applied

sciences
Review
Anaerobic Co-Digestion of Wastes: Reviewing Current Status
and Approaches for Enhancing Biogas Production
Rubén González 1 , Daniela Carrillo Peña 2 and Xiomar Gómez 2, *

1 Department of Electrical, Systems and Automatic Engineering, School of Industrial, Computer and
Aeronautical Engeneering, University of León, Campus de Vegazana, 24071 Leon, Spain
2 Chemical and Environmental Bioprocess Engineering Group, Natural Resources Institute (IRENA),
Department of Applied Chemistry and Physics, University of León, Av. de Portugal 41, 24071 Leon, Spain
* Correspondence: [email protected]

Abstract: Anaerobic digestion is one of the technologies that will play a key role in the decarbonization
of the economy, due to its capacity to treat organic waste, recover nutrients and simultaneously
produce biogas as a renewable biofuel. This feature also makes this technology a relevant partner for
approaching a circular economic model. However, the low biogas yield of traditional substrates such
as sewage sludge and livestock waste along with high installation costs limit its profitability. Further
expansion of this technology encounters several barriers, making it necessary to seek improvements to
attain a favorable financial balance. The use of co-substrates benefits the overall digestion performance
thanks to the balancing of nutrients, the enhanced conversion of organic matter and stabilization,
leading to an increase in biogas production and process economics. This article reviews the main
co-substrates used in anaerobic digestion, highlighting their characteristics in terms of methane
production, kinetic models commonly used and the synergistic effects described in the literature. The
main process parameters and their influence on digestion performance are presented, as well as the
current lines of research dedicated to improving biogas yields, focusing on the addition of hydrogen,
Citation: González, R.; Peña, D.C.; bioaugmentation, supplementation with carbon compounds and nanoparticles, the introduction of
Gómez, X. Anaerobic Co-Digestion of bioelectrodes and adsorbents. These techniques allow a significant increase in waste degradation and
Wastes: Reviewing Current Status reduce inhibitory conditions, thus favoring process outcomes. Future research should focus on global
and Approaches for Enhancing process efficiency, making particular emphasis on the extrapolation of laboratory achievements into
Biogas Production. Appl. Sci. 2022, 12, large-scale applications, by analyzing logistical issues, global energy demand and economic feasibility.
8884. https://doi.org/10.3390/
app12178884 Keywords: methane kinetic models; synergistic effects; bioaugmentation; conductive materials;
Academic Editors: Chang-Gu Lee process parameters; reactor performance
and Ramaraj Boopathy

Received: 28 July 2022

Accepted: 2 September 2022
1. Introduction
Published: 5 September 2022
Organic waste has been traditionally treated by biological processes such as com-
Publisher’s Note: MDPI stays neutral
posting and anaerobic digestion. Composting requires a supply of air to keep microbial
with regard to jurisdictional claims in
metabolisms active, whereas anaerobic digestion lacks these oxygen requirements result-
published maps and institutional affil-
ing in a less exigent energy demand. Digestion technology has been applied worldwide
iations.
because the process can deal with high organic loading and generates biogas, which can
be easily valorized for producing heat or electricity. The excellent capacity for treating a
wide variety of wastes makes anaerobic digestion a technology capable of reintroducing
Copyright: © 2022 by the authors.
low-quality materials into the production chain, attaining their transformation into energy,
Licensee MDPI, Basel, Switzerland. organic amendments or any other type of goods. Therefore, wastes can be used to generate
This article is an open access article new products and should be considered as “renewable resources” [1].
distributed under the terms and Digestion technologies have a relevant role in transforming the linear economy model
conditions of the Creative Commons by integrating circularity. Biogas is the main energetic product obtained, which is composed
Attribution (CC BY) license (https:// of methane (CH4 ) and carbon dioxide (CO2 ) as its majority constituents. Digestate is also
creativecommons.org/licenses/by/ derived from this process and contains anaerobic biomass, partially degraded organic ma-
4.0/). terials and residual components which are recalcitrant to the degradation route. Digestate

Appl. Sci. 2022, 12, 8884. https://doi.org/10.3390/app12178884 https://www.mdpi.com/journal/applsci

Appl. Sci. 2022, 12, 8884 2 of 30

is rich in humic, fulvic substances and nutrients making it suitable as raw material to
produce organic fertilizers [2]. Recently, digestate is also being considered as an organic
soil improver, growing medium or organic non-microbial plant biostimulant [3].
Digestion plants can also become an excellent ally for mitigating greenhouse gases
(GHG). The treatment of organic waste avoids uncontrolled degradation and thus the
release of methane into the atmosphere. Biogas obtained from this process is easily val-
orized for energy production (thermal or electrical) or upgraded to obtain a gaseous fuel
with similar characteristics to that of natural gas. There is a rising concern regarding the
effect of CO2 concentration in the atmosphere and how the global climate responds to
the continuous increase in CO2 levels. Research efforts should focus on attenuating these
changes, reducing the negative impact on the economy and searching for efficient ways
of producing energy from renewable sources. The current energetic crisis needs urgent
solutions provided by mature technologies capable of producing huge amounts of energy.
Anaerobic digestion is capable of generating eco-friendly energy and, at the same time,
addressing the waste management crisis [4]. However, several aspects are still pending a
solution such as the profitability of the whole treatment system and the need to increase
conversion efficiency to reduce installation costs.
Anaerobic digestion has been traditionally linked to the treatment of sewage sludge
in large-scale wastewater treatment plants and the treatment of livestock waste. Both
applications are characterized by the use of substrates with high organic content but
lacking suitable nutrient balances. The addition of a co-substrate to any of these systems
aids in balancing the C/N ratio and improving the global process performance allowing a
better economic balance by increasing profits [5]. In the case of sewage sludge digestion,
its composition based on primary and secondary sludge results in a mixture that would
demand an excessive amount of energy if stabilization is carried out by aerobic treatment.
In addition, secondary sludge or waste-activated sludge may need the application of
pre-treatments for facilitating microbial degradation under anaerobic conditions, but this
increases the energy demand. On the other hand, when dealing with livestock wastes, it is
usually accepted that the methane yield of these organic materials is not high enough to
make digestion attractive. Nitrogen-containing compounds may cause inhibition leading
to poor performance. Therefore, adding a co-substrate capable of balancing the C/N
ratio and trace element content will significantly increase methane production [6,7] and
energy valorization.
Improving the efficiency of anaerobic digestion is of great relevance when considering
this process to be a suitable alternative for energy production. This is a key aspect if
this technology is to play a relevant role in decarbonizing the economy. The valorization
of organics into energy and valuable end-products allows the reduction of the carbon
footprint of different human activities. Low-quality resources are in this way re-integrated
into the global economy as energy, nutrient cycling or valuable organics. However, not all
attempts to increase the efficiency of anaerobic digestion are to be considered adequate.
A careful evaluation of the energy demand of the whole process should be carried out.
The transformation of organics into biogas may require additional equipment based on
co-substrate characteristics and introducing pre-treatment units, which would translate into
further energy demands [8], probably making the whole treatment chain unfeasible. There
are several pre-treatment options for improving the degradation of organics and enhancing
the hydrolysis stage. Still, not all of these alternatives find commercial applications due
to their excessive energy demand, the limited capacity for recovering energy and the
detrimental effect of some chemical compounds generated during the pre-treatment.
Major achievements recently attained in anaerobic digestion deal with new technolo-
gies capable of accelerating the hydrolysis stages (thermal, mechanical pre-treatments,
ultrasound application, additions of chemicals). Novel techniques have been developed
such as the application of pulsed electrical fields, high-voltage pulsed discharges and
electrooxidation [9–11]. The success of the industrial implementation of these technolo-
gies keeps a close relationship between biogas production improvement and the energy
Appl. Sci. 2022, 12, 8884 3 of 30

demanded during pre-treatment. Other alternatives for improving anaerobic digestion are
the supplementation of carbon conductive materials, adsorbents, nanomaterials and trace
elements to enhance organic degradation [12–16]. However, any type of material added to
the system may be subsequently released into the environment, creating interactions with
biota which may result in adverse effects due to the presence of co-contaminants [17].
Although anaerobic digestion is a widely extended technology, several factors prevent
the number of installed units from growing worldwide at a higher pace. These are related
to high installation costs and operational complexities. The economy of scale favors large
industrial plants, but this option is not always possible due to social opposition and
constraints due to substrate transport. Recently, several reports have been published in
the literature regarding the costs associated with these treatment plants, the efficiency of
the process and the enhancement of biogas production in an attempt to increase economic
feasibility [18–20]. There is vast experience at a large scale in co-digestion of sewage sludge
and livestock farm wastes and extensive literature regarding research work also dealing
with this subject [21–23]. However, better performance and faster conversion rates are still
needed to improve plant financial balance and search for configurations that allow the
finding of a mid-point between process conversion efficiency and plant operating costs.
For this reason, great hope is set on the addition of supplements capable of attaining these
objectives, such as conductive carbon materials and low-cost adsorbents [24,25].
The present manuscript provides a description of the substrates suitable for the diges-
tion process. A brief review of the application of different kinetic models for predicting
cumulative methane production under batch tests is also included. The main goal of this
manuscript is to present an assessment of the different parameters affecting co-digestion
process performance and highlight the relevance between microbial interactions and reactor
operating conditions. Finally, an analysis of the current alternatives for increasing biogas
productivity is presented, setting a special focus on reactor dynamics and conversion ef-
ficiency. The present review connects the current state of the art regarding data obtained
under laboratory experimental conditions with the implications expected under large-scale
performance, setting a special focus on process efficiency and treatment capacity. The
novelty of the present document is establishing a link between the findings obtained under
laboratory scale conditions and the implications at large-scale plants.

