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 Factors Affecting Yeast 
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
 Flocculation
Fermentation 2018, 4(2), 28; https://doi.org/10.3390/fermentation4020028 Open Access
 Flocculation Gene Structure
(https://doi.org/10.3390/fermentation4020028)

 The Genetic Instability of
Review
Strains †
Yeast

Flocculation—Sedimentation
The Influence of Cell Surface
and Flotation 0
bySurface
Graham G. Stewart (https://sciprofiles.com/profile/214252)  (mailto:please_login)
Charge on Yeast
Flocculation
The International Centre for Brewing and Distilling, Heriot-Watt University, Riccarton, Edinburgh
 Premature Yeast Flocculation
EH144AS,

UK
Phenotypic Effects on Flocculation
† Adhesion andinformation
Biofilm Formation
Some of the included in this review paper was obtained (with the permission of the
 Centrifugesfrom:
to Crop Yeast G.G. Harvesting and cropping yeast: Flocculation and centrifugation.
publisher) Stewart,
 Conclusions
Brewing and Distilling Yeasts; Springer International Publishing, Cham, Switzerland, 2017; pp.
 Acknowledgments
259–308.
 References
Received: 8 March 2018 / Accepted: 10 April 2018 / Published: 16 April 2018
Abstract: Unlike most fermentation alcohol beverage production processes, brewers recycle their
yeast. This is achieved by employing a yeast culture’s: flocculation, adhesion, sedimentation,
flotation, and cropping characteristics. As a consequence of yeast recycling, the quality of the
cropped yeast culture’s characteristics is critical. However, the other major function of brewer’s yeast
is to metabolise wort into ethanol, carbon dioxide, glycerol, and other fermentation products, many of
which contribute to beer’s overall flavour characteristics. This review will only focus on brewer’s
yeast flocculation characteristics.
Keywords: centrifugation; cropping; flocculation; flotation; recycling; sedimentation
1. Introduction
Back to TopTop

 The objectives of brewer’s wort fermentations are to consistently metabolise wort constituents
Abstract (/)
into ethanol, carbon
Introduction dioxide, glycerol, and other fermentation products, many of which contribute to

beer flavour
Yeast in order to produce it with satisfactory quality, drinkability, and stability. Another important
Flocculation
fermentation
Measurement objective
of YeastisFlocculation
to produce yeast crops that can be harvested, stored, and subsequently re-
pitched intoCharacteristics
Cell Wall a later brew [1]. During brewing, overall yeast performance is controlled by a plethora of
factors that
Sexual include (in no order of priority):
Aggregation
 TheCell
Yeast yeast
Wallstrains employed
Structure and and their condition during pitching and throughout the wort
Flocculation
fermentation cycle [2];
 Factors Affecting Yeastand category of assimilable nitrogen [3];
the concentration
Flocculation
the spectrum of wort sugars [4];


Co-Flocculation
fermentation temperature [2];
yeast pitching (inoculation) rate [1];

 Flocculation
concentration of inorganic ions [5]; 
yeast tolerance to stress factors such as: heat, osmotic pressure, temperature, ethanol,

mechanical tension, desiccation, etc. [6];
wort gravity [7];
Strains
 wort dissolved oxygen concentration at pitching and during fermentation [8];
The Influence of Cell Surface 0
flocculation,(CSH)
Hydrophobicity adhesion, sedimentation, and cropping characteristics [9];
and Cell
culture
Surface characteristics
Charge on Yeast during storage between fermentations and its subsequent re-pitching into
wort [10].
Flocculation
 Many of the
Phenotypic above
Effects factors will be discussed in other parts of this monograph. They influence
on Flocculation
yeast performance
Adhesion either
and Biofilm individually
Formation or in combination with other factors. Together, they permit
definition of the
Centrifuges requirements
to Crop Yeast of an acceptable brewer’s yeast strain [10]: “in order to achieve beer of
high quality, it is axiomatic that not only must the yeast be effective in removing the required nutrients
Conclusions
from the growth/fermentation medium (the malt extract—the wort), able to tolerate the prevailing
Acknowledgments
environmental conditions (for example, ethanol toxicity) and impart the desired flavour and stability to
Conflicts of Interest
theReferences
beer, but the microorganisms themselves at the end of fermentation must be effectively removed
from the wort by flocculation (Figure 1), sedimentation, centrifugation (Figure 2), and/or filtration after
they have fulfilled their metabolic role.”
Back to TopTop

 
Abstract (/)
Introduction

 Cell Wall Characteristics
Flocculation
Flocculation


Co-Flocculation


Strains
 Figure 1. Static fermentation flocculation characteristics [1].
Flocculation
 Adhesion and Biofilm Formation
 Conclusions
 Acknowledgments
 References

Annotate
Back to TopTop
 Figure 2. A disc stack centrifuge (5 hL per hour) [2].
 Abstract (/)
 Introduction

 It is also worthy of note that brewing is the only major alcoholic beverage process that recycles its
Yeast Flocculation
yeast [2]. It is, therefore, important to jealously protect the quality of the cropped yeast because it will
Measurement of Yeast Flocculation
beCell
used to pitch subsequent wort fermentations and will therefore have a profound effect on the
Wall Characteristics
quality of the beer resulting from subsequent wort fermentations.
Sexual Aggregation
ThisCell
 Yeast review initially examines brewer’s yeast flocculation, including both nature and nurture
Wall Structure and
aspects [11]. It subsequently analyses other aspects of yeast’s sedimentation and cropping
Flocculation
characteristics
Factors Affectingduring
Yeastbrewing. In addition, yeast flocculation and adhesion, as they apply to other
industrial processes [3,12] and medical aspects [13], are briefly discussed.
Flocculation
The flocculation properties of brewer’s yeast strains are of considerable importance because the
 Co-Flocculation
number
GeneticofControl
yeast cells

suspended in wort during both primary and/or secondary fermentations affects
of Yeast

wort fermentation velocity, beer flavour, maturation, cell crop formation, and filtration [14]. A number of
hypotheses
Flocculation have
Gene been proposed to explain the mechanisms of flocculation in Saccharomyces
Structure

cerevisiae
The Genetic[9]. Instability
The prevalent
of hypothesis is the lectin-like theory of flocculation [15]. According to this
model, a specific protein (lectin) present only in flocculent yeast cells is secreted into the outer
Strains of the cell wall and the N-terminal part of this protein binds two mannose residues present
extremities
 The Influence of Cell Surface 0
in the cell walls of neighbouring flocculent and non-flocculent yeast cells; details of this follow [16]
(Figure 3). In this process, calcium ions are necessary for the activation of the lectins [17].
Flocculation
 Conclusions
 Acknowledgments
 References
Figure 3. Lectin theory of flocculation. Protein lectins on the yeast cell surface interact with
either mannose-containing and/or glucose-containing carbohydrate determinants on the cell
walls of adjacent cells only in the presence of calcium [15].
2. Yeast Flocculation
Flocculation is a complex phenomenon that is affected by genetic [16], physiological [9], and
environmental factors—the nature-nurture effects [11]. Within the two brewer’s yeast species
(Saccharomyces cerevisiae (ale) and Saccharomyces pastorianus (lager)) [17,18] there are both
flocculent and non-flocculent strains (Figure 1). These characteristics are illustrated by the
concentration of yeast in suspension during the course of a static wort fermentation [1] (Figure 1) and
overall wort sugar uptake properties [19].
Back to TopTop

 The reasons for, and advantages of, yeast flocculation (particularly, but not only, during the
Abstract (/)
brewing process)
Introduction have been extensively studied and debated. Flocculation may enhance the survival

of Yeast
yeast Flocculation
cells during adverse (for example, starvation) conditions [20]. Flocculation is an important
characteristic
Measurement inofanYeast
environment with limited nutrients because the death and autolysis of the cells
Flocculation
inside
Cell flocs can provide further nutrients to cells in the surrounding environment [21]. It has been
shown
Sexualthat some highly flocculent cultures lost their flocculation characteristic during the early stages
Aggregation
of Yeast
growth, inWall
Cell the Structure
presenceandof nutrients, and recovered it towards the end of the exponential growth
phase coinciding with nutrient depletion [22].
Flocculation
Individual
 Factors strains
Affecting of brewer’s yeast (both ale and lager) differ considerably in their flocculating
Yeast
characteristics.
Flocculation At one extreme, there are highly non-flocculent strains, often referred to as powdery
strains [23]. At the other extreme, there are very flocculent strains (Figure 4A). The latter strains tend

Co-Flocculation
to Genetic
separate early of
Control outYeast
of suspension from fermenting wort, which can result in an under-attenuated,

 Flocculation
sweeter, and less fully fermented beer. Beers of this nature, because of the presence of fermentable 
sugars (usually maltotriose and maltose), can be liable to biological instability. In contrast, poorly

flocculent (non-flocculent or powdery) yeasts (Figure 4B) produce a dry yeast, fully fermented, more
Flocculation
biologically in Brewer’s
stable beer inYeast
which clarification is slow, leading to filtration difficulties and the possible
Strains
acquisition of yeasty off-flavours.
Flocculation
 Conclusions
 Acknowledgments
 References
Figure 4. Flocculation characteristics of Saccharomyces cerevisiae strains. (A) Flocculent

culture; (B) non-flocculent culture [10].
The disadvantages presented by the two flocculation types of brewer’s yeast strains are
particularly relevant to traditional fermentation systems where the fermentation process is dependent
upon the yeast sedimentation characteristics. Contemporary brewing technology has largely reversed
this situation, as yeast sedimentation characteristics are now applied to fermenter design. The
efficiency, economy, and speed of batch fermentations have improved with the use of cylindro-conical
fermentation vessels [24] (Figure 5), which are often (but not always) employed in tandem with
flocculent cultures; details of this are discussed later. There is no doubt that differences in the
flocculation characteristics of various yeast cultures are primarily manifestations of the culture’s cell
wall structure, as also discussed later.
Back to TopTop

 
Abstract (/)
Introduction

Flocculation
Flocculation


Co-Flocculation


Strains
Flocculation
 Centrifuges to Crop Yeast Figure 5. Cylindroconical batch fermenters [10,24].
 Conclusions

