3 Biotech (2023) 13:220

https://doi.org/10.1007/s13205-023-03638-4

REVIEW ARTICLE

An insight on sources and biodegradation of bioplastics: a review

Nag Pooja1 · Ishita Chakraborty1 · Md. Hafizur Rahman2 · Nirmal Mazumder1

Received: 7 July 2022 / Accepted: 15 May 2023 / Published online: 31 May 2023
© The Author(s) 2023

Abstract
Durability and affordability are two main reasons for the widespread consumption of plastic in the world. However, the
inability of these materials to undergo degradation has become a significant threat to the environment and human health
To address this issue, bioplastics have emerged as a promising alternative. Bioplastics are obtained from renewable and
sustainable biomass and have a lower carbon footprint and emit fewer greenhouse gases than petroleum-based plastics. The
use of these bioplastics sourced from renewable biomass can also reduce the dependency on fossil fuels, which are limited
in availability. This review provides an elaborate comparison of biodegradation rates of potential bioplastics in soil from
various sources such as biomass, microorganisms, and monomers. These bioplastics show great potential as a replacement
for conventional plastics due to their biodegradable and diverse properties.

Keywords Bioplastics · Biodegradation · Starch · Cellulose · Conventional plastics

Introduction rose from 1.7 million tons in 1950 to 322 million tons in
2015 (Suman et al. 2020). It was also estimated that due to
Plastics are crucial materials in modern life, and due to the increased use of plastic personal protective gear during
their resistance to chemical, physical and biological deg- the COVID-19 pandemic, 585 million tons of single-use
radation, society relies majorly on them (Bhogayata and plastic waste would be generated by the end of 2020. The
Arora 2018). However, the overexploitation of plastics by major contributing countries to generating plastic waste are
the ever-growing human society has led to various environ- China, India, the USA, and Brazil (Benson et al. 2021). This
mental and health risks (Wright and Kelly 2017; Bradney demands an alternative to the ubiquitous plastics, and bio-
et al. 2019). Two of the main factors why plastics are so plastics have emerged as promising alternatives. ‘Biodegrad-
widely used are their durability and inexpensiveness, allow- able bioplastics’ are plastics that are derived from renewable
ing them to be used for various applications like food pres- biomass which are bio-based such as starch, cellulose, col-
ervation, packaging, transportation, building, construction, lagen, polylactic acid, and polyesteramides (Coppola et al.
etc. Unfortunately, the poor post-production management 2021). The use of bioplastics can reduce the dependency
of plastics becomes a significant problem in aggravating its on fossil fuels which are present in limited amounts. Unlike
impact on the environment. The inefficient waste manage- petroleum-based plastics, bioplastics emit a lesser amount
ment and deliberate littering have resulted in tons of plas- of greenhouse gases (Mittal et al. 2022). The term bioplastic
tic waste floating in the oceans, causing damage to marine does not necessarily mean that they are biodegradable or
ecosystems. Global petroleum-based plastic production eco-friendlier. Some of the bioplastics are not biodegradable
or may require a very long time to disintegrate. Bio-polyeth-
ylene (bio-PE), bio-polypropylene (bio-PP), bio-polyethyl-
* Nirmal Mazumder ene-terephthalate (bio-PET), bio-polytrimethylene tereph-
nirmaluva@gmail.com thalate (Bio-PTT), and bio-polyamide (bio-PA) are some of
1
the most common non-biodegradable bioplastics (Rahman
Department of Biophysics, Manipal School of Life
Sciences, Manipal Academy of Higher Education, Manipal,
and Bhoi 2021). Biodegradable polymers are defined as
Karnataka 576104, India polymers that can be degraded into carbon dioxide, water,
2
Department of Quality Control and Safety Management,
methane, and other low-molecular-weight compounds (Ishi-
Faculty of Food Sciences and Safety, Khulna Agricultural gaki et al. 2004).
University, Khulna, Bangladesh

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In biodegradation assays, two fundamental approaches are as photodegradation and chemical hydrolysis break down
used: aerobic and anaerobic digestion (Ruggero et al. 2019). the material chemically and physically at high temperatures
Biodegradation is a set of chemical reactions that take place and/or under acidic or basic pH conditions. (Polman et al.
in the presence of living organisms such as bacteria, fungi, 2021). There is a lack of reviews that study and compare
yeast, algae, and insects under optimum light, temperature, the detailed biodegradation of bioplastics from all the com-
and oxygen. The degradation process is also influenced by mon sources. This review aims to provide a comprehensive
the microstructure of the polymer. Furthermore, the rate of insight into the commonly used sources of bioplastics and
biodegradation might differ, depending on the environmental several factors affecting their biodegradation. Sources of
conditions and the polymer’s integral molecular structure Bioplastics” elaborates on the broad classification of bio-
(Scaffaro et al. 2019). Most of the plastic wastes are dumped plastics based on the source of raw materials. Some of the
as landfills and in soil, leading to the emission of green- most widely used sources for the synthesis of bioplastics
house gases and leachate. Using compost in the degrada- like polysaccharides (starch, cellulose, chitin and pectin),
tion of different biodegradable bioplastics has been studied. proteins (collagen, whey), polyhydroxyalkanoates and other
Composting is classified into three types: aerobic, anaero- sources such as polylactic acid, polyesteramides have been
bic, and vermicomposting. Aerobic composting occurs in reviewed. Table 1 summarizes the sources, advantages, and
the presence of plenty of oxygen. Aerobic microorganisms disadvantages of various native biodegradable bioplastics.
degrade organic matter, producing carbon dioxide (­ CO2), “Biodegradation of bioplastics” includes the process of bio-
ammonia, water, heat, and humus. Anaerobic composting degradation of the most commonly used bioplastics. The
involves decomposition in the absence or limited supply of microorganisms and enzymes involved in their degradation
oxygen. Anaerobic microorganisms produce intermediate are enlisted in Table 2. A summary of the percentage of
compounds such as methane, organic acids, and hydrogen degradation of bioplastics from different sources is provided
sulfide. Vermicomposting is the decomposition process that in Table 3.
involves the use of various species of earthworms to produce
a decomposition mixture (Jouhara et al. 2017; Sanchez-Her-
nandez et al. 2020). However, composting rates at home and Sources of bioplastics
in industrial conditions may differ. Home composting is a
basic process that creates nutrient-rich soil by decomposing Broadly, bioplastics are produced from biopolymers
organic waste such as vegetable waste and food leftovers in obtained from biomass, and monomers. Bioplastics are clas-
compost bins. However, the conditions and temperatures for sified based on their source of raw material as depicted in
home composting are not efficient in breaking down bioplas- Fig. 1 and are discussed below.
tics like PLA. Industrial composting on the other hand pro-
vides a strictly monitored composting process that includes Biomass
measured inputs of water, air, and carbon and nitrogen-rich
materials to ensure rapid biodegradation of organic material. Bioplastics made from renewable biomass have sparked
The residues produced can be integrated into the natural geo- curiosity in recent years. Polysaccharides including starch,
chemical cycle (Schrader et al. 2017; Narancic et al. 2018). cellulose, chitin, pectin as well as proteins are extracted from
Further, most industrial composting sites have leachate col- conventional animal and plant sources.
lection and storage systems where the stored leachate can
be pumped into the wastewater treatment plant for further Polysaccharides
processing. This will prevent the leachate from running into
the freshwater bodies and causing eutrophication. Polysaccharides, including starch and cellulose, are the
Biodegradability is the foremost important factor to be primary and most prevalent biobased polymers utilized to
considered when bioplastics are considered over regular manufacture bioplastics (Lubis and Harahap 2018; Abe and
plastics (Din et al. 2020). Bioplastics from various sources Branciforti 2021). Other lesser-known polysaccharides such
exhibit different mechanisms and rates of biodegradation. as chitin and pectin are also used. (Tables 1 and 3). Starch
Biodegradation also depends on several aspects such as is a highly degradable, most abundant homopolysaccharide
environmental factors, moisture, and microbial presence. present in plants, consisting of both linear (amylose) and
Extracellular enzymes secreted by microorganisms assist branched (amylopectin) structures. Corn, potatoes, banana,
the biodegradation of polymers in processes involving the tapioca, wheat, rice, yam, sago, and buckwheat are the tradi-
hydrolysis of ester linkages to release monomers. Both tional sources of starch used in the production of bioplastics
biotic and abiotic processes influence the total breakdown (Marichelvam et al. 2019; Abral et al. 2019; Jiménez-Rosado
of organic matter. Microorganisms break down the material et al. 2019; Asrofi et al. 2020). The synthesis of starch-based
through biotic decomposition, while abiotic processes such bioplastics involves heating native or modified starch in the

