Bioresource Technology 338 (2021) 125511

Contents lists available at ScienceDirect

Bioresource Technology
journal homepage: www.elsevier.com/locate/biortech

Review

Integrated biohydrogen production via lignocellulosic waste: Opportunity,

challenges & future prospects
Tripti Singh a, 1, Alaa Alhazmi b, c, 1, Akbar Mohammad d, 1, Neha Srivastava e, 1, Shafiul Haque f, j,
Shalini Sharma a, Rajeev Singh g, Taeho Yoon d, Vijai Kumar Gupta h, i, *
a
School of Biosciences IMS Ghaziabad UC Campus, Ghaziabad, Uttar Pradesh 201015, India
b
Medical Laboratory Technology Department Jazan University, Jazan, Saudi Arabia
c
SMIRES for Consultation in Specialized Medical Laboratories, Jazan University, Jazan, Saudi Arabia
d
School of Chemical Engineering, Yeungnam University, Gyeongsan-si, Gyeongbuk 38541, South Korea
e
Department of Chemical Engineering and Technology, Indian Institute of Technology (BHU), Varanasi 221005 India
f
Research and Scientific Studies Unit, College of Nursing and Allied Health Sciences, Jazan University, Jazan 45142, Saudi Arabia
g
Department of Environmental Studies, Satyawati College, University of Delhi, Delhi 110052, India
h
Biorefining and Advanced Materials Research Center, Scotland’s Rural College (SRUC), Kings Buildings, West Mains Road, Edinburgh EH9 3JG, UK
i
Center for Safe and Improved Food, Scotland’s Rural College (SRUC), Kings Buildings, West Mains Road, Edinburgh EH9 3JG, UK
j
Bursa Uludağ University Faculty of Medicine, Görükle Campus, 16059, Nilüfer, Bursa, Turkey

H I G H L I G H T S

• Integrated biohydrogen production have been reviewed using lignocellulosic biomass.

• Integration of dark and photo-fermentation remarkably improves bio-H2 production.
• Integrated process has been discussed in details as new sustainable approach.
• Key factors have been discussed to enhance the performance of integrated process.
• Significance of integrated process have been explored as future prospects.

A R T I C L E I N F O A B S T R A C T

Keywords: Hydrogen production through biological route is the cleanest, renewable and potential way to sustainable energy
Biohydrogen generation. Productions of hydrogen via dark and photo fermentations are considered to be more sustainable and
Dark fermentation economical approach over numerous existing biological modes. Nevertheless, both the biological modes suffer
Lignocellulosic biomass
from certain limitations like low yield and production rate, and because of these practical implementations are
Photo-fermentation
Integrated approach
still far away. Therefore, the present review provides an assessment and feasibility of integrated biohydrogen
production strategy by combining dark and photo-fermentation as an advanced biochemical processing while
using lignocellulosics biomass to improve and accelerate the biohydrogen production technology in a sustainable
manner. This review also evaluates practical viability of the integrated approach for biohydrogen production
along with the analysis of the key factors which significantly influence to elevate this technology on commercial
ground with the implementation of various environment friendly and innovative approaches.

1. Introduction serious environmental impact on the environment (Gonzalez and Rong,

2019; Duffy et al., 2018). Renewable energy production and utilization
Continuously increasing demand of energy of the contemporary so­ is one of the most attentive topics and currently seems to be potential
ciety is the consequences of tremendous changes in life style. On the alternative to replace fossil fuels (Rosen and Fayegh, 2016). In this
other hand, due to the limited life span, fossil fuels are unable to deliver context, biofuels are suggested to be highly potential and green alter­
it fully because of its limited reserve, high fluctuation in cost and their native as the renewable energy sources, and may be capable to compete

* Corresponding author.
E-mail addresses: vijai.gupta@sruc.ac.uk, vijaifzd@gmail.com (V.K. Gupta).
1
These authors have equal contribution

https://doi.org/10.1016/j.biortech.2021.125511
Received 30 May 2021; Received in revised form 30 June 2021; Accepted 2 July 2021
Available online 6 July 2021
0960-8524/© 2021 Elsevier Ltd. All rights reserved.
T. Singh et al. Bioresource Technology 338 (2021) 125511