Flocculation requires the presence of cell surface protein and mannan receptors (Figure 3). If
Acknowledgments
these are not available or are masked, blocked, inhibited, or denatured, flocculation cannot occur. The
onset of yeast flocculation is an aspect of the subject where there is great commercial interest but
 References
about which relatively little is known. As previously discussed, the ideal brewing culture remains in
suspension as fermenting single cells or very small flocs until the end of fermentation when wort
sugars (sucrose, glucose, fructose, maltose and maltotriose) are depleted by the yeast and the vicinal
diketones (VDKs) (unwanted diacetyl—butterscotch flavour) are managed appropriately [25]; only
then does the culture flocculate out of suspension. What signals the onset of flocculation or relief from
its inhibition? This is still an unanswered question that is currently being studied by a number of
research laboratories worldwide [9,10,14,25,26,27].
Brewer’s yeast flocculation is an off-cost process which does not require significant energy input.
However, cooling at the end of fermentation does facilitate yeast separation and this normally requires
some energy. It is considered that flocculation increases process efficiency and reduces the energy
consumption associated with cell separation with, for example, the use of a centrifuge—details of this
follow. Indeed, yeast flocculation in brewing is an example of Lean Manufacturing [7].
3. Measurement of Yeast Flocculation
Back to TopTop

 Although the requirement for yeast flocculation during brewer’s wort fermentation is a much-
Abstract (/)
studied phenomenon,
Introduction there is dire need for some degree of flocculation test standardisation. There

areYeast
a plethora of flocculation tests and many laboratories working in this field appear to employ their
Flocculation
own favourite method.
Measurement of YeastAs a consequence, it is difficult to translate results from one laboratory to
Flocculation
another. The
Cell Wall methods used to measure flocculation can be roughly divided into three groupings, as
Characteristics
listed below.
Sexual Aggregation
3.1. Sedimentation Methods (For Example, the Helm Sedimentation Test)
Flocculation
 In thisAffecting
Factors test, theYeast
yeast culture is removed from the growth medium and the cells are washed a
number of times with deionised water containing calcium ions (usually 80 mg/mL calcium chloride) at
Flocculation
pHCo-Flocculation
4.0 and, depending on the scale of the test, the suspension is placed in a test tube (10-mL scale)
or Genetic
a measuring cylinder
Control of Yeast(100-mL scale) (Figure 6A shows results of the Helm’s Sedimentation Test


[28] for co-flocculation—details to follow).

Strains
Flocculation
 Conclusions
 Acknowledgments
Figure 6. (A) Helm’s sedimentation flocculation test—co-flocculation [28,29]; (B) ale yeast co-
flocculation—2-L cylinder wort fermentation test [29].
 References
3.2. Direct Observation of Floc Formation in the Growth/Fermentation Medium

During this method, small inocula of the yeast strain’s being studied are seeded into 20-mL screw
capped glass bottles each containing 15 mL of fermentation media. After three days of incubation
(usually at 25 °C), the flocculation characteristics of the cultures are determined by the ratio of the
flocs subsequent to the yeast sediment being brought back into suspension by shaking the bottles.
This method permits routine flocculation determinations of a large number of cultures and has been
employed extensively for segregation studies on flocculation during FLO gene mapping studies [30]—
details to follow. An alternative measurement of flocculation is to microscopically examine the flocs
and determine the percentage of cells in a number of flocs compared to un-flocculated cells.
3.3. Static Fermentation Methods

In this method, the concentration of yeast in suspension is determined during the course of static
fermentation (Figure 1). The first two methods for measuring yeast flocculence can be viewed as
artificial tests for flocculation due to the fact that they are conducted under in vitro conditions in
Back toout
relation to the brewing process. The latter method is an in vivo style test because it is carried TopTop

under conditions
Abstract more closely akin to the static fermentation conditions encountered in a typical
(/)
brewery [31] (Figure 6B).
Introduction


4. Measurement of Yeast Flocculation
Cell Wall Characteristics
 YeastAggregation
Sexual cell surface morphology differences between flocculent and non-flocculent brewer’s yeast
cultures (another
Yeast Cell yeast species)
Wall Structure and have been extensively examined [32]. Although details of yeast cell
Flocculation
wall structure will be considered later, it is worth discussing here that the wall consists of an inner
layer composed predominantly of β-glucan and chitin together with an outer fibrilar layer on flocculent
Flocculation
cells (Figure 7) [33], consisting primarily of α-mannan (highly glycosylated) associated with
 Co-Flocculation

mannoproteins [34] (Figure 8).


Strains
Flocculation
 Conclusions
 Acknowledgments

Figure 7. Schematic diagram of the architecture yeast cell wall. Components such as

mannoprotein occur as a matrix and are distributed throughout the entire wall; therefore, the
References
layered arrangement should show zones of enrichment [15].
Figure 8. Electron photomicrograph of Saccharomyces cerevisiae flocculent and non-flocculent

strains shadow-cast with tungsten oxide [33].
Back to TopTop

 In addition to cell-cell adhesion in the formation of flocs (Figure 3), some yeast cells (both
Abstract (/)
brewing and distilling
Introduction strains) possess the capacity to adhere to abiotic (non-living) surfaces, cells

and, tissues,
Yeast resulting in biofilm formation [12] (Figure 9). The cell wall functions as a means for yeast
Flocculation
cells (and other of
Measurement microorganisms) to interact with their environment. It has already been discussed that
Yeast Flocculation
one of Wall
Cell the most critical functions of some cell surfaces is their ability to adhere to other cells (floc
Characteristics
formation) and surfaces. Adhesion prevents cells from being washed away when they are present in a
Sexual Aggregation
nourishing
Yeast Cellenvironment,
Wall Structurewhile
and it also allows them to form biofilms that protect the cells from hazardous
and stressful conditions [35]. In addition to an industrial environment, pathogenic yeasts (for example,
Flocculation
Candida
Factorsalbicans) exploit their capacity to adhere to abiotic surfaces such as plastic prosthetics in
Affecting Yeast
order to gain access to the bloodstream and the internal organs of patients [36]. It is believed that
Flocculation
adhesion of C. albicans cells to mucosal surfaces involves lectin-like interaction between the protein

Co-Flocculation
portion
Geneticof mannoproteins
Control of Yeast located in the fibrils on the cell surface and glycoside receptors on epithelial

 Flocculation
cells [13] (Figure 10). 

Strains
Flocculation
 Conclusions
Figure 9. Micrograph of a biofilm of microorganisms (yeast and bacteria) on the inner surface of
 Acknowledgments
a beer dispersing line [12].
 References
Figure 10. Electron photomicrographs of adhering and non-adhering cultures of Candida

albicans (photograph courtesy of L. J. Douglas) [13].
The word floc derives from the Latin word floccus, which means a tuft of wool, while cells that are
not able to form flocs are known as non-flocculent or powdery cells [37]. S. cerevisiae can betofound
Back TopTop

aggregated
Abstract in different ways not to be confused with flocculation, such as sexual aggregation [38], co-
(/)
flocculation—addressed
Introduction later in this paper [29] (Figure 6A and Figure 11), biofilm formation [12]

(Figure
Yeast 9), and chain (pseudohyphae) formation, where daughter cells do not separate physically
Flocculation
from their mother
Measurement of cells
Yeast[39] (Figure 12).
Flocculation
Flocculation
Flocculation


Co-Flocculation


Strains
 The Influence 0
Figure 11. of Cell Surface between an ale yeast strain and a Lactobacillus fermentum strain—
Co-flocculation
Hydrophobicity (CSH)yeast
bacterial-induced and Cell
flocculation [40].
Flocculation
 Conclusions
 Acknowledgments
 References
Figure 12. Chains of S. cerevisiae—pseudohyphae [39,41].

Back to TopTop

5. Sexual Aggregation
Abstract (/)
 Introduction

 Sexual
Yeast aggregation between haploid strains of S. cerevisiae involves complementary mating
Flocculation
types (MATα and
Measurement of MATa) which can
Yeast Flocculation occur after an exchange of pheromones (an agent secreted by an
organism
Cell Wallthat produced a change in the sexual behaviour of a related organism)—a and α,
Characteristics
respectively. This induces the appearance of complementary molecules (proteins) on the surface of
Sexual Aggregation
theYeast
cellsCell
which
Wallfacilitates
Structure the
and fusion of haploid cells—mating a cells with α cells [42]. Co-flocculation
[29], mutual aggregation [43], mutual agglutination [44], and mutual flocculation [40] all describe a
Flocculation
heterotypic aggregation
Factors Affecting Yeast process (while single-strain flocculation is homotypic) between two microbial
strains which are usually (not always) yeast [45] (Figure 6A and Figure 11). When two yeast strains
Flocculation
(one being non-flocculent and the other weakly flocculent) are mixed together, in the presence of Ca2+

Co-Flocculation
ions at pHControl
Genetic 4.0–4.5, flocs form, which rapidly settle out of suspension or rise to the surface of a wort
of Yeast

 Flocculation[46] (Figure 6A). Details of co-flocculation can be found in Section 8.
fermentation 
It has already been discussed that chain formation (also called pseudohyphal growth) occurs due

to failure of a young daughter yeast cell to separate from the mother cell. This results in an aggregate,
Flocculation
usually in Brewer’s
composed of 30–50Yeast
cells [12] (Figure 12). These cell aggregates are physically linked to each
Strains
other and, consequently, following dispersion of the cells (for example, physical disruption, sonication,
0
or enzyme treatment), cells will not be able to re-aggregate prior to a subsequent growth cycle. Chain
formation in brewing yeast strains is restricted to ale yeast species (S. cerevisiae); lager yeast strains
(S.Flocculation
pastorianus) do not usually exhibit chain formation characteristics [47]. Various effects on yeast
flocculation
Prematureand aggregation
Yeast Flocculationare discussed in greater detail later.
6. Yeast Cell Wall Structure and Flocculation
 Conclusions
It has already been stated that flocculation is a cell surface characteristic [32]. Some (not all)
 Acknowledgments
heat-treated flocculent cells retain their ability to flocculate [26], as well as isolated in vitro cell walls
 Conflictsfrom
of Interest
prepared a flocculent culture [48]. The yeast cell wall carries a negative charge due to the
 References
ionisation of carboxyl and phosphodiester groups in cell wall proteins and phosphomannans,
respectively. The repulsion of charges of the same ionisation sign prevents cells from getting
sufficiently close and acts as an effective barrier to cell aggregation [49]. Also, a positive correlation
between cell surface hydrophobicity (CSH) and flocculation has been reported [50]. CSH is partially
responsible for triggering flocculation in brewing yeast strains [51]. Based on these results, an
increase in yeast surface hydrophobicity has been described when dominant flocculation genes (FLO
genes) (details to follow) are expressed in the structure of yeast cell walls [27].
It has been proposed that lectins (carbohydrate-binding proteins) are macromolecules that are
specific for sugar moieties [52]. They are considered to be present on flocculent cell surfaces. Specific
lectin-like proteins interact with carbohydrate residues for α-mannans (receptors) on neighbouring
cells (Figure 7). Also, calcium ions enable lectins to achieve their active conformation [53] (details of
calcium effects on yeast flocculation are discussed later). Although flocculation lectins are only
present on the surface of flocculent cells, the receptors are present in both flocculent and non-
flocculent cells since, as already discussed, and the outer layers of the S. cerevisiae cell wall are in
part composed of mannan (Figure 8). Analysis of the inhibitory action of sugars and the use of
mannan synthesis mutants and concanavalin A (a lectin originally extracted from the jack-bean) have
Back to TopTop

shown that
Abstract the wall specifically consists of glycoproteins with mainly α- -mannosyl and α- -glucosyl
(/)
groups
IntroductionThis supports the theory that flocculation receptors are most likely nonreducing residues
[54].

