Saccharomyces cerevisiae (/ˌsɛrɪˈvɪsiiː/) is a species of
yeast. It has been instrumental to winemaking, baking, and brewing
since ancient times. It is believed to have been originally isolated
from the skin of grapes (one can see the yeast as a component of the
thin white film on the skins of some dark-colored fruits such as
plums; it exists among the waxes of the cuticle). It is one of the
most intensively studied eukaryotic model organisms in molecular and
cell biology, much like
Escherichia coli as the model bacterium. It is
the microorganism behind the most common type of fermentation.
S. cerevisiae cells are round to ovoid, 5–10 μm in
diameter. It reproduces by a division process known as budding.
Many proteins important in human biology were first discovered by
studying their homologs in yeast; these proteins include cell cycle
proteins, signaling proteins, and protein-processing enzymes. S.
cerevisiae is currently the only yeast cell known to have Berkeley
bodies present, which are involved in particular secretory pathways.
Antibodies against S. cerevisiae are found in 60–70% of
Crohn's disease and 10–15% of patients with ulcerative
colitis (and 8% of healthy controls).
3.3 Nutritional requirements
3.5 Cell cycle
18.104.22.168 Actomyosin ring and primary septum formation
22.214.171.124 Differences from fission yeast
4 In biological research
4.1 Model organism
4.2 In the study of aging
4.3 Meiosis, recombination and DNA repair
Gene function and interactions
4.6 Other tools in yeast research
4.7 Synthetic yeast genome project
5 In commercial applications
5.3 Uses in aquaria
6 See also
8 Further reading
9 External links
"Saccharomyces" derives from Latinized Greek and means "sugar-mold" or
"sugar-fungus", saccharon (σάκχαρον) being the combining form
"sugar" and myces (μύκης, genitive μύκητος) being
"fungus". Cerevisiae comes from Latin and means "of beer". Other names
for the organism are:
Brewer's yeast, though other species are also used in brewing
Ragi yeast, in connection to making Tapai
This species is also the main source of nutritional yeast and yeast
In the 19th century, bread bakers obtained their yeast from beer
brewers, and this led to sweet-fermented breads such as the Imperial
"Kaisersemmel" roll, which in general lacked the sourness created
by the acidification typical of Lactobacillus. However, beer brewers
slowly switched from top-fermenting (S. cerevisiae) to
bottom-fermenting (S. pastorianus) yeast and this created a shortage
of yeast for making bread, so the Vienna Process was developed in
1846. While the innovation is often popularly credited for using
steam in baking ovens, leading to a different crust characteristic, it
is notable for including procedures for high milling of grains (see
Vienna grits), cracking them incrementally instead of mashing them
with one pass; as well as better processes for growing and harvesting
top-fermenting yeasts, known as press-yeast.
Refinements in microbiology following the work of
Louis Pasteur led to
more advanced methods of culturing pure strains. In 1879, Great
Britain introduced specialized growing vats for the production of S.
cerevisiae, and in the United States around the turn of the century
centrifuges were used for concentrating the yeast, making modern
commercial yeast possible, and turning yeast production into a major
industrial endeavor. The slurry yeast made by small bakers and grocery
shops became cream yeast, a suspension of live yeast cells in growth
medium, and then compressed yeast, the fresh cake yeast that became
the standard leaven for bread bakers in much of the Westernized world
during the early 20th century.
During World War II, Fleischmann's developed a granulated active dry
yeast for the United States armed forces, which did not require
refrigeration and had a longer shelf-life and better temperature
tolerance than fresh yeast; it is still the standard yeast for US
military recipes. The company created yeast that would rise twice as
fast, cutting down on baking time. Lesaffre would later create instant
yeast in the 1970s, which has gained considerable use and market share
at the expense of both fresh and dry yeast in their various
Yeast colonies on an agar plate.
In nature, yeast cells are found primarily on ripe fruits such as
grapes (before maturation, grapes are almost free of yeasts). Since
S. cerevisiae is not airborne, it requires a vector to move.
Queens of social wasps overwintering as adults (
Vespa crabro and
Polistes spp.) can harbor yeast cells from autumn to spring and
transmit them to their progeny. The intestine of Polistes
dominula, a social wasp, hosts S. cerevisiae strains as well as S.
cerevisiae × S. paradoxus hybrids. Stefanini et al. (2016) showed
that the intestine of
Polistes dominula favors the mating of S.
cerevisiae strains, both among themselves and with S. paradoxus cells
by providing environmental conditions prompting cell sporulation and
The optimum temperature for growth of S. cerevisiae is
30–35 °C (86–95 °F).
Two forms of yeast cells can survive and grow: haploid and diploid.
