processes

Review
Waste to Energy: A Focus on the Impact of Substrate
Type in Biogas Production
Nwabunwanne Nwokolo 1, *, Patrick Mukumba 1 , KeChrist Obileke 1 and Matthew Enebe 2
1 Department of Physics, University of Fort Hare, P/Bag X1314, Alice 5700, South Africa;
[email protected] (P.M.); [email protected] (K.O.)
2 Department of Microbiology, North-West University, P/Bag X2046, Mmabatho 2735, South Africa;
[email protected]
* Correspondence: [email protected]; Tel.: +27-833433195




Received: 11 July 2020; Accepted: 3 September 2020; Published: 1 October 2020


Abstract: Anaerobic digestion is an efficient technology for a sustainable conversion of various

organic wastes such as animal manure, municipal solid waste, agricultural residues and industrial
waste into biogas. This technology offers a unique set of benefits, some of which include a good waste
management technique, enhancement in the ecology of rural areas, improvement in health through
a decrease of pathogens and optimization of the energy consumption of communities. The biogas
produced through anaerobic digestion varies in composition, but it consists mainly of carbon dioxide
methane together with a low quantity of trace gases. The variation in biogas composition are
dependent on some factors namely the substrate type being digested, pH, operating temperature,
organic loading rate, hydraulic retention time and digester design. However, the type of substrate
used is of greater interest due to the direct dependency of microorganism activities on the nutritional
composition of the substrate. Therefore, the aim of this review study is to provide a detailed analysis
of the various types of organic wastes that have been used as a substrate for the sustainable production
of biogas. Biogas formation from various substrates reported in the literature were investigated,
an analysis and characterization of these substrates provided the pro and cons associated with
each substrate. The findings obtained showed that the methane yield for all animal manure varied
from 157 to 500 mL/gVS with goat and pig manure superseding the other animal manure whereas
lignocellulose biomass varied from 160 to 212 mL/gVS. In addition, organic municipal solid waste
and industrial waste showed methane yield in the ranges of 143–516 mL/gVS and 25–429 mL/gVS
respectively. These variations in methane yield are primarily attributed to the nutritional composition
of the various substrates.

Keywords: biogas; organic waste; methane yield; microorganism; anaerobic digestion

1. Introduction
Energy is of great importance in facilitating the socioeconomic development of a country.
The increasing world population has consequently increased the demand for energy. However,
the combustion of fossil fuel resources to meet these energy needs leaves a negative footprint.
It contributes to global warming due to the emission of carbon dioxide (CO2 ), methane (CH4 ) and
nitrous oxide (N2 O) all classified as greenhouse gases (GHGs). In addition, the depletion of fossil
fuel resources due to its non-renewable nature and massive utilization has brought the need for
an alternative energy resource that are renewable, abundant and cost effective [1,2].
Biogas production from locally available renewable organic resources can be a good alternative
because it contributes to the reduction of GHG emissions. Biogas technology provides an attractive
route for the utilization of different categories of biomass for meeting energy needs [3]. This technology

Processes 2020, 8, 1224; doi:10.3390/pr8101224 www.mdpi.com/journal/processes

Processes 2020, 8, 1224 2 of 21

offer a unique set of benefits, some of which include good waste management technique, enhancement
in the ecology of rural areas, decrease in pathogenic diseases, optimization of the energy consumption
of rural communities and promotion in agricultural structure [4].
Biogas is a mixture of gases comprising mostly of methane and carbon dioxide as well as a low
quantity of other gases such as hydrogen sulphide (H2 S), ammonia (NH3 ), oxygen (O2 ), hydrogen (H2 ),
nitrogen (N2 ) and carbon monoxide (CO) [5,6]. This mixture of gases results from a biochemical process
known as anaerobic digestion (AD). Anaerobic digestion is a biochemical process that involves the
degradation of organic resources to simple substances to produce biogas and digestate through the
activities of a microorganism. Digestate is a biofertilizer, which when used in an agricultural integrated
system aids in closing the nutrient demand gap [7]. AD technology is employed in the treatment of
various organic wastes [1]. These organic wastes include animal manure, municipal solid wastes and
agroindustrial wastes. They serve as a substrate for the anaerobic digestion process.
In biogas production, various factors are of great importance, however the type of organic substrate
used has been found to play a significant role in the yield and composition of the biogas [3]. A number
of biogas digesters operating worldwide utilize different types of substrates and this results in a unique
microbial community and variation in methane composition in such digesters. Some studies have
used a sequencing technology to analyze the structure of these microorganism communities involved
in the AD process. These microorganisms consisting of bacteria, fungi and archaea are responsible for
all the reactions occurring within the digester system [8,9].
The variation in biogas compositions are dependent on the substrate types used, which is traceable
to the difference in their chemical composition as well as their biodegradability. However, there is a lack
of comprehensive studies in the literature, which evaluates the contributions of various substrates
used in biogas production and the limitations associated with each substrate. Therefore, the aim of this
review study is to provide a comprehensive analysis on the different types of organic wastes that has
been used as a substrate for the sustainable production of biogas via the anaerobic digestion process.
Biogas formation from various substrates reported in the literature will be investigated and their pros
and cons will be addressed to enable the synthesis of knowledge for proper means of maximizing
biogas yield from organic matters.

2. Anaerobic Digestion Metabolic Pathways

The biological decomposition of organic substrates during anaerobic digestion is divided into
four phases namely hydrolysis, acidogenesis, acetogenesis and methanogenesis. These biochemical
phases are directly linked in such a manner that the byproduct of one phase is the substrate of the
next phase. They occur simultaneously in an oxygen-free environment and results in the formation of
biogas and digestate as a byproduct [10]. These biochemical decomposition phases are associated with
a series of chemical reactions as shown in Equations (1)–(13) presented in Table 1.
Hydrolysis Reactions
C6 H10 O4 + 2H2 O → C6 H12 O6 + 2H2 (1)

Acidogenesis Reactions
C6 H12 O6 ↔ 2CH3 CH2 OH + 2CO2 (2)

C6 H12 O6 + 2H2 ↔ 2CH3 CH2 COOH + 2H2 O (3)

C6 H12 O6 → 3CH3 COOH (4)

C3 H7 O2 N + 2H2 O → C2 H4 O2 + NH3 + CO2 + 2H2 + ATP (5)

C4 H9 O3 N + H2 O → C3 H6 O2 + NH3 + CO2 + H2 + ATP (6)

4CH3 COCOO− + 4H2 O → 5CH3 COO− + 2HCO3 − + 3H+ (7)

Processes 2020, 8, 1224 3 of 21

Acetogenesis Reactions

CH3 CH2 COO− + 3H2 O ↔ CH3 COO− + H− + HCO3 − + 3H2 (8)

C6 H12 O6 + 2H2 O ↔ 2CH3 COOH + 2CO2 + 4H2 (9)

CH3 CH2 OH + 2H2 O ↔ CH3 COO− + 2H2 + H− (10)

Methanogenesis Reactions
CH3 COOH → CH4 + CO2 (11)

CO2 + 4H2 → CH4 + 2H2 O (12)

2CH3 CH2 OH + CO2 → CH4 + 2CH3 COOH (13)

Table 1. Biochemical reactions associated with anaerobic digestion phases.

