1 Valorization of dairy waste and by-products through microbial bioprocesses

2 Zeba Usmani1#, Minaxi Sharma1#, James Gaffey2, Monika Sharma3, Richard J. Dewhurst4,

3 Benoît Moreau5, John Newbold6, William Clark7, Vijay Kumar Thakur8,9,10, Vijai Kumar

4 Gupta8,11*

1
5 Department of Applied Biology, University of Science and Technology, Meghalaya 793101,
6 India
2
7 Circular Bioeconomy Research Group, Shannon Applied Biotechnology Centre, Munster
8 Technological University, Kerry, Ireland
3
9 Department of Botany, Shri Awadh Raz Singh Smarak Degree College, Faizabad University,
10 UP, India
4
11 Dairy Research Centre, SRUC, Kings Buildings, West Mains Road, Edinburgh EH9 3JG,
12 UK
5
13 Laboratoire de "Chimie verte et Produits Biobasés", Haute Ecole Provinciale du Hainaut-
14 Département AgroBioscience et Chimie, 11, rue de la Sucrerie, 7800 Ath, Belgique
6
15 Dairy Research Centre, SRUC, Dumfries, UK
7
16 Zero Waste Scotland, Moray House, Forthside Way, Stirling FK8 1QZ, UK
8
17 Biorefining and Advanced Materials Research Center, SRUC, Kings Buildings, West Mains
18 Road, Edinburgh EH9 3JG, UK
9
19 Department of Mechanical Engineering, School of Engineering, Shiv Nadar University,
20 Uttar Pradesh 201314, India
10
21 School of Engineering, University of Petroleum & Energy Studies (UPES), Dehradun,
22 Uttarakhand, India
11
23 Center for Safe and Improved Food, SRUC, Kings Buildings, West Mains Road, Edinburgh
24 EH9 3JG, UK (vijaifzd@gmail.com, Vijai.Gupta@sruc.ac.uk)
25 *Corresponding author
26 #Equal contributors
27

1
28 Abstract

29 Waste is an inherent and unavoidable part of any process which can be attributed to various

30 factors such as process inefficiencies, usability of resources and discarding of not so useful

31 parts of the feedstock. Dairy is a burgeoning industry following the global population growth,

32 resulting in generation of waste such as wastewater (from cleaning, processing, and

33 maintenance), whey, and sludge. These components are rich in nutrients, organic and

34 inorganic materials. Additionally, the presence of alkaline and acidic detergents along with

35 sterilizing agents in dairy waste makes it an environmental hazard. Thus, sustainable

36 valorization of dairy waste requires utilization of biological methods such as microbial

37 treatment. This review brings forward the current developments in utilization and valorization

38 of dairy waste through microbes. Aerobic and anaerobic treatment of dairy waste using

39 microbes can be a sustainable and green method to generate biofertilizers, biofuels, power,

40 and other biobased products.

41 Keywords: Dairy waste, microbial processing, biodegradation, circular economy, biomass

42 valorization

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43 1 Introduction

44 The rapid growth of global population in the last couple of decades has led to a rapid pace of

45 industrialization of the sectors such as agriculture, animal farming, food production and

46 processing. Dairy is a part of the food industry which serves the need of global populace with

47 products such as milk, cheese, butter, ghee, milk powder etc. (Jaganmai & Jinka, 2017). The

48 demand for dairy products has significantly risen across many countries resulting in setup of

49 large-scale production plants (Chokshi, Pancha, Ghosh, & Mishra, 2016). As reported by

50 FAO the global milk production stood at ~906 million tonnes led by India with a production

51 of ~196 million tonnes. Milk products such as cheese, whey, and whole milk powder, rose in

52 trading globally while butter and skim milk powder fell in trade. In European Union

53 production rose by 1.6 percent to 236 million tonnes (FAO, 2020).

54 The dairy industry while catering to the needs of the population has led to significant amount

55 of pollution due to heavy use of water and emission of effluents into the environment. The

56 annual dairy waste residue in the form of solid waste and effluents stands at ~4–11 million

57 tonnes globally. Additionally, in Europe alone, nearly 29 million tonnes of dairy product end

58 up as waste annually. This waste mainly constitutes of processing waste, spoiled products,

59 and mishandled products and materials (Mahboubi et al., 2017). The processing of every

60 volume of milk results in expulsion of 1-3 times of effluent into the environment, which

61 amounts to 3.739–11.217 million cubic meters of waste annually. A by-product of cheese

62 production is whey which is produced in large amounts i.e. 9 kg for every kg of cheese

63 (Parashar et al., 2016). Various other materials such as proteins, fats, sugars, food additives,

64 and cleaning agents are left behind either in the effluents or the sludge during dairy

65 processing. The spoilage of dairy products can be attributed to their higher level of absorption

66 capability and rich composition, which is favourable for bacterial and fungal contamination

67 resulting in off-odour and bad flavour development.

