Environ Monit Assess (2023) 195:1252

https://doi.org/10.1007/s10661-023-11863-w

RESEARCH

Yeast‑driven valorization of agro‑industrial wastewater:

an overview
Ndubuisi Ifeanyi Amara · Enemour Simeon Chukwuemeka ·
Nnabuife Obianuju Obiajulu · Ogbonna James Chukwuma

Received: 12 June 2023 / Accepted: 11 September 2023 / Published online: 28 September 2023
© The Author(s), under exclusive licence to Springer Nature Switzerland AG 2023

Abstract The intensive industrial and agricultural In addition to remediating the wastewater, numerous
activities currently on-going worldwide to feed the value-added products such as single cell oil (SCO),
growing human population have led to significant single cell proteins (SCPs), biofuels, organic acid, and
increase in the amount of wastewater produced. These aromatic compounds amongst others can be produced
effluents are high in phosphorus (P), nitrogen (N), through fermentation of wastewater by yeast cells.
chemical oxygen demand (COD), biochemical oxygen This work thus brings to limelight the potential roles
demand (BOD), and heavy metals. These compounds of yeast cells in reducing, reusing, and recycling of
can provoke imbalance in the ecosystem with grievous agro-industrial wastewaters while proffering solutions
consequences to both the environment and humans. to some of the factors that limit yeast-mediated waste-
Adequate treatment of these wastewaters is there- water valorization.
fore of utmost importance to humanity. This can be
achieved through valorization of these waste streams, Keywords Valorization · Wastewater · Yeast ·
which is based on biorefinery idea and concept of Remediation · Biorefinery
reduce, reuse, and recycle for sustainable circular
economy. This concept uses innovative processes
to produce value-added products from waste such as Introduction
wastewater. Yeast-based wastewater treatment is cur-
rently on the rise given to the many characteristics of Environmental pollution is an issue of major concern
yeast cells. Yeasts are generally fast growing, and they with profound impact on human health and ecosys-
are robust in terms of tolerance to stress and inhibi- tem. Rapid urbanization, industrialization, and popu-
tory compounds, in addition to their ability to metabo- lation growth are aggravating the pollution-related
lize a diverse range of substrates and create a diverse issues. This already bad situation is being exacerbated
range of metabolites. Therefore, yeast cells possess by poor waste and effluent management from house-
the capacity to recover and transform agro-industrial holds, industries, and agricultural fields, thus causing
wastewater nutrients into highly valuable metabolites. further damage to the already crippling ecosystem.
Aside from its effect on public health and the econ-
omy, environmental pollution is a serious threat to
N. I. Amara · E. S. Chukwuemeka · N. O. Obiajulu (*) · food and portable water security, as well as biodiver-
O. J. Chukwuma sity (Akhtar & Mannan, 2020).
Department of Microbiology, University of Nigeria,
Wastewaters are water that has been contami-
Nsukka, Enugu State, Nigeria
e-mail: [email protected] nated by human activities. The three major sources of

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wastewater are domestic, agricultural, and industrial by using either native isolates from the waste stream or
activities. These processes require enormous amount metabolically engineered hosts that use enzymes from
of good-quality water but ultimately generate large these isolates. Whatever the strategy employed, micro-
quantities of wastewaters, which are streamed directly bial fermentation generates high-value products while
with little or no treatment into large water bodies or detoxifying wastewaters, making it a critical technol-
surroundings, thereby polluting them. Wastewater is ogy for waste valorization and achieving a circular
a complex stream made up of about 99.9% of water economy (Lad et al., 2022).
and 0.1% of suspended solids (350–1200 mg/L), Yeasts are generally fast growing and exhibit
organic compounds such as body waste (faeces, toilet remarkable tolerance to low pH, high salinity, high
paper, food waste), dissolved biodegradable organics content of organics, antibiotics, heavy metals, and
(proteins, carbohydrates, and lipids), inorganic solids other inhibitors. They can metabolize a wide range of
(sediment soil, salts, and metals), micro-pollutants, substrates, including sugars (glucose, sucrose lactose,
nutrients, heavy metals, and diverse microorgan- fructose, and maltose), biopolymers (starch, cellulose,
isms (about ­109 cells/mL), with high chemical oxy- hemicellulose, and pectin), pentoses, methanol, alco-
gen demand (COD) and biochemical oxygen demand hols, polyols, hydrocarbons, fatty acids, and organic
(BOD) (Jain et al., 2021). acids (Yang & Zheng, 2014). Aside from utilizing
Owing to their harmful nature, wastewaters must soluble organic compounds, most yeast can readily
first be treated accordingly before reuse or return to the absorb ammonium ions, urea, inorganic phosphates,
environment (Jain et al., 2021). Wastewater treatment and sulphates. Therefore, treatment of these wastewa-
is a process of reducing the contaminant in wastewa- ters which are high in oil, salinity, and other organic
ter to a harmless level in accordance with acceptable compounds with yeast cells seems to be a suitable
standards before reuse or return to the environment and low cost approach. This work will thus highlight
(Jain et al., 2021). Currently, there are various meth- the sources and impact of these wastewaters on the
ods including ponding, aerobic digestion, anaerobic environment. It will also highlight the potential value-
digestion, and physical, chemical, and physicochemical added products that can be generated from these
methods available for wastewater treatment (Iwuagwu wastewaters. Finally, the work will bring to limelight
& Ugwuanyi, 2014; Mohammad et al., 2021; Nwuche the potential roles of yeast cells in reducing, reusing,
et al., 2014). In addition to being expensive and time- and recycling of wastewaters while proffering solu-
consuming, these methods entails disposing these tions to some of the factors that limit yeast-mediated
wastewaters without further exploitation for value- wastewater valorization.
added products. Over the years, a consolidated form
of waste management for the exploitation of these low-
cost feedstocks which revolves around the biorefinery Wastewater: sources, composition, and impact
concept and idea has been postulated for sustainable on the environment
economy. Valorization and recycling are novel con-
cepts that are becoming increasingly important on a Due to the harmful substance present in these waste-
worldwide scale. By-products, wastes, and effluents waters and their uncontrolled disposal, they pose
from agricultural and industrial processes typically serious threat to human health and the ecosystem
contain high levels of sugars, lipids, and proteins, as (Fig. 1). The composition and environmental impact
well as distinct aromatic and aliphatic compounds; of some selected wastewater are discussed below.
thus, they could be regarded as low-cost and readily
available resources for the synthesis of value-added Oil‑rich wastewater
chemicals and biomaterials (Ahmed et al., 2019; San-
tos et al., 2019). Given the complexity of wastewaters, These are oil-containing wastewaters generated by oil
valorization approach involving microbial fermenta- manufacturing industries. During the production of edi-
tion is preferable. More specifically, microorganisms ble oil, a water washing process involving the separation
can metabolize structurally distinct compounds under of a substantial quantity of fatty acid in raw vegetable
mild conditions and channel them to a single product oil from glyceride generates an oil-rich industrial waste-
(Lad et al., 2022). Host selection can be accomplished water (Yang & Zheng, 2014). Oil-rich wastewater can

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Fig. 1  Impact of untreated

and indiscriminate disposal
of wastewater

also be generated from other sources such as slaughter are lignin-like structure constituting the most recalci-
house, dairy manufacturing plant, and tanneries (Yang trant fraction of this effluent (Ahmed et al., 2019). As a
& Zheng, 2014). The two major contributors to this oil- result of their high toxicity and low pH, untreated OMW
rich wastewater are palm oil mill effluent (POME) and is harmful to sewage treatment plants and pose serious
olive oil mill wastewater (OMW). ecological issue when discarded to the environment
(Carrara et al., 2021; Rytwo et al., 2013). Beside its
Olive oil mill wastewater (OMW) high-polluting power, OMW typically exhibits profound
levels of phytotoxic and antibacterial activity owing to
During extraction of oil from olives, water is added to the presence of various phenolic compounds, making
obtain initial separation of the oil and the pomace. This it tough to remediate using biological technologies and
generates a three-layer fraction made up of oil, pomace, indigenous soil microbial communities (Ahmed et al.,
and a heavily polluted liquid waste, known as OMW, 2019; Ouzounidou et al., 2012). Dumping of untreated
that is rich in carbon and other valuable nutrients but at OMW in the environment causes severe environmental
the same time containing toxic compounds especially hazards including surface and ground water contamina-
phenols. OMW has variable physicochemical properties tion, changing the colour of natural water bodies, toxic-
which are influenced by factors such as the geographic ity to aquatic life, altering of soil quality, cytotoxicity,
location of the olives, prevailing climate conditions, phytotoxicity, and emission of strong offensive odour
the type of olive, the level of maturity, cultivation/pro- (Carrara et al., 2021; Dermeche et al., 2013).
cessing procedures, method of extraction, and process-
ing (batch or continuous) applied in obtaining the oil Palm oil mill effluent (POME)
(Ahmed et al., 2019). Also known as “alpechin”, OMW
is generally high in CODs and BODs, in addition to POME is the voluminous waste recovered from palm
elevated levels of total phenols, tannins, carbohydrates, oil milling processes. For every tonne of crude palm oil
pectins, polysaccharides, polyalcohols, fatty acids, produced, approximately 2.5 to 3.5 tonnes of POME
organic matter, and minerals, such as potassium and are generated (Madaki & Seng, 2013). POME is typi-
magnesium (Ahmed et al., 2019; Carrara et al., 2021). cally derived from three sources: clarification wastewa-
Ordinarily, OMW is brownish-black in appearance ter, sterilizer condensate, and hydro-cyclone wastewa-
resulting from the presence of polymeric phenols which ter (Dominic & Baidurah, 2022; Karim et al., 2021a, b;