The haploid cells undergo a simple lifecycle of mitosis and growth,
and under conditions of high stress will, in general, die. This is the
asexual form of the fungus. The diploid cells (the preferential 'form'
of yeast) similarly undergo a simple lifecycle of mitosis and growth.
The rate at which the mitotic cell cycle progresses often differs
substantially between haploid and diploid cells. Under conditions
of stress, diploid cells can undergo sporulation, entering meiosis and
producing four haploid spores, which can subsequently mate. This is
the sexual form of the fungus. Under optimal conditions, yeast cells
can double their population every 100 minutes. However,
growth rates vary enormously both between strains and between
environments. Mean replicative lifespan is about 26 cell
In the wild, recessive deleterious mutations accumulate during long
periods of asexual reproduction of diploids, and are purged during
selfing: this purging has been termed "genome renewal".
Yeast assimilable nitrogen
All strains of S. cerevisiae can grow aerobically on glucose,
maltose, and trehalose and fail to grow on lactose and cellobiose.
However, growth on other sugars is variable. Galactose and fructose
are shown to be two of the best fermenting sugars. The ability of
yeasts to use different sugars can differ depending on whether they
are grown aerobically or anaerobically. Some strains cannot grow
anaerobically on sucrose and trehalose.
All strains can use ammonia and urea as the sole nitrogen source, but
cannot use nitrate, since they lack the ability to reduce them to
ammonium ions. They can also use most amino acids, small peptides, and
nitrogen bases as nitrogen sources. Histidine, glycine, cystine, and
lysine are, however, not readily used. S. cerevisiae does not
excrete proteases, so extracellular protein cannot be metabolized.
Yeasts also have a requirement for phosphorus, which is assimilated as
a dihydrogen phosphate ion, and sulfur, which can be assimilated as a
sulfate ion or as organic sulfur compounds such as the amino acids
methionine and cysteine. Some metals, like magnesium, iron, calcium,
and zinc, are also required for good growth of the yeast.
Concerning organic requirements, most strains of S. cerevisiae require
biotin. Indeed, a S. cerevisiae-based growth assay laid the foundation
for the isolation, crystallisation, and later structural determination
of biotin. Most strains also require pantothenate for full growth. In
general, S. cerevisiae is prototrophic for vitamins.
Saccharomyces cerevisiae mating type a with a cellular bulging called
a shmoo in response to α-factor
Main article: Mating of yeast
Yeast has two mating types, a and α (alpha), which show primitive
aspects of sex differentiation. As in many other eukaryotes,
mating leads to genetic recombination, i.e. production of novel
combinations of chromosomes. Two haploid yeast cells of opposite
mating type can mate to form diploid cells that can either sporulate
to form another generation of haploid cells or continue to exist as
diploid cells. Mating has been exploited by biologists as a tool to
combine genes, plasmids, or proteins at will.
The mating pathway employs a G protein-coupled receptor, G protein,
RGS protein, and three-tiered
MAPK signaling cascade that is
homologous to those found in humans. This feature has been exploited
by biologists to investigate basic mechanisms of signal transduction
Growth in yeast is synchronised with the growth of the bud, which
reaches the size of the mature cell by the time it separates from the
parent cell. In well nourished, rapidly growing yeast cultures, all
the cells can be seen to have buds, since bud formation occupies the
whole cell cycle. Both mother and daughter cells can initiate bud
formation before cell separation has occurred. In yeast cultures
growing more slowly, cells lacking buds can be seen, and bud formation
only occupies a part of the cell cycle.
Cytokinesis enables budding yeast
Saccharomyces cerevisiae to divide
into two daughter cells. S. cerevisiae forms a bud which can grow
throughout its cell cycle and later leaves its mother cell when
mitosis has completed.
S. cerevisiae is relevant to cell cycle studies because it divides
asymmetrically by using a polarized cell to make two daughters with
different fates and sizes. Similarly, stem cells use asymmetric
division for self-renewal and differentiation.
For many cells, M phase does not happen until S phase is complete.
However, for entry into mitosis in S. cerevisiae this is not true.
Cytokinesis begins with the budding process in late G1 and is not
completed until about halfway through the next cycle. The assembly of
the spindle can happen before S phase has finished duplicating the
chromosomes. Additionally, there is a lack of clearly defined G2
in between M and S. Thus, there is a lack of extensive regulation
present in higher eukaryotes.
When the daughter emerges, the daughter is two-thirds the size of the
mother. Throughout the process, the mother displays little to no
change in size. The RAM pathway is activated in the daughter cell
immediately after cytokinesis is complete. This pathway makes sure
that the daughter has separated properly.
Actomyosin ring and primary septum formation
Two interdependent events drive cytokinesis in S. cerevisiae. The
first event is contractile actomyosin ring (AMR) constriction and the
second event is formation of the primary septum (PS), a chitinous cell
wall structure that can only be formed during cytokinesis. The PS
resembles in animals the process of extracellular matrix
remodeling. When the AMR constricts, the PS begins to grow.