with the global energy crises, originated due to the various limitations of fermentation (Venegas et al., 2015). Agro-industrial residues such as
fossil fuels (Kumar and Majid, 2020; Saravanakumar et al., 2016). sugarcane bagasse, cornstalk, corn stover, corn cobs, corn bran, wheat
Moreover, biofuels production using biomass is considered as a potential straw, sorghum rusk, sorghum leaves, sorghum stover, rice straw, rice
solution to overcome these challenges (Ahorsu et al., 2018; Ben Iwo bran, rice husk and oat straw, are well reported for the dark fermentative
et al., 2016). Biomasses are one of the most versatile carbon rich, biohydrogen production (Wei et al., 2016; Wang and Yin, 2018; Sekoai
renewable and low-cost resources, being employed to produce various and Daramola, 2018; Raghulchandrana et al., 2020; Magalhaes et al.,
kinds of biofuels like biodiesel, bioethanol, biobutanol, biogas and 2019). However, the complex structures of these biomasses require
biohydrogen (Beschkov, 2017; Busic et al., 2018). Among these, bio­ complicated pretreatment method for making it simple which are the
hydrogen is known to be an excellent renewable energy source and has factors and directly influencing the efficiency of biohydrogen produc­
received enormous attention because of its unique properties like being tion technology. Three polymers namely cellulose, hemicellulose, and
inexhaustible, renewable, pollution free and low-cost (Hroncova et al., lignin build up the complete structure of the biomass which is needed to
2016). Additionally, it also acknowledges to carry the highest energy be firstly processed to remove lignin and hemicellulose via pretreatment
density and no carbon dioxide is generated since the combustion of H2 methods (Baruah et al., 2018). Removal of lignin and hemicellulose
produces only water vapor as the by-product (Salvatore et al., 2021; creates smooth passage of cellulose, breaking down into sugar via
Billaud et al., 2016). enzymatic hydrolysis under the optimum conditions. Enzymatic hy­
Hydrogen can be produced using different techniques such as phys­ drolysis can be proceeded by mutual interaction of cellulase enzymes
ical, chemical and biological processes (Raghulchandrana et al., 2020). and lignocellulosic biomass under the optimum condition by producing
Among these, biological process of hydrogen production presents sig­ high amount of sugars (Zoghlami and Paës, 2019; Robak and Balcerek,
nificant advantages over the other processes due to the various advan­ 2018). Although, sugar production via bioconversion of lignocellulosic
tages e.g. less cost-intensive, use of diverse organic substrates and biomass is one of the main concerns of the biohydrogen production
microorganisms, meanwhile being the renewable in nature and eco- technology, the complicated pretreatment methods and utilizations of
friendly (Xia et al., 2018; Islam et al., 2021). costly cellulolytic enzymes makes the overall process less economically
Lignocellulosic biomasses have huge availability, comprising of high viable (Loow et al., 2016; Yu et al., 2018). Pretreatment of lignocellu­
cellulosic content and thus can be utilized as sustainable and renewable losic biomass (LCB) can be done by three different methods e.g. physical,
substrates for the hydrogen production through biological routes (Al- chemical and biological. Among these, chemical pretreatment method is
Battashi et al., 2019; Kucharska et al., 2018). Cellulose, hemicellulose most-effective and frequently used to remove the lignin contents.
and lignin are the main constituents of the lignocellulosic biomass, and However, due to the high cost of the chemicals and neutralizing agents,
their availability in free form is essential for the effective bioconversion this process is not always feasible for the industrial application. In
following dark fermentation (Akubude et al., 2021; Sarangi and Nanda, addition to the pretreatment cost, enzymatic hydrolysis cost of LCB via
2020). Biohydrogen production through dark fermentation (DF) using cellulase enzyme is another point of concern for its industrial and eco­
sugar hydrolyzate shows various advantages e.g. high H2 production nomic application. Moreover, at present, these cellulolytic enzymes are
rate, use of broad range of organic substrates, but suffers from the produced using synthetic substrate by implementing genetically modi­
drawback of low yield (Osman et al., 2020; Mishra et al., 2019; Bhatia fied strains separately which further increases the overall cost of the
et al., 2021). Further, it is noticed that due to the high H2 production biomass to biohydrogen production technology. In this reference, use of
rate, huge amount of low fatty acids are formed as the metabolites which low-cost sustainable technologies for the effective pretreatment and
makes the fermentation medium more acidic and thus impedes the H2 low-cost enzyme production may be useful to cut down the major
production process (Slezak et al.,2020). However, these low fatty acids challenges significantly. With the combination of suitable pretreatment
are the key substrates for photo-fermentative (PF) H2 production, but processes and low cost substrate based efficient cellulase production
this route suffers from the poor yield of H2. Nevertheless, the above either from mesophilic or thermotolreant microorganism may help to
drawbacks involved in case of DF and PF can be overcome when both the cut down the overall production cost involved in biohydrogen using
processes are combined, and this will significantly contribute to enhance LCB. Several studies have explored that the chemical pretreatment and
the H2production rate and yield both. Thus, following such an integrated thermotolreant fungi could be potential contributors for cutting down
approach high cumulative biohydrogen production can be achieved the cost involved in LCB to biohydrogen production process (Nadeem
(Ntaikou, 2021; Cai et al., 2019; Islam et al., 2021). et al., 2020). Nevertheless, inhibitors formation is one of the major
In view of the above facts, the objective of the present review is to constrains related to the pretreatment process of LCB which hinders the
provide an assessment and the feasibility of integrated biohydrogen overall productivity of the enzymatic hydrolysis and fermentation pro­
production as an advanced biochemical route to improve and accelerate cess (Bundhoo and Mohee, 2016; Srivastava et al., 2020). Kumar et al.,
the biohydrogen production technology in a sustainable manner. The (2015), reported detoxification methods for the pre-hydrolysates ob­
review also evaluates practical viability of the integrated approach of tained via pretreatment and concluded that the efficiency of any
biohydrogen production along with future analysis to elevate this detoxification method depends on both, firstly on hydrolyzate compo­
technology on the commercial ground while implementing various sition, which differs according to the raw material used and secondly on
economical and environment friendly approaches. the hydrolysis conditions employed therein. Because several inhibitory
compounds are formed during the hydrolysis, the identification of these
2. Lignocellulosic biomass to Biohydrogen: Process overview inhibitory compounds, optimization of the hydrolytic process and the
choice of the best hydrolyzate detoxification method are crucial for
Cellulose rich biomass is an ideal candidate considered nowadays for improving the efficiency of the fermentative processes. As each detoxi­
the second-generation biofuels production due to their huge advantages fication method is specific to a certain types of compounds, better results
like being renewable, broadly and easily availability, perennial nature, can be obtained by combining two or more different methods (Reddy
relatively easy bioconversion, non-toxic, environment friendly as well as et al., 2017). Furthermore, biomass to biohydrogen production process
pollution free characteristics (Ferdes, et al., 2020). Among various also depends upon the types of microorganisms employed in fermenta­
available processes of biohydrogen production, dark fermentation is tion process. Anaerobic bacteria like Enterobacter, Bacillus, and Clos­
known to be the most prominent, explores maximum possibility to uti­ tridium are well known H2 producers at large scale via dark fermentation
lize lignocellulosic substrate for the biohydrogen production (Fatma in case of both SSF [simultaneous saccharification and fermentation]
et al., 2018; Calderon and Arantes, 2019). Additionally, lignocellulosic and SHF [separate hydrolysis and fermentation] (Kucharska et al., 2018;
biomasses are the highest cellulose containing biomass and therefore Pugazhendhi et al., 2019). Based on the intense research studies, it is
most effectively studied for biohydrogen production following the dark well established that the SHF is superior to SSF because of the feasibility

2
T. Singh et al. Bioresource Technology 338 (2021) 125511

Table 1
Studies reported on dark and photo fermentative biohydrogen production using different types of lignocellulosic biomass.
Sr. Mode of Microorganisms Substrates Temperature pH H2Production Max. Hydrogen References
No. fermentation rate yield