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Mohammad et al., 2021). POME is a viscous liquid with of HCW is cassava mill wastewater (CMW). CMW
a characteristic brownish appearance attributed to its ful- is one of three major wastes generated during cassava
vic acid-like and humic acid content (Dominic & Baidu- processing. This effluent is a colloidal suspension of
rah, 2022). This brownish colour can also be attributed to fine cassava (starch) particles in water (Izah, 2019).
carotene (8 mg/L), pectin (3400 mg/L), tannin, phenolic Besides the high moisture content nature of cassava
(5800 mg/L), and 4700 mg/L of lignin (Mohammed & root, large volume of water is utilized during process-
Chong, 2014). POME is classified as a low-pH, high- ing of the roots to edible products thus generating high
strength pollutant as a result of the organic and free fatty loads of wastewater. It estimated that around 250 and
acids produced during the partial decomposition of palm 600 kg of wastewater is produced for every 1000 kg of
fruits prior to processing (Iwuagwu & Ugwuanyi, 2014). cassava processed (Oghenejoboh et al., 2021). CMW
POME is a colloidal suspension that is composed of is very acidic in nature, with a high concentration of
90–96% water, 2–4% suspended particles, and 0.6–0.7% organic materials, suspended solids, sulphur dioxide,
oils (Mohammad et al., 2021). It has also been reported and cyanide (Izah et al., 2018). CMWs have been
that in addition to other nutrients, POME is rich in miner- shown to have low pH (2.5–5.07), low dissolved oxy-
als, trace elements, and heavy metals. Although the prop- gen (1.10–2.60 mg/L), and high COD in the range of
erties of POME are determined by the raw material qual- 320–365 mg/L (Izah et al., 2015; Rim-Rukeh, 2012).
ity and the manufacturing procedures, it is generally low This effluent also has total dissolved solids of about
in pH (3.4–5.5) and high in COD (50,000 to > 100,000 mg 799 mg/L, total suspended solids of 789 mg/L, total
­L−1) and BOD (25,000–54,000 mg ­L−1) (Dominic & Bai- solids of 5600 mg/L, total hardness of 75.00 mg/L,
durah, 2022; Islam et al., 2018; Iwuagwu & Ugwuanyi, and a conductivity 1550 S/cm (Olorunfemi & Lolodi,
2014; Karim et al., 2021a, b; Mohammad et al., 2021; 2011; Orhue et al., 2014; Patrick et al., 2011). In
Nwuche et al., 2014). In addition, POME also contains addition to these, this effluent also contains alkaline
40,500–75000 mg ­L−1 of total solids, 2000–8300 mg ­L−1 earth metals and heavy metals (Izah et al., 2017). In
of oil and grease, 18,000–47,000 mg ­L−1 of suspended developing countries like Nigeria, this wastewater is
solids, total nitrogen of 400–800 mg ­L−1, and an ash con- ordinarily discharged into the environment (soil and
tent of approximately 3000–42,000 mg L ­ −1 (Dominic & water) without any prior treatment. CMW alters the
Baidurah, 2022; Iwuagwu & Ugwuanyi, 2014; Moham- physicochemical properties, heavy metal content, and
mad et al., 2021). Due to its voluminous abundance microbial characteristics of soil with the potential to
and disposal challenges, POME is a paramount chal- cause toxicological impacts on the environment and
lenge associated with oil palm factory. Direct disposal its biota (humans, fisheries, flora, and fauna) (Izah
of untreated POME on land could result to clogging and et al., 2018). Besides polluting the soil, surface, and
water logging of the soil, as well as death of vegetation ground water, when left in stagnant open channels,
(Mohammad et al., 2021). On the other hand, discharg- CMW serves as a breeding ground for mosquitoes and
ing untreated POME into water bodies causes depletion other parasitic insects, coupled with its characteristic
in the oxygen content of the water, resulting in aquatic foul odour (Oghenejoboh et al., 2021).
pollution, death of fish, and other aquatic flora (Dominic
& Baidurah, 2022; Mohammad et al., 2021). In some Distillery wastewater (DWW)
extreme cases, it brings about a colour change in the water
body with an unpleasant smell and viscous characteristic, This large volume of waste chemical liquid gener-
thereby rendering the water unsuitable for use (Iwuagwu ated in the course of ethanol distillation is called
& Ugwuanyi, 2014; Mohammad et al., 2021). distillery wastewater. With an average amount of
about 12–15 L of wastewater per litre of ethanol
High‑carbohydrate wastewater (HCW) produced, distillery industries are estimated to be
the second-highest most wastewater generating
High-carbohydrate wastewaters are produced widely agro-processing industries after paper-pulp (Wagha
in many food processing and fermentation indus- & Nemadeb, 2018).
tries. HCWs are rich in organic substances and high Distillery wastewater is dark brown in appear-
in COD. Thus, the serious environmental pollution if ance, acidic in nature, with high BOD and COD
not properly treated and disposed. A typical example (Ratna et al., 2021). It also contains various salts,

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heavy metals, endocrine-disrupting chemicals component in PMW vary from one production plant
(EDCs), and micro- and macro-minerals (Fito et al., to the other. These variations can be attributed to fac-
2019). However, the type and concentration of these tors such as the type of raw material (wood) used, the
parameters in DWW are dependent on the feedstock production process, the technology employed, the
used during fermentation (Chowdhary et al., 2018). volume of water that the mill can circulate as well
Melanoidin, a by-product of a non-enzymatic Mail- as the management techniques adopted (Dagar et al.,
lard reaction between reducing sugar and amino 2022; Islam et al., 2022; Toczyłowska-Mamińska,
compounds at high temperature, is the predomi- 2017). The effluent from the pulping processing is
nant colouring compound in DWW responsible for called “black liquor”, due to its characteristic black
its characteristic dark colour (Cooper et al., 2019). colour owing to being rich in lignin and lignin deg-
Moreover, DWWs are composed of different types radation products (Toczyłowska-Mamińska, 2017).
of carbohydrates and organic acid, thereby making Depending on the type of wood and the production
them high in organic matter (Gebreeyessus et al., process employed, this waste fluid is significantly
2019). However, the combination of organic chemi- high in BOD, COD, tannins, suspended particles
cals, salt, melanoidin, and heavy metals in the efflu- (majorly fibres), fatty acids, resin acids, lignin, and
ent makes DWWs exceedingly mutagenic, cyto- its derivatives (Dagar et al., 2022; Toczyłowska-
toxic, and genotoxic to both terrestrial and aquatic Mamińska, 2017). Currently, a number of organic and
organisms at low concentrations (Ratna et al., 2021). inorganic chemicals, ranging from adsorbable organic
DWWs are major sources of surface water pollut- halogens (AOX), lignins, stilbenes, sulphur com-
ants. Their presence in surface water has a negative pounds, dioxins, chlorides, furans, phenols, and other
impact on penetration of sunlight, dissolved oxy- compounds such as organic substances, suspended
gen (DO) level, photosynthesis, aquatic fauna, and solids, and chromophoric compounds have been iden-
flora (Ratna et al., 2021). Due to their high content tified in PMW (Bhardwaj et al., 2022; Dagar et al.,
of organic and inorganic matter as well as sufficient 2022; Toczyłowska-Mamińska, 2017). While some
amount of macronutrients, DWWs have been pro- of these compounds are natural wood extractives, oth-
posed as a source of fertilizer or to be used in fer- ers are xenobiotics generated via the pulp and paper
tigation. As stated by Umair Hassan et al. (2021) in manufacturing process that are recalcitrant in nature
their extensive review, the use of optimized doses (Bhardwaj et al., 2022; Dagar et al., 2022). Given
of DWW have been reported to improve soil enzy- their dark colour and high turbidity, their presence in
matic and microbial activities substantially as well surface water adversely affect sunlight penetration,
as other soil properties including organic carbon, dissolved oxygen (DO) content, and photosynthetic
nutrient uptake, soil porosity, water holding capac- activity. This may result in anaerobic decomposition
ity, aggregate stability, and anti-oxidant activities, under the blanketing effect releasing hydrogen sul-
which results in improved photosynthetic efficiency, phide into aquatic ecosystems, thereby affecting the
growth, and yield. Although the role of DWW in soil aquatic flora and fauna as well as loss of aesthetic
quality and crop yields improvement is well docu- beauty of the environment (Singh & Tripathi, 2020).
mented, nevertheless, there are still numerous unad- Some of the compounds in the PWM are mutagenic
dressed questions concerning its safety especially resulting in various physiological and morphological
when used untreated (Umair Hassan et al., 2021). abnormalities in fish inhabiting receiving water bod-
ies (Singh & Tripathi, 2020). The recalcitrant com-
Pulp mill wastewater (PMW) pounds bio-accumulate with negative consequences
on the ecosystem. Additionally, some of the toxins
However, the most significant sources of contami- like chlorides, transition metals, chelating agents, and
nants during paper manufacturing processes are wood dioxins found in PMW could be harmful to humans
preparation, pulping, pulp washing, screening, bleach- when ingested either through drinking contaminated
ing, papermaking, and coating processes depending water or consuming products from these contami-
on the method used (Dagar et al., 2022). The chemi- nated water bodies (Toczyłowska-Mamińska, 2017).
cal composition and concentration of individual

