KarakteristikSunting

R. oligosporus mempunyai koloni abu-abu kecoklatan dengan tinggi 1 mm atau

lebih.[4]Sporangiofor tunggal atau dalam kelompok dengan dinding halus atau agak
sedikit kasar, dengan panjang lebih dari 1000 mikro meter dan diameter 10-18 mikro
meter.[4] Sporangia globosa yang pada saat masak berwarna hitam kecoklatan, dengan
diameter 100-180 mikro meter.[4] Klamidospora banyak, tunggal atau rantaian pendek,
tidak berwarna, dengan berisi granula, terbentuk
pada hifa, sporangiofor dan sporangia.[5] Bentuk klamidospora globosa, elip atau
silindris dengan ukuran 7-30 mikro meter atau 12-45 mikro meter x 7-35 mikro
meter.[6]

Kondisi pertumbuhanSunting
R. oligosporus dapat tumbuh optimum pada suhu 30-35 °C, dengan suhu minimum
12 °C, dan suhu maksimum 42 °C.[6]

ManfaatSunting
Beberapa manfaat dari R. oligosporus antara lain meliputi aktivitas enzimatiknya,
kemampuan menghasilkan antibiotik alami yang secara khusus dapat
melawan bakterigram positif [5], biosintesa vitamin-vitamin B [5], kebutuhannya akan
senyawa sumber karbon dan nitrogen [5], perkecambahan spora [6], dan penetrisi
miselia jamur tempe ke dalam jaringan biji kedelai.[6]

izopus adalah genus fungi saprofit yang umum pada tanaman dan parasit yang
terspesialisasi pada hewan. Mereka ditemukan di berbagai substrat organik, termasuk
"buah dan sayuran matang",[2] jeli, sirup, kulit, roti, kacang tanah, dan tembakau.
Beberapa spesies Rhizopus adalah agen oportunistik dari zigomikosis manusia (infeksi
jamur) dan bisa berakibat fatal. Infeksi Rhizopus juga bisa menjadi
komplikasi ketoasidosis diabetik.[3] Genus yang tersebar luas ini mencakup setidaknya
delapan spesies.[4][5]
 Rhizopus, perbesaran 400x

Spesies Rhizopus tumbuh sebagai hifaberbentuk filamen dan bercabang yang umumnya
tidak memiliki dinding silang (yaitu koenositik). Mereka berkembang biak dengan
membentuk spora aseksual dan seksual. Dalam reproduksi
aseksual, sporangiosporadiproduksi di dalam struktur berbentuk bola,
yaitu sporangium. Sporangium didukung oleh kolumela apophysate besar di atas
tangkai yang panjang, sporangiofor. Sporangiofor muncul di antara rizoid khas yang
mirip akar. Dalam reproduksi seksual, zigospora gelap diproduksi pada titik di mana
dua miselium yang kompatibel melebur. Setelah berkecambah, zigospora menghasilkan
koloni yang secara genetis berbeda dari induk-induknya.
 R. microsporus var. oligosporus digunakan untuk membuat tempe, makanan fermentasi yang
berasal dari kedelai.
 R. oryzae digunakan dalam produksi minuman beralkohol di beberapa wilayah Asia dan Afrika.
 Rhizopus stolonifer (kapang roti hitam) menyebabkan buah membusuk pada stroberi, tomat,
dan ubi jalar dan digunakan dalam produksi komersial asam fumaratdan kortison.

Berbagai jenis, termasuk R. stolonifer, dapat menyebabkan busuk lunak pada ubi jalar
dan narsis.

