1 s2.0 S0960852422016650 Main

Bioresource Technology 368 (2023) 128332
Contents lists available at ScienceDirect
Bioresource Technology
journal homepage: www.elsevier.com/locate/biortech
Review
Algal biomass to biohydrogen: Pretreatment, influencing factors, and

conversion strategies
Shashi Kant Bhatia a, b, J. Rajesh Banu c, Vijai Singh d, Gopalakrishnan Kumar e, f,
Yung-Hun Yang a, b, *
a
Department of Biological Engineering, College of Engineering, Konkuk University, Seoul 05029, Republic of Korea
b
Institute for Ubiquitous Information Technology and Applications, Seoul 05029, Republic of Korea
c
Department of Biotechnology, Central University of Tamil Nadu, Neelakudi, Thiruvarur 610005, India
d
Department of Biosciences, School of Science, Indrashil University, Rajpur, Mehsana 382715, Gujarat, India
e
Institute of Chemistry, Bioscience and Environmental Engineering, Faculty of Science and Technology, University of Stavanger, Box 8600 Forus, 4036 Stavanger,
Norway
f
School of Civil and Environmental Engineering, Yonsei University, Seoul, 03722, Republic of Korea
H I G H L I G H T S G R A P H I C A L A B S T R A C T
• Algae capture CO2 and produce

biomass.
• Algal biomass requires mild pretreat
ment to produce bio-sugars.
• Algal hydrolysate pretreatment results
in less side products.
• It can be used as feedstock for hydrogen
production in dark fermentation.
• Dark fermentation process can be
coupled with photo fermentation and
methanogenesis.
A R T I C L E I N F O A B S T R A C T
Keywords: Hydrogen has gained attention as an alternative source of energy because of its non-polluting nature as on
Algal biomass combustion it produces only water. Biological methods are eco-friendly and have benefits in waste management
CO2 fixation and hydrogen production simultaneously. The use of algal biomass as feedstock in dark fermentation is ad
Dark fermentation
vantageous because of its low lignin content, high growth rate, and carbon-fixation ability. The major bottlenecks
Biohydrogen
Pretreatment
in biohydrogen production are its low productivity and high production costs. To overcome these issues, many
advances in the area of biomass pretreatment to increase sugar release, understanding of algal biomass
composition, and development of fermentation strategies for the complete recovery of nutrients are ongoing.
Recently, mixed substrate fermentation, multistep fermentation, and the use of nanocatalysts to improve
hydrogen production have increased. This review article evaluates the current progress in algal biomass pre
treatment, key factors, and possible solutions for increasing hydrogen production.
* Corresponding author at: Department of Biological Engineering, College of Engineering, Konkuk University, Seoul 05029, Republic of Korea.
E-mail address: seokor@konkuk.ac.kr (Y.-H. Yang).
https://doi.org/10.1016/j.biortech.2022.128332
Received 27 September 2022; Received in revised form 8 November 2022; Accepted 11 November 2022
Available online 19 November 2022
0960-8524/© 2022 Elsevier Ltd. All rights reserved.
S.K. Bhatia et al. Bioresource Technology 368 (2023) 128332
1. Introduction hydrogen production potential as compared to protein and lipids.

Polysaccharide composition varies in algae, for example, green algae are
Fossil fuels (coal, oil, and natural gas) fulfill approximately 80 % of rich in xylan and mannan, which serve as a crystalline skeleton, red
the energy demand and will become extinct in the next few decades (e. algae are rich in agar and carrageenan, and brown algae are composed of
g., oil in 40 years, natural gas in 60 years, and coal in less than 250 alginate, laminarin, and mannitol (Jung et al., 2011). Laminaria japonica
years) (Ji & Wang, 2021). The consumption of fossil fuels emits green (brown algae) is the most widely studied algae for hydrogen production
house gases (GHG) and is a major cause of global warming (Narisetty because of its high and easily digestible carbohydrate content. Hydrogen
et al., 2022). Under the United Nations Framework Convention on production is also affected by dark fermentation conditions, such as
Climate Change (UNFCCC), around 185 countries promised to limit inoculum to substrate ratio, pH, temperature, organic loading rate
global temperatures to 2 ◦ C less than before the industrialized period (OLR), and hydraulic retention time (HRT). To improve hydrogen pro
(King et al., 2020). To accomplish this aim, European Union (EU) has duction, researchers are working on different strategies, such as mixing
started a “20–20–20” project with the purpose of GHG emission reduc algal biomass with other feedstocks, use of additional materials, two-
tion (20 %), reducing fossil fuel consumption (20 %), and increasing step (dark and photofermentation/anaerobic digestion), and three-step
renewable energy utilization (20 %) (Hong & Wu, 2020). The use of fermentation (dark, photo, and anaerobic digestion).
renewable energy resources can help address the depletion of fossil- This review article focuses on the conversion of algal biomass to
based fuels and CO2 emissions (Hoang et al., 2022; Sekar et al., 2021). biohydrogen using dark fermentation. This review provides an overview
There are different renewable energy sources such as solar, wind, hydro, of recent advances in algal biomass pretreatment technologies, factors
geothermal, biogas, etc. have been explored to fulfill the energy de that affect dark fermentation, and different strategies reported to
mands. Most of these energy sources are sporadic and limit their use in a improve hydrogen production from the algal feedstock.
certain region and time frame (Bhatia et al., 2021). Another drawback is
the lack of government regulations and policies favoring the develop 2. Algal biomass as feedstock for biohydrogen
ment and adaptation of these technologies. Hydrogen is gaining atten
tion worldwide to fulfill the energy demand in the residential, industrial, The selection of the right feedstock for industrial-scale production of
and transportation sectors because of its zero-carbon characteristics and commodities is important as almost 60–70 % of the cost is due to raw
high energy content compared to fossil-based fuels (Preethi et al., 2022; materials. Furthermore, due to food security issues, feedstock for bio
Singh et al., 2022). The low heating value of hydrogen (120 MJ/kg) is energy production has shifted from the first (sugar and starch) to the
superior as compared to other fuels such as methane (50.0 MJ/kg), second generation (lignocellulosic material) and then to the third gen
gasoline (44.5 MJ/kg), and diesel (42.5 MJ/kg) (Enayatizade et al., eration (algae) (Bhatia et al., 2017). Lignocellulose is an abundantly
2019). Most hydrogen (95 %) is obtained from fossil fuels and releases available complex material composed of cellulose, lignocellulose, and
approximately 830 million tons of CO2 annually (Franchi et al., 2020). lignin as the main components (Awasthi et al., 2022). The crystalline
To overcome these issues, researchers are working on biological structure restricts the enzyme’s access to cellulose and liberates sugars,
methods of hydrogen production, such as photo fermentation, anaerobic which are the main carbon sources for microbial fermentation.
digestion, dark fermentation, and microbial electrolysis (Godvin Shar Furthermore, microbes are unable to utilize lignin, and aromatic com
mila et al., 2022; Kumar et al., 2022). Dark fermentation is the preferred pounds inhibit microbial growth and enzyme activity. Land availability
approach for hydrogen production because it does not require any light, also prevents the cultivation of energy crops. Algae are photosynthetic
has a high hydrogen production rate, and can utilize a broad range of organisms that convert CO2, water, and light into biomass, and have
feedstocks (Martins et al., 2022). A variety of raw materials such as agro- many advantages over conventional feedstocks (Table 1) (Wang & Yin,
industry waste, wastewater, olive mill waste, corn steep liquor, and 2018). Microalgae can also be used for wastewater treatment and to
cheese whey have been utilized for hydrogen production (Lovato et al., produce biomass that is rich in organic content and can be further uti
2021; Luo et al., 2022; Martinez-Burgos et al., 2021; Mugnai et al., lized as a feedstock for bioenergy and biochemical production (Chandel
2021). Lignocellulosic-based feedstocks are recalcitrant and require et al., 2022; Goswami et al., 2022). The advantages of algal biomass over
harsh pretreatment to release free sugars, which leads to the formation
of many side products that affect microbial growth and hydrogen pro
ductivity (Bhatia et al., 2021). Table 1
Algae (microalgae and macroalgae) are considered futuristic Algal biomass advantages and disadvantages as a feedstock for biohydrogen
renewable feedstocks for hydrogen production because of their superior production.
CO2 capture capacity, high growth rate, ability to accumulate high Algal feedstock advantages Algal feedstock disadvantages
carbohydrate content, and milder pretreatment due to low lignin con
• High rate of photosynthesis and 10–50 • Algal cultivation requires temperature
tent (Bhatia et al., 2022b; Pugazhendi et al., 2022). Approximately times growth compared to plants. maintenance and it’s hard to maintain
20,000 microalgal species and 10,000 macroalgae have been identified. at a large scale.
Algae also vary in composition, which is affected by various environ • Ability to utilize CO2 and water for the • Algae are prone to microbial
mental factors, such as temperature, light intensity, salt concentration, synthesis of biomass. contamination.
• No need for fertile land and can be • Require high content of nutrients and
and nutrient availability (Ganesh Saratale et al., 2018). Algae can be grown in unproductive land. water for cultivation.
used to produce a variety of commercially valuable products, and after • Most of the algae can tolerate high • Have large water content in biomass
extraction of the desired product, the residual biomass can be used for concentrations of CO2, SOx, and NOx. and the drying process is energy-
hydrogen production (Mehariya et al., 2022; Patel et al., 2021). The consuming.
• Algal biomass has high content of • Biomass concentration is low which
algal cell wall is rigid and requires disruption and different pretreatment
carbohydrates that support makes harvesting costly.
methods, including physical (autoclaving, sonication, and microwave), fermentation and increase H2
chemical (acidic and alkaline), and biological (enzymatic) methods productivity.
