Pierre BRICAGE Biologie, Faculté des Sciences, Université de Pau, FRANCE, - PDF

The Cell originated through Successive Outbreaks of Networking and Homing into Associations for the Mutual and Reciprocal Sharing of Advantages and of Disadvantages, between the Partners, with a Benefit

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The Cell originated through Successive Outbreaks of Networking and Homing into Associations for the Mutual and Reciprocal Sharing of Advantages and of Disadvantages, between the Partners, with a Benefit only for their Wholeness. 1 ( Les Associations à Avantages et Inconvénients Réciproques et Partagés. L'origine endosyncénotique de la cellule : avantages et inconvénients partagés entre partenaires indissociables et bénéfice global pour le nouveau tout émergeant ) Pierre BRICAGE Biologie, Faculté des Sciences, Université de Pau, FRANCE, Abstract : A living system is integrated into an ecoexotope. 2 The interactions between its ecoexotope and its endophysiotope shape its morphogenesis. In order to survive an organism has to transform disadvantages into advantages and to avoid that advantages become disadvantages . Alone survived Associations for the Mutual and Reciprocal Sharing of Advantages and of Disadvantages 3, in which each parcener is both a guest and an hostage. They originated from the juxtaposition and the encasement of previous systems. Each guest must maintain its identity into the new wholeness, through the fitness between the capacity of to be hosted of its endophysiotope and the capacity of hosting of its ecoexotope. This is allowed only through the simultaneous losses by each protagonist of the capacity to destroy the other one(s). Their whole's outcome results from their simultaneous metamorphoses. The comparison of the cell's organelles with the moneres and the study of the apoptosis, necrosis and tumourisation of the cell, make possible to resume the paradigm of the cell's phylogeny. The amalgamation, without membranes coalescing, of a population of side by side aggregated moneres, had made their endophysiotopes to become continuous, and was at the origin, of both the nuclear envelope and the endoplasmic reticulum. These new internal spaces were delimiting thus a continuous intermediary lumen, an ecophysiotope, which has played the role of a buffer between the ancient endophysiotope and the previous ecoexotope. It allowed a new network, the Golgi apparatus, to merge. The first need, in order to survive, is to eat and not to be eaten . Each organelle has its specific phages and descends from previous free-living moneres. The mitochondria and the plastids, which are delimited by two membranes, originated from the merging of Associations, for the Mutual and Reciprocal Sharing of Advantages and of Disadvantages, between preys and predators. The nude centrosome, which has the structure of a viral capsid, came from the melting of organisms with the same common membrane. The lysosomes and peroxisomes, with a single membrane, originated from the fusion of moneres having the same membrane interfaces. Résumé : Tout organisme est indissociable de son écoexotope dans lequel il est accueilli. Sa survie et son ontogénèse résultent des interactions entre son écoexotope et son endophysiotope. 4 Survivre c'est transformer des inconvénients en avantages et éviter que des avantages deviennent des inconvénients. Quel que soit le mode d'intégration d'un système vivant, seules survivent les Associations à Avantages et Inconvénients Réciproques et Partagés. 5 Leur mise en place émerge d'une adéquation entre la capacité d'être accueilli de l'endophysiotope et la capacité d'accueil de l'écoexotope. Elle est réalisée par la perte simultanée par chaque protagoniste de la capacité de détruire l'autre. La mise en place de l'organisation spatio-temporelle de la cellule, sa cancérisation, son apoptose ou sa nécrose, son ontogénèse par comparaison avec celle des monères, permettent de retrouver sa phylogénèse. L'accolement, puis la fusion hyaloplasmique, mais sans fusion membranaire, de monères apparentées dont l'endophysotiope devient ainsi 1 For the Complementary Data (Tables and Figures) and the Supplementary References, quoted * into the text, and for a more detailed plan, please do look at: 2 BRICAGE P. (2002a) The Evolutionary Shuttle of the Living Systems., 6 p. 5 th ECSS (Heraklion, Creta, Greece). 