Rhizopus oryzae
Rhizopus oryzae is a fungus that is a filamentous heterothallic microfungus that
occurs as a saprotroph in soil, dung, and rotting vegetation.[1] This species is very
similar to Rhizopus stolonifer, but it can be distinguished by its smaller sporangia
and air-dispersed sporangiospores. It differs from R. oligosporus and R.
microsporus by its larger collumellae and sporangiospores.[2] R. oryzaeis used
economically in the production of the enzymes, glucoamylase and lipase, in the
synthesis of organic acids, and in various fermented food applications.[1] The many
strains of R. oryzae has many capabilities of producing many enzymes like
carbohydrate digesting enzymes and polymers along with a number of organic
acids, ethanol and esters giving it useful properties within the food industries, bio-
diesel production, and pharmaceutical industries.[1] Although R. oryzae has many
commercial uses, it is also an opportunistic pathogen of humans
causing mucormycosis.
Rhizopus oryzae

Scientific classification

Kingdom: Fungi

Division: Zygomycota

Class: Zygomycetes

Order: Mucorales

Family: Mucoraceae

Genus: Rhizopus

Species: R. oryzae

Binomial name

Rhizopus oryzae
Went & H.C. Prinsen Geerligs, (1895)

Synonyms

Rhizopus arrhizus A. Fisch.,

(1892)
Rhizopus stolonifer Vuillemin,
(1902)
Rhizopus japonicus Vuill.,
(1902)
Rhizopus nodosus Namsyl.,
 (1906)
Rhizopus nodosus Hanzawa,
(1912)
History and taxonomyEdit
R. oryzae was discovered by Frits Went and Hendrik Coenraad Prinsen Geerligs in
1895.[2]The genus Rhizopus (family Mucoraceae) was erected in 1821 by the
German mycologist, Christian Gottfried Ehrenberg[3] to accommodate Mucor
stolonifer and Rhizopus nigricans as distinct from the genus Mucor.[4]The
genus Rhizopus is characterized by having stolons, rhizoids, sporangiophores
sprouting from the points of which rhizoids were attached, globose sporangia with
columellae, striated sporangiospores.[4] In mid 1960s, researchers divided the
genus based on temperature tolerance. Numerical methods were later used in early
1970s where researchers arrived at similar conclusions. R. oryzae was relegated to
a distinct section because it grew well at 37°C but failed to grow at 45°C.[5] In the
past, strains were identified through isolating active components of the species that
were commonly found in food and alcoholic drinks in Indonesia, China, and
Japan.[5] There are approximately 30 synonyms, the most common being R.
arrhizus.[6] Scholer popularized R. oryzae because he thought R.
arrhizus represented an extreme form of R. oryzae.[5]
Growth and morphologyEdit
Rhizopus oryzae is characterized to be a fast growing fungus where growth under
optimal temperatures is fast at 1.6mm per hour (nearly 0.5 µm per second - enough
to be able to directly visualize hyphal elongation in real-time under the
microscope).[2] R. oryzae can grow in temperature of 7°C to 44°C and the optimum
growth temperature is 37 °C.[2][7]There is very poor growth from 10 °C to 15
°C[4] and no growth is observed at 45°C.[3][5] In a NaCl solution, there is good
growth at a 1% NaCl concentration and there is very poor growth of the mycelia in
media containing 3% NaCl. There is also no growth seen in a 5% NaCl solution. R.
oryzae favors acidic media where good growth is observed at a pH of 6.8 and in the
range of 7.7-8.1 there is very poor growth.[4] Most amino acids, with the exception
of L-valine, promote R. oryzaegrowth with L-tryptophan and L-tyrosine being the
most effective. It also grows well on mineral nitrogen sources, except nitrate, and
can utilize urea.[8]
Rhizopus oryzae has variable sporangiosphores. They can be straight or curved,
swollen or branched, and the walls can be smooth or slightly rough. The colour of
sporangiosphores range from pale brown to brown. Sporangiosphores grow
between 210-2500 μm in length and 5-18 μm in diameter. The sporangia in R.
oryzae are globose or subglobose, wall spinous and black when mature, 60-180 μm
in diameter. They can be distinguishable from Rhizopus stolonifer as they have
smaller sporangia and spores.[2]The optimal conditions for sporangium production
are temperatures between 30 °C to 35 °C and low water levels.[8] Sporulation is
stimulated by amino acids (except L-valine) when grown in light, while in darkness
only L-tryptophan and L-methionine effect stimulation of growth. The columellae is
globose, subglobose, or oval in shape. The wall is usually smooth and the colour is
pale brown. The average diameter growth ranges from 30-110 μm.
Sporangiospores are elliptical, globose, or polygonal, they are striated and grow 5-8
μm in length. Dormant and germinated sporangiospores show deep furrows and
prominent ridges with a pattern that makes it distinguishable from that of R.
stolonifer. The germination of sporangiospores can be induced by the combined
action of L-proline and phosphate ions. L-ornithine, L-arginine, D-glucose and D-
mannose are also effective. Optimal germination occurs on media containing D-
glucose and mineral salts.[8]R. oryzae has abundant, root-
shaped rhizoids.[4] Zygosporesare produced by diploid cells when sexual