[55]. Besides
Yeast specific lectin sugar interactions, other non-specific interactions such as hydrogen bonds
Flocculation
and hydrophobicofinteractions
Measurement can occur.
Yeast Flocculation

7. Sexual
FactorsAggregation
Affecting Yeast Flocculation
Flocculation
Several factors can affect yeast flocculation [56]. The first factor is the genetic background of a
strain. Flocculin proteins are encoded by members of the FLO group of genes [23,56]. The genetic
Flocculation(nature effects) with regard to FLO genes varies greatly amongst various types of
background


Co-Flocculation
brewer’s yeast (ale or lager) and other strains. Strains contain a variety of FLO gene combinations
[57], resulting in a spectrum of flocculation characteristics [58]. The current status with respect to S.

 Flocculation
cerevisiae (and S. pastorianus) FLO genes are discussed in Section 9. 
Second, flocculation is affected by the physiological environment (nurture effects) [59]. This

includes the pH, availability of appropriate metal ions, and nutrients during the growth phase [60]. The
pHStrains
will influence the cell surface charge which will have an effect on the flocculation phenotype.
Changes in pH may also modify the ionisation of functional groups in flocculin proteins, which will
modify their conformation
Hydrophobicity (CSH) and[21].
Cell The environment is sensed by cells leading to the expression of FLO,
their translation
Surface ChargetoonFlo proteins, and their location into the cell wall, which is influenced by a number of
Yeast
environmental
Flocculation factors [27].
 Premature
Thirdly, the
Yeastphysical environment affects flocculation. The hydrodynamic (relevant to liquids in
Flocculation
motion) conditions
Phenotypic Effectsmust be favourable and promote collision rates between cells. However, agitation
on Flocculation
must not beand
Adhesion sufficiently violent to cause flocs to disperse. In addition, there must be sufficient cells in
Biofilm Formation
suspension
Centrifugesintoorder
Crop to result in cell collisions to form flocs [61]. Factors that increase cell walls’
Yeast
hydrophobic
Conclusionscharacter (cell surface hydrophobicity) and factors that decrease the repulsive negative
electrostatic charges on cell walls (cell surface change) result in stronger flocculation, because they
Acknowledgments
facilitate
Conflictscell-cell contact [62].
of Interest
 References
Flocculation, as such, is not absolutely necessary for brewer’s yeast cells to sediment out of wort
at the end of primary fermentation. This is because the size and density of yeast cells can overcome
the Brownian motion that maintains cells in suspension [26]. However, under these circumstances, the
sedimentation rate is slow, which is dependent on particle size. Smaller particles settle more slowly
than larger particles of similar density, because they are more retarded by friction (overall viscosity).
Older (larger) yeast cells settle faster than younger, smaller cells. Nevertheless, the sedimentation of
single cells is too slow to be of practical importance during wort fermentation, where flocculation is
necessary to achieve the sedimentation of most yeast cultures during the latter stages of a typical
fermentation cycle.
Flocculating yeast cultures contain some cells that are not entrapped in flocs which do not
necessarily have the ability to flocculate [63]. Single cells are set free from flocs and, at the same
time, new cells are entrapped. Loosening of cells occurs from flocs, because of hydrodynamic forces,
resulting in a dynamic equilibrium [64]. Dilution favours an increased proportion of free cells. An
appropriate concentration of yeast cells is required for effective flocculation and some cells often
remain in suspension even following the bulk sedimentation of yeast flocs. As with many aspects of
brewer’s yeast metabolism, yeast flocculation is primarily strain-dependent.
Back to TopTop

 During the later stages of fermentation, the following conditions are favourable for culture
Abstract (/)
sedimentation:
Introduction


TheFlocculation
Yeast carbon dioxide production rate is slow;

wort attenuation
Measurement is Flocculation
of Yeast approaching completion—most of the fermentable sugars in wort have been
 Cellremoved by the yeast culture, including glucose, fructose, sucrose, maltose, and finally

maltotriose
Sexual [4];
Aggregation

flocculation
Yeast ability is and
Cell Wall Structure high but not too high;
Flocculation
yeast concentration in suspension is maximal [61];
Flocculation
There are several factors that influence the rate of floc sedimentation out of the wort:
 Co-Flocculation


The way that yeast cells are packed into flocs;
Genetic Control of Yeast
the floc size, shape, and density;

 nurture factors that include wort properties and encompasses: concentration (gravity), viscosity,
density, and turbulence [65] (the issue of premature yeast flocculation (PYF) [66] is a separate,

but related,
Flocculation phenomenon
in Brewer’s Yeast which is addressed later);
higher gravity worts, following fermentation, results in “green” (immature) beers with a higher
Strains
 Theviscosity
Influenceand density.
of Cell Both these factors will retard yeast sedimentation and lead to increased
Surface 0
osmotic pressure
Hydrophobicity and Cell
(CSH) and ethanol prior to dilution to the fermented wort’s sales gravity and alcohol
concentration.
Flocculation

The floc Yeast
Premature size often decreases as settling proceeds because the concentration of yeast cells in the
Flocculation

suspension
Phenotypiccontinues
Effects onto decrease and smaller flocs form which will slowly settle out of the
Flocculation

suspension.
 Conclusions
8. Co-Flocculation
Acknowledgments
Co-flocculation, also known as mutual aggregation or mutual flocculation, is a heterotypic
 References
aggregation process (while flocculation is homotypic) amongst two separate yeast strains [67]. One
strain is non-flocculent and the other strain is weakly flocculent. When these strains are mixed
together in the presence of Ca2+ ions, flocs form and the culture rapidly settles out of suspension [29].
Heterotypic yeast non-sexual flocculation was first described by Eddy and Rudin [68] with a number of
ale strains. The focus of our research group on co-flocculation began with the top-cropping Labatt ale
yeast culture. In the 1960s and 1970s, consumption of pale ales was popular in Canada; in 1970 it
represented 60% of the beer consumed in Ontario and 80% in Quebec [69]. However, similar to the
situation in Britain, the ale yeast cultures employed (unlike lager strains) were (and many still are)
largely uncharacterised. Also, no Labatt employee (or Labatt pensioner) could inform us much about
the history of their ale culture. This ale culture possessed classical top-cropping properties. It also
exhibited intermittent premature flocculation characteristics, resulting in under-fermented worts
containing residual sugars (mainly maltotriose). This was a problem in Canada because the alcohol
specification was a legal regulation (Federal Health Protection Board Regulations) and the beer’s
alcohol composition was (and still is) specified on the label of the bottle, can, or keg with a ±0.2% (v/v)
alcohol specification permitted. This problem was exacerbated during high gravity brewing trials [69]
Back to TopTop
of

ales. Consequently, it was important to enumerate the number of strains in this ale culture and
Abstract (/)
characterise
Introductionthem.

 A suitable
Yeast method to examine yeast culture’s strain composition in the 1960s and 1970s was the
Flocculation
giant colony morphology
Measurement method initially
of Yeast Flocculation developed by Gilliland [70] and Richards [71]. The Richard’s
method involved
Cell Wall inoculating the yeast culture in question onto wort solid media and examining the
Characteristics
colonial
Sexualmorphology
Aggregation that developed after incubating under humid conditions for at least three weeks at
18Yeast
°C. ItCell
wasWall
found that gelatin,
Structure and as the solidifying matrix, tends to enhance the distinctive features of
theFlocculation
colonial morphology to a greater extent than agar and that wort, instead of a synthetic or defined
medium,
Factorsgave distinctive
Affecting Yeast and reproducible results (Figure 13). Also, lager yeast strains do not exhibit
distinctive colonial morphologies on wort gelatin media (Figure 13).
Flocculation


Co-Flocculation


Strains
Flocculation
 Conclusions
 Acknowledgments
 References
Back to TopTop

 Figure 13. Giant colony morphologies of brewer’s yeast strains (A,B) exhibit the co-flocculation
Abstract (/)
phenotype [16,29,71].
Introduction

 Analysis of the Canadian brewery’s top cropping ale yeast culture’s strain composition showed
that two morphologically different colony types were present (Figure 13A,B) [72]. On isolation, both
colony types proved to be stable respiratory sufficient individual yeast strains of the species S.
Sexual Aggregation
cerevisiae and they were coded LAB A/69 and LAB B/69, with the former strain being ~75% of the ale
Yeast Cell Wall Structure and
culture and the latter comprising ~25% of the culture. A production-scale fermentation trial with ale
Flocculation
strain LABAffecting
Factors A/69 was conducted in a 200-hL (20,000 L) open wood fermenter with a 12 °Plato wort at
Yeast
21Flocculation
°C. Although the fermentation was under-pitched (under-inoculated), it proceeded rapidly, and all of
theCo-Flocculation
wort’s fermentable sugars were metabolised in less than 96 h. It was then that problems began.
The expected
Genetic top-crop
Control of Yeast

failed to develop on the fermentation’s surface (which occurred in the original

two strain ale mixed culture), and most of the yeast culture remained in suspension. As the brewery in
question (at that
Flocculation Genetime) did not possess a centrifuge, it was not possible to collect the yeast for reuse in
Structure

