Biohydrogen

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Microbial hydrogen production. Algae hydrogen production.jpg
Microbial hydrogen production.

Biohydrogen is H2 that is produced biologically. [1] Interest is high in this technology because H2 is a clean fuel and can be readily produced from certain kinds of biomass, [2] including biological waste. [3] Furthermore some photosynthetic microorganisms are capable to produce H2 directly from water splitting using light as energy source. [4] [5]

Contents

Besides the promising possibilities of biological hydrogen production, many challenges characterize this technology. First challenges include those intrinsic to H2, such as storage and transportation of an explosive noncondensible gas. Additionally, hydrogen producing organisms are poisoned by O2 and yields of H2 are often low.

Biochemical principles

The main reactions driving hydrogen formation involve the oxidation of substrates to obtain electrons. Then, these electrons are transferred to free protons to form molecular hydrogen. This proton reduction reaction is normally performed by an enzyme family known as hydrogenases.

In heterotrophic organisms, electrons are produced during the fermentation of sugars. Hydrogen gas is produced in many types of fermentation as a way to regenerate NAD+ from NADH. Electrons are transferred to ferredoxin, or can be directly accepted from NADH by a hydrogenase, producing H2. Because of this most of the reactions start with glucose, which is converted to acetic acid. [6]

A related reaction gives formate instead of carbon dioxide:

These reactions are exergonic by 216 and 209 kcal/mol, respectively.

It has been estimated that 99% of all organisms utilize or produce dihydrogen (H2). Most of these species are microbes and their ability to use or produce H2 as a metabolite arises from the expression of H2 metalloenzymes known as hydrogenases. [7] Enzymes within this widely diverse family are commonly sub-classified into three different types based on the active site metal content: [FeFe]-hydrogenases (iron-iron), [NiFe]-hydrogenases (nickel-iron) hydrogenases, and [Fe]-hydrogenases (iron-only). [8] Many organisms express these enzymes. Notable examples are members of the genera Clostridium, Desulfovibrio, Ralstonia or the pathogen Helicobacter , being most of them strict-anaerobes or facultative microorganisms. Other microorganisms such green algae also express highly active hydrogenases, as it is the cae for members of the genera Chlamydomonas.

The active site structures of the three types of hydrogenase enzymes. ActiveSitesCorrected.png
The active site structures of the three types of hydrogenase enzymes.

Due to the extreme diversity of hydrogenase enzymes, on-going efforts are focused on screening for novel enzymes with improved features, [9] [10] [11] as well as engineering already characterized hydrogenases to confer them more desirable characteristics. [12]

Production by algae

The biological hydrogen production with algae is a method of photobiological water splitting which is done in a closed photobioreactor based on the production of hydrogen as a solar fuel by algae. [13] [14] Algae produce hydrogen under certain conditions. In 2000 it was discovered that if C. reinhardtii algae are deprived of sulfur they will switch from the production of oxygen, as in normal photosynthesis, to the production of hydrogen. [15] [16] [17]

Green algae express [FeFe] hydrogenases, being some of them considered the most efficient hydrogenases with turnover rates superior to 104 s−1. This remarkable catalytic efficiency is nonetheless shadowed by its extreme sensitivity to oxygen, being irreversibly inactivated by O2 [12] . When the cells are deprived from sulfur, oxygen evolution stops due to photo-damage of photosystem II, in this state the cells start consuming O2 and provide the ideal anaerobic environment for the native [FeFe] hydrogenases to catalyze H2 production.

Photosynthesis

Formation of Chlorella cell-based spheroids.webp
Schematic illustration showing the assembly, spatial organization and dual functionality of multicellular droplet-based living micro-reactors.webp
Algal cell-based bioreactors that can produce hydrogen [18]

Photosynthesis in cyanobacteria and green algae splits water into hydrogen ions and electrons. The electrons are transported over ferredoxins. [19] Fe-Fe-hydrogenases (enzymes) combine them into hydrogen gas. In Chlamydomonas reinhardtii Photosystem II produces in direct conversion of sunlight 80% of the electrons that end up in the hydrogen gas. [20]

In 2020 scientists reported the development of algal-cell based micro-emulsion for multicellular spheroid microbial reactors capable of producing hydrogen alongside either oxygen or CO2 via photosynthesis in daylight under air. Enclosing the microreactors with synergistic bacteria was shown to increase levels of hydrogen production via reduction of O2 concentrations. [21] [18]

Improving production by light harvesting antenna reduction

The chlorophyll (Chl) antenna size in green algae is minimized, or truncated, to maximize photobiological solar conversion efficiency and H2 production. It has been shown that Light-harvesting complex photosystem II light-harvesting protein LHCBM9 promotes efficient light energy dissipation. [22] The truncated Chl antenna size minimizes absorption and wasteful dissipation of sunlight by individual cells, resulting in better light utilization efficiency and greater photosynthetic efficiency when the green alga are grown as a mass culture in bioreactors. [23]

Economics

With current reports for algae-based biohydrogen, it would take about 25,000 square kilometre algal farming to produce biohydrogen equivalent to the energy provided by gasoline in the US alone. This area represents approximately 10% of the area devoted to growing soya in the US. [24]

Bioreactor design issues

Attempts are in progress to solve these problems via bioengineering.