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Textile wastewater these pollutants in LW depends largely on the com-

position of the animal feed, the housing method, the
Textile industries are key contributors to global envi- species of the animal species, the age of the animal,
ronmental pollution problems because they release and other prevailing environmental factors (L´opez-
undesirable and harmful dye effluents. Typically, it is S´anchez et al., 2022; Nagarajan et al., 2019). These
difficult to define textile waste effluent due to vary- compounds when they enter water bodies can pro-
ing production procedures amongst textile indus- mote cultural eutrophication of water bodies posing
tries. Generally, the textile effluent is heavily polluted hazard to both the environment and humans. Also
due to presence of mixture of reactive dyes, metals, when LWW is dumped into the environment, it emits
and other pollutants (Holkar et al., 2016; Yaseen & strong awful odour causing air pollution.
Scholz, 2019). The textile effluents contain high Furthermore, livestock feed is supplemented with
loads of organic and inorganic compounds including antibiotics and parasiticide to impede microbial infec-
colour, suspended solids, metals, and salts, in addi- tions, promote feed efficiency, and boost growth.
tion to high COD and BOD (Yaseen & Scholz, 2016, Additionally, livestock feeds are supplemented with
2019; Holkar et al., 2016; Kannan et al., 2022). Due heavy metals such as copper and zinc and, to a lesser
to their high colour content, COD, and BOD, textile extent, nickel and selenium, which are necessary for
wastewater impairs with light penetration and other several physiological processes in livestock (Hu et al.,
normal processes in aquatic environment resulting 2020; Li et al., 2020). Eventually, as much as 70% or
in an imbalance in the aquatic ecosystem (Danouche more of these antibiotics and 80–90% of these heavy
et al., 2021; Holkar et al., 2016). Owing to the pres- metals are discharged unmodified into the waste-
ence of component metals and chlorine in synthetic water (Hu et al., 2020; Li et al., 2020). Also steroid
dyes, the components of textile wastewater can have hormones are found in LWW as a result of supple-
deadly effects on certain species of marine life, and mentation of livestock feeds with steroids as well as
this toxicity of synthetic dyes may occur across food those naturally produced livestock (Hu et al., 2020; Li
chains (Danouche et al., 2021; Holkar et al., 2016). et al., 2020). By implication, these antibiotics, heavy
In addition, textile wastewater can severely affect the metals, and steroids enter the environment increas-
nutritional quality of the agricultural products and ing problems of antibiotics resistance, contaminating
animal and human health causing skin itching, blind- both surface and underground water sources and ulti-
ness, chemosis, and dermatitis (Kannan et al., 2022; mately causing havoc for humans.
Parmar & Shukla, 2018).
Wastewater remediation processes
Livestock wastewater (LWW)
Given the harmful nature of wastewater, it is essen-
Livestock effluent is one of the severe sources of pol- tial for wastewater to be treated before discharging
lution to both terrestrial and aquatic environments. it into the environment. The conventional method of
Livestock wastewater include swine effluent, cat- wastewater treatment combines physical, chemical,
tle effluent, and poultry effluent which refer to all and biological processes to remove suspended and
excrement or residual excrement produced, as well dissolved solids from wastewater, as well as biologi-
as urine, feed residue, and washing wastewater gen- cal breakdown of organic debris and nutrients. These
erated by livestock farms (Hu et al., 2020). This has three major wastewater treatment processes are com-
led to profound environmental problems, such as soil bined into an array of systems, namely, preliminary,
degradation, surface and groundwater pollution asso- primary, secondary, tertiary, and/or advanced (treat-
ciated with livestock effluent. ment of the generated sludge), so as to achieve dif-
LWW contains high loads of suspended sol- ferent levels of decontamination of the wastewater.
ids, volatile compounds, nitrogen, phosphorus, and Table 1 captures the unit of operation of each treat-
organic matter (expressed as COD and BOD) nutri- ment system as well as the limitations of the conven-
ents and faecal coliforms (Hu et al., 2020; L´opez- tional wastewater treatment. Although the conven-
S´anchez et al., 2022). However, the proportion of tional wastewater treatment has the advantages of

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being technically simple and adaptable for different the aid of microorganisms will not only contribute in
waste, well-established, and proven to be efficient waste stream control, but will also close material loops
in wastewater treatment, it does not support the cir- (Gudiukaite et al., 2021). Successful lab-scale valori-
cular economy concept. Wastewater is solely treated zation of wastewater using microorganisms including
with the aim of making it suitable for disposal with- bacteria, fungi, microalgae, yeast, or even a combina-
out actually exploiting the wastewater for recovering tion of different microorganisms has been reported
and transforming its nutrients into highly valuable (Broos et al., 2022; Kannan et al., 2022).
bioactive compounds (Fig. 2). Additionally, the con-
ventional wastewater treatment is capital-intensive
and may not be affordable to small-scale industries Characteristics of yeast that make them suitable for
especially in developing countries. In view of these wastewater valorization
limitations, microbial valorization of wastewater is
being proposed as an alternative to the conventional Yeasts are classified as either fermenting yeast (that
wastewater treatment. can utilize six-carbon sugar and produce alcohol
and carbon dioxide) or oxidizing yeast (with strong
Microbial wastewater valorization oxidation ability and low or no fermentative abil-
ity) (Eliodório et al., 2019). Yeast cells are ubiqui-
Currently, there is increasing awareness of the valuable tous in nature, generally fast growing, can utilize a
resource present in wastewater which can be recov- wide range of substrate, show substantial resistance
ered and converted to important bioactive compounds to stress and inhibitory conditions, and have found
using microorganisms. Microbial wastewater valoriza- application in many industrial processes. Some spe-
tion has also gained attention as a wastewater treatment cific characteristics that make yeast cells suitable for
alternative as a result of its natural means of waste valorization of wastewater are discussed below.
processing coupled with its autonomous growth and
environmental friendliness. During microbial treat- Fast growth rate
ment of wastewater, the contaminants are removed
while economically important compounds are syn- When grown in a liquid medium, yeast cells follow a
thesized (Fig. 2). This integrated system incorporat- well-established pattern for microbial growth. Under
ing wastewater treatment and resource recovery with favourable conditions, most wild-type yeast strains

Table 1  Unit operation, advantages, and disadvantages of conventional wastewater treatment technique
Treatment Unit operation Aim Limitations

Preliminary Screening Removal of floating materials It is capital-intensive thus may not be

Comminution Removal of heavy settleable inorganic solids affordable for small-scale industries
Skimming such as sand and oil Requires the use of chemicals which may
Primary Sedimentation Removes 60–65% of suspended solids and introduce other pollutants
Coagulation 30–35% of BODs’ large-sized settleable Little or no value-added product is generated
Flocculation organic matter and small-sized inorganic Large amount of sludge may be generated
materials which may be difficult to handle sometimes
Some of the processes requires a skilled
Secondary Aerobic processing About 95% of the total suspended solid
personnel
Anaerobic processing removal
Pond processing Reduction of BOD and COD to the lowest
acceptable concentration
Tertiary Ion exchange Removal of total suspended particles, total
membrane separation dissolved solids, organic and inorganic
techniques materials from secondary effluent
Electrochemical Removal of certain organic and inorganic
techniques elements as well as nutrients, killing of
Advanced oxidation pathogens
process