a subsequent
The Genetic fermentation
Instability of and the fermented wort (all 200 hL of it) had to be discarded into the sewer
Flocculation in
with the cost chargedBrewer’s Yeast
to the brewery’s effluent budget.
Strains
Following the above traumatic brewing-scale trial, a laboratory-scale characterisation of both ale
yeast strains isolated from the Labatt ale cultures was conducted. When the two isolated strains were
cultured alone in wort using 2-L glass cylinders, both were non-flocculent during all phases of growth.
However, when cultured together in wort in a 1:1 ratio, the culture was flocculent in the later stages of
Flocculation
fermentation and sedimented out of suspension [72]; a top crop also formed (Figure 6B). This type of
Premature Yeast Flocculation
behaviour, where two yeast strains are non-flocculent alone but flocculent when mixed together [69],
has been termed co-flocculation. As mentioned above, co-flocculation has also been termed mutual
aggregation and mutual flocculation [44]. When stationary phase cells of the two strains were mixed
Centrifuges to Crop Yeast
together in a 1:1 ratio in a solution of calcium ions at pH 4.5 [29], flocs immediately began to appear
Conclusions
and a very flocculent culture resulted that sedimented out of suspension (Figure 6A). This is the
Acknowledgments
Helm’s Sedimentation Test [28].
Protein-denaturants (for example, urea and guanidine) and several sugars (mannose, glucose,
 References
fructose, and galactose) cause the reversible inhibition of co-flocculation in the presence of Ca2+ ions.
Also, the effect of cell treatment with proteolytic enzymes (trypsin and chymotrypsin) and chemical
modification of the cell surface protein and carbohydrate components suggests that co-flocculation
results from an interaction between surface protein cellular components of one of the strains and
surface carbohydrate components of the other strain [43].
To date, co-flocculation has only been observed with ale strains [29]. There are no reports
(including extensive studies in the Labatt research laboratories) of co-flocculation between non-
flocculent lager yeast strains. Another type of co-flocculation reaction that has been described occurs
when an ale yeast strain has the ability to aggregate and co-sediment with contaminating bacteria
such as Hafnia protea [73], Lactobacillus brevis [45], Pediococcus sp., or Lactobacillus fermentum
[40] (Figure 11). This Lactobacillus fermentum strain was isolated from a fuel alcohol molasses
fermentation in Brazil and its co-flocculation characteristics were studied in Canada.
The two-strain composition of the Labatt co-flocculent production ale culture was deemed to be
undesirable, particularly because of its tendency for premature flocculation and the consequent wort
under-attenuation that occurred. This was particularly the case with high gravity worts [7] (>14 °Plato).
This often resulted in the failure to comply with the beer’s alcohol specification (this wasBack priortotoTopTop
the

introduction
Abstract of high gravity brewing procedures on a production basis, which exacerbated the
(/)
problem). Production trials with the LAB A/69 ale strain were conducted in both regular (sales) (12
Introduction

°Plato)
Yeastand higher gravity (16 °Plato) worts. This strain proved to be capable of successfully
Flocculation
fermenting
Measurementbothofwort gravities
Yeast but, because of its non-flocculent property, centrifugation was required
Flocculation
in order to harvest
Cell Wall the culture for yeast removal, beer clarification, and yeast collection for reuse. This
Characteristics
strain hasAggregation
Sexual been employed for ale production by Labatt with high gravity worts for the past 40 years
and wasCell
Yeast oneWall
of the reasons
Structure andfor the introduction of centrifuges into Labatt plants (further details on the
use of centrifuges and their effect on brewer’s yeast strains are given later in this paper).
Flocculation
Mortier
 Factors and Soares
Affecting Yeast [67] studied co-flocculation from a different perspective. They viewed co-
flocculation
Flocculationas a process to separate non-flocculent yeast cells from a fermentation medium. The
possibility of this process being employed for the cell separation of different yeast species was

Co-Flocculation
assessed. The fission
Genetic Control yeast Schizosaccharomyces pombe was used as a control, since these cells
of Yeast

 Flocculation
are unable to be aggregated by flocculent cells of S. cerevisiae, due to a lack of compatible receptors
with S. cerevisiae flocculation lectins [56]. However, the yeast Kluyveromyces marxianus can exhibit

co-flocculation, and consequently separate this culture from suspension by settling using flocculent
Flocculation
cells in Brewer’s
of S. cerevisiae Yeast
to enhance it. The different degrees of co-flocculation amongst non-flocculent
Strains
strains were the consequence of the different composition and structure of the yeast cell wall,
particularly subtle differences in its detailed mannan architecture.
Recent studies by Rossouw et al. [74] examined the relevance of different Flo proteins in
mediating differential interspecies aggregation in a “natural” yeast ecosystem in the wine industry.
Flocculation
This publication
Premature Yeastreports on co-flocculation behaviour in mixed cultures of S. cerevisiae and non-
Flocculation
Saccharomyces
Phenotypic Effectsyeaston cultures. It is suggested that adhesion phenotypes and, in particular, Flo
Flocculation
proteins,
Adhesionmayandplay roles
Biofilm in their system dynamics beyond the roles previously assigned to these
Formation
proteins. Further
Centrifuges elucidation
to Crop Yeast of the molecular mechanisms interpinning different co-flocculation yeast
behaviours
Conclusionswill be of fundamental importance to understand the role of direct cell-cell interactions
between different species in a shared environment.
Acknowledgments
Cell adhesion
 Conflicts of Interestphenotypes are complex, and are influenced not only by genetic factors which
determine
Referencesthe composition and properties of the cell wall, but also by a number of environmental
parameters which impart adhesion behaviour. The observations, under controlled laboratory
conditions, therefore presents a simplified view of interspecies co-flocculation as discussed in this
review. The data provides, for the first time, genetic insights into the phenomenon of co-flocculation in
S. cerevisiae.
Future studies should include additional yeast species and strains under various experimental
conditions in order to comprehensively investigate the yeast co-flocculation concept. In addition, the
relevance of yeast species adhesion phenotypes to microbial interactions in natural ecosystems
requires further detailed investigation [75,76].
9. Genetic Control of Yeast Flocculation
Genetic studies on yeast flocculation began over 70 years ago. The first publication on this
subject was by Pomper and Burkholder [77], who reported crossing a haploid culture that possessed a
“dispersed” character (non-flocculent) with a haploid culture that possessed a non-dispersed character
(flocculent). The “dispersed” character was reported to be dominant over the non-dispersed character.
Back to TopTop
In

the early 1950s, Gilliland (working in the Guinness laboratory in Dublin) and Thorne (working in the
Abstract (/)
Alfred Jorgensen
Introduction laboratory in Copenhagen) independently carried out extensive studies on the

genetics of yeast flocculation. Gilliland [17] studied two non-brewing strains of S. cerevisiae which
Yeast Flocculation
differed only in their
Measurement flocculation
of Yeast properties. It was proposed that a single gene was responsible for the
Flocculation
flocculent
Cell Wallphenotype.
CharacteristicsThorne [18] confirmed Gilliland’s studies and demonstrated that yeast
flocculence was an inherited characteristic and that the flocculence phenotype was dominant over the
Sexual Aggregation
non-flocculent
Yeast Cell Wall characteristic.
Structure and These studies were also confirmed by Guo et al. [41].
It has already been discussed that genetic studies, involving brewer’s yeast strains, is fraught
Flocculation
with difficulties
Factors because
Affecting Yeast of their frequent triploid, polyploid, or aneuploid ratios [78]. It was confirmed
that this yeast characteristic is controlled by dominant genes that were termed FLO genes [79]. FLO
Flocculation
genes from S. cerevisiae resemble the EPA (Epithelial Adhesin) gene family from the opportunistic

Co-Flocculation
pathogenic yeast of
Genetic Control species
Yeast Candida glabrata [80] and the ALS genes (agglutinin-like sequence) from

 Flocculation
Candida albicans [81]. C. glabrata encodes a family of at least 23 ETA genes, which encode cell 
surface proteins capable of mediating adherence to epithelial cells.

The first flocculation gene from S. cerevisiae to be studied in detail was FLO1 [79]. The early
Flocculation
studies focussed in Brewer’s Yeast haploid and diploid flocculent and non-flocculent strains. The FLO1-
on laboratory
Strains
containing haploid strain studied (coded 169) was a MATα mating type opposite to that of a non-
flocculent haploid strain (coded 168-MATα). These two strains were mated using micromanipulation
techniques and found to be flocculent, confirming previous findings that the flocculent character
(coded by FLO1) was dominant and stable [82].
Flocculation
The mapping
 Premature of the FLO1 gene employed classical gene mapping techniques (mating,
Yeast Flocculation
sporulation,
Phenotypicmicromanipulation,
Effects on Flocculationtetrad analysis, spore germination, multiple flocculation tests, etc.)
according
Adhesiontoand theBiofilm
methods discussed by Sherman and his colleagues [83]. Since the successful
Formation
mapping of FLO1,
Centrifuges to Cropnovel genetic techniques have been developed, the principal of which began with
Yeast
theConclusions
sequencing of the Saccharomyces genome [82,84] with the haploid yeast strain S288C. This has
significantly expanded our knowledge regarding the genetic control of yeast flocculation. As a result of
Acknowledgments
theConflicts
reversible inhibition of flocculation by sugars, salt, a low pH environment, or protease sensitivity,
of Interest
two main flocculation phenotypes have been distinguished: Flo1 and NewFlo phenotypes [55]. The
References
Flo1 phenotype includes strains in which flocculation is specifically inhibited by the monosaccharide
mannose and its derivatives. The NewFlo phenotype contains the majority of brewing ale strains. In
this phenotype, flocculation is reversibly inhibited by mannose, maltose, glucose, and sucrose, but not
galactose. NewFlo phenotype strains are more sensitive to inhibition by cations, low pH conditioning,
and digestion by trypsin or proteinase [22]. Also, phenotypes display different sensitivities to culture
conditions such as temperature [9], pH [85], ions, and nutrient availability [86].
It has already been discussed that there are a number of dominant and recessive genes as well
as a flocculation actuator and suppressor genes. The genes that encode flocculation lectins are called
FLO genes. FLO1 is the most studied [87]. There are at least nine FLO genes—FLO1, FLO5, FLO8,
FLO9, FLO10, FLO11, FLONL, FLONS, and Lg-FLO in both S. cerevisiae and S. pastorianus strains
that encode flocculin proteins (Table 1). The flocculin encoded by the FLO11 gene differs from the
others in that it is involved in filamentous growth (pseudomycelia), adhesion to solid surfaces, and flor
formation rather than flocculation per se [88]. Another FLO gene, FLO8, encodes for a transcription
factor regulating the expression of other FLO genes [89]. Many yeast strains usually possess several
FLO genes [90]. For example, it has already been briefly described that the first completely
sequenced haploid strain—S288C [82]—contains six FLO genes (FLO1, FLO5, FLO8, FL010, Back to and
TopTop

FLO11) as
Abstract well as four non-functional FLO pseudogenes [90]. The amino acid sequences of Flo5,
(/)
Flo9, and Flo10
Introduction proteins are 96%, 94%, and 58% identical, respectively, to Flo10p. As expected,

Flo11p
YeastisFlocculation
the most distantly related to other flocculins, being only 37% identical to Flo1p [27].

