MINIREVIEW
Occurrence and function of yeasts in Asian indigenous fermented
foods
Kofi E. Aidoo1, M. J. Rob Nout2 & Prabir K. Sarkar3
1
Food Research Laboratories, Caledonian University, Glasgow, Scotland; 2Laboratory of Food Microbiology, Wageningen University, Wageningen,
The Netherlands; and 3Microbiology Laboratory, University of North Bengal, Siliguri, India
Correspondence: M. J. Rob Nout,
Laboratory of Food Microbiology, Bomenweg
2, 6703 HD Wageningen, The Netherlands.
Tel.: 131 317 482834; fax: 131 317 484978;
e-mail: rob.nout@wur.nl
Received 10 March 2005; revised 31 August
2005; accepted 31 August 2005.
First published online 15 November 2005.
doi:10.1111/j.1567-1364.2005.00015.x
Editor: Teun Boekhout
Keywords
pancakes; bread; amylolytic starters; snacks;
beverages; condiments.
Introduction
Indigenous, also referred to as traditional, fermented foods
are those popular products that since early history have
formed an integral part of the diet and that can be prepared
in the household or in cottage industry using relatively
simple techniques and equipment. Some of these products
have undergone industrial development and are also now
manufactured on a large scale (Wood, 1998; Boekhout &
Robert, 2003; Hui et al., 2004). Yeasts occur in a wide range
of fermented foods, made from ingredients of plant as well
as animal origin. This minireview illustrates a selection of
fermented products of particular interest because of the
predominant yeasts that contribute to their attractive char-
acteristics. When yeasts are abundant, alone or in stable
mixed populations with mycelial fungi or with (usually
lactic acid) bacteria, they have a significant impact on food
quality parameters such as taste, texture, odour and nutri-
tive value. Among the Asian indigenous fermented food
products, we examined pancakes and bread, amylolytic
fermentation starters, alcoholic snacks and beverages, and
condiments. Table 1 illustrates the diversity of ingredients
used. Whereas fermentation plays an important role in the
home kitchen, several products are presently manufactured
c 2005 Federation of European Microbiological Societies
Published by Blackwell Publishing Ltd. All rights reserved
Abstract
In the Asian region, indigenous fermented foods are important in daily life. In
many of these foods, yeasts are predominant and functional during the fermenta-
tion. The diversity of foods in which yeasts predominate ranges from leavened
bread-like products such as nan and idli, to alcoholic beverages such as rice and
palm wines, and condiments such as papads and soy sauce. Although several
products are obtained by natural fermentation, the use of traditional starter
cultures is widespread. This minireview focuses on the diversity and functionality
of yeasts in these products, and on opportunities for research and development.
on medium or large industrial scale (Steinkraus, 1989). Up-
scaling of processes requires control of the operations and of
the quality and safety of ingredients and products. In
addition, new aspects of functionality are becoming more
important for exploitation. Future prospects for research
and development will be discussed at the end of this review.
Pancakes and breads
Pancakes
Idli, dosa and appam, consumed in India and Sri Lanka, are
cereallegume mixture foods which, from a nutritional
point of view, are advantageous because of an improved
balance of carbohydrates and proteins. To prepare idli, rice
(Oryza sativa) and blackgram (Phaseolus mungo) dal (de-
husked split seeds) are soaked separately in water. The
soaked rice is then coarsely ground, whereas dal is ground
to a smooth mucilaginous paste. The two slurries (2 : 1) are
mixed with salt, put in a closed container and left overnight
to allow a definite leavening (a two- to three-fold increase in
the original volume) and to develop a pleasant acid flavour.
The fermented batter is poured in cups of an idli pan, and
FEMS Yeast Res 6 (2006) 3039
 Table 1. Yeast-based traditional fermented foods and beverages of Asia
Name of the food Country Major Ingredients Functional Microflora Fermentation contributes to: References
Pancakes and leavened low-salt bread
Idli India, Sri Lanka Rice and blackgram LAB, Saccharomyces cerevisiae Flavour, texture, nutritional value Soni & Sandhu

FEMS Yeast Res 6 (2006) 3039

dal (1991)
Dhokla India Rice and Bengalgram LAB, Pichia silvicola Flavour, texture, nutritional value Kanekar &
Joshi (1993)
Nan, kulcha and India, Pakistan, Wheat flour LAB, Saccharomyces cerevisiae and other yeasts Texture, flavour Sandhu et al.
bhatura Afghanistan, Iran (1986)
Yeasts in Asian indigenous fermented foods

