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Received: 5 August 2021

DOI: 10.1049/enb2.12020

REVIEW
- -Revised: 17 January 2022 Accepted: 21 February 2022

- Engineering Biology

Biologically engineered microbes for bioremediation of


electronic waste: Wayposts, challenges and future directions

Ping Han1,2,3 | Wei Zhe Teo1,2,3 | Wen Shan Yew1,2,3

1
Synthetic Biology for Clinical and Technological Abstract
Innovation, National University of Singapore,
Singapore, Singapore
In the face of a burgeoning stream of e‐waste globally, e‐waste recycling becomes
2
increasingly imperative, not only to mitigate the environmental and health risks it poses
Synthetic Biology Translational Research
Programme, Yong Loo Lin School of Medicine, but also as an urban mining strategy for resource recovery of precious metals, rare Earth
National University of Singapore, Singapore, elements, and even plastics. As part of the continual efforts to develop greener alter-
Singapore natives to conventional approaches of e‐waste recycling, biologically assisted degradation
3
Department of Biochemistry, Yong Loo Lin School of e‐waste offers a promising recourse by capitalising on certain microorganisms' innate
of Medicine, National University of Singapore,
ability to interact with metals or degrade plastics. By harnessing emerging genetic tools in
Singapore, Singapore
synthetic biology, the evolution of novel or enhanced capabilities needed to advance
Correspondence bioremediation and resource recovery could be potentially accelerated by improving
Wen Shan Yew, Department of Biochemistry, Yong enzyme catalytic abilities, modifying substrate specificities, and increasing toxicity toler-
Loo Lin School of Medicine, National University of ance. Yet, the management of e‐waste presents formidable challenges due to its massive
Singapore, 8 Medical Drive, Singapore 117597,
volume, high component complexity, and associated toxicity. Several limitations will need
Singapore.
Email: wenshanyew@nus.edu.sg to be addressed before nascent laboratory‐scale achievements in bioremediation can be
translated to viable industrial applications. Nonetheless, vested groups, involving both
Funding information start‐up and established companies, have taken visionary steps towards deploying mi-
Agency for Science, Technology and Research, crobes for commercial implementation in e‐waste recycling.
Grant/Award Number: IAF‐ICP HH@NUS
SINERGY Laboratory
KEYWORDS
biochemical engineering, bio‐economy, biological design, microbial engineering, synthetic biology

1 | INTRODUCTION the largest proportion of e‐waste, followed by large equipment


and consumer appliances (Figure 1). [1] Although this volume
Remarkable advances in science, engineering, and technology is modest compared to the 242 million tonnes of plastic waste,
since the early 20th century have created amazing tools and which constituted the 2.01 billion tonnes of municipal solid
devices that have revolutionised our world and become argu- waste the world produced in 2016, e‐waste presents one of the
ably indispensable in our daily lives. In many developed fastest growing waste streams, with an estimated growth rate of
countries, higher levels of disposable incomes and growing 3%–5% annually. [2, 3] It was predicted that by 2030, that
urbanisation have driven a fervent pace of technological global e‐waste will increase up to 74 million tonnes, impelled
change and the adoption of electrical and electronic gadgets. by a burgeoning penetration of electric and electronic equip-
While modern technology has undeniably elevated living ment in developing countries, a foreseeable replacement
standards and ushered in lifestyle conveniences, it has, at the market in developed countries and increasing product obso-
same time, left in its wake two distressing impacts on the lescence rates. [1]
environment—pollution and depletion of natural resources. Composed of diverse metals and non‐metals, the constit-
In 2019, the Global E‐waste Monitor 2020 reported a re- uents of e‐waste differ across product lines and categories and
cord 53.6 million metric tons of electrical and electronic waste can contain, in addition to prevailing base metals and varying
(e‐waste) generated, with small electronic devices representing quantities of plastics and ceramics, both precious metals and

-
This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is
properly cited.
© 2022 The Authors. Engineering Biology published by John Wiley & Sons Ltd on behalf of The Institution of Engineering and Technology.

Eng. Biol. 2022;6:23–34. wileyonlinelibrary.com/journal/enb2 23


24
- HAN ET AL.

F I G U R E 1 Global e‐waste generated in 2019. Regionally, Asia generated the largest volume of e‐waste and Oceania, the least. In 2019, the global quantity of
e‐waste mainly comprised small equipment (17.4 Mt), large equipment (13.1 Mt), temperature exchange equipment (10.8 Mt), screens and monitors (6.7 Mt),
small IT and telecommunication equipment (4.7 Mt) and lamps (0.9 Mt). Of these, 9.3 Mt, or 17.6%, was documented to be collected and properly recycled,
while the fate of 82.6% remained uncertain. Source: Global E‐Waste Monitor 2020