reproduction occurs under nutrient poor conditions. They have colors that range
from red to brown, they are spherical or laterally flattened, and ranges from 60-
140μm in size.[3] In high nutrient levels, R. oryzaereproduces asexually,
producing azygospores.[9] The stolons found in R. oryzaeare smooth or slightly
rough, almost colorless or pale brown, 5-18 μm in diameter.
The chlamydospores are abundant, globose ranging in 10-24 μm in diameter,
elliptical, and cylindrical. Colonies of R. oryzae are white initially, becoming
brownish with age[7] and can grow to about 1 cm thick.[3]
Habitat and ecologyEdit
R. oryzae can be found in various soils across the world. For example, it has been
found in India, Pakistan, New Guinea, Taiwan, Central America, Peru, Argentina,
Namibia, South Africa, Iraq, Somalia, Egypt, Libya, Tunisia, Israel, Turkey, Spain,
Italy, Hungary, Czechoslovakia, Germany, Ukraine, British Isles, and USA. The soils
where R. oryzae has been isolated are varied ranging from grassland, cultivated
soils under lupin, corn, wheat, groundnuts, other legumes, sugar canes, rice, citrus
plantations, steppe type vegetation, alkaline soils, salt-marshes, farm manure soils,
to sewage filled soils. The pH of the soils where the species has been isolated
typically range from 6.3 to 7.2.[8]
R. oryzae is isolated from foods, often identified as R. arrhizus. It is found in rotting
fruits and vegetables where it is often called R. stolonifer. Unlike the other species
such as R. stolonifer, R. oryzae is common in tropical conditions. In East Asia, it is
common in peanuts. For instance, there was 21% isolation from peanut kernels
from Indonesia.[2] It is present in maize, beans, sorghum, and cowpeas, pecans,
hazelnuts, pistachios, wheat, barley, potatoes, sapodillas, and various other tropical
foods.[2] Maize meal on which isolates of R. oryzae had been grown was found to be
toxic to ducklings and rats, causing growth depression.[5]
PathogenicityEdit
R. oryzae commonly causes a disease known as mucormycosis characterized by
growing hyphae within and surrounding blood vessels. The causal agents of
mucormycosis is the ergot alkaloid agroclavine which is toxic to humans, sheep and
cattle.[8]This infection usually occurs in immunocompromised individuals but is
rare.[10][11] Common risk factors associated with primary cutaneous mucormycosis
is ketoacidosis, netropenia, acute lymphobloastic leukemia, lymphomas, systemic
steroids, chemotherapy, and dialysis. Treatment includes amphotericin B,
posaconazole, itraconazole, and fluconazole.[12] The majority of the cases of
infection are rhinocerebral infections. At the same time, it has been found in
literature that R. oryzae can produce antibiotic activity on some bacteria.[8]
In Indonesia, where white cakes are commonly consumed are made from coconut
and fermented with R. oryzae, traditionally called "bongkerk" caused food
poisoning. Symptoms included hypoglycemia, severe spasms, convulsions, and
death.[13]
The pathogenicity towards plants is attributed to the presence of large number of
carbohydrate digesting enzymes.[1]
Physiology and industrial usesEdit
Rhizopus oryzae is involved in steroid transformations and it produces 4-desmethyl
steroids which has been useful in the fermentation industry. The carbon sources
does influence the ratio of polar and neutral lipids. The mycelium found in R.
oryzaecontains lipids and the highest lipid content occurs when grown on fructose.
The highest unsaturated fatty acid content is observed at 30°C and lowest at 15°C.
Proteolytic properties have been observed well under the conditions of pH 7 at
35°C. Pyridozine and thiamine prefer proteinase production. R. oryzae can
degrade aflatoxin A1 to isomeric hydroxy compounds and aflatoxin G1 to fluorescent
metabolite aflatoxin A1.[8] There are various factors that influence the production of
dextro-lactic acids, fumaric acid, and metabolism of R. oryzae. For examples, in 40°C
there is more favorable growth for glucose consumption, however this influenced
production of d-lactic acid production negatively. Glucose concentration of 15% is
needed for optimal production of d-lactic acid. Fumaric acid production was
suppressed in mediums containing more than 6 grams of NH4NO3 per liter and is
favorable to d-lactic acid production.[14]
R. oryzae is considered GRAS by the FDA and thus recognized as safe to use
industrially as it can consume a range of carbon sources.[15]During fermentation. R.
oryzae produce amylase, lipase, and protease activity to increase nutrient's ability
to use many compounds as an energy and carbon source.[16] Historically, it has
been used in fermentation, specifically to ferment soybean and create tempeh in
Malaysia and Indonesia.[17] Using the same methods to create traditional
tempeh, R. oryzae can be inoculated in other cooked legumes such as peas, beans,
and fava beans. Similarly in tempeh making, there is an initial bacterial
fermentation in legumes when they are soaked for a while before being cooked.
Fermentation incubation lasts for 48 hours at 33°C. After incubation, mycelium can