Disrupting AMR misorients the PS, suggesting that both have a
dependent role. Additionally, disrupting the PS also leads to
disruptions in the AMR, suggesting both the actomyosin ring and
primary septum have an interdependent relationship.
The AMR, which is attached to the cell membrane facing the cytosol,
consists of actin and myosin II molecules that coordinate the cells to
split. The ring is thought to play an important role in ingression
of the plasma membrane as a contractile force.
Proper coordination and correct positional assembly of the contractile
ring depends on septins, which is the precursor to the septum ring.
These GTPases assemble complexes with other proteins. The septins form
a ring at the site where the bud will be created during late G1. They
help promote the formation of the actin-myosin ring, although this
mechanism is unknown. It is suggested they help provide structural
support for other necessary cytokinesis processes. After a bud
emerges, the septin ring forms an hourglass. The septin hourglass and
the myosin ring together are the beginning of the future division
The septin and AMR complex progress to form the primary septum
consisting of glucans and other chitinous molecules sent by vesicles
from the Golgi body. After AMR constriction is complete, two
secondary septums are formed by glucans. How the AMR ring dissembles
remains poorly unknown.
Microtubules do not play as significant a role in cytokinesis compared
to the AMR and septum. Disruption of microtubules did not
significantly impair polarized growth. Thus, the AMR and septum
formation are the major drivers of cytokinesis.
Differences from fission yeast
Budding yeast form a bud from the mother cell. This bud grows during
the cell cycle and detaches; fission yeast divide by forming a cell
Cytokinesis begins at G1 for budding yeast, while cytokinesis begins
at G2 for fission yeast. Fission yeast “select” the midpoint,
whereas budding yeast “select” a bud site 
During early anaphase the actomyosin ring and septum continues to
develop in budding yeast, in fission yeast during metaphase-anaphase
the actomyosin ring begins to develop 
In biological research
S. cerevisiae, differential interference contrast image
Numbered ticks are 11 micrometers apart.
When researchers look for an organism to use in their studies, they
look for several traits. Among these are size, generation time,
accessibility, manipulation, genetics, conservation of mechanisms, and
potential economic benefit. The yeast species S. pombe and S.
cerevisiae are both well studied; these two species diverged
approximately 600 to 300 million years ago, and are
significant tools in the study of DNA damage and repair
S. cerevisiae has developed as a model organism because it scores
favorably on a number of these criteria.
As a single-cell organism, S. cerevisiae is small with a short
generation time (doubling time 1.25–2 hours at 30 °C or
86 °F) and can be easily cultured. These are all positive
characteristics in that they allow for the swift production and
maintenance of multiple specimen lines at low cost.
S. cerevisiae divides with meiosis, allowing it to be a candidate for
sexual genetics research.
S. cerevisiae can be transformed allowing for either the addition of
new genes or deletion through homologous recombination. Furthermore,
the ability to grow S. cerevisiae as a haploid simplifies the
creation of gene knockout strains.
As a eukaryote, S. cerevisiae shares the complex internal cell
structure of plants and animals without the high percentage of
non-coding DNA that can confound research in higher eukaryotes.
S. cerevisiae research is a strong economic driver, at least
initially, as a result of its established use in industry.
In the study of aging
S. cerevisiae has been highly studied as a model organism to better
understand aging for more than five decades and has contributed to the
identification of more mammalian genes affecting aging than any other
model organism. Some of the topics studied using yeast are calorie
restriction, as well as in genes and cellular pathways involved in
senescence. The two most common methods of measuring aging in yeast
Life Span, which measures the number of times a cell
divides, and Chronological
Life Span, which measures how long a cell
can survive in a non-dividing stasis state. Limiting the amount of
glucose or amino acids in the growth medium has been shown to increase
RLS and CLS in yeast as well as other organisms. At first, this
was thought to increase RLS by up-regulating the sir2 enzyme, however
it was later discovered that this effect is independent of sir2.
Over-expression of the genes sir2 and fob1 has been shown to increase
RLS by preventing the accumulation of extrachromosomal rDNA circles,
which are thought to be one of the causes of senescence in yeast.
The effects of dietary restriction may be the result of a decreased
signaling in the TOR cellular pathway. This pathway modulates the
cell's response to nutrients, and mutations that decrease TOR activity
were found to increase CLS and RLS. This has also been shown
to be the case in other animals. A yeast mutant lacking the
genes sch9 and ras2 has recently been shown to have a tenfold increase
in chronological lifespan under conditions of calorie restriction and
is the largest increase achieved in any organism.