(Bhatia et al., 2022a). Among the various pretreatment methods, acid- • Algae lack lignin and require milder
–thermal treatment is the most effective for reducing sugar release. In pretreatments to release the
fermentable sugar.
dividual pretreatments of Chlorella pyrenoidosa algal biomass (steam • Algae cultivation doesn’t require any
heating or microwave) can produce hydrogen by 8.8–12.7 mL/g total herbicides or pesticides.
volatile solids (TVS), while combined pretreatment with steam/micro • Algal biomass can be used to produce
wave and diluted HCl increased hydrogen production to 75.6–83.3 mL/ many commercially valuable
compounds.
g.TVS (Xia et al., 2013). It is well reported that carbohydrate has higher
2
lignocellulosic biomass for hydrogen production are presented in Closed system: Closed systems or photobioreactors (PBR) are vessels
Table 1. made of transparent materials. Different PBR systems have been devel
The main components of algal biomass include proteins (40–60 %), oped for microalgal cultivation, such as vertical bubble columns, airlift
carbohydrates (20–30 %), and lipids (10–20 %), with minor contents of reactors, and tubular and helical photobioreactors (Shekh et al., 2022).
nucleic acids and pigments (Wang & Yin, 2018). Algal cells are Closed systems can operate under controlled conditions and are less
composed of a variety of complex polysaccharides, such as cellulose, prone to contamination. The main advantages of PBR include low har
starch, glycogen, and agarose. The composition may vary with species vesting cost, high productivity, and lower water and CO2 losses (Egbo
and is affected by cultivation conditions. Microalgae are known for their et al., 2018). The operation and maintenance costs of these types of
high lipid content, which can be increased up to 80 % by adjusting systems are high compared to those of open systems, and they are not
nutrient conditions. Under nitrogen limitation and high-light condi preferred for large-scale cultivation. Schädler et al., (2020) developed a
tions, microalgal cells prefer to produce lipids instead of starch (di Cicco high-density microalgae cultivation system using a thin-layer cascade
et al., 2021). The composition of the various algae is shown in Table 2. photobioreactor and reported that water recycling has no adverse effect
on microalgal growth if all the necessary nutrients are replenished and
3. Algae cultivation urea is employed as a nitrogen source. This process led to 30 g/L of
Microchloropsis salina production, thereby reducing nutrient consump
3.1. Microalgae tion and wastewater production.
More than 50,000 microalgae species exist, only a few of which are 3.2. Macroalgae
suitable for cultivation. Two approaches are used for microalgae culti
vation: (i) open pond systems and (ii) closed systems. Macroalgae are multicellular photosynthetic organisms found in
Open pond: The open pond system (ponds, lakes, and artificial bodies marine environments such as oceans and coastal areas. Approximately
of water) is the oldest and most widely used algal cultivation method, 10,000 macroalgal species exist and are classified as green, red, or
owing to its cheaper construction, maintenance, and operating costs. brown based on their pigments (Gordalina et al., 2021). Macroalgae are
The main challenge in an open pond system is to control growth pa composed of proteins, lipids, and polysaccharides, which are catego
rameters (temperature, light intensity), which affect growth and pro rized as storage and structural polysaccharides (Jönsson et al., 2020).
ductivity (Tan et al., 2020). Runoff water also affects the pH and Different types of systems can be used for the cultivation of macroalgae,
turbidity of open ponds and productivity. In addition, open ponds are such as marine aquaculture, land-based tanks, and closed photo
susceptible to contamination by undesired microalgae and microor bioreactor. Marine aquaculture lacks a control system and productivity
ganisms (Lam et al., 2018). The selection of suitable microalgae is depends on meteorological and oceanographic conditions (Buck &
important for addressing these challenges. Microalgae related to the Grote, 2018). The primary factors include climate change, ocean acidi
genus Galdieria are extremophilic, adapted to acidic hot springs, and fication, microbial proliferation, and herbivores. These drawbacks can
able to remove heavy metals from wastewater (di Cicco et al., 2021). be controlled in land-based systems, where these parameters can be
Model predictive control (MPC) can be used to predict algal yield, controlled to a certain extent (Schmitz & Kraft, 2022). Marine- and land-
depending on the species, weather conditions, and system design. De- based systems have to compete with fisheries, tourism, and water ac
Luca et al. (2019) used a meteorological data-based optimal control data tivities. Photobioreactors allow cultivation under controlled conditions
method for microalgae cultivation in an open pond and were able to but scaling up to an industrial scale is the main challenge. For the past
increase productivity by 2 times with a constant pond depth and dilution 50 years, many macroalgae have been investigated in field farms, but
rate. only a dozen are cultivated commercially today, such as Gracilaria,
Table 2
Composition of different algae and their CO2 tolerance and fixation rate.
Algal species CO2 tolerance CO2 fixation rate (g/L. Carbohydrate Lipids (%) Protein (%) References
(%) d) (%)
Microalgae
Anabaena 15 1.01 29.4–37.5 4.7–8.8 53.8–65.9 (Chiang et al., 2011; Xia et al., 2015)
Chlorella 15–50 2.25 14.0–43.5 0.8–63.2 22.8–69.0 (Park et al., 2021; Xia et al., 2015)
Chlamydomonas 15 – 11–38.4 15–22.2 33.4–59 (Verspreet et al., 2021; Xu et al., 2018)
Dunaliella 15 0.272 20–20.99 26–32 30–33.41 (Huo et al., 2011; Singh & Ahluwalia, 2013)
Euglena 45 15.7–23.4 15.7–26.0 50.6–68.5 (Singh & Ahluwalia, 2013; Xia et al., 2015)
Oocystis – – 33 17 54 (Ajala & Alexander, 2020)
Porphyridium 15 – 51.2 13.2 22.2 (Verspreet et al., 2021)
Nannochloropsis 10 1.7 10.2 6.20 45.6 (He et al., 2020)
Scenedesmus 18 1.84 23–38 13–17 54–65 (Ajala & Alexander, 2020; Garcia et al., 2015; Tang et al.,
2011)
Spirulina 5 0.318 25 18 70–55 (Anvar & Nowruzi, 2021; Xia et al., 2015)
Tetraselmis 14 0.270 10.8–52.2 4.3–22.5 31.1–74.3 (Schipper et al., 2019; Viswanaathan et al., 2022; Xia et al.,
2015)
Macroalgae
Codium – – 32.3 1.8 10.9 (Milledge et al., 2019)
Ecklonia 48.6 2.4 13.6 (Milledge et al., 2019)
Gelidium – – 79.24–86.78 0–1.2 2.22–3.46 (Panggabean et al., 2022)
Gracilaria – – 33.5 3.2 15.6 (Milledge et al., 2019)
Hizikia – – 28.6 0.4–1.5 5.9–13.9 (Milledge et al., 2019)
Laminaria – – 54.6 8.4 1.6 (Panggabean et al., 2022)
Porphyra – – 35.9 4.4 38.7 (Milledge et al., 2019)
Spirogyra sp. – – 27 14.2 23.2 (Pinto et al., 2018)
Ulva – – 55.40 6.67 4.24 (Panggabean et al., 2022)
Undaria – – 40.1 2.0 15.9 (Milledge et al., 2019)
3
Eucheuma, Laminaria, Porphyra, and Undaria. Under the European Blue byproduct, and 2 mol of hydrogen was produced when butyrate was
Development Initiative, many countries have been working on the produced (Chandrasekhar et al., 2015).
commercial cultivation of macroalgae such as Furcellaria lumbricalis in
Estonia, Saccharina latissimi in Sweden, Ulva in Poland, and Fucus ser i) Acetic acid pathway C6H12O6 + 2H2O → 2CH3COOH + 2CO2 +
ratus in Germany (Armoškaitė et al., 2021). Schmitz & Kraft (2022) 4H2
developed a photobioreactor for macroalgae Ulva intestinalis cultivation ii) Butyric acid pathway C6H12O6 + 2H2O → CH3CH2CH2COOH +
and were able to achieve fresh biomass of 3.0 kg/m2. 2CO2 + 2H2
iii) Propionic acid pathway C6H12O6 + 2H2 → 2CH3CH2COOH +
4. Algal biomass conversion to biohydrogen by dark 2H2O
fermentation iv) Ethanol pathway C6H12O6 → CH3CH2OH + CO2
Dark fermentation is a biological method of hydrogen production Algal biomass is deemed an alluring feedstock for biohydrogen
carried out by obligate anaerobes (Clostridium, Ethanoligenes, Desulfovi production because of the high content of carbohydrates and the pre
brio sp., etc.) and facultative anaerobes (Enterobacter, Citrobacter, Kleb requisite of mild pretreatment to release fermentable sugars. Microbes
siella, E. coli, Bacillus sp., etc.) in the absence of light (Pugazhendhi et al., are not able to utilize algal biomass directly as sugars are present in a
2019). Various organic wastes (lignocellulosic biomass, algal biomass, polymeric form in algae and require pretreatment to convert them into
wastewater, municipal waste, etc.) can be utilized as substrates for dark fermentable sugars. Various methods have been reported for algal
fermentation to produce biohydrogen (Bhatia et al., 2019; Dange et al., biomass pretreatment and hydrogen production, as discussed in the next
2021). Organic polymer hydrolysis is the first step, followed by aceto section, and are presented in Table 3.
genesis, which leads to the production of organic acids and alcohols with
the release of hydrogen. Hydrogenases are the main enzymes involved in 4.1. Algal biomass pretreatment and biohydrogen production
hydrogen production and are categorized as [Ni-Fe] hydrogenases and
[Fe-Fe] hydrogenases (Peters et al., 2015). Pyruvate, formed during The algal cell wall is rigid and requires pretreatment (acidic, basic,
carbohydrate metabolism, is a major source of hydrogen production enzymatic, thermal, and combined methods) to release free sugars. No
(Fig. 1). Pyruvate is oxidized into acetyl-CoA, which is further converted single method is universally applicable as algal characteristics are
to formate by pyruvate formatase (PFL) in facultative microbes. Formate diverse, and pretreatment conditions vary from strain to strain.
is the main product of facultative microbes and is catalyzed into CO2 and
H2 by [Ni-Fe] hydrogenase (Cao et al., 2022). In obligate anaerobes, 4.1.1. Physical pretreatment and biohydrogen production
acetyl-CoA is converted to ferredoxin by pyruvate ferredoxin oxidore Singh et al. (2022) investigated different physical methods (soni
ductase (PFOR). [Fe-Fe]-hydrogenase catalyzes the re-oxidation of cation, grinding, homogenization, and autoclaving) to extract ferment
ferredoxin and produces hydrogen (Cao et al., 2022). In the dark able sugars from deoiled Scenedesmus obliquus biomass. The sonication
fermentation process, the low yield of hydrogen is the main challenge, pre-treatment method led to a maximum carbohydrate recovery of
and most organic acids remain unutilized. Hydrogen yield depends on 33.4 %, followed by autoclaving (27 %), grinding (22 %), and homog
the type of organic acid produced during the fermentation process. enization (20 %). The poor release of carbohydrates is due to tri-laminar
Theoretically, 1 mol of glucose can produce 12 mol of hydrogen; how cell morphology, which confers high resistance to cells. The use of un
ever, researchers are not able to achieve complete oxidation of glucose treated deoiled S. obliquus biomass in dark fermentation led to 10 mL/g
and can produce only 1–4 mol H2/mol of glucose (Patel et al., 2018). A volatile solids (VS) biohydrogen indicating that recalcitrant components
maximum of 4 mol of hydrogen was produced when acetic acid was the present in cells hinder the release of sugars required for microbial
growth. Physical pretreatment improved hydrogen production to a
certain extent, and different H2 yields were recorded with various pre
treatments, for example, sonication (36.2 mL/g.VS), autoclaving (30.4
mL/g.VS), homogenization (19.6 mL/g.VS), and grinding (15.1 mL/g.