AD Phase Associated Chemical Reactions Ref.

Hydrolysis C6 H10 O4 + 2H2 O → C6 H12 O6 + 2H2 [11]
C6 H12 O6 ↔ 2CH3 CH2 OH + 2CO2
C6 H12 O6 + 2H2 ↔ 2CH3 CH2 COOH + 2H2 O
Acidogenesis C6 H12 O6 → 3CH3 COOH [12–16]
C3 H7 O2 N + 2H2 O → C2 H4 O2 + NH3 + CO2 + 2H2 + ATP
C4 H9 O3 N + H2 O → C3 H6 O2 + NH3 + CO2 + H2 + ATP
4CH3 COCOO− + 4H2 O → 5CH3 COO− + 2HCO3 − + 3H+
CH3 CH2 COO− + 3H2 O ↔ CH3 COO− + H− + HCO3 − + 3H2
Acetogenesis C6 H12 O6 + 2H2 O ↔ 2CH3 COOH + 2CO2 + 4H2 [17]
CH3 CH2 OH + 2H2 O ↔ CH3 COO− + 2H2 + H−
CH3 COOH → CH4 + CO2
Methanogenesis CO2 + 4H2 → CH4 + 2H2 O [18]
2CH3 CH2 OH + CO2 → CH4 + 2CH3 COOH

At the hydrolysis phase as shown in Table 1 complex biopolymer compounds such as carbohydrate,
protein and lipids are degraded into water-soluble compounds. A typical example is shown in
Equation (1) in which cellulose is hydrolyzed in water to produce glucose and hydrogen. This phase
prepares the substrate for further degradation by making its products available for the fermentative
microorganisms in the next phase [19]. In the next phase, known as the fermentation stage, the acidogenic
bacteria converts glucose, amino acids and lipids into organic acids, volatile fatty acids (VFAs),
carbon dioxide (CO2 ) and hydrogen gas (H2 ) as shown in Equations (2)–(7). The most significant
organic acid produced at this phase is CH3 COOH, which serves as substrates for methanogenic
microorganisms [20]. At the acetogenesis stage, the VFAs particularly acetic acid and butyric acid
are converted to acetate, H2 and CO2 as shown in Equations (8)–(10). Equation (8) shows the
conversion of the phase product to acetate (CH3 COO− ) and H2 , which are utilized in the next stage [17].
In the final phase, known as the methanogenesis stage, CH3 COOH is converted to CH4 and CO2
as depicted by Equation (11). Furthermore, the produced CO2 reacts with H2 gas to produce more
CH4 , while CH3 CH2 OH undergoes decarboxylation to yield CH4 as shown in Equations (12) and (13)
respectively. Acetophilic methanogenic bacteria are responsible for the decarboxylation of acetate
into CH4 while the hydrogenophilic methanogenic bacteria group produces CH4 through a CO2 and
H2 reaction [21]. These products form the biogas that emerges as the final product of an anaerobic
digestion process.
Processes 2020, 8, 1224 4 of 21

Microbial Groups Involved in Anaerobic Digestion

The production of biogas from organic substrates (biomass material) is a complex microbiologically
dependent process. The organisms that catalyze the conversion of organic substrates into biogas and
other inorganic subunits operate under interdependent microbial activities, involving a consortium
of different microbes namely bacteria, archaea and fungi. These complex microbial community that
facilitate the conversion of organic polymers under anaerobic condition into biogas has until now
been understudied as most of them could not be cultured in the laboratory. Recently, the advent
of high throughput sequencing has revealed the different microbial groups involved in biogas
production [6,22–25].
A recent study showed that fungi, though the group least present within the microbial community,
is the most stable member of the community within the system. Bacteria and archaea are less stable
and fluctuates as the system’s chemical conditions changes during the biogas production processes.
Methanoculleus belonging to the Methanomicrobiales order was found to be abundant at the completion
phase of the methanogenesis compared to Methanosarcinales and Methanobacteriales. This shows that
the hydrogenotrophic pathway is the main methanogenesis channel within the biogas digester system.
These microbes account for 8–15 percent of the entire microbial community and perform the most
important role that results in the production of biogas. Methylotrophic methanogenesis are carried
out mostly by species of Methanosarcinales and Methanosphaera within the order Methanobacteriales.
Acetate is used as a growth substrate for methanogens such as Methanosarcina and Methanosaeta spp.
The former has been found to be a predominant group of methanogen in the digester due to its ability
to adapt and tolerate stressful conditions such as heat, salt, ammonia and organic matter load of the
digester better than the latter [26–28].
However, bacteria are equally an important agent in the production of biogas as their activities result
in the production of substances (acetic acids), which serve as a substrate for methanogens. These bacteria
can best be described as facultative anaerobes (Clostridium, Paenibacillus, Ruminococcus, Streptococci
and Thermoanaerobacteriaceae) as well as acetogenic bacteria (Acidaminococcus and Aminobacterium) and
sulphate reducing bacteria (Desulfovibrio). Other bacteria groups identified to participate in the process
are Proteobacteria, Chloroflexi, Verrucomicrobia, Actinobacteria, Acidobacteria, Spirochaetes, Plantomycetes,
Fibrobacteres, Tenericutes and Cloacimonetes. Some of these bacteria participate in the hydrolysis of
organic molecules, while Acidaminococcus, and Desulfovibrio are responsible for the production of
acetic acids from the substrates [29–32]. In addition, syntrophic acetogens, which are responsible
for the breakdown and oxidation of alcohols and organic acids to acetate, hydrogen and carbon
dioxide, are Syntrophorhabdus, Syntrophus, Syntrophobacter and Pelobacter. They are very abundant
in the biogas digester. They are cellulolytic microbes capable of cellulose breakdown to simple
sugars [33,34]. The type of substrate used and temperature of the biogas digester will determine which
group of microbes will dominate in the system. At higher temperature, Thermotogae dominates. At the
substrate level, municipal wastewater has more of Chloroflexi than animal waste or organic manure
and Firmicutes are the most dominant bacteria in the digester containing animal waste and/or organic
manure [25,35,36].
Although, bacteria and archaea are the predominant microbes in the digester, fungi are equally
key players in the biogas formation process. They are facultative anaerobes and are capable of
converting carbohydrate molecules via fermentation to acetate, carbon dioxide, formate, ethanol,
hydrogen and lactate. They belong to the phyla Mucoromycotina, Pucciniomycotina, Agaricomycotina,
Saccharomycotina, Pezizomycotina and Neocallimastigomycota. Studies have shown that fungi
operate best in synergy with methanogens in the breakdown of cellulose [37–39]. Therefore, to achieve
a maximum yield of biogas in the digester a symbiotic relationship between these microbes mentioned
above should be maintained and efforts should be made to monitor the activities of detrimental
organisms such as bacteriophages and protozoans that prey on the bacterial community, which could
affect the performance of the system–biogas digester.
Processes 2020, 8, 1224 5 of 21