be observed between the legumes creating a larger, uniform product. Overall,
fruits, grains, nuts, and legumes mold-fermentation with R. oryzae produces
sensory changes in foods such as creating acidity, sweetness and bitterness. R.
oryzae can produce lactate from glucose at high levels, which is used as a food
additive and can also degrade plastics.[18] In enzyme modified cheese products,R.
oryzaeprovides microbial enzymes where milk fat and proteins are broken down to
create powder and paste forms of cheese. Specifically, it breaks down cheese curds
and acid casein.[19]
Among finding cellulases and hemicellulases, other enzymes such
as protease, urease, ribonuclease, pectate lyase, and polygalacturonase are found in
cultural mediums of R. oryzae. Besides producing a number of enzymes, it can also
produce a number of organic acids, alcohol, and esters. Cellulases in R. oryzae can
be applied to biotechnology, in food, brewery and wine, animal feed, textiles and
laundry, pulp and paper industries, and agriculture. R. oryzaecan convert both
glucose and xylose under aerobic conditions into pure L (+)-lactic acids with by-
products such as xylitol, glycerol, ethanol, carbon dioxide and fungal biomass.
Endo-xylanase is a key enzyme for xylandepolymerization and was produced by R.
oryzae fermentation from different xylan-containing agricultural by-products such
as wheat straw, wheat stems, cottons bagasse, hazelnut shells, corn cobs, and oat
sawdust. Pectinases are required for extraction and clarification of fruit juices and
wines, extraction of oils, flavors and pigmentation from plant material, preparation
of cellulose fibers for linen, jute and hemp manufacture as well as, coffee and tea
fermentations. R. oryzae can break down starch content in rice plants and therefore
shows amylolytic activities. Also, it has been reported to produce extra
cellular isoamylase which is used in food industries. Isoamylase was found
to saccharify potato starch, arrow root, tamarind kernel, tapioca, and oat. The
saccharifying ability of the enzyme is highly applicable in sugar production
industries. Proteases, which can be found in R. oryzaeare highly useful in
commercial industries. For instance, it has increased application in food,
pharmaceutical, detergent, leather, tanning industries. It is also involved in silver
recovery and peptide synthesis. One strain of R. oryzaewas found to secrete
alkaline serine protease which shows high pH stability within 3 to 6 and poor
thermos-stability. Lipase that is extracted from R. oryzae have been consumed as
digestive aids without adverse reactions. Lipases hydrolyze fats and oils with
subsequent release of free fatty acids such as diacylglycerols, monoacylglycerols
and glycerol. Lipases have been involved in biotechnology applications because of
its ability to catalyze synthetic reactions in non-aqueous solutions. One study has
reported the expression of a fungal 11 alpha-steroid hydroxylase from R.
oryzae which can be used to perform the 11 alpha-hydroxylation of the steroid
skeleton which has simplified steroid drug production.[20]R. oryzae can produce
intracellular ribonuclease in a metal ion-regulated liquid medium with the addition
of calcium and molybdenum stimulating ribonuclease production. R. oryzae strain
ENHE isolated from contaminated soil was found to be capable of tolerating and
removing pentachlorophenol. R. oryzae is known to produce L (+)-lactic acid
because the fungus cells possess better resistance to high concentration of
accumulated lactic acid and lower content of nutrient requirement compared to the
commonly used bacterial procedures. Thus, R. oryzae is the most efficient
approached to improve lactic acid production process that facilitates multiple
reuses of fungal cells for long-term lactic acid production. Ethanol is the main by-
product in the fermentation process of R. oryzae during the production of L-lactic
acid. R. oryzae can be used as a biocatalyst for ester production in organic solvent.
Dry mycelium of four R. oryzae strains proved effective for catalysing the synthesis
of different flavor esters. For example, the pineapple flavour or butyl acetate esters
was produced by the esterification reactions between acetic acid and butanol by R.
oryzae. This flavor compound can be used in food, cosmetic and pharmaceutical
industries. Within the biodiesel industry, biodiesel fuel as fatty acid methyl ester is
produced by the esterification of plant oil or animal fat with methanol. This is a
renewable fuel resource compared to the traditional petroleum-based fuels.
Production of biodiesel fuel from plant oils from cells of R. oryzae immobilized
within biomass support particles were investigated for the methanolysis of
soybean oil. Olive oil or oleic acid was found to be effective for enhancing
methanolysis activity which is a promising results within the biodiesel industry
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Saccharomyces cerevisiae
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 μmin diameter. It reproduces by a division process known as budding.[1]
Saccharomyces cerevisiae