VS) (Singh et al., 2022). Batista et al. used the autoclaving method for
sterilization and sugar extraction from S. obliquus. The hydrolysate was
used for dark fermentation with Enterobacter aerogenes, and the process
stemmed in hydrogen at 40.9 mL/g algae (Batista et al., 2018). Kumar
et al. explored different pretreatment methods (autoclave, ultra
sonication, electrolysis, etc.) for mixed microalgae (Scenedesmus and
Chlorella) from the natural ecological niche. The electrolysis method was
reported to be the most effective method compared to the other tested
methods, with the highest total chemical oxygen demand (t-COD)
release (20 % increase) of 236 mL/L.d hydrogen production (Kumar
et al., 2016). The optimum temperature and time for the hydrothermal
process of algal biomass pretreatment were reported as 135–140 ◦ C and
15–20 min respectively. When the pretreatment temperature was below
135 ◦ C and the time was less than 15 min, most of the recalcitrant
components, such as cellulose, were not hydrolyzed efficiently to
smaller sugar molecules. The application of temperatures above 140 ◦ C
for 20 min leads to the Maillard reaction between reducing sugars and
proteins, resulting in low free sugar release (Cheng et al., 2019). The
microwave method is a widely used pre-treatment technique owing to its
high efficiency and ease of use. Yin et al. (2019) subjected L. japonica to
microwave pretreatment at temperatures of 100–180 ◦ C and reported an
increase in disruption with an increase in temperature. The highest
hydrogen production of 15.8 mL/g total solids (TS) was recorded in the
Fig. 1. Fermentative pathway for biohydrogen production. biomass treated at 160 ◦ C (Yin et al., 2019). The use of gamma
4
Table 3
Algal biomass pretreatment strategies and its dark fermentation to produce biohydrogen.
Microalgae Pretreatment Fermentative microbes Comments Biohydrogen Reference
production
Microalgae
Anabaena sp. Heat-HCl pretreatment (2 % v/v Mixed consortia of Pretreatment leads to the production of 1230 mL/L (Nayak et al.,
HCl, 100 ◦ C, 1 h) thermophilic bacteria 6.5 g/L glucose with furfural as the main 2014)
side product
Autoclaving (121 ◦ C, 1 atm for Mixed consortia of Pretreatment leads to 4.2 g/L glucose 600 mL/L (Nayak et al.,
20 min) thermophilic bacteria 2014)
Chlorella fusca Enzymatic treatment (cellulase Rhodobacter sp. The combined use of enzymes increases 2350 mL/L (Abdel-Kader
and maceroenzyme) at room biomass hydrolysis. (cumulative) et al., 2022)
temperature
Sonicated yeast enzyme pool Rhodobacter sp. Enzymes released by sonicated yeast 4200 mL/L (Abdel-Kader
hydrolyze Chlorella mats to produce et al., 2022)
sugar
Chlorella pyrenoidosa Steam heating and dilute acid Microbes isolated from sludge Without pretreatment only 8.8 mL/g. 198.3 mL/g.TVS (Xia et al.,
TVS hydrogen production was observed 2013)
Microwave heating and dilute 83.3 mL/g. TVS
acid
Chlorella pyrenoidosa Steam-dilute H2SO4 Anaerobic digestate Cassava starch was mixed to fix C/N 276.2 mL/g.TVS (Xia et al.,
ratio 25.3. Three step process was used. 2014)
Chlorella vulgaris Thermal treatment 100 ◦ C for 60 Digested sludge A substrate/inoculum ration of 8 leads to 190.90 mL H2/g. (Stanislaus
min higher H2 production. VS et al., 2018)
Enzymatic pretreatment Wastewater treatment plant The enzymatic treatment effectively 135 mL H2/g.VS (Wieczorek
(Onozuka R-10 and Macerozyme sludge enhanced H2 production by 9.1 fold et al., 2014)
R-10)
Dunaliella primolecta 6 M HCl and boiling Thermococcus eurythermalis Deoiled algal biomass was used as a 192.35 mL/g.VS (Chen et al.,
A501 feedstock. 2020b)
Nannochloropsis Microwave Mixed culture Three-stage fermentation i.e., dark, 183.9 mL/g.VS (Xia et al.,
oceanica photo, and anaerobic digestion 2013)
Oscillatoria angustissmia 0.5 % (V/V) H2SO4 Brevibacillus invocatus SAR Iron addition enhanced hydrogenase 3.8 mol H2/mole (Gad El-Rab
and Synchocystis sp activity and accumulative hydrogen of sugar et al., 2018)
production
Scenedesmus obliquus Autoclave Enterobacter aerogenes ATCC Facultative anaerobe requires low 57.6 mL/g.VS (Batista et al.,
1304 maintenance and operating cost 2014)
Autoclave Clostridium butyr- Require anoxic conditions for growth 113.1 mL/g.VS
icum DSM 10702 and hydrogen production
Scenedesmus Hydrothermal/HCl treatment (3 Granular sludge Biomass solubilization efficiency 48 mL/g.VS (Rincón-Pérez
obtusiusculus % HCl, 100 ◦ C, 1.7 h) depends on temperature and acid et al., 2020)
concentration and their interactions
Spirulina platensis 0.5 % H2SO4 Co-culture of Rhodobacter Two-stage fermentation strategy was 1510 mL/L (Pandey et al.,
sphaeroides NMBL-01 and used 2021)
Bacillus firmus NMBL-03
Macroalgae
Gelidium amansii 2 % w/v H2SO4 Sludge collected from the Activated carbon was used to detoxify 53.5 mL/g (Park et al.,
wastewater treatment plant. algal biomass hydrolysate 2011)
Thermal treatment, 170 ◦ C, 20 Anaerobic digestate from 34.2 mL/g.COD (Jung et al.,
min wastewater treatment plant – 2011)
Laminaria japonica 2 % NaOH, 121 ◦ C, 30 min Anaerobic sludge Cofermentation of sewage sludge and 14.8 mL /g.VS (Yin et al.,
algal biomass was performed and Fe2+ 2021)
was added
Drying and grinding Anaerobic sludge After dark fermentation digestate was 761.4 mL/g.TS (Shi et al.,
subjected to anaerobic digestion. 2011)
Spirogyra sp. Homogenization + bead beating Clostridium butyricum DSM Macronutrient stress was applied to 146.3 mL H2/g (Pinto et al.,
+ 1 N H2SO4 at 121 ◦ C for 1 h 10702 increase the carbohydrate content in 2018)
Spirogyra
Undaria pinnatifida Thermal treatment, 170 ◦ C, 20 Anaerobic digestate from 22.6 mL/g.COD (Jung et al.,
min wastewater treatment plant – 2011)
Ulva reticulate Microwave-H2O2 treatment Anaerobic sludge Alkaline conditions promote 87.5 mL H2/g. (Dinesh Kumar
decomposition of H2O2 to OH− and COD et al., 2020)
increase biomass solubilization
irradiation has also been described for the pretreatment of algal 4.1.2. Chemical pretreatment and biohydrogen production
biomass. Chen et al. (2022) pretreated L. japonica biomass by applying Different acids and bases have been used for the hydrolysis of algal
gamma irradiation of 10–30 kGy and effectively destroyed the algal cell biomass. Acidic environments break down interchain hydrogen bonds
structure, with maximum organic material release at 20 kGy and a 71.4 and cause swelling of the organic structure, making it more susceptible
% increase in hydrogen production. The use of gamma irradiation to hydrolysis (Giang et al., 2019). The efficiency depends on biomass
beyond 39 kGy causes the degradation of organic compounds and has a loading, treatment temperature, acid concentration, and hydrolysis
negative effect on hydrogen production (Chen et al., 2022). Ultrasonic time. Giang et al. (2019) collected Chlorella sp. biomass grown using
pretreatment is considered a powerful method for degradability; how coal-fixed flue gas from an industrial plant. Chlorella sp. biomass was
ever, it is energy intensive which restricts its application. It can be subjected to acidic treatment (HCl, H2SO4, HNO3, and H3PO4), and
combined with other pretreatment methods to improve hydrolysis by various parameters, such as acid and biomass concentrations and pre
reducing the production of inhibitors. treatment time, were optimized. The acid H2SO4-thermal treatment was
5
highly efficient in terms of reducing sugar release, followed by HCL and Dianchi Lake. Hydrothermal/acid treatment was performed in a hy
HNO3, while no sugar was produced in the case of H3PO4. An increase in drothermal reactor, and steam/acid treatment was performed in an
acid concentration, biomass loading, and pretreatment time led to an autoclave with 2 % H2SO4 for 15 min at 135 ◦ C. Hydrothermal and
increase in reducing sugar concentration, but an increase in inhibitor steam pretreatments significantly increased the sugar yield by hydro
concentration was also reported. The optimum conditions for maximum lyzing macromolecules. Hydrothermal treatment was more efficient
sugar release (151.8 mg/g biomass) from Chlorella sp. were 0.75 % v/v than steam treatment because of the higher pressure and rapid heat
H2SO4, 160 ◦ C, 30 min, and 40 g biomass/L. Pretreated biomass was transfer in smaller-volume hydrothermal reactors, which promote cell
subjected to anaerobic digestion, resulting in 1,079 mL/L of hydrogen disruption. Dark fermentation of hydrothermal and steam hydrolysate
(Giang et al., 2019). Sivagurunathan et. al. also investigated different with sludge collected from anaerobic digestate resulted in 24.96 mL/g.
dilute acids (HCl, H2SO4, HNO3, and H3PO4) for the treatment of Geli TVS and 18.63 mL/g.TVS hydrogen production, respectively (Cheng
dium amansii biomass and found that 1 % H2SO4 was most effective, et al., 2019). Yun et al. (2013) studied acid + ultrasonic combined
releasing 19.4 g/L of reducing sugar. The hydrolysate was subjected to pretreatment of microalgal biomass and used response surface design to
anaerobic digestion, and the process resulted in 52 mL/g dry biomass optimize optimum conditions. Combined pretreatment resulted in
hydrogen, which was almost 2 fold high than that of the control improved hydrolysis and fermentation leading to 42.1 mL/g hydrogen
experiment (Sivagurunathan et al., 2017). production (Yun et al., 2013). The combined treatment follows a
In alkali hydrolysis solubilisation of biomass occurs by reaction be different disruption mechanism: after acid treatment, algal cell walls
tween alkali reagents and biomass which includes saponification of become more vulnerable to shear stress generated by ultrasonication,
uronic acids and esters and neutralization of acid intermediates. which elevates disruption and makes the overall process more
Monovalent alkalis lead to better solubilisation as compared to divalent economical. The use of surfactants during ultrasonication has also been
alkalis since divalent ionize only partially in aqueous media (de Oliveira reported, as it helps to reduce the interfacial tension due to hydrophobic
et al., 2022). Yang et al., explored thermos-alkaline pretreatment for and hydrophilic impacts with an expansion in the cavitation bubble.
lipid-extracted algal biomass and its fermentation leads to 45.54 H2 mL/ Snehya et al., (2022) used surfactant-mediated ultrasonic pretreatment
g.VS (Yang et al., 2010). Yin et al., used 2 % NaOH for the pretreatment for Sargassum tennerimum collected from Rameswaram Beach, India, and
of S. japonica at room temperature and 121 ◦ C. High polysaccharides and reported 86 mL/g.COD hydrogen production.
protein release in thermo-alkaline pretreated samples were recorded
and further anaerobic digestion with digested sludge inocula resulted in 5. Factors affecting biohydrogen production
higher hydrogen yield (17.5 mL/g.TS) (Yin & Wang, 2019).
