Plant growth-promoting yeasts

(PGPY) in sustainable agriculture

Under supervision :
Prof. Dr Mohamed El- Sayed Osman

Done by :
Nesma Mamdouh Abd El-Ghany

2023- 2024
 1. The potential of yeasts in promoting plant
growth

Yeasts are eukaryotic, unicellular microorganisms, classified under

the kingdom Fungi and grouped either in phylum Ascomycetes or
Basidiomycetes. The term ―yeast‖ has been derived from the Dutch
word gist and the German word gischt meaning to boil or bubble
[1].Yeasts are ubiquitous in occurrence, found almost everywhere in
the terrestrial (soil, vegetation) and aquatic (ocean, sea, estuary,
mangrove, lakes, and river) ecosystems. Around 100 genera and 800
species of yeasts have been described so far, which is a very small
percentage of yeast species that actually exist in nature, and the rest
are nonculturable forms [2]. Yeasts take part in a range of
ecologically significant processes via decomposition of plant
substrates, nutrient-recycling, biodegradation of hydrocarbons, and
microhabitat colonization. Yeasts have a symbiotic, parasitic, or
saprophytic association with their hosts. In recent years, they have
gained importance in the field of agriculture as agents of plant
growth promotion (PGP) and are aptly named plant growth-
promoting yeasts (PGPY). The PGPY are able to colonize plant
parts and benefit their hosts by modulating phytohormone
production, improving soil fertility, increasing nutrient availability,
enhancing resistance to abiotic stress, and inhibiting phytopathogens
There is a dire need to increase agricultural food production to meet
the food requirements of a rapidly growing population. This has led
to the extensive use of synthetic fertilizers and pesticides in
agriculture. This has negatively affected the soil microbiota, which
in turn has led to a reduction in agricultural production. Use of
chemical fertilizers and pesticides not only failed to produce the
desired results but also negatively impacted the environment. A
sustainable alternative to this problem is the use of plant growth
promoting microbes (PGPM). Plant growth-promoting bacteria
(PGPB) lead the list of PGPM as they possess the ability to increase
plant growth and protect plants from disease and abiotic stress. The
major bacterial candidates advocated for PGP include Azospirillum,
Bacillus, and Rhizobium. Though the role of PGPB in agriculture
was recognized quite early in science, the potential of yeasts in
promoting plant growth have been recognized only recently. As
yeasts are Generally Recognized as Safe (GRAS) for field
application, their use as PGP agents has received much attention
among researchers advocating sustainable agriculture. It is critical to
understand the plant growth-promoting traits of yeasts and their
interactions with plants for their successful application in the field.

2. Sources of PGPY and Isolation

Yeasts are a group of microbes with a cosmopolitan distribution in

aquatic and terrestrial ecosystems. Though yeasts may not be the
dominant group in the environment, they play a vital role in
biogeochemical cycling and ecosystem maintenance. PGPY are a
beneficial group of yeasts, and they have been mainly isolated from
the rhizosphere and phyllosphere of a wide variety of plants. In
order to increase the yield of economically important plants and
ensuring food security, most of the studies concentrated on the
yeasts associated with crop plants like maize [3], wheat [4], rice[5],
corn [6], black gram [7], kidney beans [8], tomatoes [9], grapes, and
pomegranate [7]. In addition to these, PGPY has also been
interestingly reported from the threatened, medicinal, insectivorous
plant Drosera [10] and from the marine environment [11]. For the
isolation of PGPY, researchers have mainly used sediment and root
samples from the rhizosphere and leaves from the phyllosphere.
Several workers have opted for surface disinfection of plant parts
using one of these agents namely: propylene oxide vapour, 70%
ethyl alcohol, Tween-40, or chloramines prior to isolation [12]. The
samples are ground, serially diluted, and plated on solid media.
Instead of direct plating of the sample, some researchers have
preferred to enrich and then plate it [10]. Yeast Peptone Dextrose
(YPD) medium is widely used [13]. After the isolation of PGPY,
the phytohormone production is ascertained by various biochemical
analyses and quantified by using LC–MS and HPLC. The media
employed for the detection of gibberellin production by yeasts,
including Potato Dextrose [14], GPI medium [15], Czapek’s-Dox
medium [16], GPII media [17], Yeast extract Peptone Dextrose
(YPD), and synthetic drop-out media (SC) [5], The YPD medium
supplemented with L-tryptophan is used for the detection of indole
acetic acid (IAA) production [18]. Pikovskaya’s agar, zinc oxide
(ZO) agar, and chrome azurol S (CAS) agar are widely used for the
detection of phosphate solubilization [5], zinc solubilizing [19], and
siderophore production [20], respectively.