Amylolytic fermentation starters

Ragi Indonesia Rice Amylomyces rouxii, Hansenula spp., Saccharomycopsis Starch degradation, alcoholic Hesseltine
fibuligera fermentation et al. (1988);
Saono et al.
(1996)
Murcha/marcha India, Nepal Rice Mucor spp., Rhizopus spp., Pichia burtonii, Saccharomyces Starch degradation, alcoholic Shrestha et al.
cerevisiae fermentation (2002);
Tsuyoshi et al.
(2005)
Loog-pang Thailand Rice Saccharomycopsis fibuligera, Pichia anomala and other Starch degradation, alcoholic Limtong et al.
yeasts fermentation (2002)
Tane koji Japan Rice Aspergillus oryzae, Aspergillus usamii Starch and protein degrading
enzymes for rice wine making
Men Vietnam Rice Amylomyces rouxii, Saccharomyces cerevisiae Starch degradation, alcoholic Dung et al.
fermentation (2005)
Sweet low alcoholic snacks fermented with amylolytic starters
Tape ketan, Tape Indonesia Rice, cassava tubers Amylomyces rouxii, Mucor spp., Rhizopus spp., Starch degradation, glucose Ko (1986)
ketella/peujeum Saccharomycopsis fibuligera, Pichia burtonii, Pichia anomala formation, alcoholic Boon-Long

c

fermentation, flavour (1986)
Kao-mark Thailand Rice
Wines brewed with amylolytic starters
Sake Japan Rice Aspergillus oryzae, Saccharomyces sake, Hansenula anomala Saccharification, alcohol, flavour
Yakju and takju Korea Rice, wheat, barley, Aspergillus oryzae, Aspergillus sojae, Rhizopus spp., Saccharification, alcohol, flavour Rhee et al.
maize, millet Saccharomyces cerevisiae, Hansenula anomala, Hansenula (2003)
subpelliculosa, Candida sake, Torulaspora inconspicua, Pichia
polymorpha
Tapuy The Philippines Rice Saccharomycopsis fibuligera, Rhodotorula glutinis, Saccharification, alcohol, flavour Kozaki &
Debaryomyces hansenii, Candida parapsilosis, Trichosporon Uchimura
fennicum (1990)
Ruou nep than Vietnam Purple glutinous rice Amylomyces rouxii, Saccharomyces cerevisiae Saccharification, alcohol, flavour Dung (2004)
Jnard/jaanr/thumba Nepal, India, Finger millet/rice/ Mucor spp., Rhizopus spp., Saccharomycopsis fibuligera, Saccharification, alcohol, flavour Tamang et al.
Bhutan maize/wheat Pichia anomala, Saccharomyces cerevisiae, LAB (1988)
31

2005 Federation of European Microbiological Societies

Published by Blackwell Publishing Ltd. All rights reserved
Continued
32 K. E. Aidoo et al.

Batra & Millner

steamed until the idli cakes (Fig. 1a) are soft and spongy

Sandhu & Soni

Mayser et al.

Ebine (1989)
(1974); Joshi
et al. (1999)

Shurpalekar

Aidoo et al.
References with a honeycomb structure inside. A slightly modified
version of this batter, where the proportion of rice is

(1995)

(1989)

(1986)

(1994)
increased and both the ingredients are finely ground, is used
to prepare two additional foods: dosa (highly seasoned
griddled pancake) and appam (poached egg-like pancake).

and proteins, formation of flavour

Idli is a natural fermented food. Both bacteria and yeasts
Alcohol and flavour production

Organic acids, vitamins, health

Enzymic degradation of starch

Fermentation contributes to:

are generally introduced by the two main ingredients and

Acidification, leavening,
participate in the fermentation. A bit of freshly fermented
batter (backslop) is often added to the newly ground batter.

nutritional value
As the fermentation progresses, both bacterial and yeast cell

Texture, flavour
numbers increase significantly with a concomitant decrease
beverage

in pH, and an increased volume of batter, amylase and

Flavour
protease activity. Leuconostoc mesenteroides is the most
commonly encountered bacterium (Nout & Sarkar, 1999).
During fermentation, along with L. mesenteroides, Sacchar-
LAB, AAB, Saccharomyces cerevisiae, Schizosaccharomyces

omyces cerevisiae, Debaryomyces hansenii, Pichia anomala

Aspergillus oryzae, Aspergillus sojae, Zygosaccharomyces

and Guehomyces pullulans are predominant among the yeasts

appearing first, and Trichosporon cutaneum develops subse-
Aspergillus oryzae, Zygosaccharomyces rouxii, LAB

quently. Eventually, only S. cerevisiae persists (Soni & Sandhu,

AAB, Brettanomyces spp., Zygosaccharomyces
pombe, Kodamaea ohmeri and other yeasts

1991).
Candida krusei, Saccharomyces cerevisiae

The role of yeasts in idli batter fermentation is contro-

kombuchaensis, Saccharomyces spp.

versial. Although the fermentation was reported (Ramak-

rishnan, 1979) to be entirely due to heterofermentative L.
mesenteroides, later work has shown yeast involvement in the
fermentation (Venkatasubbaiah et al., 1984). The major
Functional Microflora

functions of the fermentation include the leavening of the

Candida krusei, LAB

rouxii, Candida spp.

batter and the improvement of taste and nutritional value of

idli. The role of lactic acid bacteria is to reduce the pH of the
batter to an optimum level (pH 4.14.5) for yeast activity.
Yeasts help in the degradation of starch (which cannot be
carried out by L. mesenteroides) into maltose and glucose by
producing extracellular amylolytic enzymes. They also pro-
duce carbon dioxide and play a significant role in leavening.
Sap of coconut, date

