Electron Transport Generates a Proton Gradient Across the Membrane
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4.3 the Light-Dependent 4B, 9B Photosynthesis Indetail 9B Transfers Energy
DO NOT EDIT--Changes must be made through “File info” CorrectionKey=B 4.3 Photosynthesis in Detail 4B, 9B KEY CONCEPT Photosynthesis requires a series of chemical reactions. VOCABULARY MAIN IDEAS photosystem The first stage of photosynthesis captures and transfers energy. electron transport chain The second stage of photosynthesis uses energy from the first stage to make sugars. ATP synthase Calvin cycle Connect to Your World In a way, the sugar-producing cells in leaves are like tiny factories with assembly lines. 4B investigate and explain cellular processes, including In a factory, different workers with separate jobs have to work together to put homeostasis, energy conversions, together a finished product. Similarly, in photosynthesis many different chemical transport of molecules, and synthesis of new molecules and 9B reactions, enzymes, and ions work together in a precise order to make the sugars compare the reactants and products that are the finished product. of photosynthesis and cellular respiration in terms of energy and matter MaiN IDEA 4B, 9B The first stage of photosynthesis captures and transfers energy. In Section 2, you read a summary of photosynthesis. However, the process is much more involved than that general description might suggest. For exam- ple, during the light-dependent reactions, light energy is captured and trans- ferred in the thylakoid membranes by two groups of molecules called photosystems. The two photosystems are called photosystem I and photosys- tem II. Overview of the Light-Dependent Reactions FIGURE 3.1 The light-dependent The light-dependent reactions are the photo- part of photosynthesis. During reactions capture energy from sun- light and transfer energy through the light-dependent reactions, chlorophyll and other light-absorbing electrons. -
Chapter 3 the Title and Subtitle of This Chapter Convey a Dual Meaning
3.1. Introduction Chapter 3 The title and subtitle of this chapter convey a dual meaning. At first reading, the subtitle Photosynthetic Reaction might seem to indicate that the topic of the structure, function and organization of Centers: photosynthetic reaction centers is So little time, so much to do exceedingly complex and that there is simply insufficient time or space in this brief article to cover the details. While this is John H. Golbeck certainly the case, the subtitle is Department of Biochemistry additionally meant to convey the idea that there is precious little time after the and absorption of a photon to accomplish the Molecular Biology task of preserving the energy in the form of The Pennsylvania State University stable charge separation. University Park, PA 16802 USA The difficulty is there exists a fundamental physical limitation in the amount of time available so that a photochemically induced excited state can be utilized before the energy is invariably wasted. Indeed, the entire design philosophy of biological reaction centers is centered on overcoming this physical, rather than chemical or biological, limitation. In this chapter, I will outline the problem of conserving the free energy of light-induced charge separation by focusing on the following topics: 3.2. Definition of the problem: the need to stabilize a charge-separated state. 3.3. The bacterial reaction center: how the cofactors and proteins cope with this problem in a model system. 3.4. Review of Marcus theory: what governs the rate of electron transfer in proteins? 3.5. Photosystem II: a variation on a theme of the bacterial reaction center. -
Cellular Biology 1
Cellular biology 1 INTRODUCTION • Specialized intracellular membrane-bound organelles (Fig. 1.2), such as mitochondria, Golgi apparatus, endoplasmic reticulum (ER). This chapter is an overview of eukaryotic cells, addressing • Large size (relative to prokaryotic cells). their intracellular organelles and structural components. A basic appreciation of cellular structure and function is important for an understanding of the following chapters’ information concerning metabolism and nutrition. For fur- ther detailed information in this subject area, please refer to EUKARYOTIC ORGANELLES a reference textbook. Nucleus The eukaryotic cell The nucleus is surrounded by a double membrane (nuclear Humans are multicellular eukaryotic organisms. All eukary- envelope). The envelope has