3. Generic diversity of PGPY

PGPY mainly belong to the phylum Ascomycetes and the majority

of them are isolated from the rhizosphere. There are only limited
reports on PGPY from the phyllosphere. There are 23 genera of
yeasts reported to have plant growthpromoting capabilities and the
dominant ones are Candida spp. (15%), Rhodotorula spp. (14%),
Cryptococcus spp. (13%), and Saccharomyces sp. (9%) (Fig.1).
PGPY from the Ascomycetous phylum includes Aureobasidium sp.,
Metschnikowia sp., Candida spp., Meyerozyma sp., Pichia sp.,
Yarrowia sp., Williopsis sp., Debaromyces sp., Torulospora sp.,
Wickerhamomyces sp., Hyphopichia sp., Saccharomyces sp.,
Leucosporidium sp., Issatchenkia sp., and Kluyveromyces sp.
PGPY from the Basidiomycetous phylum includes Hannaella sp.,
Rhodotorula spp., Sporidiobolus sp., Trichosporon sp.,
Cryptococcus sp., Leucosporidium sp., and Rhodosporidium sp.
PGPY traits reported from yeasts include phytohormone production,
phosphate solubilization, siderophore production, increase tolerance
to abiotic stress, and plant pathogen inhibition. Some of the
pigmented yeasts demonstrating PGP activity are R. mucilaginosa,
R. paludegina, and A. pullulans. The dominant PGPY isolated from
the rhizosphere are Candida, Rhodotorula, and Cryptococcus, and
those from the phyllosphere are Psuedozyma and Leucosporidium.
Candida, Rhodotorula, and Cryptococcus are of cosmopolitan
distribution and their isolation is independent of the plant species
with which they are associated. In the case of marine isolates,
Rhodotorula is a major PGPY genus, followed by Kazhakistania
and Rhodosporidium is reported from the Bromeliad tank water.
Genetically improved Yarrowia lipolytica strain has also been
employed for PGP.

Figure 1. Generic diversity of PGPY.

 4. Plant growth-promoting traits of yeasts
Yeasts are as a source of phytohormones, enzymes, and amino
acids, which significantly promote plant growth and biomass
production even under conditions of water stress [21]. When used
with mineral fertilizers (NPK), the yeasts significantly increase leaf
number, leaf area, and chlorophyll concentrations [22]. PGPY are
capable of promoting plant growth, right from the tip of the root to
the topmost portion of the shoot (Table 1). It is evident from the
literature that, soil yeasts especially possess traits that enhance plant
growth, which include mineral solubilization, production of
phytohormone, siderophore, and enzymes, stimulation of
mycorrhizal root colonization, plant pathogen inhibition, and
enhancing soil fertility [23] (Fig. 2).

4.1. Phytohormones

Phytohormones are organic compounds that are responsible for

plant growth. The effect of phytohormone depends on the
inoculation methods and the abiotic factors. Phytohormones such as
abscisic acid (ABA), auxins, cytokinins, and gibberellins (GA)
control the growth and development of root, shoot, and fruit in a
plant. The effect of stress on the plant can be mitigated by these
phytohormones. The yeast cells when used in combination with the
phytohormones promote better plant growth than when either is
used alone [32].
Figure 2. Mechanism of PGP by PGPY.
 Table 1. Effect of PGPY on plants.