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Fig. 2  Conventional vs. microbial valorization of wastewater

have a doubling time in the 90 to 120 min. This may metabolized by yeast cells (Yang & Zheng, 2014).
however be affected by factors such as the specie of As recently reviewed by Baptista and Domingues
the yeast and growth conditions. With the exception (2022), it was shown that the yeast Kluyveromy-
of bacteria, yeast cells grow faster than most other ces marxianus could be a microbial cell factory
group of microorganisms. Furthermore, product for- for lignocellulosic biomass valorization owing to
mation in yeast cells can occur during the exponential but not limited to its ability to utilize wide range of
growth and stationary phases. Moreover, yeast cells substrate. In another recent review, Wierzchowska
can stay in the stationary phase of growth for a pro- et al. (2022) also revealed that the yeast Yarrowia
longed period of time and can resume growth if con- lipolytica is suitable for valorization of hydrophobic
dition becomes favourable again. industrial waste due to its ability to utilize a wide
range of carbon source amongst other traits. There
Utilization of wide range of substrate are also several reports on different substrate utili-
zation by both wild-type and engineered strains of
Most yeast cells are not fastidious in their growth Saccharomyces cerevisiae. It is likely possible that
requirements. They can grow on cheap available numerous other yeast species have this potential but
nutrients, and that is part of the reason why yeast are yet to be brought to limelight.
cells have found application in many industrial pro-
cesses (Yang & Zheng, 2014). This makes them Tolerance to stress and inhibitor
good candidates for valorization of wastewater
which is laden with numerous substrates. Sugars Considering the characteristics of wastewater, it poses
(glucose, sucrose, lactose, fructose, and maltose), a great stress and inhibitory conditions for microbes.
biopolymers (starch, cellulose, hemicellulose, and Yeast cells are known to have mechanisms that help
pectin), pentoses, methanol, alcohols, polyols, them cope with these stressors and inhibitory com-
hydrocarbons, fatty acids, and organic acids are all pounds. Yeast cells have been reported to grow at

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low pH, high temperature, and elevated salt and sugar discolouration of wastewaters (Tables 2 and 3). Yeast
conditions. In addition, they have also been reported cells were also effective in the reduction of oil con-
to tolerate heavy metals and lignocellulosic inhibitory tent, total nitrogen content, total solid content, and
compounds (furfural derivatives, organic acids, and total phosphorus content of wastewater. Dias et al.
phenolics). For instance, the yeast Pichia kudriavzevii (2020) reported 45.8% total Kjeldahl nitrogen reduc-
has been reported to tolerate low pH (as low as pH tion from secondary brewery wastewater mixed with
2.5), high temperature (above 45 ℃), high sugar con- sugarcane molasses (SCM) by the yeast Rhodosporid-
centration (up to 20%), and high ethanol concentra- ium toruloides. Removal efficiencies of 67.69% total
tion (about 13%), in addition to high concentrations phosphorus (TP) and 57.81% total nitrogen (TN) was
of furfurals, organic acids, heavy metals, and pheno- achieved within three cultivation days in distillery
lics (Hisamatsu et al., 2006; Mukherjee et al., 2017; wastewater with the yeast Rhodosporidium toruloides
Ndubuisi et al., 2020). Other yeast species such as (Ling et al., 2013). The co-culture of C. rugosa and Y.
Kluyveromyces marxianus, Yarrowia lipolytica, Wick- lipolytica effectively eliminated up to 98.5% triglycer-
erhamomyces anomalus, and Torulaspora delbrueckii ides from the undiluted POME within 120-h culture,
have been documented to have similar stress and while 94.5% and 83.9% removal were achieved with
inhibitor tolerance abilities. monocultures of C. rugosa and Y. lipolytica (Theera-
chat et al., 2017). Yousuf et al. (2010) recorded 86%
Genetically modifiable and 98% sugar content and protein content removal,
respectively, from olive oil mill effluent by the yeast
Yeast cells like Pichia pastoris and Saccharomyces L. starkeyi. The removals of oil content, total phos-
cerevisiae are the most engineered of all the yeast phorus, and N ­ H4+-N from refined soybean oil waste-
species. This shows that these yeast strains can be water by the yeast Trichosporon fermentans were
modified to improve their wastewater valorization 90.33%, 89.7%, and 82.3%, respectively (Yu et al.,
potential. Currently, some of the nonconventional 2018). Rhodotorula kratochvilovae EXF7516 and
yeasts are not easily engineered due to poor under- Cutaneotrichosporon oleaginosum ATCC 20509 con-
standing of their genetic makeup. However, there is sumed approximately 87% of monomeric substrates
high expectation that with the fast developing molec- (sugars, aromatics, and organics acids) and about
ular biology tool kit and bioengineering technologies, 23% of non-monomeric substrates (partially degraded
this problem will be solved and scientists could fur- xylan, lignin, cellulose) in lignocellulose, phenolic
ther augment microbial wastewater remediation and wastewaters (Broos et al., 2022). Rhodotorula glutinis
conversion to useful metabolites (Sun et al., 2020). achieved 69.12% and 73.85% removal of TN and TP,
Beyond this, they can also serve as source of neces- respectively, from mixed corn steep water/corn gluten
sary genes for modification of other organisms for water effluent (Liu et al., 2016). Zhang et al. (2018)
wastewater valorization. reported that R. glutinis cultivation in cellulosic etha-
nol wastewater resulted in TOC (total organic carbon),
­NH4+-N, TN, and TP removal rates reaching 81.81%,
Yeast‑mediated valorization of wastewater 85.49%, 70.52%, and 67.46%, respectively. There was
significant reduction in the toxic components includ-
Bioremediation of agro‑industrial wastewater by ing 89% colour, 94.27% lignin, and 99.60% phenol
yeast cells from pulp and paper industry effluent when used for
the cultivation of Rhodosporidium kratochvilovae
Yeast cells have been used successfully to clean up HIMPA1 (Patel et al., 2017). Significant reduction
different types of wastewater. Although most of the in TOC (54%) and 98% ADMI colour removal were
reports are at lab scale, they are promising. Yeast cells obtained when textile dyeing wastewaters was treated
either singly or in consortium have been reported with oleaginous consortium of Barnettozyma cali-
to bring about 18–95% reduction in COD from dif- fornica SSA1518, Yarrowia sp. SSA1642, and Ster-
ferent wastewater (Tables 2 and 3). They were also igmatomyces halophilus SSA1511 (Ali et al., 2021).
reported to bring about 35–77% reduction in BOD, Saccharomyces cerevisiae removed 98.1%, 83.0%,
as high as 72% reduction in total phenolics and 60% and 60.5% of turbidity, sulphates, and phosphate

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Table 2  Remediation of oil-based wastewater by yeast cells

Wastewater Yeast strain COD Total phenol Decolourization Ref
reduction reduction (%)
(%) (%)

Olive oil mill Yarrowia lipolytica 80 70 - Araujo et al., 2005

molasses and olive mill Saccharomyces cerevisiae - 28 60 Sarris et al., 2014
MAK-1
Olive oil mill Candida diddensiae YB37 - 72 - Chakri et al., 2007
Olive oil mill Lipomyces starkeyi 48 43 - Yousuf et al., 2010
Olive oil mill S. cerevisiae strain MAK-1 - 34 63 Sarris et al., 2014
Olive oil mill Yarrowia lipolytica ACA-YC - 51 58 Sarris et al., 2017
5033
Olive oil mill Y. lipolytica PO1 41.2 - - Lanciotti et al., 2005
Olive oil mill Geotrichum candidum 10,457 22 31 6 Bleve et al., 2011
G. candidum 10,461 23 30 3
Pichia fermentans 10,462 18 26 3
Candida tropicalis 10,456 18 25 0
Bead immobilized G. candidum 52.3 49.1 26
Palm oil mill Y. lipolytica 48 64 Theerachat et al.,
Candida rugosa 54.6 71.7 2017
Co-culture of C. rugosa and Y. 60.3 68.3
lipolytica
Olive oil mill blended with Y. lipolytica 10 30 Sarris et al., 2017
Crude Glycerol
Olive oil mill Candida diddensiae - 44 - Abdelhadi et al.,
2010
Palm oil mill Y. lipolytica NCIM 3589 95 - - Oswal et al., 2002
Olive oil mill C. diddensiae ymc78 64.8 32 4 Sassi et al., 2008
Pichia guilliermondii ymc85 53 25 7
C. ernobii ymt95 62 34 0
C. diddensiae ymc98 55 43 2
Candida holstii ymc73 57.9 39 0
Palm oil mill Saccharomyces sp L3 83 - - Iwuagwu &
Ugwuanyi, 2014
Palm oil mill Rhodotorula glutinis 66 - - Saenge et al., 2011
Olive oil mill Rhodotorula mucilaginosa 56.9 34.8 56.08 Jarboui et al., 2012
Palm oil mill Y. lipolytica TISTR 5151 81.2 - - Louhasakul et al.,
2016
62.4% expired soft D. etchellsii BM1 41.3 Arous et al., 2016
drinks + 37.6% olive mill
wastewater
Palm oil mill L. starkeyi 74.35 Karim et al., 2021a,
b
Refined soybean oil wastewater Trichosporon fermentans 94.7 Yu et al., 2018

content, respectively, while Torulaspora delbrueckii phosphate content, respectively, during growth in tan-
reduced 92.9% and 61.9% of total sulphates and nery effluent (Okoduwa et al., 2017).