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Table 1  Summary of sources, advantages, and disadvantages of various native biodegradable bioplastics
Raw material Origin Advantages Disadvantages References

Polysaccharides
Starch Corn, potato, rice, tapioca, • Good film forming proper- • Brittleness (Thakur et al. 2019)
tapioca/cassava, banana, ties, oxygen and aroma • Poor mechanical strength
wheat, yam, sago and buck- permeability • Hydrophilicity
wheat
Cellulose Cotton, wood, sisal, flax, • Ease of availability • Susceptible to moisture (Silva et al. 2016)
hemp, jute, sugarcane • Low cost of production
bagasse • Transparency
Lignocellulose Wood pulp, jute, hemp, cotton • Antimicrobial properties • Hydrophilicity (Wang et al. 2011)
• Good viscoelastic, film- • Incompatibility with many
forming capacity polymers
• Brittleness
Pectin Apples, guavas, citrus fruits, • Ease of availability • Poor mechanical properties (Liu et al. 2005)
plums • Hydrophilicity
Chitosan Chitin (exoskeleton of crus- • Inherent antimicrobial and • High water sensitivity (Martínez-Camacho et al.
taceans like crabs, lobsters, antifungal activity 2013)
prawns, shrimps) • Good mechanical strength
• Low oxygen and carbon
dioxide permeability
Proteins
Casein Milk, cheese, yogurt, and • High thermal stability • Difficulty in moldability (Dodd 2010)
other dairy products
Whey protein Waste stream of the cheese • Good oxygen and aroma • Moderate moisture (Galus and Kadzińska 2016)
isolate industry permeability • barrier capacity
• Require plasticizers to cre-
ate easy to handle films
Collagen Fish, chicken, egg whites, • Good oxygen and aroma • Relatively low water barrier (Ma et al. 2018)
seafood barrier capacities capacity
• Poor mechanical strength
Zein Corn • Good film-forming proper- • Brittleness (Ghanbarzadeh et al. 2006)
ties after dissolving in
ethanol and acetone
• Good tensile strength and
moisture barrier properties
Soy protein Soybean • Transparent and flexible • Poor mechanical properties (Zheng et al. 2017)
isolate films • High sensitivity to water
• Gas barrier properties
Gluten Waste stream of the wheat • Low cost • High moisture sensitivity (Zhang et al. 1996)
starch industry • Good oxygen barrier capac- • Brittleness
ity
• Good film-forming proper-
ties
Microorganisms
PHAs Ralstonia eutropha • Excellent barrier capacity • Difficulty in sustaining opti- (Zakaria Gomaa 2014)
to carbon dioxide, oxygen, mal growth conditions
and water • High cost of recovery
• Good water resistance
Biobased monomers
PLA Lactic acid • Excellent film-forming • Brittleness and rigidity (Elsawy et al. 2017)
properties • Incompatibility with certain
• Hydrophobic polymers
Synthetic monomer
PCL ε-caprolactone • Hydrophobic • Low melting point (Dong and Walker 2012)
• Oil, solvent, and chlorine
resistance
PBAT Adipic acid, 1,4-butanediol, • Flexible • Low thermal stability (Al-Itry et al. 2012)
and terephthalic acid • Good tensile strength • Stiffness

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Table 1  (continued)
Raw material Origin Advantages Disadvantages References
PEA Hydrophobic α-amino acids, • High thermal stability and • Expensive (Zou et al. 2004)
α, ω-diols, aliphatic dicar- tensile strength
boxylic acids and dianhy-
drohexitoles

PLA Polylactic acid; PHA Polyhydroxyalkanoates; PCL Polycaprolactone; PBAT Polybutylene adipate terephthalate; PEA Polyesteramides

presence of water to initiate gelatinization. As much as fifty linear molecule composed of β- (1 → 4) -linked d-glucosa-
percent of commercially available bioplastics are produced mine and N-acetyl-d-glucosamine. Owing to its non-toxicity
from starch (Erabela et al. 2021). Due to their simple and and biodegradability, chitosan was extensively studied for
cost-effective production process, starch-based bioplastics its ability to form environment-friendly plastics (Table 1). It
have a broad range of applications in the packaging industry is one of the few polysaccharides known for its native anti-
(Liang and Wang 2018; Meereboer et al. 2020). Cellulose, microbial properties against gram-negative, gram-positive
another highly abundant homopolysaccharide consisting of bacteria, algae, and fungi. Temperature is also a crucial fac-
repeating units of D-glucose bound together by β (1 → 4) tor in evaluating the antimicrobial activity of chitosan-based
glycosidic bonds is typically added to starch to enhance its films/bioplastics, according to a report (Leceta et al. 2013).
mechanical properties, gas permeability, and water resist- The film-forming solution showed antibacterial activity but
ance. The most popular raw material used to produce cellu- was only found to be bacteriostatic after being dried into
lose-based bioplastics is softwood (Liu et al. 2019b). Cel- films. The use of various polyols, like glycerol, sorbitol, and
lulose is also used in different forms as a reinforcement filler polyethylene glycol as plasticizers for chitosan is studied
in other biodegradable polymer matrices to improve their extensively. The physicochemical properties of the film are
mechanical and physical properties. Lignin is a complex influenced by the type and volume of plasticizer used (Ma
plant-based polymer found in vascular tissues as a support- et al. 2019; Jha 2020).
ing structure. It is called lignocellulose since it is found in
cell walls alongside cellulose and hemicellulose. The com- Proteins
position of lignin in the lignocellulose varies according to
the botanical source (Le Digabel and Avérous 2006). Lately, Many proteins, of both plant and animal origin, are used as
lignocellulosic fibers are being utilized as bioplastic rein- viable options for the creation of packaging materials.
forcements in place of synthetic fibers due to their ability to
be biodegraded and renewable. A significant number of stud- Proteins of animal origin Casein is a milk protein belonging
ies have focused on combining lignin with other bio-based to the family of phosphoproteins and is majorly found in
polymers like starch, cellulose, and polylactic acid (PLA) to mammalian milk (Shivani et al. 2021). To form a bioplastic,
synthesize biodegradable bioplastics (Kai et al. 2016; Shi casein is dispersed in an aqueous alkali solution, followed
and Li 2016; Brodin et al. 2017). Pectin is a heteropolysac- by coagulation with an acid or salt of an acid (Yong et al.
charide distributed in the primary lamella, middle lamella, 2021). The resulting coagulum is pressed into plates and
and cell walls of plants. It is made up of esterified d-galactu- soaked in formaldehyde until it hardens to obtain the casein-
ronic acid units bound together by α-(1–4) glycosidic bonds based bioplastic. Rennet casein was extruded with water in
(Ropartz and Ralet 2020). Citrus fruits like oranges, lemons, the presence of high moisture and pressure to obtain pliable
gooseberries, and strawberries are rich in pectin, which was films. Despite its low cost, the film was not easily moldable,
used to develop biodegradable bioplastics for the food and and as a result, it was phased out of commercial use (Dodd
packaging industries. However, its applications are limited 2010). Furthermore, the formaldehyde used was a prob-
due to its inadequate mechanical properties and hydrophilic able carcinogen that can cause genetic abnormalities, mak-
nature. (Shrestha et al. 2021). Like other polymers, several ing its use hazardous (Jefferson et al. 2020). Whey protein
studies reported the addition of plasticizers like polyols to is the by-product of cheese manufacturing, i.e., the liquid
help in tackling the brittleness of the films. Also, pectin portion of milk after it is curdled and strained. The ability
is blended with different additives like sodium alginate, of whey to produce biodegradable, odorless, and transpar-
starch, cellulose, and chitosan to improve the properties of ent films are being extensively researched (Oliveira et al.
the resulting bioplastic (Nešić et al. 2017; Younis and Zhao 2017; Schmid and Müller 2018). The films are formed by
2019). Chitin makes up the exoskeleton of crustaceans and is casting and drying the whey protein isolate. But these films
one of the most abundant polysaccharides. Chitin undergoes are highly brittle due to their disulfide crosslinking and are
N-deacetylation in an alkaline medium to form chitosan, a hence hydrophobic. To reduce the intermolecular interac-

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 Table 2  Summary of microbes and enzymes responsible for degradation of biodegradable biopolymers
Biodegradable polymer source Microorganisms Enzymes Reference
Bacteria Fungi