Mother cells give rise to progeny buds by mitotic divisions, but
undergo replicative aging over successive generations and ultimately
die. However, when a mother cell undergoes meiosis and gametogenesis,
lifespan is reset. The replicative potential of gametes (spores)
formed by aged cells is the same as gametes formed by young cells,
indicating that age-associated damage is removed by meiosis from aged
mother cells. This observation suggests that during meiosis removal of
age-associated damages leads to rejuvenation. However, the nature of
these damages remains to be established.
Meiosis, recombination and DNA repair
S. cerevisiae reproduces by mitosis as diploid cells when nutrients
are abundant. However, when starved, these cells undergo meiosis to
form haploid spores.
Evidence from studies of S. cerevisiae bear on the adaptive function
of meiosis and recombination. Mutations defective in genes essential
for meiotic and mitotic recombination in S. cerevisiae cause increased
sensitivity to radiation or DNA damaging chemicals. For
instance, gene rad52 is required for both meiotic recombination
and mitotic recombination. Rad52 mutants have increased
sensitivity to killing by X-rays,
Methyl methanesulfonate and the DNA
cross-linking agent 8-methoxypsoralen-plus-UVA, and show reduced
meiotic recombination. These findings suggest that
recombination repair during meiosis and mitosis is needed for repair
of the different damages caused by these agents.
Ruderfer et al. (2006) analyzed the ancestry of natural S.
cerevisiae strains and concluded that outcrossing occurs only about
once every 50,000 cell divisions. Thus, it appears that in nature,
mating is likely most often between closely related yeast cells.
Mating occurs when haploid cells of opposite mating type MATa and
MATα come into contact. Ruderfer et al. pointed out that such
contacts are frequent between closely related yeast cells for two
reasons. The first is that cells of opposite mating type are present
together in the same ascus, the sac that contains the cells directly
produced by a single meiosis, and these cells can mate with each
other. The second reason is that haploid cells of one mating type,
upon cell division, often produce cells of the opposite mating type
with which they can mate. The relative rarity in nature of meiotic
events that result from outcrossing is inconsistent with the idea that
production of genetic variation is the main selective force
maintaining meiosis in this organism. However, this finding is
consistent with the alternative idea that the main selective force
maintaining meiosis is enhanced recombinational repair of DNA
damage, since this benefit is realized during each
meiosis, whether or not out-crossing occurs.
S. cerevisiae was the first eukaryotic genome to be completely
sequenced. The genome sequence was released to the public domain
on April 24, 1996. Since then, regular updates have been maintained at
Genome Database. This database is a highly annotated
and cross-referenced database for yeast researchers. Another important
S. cerevisiae database is maintained by the Munich Information
Center for Protein Sequences (MIPS). The S. cerevisiae genome is
composed of about 12,156,677 base pairs and 6,275 genes, compactly
organized on 16 chromosomes. Only about 5,800 of these genes are
believed to be functional. It is estimated at least 31% of yeast genes
have homologs in the human genome.
Yeast genes are classified
using gene symbols (such as sch9) or systematic names. In the latter
case the 16 chromosomes of yeast are represented by the letters A to
P, then the gene is further classified by a sequence number on the
left or right arm of the chromosome, and a letter showing which of the
two DNA strands contains its coding sequence.
Systematic gene names for Baker's yeast
Example gene name
the Y to show this is a yeast gene
chromosome on which the gene is located
left or right arm of the chromosome
sequence number of the gene/ORF on this arm, starting at the
whether the coding sequence is on the Watson or Crick strand
YBR134C (aka SUP45 encoding eRF1, a translation termination factor) is
located on the right arm of chromosome 2 and is the 134th open reading
frame (ORF) on that arm, starting from the centromere. The coding
sequence is on the Crick strand of the DNA.
YDL102W (aka POL3 encoding a subunit of DNA polymerase delta) is
located on the left arm of chromosome 4; it is the 102nd ORF from the
centromere and codes from the Watson strand of the DNA.
Gene function and interactions
The availability of the S. cerevisiae genome sequence and a set
of deletion mutants covering 90% of the yeast genome has further
enhanced the power of S. cerevisiae as a model for understanding
the regulation of eukaryotic cells. A project underway to analyze the
genetic interactions of all double-deletion mutants through synthetic
genetic array analysis will take this research one step further. The
goal is to form a functional map of the cell's processes. As of 2010 a
model of genetic interactions is most comprehensive yet to be
constructed, containing "the interaction profiles for ~75% of all
genes in the
Budding yeast". This model was made from 5.4 million
two-gene comparisons in which a double gene knockout for each
combination of the genes studied was performed. The effect of the
double knockout on the fitness of the cell was compared to the
expected fitness. Expected fitness is determined from the sum of the
results on fitness of single-gene knockouts for each compared gene.