3 BRICAGE P. (2002b) Only sustainable development can ensure both care of the environment and intra-generational equity., 1p. Global Ethics for a Humane World, UNO & IISD globus conference (Utrecht, Netherlands). 4 BRICAGE P. (2001) La nature de la décision dans la nature? Systèmes biologiques: production, consommation, croissance et survie. Quelles règles? Quels degrés d exigence? Quels bilans?, 16 p. La décision. Coll. Afscet (Andé, France). 5 BRICAGE P. (2002c) Héritage génétique, héritage épigénétique et héritage environnemental: de la bactérie à l homme, le transformisme, une systémique du vivant., 28 p. Évolution du vivant & du social Coll. Afscet (Andé, France). 1 continu, est à l'origine, à la fois, de l'enveloppe nucléaire et du réticulum endoplasmique. 6 Leurs espaces internes, en continuité, délimitent un espace intermédiaire nouveau, qui joue le rôle d'un tampon entre le nouvel endophysiotope et le nouvel écoexotope: l'écophysiotope. La fusion résulte de leurs métamorphoses simultanées. L'appareil de Golgi, structure émergente unique, en constitue le réseau de coordination. Survivre c'est manger et ne pas être mangé pour se survivre. 7 Tout organite descend de monères à vie libre avec ses phages spécifiques. Délimités par une double membrane, mitochondries et plastes ont pour origine la mise en place d'associations à avantages et inconvénients réciproques et partagés entre proies et prédateurs, par pertes simultanées par chacun de la capacité de détruire l'autre. Non délimité par une membrane, le centrosome, dont la structure est celle d'une capside virale, provient de la fusion d'organismes délimités par la même membrane. Organites à une seule membrane, lysosomes et peroxysomes proviennent de la fusion de monères délimitées par la même interface membranaire ProKarya, the initial old Compartments 9 : Archaea and EuBacteria, the Moneres. A MON is made of a single CPT limited by 1 MB. MONs are able to resist to difficult EXOs, if agglomerated. An hyperosmotic stress induces the acidification of their ENDO and their sporulation. 1a. The membrane, interface between endophysiotope and ecoexotope, and the wall. The MB isolates from the EXO. Controlling the exchanges, it allows to avoid its DISs, while keeping its ADVs.* The W* doubles the MB and determines the form. Protoplasts, without W*, are very little resistant to osmotic shocks (what is a DIS), but their nude L deforms itself easily (what is an ADV). Aggregation or fusion between protoplasts* is triggered by a high ph, an osmotic shock or a VIR.* Gram- BACTs have one more MB containing POR.* Gram+ BACTs have only a 1X MB.* ARKs* do not sporulate. Only 1 PROT can control both the MB structure and the transfers of genes.* 1b. To survive it is To eat and not to be eaten . Ancient MONs survived from early organic compounds. Their first metamorphosis resulted from the depletion of nutriments and accumulation of waste products. BACT are eaten by phage VIRs. 1c. Emergence and level of organisation: the EoMonere, biodiversity and exaptation.* Each reaction is reversible.* The same structure can have several functions.* The phenotype is changing with the EXO changes. With the time, the common becomes particular and the rare common. Genes controlling metabolism and division of ARKs, BACTs and EUKs have a common ancestor (Figure 3).* ARKs* and MITs have cytochromes. ARKs and BACTs* have ferredoxines. ARKs, MITs and cbacts have a common ancestor. The EoMON was anaerobic and survived in a hot reducing EXO. Its progeny survived by acquiring the capacity to use O2. BACTs merged in a milder EXO.* The lack of W, a DIS in an hypoosmotic EXO, allows also the ingestion of preys that is an ADV.