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68 1.1 Dairy waste categories

69 There are two broad categories of dairy waste viz. effluent (wastewater) and solid waste.

70 Every stage in the dairy production and processing contributes to a significant amount of

71 waste generation i.e., from dairy farms to transportation to processing and packaging of

72 products. The human consumption of dairy products necessitates cleanliness in dairy

73 processing units which leads to usage of cleaning agents that eventually end up in the waste

74 effluents and solid wastes. The breakdown of the milk proteins in waste raw milk which

75 contains various organic and inorganic functional groups of nitrogen including NO −2, NO−3,

76 NH+4 (14-830 mg/L of total nitrogen concentration) gets converted to nitrite thus

77 contaminating groundwater causing methemoglobinemia (Kushwaha et al., 2011; Kavitha et

78 al., 2019). The dairy effluent is highly rich in organic matter which is favourable for the

79 sustenance of microbial communities thus facilitating the production of various useful

80 products through microbial treatments (Lappa et al., 2019). Whey is a major component in

81 dairy effluent and is primarily composed of lactose (4-5%) (Slavov, 2017). Dairy wastewater

82 is typically characterized by high chemical oxygen demand (COD) and biological oxygen

83 demand (BOD) in the range of 1-10 g/L and 0.3-5.9 g/L, respectively (Kothari et al., 2017;

84 Yonar et al., 2018). This is the side effect of high organic content which leads to a rapid

85 depletion of dissolved oxygen when the effluent is discharged directly into aqueous systems,

86 which inadvertently harms the aquatic lifeforms and in turn the overall environment. The

87 high BOD and COD of the dairy effluent also makes the lactose from whey highly harmful to

88 the environment (Slavov, 2017). The oily and greasy fat effluents form a surface film on

89 water bodies preventing the oxygen transfer thus negatively impacting the aquatic flora and

90 fauna (Kavitha et al., 2019). A fully scaled dairy production unit can generate around 200–

91 350 kg of sludge by processing approximately 5 hundred thousand litres of milk daily

92 (Kavitha et al., 2019). This sludge is broadly classified as biological and chemical sludge and

4
 93 consists of non-biodegradable solids and biodegradable organic materials. Handling this solid

94 waste residue is a huge challenge and can cost up to 60% of total treatment cost in the

95 processing unit (Kwapinska et al., 2020).

96 1.2 Treatment of dairy waste

97 The general routes for treatment of dairy waste include physicochemical and biological

98 methods, but the cost-effective and environment-friendly nature of biological methods make

99 them preferable over physicochemical techniques. There has been significant growth in the

100 number of studies that have explored the valorization of dairy waste into useful products and

101 they have been able to successfully demonstrate the effectiveness of bio-based techniques in

102 establishing a biorefinery (Wong et al., 2019; Chandra et al., 2018; Ganju & Gogate, 2017).

103 In fact, the rich organic nature of dairy waste makes it a valuable feedstock for production of

104 value-added products (Table 1). Various anaerobic and aerobic treatment techniques such as

105 Activated Sludge Process, Sequential Batch Reactors (SBR), Moving Bed Biological Reactor

106 (MBBR), Completely Stirred Tank Reactor (CSTR), Up-flow Sludge Anaerobic Blanket

107 (USAB), Fluidized Bed Reactor (FBR), wetlands, and co-composting have been deployed to

108 biologically treat dairy waste (Kwapinska et al., 2020; Ahmad et al., 2019; Zhou et al., 2015).

109 Many factors determine the efficacy of the biobased techniques, including substrate type and

110 characterization, operational parameters and the performance of the microbes themselves. In

111 fact, anaerobic conversion of dairy waste is significantly impacted by the syntrophic activity

112 of the microbes active in different phases of anaerobic digestion (Pagliano et al., 2019). Dairy

113 effluent has been demonstrated to be a beneficial substrate for enzymatic hydrolysis using

114 Saccharomyces cerevisiae to produce ethanol (Parashar et al., 2016). The enzymes produced

115 by filamentous fungi can hydrolyse complex carbohydrates present in dairy waste into simple

116 sugars aiding in production of high-quality biomass suitable as animal/fish or even human

117 consumption (Mahboubi et al., 2017).

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118 Table 1: Production of bio-based products from dairy waste

119 However, the formation of sludge in anaerobic treatments can be a potential inhibitor to

120 its large-scale implementation as the remediation cost becomes higher along with its ability to

121 absorb toxic and organic material from the feedstock itself (Dabrowski et al., 2017). Based on

122 the oxygen requirement of the process biobased treatments can be categorized as anaerobic or

123 aerobic. Most of the dairy effluent treatments are aerobic in nature, although they are not the

124 most efficient processes due to high acidification and growth of filamentous fungi (Ahmad et

125 al., 2019). The most commonly used reactor for dairy wastewater processing is SBR where

126 the entire process is carried out in a single tank. But the drawback of this reactor type is the

127 inhibition of the overall process due to presence of nutrients such as phosphorus and nitrogen

128 (Slavov, 2017). An alternate approach suggested by Gil-Pulido et al. (2018) was to

129 intermittently aerate the SBR to remove the nutrients at various time intervals thus improving

130 the overall efficiency of the process. Anaerobic digestion systems have proven to be an

131 effective method in this regard with two phase anaerobic digestion taking the lead in being

132 highly efficient at both degradation and energy recovery (Cydzik-Kwiatkowsk and Zielinsk,

133 2016). But hydrolysis of lipids during the anaerobic digestion of fat/oil rich dairy waste

134 inhibits the overall process, which can be overcome by addition of enzymes (Meng et al.,

135 2017). Overall, a proper combination of anaerobic and aerobic treatments can help in

136 efficient valorization of dairy waste.

137 This review paper brings together current developments in the valorization of dairy

138 waste into value-added products. There are several microbial techniques and strains that have

139 emerged in the recent past to develop bio-based products such as bioplastics, biosurfactants,

140 bioenergy etc. This review summarizes the challenges and future perspectives in deployment

141 of dairy waste and can be useful to suggest the research opportunities to develop the

142 microbial techniques further for scaled-up industrial application.