5.1. Inoculum/substrate ratio
4.1.3. Biological pretreatment method
The main drawback of chemical pretreatment methods is the for Insufficient substrate availability may cause an imbalance between
mation of furfurals, which affects microbial growth and hydrogen pro anabolic and catabolic reactions, ultimately affecting productivity.
duction. Biological methods involve the use of purified enzymes and a Taskan et al. (2022) studied the effect of inoculum/substrate ratio in a
cocktail of enzymes or microorganisms for the algal cell wall lysis. These laboratory-scale batch reactor for hydrogen production. Cladophora sp.
methods prevent the production of furfurals. The main enzymes used in biomass gathered from Hazar Lake, Turkey, was dried, milled into
pretreatment include cellulase, hemicellulase, pectinase, protease, powder, and used as feedstock. An anaerobic culture from a wastewater
amylase, etc. Cyanobacterial biomass is rich in glycogen, and amylase treatment plant was treated at 100 ◦ C for 30 min to incapacitate
can cleave (1, 4) glycosidic bonds and release sugar. Nayak et al. applied hydrogen consumers and used as an inoculum. Fermentation experi
an enzymatic treatment to Anabaena sp. PCC 7120 and were able to ments were performed at 100 mL capacity in a 120 mL capacity serum
achieve free sugar of 7.6 g/L without the production of any inhibitor bottle using different inoculum/feedstock ratios (0.0625, 0.125, 0.25,
(furfural). Fermentation of enzymatically pretreated hydrolysate resul 0.5, 1, 2, and 4). The maximum cumulative hydrogen (54.7 mL H2/g.VS)
ted in the production of 1600 mL/L of hydrogen, whereas untreated production was reported at I/S ratio of 4, which was 1.3 times as
algal biomass was unable to produce any hydrogen (Nayak et al., 2014). compared to I/S of 0.0625 (40.8 mL H2/g.VS). Decreases in the I/S ratio
The use of cellulase and macerozymes for the hydrolysis of algal mats cause an adverse effect on biohydrogen, probably because of the shock
has also been reported. Abdel-Kader et. al. (2022) treated Chlorella loading of organic biomass, high concentration of volatile fatty acids
biomass using cellulase and macerozymes in different combinations and (VFA), and low pH at low I/S ratios (Taşkan et al., 2022).
sonicated yeast hydrolases (amylase and pectinase). The use of cellulase-
processed hydrolysate resulted in 1300–1700 mL/L of hydrogen, which 5.2. pH
was lower than that of macerozyme-pretreated algal biomass
(1900–1915 mL/L). Combined pretreatment with cellulase and macer The pH of the medium is an important parameter, and microbes react
ozymes leads to higher production of hydrogen (2159–2359 mL/L) to changes in internal and external pH, shifting their metabolic activity,
(Abdel-Kader et al., 2022). Sonicated yeast enzyme pool pretreated algal which affects hydrogen production. During the fermentation process,
biomass led to higher hydrogen production of 2500–4200 mL/L. VFA are produced owing to a reduction in pH (4) and may cease
Ruminal microorganisms can degrade cellulosic and hemicellulosic hydrogen production. With an increase in pH up to a certain extent, an
components and are used to develop a consolidated process for bio increase in hydrogen production is reported due to increased microbial
ethanol, methane, etc. (Bhujbal et al., 2022). In a study Botta et al., growth, but beyond certain limits, an increase in pH causes a decrease in
(2020) isolated fermentative nonmethanogenic consortium from rumen ATP levels and reduces growth. The optimum pH range for hydrogen
fluid and further utilized it for hydrolysis and fermentation of paper for production depends on the inoculum, temperature, reactor type, feed
hydrogen production (8.2 H2 mL/g). Thongbunrod & Chaiprasert used stock, etc. (Arslan et al., 2015). The most favorable pH for hydrogen
rumen fluid for anaerobic digestion of rice straw and were able to pro generation is in the range 5–7 which corresponds to the optimum mi
duce methane up to 293 mL/g.VS. Rumen fluid can also be used for algal crobial growth temperature. Kumar et al., (2013) used Chlorella sor
biomass hydrolysis and the production of hydrogen and other products. okiniana as feedstock for E. cloacae IIT-BT 08 fermentation and studied
the pH profile, and a continuous decrease in pH was reported with op
4.1.4. Combined pretreatment timum pH production around pH 5.5. Liu et al., (2012) used C. butyricum
Combined pre-treatment methods can increase the hydrolysis of for dark fermentation of C. vulgaris biomass, adjusting the initial pH to 7;
algal biomass. Cheng et al. (2019) explored a hydrothermal/steam acid during the fermentation, the pH was controlled at 5.5 using an auto
treatment method for the pretreatment of algal biomass collected from matic titration method with 1 N NaOH, resulting in 1476 mL/L
6
hydrogen production. 6.2. Nanocatalysts addition
5.3. Temperature The addition of various catalysts has been reported to improve
hydrogen production. Hydrogenases are the main enzymes involved in
Temperature is also an important factor that determines microbial hydrogen production in dark fermentation. The addition of different
community structure, liquid metabolite distribution, and hydrogen types of nanocatalysts, such as nickel, iron, gold, copper, silver, and ti
production activity. The mixed microbial community can work in a wide tanium oxide has been reported. Nanocatalysts act as oxygen scavengers
pH range and produce hydrogen (Qiu et al., 2017). Gadow et al. (2012) and cofactors for hydrogenase and nitrogenase enzymes and enable
studied hydrogen production under mesophilic (37 ◦ C), thermophilic rapid electron transfer from the donor to the acceptor. Nanocatalysts
(55 ◦ C), and hyperthermophilic (80 ◦ C) conditions. Under mesophilic also promote microbial growth as they can also act as micronutrients.
conditions, the gas contained methane (26 %) as the main component, Srivastava et. al. studied the effect of Fe3O4 nanoparticles under dark
whereas no methane was identified under thermophilic and hyperther fermentation of algal biomass feedstock to increase hydrogen produc
mophilic conditions (Gadow et al., 2012). Thermophilic microbes are tion. Nanoparticle addition resulted in a 37.14 % higher cumulative H2
promising candidates for hydrogen production because of their capa production than that of the control (Srivastava et al., 2021). In another
bility to utilize a wide variety of raw materials and produce a small study, Kim et al. (2021) used conductive materials (Fe3O4) during
variety of end products. Clostridium, Thermoanaerobacterium, Caldi fermentation to overcome the inhibitory effect of red algae substrates
cellulosiruptor, and Thermoanaerobacter are well-reported microbes used and the process resulted in 20 % enhanced hydrogen production in the
for hydrogen production under thermophilic conditions (O-Thong et al., presence of all types of inhibitors (formic, levulinic acid, and HMF).
2011).
5.4. Organic loading rate (OLR) and hydraulic retention time (HRT) 6.3. Co-fermentation of substrates
HRT is the measurement of the time of contact between the substrate Another main issue in hydrogen production is the carbon/nitrogen
and fermentative microbes. OLR quantifies the amount of feedstock (C/N) ratio and low biodegradability of the biomass. The high C/N ratio
added every day per unit volume of the bioreactor. As the HRT increases, in macroalgae promotes VFA accumulation in microbes, whereas
the OLR decreases and vice versa, and the optimum values of these microalgae have a low C/N ratio, which increases ammonia levels and
parameters depend on the feedstock composition and microbial popu inhibits anaerobic fermentation. The use of mixed microalgae and
lation (Martins et al., 2022). With a short HRT, it is difficult to attain macroalgal feedstock can help improve the fermentation process.
sufficient microbial populations, while a higher microbial density can Microalgae Arthrospira platensis is rich in protein contents (low C/N),
resist load impact and inhibiting substances present in the influent (Su whereas macroalgae L. digitata has a high C/N ratio, and their co-
et al., 2020). Martins et al. (2022) studied the HRT and OLR during fermentation leads to an improved hydrogen production of 85.0 mL/g
continuous hydrogen production in a mesophilic continuous stirred tank vS (Xia et al., 2016). Ding et al. (2016) used L. digitata, Chlorella pyr
bioreactor (CSTR). The HRTs of 4, 5, and 6 days, corresponding to OLRs enoidosa, and Nannochloropsis oceanica biomass to maintain C/N (20) in
of 23.6, 18.0, and 10.6 g.VS/L/D, respectively, led to biohydrogen yield a two-stage process for hydrogen production. Co-fermentation resulted
of 8.8, 18.2, and 1.64 L/kg.vS respectively. In short HRT content of in a hydrogen yield of 94.5–97.0 mL/g VS which was 15.5–18 % greater
organic acids was higher, which hindered biohydrogen production and 5 than monofermentation of L. digitata (Ding et al., 2016). In another
d was reported as the most favorable HRT for hydrogen production. study, Jehlee et al., (2019) explored Chlorella sp. digestion with molasses
and POME and achieved hydrogen production of 44–75 and 40–48 mL
6. Strategy to increase biohydrogen production H2/g.VS respectively. The rigid cell wall and low carbohydrate content
of Taihu algae restrict its use in dark fermentation. Food waste is
6.1. Detoxification of algal biomass hydrolysate considered as a suitable cosubstrate due to its high organic content and
easy hydrolysis. Xu et al., performed co-fermentation of food waste with
During the pretreatment process side products, such as 5-hydroxy algae and investigated the effect of wet and dried algal biomass on
methylfurfural (5-HMF), levulinic acid, and formic acid are produced hydrogen production. Co-fermentation of food waste with dried algal
which inhibit microbial growth and affect hydrogen productivity biomass resulted in 12 times higher hydrogen production (31.42 mL H2/
(Kumar et al., 2015). 5-HMF is the major byproduct derived from hexose g.VS) as compared to fermentation with wet biomass (Xu et al., 2019).
sugars, and levulinic acid and formic acid are the degradation products
of 5-HMF. The key challenge of algal biomass hydrolysate is to alleviate
inhibitors to avoid negative effects. The presence of 1.37 g/L 5-HMF in 6.4. Two-step fermentation
hydrolysate inhibited hydrogen production by up to 50 %. Kumar et al.
(2015) developed a hybrid immobilized cell method for the simulta Low hydrogen yield and management of the residue left over after
neous removal of inhibitors and hydrogen production. The hybrid dark fermentation are major issues because they are rich in various
immobilized system was prepared by mixing sludge powder with so organic acids and cause environmental pollution. A two-step fermenta
dium alginate (2 % v/v), chitosan (1 % w/v), and activated carbon (2 % tion procedure combining dark fermentation and photofermentation
w/v). Hydrogen production was performed using algal hydrolysate as may increase hydrogen production and hinder waste residue manage
the carbon source in a batch vial, and 10 g of immobilized beads were ment (Fig. 2). The main challenge in two-step fermentation is the high
added. The process achieved 100 % 5-HMF removal and a hydrogen NH+4 content in the residue of dark fermentation, which inhibits nitro
yield of 4.8 L/L.d (Kumar et al., 2015). Gonzales et al. studied the ki genase (Zheng et al., 2010). Cheng et al. (2012) used Arthrospira platensis
netics of 5-HMF removal from algal hydrolysate using granular activated wet biomass for combined fermentation and pretreated the biomass with
carbon and reported that the removal was chemical adsorption-based microwave-aided dilute acid and glucoamylase. The hydrolysate was
and followed pseudo-second-order kinetics (Rolly Gonzales et al., subjected to dark fermentation, and the residue solution of dark
2016). Park et al., (2011) also used activated carbon to detoxify red algal fermentation containing acetate and butyrate was treated with zeolite to
hydrolysate and to produce 53.5 mL/g of algal biomass hydrogen. remove ammonia. Zeolite-treated dark fermentation residue was used
for photofermentation. Combined dark fermentation and photo
fermentation ensued an enhancement of hydrogen production by 96.6 to
337.0 mL H2/g.DW (Cheng et al., 2012).