PGPY Phylum Source Effect on plant References

Plant
Hannaella Basidiomycete Leaves of Increased number lateral root [24]
coprosmaensis Drosera indica phytohormone production (IAA)
Inhibited fungal pathogens

A. pullulans Ascomycete Vineyard of Enhancement of shoot and root [25]

Pseudozyma aphidis Basidiomycete Grapes weight
R. dairenensis Basidiomycete Long root hair formation and
better branching

P. dianae Ascomycete Leaves of Increased lateral root number, leaf [10]

Sporidiobolus Basidiomycete Drosera number and primary root
ruineniae spatulata formation
increased stem length, increased
chlorophyll content

R. mucilaginosa Basidiomycete water of Red Antifungal activity [11]

sea Enhancement of shoot and root
growth

Rhodotorula sp. Basidiomycete Rhizosphere Gains fruit weight [26]

Tomato plants
M. guilliermondii Ascomycete Rhizosphere of Increase the nutrient uptake, [27]
M. caribbica Soil growth and yield.
R. mucilaginosa Basidiomycete

S. occidentalis Ascomycete Rhizosphere of Promote root growth and yield.in [28]

C. saturnus Soil plant
C. tropicalis
Saccharomyces sp., Ascomycete Rhizosphere Promote root growth in plant [29]
C. atropicalis Soil
A. pullulans
W. california
W. saturnus
R. mucilaginosa Basidiomycete
A. pullulans Ascomycete Rhizosphere of Enhance plant growth by [30]
Alfalfa, clover, producing
sweet clover, IAA, antifungal activities
soybean

S. cerevisiae Ascomycete Tomato Increase dry weight of shoots, [9]

crop growth, and nutrient supply
of sugarcane and tomato by N and
P nutrition

R. mucilaginosa Basidiomycete Phyllosphere Enhance plant growth by [31]

L. golubevii of Arabidopsis producing indolic compounds
Microbotryozyma
collariae

4.1.1. Abscisic acid

ABA is a sesquiterpenoid phytohormone involved in
phytobeneficial processes, such as seed dormancy, cell elongation,
and flower induction [33]. Currently, ABA from yeast is used in
agriculture to enhance the color of red table grapes, produce hybrid
seeds, and induce salt tolerance in citrus, drought tolerance in
chickpeas, and cold/drought tolerance in wheat [34]. Genetically
engineered oleaginous yeast Y. lipolytica is reported to be a
promising host for heterologous ABA production [33]. The
ameliorative effects of ABA and yeast on maize plants are reported
to reduce the deleterious effects of water deficit and enhance growth
and productivity [32].

4.1.2. Auxin

The phytohormone auxin, through its main representative IAA, a

derivative of an aromatic heterocyclic organic compound containing
a carboxymethyl substituent, stimulates rapid and sustained
response by plants, by promoting the elongation of lateral roots
[35]. Several microorganisms, including yeast are able to produce
IAA and significantly influence plant growth and development [36].
Levels of IAA production is found to vary with the yeast strains.
The IAA-producing yeast has recently received attention as a
favorable candidate for its use as a biofertilizer, as it facilitates the
uptake of nutrients from the surroundings to increase the root length
and surface area. The soil yeast Candida tropicalis HY (CtHY)
stimulated rice seedling growth [5]. Five genera namely
Cryptococcus flavus, Hannaella coprosmaensis, Pseudozyma
aphidis, Sporisorium reilianum, and Ustilago esculenta of the
phylum Basidiomycota isolated from green and undamaged
insectivorous Drosera indica plant leaves, possessed the capacity
for IAA-production [24]. Aureobasidium pullulans isolated from the
phyllosphere and rhizosphere of the medicinally important plant,
Drosera spathulata exhibited the production of indole-3-acetic acid,
ammonia, and 26 polyamines in addition to the ability of
solubilizing calcium phosphate and zinc oxide and catalase enzyme
activity [10]. The IAA produced by the yeasts Torulaspora globosa,
Meyerozyma guilliermondii, and Rhodotorula mucilaginosa
promoted the development of tomato seedlings [37]. The
inoculation of seeds and seedlings of lettuce, cv. Crocantela, with a
rhizosphere yeast T. globosa reportedly increased the root dry mass
production under field conditions [38]. The inoculation methods and
formulation standardized are expected to promote the growth and
development of other plants as well as seeds [38].