2. Common Substrates Used in Anaerobic Digestion

Animal manures are residues characterized by high nitrogen (N) and organic content.
Ammonia is released during the degradation of proteins, reaching a high concentration in
the reactor that may inhibit methanogens. This reason explains this fact for the extended
application of co-digestion in livestock farms. The high ammonia content reached in
reactors also affects the equilibrium between different chemical species such as carbonates
and volatile fatty acids (VFAs) derived from the sequential transformation of organics.
Ammonia in the digester liquor is present as free ammonia (NH3 ) and ionized ammonium
(NH4 + ), with the first being considered the most toxic form [26]. The buffering system
created by the presence of these compounds produces an environment where pH is kept at
levels higher than 6.4 units, ensuring suitable acid–base environments for methanogens [27].
Sewage sludge is another common waste traditionally treated by anaerobic digestion.
The treatment of urban wastewaters leads to a rejected stream with a high organic content
and a significant amount of water. Sludge obtained from the primary settler receives
the denomination of primary sludge. The aerobic treatment of wastewater by the waste-
activated sludge process also gives rise to a sludge stream mainly containing microbial
biomass. The rapid growth of this biomass makes the extraction from the biological system
imperative, thus producing a secondary sludge or a waste-activated sludge (WAS). Large-
scale wastewater treatment plants (WWTPs) generally have a sludge line dedicated to the
exclusive treatment of sewage sludge, which is composed of a mixture of the above streams.
Thus, the digestibility of sludge depends on the characteristics of WAS, which is recognized
Appl. Sci. 2022, 12, 8884 4 of 30

as having a limited degradation because cellular material needs to be hydrolyzed prior to

the release of its internal content to make it accessible to the anaerobic microflora.
Anaerobic digestion of food wastes or the organic fraction of municipal solid wastes
(OFMSW) has gained popularity in recent years, with several plants being installed for
treating this material in urban waste treatment centers. Source-sorted separated material
is usually preferred for its higher quality due to the lower presence of inert components.
On the other hand, mechanically separated food waste generates a lower quality material,
needing several additional pieces of equipment to handle the slurry produced. In this latter
case, grit and contaminants contained in the feed need to be removed before introducing
the slurry into the digester. In addition to these inconveniences, the seasonal fluctuation of
this type of waste should be noted, which highly influences biogas production [28]. The
presence of heavy metals is another factor that may also add complexity to pre-treatment
operations. The difficulty encountered when attempting the removal of inert materials
and the risk of obtaining digestate with undesirable levels of toxic compounds make this
digested slurry not suitable for agronomic use.
The application of anaerobic digestion to the conversion of crop wastes and agro-
industrial wastes is another field where this technology finds excellent results. However,
when considering this type of substrate, the seasonal availability should be carefully
evaluated along with the lignocellulosic content and high C/N ratio, which translates into
excessively long digestion times and incomplete degradation. In addition, the low nitrogen
levels and the lack of enough trace nutrients may hinder the successful performance and
proper development of the anaerobic microflora.
Given the different characteristics of these individual substrates, co-digestion of the
above materials becomes the obvious solution for balancing nutrients and reducing the dis-
advantages associated with mono-digestion. The mixture of different wastes and biomasses
allows the adjustment in nutrients, improves the stabilization and conversion of the organic
matter, and results in cost-effective use of installations because a single plant is used for
treating a diversity of organics obtaining higher methane yields from the feeding mix-
ture [29,30]. However, the composition of substrates is not the only factor influencing
the global performance of a digestion plant; other parameters such as seasonal availabil-
ity, transport distance and collecting costs have great relevance in the final decision for
considering whether a material is a suitable co-substrate.
Biogas yields from the co-digestion of food wastes have a range of 0.31–0.88 L/g vs.
(volatile solids) with methane contents in the range of 53–70%, whereas these values are usu-
ally lower for the single digestion of manures [31]. The improvement in process efficiency is
expected to be in the range of 25 to 400%, thanks to the increase in organic loading and the
enhanced degradation of volatile solids [32,33]. The composition of substrates significantly
affects reactor performance. Carbohydrates, proteins, cellulose, hemicellulose, lignin and
lipids present different degradation rates and releases of intermediary compounds exerting
in some cases negative effects in fermentation development. Figure 1 presents a schematic
description of the different substrates frequently used in digestion plants.
Appl. Sci.
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2022, 12,
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Figure 1. Schematic
Figure 1. Schematic representation
representation of different
different substrates
substrates suitable
suitable for
for anaerobic
anaerobic co-digestion
co-digestion and
and
valorization of
valorization of main
main process
process products
products(biogas
(biogasand
anddigestate).
digestate).

2.1.
2.1. Carbohydrate-Rich
Carbohydrate-Rich Substrates
Substrates
Food wastes, wastes from the food processing industry, catering wastes and source-
Food wastes, wastes from the food processing industry, catering wastes and source-
separated wastes from residential homes are characterized by a high carbohydrate content.
separated wastes from residential homes are characterized by a high carbohydrate con-
Saccharides and disaccharides are the main components of fruit and vegetable wastes.
tent. Saccharides and disaccharides are the main components of fruit and vegetable
These compounds are easily converted into fatty acid intermediaries by the anaerobic
wastes. These compounds are easily converted into fatty acid intermediaries by the anaer-
microflora, giving rise to pH changes if the accumulation of these acids overcomes the
obic microflora, giving rise to pH changes if the accumulation of these acids overcomes
buffer capacity of the fermentation media [23]. The VFA imbalance may adversely affect
the buffer capacity of the fermentation media [23]. The VFA imbalance may adversely
the production rate of biogas. Accumulation of these intermediaries is commonly observed
affect the production rate of biogas. Accumulation of these intermediaries is commonly
during the anaerobic conversion of easily degradable wastes. Wastes from the food pro-
observed during the anaerobic conversion of easily degradable wastes. Wastes from the
cessing industry, such as cheese whey or fruit wastes, also have a low nitrogen content
food processing industry, such as cheese whey or fruit wastes, also have a low nitrogen
leading to poor buffering characteristics of the fermentation liquor. The summation of these
content leading to poor buffering characteristics of the fermentation liquor. The summa-
features results in inhibitory levels of acetic and propionic acids. When severe digestion
tion of theseare
imbalances features
present,results in inhibitory
higher levels
carbon chain (C4of
–Cacetic and propionic acids. When severe
5 ) acids and iso-forms can be measured
digestion imbalances
in the fermenting slurry. are present, higher carbon chain (C 4–C5) acids and iso-forms can be

measured in the fermenting slurry.

The anaerobic digestion of cheese whey has been studied under different reactor
The anaerobic
configurations suchdigestion
as up-flow ofanaerobic
cheese whey has blanket
sludge been studied(UASB)under different
reactors reactor con-
and sequencing-
figurations such as up-flow anaerobic sludge blanket (UASB) reactors
batch anaerobic reactors (SBR) [34,35]. Cheese whey is a high organic content stream and sequencing-
batchsoluble
with anaerobic reactors
sugars, (SBR)
which are [34,35].
derivedCheese whey manufacturing.
from cheese is a high organicThecontent stream
digestion ofwith
this
substrate has proven challenging due to the lack of sufficient nutrients to keep a balanced
Appl. Sci. 2022, 12, 8884 6 of 30

microflora and the low buffering capacity of the digestion system. In addition, the presence
of soluble sugars aggravates the reaction imbalance, with acidification outcompeting the
subsequent degradation stages. The application of high organic loading is attained by
retaining anaerobic biomass inside the system. Mesophilic and thermophilic digestion of
this single substrate have been studied by Treu et al. [36] and Fernández et al. [37], indicating
the accumulation of VFA and proposing two-stage systems as a way of overcoming the
acidification problems. Another solution proposed for stabilizing the fermentation is
the addition of different nitrogen-containing substrates such as manures and sewage
sludge [38,39].
Another high sugar-containing substrate is sugar molasses. The digestion of this
material shows similar behavior to that of cheese whey. Therefore, two-stage configurations
where acidification and methanogenesis, or hydrogen production and methanogenesis,
have been proposed to overcome the problems associated with the rapid evolution of VFA
and slow degradation of the acid intermediate stream [40–42]. The use of rich-carbohydrate
substrates may be interesting in co-digestion systems, but the availability of these substrates
is usually determined by their use in animal feeding. Therefore, an increase in the demand
for these by-products will ultimately affect market prices and probably create adverse effects
on the economy. Market distortions should be avoided either by the use of specific energy
crops or by the application of specific policies intended to attenuate market deviations.

