laboratory strains just like the ones we are seeing

play an essential role into biological research

p<when laced under a proper growth medium yeast cells divide by budding

a protuberance appears on the surface of the cells and growths in to a spherical bud

the bud neck is pinch off

both the old and the new cells continue budding

in real life the cell cycle is not a matter of seconds it takes about two hours

a bud starts its separate life in stage g1

each cell has a nucleus invisible without special stains

dna synthesis begins at the time of budding initiation or a little before it

the s period of dna synthesis extends over one third to one half of the cell cycle

the nucleus stretches out through the neck in to the bud

then divides into equal nuclei one on each side of the neck

the cell cycle is completed with the formation of a wall between mother and daughter cells

in mother and daughter cells and all their descendants are genetically identical and constitute a
clone

the yeast cell wall is largely made of glucan and mannan

the scar tissue form on the mother cell after budding contain another polymer chitin

the scar tissue is unable to bud again

each new bud appears on a different spot of the mother cell circus

after the mother cell has produced a score daughters it cannot bud anymore and dies

yeast is thus among the simplest organism whose cells cannot divide indefinitely but have a
limited life spam
on solid food surfaces the progeny cells accumulate on top of each other and in a couple of days
they form colonies visible to the naked eye

a single cell may thus form a colony containing many millions of cells

which except for rare mutation occurring during growth are all genetically identical

the cells in nature and in laboratory however form a vary lot

many variants are available with different metabolic characteristics

for example, a single small change in the genetic information have made these cells accumulate a
red pigment

and require adenine for growth

besides vegetative reproduction

yeast can go through a sexual process

these two yeast cells are genetically different

scientist usually work with haploid yeast strains

there are two matting types or sexes of haploid cells called a and alpha

separated from each other they can be maintained indefinitely in culture

when two haploid cells of opposite mating type growth together

they exchange chemical signals

which stops their budding,

render them sticky

and change their form

the two haploid cells then fused to form a single diploid cell the zygote

here is a mixture of haploid cells of opposite sex t

they have form clamps under the influence of sex specific chemicals
called pheromones which are small peptides

pairs of haploid cells of opposite sex fused to form diploids zygotes

with a very peculiar dumbbell look

many haploid cells do not fused

yeast have no active way to look for partners

now let’s watch the congregations of these 3 cell pairs

the pheromones stops the cell cycle at stage g1 and inhibits budding

budding resumes some two hours after matting

the result is a mixture of diploid and haploid cells

the haploid cells tend to be smaller and rounder than the diploid cells

just before mating the congregating cells change their form from robust to para---

under the influence of the pheromones

the fusion of the cells is immediately followed by the fusion of their nuclei to form a diploid
nucleus

the first bud usually appears in the middle of the zygote which then looks like a clover leaf

the zygote and its daughter continue to bud

a clone of identical diploid cells is form

the cell cycle of the diploid is very similar to that of the haploids

diploid may thus be kept indefinitely in culture

this diploid is on a starvation diet

then it under goes myoisis and form 4 spores

they are haploid

the genes of the diploid are distributed to the 4 spores

for each character 2 of the four spores will resemble each of the haploid ancestors of the diploid

thus 2 of the spores are a and 2 alpha

these diploid cells have been feed an acetate and starved for nitrogen

they look a little different

have accumulated lipid droplets and have stop budding

they made a round of dna synthesis and enter myosis

in myotic prophase active genetic recombination takes place

the diploid cells convert to an ascus containing 4 haploid cells called ascospores

more asci are now being form

some cells do not sporulate and remain diploid

with sporulation each of the four nuclei produce by myosis

ends up in a separate ascospores and the 4 ascospores in an ascus

the cell walls of the ascus and the ascospores

are chemical different

it is thus possibly to dissolve the ascus cell wall without damaging the ascospores

the ascospores may then be separated from each other

when the ascuspores are taking apart and allowed to grow separately

each forms a haploid clone

the ascus thus gives rise to 4 clones

with the help of a micromanipulator the ascospores are distributed at will on a solid medium

and allow to form separate colonies

tetrad analysis is the genetic study of the 4 clones produce by each ascus

it is a powerful tool of yeast genetics

which finds no equivalent in the higher plants and animals

the 2 2 distribution is characteristic of nuclear genes inherted in zygote diploids

here is the gemanation of an undisrupted ascus whose wall is breaking up

continguous cells of opposite cells fused to form new diploid zygotes

the result is a complex mixture of diploid and haploid cells of different genetic constitutions

in these other case the 2 pairs of brothers and sisters immediately react to each other and form 2
diploid zygotes

the resulting population will all be diploid but not necessarily genetically uniform

so far we have been looking at hetero yeast the most useful in laboratories

but no all yeast are heterothallic

in fact, most of the wine yeast and other natural strains are homothallic

haploid homothallic strains easily change sex

cells that have divided may change their sex in the second generation

mother and daughter

oh no

daughter and father may fuse to form a diploid zygote

sex switching is characteristic of homothalism

here are 4 homothallic ascospores

mother and daughter do not react to each other

it is only in the second generation that sex switching lead to cell fusion

here are 2 other homothallic ascospores

first budding, second budding, cell fusion, ... upper left

the diploids continue to grow

if the homothallic spores germanate in the ascus

a brother and a sister can fuse immediately without sex switching

and form a zygote

the progeny of the other ascospore after 2 budding cycles change sex and form another diploid

no sex switching is observed in these homothallic ascus because the 2 pairs of ascospores

immediately form 2 zygotes

these double congregation was also seen in the previous intact heterothallic ascus

the resulting population are only diploid but may not be genetically uniform

the life of homothallic yeast is similar to that of higher animal and plants

in that they spent most of their time in the diploid stage

with only brief excursion in to the haploid stage to increase genetic variability

the heterothallic yeast has a possibility of remaining indefinitely haploid

these facilitates many forms of genetic manipulation

and that is why heterothallic yeast are preferred by scientists