When there is a change in fitness from what is expected, the genes are
presumed to interact with each other. This was tested by comparing the
results to what was previously known. For example, the genes Par32,
Ecm30, and Ubp15 had similar interaction profiles to genes involved in
the Gap1-sorting module cellular process. Consistent with the results,
these genes, when knocked out, disrupted that process, confirming that
they are part of it. From this, 170,000 gene interactions were
found and genes with similar interaction patterns were grouped
together. Genes with similar genetic interaction profiles tend to be
part of the same pathway or biological process. This information
was used to construct a global network of gene interactions organized
by function. This network can be used to predict the function of
uncharacterized genes based on the functions of genes they are grouped
Other tools in yeast research
Approaches that can be applied in many different fields of biological
and medicinal science have been developed by yeast scientists. These
include yeast two-hybrid for studying protein interactions and tetrad
analysis. Other resources, include a gene deletion library including
~4,700 viable haploid single gene deletion strains. A GFP fusion
strain library used to study protein localisation and a TAP tag
library used to purify protein from yeast cell extracts.[citation
Synthetic yeast genome project
The international Synthetic
Genome Project (Sc2.0 or
Saccharomyces cerevisiae version 2.0) aims to build an entirely
designer, customizable, synthetic S. cerevisiae genome from scratch
that is more stable than the wild type. In the synthetic genome all
transposons, repetitive elements and many introns are removed, all UAG
stop codons are replaced with UAA, and transfer RNA genes are moved to
a novel neochromosome. As of March 2017[update], 6 of the 16
chromosomes have been synthesized and tested. No significant fitness
defects have been found.
Among other microorganisms, a sample of living S. cerevisiae was
included in the Living Interplanetary Flight Experiment, which would
have completed a three-year interplanetary round-trip in a small
capsule aboard the Russian
Fobos-Grunt spacecraft, launched in late
2011. The goal was to test whether selected organisms could
survive a few years in deep space by flying them through
interplanetary space. The experiment would have tested one aspect of
transpermia, the hypothesis that life could survive space travel, if
protected inside rocks blasted by impact off one planet to land on
another. Fobos-Grunt's mission ended unsuccessfully,
however, when it failed to escape low Earth orbit. The spacecraft
along with its instruments fell into the Pacific Ocean in an
uncontrolled re-entry on January 15, 2012. The next planned exposure
mission in deep space using S. cerevisiae is BioSentinel. (see: List
of microorganisms tested in outer space)
In commercial applications
Yeast in winemaking
Saccharomyces cerevisiae is used in brewing beer, when it is sometimes
called a top-fermenting or top-cropping yeast. It is so called because
during the fermentation process its hydrophobic surface causes the
flocs to adhere to CO2 and rise to the top of the fermentation vessel.
Top-fermenting yeasts are fermented at higher temperatures than the
Saccharomyces pastorianus, and the resulting beers have a
different flavor than the same beverage fermented with a lager yeast.
"Fruity esters" may be formed if the yeast undergoes temperatures near
21 °C (70 °F), or if the fermentation temperature of the
beverage fluctuates during the process. Lager yeast normally ferments
at a temperature of approximately 5 °C (41 °F), where
Saccharomyces cerevisiae becomes dormant.
In May 2013, the
Oregon legislature made S. cerevisiae the official
state microbe in recognition of the impact craft beer brewing has had
on the state economy and the state's identity as the craft
beer-brewing capital of the United States.
Main article: Baker's yeast
S. cerevisiae is used in baking; the carbon dioxide generated by the
fermentation is used as a leavening agent in bread and other baked
goods. Historically, this use was closely linked to the brewing
industry's use of yeast, as bakers took or bought the barm or
yeast-filled foam from brewing ale from the brewers (producing the
barm cake); today, brewing and baking yeast strains are somewhat
Uses in aquaria
Owing to the high cost of commercial CO2 cylinder systems, CO2
injection by yeast is one of the most popular DIY approaches followed
by aquaculturists for providing CO2 to underwater aquatic plants. The
yeast culture is, in general, maintained in plastic bottles, and
typical systems provide one bubble every 3–7 seconds. Various
approaches have been devised to allow proper absorption of the gas
into the water.
Saccharomyces cerevisiae extracts: Vegemite, Marmite, Cenovis,
Yeast Extract, mannan oligosaccharides, pgg-glucan, zymosan
Saccharomyces cerevisiae boulardii (
Fungus – Cofactor Ora". cofactor.io.
Retrieved December 19, 2017.
^ Feldmann, Horst (2010). Yeast. Molecular and Cell bio.
