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Mitochondrial Physiology 1. Mitochondria and Bioblasts MiP1.Mitochondria and bioblasts_BEC-pre01 (2020-05-05) https://doi:10.26124/bec:2020-0001.vnn 1 2 Consortium communication 3 4 Mitochondrial physiology 5 1. Mitochondria and bioblasts 6 7 MitoEAGLE Task Group* 8 9 Living Communication: from Gnaiger et al (2020) Bioenerg Commun 2020.1. 10 11 Overview 12 Richard Altmann (1894): 13 The protoplasm is a colony of bioblasts. 14 Microorganisms and granula are at an 15 equivalent level and represent elementary 16 organisms, which are found wherever living 17 forces are acting, thus we want to describe 18 them by the common term bioblasts. In the 19 bioblast, that morphological unit of living 20 matter appears to be found. 21 Mitochondria are oxygen-consuming 22 electrochemical generators that evolved 23 from the endosymbiotic 24 alphaproteobacteria which became 25 integrated into a host cell related to Asgard 26 Archaea (Margulis 1970; Lane 2005; Roger 27 et al 2017). Richard Altmann (1894) 28 described the ‘bioblasts’, which include not 29 only the mitochondria as presently defined, 30 but also symbiotic and free-living bacteria. 31 The word ‘mitochondria’ (Greek mitos: 32 thread; chondros: granule) was introduced 33 by Carl Benda (1898). Mitochondrion is 34 singular and mitochondria is plural. 35 Abbreviation: mt, as generally used in 36 mtDNA. 37 Given the multiple roles of 38 mitochondria, it is perhaps not surprising 39 that mitochondrial dysfunction is 40 associated with a wide variety of genetic 41 and degenerative diseases. Robust 42 mitochondrial function is supported by 43 physical exercise and caloric balance, and is 44 central for sustained metabolic health 45 throughout life. Therefore, a more 46 consistent set of definitions for 47 mitochondrial physiology will increase our 48 understanding of the etiology of disease and 49 improve the diagnostic repertoire of mitochondrial medicine with a focus on protective medicine, 50 evolution, lifestyle, environment, and healthy aging. 51 52 Updates: 53 https://www.bioenergetics-communications.org/index.php/BEC2020.1_doi10.26124bec2020-0001.v1 54 www.bioenergetics-communications.org 1 of 10 Consortium Communication 55 Abstract 56 57 58 Keywords: 59 60 61 Reference: Mitochondrial physiology. 2. Respiratory states and rates 62 63 64 Mitochondrial structure-function relationships 65 ‘For the physiologist, mitochondria afforded the first opportunity for an experimental approach 66 to structure-function relationships, in particular those involved in active transport, vectorial 67 metabolism, and metabolic control mechanisms on a subcellular level’ (Ernster and Schatz 1981). 68 69 Contrary to current textbook dogma, which describes mitochondria as individual organelles, 70 mitochondria form dynamic networks within eukaryotic cells. Mitochondrial movement is 71 supported by microtubules. Mitochondrial size and number can change in response to energy 72 requirements of the cell via processes known as fusion and fission; these interactions allow 73 mitochondria to communicate within a network (Chan 2006). Mitochondria can even traverse cell 74 boundaries in a process known as horizontal mitochondrial transfer (Torralba et al 2016). 75 Another defining morphological characteristic of mitochondria is the double membrane. The 76 mitochondrial inner membrane (mtIM) forms dynamic tubular to disk-shaped cristae that 77 separate the mitochondrial matrix, i.e., the negatively charged internal mitochondrial 78 compartment, from the intermembrane space; the latter being enclosed by the mitochondrial 79 outer membrane (mtOM) and positively charged with respect to the matrix. 80 Intracellular stress factors may cause shrinking or swelling of the mitochondrial matrix that 81 can ultimately result in permeability transition (mtPT; Lemasters et al 1998). The mtIM contains 82 the non-bilayer phospholipid cardiolipin, which is also involved in the mtOM (Gebert et al 2009) 83 but is not present in any other eukaryotic cellular membrane. Cardiolipin has many regulatory 84 functions (Oemer et al 2018); it promotes and stabilizes the formation of supercomplexes 85 (SCInIIInIVn) based on dynamic interactions between specific respiratory complexes (McKenzie et 86 al 2006; Greggio et al 2017; Lenaz et al 2017), and it supports proton transfer on the mtIM from 87 the electron transfer system to F1FO-ATPase (ATP synthase; Yoshinaga et al 2016). The mtIM is 88 plastic and exerts an influence on the functional properties of incorporated proteins (Waczulikova 89 et al 2007). 