2.2. Lignocellulosic Biomass

Other relevant substrates treated by anaerobic digestion are crop wastes, energy crops
and any type of high cellulosic-containing material, such as cellulose pulp mill effluent.
The material conforming plant cell walls (cellulose, hemicellulose and lignin) is a complex
structure with different levels of heterogeneity based on the biological function, age and
type of tissue [43]. Cellulose is a component suitable for valorization through anaerobic
digestion. However, substrates containing cellulose may also have a fraction of the lignin
structure linked to the cellulosic material, making its access difficult to the anaerobic
microflora. Therefore, this is the reason for denoting this biomass as lignocellulosic material.
Cellulose is an insoluble polymer with a high molecular weight having a main struc-
ture formed of D-glucopyranose units linked by β-1,4-glycosidic bonds and cellobiose
repetitive units. Two forms of cellulose are generally considered, a crystalline and an
amorphous structure, which are easier to degrade by enzyme complexes [44]. For cellulose
to be assimilated by microorganisms, the degradation should be initiated exocellularly,
either completely extracellularly with the aid of specific enzymes or in association with
the outer cell envelop layer [45]. Anaerobic bacteria possess cellulosome, which is an
extracellular multi-enzyme complex. This complex attaches to the cell envelope and the
substrate, starting the degradation of cellulose [46].
Hemicellulose is the other main component of lignocellulosic biomass, with a lower
molecular weight than cellulose. Hemicellulose forms together with lignin in a covering
structure of cellulose fibers. Hemicellulose is a polysaccharide containing different types of
sugars linked by β-1,4- and, less frequently, by β 1,3-glycosidic bonds [47]. Hardwoods and
straw contain xylans as the predominant hemicellulose constituent, whereas galactoman-
nans are the largest hemicellulose fraction in softwoods [48]. Cellulose and hemicellulose
can be degraded by anaerobic microflora resulting in the accumulation of recalcitrant
aliphatic molecules [49].
The degradation of cellulose was studied by Yamazawa et al. [50] and Li et al. [51]. Its
degradation produces short-chain components, such as acetic and propionic acid, which
are mainly metabolized by clostridial species. However, this conversion takes a long time
(40–50 days) compared with that of carbohydrates under the same anaerobic conditions [48].
This is the main reason for proposing the application of pre-treatments when biogas
production is intended [52]. Thus, accelerating the initial stages of this degradation process
leads to a significant reduction in digester volume and therefore in plant installation costs.
Appl. Sci. 2022, 12, 8884 7 of 30

Spectroscopic techniques have been used as a tool for evaluating the degradation of
different substrates under anaerobic digestion [53,54]. Techniques such as nuclear mag-
netic resonance (NMR) and Fourier transform infrared (FTIR) spectroscopy allow for the
evaluating of the fate of the process and characteristics of digestates in order to study its
adequacy as an organic amendment and act as a soil improver when analyzing agronomic
benefits [55,56]. Under anaerobic conditions, a preferential degradation of carbohydrates,
cellulose and hemicellulose takes place, thus concentrating chemically recalcitrant aliphatic
structures [57]. Aromatic structures originally present in the substrate may be partially de-
graded, causing also the accumulation of this material. The previous features translate into
large digester volumes and therefore high capital costs. The accumulation of recalcitrant
materials affects the final amount of digestate to be disposed of and becomes a problem if
there is not enough land nearby.
Higher methane yields have been reported for cellulose, but a faster conversion was
found for hemicellulose under mesophilic conditions [58,59]. Lignin structures, on the con-
trary, are scarcely affected during anaerobic digestion, hardly experiencing small changes
in their native structure when extended digestion studies were performed [60]. Due to the
recalcitrance of lignin structures, lignocellulosic biomass is usually used as a structuring
agent during solid-phase fermentation under percolating leachate configurations. The
poor degradation rate of lignocellulosics under anaerobic conditions here becomes an
advantage since the porosity of the percolating bed is desirable to allow the circulation
and homogenization of soluble compounds by leachate recirculation. However, if this type
of biomass is added as a co-substrate, then pre-treatments are recommended to facilitate
access to the microflora. Several value-added products can be obtained from the fractiona-
tion and conversion of this raw material by means of a concatenation of different processes
capable of a sequential transformation, always keeping in mind the global efficiency of the
production line.
The coupling of different processes leads to new developments integrated into the
biorefinery concept, where a set of conversion platforms are available for obtaining green
chemicals and recovering energy. Second-generation biofuels, such as biogas from lig-
nocellulosic biomass, have great potential because of their plentiful abundance, offering
no interference with other commercial activities such as animal feed or crops for human
consumption. Still, the heterogeneous structure of this material and its recalcitrant nature
adversely affect its use as a substrate in biogas plants [61]. Pre-treatment stages consider-
ably increase the energy demand of the installation. Careful analysis should be performed
regarding improvement obtained in biogas production after pre-treatment application and
the energy required in the process.
Thermal pre-treatments are widely extended at an industrial scale due to the ex-
perience gained in pre-treating sewage sludge and the unique feature of recovering
energy from high-quality lateral streams. The hydrolysis of hemicellulose produces
oligosaccharides such as pentose (xylose and arabinose), hexose (glucose, mannose, and
galactose), acids (acetic acid, formic acid, and levulinic acid) and furans (furfural and
5-hydroxymethylfurfural). Insoluble humins are also obtained as products under harsh
hydrolysis conditions [62]. However, the high temperature and high pressure under which
hydrolysis is carried out may release some compounds that can behave as inhibitors [63].
Recalcitrant inhibitory substances, such as furfurals and hydroxyl methyl furfural can be
produced at high temperatures [64,65], thus introducing new complexities into the valoriza-
tion process due to the additional stages necessary for removing these toxic compounds.
Another relevant fact that should be noted is the high installation and operating costs
associated with pre-treatment units, which sum up to the already high capital investments
of digestion plants.
Table 1 reports on the different methane yield values found in the literature for a variety
of substrates. Some of these results present a wide range of variability since methane yields
are highly dependent on the characteristic of the substrate, experimental conditions and
the presence of inhibitory compounds. Another parameter of relevance is the time needed
Appl. Sci. 2022, 12, 8884 8 of 30

for degrading the organic material, which translates into high retention times and therefore
large digester volumes.

Table 1. Methane yields reported by different authors when evaluating individual substrates.

Organic Substrates Methane Yield References

Sewage sludge 0.13–0.45 [66–72]
Food wastes 0.33–0.5 [69,72,73]
Pig, swine manure 0.3–0.5 [19,74–78]
Poultry manure 0.03–0.11 [19,74,78,79]
0.52 [80]
Chicken manure
0.053–0.75 [19]
Cattle manure 0.11–0.54 [12,81]
Slaughterhouse waste 0.2–0.8 [82,83]
Brewery waste 0.3–0.51 [84,85]
Residual glycerine 0.56 [86]
Corn stover 0.3–0.4 [75,87]
Sunflower crop wastes 0.2–0.4 [88,89]
Rapeseed crop wastes 0.25 [75]
Wheat straw (steam explosion pretreatment) 0.25–0.35 [90,91]
Rice straw 0.26 [92]
Grass: Napier grass, Canary grass, King grass 0.15–0.60 [93–95]
Meadow grass 0.39 [96]
Microalgae Chlorella sp. 0.23–0.26 [77,97]
Microalgae Nannochloropsis oculata 0.3–0.35 [98]