Soybeans and wheat

Tea liquor and sugar

The sources of the yeast strains are the surface of stone

Rice and soybeans
Major Ingredients

or palmyra palm

grinders used for preparation of batter, and the rice used in

Blackgram dal

Blackgram dal

batter preparations. Not only the load but also the diversity
of yeasts in fermenting batter is greater in winter than in
summer. Fermentation of batter by inoculating the ingredi-
ents with individual yeasts and in combination with L.
AAB, acetic acid bacteria; LAB, lactic acid bacteria.

mesenteroides reveals that yeasts contribute not only to gas

India, Bangladesh,

Japan, Indonesia,

production, resulting in good texture, but also towards

India, Pakistan
Indonesia, the

China, Russia

Japan, China

sensory qualities. The higher activity of amylases, levels of

Beverages fermented from sugary juices

Philippines
Sri Lanka,

Malaysia,
Thailand,

B vitamins and free amino acids attained in yeast-enriched

Country

Japan
India

fermentations suggest the positive contribution of yeasts to

these constituents (Venkatasubbaiah et al., 1984, 1985). It
can be said that even if the yeasts are not essential for the
Palm wines (Toddy/

leavening of idli batter, they are certainly important for the

Table 1. Continued.

tari, Tuack, Tuba)

Papad/papadam
Kombucha/tea-
Name of the food

desirable organoleptic qualities and on nutritional grounds.