Polysaccharides
Starch Bacillus sp: B. amyloliquefaciens, Clonostachys rosea, Trichoderma sp Amylase (Alariya et al. 2013; Urbanek et al.
3 Biotech (2023) 13:220

B. subtilis, • α amylase; endoamylase (EC 2017; Luang-in et al. 2019)

B. licheniformis, 3.2.1.1)
B. stearothermophilus, • β amylase; exoamylase
B. megaterium, B. circulans (EC 3.2.1.2)
• Glucoamylase (EC 3.2.1.3)
Cellulose Clostridium, Cellulomonas, Cel- Trichoderma, Aspergillus, Penicil- Cellulases: (Liang et al. 2014; Fernandes et al.
lulosimicrobium, Thermo- lium, Phanerochaete, and Fomi- • β-1,4-endoglucanase (EC 3.2.1.4) 2018)
monospora, Bacillus, Rumi- topsis • Cellobiohydrolase or exoglucanase
nococcus, Erwinia, Bac- (EC. 3.2.1.91)
teriodes, Acetovibrio, • β-glucosidase or cellobiase (EC.
Streptomyces, Microbispora, Fibro- 3.2.1.21
bacter,
and Paenibacillus
Lignocellulose Streptomycesviridospo- Basidiomycetes, Ascomycetes Ligninolytic enzymes: (Bugg et al. 2011; Kumar and Chandra
rus T7A, Nocardia autotroph- • Laccase (EC 1.10.3.2) 2020)
ica, Sphingobium sp. SYK-6, • Lignin peroxidase
Pseudomonas putida mt2, Rhodococ- (EC 1.11.1.14)
cus sp., • Manganese peroxidase
Burkholderia cepacia, Microbacte- (EC 1.11.1.13)
rium sp., and Citrobacter sp
Chitin Streptomyces, Alteromonas Escheri- A. nidulans, N. crassa, M. oryzae, Chitinases: (Yang and Zhang; Beier and Bertilsson
chia, Aeromonas, Serratia marces- Schizosaccharomyces pombe, S. Glycosyl hydrolases 2013; Hamid et al. 2013)
cens cerevisiae, A. oligospora (E.C 3.2.2.14)
Pectin Streptomyces sp, Cellulomonas sp, Aspergillus fumigatus, A. flavus, A. Pectinases (EC 3.2.1.15): (Ramos 2011; Shrestha et al. 2021)
Bacillus sp niger, A. ochraceus, A. oryzae, A. • Pectin methyl esterase (EC
sydowii, Trichothecium sp., Penicil- 3.1.1.11)
lium sp., Trichoderma harzianum • Polygalacturonases (endo-, EC
and T. viride 3.2.1.15; exo-, EC 3.2.1.67)
• Pectin transeliminases
(EC 4.2.2.10)
Proteins
Casein, whey, Bacillus sp Penicillium, Aspergillus, Rhizopus, Proteases /Proteolytic enzymes (Razzaq et al. 2019)
soy protein, gluten Endothia, Mucor
Microorganisms

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Page 5 of 22 220
 Table 2  (continued)
220

Biodegradable polymer source Microorganisms Enzymes Reference

Bacteria Fungi

PHAs Enterobacter sp, Bacillus sp, Gracili- Penicillium pinophilium, Penicil- Polyhydroxyalkanoate (PHA) depoly- (Volova et al. 2010; Singh et al. 2020)
Page 6 of 22

bacillus sp., lium funiculosum, Paecilomyces merase (EC 3.1.1.75) (Tokiwa et al. 2009)
Pseudomonas lemoigne, Coma- lilacinus, Aspergillus fumigatus,
monas sp. Emericellopsis minima

13
Acidovorax faecalis, Aspergillus
fumigatus,
Variovorax paradoxus,
Streptomyces sp,
Aspergillus sp.
Bioderived monomers
PLA Bacillus pumilus Tritirachium album, • Carboxylesterase (EC 3.1.1.1) (Tokiwa et al. 2009; Bonifer et al.
Fusarium moniliforme, Penicillium • Cutinases (EC 3.1.1.74) 2019; Butbunchu and Pathom-aree
roqueforti, Clonostachys rosea and • Lipases (E.C. 3.1.1.3) 2019)
Trichoderma sp • Serine proteases
(EC 3.4.21.14)
PBAT Thermobifida fusca, Pelosinus • Carboxylesterase (EC 3.1.1.1) (Wallace et al. 2017)
fermentan,s • Lipases (E.C. 3.1.1.3) (Tokiwa et al. 2009; Singh et al. 2020)
lostridium botulinum, Pseudomonas
sp
PEA Aspergillus, Aureobasidium, Penicil- Penicillium sp, R. arrizus, R.delemar,
lium, Pullularia Achromobacter sp. and Candida
cylindracea
PCL Clostridium sp Aureobasidium sp., Cryptococcus
sp., Aspergillus flavus, A. niger, A.
fumigatus, Chaetomium globosum,
Pencillium funiculosum, Fusarium
sp.
Clonostachys rosea, Trichoderma sp.

PHA Polyhydroxyalkanoates, PLA Polylactic acid, PBAT Polybutylene adipate terephthalate, PEA Polyesteramides, PCL Polycaprolactone
3 Biotech (2023) 13:220
3 Biotech (2023) 13:220 Page 7 of 22 220

Table 3  Table comparing the percentage of biodegradation of some bioplastics from various source
Sources Mass loss (%) Conditions of biodegradation Approx. Reference
Duration
(days)

Biomass
Polysaccharides
Starch ̴ 90 Organic compost Moisture content: 50% 31 (Torres et al. 2011)
Cellulose ̴ 30 pH: 7.0–8.0 31
Chitin 83.8 Burial in red clay 31 (NAKASHIMA et al. 2005)
Lignin 19–60 Laboratory incubation 91- 730 (Zabel and Morrell 2020)
Pectin-cellulose composite 90 Anaerobic digestion in batch reactors 15 (Bátori et al. 2017)
maintained at 55 °C
Protein
Soy protein 30 Aqueous solution containing pronase 6 (Yamada et al. 2020)
(proteolytic enzyme)
Casein 20 Loam soil 7 (Bagares et al. 2020)
Zein 58 Saturated field soil with Fafard® 52 84 (Helgeson et al. 2009)
medium
Soy protein and whey protein isolate 36 Composting 7 (Li and Chen 2000)
composite
Microorganisms
PHA 88–99 Under optimum static laboratory condi- 49 (Siracusa et al. 2008)
tions
PHB 62 Fertile garden soil with pH 7.30 and 62 (Altaee et al. 2016)
PHB–TiO2 composite nanofiber films 100 humidity of 80% at 30 °C 21
PHBV 100 Lab-scale composting 60–80 (Salomez et al. 2019)
Monomers
Bio-derived monomers
PLA 88 Composting 90 (Kawashima et al. 2021)
Synthetic monomers
PCL 92 pH 13 63 (Sailema-Palate et al. 2016)
80 pH 1 69
100 Composting 1460 (Manoukian et al. 2019)
PBAT 2.38 Real soil burial 91 (Wang et al. 2015)

PHA Polyhydroxyalkanoate, PHB Polyhydroxybutyrate, TiO2 Titanium dioxide, PHBV Poly(3-hydroxybutyrate-co-3-hydroxy valerate, PLA Pol-
ylactic acid, PCL Polycaprolactone, PBAT Polybutylene adipate terephthalate

Fig. 1  Classification of bioplas-

tics based on sources (biomass,
microorganisms, and mono-
mers). The figure is adapted
from (Avérous 2004) with kind
permission from Taylor &
Francis

tions and overcome brittleness, it is treated with various gradually, while mechanical resistance, Young’s modulus,
plasticizers like glycerol and sorbitol. With the addition of and glass transition temperature decreased (Galietta et al.
plasticizers, the percentage of water solubility improved 1998). Collagen is another protein widely studied for its