Table 1. Current view of flocculation phenotypes [89].
Flocculation
Flocculation


Co-Flocculation
 The DNA
Genetic sequences
Control of Yeast of FLONL and FLONS are very similar to that of FLO1, but compared to this

 Flocculation
gene they have lost some of the internal tandem repeats. More tandem repeats were lost in FLONL 
 Flocculation Gene
than in FLONS. TheStructure
deletion of these repeats appears to have converted the flocculation phenotype

from Flo1 to the NewFlo phenotype [91]. The NewFlo phenotype includes the majority of brewing ale
Flocculation
yeast in Brewer’s
strains. In Yeast flocculation is reversibly inhibited by mannose, maltose, glucose, and
this phenotype,
Strains
sucrose, but not galactose [26,55]. Moreover, the NewFlo flocculation phenotype conferred by FLONS0
and FLONL is also inhibited by galactose [92], whereas the usual NewFlo phenotype is insensitive to
galactose. This indicates that the sugar-binding properties of the flocculins are dependent upon the
number of tandem repeats present in the flocculation gene. The different flocculent phenotypes are
Flocculation
summarised in Table
Premature Yeast 1.
Flocculation
10. Flocculation Gene Structure
 Conclusions
FLO genes are very long (up to 4.6 Kbp), whereas the average gene length in the
 Acknowledgments
Saccharomyces genome is only 1.0 Kbp because the number of tandem repeated DNA sequences of
about 100 nucleotides are repeated 10–20 times in each gene [93]. Although tandem repeated DNA
 References
sequences in FLO genes are highly dynamic components of the genome, they change more rapidly
than other parts of the Saccharomyces genome [27]. They enable rearrangements both between and
within flocculin genes. The longer the Flo protein, the stronger the yeast strain’s flocculation ability
[94]. FLO1 has been identified as the longest FLO1 gene, with the most repeats, and it confers the
strongest flocculation phenotype [95].
Also, all FLO genes except FLO11 are located close to telomeres (disposable buffers at the end
of chromosomes). Consequently, they are prone to rearrangements such as deletions, duplications,
and translocations. Indeed, the Lg-FLO1 gene itself represents a translocation event between FLO
genes located on different chromosomes. In addition, near-telometric genes (as well as FLO11) [96]
can become transcriptionally repressed by an epigenetic process known as telomeric silencing. This is
caused by an alteration in chromatin structure near the telomeres, leading to the silencing of genes
located in that region. This effect can be long-lasting. For example, the epigenetic state of FLO11 is
inheritable for many generations. The strength of this telomeric silencing varies greatly between
different telomeres in yeast. Also, significant variation has also been detected between strains. The
series of proteins associated with a trithorax-related SET domain protein (COMPASS) complex is
involved in telomeric silencing in yeast [97]. FLO and MAL (a gene complex that controls maltose
Back to TopTop

metabolism
Abstract in yeast) genes near to telomeres were found to be silenced in some yeast strains,
(/)
whereas in
Introductionother strains with an inactivated COMPASS complex the expression of these genes was

notYeast
affected. This is consistent with the finding of significant variation in the strength of the silencing
Flocculation
effect between different
Measurement chromosome ends and in different strains. In strains where the COMPASS
of Yeast Flocculation
complex
Cell Wallhad a strong silencing effect, genetic inactivation of this complex increased the expression of
Characteristics
FLO1,
SexualFLO5, and FLO9 genes. Compared to wild cells, these mutants displayed enhanced
Aggregation
flocculation
Yeast Cell properties during
Wall Structure and high gravity wort fermentation. Flocculation occurred earlier and formed
much larger aggregates [96]. These findings are of considerable interest concerning the regulation of
Flocculation
flocculation.
Factors Affecting Yeast
Flocculation


Co-Flocculation
11. The Genetic Instability of Flocculation in Brewer’s Yeast Strains

 Flocculation
The sedimentation performance of a brewer’s yeast strain very often changes during repeated 
cropping and re-pitching in a brewery [98]. In principle, this could be due to irreversible or reversible

genetic change [98]. Alternatively, it could be due to long-lasting physiological, perhaps epigenetic
(the study of cellular or physiological phenotypic traits) effects caused by external or environmental
Strains
factors that switch genes on and affect how cells read genes [99] and respond to modifications 0in
The Influence of Cell Surface
yeast-handling
Hydrophobicityand fermentation
(CSH) and Cell environments (for example, high gravity worts [7]). When a genetic
change,
Surfaceconferring
Charge onaYeastnon-flocculent phenotype, occurs in a yeast culture, the culture gradually
becomes a mixture of flocculent and non-flocculent cells [100]. Often within a production lager yeast
Flocculation
population
Prematureexhibiting moderate flocculent characteristics, a more flocculent variant within the culture
Yeast Flocculation
can be isolated.
Phenotypic An example
Effects of this development occurred when a Canadian brewing company began
on Flocculation
brewing
Adhesionits lager beer, Formation
and Biofilm under contract, in breweries located in the United Kingdom. Most of the
contracted
Centrifuges UKtobreweries
Crop Yeastemployed vertical fermenters (Figure 5). However, at that time, Canadian
breweries
Conclusionspredominantly employed horizontal tanks (as both fermentation and maturation vessels).
This difference in tank geometry influenced the yeast culture’s sedimentation characteristics. In
Acknowledgments
vertical fermenters,
Conflicts of Interestthis yeast culture was too non-flocculent (powdery), with considerable yeast cells
remaining
References in suspension at the end of fermentation (centrifuges were not available in the UK
breweries at the time). It was thought that possibly this culture contained a spectrum of isolates that
exhibited differing flocculation intensities. Consequently, one of the variants from this strain, with more
intensive flocculation characteristics, was successfully employed in the vertical fermenters. The result
was less yeast in suspension at the end of fermentation. However, care had to be taken to ensure that
the flocculent variant used was not too flocculent because under-fermented wort (unfermented
maltose and maltotriose) and residual undesirable flavours (for example, diacetyl and other vicinal
diketones) could have been the result [69].
PCR-based methodology (a technique that amplifies a single copy or a few copies of DNA in
order to generate millions of copies of a particular DNA sequence) has been employed to detect FLO5
genes [101]. All 48 single colonies isolated from the stock culture of a lager strain exhibited this PCR
band. After repeated recycling of this yeast culture in a brewery, single cells were again isolated from
batches of the yeast that exhibited poor sedimentation performance. In one case, most (75%) single-
cell colonies failed to yield the FLO5 PCR band, indicating that a genetic mutation (loss of intact
FLO5) had occurred and spread throughout the population, and it was necessary to replace the
brewery yeast with a fresh culture [102]. In the other case, nearly all (>90%) of the single colonies still
Back to TopTop

showed the
Abstract FLO5 PCR band, indicating that the poor sedimentation behaviour did not exclusively
(/)
have
Introduction basis, and was due in many instances to the culture’s physiological condition. In this
a genetic

case,
Yeast theFlocculation
flocculation ability gradually recovered during repeated recycling of the yeast in the brewery.
Spontaneous
 Measurement changes
of Yeast in flocculation behaviour as a result of repeated re-pitching in wort during
Flocculation
brewery
Cell Wallfermentations
Characteristics have been studied by a number of research groups [103,104] and changes in
flocculation intensity [76] have been observed. Cropped ale yeast, during 30 successive
Sexual Aggregation
fermentations,
Yeast Cell Wall has been studied
Structure and [105]. During the first seven cycles, flocculation intensity increased
from 50% to 100% of the original culture. Between the ninth and 23rd cycle, flocculation remained
Flocculation
high. Then,
Factors between
Affecting the 24th and 32nd cycles, flocculation ability and cell viability diminished. Sato
Yeast
and colleagues [100] reported that the flocculation intensity of a lager strain decreased after serial re-
Flocculation
pitching. A long-term study in a number of breweries showed that flocculation intensity tended to

Co-Flocculation
decrease
Genetic whilst
Controlatofthe same time other fermentation parameters (wort fermentation rate, ester, and
Yeast

 Flocculation
higher alcohol production, etc.) usually remained constant. However, studies in the author’s breweries 
with high gravity worts (16–18 °Plato) do not confirm this fact—rather, they proposed that many

observations of this nature must be yeast strain-dependent [7].
Flocculation
Cropping in Brewer’sfavour
methods Yeast the enrichment of certain cell types. Few modern lager breweries
Strains
currently recycle their yeast culture more than 20 times (many breweries less than 10 times) [1].
However, as the original wort concentration (gravity) increased, in the situation that occurs with high
gravity brewing (HGB), the number of yeast cycles has been significantly reduced. Typically, with a 16
°Plato wort, the number of cycles is 10 times or less [7]. The results of these studies suggest that it is
Flocculation
more likely that
Premature Yeasta change in flocculation behaviour is due to a modification in process conditions or
Flocculation
raw materialsEffects
Phenotypic (for example, a malt leading to changes in wort composition also increased the adjunct
on Flocculation
(un-malted
Adhesion cereal levels)),
and Biofilm especially if the change persists when freshly propagated yeast is
Formation
introduced
Centrifuges intoto the
Crop process.
Yeast
It has been suggested [103] that the PCR-based method to detect FLO5 genes (discussed
 Conclusions
above) can be used for the early detection of non-flocculent mutants in brewery fermentations. More
Acknowledgments
than 30 different
Conflicts production lager strains with PCR primers designed to detect FLO5 have been
of Interest
detected.
References The PCR product varies in size from 4.8 to 2.3 Kb, with more flocculent strains showing
larger gene products (>4 Kb) and less flocculent strains showing smaller products (2.3 Kb in a non-
flocculent strain). Cell surface hydrophobicity also decreased with the reduced size of the PCR
product. Consequently, the appearance of non-flocculent mutants during a series of brewery
fermentations could be tested by PCR analyses. Interestingly, PCR primers designed for Lg-FLO1,
FLO1, or the regulatory gene, FL08, did not exhibit this predictive power. In strains failing to form a
PCR product with the PCR primers, Southern blot hybridisation (a method to detect a specific DNA
sequence in DNA samples [106]) after chromosome fingerprinting using a FLO5 fragment as a probe
showed that FLO5 was missing from chromosome VIII, whereas hybridisation to chromosome I (the
location of FLO1) still occurred. The sequences of the successful FLO5 PCR primers were not
revealed. Consequently, we do not know whether these primers only recognise FLO5 (which is 96%
identical to FLO1).
12. The Influence of Cell Surface Hydrophobicity (CSH) and Cell Surface Charge on Yeast
Flocculation
Back to TopTop