Production by cyanobacteria

Biological hydrogen production is also observed in nitrogen-fixing cyanobacteria. This microorganisms can grow forming filaments. Under nitrogen-limited conditions some cells can specialize and form heterocysts, which ensures an anaerobic intracellular space to ease N2 fixation by the nitrogenase enzyme expressed also inside.

Under nitrogen-fixation conditions, the nitrogenase enzyme accepts electrons and consume ATP to break the triple dinitrogen bond and reduce it to ammonia. [25] During the catalytic cycle of the nitrogenase enzyme, molecular hydrogen is also produced.

Nevertheless, since the production of H2 is an important loss of energy for the cells, most of nitrogen fixing cyanobacteria also feature at least one uptake hydrogenase. [26] Uptake hydrogenases exhibit a catalytic bias towards oxygen oxidation, thus can assimilate the produced H2 as a way to recover part of the energy invested during the nitrogen fixation process.

History

In 1933, Marjory Stephenson and her student Stickland reported that cell suspensions catalysed the reduction of methylene blue with H2. Six years later, Hans Gaffron observed that the green photosynthetic alga Chlamydomonas reinhardtii , would sometimes produce hydrogen. [27] In the late 1990s Anastasios Melis discovered that deprivation of sulfur induces the alga to switch from the production of oxygen (normal photosynthesis) to the production of hydrogen. He found that the enzyme responsible for this reaction is hydrogenase, but that the hydrogenase lost this function in the presence of oxygen. Melis also discovered that depleting the amount of sulfur available to the algae interrupted their internal oxygen flow, allowing the hydrogenase an environment in which it can react, causing the algae to produce hydrogen. [28] Chlamydomonas moewusii is also a promising strain for the production of hydrogen. [29] [30]

Industrial hydrogen

Competing for biohydrogen, at least for commercial applications, are many mature industrial processes. Steam reforming of natural gas - sometimes referred to as steam methane reforming (SMR) - is the most common method of producing bulk hydrogen at about 95% of the world production. [31] [32] [33]

See also

Related Research Articles

<span class="mw-page-title-main">Photosynthesis</span> Biological process to convert light into chemical energy

Photosynthesis is a biological process used by many cellular organisms to convert light energy into chemical energy, which is stored in organic compounds that can later be metabolized through cellular respiration to fuel the organism's activities. The term usually refers to oxygenic photosynthesis, where oxygen is produced as a byproduct and some of the chemical energy produced is stored in carbohydrate molecules such as sugars, starch, glycogen and cellulose, which are synthesized from endergonic reaction of carbon dioxide with water. Most plants, algae and cyanobacteria perform photosynthesis; such organisms are called photoautotrophs. Photosynthesis is largely responsible for producing and maintaining the oxygen content of the Earth's atmosphere, and supplies most of the biological energy necessary for complex life on Earth.

<i>Chlamydomonas reinhardtii</i> Species of alga

Chlamydomonas reinhardtii is a single-cell green alga about 10 micrometres in diameter that swims with two flagella. It has a cell wall made of hydroxyproline-rich glycoproteins, a large cup-shaped chloroplast, a large pyrenoid, and an eyespot that senses light.

<span class="mw-page-title-main">Photosystem</span> Structural units of protein involved in photosynthesis

Photosystems are functional and structural units of protein complexes involved in photosynthesis. Together they carry out the primary photochemistry of photosynthesis: the absorption of light and the transfer of energy and electrons. Photosystems are found in the thylakoid membranes of plants, algae, and cyanobacteria. These membranes are located inside the chloroplasts of plants and algae, and in the cytoplasmic membrane of photosynthetic bacteria. There are two kinds of photosystems: PSI and PSII.

<span class="mw-page-title-main">Photosystem I</span> Second protein complex in photosynthetic light reactions

Photosystem I is one of two photosystems in the photosynthetic light reactions of algae, plants, and cyanobacteria. Photosystem I is an integral membrane protein complex that uses light energy to catalyze the transfer of electrons across the thylakoid membrane from plastocyanin to ferredoxin. Ultimately, the electrons that are transferred by Photosystem I are used to produce the moderate-energy hydrogen carrier NADPH. The photon energy absorbed by Photosystem I also produces a proton-motive force that is used to generate ATP. PSI is composed of more than 110 cofactors, significantly more than Photosystem II.

Artificial photosynthesis is a chemical process that biomimics the natural process of photosynthesis to convert sunlight, water, and carbon dioxide into carbohydrates and oxygen. The term artificial photosynthesis is commonly used to refer to any scheme for capturing and storing the energy from sunlight in the chemical bonds of a fuel. Photocatalytic water splitting converts water into hydrogen and oxygen and is a major research topic of artificial photosynthesis. Light-driven carbon dioxide reduction is another process studied that replicates natural carbon fixation.