7
Fig. 2. Multistep fermentation process for biohydrogen production.
6.5. Co-production of methane ($2.83), and dark fermentation ($2.57–6.98) (Ji & Wang, 2021). Most of
the reported technologies have only been studied at the lab scale, few at
To efficiently utilize algal biomass, researchers have also worked on the pilot-scale level, and there are no reports on hydrogen production at
the co-production of hydrogen and methane (Fig. 2). First, algal biomass the commercial level using biological methods. Tredici et al., (2016)
is fermented under thermophilic conditions via dark fermentation into performed a techno-economic analysis (TEA) for microalga Tetraselmis
H2, acetate, and CO2. In the next step, the leftover broth is subjected to suecica production in a 1 ha plant “Green Wall Panel-II photobioreactor
methane production. Wieczorek et al. (2014) investigated hydrogen and considering steps from inoculation to preparation of algal paste. The
methane production from C. vulgaris using two-stage fermentation, study concluded that locating the plant under favorable conditions al
combining dark fermentation for H2 and photo fermentation of residual lows algal production to reach around 54 tons per hectare at a cost of 6.2
broth for CH4 production. This process resulted in a production of 19 mL $/kg at 1 ha scale and the cost can be further reduced to 3.2 $/kg at a
H2/g.VS and 414 mL CH4/g.VS (Wieczorek et al., 2014). In another scale of 100 ha (Tredici et al., 2016). In another study, Vazquez-Romero
study, Ding et al. (2020) used L. digitata biomass pretreated hydrother et al., performed TEA comparing operation strategies including pro
mally and subjected it to two-step fermentation. Dark fermentation led duction scales (1 to 10 ha), harvesting (centrifugation or ultracentrifu
to 57.4 mL H2/g.VS and anaerobic digestion resulted in a yield of 282 gation), and drying methods (freeze-drying or spray drying). This study
mL/g.VS methane production (Ding et al., 2020). A three-stage strategy concluded that production cost can be reduced by 7.03 % using a
was also explored to achieve a high energy yield. Xia et al., (2013) combination of ultrafiltration and spray drying and up to 17.99 % by
developed a three-stage process for efficient conversion of Chlorella scale-up at 10 ha (Vázquez-Romero et al., 2022). Vatsala et al. (2008)
prrenoidosa, where biomass was first exposed for dark fermentation and used distillery effluent for hydrogen production at the industrial level by
photofermentation and then to the methanogenesis process which leads utilizing mixed co-culture of Citrobacter freundii 01, Enterobacter aero
to a total hydrogen yield of 193.3 mL/g.TVS and 186.2 mL/g.TVS genes E10, and Rhodopseudomonas palustris P at a 100 m3 scale and re
methane. Algal biomass collected from a freshwater lake was pretreated ported 21.38 kg of H2 production. In another study, Balachandar et al.,
with 1 % HCl and subjected to two-step fermentation, resulting in (2020) studied hydrogen production using E. claceae IIT-BT 08 from
hydrogen production of 39.4 mL/g.VS and 177.1–276.8 mL/g.VS different organic wastes at the lab scale (2 L) to pilot scale (10000 L) and
methane production (Chen et al., 2020a). reported a 1.62-fold in hydrogen productivity with an increase in scale.
Despite many studies related to biohydrogen production using various
7. Biohydrogen production scenario, feasibility, and future technologies, the literature related to the TEA of the process is limited.
perspectives Feedstock costs and low hydrogen yields are major issues. The produc
tion cost varies with the feedstock, for example, whole tree biomass
Various biomass-to-hydrogen technologies have been reported, and ($2.40/kg), forest residue ($3.00/kg), and agricultural residue (4.55/
the associated cost per kg of hydrogen ($/kg H2) reported for some of the kg) is reported. The utilization of algal biomass can provide a continuous
technologies are: biomass gasification ($1.77–2.77), direct bio supply of hydrogen at a lower price, with reduced global warming.
photolysis ($2.13), indirect biophotolysis ($1.42), photofermentation Ferreira et al., (2013) studied scaled-up scenarios for hydrogen
8
production from algal biomass. S. obliquus was cultivated in a raceway economics of the process.
pond, the collected biomass was subjected to dark fermentation using
C. butyricum and 7.3 H2/kg of biomass was produced by expending 8. Conclusions
71–100 MJ/MJH2 energy and emitting 5–6 kg CO2/MJH2.
Algal-based biorefineries have significant commercial potential, as Algae is a futuristic feedstock for biohydrogen production, as it re
they can be used for economic wastewater treatment, biofuel produc quires mild pretreatment compared to lignocellulosic material, is rich in
tion, and biochemical production, and have applications in cosmetics, carbohydrates, and results in higher hydrogen productivity. Although
nutraceuticals, and pharmaceuticals. However, there are many chal the algal cultivation is simple, there is a need to identify algal species
lenges that restrict its use on a commercial scale, such as high energy with high carbon capture capacity and biomass production potential.
demand for cultivation, low algal density, inefficient harvesting process, Further, dark fermentation has its limitations; therefore, the co-
the requirement of pretreatment, byproduct production, and low yield fermentation of algal biomass with other feedstock and the use of
of hydrogen. To make the cultivation process more economical, re multistep fermentation should be explored for efficient utilization of
searchers are working on the utilization of waste materials as feedstock. feedstock. Integration of hydrogen production with the co-production of
Carbohydrate is a component involved in hydrogen production, and other processes should also be explored to make the overall process
there is a need to identify algal species capable of accumulating high more economical.
carbohydrate content. Genetic engineering, mutagenesis, and cultiva
tion under stress conditions can be used to improve the growth and
CRediT authorship contribution statement
accumulation of carbohydrates in algal species. Enhanced CO2 fixation
via photosynthesis may lead to an increase in biomass production. In one
Shashi Kant Bhatia: Conceptualization, Writing – review & editing.
study, RuBisCo activase was transcriptionally upregulated in Nanno
J. Rajesh Banu: Writing – review & editing. Vijai Singh: Writing –
chloropsis aceanica IMET1, and the improved strain showed enhanced
review & editing. Gopalakrishnan Kumar: Conceptualization. Yung-
growth rate (32 %), biomass production (46 %), and lipid accumulation
Hun Yang: Conceptualization, Writing – review & editing.
(41 %) (Wei et al., 2017). All these methods are costly, and there is a
need to find economical and easy to scale up methods. Various methods
of algal harvesting have been reported, including flocculation, centri Declaration of Competing Interest
fugation, and flotation (Huy et al., 2022). Castellaños-Estupiñan et al.
(2018) designed an electro-flotation system for the concentration and The authors declare that they have no known competing financial
harvesting of freshwater algae and found it to be a good material with interests or personal relationships that could have appeared to influence
high efficiency (greater than95 %). Evaluation of the pretreatment the work reported in this paper.
method is also important, as it adds to the cost. The pretreatment
method should be efficient, low energy consuming, easy to scale up, easy Data availability
to perform, produce minimal side products, be environmentally
friendly, and cost-effective. Owing to morphological and physiological Data will be made available on request.
variations in algal species, it is not possible to use the same method of
pretreatment of algae, and continuous research efforts are needed to
Acknowledgment
develop a combined method of pretreatment. Understanding the cell
wall structure and components helps in designing better pretreatment
The authors acknowledge the KU Research Professor Program of
methods. Halim et al. (2019) demonstrated induction of autolytic cell
Konkuk University, Seoul, South Korea. This research was supported by
wall thinning in Nannochloropsis cells incubated at 38 ◦ C for 24 h. Under
the C1 Gas Refinery Program through the National Research Foundation
dark anoxic conditions, cells activate anaerobic metabolism, which,
of Korea (NRF), funded by the Ministry of Science and ICT
depletes polysaccharide reserves, and causes cell wall thinning. Cell wall
(2015M3D3A1A01064882), and by the National Research Foundation
thinning causes the weakening of cells, and a range of methods can be
of Korea (NRF) [NRF-NRF-2022M3I3A1082545, NRF-
used for cell disruption (Halim et al., 2019). The cost of concentrated
2022M3J4A1053702, NRF-2022R1A2C2003138 and NRF-
acids, alkalis, and other reagents used in pretreatment with energy
2021R1F1A1050325].
consumption used in steam pretreatment is high as compared to the
sugars produced. The cost of hydroxyl radical-aided thermal treatment
for algal biomass is reported around 22.52 $/ton biomass (Gao et al., References
2015). To improve productivity detoxification of hydrolysate is also
Abdel-Kader, H.A.A., Abdel-Basset, R., Danial, A.W., 2022. Yeast and enzymatic
considered an important step. According to a report from a technical hydrolysis in converting Chlorella biomass into hydrogen gas by Rhodobacter sp. and
perspective, the anionic resin is the best alternative, from techno- Rhodopseudomonas palustris. Int. J. Hydrog. Energy. 47, 1516–1528.
economic point overliming is the preferred choice while for a socio- Ajala, S.O., Alexander, M.L., 2020. Assessment of Chlorella vulgaris, Scenedesmus obliquus,
and Oocystis minuta for removal of sulfate, nitrate, and phosphate in wastewater. Int.
economic reason overliming and ultrafiltration are the best options J. Energy Environ. Eng. 11, 311–326.
(Llano et al., 2021). Along with the improvement of algal cultivation and Anvar, A., Nowruzi, B., 2021. Bioactive Properties of Spirulina: A Review. Microbial
pretreatment methods, the selection of suitable microbes capable of Bioactives. 4, 134–142.
Armoškaitė, A., Bārda, I., Andersone, I., Bonnevie, I.M., Ikauniece, A., Kotta, J.,
utilizing algal biomass and producing high hydrogen content is also
Kõivupuu, A., Lees, L., Psuty, I., Strāķe, S., Sprukta, S., Szymanek, L., von
important. Thus, there is a need to identify microbes capable of toler Thenen, M., Schrøder, L., Hansen, H.S., 2021. Considerations of use-use interactions
ating high concentrations of algal biomass-derived inhibitory products between macroalgae cultivation and other maritime sectors: an eastern baltic msp
case study. Sustainability. 13, 13888.
with enhanced hydrogen production. Furthermore, hydrogen produc
Arslan, C., Sattar, A., Changying, J., Nasir, A., Ali Mari, I., Zia Bakht, M., 2015. Impact of
tion is mostly carried out using mixed culture as it does not require ph management interval on biohydrogen production from organic fraction of
aseptic conditions and is least affected by the type of feedstock (Moussa municipal solid wastes by mesophilic thermophilic anaerobic codigestion. Biomed
et al., 2022). The use of mixed-culture microbes that consume hydrogen Res. Int. 2015, 590753.