2.3. Protein-Rich Substrates

Proteins are also abundant in organic substrates, particularly in those derived from
animal wastes. Slaughterhouse wastes, pig, cattle, chicken manure and any other type of
manure from livestock farms are residues with a high protein content. When dealing with
this material, ammonia accumulation may cause problems in the reactor performance if
an equilibrating carbon source is not added to balance the C/N ratio of the feeding recipe.
Another residue that has been studied recently as a suitable co-substrate is animal carcasses.
This waste is subject to strict regulations, but livestock farms must confront a significant risk
associated with the transport of animal carcasses due to the possible cross-contamination
that may take place because of the route the transport truck must follow during collection
operations, with a risk of failure in decontamination when traveling from one farm to the
other always existing. This risk could be reduced if alternatives are allowed in situ in
compliance with Regulation (EC) 1069/2009 and (EU) 142/2011 for animal by-products, so
these farms could safely pre-treat this material to make it a suitable co-substrate [99].
Arenas et al. [24] studied biogas production from animal carcasses, reporting a
methane yield of 0.47 m3 CH4 /kg vs. from biochemical methane potential (BMP) tests.
Tápparo et al. [100] reported a doubling in gas production when studying the co-digestion
of swine manure along with animal carcasses, and Xu et al. [101] proposed the optimization
of the hydrothermal pretreatment of animal carcasses for increasing biogas production,
given the regulation requirement already established for category 2 material.
Ammonium ions released from the degradation of proteins inhibit anaerobic activity
at values close to 4.0 g/L [102]. However, several factors are relevant in the response of
the microflora to ammonium. Acclimation is crucial for tolerating high levels of this cation
in the reactor liquor, along with temperature and pH. Co-digestion of substrates, with
different C/N ratios, is a proper strategy for enhancing degradation performance [103] and
avoiding toxic ammonia concentrations. Thus, the use of grass, straw and lignocellulosic
biomass in general, or micro-algae biomass as co-substrates are suitable options that have
been evaluated under small laboratory conditions in many cases. However, the availability
of these materials should be carefully considered, with this not being always possible due
to the limitations imposed by transport distances. The use of energy crops as co-substrates
is currently a feasible option in some European countries, with maize silage being widely
Appl. Sci. 2022, 12, 8884 9 of 30

used as a co-substrate. However, this option is not adequate for many countries and may be
undesirable due to the rise of market distortions. Digestion technology should be integrated
into the economic cycle without requiring additional incentives. Otherwise, the process
would not become a sustainable alternative for energy production.

2.4. Lipid-Containing Materials

Lipid materials are a type of waste coming mainly from the food processing industries,
slaughterhouses, palm oil industry and grease traps present in different industrial and
commercial activities which have this type of collector in their sewage system. This
substrate presents an extremely high biogas potential with a value of 1014 m3 /kg vs. [104],
although its addition to a digester needs careful control of organic loading due to problems
associated with stratification and the formation of long-chain fatty acids which inhibit
methanogenic metabolism [105]. This is particularly true when short hydraulic retention
times (HRTs) are applied [106]. Other problems are associated with technical constraints.
Several authors have reported pipe clogging and foaming problems in digester gas pipelines
along with severe fouling of these lines [107,108]. If all these problems are overcome, this
material could significantly enhance biogas production, almost doubling methane yields
with a small supplementation in the feeding mixture [109].
The digestion of lipids requires hydrolysis into long-chain fatty acids (LCFAs) and
glycerol as a first stage. Glycerol is transformed into an intermediate compound (glycer-
aldehyde 3-phosphate) by a set of enzymes before entering the glycolysis pathway, with
this route being more complex than that followed by glucose [110], causing, in some cases,
the accumulation of propionic acid. LCFAs are subsequently degraded via β-oxidation,
requiring an external electron acceptor for oxidation [111]. Pre-treatments of lipid sub-
strates are considered necessary for enhancing degradation through the increase in surface
area and reducing the formation of conglomerates which affects the hydrolysis stage in a
negative way [112].
Several reports have dealt with the successful digestion of lipid-rich wastes, indicating
increments in methane production between 10 and 200% [113–115] and methane yields
as high as 999.2 mL CH4 /g vs. for slaughterhouse wastes [116] obtained from BMP
tests. The use of lipids as co-substrates in anaerobic digestion and the application of
different strategies to avoid operational problems, such as acclimatization of the microbial
consortium and developments of new reactor configurations, could be a potential approach
for maximizing the valorization of these streams for producing biomethane [117].

3. Modelling Cumulative Methane Production

Labatut et al. [118] studied the behavior of several substrates using BMP tests and
reported differences in biogas evolution behaviors based on the ease of degradation of the
substrate by the microflora. Edwiges et al. [119] also studied the biogas production from
several substrates and the effect of their composition. It is reasonable to assume that the
main substrate constituents will keep a close relationship with the methane yield obtained
and the evolution of the gas production curve. However, factors such as accessibility to the
degradable components and inhibitory conditions created during their transformation may
alter the final outcome. Thus, carbohydrate-rich substrates are usually well-fitted to first-
order decay models (Equation (1)) whereas, more complex substrates or those experiencing
inhibitory conditions are fitted to different models capable of predicting these effects.
 
B(t) = B0 1 − ek·t (1)

where B(t) represents the cumulative methane yield at any time, t is the time of the batch
assay, B0 is the maximum gas produced by the substrate and k is the first-order con-
stant. The value of the first-order constant gives an indication of the easiness of degrada-
tion. Thus, substrates rich in carbohydrates are characterized by high k values between
0.39–0.66 1/d [119]. This model gives curves with a fast evolution of biogas during the
Appl. Sci. 2022, 12, 8884 10 of 30

first days of the batch assay, usually ending the fermentation in a short time and reach-
ing a plateau very soon, coinciding with the end of the degradation. However, this fast
degradation may cause operational problems in digesters and also when evaluating BMP
tests. The inoculum-to-substrate ratio used for starting up the tests greatly affects the
ultimate methane production. When acidification is expected to become a problem, adding
a greater amount of inoculum, and if necessary alkaline solutions, would aid in obtaining
the desired results.
The addition of rich-carbohydrate substrates as feed to a reactor operating under
a continuous mode may cause localized acidification resulting in pH excursions. Many
reactors work using a feeding recipe where the substrate is introduced at specific hours of
the day. These substrates may cause a temporal variation in methane content, increasing
CO2 levels in biogas and leading to a greater gas production rate right after feeding.
Special care should be taken regarding the type of reactor configuration based on
operating conditions and substrate composition. Continuous stirred tank reactors (CSTRs)
run under equivalent values of HRT and cell retention time (CRT). Therefore, imbalances
may appear due to the lack of enough methanogenic microorganisms. The lower growth
rate of these organisms compared with that of acidogenic bacteria may cause a washout
of the first ones. Other configurations capable of retaining biomass in the reactor may be
more suitable for dealing with substrates that are easily degraded, thus needing a shorter
retention time such as UASB, anaerobic filters or anaerobic sequencing batch reactors
(ASBR). Recent strategies for attaining biomass retention and improving degradation rates
are the introduction of active filling to promote biogas production and attain higher-quality
effluents thanks to the presence of magnetically active filling layers inside the reactor
which are capable of reducing the nitrogen and phosphorus concentration [120]. The use
of active filling containing metals (copper and iron) has also proven effective with similar
results [121].
Table 2 presents the k values obtained by different authors under mesophilic conditions.
There is a wide range of values reported in the literature for this parameter which is highly
dependent on the characteristics of the substrate and the experimental conditions [122],
such as substrate concentration, inoculum-to-substrate ratio, temperature and particle size.
As a matter of example, a decrease in the value of the disintegration rate constant was
reported by Liotta et al. [123] with the increase in solid content and particle size of the
substrate. Any increase in temperature leads to a rise in the reaction rate, whereas reducing
particle size increases the specific surface area making it more accessible to the enzyme
attack. Aldin et al. [124] studied the effect of particle size using casein as model protein
material. These authors found an increase in the hydrolysis constant from 0.034 to 0.298 1/d
by decreasing this parameter.

Table 2. Values of first-order decay constant available in the literature at mesophilic conditions.

Substrate k (1/d) Methane Yield (L/g VS) Reference

Cattle, pig manure 0.106–0.149 0.217–0.287 [125]
Pig manure, swine 0.213 0.202 [86]
manure 0.11 0.161 [126]
0.037–0.086 0.254–0.290 1 [127]
0.069–0.278 - [128]
Cattle manure
0.082 0.239 [129]
0.19 0.238 [130]
Chicken manure 0.07–0.12 0.298–0.351 [131]
Grass 0.107 0.400 [129]
Two-phase olive
0.054 - [132]
pomace
Appl. Sci. 2022, 12, 8884 11 of 30

Table 2. Cont.

Substrate k (1/d) Methane Yield (L/g VS) Reference

0.55 - [133]
Food wastes
0.2 0.524 [126]
Organic fraction
municipal solid 0.0061 - [134]
wastes
Fruit and vegetable
0.34 0.350 2 [130]
wastes
Vegetable crop
0.094–0.167 0.094–0.147 [59]
residues
Food waste
leachate–sewage 0.08 0.343 [135]
sludge
0.197 0.0008–0.0023 [136]
Corn stover
0.06–0.11 0.218–0.300 [131]
Green corn stover 0.159 0.347 [137]
Air-dried corn stover 0.0624 0.319 [137]
0.123 0.348 [129]
Cellulose
0.32 0.353 [130]
1 Calculated from gas and vs. data reported. 2 Digitized from graph reported.