3. Substrates for Biogas Production

The suitability of a biomass as a substrate for the production of biogas is majorly dependent
on its nutritional composition. These compositions influence the biogas yield, methane content,
biodegradability and degradation kinetics of the biomass involved. The major nutritional composition
of interest in substrates include carbohydrate, protein and fats [10]. Previous studies showed the
theoretical estimation of the possible methane yield and biogas percentage realizable from these
nutrients using the Buswell formula as shown in Table 2.

Table 2. Estimation of the maximum theoretical methane yield and biogas percentage composition.

Nutrient Methane Yield (m3 /kg VS) CH4 (%) CO2 (%) Reference
Carbohydrate 0.42 50 50 [24]
Protein 0.50 50 50 [24]
Lipid 1.01 70 30 [40]

As observed in Table 2, substrates rich in lipids such as fats hold a greater potential for methane
yield. Nevertheless, its degradation releases long-chain fatty acids that could be toxic to the microbial
community and causes a drop in pH. This could be minimized, by using a start-up strategy that
enhances the development of a special group of microorganism that is resistant to toxicity [41–43].
Protein rich substrates also have a high potential for methane yield. Degradation of such substrates
releases ammonium (NH4 + ) that could increase the alkalinity of the anaerobic digestion process.
This will consequently enhance the digestate value as a fertilizer as well as inhibit the activities of the
methanogens. This inhibition occurs because of a shift in equilibrium from ammonium to ammonia
(NH3 ) usually in a varying concentration ranging from 53 to 1450 mg/L [44]. Previous studies have
indicated that microorganisms can acclimatize in an environment of high NH3 concentration and still
produce biogas efficiently [45].
Besides protein and lipid-rich substrates, some other biomass with a high degree of lignocellulose
are used as a substrate for the production of biogas. Although this class of biomass is usually
difficult to degrade because of their heterogeneous structure, recalcitrant nature and low accessibility
by enzymes (carbohydrate polymers) [46–48]. Different pretreatment mechanisms are adopted in
breaking down the heterogeneous matrix, increasing the surface area and porosity of the lignocellulose
biomass for enhanced biogas production. For efficient production of biogas from any given substrate,
it is paramount to supply sufficient nutrients in the right proportion to meet the microorganisms’
nutritional requirements and sustain an optimal growth of these microbial communities. However,
these requirements are usually difficult to meet with one type of substrate explaining the reason for
exploration of various types of substrates in a monodigestion process as well as codigestion of two or
more substrates. Substrates for the production of biogas can be broadly divided into three groups,
namely agricultural waste, municipal organic waste and industrial waste.

3.1. Agricultural Waste

Agricultural waste emanates from different agricultural activities. They include animal waste,
lignocellulosic biomass usually in the form of crop residue, forest residue and energy crop.

3.1.1. Animal Waste

Animal waste is a renewable and cost-effective substrate for the production of biogas through
anaerobic digestion. This organic waste if not properly handled and disposed poses a threat to the
environment because of its high concentration of nitrogen and phosphorus. In addition, the presence
of pathogens, antibiotics, heavy metals and microorganisms in animal waste can contaminate water
bodies, air and the soil [1]. However, the transformation of these animal wastes using anaerobic
digestion technology could offer the dual benefits of producing sustainable biogas energy and good
Processes 2020, 8, 1224 6 of 21

waste management technique. Animal wastes identified as a viable substrate for biogas production are
manure derived from cattle, pig, sheep, goat and poultry. This is due to its high nutrient content, high
organic matter concentration and high buffering capacity. Typically, the amount of animal waste that
can be derived from a given farm varies based on type, size and age of the animal, feed and feeding
mechanism and type of breeding [49].
Generally, manure from different animals show diverse characteristics, which can be a consequence
of their management system, diet, digestive system and animal type. Apart from feces and urine
that make up the major part of animal slurry, materials such as sand, straw from bedding material,
water from cleaning and small branches are also found in it. All these constitute variation in the
characteristics of animal waste. The volatile solid (VS), total solid (TS), pH and carbon to nitrogen ratio
are important quantitative characteristic parameters for determining the methane production potential
of animal waste. Table 3 presents the physiochemical characteristics of various animal manure and
their corresponding methane yield.

Table 3. Physiochemical characteristics and methane yield of various animal manure.

Animal Manure pH TS (%) VS (%) C/N Ratio CH4 Yield (mL/gVS) Reference
Cattle Manure 7.1–8.6 14.5–22.7 11.9–72.0 14.59–18.9 157.0–395.0 [50–54]
Pig Manure 6.4–7.5 8.2–36.7 6.2–82.8 5.7–13.5 204–438.4 [53,55–60]
Chicken Manure 6.9–7.4 20.0–92.6 18.3–84.1 7.5–9.75 160.0–396.0 [53,61–65]
Sheep Manure 7.16–8.1 22.3–40.0 18.7–72.7 11.3–14.7 207.0–357.0 [50,66,67]
Goat Manure 7.9 33.7–55.5 27.7–89.4 18.0 402–500 [66–68]
Donkey Manure 6.8 19.8 14.4 - 380 [69]