S. cerevisiae, electron micrograph

Scientific classification

Kingdom: Fungi

Division: Ascomycota
Class: Saccharomycetes

Order: Saccharomycetales

Family: Saccharomycetaceae
Genus: Saccharomyces
 Species: S. cerevisiae

Binomial name

Saccharomyces cerevisiae
Meyen ex E.C. Hansen
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. Antibodiesagainst S. cerevisiae are found in 60–70% of
patients with Crohn's disease and 10–15% of patients with ulcerative colitis (and
8% of healthy controls).[2]
EtymologyEdit
"Saccharomyces" derives from Latinized Greekand 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[3]
 Ale yeast
 Top-fermenting yeast
 Baker's yeast[3]
 Ragi yeast, in connection to making tapai
 Budding yeast

This species is also the main source of nutritional yeast and yeast extract.
HistoryEdit
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,[4] 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.[5] 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[6]), 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,[7] 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 applications.
BiologyEdit

Yeast colonies on an agar plate.

EcologyEdit
In nature, yeast cells are found primarily on ripe fruits such as grapes (before
maturation, grapes are almost free of yeasts).[8] 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.[9] 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 dominulafavors the mating of S.
cerevisiae strains, both among themselves and with S. paradoxuscells by providing
environmental conditions prompting cell sporulation and spores germination.[10]
The optimum temperature for growth of S. cerevisiae is 30–35 °C (86–95 °F).[9]
Life cycleEdit
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.[11] 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.[12][13] However, growth rates vary enormously both between strains
and between environments.[14] Mean replicative lifespan is about 26 cell
divisions.[15][16]
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".[17][18]
Nutritional requirementsEdit
See also: Yeast assimilable nitrogen
All strains of S. cerevisiae can grow aerobically on glucose, maltose,
and trehaloseand 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.
MatingEdit