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Table 3  Remediation of other wastewater by yeast cells

Wastewater Yeast strain COD BOD Ref
reduction reduction
(%) (%)

Corn steep water/corn gluten water R. glutinis 77.4 Liu et al., 2016
50% cheese whey + 50% expired soft D. etchellsii BM1 58.35 Arous et al., 2016
drinks
Cellulosic ethanol wastewater R. glutinis 83.15 Zhang et al., 2018
S. cerevisiae CCMA 0188 and CCMA Vinasse 29.29 51.56 Pires et al., 2016
0137
Tannery effluent Torulaspora delbrueckii 90.6 48 Okoduwa et al., 2017
Saccharomyces cerevisiae 54.2 60.7
Textile effluent Candida zeylanoides 57 57 Abioye et al., 2014
Saccharomyces cerevisiae 66 66
Pharmaceutical effluent Torulaspora delbrueckii 38.3 38.3 Abioye et al., 2015
Saccharomyces cerevisiae 52.5 52.5
Cassava mill effluents Saccharomyces cerevisiae 74.48 Izah et al., 2017
Potato wastewater and 5% glycerol Candida utilis ATCC 9950 91 Kurcz et al., 2018
Bioethanol wastewater Rhodosporidium toruloides Y2 72.3 Zhou et al., 2013
Distillery wastewater Cryptococcus curvatus 78.98 Gonzalez-Garcia et al., 2011
Rhodotorula glutinis 84.44
Dairy waste Yarrowia lipolytica ATCC 9773 44.3 43.3 Dunoyer et al., 2020
Secondary brewery wastewater mixed Rhodosporidium toruloides 81.7 Dias et al., 2020
with sugarcane molasses
Mixed distillery and domestic waste- R. toruloides 91.54 Ling. et al., 2014
water
Mixed distillery and local municipal R. toruloides 86 Ling et al., 2013
wastewater
Cellulosic ethanol wastewater + 40 g/L Rhodotorula glutinis 83.15 Zhang et al., 2018
glucose
Acetone-butanol-ethanol (ABE) Trichosporon cutaneum 68 Xiong et al., 2015
fermentation wastewater
Fishmeal wastewater + 20 g/L glucose Lipomyces starkeyi 43.4 Huang et al., 2011
Butanol fermentation wastewater Trichosporon dermatis 68 Peng et al., 2013
Pulp and paper industry effluent Rhodosporidium kratochvilovae 94.22 77.36 Patel et al., 2017
HIMPA1
Cellulosic ethanol fermentation T. cutaneum ACCC 20271 55.05 Wang et al., 2017
wastewater
Textile dyeing effluent Yarrowia sp. SSA1642 81 67 Ali et al., 2021
S. halophilus SSA1511 87 60
Mixed culture of Yarrowia sp. 95 74
SSA1642, Barnettozyma californica
SSA1518, and Sterigmatomyces
halophilus SSA1511

Heavy metal wastewater some are carcinogenic (Mersin & Açikel, 2021). At
high concentrations, these metals are hazardous, caus-
Heavy metal contamination has emerged as one of ing physiological and neurological hazards (Mersin &
today’s most serious environmental issues. These Açikel, 2021). Several processes are used to remove
heavy metals are associated with various diseases and heavy metals from wastewater, including chemical

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precipitation, ion exchange, adsorption, membrane for removal of harmful or important. Yeast cells both
filtration, reverse osmosis, and solvent extraction. dead and alive have been studied for their biosorp-
Also reported as affordable adsorbent materials for tion potentials. Saccharomyces cerevisiae and Can-
the removal of heavy metals from contaminated water dida species have received increasing attention due to
are clay, zeolite, chitin, and agro wastes. Removal distinct metal sorption capacity. Nevertheless, other
of these metals using biological approaches such as yeast cells with metal sorption capacity have been
biosorption or bioaccumulation strategies are attrac- reported (Table 4). Biosurfactant from the yeast Can-
tive options to be explored. Microorganisms, as heavy dida sphaerica UCP0995 was reported to have 75%
metal bioadsorbents, are being explored as alternative Pb and 87% Cd removal efficiencies from aqueous

Table 4  Remediation of heavy metals wastewater by yeast cells

Yeast strain Wastewater Initial heavy metal Result Ref
concentration

Saccharomyces Pb2 + containing aqueous < 1 ppm Achieved an uptake of up Stathatou et al., 2021
cerevisiae solutions to 12 mg lead per gram of
biomass
Engineered Saccharomy- Athabasca oil 1–2 ppm or mg/L Yeast-induced chemical Sun et al., 2020
ces cerevisiae Sand effluent containing precipitation removed more
copper, cadmium, than 85% of copper, mercury,
mercury, lead, and zinc and lead. Cadmium and zinc
were also removed to the
tune of 30–50%
Aureobasidium pullulans Zn-containing minimal 1 mg/L Over 90% detoxification of Zn García-Béjar et al., 2020
H1 salt medium
D. rugosa FP5 98% detoxification
P. kudriavzevii Ak9 77% detoxification
S. cerevisiae EB21 95.56% detoxification
P. fermentans 98% detoxification
Aureobasidium pullulans AFB1-containing mini- 0.04 mg/L Almost 70% of AFB1
H1 mal salt medium detoxification
K. pastoris 71% AFB1 detoxification
S. cerevisiae Artificial aqueous solu- 2.6 mg Mn2 + /l S. cerevisiae F-25 living Fadel et al., 2017
tion of Mn2 + cells exhibited a maximum
biosorption efficiency of
22.5 mg Mn2 + /g yeast
biomass, suggesting an
86.53% efficiency
Candida Aqueous solution 25 mg/L Cu(II) or 68.4% Cu(II) bioaccumulation Mersin & Açikel, 2021
membranifaciens containing either Cu(II) 25 mg/L Ni(II) efficiency
or Ni(II) 58.8% Ni(II) bioaccumulation
efficiency
C. utilis 60.4% Cu(II) bioaccumulation
efficiency
48.3% Ni(II) bioaccumulation
efficiency
C. tropicalis 78.7% Cu(II) bioaccumulation
efficiency
64.5% Ni(II) bioaccumulation
efficiency
C. lipolytica 88.8% Cu(II) bioaccumulation
efficiency
81.7% Ni(II) bioaccumulation
efficiency

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solution (Luna et al., 2016). Yeast has also been stud- being used in the production of chocolate (Caporusso
ied for remediation of fossil fuel–contaminated water. et al., 2021). SCOs can be converted to other high-
In a study by Gargouri et al. (2015), Candida tropica- value products via a various chemical, physical, and
lis and Trichosporon asahii were shown to be capable biochemical techniques. The most common of them
of degrading a wide spectrum of hydrocarbons and all being the transesterification of SCOs to biodiesel.
totally metabolizing n-alkanes. As the sole carbon SCOs have been produced from various wastewaters
source, they were unable to completely break down including OMW, POME, cassava wastewater, live-
aromatic hydrocarbons (phenanthrene, anthracene, stock wastewater, and distillery wastewater by yeast
fluoranthene, and pyrene). (Table 5). Lipid accumulation as high as close to 70%
of dry weight by yeast grown in wastewater has been
Production of value‑added products from wastewater reported (Ali et al., 2021). Interestingly, there are var-
by yeast cells iations in the amount of lipid accumulated as well as
the lipid profile from different wastewaters as well as
Single cell oil different yeast strains (Table 5).

