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Yeast : Cell Cycle and

Reproduction
Augustine Natasha
PPDS SEMESTER 2
Introduction
• Phylum : Ascomycota -> subphylum : Saccharomycotina ->
class : Saccharomycetes or “true yeasts” -> order : the
Saccharomycetales.
• Pathogenic yeasts are found within three clades :
– Clade 1. Candida albicans, C. dubliniensis, C. tropicalis, C. viswanathii, C.
parapsilosis, C. orthopsilosis, C. metapsilosis.
– Clade 2. Candida guilliermondii, and the teleomorph Pichia guilliermondii,
Clavispora (Candida) lusitaniae, Candida zeylanoides, Pichia (Candida)
norvegensis.
– Clade 3. Candida glabrata and Issatchenkia orientalis (anamorph: Candida
krusei). Saccharomyces cerevisiae is also in Clade 3
Introduction

• Source : boyce et al.


Yeast : organelles

Source : Uchida M, et al. 2010


Yeast : Nutrients for living
Yeast : Nutrients for living
Yeast : Nutrients for living
Yeast : Nutrients for living

Most yeast and fungal species thrive in warm,


sugary, acidic, and aerobic conditions.

most fungi are acidophilic and grow well


between pH 4 and 6, but many species are able
to grow, albeit to a lesser extent, in more acidic
or alkaline conditions
Fungi life-cycle
Yeast : Budding and Mating

Source : Slaughter et al. 2009.


Yeast Ploidy Changes : Diploid, haploid, polyploid

• Eukaryotes alternate the diploid (two chromosome sets)


and haploid (one set) phases during their life cycles.

Yeasts, live naturally in both haploidy and diploidy

• the number of chromosomes in one set appears to be markedly stable


under optimal conditions

Nonetheless, polyploid yeasts are common:

• autopolyploidy (hereafter called polyploidy) arises by the duplication of


basically identical genomes, either from the same individual or from
the same species;
• allopolyploid cells arise by the fusion of two or more cells of closely
related but non-identical species.
• Yeasts proliferate in either haploid or diploid cycles. The preference for diploidy or
haploidy depends on the species and the environmental context.

Source : Storchova Z. 2014


Diploid vs haploid yeast
Diploid Haploid
5 x 6 µm ellipsoids 4 µm diameter
spheroids
cultures tend to cultures tend to
have lower have higher
numbers of cells numbers of cells
per cluster (during per cluster (during
exponential exponential
growth) growth)

Axial Budding Radial Budding

Source : Uchida M, et al. 2010


Yeast cell cycle
• Vegetative cell division of A daughter is initiated
yeast characteristically occurs as an out growth
by budding from the mother cell
• Under optimized growth
conditions, budding yeasts,
typified by S. cerevisiae, can nuclear division
complete their budding cell
division cycle in around 2
hours
• Each mother cell usually forms cell-wall formation
no more than 20-30 buds, and
its age can be determined by
the number of bud scars left
on the cell wall (calcufluor
staining) cell separation.
Yeast cell cycle
Yeast cell cycle
Yeast cell cycle
CELL CYCLE: YEAST VS
PSEUDOHYPHAE
Cell Cycle : Yeast
•Yeast cells grow by Selection of bud sites in C.
asymmetric budding, albicans yeast
forming smooth, round cells is temperature-
colonies dependent.

Cultures generally contain


Septin rings appear before
a mixture of cells with
bud emergence, and nuclei
more cells exhibiting an
divide across the
axial pattern at lower
motherbud neck.
temperatures.
Cell Cycle : Yeast
At START, the transition from
G1 to S phase of the cell cycle,
bud emergence is coordinated
with the onset of DNA
replication and spindle pole
body duplication.

Yeast cells separate after


cytokinesis, when daughter
cells have not yet reached the
size of their mother cells.

Daughters enter the next cell


cycle slightly later than their
mothers, consistent with the
idea that a cell size threshold
affects the timing of START

Source : Futcher B. 1996


Models for cell cycle progression in
yeast

• Source : berman J, 2006


Yeast budding pattern

Source : Slaughter et al. 2009.


Cell Cycle : Pseudohyphae
The cells remain
attached after
cytokinesis,
forming branched
C. albicans pseudohyphal cells bud in a unipolar pattern. chains of elongated
buds and colonies
that are fibrous or
rough.