Wiley-Blackwell. ISBN 352732609X. [page needed]
^ Walker LJ, Aldhous MC, Drummond HE, Smith BR, Nimmo ER, Arnott ID,
Satsangi J (2004). "Anti-
Saccharomyces cerevisiae antibodies (ASCA) in
Crohn's disease are associated with disease severity but not
NOD2/CARD15 mutations". Clin. Exp. Immunol. 135 (3): 490–96.
doi:10.1111/j.1365-2249.2003.02392.x. PMC 1808965 .
^ a b Moyad MA (2008). "Brewer's/baker's yeast (Saccharomyces
cerevisiae) and preventive medicine: Part II". Urol Nurs. 28 (1):
73–75. PMID 18335702.
^ Eben Norton Horsford (1875). Report on Vienna bread. U.S. Government
Printing Office. p. 86.
^ Kristiansen, B.; Ratledge, Colin (2001). Basic biotechnology.
Cambridge, UK: Cambridge University Press. p. 378.
^ Eben Norton Horsford (1875). Report on Vienna bread. U.S. Government
Printing Office. pp. 31–32.
^ Marx, Jean & Litchfield, John H. (1989). A Revolution in
biotechnology. Cambridge, UK: Cambridge University Press. p. 71.
^ Marshall, Charles, ed. (June 1912). Microbiology. P. Blakiston's son
& Company. p. 420. Retrieved November 5, 2014.
^ a b Stefanini I, Dapporto L, Legras JL, Calabretta A, Di Paola M, De
Filippo C, Viola R, Capretti P, Polsinelli M, Turillazzi S, Cavalieri
D (2012). "Role of social wasps in
Saccharomyces cerevisiae ecology
and evolution". Proc. Natl. Acad. Sci. U.S.A. 109 (33): 13398–403.
PMC 3421210 . PMID 22847440.
^ Stefanini I, Dapporto L, Berná L, Polsinelli M, Turillazzi S,
Cavalieri D (2016). "Social wasps are a
Saccharomyces mating nest".
Proc. Natl. Acad. Sci. U.S.A. 113 (8): 2247–51.
PMC 4776513 . PMID 26787874.
^ Zörgö E, Chwialkowska K, Gjuvsland AB, Garré E, Sunnerhagen P,
Liti G, Blomberg A, Omholt SW, Warringer J (2013). "Ancient
evolutionary trade-offs between yeast ploidy states". PLoS Genet. 9
(3): e1003388. doi:10.1371/journal.pgen.1003388. PMC 3605057 .
^ Herskowitz I (1988). "
Life cycle of the budding yeast Saccharomyces
cerevisiae". Microbiol. Rev. 52 (4): 536–53. PMC 373162 .
^ Friedman, Nir (January 3, 2011). "The Friedman Lab Chronicles".
Growing yeasts (Robotically).
Nir Friedman Lab. Retrieved
^ Warringer J, Zörgö E, Cubillos FA, Zia A, Gjuvsland A, Simpson JT,
Forsmark A, Durbin R, Omholt SW, Louis EJ, Liti G, Moses A, Blomberg A
(2011). "Trait variation in yeast is defined by population history".
PLoS Genet. 7 (6): e1002111. doi:10.1371/journal.pgen.1002111.
PMC 3116910 . PMID 21698134.
^ Kaeberlein M, Powers RW, Steffen KK, Westman EA, Hu D, Dang N, Kerr
EO, Kirkland KT, Fields S, Kennedy BK (2005). "Regulation of yeast
replicative life span by TOR and Sch9 in response to nutrients".
Science. 310 (5751): 1193–96. Bibcode:2005Sci...310.1193K.
doi:10.1126/science.1115535. PMID 16293764.
^ Kaeberlein M (2010). "Lessons on longevity from budding yeast".
Nature. 464 (7288): 513–19. Bibcode:2010Natur.464..513K.
doi:10.1038/nature08981. PMC 3696189 .
^ Mortimer, Robert K.; Romano, Patrizia; Suzzi, Giovanna; Polsinelli,
Mario (December 1994). "
Genome renewal: A new phenomenon revealed from
a genetic study of 43 strains of
Saccharomyces cerevisiae derived from
natural fermentation of grape musts". Yeast. 10 (12): 1543–52.
doi:10.1002/yea.320101203. PMID 7725789.
^ Masel, Joanna; Lyttle, David N. (December 2011). "The consequences
of rare sexual reproduction by means of selfing in an otherwise
clonally reproducing species". Theoretical Population Biology. 80 (4):
317–22. doi:10.1016/j.tpb.2011.08.004. PMC 3218209 .
^ a b c d e f Morgan, David (2007). The Cell Cycle: Principles of
Control. Sinauer Associates.
^ a b Bi, Erfei (2017). "Mechanics and regulation of cytokinesis in
budding yeast", Seminars in Cell & Developmental Biology, 66:
^ a b c Wloka, Carsten (2012). "Mechanisms of cytokinesis in budding
yeast", Cytoskeleton, 69(10): 710–26.