 There have been a number of published studies that indicate that an increase of CSH and a
Abstract (/)
decrease of cell
Introduction surface charge occur at the outset of flocculation [107]. CSH increases rapidly as

cells pass
Yeast through the exponential growth phase and reach higher and more stable levels during the
Flocculation
stationary phaseof[63].
Measurement YeastLow CSH in an exponential phase culture is due to the presence of many
Flocculation
daughter
Cell Wallcells, which are significantly less hydrophobic than older (more mature) cells [69]. It is
Characteristics
considered that CSH plays an important role in maintaining the correct conformation of flocculin
Sexual Aggregation
molecules
Yeast Cell[50],
Wallso that theand
Structure flocculins located in a stationary phase cell are more active [108]. It has
already been discussed that the yeast cell surface has an overall negative charge. As a result, the cell
Flocculation
wall phosphate
Factors Affectinggroups
Yeastand mannoproteins [109] are greater when the pH of the fermented medium is
higher [110]. The cell surface charge of brewer’s yeast strains has been shown to vary during growth
Flocculation
[111]. However, no clear relationship between cell surface charge and flocculation onset has been

Co-Flocculation
observed
Genetic [112].
ControlAofnumber
Yeast of environmental factors have been observed to affect CSH. Increased

 Flocculation
CSH can be stimulated by higher ethanol concentrations [21], lower temperatures [113], and higher 
pitching (inoculation) rates [21]. It has been proposed that hydrophobic oxylipins located at the cell

surface of flocculent cells are a cause of increased CSH [91]. However, a recent publication [114]
Flocculation
suggested thatinthe
Brewer’s Yeast
precise rate of 3-OH oxylipin formation is still unclear and detection methods must
Strains
be combined with novel techniques to target the cell wall architecture.
Ale strains (S. cerevisiae) have been found to be more hydrophobic and less negatively charged
than lager (S. pastorianus) strains [107]. As well as contributing to floc formation, the greater
hydrophobicity
Flocculation of ale strains (not definite) explains why the flocs of this species associate with CO2
bubbles and Yeast
Premature rise toFlocculation
the beer surface during traditional top-cropping ale fermentations, whereas the
flocs of lager Effects
Phenotypic strainsondescend to the bottom of the fermenter (bottom cropping) [1].
Flocculation

13.Centrifuges
Premature to Yeast
Crop Yeast
Flocculation
 Conclusions
 It is worth repeating that the timing of yeast flocculation during wort fermentation is crucial for
Acknowledgments
beer production
Conflicts with
of Interestthe necessary quality characteristics. Occasionally, certain malts can cause
premature or heavy flocculation, leaving the wort under-fermented with sugars still in solution and
References
preventing the achievement of the alcohol specification [115]. This phenomenon has been termed
premature yeast flocculation (PYF) [116] and is detected by way of a fermentability test [117].
Most of the PYF studies have not focussed on yeast per se. They have focussed on the malt
employed to produce the wort used during the fermentation studies. Nevertheless, nature-nurture
interactions are critical [11]. Also, the need for future detailed PYF studies with a number of brewer’s
yeast strains (both ale and lager) are emphasised later.
Depending on the local national beer regulations, often a positive result with the fermentability
test does not always translate into a problem because this discrepancy is not questioned. However, it
has already been discussed (Section 8) that some countries (for example, Canada) enforce a rigid
beer alcohol specification (±0.2 v/v) which is listed on every bottle, can, or keg.
During the past 50 years, more than 30 papers have discussed the PYF phenomenon. In the last
two decades, South African Breweries Research Group in Johannesburg devoted considerable
attention to PYF, which has considerably extended our understanding of the problem [117]. It should
be emphasised that PYF is distinct from typical yeast flocculation, which discussed in other sections of
this paper.
Back to TopTop

 What exactly is PYF [66,116]? The published literature contains a number of definitions [118].
Abstract (/)
Although the details
Introduction vary, the definitions are similar. Lake and Speers [119] noted that PYF behaviour

results
Yeastin:
Flocculation
 Rapid yeast
Measurement of flocculation out of suspension, resulting in elevated apparent extract values relative
Yeast Flocculation
 CelltoWall
a “normal” malt (depending on the fermentation vessel employed);
Characteristics
 typically,
Sexual normal fermentations exhibit parabolic yeast-in-suspension trends while PYF yeast-in-
Aggregation
 suspension
Yeast curves proceed
Cell Wall Structure and in a normal and parabolic manner to a peak and then decline in a
Flocculation
concave fashion (Figure 14).
Flocculation


Co-Flocculation


Strains
Flocculation

Figure 14. Test tube-sized (15 mL) fermentations of a premature yeast flocculation (PYF) and

control malt yeast in suspension continuously measured [119].
Conclusions
Acknowledgments
Two theories dominate the literature with respect to the development of PYF factors in barley and
Conflicts of Interest
malt. PYF tends to be a sporadic phenomenon that occurs concurrently with wet and rainy seasons. In

theReferences
first theory, it is believed that increased microbial loads during wet seasons lead to the production
of PYF factors [120]. Barley husk is the main carrier of microorganisms [121]. The microflora consists
of bacteria, wild yeasts, and filamentous fungi.
The second theory regarding PYF mechanisms is that antimicrobial peptide factors inhibit or
negatively affect yeast metabolism. This would initiate flocculation earlier and/or to a greater extent
than normal. It has been shown that a PYF-positive malt leads to a minor decrease in wort sugar
(particularly maltose and maltotriose) metabolism by yeast [10]. However, it is unclear whether this
reduction in sugar metabolism is due to insufficient yeast in suspension (due to PYF) or a direct
inhibition of yeast metabolism. However, Lake and colleagues [119], who employed a miniature
fermentation assay, did not detect metabolic differences between PYF and a control malt. This course
of events does not preclude antimicrobials associating with yeast cells, which would lead to increased
flocculation.
As would be expected, yeast strains show different susceptibility to PYF malts [119]. It is unclear
whether strain susceptibility is due to differing mechanisms, categories of PYF, or whether variability is
related to the differing flocculation potential of various yeast strains. Also, it is unclear whether ale
Back to TopTop

yeast (S. cerevisiae)
Abstract strains are less susceptible or if the effects of PYF are masked by the
(/)
flocculation behaviour of hydrophobic ale yeasts.
Introduction

 YeastStudies by Chinese maltsters have conducted small-scale fermentations with worts produced
Flocculation
from seven malts.
Measurement These
of Yeast fermentations monitored their PYF potential and showed that PYF factors
Flocculation
were
Cellpresent in both the malt husk and non-husk portions. Also, it was shown that antimicrobial
substances that damage yeast cells were present in the non-husk portion [118]. Further research is
Sexual Aggregation
required on the
Yeast Cell Wallmechanisms
Structure and of PYF activity. No single theory supports or challenges any of the
currently proposed mechanisms. Most publications have reported on the gross chemical ratio of PYF
Flocculation
factors and
Factors have avoided
Affecting Yeast discussing or investigating PYF mechanisms.
Flocculation


Co-Flocculation
14. Phenotypic Effects on Flocculation

 Flocculation
Yeast flocculation complexities cannot be overlooked. In addition to the genetic characteristics 
(nature) of brewing strains (FLO genes together with their suppressors and activators), a number of

nurture parameters will also affect yeast flocculation [11,65].
Strains
14.1. Cations
Cations have(CSH)
Hydrophobicity similar
androles
Cell in both ale and lager yeast flocculation. However, calcium ions are
recognised as theon
Surface Charge most
Yeasteffective ions for the promotion of flocculation [53]. Although details of this
importance have been known for a long time [46], the interaction with the yeast cell surface requires
Flocculation
further elaboration.
Premature It is important
Yeast Flocculation to reiterate the importance of calcium in yeast flocculation. Calcium
can be removed
Phenotypic from
Effects onthe yeast cell surface by washing with deionised water, through which many
Flocculation
flocculent
Adhesioncultures will Formation
and Biofilm be de-flocculated. A de-flocculated culture will become flocculent again when
calcium ions are
Centrifuges added
to Crop (Figure 15). Some flocculent yeast strains are not de-flocculated by washing
Yeast
with water, but rather the cells need to be treated with a solution of a chelating agent such as EDTA—
Conclusions
10Acknowledgments
mM [1] followed by washing with water to remove the EDTA. This treatment de-flocculates these
cultures, and
Conflicts the flocculation phenotype is restored upon the re-addition of calcium ions.
of Interest
 References
Back to TopTop

 Figure 15. De-flocculation of yeast as a result of repeated water washing and re-flocculation
Abstract (/)
following the addition of calcium ions [60].
Introduction

 Employing radio-labelled Ca45, it has been shown that flocculent cultures bind more Ca2+ ions
than non-flocculent cultures [60]. There was no direct correlation found between the total calcium
adsorbed and the strain’s flocculation phenotype. However, there is strain-to-strain variation in calcium
Sexual Aggregation
binding, and furthermore this variation does not correlate with flocculation and non-flocculation when
one strain was compared to another. With the knowledge that many flocculent yeast cultures can be
Flocculation
de-flocculated by washing
Factors Affecting Yeast with deionised water, it was of interest to see if the amount of calcium
washed off a yeast culture could be correlated with the visible loss of floc formation.
Flocculation
As a result of this calcium adsorption study, an improved perspective of calcium-binding
 Co-Flocculation
behaviour in yeastofand
Genetic Control

its relationship to flocculation might be obtained to test this hypothesis.
Yeast
of flocculent and non-flocculent yeast suspensions were taken and incubated with the Ca45 