As with S. cerevisiae
These daughters and
pseudohyphae, C.
mothers also reach
As in yeast cells, albicans Filaments invade the
START when they are
septin rings form pseudohyphal cells agar below the colony
a similar size and thus
before bud spend more time and extend across the
enter the next cell
emergence, and nuclei growing in a polarized agar from the colony
cycle with more
divide across the neck. manner and remain in edge.
synchrony than do
G2 longer than do
yeast cells
yeast cells.
Models for cell cycle progression in
pseudohyphae

• Source : berman J, 2006


Cell Cycle : Hyphae
• Hyphal growth has properties distinct from those of pseudohyphae,
and C. albicans hyphae resemble the hyphae of filamentous fungi.
• Hyphae are narrower than pseudohyphal cells ("2 mm) and have
parallel walls with no obvious constriction at the site of septation
• The crucial transition point occurred when buds reached a size at
which they normally switch from polarized growth to isotropic
growth, suggesting that buds that have switched to isotropic
growth can no longer form hyphae.
• In filamentous fungi, the Spitzenkӧrper, or ‘tip body’, is a structure
just behind the hyphal tip, that mediates growth directionality and
hyphal tip morphogenesis.
• Continuous polarized tip grow this associated with the presence of
the Spitzenkӧrper, whereas cell cycle dependent polarized grow this
associated with the presence of the polarisome.
Models for cell cycle progression in
hyphae
Yeast, pseudohyphae, and true hyphae

• Source : berman J, 2006


YEAST REPRODUCTION
Sex in Yeast

inbreeding/selfing (homothallism)
versus outbreeding (heterothallism)
modes of reproduction;

There are three central


dichotomies : the sex determinants encoded by the
mating-type (MAT) locus;

sex systems with a single biallelic locus


(bipolar) versus systems with two
unlinked, multiallelic sex loci
(tetrapolar)
Sex in Yeast : Pheromone
• An early key step in sexual reproduction is mate recognition

α cells produce
the α factor
peptide
pheromone

signal α cells Signal a cells,

a cells produce a
factor, a lipid-
modified peptide
pheromone
Sex in Yeast
• The S. cerevisiae paradigm serves as a basic model to
understand the process of mating when there is no structural
difference between the gametes of each mating type (termed
isogamy).
• This system is common in the closely related Candida genus
and in the rather distantly related archiascomycete
Schizosaccharomyces pombe, where both haploid and diploid
states of the sexual cycle are characterized by yeast cells
Homothallism VERSUS Heterothallism


Heterothallic fungi

• require two partners of opposite mating types with compatible MAT idiomorphs, which
contain genes controlling cell identity, cell fusion, and the formation of the dikaryotic
zygote state that leads to nuclear fusion, meiosis, and sporulation.

Homothallic fungi

• self fertile with a single individual capable of sexual reproduction even in solo culture

Both modes of sexual reproduction share key features (e.g., ploidy


changes, meiosis, production of recombinant progeny) but differ in
other key features involving aspects of cell or hyphal fusion.
Homothallism VERSUS Heterothallism
Homothallism VERSUS Heterothallism
Homothallism VERSUS Heterothallism
• Some heterothallic fungi exhibit homothallism under specific
environmental conditions.
• The heterothallic basidiomycete C. neoformans can undergo a
transition from yeast to hyphae and complete opposite-sex mating
or, in the absence of a compatible mating partner, undergo same-
sex mating.
• The heterothallic ascomycete C. albicans, in the absence of the Bar1
protease or the presence of α pheromone produced by α cells in
mèenage trois à matings, can also undergo autocrine or paracrine
pheromone signaling and
same-sex mating
MATING-TYPE SWITCHING
• Sexual identity in fungi is controlled by the MAT locus, which
encodes key regulators of mating.
• In haploid cells, MAT is defined by two alleles: MATα and MATa.
• Two cells of opposite mating type are able to fuse and mate in
response to pheromone and environmental cues.
• Some fungal species, specifically the ascomycetes S. cerevisiae, S.
pombe, and K. lactis, exhibit the unusual property of being able to
undergo mating-type switching, a process in which a haploid cell
gives rise to a cell of opposite mating type (a→α or α→a).
• These systems involve
– a cassette mechanism with one active expression locus,
– two silent MAT allele copies, and
– machinery for DNA lesion-promoted recombination
sex systems : bipolar vs tetrapolar

In basidiomycetes, there exist both


bipolar and tetrapolar mating systems

Bipolar mating tetrapolar mating


system : systems :
Mating type is determined by one Mating type is determined by two
locus, and two strains must possess loci, and two strains must possess
different alleles at the locus to be different alleles at both loci to be
sexually compatible sexually compatible.
Sources
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morphogenetic adaptation allowing colonization of a host. FEMS microbiology reviews. 2015 Aug 7;39(6):797-
811.
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analysis of yeast internal architecture using soft X‐ray tomography. Yeast. 2011 Mar;28(3):227-36.
6. Storchova Z. Ploidy changes and genome stability in yeast. Yeast. 2014 Nov;31(11):421-30.
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8. Berman J. Morphogenesis and cell cycle progression in Candida albicans. Current opinion in microbiology. 2006
Dec 1;9(6):595-601.
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10. Kashem SW, Igyártó BZ, Gerami-Nejad M, Kumamoto Y, Mohammed J, Jarrett E, Drummond RA, Zurawski SM,
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