It was observed from the literature that the methane yield of various animal manure varies
significantly worldwide as evident in Table 3. This can be attributed to a range of factors such
as a difference in the origin of the animal manure, animal diet, variation in animal digestion,
management system, manure storage mechanism prior to the anaerobic digestion process and intestinal
microorganism [70]. The methane yield from pig manure (204.0–438.4 mL/gVS) and goat manure
(402–500 mL/gVS) were found to supersede that of other manure. The high methane yield in pig
manure can be related to its high buffering capacity that protects the digestion process against problems
such as acidification associated with high TS. The accumulation of volatile fatty acids usually causes
instability and AD system failure particularly at a high organic loading rate [57]. However, with good
buffering capacity of the substrate an adequate environment is provided for optimum functioning
of microorganisms. The TS and VS concentration of the animal manure has its contribution towards
the methane yield. As observed from Table 3, chicken manure gave the highest TS content of 92.6%
obtained in Bojti et al.’s [62] study. This resulted in a lower methane yield of approximately 160 mL/gVS,
which is comparable to the lowest methane yield of cattle manure. With a higher percentage of TS
in a substrate, the amount of water decreases and this reduces the activities of the microorganism.
This is because of the main contribution of water in the growth of microorganisms that facilitates
the dissolution and transport of nutrients [71]. Cattle manure with the reported TS content that
ranged from 14.5 to 22.7% and VS content that ranged from 11.9 to 72.0% gave the lowest methane
yield. This could be traceable to a higher lignin content in cattle manure originating from their feed
that contains a structural carbohydrate (cell wall components) as compared to other farm animal
manure [72]. In a recent review study, it was shown that the highest lignin content was recorded
in cattle manure (11.5%) as compared to pig manure (8.5%) and poultry manure (4.2%). This lignin
content inhibited methane production in cow manure as evident in the average methane yield of
168.0 mL/gVS reported compared to 215.0 mL/gVS and 255.0 mL/gVS reported for pig and poultry
manure respectively [73]. Li et al.’s [2020] study further showed a higher lignin content (14.0%) in
cattle manure as compared to sheep manure (8.6%), which resulted in a theoretical methane yield
(357 mL/gVS) of cattle being lower than that of sheep (395.0 mL/gVS). The pH for all the animal manure
as gathered from the literature presented in Table 3 ranged from 6.4 to 8.6, where cattle manure and
Processes 2020, 8, 1224 7 of 21

sheep manure gave the highest pH value of 8.6 and 8.1 as recorded in a recent study [50]. The reported
pH range efficiently supports the methanogenesis process and enhances the activities of hydrolytic
enzymes [74].

3.1.2. Lignocellulosic Biomass

The abundant production of lignocellulosic biomass, which approximates to 200 billon tons per
year, made it a viable resource for sustainable energy production. Anaerobic digestion identified as
a less energy intensive process is a sustainable route for the conversion of a variety of lignocellulosic
biomass into biogas [75]. Lignocellulosic biomass is comprised of three major components namely
cellulose, hemicellulose and lignin that primarily influences its biodegradation. Cellulose, being the
most abundant and main component of all plant cell walls, usually constitutes about 35–50% of
the entire composition. It is a linear polymer formed by D-glucose subunits linked through β-(1-4)
covalent bonds [76]. The β configuration gives cellulose a structure, which could be in the crystalline
or amorphous form. Its biodegradation particularly crystalline cellulose involves a number of
enzymes namely endo-cellulases, exo-cellulases and β-glucosidases. These enzymes are responsible
for splitting of β-1, 4 glucosidic bonds, degradation of polymers and cellobiose to glucose. In addition,
an extracellular multi-enzyme called cellulosome performs the complete degradation by binding to the
substrate [77].
Hemicellulose, the second most dominant fraction of the lignocellulosic biomass, accounts for
about 20–35% of the total composition. It consists of a variety of polysaccharides including xylan,
glucomannan, glucuronoxylan, xyloglucan and arabinoxylan depending on the plant tissue and
species [78]. Hemicellulose degradation are majorly attributed to a key enzyme called xylanases that
breaks β-1,4 backbone of the xylan polymers. This key enzyme exists in a variety of forms depending
on substrate specificities, action mechanisms and hydrolytic activity. The third component known as
lignin, encapsulates hemicellulose and cellulose to produce a hydrophobic 3-dimensional structure
called “Lignin-Carbohydrate Complexes” (LCC). Lignin recalcitrant nature limits its degradation in
an anaerobic environment [75]. However, various pretreatment mechanisms are usually adopted
to break the lignin and polysaccharides linkage, thus, making the hemicellulose and cellulose more
accessible to microorganisms and hydrolytic enzymes. Some lignocellulosic biomass with their
respective hemicellulose, cellulose and lignin composition are shown in Table 4.

Table 4. Composition of some lignocellulosic biomass for anaerobic digestion.

% of Dry Weight
Biomass Type Reference
Cellulose Hemicellulose Lignin
Eucalyptus 38.0–45.0 12.0–13.0 25.0–37.0 [79]
Switch grass 43.1 31.7 11.3 [80]
Nut shell 25.0–30.0 25.0–30.0 30.0–40.0 [81]
Grasses 25.0–40.0 35.0–50.0 10.0–30.0 [81]
Corn Stover 33.7 19.1 15.2 [82]
Bagasse 38.2 27.1 20.2 [79]
Rice straw 37.8 29.6 14.8 [54]
Cotton stalk 50.4 15.6 16.3 [83]
Wheat straw 48.6 29.4 7.3 [84]
Corn cob 45.0 35.0 1.05 [81]
Rice Husk 41.4 18.0 20.4 [80]
Pineapple leaves 30.0 37.0 22.0 [85]
Pineapple stem 37.0 34.0 20.0 [85]
Pineapple root 42.0 32.0 19.0 [85]

From Table 4, it can be observed that cellulose dominated in the composition for all lignocellulosic
biomass particularly in the cotton stalk, wheat straw and corncob, with lignin accounting for the
Processes 2020, 8, 1224 8 of 21

lowest composition with an exception of Eucalyptus and nutshell. The high lignin content in Eucalyptus
is attributed to it been a woody biomass that derives its high thermal stability from high lignin
content. It is evident from Table 4 that the total composition of cellulose, hemicellulose and lignin in all
biomass did not amount to 100%. The remaining percentage is a summation of organic, and inorganic
compounds such as lipids, protein and other extractives.
Extractives are classified as plant cell wall chemicals constituting of fatty alcohols, fatty acids,
phenols, resin acid, terpenes, rosin, steroids and waxes. These extractives increases the biodegradability
of lignocellulose and are responsible for its durability, color as well as smell [85,86]. It is important
to highlight that cellulose, hemicellulose and lignin composition does not only vary with biomass
species as observed in Table 4, but also varies based on biomass maturity and growth conditions.
These variations in composition influence the biogas production potentials of lignocellulosic biomass.
A recent study focused on AD performance of cellulose, hemicellulose, lignin and their mixtures to
understand their individual contribution towards biomethane production. Table 5 presents the AD
performance of these components as reported.

Table 5. Anaerobic digestion performance of lignocellulose compositions. Adapted from [75].

Substrates TMMY (mL/gVS) EMY (mL/gVS) BD (%)

MC 407.8 211.8 51.9
XY 382.5 133.9 35.0
MC + XY 394.6 185.1 46.9
MC + XY + AL 414.8 178.1 42.9
MC + XY + SL 450.2 160.1 35.6
MC: microcrystalline cellulose, XY: xylan (hemicellulose), AL: alkali lignin and SL: sodium lignosulfonate.