Single cell oils (SCOs) are intracellular storage lipids Biodiesel

consisting of triacylglycerols (TAGs), free fatty acids,
polar lipids, sterols, hydrocarbons, and pigments. Biodiesel is considered an alternative to fossil-based
SCOs have similar composition with vegetable oils diesel with high energy density, highly biodegrad-
and animal fats (Mhlongo et al., 2021). Neverthe- able, and low in greenhouse gas emissions. Biodiesel
less, single cell oils are preferred to plant- and ani- is currently mass-produced from conventional feed-
mal-derived oils because it is easy to scale up their stocks, making it expensive. SCOs which have similar
production. Their production is not affected by fac- properties to animal fats and vegetable oils are poten-
tors such as seasonal changes, geographic location, tial alternatives for the production of biodiesel. SCO
harvest time, and transport, which are of concern produced from wastewater by yeast has been explored
when using plant and animal materials (Mhlongo for biodiesel production. Metschnikowia pulcherrima
et al., 2021). SCOs are produced by all organisms MTCC 632 produced 10.8 ml of FAME for every
as part of their structural and functional processes; litre of culture media from a mixture of distillery
however, their capacity to produce SCOs can vary. spent wash lignocellulosic hydrolysate (Tamilalagan
Oleaginous microorganisms possess the ability to et al., 2019). In the same study, fossil diesel blended
accumulate lipids over 20% (w/w) of their dry body with 20% of produced biodiesel resulted in the best of
weight. Oleaginous yeasts are intriguing microbial reduced hydrocarbon and carbon monoxide emissions,
factories. They have the ability to produce and accu- whilst also delivering the highest brake thermal effi-
mulate a significant quantity of SCOs. Oleaginous ciency and the lowest brake specific fuel consumption.
yeasts accumulate lipids, in its neutral forms, namely, Wang et al. (2017) stated that the fatty acid composi-
monoacylglycerols, diacylglycerols, and triacylglyc- tion of the SCO derived during the cultivation of cel-
erols (Caporusso et al., 2021; Mhlongo et al., 2021). lulosic ethanol fermentation wastewater with Trichos-
Generally, the most regularly accumulated fatty poron cutaneum ACCC 20271 was comparable with
acids by oleaginous yeasts are C16:0 (palmitic acid), vegetable oil, thus could be used as a feedstock for the
C16:1 (palmitoleic acid), C18:0 (stearic acid), C18:1 production of biodiesel. In a direct transesterification
(oleic acid), and C18:2 (linoleic acid), although fatty reaction, lipid from the wet cells of Y. lipolytica cul-
acids such as C14:0 (myristic acid) and C18:3 (lino- tivated in POME produced 40.9% fatty acid methyl
lenic acid) are produced in relatively small quantities esters that are quite similar to plant oil, demonstrating
(Caporusso et al., 2021). In food industries, SCOs are its prospect as a feedstock for the production of bio-
proposed as sources of polyunsaturated fatty acids diesel (Louhasakul et al., 2016). Saenge et al. (2011)
(PUFAs) which are required for maintaining bio- reported that the biodiesel produced by POME grown
functions in mammals especially humans. They are R. glutinis TISTR 5159 had CN values above 54 which
used as substitutes for cocoa butter which has many meets the minimum requirement standards, for CN
applications in the cosmetic industries in addition to values set by DIN 51606 (Germany), ASTMD 6751

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Table 5  Lipid accumulation and lipid profile of SCO produced from wastewater by yeast
Yeast cell wastewater Lipid Lipid Lipid profile Ref
accumulated yield
(%DCW) (g/L)

C. curvatus Tequila vinasses 25.2 1.49 Myristic, palmitic, Gonzalez-Garcia et al.,

stearic, oleic, and 2011
linoleic
R. glutinis 27.02 1.86 Myristic, palmitic,
stearic, oleic, linoleic,
and linolenic
Trichosporon fermentans Refined soybean oil 43 3.4 Palmitic acid (19.6%), Yu et al., 2018
wastewater linoleic acid (36.4%),
oleic acid (35.5%), and
stearic acid (8.5%)
T. cutaneum ACCC​ Cellulosic ethanol - 2.16 Oleate (67.8%) Wang et al., 2017
20271 fermentation waste- Palmitate (14.2%)
water Stearate (10.12%)
Margarate (1.88%)
Lignocerate (1.65%)
Pentadecanoate (1.09%)
Others (3.25%)
Metschnikowia Distillery spent 37.2 - - Tamilalagan et al., 2019
pulcherrima MTCC wash + lignocellulosic
632 hydrolysate
Rhodosporidium Secondary brewery 29.9 SFA (32%/w TFA) Dias et al., 2020
toruloides wastewater + with MUFA (48.8%/w TFA)
sugarcane molasses PUFA (19%w/w TFA)
α-linolenic acid
(3.4w/%w TFA)
Yarrowia lipolytica Olive mill wastewaters 22.4% 2.54 Oleic acid (63%) Sarris et al., 2019
ACA-DC 5029 blended with Linoleic acid (52%)
crude glycerol Palmitoleic acid (36%)
Palmitic acid (18%)
Stearic acid (8.5%)
R. toruloides Distillery wastewater 43.65 3.54 - Ling et al., 2013
Yarrowia lipolytica Palm oil mill effluent 1.64 - Louhasakul et al., 2016
TISTR 5151
Rhodotorula glutinis Palm oil mill effluent 51.85 4.58 Saenge et al., 2011
Lipomyces starkeyi Olive oil mill 29.5 Palmitic acid (19.1%) Yousuf et al., 2010
wastewaters Stearic acid (8.5%) oleic
acid (49.1)
Linoleic acid (18.8%)
Linolenic acid (3.5%)
Free fatty acids (14.5%)
Yarrowia lipolytica OMW/glucose blend 48 Oleic acid (71.7%) Sarris et al., 2017
ACA-YC5033 Linoleic acid (22.5%)
Palmitoleic acid (18.2)
Rhodococcus opacus Pine organosolv pretreat- 26.99 30% oleic, 20% palmitic, Wells Jr et al., 2015
DSM 1069 ment effluent and 17% stearic fatty
acids
C. pseudolambica 35.3 Palmitic acid (16.9%)
Stearic acid (9.9)
Oleic acid (52%)
Linoleic acid (8.7%)
Palmitoleic acid (6.9)

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Table 5  (continued)
Yeast cell wastewater Lipid Lipid Lipid profile Ref
accumulated yield
(%DCW) (g/L)

I. occidentalis Livestock wastewater 28.9 Palmitic acid (11.9) Chung et al., 2016
Stearic acid (3.9%)
Oleic acid (58.9%)
Linoleic acid (18.4%)
Palmitoleic acid (9.6)
Rhodotorula Palmitic acid (23%)
kratochvilovae Stearic acid (12%)
EXF7516 Oleic acid (44%)
Linoleic acid (19%)
Palmitoleic acid (1%)
Cutaneotrichosporon Phenolic wastewaters 30 5.4 Palmitic acid (2%) Broos et al., 2022
oleaginosum ATCC Stearic acid (72%)
20509 Oleic acid (10%)
Linoleic acid (2%)
Palmitoleic acid (14%)
Rhodotorula glutinis Cassava waste water 1.34 Oleic acid (59.76%) Ribeiro et al., 2019
linoleic acid (7.59%)
Rhodotorula glutinis Corn steep water/corn 28.90 7.62 Liu et al., 2016
gluten water
Rhodotorula glutinis Cellulosic ethanol 18.35 2.08 Palmitic acid (10.62%) Zhang et al., 2018
wastewater Linoleic acid (4.5%)
Oleic acid (79.36%)
Stearic acid (5.52%)
Debaryomyces etchellsii 62.4% of expired soft 28.9% 1.2 Arous et al., 2016
drinks and 37.6% of
OMW (25%, v/v)
R. kratochvilovae Pulp and paper industry 61.71 8.56 Palmitic acid (21.86%) Patel et al., 2017
HIMPA1 effluent + glucose Linoleic acid (15.9%)
Oleic acid (45.43%)
Consortium of Yarrowia Dyestuff and textile 68.6 7.81 Oleic acid (45%) Ali et al.,2021
sp. SSA1642, effluent Stearic acid (4.79%)
Barnettozyma califor- Linoleic acid (18.04)
nica SSA1518, and
Sterigmatomyces halo-
philus SSA1511
L. starkeyi Palm oil mill effluent 22.8 1.6 Oleic acid (13.8%) Karim et al., 2021a, b
Stearic acid (21.3%)
Linoleic acid (4.6%)
Palmitic acid (28. 6%)
Myristic (10.3%)
Pentadecanoic (7.2%)
Trichosporon dermatis Butanol fermentation 13.5% Oleic acid (39.9%) Peng et al., 2013
wastewater Stearic acid (10.6%)
Linoleic acid (21.7%)
Palmitic acid (19.4%)
Others (8.4%)
Trichosporon Deproteinated potato 33 4.8 Linoleic acid (36%) Gientka et al., 2019
domesticum PCM 2960 wastewater Oleic acid (26%)
Palmitic acid (19.5%)
Stearic acid (7%)
α-Linolenic acid (5%)

