|
REVIEW ARTICLE
Flocculation in Saccharomyces cerevisiae: a review
E.V. Soares1,2
1 Bioengineering Laboratory, Chemical Engineering Department, Superior Institute of Engineering from Porto Polytechnic Institute, Rua Dr
Anto nio Bernardino de Almeida, Porto, Portugal
2 IBB-Institute for Biotechnology and Bioengineering, Centre for Biological Engineering, Universidade do Minho, Campus de Gualtar, Braga,
Portugal
Introduction
Yeast flocculation can be defined as a nonsexual, homotypic
(involving only one type of cell in the interactions),
reversible (flocs can be reversible dispersed by the action
of EDTA or specific sugars, like mannose) and multivalent
process of aggregation of yeast cells into multicellular
masses (composed by thousands or even millions of
cells), called flocs, with the subsequent rapid sedimentation
from the medium in which they are suspended
(Fig. 1) (Stratford 1992b; Stewart 2009). The word floc
derives from the Latin word floccus, which means a tuft
of wool. The cells with the ability to form flocs are called
flocculants and look like tufts of wool (Fig. 1a), while the
cells that are not able to form flocs are usually known as
powdery. Saccharomyces cerevisiae cells can be found
aggregated in different ways, which should not be
confused with flocculation, such as sexual aggregation,
co-flocculation and chain formation. Sexual aggregation in
haploid strains of S. cerevisiae of complementary mating
types (a and a) can occur after exchange of pheromones
a and a, respectively, which induces the appearance of
complementary molecules (proteins) on surface of cells
and facilitates the fusion of the haploid cells (Chen et al.
2007). Co-flocculation or mutual flocculation is a heterotypic
aggregation process (while flocculation is homotypic)
among two strains: one of them is nonflocculent and
the other weakly flocculent; when mixed, in the presence
of Ca2+, the strains rapidly settle (Stewart 2009). Chain
formation occurs because of a failure of the younger bud
to separate from the mother cell; this results in an aggregate
formation composed of 30–50 cells. These aggregates
are covalently linked; thus, after mechanical dispersion,
cells will not be able to re-aggregate (Stewart 2009). Other
aggregation phenomena in S. cerevisiae include biofilm
(Verstrepen et al. 2004) and pseudohyphae formation,
which grow and invade the agar medium (Gimeno et al.
1992).
Saccharomyces sensu stricto complex had a long and
profitable association with food, biotechnology and pharmaceutical
industries; it is the micro-organism most
industrially exploited by the man (Vaughan-Martini and
Keywords
aggregation, alcoholic beverages, bio-ethanol,
brewing yeast, cell separation, FLO genes,
flocculation.
Correspondence
Soares Eduardo, Bioengineering Laboratory,
Chemical Engineering Department, Superior
Institute of Engineering from Porto
Polytechnic Institute, Rua Dr Anto nio
Bernardino de Almeida, 431, 4200-072 Porto,
Portugal. E-mail: evs@isep.ipp.pt
2010 0514: received 27 April 2010, revised
and accepted 26 October 2010.
doi:10.1111/j.1365-2672.2010.04897.x
Summary
The present work reviews and critically discusses the aspects that influence
yeast flocculation, namely the chemical characteristics of the medium (pH and
the presence of bivalent ions), fermentation conditions (oxygen, sugars, growth
temperature and ethanol concentration) and the expression of specific genes
such as FLO1, Lg-FLO1, FLO5, FLO8, FLO9 and FLO10. In addition, the metabolic
control of loss and onset of flocculation is reviewed and updated. Flocculation
has been traditionally used in brewing production as an easy and
off-cost cell-broth separation process. The advantages of using flocculent yeast
strains in the production of other alcoholic beverages (wine, cachac愀 and
sparkling wine), in the production of renewal fuels (bio-eth |
|