In Table 5, the theoretical maximum methane yield (TMMY), experimental methane yield (EMY)
and biodegradability (BD) of cellulose, hemicellulose, lignin and their combination are presented.
TMMY and BD were determined using Equations (14)–(16) along with the elemental compositions of
cellulose, hemicellulose and lignin.
     
b c 3d a b c 3d a b
C a H b O c Nd + a − 4 − 2 + 4 H2 O → 2 + 8 − 4 − 8 CH4 + 2 − 8 + 4c + 3d
8 CO2 + dNH3 (14)

22. 4 ×1000 × (a/2 + b/8 − c/4−3d/8)

TMMY = (15)
12a + b + 16c + 14d
EMY
 
BD = × 100 (16)
TMMY
where Ca Hb Oc Nd is the chemical formula for the substrate derived experimentally. As observed
from Table 5, hemicellulose gave the lowest experimental methane yield when fermented separately.
This can be attributed to acidification of hemicellulose that inhibits the activities of methanogens
responsible for methane production. As hemicellulose was codigested with microcrystalline cellulose,
its biodegradability was enhanced from 35.0 to 46.9%. This is because of an increase in the nutritional
balance brought about by the addition of cellulose. Contrarily, the addition of alkali lignin (AL)
and sodium lignosulfonate (SL) to the fermentation system of hemicellulose and cellulose caused
a decrease in EMY. This in turn resulted in a decrease in biodegradability from 46.9 to 42.9% and
35.6% for AL and SL respectively. This decrease is traceable to the inhibition effect of lignin and the
presence of sulphur. Similarly, in another study, lignin showed the lowest biodegradability of 0.1%
as compared to cellulose (57.8 and 59.9%) and hemicellulose (43.6–55.3%). The methane production
characteristics of these lignocellulosic components were equally investigated in the study. Lignin gave
the least EMY of 0.7 mL/gVS followed by hemicellulose with EMY of 178.6–223.5 mL/gVS and cellulose
giving the highest EMY of 241.7 mL/gVS and 251.4 mL/gVS [61]. Comparably, Xu et al.’s [87] study
evaluated the methane potential of different parts of corn stover namely stem bark, stem pith and
Processes 2020, 8, 1224 9 of 21

leaves with varying compositions of cellulose, hemicellulose and lignin. The cumulative methane
yields obtained for corn stover stem bark, stem pith and leaves were 0.194 L/gVS, 0.210L/gVS and
0.198 L/gVS respectively. The lower methane yield of stem bark is attributed to its lignin content
(17.61%), which is higher compared to the lignin content of 7.16% and 5.16% for stem pith and leaves
respectively. The higher lignin content of stem bark created a stronger binding for its cellulose and
hemicellulose content thus resulting in a lower degradation rate for both compositions. As observed
from the literature, lignin is confirmed to contribute the most in limiting the biodegradability of
lignocellulosic biomass. Previous studies showed that an increase in lignin content by 1% would
cause a reduction of 7.49 L CH4 /kg total solid on average [88,89]. In addition, a lignin content of
any lignocellulose biomass in excess of >100 g/kg VS will result in a significantly lower yield of
methane [90]. Li et al.’s [91] study further indicated that lignin content of 15% could be a critical point
in AD of lignocellulosic biomass.
Comparing only the cellulose and hemicellulose degradation rate, Zhao et al.’s [92] study showed
a significant difference in the degradation of the hemicellulose and cellulose composition of oat straw
when digested as a monosubstrate. Hemicellulose degradation (68.85–81.44%) was observed to be
greater than cellulose degradation (28.06–33.61%) as the total solid of the oat straw varied from 2 to
10% in steps of two. Additionally, in Song and Zhang’s [84] study, hemicellulose was reported to have
decomposed more compared to cellulose and lignin. According to the study, their decomposition rates
were recorded to be 12.5–45.2% for hemicellulose, 9.3–30.2% for cellulose and 5.4–21.9% for lignin.
Recalling that hemicellulose hydrolyzes more easily compared to cellulose and has a lower degree of
polymerization than cellulose; hemicellulose can be expected to degrade faster. Although, some studies
are of the view that cellulose has a higher biodegradability and yields more methane than hemicellulose,
some other studies have a different view where the reverse is the case. Both cellulose and hemicellulose
contain D-glucose molecules but hemicellulose is easily hydrolyzable than cellulose, which forms
an association with other plant substances that limit its biodegradability. In addition, the ease of
hemicellulose biodegradability could be attributed to its composition of short side chains consisting of
a variety of sugars and its inability to form an aggregate when crystallized with cellulose. However,
due to cellulose, dominating in the percentage composition (35–50%) compared to hemicellulose
(20–35%), [93,94] cellulose methane yield per unit biomass under complete degradation will be higher
compared to hemicellulose. This conclusion might not entirely hold for all biomass as the high rate of
biodegradation of hemicellulose could result in higher yield of methane than cellulose. Hence, further
studies are required to ascertain between cellulose and hemicellulose the one that yields more methane.

3.2. Codigestion of Lignocellulose and Animal Manure Studies

Anaerobic microorganisms differ in their requirements of organic and micronutrients needed for
their growth and degradation of substrates for biogas production. To satisfy these nutritional
requirements of microorganisms, a combination of substrates are codigested to improve their
characteristics. For instance, the low and imbalanced carbon to nitrogen ratio in animal manure can be
compensated through codigestion with carbon-rich substrates. As it has been noted that the protein,
fats and carbohydrate content of a given substrate influence the percentage of methane in biogas.
Substrates that are rich in these compositions can be combined with other substrates to achieve process
stability and subsequently improve biogas production. This section presents an overview on anaerobic
codigestion of a variety of substrates particularly lignocellulose and animal manure. Table 6 presents
a summarized overview of some studies on codigestion of lignocellulosic biomass and animal manure.
Processes 2020, 8, 1224 10 of 21

Table 6. A summary on some studies of codigestion of lignocellulose and animal manure.

Substrate Type Mixing Ratio pH Pretreatment CH4 Yield (mL/gVS) Ref.