Awasthi, M.K., Sindhu, R., Sirohi, R., Kumar, V., Ahluwalia, V., Binod, P., Juneja, A.,
should be avoided. The major bottle neck in algal-based hydrogen pro Kumar, D., Yan, B., Sarsaiya, S., Zhang, Z., Pandey, A., Taherzadeh, M.J., 2022.
duction is high cost and rigorous TEA and life cycle analyses (LCA) with Agricultural waste biorefinery development towards circular bioeconomy. Renew.
lab and pilot scale demonstration are required to confirm positive eco Sust. Energy. Rev. 158, 112122.
Balachandar, G., Varanasi, J.L., Singh, V., Singh, H., Das, D., 2020. Biological hydrogen
nomic returns. The production of other industrially important products, production via dark fermentation: A holistic approach from lab-scale to pilot-scale.
such as polyhydroxyalkanoates, can also help to improve the overall Int. J. Hydrog. Energy. 45, 5202–5215.
9
Batista, A.P., Moura, P., Marques, P.A.S.S., Ortigueira, J., Alves, L., Gouveia, L., 2014. Ding, L., Cheng, J., Xia, A., Jacob, A., Voelklein, M., Murphy, J.D., 2016. Co-generation
S. obliquus as feedstock for biohydrogen production by Enterobacter aerogenes and of biohydrogen and biomethane through two-stage batch co-fermentation of macro-
Clostridium butyricum. Fuel 117, 537–543. and micro-algal biomass. Bioresour. Technol. 218, 224–231.
Batista, A.P., Gouveia, L., Marques, P.A.S.S., 2018. Fermentative hydrogen production Ding, L., Cheng, J., Lin, R., Deng, C., Zhou, J., Murphy, J.D., 2020. Improving
from microalgal biomass by a single strain of bacterium Enterobacter aerogenes – biohydrogen and biomethane co-production via two-stage dark fermentation and
Effect of operational conditions and fermentation kinetics. Renew. Energy 119, anaerobic digestion of the pretreated seaweed Laminaria digitata. J. Clean. Prod. 251,
203–209. 119666.
Bhatia, S.K., Kim, S.-H., Yoon, J.-J., Yang, Y.-H., 2017. Current status and strategies for Egbo, M., Okoani, A., Okoh, I., 2018. Photobioreactors for microalgae cultivation–An
second generation biofuel production using microbial systems. Energ. Conver. overview. Int. J. Sci. Eng. Res. 9, 65–74.
Manage. 148, 1142–1156. Enayatizade, H., Chahartaghi, M., Hashemian, S.M., Arjomand, A., Ahmadi, M.H., 2019.
Bhatia, S.K., Gurav, R., Choi, T.-R., Jung, H.-R., Yang, S.-Y., Moon, Y.-M., Song, H.-S., Techno-economic evaluation of a new CCHP system with a hydrogen production
Jeon, J.-M., Choi, K.-Y., Yang, Y.-H., 2019. Bioconversion of plant biomass unit. Int. J. Low-Carbon. Technol. 14, 170–186.
hydrolysate into bioplastic (polyhydroxyalkanoates) using Ralstonia eutropha 5119. Franchi, G., Capocelli, M., De Falco, M., Piemonte, V., Barba, D., 2020. Hydrogen
Bioresour. Technol. 271, 306–315. production via steam reforming: a critical analysis of mr and rmm technologies.
Bhatia, S.K., Jagtap, S.S., Bedekar, A.A., Bhatia, R.K., Rajendran, K., Pugazhendhi, A., Membranes 10, 10.
Rao, C.V., Atabani, A.E., Kumar, G., Yang, Y.-H., 2021. Renewable biohydrogen Gad El-Rab, S.M.F., Hifney, A.F., Abdel-Basset, R., 2018. Costless and huge hydrogen
production from lignocellulosic biomass using fermentation and integration of yield by manipulation of iron concentrations in the new bacterial strain Brevibacillus
systems with other energy generation technologies. Sci. Total Environ. 765, 144429. invocatus SAR grown on algal biomass. Int. J. Hydrog. Energy. 43, 18896–18907.
Bhatia, S.K., Ahuja, V., Chandel, N., Gurav, R., Kant Bhatia, R., Govarthanan, M., Kumar Gadow, S.I., Li, Y.-Y., Liu, Y., 2012. Effect of temperature on continuous hydrogen
Tyagi, V., Kumar, V., Pugazendhi, A., Rajesh Banu, J., Yang, Y.-H., 2022a. Advances production of cellulose. Int. J. Hydrog. Energy. 37, 15465–15472.
in algal biomass pretreatment and its valorisation into biochemical and bioenergy by Ganesh Saratale, R., Kumar, G., Banu, R., Xia, A., Periyasamy, S., Dattatraya Saratale, G.,
the microbial processes. Bioresour. Technol. 358, 127437. 2018. A critical review on anaerobic digestion of microalgae and macroalgae and co-
Bhatia, S.K., Ahuja, V., Chandel, N., Mehariya, S., Kumar, P., Vinayak, V., Saratale, G.D., digestion of biomass for enhanced methane generation. Bioresour. Technol. 262,
Raj, T., Kim, S.-H., Yang, Y.-H., 2022b. An overview on microalgal-bacterial granular 319–332.
consortia for resource recovery and wastewater treatment. Bioresour. Technol. 351, Gao, L., Li, D., Gao, F., Liu, Z., Hou, Y., Chen, S., Zhang, D., 2015. Hydroxyl radical-aided
127028. thermal pretreatment of algal biomass for enhanced biodegradability. Biotechnol.
Bhujbal, S.K., Ghosh, P., Vijay, V.K., Rathour, R., Kumar, M., Singh, L., Kapley, A., 2022. Biofuels 8, 194.
Biotechnological potential of rumen microbiota for sustainable bioconversion of Garcia, J., Teymouri, A., Kumar, S. 2015. Kinetics of peptides and arginine production
lignocellulosic waste to biofuels and value-added products. Sci. Total Environ. 814, from microalgae (Scenedesmus sp .) by flash hydrolysis. Ind. Eng. Chem. Res. 54,
152773. 150209125521000.
Botta, L.S., Delforno, T.P., Rabelo, C.A.B.S., Silva, E.L., Varesche, M.B.A., 2020. Giang, T.T., Lunprom, S., Liao, Q., Reungsang, A., Salakkam, A., 2019. Improvement of
Microbial community analyses by high-throughput sequencing of rumen hydrogen production from Chlorella sp. biomass by acid-thermal pretreatment. PeerJ
microorganisms fermenting office paper in mesophilic and thermophilic lysimeters. 7, e6637–e.
Process Saf. Environ. Prot. 136, 182–193. Godvin Sharmila, V., Tamilarasan, K., Dinesh Kumar, M., Gopalakrishnan, K., Sunita, V.,
Buck, B., Grote, B. 2018. Seaweed in high-energy environments. Protocols for Adish Kumar, S., Rajesh Banu, J., 2022. Trends in dark biohydrogen production
macroalgae research. CRC Press, Boca Raton, 3-36. strategy and linkages with transition towards low carbon economy: An outlook, cost-
Cao, Y., Liu, H., Liu, W., Guo, J., Xian, M., 2022. Debottlenecking the biological effectiveness, bottlenecks and future scope. Int. J. Hydrog. Energy. 47,
hydrogen production pathway of dark fermentation: insight into the impact of strain 15309–15332.
improvement. Microb. Cell Fact. 21, 166. Gordalina, M., Pinheiro, H.M., Mateus, M., da Fonseca, M.M.R., Cesário, M.T., 2021.
Castellaños-Estupiñan, M., Sánchez-Galvis, M., Garcia Martinez, J., Barajas-Ferreira, C., Macroalgae as protein sources—a review on protein bioactivity, extraction,
Zuorro, A., Barajas-Solano, A., 2018. Design of an electroflotation system for the purification and characterization. Appl. Sci. 11, 7969.
concentration and harvesting of freshwater microalgae. Chem. Eng. Trans. 64. Goswami, R.K., Mehariya, S., Karthikeyan, O.P., Gupta, V.K., Verma, P., 2022.
Chandel, N., Ahuja, V., Gurav, R., Kumar, V., Tyagi, V.K., Pugazhendhi, A., Kumar, G., Multifaceted application of microalgal biomass integrated with carbon dioxide
Kumar, D., Yang, Y.-H., Bhatia, S.K., 2022. Progress in microalgal mediated reduction and wastewater remediation: A flexible concept for sustainable
bioremediation systems for the removal of antibiotics and pharmaceuticals from environment. J. Clean. Prod. 339, 130654.
wastewater. Sci. Total Environ. 825, 153895. Halim, R., Hill, D.R.A., Hanssen, E., Webley, P.A., Blackburn, S., Grossman, A.R.,
Chandrasekhar, K., Lee, Y.-J., Lee, D.-W., 2015. Biohydrogen production: strategies to Posten, C., Martin, G.J.O., 2019. Towards sustainable microalgal biomass
improve process efficiency through microbial routes. Int. J. Mole. Sci. 16, processing: anaerobic induction of autolytic cell-wall self-ingestion in lipid-rich
8266–8293. Nannochloropsis slurries. Green Chem. 21 (11), 2967–2982.
Chen, S., Qu, D., Xiao, X., Miao, X., 2020b. Biohydrogen production with lipid-extracted He, S., Zhao, M., Wang, J., Cheng, Z., Yan, B., Chen, G., 2020. Hydrothermal liquefaction
Dunaliella biomass and a new strain of hyper-thermophilic archaeon Thermococcus of low-lipid algae Nannochloropsis sp. and Sargassum sp.: Effect of feedstock
eurythermalis A501. Int. J. Hydrog. Energy. 45, 12721–12730. composition and temperature. Sci. Total Environ. 712, 135677.
Chen, C., Sun, C., Xia, A., Liao, Q., Guo, X., Huang, Y., Fu, Q., Zhu, X., Zhu, X., 2020a. Hoang, A.T., Sirohi, R., Pandey, A., Nižetić, S., Lam, S.S., Chen, W.-H., Luque, R.,
Sustainable biohythane production from algal bloom biomass through two-stage Thomas, S., Arıcı, M., Pham, V.V., 2022. Biofuel production from microalgae:
fermentation: Impacts of the physicochemical characteristics and fermentation challenges and chances. Phytochem. Rev. https://link.springer.com/article/
performance. Int. J. Hydrog. Energy. 45, 34461–34472. 10.1007/s11101-022-09819-y.
Chen, Y., Yin, Y., Wang, J., 2022. Biohydrogen production using macroalgal biomass of Hong, Y., Wu, Y.-R., 2020. Acidolysis as a biorefinery approach to producing advanced
Laminaria japonica pretreated by gamma irradiation as substrate. Fuel 309, 122179. bioenergy from macroalgal biomass: A state-of-the-art review. Bioresour. Technol.
Cheng, J., Xia, A., Liu, Y., Lin, R., Zhou, J., Cen, K., 2012. Combination of dark- and 318, 124080.
photo-fermentation to improve hydrogen production from Arthrospira platensis wet Huo, S., Dong, R., Wang, Z., Pang, C., Yuan, Z., Zhu, S., Chen, L., 2011. Available
biomass with ammonium removal by zeolite. Int. J. Hydrog. Energy. 37, resources for algal biofuel development in China. Energies 4, 1321–1335.