The possible synergism between bacteria and yeasts remains
Soy sauces
Condiments

to be determined.
fungus

Wadi

Miso

Dhokla, popular all over India, is similar to idli except

that the dal used is of Bengalgram (Cicer arietinum). During

c 2005 Federation of European Microbiological Societies FEMS Yeast Res 6 (2006) 3039
Published by Blackwell Publishing Ltd. All rights reserved
Yeasts in Asian indigenous fermented foods
Fig. 1. (a) Idli (centre) served hot, with sambar (left) and chutney (right). (b) Amylolytic fermentation starters: Men (left) and Ragi (right). (c)
Condiments: wadis of Punjab (left) and Bengal (top right), showing cavities in some inverted wadis. Middle-sized wadis (bottom right) are made of lentil.
(d) Condiments: Papads (top) as marketed, (bottom) baked.
fermentation, there is an almost two-fold rise in the batter
volume and a drop of pH from 5.2 to 4.5. The microorgan-
isms involved in the fermentation are Lactobacillus fermen-
tum, L. mesenteroides and Pichia silvicola (up to 107 g 1)
(Joshi et al., 1989). The lactic acid bacteria contribute lactic
acid and acetoin, imparting a sour taste and a pleasant
flavour. The yeast produces folic acid and raises the volume
of the batter, imparting sponginess to the product (Kanekar
& Joshi, 1993).
Breads
Nan (naan) is generally consumed as a staple food by the
people of Afghanistan, Iran, India and Pakistan. It is a flat
leavened bread, made by mixing white wheat flour with
sugar, salt, backslop and water. The dough is left for 1224 h,
formed into balls and flattened. Smoothly flattened dough is
slapped onto the inner wall of the clay-clad brick oven,
called tandoor, where it sticks while baking until the dough
is puffed-up and light brown.
From a new dough (pH 5.9) for making nan, 105 yeast
cfu g 1 and 102 lactic acid bacteria cfu g 1 were obtained, as
compared with respective counts of 108 and 109 from ripe,
fermented dough (pH 4.8). Saccharomyces cerevisiae is the
predominant yeast (Sandhu et al., 1986).
Bhaturas (pathuras) and kulchas of northern India and
Pakistan are prepared from similar leavened doughs; they
are respectively deep-fried in oil or prepared on a griddle.
Kulcha dough contains yeasts and lactic acid bacteria. The
yeasts belong mainly to the genera Saccharomyces, Candida,
Hansenula, Saccharomycopsis, Kluyveromyces, Rhodotorula,
Pichia, Torulopsis, Trichosporon and Debaryomyces (Sandhu
et al., 1986).
FEMS Yeast Res 6 (2006) 3039
33
Amylolytic fermentation starters
Amylolytic starters are used in the form of starchy tablets
containing mixed cultures of starch-degrading moulds and
fermenting yeasts. They are used for the manufacture of
beers, wines and pasty snacks from various kinds of cereals
and starchy crops, such as rice, sorghum, millet and cassava.
Table 1 shows some of the starters that have been described
and analyzed. The major principle of their manufacture is to
prepare an uncooked dough of rice or wheat flour (some-
times mixed with cassava meal), water and a variety of herbs
and spices. This dough is inoculated with a backslop and
shaped into small balls or flattened tablets (cakes), which are
incubated at around 30 1C for about a week. During this
period the niche microfloras develop and, simultaneously,
the tablets dehydrate. These tablets can be stored at ambient
temperatures without significant loss of viability for at least
6 months.
Studies on the microflora of starters such as Indonesian
ragi (Fig. 1b) and Chinese chiu-chu have been reported as
early as the end of the 19th century. The principal amylolytic
moulds are Amylomyces rouxii, Rhizopus spp., Mucor spp.
and Aspergillus spp. Common yeasts in many starter tablets
are Hansenula spp., Saccharomycopsis fibuligera, Candida
spp. and Sm. cerevisiae (Saono et al., 1996). The exact
mycoflora of ragi varies with location and the particular
fermentation for which the starter is to be used. A study on
nearly 100 amylolytic yeast strains isolated from ragi and
other starters revealed that the predominant amylolytic
yeasts are Sm. fibuligera and, to a lesser extent, Saccharomy-
copsis malanga (Hesseltine & Kurtzman, 1990).
Selected strains of Aspergillus oryzae are used in the
preparation of tane-koji in the manufacture of sake. The
c 2005 Federation of European Microbiological Societies
Published by Blackwell Publishing Ltd. All rights reserved
34
mould produces a-amylase and amyloglucosidase, which
hydrolyze starches to dextrin, maltotriose, maltose and
glucose, and acid and alkaline proteases, which hydrolyze
proteins to peptides and amino acids. Other essential
moulds in the production of rice wine include Aspergillus
usamii and A. rouxii.
Moulds belonging to the genera Mucor and Rhizopus are
usually the main enzyme producers for the production of
rice wines in India and Nepal (Shrestha et al., 2002). The
main yeasts which ferment saccharified rice starch to alcohol
are Pichia burtonii, Sm. fibuligera, S. cerevisiae, Candida
glabrata and Candida lactosa, while Sm. fibuligera produces
amylolytic enzymes as well (Tsuyoshi et al., 2005). In