Wheat Straw + Cattle manure 40:60 6.5–7.0 3% w/w H2 O2 320.8 [84]
Wheat Straw + Cattle manure 30:70 6.8–7.1 None 254.6 [84]
ASW + Cow dung 60:40 8.1 None 297.7 [95]
ASW + Cow dung 60:40 8.1 1 g NaHCO3 /g of VS 386.3 [95]
Corn stover + Chicken manure 3:1 6.9–8.2 Wet-AD 218.8 [91]
Corn stover + Chicken manure 3:1 7.4–8.3 HSS-AD 208.2 [91]
Corn stover + Chicken manure 1:1 8.0–9.3 SS-AD 147.8 [91]
Goat manure + Wheat straw 30:70 6.5–7.5 None 12.8 [67]
Goat manure + Corn Stalk 70:30 6.5–7.5 None 16.0 [67]
Goat manure + Rice Straw 50:50 6.5–6.8 None 15.7 [67]
Sugarcane Bagasse + Cow dung 1:2 6.8 2% w/w NaOH 386 [96]
Sugarcane Bagasse + Cow dung 1:2 6.8 2% w/w Ca(OH)2 334 [96]
Sugarcane Bagasse + Cow dung 1:2 6.8 None 322 [96]
Animal manure+ Grape by-products 1:2 4.3–8.8 None 348 [97]
Animal manure+ Tomato pulp 1:2 3.6–8.8 None 404 [97]
Animal manure + Olive by-product 1:2 4.6–8.8 None 398 [97]

A summary of the discussed studies on codigestion of lignocellulosic biomass and animal manure
presented below are provided in Table 6. All studies presented in Table 6 were conducted under the
mesophilic temperature (35–38 ◦ C) condition but at varying retention times.
Wheat straw, a typical example of an agricultural waste presented in Table 6, is a suitable substrate
for the production of biogas, although its lignocellulose content slows down the degradation process.
Song and Zhang’s [84] study presented in Table 6 explored the monodigestion and codigestion
of pretreated wheat straw with cattle manure. In the study, wheat straw was pretreated with
four concentrations of H2 O2 (1%, 2%, 3% and 4%) prior to its digestion as a monosubstrate and
cosubstrate in varying ratios of dairy cattle manure. The methane yield for monodigestion of
H2 O2 -treated wheat straw were 94.8, 108.5, 128.4 and 118.7 mL/gVS for the 1%, 2%, 3% and 4%
pretreatment and 84.3 mL/gVS for untreated wheat straw. A significant improvement in the methane
yield was observed as H2 O2 -treated wheat straw and cattle manure were codigested. The mixing
ratio of 40:60 for H2 O2 -treated wheat straw and cattle manure gave the highest methane yield of
320.8 mL/gVS while codigestion of untreated wheat straw/cattle manure gave an optimum methane
yield of 257.6 mL/gVS at a 30:70 mixing ratio. Additionally, with codigestion the methanogenic
community moved from acetoclastic methanogens to hydrogenotrophic methanogens.
Almomani and Bhosale’s [95] study proposed a means of optimizing the biogas yield of some
agricultural solid wastes (clover, grass and wheat straw) by addition of cow dung. A maximum
cumulative methane production (CMP) of 297.99 NL/kgVS was recorded at a mixing ratio of 60:40 for
the agricultural solid wastes and cow dung as shown in Table 6. The CMP of the codigested substrates
were further enhanced through chemical pretreatment by addition of different doses (D1 = 0.25, D2 = 0.5,
D3 = 0.75, D4 = 1.0, D5 = 1.25 and D6 = 1.5 g) of NaHCO3 . A maximum CMP of 386.3 NL/kgVS was
obtained with 1.0 g of NaHCO3 /gVS (D4) at a mixing ratio of 60:40. Another study investigated the
contribution of the organic loading rate in codigestion of rice straw and cow manure using a continuous
feeding mechanism. An optimal mixing ratio of 1:1 for the volatile solids was obtained through
a batch test analysis prior to the continuous experiment. An organic loading rate of 6 g/Ld resulted in
an efficient and stable codigestion with an average biogas production and daily volumetric biogas
production rate of 383.5 L/kgVS and 2.3 m3 /day respectively. A further increase in organic loading led
to accumulation of VFA, which in turn caused a severe inhibition to the codigestion process [32].
Corn stover is a potential substrate for biogas production that usually results as a leftover from
maize harvest. In Li et al.’s [91] study shown in Table 6, corn stover was codigested with chicken
manure under three conditions namely: the hemi-solid state (HSS-AD), wet (W-AD) and solid-state
(SS-AD) anaerobic digestion. The study was focused on determining the best mixing ratio for the
optimum methane yield and achieving process stability under the three anaerobic digestion conditions.
An optimum methane yield of 218.8 mL/gVS and 208.2 mL/gVS occurred at a substrate-mixing ratio of
3:1 for corn stover and chicken manure under wet and hemi-solid state anaerobic digestion conditions
Processes 2020, 8, 1224 11 of 21

respectively. In addition, a mixing ratio of 1:1 gave the maximum volumetric methane productivity
of 14.2 Lmethane /Lreactor volume . Moreover, a synergistic effect was observed when the substrates were
mixed in the ratio of 3:1 and 1:1 under the solid-state condition. Zhang et al. [67] investigated the
codigestion of goat manure with three crop residues namely corn stalk, rice straw and wheat straw
under mesophilic conditions and at varying mixing ratios for efficient biogas production. An optimum
biogas yield presented in Table 6 were obtained at mixing ratios of 30:70, 70:30 and 50:50 for goat
manure/wheat straw, goat manure/corn stalk and goat manure/rice stalk respectively. The total biogas
yield of 12.8 L/kgVS from the codigestion of goat manure and wheat straw at a mixing ratio of 30:70
were 62.1% and 23.0% higher, compared to their monodigestion. On the other hand, at a mixing ratio of
50:50, the codigestion of goat manure/rice straw gave a total biogas yield of 15.7 L/kgVS that is 111.28%
and 51.31% higher than digesting rice straw and goat manure separately. Whereas, the total biogas
yield of 16.0 L/kgVS for goat manure/corn stalk when codigested is 83.02% and 54.44% higher than that
of corn stalk and goat manure alone. A significant improvement in biogas production was achieved
through codigestion by overcoming the carbon to nitrogen imbalance associated with single substrates.
Sugarcane bagasse is another agricultural waste that can serve as a substrate for codigestion
purposes due to its energy potential. It is a byproduct of the sugar milling industry that results from
milling of the sugarcane crop. A recent study investigated the anaerobic codigestion of pretreated
sugarcane bagasse with cow dung. The bagasse was pretreated with a solution of NaOH and Ca(OH)2
for one day before mixing it with cow dung in the ratio of 1:2. As presented in Table 6, NaOH treated
bagasse gave a maximum biogas yield of 386 mL/gVS, Ca(OH)2 treated bagasse gave about 334 mL/gVS
and pure bagasse (untreated) yielded about 322 mL/gVS of biogas at 35 ◦ C. With the addition of cow
dung to the pure bagasse and an increase in temperature from 35 to 55 ◦ C, an increase in biogas yield
of 27 mL/gVS was observed. This could be attributed to the shift in the carbon/nitrogen ratio (130:1)
of pure sugarcane bagasse to 29:1 when cow dung was added [96]. The higher biogas yield in the
codigestion of pretreated sugarcane bagasse with cow dung as compared to pure sugarcane bagasse
is due to increase in the internal surface area and decrease in the degree of polymerization of the
lignocellulose material. This in turn breaks the linkage between carbohydrate and lignin due to applied
alkaline pretreatment.
A study developed an assay for codigestion of animal manure (a mixture of 45% VS of calf
manure, 41% VS of lamb manure and 2% VS of pig manure) with agro-food byproduct silages (grape
byproduct, tomato pulp and olive agro-food byproduct). The study showed that a more synergetic
effect existed in the codigestion of animal manure and tomato pulp due to a higher methane yield
of 404 mL/gVS attained, compared to the grape byproduct and olive agro-food byproduct shown in
Table 6. The highest methane yield was recorded when the animal manure was higher in proportion
compared to the agro-food byproduct. This could be attributed to an elevation in the chemical oxygen
demand of the assay. Additionally, a correlation was observed with alkaline parameters and ammonia
nitrogen at a higher ratio of animal waste [97]. Mukumba et al.’s [98] study mathematically modeled
the performance potential of a biogas digester fed with selected types of a substrate. An optimal
methane percentage composition of 75% was reported with an equal mixture of cow dung, goat dung,
donkey dung and horse dung. It has been shown from the reviewed studies that codigestion of animal
manure with agricultural residues leads to the optimization of biogas production and stabilization
of the anaerobic digestion process. This is attributed to more balanced nutrients achieved through
codigestion particularly between carbon and nitrogen.