13330–13337. Huy, M., Kumar, G., Sharma, P., Sirohi, R., Pandey, A., Kim, S.-H., 2022. Effective
Cheng, J., Yue, L., Ding, L., Li, Y.-Y., Ye, Q., Zhou, J., Cen, K., Lin, R., 2019. Improving recovery of microalgal biomass using various types of emulsion polymers.
fermentative hydrogen and methane production from an algal bloom through J. Biotechnol. 358, 25–32.
hydrothermal/steam acid pretreatment. Int. J. Hydrog. Energy. 44, 5812–5820. Jehlee, A., Rodjaroen, S., Waewsak, J., Reungsang, A., O-Thong, S., 2019. Improvement
Chiang, C.-L., Lee, C.-M., Chen, P.-C., 2011. Utilization of the cyanobacteria Anabaena sp. of biohythane production from Chlorella sp. TISTR 8411 biomass by co-digestion
CH1 in biological carbon dioxide mitigation processes. Bioresour. Technol. 102, with organic wastes in a two-stage fermentation. Int. J. Hydrog. Energy. 44,
5400–5405. 17238–17247.
Dange, P., Pandit, S., Jadhav, D., Shanmugam, P., Gupta, P.K., Kumar, S., Kumar, M., Ji, M., Wang, J., 2021. Review and comparison of various hydrogen production methods
Yang, Y.-H., Bhatia, S.K., 2021. Recent developments in microbial electrolysis cell- based on costs and life cycle impact assessment indicators. Int. J. Hydrog. Energy.
based biohydrogen production utilizing wastewater as a feedstock. Sustainability. 46, 38612–38635.
13, 8796. Jönsson, M., Allahgholi, L., Sardari, R.R.R., Hreggviðsson, G.O., Nordberg Karlsson, E.,
de Oliveira, M.C., Bassin, I.D., Cammarota, M.C., 2022. Microalgae and cyanobacteria 2020. Extraction and modification of macroalgal polysaccharides for current and
biomass pretreatment methods: a comparative analysis of chemical and next-generation applications. Mole. 25, 930.
thermochemical pretreatment methods aimed at methane production. Fermentation. Jung, K.-W., Kim, D.-H., Shin, H.-S., 2011. Fermentative hydrogen production from
8, 497. Laminaria japonica and optimization of thermal pretreatment conditions. Bioresour.
De-Luca, R., Bezzo, F., Béchet, Q., Bernard, O., 2019. Meteorological data-based optimal Technol. 102, 2745–2750.
control strategy for microalgae cultivation in open pond systems. Complexity 2019, Kim, D.-H., Yoon, J.-J., Kim, S.-H., Park, J.-H., 2021. Effect of conductive material for
4363895. overcoming inhibitory conditions derived from red algae-based substrate on
di Cicco, M.R., Iovinella, M., Palmieri, M., Lubritto, C., Ciniglia, C., 2021. Extremophilic biohydrogen production. Fuel 285, 119059.
microalgae Galdieria gen. for urban wastewater treatment: current state, the case of King, A.D., Lane, T.P., Henley, B.J., Brown, J.R., 2020. Global and regional impacts differ
“power” system, and future prospects. Plants 10, 2343. between transient and equilibrium warmer worlds. Nat. Clim. Chang. 10, 42–47.
Dinesh Kumar, M., Yukesh Kannah, R., Kumar, G., Sivashanmugam, P., Rajesh Banu, J., Kumar, M.D., Kavitha, S., Tyagi, V.K., Rajkumar, M., Bhatia, S.K., Kumar, G., Banu, J.R.,
2020. A novel energetically efficient combinative microwave pretreatment for 2022. Macroalgae-derived biohydrogen production: biorefinery and circular
achieving profitable hydrogen production from marine macro algae (Ulva reticulate). bioeconomy. Biomass Convers. Biorefine. 12, 769–791.
Bioresour. Technol. 301, 122759.
10
Kumar, K., Roy, S., Das, D., 2013. Continuous mode of carbon dioxide sequestration by Pugazhendi, A., Jamal, M.T., Al-Mur, B.A., Jeyakumar, R.B., Kumar, G., 2022.
C. sorokiniana and subsequent use of its biomass for hydrogen production by Macroalgae (Ulva reticulata) derived biohydrogen recovery through mild surfactant
E. cloacae IIT-BT 08. Bioresour. Technol. 145, 116–122. induced energy and cost efficient dispersion pretreatment technology. Chemosphere
Kumar, G., Sivagurunathan, P., Kobayashi, T., Xu, K.Q., Kim, S.-H., 2015. Simultaneous 288, 132463.
removal of 5-hydroxy methyl furfural (5-HMF) and hydrogen production from acid Qiu, C., Yuan, P., Sun, L., Wang, S., Lo, S., Zhang, D., 2017. Effect of fermentation
(H2SO4) pretreated red-algal hydrolysate via hybrid immobilized cells. Algal Res. temperature on hydrogen production from xylose and the succession of hydrogen-
11, 326–333. producing microflora. J. Chem. Technol. Biotechnol. 92, 1990–1997.
Kumar, G., Sivagurunathan, P., Thi, N.B.D., Zhen, G., Kobayashi, T., Kim, S.-H., Xu, K., Rincón-Pérez, J., Razo-Flores, E., Morales, M., Alatriste-Mondragón, F., Celis, L.B., 2020.
2016. Evaluation of different pretreatments on organic matter solubilization and Improving the biodegradability of Scenedesmus obtusiusculus by thermochemical
hydrogen fermentation of mixed microalgae consortia. Int. J. Hydrog. Energy. 41, pretreatment to produce hydrogen and methane. Bioenergy Res. 13, 477–486.
21628–21640. Rolly Gonzales, R., Hong, Y., Park, J.-H., Kumar, G., Kim, S.-H., 2016. Kinetics and
Lam, T.P., Lee, T.-M., Chen, C.-Y., Chang, J.-S., 2018. Strategies to control biological equilibria of 5-hydroxymethylfurfural (5-HMF) sequestration from algal hydrolyzate
contaminants during microalgal cultivation in open ponds. Bioresour. Technol. 252, using granular activated carbon. J. Chem. Technol. Biotechnol. 91, 1157–1163.
180–187. Schädler, T., Neumann-Cip, A.-C., Wieland, K., Glöckler, D., Haisch, C., Brück, T.,
Liu, C.-H., Chang, C.-Y., Cheng, C.-L., Lee, D.-J., Chang, J.-S., 2012. Fermentative Weuster-Botz, D., 2020. High-density microalgae cultivation in open thin-layer
hydrogen production by Clostridium butyricum CGS5 using carbohydrate-rich cascade photobioreactors with water recycling. Appl. Sci. 10, 3883.
microalgal biomass as feedstock. Int. J. Hydrog. Energy. 37, 15458–15464. Schipper, K., Al Muraikhi, M., Alghasal, G.S.H.S., Saadaoui, I., Bounnit, T., Rasheed, R.,
Llano, T., Rueda, C., Dosal, E., Andrés, A., Coz, A., 2021. Multi-criteria analysis of Dalgamouni, T., Al Jabri, H.M.S.J., Wijffels, R.H., Barbosa, M.J., 2019. Potential of
detoxification alternatives: Techno-economic and socio-environmental assessment. novel desert microalgae and cyanobacteria for commercial applications and CO2
Biomass Bioenergy 154, 106274. sequestration. J. Appl. Phycol. 31, 2231–2243.
Lovato, G., Augusto, I.M.G., Ferraz Júnior, A.D.N., Albanez, R., Ratusznei, S.M., Schmitz, T.A., Kraft, E., 2022. Pilot scale photobioreactor system for land-based
Etchebehere, C., Zaiat, M., Rodrigues, J.A.D., 2021. Reactor start-up strategy as key macroalgae cultivation. J. Appl. Phycol. 34, 507–516.
for high and stable hydrogen production from cheese whey thermophilic dark Sekar, M., Mathimani, T., Alagumalai, A., Chi, N.T.L., Duc, P.A., Bhatia, S.K.,
fermentation. Int. J. Hydrog. Energy. 46, 27364–27379. Brindhadevi, K., Pugazhendhi, A., 2021. A review on the pyrolysis of algal biomass
Luo, L., Sriram, S., Johnravindar, D., Louis Philippe Martin, T., Wong, J.W.C., for biochar and bio-oil – Bottlenecks and scope. Fuel 283, 119190.
Pradhan, N., 2022. Effect of inoculum pretreatment on the microbial and metabolic Shekh, A., Sharma, A., Schenk, P.M., Kumar, G., Mudliar, S., 2022. Microalgae
dynamics of food waste dark fermentation. Bioresour. Technol. 358, 127404. cultivation: photobioreactors, CO2 utilization, and value-added products of
Martinez-Burgos, W.J., Sydney, E.B., de Paula, D.R., Medeiros, A.B.P., de Carvalho, J.C., industrial importance. J. Chem. Technol. Biotechnol. 97, 1064–1085.
Molina, D., Soccol, C.R., 2021. Hydrogen production by dark fermentation using a Shi, X., Jung, K.-W., Kim, D.-H., Ahn, Y.-T., Shin, H.-S., 2011. Direct fermentation of
new low-cost culture medium composed of corn steep liquor and cassava processing Laminaria japonica for biohydrogen production by anaerobic mixed cultures. Int. J.
water: Process optimization and scale-up. Bioresour. Technol. 320, 124370. Hydrog. Energy. 36, 5857–5864.
Martins, I., Surra, E., Ventura, M., Lapa, N., 2022. BioH2 from dark fermentation of Singh, U.B., Ahluwalia, A., 2013. Microalgae: A promising tool for carbon sequestration.
OFMSW: Effect of the hydraulic retention time and organic loading rate. Appl. Sci. Mitig. Adapt. Strateg. Global. Change. 18, 73–95.
12, 4240. Singh, H., Rout, S., Das, D., 2022. Dark fermentative biohydrogen production using
Mehariya, S., Plöhn, M., Leon-Vaz, A., Patel, A., Funk, C., 2022. Improving the content of pretreated Scenedesmus obliquus biomass under an integrated paradigm of
high value compounds in Nordic Desmodesmus microalgal strains. Bioresour. Technol. biorefinery. Int. J. Hydrog. Energy. 47, 102–116.
359, 127445. Sivagurunathan, P., Kumar, G., Kobayashi, T., Xu, K., Kim, S.-H., 2017. Effects of various
Milledge, J.J., Nielsen, B.V., Maneein, S., Harvey, P.J., 2019. A brief review of anaerobic dilute acid pretreatments on the biochemical hydrogen production potential of
digestion of algae for bioenergy. Energies 12, 1166. marine macroalgal biomass. Int. J. Hydrog. Energy. 42, 27600–27606.
Moussa, R.N., Moussa, N., Dionisi, D., 2022. Hydrogen production from biomass and Snehya, A.V., Sundaramahalingam, M.A., Rajeshbanu, J., Sivashanmugam, P., 2022.
organic waste using dark fermentation: an analysis of literature data on the effect of Studies on evaluation of pretreatment efficiency of surfactant mediated ultrasonic
operating parameters on process performance. Processes. 10, 156. pretreatment of Sargassum tennerimum (marine macroalgae) for achieving profitable
Mugnai, G., Borruso, L., Mimmo, T., Cesco, S., Luongo, V., Frunzo, L., Fabbricino, M., biohydrogen production. Int. J. Hydrog. Energy. 47 (39), 17184–17193.