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143 2 Value-added products from dairy waste

144 Almost entire amount of organic matter present in dairy effluent is biodegradable. But

145 individual biological treatments when applied to the effluent do not completely degrade the

146 organic fraction, leaving behind some residue which allows for insects and pathogens to

147 breed while also causing bad smell (Kothari et al., 2017). Thus, deploying integrated

148 bioremediation strategies to degrade the dairy waste along with efficient production of energy

149 rich products such as biohydrogen, biodiesel and biomethane is important. Bioconversion of

150 dairy waste into value-added products also aligns with current European policies on circular

151 economy to promote sustainable production and waste reduction. There is a growing interest

152 among researchers and industries alike on deploying biological technologies for the

153 production of products such as biofuels, bioplastics, biosurfactants and other biobased

154 platform chemicals. Figure 1 gives an overview of few of the prominent microorganism

155 species that have been deployed in this regard.

156 Figure 1: Prominent microorganism species deployed in bioconversion of dairy waste

157 2.1 Biohydrogen

158 The zero-emission nature of hydrogen has earned it the title of clean and green energy with

159 water being its only combustion by-product. The application of hydrogen ranges from power

160 generation through fuel cells to be used as fuel in vehicles (Chan et al., 2016). Another area

161 that has been explored is the combination of hydrogen production with other bio-based

162 technologies to sustainably produce energy. For instance, integrating the production of

163 methane with hydrogen from the biodegradation of organic waste such as dairy waste, food

164 waste and other organic industrial waste has been touted as the next stage in developing green

165 energy from waste (Aziz, 2016; Kothari et al., 2017). The development of the two-stage

166 approach integrating reduction of dairy wastewater pollution with bioenergy generation

167 (methane and hydrogen) via two-phase process has been shown to be a cost-effective and

7
168 highly productive process in energy recovery (Zhong et al., 2015; Chu and Wang, 2017).

169 Despite these obvious advantages, few downstream and upstream parameters such as the

170 choice of bacterial culture for the dairy waste treatment and their biohydrogen production

171 efficiency still need to be studied in depth to test the large-scale feasibility and setup of these

172 integrated techniques.

173 Microbial bioprocesses effectiveness in dairy waste remediation principally depends on the

174 substrate utilization pattern and the kinetics involved in the dairy biomass utilization which

175 delivers the key evidence during the dairy waste remediation (Kumar et al., 2016). Different

176 types of microbial populations including bacteria, fungi, algae are known to produce

177 biohydrogen using dairy waste biomass as their carbon source in anaerobic conditions. The

178 microbial bioprocesses for biohydrogen production are generally of two types: photo

179 fermentation (PF) and dark fermentation (DF). The dark fermentation (DF) is most preferred

180 technology for the biohydrogen production as the by-products (such as volatile fatty acids)

181 obtained from these fermentations are quickly converted into biohydrogen via photo-

182 fermentation (Montiel Corona et al., 2017). The sequential integration of PF and DF with a

183 targeted microbe concurrently generates biohydrogen, CO 2, organic acids, soluble

184 metabolites, and alcohols during fermentation. From these, the soluble metabolites are key

185 substrate which is further utilized by photosynthetic bacteria to produce biohydrogen

186 (Ghimire et al., 2015). Dairy wastewater from cheese industry is an enriched source of carbon

187 can be a prominent substitute for microbial fermentation as it contains 5% lactose (Chong et

188 al., 2009). Several studies (Carrillo-Reyes et al., 2012; Nath et al., 2008) have been

189 conducted to check the biohydrogen production potential of wastewater from dairy industry

190 using different microbial populations. Collet et al. (2004) investigated the biohydrogen

191 production with an increased production about 5 mmol H 2 (g cell)-1 h-1 using Clostridium

192 thermolacticum. The potential of biohydrogen production of dairy effluent coupled with

8
193 effluent remediation was investigated by Vekata Mohan et al (2007). They observed that the

194 biohydrogen production was highest about 1.105 mmol H2/m3/min at organic loading rate 3.5

195 kg COD/m3 day with the removal of 64.7% COD. Yang et al. (2007) reported the maximum

196 biohydrogen production (1.8 and 2.3 mM/g COD) with various loading rates at 24 h

197 hydraulic retention time (HRT) using mixed microbial culture. However, the production of

198 biohydrogen is controlled in various ways such as processing conditions, nature of substrate,

199 organic loading rate, nature and types of microbes, reactors used for fermentation etc. In a

200 study Silva et al. (2019) examined the effect of organic loading rate (OLR) and pH on the

201 production of biohydrogen from dairy effluent via natural fermentation in an anaerobic

202 fluidized bed reactor. According to their study, pH range was inversely related to the

203 biohydrogen production yield. It is also reported that temperature has a significant effect on

204 biohydrogen yield, degradation of substrate, formation of subproducts and growth of the

205 microbial community (Perna et al., 2013). Mu et al. (2006) reported that yield of biohydrogen

206 and degradation of substrate accelerated with rise in temperature but very high temperature

207 than an average (ambient temperature) is not favourable for either microbial growth or

208 recovery of energy. Therefore, to establish a balance between biohydrogen production and

209 energy recovery, a reasonable temperature i.e. ambient temperature is required for

210 fermentation. The temperature also affecting the solubility of biohydrogen (Silva et al.,

211 2019). Gavala et al., (2006) reported an increase in biohydrogen production (131.5 mL H2

212 g/CODremoved) at 60℃ when compared at 40 ℃ (116.5 mL H2 g/CODremoved). Methane

213 production is also playing an inhibitory step on the biohydrogen production in the

214 fermentation processes (Silva et al., 2019).