murcha from India, Saccharomycopsis capsularis and P.
burtonii also contribute to the degradation of starch
(Tsuyoshi et al., 2005). Other yeast species, namely Hanse-
nula spp., Pichia spp. and Torulopsis spp., have also been
isolated from rice wine. The role of Sm. fibuligera is two-
fold: it is an important amylolytic species (Dansakul et al.,
2004) and it also produces alcohol, albeit at relatively low
levels (Limtong et al., 2002).
In a study of yeast diversity in Thai traditional alcohol
starters, 43 yeasts from 38 samples of loog-pangkao-mag
(starters for alcoholic sweetened rice) and 49 from 19
samples of loog-pang-lao (starters for rice wine) were
investigated (Limtong et al., 2002). Saccharomycopsis fibuli-
gera predominated (31 isolates) in both types of loog-pang;
the organism showed strong amylolytic activity but was a
poor alcohol producer. Other species identified include P.
anomala (8), Issatchenkia orientalis (6), P. burtonii (6),
Pichia fabianii (4), Candida rhagii (4), C. glabrata (3),
Torulaspora globosa (3), Pichia mexicana (2) and single
isolates of Pichia heimii, Rhodotorula philyla, S. cerevisiae,
Torulaspora delbrueckii and Trichosporon faecale. These have
low amylolytic activity but possess high or moderately high
alcoholic fermentation.
An interesting functional aspect of these fungi is their
formation of functional enzymes to release assimilable
carbon sources. These enzymes are valuable in brewing and
flavour development. For example, glucoamylase (glucan
1,4-a-glucosidase) [EC 3.2.1.3] is a key enzyme in rice wine
fermentation, converting starch directly into glucose. Glu-
coamylase is inducible by glucose, starch, maltose and
glycerol and can degrade a variety of polysaccharides. This
enables the yeast to mobilize assimilable carbon sources. On
the other hand, glucosidase activity can also contribute to
flavour development as a number of flavour precursors in
fruits are glycosides. Esters, fusel alcohols, acids and other
compounds which contribute to flavour are also produced.
Based on abilities to produce high amylolytic activities
(mainly due to a-amylase and/or amyloglucosidase), a
number of strains of A. rouxii, Rhizopus spp. and Sacchar-
omycopsis spp. have been selected. For instance, a combina-
c 2005 Federation of European Microbiological Societies
Published by Blackwell Publishing Ltd. All rights reserved
K. E. Aidoo et al.
tion of A. rouxii and Sm. fibuligera produces a good quality
tape . Similarly, defined granulated starters containing
A. rouxii and Sm. cerevisiae make high-quality Vietnamese
rice wine (Dung et al., 2005).
Snacks and beverages
Sweet, low alcoholic snacks
A popular type of indigenous fermented snack can be
encountered in a large part of East Asia, e.g. tape ketan,
kao-mark (from glutinous rice), and tape ketella or peujeum
(from cassava roots) (see Table 1). Common features are the
semi-solid consistency, the distinct sweet, mild-sour taste
and the fruity flavour of alcoholic fermentation that has just
started. They can be consumed as such or used as an
ingredient in homecooking and baking. The principle of
their manufacture involves the preparation of the substrate
(glutinous rice or peeled, chopped cassava) by washing,
soaking and steaming until soft; followed by cooling,
inoculating with powdered starter and fermenting under
cover for 13 days. In Indonesia, ragi tape would be used
(Ko, 1986) and in Thailand, loog-pang starter is used to
prepare kao-mark (Boon-Long, 1986). The major modifica-
tions of the substrate, i.e. release of glucose from cooked
starch by fungal glucoamylase, and alcoholic-organic acids
fermentation and flavour formation are ascribed to a limited
selection of fungi. These include A. rouxii, Rhizopus spp.,
Mucor spp., Sm. fibuligera, P. burtonii, P. anomala and S.
cerevisiae (Boon-Long, 1986; Ko, 1986). Pichia anomala is
required to develop a rich aroma and the typical flavour of
tape (Ko, 1986) by formation, for example, of isobutanol,
isoamyl alcohol and their esters. The ethanol (up to 10%
v/v) serves as a source of calories and helps prevent growth
of disease- or toxin-producing microorganisms in the
products. Although bacteria, particularly Bacillus spp. and
Acetobacter spp., are present in relatively low numbers
(o 105 cfu g 1), they nevertheless are considered to contri-
bute to the quality and flavour of tape ketan (Ardhana &
Fleet, 1989). In particular, the ability of Bacillus spp. to
produce amylolytic and proteolytic enzymes would make a
significant contribution to the degradation of macromole-
cular substrates, leading to its desired moist appearance and
soft texture (Ardhana & Fleet, 1989).
Rice wines
Rice wines are produced from the hydrolytic breakdown
products of cereal starches and other polysaccharides. They
range from simple Thai rice wine to highly sophisticated
Japanese sake . The moulds involved in alcohol production
of Asian rice wines include A. rouxii, and the yeasts
S. cerevisiae, Saccharomycopsis burtonii, Sm. fibuligera,
FEMS Yeast Res 6 (2006) 3039
Yeasts in Asian indigenous fermented foods 35