The composite structure of lignocellulosic materials translates into long degradation

times, making inadequate the use of the first-order decay model to describe cumulative
biogas evolution from batch tests. The slow degradation of cellulose and hemicellulose,
contrary to what is observed in carbohydrates, makes the modified Gompertz model more
suitable (Equation (2)). This model has demonstrated adequacy to evolved biogas data
from substrates such as manures and agronomic wastes [138,139].

( Rmax
B0
e (λ−t)+1)

B(t) = B0 ·e−e (2)

In this model, two additional parameters are introduced with regard to the first-order
one. Rmax represents the maximum methane production rate. λ represents the delay
associated with the acclimation of the microflora to a different substrate and environment.
The e number (2.718) is also used in this equation. The modified Gompertz model presents
the advantage of fitting biogas evolution from complex substrates or those containing
partial inhibitory compounds. Thus, λ values describe the delay in biogas production
during the initial stage of the fermentation. Other models containing a lag phase have also
been evaluated with good results, such as the transfer function model, logistic function,
cone model and Richards’ model [140–142]. Diauxic metabolism has also been considered
appropriate for the description of cumulative biogas curves using the modified Gompertz
and the logistic model for describing each sequential degradation stage [143,144].
Table 3 shows a list of different values reported in the literature for λ and Rmax . Values
of λ are usually in the range of 1.5–9.4 days [145,146]. This parameter, as well as the
maximum production rate, is affected by inhibitory substances either already present in
the substrate or produced during the fermentation, as would be the accumulation of VFAs,
long chain fatty acids and ammonia. Sánchez et al. [147] showed that an increase in the
lipid fraction during anaerobic co-digestion of slaughter wastewater caused an increase in
the lag phase. Similar results were also reported by Andriamanohiarisoamanana et al. [148]
when evaluating the co-digestion of manure, slaughter wastes and glycerol. Increasing
the content of glycerol in the mixture also caused a greater delay due to the fast initial
conversion and accumulation of propionic acid.
Appl. Sci. 2022, 12, 8884 12 of 30

Table 3. Values reported in the literature for different substrates under mesophilic conditions regard-
ing kinetic parameters from the Gompertz model.

Rmax Methane Yield

Substrate λ (Days) Reference
(mL CH4 /g vs. d) (L/g VS)
0 25.2 0.322 [149]
Swine manure
0.5 12.8 0.161 [126]
2.45 15.7 0.239 [129]
Cattle manure
0 11.9 0.202 [149]
0.3–2.8 19.4–48.9 0.180 1 [136]
Chicken manure
0 19.2 0.258 [149]
Food wastes 0.5 72.3 0.524 [126]
Food waste
leachate–sewage 1.98 28.4 0.343 [135]
sludge
Waste activated
5.4 19.2 0.253 [150]
sludge
Grass 1.94 34.5 0.400 [129]
Corn stover 0.9–1.9 16–32.1 0.218–0.300 [131]
Liquid effluent
from Biorefinery
3.9–10.2 44.7–66.3 0.459–0.505 [151]
(treating grass
material)
Cellulose 2.93 42.0 0.348 [129]
1 Digitized from graph reported.

4. Taking Advantage of Process Synergies

Digesting a mixture of several substrates is an efficient way of enhancing reactor
performance and increasing methane yields. The main advantages of co-digestion, as
already stated, are associated with the supply of nutrients lacking in single components,
thus equilibrating the feeding recipe. Different values of biogas yields obtained from
co-digestion experiments are listed in Table 4. The greater yields obtained for the mixture
than from the individual digestion of substrates are usually explained by a better balance
of nutrients and therefore are represented as a positive synergy. However, if this is not the
case, then a summation effect is still of interest because of the increase in organic loading
provided, which increases reactor productivity.

Table 4. Methane yields found in the literature for different co-digestion mixtures obtained from
batch tests.

Methane Yield
Digestion Mixture Reference
(L CH4 /g VS)
Sewage sludge + food wastes 0.293–0.365 [72]
Waste activated sludge + organic fraction of
0.162–0.243 [152]
municipal solid wastes
Sewage sludge + sludge from brewery 0.176–0.263 [135]
Sewage sludge + food waste leachate 0.233–0.344 [135]
Sewage sludge + maize straw 0.336–0.472 [71]
Sewage sludge + cattle manure 0.352–0.470 [71]
Swine Manure + glycerine 0.349–0.467 [86]
Pig manure + ESBP 1 0.212 [153]
1 ESBP: exhausted sugar beet pulp at 25:75 mixture ratio.

Some authors reported an additive effect in biogas production when studying the
mixture of agricultural by-products and manures [75,96]. On the contrary, Li et al. [154]
Appl. Sci. 2022, 12, 8884 13 of 30

reported a higher biogas production when co-digesting sewage sludge and leachate derived
from food wastes in MSW incineration plants than from the individual substrates. In this
latter case, the greater production was explained by the enhancement in solid removal
which brings a higher methane evolution as a result. Similar performance was reported
by Anjum et al. [155] and Ghaleb et al. [156] when studying co-digestion at different C/N
ratios, indicating that the modification of the C/N ratio favors microbial activity and
improves solid removal. The benefit associated with the addition of the co-substrate is
considered as a priming effect by Insam and Markt [157] in resemblance to the enhanced
organic matter decomposition that takes place in other habitats such as soils and sediments.
This effect would explain the greater removal of volatiles frequently reported by several
authors. However, the addition of a readily degradable substrate can affect the outcome of
the digestion system; not always getting benefits.
Tambone et al. [49] indicated that anaerobic digestion proceeds through preferential
degradation, accumulating complex structures in the remaining solids. Therefore, the
co-digestion may lead to a degradation of carbohydrates contained in the co-substrate,
hardly modifying the other components of the mixture [23]. In addition, not all BMP
results can be directly extrapolated to a continuously operating system, as demonstrated
by González et al. [86]. These authors indicated the successful performance of batch
co-digestion tests under different co-digestion ratios but failures when attempting the
semi-continuous operation of these same mixtures. Seekao and co-workers [158] set a
mathematical connection between BMP and continuous operation using Monod kinetics.
These authors indicated that although these tests give no clue regarding chronic toxicity,
there is an evident link associated with microbial kinetics for both modes of operation
which set the optimum operating conditions regarding the organic loading rate (OLR), HRT
and methane production. Therefore, the different constituents of the feeding mixture may
not be fully degraded if the OLR and HRT of the reactors are not in accordance with the
microbial dynamics, although BMP tests predicted successful results.
Co-digestion allows the treatment of substrates that otherwise would not be possible
or would lead to extremely low methane yields. This is the case in the study performed
by Zahedi et al. [159] when co-digesting a mixture composed of chicken manure, sewage
sludge and wine distillery wastewater. The results reported by Porselvam et al. [115] and
Cuetos et al. [160,161] are similar, who studied the digestion of slaughterhouse wastes.
The high lipid content in the first case and the high protein content in the second, where
residual blood from slaughterhouses was studied, prevented the correct development of
the fermentation when attempting the mono-digestion of these substrates.
The benefits of co-digestion are undeniable, but high installation costs along with the
need for high-skill personnel for the operation and maintenance tasks are two important
barriers to be overcome. Policies should focus on solving these issues, proposing solutions
for small- and mid-size treatment waste systems which present serious difficulties in
attaining proper waste valorization at a reasonable cost. Partial decentralization may
become a practical solution if small digestion units are dedicated to treating local wastes,
whereas raw biogas may be transported and upgraded at a centralized treatment plant.
Figure 2 shows a schematization of different substrates commonly used in anaerobic co-
digestion focusing on expected inhibitory effects.
Appl. Sci. 2022, 12, 8884 14 of 30
Appl. Sci. 2022, 12, x FOR PEER REVIEW 14 of 32

Figure2.2.Co-substrates
Figure Co-substratesused
usedininanaerobic
anaerobicdigestion
digestionand
andexpected
expectedproblems
problemsininreactor
reactordynamics.
dynamics.