4.1.3. Cytokinins

Cytokinins are a class of phytohormones which is a N6 substituted

adenine derivative, which influences all aspects of plant growth,
including cell division, cytokinesis, apical dominance, axillary bud
growth, photomorphogenic development, and leaf senescence [39].
Zeatin is one of the most common adenine-derived cytokinins
synthesized in plant roots [40]. Kluyveromyces lactis,
Schizosaccharomyces pombe, and Saccharomyces cerevisiae
synthesize cytokinins [41]. The diversity of yeast species, that
produce cytokinins is largely unknown. Metschnikowia pulcherrima
and Rodotorula mucilaginosa have been found to synthesize zeatin
in their exponential growth phase unlike bacteria [42].
Moesziomyces antarcticus apart from being a commercial producer
of enzymes and glycolipid surfactants, are also capable of
synthesizing zeatin [43].

4.1.4. Gibberellins

GA are weakly acidic growth hormones involved in stem

elongation, germination, flowering, and senescence etc [44].
Yarrowia lipolytica olegaceous yeast has been reported to produce
GA either by genetically engineering (by the recombination of
expressing tesA gene) or by pretreatment of cells [45].
Phosphate solubilization Phosphate is a limiting nutrient for plant
growth due to its limited availability and mobility in soil. Its
deficiency leads to purple spots on leaves and stems or inhibition of
development and maturation. Several groups of microorganisms
such as actinomycetes, bacteria, fungi, and yeast are known as
efficient P solubilizers. Phosphate solubilizing microorganisms can
play a significant role in dissolving both fertilizer phosphorus and
bound phosphorus in the soil in an ecofriendly and sustainable
manner [46]. The yeast species that are capable of solubilizing
phosphate are Schwanniomyces occidentalis, Oideodendron sp.,
Pseudonymnoascus sp., Candida tropicalis, Geotrichum candidum,
Geotrichum capitatum, Rhodotorula minuta, Rhodotorula rubra,
Saccharomyces sp., Hansenula sp., Klockera sp., and
Debaryomyces spp. [47]. A field study on the phosphate
solubilization capacity of M. guilliermondii CC1 has proved to
reduce the essential use of chemical fertilizers without affecting the
optimal productivity in maize (Zea maize L.) [48]. A study from the
rhizosphere soil in the region of east and west Gojam, Ethiopia has
identified nine yeast species with phosphate solubilizing ability and
these are Pichia norvegensis, Cryptococcus albidus var aerius,
Candida etchellsii, Cryptococcus albidus var albidus, Rhodotrula
aurantiaca, Cryptococcus luteolus, Cryptococcus albidus var
diffluens, Cryptococcus terreus [49]. Arabidopsis thaliana
inoculated with selected yeast Cryptococcus laurentii promoted
plant growth in inorganic phosphate (Pi)-deficient medium
supplemented with calcium phosphate dibasic dihydrate. The higher
levels of cellular inorganic phosphate noted in plants treated with
yeast reveals its potential for use as a commercial biofertilizer [50].