Dark fermentation Clostridium termitidis ATCC-21846 Rice husk 37 ◦ C 7.5 0.023 mL H2 g- 5.59 mL g-1dw Tosuner et al.
1
and Clostridium intestinale ATCC-BAA dw rice husk rice husk 2019
1027 h− 1
Dark fermentation Mixed culture Oil palm 35 ◦ C 5.0 3266.86 mL H2 283.91 mL H2 Gonzalez and
empty fruit L-1d-1 g-1TS Rong, 2019
brunch
Dark fermentation Escherichia coli Brewery spent 37 ◦ C 7.5 _ 0.75 ± 0.03 Poladyan et al.
grain mmol H2 L− 1 2018
Dark fermentation Caldicellulosiruptorspecies and Wheat straw 30 C◦
6.5 _ 134.0 mmol Soto et al., 2019
Ralstoniaeutropha H2/L
Dark fermentation Clostridium pasteurianum(MTCC116) and Sugarcane 37 ◦ C 7.0 31.66 mL/L/h 2870 mL/L Srivastava
Bacillus subtilisPF_1 bagasse et al., 2018
Dark fermentation Clostridium roseum Cashew apple 37 ◦ C 5.5 12.57 mL H2/L/ 0.08–1.89 mL Silva et al. 2018
bagasse h H2/g biomass
T
Dark fermentation Clostridium guangxienseZGM211 Peanut shells 35 C◦
5.5 _ 6.4–39.9 mL Qi et al. 2018
H2/g biomass
Dark fermentation Trichoderma asperellumBPLMBT1 Sweet 37 ◦ C 7.0 – 402.01 mL Shanmugam
sorghum et al., 2020
stover
Dark fermentation Mixed culture (Anaerobic sludge from a Sugarcane 37 ◦ C 6.5 – 18.6 mLH2 g_1 Moodley and
wastewater treatment plant). leaf waste Gueguim Kana,
2018
Dark fermentation Clostridium pasteurianum(MTCC116) Rice straw 37 ± 2 ◦ C 7.5 23.96 mL/L/h 2580 mL/L Srivastava et al.
2017
Dark fermentation Mixed Culture Sugarcane _ 5.0 _ 0.874 mol (mol Reddy et al.
bagasse glucose − 1) 2017
Dark fermentation Clostridium sp. T2 Corn stalk 37 ◦ C 6.5–7.2 14.2 mmol H2 _ Zhao et al.,
hydrolysate /L/ h 2017
Dark fermentation Mixed culture (Anaerobic sludge from an Empty fruit 35 C◦
7.0–7.5 2061 mL H2 L-1 275.75 mL H2 Gonzalez and
UASB reactor treating brewery bunch of oil d-1 g-1TS Rong, 2019
wastewater) palm
1
Dark fermentation Clostridium butyricum Sugarcane 35 ◦ C 5.5 1611 mL H2/L/ 1.73 molmol− Pattra et al.,
bagasse day 2016
hydrolysate
Dark fermentation Mixed culture (Sludge from a settling Rice Crop 37 ◦ C 6 to 7 97.08 mL/h 30.37 mL/ Sattar et al.
channel) VSrem 2016
Dark fermentation Clostridium butyricum Jatropha hull 35 C◦
5.5 _ 1.95molH2/ Jiang et al.,
mol-TRS 2016
Dark fermentation Clostridium thermocellumATCC 27,405 Sugarcane 55 ◦ C _ _ 109.6 mL/g Tian et al., 2015
bagasse
Dark fermentation Mixed culture Oat Straw 35 ◦ C 7.0 110 mL H2/L/h 2.39 mol H2/ Arreola-Vargas
mol reducing et al. 2015
sugars
Dark fermentation Clostridium beijerinckii KCTC 1785 Sorghum rusk 35 ◦ C 6.0–7.5 46.54 mL/L/h 1,117 mL/L Saratale et al.
(2015)
Photofermentation Spirulina platensis Wet biomass 32 ± 2 ◦ C 5.95 _ 1.92 ± 0.20 Pandey et al.,
mmolH2/ mol 2021
COD red.
Photofermentation HAU-M1 Corn stalk 30 ◦ C 6.0 _ 119.3 mL/g Quanguo
et al.,2017
Photofermentation Mixed bacteria strain (HAU-M1) Corncob – 7.39 _ 84.7 mL/g TS Zhang et al.
2020
Photofermentation Rhodospirillumrubrum, Corn stalk 30 ◦ C 7.3 38.4 mL/L/h 211.9 mL/L- Jianga et al.,
Rhodopseudomonascapsulata, medium 2020
Rhodopseudomonaspulastris, Rhodobacter
sphaeroidesand Rhodobactercapsulatus
Photofermentation Mixed culture Corn stalk 30 ◦ C 7.0 96 mL/L/h 2.6 ± 0.3 mol Jiang et al.,
H2/mol sugar 2016
consumed

to avoid the harmful impact of the enzymatic hydrolysis such negative et al., 2018). Nevertheless, these metabolites effluent i.e. volatile fatty
impacts of inhibitors, raised temperature due to the shaking conditions acids (VFAs) are the potential substrate for the photo-fermentative
and the pH change which directly affect the fermentative microorgan­ bacteria to produce H2 via photo-fermentation under the optimized
isms and consequently biohydrogen production (Shanmugam et al., conditions (Liu et al., 2015; Cai et al., 2019; Ghosh et al., 2018). Addi­
2020). Nevertheless, it usually noticed that the high H2 production rate tionally, by consuming these VFAs effluent the overall biohydrogen
may lead dark fermentative medium highly acidic soon due to the large production can be significantly enhanced by overcoming the challenge
amount of the produced acidic metabolites e.g. acetic acid, butyric acid, of low H2 production rate in case of photo-fermentation. Therefore,
malic, propionic, fumaric and succinic acid (Wainaina et al., 2019; Petra following the integrated approach as presented in this review that
Muri et al., 2016; Baumann and Westermann, 2016). Because of the high combines dark and photo fermentation (DF + PF) while using LCB as
production of these metabolites as the byproducts, hydrogen production substrate can further improve the biomass to biohydrogen production
ceases soon and thereby hinders the overall H2 production yield (Nagy process, significantly. Table 1 describes various studies reported on the

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T. Singh et al. Bioresource Technology 338 (2021) 125511

Fig. 1. Mechanism of Dark fermentative biohydrogen production [Adapted with permission from Hallenbeck et al., 2012].