^ a b Bi, Erfei (2002). "Cytokinesis in
Budding Yeast: the
Relationship between Actomyosin Ring Function and Septum Formation",
Cell Structure and Function, 26(6): 529–37
^ Fang, X (2010). "Biphasic targeting and cleavage furrow ingression
directed by the tail of a myosin-II", J Cell Biol 191: 1333–50.
^ VerPlank, Lynn (2005). "Cell cycle-regulated trafficking of Chs2
controls actomyosin ring stability during cytokinesis", Mol. Biol.
Cell, 16: 2529–43
^ Adams, A (1984). “Relationship of actin and tubulin distribution
to bud growth in wild-type and morphogenetic-mutant Saccharomyces
cerevisiae”, ‘’J. Cell Biol.’’ 98: 934–945
^ a b Balasubramanian, Mohan (2004). "Comparative Analysis of
Budding Yeast, Fission
Yeast and Animal Cells", Curr.
Biology, 14(18): R806–18.
^ Nickoloff, Jac A.; Haber, James E. (2011). "Mating-Type Control of
DNA Repair and Recombination in
Saccharomyces cerevisiae". In
Nickoloff, Jac A.; Hoekstra, Merl F. DNA Damage and Repair.
Contemporary Cancer Research. pp. 107–24.
doi:10.1007/978-1-59259-095-7_5. ISBN 978-1-59259-095-7.
^ Boekhout, T.; Robert, V., eds. (2003).
Yeasts in Food: Beneficial
and Detrimental aspects. Behr's Verlag. p. 322.
ISBN 978-3-86022-961-3. Retrieved January 10, 2011.
^ a b c d e Longo VD, Shadel GS, Kaeberlein M, Kennedy B (2012).
"Replicative and chronological aging in
Cell Metab. 16 (1): 18–31. doi:10.1016/j.cmet.2012.06.002.
PMC 3392685 . PMID 22768836.
^ a b c d Kaeberlein M, Burtner CR, Kennedy BK (2007). "Recent
developments in yeast aging". PLoS Genet. 3 (5): 655–60.
doi:10.1371/journal.pgen.0030084. PMC 1877880 .
^ Wei M, Fabrizio P, Hu J, Ge H, Cheng C, Li L, Longo VD (2008). "Life
span extension by calorie restriction depends on Rim15 and
transcription factors downstream of Ras/PKA, Tor, and Sch9". PLoS
Genet. 4 (1): 139–49. doi:10.1371/journal.pgen.0040013.
PMC 2213705 . PMID 18225956.
Life Span Extension Reported". University of Southern
California. Archived from the original on 2016-03-04.
^ Unal E, Kinde B, Amon A (2011). "
age-induced cellular damage and resets life span in yeast". Science.
332 (6037): 1554–57. Bibcode:2011Sci...332.1554U.
doi:10.1126/science.1204349. PMC 3923466 .
^ Herskowitz I (1988). "
Life cycle of the budding yeast Saccharomyces
cerevisiae". Microbiol. Rev. 52 (4): 536–53. PMC 373162 .
^ a b c Ruderfer DM, Pratt SC, Seidel HS, Kruglyak L (2006).
"Population genomic analysis of outcrossing and recombination in
yeast". Nat. Genet. 38 (9): 1077–81. doi:10.1038/ng1859.
^ a b Haynes, Robert H.; Kunz, Bernard A. (1981). "
DNA repair and
mutagenesis in yeast". In Strathern, Jeffrey N.; Jones, Elizabeth W.;
Broach, James R. The Molecular Biology of the
Life Cycle and Inheritance. Cold Spring Harbor, N.Y.: Cold Spring
Harbor Laboratory. pp. 371–414.
^ a b Game JC, Zamb TJ, Braun RJ, Resnick M, Roth RM (1980). "The Role
Radiation (rad) Genes in Meiotic Recombination in Yeast". Genetics.
94 (1): 51–68. PMC 1214137 . PMID 17248996.
^ Malone RE, Esposito RE (1980). "The
RAD52 gene is required for
homothallic interconversion of mating types and spontaneous mitotic
recombination in yeast". Proc. Natl. Acad. Sci. U.S.A. 77 (1):
503–07. Bibcode:1980PNAS...77..503M. doi:10.1073/pnas.77.1.503.
PMC 348300 . PMID 6987653.
^ Henriques, J. A. P.; Moustacchi, E. (1980). "Sensitivity to
Photoaddition of Mono-And Bifunctional Furocoumarins of X-Ray
Sensitive Mutants of
Saccharomyces Cerevisiae". Photochemistry and
Photobiology. 31 (6): 557–63.