* The entry of H2O, by increasing the volume, is a simple means to escape the ingestion by a predator.* 6 BRICAGE P. (2003) Organisation, intégration et espace-temps des systèmes vivants., 31 p. Coll. Afscet (Andé, France). 7 BRICAGE P. (2000) La Survie des Organismes Vivants. Afscet Systémique & Biologie., 44 p. Fac. Médecine St Pères (Paris, France). 8 Mots clés: capacité d'accueil , capacité d'être accueilli , cellule, écoexotope, émergence, endophysiotope, métamorphose, mode d'intégration, monère, niveau d'organisation, ontogénèse et phylogénèse, virus, acteur clé-de-voûte : BRICAGE P. (2004) La gouvernance du vivant: les acteurs et les systèmes., 26 p. La gouvernance. Coll. Afscet (Andé, France). 9 Terms in use: abbreviations, definitions, & word roots according to CAILLEUX A. & KOMORN J. (1981) Dictionnaire des Racines Scientifiques., 263 p. CDU-SEDES (Paris, France). Archaea (ARK) from the Greek: archo first, archeo primitive, Bacterium (BACT) from the Greek: bacter baton, ecoexotope (EXO) from the Greek: exo external, tope space, eco of habitat, of welcome (BRICAGE P. 2002a), endophysiotope (ENDO) from the Greek: endo internal, tope space, physio of that grows and reproduces (Figure 1.) ecophysiotope (ECO): the space of interactions between the ecoexotope and the endophysiotope, (eo)monere (MON) from the Greek: mono unit, moner the simplest (HAECKEL 1866), eo of the early beginning, hyaloplasm & hyaloplasmic (HYA) from the Greek: plasm what is fashioned, integration from the Greek: integer whole, that makes only one with its ecoexotope, that is inseparable from it, Prokarya, prokaryotes (PROK), Eukarya, eukaryotes (EUK) from the Greek: pro before, karyo nucleus, eu true, symbiotes: the joined partners in a symbiotic association (symbiosis SYM), symbionte: the whole, the association. advantage: ADV, disadvantage: DIS, compartment: CPT 2 Without W, the hyperosmoticity of the EXO avoids the bursting and becomes an ADV. The acquisition of photosynthesis, metamorphosis that has allowed the surviving in an organic matter deprived EXO, is at the origin of cbacts. 10 This huge ADV loads a huge DIS, the release of O2. Its accumulation has forced the MONs to escape in anoxic EXOs or to new metamorphoses for the utilisation of this toxic waste. The acetic fermentation 11 and oxidations* have transformed 2 DISs, alcohol and toxic O2, into 1 ADV.* To survive (to grow) it is to transform disadvantages into advantages and to avoid that advantages become disadvantages , in order to survive itself (to reproduce).* 2. Simultaneous merging of the nucleus and the reticulum: the cell, a network of moneres. The CEL* is built with modules, limited by MBs, the organelles.* It merged from the fusion, without membranes coalescing, of a population of side by side aggregated moneres. The HYA, a reducing CPT, is their EXO. The today L contains a redox system, consumer of O2, which acidifies the HYA.* 2a. Organisation and integration: the cell is an endosyncoenosis. The HYA is specialised in the metabolism of sugars without O2.* The PER is an ubiquitous and pluripotent MON (see 4).* The MIT (see 3b) is a specialised MON that converts lipids and sugars in ATP.* The cplst (see 3c) is a MON that is specialised in sugars synthesis.* As MIT, it owns its nucleotides, proteins and lipids metabolisms. The NY is an emergent CPT (see 2c) that is specialised in nucleic acids synthesis and storage.* The ER and the GA (see 2d) are specialised in PROT metabolism.* 2b. Metabolism flows and membranes: the merging properties. The inhabitant organelles are protected form VIR* by the HYA, their inhabited MON of welcome* in which the abundance of H2O favours the hydrolysis, not the dehydrating syntheses. The metabolic pathways are juxtaposed and encased in the same manner that CPTs that contain them are. 12 2b1. The basic dynamics of predator/prey interactions. The phagocytosis is an emergent process associate to the SQ, lacking in PROK.