Abstract

<p indent="0mm">The thylakoid membrane, which contains photosynthetic pigments, lipids and proteins, is the main site where the photosynthetic light reaction of green plants, eukaryotic algae and cyanobacteria occurs. Water photolysis and photosynthetic electron transport are two core processes of the light reaction. The former is responsible for converting photons into electrons, and the latter then condenses the energy from the high-potential electrons into energy carriers such as adenosine triphosphate (ATP) and reduced nicotinamide adenine dinucleotide phosphate (NADPH). Due to the electron transport and energy conversion properties of the light reaction, thylakoid membrane can be used in biophotoelectrochemical and biophotoelectrocatalytic systems. Biophotoelectrochemical systems refer to direct or indirect extraction of electrons from the photosynthetic electron transfer chain (PETC) of the thylakoid membrane and transfer the electrons into an external electrode, eventually converting light energy into electric current. Biophotoelectrocatalytic systems refer to the use of ATP and NADPH generated in the light reaction of the thylakoid membrane to drive a series of ATP- or NADPH-dependent biocatalytic reactions. These thylakoid membrane-based light energy utilization systems are featured with material renewability and environmental compatibility. In addition, they show higher energy efficiency than that of the whole photosynthetic-cell systems. Furthermore, thylakoid membranes also hold clear advantages over sub-cellular photosystems, including retention of photosynthetic complexes in their natural environment and relatively simple and fast procedures for their extraction and purification. For biophotoelectrochemical systems, scientists have optimized the electron transfer between the thylakoid membrane and electrode, for example by developing different immobilization methods, or modifying electrode surface with redox polymers and nanomaterials. As an effective platform for cofactor regeneration, the thylakoid membrane-based biophotoelectrocatalytic systems have been used to drive the enzymatic synthesis of poly(3-hydroxybutyrate) (PHB) from acetate, as well as CO<sub>2</sub> fixation through a synthetic crotonyl-coenzyme A (CoA)/ethylmalonyl-CoA/hydroxybutyryl-CoA (CETCH) cycle. In addition to light energy utilization, thylakoid membranes can also be used in photodynamic therapy (PDT) of tumors. Firstly, the photosensitizer porphyrin in the chlorophyll molecules can produce singlet oxygen (<sup>1</sup>O<sub>2</sub>) from O<sub>2</sub> under visible light irradiation, which in turn kills the tumor cells. Secondly, the photosynthetic oxygen evolution of thylakoid membrane can rapidly supply O<sub>2</sub> <italic>in situ</italic>. Moreover, thylakoid membranes contain abundant catalase, which catalyzes the decomposition of H<sub>2</sub>O<sub>2</sub> into O<sub>2</sub>, thus further increasing the supply of O<sub>2</sub>. This paper summarizes recent progress in the research of thylakoid membranes in these three fields, focusing on the application forms and underlying mechanisms. Moreover, the difficulties and challenges in the application of thylakoid membrane are discussed, and future research directions are prospected. Specifically, developing protective structures such as hydrogels or artificial vesicles to address the issue of short half-life of isolated thylakoid membranes will be crucial. In light of their ingenious energy conversion characteristics and amenability for potential commercialization, thylakoid membrane-based biotechnological systems will have a bright future in the fields of bioenergy, biocatalysis and biomedicine.

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