215 2.2 Biofuels

216 Biofuels have emerged as a promising alternative to non-renewables due to their better

217 emission profile, biodegradability, and overall contribution to sustainability. Biodiesel for

9
218 instance is a clean fuel that contains no sulphur or other toxic compounds, and its combustion

219 causes much lower emission of particulate matter, hydrocarbons, and carbon monoxide

220 (Bhatia et al., 2018). Production of ethanol from dairy waste as a substrate can be carried out

221 using a range of yeasts such as Kluyveromyces marxianus URM 7404 (Murari et al., 2019),

222 Candida inconspicua W16 (Eloutassi et al., 2018) and Kluyveromyces lactis CBS2359

223 (Sampaio et al., 2020). Kluyveromyces sp. yeast has shown its capability to directly convert

224 the lactose present in dairy waste specially cheese whey permeates and wastewater into

225 ethanol. Additionally, it also presents the advantage of high fermentation capability on a

226 range of carbohydrates e.g. pentoses, hexoses, and disaccharides as well as high metabolic

227 activity, faster growth, and high temperature resistance making it particularly suitable for

228 industrial fermentation application (Murari et al., 2019a). Kluyveromyces marxianus can

229 hydrolyse the lactose present in the dairy wastewater to produce 20.6 g/L of ethanol under

230 optimal conditions of 32-35oC and pH range 4.8-5.3 (Murari et al., 2019b). It was also

231 reported that operational parameters such as pH, temperature and lactose concentration in the

232 substrate determine the ethanol production of which temperature was the most important

233 factor at controlling the final output. Another way is to utilize dairy waste as a substrate and

234 growth media for microbes such as oleaginous yeasts in the production of microbial

235 biodiesel. It is owing to the high carbohydrates content along with low nitrogen present in the

236 dairy waste. Taskin et al. (2015) were the first to report culturing of cold-adapted oleaginous

237 yeast Yarrowia lipolytica B9 in a non-sterile whey media. They were able to attain a lipid

238 production of 4.29 g/L with 80.54% of the lipids being monounsaturated fatty acids, making

239 it highly suitable for biodiesel production. In another study, ricotta cheese whey was used by

240 Carota et al. (2017) as a substrate to grow Cryptococcus curvatus NRRL Y-1511 and

241 Cryptococcus laurentii UCD 68-201 yeasts which produced high amount of lipids (6.8 and

242 5.1 g/L) and oleic acid (52.8 vs. 48.7%), respectively. Another yeast Aspergillus candidus

10
243 when grown on dairy waste whey, produced biomass with the required polyunsaturated

244 (38.3%), monounsaturated (31.5%) and saturated methyl esters (30.2%) profile making dairy

245 waste a suitable substrate for biodiesel production (Kakkad et al., 2015). On the other hand,

246 Chan et al. (2018) cultivated Mucor circinelloides, a filamentous fungus, on cheese whey

247 permeate and reported that hydrolysis of lactose in the permeate was favourable to biomass

248 production which increased from 2.4 g/L to 7.8 g/L. Chowdhury and Franchetti (2017)

249 developed a lifecycle assessment (LCA) based approach to evaluate the feasibility of using

250 algal biomass grown on dairy industry waste for bioenergy production. They concluded that

251 biooils produced by growing microalgae on dairy wastewater can produce approximately

252 1.12×109 Gigajoules of energy per year through a mix of other energy extraction

253 technologies. It can thus be safely concluded that biofuel production from microbial

254 cultivation on dairy waste is a feasible technique, but the dependence on temperature, pH,

255 lipid production and further treatment of the biomass thus produced needs to be further

256 explored.

257 2.3 Biosurfactants

258 Various microbes such as bacteria, fungi and yeast are capable of producing biosurfactants

259 also known as surface-active agents which can be used in place of other organic surfactants

260 (Gaur et al., 2019). The global market for biosurfactants has been estimated to stand at USD

261 30.64 billion (Singh et al., 2019). Pseudomonas, Candida, Rhodococcus, and Bacillus genera

262 of microbes have been majorly used for biosurfactant production in various studies and pilot

263 scale implementations (Santos et al., 2016). Typically, a biosurfactant consists of a

264 hydrophobic and a hydrophilic component (Almeida et al., 2018; Otzen, 2017).

265 Biosurfactants can be classified into two types viz. high and low molecular weight

266 biosurfactants depending upon their molecular weight, physicochemical properties, and mode

267 of action (Jimoh and Lin, 2019). The high-molecular-weight biosurfactants can stabilize oil-

11
268 in-water emulsions and are used as bioemulsifiers whereas low-molecular-weight ones can

269 reduce interfacial and surface tension (Sałek and Euston, 2019; Singh et al., 2020). Among all

270 currently recognised biosurfactants, Pseudomonas species and Pseudomonas aeruginosa in

271 particular has proven to be the most beneficial genera in the production of biosurfactants.

272 This could be attributed to their high tolerance to extreme environmental conditions (Varjani

273 et al., 2020). Biosurfactants such as Sophorolipids, Rhamnolipids, Trehalose lipids, and

274 Ornithine lipids have all been demonstrated to be produced by various microbial cultures

275 using organic waste as substrates (Varjani et al., 2020).