Pirozzi, F., Cappitelli, F., Villa, F., 2021. Dynamics of bacterial communities and Srivastava, N., Srivastava, M., Singh, R., Syed, A., Bahadur Pal, D., Elgorban, A.M.,
substrate conversion during olive-mill waste dark fermentation: Prediction of the Kushwaha, D., Mishra, P.K., Gupta, V.K., 2021. Co-fermentation of residual algal
metabolic routes for hydrogen production. Bioresour. Technol. 319, 124157. biomass and glucose under the influence of Fe3O4 nanoparticles to enhance
Narisetty, V., Nagarajan, S., Gadkari, S., Ranade, V.V., Zhang, J., Patchigolla, K., biohydrogen production under dark mode. Bioresour. Technol. 342, 126034.
Bhatnagar, A., Kumar Awasthi, M., Pandey, A., Kumar, V., 2022. Process Stanislaus, M.S., Zhang, N., Yuan, Y., Zheng, H., Zhao, C., Hu, X., Zhu, Q., Yang, Y., 2018.
optimization for recycling of bread waste into bioethanol and biomethane: A circular Improvement of biohydrogen production by optimization of pretreatment method
economy approach. Energ. Conver. Manage. 266, 115784. and substrate to inoculum ratio from microalgal biomass and digested sludge.
Nayak, B.K., Roy, S., Das, D., 2014. Biohydrogen production from algal biomass Renew. Energy 127, 670–677.
(Anabaena sp. PCC 7120) cultivated in airlift photobioreactor. Int. J. Hydrog. Su, C., Liu, Y., Yang, X., Li, H., 2020. Effect of hydraulic retention time on biohydrogen
Energy. 39, 7553–7560. production from glucose in an internal circulation reactor. Energy Fuel 34,
Pandey, A., Sinha, P., Pandey, A., 2021. Hydrogen production by sequential dark and 3244–3249.
photofermentation using wet biomass hydrolysate of Spirulina platensis: Response Tan, J.S., Lee, S.Y., Chew, K.W., Lam, M.K., Lim, J.W., Ho, S.-H., Show, P.L., 2020.
surface methodological approach. Int. J. Hydrog. Energy. 46, 7137–7146. A review on microalgae cultivation and harvesting, and their biomass extraction
Panggabean, J.A., Adiguna, S.P.P., Rahmawati, S.I., Ahmadi, P., Zainuddin, E.N., processing using ionic liquids. Bioengineered. 11, 116–129.
Bayu, A., Putra, M.Y., 2022. Antiviral activities of algal-based sulfated Tang, D., Han, W., Li, P., Miao, X., Zhong, J., 2011. CO2 biofixation and fatty acid
polysaccharides. Molecules 27, 1178. composition of Scenedesmus obliquus and Chlorella pyrenoidosa in response to
Park, J., Kumar, G., Bakonyi, P., Peter, J., Nemestóthy, N., Koter, S., Kujawski, W., Bélafi- different CO2 levels. Bioresour. Technol. 102, 3071–3076.
Bakó, K., Pientka, Z., Muñoz, R., Kim, S.-H., 2021. Comparative evaluation of CO2 Taşkan, B., Köroğlu, E.O., Taşkan, E., 2022. Cladophora sp. as a sustainable feedstock for
fixation of microalgae strains at various CO2 aeration conditions. Waste Biomass dark fermentative biohydrogen production. Int. J. Hydrog. Energy. 47,
Valoriz. 12, 1–9. 15410–15418.
Park, J.-H., Yoon, J.-J., Park, H.-D., Kim, Y.J., Lim, D.J., Kim, S.-H., 2011. Feasibility of Thong, S., Mamimin, C., Prasertsan, 2011. Effect of temperature and initial pH on
biohydrogen production from Gelidium amansii. Int. J. Hydrog. Energy. 36, biohydrogen production from palm oil mill effluent: Long-term evaluation and
13997–14003. microbial community analysis. Electron. J. Biotechnol. 14.
Patel, S.K.S., Lee, J.-K., Kalia, V.C., 2018. Beyond the theoretical yields of dark- Thongbunrod, N., Chaiprasert, P., 2021. Efficacy and metagenomic analysis of the
fermentative biohydrogen. Ind. J. Microbiol. 58, 529–530. stabilized anaerobic lignocellulolytic microbial consortium from bubalus bubalis
Patel, A.K., Singhania, R.R., Awasthi, M.K., Varjani, S., Bhatia, S.K., Tsai, M.-L., Hsieh, S.- rumen with rice straw enrichment for methane production. Bioenergy Res. 14 (3),
L., Chen, C.-W., Dong, C.-D., 2021. Emerging prospects of macro- and microalgae as 870–890.
prebiotic. Microb. Cell Fact. 20, 112. Tredici, M.R., Rodolfi, L., Biondi, N., Bassi, N., Sampietro, G., 2016. Techno-economic
Peters, J.W., Schut, G.J., Boyd, E.S., Mulder, D.W., Shepard, E.M., Broderick, J.B., analysis of microalgal biomass production in a 1-ha Green Wall Panel (GWP®) plant.
King, P.W., Adams, M.W.W., 2015. [FeFe]- and [NiFe]-hydrogenase diversity, Algal Res. 19, 253–263.
mechanism, and maturation. Biochim. Biophys. Acta – Mole. Cell Res. 1853, Vatsala, T.M., Raj, S.M., Manimaran, A., 2008. A pilot-scale study of biohydrogen
1350–1369. production from distillery effluent using defined bacterial co-culture. Int. J. Hydrog.
Pinto, T., Gouveia, L., Ortigueira, J., Saratale, G.D., Moura, P., 2018. Enhancement of Energy. 33, 5404–5415.
fermentative hydrogen production from Spirogyra sp. by increased carbohydrate Vázquez-Romero, B., Perales, J.A., Pereira, H., Barbosa, M., Ruiz, J., 2022. Techno-
accumulation and selection of the biomass pretreatment under a biorefinery model. economic assessment of microalgae production, harvesting and drying for food, feed,
J. Biosci. Bioeng. 126 (2), 226–234. cosmetics, and agriculture. Sci. Total Environ. 837, 155742.
Preethi, R.B., Rajesh Banu, J., Gopalakrishnan, K., Tyagi, V.K., Bajhaiya, A.K., Verspreet, J., Soetemans, L., Gargan, C., Hayes, M., Bastiaens, L., 2021. Nutritional
Gugulothu, P., Gunasekaran, M., 2022. Biohydrogen production from waste profiling and preliminary bioactivity screening of five micro-algae strains cultivated
activated sludge through thermochemical mechanical pretreatment. Bioresour. in Northwest Europe. Foods. 10, 1516.
Technol. 358, 127301. Viswanaathan, S., Perumal, P.K., Sundaram, S., 2022. Integrated approach for carbon
Pugazhendhi, A., Kumar, G., Sivagurunathan, P., 2019. Microbiome involved in sequestration and wastewater treatment using algal and bacterial consortia:
anaerobic hydrogen producing granules: A mini review. Biotechnol. Rep, 21, opportunities and challenges. Sustainability. 14, 1075.
e00301.
11
Wang, J., Yin, Y., 2018. Fermentative hydrogen production using pretreated microalgal Xu, L., Cheng, X., Wang, Q., 2018a. Enhanced lipid production in Chlamydomonas
biomass as feedstock. Microb. Cell Fact. 17, 22. reinhardtii by co-culturing with Azotobacter chroococcum. Front. Plant, Sci, p. 9.
Wei, L., Wang, Q., Xin, Y., Lu, Y., Xu, J., 2017. Enhancing photosynthetic biomass Xu, L., Cheng, X., Wang, Q., 2018b. enhanced lipid production in chlamydomonas
productivity of industrial oleaginous microalgae by overexpression of RuBisCO reinhardtii by co-culturing with Azotobacter chroococcum. Front. Plant Sci. 9.
activase. Algal Res. 27, 366–375. Yang, Z., Guo, R., Xu, X., Fan, X., Li, X., 2010. Enhanced hydrogen production from lipid-
Wieczorek, N., Kucuker, M.A., Kuchta, K., 2014. Fermentative hydrogen and methane extracted microalgal biomass residues through pretreatment. Int. J. Hydrog. Energy.
production from microalgal biomass (Chlorella vulgaris) in a two-stage combined 35, 9618–9623.
process. Appl. Energy 132, 108–117. Yin, Y., Hu, J., Wang, J., 2019. Fermentative hydrogen production from macroalgae
Xia, A., Cheng, J., Ding, L., Lin, R., Huang, R., Zhou, J., Cen, K., 2013. Improvement of Laminaria japonica pretreated by microwave irradiation. Int. J. Hydrog. Energy. 44,
the energy conversion efficiency of Chlorella pyrenoidosa biomass by a three-stage 10398–10406.
process comprising dark fermentation, photofermentation, and methanogenesis. Yin, Y., Chen, Y., Wang, J., 2021. Co-fermentation of sewage sludge and algae and Fe2+
Bioresour. Technol. 146, 436–443. addition for enhancing hydrogen production. Int. J. Hydrog. Energy. 46, 8950–8960.
Xia, A., Cheng, J., Ding, L., Lin, R., Song, W., Zhou, J., Cen, K., 2014. Enhancement of Yin, Y., Wang, J., 2019. Hydrogen production and energy recovery from macroalgae
energy production efficiency from mixed biomass of Chlorella pyrenoidosa and Saccharina japonica by different pretreatment methods. Renew. Energy 141, 1–8.
cassava starch through combined hydrogen fermentation and methanogenesis. Appl. Yun, Y.-M., Jung, K.-W., Kim, D.-H., Oh, Y.-K., Cho, S.-K., Shin, H.-S., 2013. Optimization
Energy 120, 23–30. of dark fermentative H2 production from microalgal biomass by combined (acid+
Xia, A., Cheng, J., Song, W., Su, H., Ding, L., Lin, R., Lu, H., Liu, J., Zhou, J., Cen, K., ultrasonic) pretreatment. Bioresour. Technol. 141, 220–226.
2015. Fermentative hydrogen production using algal biomass as feedstock. Renew. Zheng, G.H., Wang, L., Kang, Z.H., 2010. Feasibility of biohydrogen production from tofu
Sust. Energy. Rev. 51, 209–230. wastewater with glutamine auxotrophic mutant of Rhodobacter sphaeroides. Renew.
Xia, A., Jacob, A., Tabassum, M.R., Herrmann, C., Murphy, J.D., 2016. Production of Energy 35, 2910–2913.
hydrogen, ethanol and volatile fatty acids through co-fermentation of macro- and
micro-algae. Bioresour. Technol. 205, 118–125.
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Bioresource Technology 368 (2023) 128332

Contents lists available at ScienceDirect

Algal biomass to biohydrogen: Pretreatment, influencing factors, and

• Algae capture CO2 and produce

1. Introduction hydrogen production potential as compared to protein and lipids.

hydrogen production. 6.2. Nanocatalysts addition

Fig. 2. Multistep fermentation process for biohydrogen production.

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