rare Earth elements. [2, 4] Waste electrical and electronic the maturing field of bioleaching has fostered the inception of
equipment (WEEE), however, also often contain hazardous nascent industry players that tap into the use of microorgan-
materials including heavy metals such as lead, nickel, chro- isms for metal recovery, limitations of the bioleaching process
mium, and Mercury, and persistent organic pollutants (POPs) are described. Ensuing are discussions on the potential of
such as polychlorinated biphenyls (PCBs) and brominated biological engineering of microorganisms to further advance
flame retardants (BFRs). [5, 6] When improperly disposed of, the bioremediation of e‐waste not only for metal recovery but
these substances can either be directly released or act as a also for plastic recycling. Next, the technical challenges that
precursor for the generation of toxic byproducts, resulting in need to be overcome in order to translate the technology into
environmental pollution and severe health risks. [7, 8] Indeed, industrial implementation are addressed. Finally, open issues
the management of e‐waste presents formidable challenges due and the future prospects of e‐waste recycling using biologically
to its massive volume, high component complexity, and asso- modified microorganisms are outlined in the last part of the
ciated toxicity. Yet, with as many as 69 elements from the review.
periodic table found in complex electronics including precious
metals such as gold, platinum, palladium, ruthenium, and silver,
e‐waste presents an alternative or secondary source of raw 2 | BIOTECHNOLOGY: A GREENER
materials in a circular economy (Figure 2). [4, 9] However, APPROACH TO E‐WASTE RECYCLING
despite the potential for resource recovery, only 17.4% of e‐
waste was formally collected and recycled globally in 2019, E‐waste recycling encompasses the collection of WEEE, its
inferring a loss of high‐value commodities conservatively processing, and finally the recovery of valuable materials. In
valued at $57 billion through disposal. [1] In the face of current industrial practices, the initial stage involves disman-
insatiable global demand and depleting geologic stores of tling and granulation of WEEE using physical–mechanical
extractable raw materials, urban mining—the extraction and methods and separation of recyclable fractions from the
purification of precious metals from waste streams—is non‐essential components via density, magnetic, or current‐
increasingly being recognised to play a strategic role in based sorting techniques. [11, 12] Plastics, which account for
resource security. It is argued that a shift to resource recovery almost 20% of e‐waste, are mostly given a second lease of
through e‐waste recycling is becoming more cost‐effective utility as recycled polymers, although emerging technologies
compared to conventional virgin mining and that the grade aim to transform e‐waste plastics into higher value products
and concentration of metals in e‐waste can be higher than that through microfactories. [13] For the majority of industrial scale
extracted from ores. [10]. e‐waste recycling, metals are recovered using pyrometallurgical
Concomitant with the shifting mindset on the importance and hydrometallurgical processes; the former involves thermal
of e‐waste management is the impetus to explore greener ap- treatment, smelting in furnaces or alkali chemicals, and other
proaches to e‐waste recycling compared to conventional solid‐liquid‐gas reactions at high temperature, while the latter
chemical methods. In this review, research developments in the uses solvent extraction, leaching, adsorption, and ion exchange
biologically assisted degradation of e‐waste is presented. While processes. [14] Unfortunately, such traditional methods of
HAN ET AL.
- 25

F I G U R E 2 Potential value and volume of raw


materials in e‐waste worldwide in 2016. E‐waste
contains not only precious metals including gold,
silver, copper, platinum, and palladium but also
valuable bulky materials such as iron and aluminium
and plastics that are recyclable. Secondary raw
materials of e‐waste are estimated to be worth
55 Billion Euros. Source: Global E‐Waste Monitor
2017