Co-Digestion
Co-DigestionatatLarge
LargeScale
Scale
Another
Another relevantadvantage
relevant advantageofofco-digestion
co-digestionisisthat
thatititattains
attainsaabetter
betteruse
useofofequipment
equipment
and cost-sharing because existing facilities can be adapted, or
and cost-sharing because existing facilities can be adapted, or new ones can new ones canbebebuilt
builtto
toprocess
process multiple waste streams in a single unit [162,163]. The biogas production
multiple waste streams in a single unit [162,163]. The biogas production of exist- of
existing plants can be increased by treating other materials from different industrial
ing plants can be increased by treating other materials from different industrial and agro- and
agronomic sectors.
nomic sectors. However,
However, implementing
implementing thisthis approach
approach for for already-operating
already-operating plants
plants im-
implies the introduction of several modifications in the facility to endow the system
plies the introduction of several modifications in the facility to endow the system with the with
the necessary
necessary flexibility
flexibility for storing,
for storing, pre-treating
pre-treating and feeding
and feeding the co-substrate
the co-substrate into theinto the
reactor.
reactor. The plant must cope with the available co-substrates found in the
The plant must cope with the available co-substrates found in the surroundings, otherwisesurroundings,
otherwise transportation
transportation costs wouldcosts would
cancel outcancel out any
any benefit benefit associated
associated with the
with the higher higher
biogas pro-
biogas production. The integration of co-digestion in WWTPs can be a solution
duction. The integration of co-digestion in WWTPs can be a solution for reducing the op- for reducing
the operating costs and also for increasing the electricity produced by the plant and the
erating costs and also for increasing the electricity produced by the plant and the share
share dedicated for self-consumption.
dedicated for self-consumption.
Digestion plants have a delicate balance between waste treatment, energy production
Digestion plants have a delicate balance between waste treatment, energy production
and economic feasibility, with the latter being dependent on the plant scale and revenues
and economic feasibility, with the latter being dependent on the plant scale and revenues
obtained from the different products. In recent years, the number of large-scale diges-
obtained from the different products. In recent years, the number of large-scale digestion
tion plants installed has increased significantly, providing economic and environmental
plants installed has increased significantly, providing economic and environmental ben-
benefits [153]. There exist many literature reports about the successful performance of
efits [153]. There exist many literature reports about the successful performance of co-
co-digestion with sewage sludge and manures under a laboratory scale [22,68,75,164].
digestion with sewage sludge and manures under a laboratory scale [22,68,75,164]. Large-
Large-scale reports are less abundant, and therefore those found in the literature such as
scale reports are less abundant, and therefore those found in the literature such as Bol-
zonella et al. [165], Sembera et al. [166] and Koch et al. [167] represent a valuable source
of knowledge.
Appl. Sci. 2022, 12, 8884 15 of 30

Bolzonella et al. [165], Sembera et al. [166] and Koch et al. [167] represent a valuable source
of knowledge.
The benefits of co-digestion are associated with greater energy production thanks
to the higher treatment capacity, organic loading increase and higher methane yields [5].
The increase obtained in biogas production may cover the whole energy demand of the
WWTP [168,169] and can be high enough, depending on the type and amount of co-
substrate added, to become an efficient way of obtaining surplus energy and be considered
as an eco-friendly and economically viable approach [170]. However, when attempting
large-scale co-digestion strategies in already existing plants such as digesters in WWTP,
operating problems become frequent as they are usually related to the high variability of
co-substrate composition, changes in technical routines, maintenance and the installation
of additional equipment to deal with these materials [171].
A co-digestion experience at the WWTP of Viareggio and Treviso (Italy) was described
by Bolzonella et al. [165], where the source-sorted OFMSW was treated with sewage
sludge, reporting a 50% increase in biogas production when increasing the organic loading
of the reactor from 1.0 to 1.2 kg VS/m3 d, and a five-fold increase in monthly biogas
production in the second plant studied. However, not all reports present this significant
of an improvement. At the Lansdowne WWTP in the municipality of Prince George,
British Columbia, Canada, Park et al. [172] reported the results of a short co-digestion assay
with source-sorted food wastes from supermarkets. The average daily biogas production
was increased by just 8–10%, but several operational problems were highlighted and
associated with this practice, such as the clogging of the hose connecting the chopper pump
and the sludge recirculation line, needing manual maintenance for clearing up the line.
Accumulation of fibrous scum was also reported near the digester floating roof and the
visual presence of impurities in the biosolids was also indicated.
In another large-scale study, a two-year experience was reported by Mattioli et al. [173]
using OFMSW separately collected for this aim. These authors performed their co-digestion
study with sewage sludge at the Rovereto WWTP. The waste was submitted to a specific
pre-treatment to remove any inert material and obtain a high-quality slurry. However,
the accumulation of floating material was observed on the top layer of the digester and
impurities were detected in dewatered sludge such as plastics, elastic bands and seeds.
Even with these disadvantages, the authors also indicated that the addition of the co-
substrate attained 85% electricity self-generation, whereas in the previous conditions, the
electricity produced from single digestion of sludge only covered 50% of the WWTP energy
demand. Table 5 present a list of different co-digestion studies regarding enhanced energy
performance at a large scale. The homogenization of the results is not possible due to the
different ways and measurement units the authors used for reporting results. However, the
table contains the main characteristics of their studies.

Table 5. Benefits reported by large-scale co-digestion studies found in the literature.

Co-Digestion Amount Added Benefits Disadvantage Reference

SS * + food waste
16% increase in
Grüneck WWTP 5.5 t/d Poor dewaterability [174]
energy production
(Munich, Germany)
SS + organic solid 174% increase in biogas 33% increase in
waste Increase in OLR from production. Energy obtained digestate production
[175]
Zirl WWTP (Tyrol, 1.17 to 2.18 kg VS/m3 d was 115% of the plant and nitrogen back load
Austria) energy demand was doubled
Enhanced methane yield
SS + food waste High nitrogen load in
reporting synergism (12%
WWTP Garching/Alz 10% (w/w) reject water. Reduced [167]
increase). Biogas
(Germany) dewaterability
production doubled
Appl. Sci. 2022, 12, 8884 16 of 30

Table 5. Cont.

Co-Digestion Amount Added Benefits Disadvantage Reference

SS + organic waste
from domestic refuse 80% biogas increase.
Increase in OLR by 25% No reported [176]
Velenje WWTP Increase in vs. degradation
(Slovenia)
Variable 82% biogas increase
SS + fat-waste amount-Added to set 29% vs. removal enhancement
No reported [177]
Iława WWTP (Poland) the OLR at a value of Attain close to total energy
4.8 g/L d as maximum consumption
Slaughterhouse waste +
High ammonia load.
mixture of substrates
Need addition of
Co-digestion plant Energy savings, better odor
ferrous chloride and
operated by the 35–75% (w/w) control, higher gas quality and [178]
hydrochloric acid to
company Svensk production
increase process
Biogas AB (SvB).
stability
Linköping (Sweden)
SS + mixture (milk Solid accumulation
processing industry inside the digester.
wastes and fat from Nitrogen backload.
186% OLR increase 300% CH4 increase [166]
grease traps) Decrease in retention
WWTP Moosburg time and lower sludge
(Germany) dewaterability.
Lower TS removal,
SS and mixture of food Increase in methane yield: higher amount of
waste-garden waste PS: 302 m3 /t TS added dewatered sludge and
Amount of SS: 850
(95:5% based on fresh WS: 133 m3 /t TS added increase demand of
t/year
mass) and grease trap Co-digestion: 627 m3 /t TS flocculants. [179]
Amount of co-substrate:
sludge added (plant data) Sludge disposal
397 t/year
Grossache-Nord Benefit to cost ratio greater represented 64% of
WWTP, Tyrol (Austria) than one overall costs (plant
data)
Kitchen waste added Additional amount of
SS + mixture of wastes Higher nitrogen input,
329 g DM/m3 treated electricity produced:
(kitchen wastes and requiring a more
SS 0.035–0.041 kWh/m3 [180]
fats) efficient denitrification
Fat added Energy self-sufficiency
WWTP Strass (Austria) stage.
9 g DM/m3 treated SS achieved for the WWTP
* SS: Sewage sludge; DM: dry matter, PS: primary sludge, WS: waste-activated sludge.

Co-digestion performed in large-scale plants offers several advantages. Nevertheless,

digestate characteristics may be adversely affected and process modifications may be nec-
essary, resulting in additional complexities in plant operation and maintenance. Research
dealing with these aspects is necessary to properly balance the benefits and inconveniences,
therefore, real practical solutions can be implemented without risking current process
operation. Policies dedicated to favor the flexible treatment of wastes, thereby facilitating
solutions to the final disposal of digestate derived from co-digestion systems are necessary.

5. Improving Reactor Performance

The increase in reactor performance is the best way for improving plant economic
feasibility because of the greater capacity for treating biowastes and improving degradation
rates, which bring along a significant enhancement in reducing the amount of digested
material needing final disposal. Attaining a stable digestion process involves the control
of biological parameters and reactor operating conditions. The dynamics of the process
are very complex because the reactor configuration and feeding rate have a marked effect
on microbial performance and, in turn, the predominant microflora greatly influences the
Appl. Sci. 2022, 12, 8884 17 of 30

process outcome. Table 6 presents a list of the main process parameters influencing the
digestion process.

Table 6. Process parameter influencing anaerobic digestion performance.