4.2. Plant pathogen inhibition

Plant diseases caused by fungal pathogens are responsible for major

crop losses world over, with a significant socioeconomic impact.
The fungal pathogens are increasingly recognized as a global threat
to our agriculture. The usage of chemical fungicides leads to
environmental pollution, has an adverse effect on human health, and
diminishes soil biodiversity. To overcome this hurdle, yeasts are
used as one of the microbial tools to control fungal pathogens as the
yeasts are GRAS for field application. The antagonistic action of
yeast against plant pathogens can be due to different mechanisms
like colonization, production of antifungal compounds, induced
systemic resistance, and mycoparasitism process [51]. As yeasts
grow much faster than filamentous fungi, they pose a competition,
thereby reducing the availability of nutrients to fungi and quickly
colonizing the plant niches [52]. A phyto-beneficial attribute of
helpful microorganisms is the release of volatile organic compounds
(VOCs) like alcohols, esters, aldehydes, ketones, and lactones,
which increases disease resistance and abiotic stress tolerance.
VOCs emitted by yeasts strains Pichia kudriavzevii, P. occidentalis,
and Meyerozyma guilliermondii in- hibit common plant pathogens
representing Mucor spp., Penicillium spp., Aspergillus sp.,
Fusarium spp., as well as Botrytis cinerea [53]. ABA can act as a
potential antifungal agent by its inhibitory effect on
chorismatemutase through the reduction of phenyl pyruvate
production, an important metabolite in beta-glucan synthesis [54].
The Killer Toxins produced by the yeast strains can damage the cell
wall and cell membrane by triggering apoptosis and beta-glucan
synthesis inhibition [55]. Yeast is also known as a potential
biocontrol agent of Cacao witches broom disease caused by
Moniliophthora perniciosa [55].
Yeasts that can trigger antifungal activity include Pichia kluyveri, P.
membranifaciens, P. farinosa, P. acacia, Zygosaccharomyces bailii,
Hansenula mrakii, Wickerhamomyces anomalus, Williopsis
saturnus, Kluyveromyces phaffii, K. lactis, and Wingearo bertsiae
[55].Saccharomyces and non-Saccharomyces yeasts such as
Trichoderma harzianum, Pichia membranifaciens, Kloeckera
apiculata, Candida guilliermondii, Cryptococcus laurentii,
Cryptococcus flavus, Cryptococcus albicus, Candida pyralidae,
Pichia kluyveri, and P. expansum are potential antagonists against
phytopathogenic fungi Penicillium, Aspergillus, and Botrytis of
table grapes, wine grapes, and raisins [56].

4.3. Siderophore production

Siderophore is a low molecular weight, high affinity ferric iron
chelating compound secreted by organisms. Siderophore producing
microorganisms are important for enhancing metal tolerance in
plants by decreasing the availability and toxicity of metals.
Pseudozyma aphidis is known to be the highest in siderophore
production among the 13 species of yeasts studied [10]. Endophytic
yeasts such as R. graminis and Cryptococcus sp. exhibit PGP by the
production of siderophores through the assimilation of sugars and
amino acids produced from plants [57]. Phytochemical constituent
and antioxidant activity of Saccharomyces cerevisiae promote the
plant growth in Sorghum bicolor and Arachis hypogea [58].
Trichosporon ovoides and Saccharomyces cerevisiae are shown to
exhibit a high production of siderophore and sidD gene expression
under different levels of Cd2+ and Pb2+ toxicity stress [59].

4.4. Stress response

The growth and yield of plants are seriously affected by

environmental stress. The conditions of stresses such as drought,
salinity, and nutrient depletion lead to a decrease in chlorophyll
content and finally cell death. Under situations of drought and
salinity, the yeasts are capable of increasing water content and
promoting growth by reducing oxidative stress. ABA apart from its
role as phytohormone, it also helps to tolerate drought conditions by
closing stomata, accumulating glycine betaine as well as enhancing
relative water content [60]. The beneficial effects of the
combination of ABA and yeasts on yield in maize, faba bean,
wheat, and sugar beet plants are attributed to their ability to improve
water content, enzymes activity, and photosynthetic rate as well as
amino acid concentration [61]. Under conditions of stress induced
synthesis of defence-related enzymes such as peroxidases,
chitinases, catalase, and polyphenol oxidases by the yeast strains
Pichia kudriavzevii POY5 and Issatchenkia terricola GRY4 are
reported to improve the seed germination of black gram Vigna
mungo L. [7]. The role of yeast Saccharomyces cerevisiae as
biofertilizer is well established for its plant growth enhancement,
phytochemical constituent changes, and increased antioxidant
activity in Sorghum bicolor and Arachis hypogeal [58].

4.5. Nitrogen fixation

During nitrogen fixation, an oxygen sensitive process, atmospheric
nitrogen is assimilated as ammonia and other nitrogenous
compounds by the metalloenzyme nitrogenase. Early research
indicated that the ability of N2 fixation was restricted to a selected
group of microorganisms mainly bacteria and doubted the capability
of eukaryotic cell to fix nitrogen. However, some workers have
explored the nitrogen fixing power of certain yeast groups. Reports
have claimed that many yeast genera like Torula, Pullularia,
Saccharomyces spp., Candida, Rhodotorula, and Lipomyces have
the ability to accumulate ammonium in nitrogen fixing yeast broth,
indicating their potential to fix N2 [62]. However, these claims are
not yet been substantiated and still remains disputed. There are
works that suggest that though yeast may not fix nitrogen on their
own, they may do so in syntrophic association with bacteria like
Clostridium or even stimulate the nitrogen fixation by Azotobacter
and Beijerinckia. To check the expression of prokaryotic nif genes
(nifH, nifD, nifJ, nifK, nifU, nifS, nifM, nifE, nifN, nifB, nifV, nifX,
hesA, groES, and groEL) in eukaryotic system, Saccharomyces
cerevisiae has been widely used as a model organism due to its
inherent safety and ease of genetic manipulation. These studies have
opened up the possibility of using genetic engineered yeast in PGP.