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220 Page 8 of 22 3 Biotech (2023) 13:220

film-forming properties (Table 1). It is a structural pro- by alkaline suspension, followed by isoelectric precipi-
tein present in the extracellular matrix of connective tissues tation at pH 4.5 (Chove et al. 2001). Attributed to the
like skin, cartilage, bones, and tendons. Collagen undergoes prevalence of polar functional groups such as hydroxyl,
partial hydrolysis to form gelatin. It is commonly used in thiol, carboxyl, and amine in the molecule, the films are
the fabrication of scaffolds, wound dressings, drug delivery hydrophilic and have poor mechanical strength (Ye et al.
systems, and packaging materials (Ge et al. 2018; Irawan 2019). This limitation can be overcome by performing
et al. 2018; Lin et al. 2019; Bhuimbar et al. 2019). Collagen- certain surface modifications such as crosslinking with
based films are known for their excellent biocompatibility, chemical agents like formaldehyde, glutaraldehyde, and
biodegradability, and oxygen and aroma barrier capacities. phenolic compounds (Insaward et al. 2015). The resulting
However, the applications of these collagen-based films are films from soy protein are brittle, hence various plasticiz-
limited in packaging due to their relatively low water barrier ers are used to reduce this and improve the processability
capacity and mechanical strength. To improve these proper- and flowability of the films (Božič et al. 2015). Improve-
ties, collagen is often combined with chitosan, soy protein, ment of SPI-based bioplastic properties is reported to be
and reinforcement fillers such as zinc oxide nanocrystals, dependent on characteristics such as the size and polar-
cellulose, and magnesium oxide nanoparticles (Ahmad ity of the plasticizer used. Microbial contamination is
et al. 2016; De Silva et al. 2017; Andonegi et al. 2020; Jiang another downside of the material. Several studies have
et al. 2020). used different nanoparticles and metal ions to impart anti-
microbial activity to SPI-based films (Jin et al. 2020). A
Proteins of plant origin Zein belongs to a group of plant self-healing, antimicrobial, SPI bioplastic was developed
storage proteins called prolamins, found majorly in the with good mechanical strength using polyethyleneimine
endosperm of maize (corn), accounting for 47% of the (PEI) and metal ions such as ­Cu2+ or ­Zn2+. It is believed
total endosperm content (dry basis) in corn (Shukla and that the polycationic property of PEI is responsible for
Cheryan 2001). It is used in textile industries to impart disrupting bacterial membranes through ion exchange (Li
hydrophobicity to the fabric. The hydrophobicity of the et al. 2019). Several other studies have used different addi-
molecule is attributed to the recurrence of many non- tives like cellulose nanocrystals, zinc oxide nanoparticles
polar amino acids like alanine, leucine, and proline in the (Xiao et al. 2020), cortex philodendron extract (Liang and
structure. The fabric can also be rendered antimicrobial by Wang 2018), grape seed extract, nisin, EDTA (Sivarooban
encapsulating ellagic acid in zein molecules (Gonçalves et al. 2008), and organic acids like citric, lactic, maleic
et al. 2020). Zein also finds its applications in the produc- acids (Eswaranandam et al. 2004), etc. Gluten belongs to
tion of ceramics for bone tissue repair (Hum et al. 2018), a family of seed proteins found in cereal grains like barley,
drug-delivery systems (Labib 2018), cosmetics (Tinoco rye, and majorly in wheat. The total wet and dry gluten
et al. 2021), adhesives (Wei et al. 2020), etc. The plasticiz- content of wheat ranges from 17.8 to 47.23% and 5.9 to
ing property of zein is widely studied and used in prepar- 10.1% respectively. The viscoelastic property of gluten is
ing thermostable, biodegradable zein-based bioplastics. exploited in the synthesis of biodegradable plastics. Fac-
The resulting zein-based bioplastic exhibits excellent gas tors like the production technique, working duration, tem-
barrier properties and hydrophobicity and is, therefore, perature, pH, plasticizer, and additive content can influ-
used in modified atmosphere packaging (MAP) and coat- ence the properties of the film (Yu et al. 2016). It was
ing on fruits and vegetables to preserve their freshness. reported that gluten-based films obtained through extru-
MAP is a technique that involves the utilization of pack- sion were found to have better plasticizer–gluten inter-
aging films to control and modify the atmosphere around action and water uptake capacity when compared to that
the packed product. The food is either sprayed, brushed, produced through compression molding. The study also
or dipped in the plasticized zein solution and allowed to reported that an increase in pH toward the alkaline side
solidify to form a thin coat (Neo et al. 2013). These bio- improved the water uptake capacity of the wheat gluten
plastic coatings improve the shelf life of the food products films whereas those with additives like xanthan gum and
like apples, pear, mango, tomato, broccoli, rice, cheese, glyoxal showed lower water uptake (Jiménez-Rosado et al.
and roasted peanuts by altering certain processes like 2019). Proteins are heteropolymers, unlike many other
delaying respiratory rate, sprouting, germination, ripen- biodegradable polymers discussed above. The presence of
ing, rancidity, reducing loss of moisture, and inhibiting various amino acids provides a wide range of chemical
microbial growth (Koh et al. 2018; Santos et al. 2018b, a; functions, which can result in a broad spectrum of poly-
Zhang et al. 2019). Soy protein is a protein that is extracted mer network architectures. Proteins such as gluten provide
from dehulled and defatted soybeans. The purified form of unique and favorable properties like viscoelasticity and
soy protein called the soy protein isolate (SPI) is obtained flow properties to the developed bioplastic (Table 1).

13
3 Biotech (2023) 13:220 Page 9 of 22 220

Polyesters of bacterial origin monomer depends on factors such as the properties required
for the final product, the cost of production, and the avail-
Polyesters are polymers made up of a carboxylic acid and ability of renewable resources.
diol group (Sudesh et al. 2000). A variety of bacteria that
have been grown under various nutrient and environmen- Bio‑derived monomers
tal conditions, produce polyesters. These molecules, which
are mainly lipid-based, are collected as storage resources, Polylactic acid (PLA) is an aliphatic polyester formed by
facilitating microbial survival in stressful situations. Based condensation of lactic acid. It can also be obtained by the
on the source organism, the granular size and number, mac- ring-opening polymerization of lactide (Mehta et al. 2005).
romolecular structure, monomer composition, and phys- PLA has been hailed as a promising, environment-friendly
icochemical characteristics change (Luengo et al. 2003). polymer due to its low toxicity and sustainability (Mulvihill
Most of these substances have the ability to break down et al. 2011). Like any other polymers, PLA films have a
naturally and are compatible with living organisms, making high modulus and are brittle (Xu et al. 2020). The use of
them highly interesting to the biotechnology industry. When plasticizers reduces the intermolecular forces of the poly-
provided with vital nutrients such as oxygen, phosphorus, meric chains, resulting in better processability, flexibility,
nitrogen, sulfur, and magnesium, bacteria can proliferate and ductility. Citrate, adipate, and oligomeric lactic acid
rapidly. However, during the prevalence of imbalance in the are by far the most widely used plasticizers for PLA (Shirai
growth environment, i.e., in the excess of carbon source and et al. 2015; Singh et al. 2019). Plasticizers from packaging
deficiency of essential nutrients, mainly phosphorus, and materials tend to migrate into food over time, posing a risk
nitrogen, a variety of bacterial and archaeal genera accumu- to one's health. To delay the migration of plasticizers, fillers
late storage polymers named polyhydroxyalkanoates (PHAs) like wheat bran and nano-additives such as chitin nanofi-
(Wong et al. 2012). One of the most thoroughly researched brils (CN) are added to the blend. A study was carried out
microorganisms for the manufacture of PHA is Ralstonia to investigate the ability of chitin nanofibrils and calcium
eutropha or Alcaligenes eutrophus. They are synthesized and carbonate (both micrometric and nanometric) in preventing
accumulated as highly refractive granules in the cytoplasm or controlling the leaching of the plasticizer acetyl n-tributyl
of the microorganisms. PHA acts as a carbon and energy citrate (ATBC) in PLA/PBS blends. It was found that the
reserve helping in the survival of bacteria under nutrient- addition of both chitin nanofibrils and micrometric calcium
deficient conditions (Sirohi et al. 2021). The bacteria also carbonate helped to slow down the leaching of the plasticizer
show enhanced stress tolerance toward ultraviolet (UV) irra- (Aliotta et al. 2020, 2022). Melt mixing PLA with other
diation, heat, and osmotic shock (Kadouri et al. 2005). Per- polymers such as polycaprolactone (PCL), polybutylene suc-
taining to the microbe and growth conditions, different types cinate (PBS), starch, polyhydroxy butyrate (PHB), polyvi-
of PHAs are synthesized. More than 150 types of PHAs nyl alcohol (PVOH), and polybutylene adipate terephthalate
have been identified so far. Polyhydroxybutyrate (PHB) is (PBAT) is another way to boost its mechanical properties
a well-known polymer belonging to the PHA family. Other (Anna and Arrigo 2019; Olaiya et al. 2019; Kahvand and
members include polyhydroxy valerate (PHV) and poly Fasihi 2019; Xiang et al. 2020). In conclusion, one of the
(3-hydroxybutyrate-co-3-hydroxy valerate) (PHBV). PHA most appealing features of PLA plastic is that its properties
is recovered from the bacterial cell through lysis of the cell can be modified by additives, expanding its potential appli-
wall, followed by solubilization, purification, and precipita- cations (Table 1). Additionally, PLA is biodegradable under
tion of PHA polymer (Kunasundari and Sudesh 2011). PHAs environmental conditions. However, PLA polymers do have
are biodegradable, non-toxic, biocompatible, water-resistant, some limitations, such as softening at 60 °C. Copolymeriza-
and highly crystalline. Owing to their novel features, PHA tion with a more heat-resistant polymer or the incorporation
bioplastics are used in the manufacture of biodegradable of fillers can aid in enhancing its thermal stability.
packaging materials (Meereboer et al. 2020). The need to
maintain optimal growth conditions and the high cost of 2.2.2. Synthetic monomers
recovery are the limitations of PHA bioplastics. However,
the utilization of recombinant microorganisms can overcome Polycaprolactone (PCL) is a biodegradable polyester derived
these disadvantages (Carpine et al. 2020). from fossil or crude oil. It was one of the first synthetic
polymers to be developed. It is obtained from the monomer
Monomers ε-caprolactone employing ring-opening polymerization.
PCL is known for its hydrophobicity, flexibility, mechanical
Several monomers (both naturally occurring (bio-derived) strength, excellent blend compatibility, and biodegradability
and synthetic) are combined by the process of polymeriza- (Woodruff and Hutmacher 2010). It is used in the synthesis
tion to produce efficient bioplastics. Overall, the choice of of polyurethanes and as biodegradable, hot melt adhesives