dark and photo-fermentation using LCB for biohydrogen production. of two major H2-producing enzymes; hydrogenase and nitrogenase
(Mishra et al., 2019; Najafpour et al., 2016). Hydrogenase catalyzes the
3. Dark fermentation: Concept & mechanism inter-conversion of hydrogen to protons and electrons, and are catego­
rized into three classes namely; [NiFe]-hydrogenase, [FeFe]-hydroge­
3.1. Concept nase, and [Fe]-hydrogenase (Ogata and Lubitz, 2020). Further, these
enzymes are participated into two pathway of dark fermentation known
Biological fermentation mode of H2 production in absence of light is as; the acetate pathway which gives theoretical yield of 4 mol H2 from 1
regarded as dark fermentation which also known to be the simplest H2 mol glucose whereas the other pathway called as the butyrate pathway
production methods at ambient condition while using organic substrates and gives the theoretical yield of 2 mol H2 from 1 mol glucose (Islam
(Soares et al., 2020; Mishra et al., 2019; Zhang et al., 2019a). This is the et al., 2021; Sarangi and Nanda, 2020; Sekoai and Daramola, 2018). In
very commonly used biological method of H2 production using versatile the first stage of dark fermentation, NADH is produced viaoxidation of
range of organic and cellulosic wastes (Rosa and Silva, 2017; Hajizadeh substrate into pyruvate and may be utilized by microorganisms having
et al., 2021). Apart from the substrate versatility and simple operation NADH: ferredoxinoxidoreductase (NFOR) to produce reduced ferre­
mode, H2 production rate is much higher in this mode when compared to doxin (Pandey et al., 2021; Preethi et al., 2019).While in the second
other known fermentative methods along with the feasibility to utilize stage, pyruvate can be converted in to acetyl-CoA and formate by py­
various fermentative microorganisms (Preethi et al., 2019; Sivagur­ ruvate formatelyase (PFL) or acetyl-CoA and reduced
unathan et al., 2017). Clostridium beijerinckii, Clostridium butyricum, ferredoxinviapyruvate-ferredo; xinoxidoreductase (PFOR) to produce
Enterobacter aerogenes and Enterobacter asburiae are some well known H2 (Zhou et al., 2015; Sağır and Hallenbeck, 2019). Nevertheless, high
potential producers of H2 in dark fermentation process using versatile production of acids during this process lowers the yield of H2and is
range of substrates (Wang and Yin, 2019). Moreover, potential con­ considered as one of the biggest challenges required to be addressed by
sumptions of different lignocellulosic biomass by these microorganisms researchers. The acids metabolites like ethanol, acetic acid, fumaric,
have been well reported in literatures (Ding et al., 2016; Ferdes, et al., lactic, butyric acid, and propionic acid are produced along with the
2020). Furthermore, like any other biochemical process, dark fermen­ dominant end product PHB. The overall process of dark fermentative
tative biohydrogen production is also influenced greatly by its physi­ reaction can be presented by following equations (Sarangi and Nanda,
cochemical parameters like temperature, pH, hydraulic retention time, 2020):
inoculum amount and size (Preethi et al., 2019; Sivagurunathan et al.,
2017). In addition, H2 production is completely driven by the presence C6H12O6 + 2NAD+→2CH3COCOO− +4H++ 2NADH (1)

4
T. Singh et al. Bioresource Technology 338 (2021) 125511
CH3COCOO− +CoA-H → acetylCoA + HCOO− (PFLP) (2)
have the capability to utilize broad range of organic substrates while
HCOO− +H+→CO2 + H2 (3)
using energy in form of light to produce H2 efficiently. Further, Rhodo­
CH3COCOO− + CoA + Ferredoxin (Fd)ox → acetyl CoA + Fdred + CO2 bacter sphaeroides, Rhodobacter capsulatus, Rhodovulums ulfidophilumW-
(PFORP) (4) 1S and Rhodopseudomonas palustris, Rhodobacter species are the group of
PNS bacteria and well known for H2 production via photo-fermentation
Fdred + 2H+→ Fdox + 2H2 (5) with high yield using VFAs as substrate (Kessi and Hörtensteiner, 2018;
Mishra et al., 2019). Organic acids, such as acetic, butyric, propionic,
Acetyl CoA + H2O → CH3COO− + H+ + CoA-H (6)
malic, and lactic are potential VFAs substrates that have been reported
Acetyl CoA + 2NADH + 2H+ → CH3CH2OH + CoA-H + 2NAD+ (7) for the significant photo-fermentative H2 production (Pandey et al.,
2021; Assawamongkholsiri et al., 2019; Mishra et al., 2019). While some
NADH + H →NAD + + +
H2 of the investigations have also reported photo-fermentative H2 produc­
Efficiency of the dark fermentative biohydrogen productions have tion directly from the sugars like glucose and sucrose showing their
been investigated in many studies (Pandey et al., 2021; Preethi promising nature towards the H2 production (Rai et al., 2014; Pandey
et al.,2019). Reddy et al., (2017), investigated biohydrogen production et al., 2021). Along with substrate and microbes, physicochemical pa­
using sugarcane bagasse under the dark fermentation. About 69.6% rameters optimization is also crucial for the high yield of H2in case of
biohydrogen production was found in presence of magnetite nano­ photo-fermentation (Soto et al., 2019; Pandey et al., 2021). Most of the
particles using Enterobacteriaceae, Clostridium spp. Whereas control photo-fermentative bacterial cell growth and hydrogen production are
system (absence of nanoparticles) showed maximum hydrogen yield achieved at medium temperature range 30–40 ◦ C, whereas pH in a range
(HY) of 0.874 mol (mol glucose− 1) at the optimum pH 0.5.0 Table 1 of 5.0–7.0 and at normal atmospheric pressure with different substrate
summarizes numerous studies reported on the dark-fermentative bio­ and using different light source having wavelength 522, 805, and 850
hydrogen production using different types of lignocellulosics biomass. nm (Liu et al., 2015). Additionally, effluent of the dark fermentation can
be directly used for the photo-fermentation in the sequential process
because no pH change is required (Soto et al., 2019). Substrate type and
3.2. Mechanism conversion efficiency may also contribute significantly to improve the
photo-fermentation mediated H2 production (Zhang and Zhang, 2018).
The fermentative H2 production pathway adaptation depends upon In case of photo-fermentation reaction; substrates’ conversion efficiency
the types of microorganism responsible for performing the dark of malate, lactate, and acetate can be given as following equations
fermentation (Li et al., 2019). Microorganism PFL pathway was not able (Assawamongkholsiri et al., 2019):
to utilize NADH for H2 generation, resulting theoretical value limited to
2 mol of H2per mol of glucose (Srivastava et al., 2017). Further, in the Malate C4H6O5 + 3H2O ———— 6H2 + 4CO2
intermediate stage, Acetyl-CoA is ultimately transforming into lower Lactate C3H6O3 + 3H2O ——— 6H2 + 3CO2
VFAs salt like acetate, butyrate as well as ethanol. The acetate pathway
offers the highest yield of 4 mol H2/mol glucose (4 mol of H2 from 1 mol Acetate C2H4O2 + 2H2O ——— 4H2 + 2CO2
of glucose, 33% of theoretical yield 12H2) and this happens due to the
avoiding of additional consumption of NADH by microorganism while
H2is produced. Moreover, only 4 mol of H2/mol of glucose is considered
as the maximum hydrogen yield via direct dark fermentation of glucose 4.2. Mechanism
and refer as the Thauer limit (Thauer et al., 1977). Whereas, practi­
callyH2 yields are lower than the theoretical value being in the range of Photo-fermentative H2 production via PNS is also an enzyme system
1.2–2.3 mol H2 /mol glucose. In contrast, the butyrate pathway is called as nitrogenase mediated biochemical process which can be con­
limited to 2 mol H2/mol glucose [Fig. 1]. Osman et.al (2020) studied the ducted using variety of different organic acids as electron donor
biohydrogen production via dark fermentation using rice straw as sub­ (Higuchi-Takeuchi and Numata, 2019). The electrons released by the
strate and Clostridium pasteurianum employed as the fermentative carbon substrate are pumped via large number of electron carriers while
microorganism. A maximum 47.6 mL/g hydrogen yield was reported at protons are pumped over the membrane forming proton gradient,
initial pH 7.5 and temperature 37 ◦ C (Osman et.al., 2020). Following the generating ATP in the presence of ATP synthase. The excess ATP formed
similar pattern Soares et. al, reported biohydrogen production using could be utilized to transport the additional electrons to the electron
Rice, corn and wheat residues employed as the lignocellulosic biomass acceptor ferredoxin (Fd) (Jianga et al., 2020). Additionally, in the
(Soares et. al, 2020). The result so obtained revealed that biohydrogen absence of molecular nitrogen, the electrons placed from ferredoxin are
production were ranging from 12 to 7019 mL H2/L via mixed culture used by nitrogenase to reduce protons into hydrogen. Photo-
under the optimum condition of temperature 37 ◦ C, pH 7.0 and the fermentation has been reported to be high theoretical hydrogen yield
hydraulic retention time 72 h (Soares et al., 2020). Similarly, Bhatia which is 8 mol of H2/mol of glucose via PNS bacteria (Mishra et al.,
et al. (2021) recorded the highest specific H2 yield of 4 mol H2/mol 2019). Whereas, the key drawbacks associated with this process include
glucose or 497.8 mL H2 /g glucose via the acetic acid pathway (Bhatia high-energy consumption, low solar energy conversion efficiencies, and
et al., 2021). the low volumetric production rates (Arimi, et al., 2015). It is reported
that the maximum H2 yield is dependent upon the kind of organic acids
4. Photo-fermentation: Concept & mechanism chosen, for an example butyric acid gives the theoretical highest yield
(HY) of 10 mol H2/mol substrate, while acetic acid yields the lowest
4.1. Concept value of 4 mol H2/mol substrate (Zhang et al., 2017) [Fig. 2].
The chemical equation for photo-fermentative hydrogen production
Photo-fermentation is another most popular and adopted mode of from acetate is given in the following equations.
biological hydrogen production after the dark fermentation, which is
2CH3COOH + 2H2O → 4H2 + 2CO2, ΔGo = + 104 kJ
facilitated via the photosynthetic purple non-sulphur (PNS) bacterial
group (Reungsang et al., 2018). These bacteria may utilize long and In biological photo-fermentative hydrogen production process
short chain of volatile fatty acids which can be efficiently converted into practical HY is always less than the theoretical HY and this might be due
H2 and the related metabolites via nitrogenase enzyme under the pres­ to the various environmental impact such as temperature, pH fluctua­
ence of light source (Pandey et al., 2021). Additionally, these bacteria tions, viability rate of microorganism in the fermentative medium,