multiple pores to allow tran- otic organisms are composed of eukaryotic cells. Eukaryotic sit of material between the nucleus and the cytoplasm. The cells (Fig. 1.1) are defined by the following features: nucleus contains the cell’s genetic material, DNA, organized • A membrane-limited nucleus (the key feature into linear structures known as chromosomes. As well as differentiating eukaryotic cells from prokaryotic cells) chromosomes, irregular zones of densely staining material that contains the cell’s genetic material. are also present. These are the nucleoli, which are responsible Inner nuclear Nucleus membrane Nucleolus Inner Outer Outer mitochondrial nuclear mitochondrial membrane membrane membrane Ribosome Intermembrane space Chromatin Mitochondrial Rough matrix Mitochondrial Nuclear endoplasmic ribosome pore reticulum Crista Mitochondrial mRNA Smooth Vesicle endoplasmic Mitochondrion Circular reticulum mitochondrial Proteins of the DNA Vesicle budding electron transport off rough ER Vesicles fusing system with trans face of Cytoplasm Golgi apparatus ‘Cis’ face + discharging protein/lipid Golgi apparatus ‘Trans’ face Lysosome Vesicles leaving Golgi with modified protein/lipid cargo Cell membrane Fig. -
Thylakoid Membrane Architecture in Cyanobacteria
THYLAKOID MEMBRANE ARCHITECTURE IN CYANOBACTERIA Dissertation zur Erlangung des Grades eines Doktors der Naturwissenschaften an der Fakultät für Biologie der Ludwig-Maximilians-Universität München vorgelegt von Anna Margareta Rast München, 03. Mai 2018 1. Gutachter: Prof. Dr. Jörg Nickelsen 2. Gutachter: Prof. Dr. Andreas Klingl Tag der Abgabe: 03.05.2018 Tag der mündlichen Prüfung: 22.06.2018 2 TABLE OF CONTENT TABLE OF CONTENT SUMMARY 5 ZUSAMMENFASSUNG 7 1 INTRODUCTION 9 1.1 Evolution of oxygenic photosynthesis 9 1.2 Thylakoid structure 10 1.2.1 Thylakoid structure in plastids 10 1.2.2 Thylakoid structure in cyanobacteria 11 1.3 Thylakoid membrane shaping factors 13 1.4 Photosynthesis in cyanobacteria 17 1.4.1 Electron transport chain 17 1.4.2 Light harvesting 19 1.4.3 Lateral heterogeneity 20 1.5 PSII biogenesis and repair in cyanobacteria 21 1.6 Cryo-electron tomography 23 2 AIMS OF THIS WORK 27 3 RESULTS 28 3.1 The role of Slr0151, a tetratricopeptide repeat protein from Synechocystis sp. PCC 6803, during photosystem II assembly and repair 28 3.2 Thylakoid membrane architecture in Synechocystis depends on CurT, a homolog of the granal CURVATURE THYLAKOID1 proteins 41 3.3 In situ cryo-electron tomography of cyanobacterial thylakoid convergence zones reveals a biogenic membrane connecting thylakoids to the plasma membrane. 65 4 DISCUSSION 86 4.1 Structural and photosystem II specific role of Slr0151 and CurT 86 4.1.1 The role of Slr0151 86 4.1.2 The role of CurT 88 4.1.3 Connection between Slr0151 and CurT via phosphorylation? 90 4.2 The thylapse – a contact area between thylakoids and plasma membrane 91 4.2.1 Factors possibly involved in thylapse formation 93 4.2.2 Membrane contact – a common feature 94 4.3 Lateral heterogeneity in cyanobacteria 95 4.4 Conclusions and future perspectives 97 3 TABLE OF CONTENT 5 REFERENCES 99 6 APPENDIX 112 6.1 Biogenesis of thylakoid membranes 112 6.2 Supplementary Material - The role of Slr0151, a tetratricopeptide repeat protein from Synechocystis sp. -
Passive and Active Transport
Passive and Active Transport 1. Thermodynamics of transport 2. Passive-mediated transport 3. Active transport neuron, membrane potential, ion transport Membranes • Provide barrier function – Extracellular – Organelles • Barrier can be overcome by „transport proteins“ – To mediate transmembrane movements of ions, Na+, K+ – Nutrients, glucose, amino acids etc. – Water (aquaporins) 1) Thermodynamics of Transport • Aout <-> Ain (ressembles a chemical equilibration) o‘ • GA - G A = RT ln [A] • ∆GA = GA(in) - GA(out) = RT ln ([A]in/[A]out) • GA: chemical potential of A o‘ • G A: chemical potential of standard state of A • If membrane has a potential, i.e., plasma membrane: -100mV (inside negative) then GA is termed the electrochemical potential of A Two types of transport across a membrane: o Nonmediated transport occurs by passive diffusion, i.e., O2, CO2 driven by chemical potential gradient, i.e. cannot occur against a concentration gradient o Mediated transport occurs by dedicated transport proteins 1. Passive-mediated transport/facilitated diffusion: [high] -> [low] 2. Active transport: [low] -> [high] May require energy in form of ATP or in form of a membrane potential 2) Passive-mediated transport Substances that are too large or too polar to diffuse across the bilayer must be transported by proteins: carriers, permeases, channels and transporters A) Ionophores B) Porins C) Ion Channels D) Aquaporins E) Transport Proteins A) Ionophores Organic molecules of divers types, often of bacterial origin => Increase the permeability of a target membrane for ions, frequently antibiotic, result in collapse of target membrane potential by ion equilibration 1. Carrier Ionophore, make ion soluble in membrane, i.e. valinomycin, 104 K+/sec 2. -