5. Methods adopted for inoculation of PGPY

The inoculation methods adopted to introduce the PGPY will

influence the establishment and persistence of yeast population
either in plant parts or rhizosphere. Combined or separate inoculum
of yeast species can generate plant promoting activity [63].The
inoculation method selected is usually on the basis of the host plant
and inoculum. There are different inoculation methods for PGPY
such as seed, root, soil, and foliar inoculation. The seed inoculation
method is the most useful method. In the case of soil inoculation,
the application of PGPY is done as close as possible to the
rhizosphere. In this regard, binders can help to bind PGPY on to
the substrate and co-inoculation can increase the efficiency of
inoculation methods [64]. On co-inoculation of yeast with other
microbes synergistically, they interact with each other and promote
plant growth. Most of the studies carried out to determine the ideal
method for inoculation of PGPY are in-vitro or in pot culture. There
are only negligible field trail reports using PGPY. The most often
adopted method for in-vitro studies is cocultivation, which involves
cultivating selected plants along with yeast species. Co-cultivation
of Nicotiana benthamiana seedlings with the yeast Sporidiobolus
ruineniae was shown to result in good development of lateral roots
and root hairs [10]. In-vitro analysis of co-cultivation of
Arabidopsis thaliana with Cryptococcus laurentii revealed the PGP
on agar plates filled with an inorganic phosphate (Pi)-deficient
medium supplemented with calcium phosphate dibasic dehydrate
[50]. Seed inoculation is generally carried out in the sterile plastic
bags, into which the seed, inoculum, and binder are added and
mixed. This helps in the uniform distribution of inoculum onto the
seeds. The seed germination process releases many exudates into
the soil and yeast present on the inoculated seeds use these exudates
as a nutritional source and colonize roots [65], which in turn aids
plant growth. Enhanced plant growth in maize is observed on
inoculation of its seeds with W. saturnus and R. glutinis [3].
The phosphate solubilization efficiency of yeast inoculated
Arabidopsis thaliana varies with the yeast strains used in the
inoculum, i.e. seeds coated with M. caribbica (JYC358), Candida
sp. (JYC363), and A. pullulans (JYC375) exhibit a high phosphate-
solubilizing ability, whereas Torulaspora sp. (JYC369), C. laurentii
(JYC370), and Pseudozyma sp. (JYC372) show low phosphate-
solubilization [50]. In another study, both seed inoculation and the
inoculation of PGPY near the root tips displayed a good shoot
growth in terms of stem elongation and leaf number [24].The seeds
of Lettuce (Lactuca sativa) inoculated with T. globosa (5S55) under
greenhouse condition is shown to enhance the seedling root–shoot
length and dry weight [38].The pot culture assay with black gram
seed inoculated with Issatchenkia terricola and Pichia kudriavzevii
is reported to have increased growth rate[7]. Thus, seed inoculation
was found to be an efficient method of introducing PGPY into the
field. Spraying is another method of PGPY inoculation. Spraying
of T. globosa inoculums near the root has improved the
development of tomato seedlings under greenhouse conditions [37].
This mode of inoculation helps the yeasts to easily colonize the
plant roots. Foliar application of active dry yeast has also increased
growth characters and yield of common bean, and the application of
yeast combined with mineral fertilizers (NPK) has significantly
increased leaf number, area, and chlorophyll concentrations in sugar
beet plants [61].However, the foliar inoculation is the least used
method. The pruned-root dip method by [66] has suggested the use
of healthy surface disinfested pregerminated seeds for root tip
cutting and dipping purpose. In this method of inoculation, the roots
of germinated seedlings are cut aseptically and dipped into the
inoculum. Choice of the inoculation method is an important factor
that determines the effect of the plant growth-promoting traits of
yeasts on host plants. The ideal method would be seed and/or soil
inoculation. The selection of the inoculation method must be based
on sound knowledge of the plant growth stages and species. The
successful inoculation also depends on the environmental
conditions, quality and quantity of the inoculum, and microbial
adaptations to the new environment.