13
220 Page 10 of 22 3 Biotech (2023) 13:220

(HMAs) in food packaging. It is also gaining popularity in bioplastics can be used for various applications from food
the biomedical field in the fabrication of biodegradable scaf- packaging to textiles and biomedical applications. Thermo-
folds, drug delivery systems, wound dressings, and contra- plastic starch is used to synthesize food packaging materi-
ceptive devices (Kim and Kim 2014; Siddiqui et al. 2018). als, disposable utensils, and compostable trash bags. PHAs
Among the aromatic co-polyesters, polybutylene adipate exhibit immense applications in medical applications includ-
terephthalate (PBAT) is one of the most promising materi- ing the development of long-term drug release capsules, and
als with great potential in various applications (Table 1). tissue scaffolds for neural regeneration (Meereboer et al.
It is a biodegradable, synthetic, block copolymer of adipic 2020). However, certain detailed advantages and disadvan-
acid, 1,4-butanediol, and terephthalic acid (Jian et al. 2020). tages of bioplastics from various sources are mentioned in
It is manufactured under different commercial names such Table 1.
as Ecoflex, Origo-Bi, Ecoword, and Wango (Siegenthaler
et al. 2012; Jacquel et al. 2015). PBAT has a random struc-
ture, hence the name random copolymer. The lack of struc- Biodegradation of bioplastics
tural order makes the material very stiff and of low elastic
modulus; however, it is tough and flexible. The mechanical The biodegradation of bioplastics from various sources
strength of the film depends on the composition of mono- depends on their physicochemical properties as well as envi-
meric units and their molecular weight. Young's modulus ronmental factors, such as the soil and its essential micro-
increases with increasing terephthalate units while decreas- bial diversity. Studies have shown that regardless of the
ing the elongation at break. Similarly, the tensile strength source and type of bioplastic, soil enrichment significantly
is positively correlated to the molecular weight while the enhances the rate of biodegradation. Additionally, factors
elongation at break shows a negative correlation (Jian et al. such as humidity and temperature play an important role in
2020). The blend of PBAT and other strong, biodegrad- the biodegradation of bioplastics. (Zoungranan et al. 2020).
able polymers can yield robust and rigid films. Over the Further, several components, especially plasticizers, are a
years, PBAT has found its use in cling films for packaging, necessity while synthesizing bioplastics. Water is the major
mulch films, water-resistant coatings (Souza et al. 2020), solvent and plasticizer in biopolymer technology. Apart
etc. Polyester amides (PEA) are a new class of biodegrad- from water, polyols, mono, di-, and oligosaccharides are the
able polymers that are used in diverse industrial applications most used plasticizers. Polyols such as glycerol, erythritol,
(Table 1). PEA can be produced from different monomers and sorbitol have 3, 4, and 6 carbons and hydroxyl groups,
using a variety of synthesis techniques like ring-opening respectively (Lang et al. 2020). It is reasonable to use these
and polycondensation methods (melt, solution, and interfa- organic compounds as plasticizers for biopolymers given
cial polycondensation (Rodriguez-Galan et al. 2011). These their completely biodegradable nature. In addition, the use of
monomers have ester and amide linkages in their chemical plasticizers also assists the quick degradation of bioplastics.
structure that can be easily degraded by bacteria. Hydropho- In a study, it was revealed that the weight loss of unplasti-
bic α-amino acids, α, 𝜔-diols, aliphatic dicarboxylic acids, cized PVOH/RWF films was lower than glycerol- and sorb-
and dianhydrohexitoles are some of the monomers used in itol-plasticized polyvinyl alcohol/rambutan skin waste flour
the synthesis of PEA films (Gomurashvili et al. 2000). PEA films (Ooi et al. 2012).
combines the benefits of polyesters and polyamides in a pol- The degradation of the bioplastics with the aid of using
ymer. The synthesized polymeric films possess good ther- soil microorganisms is evident, and numerous microorgan-
mal, chemical resistance, and mechanical properties. Over isms including Bacillus sp. and Aspergillus sp., are isolated
the years, various fillers have been added to enhance the and recognized as degraders of bioplastics in soil (Adhikari
performance and durability of PEA films. A study reported et al. 2016). Table 2 summarizes the common microbes and
that melt-blending of octadecyl amine-treated montmoril- enzymes responsible for degrading biodegradable bioplas-
lonite clay with PEA showed a slight improvement in the tics. Complete biodegradation (≥ 90%) prevents accumula-
oxygen and water vapor barrier properties (Krook et al. tion in soil, which is the initial level of testing biodegra-
2002). Another study reported that the addition of nano- dability (Ardisson et al. 2014). The steps involved in the
silicon dioxide and nano-calcium carbonate as fillers for microbial degradation of polymers have been depicted in
PEA showed a significant increase in tensile strength and Fig. 2. However, the rate of biodegradation of bioplastic
reduction in the rate of hydrolysis (Liu et al. 2007). depends on the soil conditions. The most widespread pro-
Biodegradable bioplastics from various sources men- cedures used to examine the biodegradation of bioplastics
tioned above may have a wide variety of uses based on their in soil are visual analysis, detecting mass loss, spectroscopy,
respective properties such as texture, mechanical strength, and ­CO2 emission (Zilliges and Damrow 2017; Ruggero
hydrophilicity, gas barrier properties, biodegradability, et al. 2019; Folino et al. 2020). The volume of C­ O2 produced
and many more. Based on these properties, the developed during biodegradation is used as the index of microbial

13
3 Biotech (2023) 13:220 Page 11 of 22 220

Fig. 2  Sequential steps of poly-

mer degradation by microbes in
the soil. The figure is adapted
from (Kumar Tiwari et al. 2018)
with kind permission from
Granthaalayah publications