276 Dairy waste is rich in proteins and moisture allowing for the growth of bacterial

277 communities which can produce biosurfactants by using the waste as the source of carbon.

278 Panner (Indian cheese) production is a major waste generator in the Indian dairy industry

279 with 20 kg of waste whey produced for every 1.5 kg of paneer produced, which is usually left

280 untreated and released from the production facility as waste into the environment harming the

281 surrounding land and water bodies (Parashar et al., 2016). Patowary et al. (2016) isolated

282 Pseudomonas aeruginosa SR17 from a hydrocarbon-contaminated soil and used it to produce

283 biosurfactant from cheese (paneer) whey. They found that under regular treatment without

284 any additional co-substrate the biosurfactant yield stood at 2.7 g/L whereas the addition of

285 mineral salts and glucose increased the output to 4.8 g/L. Application of Lactococcus lactis

286 CECT-4434 on 15% whey by Vera et al. (2018) resulted in 8.9 mg/L production of

287 biosurfactant, and surface tension reduction by ~18.1 mN/m. Dharmik et al. (2017) tested the

288 production of biosurfactants from agro-industrial & dairy wastes using Pseudomonas

289 aeruginosa and Bacillus subtilis. They were able to produce surfactant using Bacillus

290 subtilis (10% inoculum) with highest surface tension reduction of ~50% (against 5% and 7%

291 inoculation which showed 14% and 20% reduction). Using whey waste as a growth media,

292 lactic acid bacterium Lactobacillus acidophilus was able to produce biosurfactant with an

12
293 emulsification index of 58% and a 10 mg/mL dose of biosurfactant produced was able to

294 prevent the adhesion of Streptococcus aureus, Pseudomonas aeruginosa, and Escherichia

295 coli pathogens (Alkan et al., 2019). Daverey and Pakshirajan (2016) used Candida

296 bombicola yeast to produce sophorolipids using dairy wastewater as the substrate and were

297 able to successfully remove the fats and oils from the wastewater (>95% COD removal).

298 Starmerella Bombicola is another microorganism which has been shown to effectively

299 produce biosurfactants from dairy wastewater (Jiménez-Peñalver et al., 2019).

300 While there is a significant market for biosurfactants, its industrial production from

301 dairy waste is posed with challenges such as low yield, high cost, feedstock availability, and

302 downstream purification and processing (Singh et al., 2019; Bertrand et al., 2018; Schultz and

303 Rosado, 2020). But the silver lining of utilizing dairy waste and integrating the production of

304 biosurfactants with dairy production plants is two-fold. First, it reduces the overall

305 biosurfactant production cost by 10-30% as the feedstock is easily available on premise,

306 second the biosurfactant produced could either serve as an additional source of income or

307 could be reutilized in the dairy processing plant thus bringing further economic advantages.

308 2.4 Bioplastics

309 Bioplastics are biopolymers which signifies their biodegradable nature and their production

310 from biological sources (Shankar and Rim, 2018). Naturally occurring molecules such as

311 proteins, amino acids, lipids and polysaccharides make up bioplastics (Sirohi et al., 2020). Its

312 similarity in characteristics with polypropylene makes it an excellent alternative to plastic

313 while also providing the additional advantage of biodegradation using enzymes or

314 microorganisms such as bacteria, fungi and yeasts. This prevents the hazardous effect of

315 plastic disposal in the environment with the final degradation products being water, CO 2, and

316 biomass. Poly-3-hydroxybutyrate (PHB) has gained significant interest from researchers and

317 industries as the biodegradable green alternate to plastics due to its excellent physical

13
318 characteristics, great blockage against gases, and high crystallinity (50-70%) (Yousuf, 2017).

319 But, their brittle, fragile and thermally instable nature limits their applicability in large-scale

320 application. There have been attempts to improve upon these drawbacks by use of chemical

321 grafting, polymer blending and introduction of co-polymers (Lamm et al., 2019; Rahman et

322 al., 2017). Another major roadblock is the high raw material cost making up for >50% of

323 total production cost. This could be alleviated by the use of waste materials from industries

324 such as food and dairy. A route for PHB production from dairy waste feedstock is to pre-treat

325 it using either biological, biochemical or physico-chemical methods. This is followed by

326 enzymatic hydrolysis and fermentation using microbes such as Bacillus subtilis, Ralstonia

327 eutropha and Azohydromonas australica etc to yield PHB.

328 In an early experiment Pais et al. (2014) investigated the usage of cheese whey in PHB

329 production. They inserted Cupriavidus necator P(3HB)-synthesis genes in Escherichia coli

330 strain CML3-1 to achieve very high biopolymer concentration of 7.8 g/L from cheese whey

331 making a viable alternate carbon source to enhance biopolymer production. On the other

332 hand, Colombo et al. (2016) were able to extract a high amount of polyhydroxyalkanoate

333 (PHA) bioplastic (60 g/kg) from whey using a mixed culture media. Sharma et al., (2017)

334 also demonstrated that PHB can be produced by the action of Azohydromonas australica

335 DSM 1124 on lactose sugar (2% w/v) forming around 2.07±0.2 g/L PHB in 72 hours. While

336 the pretreatment of waste before being subjected to bioplastics production will enhance the

337 quality of products, the cost associated with it is a blocker. But Israni et al. (2020) showed

338 that protease and lipase enzymes can valorize unprocessed dairy waste into valuable

339 bioplastics viz. PHA (2.20 ± 0.11 g/L). However, sustainable valorization of dairy waste into

340 PHB at low cost requires process optimization and tuning the process conditions. Biobased

341 production of PHA can be optimized for cost through waste recycling in an open loop

342 process. The residues (fermentation broth and cell debris) from the first processing cycle can

14
343 be reused in the next cycle as the started culture for fermentation thereby reducing the overall

344 cost of the process (Koller, 2015). In fact, whey from cheese production could be the most

345 cost-effective fermentation substrate to produce PHB bioplastic according to Pena-Jurado et

346 al. (2019). They simulated the production of PHB using Bacillus subtilis EPAH18 on cheese

347 whey at optimal operational conditions (37°C, 24 hours, 200 rpm agitation, 5% v/v inoculum

348 and 0.5% v/v ammonium sulphate) which resulted in the highest production of 0.54 g/L PHB.