90 Mitochondria constitute the structural and functional elementary components of cell 91 respiration (Figure 1). Mitochondrial respiration is the reduction of molecular oxygen by electron 92 transfer coupled to electrochemical proton translocation across the mtIM. In the process of 93 OXPHOS, the catabolic reaction of oxygen consumption is electrochemically coupled to the 94 transformation of energy in the phosphorylation of ADP to adenosine triphosphate (ATP; Mitchell 95 1961, 2011). Mitochondria are the powerhouses of the cell that contain the machinery of the 96 OXPHOS-pathways, including transmembrane respiratory complexes (proton pumps with FMN, 97 Fe-S and cytochrome b, c, aa3 redox systems); alternative dehydrogenases and oxidases; the 98 coenzyme ubiquinone (Q); F1FO-ATPase or ATP synthase; the enzymes of the tricarboxylic acid 99 cycle (TCA), fatty acid and amino acid oxidation; transporters of ions, metabolites and co-factors; 100 iron/sulphur cluster synthesis; and mitochondrial kinases related to catabolic pathways. TCA 101 cycle intermediates are vital precursors for macromolecule biosynthesis (Diebold et al 2019). The 102 mitochondrial proteome comprises over 1,200 proteins (Calvo et al 2015; 2017), mostly encoded 103 by nuclear DNA (nDNA), with a variety of functions, many of which are relatively well known, e.g., 104 proteins regulating mitochondrial biogenesis or apoptosis, while others are still under 105 investigation, or need to be identified, e.g., mtPT pore and alanine transporter. The mammalian 106 mitochondrial proteome can be used to discover and characterize the genetic basis of 107 mitochondrial diseases (Williams et al 2016; Palmfeldt and Bross 2017). 108 2 of 10 NN et al (202n) Bioenerg Commun 202n.nn BEC202n.nn doi:10.26124/bec:2020-0001.vnn 109 Mitochondrial crosstalk 110 111 Numerous cellular processes are orchestrated by a constant crosstalk between mitochondria and 112 other cellular components. For example, the crosstalk between mitochondria and the endoplasmic 113 reticulum is involved in the regulation of calcium homeostasis, cell division, autophagy, 114 differentiation, and anti-viral signaling (Murley and Nunnari 2016). Mitochondria contribute to 115 the formation of peroxisomes, which are hybrids of mitochondrial and ER-derived precursors 116 (Sugiura et al 2017). Cellular mitochondrial homeostasis (mitostasis) is maintained through 117 regulation at transcriptional, post-translational and epigenetic levels (Ling and Rönn 2018; 118 Lisowski et al 2018), resulting in dynamic regulation of mitochondrial turnover by biogenesis of 119 new mitochondria and removal of damaged mitochondria by fusion, fission and mitophagy (Singh 120 et al 2018). Cell signalling modules contribute to homeostatic regulation throughout the cell cycle 121 or even cell death by activating proteostatic modules, e.g., the ubiquitin-proteasome and 122 autophagy-lysosome/vacuole pathways; specific proteases like LON, and genome stability 123 modules in response to varying energy demands and stress cues (Quiros et al 2016). In addition, 124 several post-translational modifications, including acetylation and nitrosylation, are capable of 125 influencing the bioenergetic response, with clinically significant implications for health and 126 disease (Carrico et al 2018). 127 128 The mitochondrial genome 129 130 Mitochondria of higher eukaryotes typically maintain several copies of their own circular genome 131 known as mitochondrial DNA (mtDNA; hundred to thousands per cell; Cummins 1998), which is 132 maternally inherited in many species. However, biparental mitochondrial inheritance is 133 documented in some exceptional cases in humans (Luo et al 2018), is widespread in birds, fish, 134 reptiles and invertebrate groups, and is even the norm in some bivalve taxonomic groups (Breton 135 et al 2007; White et al 2008). The mitochondrial genome of the angiosperm Amborella contains a 136 record of six mitochondrial genome equivalents acquired by horizontal transfer of entire 137 genomes, two from angiosperms, three from algae and one from mosses (Rice et al 2016). In 138 unicellular organisms, i.e., protists, the structural organization of mitochondrial genomes is highly 139 variable and includes circular and linear DNA (Zíková et al 2016). While some of the free-living 140 flagellates exhibit the largest known gene coding capacity, e.g., jakobid Andalucia godoyi mtDNA 141 codes for 106 genes (Burger et al 2013), some protist groups, e.g., alveolates, possess 142 mitochondrial genomes with only three protein-coding genes and two rRNAs (Feagin et al 2012). 143 The complete loss of mitochondrial genome is observed in the highly reduced mitochondria of 144 Cryptosporidium species (Liu et al 2016). Reaching the final extreme, the microbial eukaryote, 145 oxymonad Monocercomonoides, has no mitochondrion whatsoever and lacks all typical nuclear- 146 encoded mitochondrial proteins, showing that while in 99 % of organisms mitochondria play a 147 vital role, this organelle is not indispensable (Karnkowska
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