Cm. lactosa and related yeasts. Rice and/or cereal wines are Jnard
produced on both a cottage and a commercial scale in most
Jnard, or jaanr (Tamang et al., 1988), is a common name for
Asian countries, especially Japan, China, Korea, Thailand,
some traditional alcoholic fermented beverages in the east-
the Philippines and Vietnam (Nout & Aidoo, 2002).
ern Himalayas in India, generally produced by using murcha
Wines may be distilled to obtain a liquor or spirit, for
as a starter. Seeds of finger millet (Eleusine coracana)
instance the famous Indonesian brem bali, an alcoholic
are boiled, drained, cooled, mixed with powdered murcha
liquor produced in Bali from the liquid portions of tape
(Tamang & Sarkar, 1995) (12%, w/w), packed in a bamboo
ketan. These liquors can also be used to fortify rice wine.
basket, covered and left for 24 days. At the
Sake is a pale yellow rice wine of Japanese origin with an
end of saccharification, a sweet aroma is emitted when
alcohol content of 1516% (weight in volume, w/v) or
the mass is transferred into an earthen pot and covered
higher. Steamed rice is mixed with tane-koji (A. oryzae )
to make it airtight. After fermentation, the seeds are
and allowed to ferment for 56 days, after which it is mixed
kneaded to remove seed coats, put into a bamboo/wooden
with yeast moto or ragi starter and water to form the main
vessel and lukewarm water is added. After about 10 min,
mash, or moromi. Moto dominates the moromi fermenta-
the beverage is ready to drink. Good quality jnard has a
tion. Wild yeasts tend to die off at the early stages of moromi
pleasant sweet aroma blended with mild alcohol (Tamang
fermentation due to nitrite produced by nitrate-reducing
et al., 1988).
bacteria.
The mixed population of moulds, yeasts and lactic acid
Malaysian rice wine or tapai is lighter in colour, ranging
bacteria in murcha starter (Tamang et al., 1988) soon
from red to pink. It is made from cooked gelatinized rice
becomes active, bringing about changes in the substrate,
and red pulverized ragi (yeast cake, or jui-piang).
increasing the temperature of the fermenting mass by 4 1C
Yakju and takju are Korean alcoholic beverages originally
over the ambient (2025 1C) within 2 days of fermentation.
made from rice, but which are now made from wheat,
Amylase activity reaches its peak on the second day of
barley, corn or millet. In the traditional yakju process,
fermentation; mucoraceous fungi have an active role in
steamed, cooled rice is mixed with nuruk amylolytic starter,
saccharification and liquefaction of starch. The moulds
and yeast inoculum is added. Takju is made by diluting fresh
Mucor circinelloides and Rhizopus chinensis, and the yeast
yakju liquor prior to filtration. In comparing the traditional
Sm. fibuligera, which are dominant at the start, disappear
and industrial rice wine brewing processes in Korea, it was
within 12 h of fermentation. Mature jnard contains P.
noted (Rhee et al., 2003) that nuruk is used in the traditional
anomala, S. cerevisiae and C. glabrata yeasts and lactic acid
method, normally carried out at low temperatures
bacteria (105106 cfu g 1), which include Pediococcus pento-
(510oC). The industrial brewing follows the Japanese sake
saceus and Lactobacillus bifermentans. These three yeast
brewing method and undergoes relatively high temperature
species increase from 105 to 107 cfu g 1 within 2 days, and
fermentation (25oC). The alcohol content in the industrial
the population then remains the same till the end of
rice wine tends to be higher but the wine is lower in esters
fermentation. The titratable acidity and alcohol content of
than the traditional brew.
the fermenting mass increase significantly during the fer-
In the Philippines, tapuy (Kozaki & Uchimura, 1990)
mentation.
(Igorot ethnic group) is an acidic but sweet alcoholic
rice wine and is known by other names such as binubudan
(Ifugao), binuburan (Ilocano) or purad (Tagalog). The Thai
Palm wines
rice wines such as sato and krachae are cloudy yellow liquids
made from glutinous rice (Vachanavinich et al., 1994). In almost all tropical places in Asia where palm trees grow,
In Vietnam, ruou nep and ruou nep than are made by the sap obtained from the decapitated inflorescence of
steaming white or purple glutinous rice, respectively, various species of palms is fermented to produce an
and inoculation with men, an amylolytic starter (Table 1, alcoholic beverage called palm wine, or toddy. In India there
Fig. 1b). Fermentation is carried out in a two-stage are three types of toddy: sendi, obtained from wild date
process of which the first stage is aerobic mould fermenta- palm (Phoenix sylvestris); tari, from palmyra palm (Borassus
tion in a solid-state condition (Dung, 2004). The main flabellifer) and date palm (Phoenix dactylifera); and nareli
alcoholic fermentation occurs during the second stage after (nira), from coconut palm (Cocos nucifera) (Batra & Mill-
water has been added and lasts for approximately 5 days. ner, 1974). The toddy from coconut flower juice is known as
At the final stage, the wine is a turbid suspension of pink ra and panam culloo in Sri Lanka, nuoudua in Vietnam,
red colour, containing 814% (w/v) alcohol and some tuak (tuack) in Malaysia and Indonesia, and tuba in the
residual sugars. The wine may be clarified and/or strength- Philippines.
ened by blending with distilled alcohol, depending on local There is an art in binding the flower spathes, pounding
demand. them to cause the sap to flow properly by cutting the spathe