metal reclamation from e‐waste tend to use, or release as part metals, which account for more than 95% of the recovery
of the extraction process, toxic chemicals such as highly acidic/ value. As integral components in innumerable electronic de-
alkaline reagents, cyanide, and dioxins that are detrimental to vices including the burgeoning end‐of‐life mobile phones,
human and ecological health. As part of efforts to minimise the PCBs are composed of around 28%–30% metal, with up to
emission of hazardous pollutants during e‐waste recycling, 20% copper and precious metals such as silver, gold, and
advanced technologies, such as electrochemical, supercritical, platinum making up 0.3%–0.4%. [24] Yet, despite the many
and vacuum metallurgical technology, as well as novel ones like studies, the commercial application of bioleaching of PCBs
ultrasound technology, are being developed often in combi- remains elusive for the reasons of low leaching efficiency and
nation with improved conventional methods. [15]. long leaching time. Besides component heterogeneity, the
In the exploration of greener alternatives to conventional toxicity of the PCBs, which can affect the growth of micro-
approaches, biologically assisted degradation of e‐waste offers organisms as well as interference by exogenous metals, have
a promising way to metal recovery. Microbe‐metal interaction been thought to be contributing factors to low efficiency in
is a naturally occurring phenomenon in which microorgan- bioleaching.
isms, both prokaryotic and eukaryotic, acquire metal species Research efforts have hence been increasingly directed
from the external environment that are essential for various towards improving bioleaching processes. One of the widely
structural or catalytic functions. [16] A diverse group of deployed methods to improve cell tolerance against e‐waste
microorganisms that includes heterotrophic and chemo- toxicity is the preadaptation of microorganisms through
lithotrophic bacteria and fungi have an innate ability to acclimatisation to increasing PCBs concentrations prior to
convert insoluble metals to their soluble and extractable form bioleaching. [25–28] A two‐step process whereby microor-
—either by immediate surface attachment and directly oxi- ganisms are allowed to grow before exposure to the target
dising minerals and solubilising the metals, or through an waste was also found to be crucial in mitigating the inhibitory
indirect action of regenerating of ferric ions, which serve as effects and additional extension to three‐ or four‐steps posi-
oxidising agents. [17, 18] This process of bioleaching, also tively affected metal recovery. [28–30] In a further move to
known as biohydrometallurgy, involves various mechanisms decouple cell viability from bioleaching activity, an indirect,
such as complexolysis, acidolysis, redoxolysis, and bio- non‐contact mechanism was investigated using bacteria‐free
accumulation. [19]. cultural supernatant. [31, 32].
In the initial phase, potential candidates for biorecovery The complex compositions of e‐waste make it challenging
of metals are often prospected from microbial communities for single microorganisms to completely utilise the substrates
inhabiting natural environmental niches or indigenous to and efficiently recover multiple metals in one process. As such,
polluted sites or operation facilities. As comprehensively the use of mixed cultures or the development of a microbial
reviewed by Valix [20] and Islam et al. [21], several mi- consortium is gaining traction. [20, 33] Bioleaching of base
croorganisms including bacteria (e.g. Acidithiobacillus spp., metals requires different microbial strains from those for
Sulfobacillus spp., Pseudomonas spp.) and fungi (e.g. precious metals. Thus, using co‐ or mixed‐cultures can
Aspergillus spp., Purpureocillium lilacinum) have been potentially harness synergy between groups of microorganisms
reported for recovery of base (Cu, Fe, Ni, Pb, Zn) and not only in accessing nutrients but also in functional robust-
precious metals (Ag, Au, Co, Pd, Pt) from e‐waste. In ness. For example, copper and gold recovery efficiencies
particular, the acidophilic group of bacteria plays a prom- reached 98% and 44%, respectively, when copper leaching was
inent role in bioleaching in e‐waste due to their lenience carried out using chemolithotrophic acidophilic Acid-
towards heavy metals. [22, 23]. ithiobacillus ferrivorans and Acidithiobacillus thiooxidans
Among WEEE, printed circuit boards (PCBs) are the followed by the use of cyanide‐producing heterotrophic Pseu-
predominant targets for recycling because of their valuable domonas fluorescens and Pseudomonas putida for gold
26
- HAN ET AL.

