s: by Urs Leupold for studying genetics,[2][3] and by Murdoch Mitchison for studying the cell cycle.[4][5]

Paul Nurse, a fission yeast researcher, successfully merged the independent schools of fission yeast genetics and cell cycle research. Together with Lee Hartwell and Tim Hunt, Nurse won the 2001 Nobel Prize in Physiology or Medicine for their work on cell cycle regulation.

The sequence of the S. pombe genome was published in 2002, by a consortium led by the Sanger Institute, becoming the sixth model eukaryotic organism whose genome has been fully sequenced. This has fully unlocked the power of this organism, with many genes homologous to human disease genes being identified. In 2006, sub-cellular localization of all the proteins in S. pombe was published using green fluorescent protein as a molecular tag.

S. pombe has also become an important organism in studying the cellular responses to DNA damage and the process of DNA replication.

Approximately 120 natural strains of S. pombe have been isolated. These have been collected from a variety of locations including Europe, North and South America and Asia. The majority of these strains have been collected from cultivated fruits such as apples and grapes, or from the various alcoholic beverages, such as Brazilian Cachaça. It is not clear at present whether S. pombe is the major fermenter or a contaminant in such brews. The natural ecology of Schizosaccharomyces yeasts is not well studied.

Contents

History [edit]

S. pombe was first discovered in 1893 when a group working in a Brewery Association Laboratory in Germany was looking at sediment found in millet beer imported from East Africa that gave it an unsavory acidic taste. The term Schizo, meaning "different", had previously been used to describe other fission species such as the fission fungi. The addition of the word pombe was due to its isolation from beer, as pombe essentially means "from beer", in Swahili. The standard S. pombe strains were isolated by Urs Leupold in 1946 and 1947 from a culture that he obtained from the yeast collection in Delft, The Netherlands. It was deposited there by A. Osterwalder under the name S. pombe var liquefaciens, after he isolated it in 1924 from French wine (most probably rancid) at the Federal Experimental Station of Vini- and Horticulture in WŠdenswil, Switzerland. The culture used by Urs Leupold contained (besides others) cells with the mating types h90 (strain 968), h- (strain 972), and h+ (strain 975). Subsequently there have been two large efforts to isolate S. pombe from fruit, nectar, or fermentations: one by Florenzano et al.[6] in the vineyards of Western Sicily, and the other by Gomes et al. (2002) in four regions of Southeast Brazil.[7]

Ecology [edit]

The fission yeast S. pombe belong to the phylum Ascomycota which represents the largest and most diverse group of fungi. Free-living ascomycetes are commonly found in tree exudates, plant roots and surrounding soil, on ripe and rotting fruits, and in association with insect vectors that transport them between substrates. Many of these associations are symbiotic or saprophytic, although numerous ascomycetes (and their basidiomycete cousins) represent important plant pathogens that target a myriad of plant species, including commercial crops. Among the ascomycetous yeast genera, the fission yeast Schizosaccharomyces is unique because of the deposition of a-(1,3)- glucan or pseudonigeran in the cell wall in addition to the better known b-glucans and the virtual lack of chitin. Species of this genus also differ in mannan composition,which shows terminal d-galactose sugars in the side-chains of their mannans. S. pombe undergo aerobic fermentation in the presence of excess sugar.[8] S. pombe can degrade L malic acid, one of the dominant organic acid in wine which makes then diverse among other Saccharomyces strain.

Comparison with budding yeast (Saccharomyces cerevisiae) [edit]

The yeast species S. pombe and S. cerevisiae are both extensively studied; these two species diverged approximately 300 to 600 million years before present,[citation needed] and are significant tools in molecular and cellular biology. Some of the technical discriminants between these two species are:

  • S. cerevisiae has approximately 5,600 open reading frames; S. pombe has approximately 4,970 open reading frames.
  • Despite similar gene numbers, S. cerevisiae has only about 250 introns, while S. pombe has nearly 5,000.
  • S. cerevisiae has 16 chromosomes, S. pombe has 3.
  • S. cerevisiae is often diploid while S. pombe is usually haploid.
  • S. cerevisiae is in the G1 phase of the cell cycle for an extended period (consequently, G1-S transition is tightly controlled) while S. pombe remains in the G2 phase of the cell cycle for an extended period (consequently, G2-M transition is under tight control).
  • Both species share genes with higher eukaryotes that they do not share with each other. S. pombe has RNAi machinery genes like those in vertebrates, while this is missing from S. cerevisiae. S. cerevisiae also has greatly simplified heterochromatin compared to S. pombe.[9] Conversely, S. cerevisiae has well-developed peroxisomes, while S. pombe does not.
  • S. cerevisiae has small point centromere of 125 bp, and sequence-defined replication origins of about the same size. Conversely, S. pombe has large, repetitive centromeres (40–100 kb) more similar to mammalian centromeres, and degenerate replication origins of at least 1kb.

Life cycle[10] [edit]

Centrosome of S. pombe.

The fission yeast is a single-celled fungus with simple, fully characterized genome and a rapid growth rate. It has long since been used in brewing, baking and molecular genetics. S. pombe is a rod-shaped cell, approximately 3 µm in diameter, that grows entirely by elongation at the ends. After mitosis, division occurs by the formation of a septum, or cell plate, that cleaves the cell at its midpoint.

The central events of cell reproduction are chromosome duplication, which takes place in S (Synthetic) phase, followed by chromosome segregation and nuclear division (mitosis) and cell division (cytokinesis), which are collectively called M (Mitotic) phase.G1 is the gap between M and S phases, and G2 is the gap between S and M phases. In the budding yeast, the G1 phase is particularly extended, and cytokinesis (daughter-cell segregation) does not happen until a new S (Synthetic) phase is launched.