Aliquots
 Flocculation
solution. The yeast pellet was then washed four times with deionised water and the activity of each

centrifuged supernatant was determined with a scintillation counter. In addition, the amount of calcium
Flocculation
removed with in Brewer’s
each washingYeast
was determined. The first wash did not de-flocculate the flocculent yeast
Strains
cultures but did remove adhering calcium around and in the interstitial spaces between the yeast cells.
This source of calcium was expected to be relatively the same percentage of the total calcium bound
for each yeast culture and was in all probability not related to flocculation, since the visible observation
of Flocculation
flocculation did not disappear during this first wash.
Subsequent
 Premature washings gradually dispersed any flocculation characteristic. The sum of the calcium
Yeast Flocculation
removed in washings 2 to 4 were expressed as a percentage of the total calcium removed during
washing. When the results were expressed in these terms (Table 2) for both flocculent and non-
flocculent cultures, the flocculent cultures were found to have bound 28–48% more calcium after four
washings than did non-flocculent cultures. As would be expected, there is strain-to-strain variation in
Conclusions
calcium adsorption [60]. This variation is in all likelihood a reflection of diversities in cell wall structure
Acknowledgments
strain to strain. In addition, this strain-to-strain variation in calcium adsorption per se does not
correlate with the flocculation phenotype when one strain is compared to another. The only meaningful
References
measure of calcium behaviour that correlated with flocculation was the case in which calcium was
washed off the cell, and this coincided with the visible loss of flocculation.
Table 2. Labelled (*) calcium removed from co-flocculent and non-flocculent cultures during de-
flocculation washings [60].
Zn2+, Mg2+, and Mn2+ have also been described as inducers of flocculation [56,122]. Second to
calcium, the effect of zinc on flocculation has received the greatest attention regarding yeast
flocculation and there are a number of publications considering this parameter [123,124]. In addition, a
great deal of research has been conducted studying the effect of Zn2+ on enzymatic activity
(particularly—but not only—alcohol dehydrogenase) and fermentation efficiency [125]. The Back to TopTop

flocculation-de-flocculation
Abstract behaviour of S. cerevisiae is strongly dependent on the concentration of
(/)
++ ions in the fermentation medium (for example, wort) and is yeast strain-specific. However, S.
ZnIntroduction

pastorianus is not affected by the presence of Zn2+, which suggests another useful method for
Yeast Flocculation
distinguishing
Measurementbetween
of Yeast lager and ale flocculent yeast strains [124]. However, the zinc effect requires
Flocculation
considerable further study [126].
Cations
 Sexual such as Ba2+, Sr2+, and Ph2+ competitively inhibit yeast flocculation because of the
Aggregation
similarity of their ionic ratios 2+
Yeast Cell Wall Structure andto Ca [43]. It is possible that these cations compete for the same
“calcium site” of flocculation lectins, but are not able to induce the appropriate conformation of the
Flocculation
lectins. At Affecting
Factors low concentrations,
Yeast Na+ and K+ induce flocculation most likely because of a reduction in the
Flocculationrepulsive forces with the yeasts and/or they stimulate the leakage of intracellular Ca2+
electrostatic
[122,127].

Co-Flocculation
14.2. Medium pH

Flocculation
 
 Medium pH
Flocculation canStructure
Gene have a profound effect on the yeast flocculation phenotype. With many laboratory

and industrial
The Genetic strains,
Instabilityflocculation
of occurs over a wide pH range (2.5–9.0), while many brewing strains
(a Flocculation
sub-group of in the
Brewer’s Yeast
NewFlo phenotype) only flocculate within a narrow pH range (3.9–5.5) [86]. In
Strains
both cases, the optimum pH value takes place between 3.0 and 5.0, according to the yeast strain
being studied. Extreme pH values promote a reversible dispersion of flocs. The modification of the pH
value affects the ionisation of lectin amino acids with the consequent change in its conformation [52].
Most ale strains (S. cerevisiae) do not flocculate following growth in a chemically defined medium
Flocculation
such as yeast nitrogen base (YNB) [11]. It could be that these strains also exhibit a narrow pH range
of Phenotypic
flocculation [86]. However, peptone, certain peptides, and wort also plays an important role
Effects on Flocculation
because ale cells grown in a peptone-containing medium or wort exhibit the flocculation phenotype.
This does not only occur in the culture medium. Following cell harvesting and washing, the cells were
still flocculent in an in vitro flocculation test such as the Helm’s Flocculation Test [28] (Figure 6).
Conclusions
 Acknowledgments
14.3. Temperature

Incubation temperature at different levels can affect the expression of the yeast flocculation
References
phenotype. The lowering of growth and fermentation temperature results in a decrease in yeast
metabolism and CO2 production. Consequently, there is a reduction in turbulence which favours yeast
sedimentation. During beer fermentation, the agitation (shear force—as discussed later) temperature
can also affect yeast flocculation by acting on cell-cell interactions. A rise in temperature to 50–60 °C,
for only a few minutes, promotes the reversible dispersion of flocs [117], probably because of
denatured flocculation lectins. Indeed, the incubation of yeast strains above the optimum growth (35–
37 °C) leads to a reduction [22] or impairment of yeast flocculation [35]. It is probable that heat stress
(<37 °C) acts directly on mitochondrial activity [128] and indirectly on cell membrane structure, which
affects lectin secretion with a consequent reduction in flocculation.
14.4. Oxygen
The influence of wort oxygenation (aeration) in the early stages of both brewing and distilling
fermentation is well documented [2]. In addition, there are a number of publications that discuss why
moderate aeration is beneficial for yeast flocculation [129]. However, the principal role of oxygen
during the initial stages of wort fermentation does not focus on yeast flocculation. It acts as a catalyst
during the synthesis of unsaturated fatty acids and sterols, whose primary role is to participate in the
Back to TopTop

structure
Abstractof membranes, particularly the plasma and mitochondrial membranes [128]. The membrane
(/)
structure is
Introductionimportant for the stress protection of yeast cultures during adverse environmental

conditions that prevail such as high gravity wort fermentations [7].
Yeast Flocculation
Cell wall mannoproteins
 Measurement are differently expressed under aerobic and anaerobic conditions [130].
The transition
Cell from semi-aerobic to anaerobic conditions, which occurs during the course of brewing
fermentations,
Sexual Aggregationis probably associated with the expression of genes that regulate or encode
flocculation
Yeast Cell lectins. Cells pitched
Wall Structure and into oxygen-depleted wort flocculated relatively early during the wort
fermentation
Flocculationcycle, but to a limited extent. The addition of ergosterol or Tween80 (a non-ionic
surfactant and emulsifier
Factors Affecting Yeast often used in foods and cosmetics) to the same wort restored the normal
flocculation
Flocculation behaviour of the culture. This study concluded that lack of oxygen inhibited the synthesis
of Co-Flocculation
ergosterol and unsaturated fatty acids (UFAs) [128,129]. This limits cell growth and results in the
early onsetControl
Genetic of stationary
of Yeastgrowth phase and cell flocculation [131].


14.5. Sugars

 The
The presence
Genetic or absence
Instability of of fermentable sugars is a major factor influencing flocculation by
NewFlo phenotypic strains [55]. It has long been discussed that as long as glucose, maltose, or
Strains are present in sufficient amounts, flocculation is inhibited because these sugars occupy
maltotriose
the flocculins. This inhibits binding to the mannose residues of adjacent cells. The addition of glucose
(20 g/L) rapidly dissociated the flocs of a starved ale yeast culture containing the NewFlo phenotype
[26]. This is a result that might be expected because glucose blocks the mannose-binding sites of
Flocculation
NewFlo-type flocculins. However, it has been suggested that the loss of flocculation requires energy.
Soares and Durante [132] examined the effect of nutrients on the loss of flocculation and presented
evidence of six aspects regarding the stimulation of flocculation loss with an ale NewFlo phenotype
strain under growing conditions:
 Carbohydrate sources are nutrients that stimulate the loss of flocculation in a defined growth
Conclusions
 medium (for example, yeast nitrogen base (YNB)) [60];
Acknowledgments
 all metabolisable
Conflicts of Interest carbon sources (for example, glucose, fructose, galactose, maltose, and
 sucrose) induce the loss of flocculation in YNB, which ethanol does not—details to follow [4];
References
the rate of sugar-induced flocculation appears to be associated with the rate of sugar
metabolism;
the rate of sugar-induced flocculation loss most likely requires energy and this process is blocked
by ethanol;
growth does not always trigger flocculation loss because cells grown in a medium containing
ethanol remained flocculent;
glucose-induced loss of flocculation requires de novo protein synthesis—cycloheximide addition
(an inhibitor of protein synthesis) to glucose-growing cells impairs the loss of flocculation [133].
15. Adhesion and Biofilm Formation
Yeast cells (including brewing and distilling strains) possess a capacity to adhere to abiotic (non-
living) surfaces, cells, and tissues [9,10]. The cell wall serves as a means for yeast cultures (and other
microbes) to interact with their environment. One of the most critical functions of the cell surface is its
ability to adhere to other cells (floc formation) (Figure 4) and inert surfaces (Figure 9). Adhesion
Back to TopTop

prevents
Abstractcells from being washed away when they are present in a nourishing environment, and
(/)
allows them to form biofilms that protect the cells from hazardous and stressful conditions. In addition
Introduction

to Yeast
industrial environments, pathogenic yeasts exploit their capacity to adhere to abiotic surfaces such
Flocculation
as Measurement
plastic prosthetics in Flocculation
of Yeast order to gain access to the bloodstream and internal organs of patients
(Figure 10)Characteristics
Cell Wall [13].
YeastAggregation
 Sexual cell adhesion is of considerable economic importance for food and beverage processing
companies
Yeast Cellbecause adherent
Wall Structure and yeasts (and other fungi) can form highly resistant biofilms. As well as
cell-cell adhesion (flocculation) in brewing (and wine) yeast cultures, biofilm formation is often
Flocculation
exploited
Factors as a convenient
Affecting Yeast and cost-effective way to separate biomass from fermentation products (for
example, fermented wort and must). The fermentation of biofilms appears to be an adaptive
Flocculation
mechanism because it usually ensures access to oxygen and permits continued growth on substrates

Co-Flocculation
such as nonfermentable
Genetic Control of Yeast ethanol. Biofilm-adhering cells have been shown to have elevated and/or

 Flocculation
modified lipid content as well as increased surface hydrophobicity [12]. 