6. Yeast as biofertilizer
Microbial biofertilizers are able to increase yield and quality of
crops without a large investment of money and labour, this has led
to increase in their popularity globally. It is clear that several yeast
genera exhibit phyto-beneficial traits and that has made them gain
entry in to the list of microbial agents suitable for application as
biofertilizers. Yeast like Candida, Geotrichum, Rhodotorula,
Saccharomyces, and Williopsis have been identified for use in
commercial biofertilizers as they are able to decompose plant
residues and release essential elements such as N, P, and K. When
yeasts cocultivated with other PGPB, they were found to enhance
their ability to promote cell activation and root division. The use of
yeast as a biofertilizer in agriculture has received considerable
attention because as they are generally recognized as safe for use in
the field and their bioactivity [67].

7. Agricultural application of yeast

Use of yeast in the industry is well known, but the
implementation in agriculture is not very common. Yeast has
been used to produce some alcoholic beverages and glycerol, and
frequent use of Saccharomyces cerevisiae makes it most
domesticated organism. But now, yeast is one of the most
important microorganisms that can be used in different areas,
including food industry, medical, agriculture, biotechnological
industry and bio-fuel production industry. Yeast contains a large
chromosome that will have a lot of functional genes that may
help in pesticide degradation and induce plant growth by
secretions of growth-regulating hormones. Till now, the study
shows that plant growth induced either direct and/or indirect
mechanisms by the bio-agents. Some literature indicates that
yeast has these two properties which help to promote the plant
growth attributes [69].
In the direct mechanism, plant growth depends on the production
of different types of hormones and enzyme that directly influence
the plant growth attributes and productivity. Besides IAA and
cytokinin and other hormones, plants required some micro and
macronutrients such as phosphate, potassium, zinc, manganese and
iron. The role of yeast on plant growth is still not very clear, but
studies suggested that yeast can be a potential PGP bio-agent. It has
been reported that yeast produces indole-3-acetic acid (IAA) and
indole-3- pyruvic acid (IPYA). These compounds are major plant
growth hormones produced by endophytic yeast (Williopsis
saturninus) isolated from the root of maize [3]. Cytokinins are other
hormones that help directly to promote plant growth by facilitated
cellular division. Zeatin is the most common type of cytokinin that
promotes plant cell growth. The yeast isolated from rhizosphere
such as Sporobolomyces roseus KBPY-5472 and Sporobolomyces
roseus KBP Y-5432, Metschnikowia pulcherrima KBP Y- 6020 and
Aureobasidium pullulans KBP Y-5404 were reported to synthesized
zeatin (another form of cytokinin) and help in plant growth [69].
Both, phosphate and potassium (P. anomala KSY29, R. glutinis
KSY33) solubilization ability was also observed in yeast [6].
Polyamines are the natural components that are involved in the plant
growth via germination, dormancy breaking, cell division, fruit set,
growth and ripening of fruit and growth, and the most important
role of polyamine is helping to induce the resistance against abiotic
stress mainly chilling injury. Studies showed that C. tropicalis HY,
Hansenula saturnus and Issatchenkia occidentalis produce different
types of polyamine (spermidine, putrescine, spermine etc.) and
protect the plants in different ways, and it also shows high ACC
deaminase activity, which leads to enhancing plant growth by
lowering the levels ethylene production [5].
Another mechanism to develop plant growth by protecting them
from both biotic and abiotic stress is known as an indirect
mechanism of plant growth. Some studies showed that yeast has the
property to inhibit the potential pathogen and induce the stress
tolerance ability of plant [70]. Protein analysis of P.
membranaefaciens indicated that it induces the level of different
antioxidant components (catalase, peroxiredoxin, glutathione
peroxidize, peroxidizes and superoxide dismutase) in peach and
grapefruit; this antioxidant enzyme helps to resist from different
stress [71]. It has been reported that Rhodotorula glutinis, Candida