Fission yeast governs mitosis by mechanisms that are similar to those in multicellular animals. It normally proliferates in a haploid state. When starved, cells of opposite mating types (P and M) fuse to form a diploid zygote that immediately enters meiosis to generate four haploid spores. When conditions improve, these spores germinate to produce proliferating haploid cells.

  • General features of the cell cycle.

  • The particular cell cycle of a fission yeast.

  • A more detailed version of the cell cycle of a fission yeast.

Cytokinesis in fission yeast [edit]

Cytokinesis of the fission yeast.

The general features of cytokinesis are shown here. The site of cell division is determined before anaphase. The anaphase spindle (in green on the figure) is then positioned so that the segregated chromosomes are on opposite sides of the predetermined cleavage plane.

Size control in fission yeast [edit]

Cell-cycle length of the fission yeast depends on nutrient conditions.

In fission yeast, where growth governs progression through G2/M, a wee1 mutation causes entry into mitosis at an abnormally small size, resulting in a shorter G2. G1 is lengthened, suggesting that progression through Start (beginning of cell cycle) is responsive to growth when the G2/M control is lost. Furthermore, cells in poor nutrient conditions grow slowly and therefore take longer to double in size and divide. Low nutrient levels also reset the growth threshold so that cell progresses through the cell cycle at a smaller size. Finally, wee1 mutant fission yeast cells are smaller than wild-type cells, but take just as long to go through the cell cycle. This is possible because small yeast cells grow slower, that is, their added total mass per unit time is smaller than that of normal cells.

A spatial gradient is thought to coordinate cell size and mitotic entry in fission yeast.[11][12][13] The Pom1 protein kinase (green) is localized to the cell cortex, with the highest concentration at the cell tips. The cell-cycle regulators Cdr2, Cdr1 and Wee1 are present in cortical nodes in the middle of the cell (blue and red dots). a, In small cells, the Pom1 gradient reaches most of the cortical nodes (blue dots). Pom1 inhibits Cdr2, preventing Cdr2 and Cdr1 from inhibiting Wee1, and allowing Wee1 to phosphorylate Cdk1, thus inactivating cyclin-dependent kinase (CDK) activity and preventing entry into mitosis. b, In long cells, the Pom1 gradient does not reach the cortical nodes (red dots), and therefore Cdr2 and Cdr1 remain active in the nodes. Cdr2 and Cdr1 inhibit Wee1, preventing phosphorylation of Cdk1 and thereby leading to activation of CDK and mitotic entry. (This simplified diagram omits several other regulators of CDK activity.)

Mating-type switching in fission yeast[14] [edit]

Fission yeast switches mating type by a replication-coupled recombination event, which takes place during S phase of the cell cycle. Fission yeast uses intrinsic asymmetry of the DNA replication process to switch the mating type; it was the first system, where the direction of replication was shown to be required for the change of the cell type. Studies of the mating-type switching system lead to a discovery and characterization of a site-specific replication termination site RTS1, a site-specific replication pause site MPS1, and a novel type of chromosomal imprint, marking one of the sister chromatids at the mating-type locus mat1. In addition, work on the silenced donor region has led to great advances in understanding formation and maintenance of heterochromatin.

References [edit]

  1. ^ Wilhelm et al. Dynamic repertoire of a eukaryotic transcriptome surveyed at single-nucleotide resolution. Nature (2008) vol. 453 (7199) pp. 1239–1243
  2. ^ Leupold U. (1950) Die Vererbung von Homothallie und Heterothallie bei Schizosaccharomyces pombe. CR Trav Lab Carlsberg Ser Physiol 24:381–480.
  3. ^ Leupold U. (1993) The origins of Schizosaccharomyces pombe genetics. In: Hall MN, Linder P. eds. The Early Days of Yeast Genetics . New York. Cold Spring Harbor Laboratory Press. p 125–128.
  4. ^ Mitchison JM. (1957) The growth of single cells. I. Schizosaccharomyces pombe. Exp Cell Res 13:244–262.
  5. ^ Mitchison JM. (1990) My favourite cell: The fission yeast, Schizosaccharomyces pombe. Bioessays 4:189–191.
  6. ^ Florenzano et al. Contributo alla ecologia dei lieviti Schizosaccharomyces sulle uve. Vitis (1977) 16; 38–44
  7. ^ Gómez et al. (2002) Fission yeast enters a joyful new era. Genome Biol. 3(6):REPORTS 4017
  8. ^ Lin et al. (2011). The evolution of aerobic fermentation in Schizosaccharomyces pombe was associated with regulatory reprogramming but not nucleosome reorganization. Mol Biol Evol. 28; 1407-13
  9. ^ Grunstein, Michael, and Susan Gasser. "Epigenetics in Saccharomyces cerevisiae." Epigenetics. 1. Cold Spring Harbor Press, 2007.
  10. ^ Cell Cycle. Principles of Control” by David O Morgan, Primers in Biology
  11. ^ A spatial gradient coordinates cell size and mitotic entry in fission yeast by James B. Moseley, Adeline Mayeux, Anne Paoletti & Paul Nurse, Nature, 11 June 2009
  12. ^ Polar gradients of the DYRK-family kinase Pom1 couple cell length with the cell cycle. Sophie G Martin and Martine Berthelot-Grosjean. Nature. 2009
  13. ^ Cell cycle: Cell division brought down to size” by Kenneth E. Sawin, Nature 459, 782-783(11 June 2009)
  14. ^ Lessons learned from studies of fission yeast mating-type switching and silencing by Amar J.S. Klar, Annual Review of Genetics, 5 July 2007

External links [edit]
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