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T. Singh et al. Bioresource Technology 338 (2021) 125511

Fig. 2. Central metabolism mechanism of Photoferentation in purple non-sulfur bacteria [, ()]. (For interpretation of the references to colour in this figure legend, the
reader is referred to the web version of this article.)
Adapted from Hadicke et al., 2011http://creativecommons.org/licenses/by/2.0

substrate-production inhibition and also the impact of intermediate in­ unavailability of the potential substrates and the high production cost
hibitors (Zhang et al., 2017). Liu et al., (2015) recorded hydrogen yield are the main drawbacks of DF process whereas poor H2production rate is
of 160 mL-H2/60 mL using acetic acid as substrate, at pH 7.0 and with the main constrain of PF process. In this reference, integration of both
inoculum time of 14 h using bacterium Rhodopseudomonas sp. nov. the processes while using LCB as the substrate may help to overcome
strainA7 (Liu et al., 2015). Mishra et al. (2016), performed photo- these challenges (Cheng et al., 2011a). Further, utilization of the VFAs as
fermentative H2 production and recorded an yield of 2.86 mL H2/mL the effluent of DF which are produced in large amount can be exploited
POME using isolated Rhodopseudomonas palustris under the light illu­ as initial substrate for PF process either by directly or by incorporating
mination of 4 flux, temperature 30 ◦ C, in a batch operational mode with minor physicochemical parameters changes. This strategy may
(Mishra et al., 2016). Lu et al., (2016) also investigated the maximum likely to improve the overall yield of H2 (Patel et al., 2021). Thus, by
specific hydrogen yield (SHY) of 111.85 ± 1 mLH2/g TS using apple using residual organic VFAs like acetic acid, butyric acids, lactate,
waste as substrate via photo-synthetic bacteria (PSB) HAU-M1 at initial formate and propionic acids in PF process is an excellent approach and
pH 7.14, light intensity of 3029.67 lx and temperature 30.46 ◦ C has enormous potential to be employed in the integrated fermentation
following the RSM method (Lu et al., 2016). technology (Liu and Smith, 2020; Mohsenpour et al., 2020). Moreover,
this process does not require any external pH adjustment, substrate in­
5. Integrated biohydrogen production via dark and photo- hibition and also reduces the operational time due to the running
fermentation operation in a single stage (Kehrein et al., 2020). The maximum con­
version of substrate in to H2 can be achieved directly by combined DF
In spite of being the most advantageous and sustainable method, and PF process at minimum physicochemical changes. Moreover, the
both the fermentative biohydrogen production methods (DF and PF) highest theoretical hydrogen yield up to 12 mol H2/mol glucose can be
suffer fromcertain drawbacks which creates main hurdles in the way of achieved via this integrated approach (Patel et al., 2018). The integrated
sustainable and economic biohydrogen production technology (Islam biohydrogen production can be operated in a single batch system or in a
et al., 2021; Ghosh et al., 2018; Cabrol et al., 2017). Low yield, sequential two step batch system. In the single stage batch system of H2