Evidence for a Respiratory Chain in the Chloroplast
Proc. NatL Acad. Sci. USA Vol. 79, pp. 4352-4356, July 1982 Cell Biology Evidence for a respiratory chain in the chloroplast (photosynthesis/respiration/starch degradation/evolution) PIERRE BENNOUN Institut de Biologie Physico-Chimique, 13, rue Pierre et Marie Curie, 75005, Paris, France Communicated by Pierre Joliot, April 12, 1982 ABSTRACT Evidence is given for the existence ofan electron in 20 ml of 20 mM N-tris(hydroxymethyl)methylglycine(Tri- transport pathway to oxygen in the thylakoid membranes ofchlo- cine)/KOH, pH 7.8/10 mM NaCl/10 mM MgCl2/1 mM K2- roplasts (chlororespiration). Plastoquinone is shown to be a redox HPO4/0.1 M sucrose/5% Ficoll. The cell suspension was carrier common to both photosynthetic and chlororespiratory passed through a Yeda press operated at 90 kg/cm2, diluted pathways. It is shown that, in dark-adapted chloroplasts, an elec- with 200 ml of Ficoll-lacking buffer, and centrifuged, and the trochemical gradient is built up across the thylakoid membrane pellet was suspended in the same buffer. by transfer of electrons through the chlororespiratory chain as Chlorophyll fluorescence kinetics and luminescence mea- well as by reverse functioning of the chloroplast ATPases. It is surements were performed as described (9). proposed that these mechanisms ensure recycling ofthe ATP and NAD(P)H generated by the glycolytic pathway converting starch into triose phosphates. Chlororespiration is thus an 02-uptake RESULTS process distinct from photorespiration and the Mehler reaction. The plastoquinone (PQ) pool ofchloroplast is a redox carrier of The evolutionary significance of chlororespiration is discussed. the photosynthetic electron transport chain. -
The Electrochemical Gradient of Protons and Its Relationship to Active Transport in Escherichia Coli Membrane Vesicles
Proc. Natl. Acad. Sci. USA Vol. 73, No. 6, pp. 1892-1896, June 1976 Biochemistry The electrochemical gradient of protons and its relationship to active transport in Escherichia coli membrane vesicles (flow dialysis/membrane potential/energy transduction/lipophilic cations/weak acids) SOFIA RAMOS, SHIMON SCHULDINER*, AND H. RONALD KABACK The Roche Institute of Molecular Biology, Nutley, New Jersey 07110 Communicated by B. L. Horecker, March 17, 1976 ABSTRACT Membrane vesicles isolated from E. coli gen- presence of valinomycin), a respiration-dependent membrane erate a trans-membrane proton gradient of 2 pH units under potential (AI, interior negative) of approximately -75 mV in appropriate conditions when assayed by flow dialysis. Using E. coli membrane vesicles has been documented (6, 13, 14). the distribution of weak acids to measure the proton gradient (ApH) and the distribution of the lipophilic cation triphenyl- Moreover it has been shown that the potential causes the ap- methylphosphonium to measure the electrical potential across pearance of high affinity binding sites for dansyl- and azido- the membrane (AI), the vesicles are shown to generate an phenylgalactosides on the outer surface of the membrane (4, electrochemical proton gradient (AiH+) of approximately -180 15) and that the potential is partially dissipated as a result of mV at pH 5.5 in the presence of ascorbate and phenazine lactose accumulation (6). Although these findings provide ev- methosulfate, the major component of which is a ApH of about idence for the chemiosmotic hypothesis, it has also been dem- -110 mV. As external pH is increased, ApH decreases, reaching o at pH 7.5 and above, while AI remains at about -75 mV and onstrated (6, 16) that vesicles are able to accumulate lactose and internal pH remains at pH 7.5. -
Chloroplast Genes Are Expressed During Intracellular Symbiotic