3.3. Municipal Solid Waste

Municipal solid waste (MSW) consists of solid wastes generated within a municipality by
households, industries and a commercial setting. Its composition and amount are variable with
organic matter constituting about 25–75% of the total MSW. The MSW production rate typically varies
between 1.1 and 2.2 kg/person/day based on a country’s income [10]. On a global scale, approximately
1.3 tonnes of MSW are generated annually. These MSW compositions consist of 46% of organic waste,
Processes 2020, 8, 1224 12 of 21

17% paper waste, 10% plastic waste, 5% glass waste, 4% metal waste, 3% textile waste, 13% inert and
2% miscellaneous waste [99]. The organic waste accounts for the highest composition among other
waste types. Their percentage composition also varies from continent to continent with Africa having
the highest composition of 66% and Australia having the least of about 25% as present in Table 7.

Table 7. Organic matter composition of municipal solid waste (MSW) by continent. Adapted from [100].

Continent Organic Percentage Composition of MSW (%)

Africa 66
Asia 47
Europe 54
North America 26
South America 53
Australia 25

For low-income countries, the organic portion of the waste ranges from 50 to 70% while that of
high-income countries ranges from 20 to 40%. The high rate of the organic fraction of MSW generated
in Africa is traceable to urbanization due to rural–urban migration, rapid population growth and
industrialization [101]. Effective conversion of this waste through innovative ways can be a sustainable
waste management technique to adopt. Two major conversion pathways exist for MSW and they are the
biochemical conversion process (anaerobic digestion) and thermochemical conversion (incineration).
The thermochemical conversion mechanism is rarely used because of the low calorific value and high
moisture content of MSW. Anaerobic digestion of MSW has attracted more attention because of the
possibility of separating the organic biodegradable fraction from the total MSW [102]. Additionally,
the generation of renewable energy, reduction of landfilling and mitigation of pollution are some
other factors that steer the public interest in AD of MSW. Food waste makes up a greater proportion
of the organic waste stream and has the highest biogas potential compared to other biodegradable
stream. This is because of the high concentration of carbohydrates, proteins, fats and the absence of
heavy metals. Table 8 presents the characteristics of different organic fractions of MSW as reported in
the literature.

Table 8. Physicochemical characteristics of organic municipal solid waste.

Substrate TS (%) VS (%) C/N Ratio pH Reference

Food waste 10.8 10.2 15.2 4.2 [103]
Kitchen waste 19.1 17.8 14.4 4.5 [104]
OFMSW 14.1 10.33 32.5 - [105]
Fruit and Veg waste 13.8 12.88 - 4.5 [106]
Catering food waste 23.0 18.5 20 4.9 [107]
SOW 12.8 11.3 13.6 4.3 [108]
Brown grease 43.8 37.2 - 6.1 [109]
OFMSW—organic fraction of municipal solid waste and SOW—synthetic organic waste.

The synthetic organic waste (SOW) reported in Table 8 represents a typical composition of organic
waste disposed in a landfill. It is comprised of leftover food such as meat, rice and beans accounting
for about 79%, fruit and vegetable waste such as orange, banana and apple accounting for about
20% and 1% cardboard. As shown in Table 8 the TS and VS content of the organic MSW are in the
ranges of 10.8–23.0 and 10.2–18.5 respectively, indicating that water accounts for about 80% of the
total composition. This is acceptable, as moisture content is recommended in the literature to be as
high as 90% to support a high methanogenesis rate [110]. The pH of the substrates as seen are low,
thus a proper control is needed during AD of these substrates for optimal performance. The carbon
to nitrogen ratio of organic municipal waste as observed ranged from 13.6 to 32.5, of which some
wastes are within the optimum recommended ratio of 20–30:1 [111]. However, for substrates with
Processes 2020, 8, 1224 13 of 21

a low C/N ratio, codigestion is suggested as a good approach to attain the optimal ratio. Generally,
the microorganisms responsible for the AD process utilize carbon more compared to nitrogen. Hence,
nitrogen concentration should not exceed the quantity needed by the microbial community to avoid
ammonia inhibition [112]. Table 9 presents the methane potentials of the organic fraction of MSW
reported in the literature.

Table 9. Organic municipal solid waste and their methane yield.

Substrate Type Methane Yield (mL/gVS) Reference

Food Waste 460.0 [103]
OFMSW 220.5 [105]
Fruit and vegetable waste 516.0 [106]
Kitchen Waste 501.0 [113]
Vegetable food waste 425.0 [114]
Catering food waste 287.0 [107]
Cucumber waste 143.0 [115]
Onion skin 400.0 [116]
Potato skin 267.0 [116]
Banana skin 277.0 [116]
Carrot petioles 309.0 [116]
Orange peel 230–332 [117]
Strawberry Extrudate 285–339 [118]
Fluted pumpkin peel 161–164 [119]
Brown Grease 400–490 [109]

The methane yield of all organic wastes presented in Table 9 fall in the range of 143–516 mL/gVS.
Food waste, kitchen waste, fruit and vegetable waste showed the highest methane yield. This could
be because of high lipid content of food waste and kitchen waste, which could be directly linked
to the presence of animal fat and oil in the waste stream. Studies have shown the lipid content of
food waste and the kitchen to be about 33.22% and 21.6% respectively [120,121]. As noted previously,
substrates rich in lipids have a greater potential for higher methane yield compared to protein and
carbohydrates. Although, high lipid content usually results in the formation of long-chain fatty
acids that could cause system failure [111]. Additionally, the high moisture content in fruit and
vegetables may have contributed to higher degradability of these substrates that led to high methane
yield. Cucumber waste showed the lowest methane yield, which could be attributed to its lignin
content as phytonutrients. Besides the nutritional composition of the substrates, other factors such as
temperature, pH, C/N ratio, organic loading rate and hydraulic retention time influence their methane
yield. However, temperature, organic loading rate and hydraulic retention time were not considered
in the present review. In summary the organic fraction of MSW substrates shows some variation in
characteristics as well as methane yield. Geographical change, seasonal variations as well as the type
of collection determine the final characteristics of the waste stream.