* The integration of the MIT into the CEL is posterior to SQ acquisition. Regularly, protagonists reply to an attack by a defence and to a stronger defence by an again stronger attack. The no-resolution of the conflict, cause and consequence of the natural selection *, starts an escalation of ripostes with a cost always greater.* How to end to a cheap stable steady state which is imperative to optimise the survival? 2b2. The control of proteins flows between the hyaloplasm and the organelles. A PROT, which is aimed at an organelle, contains a signal of transportation and entry. A MB translocator controls its passage. A protease removes the signal, which is a source of AAs that allows the organelle to eat even in hyperosmoticity.* VIR PROTs are the same way sent to NY and MITs. 2b3. The internal membranes originated from moneres plasma membranes.* MIT OMB and ER MB have the same lipid contents.* NY MB and ER MB have a common ancestor, GA MB and ER MB also, MIT IMB and L of Gram- BACTs have another one, LYS MB and EUK L have a common ancestor.* NY IMB and EUK L* have the same properties of endocytosis, MIT IMB and VAC MB have the same ATPases VIR receptors* EUK L, ER MB, and NY OMB carry the same ATP dependent proteases, absent of cplst and MIT OMBs (Tables* A1 & B1) 2c. The nucleus: the actors and the system.* 10 Other abbreviations of terms in use: cellular level: apoptosis/tic APO, cell/cellular CEL, centrosome CENT, Cyanobacteria cbacts, cytoskeleton SQ, endosome END, envelope ENV, Golgi apparatus GA, glyoxysome GLY, lysosome LYS, stroma MAT, membrane MB, outer MB OMB, inner MB IMB, single membrane 1X MB, double membrane 2X MB, membrane of sequestration SQMB, mitochondria MITs, nucleus/ar NY, nucleolus/ar NU, skeletal wall W, peroxysome PER, plasma membrane L, vacuolar membrane T, microsome MICR, microtubule TUB, phagosome PHAG, plastid PLST, amyloplaste aplst, chloroplast cplst, perforine PERF, porine POR, aquaporine apor, endoplasmic reticulum ER (smooth LER or granular RER), thylakoïd THYL, vacuole/ar VAC, virus/al VIR molecular level: amino acid AA, calcium Ca++, carbon dioxide gas CO2, electron e-, enzyme/atic ENZ (CAT catalase, OXY oxidase, PEROX peroxidase, SuperOxydeDismutase SOD), proton(s) H+, water H2O, hydrogen peroxide H2O2, dioxygen gas O2, protein/aceous PROT. 11 TORTORA G.J. & al. (2004) Microbiology., 924 p. Pearson (San Francisco, USA). 12 De DUVE C. (1987) Une visite guidée de la cellule vivante., 437 p. De Boeck Université (Bruxelles, Belgique).* 3 Containing at least 1 NU, that can reach the size of a MON, the NY is delimited with a 2X MB pierced with pores. Each pore is a place of junction between OMB and IMB and a place of exchanges between NY ENDO and HYA ENDO. With its symmetry and dimensions it looks like a VIR capsid. 2c1. The membrane interface between endophysiotope and ecoexotope. The NY OMB is in continuity with the ER MBs.* The NY MBs and the L are homologous. The periny and intraer spaces are equivalent.* They built an interactive space in continuity between the EXO and the ENDO, the ECO, which is both more and less than the ENDO and EXO. The common origin of NY IMB and L, and of NY OMB, ER MBs and L is proved by their reassembly from the ER 13 after mitosis.* They carry the same VIR domains (Table B2*). The NY OMB and L behave the same manner during the HIV induced fusion.* The NY pores are intermb specialised opened 14 junctions, through which actin 15 filaments extend. The NY and HYA contain the same SQ. TUBs are attached to the L and to the NY OMB. Nevertheless, the interface is mobile. It is a slipping surface that allows the NY, coated with its IMB alone, to turn on itself. The pores can be broken and rebuilt. The intraer space is a slipping surface also (Figure 2a). The simultaneous emergence of the central NY and the outlying ER results from a position effect within an aggregate of prearks. 