349 This efficiency was achieved when the production of PHB was simulated for a 1000

350 tonne/hour plant. In another study, Prabisha et al. (2015) optimized the production of PHB

351 from dairy waste using Comamonas sp. isolated from a dairy wastewater and were able to

352 produce 0.523 g/g of PHB with an inoculum concentration of 5.5% v/v. The issue of

353 brittleness in the bioplastics can be addressed by creating co-product composite solutions in

354 bioplastic films. For instance, when κ-carrageenan, sodium carboxymethylcellulose

355 (NaCMC) and gelatin were added to casein from dairy wastewater, the resultant biopolymer

356 film was shown to exhibit enhanced physical properties over casein-only biofilm as well as

357 better degradability (Ryder et al., 2020). While further research is required to make

358 bioplastics an economically viable alternate to traditional plastics, the environmental impact

359 justifies the investment into developing these methodologies. In fact, a recent lifecycle

360 assessment (LCA) analysis of bioplastic production from whey protein projected the energy

361 consumption of 2900 MJ, CO2 emissions of 115.3 kg, NOx emission of 82.5 kg, making it

362 least negatively impactful to global temperature rise, and least toxicity to aquamarine

363 ecosystem when compared with traditional plastic (Chalermthai et al., 2021).

364 2.5 Organic acids

365 There is a consistent demand for organic chemicals as additives in the process of industrial

366 processing of food, beverages, and pharmaceuticals. They are used to preserve, stabilize,

367 emulsify, acidify and enhance flavours of consumables. Dairy waste such as whey in

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368 different forms can be successfully used to derive organic acids such as acetic, lactic,

369 succinic, citric and propionic acid (De Jesus et al., 2015). Table 2 gives an overview of some

370 of the microbial communities utilized in converting whey waste into organic acids. For

371 instance, Yarrowia lipolytica B9 yeast was used to produce citric acid from deproteinized

372 milk whey yielding 33.3 g/L at 20oC and pH 5.5 (Arslan et al., 2016). In another study

373 Louasté and Eloutassi (2020) performed a batch fermentation to produce succinic acid using

374 milk lactose and whey and Actinobacillus succinogenes 130Z. They were able to derive 35

375 g/L and 25 g/L of succinic acid using whey and lactose from dairy waste successfully.

376 Similarly, Propionibacterium could be used to produce propionic acid which is an important

377 chemical to preserve vitamin B12 during the production process (De Jesus et al., 2015). Jiang

378 et al. (2015) used immobilized Propionibacterium acidipropionici on whey lactose and were

379 able to achieve a propionic acid yield of 135±6.5 g/L in a fed-batch fermentation method.

380 They also found that genetically modifying the bacteria with enhanced trehalose synthesis

381 mutant gene further enhanced the propionic acid production. Another important organic acid

382 is lactic acid which can be produced from salted whey and whey permeates by the application

383 of Lactobacillus casei and Lactobacillus rhamnosus B-445. Dosuky Atiat et al. (2019) were

384 able to achieve maximum lactic acid production of 38 g/L at 37 oC. Acetic acid bacteria are

385 also able to convert whey-based ethanol into acetic acid by undergoing fermentation. But the

386 amount of acetic acid produced can vary significantly depending on the fermentation setup

387 used and the microbial strain used (Gullo et al., 2014). Cheese whey when treated with

388 Lactobacillus acidophilus to produce biohydrogen, also results in production of valuable

389 organic acids such as pyruvic, propionic, acetic, lactic, formic and butyric acid (Pandey et al.,

390 2019).

391 Table 2: Production of organic chemicals from dairy waste using microbial fermentations

16
392 Aspergillus niger fungus has the capability to produce oxalic acid when grown over a lactose

393 substrate. When Brown et al. (2018) used Aspergillus niger ATCC 6275 and ATCC 9029

394 strains on lactose whey permeate they were able to derive oxalic acid, but in very low

395 quantities. Co-fermentation of cheese whey has been shown to be highly effective at

396 enhancing its valorization capacity. It has been reported that addition of whey permeates into

397 wheat fermentation for ethanol production improved not only the ethanol yield but also

398 produced lactic and acetic acid (Parashar et al., 2016). Whey alone has lower ethanol yield

399 but when used along with wheat waste, the glucose from the wheat as the carbon source and

400 the action of Saccharomyces cerevisiae results in the lactic acid formation. On the other

401 hand, when thermal-alkaline pretreated Yerba Mate (Ilex paraguariensis) waste was co-

402 fermented with cheese whey it not only increased the hydrogen production by 7.5 times but

403 also produced noticeable amounts of lactic acid (∼140 mM), caproic acid (∼45 mM), and

404 butyric acid (∼135 mM) (Júnior et al., 2021).