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T. Singh et al. Bioresource Technology 338 (2021) 125511

Table 2
Studies on biohydrogen production via integrated (dark and photo) fermentation using different types of lignocellulosic biomass.
Sr. Mode of integrated process Substrate Optimized conditions Hydrogen Production References
No.
1
1. Single stage fermentation Food waste pH5.35–6.85, Temp 32 ◦ C and Light 197.15 ± 2.9 mlg− VS Rezaeitavabe et al.,
intensity 5000 lx. 2020
-1
2. Two stage fermentation (Co- Corn stover pH6.5, Temp. 30 ◦ C and Light intensity 90.13 mL.(g TS) Zhang et al., 2019
fermentation) 3500 lx
3. Sequential Dark and Photo Corn stover pH 4.5 and 35 ◦ C (Dark) pH 7.0 and 30 ◦ C 87.8 ± 3.8 m3/d with 68% Zhang et al., 2018
fementation (Photo) H2 content
4. Two-stage fermentation Platanusorientalis pH 7, Temp. 30 ◦ C, and Light intensity 368.8 mL of substrate Zhang et al., 2017
eaves 3,000 lx
5. Integrated Dark and Corn stover pH 6.5, Temp. 30 ◦ C and Light intensity 332.6 mL for 72 h. Quanguo et al.,2017
Photofermentation 3500 lx
6. Two-stage fermentation Corn stalk pH 7.0 and Temp. 37 ◦ C 339.5 mL-H2/g-corn stalk Yang et al.,2015
7. Mixed dark and photo Sorghum starch pH 7.3, Temp.30 ◦ C, Light intensity Morsy,2015
fermentation 150–200 W/m2
8. Two stage fermentation Sugarcane bagasse pH 6.8, Temp. 34 ◦ C and Light intensity 1000 mL/L (Dark) 755 mL/ Rai et al., 2014
(SCB) 8.5 W/m2 H2(Photo)
9. Mixed dark and photo Sugarcane waste pH 7.0, Temp. 32 ◦ C and Light intensity 0.2, 0.35 and 0.16 mol/kg of Hema and Pushpa,
fermentation (bagasse) 3.75 W m− 2 substrate 2012
10 Two-stage fermentation Rice straw pH 7, Temp. 30 ◦ C and Light intensity 200 463 mL/g TVS Cheng et al. 2011
W/m2 (6000 lx)
11. Two stage fermentation Corn stalk pH 7.0, Temp. 37 ◦ C and Light intensity 294 mL/g corn stalk Ma et al., 2010
4000Lx

production, both types of microorganisms responsible for the DF and PF cellulosic contents. High cellulose and less lignin substrate may reduce
are utilized in mixed form under the certain optimized conditions, but the pretreatment time and the cost, meanwhile improve the sugar con­
the pH control is difficult. Additionally, microorganisms performing tents, required for the fermentation to generate H2 (Bhatia et al., 2021;
both the processes may also have to face mutual metabolites inhibitory Kucharska et al., 2018). Additionally, lignocellulosic biomasses are
effect along with the substrate and the product inhibition factors. On the known for the highest cellulosic biomass to produce biohydrogen with
other hand, in case of sequential two stage integrated H2 production the generated metabolites. Moreover, high sugar content will also
system, microorganisms of PF process are introduced to the reaction ensure the production of a huge amount of VFAs as the byproduct of
medium after the completion of DF meanwhile with the possible dark fermentation and this may be potential as the initial substrate for
adjustment of the pH if required (Tawfik and El-Qelish, 2012). In photo-fermentation (Zagrodnik and Seifert, 2020; Mishra et al., 2019).
addition, temperature of this system is best suited in the range 28–30 ◦ C, Bioconversion of lignocellulosic biomass via cellulolytic enzymes will
which is also the common temperature range of DF and PF mode of H2 also help to reduce the production of inhibitors for continuous produc­
production. Moreover, the optimum experimental conditions would not tion of H2 in DF and PF processes simultaneously. Although, there are a
be compromised in case of two stage sequential integrated H2 produc­ number of lignocellulosic biomass yet needed to be more explored to
tion system (Yang et al. 2015). The positive impact of the integrated find out the optimum feed-stocks for improving the H2production rate
process that combines DF and PF exhibited improved H2 production, and and the yield and for the sustainable process economy of the integrated
has been confirmed by previous studies (Cheng et al., 2011b; Tawfik and H2 production technology (Sivagurunathan et al., 2017; 2017;; Peri­
El-Qelish, 2012; Ghimire et al., 2016; Mishra et al., 2019). Ghimire yasamy et al., 2018). In addition, it is also confirmed from many studies
et al., (2016), investigated an integrated biohydrogen strategy while that the organic-rich biomass such as rice straw, wheat straw and sug­
using effluent of DF in form of organic acids and alcohol under the op­ arcane bagasse have huge potential to be utilized as substrate for the
timum conditions e.g. pH 6.5, and temperature 24 ± 2 ◦ C. The total 256 biological mode of hydrogen production (Rai and Singh, 2016; Sołowski
(±2) N mL H2/g chemical oxygen demand (COD) could be achieved in a et al., 2020; Venegas et al., 2015). Therefore, it may be concluded that
sustainable manner. Further, Mishra et al. (2016), conducted a single the substrate selection directly influence the overall H2production
stage sequential H2 production by integrating DF and PF process. The following integrated technique that combines DF and PF process (Osman
maximum H2 yield could be gained in the first stage of DF was recorded et. al., 2020). A simple and low-cost pretreatment technology and
to be 0.78 mL H2/mL POME, which further could be enhanced up to cellulolytic enzymes production may also improve the sustainability of
2.86 mL H2/mL POME after the photo-fermentation via lab isolate the overall integrated biohydrogen production process along with the
Rhodopseudomonas palustris (Mishra et al., 2019). Pandey et al., (2021) rate and yield of H2 (Li et al., 2018; Islam et al., 2021). However, this
investigated the efficiency of Spirulinaplatensis hydrolysateas substrate research area requires much attention in form of more intense studies for
in DF process mediated by Bacillus firmus NMBL-03 and produced 23.06 the better exposure at practical ground. Nevertheless, till date limited
± 3.63 mmol of H2 with yield of 1.92 ± 0.20 mmol H2/g COD reduction. studies have been devoted on integrated biohydrogen production pro­
Whereas, in the second stage of PF process, 1510 ± 35 mL/L H2 was cess using lignocellulosics biomass [Table 2].
recorded using Rhodobacter sphaeroides NMBL-01 under the optimum
conditions (Pandey et al., 2021). 5.1.2. Nanocatalyst approach
Nowadays, application of nanomaterials is considered as a potential
approach to improve the overall biomass to biohydrogen production
5.1. Key factors influencing the integrated biohydrogen production: process when employed as catalyst or co-factor. The unique physico­
Future prospects chemical properties of nanomaterials along with feasibility to recycle
and long lasting impact can elevate every steps of the biohydrogen
5.1.1. Substrate production process (Bhatia et al., 2021; Patel et al., 2018; Kwona et al.,
Substrate and its composition play a key role to improve the inte­ 2020). In general, use of nanomaterials can improve the efficiency of the
grated biohydrogen production process, especially in case of single stage chemical pretreatment of biomass, enzyme production and stability in
sequential integrated method (Kuppam et al., 2017). For the enhance­ the enzymatic hydrolysis, speed up the fermentation process, product
ment of biohydrogen yield, initial substrate should be highly rich of