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(USA), and EN 14214 (European Organization), thus rich in variety of essential nutrients. However, for a
making it an excellent feedstock for biodiesel pro- microbe to be used for SCP production, it must be
duction. Issatchenkia occidentalis grown in livestock certified as generally regarded as safe (GRAS) organ-
wastewater–produced lipids meets all the European ism. Among the SCP-producing organisms (bacte-
(EN) standards for biodiesel in relation to cetane num- ria, yeasts, microalgae, and mould), yeast is the most
ber (CN), cold filter plugging point (CFPP), oxidation extensively used. Yeast is bulky, fast growing (with a
stability (OS), and iodine value (Chung et al., 2016). generation time of 1 to 3 h), and high in protein con-
The composition of FAMEs in the lipid accumulated tent which is 45 to 55% (Zhang et al., 2021). Addi-
by Rhodococcus opacus DSM 1069 cultivated with tionally, yeast can alter the microbial flora in animals,
pine organosolv pretreatment effluent were largely improve production performance, modulate immuno-
even numbered monounsaturated and saturated fatty logical function, and increase immunity to infections
acids with potentials as a raw-material for biofuel pro- (Zhang et al., 2021). Various agro-industrial effluents
duction (Wells Jr. et al., 2015). Zhang et al. (2018) have been explored for their ability to support yeast
concluded that the lipid obtained from fermentation growth and biomass accumulation (Table 6) with
cellulosic ethanol wastewater by Rhodotorula glutinis success. They have also been investigated as feed-
could be used as substrates for biodiesel production. stock for production of SCP with success. Crude
Fatty acid methyl esters spectrum of lipid accumulated protein content as high as 35.96% of dry cell weight
by Rhodosporidium kratochvilovae HIMPA1 dur- was recorded (Zhang et al., 2021). The SCP amount
ing fermentation of pulp and paper industry revealed and content produced varied with varying feedstock
45.43% of high quantity of long-chain monounsatu- and yeast strain. Iwuagwu and Ugwuanyi (2014) also
rated fatty acids and 15.91% of polyunsaturated fatty reported that the amino acid content of the Saccha-
acids (Patel et al., 2017). These types of fatty acids romyces sp. L31 biomass cultivated in POME was
improve biodiesel quality under low-temperature con- equivalent or even superior to the recommended FAO
dition in terms of low cold filter plugging, point along specification for SCP protein proposed for use as feed
with good oxidative stability, and cetane number (Patel for livestock.
et al., 2017). When dyestuff and textile effluent were
fermented with a consortium of Barnettozyma califor- Enzymes
nica SSA1518 and Yarrowia sp. SSA1642, the total
saturated fatty acid content of 36.09% was achieved, Enzymes are important catalyst used in several indus-
while the mono-unsaturated and poly-unsaturated fatty trial processes. Wastewaters with different composi-
acids were 45.44 and 18.30%, respectively, making the tion encourage the growth of microbes as well as
accumulated lipid valuable for biodiesel production induce enzyme production in them. Table 7 shows
(Ali et al., 2021). From these reports, fermentation of the enzymes produced from different wastewater.
wastewater by oleaginous yeast cells could generate The content of the wastewater and the fermenting
SCOs that are suitable for biodiesel production. yeast are some factors that determine the enzyme pro-
duced. Oil-based wastewater induced the production
Single cell proteins of lipase, while dyestuff and textile effluent induced
the production of CMCase and xylanase in addition
Single cell protein (SCP) is a protein derived from to lipase (Table 7). Al-Tohamy et al. (2020) produced
pure or mixed microbiological cultures such as algae, 8 different enzymes including laccase, xylanase, and
yeasts, moulds, or bacteria that can be used as feed for CMCase from salty azo dye wastewater using the
fish and animals as well as food for humans. SCP has yeast Sterigmatomyces halophilus SSA-1575. In a
been utilized extensively as a protein alternative for study by Broos et al. (2022), Rhodotorula kratoch-
traditional protein, particularly in animal feed. Pro- vilovae EXF7516 and Cutaneotrichosporon oleagi-
teins of microbial origin are generally highly nutri- nosum ATCC 20509 produced lignin peroxidase and
tious, cheap, and rapidly synthesized (Zhang et al., manganese peroxidase from phenolic wastewater.
2021). Microbes can ferment an array of agricultural Bleve et al. (2011) reported that three different strains
and industrial wastes with high biomass yield that is of Geotrichum candidum produced five different
enzymes including tyrosinase, cellulase, and laccase

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Table 6  Biomass yield and protein content of yeast cells cultivated in wastewater
Yeast strain Wastewater Biomass SCP profile Ref
accumulated
(g/L)

C. curvatus Tequila vinasses 5.19 Gonzalez-Garcia et al., 2011

R. glutinis 6.06
Trichosporon fermentans Refined soybean oil 7.9 Yu et al., 2018
wastewater
Rhodotorula glutinis Brewery 7.38 Schneider et al., 2013
wastewater + glucose
Metschnikowia pulcher- Distillery spent 34 Tamilalagan et al., 2019
rima MTCC 632 wash + lignocellulosic
hydrolysate
Nectaromyces rattus Biogas slurry 12.58 Protein content (35.96%) Zhang et al., 2021
19 distinct amino acids
made up 46.85% of cell
dry weight, with proline
accounting for 12.0%
Rhodosporidium toruloides Secondary brewery waste- 42.5 Dias et al., 2020
water + with sugarcane
molasses
R. toruloides Distillery wastewater 8.12 Ling et al., 2013
Yarrowia lipolytica TISTR Palm oil mill effluent 7.22 Louhasakul et al., 2016
5151
Rhodotorula glutinis Palm oil mill effluent 8.82 Saenge et al., 2011
S. cerevisiae MAK-1 Molasses and 10% OMW 7.3 Sarris et al., 2014
Yarrowia lipolytica ACA- OMW/glucose blend 13.3 Sarris et al., 2017
YC5033
Yarrowia lipolytica ACA- Olive mill wastewaters 12.48 Sarris et al., 2019
DC 5029 blended with crude
glycerol
C. pseudolambica 2.19
Candida utilis Potato wastewater with 5% 33.25 Protein yield (12.2 g/L) Kurcz et al., 2018
glycerol
I. occidentalis Livestock wastewater 7.3 Chung et al., 2016
Rhodotorula glutinis Cassava waste water 10.28 Ribeiro et al., 2019
Rhodotorula glutinis Corn steep water/corn 26.38 Liu et al., 2016
gluten water
Rhodotorula glutinis Cellulosic ethanol waste- 11.31 Zhang et al., 2018
water
Candida tropicalis Soy molasses 10.83 6.11 g/L of total protein Gao et al., 2012
SCP comprised 56.42%
crude protein and 5.28%
nucleic acid, respectively,
and was rich in vital
amino acids
R. kratochvilovae HIMPA1 Pulp and paper industry 13.87 Patel et al., 2017
effluent + glucose
Consortium of Yarrowia sp. Dyestuff and textile effluent 11.38 Ali et al., 2021
SSA1642, Barnettozyma
californica SSA1518,
and Sterigmatomyces
halophilus SSA1511

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Table 6  (continued)
Yeast strain Wastewater Biomass SCP profile Ref
accumulated
(g/L)

R. opacus PD630 Corn stover effluent 2.20 1.16 g/L total protein Mahan et al., 2018
equivalent to 52.7%
protein content
R. opacus DSM 1069 2.66 1.25 g/L total protein
equivalent to 47% protein
content
Trichosporon dermatis Butanol fermentation 7.4 Peng et al., 2013
wastewater
S. cerevisiae CCMA 0188 Vinasse 73.2 53.31% protein and 2.3% Pires et al., 2016
and CCMA 0137 nucleic acid
L. starkeyi Palm oil mill effluent 7.08 Karim et al., 2021a, b
Saccharomyces sp. L3 Palm oil mill effluent 4.42 27% crude protein content Iwuagwu & Ugwuanyi, 2014
Trichosporon domesticum Deproteinated potato 14.42 Gientka et al., 2019
PCM 2960 wastewater
C. parapsilosis CCMA Spirit beverages vinasse 8.89 26.93% total protein Coimbra et al., 2021
0544
Meyerozyma caribbica 5.12 27.30% total protein
CCMA 0198

from oil mill wastewater (Table 7). Wastewater could biosurfactant from wastewater offers a cheap substrate
therefore constitute cheap raw material for the pro- as well as remediation of the wastewater and produc-
duction of so many important enzymes and thus tion of value-added product. As recently reviewed by
should be explored for enzymes with novel qualities. Fernandes et al. (2023), yeast cells have potential to
produce biosurfactant from various substrate. Carote-
Other metabolites noids, organic acid (citric acid), aromatic compounds
(2-phenylethanol), and alcohols (ethanol, erythritol,
Table 8 shows some other important metabolites that mannitol, and arabitol) are important metabolite that
have been produced from wastewater fermentation by are used in various industries including health, cos-
yeast. Biosurfactant, carotenoids, organic acid (citric metics, biochemical, food and animal feed, biomate-
acid), aromatic compounds (2-phenylethanol), alco- rials, and biofuels industries. As detailed in Table 8,
hols (ethanol, erythritol, mannitol, and arabitol), and only limited studies are available on the produc-
others are some of the important metabolites gener- tion of these important metabolites from wastewater
ated from wastewater fermentation by yeast. Bio- using yeast strains. Thus, there is need for research to
surfactants are naturally occurring molecules that explore wastewater as substrate for the production of
have hydrophilic and hydrophobic chemical moieties these metabolites. If well harnessed, yeast-mediated
which enable them to reduce the surface tension and valorization of wastewater could be an affordable
form micelles (Gudiukaite et al., 2021). They are a technology for synthesis of some otherwise expensive
versatile group of chemicals with potential applica- metabolites.
tion in the production of paints, detergents, pharma-
ceuticals, paper products, cosmetics, petroleum, food, Limitations and way forward
water recycling, and remediation of environmental
pollution (Akbari et al., 2018). Biosurfactants are One major limitation of wastewater valorization by
environmental-friendly, biodegradable, and lower yeast cells is the toxic nature of this waste. Although
in toxicity compared to other chemical compounds yeast cells are known to have good tolerance to inhib-
of same function (Pi et al., 2017). Production of itors, nevertheless, when there is a combination of