Proc. Natl. Acad. Sci. USA Vol. 93, pp. 12333-12338, October 1996 Cell Biology Chloroplast genes are expressed during intracellular symbiotic association of Vaucheria litorea plastids with the sea slug Elysia chlorotica (photosystem II reaction center/photosynthesis/chromophytic alga/ascoglossan mollusc/gene expression) CESAR V. MUJER*t, DAVID L. ANDREWS*t, JAMES R. MANHART§, SIDNEY K. PIERCES, AND MARY E. RUMPHO*II Departments of *Horticultural Sciences and §Biology, Texas A & M University, College Station, TX 77843; and IDepartment of Zoology, University of Maryland, College Park, MD 20742 Communicated by Martin Gibbs, Brandeis University, Waltham, MA, August 16, 1996 (received for review January 26, 1996) ABSTRACT The marine slug Elysia chlorotica (Gould) lowing metamorphosis from the veliger stage when juvenile forms an intracellular symbiosis with photosynthetically ac- sea slugs begin to feed on V litorea cells (1, 2). Once ingested, tive chloroplasts from the chromophytic alga Vaucheria litorea the chloroplasts are phagocytically incorporated into the cy- (C. Agardh). This symbiotic association was characterized toplasm of one of two morphologically distinct, epithelial cells over a period of 8 months during which E. chlorotica was (3) and maintain their photosynthetic function (1, 3). The deprived of V. litorea but provided with light and CO2. The fine plastids are frequently found in direct contact with the host structure of the symbiotic chloroplasts remained intact in E. cytoplasm as revealed by ultrastructural studies (3). In nature, chlorotica even after 8 months of starvation as revealed by the adult animal feeds on algae only sporadically, obtaining electron microscopy. Southern blot analysis of total DNA metabolic energy from the photosynthetic activity of the from E. -
Photosynthesis and Respiration
18 Photosynthesis and Respiration ATP is the energy currency of the cell Goal To understand how energy from sunlight is harnessed to Cells need to carry out many reactions that are energetically unfavorable. generate chemical energy by photosynthesis and You have seen some examples of these non-spontaneous reactions in respiration. earlier chapters: the synthesis of nucleic acids and proteins from their corresponding nucleotide and amino acid building blocks and the transport Objectives of certain ions against concentration gradients across a membrane. In many cases, unfavorable reactions like these are coupled to the hydrolysis of ATP After this chapter, you should be able to: in order to make them energetically favorable under cellular conditions; we • Explain the concepts of oxidation and have learned that for these reactions the free energy released in breaking reduction. the phosphodiester bonds in ATP exceeds the energy consumed by the • Explain how light energy generates an uphill reaction such that the sum of the free energy of the two reactions is electrochemical gradient. negative (ΔG < 0). To perform these reactions, cells must then have a way • Explain how an electrochemical of generating ATP efficiently so that a sufficient supply is always available. gradient generates chemical energy. The amount of ATP used by a mammalian cell has been estimated to be on the order of 109 molecules per second. In other words, ATP is the principal • Explain how chemical energy is harnessed to fix carbon dioxide. energy currency of the cell. • Explain how glucose is used to generate How does the cell produce enough ATP to sustain life and what is the source ATP anaerobically. -
Chloroplast Is the "Proteinaceous Shield" Regulating Photosystemii Electron Transport and Mediating Diuron Herbicide Sensitivity
Proc. Nati. Acad. Sci. USA Vol. 78, No. 3, pp. 1572-1576, March 1981 Biochemistry The rapidly metabolized 32,000-dalton polypeptide of the chloroplast is the "proteinaceous shield" regulating photosystem II electron transport and mediating diuron herbicide sensitivity (Spirodela/thylakoids/triazine/photosynthesis) AUTAR K. MATTOO*t, URI PICKO, HEDDA HOFFMAN-FALK*, AND MARVIN EDELMAN* Departments of *Plant Genetics and tBiochemistry, The Weizmann Institute of Science, Rehovot, Israel Communicated by Martin Gibbs, December 5, 1980 ABSTRACT Mild trypsin treatment of Spirodela oligorrhiza plex (LHCP) (14). In Spirodela this rapidly metabolized thy- thylakoid membranes leaks to partial digestion of the rapidly me- lakoid protein is translated by a discrete poly(A)- plastid mes- tabolized, surface-exposed, 32,000-dalton protein. Under these senger RNA of =500 conditions, photoreduction of ferricyanide becomes insensitive to kDal (15) into a 33.5-kDal precursor diuron [3-(3,4-dichlorophenyl)-1,1-dimethylurea], an inhibitor of molecule, which is speedily processed into the mature 32-kDal