2c2. Like the cell, the nucleus is also a polymonere endosyncoenosis. The NY of the Echinides coelomocytes* contains inclusions of MON type: - organising centres with a 2X MB, a PROT canvas, crystalline like a PER, containing RNA, that migrate between NY and HYA, - globules analogous to the promits, forming in the NY and migrating to the HYA, by dilations of the 2X MB NY ENV. The NU with its RNA nucleoid and its plasmatic heterogeneity, has also a PROK organisation, but without MB.* It happens as if the NU was an eomonere with an RNA genome and the NY a polyeomonere endosyncoenosis, with its own phages*, initially of the RNA sort. 16 2d. Endoplasmic Reticulum, Golgi Apparatus and Centrosome are indivisible. The ER represents 50 % of the cell s MB surface. Like the MIT, it insures the sequestration of Ca++ and detoxication processes through OXY. It surrounds, and isolates from the ENDO, the senescent organelles, in digestive VACs. The GA and ER*, specialised parts* of the ECO, have the same role of convergence and distribution 17 like the intermb space of Gram-. The GA grows by extension and fission of a previous GA. During the fusion of 2 CELs, the GAs fuse in only 1 structure, as CENTs do. 18 The NY OMB, ER and GA have the same dynamics.* The GA, PER and CENT may be formed de novo, while being selfdividing. Their enclosed dangers * reveal the origin of the organelles.* Each one has its own phages. VIRs built and bud on NY IMB, ER MB and GA. A unique PROT allows the MB fusion of VIRs and the fusion of CELs into a syncytium with only 1 L*, this indicating the common origin of these MBs. There are so much potential hosts for a VIR that CPTs are (Table C).* The integrity of GA, CENT and NY is depending on the SQ. 19 Their movements and the maintenance of the shape and movement of the CEL is the duty of actin or tubulin filaments. The TUBs are formed of PROT fibres arranged in a repeated motive delimiting an internal canal. This topology is that of a VIR, like the tobacco mosaic VIR. The CENT resembles a VIR with 2 nude empty capsides.* 3. The Actors and the System: About the Origin of the Mitochondria and Plastids. MITs and PLSTs possess a genome. Their OMB and IMB contain the same PROTs, but more concentrated in the simplified OMBs, specialised in the control of H2O and solutes flows. 3a. The membrane, interface of confrontation and reciprocal constrained exchanges. In anaerobiosis, the ADV of the absorption of sugars by the BACT* L or CEL* L, is associated with a DIS, an entering flow of deleterious H+. the excretion of which is ATP consuming. 13 BURKE B. & ELLENBERG J. (2002) Remodelling the walls of the nucleus., p Nature Rev. Mol. Cell Biol. n HOELZ A. & BLOBEL G. (2004) Cell biology: popping out of the nucleus., p Nature n LÉVY N. & CAU P. (2003) Anomalies du noyau et maladies génétiques., p Pour La Science n Do look at: and at 17 SMALLRIDGE R. (2002) Endocytosis. A new recycling route., p 315. Nature Rev. Mol. Cell Biol. n PELLETIER L. & al. (2002) Golgi biogenesis in Toxoplasma gondii., p Nature n CAMERA P. & al. (2003) Citron-N is a neuronal Rho-associated protein involved in Golgi organization through actin cytoskeleton regulation., p Nature Cell Biol. n 5. 4 3a1. The mitochondrion is an hostage trapped into the hyaloplasm. The LYS, END or MIT ph regulative ATPases/synthases are always turned inwards the ENDO. They have the same reversible functioning, whatever is the MB.* Alone factors of regulation or modulation differ. But, if the Ca++ATPase of the L ejects outside in the EXO the toxic Ca++, that of the MIT is forced to pump it in its own ENDO.* That is a DIS. But, in the presence of O2, the excretion of H+ is linked with the IMB chain of e- transport and allows the synthesis of ATP. 2 DISs become 1 ADV. To survive in oxygenated and acidic EXO, alone the IMB is necessary. To what serves the OMB? The MIT ATP is exchanged against carbohydrates, lipids and PROT from the HYA. The preprots entry, through sites of contact between OMB and IMB equalling to NY pores, and their activation is ATP consuming, but this DIS is compensated by the ADV of the digestion of PROTs cleaved part. PROTs analysis indi
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