16.Flocculation
Centrifuges to CropYeast
in Brewer’s Yeast
Strains
 The incentive to optimise brewing operating costs while reducing processing times is imperative
forHydrophobicity
the commercial survival
(CSH) of the process. Breweries (and many other manufacturing industries)
and Cell
continuously search
Surface Charge for ways to exploit production efficiency [134] and lean technology concepts
on Yeast
[135]. As a consequence, the disc stack centrifuge has become a popular component of yeast
Flocculation
process management
Premature systems
Yeast Flocculation in order to reduce fermentation, maturation, and clarification times as
well as to control
Phenotypic effluent
Effects treatment costs [136] (Figure 2).
on Flocculation
 Adhesion
The useandof Biofilm
centrifuges in breweries was initially viewed with misgivings by many brewers [137].
Formation
However, the to
Centrifuges advantages
Crop Yeast and disadvantages are now much clearer [138].
 Conclusions
Modern centrifuges produce gravitational forces in excess of 10,000 times atmospheric pressure,
achieving solid separation in seconds with reduced equipment volume. Centrifuges have a number of
Acknowledgments
applications.
Conflicts of They can be used with brewing yeast cultures for:
Interest
 References
Cropping of non-flocculent yeast cultures at the end of primary fermentation;
reducing the yeast quantity from green beer before the start of secondary
fermentation/maturation;
beer recovery from cropped yeast [138];
removal of cold break (precipitated protein, etc.) and yeast at the end of maturation;
separation of the hot break after wort boiling.
Yeast management and handling systems are influential in determining the physiological status of
yeasts’ subsequent fermentation performance activity [138] and clarification during
aging/lagering/maturation [137]. It has been documented that beer haze can result from the yeast cell
wall releasing mannan as it is processed during agitation in storage and centrifugation [139].
Hydrodynamic stresses can have the potential to inflict damage to the yeast cell wall during beer
production and may lead to unfilterable beer haze formation. Intermediate haze formation from yeast
(there are also other sources of beer haze—a discussion of which is beyond the scope of this review
[140]).
Back to TopTop

 The uses of centrifuges during brewing are diverse (discussed above). This section concentrates
Abstract (/)
onIntroduction
the effect that centrifugation has on brewer’s yeast, particularly regarding yeast cell wall damage

and the resulting
Yeast impact on beer quality and stability together with yeast viability and vitality. Studies
Flocculation
were conductedofatYeast
Measurement Heriot-Watt University during the first decade of this century and focused on a
Flocculation
better
Cell understanding of the effects of passing brewing yeast cultures through a disc stack centrifuge
[136] as part
Sexual of yeast cropping. In order to confirm that the effects on yeast was from centrifugation,
Aggregation
anYeast
extensive number
Cell Wall of centrifugation
Structure and cycles operating at two different G-forces (high and low) was
employed.
Flocculation
The passage
 Factors Affecting of yeast through a disc stack centrifuge exposes cells to mechanical and
Yeast
hydrodynamic
Flocculation shear stresses [138]. These stresses can result in a decrease in cell viability and
vitality, reduced flocculation intensity, cell wall damage, increased yeast extracellular proteinase A

Co-Flocculation
(PrA) levels,
Genetic hazier
Control beers, and poorer beer foam stability [2]. In a more recent study, biological
of Yeast

 Flocculation
indicators of yeast physiology such as viable and damaged cells, intracellular pH, glycogen levels, and 
trehalose levels, as well as beer physical stability parameters including mannan residues, particle

size, and beer haze have been employed to quantify the damage to yeast cells as a function of the
Flocculation
number in Brewer’s
of cycles throughYeast
a centrifuge operating at high G-force (Table 3) [141].
Strains
 The Influence
Table of Cell Surface of yeast cells recycled nine times in 20 °P wort and beer before0 and
3. Characteristics
Hydrophobicity (CSH) and
after centrifugation Cell G-force [138].
at high
Flocculation
 Conclusions
 A disc stack centrifuge (Figure 2) is typically made up of 50 to 150 discs (Figure 16); the number
Acknowledgments
of discs is determined
Conflicts of Interest by process requirements. The discs are truncated cones and flanged at the
inner and outer
References diameters (Figure 16). The close proximity of the discs to one another reduces the
sedimentation distance for yeast cells. Due to centrifugal forces within the centrifuge, numerous
reactionary forces take place between liquid, solid, and discs with shear and beer physical instability
stress occurring on the yeast cell surface (Figure 17).
Back to TopTop
 Figure 16. The disc stack in a centrifuge and the flow pattern [138].
 Abstract (/)
 Introduction

Flocculation
Flocculation


Figure 17. Environment scanning electron microscope (ESEM) of ale yeast strain damage: (A)
Co-Flocculation
 Cells before
Genetic Control passage
of Yeast through a disc centrifuge, and (B) cells following passage through a disc

 Flocculation
centrifuge operating at high G-force [141]. 

 It has
The already
Genetic beenofdiscussed here that in the late 1970s and early 1980s, Labatt in Canada
Instability
(along with a number
Flocculation in Brewer’sof other
Yeast brewing companies) began installing centrifuges in their breweries. This
wasStrains
to harvest ale and lager yeasts at the end of fermentation and, for environmental reasons, to
defer sewer surcharges. All of a sudden in 1988, one of the Labatt breweries with a centrifuge
Hydrophobicity
(operating at 300(CSH)
hL/h)and Cell that its ale fermentations were exhibiting slower and incomplete
reported
fermentations with a 16 °Plato OG wort. Closer study of the fermented wort revealed reduced wort
Flocculation
maltose and particularly maltotriose uptake rates, with residual sugars when fermentation ceased.
Consequently, the fermented wort’s alcohol specification was not achieved. In addition, diacetyl
(butterscotch-like flavour) and other vicinal diketone (VDK) levels were elevated at the end of
fermentation because of difficulties with VDK reabsorption by yeast at the end of primary fermentation
and during maturation [1]. Also, yeast autolysis occurred, resulting in reduced foam stability due to
 Conclusions
excreted intracellular proteinase, elevated unfilterable haze consisting mainly of mannoproteins from
 Acknowledgments
disrupted cell walls, and autolysed yeast off-flavours [141].
The centrifuged ale yeast exhibited decreasing cell viability (determined with methylene blue
 References
staining) during repeated cycles. Also, the same cultures had a higher percentage of respiratory
deficient (RD) petite mutants, determined with the triphenyl tehazolium chloride overlay method [142].
This increasing RD level and decreasing viability were due to centrifugation when the exit temperature
was 30 °C [128]. When the bowl of the centrifuge was cooled and the exit temperature reduced to 20
°C, the cell viability increased, the RD level decreased, the wort fermentation characteristics returned
to normal, and the beer was drinkable again [141].
Studies at Heriot Watt University with a 5 hL/h centrifuge (Figure 2) quantified the damage that
occurs to brewer’s yeast strains as a function of cycles through the centrifuge operating at low and
high G-forces [136]. Biological indicators of yeast physiology as a result of centrifugation include non-
viable and damaged cells, intracellular pH, glycogen and trehalose levels, as well as beer physical
stability parameters such as mannan residues and particle size distribution. Many of these
measurements were conducted with a flow cytometer (Table 3) [143,144]. A detailed discussion of the
principles of flow cytometry is beyond the scope of this paper, but it has been developed to measure
yeast cell viability, damaged cells, intracellular pH (pHi), extracellular PrA levels, mannose residues,
and intracellular glycogen and trehalose (Table 3). When yeast cultures are passed through a disc
stack centrifuge they exhibit lower cell viabilities when compared with cells that had not Back
beento TopTop

centrifuged.
Abstract In addition, the effects of G-force upon yeast cells as a function of centrifugation cycles
(/)
have been studied [141]. In order to establish a relationship between a centrifuge G-force and effects
Introduction

onYeast
the physiological
Flocculationstate of yeast, it was necessary for cells to possess an analogous history and a
similar physiological
Measurement state.
of Yeast The yeast strain employed in this study, S. cerevisiae (an ale strain), was
Flocculation
subjected
Cell Wallto similar fermentation conditions. The yeast culture was exposed to a high G-force (12,000
Characteristics
rpm ≡ 20,000
Sexual G) and the centrifuged cells were subsequently conditioned for an additional 48 h.
Aggregation
 YeastCentrifugation of cultures
Cell Wall Structure and after nine times recycling through 20 °Plato worts demonstrated that the
yeast had been damaged in a number of ways (Table 3). These included a reduction in cell viability as
Flocculation
well as glycogen
Factors Affectingand trehalose levels. Also, the hydrodynamic shear generated by the centrifuge
Yeast
resulted in cell surface components being released (mannoprotein) (Figure 17), together with a PrA
Flocculation
increase, resulting in a reduction in wort hydrophobic polypeptides and poor beer foam stability [141].

Co-Flocculation
 Genetic
Although centrifugation
Control of Yeast can exhibit negative effects, the positive effects of controlled

 Flocculation on beer production and effluent control cannot be overstated. However, yeast is
centrifugation 
subjected to numerous factors that individually and collectively impose stresses on yeast cells (Figure

18) [2]. The effects of environmental conditions and beer production equipment may have been
Flocculation in Brewer’s
underestimated (and even Yeast
ignored). A more complete understanding of yeast’s biological response to
Strains
interactions with cell physiology and brewing equipment is an important criterion in maintaining
process efficiency and beer quality. It is worth noting that the advent of flow cytometry [143,144] and
confocal imaging [145] has introduced analytical methods to yeast research that have broadened the
scope of research in this area.
Flocculation
 Conclusions
 Acknowledgments
 References
Figure 18. Factors which promote yeast stress [7].
17. Conclusions
Brewers employ a number of methods to crop their yeast cultures at the end of a typical wort
fermentation, which vary depending on whether one is dealing with traditional top-cropping, accepted
lager bottom-cropping, cylindroconical fermentation systems, a non-flocculent culture where the yeast,
still in suspension, is cropped with a centrifuge or a portion of the yeast, still in suspension, is blended
into the fresh wort of a subsequent fermentation. Yeast quality is influenced by the manner that the
yeast is cropped and centrifuges play an important part in this regard. The cell wall structure is a
critical parameter involved in yeast flocculation. The wall consists of an inner layer composed
predominantly of β-glucose and chitin with a fibrillar outer layer consisting primarily of α-mannan
Back to TopTop

associated
Abstract with mannoproteins (Figure 8). Yeast management and handling systems, including
(/)
culture harvesting,
Introduction are critical in determining the yeast’s physiological status.

Acknowledgments

The invaluable
Sexual Aggregationassistance and support of Anne Anstruther in developing this review paper is

gratefully acknowledged.
Flocculation
Flocculation


Co-Flocculation
 The author declare no conflicts of interest.
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Keywords: centrifugation; cropping; flocculation; flotation; recycling; sedimentation

Figure 4. Flocculation characteristics of Saccharomyces cerevisiae strains. (A) Flocculent

3. Measurement of Yeast Flocculation

3.2. Direct Observation of Floc Formation in the Growth/Fermentation Medium

3.3. Static Fermentation Methods

Figure 8. Electron photomicrograph of Saccharomyces cerevisiae flocculent and non-flocculent

Figure 10. Electron photomicrographs of adhering and non-adhering cultures of Candida

Figure 12. Chains of S. cerevisiae—pseudohyphae [39,41].

9. Genetic Control of Yeast Flocculation

15. Adhesion and Biofilm Formation

Figure 18. Factors which promote yeast stress [7].

(https://scholar.google.com/scholar_lookup?title=Effects of fermentation parameters

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