leaching. [34] And chromium recovery was reported to oxidants to leach out the copper, tin, and other less valuable
improve when co‐cultures of biosurfactant‐producing bacteria metals, leaving behind precious metals such as gold and
and sulphur‐oxidising bacteria were used. [35]. palladium in the solid remains. The solid e‐waste then un-
Besides biotic determinants to the bioleaching processes, dergoes another round of acid/oxidants leaching to obtain
several abiotic factors, such as initial pH, carbon source, pulp gold ions, which are subsequently exposed to the bacterium
density, and particle size, and their interaction can also influ- Cupriavidus metallidurans to absorb the gold ions and
ence bioleaching efficiency. To maximise metal recovery, these recover the precious metal. The company has been able to
parameters can be optimised under a multi‐objective strategy process around 1 metric ton of circuit boards in a week in their
using response surface methodology in order to determine demonstration plant, recovering about 150 g of gold in the
optimal bioleaching conditions. [36, 37] Bioleaching offers a process. Mint Innovation is in the midst of building a com-
promising route in the recovery of the metallic fractions from mercial plant that is set to improve the throughput to 10 metric
e‐waste and is considered more environmentally benign than tonnes of circuit boards per day.
physicochemical approaches. In the light of its importance, it is In 2018, the IT asset lifecycle management company N2S
critical that the processes involved are optimised to attain (www.n2s.co.uk) collaborated with research groups from
levels of recovery efficiencies that enable the translation of Coventry University, UK to develop a pilot bioleaching plant
research to sustainable, commercial application. to process e‐waste. They have demonstrated, for the first time,
the direct application of electrowinning to PCBs bioleachate
for the recovery of copper. [45] Shredded PCBs were intro-
3 | BIOLEACHING: FROM duced to the bacteria A. ferrooxidans after their growth phase
EXPLORATION TO IMPLEMENTATION and the electrowinning process was then carried out once the
bioleachate reached the optimum condition for copper metal
Fostered by years of research and development, microbial recovery. Characterisation of the recovered copper foil revealed
leaching has been successfully applied at industrial scale bio- a high purity (>99%), demonstrating the selectivity of the re-
mining operations for extraction of base and precious metals covery process. This approach of copper separation from e‐
from sulfidic ores. [38] Biohydrometallurgical extraction of waste could potentially serve as a pre‐treatment method to
metals such as cobalt and nickel has been technically estab- improve the recovery of precious metals from e‐waste.
lished and have progressed into commercial implementation.
Today, metal recovery from low‐grade copper ores and re-
fractory gold ores are the two dominant industrial applications 4 | SYNTHETIC BIOLOGY: A
and account for about 20% and 5% of global production of PROMISING TOOL IN ADVANCING
copper and gold, respectively. [33, 39–41] Although the use of BIOHYDROMETALLURGY
microorganisms in metal recovery have found profitable
actualisation in biomining, their application in the recycling of The use of microorganisms for e‐waste management opens up
waste materials such as e‐waste is still emerging and remains avenues for investigation using synthetic biology. Through the
largely confined to laboratory‐scale studies. application of engineering principles to molecular biology,
Despite the challenges, several groups have managed to synthetic biology offers the tools to improve enzyme catalytic
move a step closer towards commercialising their approaches. abilities, modify substrate specificities and raise toxicity toler-
One such example is Mintek (www.mintek.co.za), an estab- ance, potentially accelerating the evolution of novel or
lished company with vast experience in bioleaching of gold enhanced capabilities needed to advance bioremediation and
from ores. [42] In partnership with BacTech, the company's resource recovery. [46, 47] Shortcomings in microbial reme-
commercial refractory gold bioleaching technology, BacoxTM, diation have been the impetus in driving vast research into the
is currently applied at the Beaconsfield Gold Mine in Tasmania development of genetically engineered microorganisms for
and the BioGold toll treatment facility located near the city of bioremediative applications targeting all mechanisms—
Laizhou, Shandong province, PR China. These facilities pro- detection, adsorption‐chelation, absorption, and bioconver-
cess between 70 and 100 tonnes of refractory gold concen- sion— of metal capture and removal as extensively reviewed by
trates a day and the bioleaching take place in large aerated Diep [48] and Sharma et al. [49].
stirred‐tank reactors where thermophilic iron‐oxidising bacte- Of these, the bulk of the studies related to metal accu-
ria are mixed with the refractory gold concentrates to leach out mulation and recovery remains focussed on using biologically
the gold. [43] Riding on its success in the use of biotechnology engineered microorganisms to elucidate the molecular mech-
for the extraction of gold from ores, the company is now anisms behind their metal tolerance, oxidation, or degradation
working on developing and optimising processes for the abilities, such as the discovery of delftibactin, a siderophore‐
treatment of e‐waste. like non‐ribosomal peptide from Delftia acidovorans that
Another example is Mint Innovation (www.mint.bio), a can detoxify soluble gold, offering a mechanism for gold
biometallurgy company based in Auckland, New Zealand. To biomineralisation. [50] Several interesting studies, however, rise
recycle e‐waste, the team behind Mint Innovation combined to the fore for their strong relevance in the potential applica-
both chemical and biological remediation steps in their work- tion for bioleaching of e‐waste. In one study, Pseudomonas
flow [44]. Grounded e‐waste is first mixed with acids and putida 06909, with characterised recombineering tools, was
HAN ET AL.
- 27