photosystem II electron transport. Preincubation of thylakoids form (13). Differentiated thylakoids are a prerequisite for 33.5- with diuron leads to a conformational change in the 32,000-dalton kDal protein synthesis (16). In addition, the whole process of protein, modifying its trypsin digestion and preventing expression 33.5/32-kDal synthesis, maturation, and degradation is under of diuron insensitivity. Finally, light affects the susceptibility of tight inductive control by light (17, 18). the 32,000-dalton protein to digestion by trypsin. In other exper- The iments, thylakoids specifically depleted in the 32,000-dalton pro- -rapidly metabolized 32-kDal thylakoid protein of Spi- tein were found to be deficient in electron transport at the re- rodela has its counterpart in other higher plants and algae. -
3. Transport Can Be Active Or Passive. •Passive Transport Is Movement
3. Transport can be active or passive. F 6-3 Taiz. Microelectrodes are used to measure membrane •Passive transport is movement down an electrochemical potentials across cell membrane gradient. •Active transport is movement against an electrochemical gradient. What is an electrochemical gradient? How is it formed? Passive and active transport of ions result in electric potential difference across membranes. •Movement of an uncharged mol Is dependent on conc. gradient alone. •Movement of an ion depends on the electric gradient and the conc. gradient. •Diffusion potential- Pump potential- How do you know if an ion is moving uphill or downhill? Nernst Eq What is the driving force for uphill movement? A) ATP ; b) H+ gradient 6-5. Pump potential and diffusion potential. How can we determine whether an ion moves in or out by active or passive transport? Nernst equation states that at equilibrium the difference in concentration of an ion between two compartments is balanced by the voltage difference. Thus it can predict the ion conc at equilibrium at a certain ΔE. Very useful to predict active or passive transport of an ion. Fig. 6-4, Taiz. Passive and active transporters. Tab 6-1, Taiz . Using the Nernst equation to predict ion conc. at equilibrium when the Cell electrical potential, Δψ = -110 mV ---------------------------------------------------------------------------------------- Ext Conc. Ion Internal concentration (mM) Summary: In general observed Nernst (Predicted) ---------------------------------------------------------------------------------------- Cation uptake: passive 1 mM K+ 75 mM 74 Cation efflux: active 1 mM Na+ 8 mM 74 1 mM Ca2+ 2 mM 5,000 Anion uptake: active 0.2 mM Mg2+ 3 1,340 Anion release: passive - 2 mM NO3 5 mM 0.02 1 Cl- 10 mM 0.01 - 1H2PO4 21 0.01 ---------------------------------------------------------------------------------------- 1 6-10. -
Inactivation of Mitochondrial Complex I Stimulates Chloroplast Atpase in Physcomitrella (Physcomitrium Patens)
bioRxiv preprint doi: https://doi.org/10.1101/2020.11.20.390153; this version posted November 20, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. Inactivation of mitochondrial Complex I stimulates chloroplast ATPase in Physcomitrella (Physcomitrium patens). Marco Mellon a, Mattia Storti a, Antoni Mateu Vera Vives a, David M. Kramer b, Alessandro Alboresi a and Tomas Morosinotto a a. Department of Biology, University of Padova, 35121 Padova, Italy b. MSU-DOE Plant Research Laboratory, Michigan State University, East Lansing, MI 48824, United States of America; Biochemistry and Molecular Biology, Michigan State University, East Lansing, MI 48824, United States of America. Corresponding author: Tomas Morosinotto, Dipartimento di Biologia, Università di Padova, Via Ugo Bassi 58B, 35121 Padova, Italy. Tel. +390498277484, Email: [email protected] Abstract While light is the ultimate source of energy for photosynthetic organisms, mitochondrial respiration is still fundamental for supporting metabolism demand during the night or in heterotrophic tissues. Respiration is also important for the metabolism of photosynthetically active cells, acting as a sink for excess reduced molecules and source of substrates for anabolic pathways. In this work, we isolated Physcomitrella (Physcomitrium patens) plants with altered respiration by inactivating Complex I of the mitochondrial electron transport chain by independent targeting of two essential subunits. Results show that the inactivation of Complex I causes a strong growth impairment even in fully autotrophic conditions in tissues where all cells are photosynthetically active.