FEMS Yeast Res 6 (2006) 3039

c 2005 Federation of European Microbiological Societies
Published by Blackwell Publishing Ltd. All rights reserved
36
tip and collecting the sap into the earthen pitchers which
contain yeasts and bacteria in the left-over toddy from the
previous lots. The fermentation starts as soon as the sap
flows into the pitcher. Palm wine is either consumed fresh as
it is brought down from the tree or fermented for up to 24 h.
The freshly cut sap is generally a dirty brown sweet liquid
having 1018% w/w sugar, which after fermentation results
in the formation of a product containing as much as 9%
(volume in volume, v/v) ethanol (Steinkraus, 1996; Joshi
et al., 1999).
The palm wine fermentation is always an alcoholiclacti-
cacetic acid fermentation, involving mainly yeasts and
lactic acid bacteria. In the fermenting sap, S. cerevisiae is
invariably present but lactic acid bacteria such as Lactoba-
cillus plantarum, L. mesenteroides or other species of bacteria
like Zymomonas mobilis and Acetobacter spp. vary. The other
yeast types include Schizosaccharomyces pombe, Saccharo-
myces chevalieri, Saccharomyces exiguus, Candida spp. and
Saccharomycodes ludwigii in the samples of coconut palm
wine (toddy). Saccharomyces cerevisiae, Sch. pombe, Koda-
maea ohmeri and Hanseniaspora occidentalis are character-
ized as maximum ethanol producers in toddy (Joshi et al.,
1999). The yeasts, especially Saccharomyces spp., are largely
responsible for the characteristic aroma of palm wine
(Uzochukwu et al., 1999). During fermentation, there is
continuous effervescence as a result of the production of
carbon dioxide. A yeasty odour develops, and after a couple
of hours yeasts form a sediment at the bottom of the
container. The main ingredient of the fresh sap (pH 7.2)
is sucrose (1215%, w/w). During the first 24 h, more than
half of the total sugars are fermented. Palm wine is a good
source of B vitamins.
Kombucha
Kombucha from Central and East Asia is a beverage
obtained by fermentation of sweetened boiled tea with a
mixed culture of yeasts and acetic acid bacteria (Campbell-
Platt, 1987). Other names for kombucha, or tea fungus,
include fungus japonicus, tee kwass, tea kvass, cham- being more favourable for bacteria and winter for yeasts.
pignon de longue vie, Indo-Japanese tea fungus and
Manchurian mushroom. Kombucha is a symbiosis of
Acetobacter spp. mainly Acetobacter xylinum and various
yeasts. The mixed yeast-bacterial culture growing on sugary
tea extract accumulates lactic (0.1%), acetic (traces) and
gluconic (0.010.3%) acids, and some ethanol (0.3%). The
pH decreases steadily to about 2.5. The resulting beverage
also contains vitamins and minerals and is considered to be
a healthy product.
The yeast flora of commercial tea fungus includes the
genera Brettanomyces (56%), Zygosaccharomyces (29%) and
Saccharomyces (26%) (Mayser et al., 1995). Saccharomycodes
ludwigii and Candida kefyr (anamorph of Kluyveromyces
c 2005 Federation of European Microbiological Societies
Published by Blackwell Publishing Ltd. All rights reserved
K. E. Aidoo et al.
marxianus) in isolated cases, as well as a pellicle-forming
yeast, C. krusei and apiculate yeasts, Kloeckera spp. and
Hanseniaspora spp. have also been reported. However,
Zygosaccharomyces kombuchaensis is the dominant yeast
species now known to be commonly associated with kom-
bucha tea (Kurtzman et al., 2001). In another study of four
commercial kombucha products (Teoh et al., 2004), the
yeasts found included Brettanomyces bruxellensis (anamorph
of Dekkera bruxellensis), Candida stellata, Sch. pombe,
T. delbrueckii and Zygosaccharomyces bailii. Comparing
these findings, it appears that the fermentation is initiated
by osmotolerant yeasts and is then succeeded and ultimately
dominated by acid-tolerant species.
Condiments
Wadis
Wadis, traditionally consumed in Punjab and Bengal of
India, are now popular in many places of India, Pakistan
and Bangladesh. These dried, hollow, brittle cones or balls
(38 cm diameter, 1540 g in weight) (Fig. 1c) are used as a
spicy condiment or an adjunct for cooking vegetables, grain
legumes or rice.
To prepare wadi, dals, generally of blackgram, are soaked,
drained, ground into a smooth soft dough, left to ferment
for 13 days, moulded into cones or balls, deposited on
bamboo or palm mats smeared with oil, and sun-dried for
48 h. The surface of the cones or balls becomes covered
with a mucilaginous coating which helps to retain the gas
formed during their fermentation. The wadis look hollow,
with many air pockets and yeast spherules in the interior and
a characteristic surface crust.
Initially the microflora is diverse and contains lactic acid
bacteria, bacilli, flavobacteria and yeasts. Gradually,
L. mesenteroides, L. fermentum, S. cerevisiae and Tr. cuta-
neum become dominant. Candida vartiovaarae and K.
marxianus are also often found. The development and
prevalence of microflora are affected by the seasons, summer
The production of acid and gas results in a fall of pH from
5.6 to 3.2 and two-fold rise in the volume of the dough. The
lactic acid bacteria are mainly responsible for the acidifica-
tion of dough, favourable conditions for the yeasts to grow
and become active for leavening. The fermentation brings
about a significant increase in soluble solids, non-protein
nitrogen, soluble nitrogen, free amino acids, proteolytic
activity and B vitamins including thiamine, riboflavin and
cyanocobalamine. On the other hand, the levels of reducing
sugars and soluble protein decrease. Amylase activity in-
creases initially, but declines thereafter (Batra & Millner,
1974; Sandhu et al., 1986; Sandhu & Soni, 1989). Wadis
prepared by inoculating sterilized ingredients with a mixed
FEMS Yeast Res 6 (2006) 3039
Yeasts in Asian indigenous fermented foods
culture of C. krusei (anamorph of Issatchenkia orientalis)
and L. mesenteroides resemble the marketed ones. In con-
trast, the uninoculated controls were hard and compact and,
when broken, had a glistening surface (Batra & Millner,
1974).
Papads
Papad (papadam) is another important condiment or
savoury food in India, Pakistan and Bangladesh. This thin,
usually circular, wafer-like product (Fig. 1d) is used to
prepare curry or is eaten by itself as a crackly snack or
appetizer with meals after roasting or deep-frying in oil.
Papad-making under controlled conditions has already
developed into a cottage or small-scale industry.
Blackgram flour or a blend of blackgram with Bengal-
gram, lentil (Lens culinaris), redgram or greengram (Vigna
radiata) flour is hand-kneaded with a small quantity of
peanut oil, common salt (about 8%, w/w), papad khar
(saltworts produced by burning a variety of plant species, or
from very alkaline deposits in the soil) and water, and then
pounded into a stiff paste. The dough (sometimes with a