decomposition. ­CO2 emission can be analyzed using two smaller fragments that are further hydrolyzed by an
methods namely the cumulative measurement respirometry array of enzymes including glucoamylase, β-amylase,
(CMR) and gravimetric measurement respirometry (GMR) and α-glucosidase. These enzymes further hydrolyze
(Ruggero et al. 2019). Visual analysis techniques include the α-glycosidic linkages (Encalada 2018). Enzymes
scanning electron microscopy (SEM), macroscopic pho- that can accomplish starch hydrolysis are present in the
tographs, and atomic force microscopy (AFM) to examine soil. Several fungi including Aspergillus oryzae and bac-
surface changes in the material. The visual evaluation com- teria such as Klebsiella pneumonia, Bacillus circulans,
monly includes checking for the distribution of particle size and Bacillus stearothermophilus synthesize an array of
in residual bioplastic and the presence of microbial coloni- starch hydrolyzing enzymes. Further lytic polysaccharide
zation (Marga et al. 2005). Other factors include noticeable monooxygenases (LPMOs) produced by a variety of bac-
degradation phenomena such as loss of consistency, thick- teria like Escherichia coli and fungi such as Thermoascus
ness, discoloration, and the presence of physical disintegra- aurantiacus, and glycoside hydrolases are also efficient in
tion like holes or cracks. Further, SEM and AFM are used breaking glucose bonds by oxidative cleavage. The bio-
to thoroughly observe the morphological changes occurring degradation of thermoplastic starch (TPS) was compared
during biodegradation like crack formation, surface rough- under laboratory conditions (soil and compost) and in the
ness, and corrosive degradation (Folino et al. 2020). Another field (soil). Based on microbial activity, it was concluded
index of studying biodegradation is the percentage of mass that fungi have a greater ability to biodegrade TPS than
loss. It includes measurement of mass loss for pieces of bacteria. It was also observed that both the crystalline
bioplastics extracted after soil burial post the experimental and molecular structures of TPS films are factors that
period. The standardized procedure includes sample screen- influence the enzymatic degradation of TPS by fungal
ing, washing with distilled water, followed by drying and α-amylase. Absolute degradation of TPS films containing
weighing (Salomez et al. 2019). higher moisture levels and buried 20 cm deep in soil was
observed in 4–6 months (Polman et al. 2021). In another
Biomass study conducted, two batches of cassava starch films were
synthesized, one set crosslinked with citric acid (TPS75-C,
Polysaccharides heated at 75 ℃ and TPS85-C, heated at 85 ℃) and another
without any citric acid crosslinking (TPS7, heated at 75 ℃
Starch is primarily degraded by glycoside hydrolases, and TPS85, heated at 8 ℃) were subjected to biodegrada-
enzymes that hydrolyze the glycosidic bonds. The enzyme tion for 30 days. The degradation process of TPS75 and
α-amylase cleaves the long starch polymers, producing TPS85 was considerably high in 12 days, while the citric

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220 Page 12 of 22 3 Biotech (2023) 13:220

acid cross-linked films (TPS75-C and TPS85-C) required polyethylene plastic film which demonstrated no indications
18 days (Seligra et al. 2016). of degradation (Ai et al. 2021).
Fungi form the major share of cellulolytic (cellulose- In a study, it was found that a thermoplastic starch
degrading) microorganisms (Tian et al. 2017). Bacteria such strengthened with 50% wt. lignocellulosic fibers and flax
as Pedobacter and Mucilaginibacter revealed intricate enzy- fibers degraded considerably slower than those of native
matic systems for the degradation of polysaccharides includ- palm, banana, and bagasse (sugar cane residue) (Jumai-
ing cellulose and hemicellulose. Cellulases are responsible din et al. 2021). Pectin is degraded by a group of enzymes
for extracellular cellulose degradation. These enzymes called pectinases (Table 2). These enzymes work by depo-
belong to the glycoside hydrolases and are adept at break- lymerization and de-esterification of the pectin. Pectinolytic
ing the β-glycosidic bonds. LPMOs are involved in the initial bacteria such as Erwinia spp. degrade pectin by producing
phase of cellulose biodegradation (Polman et al. 2021). The such enzymes (Abbott and Boraston 2008). The anaerobic
degradation of cellulose produces cellobiose, a disaccharide degradation study of pectin-cellulose biofilms from orange
of glucose. Depending on the soil condition, cellulose deg- peel wastes exhibited 90% degradation in about 15 days. A
radation may require 81–495 days. Cellulose acetate (CA) highly biodegradable high-methoxy pectin (HMP) film with
is generated by the acetylation of cellulose. These acetyl dialdehyde starch (DS) (0%, 25%, 50%, 75%, 100%) was
groups hinder the microbial and enzymatic attack on the CA- developed. Increasing the content of DS caused a decrease
based bioplastics and prolong their time of biodegradation in the biodegradation percentage (Fig. 4) (Bátori et al. 2017).
(Hayakawa et al. 2014). However, biodegradation may take Chitin and chitin-based bioplastics are degraded by the
place through oxidative cleavage by LPMOs and hydroly- action of bacteria and fungi in soil by a process called,
sis by cellulases. The collaboration of cellulases, LPMOs, chitinolysis via chitinases. Chitin undergoes deacetylation
and acetyl esterases is known to degrade CA with a degree
of substitution of less than 1.8. The faster degradation of
flax fibers was attributed to slight chemical differences
between the flax and cotton-based cellulose acetate (CA).
Complete breakdown of CA from rice straw was observed
after 105 days in the soil. During biodegradation, the degree
of substitution of the residual CA remained unaffected, as
enzymatic hydrolysis occurs at a higher rate than deacetyla-
tion (López-Mondéjar et al. 2016). In a study, regenerated
cellulose films were prepared from cellulose solution from
pulps of cotton stalk, cotton linter, and wheat straw. They
were buried in the soil to assess biodegradability (Fig. 3a).
Results indicated 10% weight loss for film in 16 days (Zhang Fig. 4  Digital macroscopic images for high-methoxy pectin (HMP)
et al. 1996). In another study, cellulose film was observed to film with varying concentrations of DS films after burial in soil for
15 days: a 100HMP/0DS, b 75HMP/25DS, c 50HMP/50DS, d
be decomposed in soils within only 4 weeks (Fig. 3b), sug- 25HMP/75DS, and e 0HMP/100DS. The image has been adapted
gesting an exceptional biodegradability as compared to the from (Bátori et al. 2017) with permission from Hindawi

Fig. 3  Biodegradability test

showing changes in the mor-
phology of cellulose films: a
Macroscopic images showing
degradation of cellulose film
over 4 weeks. The image has
been reproduced from (Ai et al.
2021) published on Frontiers. b
SEM images showing biodegra-
dation of regenerated cellulose
films from pulps of cotton linter,
cotton stalk, and wheat straw
over 40 days. The image has
been reproduced from (Zhang
et al. 1996) with permission
from the American Chemical
Society

13
3 Biotech (2023) 13:220 Page 13 of 22 220

to produce chitosan, which is hydrolyzed by chitosanases of 180° from each other forming a rigid elongated struc-
(Gooday 1994). Bacterial species ofAeromonas, Vibrio, ture (Conley et al. 2016). However, the rate of bioplastic
Cytophaga, Photobacterium, Streptomyces, Bacillus, Chro- biodegradation is faster in a composting plant compared to
mobacterium, and Clostridium are well-known chitinolytic natural conditions due to the continuous supply of air as well
bacteria. Chitinolytic fungal species include Mucorales as humidity and temperature control. The biodegradation
like Deuteromycetes and Mortierella spp, and Ascomycetes of starch-based bioplastics in the soil is depicted in Fig. 6.
like Aspergillus, Thielavia, Trichoderma, Humicola Penicil- However, several factors such as the presence of additives
lium, and Verticillium (Moussian 2019). The biodegradation affect the time taken for the degradation of bioplastics.
of chitin and chitosan films revealed that after 1, 1.5, and
2 months, chitin films buried in red clay achieved 83.8%, Proteins of plant and animal origin
99%, and 100% weight loss, respectively. In the case of chi-
tosan films, the weight loss was 79.2%, 98.9%, and 100% for According to a study conducted by Bagares et al., the deg-
the same periods and conditions. (Nakashima et al. 2005). radation of rapeseed protein-based bioplastic was 57% and
Figure 5 exhibits the SEM images of chitin and chitosan 74% in the soil and liquid medium, respectively. The bio-
films before and after biodegradation. In another study, the plastic produced from casein-based bioplastic from spoiled
biodegradation of two composite films polyethylene-chitin processed cow’s (Bos taurus) milk took seven days to be
(PE-chitin) and polyethylene-chitosan (PE-chitosan) films, degraded with the help of catalysts such as scavenger insects
containing 10% chitin or chitosan, by pure microbial cultures in the presence of sunlight and rain (Bagares et al. 2020).
in a soil environment was studied and compared to commer- Further, the transparent whey protein isolate films began to
cial starch films. In soil, 73.4% of the PE-chitosan and 84.7% degrade within only 2 days and completely degraded over
of PE-chitin degraded, while only 46.5% of the commercial 7 days with more than 80% of weight loss. Figure 7a shows
starch film degraded after six months (Lee 1995). a scanned image of whey protein isolate films after 10 days
It was observed that starch-based bioplastic biodegrades of soil burial. Similarly, there are reports of soy protein and
much faster compared to bioplastics developed from other whey protein isolate composite films being degraded in
polysaccharides such as cellulose (Table 3) based on the about 7 days with 36% of weight loss (Li and Chen 2000).
prevailing conditions including the microbial flora. This may In another study, hydrophobic zein-based bioplastic films
be attributed to the fact that cellulose as a polysaccharide is containing licorice essential oil were developed (Luís et al.
much more stable compared to starch due to its molecular 2019). The biodegradability of the films was analyzed by
orientation where opposite molecules are placed at a rotation soil burial degradation test for 10 days (Fig. 7b). The films