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.[19] As in many other eukaryotes, mating leads to genetic
recombination, i.e. production of novel combinations of chromosomes.
Two haploidyeast cells of opposite mating type can mate to form diploid cells that
can either sporulateto 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 and desensitization.
Cell cycleEdit
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.
CytokinesisEdit
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.[20]
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.[21]
TimingEdit
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.[20]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.[20]
When the daughter emerges, the daughter is two-thirds the size of the
mother.[22]Throughout the process, the mother displays little to no change in
size.[23] The RAM pathway is activated in the daughter cell immediately after
cytokinesis is complete. This pathway makes sure that the daughter has separated
properly.[22]
Actomyosin ring and primary septum formationEdit
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.[22] 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.[24][23]
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.[20] 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.[20] 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 site.
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.[25] After AMR constriction is complete, two secondary septums are formed by
glucans. How the AMR ring dissembles remains poorly unknown.[21]
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.[26] Thus, the AMR and septum formation are the major drivers of
cytokinesis.
Differences from fission yeastEdit
 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 wall [20]
 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 [27]
 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 [27]

In biological researchEdit
Model organismEdit

S. cerevisiae, differential interference contrast image

Saccharomyces cerevisiae
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 damageand repair mechanisms.[28]
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[29] 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 agingEdit

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.[30] 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 are Replicative 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.[30] Limiting the amount of glucose or amino acids in
the growth mediumhas been shown to increase RLS and CLS in yeast as well as
other organisms.[31] 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.[31] The effects of dietary restriction
may be the result of a decreased signaling in the TOR cellular pathway.[30] This
pathway modulates the cell's response to nutrients, and mutations that decrease
TOR activity were found to increase CLS and RLS.[30][31] This has also been shown
to be the case in other animals.[30][31] 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.[32][33]
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.[34] 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 repairEdit
S. cerevisiae reproduces by mitosis as diploid cells when nutrients are abundant.
However, when starved, these cells undergo meiosis to form haploid spores.[35]
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.[36][37] For instance, gene rad52 is required for both meiotic
recombination[38] and mitotic recombination.[39] 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.[37][38][40] 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.[36] (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.[36] 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,[41][42][43] since this benefit is realized
during each meiosis, whether or not out-crossing occurs.
Genome sequencingEdit
S. cerevisiae was the first eukaryotic genometo be completely sequenced.[44] The
genome sequence was released to the public domainon April 24, 1996. Since then,
regular updates have been maintained at the Saccharomyces 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 pairsand 6,275 genes, compactly organized on
16 chromosomes.[44] 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.[45] 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.[citation needed]

Systematic gene names for Baker's yeast

Example
YGL118W
gene name

Y the Y to show this is a yeast gene

chromosome on which the gene is

G
located

L left or right arm of the chromosome

sequence number of the gene/ORF

118 on this arm, starting at the
centromere

whether the coding sequence is on

W
the Watson or Crick strand

 Examples
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 interactionsEdit
The availability of the S. cerevisiae genome sequence and a set of deletion mutants
covering 90% of the yeast genome[46] 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".[47] 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.[47] 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.[48]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 with.[47]
Other tools in yeast researchEdit
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 needed]
Synthetic yeast genome projectEdit
The international Synthetic Yeast 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, 6 of the 16 chromosomes
have been synthesized and tested. No significant fitness defects have been
found.[49]
AstrobiologyEdit
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.[50][51] 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.[50][51][52] 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 applicationsEdit
Further information: Yeast in winemaking
BrewingEdit
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 lager yeast 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. A
variant yeast known as Saccharomyces cerevisiae var. diastaticus is a beer spoiler
which can cause secondary fermentations in packaged products.[53]
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.[54]
BakingEdit
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 different.
Uses in aquariaEdit
Owing to the high cost of commercial CO2cylinder systems, CO2 injection by yeast is
one of the most popular DIY approaches followed by aquaculturists for providing
CO2to 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