engineered to produce a high Cd‐affinity metal binding peptide with this in mind, a systematic genetic toolkit was developed to
EC20 that not only improved cadmium binding but also alle- facilitate future engineering for biotechnological applications.
viated the cellular toxicity of cadmium. [51] In another study [61].
using an in silico genome‐scale metabolic model (GEM)‐based While laboratory scale research provides insight into the
approach, gene targets that redirect the metabolic flux towards tremendous potential of microorganisms in remediation and
the production of cyanide were identified, leading to over- resource recovery, bold claims on their efficiency or cost‐
production of the bioleaching agent and a concomitant effectiveness remain premature if the studies are not progres-
improvement in gold and copper recovery. [52] Employing sively scaled up to bioreactor configurations and onward to
both genetic and chemical engineering, Zhu et al. constructed pilot scale setups and field tests. For the successful translation
novel biotic‐abiotic coassemblies, comprising synthetic bacte- to industrial‐scale bioleaching processes, there is clearly a need
rial cells displaying the heavy‐metal‐capturing SynHMB on the to implement scale‐ups and validation tests, which could reveal
cell surface and magnetic nanoparticles, which could remove bottlenecks such as decreasing bioleaching efficacy with
cadmium and lead at high efficiencies. [53]. increasing pulp density, pH fluctuations that affect the growth
While an ever‐expanding range of toolkits has become and activity of the acidophilic microorganism, or toxicity of the
available for a variety of microbial chassis, molecular tools for non‐metallic fraction of the WEEE, to name a few.
non‐model organisms, such as the bioleaching bacteria and
archaea, are notably lacking. For example, despite their pre-
eminent role in the biomining industry, Acidothiobacillus sp. 5 | SYNTHETIC BIOLOGY: A
and Sulfolobus sp. have rudimentary genetic tools for molec- POTENTIALITY FOR PLASTIC
ular manipulation. [35] Initial efforts in engineering these mi- BIODEGRADATION
croorganisms for improved performance in bioleaching show
promise. The expression of arsenic resistance genes in A. fer- A greater part of the focus on e‐waste recycling lies in the
rooxidans generated strains with increased tolerance to arsenic recovery of precious metals, although not to be disregarded are
while expression of the mer operon from A. ferrooxidans the bulky plastics that are comparably valuable (Figure 2). [9]
resulted in Mercury resistance in mercury‐ion sensitive vari- Plastics account for almost 37% of WEEE, but there are
ants. [54, 55] Although still far lagging than their model formidable hurdles to the recycling of e‐waste plastics mainly
counterparts, importation of established toolkits, such as the due to the mixed material composition and recalcitrant prop-
shuttle vectors and genetic regulatory elements from E. coli erties of synthetic polymers. [62] The constituents of e‐waste
into these acidophiles, serve as a starting point for establishing plastics include some 15 different polymer types with the
platforms for future work on metabolic or microbial engi- typical composition comprising high impact polystyrene
neering of biomining microorganisms for desired traits and, (HIPS, 27%), acrylonitrile butadiene styrene (ABS, 24%),
ultimately, improved bioleaching efficiencies. [56]. polycarbonate (PC, PC‐ABS, 7%), polypropylene (PP, 5%) and
Another non‐model microorganism of interest is the polyethylene (PE, 2%), and the remaining consisting of poly-
cyanogenic bacteria Chromobacterium violaceum found to be mer blends and other thermoplastics. [63] Compounding the
one of the most effective microorganisms for the bio‐ problem is the presence of harmful additives such as BFRs and
dissolution of gold. [57] Cyanogenic bacteria produces cya- POPs. [64].
nide lixiviant, which complexes with gold facilitating the bio- Even though established industrial processes for plastic
leaching process. The cyanide lixiviant is generated from the recycling exist, the biological degradation of plastics could
secondary metabolite hydrogen cyanide (HCN) produced by offer an ecologically friendlier strategy. The capability to
oxidative decarboxylation of glycine in a reaction catalysed by degrade plastics has been reported in several microorganisms
the enzyme (HCN) synthase encoded by the hcnABC operon. as extensively reviewed by Ru et al. [65] and Sameh et al. [66].
Regulation of this operon under quorum control limits the Isolated from a variety of environments, including soil, marine
production of cyanide to 20 mg/L at the onset of the sta- waters, landfill sites and waste sludge, these microorganisms
tionary phase. [58] To circumvent the limited cyanogenic have shown to have the ability to degrade synthetic polymers,
capability of wild‐type C. violaceum, lixiviant production was such as PP, PE, polystyrene (PS), polyvinyl chloride, poly-
decoupled from quorum control through the use of exogenous urethane (PU) and polyethylene terephthalate (PET), some
promoters (pBAD and pTAC) resulting in about 70% more even down to the respective simple monomeric units. [67] The
cyanide production in the engineered strains and more than degradation is based on the structural similarity of these
twice the recovery of gold. [59] An extension of the study plastics with natural polymers such as lignin present in the
introduced an additional copy of the cyanide producing operon environment. Depolymerisation of synthetic plastics often in-
to produce higher cyanide lixiviants was shown to boost gold volves microbial enzymes that target oxidisable C–C bonds,
recovery from 11% to 30%. [60] The cyanogenic capabilities of albeit with much lower efficiency. These naturally occurring
C. violaceum and its promising role in the targeted application enzymes include peroxidases, laccases, cutinases, esterases,
of gold biorecovery from e‐waste makes it an attractive bio- hydrolases and lipases as detailed in Amobonye et al. [68] and
logical chassis for synthetic biology. Comparative proteomics Othman et al. [69].
analyses suggest that further increase in cyanogenesis is To date, PET degradation by thermophilic actinobacteria
possible through lixiviant metabolic engineering and hence, via two key enzymes PETase and MHET hydrolase is arguably
28
- HAN ET AL.