valida and Trichosporon asahii possess a significant biocontrol
activity against soil-borne fungal plant pathogen Rhizoctonia solani
that causes root diseases [72]. Rhodotorula glutinis helps indirectly
to promote plant growth by showing antagonistic effect against
pathogen Penicillium expansum by producing siderophores
[73].Another yeast A. pullulans is used to promote plant growth and
suppress growth fungal plant pathogens G. cingulated [74]. Some
yeast strains R. mucilaginosa (KKU-M12c), C. albidus (KKU-
M13c), P. membranifaciens (KKU-M18c), H. uvarum (KKU-M19c)
and C. alifornica (KKU-M20c) showed protease enzyme activity
which plays a major role in plant defence by killing pathogen and
supplying protein-degrading components [75]. And also, these
strains are isolated from natural environmental samples. Recently,
Some yeast strains namely Rhodotorula lactose (KKU-A5), R.
nymphaeae (KKU-A6), R. graminis (KKU-A12), A. minuta (KKU-
A20), Exophiala dermatitidis (KKU-A23), Candida davisiana
(KKU-A24), R. slooffiae (KKU-A26), R. mucilaginosa (KKU-A29)
and Rhodosporidium diobovatum (KKU-A38) have the ability to be
applicable for degradation of petroleum components like toluene,
benzene and xylene compounds from the environmental soil as sole
source of carbon for growth and development.
S. cerevisiae synthesizes three important cellulose-degrading
enzymes like cellobiohydrolase, endoglucanase and β-glucosidase
on their cell surface. So that this may be useful for cellulose
degradation of crop residues in soils to enhance the organic content
in soils [76]. Several scientific studies suggested that the bio-agents
(bacteria, fungi) induce plant growth either by providing essential
nutrients and hormones or by inducing stress resistance by directly
killing the pathogen or inducing plant immunity through ISR/SAR.
These characteristics are present in the yeast that induces plant
growth. Themost important and challenging task for agricultural
crops is to maintain the crops for long time after harvesting.
Because, post harvesting protection of crops is very important. A lot
of microorganisms including yeast have been used for a long time
in laboratory and some commercial sector as an antagonist’s agent
against different pathogen [77].
Fig. 3 Diagrammatic presentation of isolation and application of beneficial
yeast in different aspect of sustainable agriculture
Two different groups of microbes can be used as post-harvest
disease-managing bio-agent. The first group is where microbes can
be found naturally in the fruits or vegetables. The second group is
artificial mechanical introduction of microbes to the host that has
pathogen-controlling ability. Among all the microbes,
Saccharomyces cerevisiae is the most common and helpful bioagent
for post-harvest disease control of agricultural products.
Saccharomyces cerevisiae is the major and dominating microbe in
the microbial community that can easily be found on the leaf, fruit
and vegetables surfaces [78].
Yeasts will be an effective biocontrolling agent, because yeasts
genotypically and phenotypically are more suitable to the particular
environment and are well skilled in colonization and competition
for space and nutrients than other groups of microbes on the surface
of fruit and vegetables [79].
In the postharvest disease control property, yeasts use some
specific property such as mycoparasitism, release some lytic
enzyme and compete for nutrients and space, altering the pH to
reduce the growth of the pathogen on fruit surface [77]. Among all
different types of property, competition for the nutrient and space
by yeast is the major factor which suppresses the growth of
pathogen on fruits and vegetables. Commercial synthesis of a
potential biocontrol agent needs to maintain efficacy and improve
shelf life for long-term preservation. Scientist developed liquid and
solid bio-formulations of yeast as potential strains or their
secondary metabolites for post-harvest disease management in
agriculture [80]. Both solid and liquid bioformulations of yeast have
some advantage like solid bioformulation can be stored for a long
term without cross contamination without refrigerator, and it can
transport easily while in the case of liquid bioformulation, it is cost
effective and has more cell count than solid bioformulation [80].

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