Process Parameter Effect

The increase in temperature accelerates degradation rates, fluid dynamics and settling
Temperature characteristics of particles [181]. The improvement in microbial activity increases the reactor
treatment capacity of organics reducing the digester volume needed [182].
pH values should be close to neutral conditions. The stability of the digestion is closely
related to the capacity of buffering acid intermediaries, the release of CO2 and the presence
pH and alkalinity
of ammonia. The interaction between the ionic species and free forms attenuates pH
deviations making the process more robust to organic loading fluctuations [27].
Represents the amount of organic material entering into the digester with the influent.
Increasing the volumetric flow or increasing the solid content of the feeding material leads
to an increase in organic loading. Biogas production is directly associated with the amount
of organics fed into the reactor, and any increment in OLR is usually associated with an
Organic loading rate (OLR) improvement in the biogas production rate. The increase in solid content attained by
adding a co-substrate in anaerobic digestion is one of the main reasons for obtaining a better
volumetric efficiency of the reactor. However, an excess in OLR may also cause process
imbalances due to the accumulation of acid intermediaries associated with disturbances in
the acidogenic and methanogenic phases.
Refers to the time the fluid spends in the reactor. This time is calculated as the ratio between
the volume of the reactor and the volumetric flow applied. HRT and OLR are linked by the
volumetric flow, thus increasing the incoming flow also leads to an increase in OLR and a
decrease in HRT. The time needed for the substrate to be fully degraded depends on the
Hydraulic retention time (HRT) characteristics of the material, complexity in the structure of organic compounds and the
activity of the microflora. Co-substrates characterized by a limited hydrolysis phase will
need a higher retention time in the anaerobic reactor. Inhibitory conditions lead to poor
performance of the microbial activity, with the digestion system not being able to degrade
organics in the time given by the HRT.
Short-chain fatty acids are produced as intermediary compounds during the anaerobic
conversion of organics. Process imbalances lead to the accumulation of these acids,
inhibition of methanogens, and therefore a decline in biogas evolution along with pH
variations when the buffer capacity of the system is surpassed [183].
Process inhibition has been reported to occur at VFA concentrations in the range of
Volatile fatty acids (VFAs)
2000–4000 mg/L [184] depending on the type of substrate evaluated. However, co-digestion
with high N-containing organics allows the maintenance of process stability even though
high levels of VFA may be present. Stable performance was reported by Jiang et al. [185]
when studying co-digestion of pig manure, reporting as inhibitory the VFA range of
16.5–18.0 g/L
This compound is derived from the conversion of protein-rich material. The toxicity of
ammonia in the digester is linked to the level of free ammonia, which is dependent on the
system pH. Nitrogen is an essential nutrient for the process, but excessive levels lead to
methanogenic inhibitory conditions. The ammonium concentration found in the reactor
liquor depends on substrate C/N ratio, HRT and OLR applied to the reactor, and the
degradability of the substrates (hydrolysis performance).
Ammonia also plays a relevant role in the buffer capacity of the system by attenuating pH
drops through the equilibrium ammonia–ammonium reaction. However, a high
Ammonium concentration of ammonium ions may be detrimental to the anaerobic microorganisms.
Moestedt et al. [186] reported that a threshold for stability is found at 1 g NH3 -N/L (free
ammonia), irrespective of the OLR studied.
Acclimation of the microflora to high ammonia levels may attain stable performance when
treating high-nitrogen-containing wastes. Bi et al. [187] reported stable performance of a
full-scale chicken manure digestion plant under ammonium-N levels of 6.2 g/L and Yan
et al. [188] indicated that 8.5 g NH4 + -N/L was the threshold for experiencing inhibitory
conditions, with this value being associated with free ammonia nitrogen (FAN) values
greater than 800 mg NH3 -N/L.
 Appl. Sci. 2022, 12, 8884 18 of 30

Several authors have proposed different alternatives for enhancing methane produc-
tion; among these strategies is worth mentioning the addition of hydrogen gas into the
reactor or introducing a hydrogen-producing culture favors higher levels of this gas in
the reactor liquor [189]. It is widely known that the digestion process is a sequential one
where a delicate balance between the different intermediary species is necessary. The great
capacity for transforming hydrogen gas into methane as described by Martínez et al. [68]
and Zhu et al. [190] has been widely reported, who indicated that the activity of homoace-
togenic microbes was enhanced when continuously feeding hydrogen into the reactor,
increasing the levels of acetate and subsequently favoring the acetoclastic pathway to end
in an increased production of methane. This same idea can be applied to the conversion
of syngas into methane, thus facilitating the treatment of this low-energy content gas and
reducing syngas handling problems.

5.1. Bioaugmentation
Bioaugmentation has also been proposed as an alternative for improving the perfor-
mance of methanogens. Ács et al. [191] studied the performance of digestion systems
inoculated with Enterobacter cloacae cells. These authors obtained a 20% increase in biogas
production after running the inoculated reactor for 6 weeks, using continuously stirred
reactors under a fed-batch configuration. Kovács and co-workers [192] also studied the
performance of mesophilic digestion systems with the inoculation of the same organism (E.
cloacae) and that of a thermophilic reactor using, in this case, Caldicellulosiruptor saccharolyti-
cus. However, they reported that inoculated microflora was washed out from the systems,
with them being incapable of competing with the microbial consortium. Therefore, the
Sci. 2022, 12, x FOR PEER REVIEW 19 of 32
improvements obtained were limited in time. Figure 3 shows several techniques available
for improving anaerobic digestion performance.

Figure 3. Techniques
Figure for improving for
3. Techniques anaerobic digestion
improving performance.
anaerobic digestion performance.

5.2. Operating5.2.
at aOperating at aContent
Higher Solid Higher Solid Content
The increase The increase
in solid in solid
content content
as a way as a way of
of increasing increasing
organic organic
loading, and loading,
thereforeand therefore
digester productivity,
digester productivity, may cause amay cause a detrimental
detrimental effect,
effect, which whichtoisthe
is linked linked to thebal-
delicate delicate balance
ance between VFA, ammonia release, pH and the lower dilution capacity of the system.
High-solid anaerobic digestion and solid-state anaerobic digestion are two forms of car-
rying out the process at high organic loadings. This strategy has as a main advantage the
downsizing of the biological system, but adverse effects may result from the lower water
content. Solid-state digestion is a term used to describe digestion carried out at a solid
Appl. Sci. 2022, 12, 8884 19 of 30

between VFA, ammonia release, pH and the lower dilution capacity of the system. High-
solid anaerobic digestion and solid-state anaerobic digestion are two forms of carrying out
the process at high organic loadings. This strategy has as a main advantage the downsizing
of the biological system, but adverse effects may result from the lower water content.
Solid-state digestion is a term used to describe digestion carried out at a solid content
greater than 15% [193]. The term high-solid digestion is used when carrying out the
fermentation at a solid content greater than 6%, a threshold for the appearance of diffusion
limitations [194,195].
Xu et al. [196] reviewed the performance of digestion reactors treating sewage sludge
under high-solid conditions. These authors reported that the main limitation of these
systems was associated with process instabilities, high viscosity and high concentration of
ammonia and acid intermediaries. The increase in solid content causes the accumulation
of VFAs and ammonia, thus leading to a decrease in methane production [197]. Pastor-
Poquet et al. [198] studied the digestion of the OFMSW under a high-solid configuration
reactor, obtaining 40% less methane yield at a total solid (TS) content of 15.0% and an
NH3 concentration greater than 2.3 g N–NH3 /kg. The performance of the process could
be improved by adding an inert material to exert a diluting effect and decrease the high
localized nitrogen levels. It becomes evident that any attempt to increase reactor treatment
capacity will need to deal with the attenuation of inhibitory conditions.