the most efficient among the currently known plastic degra- production but have instead pivoted towards engineering
dation processes. [70] PET, however, is not a constituent of the metabolic pathways in microbes to bio‐upcycle synthetic
e‐waste plastic stream. Hence, to strive towards a biological polymers to produce value‐added products, such as high‐value
approach in e‐waste plastic recycling, it is imperative that the aromatics and innovative chemicals for novel materials. [65]
search and isolation persist for novel enzymes active on Riding on this approach is the EU Horizon 2020 research
recalcitrant polymers relevant to e‐waste. [71] This is particu- project MIX‐UP—Mixed plastics biodegradation and upcy-
larly pressing in the face of a dearth of reports on microbial cling using microbial communities, which ambitions to explore
biodegradation of high‐strength polymers ABS and HIPS, novel sustainable routes to valorise plastic waste streams. [82]
which constitute about 50% of e‐waste plastics. [72, 73]. The project aims to work towards controlled enzymatic and
Biodegradation of plastics is a very slow process under microbial degradation of pre‐treated plastic waste using heavily
normal conditions. To overcome inherent limitations, the use engineered, plastic type‐specific enzyme blends, and subse-
of biological engineering would thus be pivotal in improving quent microbial conversion to polymers and value‐added
the efficiency of polymer degradation. As exemplified by the chemicals by self‐sustaining microbiomes.
enhancement of PET hydrolytic activity by rational design,
protein engineering efforts can be directed to the development
of modified enzymes with the aim of improving specific 6 | CHALLENGES IN INDUSTRIAL
binding capacities and catalytic activities towards a broader IMPLEMENTATION
range of synthetic polymers. [74–76] This is particularly crucial
when dealing with plastic material heterogeneity in e‐waste. The utilisation of microbes for the remediation of e‐waste
The often time consuming process of molecular modifications provides a greener, less expensive, and less toxic avenue for
towards gain‐of‐function or activity augmentation can be e‐waste recycling. However, there are several limitations that
bolstered by maturing synthetic biology approaches such as need to be addressed before this process can be implemented
whole‐genome sequencing, multi‐omics, automated high‐ on an industrial scale (Table 1). [22, 83].
throughput mutant screening and novel strain construction
using genome‐editing tools. [77] The growing availability of
bioinformatics resources and computational biology tools can 6.1 | Toxicity of organic and inorganic
also aid in the advancement of plastic biodegradation by components in e‐waste
facilitating in silico research such as computer‐aided targeted
mutagenesis and model‐based predictions of mechanistic E‐waste contains diverse materials, including hazardous
enzyme‐polymer binding. [78]. organic substances such as PCBs, polybrominated biphenyls,
The use of biologically engineered microorganisms has polybrominated diphenyl ethers, and BFRs, and inorganic
been the traditional strategy for achieving consolidated bio- metals such as lead, Mercury, chromium, and cadmium. [5, 84–
processing (CBP) by integrating all bioconversion reactions in 87] Concentrations of these components are likely to differ
a one‐step biological process. [79] In the wake of the huge from batch to batch depending on the myriad types of e‐waste.
challenges faced in designing and optimising all desired func- This contrasts with most laboratory‐based studies where the
tional genes in one strain even with the advances in synthetic bioremediation activity of the microbes is usually optimised
biology toolkits, there is growing recognition, particularly in e‐ using a uniform source of e‐waste, and the microbes' tolerance
waste recycling, that engineering of mixed cultures and even towards the hazardous components is well‐defined. As a result,
microbial consortia to compartmentalise complex tasks and transiting from lab‐scale to industrial‐scale bioremediation
functionalities would be the way forward (Figure 3). [80] poses a huge challenge for consistency in bioremediation ef-
Notably, the use of tailored microbial consortia has shown ficiency as the viability of microbes, and consequently their
higher biodegradation efficiencies of PE, PU, and PET activities, are likely to vary from batch to batch, thus limiting
compared to individual strains. [77, 78] With complex micro- their potential for application in commercial scale e‐waste
bial communities, it is postulated that different microorganisms recycling.
may exploit metabolic cross‐feeding and express distinct
degrading enzymes when cultured with the different plastic
materials, and that combinations of selected bacteria may have 6.2 | Slow process in comparison to physical
synergistic effects on biodegradation. and chemical remediation
A remarkable characteristic of the PET‐degrading bacte-
rium Ideonella sakaiensis is its innate ability to utilise the Even though bioremediation is an effective and less hazardous
degradation products as a major carbon and energy source for method for e‐waste recycling, its sluggish pace makes the
growth. [70] Tapping on the microorganism's ability to process a weak contender as compared to the use of physical
assimilate PET degradation products as building blocks for and chemical means to recycle e‐waste. To gain ground, the
growth opens the possibility of plastic waste as a novel feed- rate of bioremediation needs to be drastically improved, and
stock for microbial biotransformations. [81] Challenged by the this often requires an in‐depth study on the mechanism of the
low cost of virgin plastics, research efforts have been directed bioremediation process as well as the genetic engineering of
away from recycling the recovered monomers for bio‐plastics microbes for better performance. In addition, the composition
HAN ET AL.
- 29

F I G U R E 3 Biologically engineered microbes for the bioremediation of e‐waste. In 2016, the proportion of raw materials in e‐waste worldwide comprised
base metals (aluminium, iron, copper) and precious metals (silver, gold, and palladium), and plastics. [9] Synthetic biology can be used to promote metal recovery
by enhancing metal tolerance or increasing bioleaching efficiencies in iron‐ or sulphur‐oxidising microorganisms, or improving lixiviants bioproduction in
cyanogenic bacteria. As more novel plastic‐degrading enzymes are uncovered, microorganisms can also be engineered to acquire plastic degradation abilities. In
addition, metabolic engineering could facilitate the potential use of polymers as a feedstock in biotransformation for plastic upcycling. Future directions will be
directed towards the use of mixed cultures or microbial consortia to tap on diversification of the function for greater efficacy against substrate heterogeneity. The
prospective use of biofilm‐forming microbial chassis could also boost the colonisation and degradation of polymers. Besides biotic enhancements, a
multipronged approach to maximise resource recovery in e‐waste can involve the application of response surface methodology (RSM) to determine optimal
bioleaching parameters. M represents metal ions (Mx) or its reduced form (M0)

TABLE 1 Advantages and disadvantages of using microbes for bioremediation of electronic waste. [22, 83]

Advantages Disadvantages
1. Simple and inexpensive 1. Bioremediation activity can be easily affected by microbes' tolerance towards
toxic e‐waste components

2. Eco‐friendly technique 2. Bioremediation process is usually slower compared to remediation through


physical and chemical processes.