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T. Singh et al. Bioresource Technology 338 (2021) 125511

Table 3
Studies on the effect of nanocatalysts to improve the biohydrogen production using different types of lignocellulosic biomass.
Sr. no Mode of fermentation Nanomaterials Substrates Hydrogen Production References

1. Photo fermentation SnO2 Corn Stover 345 mL Nadeem et al., 2020

2. Dark Fermentation Fe3O4@SiO2chitosan Sweet Sorghum stover 2.8 molH2/mol substrate Shanmugam et al., 2020
3. Dark fermentation Graphene oxide (GO) Sugarcane bagasse 2870 mL/L Srivastava et al., 2018
4. Dark Fermentation Fe3O4 NPs (magnetite) Sugarcane bagasse 0.874 mol(mol glucose-1) Reddy et al., 2017
5. Dark Fermentation TiO2 NPs Sugarcane bagasse 101.5 mL/g VS Jafari and Zilouei, 2016
6. Dark Fermentation Fe2O3 NPs Cassava Starch 192.4 mL/g Lin et al., 2016
7. Dark fermentation NiO NPs and Fe2O3 NPs Molasses 8.83 mmol/g COD Gadhe et al., 2015
8. Dark Fermentation Fe3O4/Alginate nanocomposite Rice Straw 2580 mL/L Srivastava et al., 2018

recovery and microorganism growth, and for these reasons use of nano- production significantly (Song et al., 2017; Patel et al., 2018). Moreover,
additive can be considered as advanced area of research (Kayfeci et al., genetic manipulation in H2 producing enzymes in both the processes (DF
2019; Kuppam et al., 2017; Mishra et al., 2019). Moreover, utilization of and PF) like knock out technology is very commonly used method to
nanomaterials such as, Fe2O3, Fe3O4, NiCo2O4, CuO, NiO, CoO, ZnO enhance the biohydrogen production yield (Peters et al., 2015). Addi­
have been confirmed to improve the biohydrogen production since they tionally, these tools should be used separately for both the processes
function as a co-factor of enzymes that help to enhance the efficiency of before performing the integrated technique. Wu et al. (2018), recorded
the enzymes involved in both DF and PF process (Srivastava, et al., improved H2 production by improving gene expression of NiFe-
2019). For an example, Fe is the key co-factor to influence the enzymatic hydrogenase in Pyrococcusfuriosus. These authors modified three new
activity of hydrogenase enzyme in DF whereas nitrogenase enzyme in strains for the determination of the specific hydrogen production ac­
case of PF process, as well as they are an important nutrient element of tivity by reducing methylviologen in form of artificial electron donor.
the microbial fermentative media for both the process (Teng et al., 2019; Based on the obtained results, it was concluded that the specific activity
Taherdanak et al., 2016). Further, Fe-based nanomaterials may be of the hydrogen production was 1.14-fold higher in case of genetically
exploited for multipurpose in the complete process of biomass to bio­ manipulated strain as compared to control (Wu et al., 2018). An incre­
hydrogen production following the integrated fermentation. Addition­ ment in 2-fold specific hydrogen production rate with unaffected oxygen
ally, being the catalyst these nanomaterials cannot be spent in the tolerance were reported in site-directed mutagenesis of [NiFe]-
process and hence are required in very less amount for the long term hydrogenase of E. coli (Flanagan et al., 2016). Further, a number of
utilizations with no toxic effect in the fermentation medium under the studies are available on improving the H2 production, employed genetic
optimal concentration and also participate to improve the metabolite modification in the size of light harvesting antenna of the photosynthetic
reactions of microorganisms (Jayasekara and Ratnayake, 2019). apparatus in case of photo-fermentation process (Wu et al., 2018; Song
Further, recycling of the used nanomaterials at each step in the process et al., 2017; Patel et al., 2018). Additionally, genetic manipulation in
are possible for their re-utilization again and again at pilot scale which nitrogenase enzymes is also reported in photo-fermentative H2 produc­
may offer an economic path to achieve its pilot-scale production goal. tion either by suppressing, introducing new or over expressing existing
Concepts of nanomaterials on influencing the DF and PF mediated bio­ genes (SundaraSekar et al., 2017). Zhang et al. (2018), conducted a pilot
hydrogen production have been reported in many studies (Trchounian scale study on sequential integrated biohydrogen production using 11
et al., 2017; Zhang et al., 2015; Preethi et al., 2019; Srivastava et al., m3 reactor in which 3 m3 volume was covered by dark whereas 8 m3 by
2019). Zhang et al. (2015) studied the effect of zero-valent iron acti­ photo compartments. These authors performed dark fermentation at pH
vated carbon (ZVIAC) micro-electrolysis system following dark 4.5 and 35 ◦ C whereas photo fermentation at pH 7.0 and 30 ◦ C using
fermentative H2production using glucose and the mixed bacterial con­ corn stover employed as substrate. It was noticed that out of the 87.8 ±
sortium. These authors observed positive results with the addition of ZVI 3.8 m3/d produced biogas, dark fermentation unit contributed 68%.
that helped to obtain ~ 38.2% more hydrogen yield, and ZVI-AC micro- However, challenges like poor mixing, lack of homogeneity were asso­
electrolysis could further enhance the hydrogen yield (improving rate ciated with the large-scale operational condition, caused large variation
50.2％). In a recent study, Zhang et al., 2021 evaluated DF process using in hydrogen production rate in case of dark and photo-fermentation. In
nickel ferrite nanoparticles and maximum H2 yield of 222 and 130 mL/g this context, for the industrial application point of view focus should be
of glucose were recorded in presence of 200 mg/L NiFe2O4 NPs (55 ◦ C) given on the selection of both types of fermentative microorganisms
groups, respectively, which were much higher than control. It was (responsible for dark/photo) which may capable to work on either
concluded that NiFe2O4 NPs might functioned as carriers in the intra­ similar or closer pH with efficient H2 production proficiency in order to
cellular electron transferring during DF mediated H2 production (Zhang avoid pH shifting, wastage of resource utilization at large-scale. Further,
et al., 2021). Henceforth, integration of DF and PF while implementing H2 production can be significantly enhanced by reducing or eliminating
nanomaterials as catalyst can be more effective and sustainable way in the various competing biochemical pathways following the integrated
the hybrid stage system for improving the process economy of the H2 biohydrogen production technology using lignocellulosics biomass as
production using lignocellulosic biomass (Mishra et al., 2019; Nem­ potential substrate. Nevertheless, rigorous studies are needed to be
estóthy et al., 2020). Table 3 also summarizes various studies on the devoted to find out novel strategies to further achieve high H2 produc­
applications of nanomaterials for enhanced biohydrogen production tion rate and yield using lignocellulosic biomass for its practical appli­
using lignocellulosics biomass. cation in near future [Fig. 3].