^ Birdsell, John A.; Wills, Christopher (2003). "The Evolutionary
Origin and Maintenance of Sexual Recombination: A Review of
Contemporary Models". Evolutionary Biology. pp. 27–138.
doi:10.1007/978-1-4757-5190-1_2. ISBN 978-1-4419-3385-0.
^ Bernstein, Harris; Bernstei, Carol (2013). "Evolutionary Origin and
Adaptive Function of Meiosis". Meiosis. doi:10.5772/56557.
^ Hrandl, Elvira (2013). "
Meiosis and the Paradox of Sex in Nature".
Meiosis. doi:10.5772/56542. ISBN 978-953-51-1197-9.
^ Goffeau A, Barrell BG, Bussey H, Davis RW, Dujon B, Feldmann H,
Galibert F, Hoheisel JD, Jacq C, Johnston M, Louis EJ, Mewes HW,
Murakami Y, Philippsen P, Tettelin H, Oliver SG (1996). "
6000 genes". Science. 274 (5287): 546, 563–67.
^ Botstein D, Chervitz SA, Cherry JM (1997). "
Yeast as a model
organism". Science. 277 (5330): 1259–60.
doi:10.1126/science.277.5330.1259. PMC 3039837 .
^ "YeastDeletionWeb". Retrieved 2013-05-25.
^ a b c Costanzo M, Baryshnikova A, Bellay J, Kim Y, Spear ED, Sevier
CS, Ding H, Koh JL, Toufighi K, Mostafavi S, Prinz J, St Onge RP,
VanderSluis B, Makhnevych T, Vizeacoumar FJ, Alizadeh S, Bahr S, Brost
RL, Chen Y, Cokol M, Deshpande R, Li Z, Lin ZY, Liang W, Marback M,
Paw J, San Luis BJ, Shuteriqi E, Tong AH, van Dyk N, Wallace IM,
Whitney JA, Weirauch MT, Zhong G, Zhu H, Houry WA, Brudno M,
Ragibizadeh S, Papp B, Pál C, Roth FP, Giaever G, Nislow C,
Troyanskaya OG, Bussey H, Bader GD, Gingras AC, Morris QD, Kim PM,
Kaiser CA, Myers CL, Andrews BJ, Boone C (2010). "The genetic
landscape of a cell". Science. 327 (5964): 425–31.
PMC 5600254 . PMID 20093466.
^ Tong AH, Lesage G, Bader GD, Ding H, Xu H, Xin X, Young J, Berriz
GF, Brost RL, Chang M, Chen Y, Cheng X, Chua G, Friesen H, Goldberg
DS, Haynes J, Humphries C, He G, Hussein S, Ke L, Krogan N, Li Z,
Levinson JN, Lu H, Ménard P, Munyana C, Parsons AB, Ryan O, Tonikian
R, Roberts T, Sdicu AM, Shapiro J, Sheikh B, Suter B, Wong SL, Zhang
LV, Zhu H, Burd CG, Munro S, Sander C, Rine J, Greenblatt J, Peter M,
Bretscher A, Bell G, Roth FP, Brown GW, Andrews B, Bussey H, Boone C
(2004). "Global mapping of the yeast genetic interaction network".
Science. 303 (5659): 808–13. Bibcode:2004Sci...303..808T.
doi:10.1126/science.1091317. PMID 14764870.
Special Issue Synthetic
Yeast Genome", Science, 10 March 2017 Vol
355, Issue 6329
^ a b Warmflash, David; Ciftcioglu, Neva; Fox, George; McKay, David
S.; Friedman, Louis; Betts, Bruce; Kirschvink, Joseph (November 5–7,
2007). Living interplanetary flight experiment (LIFE): An experiment
on the survivalability of microorganisms during interplanetary travel
(PDF). Workshop on the Exploration of Phobos and Deimos. Ames Research
^ a b "Projects: LIFE Experiment: Phobos". The Planetary Society.
Archived from the original on 16 March 2011. Retrieved 2 April
^ Anatoly Zak (1 September 2008). "Mission Possible". Air & Space
Magazine. Smithsonian Institution. Retrieved 26 May 2009.
Saccharomyces cerevisiae as official microbe of State of
Oregon State Legislature.
^ "CO2 Injection: The
Yeast Method". www.thekrib.com. Retrieved
Arroyo-López FN, Orlić S, Querol A, Barrio E (2009). "Effects of
temperature, pH and sugar concentration on the growth parameters of
Saccharomyces cerevisiae, S. kudriavzevii and their interspecific
hybrid" (PDF). Int. J. Food Microbiol. 131 (2–3): 120–27.
doi:10.1016/j.ijfoodmicro.2009.01.035. PMID 19246112.
Jansma, David B. (1999). Regulation and variation of subunits of RNA
polymerase II in
Saccharomyces cerevisiae (PDF) (Ph.D.). University of
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