5.3. Thermophilic Regimen to Increase Reactor Treatment Capacity

The increase in digestion temperature is also an evident way of increasing the degrada-
tion rate and therefore reactor productivity. Anaerobic digestion is strongly influenced by
temperature [199]. Operating at thermophilic conditions and attaining stable performance
may not be exempt from complications. Increasing the temperature of the digestion system
from a mesophilic to thermophilic regimen allows for an existent installation to treat a
significantly greater amount of organics. The biogas yield obtained under thermophilic con-
ditions has been reported by some authors to be similar to that obtained under mesophilic
conditions [200,201], but the main advantage resides in the lower time needed to complete
the full degradation.
Thermophilic regimen may also bring, as a consequence, a lower quality of digested
material. Thermophilic reactor liquor has been reported to contain higher levels of VFA and
ammonia [202–204], which adversely affects its organic quality. Gómez et al. [205] studied
the organic characteristics of cattle manure digestates, indicating a better quality than that
obtained under mesophilic conditions. On the contrary, Provenzano et al. [206], studying
the digestion of sewage sludge and municipal solid wastes, reported a better performance
for the thermophilic system. The apparent inconsistency in results may be explained by the
different characteristics of substrates and nitrogen content. The higher nitrogen content
of manures leads to higher ammonia levels during digestion, having, therefore, a greater
adverse effect on the process outcomes when the temperature is increased.
Yenigün and Demirel [207] performed a literature review regarding the effect of am-
monia in mesophilic and thermophilic digestion. These authors explained the discrepancies
found in numerous studies due to the different levels of free ammonia reached in the
digestion assay, the values of which depend greatly on temperature, pH conditions and
ammonium concentration. Thus, higher temperatures favor the degradation rate, reaching
higher ammonium levels in a lower period, affecting the pH of the system and creating a
toxic environment because of the high free ammonia content in the reactor. The increase in
temperature may need specific adaptation protocols for the microbial biomass based on
the intrinsic characteristics of the substrate and the OLR at which the reactor is expected
to operate.
Takashima and Yaguchi [208] studied the digestion of sewage sludge under ther-
mophilic conditions in a high-solid configuration (9–10% total solids). These authors
included in the digestion system an ammonia stripping stage to remove the excess am-
monia produced. This way, the digestion could proceed at low levels of total ammonium
Appl. Sci. 2022, 12, 8884 20 of 30

nitrogen of 1720 mg N/L (below the value of 2500 mg N/L reported as inhibitory by the
authors) proving that this strategy is an efficient way of attaining higher gas production
rates at high loadings.

5.4. Addition of Adsorbents, Conductive Materials and Nanoparticles

The addition of certain compounds to attenuate the negative effects associated with
acid intermediaries and microbial products may aid in obtaining a better performance of
high-solid and solid-phase digestion systems. Carbon conductive materials, bio-electrodes
and adsorbents may provide alternative routes or protective sites for microorganisms,
thus allowing anaerobic degradation to proceed under highly inhibitory conditions. Wang
et al. [209] reported an outstanding capacity for thermophilic reactors working at high
OLR when biochar was added. These authors attributed this excellent performance to the
ability of char particles to favor VFA syntrophic oxidation thanks to the electron-accepting
capacity of the carbon particles. Petracchini et al. [210] also studied high-solid digestion of
food waste and cow manure. To prevent negative effects, natural zeolites were added to
the reactor, thus obtaining a biogas yield in the range of 680 and 920 mL/g VS. The use of
adsorbents in digestion reactors temporarily reduces the level of ammonia and total VFA
allowing for the process to proceed steadily [211,212].
The experiments carried out by Cuetos et al. [12] clearly demonstrated this fact. These
authors studied the digestion of poultry blood as a single substrate. In this study, the
addition of activated carbon allowed the digestion to be completed, whereas in the control
reactor with no adsorbent addition total inhibition took place. Adsorption is not the only
mechanism justifying the better performance of the biological process, as the presence of
the adsorbent creates protecting sites to the microflora and mass transfer limitations, thus
microorganisms attached to these particles experience a lower concentration of inhibitory
compounds. Several authors have also proposed the mechanism of direct interspecies
electron transfer (DIET) as a phenomenon responsible for explaining the greater capac-
ity of these anaerobic systems to degrade short-chain fatty acids and enhanced biogas
production [213–216].
The addition of biochar particles and the presence of bioanodes in anaerobic reactors
have similar effects to that reported for activated carbon and adsorbents [24,217,218]. Cui
et al. [219] studied the behavior of digestion systems under high-solid conditions and
biochar addition. The improvement in digestion obtained was attributable to the presence
of biochar causing an enhancement in food waste hydrolysis thanks to the promotion of
butyric acid degradation pathways. These authors demonstrated the involvement of the
DIET mechanism by proving a relationship between the Syntrophomonas and Methanosarcina
species. The introduction of bio-electrodes also provides a similar effect in digestion
systems. Moreno et al. [220] described the improvements in VFA degradation thanks to the
role played by soft-carbon-felt electrodes in overloaded batch reactors. However, adding
just a conductive material to create a biofilm attached to its surface is an even more efficient
way of increasing digestion performance as Baek et al. [83] were able to prove. These
authors evaluated anaerobic digestion under fed-batch mode operation, introducing a
large carbon brush device, electrodes and a combination of brushes and electrodes. As
a result, it was observed that a completely different microbial community structure was
formed in the large-size brushes, with Methanothrix being predominant in the biofilm. The
reactor containing these brushes was highly effective in improving digestion performance,
demonstrating a superior efficiency compared with the system using microbial electrodes
with an applied voltage.
Nanoparticles have also been studied as a way for improving biogas production. Ali
et al. [221] studied the addition of three different types of iron oxide nanoparticles report-
ing increments in biogas production, doubling the yields obtained from the conventional
system. The combination of nanoparticles and pre-treatments or their use with bioelec-
trochemical systems has also reported promising results [222,223]. The addition of this
material to co-digestion reactors enhances biodegradability, allowing digestion to proceed
Appl. Sci. 2022, 12, 8884 21 of 30

under conditions that otherwise would lead to upsetting results, as was demonstrated by
Samer et al. [224]. These authors evaluated dry anaerobic co-digestion of manure and whey
supplemented with photoactivated cobalt oxide nanoparticles. The control system showed
a methane yield of 28.01 mL CH4 /g vs. whereas the fermenter containing nanoparticles
gave a result of 169 mL CH4 /g VS. The relevance of these findings lay in the benefits
associated with reactor downsizing due to the higher organic content of the feed and the
greater capacity for avoiding acute acidification. Dry anaerobic digestion and high-solid
anaerobic digestion are biological processes that must affront localized inhibitory levels of
acid intermediaries. The study performed by Ajayi-Banji and Rahman [225] demonstrated
that the use of magnetic nanoparticles (nFe3 O4 )—when evaluating batch digestion of pre-
treated corn stover and dairy manure also using a solid-state system—enhanced reactor
stability by facilitating acid conversion, thus reducing the initial lag phase and increasing
the degradation rate of the different substrate components. Therefore, higher methane
yields were obtained in a much shorter period.
Although the use of nanoparticles may represent a promising technology for improv-
ing biological reaction rates, their final disposal and the possible interactions with microbial
ecosystems should be kept in mind. These particles may finally be released in aquatic
environments and on the soil matrix affecting the biogeochemical cycling of nutrients [226]
(Donia & Carbone, 2019). Future applications of this technology should consider any
negative impacts on the environment by monitoring long-term effects.

6. Conclusions
Anaerobic digestion is a technology capable of converting organic materials into
energy, stabilizing organic matter and recovering nutrients. There is plenty of room to
improve process performance and increase the economic feasibility of digestion plants. Co-
digestion is an efficient way to increase reactor productivity, but factors related to operating
at higher organic loads need to be addressed to improve process economics. Greater plant
flexibility is needed, and economy of scale needs to be carefully evaluated to take advantage
of process synergies and benefit from an enhanced removal of volatile solids.
The co-digestion process has been widely studied in the past and it is expected
that future work will deal with bioaugmentation by inoculating anaerobic reactors with
specific microbiota, allowing an increase in degradation rates and enhancing the removal
of organic components under environments that may currently be considered as inhibitory.
The addition of supplements such as carbon-conductive materials and nanoparticles, the
introduction of bioelectrodes or the development of internal biofilms capable of increasing
the degradation rate of acid intermediaries is another field of research with great expectation
of implementation in the near-term given the significant improvement obtained in methane
yields and reactor performance. More research activities are needed regarding the feasibility
of extrapolating different methodologies that may prove successful on a small scale but
implementation at a larger scale may give rise to serious doubts. Therefore, further research
integrating global process efficiency, by considering the improvement in biogas yields,
along with energy demand and logistical issues are needed. There is an urgent need to
produce huge amounts of energy, which is being exacerbated by the current war crisis.
Production of bioenergy is imperative and attaining this goal is only possible if tech-
nologies involved present clear profitability. The productivity of digesters needs significant
improvements if this technology is to play a relevant role in the development of circular
economy models. Experimental research regarding laboratory conditions under batch tests
and small scales is extensive. However, there is a lack of reports regarding the large-scale
implementation and description of the necessary modifications of equipment and energy
demand associated with the auxiliary equipment involved when co-substrates are added
to conventional digestion units.
Appl. Sci. 2022, 12, 8884 22 of 30

Author Contributions: Conceptualization, X.G. and R.G.; methodology, D.C.P.; formal analysis, X.G.;
resources, X.G. and R.G.; data curation, X.G.; writing—original draft preparation, X.G. and R.G.;
writing—review and editing, D.C.P.; visualization, X.G.; supervision, X.G. All authors have read and
agreed to the published version of the manuscript.
Funding: This research received no external funding.
Data Availability Statement: Not applicable.
Conflicts of Interest: The authors declare no conflict of interest.

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