backslop and spices added) is left to ferment for 1-6 h. The
fermented dough is shaped into small balls which are rolled
into thin, circular flat sheets (1024 cm diameter,
0.21.2 mm thick) and generally dried in the shade to
1217% (w/w) moisture content. Candida krusei and
S. cerevisiae are involved in the preparation of papad
(Shurpalekar, 1986).
Soy sauces
Soy sauces are light to dark brown liquids with a meat-like
salty flavour used in cooking and as a table condiment.
Traditionally made in China, Japan, Korea, Thailand, the
Philippines, Indonesia and Malaysia, soy sauce is now also
produced in Europe and the Americas. There are two
specific fermentation stages involved in soy sauce produc-
tion: aerobic koji fermentation, which involves the use of A.
oryzae or Aspergillus sojae, and an anaerobic moromi or salt
mash, which undergoes lactic acid bacteria and yeast (Zygo-
saccharomyces rouxii) fermentations.
The two main groups of enzymes produced by A. oryzae
during koji fermentation are carbohydrases (a-amylases,
amyloglucosidase, maltase, sucrase, pectinase, b-galactosi-
dase, cellulase, hemicellulase and pentosan-degrading en-
zymes) and proteinases. Lipase activity has also been
reported. These major enzymes hydrolyze carbohydrates
and proteins to sugars and amino acids and low molecular
weight peptides, respectively. These soluble products are
essential for yeast and bacterial activities during moromi
fermentation (Aidoo et al., 1994; Chou & Rwan, 1995).
In this fermentation Tetragenococcus halophila initially
FEMS Yeast Res 6 (2006) 3039
37
proliferates and produces lactic acid, which lowers the pH
to 5.5 or less. Acid-tolerant dominant yeasts, notably Z.
rouxii, grow and produce about 3% (w/v) alcohol and
several compounds which add characteristic aroma to soy
sauce.
Although Z. rouxii is the dominant moromi yeast which
produces alcohol and several compounds that add charac-
teristic aromas to soy sauce, other yeasts such as Candida
versatilis and Candida etchellsii also produce phenolic com-
pounds, i.e. 4-ethylguaiacol and 4-ethylphenol, which con-
tribute to soy sauce aroma. Nearly 300 types of flavour
compounds have been identified in Japanese soy sauce
(Nunomura & Sasaki, 1992). Zygosaccharomyces rouxii pro-
duces flavour compounds including alcohols, glycerol,
esters, 4-hydroxy-5-methyl-3(3 H)-furanone (HMMF),
4-hydroxy-2 (or 5)-ethyl-5(or 2) -methyl-3(2 H)-furanone
(HEMF) and 4-hydroxy-2,5-dimethyl-3(2 H)-furanone
(HDMF). Of the furanones, HEMF produced by Z. rouxii
and Candida spp. gives Japanese-type soy sauce its
characteristic flavour (Hanya & Nakadai, 2003). This com-
pound is also reported to have antitumour and antioxidative
properties (Nagahara et al., 1992; Koga et al., 1998). Higher
alcohols such as isobutyl alcohol, isoamyl alcohol and
2-phenyl ethanol, produced by C. versatilis, are also impor-
tant flavour constituents of soy sauce. Certain strains
of yeasts have deleterious effects on soy sauce. Film-
forming yeasts, mainly belonging to the genera Zygosacchar-
omyces, Hansenula and Pichia, cause spoilage in moromi
fermentation.
Miso
Miso is a salty paste with a meat-like flavour made by
fermenting soybean, with or without the addition of rice or
barley, using A. oryzae and a yeast, Z. rouxii. Sometimes,
Tetragenococcus halophila and Enterococcus faecalis are also
involved in the fermentation. Miso is a seasoning agent and
is also used in the preparation of miso soup. Heat-treated
rice and/or soybeans are used to prepare shinshu or rice-
soybean miso. After the initial solid-substrate fermentation
dominated by A. oryzae, salt (38% of the original weight of
dry soybeans) is added to the koji and mixed thoroughly.
The mixture is backslopped or inoculated with Z. rouxii and
allowed to ferment for up to 15 days for sweet miso and
212 months for salty miso. Although other halophilic
yeasts such as Torulopsis versatilis may be present, only Z.
rouxii produces the desired metabolites for an acceptable
product (Ebine, 1989). Flavour components in miso are
similar to those of soy sauce. Furanones, HEMF and HDMF,
produced by Z. rouxii, have been identified as important
flavour components in miso. Miso also contains B vitamins
(riboflavin and cyanocobalamine) as a result of yeast
fermentation.
c2005 Federation of European Microbiological Societies
Published by Blackwell Publishing Ltd. All rights reserved
38
Future perspectives
Upgrading of traditional home-scale processes is needed so
that they can continue to maintain and strengthen the
cultural heritage and can compete successfully with im-
ported products. Whereas small-scale manufacture has the
advantages of short distribution lines, income generation for
families, etc., urbanization and the resulting growing de-
mand for ready-to-consume high-quality foods requires
larger-scale controlled industrial production. Examples of
industrialized traditional fermented foods (Steinkraus,
1989) are:
i alcoholic snacks such as tapai, which are now produced at
a small cottage scale in Malaysia using commercially
available pure culture starters of the starch-degrading
mould A. rouxii and the yeast Sm. fibuligera;
ii rice wines such as Japanese sake , using A. oryzae for rice
saccharification and sake yeasts (S. cerevisiae strains
selected for reduced foam production, or killer properties
if required);
iii condiments such as soy sauce inoculated with Z. rouxii
and Candida spp. and miso in which similar halotolerant
yeasts are used for flavour development.
Yeast products such as enzymes, B vitamins, trace ele-
ments (selenium, chromium), glycans, flavour components
and carotenoid pigments occur in traditional foods, but
could be exploited more effectively as purified substances
and food ingredients. Yeasts have a relatively high content of
protein, lipids and micronutrients. In view of the wide-
spread micronutrient deficiencies in regions that depend
predominantly on plant-based diets, the addition of yeast-
derived food products could contribute to improved nutri-
tional status (Mai et al., 2002).
In conclusion, a wide variety of yeasts are involved in
traditional fermented foods. Although the occurrence of
various yeasts has been reported, knowledge and under-
standing of their ecology including aspects such as microbial
successions and competitiveness, and of their genetic and
physiological properties remain to be acquired. In particu-
lar, yeasts that contribute to desirable product properties
require more precise characterization, using genomics,
proteomics and physiological approaches for more efficient
identification and exploitation, while developing consumer-
friendly strategies to control fermentations and safeguard
hygiene.
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c 2005 Federation of European Microbiological Societies
Published by Blackwell Publishing Ltd. All rights reserved