3. Avoid the use of harsh chemicals, high temperature, and pressure 3. Limited availability of molecular toolkit to improve the rate of bioremediation
and storage of microbes

4. Permanently eliminate contaminants through biochemical transformation or 4. Scaling up of the bioremediation process is complex and time‐consuming
mineralisation

of e‐waste can affect the speed of the bioremediation process, bacteria, acidophilic bacteria) often have very limited molecular
with reports demonstrating inhibitory effects on bioleaching by toolkits available for genetic manipulation. [35] This makes the
competing metals in the e‐waste. [20, 22]. This inhibition could engineering of these species to enhance their bioremediation
make the bioleaching of PCBs 20 times slower than its po- activities difficult and slow. Besides the lack of molecular
tential rate. toolkits, preservation of these microbes is also a challenge as
some of these microbes exhibit poor viability following pres-
ervation. [35, 88] Differences in the microbes' cell structure
6.3 | Limited availability of molecular and their overall sensitivity towards ultra‐low temperatures and
toolkits desiccation processes often result in failure of preservation
using standard methods as observed in acidophilic thermo-
Unlike common microbes such as E. coli, Saccharomyces cer- philic microorganisms. [89, 90] To circumvent the problem,
evisiae, and Pichia pastoris, microbes that display great po- these microbes are maintained as live cultures requiring
tential for e‐waste bioremediation (Chromobacterium continual subculturing and storage at temperatures below
violaceum, Acidithiobacillus ferooxidans, thermophilic optimal growth requirements, and with slow release substrates
30
- HAN ET AL.

to prolong viability. [91] Hence, with the push to discover on specialised facilities and skills and a possible workaround
novel microorganisms for bioremediation, tandem efforts must will be for companies and the scientific community to
be made to develop methods for the perseveration of these collaborate with biofoundries equipped with automation sys-
microbes for ease of use. tems and high‐throughput analytical equipment to accelerate
the screening process. [95].
Indisputably, the most pressing issue that needs to be
6.4 | Difficulty in scaling up of the resolved is the efficiency, and consequently speed, of the
bioremediation process bioremediation process. Two aspects, namely enzymatic per-
formance and toxicity tolerance, are important determinants
Moving an idea from laboratory‐scale success to a pilot‐scale affecting bioremediation efficiency. The way forward to
demonstration plant and onwards to an economically profit- achieving significant improvements in enzymatic performance
able commercial scale is a road fraught with challenges. While will likely rely on the use of biologically modified microbes.
physical–chemical technologies can mostly be scaled up with The Chinese–European MIX‐UP project, for example, en-
relatively high predictive reliability, biological applications, on deavours to create a novel workflow to recycle plastic waste
the other hand, are more problematic in which how well a through the use of heavily engineered enzyme cocktails. [82]
remediation treatment will work and how long that treatment Through the application of the ‘design, build, test, and learn’
will require is plagued with uncertainty as a result of the use of strategy in the engineering of the microbes, synthetic biology
live organisms. It has been reported that in bioremediation can enable a systematic and iterative construction of the
scale‐ups, laboratory‐scale kinetics can exceed field‐scale ki- engineered microbes and feedback loop analyses. [47] To
netics and the duration of treatment under‐predicted in excess facilitate genetic engineering, toolkits containing standardised
of 100% and by as much as 11,900%. [92] Aside from the genetic parts for building transcriptional units and operons are
complex, capricious nature of microorganisms, biological needed to generate mutant libraries in vitro for the selection of
treatment processes are also sensitive towards the physical (e.g. desirable phenotypes. [96–99] For this purpose, better efforts
temperature, oxygen level) and chemical (e.g. pH, humidity, will be needed to develop molecular toolkits for the non‐
nutrition) conditions of the reaction setup, and these condi- conventional microbes naturally adept with niche capabilities
tions are often affected when translating from laboratory‐scale in bioremediation. This can be done by tapping on the
systems to field‐scale designs. [93] This, in addition to the established BioBrick system for model organisms in the
introduction of spatial heterogeneities, and additional mass bioremediation of gold, cobalt, nickel, and Mercury. [100–102]
transport mechanisms and limitations, render the engineering Biofoundries can again play a vital role in screening exhaustive
of bioremediation industrial scale‐ups an arduous task. mutant libraries. For instance, Allonnia, a spin‐off venture of
the leading biotech company Ginkgo Bioworks, has plans to
engineer microbes through synthetic biology to remediate
7 | THE ROAD AHEAD: THE FUTURE organic pollutants in waste. Maturing fields in computer‐
OF E‐WASTE BIOREMEDIATION assisted protein engineering and droplet‐based microfluidics
can also enable improvements in directed evolution of en-
Despite being cumbered by limitations that are not trivial, zymes of interest, potentially hastening research efforts in
commercial implementation of e‐waste bioremediation and targeted improvements of bioremediation capabilities of
resource recovery remains highly desired goals in a global drive modified microbes. [103].
towards a zero‐waste circular economy. To advance the bio- Concomitant with growing research studies is the realisa-
logical approach to e‐waste management, gaps or inadequacies tion that dependence on a singular biological workhorse, even
in our current knowledge or methods have to be identified and if highly engineered, to remediate complex e‐waste is a tall
addressed by directing rigorous research efforts to overcome order. Research directions have thus been re‐modelled to
technological hurdles (Figure 4). include the exploitation of microbial communities to better
One of the utmost research needs is the expansion of the address the material heterogeneities of e‐waste, whether metals
biological arsenal capable of driving the enzymatic remediation or plastics. While holding much potential for dealing with the
of e‐waste, particularly in the area of plastic degradation where complex compositions of e‐waste, mixed cultures are, in reality,
enzymes specific for diverse polymers are sorely lacking. As difficult to implement as maintaining favourable growth con-
exemplified by the uncovering of more than 500 potential PET ditions for different microbes and ensuring the cellular envi-
hydrolases globally distributed in marine and terrestrial meta- ronment optimal for different enzymes are daunting
genomes, and the anomalous discovery of PR‐degrading bac- challenges. Equally important is the need for the process scale‐
teria in insect guts, efforts to prospect from microbial up from laboratory to pilot and field operations. This is a
communities in natural environmental niches or from indige- crucial step in the progress from proof‐of concepts to indus-
nous microbes inhabiting polluted sites or operation facilities trially applicable technologies. Consideration of the above is-
should continue. [94] Here, automation provides a means to sues in the early design stages would be useful in the path to
perform the tedious task of conducting high throughput developing a commercial bioremediation process.
screening of huge gene pools for desirable novel functions. Besides technical challenges, complementary e‐waste lo-
Automation, however, requires substantial capital investment gistics and management are also critical gaps to close in the
HAN ET AL.
- 31