<span class="mw-page-title-main">Water splitting</span> Chemical reaction

Water splitting is the chemical reaction in which water is broken down into oxygen and hydrogen:

A hydrogenase is an enzyme that catalyses the reversible oxidation of molecular hydrogen (H2), as shown below:

<span class="mw-page-title-main">Photoinhibition</span>

Photoinhibition is light-induced reduction in the photosynthetic capacity of a plant, alga, or cyanobacterium. Photosystem II (PSII) is more sensitive to light than the rest of the photosynthetic machinery, and most researchers define the term as light-induced damage to PSII. In living organisms, photoinhibited PSII centres are continuously repaired via degradation and synthesis of the D1 protein of the photosynthetic reaction center of PSII. Photoinhibition is also used in a wider sense, as dynamic photoinhibition, to describe all reactions that decrease the efficiency of photosynthesis when plants are exposed to light.

Anastasios Melis is a Greek-American biologist at the University of California, Berkeley who elucidated the possibility of creating hydrogen from algae. He is currently Professor of Plant & Microbial Biology in the institution and Editor-in-Chief of the Planta journal.

<i>Cupriavidus necator</i> Species of bacterium

Cupriavidus necator is a Gram-negative soil bacterium of the class Betaproteobacteria.

Hydrogen-oxidizing bacteria are a group of facultative autotrophs that can use hydrogen as an electron donor. They can be divided into aerobes and anaerobes. The former use hydrogen as an electron donor and oxygen as an acceptor while the latter use sulphate or nitrogen dioxide as electron acceptors. Species of both types have been isolated from a variety of environments, including fresh waters, sediments, soils, activated sludge, hot springs, hydrothermal vents and percolating water.

<i>Scenedesmus</i> Genus of green algae

Scenedesmus is a genus of green algae, in the class Chlorophyceae. They are colonial and non-motile. They are one of the most common components of phytoplankton in freshwater habitats worldwide.

<span class="mw-page-title-main">Ferredoxin hydrogenase</span> Class of enzymes

In enzymology, ferredoxin hydrogenase, also referred to as [Fe-Fe]hydrogenase, H2 oxidizing hydrogenase, H2 producing hydrogenase, bidirectional hydrogenase, hydrogenase (ferredoxin), hydrogenlyase, and uptake hydrogenase, is found in Clostridium pasteurianum, Clostridium acetobutylicum,Chlamydomonas reinhardtii, and other organisms. The systematic name of this enzyme is hydrogen:ferredoxin oxidoreductase

In enzymology, a hydrogen:quinone oxidoreductase (EC 1.12.5.1) is an enzyme that catalyzes the chemical reaction

<span class="mw-page-title-main">Photofermentation</span>

Photofermentation is the fermentative conversion of organic substrate to biohydrogen manifested by a diverse group of photosynthetic bacteria by a series of biochemical reactions involving three steps similar to anaerobic conversion. Photofermentation differs from dark fermentation because it only proceeds in the presence of light.

[NiFe] hydrogenase is a type of hydrogenase, which is an oxidative enzyme that reversibly converts molecular hydrogen in prokaryotes including Bacteria and Archaea. The catalytic site on the enzyme provides simple hydrogen-metabolizing microorganisms a redox mechanism by which to store and utilize energy via the reaction

<span class="mw-page-title-main">Chlororespiration</span>

Chlororespiration is a respiratory process that takes place within plants. Inside plant cells there is an organelle called the chloroplast which is surrounded by the thylakoid membrane. This membrane contains an enzyme called NAD(P)H dehydrogenase which transfers electrons in a linear chain to oxygen molecules. This electron transport chain (ETC) within the chloroplast also interacts with those in the mitochondria where respiration takes place. Photosynthesis is also a process that Chlororespiration interacts with. If photosynthesis is inhibited by environmental stressors like water deficit, increased heat, and/or increased/decreased light exposure, or even chilling stress then chlororespiration is one of the crucial ways that plants use to compensate for chemical energy synthesis.

Biological photovoltaics, also called biophotovoltaics or BPV, is an energy-generating technology which uses oxygenic photoautotrophic organisms, or fractions thereof, to harvest light energy and produce electrical power. Biological photovoltaic devices are a type of biological electrochemical system, or microbial fuel cell, and are sometimes also called photo-microbial fuel cells or “living solar cells”. In a biological photovoltaic system, electrons generated by photolysis of water are transferred to an anode. A relatively high-potential reaction takes place at the cathode, and the resulting potential difference drives current through an external circuit to do useful work. It is hoped that using a living organism as the light harvesting material, will make biological photovoltaics a cost-effective alternative to synthetic light-energy-transduction technologies such as silicon-based photovoltaics.

<i>Cyanothece</i> Genus of bacteria

Cyanothece is a genus of unicellular, diazotrophic, oxygenic photosynthesizing cyanobacteria.

<span class="mw-page-title-main">Wolfgang Lubitz</span> German chemist and biophysicist

Wolfgang Lubitz is a German chemist and biophysicist. He is currently a director emeritus at the Max Planck Institute for Chemical Energy Conversion. He is well known for his work on bacterial photosynthetic reaction centres, hydrogenase enzymes, and the oxygen-evolving complex using a variety of biophysical techniques. He has been recognized by a Festschrift for his contributions to electron paramagnetic resonance (EPR) and its applications to chemical and biological systems.

References

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