3.4. Industrial Waste

Industrial wastes are byproducts, residues and wastes that result from various industrial activities.
They include waste from the pulp and paper industry, food industry, petrochemical refinery waste, textile
industry and liquid biofuel production waste. Besides waste from the food industry, other industrial
wastes have not been widely used as a substrate in anaerobic digestion due to their recalcitrant chemical
properties and low biodegradability of about 30–50% [122]. In the pulp and paper industry, a large
quantity of wastewater with a high organic load is produced during the paper manufacturing process.
These wastewaters typically have high chemical oxygen demand (COD) in the range of 800–4400 m/g,
high biological oxygen demand (BOD) of about 300–2800 m/g and high dye content in the range of
1200–6500 color units [109]. Anaerobic treatment of this wastewater has an added benefit of lower
treatment cost because of the possibility of utilizing the biogas produced for energy generation. Due to
Processes 2020, 8, 1224 14 of 21

low solid content (TS < 1%) of this wastewater, they are codigested with other substrates or pretreated
to improve their biodegradability. The textile industry is another industry that generates a significant
quantity of wastewater through the production process of washing, dyeing and finishing. Some studies
have reported on the anaerobic treatment of textile wastewater for the production of biogas. Table 10
presents the methane yield of different industrial wastes.

Table 10. Methane potential of industrial waste.

Industrial Waste Methane Yield (mL/gVS) Reference

Textile wastewater 200–400 [123]
Paper mill effluent 220–340 [109]
Cheese whey and diary 280–350 [124]
Cane Molasses stillage 168 [124]
Barley waste 25 [125]
Biodiesel wastewater 161 [126]
Sunflower oil cake 195 [127]
Rapeseed oilcake 310 [128]
Pulp and paper mill sludge 429 [129]
Jatropha curcas oil cake 250 [130]
Sugarcane vinasse 350 [131]

As observed from Table 10, the methane yield of all wastewater streams gave a significant methane
yield that ranged from 25 to 429 mL/gVS. The paper mill effluent with an average methane yield of
280 mL/gVS resulted from a COD removal efficiency of 50–65%. The reported paper mill effluent
is a mixture of four wastewater streams namely foul condensate milling from the chemical pulping
process, liquid waste from the bleaching process, screw press liquor and alkaline extraction operating
liquid [109]. Barley waste that results from the production of an instant coffee substitute gave the
lowest methane yield of 25 mL/gVS with a TS and VS reduction of 31% and 40%. This low yield is
due to the existence of complex heterocyclic compounds that emanates from the hydrolysis phase
of the anaerobic digestion process. Moreover, knowing that the chemical structure of compounds
has been found to have a direct influence on the methanogenic degradation mechanism, the alkaline
hydrolysis pretreatment was applied, which resulted in an increase in methane yield (222 mL/gVS) [125].
Sugarcane vinasse is a recalcitrant wastewater that emerges from ethanoic distillation of sugarcane.
The high methane yield (350 mL/gVS) of sugarcane vinasse could be traced to the organic matter
and solid minerals content. Some of these organic compounds include alcohols, esters, aldehydes,
acids, ketones and sugars. While the solid minerals are majorly potassium, calcium, sulphate ion,
magnesium as well as phosphorus [131]. As noted in the literature calcium and magnesium are
essential macronutrients that enhance the anaerobic digestion process.

4. Conclusions
Anaerobic digestion is a suitable technology for the efficient management of organic waste that
contributes to the uncontrolled emission of methane and carbon dioxide when dumped in landfill sites.
Biogas produced through AD has a wide range of applications; it can serve as a cooking fuel as well as
being a good substitute for other cooking fuels such as coal gas, kerosene, charcoal, cow dung and
firewood. In addition, biogas could be used in a direct combustion system for power generation, space
heating, drying, water heating and fuel for vehicles. This review study has highlighted the various
substrates used in biogas production and their limitations such as longer retention time for industrial
wastes, slow hydrolysis for lignocellulosic materials and high nitrogen concentration of animal manure,
which affect the overall yield of biogas. However, codigestion of these substrates is a good approach
for the simultaneous treatment of solid and liquid organic waste. This will aid in solving the nutrient
imbalance problem associated with the anaerobic treatment of single substrates and the effects of toxic
Processes 2020, 8, 1224 15 of 21

compound build-up during the digestion process. Consequently, biogas production will be greatly
enhanced by codigestion as compared to monodigestion.

5. Recommendation
The type of substrate digested, operating pH, temperature, organic loading rate, hydraulic retention
time and digester design controls the efficient production of biogas through AD. For maximum growth
and activities of microorganisms that enhances efficient biogas production, essential organic and
mineral nutrients are needed in the substrate. The concentration and availability of these macro
substances, microelements and vitamins in substrates vary. Lipid-rich and protein-rich substrates
result in a higher yield of methane compared to carbohydrate-rich substrates. However, high lipid
content in substrates results in the formation of long-chain fatty acids that causes anaerobic digestive
system failure. Additionally, carbohydrate rich substrates can affect the C/N ratio, which results in
nutrient restriction and quick acidification. Hence, a mixture of substrates is recommended to achieve
nutrient balance, process stability and enhanced biogas yield. The study recommends that the choice
of substrates for anaerobic digestion should be guided by their nutrient composition, availability,
cost and pretreatment availability. Finally, future researchers should consider supplementing the
organic wastes with a little quantity of inorganic fertilizer that is small enough to enhance microbial
growth and activities as well as not cause salinization effects. This could enhance the yield of biogas
per given digested organic waste substances.

Author Contributions: Conceptualizing, and writing—original drafting of the article was: N.N., writing a section
of the article and editing: M.E., Proof reading of manuscript: P.M. and K.O. All authors have read and agreed to
the published version of the manuscript.
Funding: This article was made possible through the financial support received from GMRDC of University of
Fort Hare, South Africa.
Acknowledgments: The authors would like to acknowledge the support received from the biogas/biomass
research group of the Department of physics, University of Fort Hare.
Conflicts of Interest: The authors declare no conflict of interest. The funders had no role in the design of the
study and drafting of the manuscript.

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