Fig. 5  SEM images of a, b, c

chitin films and d, e, f chitosan
films buried for 1 month in
red clay. a and d SEM images
of chitin and chitosan films,
respectively, before soil burial.
b and c SEM images of chitin
films; e and f SEM images of
chitosan films after a month of
burial in red clay. The images
have been reproduced from
(Nakashima et al. 2005) with
kind permission from J-Stage

13
220 Page 14 of 22 3 Biotech (2023) 13:220

Fig. 7  a Digital images of whey protein isolate films before and
after soil composting burial test over 10 days. The images have been
Fig. 6  a Macroscopic appearance of thermoplastic starch (TPS) films
reproduced from (Li and Chen 2000) with permission from Springer
biodegradation in soil for 30 days. The figure has been reproduced
Nature. b Macroscopic digital images showing biodegradation of
from (Seligra et al. 2016) with kind permission from Elsevier. b
hydrophobic zein-based functional films over 10 days. The images
Macroscopic images of soil burial test of starch-based bioplastic over
have been reproduced from (Luís et al. 2019) with kind permission
10 days indicating respective mass loss%: (i) Day 0, (ii) Increasing
from MDPI
size of bioplastic on day 2, (iii) More soil water and the soil itself
have entered the bioplastic pores on day 4, (iv) Wider cracking area
on bioplastic sample on day 6, (v) Some parts of bioplastic were
destroyed on day 8, (vi) Wider parts of the bioplastic were destroyed is generally observed that animal-based protein bioplastics
on day 10. The figure has been reproduced from (Nissa et al. 2019) are easier to break down compared to plant-based protein
with kind permission from IOP science bioplastics since plant proteins lack branched chain amino
acids (BCAA).
lost about 50–60% of their weight, which is a strong indi-
cation of the biodegradation process carried out by the Polyesters of bacterial origin
microorganisms in the soil. The visual analysis confirmed
that, after 10 days in the soil, the films appeared thinner Microbes that produce extracellular PHA-degrading
and more fragmented compared to the initial samples (Luís enzymes are widespread in the soil and marine environment.
et al. 2019). Wheat gluten-based bioplastics were subjected PHAs are biodegraded by the action of bacteria present in the
to biodegradation in farmland soil. All gluten-based materi- soil. These bacteria belong to the following genera: Arthro-
als were entirely degraded within 50 days in farmland soil bacter, Corynebacterium, Actinomyces, Acinetobacter, Alca-
(Domenek et al. 2004). ligenes, Aspergillus, Bacillus, Burkholderia, Clostridium,
The study of biodegradation of bioplastics from bio- Comamonas, Cupriavidus, Mycobacterium, Pimelobacter,
mass such as proteins, polysaccharides (starch, cellulose, Enterobacter, Gracilibacillus, Planococcus, Nocardia, Pseu-
lignin chitin, and pectin), microorganisms, and bioderived doalteromonas, Staphylococcus, Micrococcus, Klebsiella,
as well as synthetic monomers receive less attention when Streptomyces, Pseudomonas, Stenotrophomonas, and Vari-
compared to physicochemical properties of these materi- ovorax (Trivedi et al. 2016). PHA-degrading fungi are
als. Further, the comparison of biodegradation of modified reported to be more efficient and belong to the divisions
composite polymers with one or more than one component Ascomycota, Basidiomycetes, Deuteromycetes, and Zygo-
from biomass has been studied in this review (Table 3). It mycotina. Biodegradation of PHA in soil is facilitated by

13
3 Biotech (2023) 13:220 Page 15 of 22 220

lipases and hydrolases (Table 2). The process involves Monomers

breaking down the polymer into oligomers by hydrolytic
depolymerase in the presence of water, converting PHAs Bio‑derived monomers
into trimer and dimer units, which are then treated by lipases
and hydrolases. (Meereboer et al. 2020). PHB depolymer- PLA has garnered significant interest in biomedical and
ases can be secreted by several microorganisms. Some PHB packaging applications due to its biodegradable and hydro-
depolymerases were isolated and purified from microorgan- phobic characteristics. The cleavage of ester bonds is the
ism species Alcaligenes, Comamonas, and Pseudomonas primary reason behind PLA degradation. Polymer degrada-
(Suzuki et al. 2021). The biodegradation of starch/PHB-V tion can be further caused by an array of factors, includ-
blends has been studied on several occasions. Accinelli et al. ing oxidation, photodegradation, thermolysis, hydrolysis,
developed a composite bioplastic synthesized from a combi- and biodegradation. The biodegradability of PLA in soil is
nation of corn starch and poly(3-hydroxybutyrate)-co-poly dependent on microorganisms (bacteria, fungus) and bio-
(3-hydroxy valerate) (PHB-V) in various proportions. The chemical processes of degradation (Weng et al. 2013). Dur-
biodegradability of the developed composite bioplastics in ing biodegradation, PLA is first hydrolyzed and then decom-
soil compost was assessed for 10 months. PHB-V blends posed by microorganisms into carbon dioxide and water.
with 50% of starch enhanced the biodegradability causing Primary PLA degrading enzymes include lipase, esterase,
it to degrade within 50% of the time required for PHB-V and alcalase (protease). PLA can be degraded in soil, due
bioplastic without any starch (Accinelli et al. 2012). Further, to the presence of many bacterial species such as Saccha-
three types of TPS synthesized from potato, corn, and water- rothrix, Kibdelosporangium, Pseudonocardia, Lentzea,
soluble potato starch, blended with PHB were evaluated in a and Amycolatopsi. In addition, studies reported that under
soil burial test. The mass loss increased with increasing time composting conditions in forest soil consisting of a mix-
and increasing glycerol content added during the synthesis ture of Actinomadura keratinilytica and Thermopolyspora
of bioplastic but decreased with increasing content of PHB sp, PLA film samples in soil were degraded completely in
(dos Santos et al. 2018b, a). The biodegradation of PHB three weeks (Zaaba and Jaafar 2020). Researchers found that
films in soil was studied including and excluding nitrate and the addition of starch augmented the biodegradation of the
at varying concentrations of oxygen. It was observed that the PLA-based bioplastics. SEM study revealed crack formation
PHB film completely degraded in two months under aero- after biodegradation caused due to the action of microbes
bic conditions in the presence of nitrate in the soil. (Fig. 8) on the surface of the polymers (Qi et al. 2017; Kalita et al.
(Bonartseva et al. 2002). 2020). In another study, the field emission scanning elec-
tron microscope (FESEM) images of PLA-chitosan com-
posite film after composting showed massive morphological
changes on the 40th and 80th days of composting process in
soil. Microbial assimilation during the biodegradation devel-
oped holes and cracks on the polymer surface, cross-sec-
tion, and core (Kalita et al. 2021) (Fig. 9). PLA bioplastics

Fig. 8  a Undegraded PHB film and PHB films with different degrees
of degradation after 2 months of incubation in experimental flasks b Fig. 9  FESEM images representing a PLA and b PLA-Chitosan com-
0% ­O2 and 0 g/L NO−3 c 10% O ­ 2 and 0 g/LNO−3 , and d 10% O­ 2 and posite films before and after biodegradation under composting condi-
5 g/L NO3 . The image has been reproduced from (Bonartseva et al.
−
tions over 100 and 80 days, respectively The images have been repro-
2002) with permission from Springer Nature duced from (Kalita et al. 2020, 2021) with permission from Elsevier

13
220 Page 16 of 22 3 Biotech (2023) 13:220

degrade extremely slowly, due to increased hydrophobicity

of the methyl group on the lactide monomer and prevent the
degrading enzymes from penetrating the polymer and break-
ing the existing ester bond (Farah et al. 2016).