5.1.3. Genetic engineering 6. Conclusion

Genetic engineering is always considered as booming technology for
improving the dark and photo fermentative H2 production process. This review has been devoted on the development of integrated
Metabolic manipulation in microorganisms for both the processes are biohydrogen production process as an advanced biochemical technology
the most innovative tool, and are being employed from past many years for the economical production of H2 using lignocellulosic biomass. It
for enhancing the H2 production yield (Preethi et al., 2019). Elimination was noticed that in spite of a number of advantages, very limited
of H2 uptake gene, modification of homologous or heterologous gene, research studies were performed and therefore this technology is needed
development of synthetic metabolic reactions and association of various to be more explored. This review emphasized on numerous aspects like
methods are frequently used approaches which may enhance the H2 potential utilizations of lignocellulosic biomass as low-cost substrate,

8
T. Singh et al. Bioresource Technology 338 (2021) 125511

Fig. 3. Schematic diagram shows strategy to improve biohydrogen production from lignocellulosic biomass following integrated fermentation.

implementation of nanocatalyst and genetic manipulation as the Al-Battashi, H. S., Annamalai, N., Sivakumar, N., Al-Bahry, S., Tripathi, B.N., Nguyen, Q.
D., Gupta, V.K., 2019. Lignocellulosic biomass (LCB): a potential alternative
possible approaches to elevate the integrated technology of biohydrogen
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production in a sustainable way for its practical applications in near SciBiotechnol 18, 183-205.
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hydrolysis of oat straw and hydrogen production from hydrolysates: Role of
hydrolysates constituents. Int. J. Hydrogen Energy 40 (34), 10756–10765.
CRediT authorship contribution statement Assawamongkholsiri, T., Reungsang, A., and Sittijunda, S., 2019 Photo-hydrogen and
lipid production from lactate, acetate, butyrate, and sugar manufacturing
wastewater with an alternative nitrogen source by Rhodobacter sp. KKU-PS1.
Tripti Singh, Alaa Alhazmi, Akbar Mohammad, Neha Srivastava: PeerJ. 2019; 7: e6653.
Conceptualization, curation, formal analysis, writing original draft. Baruah, J., Nath, B. K., Sharma, R., Kumar, S., Deka, R. C., Baruah, D. C., Kalita, E., 2018.
Shafiul Haque, Shalini Sharma, Rajeev Singh, Taeho Yoon: Inves­ Recent Trends in the Pretreatment of Lignocellulosic Biomass for Value-Added
Products.Front. Energy Res., 6.
tigation methodology, validation, review editing. Vijai Kumar Gupta: Baumann, I., Westermann, P., 2016. Microbial production of short chain fatty acids from
Supervision, Writing - review & editing. lignocellulosic biomass: Current processes and market. BioMed Res. Int 1–15.
Ben Iwo, J., Manovic, V., Longhurst, P., 2016. Biomass resources and biofuels potential
for the production of transportation fuels in Nigeria. Renew Sustain Energy Rev. 63,
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Declaration of Competing Interest Beschkov, V., 2017.Biogas, Biodiesel and Bioethanol as Multifunctional Renewable Fuels
and Raw Materials. Chapter 10.
The authors declare that they have no known competing financial Bhatia, S.K., Jagtap, S.S., Bedekar, A.A., Bhatia, R.K., Rajendran, K., Pugazhendhi, R.,
Rao, C.V., Atabani, A.E., Kumar, G., Yang, Y.H., 2021. Renewable biohydrogen
interests or personal relationships that could have appeared to influence production from lignocellulosic biomass using fermentation and integration of
the work reported in this paper. systems with other energy generation technologies. Sci. Total Environ. 765, 144429.
Billaud, J., Valin, S., Peyrot, M., Salvador, S., 2016. Influence of H2O, CO2 and O-2
addition on biomass gasification in entrained flow reactor conditions: Experiments
Acknowledgements and modelling. Fuel, Elsevier 166, 166–178.
Bundhoo, M.A.Z., Mohee, R., 2016. Inhibition of dark fermentative bio-hydrogen
Author N.S. thankfully acknowledges Department of Chemical En­ production: A review. Int. J. Hydrog. Energy 41 (16), 6713–6733.
Busic, A., Kundas, S., Morzak, G., Belskaya, H., Mardetko, N., Santek, M.I., Komes, D.,
gineering and Technology, IIT (BHU) Varanasi, India. V.K.G. would like Novak, S., Santek, B., 2018. Recent trends in biodiesel and biogas production. Food
to acknowledge the institutional research funding supported by the Technol. Biotechnol. 56 (2), 152–173.
Scotland’s Rural College (SRUC), UK. Cabrol, L., Marone, A., Tapia-Venegas, E., Steyer, J.-P., Ruiz-Filippi, G., Trably, E., 2017.
Microbial ecology of fermentative hydrogen producing bioprocesses: useful insights
for driving the ecosystem function. FEMS Microbiol. Rev. 41 (2), 158–181.
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