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Table 7  Enzyme production by yeast cultivated in wastewater

Yeast strain Wastewater Enzyme produced Enzyme activity Ref

Yarrowia lipolytica TISTR POME Cell-bound lipase 4081 (U/L) Louhasakul et al., 2016
5151
Yarrowia lipolytica W29-N6 Olive oil mill wastewater Lipase 77.7 (U/L) Araujo et al., 2005
C. rugosa and Y. lipolytica POME Lipase - Theerachat et al., 2017
Lacasse
62 different strains of Y. Olive mill wastewater lipase 0U to 2315 U Lanciotti et al., 2005
lipolytica
3 different strains of Olive mill wastewater Cellulase-, xylanase-, - Bleve et al., 2011
Geotrichum candidum lignin-, and manganese-
dependent peroxidase;
laccase; Tyrosinase
C. tropicalis10456 Cellulase
Lipomyces starkeyi Olive oil mill wastewaters Lipase 1040 U/mL Yousuf et al., 2010
Rhodotorula kratochvilovae phenolic wastewaters Lignin peroxidase 3.9 U/L Broos et al., 2022
EXF7516 Manganese peroxidase 0.37
Cutaneotrichosporon Lignin peroxidase 5.5 U/L
oleaginosum ATCC 20509 Manganese peroxidase 0.25 U/L
Sterigmatomyces halophilus Salty azo dye wastewater NADH-DCIP reductase 43 U Al-Tohamy et al., 2020
SSA-1575 Azoreductase 4.98 U
Lignin peroxidase 0.452 U
Laccase 0.367 U
Manganese peroxidase 0.17 U
CMCase 8.81 U
Xylanase 7.378 U
16 different yeast strains Municipal, inosine Lipase 500 to 2500 U Yang et al.,
14 different yeast strains fermentation, papermak- Protease 350 to 12,140 U 2013
ing, antibiotic fermenta-
55 different yeast strains Manganese-dependent 19 to 2154 U
tion, printing, and dyeing
peroxidase
wastewater
11 different yeast strains Lignin peroxidase 36 to 231 U
Consortium of Yarrowia sp. Dyestuff and textile effluent Lipase 170.3 U/mL Ali et al., 2021
SSA1642, Barnettozyma CMCase 5.41 U/mg
californica SSA1518, Xylanase 7.6 U/mg
and Sterigmatomyces
halophilus SSA1511

these inhibitors just like in wastewater, it exerts more wastewaters. In view of this, a mixture of wastewa-
inhibitory effects on the fermenting yeast. In order to ters may be a solution to this limitation. There have
overcome this limitation, a yeast strain with multi- been several reports on increase in bioremediation
stress tolerant ability is necessary. Also the use of efficiency as well as product yield when mixtures of
yeast consortium may also be beneficial to overcom- wastewaters are used.
ing this limitation. Furthermore, immobilization of Although yeast cells have been used successfully
yeast especially by entrapment on a matrix can also for valorization of wastewaters, some of the results
be explored given to the fact that entrapment protects are characterized with low remediation efficiency as
cells from inhibitory compounds. well as low product yield. This could be as a result of
Some of these wastewaters are lean in nutrient several factors including nutrient and inhibitor con-
content such that the available essential nutrient may tent of the wastewater, bioremediation ability of the
not support the proper growth and metabolic activi- yeast strain, fermentation condition, and other prevail-
ties of these yeast strains. Nutrient supplementation ing conditions. It is thus important to put these factors
is therefore needed for effective fermentation of these into consideration while choosing a yeast strain for

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Table 8  Other metabolites produced from wastewater fermentation by yeast cells

Yeast strain Wastewater Metabolite Concentration Reference

Rhodotorula glutinis Brewery wastewater + glucose Carotenoid torularhodin 1.2 mg/L Schneider et al., 2013
(47.7%) b-carotene (51.6%)
Rhodosporidium toruloides Secondary brewery Carotenoid 0.23 mg/g Dias et al., 2020
wastewater + with sugarcane
molasses
Rhodotorula glutinis Palm oil mill effluent Carotenoid 176.45 mg/L Saenge et al., 2011
S. cerevisiae MAK-1 Molasses and 10% OMW Ethanol 34.3 g/L Sarris et al., 2014
S. cerevisiae MAK-1 Olive mill wastewater Ethanol 52 g/L Sarris et al., 2013
Yarrowia lipolytica ACA-DC Olive-Mill wastewaters Erythritol 66 g/L Sarris et al., 2019
5029 blended with Mannitol 10.1 g/L
crude glycerol
Arabitol 4.5 g/L
Citric acid 79 g/L
Rhodotorula glutinis Cassava waste water Carotenoid 0.98 mg/L Ribeiro et al., 2019
Saccharomyces cerevisiae Cassava wastewater + 20 g/L 2-Phenylethanol 1.33 g/L Oliveira et al., 2015
glucose
Candida bombicola Cassava wastewater Sophorolipids (biosurfactant) 300 g/L John & John, 2015
C. parapsilosis CCMA 0544 Spirit beverages vinasse 2-phenylethanol 3.87 g/L Coimbra et al., 2021
Yarrowia lipolytica ACA-YC OMW/glucose blend Citric acid 52 g/L Sarris et al., 2017
5033
20 different strains of Y. Olive mill wastewater Citric acid 0.8–5.2 g/L Lanciotti et al., 2005
lipolytica

the valorization process. The use of yeast consortium wastewater. In a study by Theerachat et al. (2017), the
may be more beneficial than using single cells. Yeast mixture of crude extracellular lipase and laccase prep-
can be used in combination with other yeast cells or arations showed the maximum COD removal efficacy
other microbes such as bacteria, mould, and microal- of 64.1% from POME, followed by crude lipase alone
gae. For instance, Ling et al. (2014) reported that a (61.3%) and crude laccase alone (59.4%). The whole
co-culture of the yeast (Rhodosporidium toruloides) cell in combination with these products can also be
and the microalgae (Chlorella pyrenoidosa) achieved explored for effective valorization of wastewater. It is
higher lipid content as well as improved reduction important however to consider microbial interactions
efficiency of total nitrogen, COD, and total phospho- when constructing yeast consortium to avoid antago-
rus compared to pure culture of the yeast. According nistic interaction.
to Theerachat et al. (2017), the co-culture of Y. lipo- Development of robust strains capable of effi-
lytica and C. rugosa removed the most triglyceride ciently valorizing wastewater with the use of biologi-
(98.5%) and COD (60.3%) from undiluted POME. cal tools should be intensified. This can be achieved
A maximum biomass, lipid production, and COD either by modifying these yeast cells that have shown
reduction were achieved when POME was fermented good potential to do better or using them as cell fac-
with co-culture of the yeast (Lipomyces starkeyi) tory for modifying other yeast strains with desir-
and the bacterium (Bacillus cereus) which were sub- able traits. Beyond genetic modification, evolutionary
stantially higher than those of the mono-cultures studies can probably be used to develop robust strains
(Karim et al., 2021a, b). Yeast products such as or yeast consortium for valorization of wastewater.
extracellular enzymes or biosurfactant can be used Furthermore, nature should also be continuously
instead of whole cell. Tsilo et al. (2022) reported explored for yeast cells with good potentials for val-
that the bioflocculant produced by Pichia kudria- orization of wastewater.
vzevii MH545928.1 achieved 43% COD, 64% BOD, The GRAS status of some of the non-conven-
73% P, and 50% N removal efficiencies in coal mine tional yeast strains is a major hindrance to their use

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especially as single cell protein for food and feed as Authors’ contributions Ndubuisi, Ifeanyi Amara: Conceptu-
well as production of valuable metabolites for other alization, Paper Resources, Data curation, Writing- Original draft,
Writing- editing, Validation. Enemour, Simeon Chukwuemeka:
industrial processes. Therefore, there is need to ascer- Conceptualization, Paper Resources, Writing- Original draft,
tain the GRAS status of as many non-conventional Validation. Nnabuife, Obianuju Obiajulu: Paper Resources,
yeast strains as possible. Writing- review and editing, Validation. Ogbonna, James Chuk‑
Wastewater valorization with yeasts is a low-cost, wuma: Paper Resources, Writing- review and editing, Validation.
sustainable method of eliminating contaminants from
Availability of data and material All data stated in this
wastewater while also providing value-added prod- review have been referenced.
ucts. Despite its potential at the lab scale, no scale-
up study has been published. Larger-scale wastewater Declarations
valorization employing yeast strains may be depend-
ent on a variety of parameters that go beyond what is Competing Interests The authors declare no competing interests.
available at the lab scale. Thus, studies on optimiza-
tion as well as implementation of various strategies
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