Synthetic monomers

Aliphatic-co-aromatic polyesters are hard to be degraded by

enzymatic hydrolysis, but many enzymes that degrade PBAT
have been found among carboxylic ester hydrolases, includ-
ing carboxylesterase, arylesterase, triacylglycerol lipase, and
cutinases (Table 2) (Kawai et al. 2019). The rate of biodeg-
radation of PBAT can fluctuate from 5 to 95% subject to
Fig. 10  a SEM images of PBAT film surface before and after the soil
conditions of biodegradation used (composting, soil, enzy-
burial test: (i) 0 months (ii) after 1 month (iii) after 2 months (iv)
matic tests, etc.) (Feuilloley et al. 2005). PBAT films were after 3 months. This image has been reproduced from (Wang et al.
reported to biodegrade at unique rates in food, manure, and 2015) with kind permission from Elsevier. b FESEM images repre-
yard compost with different microbial activities. The maxi- senting the PCL film samples before and after biodegradation over
100 days under composting conditions. The image has been repro-
mum biodegradation rate was found in manure compost,
duced from (Kalita et al. 2020) with permission from Elsevier
resulting in the highest ­CO2 emission. SEM studies reveal
that undegraded PBAT film shows a smooth surface (Weng
et al. 2013). With increasing time of soil burial and the reinforcements for bioplastic polymers, affecting their rate
effects of microorganism activities, microbe-created cavities of biodegradation while also improving their functional
were observed on the surfaces (Wang et al. 2015). PCL sur- properties.
face morphology was rough with certain fibrillar structures
(Kalita et al. 2020). PCL is one of the few synthetic biode-
gradable polymers available and is used by microorganisms Conclusion
as an energy and carbon source (Chu and Wang 2013). The
enzymes, lipase, and cutinase belong to the esterases under Plastics are used in our daily lives as packaging materials,
the hydrolytic enzymes. Recently, a combination of these scientific gadgets, and in a variety of other applications.
two enzymes proved to be appropriate for the biodegradation However, traditional plastics are non-biodegradable, and
of PCL (Liu et al. 2019a). PCL is biodegraded by cutinolytic their rapid buildup in the soil ecosystem poses a hazard
enzymes secreting bacteria like Pseudozyma japonica-Y7-09 to the environment. Although plastics have significantly
(Abdel-Motaal et al. 2014). Further, a thermophilic Strep- improved our quality of life, it is crucial to shift toward sus-
tomyces thermoviolaceus subsp. thermoviolaceus isolate tainable alternatives, such as bio-based biodegradable plas-
76 T-2 isolated from soil in Taiwan is reported to cause tics. Biodegradable plastics are gaining popularity, and for
biodegradation of PCL (Chua et al. 2013). An early study the sake of the environment, plastics could be replaced by
reported that soil burial and compost cause chain scission biodegradable bioplastics derived from various sources. The
of the PCL backbone. This results in loss of mechanical biodegradation of bioplastics in the soil can be affected by
properties and significant weight loss of PCL bioplastics in numerous elements which include microbial diversity, tem-
a short time (Ardisson et al. 2014). Lipases from R. delemar, perature, humidity, and pH of the soil. The most common
R. arrizus, Candida cylindracea, and Achromobacter sp. and techniques in the literature to study the biodegradation of
esterase from hog liver showed degrading activities on PEA plastics are mass loss and visible evaluation via macroscopic
and PCL. Studies showed that PCL completely degraded in digital and SEM images. Biomass, particularly polysaccha-
12 days using Penicillium sp. strain 26–1 (ATCC 36507) rides like starch and cellulose, is widely preferred for the
isolated from soil. Figure 10 depicts FESEM images of bio- synthesis of bioplastics. However, the increasing demand
degradation of PCL film. for such bioplastics may create heavy competition for food
The most common approach in the literature is to examine sources. Starch and cellulose are commonly used polysac-
the total mass loss. Table 3 compares the percentage mass charides. Also, starch is more hygroscopic than cellulose,
loss of bioplastics after soil burial under different conditions. making starch-based bioplastics more susceptible to degra-
For a bioplastic to be considered truly biodegradable, it must dation. Additionally, discarded bioplastics in landfills release
degrade into carbon dioxide, water, biomass, and/or min- methane, a potent greenhouse gas, which could negatively
eral salts when exposed to air, moisture, soil, and microbes. impact the ecosystem. Bioplastics of microbial origin have
Bio-based materials can also be used as plasticizers and a significant constraint in the form of bulk manufacturing

13
3 Biotech (2023) 13:220 Page 17 of 22 220

and high recovery costs. Bioplastics may be implemented by Pseudozyma japonica sp. Nov., a novel cutinolytic ustilagino-
in the manufacturing of scientific devices and various other mycetous yeast species. 3 Biotech 4:507–512. https://​doi.​org/​10.​
1007/​s13205-​013-​0182-9
important applications including flexible packaging. Biodeg- Abe MM, Branciforti MC (2021) Biodegradation of hemicellulose-
radability is a characteristic regularly sought in terms of food cellulose-starch-based bioplastics and microbial polyesters.
packaging. Due to the extensive variety of bioplastics from Recycling. https://​doi.​org/​10.​3390/​recyc​ling6​010022
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gen/soy protein isolate for food packaging applications. RSC
Acknowledgements We thank the Department of Biotechnology Adv 6:82191–82204. https://​doi.​org/​10.​1039/​c6ra1​3043g
(DBT) and Department of Science and Technology (DST), Govern- Ai B, Zheng L, Li W et al (2021) Biodegradable cellulose film
ment of India, for the financial support (project number: BT/PR25099/ prepared from banana pseudo-stem using an ionic liquid for
NER/95/1014/2017 and DST/INT/Thai/P-10/2019). We thank the mango preservation. Front Plant Sci 12:1–10. https://​doi.​org/​
Manipal School of Life Sciences, Manipal Academy of Higher Edu- 10.​3389/​fpls.​2021.​625878
cation (MAHE), Manipal, Karnataka, India for providing the infra- Al-Itry R, Lamnawar K, Maazouz A (2012) Improvement of ther-
structure and facilities. Ms. Pooja N and Ms. Ishita Chakraborty mal stability, rheological and mechanical properties of PLA,
thank MAHE, Manipal, Karnataka, India for the Dr. T.M.A. Pai Ph.D. PBAT and their blends by reactive extrusion with functional-
fellowship. ized epoxy. Polymer degradation and stability. Elsevier, New
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Funding Open access funding provided by Manipal Academy of Alariya SS, Sethi S, Gupta S et al (2013) Amylase activity of a
Higher Education, Manipal. We thank the Department of Biotech- starch degrading bacteria isolated from soil. Arch Appl Sci
nology (DBT) and Department of Science and Technology (DST), Res 5:15–24
Government of India, for the financial support (Project number: BT/ Aliotta L, Vannozzi A, Cinelli P et al (2022) Wheat bran addition
PR25099/NER/95/1014/2017 and DST/INT/Thai/P-10/2019). as potential alternative to control the plasticizer migration into
PLA/PBSA blends. J Mater Sci 57:14511–14527. https://d​ oi.o​ rg/​
Data availability statement Data availability not applicable. 10.​1007/​s10853-​022-​07534-9
Aliotta L, Vannozzi A, Panariello L et al (2020) Sustainable micro
Declarations and nano additives for controlling the migration of a biobased
plasticizer from PLA-based flexible films. Polymers 12:1366.
Conflict of interest The authors declare no conflict of interest. https://​doi.​org/​10.​3390/​POLYM​12061​366
Altaee N, El-Hiti GA, Fahdil A et al (2016) Biodegradation of different
Open Access This article is licensed under a Creative Commons Attri- formulations of polyhydroxybutyrate films in soil. Springerplus.
bution 4.0 International License, which permits use, sharing, adapta- https://​doi.​org/​10.​1186/​s40064-​016-​2480-2
tion, distribution and reproduction in any medium or format, as long Andonegi M, Peñalba M, de la Caba K, Guerrero P (2020) ZnO nano-
as you give appropriate credit to the original author(s) and the source, particle-incorporated native collagen films with electro-conduc-
provide a link to the Creative Commons licence, and indicate if changes tive properties. Mater Sci Eng C 108:110394. https://​doi.​org/​10.​
were made. The images or other third party material in this article are 1016/j.​msec.​2019.​110394
included in the article's Creative Commons licence, unless indicated Anna AD, Arrigo R (2019) PLA/PHB blends: biocompatibilizer E ff
otherwise in a credit line to the material. If material is not included in ects. Polymers 11:1–12
the article's Creative Commons licence and your intended use is not Ardisson GB, Tosin M, Barbale M, Degli-Innocenti F (2014) Biodeg-
permitted by statutory regulation or exceeds the permitted use, you will radation of plastics in soil and effects on nitrification activity. A
need to obtain permission directly from the copyright holder. To view a laboratory approach. Front Microbiol 5:1–7. https://​doi.​org/​10.​
copy of this licence, visit http://​creat​iveco​mmons.​org/​licen​ses/​by/4.​0/. 3389/​fmicb.​2014.​00710
Asrofi M, Sapuan SM, Ilyas RA, Ramesh M (2020) Characteristic of
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