405 2.6 Other bioproducts

406 The application of biorefinery principles on dairy waste has also brought attention to the

407 production of galacto-oligosaccharides (GOS), microbial pigments and industrial enzymes

408 from dairy waste. GOS are non-digestible oligosaccharides with Glu α1–4(β Gal 1–6) n as the

409 terminal glucose molecule and polymerization degree between 2–9 (Sakarika et al., 2020). β-

410 galactosidase (EC 3.2.1.23) acts upon dairy waste to carry out a trans-galactosylation reaction

411 forming a mix of mono-, di- and oligosaccharides. Another medicinally valuable prebiotic,

412 that is used in baby-formula feed, is lactulose (4-O-β-d-galactopyranosyl-d-fructose) which is

413 useful in improving the gut microbiota (Ottaviano et al., 2017; Nooshkam et al., 2018).

414 Various microbes such as Streptococcus thermophiles, Aspergillus oryzae, Bifidobacterium

415 bifidum, Bacillus circulans, and Kluyveromyces lactis have been used to produce β-

416 galactosidase biocatalyst (Contesini et al., 2019; Rico-Rodriguez et al., 2021). In an earlier

17
417 study, Sangwan et al. (2015) applied Streptococcus thermophilus on Lactose whey and

418 catalysis action of β-galactosidase resulted in the production of 53.45 g/L GOS. Whey waste

419 when hydrolysed in the presence of Lactobacillus sp. formed GOS (28%) due to the action of

420 β-galactosidase on reducing sugars (Sivakumar, 2021). β-galactosidase from Bacillus

421 circulans was demonstrated to result in even higher GOS production (77%) from whey (Das

422 et al., 2016). Ghimire et al. (2017) established a pilot scale membrane reactor plant and used

423 Kluyveromyces lactis to extract oligosaccharides from whey. They reported that the action of

424 β-galactosidase produced 31% oligosaccharides in the pilot setup.

425 Another important area of interest is the production of bioactive peptides from dairy

426 whey using methods such as proteolytic activity of microorganisms and starter cultures and,

427 enzymatic hydrolysis by digestive enzymes (Mohan et al., 2015). Different biomolecules

428 have been reported as the result of enzymatic hydrolysis of cheese whey proteins. For

429 instance, the microbial fermentation can form biopeptides with opioid characteristics (Garg et

430 al., 2018) anticancer (Hernández-Ledesma et al., 2017) and antioxidant (Rocha et al., 2017)

431 properties. In an attempt to observe the angiotensin-converting enzyme activity in

432 gastrointestinal digestion, Alvarado et al. (2019) used whey protein hydrolysate to produce

433 antihypertensive peptides and observed a 10% increase in enzyme activity.

434 Carotenoids have also been produced from dairy waste, and Roukas et al. (2015) used

435 a bubble column reactor with a range of microbes and concluded that Blakeslea trispora

436 acted on deproteinized hydrolysed whey to produce 405.5 mg/L carotenoids (β-carotene, γ-

437 carotene, and lycopene). Several filamentous fungal species such as Penicillium

438 chrysogenum and Penicillium purpurogenum have been reported to produce pigments by

439 using cheese whey as the sole carbon source. This is attributed to their ability to thrive in high

440 lactose substrates to produce pigments (Basto et al., 2019). Similar results were reported by

18
441 da Costa and Vendruscolo (2017) to produce red pigments by using Monascus ruber CCT

442 3802 on cheese whey as the lactose source.

443 However, these bioprocesses require further exploration to identify the metabolic

444 pathways involved in the production of useful biomolecules and there are several challenges

445 that need to be overcome for in vitro and in vivo production of these molecules. For instance,

446 development of new bioprocessing techniques using omics approaches such as nutrigenomic,

447 proteomics, and peptidomics, are being progressed for bioactive peptides production (Dupont

448 2017; Valdés et al., 2017; Agyei et al., 2018).

449 3 Research gaps and future research directions

450 Although bioenergy production from dairy wastewater is a sustainable route for its

451 valorization, the efficiency and yield of bioenergy is very dependent on the process

452 parameters such as temperature, pH, co-substrates, mode of operation, and reactor

453 configuration. One way to tackle few of these variations is to develop an integrated waste

454 treatment plant with bioenergy recovery system. The dairy wastewater can be a continuous

455 feedstock available to produce hydrogen while the effluent after hydrogen extraction can be

456 mixed with the solid fraction of dairy waste and be further exposed to microbial degradation

457 in anaerobic composting to produce biogas. This, multi-channel approach would not only

458 reduce the burden of waste disposal at an industry level, but also provide additional source of

459 income for the plant. Another lesser explored approach is the use of nanoparticles for

460 enhancing process efficiency (Gadhe et al., 2015) by affecting the activity of microbes

461 through inoculum immobilization and providing a larger reaction surface. But the effect of

462 nanoparticle addition varies largely with the choice of microbes as well as the substrate,

463 which makes it important to further explore their applications and develop standards that can

464 be applied to a wide range of waste substrates.

19
465 When it comes to the production of organic acids from dairy waste, there is a lot of

466 variability in the process and a large dependency on process parameters to successfully

467 produce acids. Additionally, the type and strain of microbe used also determines the final

468 yield of the products. This makes the whole process of organic acid production non-scalable.