F I G U R E 4 Roadmap to addressing the research needs to overcome technological gaps in the advancement of bioremediation of e‐waste using biologically
modified microorganisms

bid towards eventual commercialisation of bioremediation of 8 | CONCLUDING REMARKS


e‐waste. For instance, segregating e‐waste based on their
composition can help with material homogeneity, thereby The use of biologically engineered microbes for bioremedia-
alleviating the issue of bioremediation activity inconsistency tion of e‐waste, although promising, is still in its infancy and
caused by interfering components in the e‐waste. In addition, there remains a lot more research and development to be
the inclusion of a pre‐treatment step before introducing the committed before the technology can be adopted commer-
e‐waste to the microbes could also help to improve the rate cially. Progress has been made to improve cell tolerance against
of biodegradation. [104] Another possible workflow will be e‐waste toxicity, to tackle the need to recover multiple metals in
the use of a two‐step bioleaching approach, such as that a single process, and to increase the recovery efficiencies of
demonstrated by Shah et. al. where the biomass was allowed bioleaching. However, studies on scaling up these laboratory
to grow in the first step prior to the addition of PCBs, scale successes are lacking in the scientific field, restricting the
resulting in a 2–3.12 fold increase in copper, zinc, and nickel commercial implementation of e‐waste bioremediation.
extraction to recover copper, zinc, and nickel from PCBs. Therefore, research communities and industries should work
[105]. together to allow a smooth transition from the laboratory scale
Finally, the following concerns ought to be deliberated for bioremediation process to the field operational scale. In addi-
future recycling technological innovations: [1] In the drive to tion, the limited availability of molecular toolkits for genetic
achieve optimal bioremediation efficacies and greater resource engineering of uncommon microbes that display great poten-
recovery rates, the bioremediation workflow should be sus- tial towards e‐waste bioremediation is slowing down the pro-
tainable in terms of cost and ecological footprints; [2] To gear cess of improving their metal recovery efficiencies. Through
towards accessible commercialisation, the technology involved the application of the ‘design, build, test, and learn’ strategy in
should be suitable for adoption by small and medium‐sized synthetic biology and automation, these much‐needed molec-
enterprises in terms of investment or infrastructure. The ular toolkits can be developed at a faster pace. Proper e‐waste
operational technicalities and costs should also be put forward management and planning of the bioremediation workflow are
through the development of pilot scale tests by the scientific also critical gaps to close in the progress towards eventual
community; [3] Due to the complex nature of e‐waste, a single commercialisation of the bioremediation of e‐waste. Once
bioremediation technology is unlikely to be sufficient to these technical hurdles are crossed, it is important to deliberate
accomplish complete e‐waste recycling. Therefore, stake- whether the technology requires integration with other sec-
holders should remain open to integrating various recycling ondary or even tertiary recycling processes for complete e‐
technologies, including physicochemical treatments, for the waste remediation, and if the technological workflow is
complete remediation of e‐waste. [15]. ecologically and economically sustainable and suitable for
32
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