469 To tackle these challenges there need to be further studies performed using genetically

470 modified organisms with gene overexpression to handle a wide range of substrates and also

471 successfully grow on the waste media. This also means that application of omics studies is

472 important to be explored for further elucidation of kinetic models and identification of better

473 strains that can produce organic acids in large quantities by completely degrading the

474 substrate.

475 Nearly half of the processing cost during bioplastic production is attributed to

476 sourcing carbon source substrates/feedstock. Thus, dairy waste provides the opportunity to

477 develop a low-cost solution. But the identification of an appropriate microbial strain in

478 bioplastic production that produces higher yields and also high-quality product remains a

479 challenge. Although few strains belonging to Bacillus sp, Cupriavidus sp. and Ralstonia sp.

480 have demonstrated better PHB production performance, they are still limited by their

481 adaptability to digest a range of waste present in the dairy effluent. One interesting approach

482 to overcome this could be the usage of mixed cultures to cover a wide range of waste present

483 in the wastewater while also reducing overall production cost. However, not much progress

484 has been made in scaling up the bioplastic production from dairy waste due to inherent

485 technical challenges of producing biofilms in a cost-effective industrially scalable method.

486 Another route could be to bioengineer the strains to withstand more stress of the harsh

487 wastewater environment and also degrade a wide range of waste. This would make it easy to

488 handle large volumes of wastewater without the need of specifically handling every waste

489 component separately. Equally important is the streamlining of downstream processing where

20
490 bioplastic will be finally isolated and purified for other applications. This is required to

491 dissociate the final product quality from dairy waste variability. Also important is to engineer

492 microbes with higher carbon consumption efficiency and the capability to produce a single

493 product i.e., bioplastic. This would further help in reducing the cost of downstream

494 processing and improve quality.

495 4 Conclusion

496 The increasing demand on dairy industry from direct consumption to production has resulted

497 in utilization of huge amount of resources and release of harmful effluents in the

498 environment. But various microbial treatment techniques developed in the recent past are

499 progressing the conversion of dairy waste into useful products. Anaerobic treatment can

500 directly convert the waste into valuable energy resource, whereas other biotechnological

501 methods can use it as a substrate for production of other value-added products such as

502 biosurfactants, bioplastics and biomolecules. These strategies are promoting circular

503 economy and reducing the impact of waste disposal on the environment.

504 Acknowledgement

505 Authors would like to acknowledge Scotland’s Rural College (SRUC), UK for research

506 facilities and the support under the respective ongoing projects.

507 Conflict of Interest

508 The authors declare no conflict of interest.

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907

38
908

909 Figure 1: Prominent microorganism species deployed in bioconversion of dairy waste

910

39
911 Table 1: Production of bio-based products from dairy waste

Microorganism/
Substrate Product Yield Reference
biomolecule

113.2 ± 2.9 mmol Landfill leachate Wong et al.

Dairy wastewater Biohydrogen
H2/g COD sludge (2019)

Scotta feedstock 60 g BSA eq/L

Papain and Monari et al.
from ricotta Bioactive Peptide and to 1.7 g AA
pancreatic enzymes (2019)
cheese eq/L

Rhodopseudomona
Dairy wastewater Bioplastic 7.23 ± 0.08 g/L Dinesh (2020)
s rutilaI

Saccharomyces Gaur et al.

Dairy waste Biomass 0.32 g/L
cereviesae (2017)

Saccharomyces Parashar et al.

Whey permeates Bioethanol 14.9-15.9% v/v
cerevisiae (2016)

Panner (Cheese) Pseudomonas Patowary et

Biosurfactant 2.7-4.8 g/L
whey aeruginosa SR17 al. (2016)

Bioenergy 850 mV and 28 Saccharomyces Al-saned et

Dairy waste
(electricity) µA cereviesae al. (2021)

Saccharomyces Beniwal et al.

Whey permeates Bioethanol 0.32 ± 0.007 g/g
cerevisiae (2021)

Dairy whey and

Pseudomonas Ibrahim et al.
vegetable oil Biosurfactant 5.72 g/L
aeruginosa (2021)
waste

912

913

40
914 Table 2: Production of organic chemicals from dairy waste using microbial fermentations

Substrate Microorganism Product Yield Reference

De-proteinated Actinobacillus 0.72 g/L per Longanesi et al.

Succinic acid
cheese whey succinogenes ATCC 55618 hour (2018)

Lactobacillus acidophilus 13.27 g/L Pandey et al.

Propionibacterium (2019)
Cheese whey Propionic acid
Acidipropionici ATCC 3.3 g/L Ahmadi et al.

4875 (2017)

Pescuma et al.
Lactobacillus casei 33.7 g/L
Whey (2015)
Lactic acid
permeates Ricciardi et al.
Lactobacillus casei N87
(2019)

Pandey et al.
Whey waste Lactobacillus acidophilus Acetic acid 7 g/L
(2019)

Pseudomonas taetrolens De Giorgi et al.

42.4 g/L
Ricotta cheese Zymomonas mobilis (2018)
Lactobionic acid
whey expressed in Escherichia Goderska et al.
3.2 g/L
coli (2015)

Quinoprotein glucose

dehydrogenase (GDH)
Cheese whey Lactobionic acid 8.70 g/L Oh et al. (2020)
expressed in Pseudomonas

taetrolens

Chwialkowska et
Whey isolate Saccharomyces cerevisiae Caproic Acid 5.0 g/L
al. (2019)
Cheese whey 0.36 g
Iglesias-Iglesias et
and Sewage Microbial consortia Caproic Acid HCap_COD
al. (2021)
sludge L-1 d
915

41