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10/5/2013 LECTURE PRESENTATIONS For BROCK BIOLOGY OF MICROORGANISMS, THIRTEENTH EDITION Michael T. Madigan, John M. Martinko, David A. Stahl, David P. Clark Chapter 3 Lectures by John Zamora Middle Tennessee State University Cell Structure and Function in Bacteria and Archaea © 2012 Pearson Education, Inc. I. Cell Shape and Size • 3.1 Cell Morphology • 3.2 Cell Size and the Significance of Smallness © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 1 10/5/2013 3.1 Cell Morphology • Morphology = cell shape • Major cell morphologies (Figure 3.1) – Coccus (pl. cocci): spherical or ovoid – Rod: cylindrical shape – Spirillum: spiral shape • Cells with unusual shapes – Spirochetes, appendaged bacteria, and filamentous bacteria • Many variations on basic morphological types © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI Figure 3.1 Representative cell morphologies of prokaryotes Spirochete Coccus Coccus cells may also exist as short chains or grapelike clusters Stalk Rod Hypha Budding and appendaged bacteria Spirillum Filamentous bacteria © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 2 10/5/2013 3.1 Cell Morphology • Morphology typically does not predict physiology1, ecology, phylogeny2, etc. of a prokaryotic cell • Selective forces may be involved in setting the morphology – Optimization for nutrient uptake (small cells and those with high surface-to-volume ratio) – Swimming motility in viscous environments or near surfaces (helical or spiral-shaped cells) – Gliding motility (filamentous bacteria) 1 2 functions and activities of living organisms and their parts the evolutionary history of a group of organisms © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 3.2 Cell Size and the Significance of Smallness • Size range for prokaryotes: 0.2 µm to >700 µm in diameter – Most cultured rod-shaped bacteria are between 0.5 and 4.0 µm wide and <15 µm long – Examples of very large prokaryotes • Epulopiscium fishelsoni (Figure 3.2a) • Thiomargarita namibiensis (Figure 3.2b) • Size range for eukaryotic cells: 10 to >200 µm in diameter © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 3 10/5/2013 Figure 3.2 Some very large prokaryotes Epulopiscium fishelsoni Thiomargarita namibiensis © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 3.2 Cell Size and the Significance of Smallness • Surface-to-Volume Ratios, Growth Rates, and Evolution – Advantages to being small (Figure 3.3) • Small cells have more surface area relative to cell volume than large cells (i.e., higher S/V) – support greater nutrient exchange per unit cell volume – tend to grow faster than larger cells © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 4 10/5/2013 Figure 3.3 Surface area and volume relationships in cells r = 1 m r = 1 m Surface area (4r2) = 12.6 m2 4 Volume ( 3 r3) = 4.2 m3 Surface =3 Volume r = 2 m r = 2 m Surface area = 50.3 m2 Volume = 33.5 m3 Surface = 1.5 Volume © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 3.2 Cell Size and the Significance of Smallness • Lower Limits of Cell Size – Cellular organisms <0.15 µm in diameter are unlikely – Open oceans tend to contain small cells (0.2– 0.4 µm in diameter) © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 5 10/5/2013 II. The Cytoplasmic Membrane and Transport • 3.3 The Cytoplasmic Membrane • 3.4 Functions of the Cytoplasmic Membrane • 3.5 Transport and Transport Systems © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 3.3 The Cytoplasmic Membrane in Bacteria and Archaea • Cytoplasmic membrane: – Thin structure that surrounds the cell – 6-8 nm thick – Vital barrier that separates cytoplasm from environment – Highly selective permeable barrier; enables concentration of specific metabolites and excretion of waste products © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 6 10/5/2013 3.3 The Cytoplasmic Membrane • Composition of Membranes – General structure is phospholipid bilayer (Figure 3.4) • Contain both hydrophobic and hydrophilic components – Can exist in many different chemical forms as a result of variation in the groups attached to the glycerol backbone – Fatty acids point inward to form hydrophobic environment; hydrophilic portions remain exposed to external environment or the cytoplasm Animation: Membrane Structure Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI © 2012 Pearson Education, Inc. Figure 3.4 Phospholipid bilayer membrane Glycerol Fatty acids Phosphate Ethanolamine Hydrophilic General architecture region of a bilayer membrane; the blue Fatty acids Hydrophobic balls depict glycerol region with phosphate and Hydrophilic (or) other hydrophilic region groups. Glycerophosphates Fatty acids © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 7 10/5/2013 3.3 The Cytoplasmic Membrane • Cytoplasmic Membrane (Figure 3.5) – 6-8 nm wide – Embedded proteins – Stabilized by hydrogen bonds and hydrophobic interactions – Mg2+ and Ca2+ help stabilize membrane by forming ionic bonds with negative charges on the phospholipids – Somewhat fluid © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI Figure 3.5 Structure of the cytoplasmic membrane Out Phospholipids Hydrophilic groups 6–8 nm Hydrophobic groups In Integral membrane proteins © 2012 Pearson Education, Inc. Phospholipid molecule Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 8 10/5/2013 3.3 The Cytoplasmic Membrane • Membrane Proteins – Outer surface of cytoplasmic membrane can interact with a variety of proteins that bind substrates or process large molecules for transport – Inner surface of cytoplasmic membrane interacts with proteins involved in energy-yielding reactions and other important cellular functions © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 3.3 The Cytoplasmic Membrane • Membrane Proteins (cont’d) – Integral membrane proteins • Firmly embedded in the membrane – Peripheral membrane proteins • One portion anchored in the membrane © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 9 10/5/2013 3.3 The Cytoplasmic Membrane • Membrane-Strengthening Agents – Sterols • Rigid, planar lipids found in eukaryotic membranes Strengthen and stabilize membranes – Hopanoids • Structurally similar to sterols • Present in membranes of many Bacteria © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 3.3 The Cytoplasmic Membrane • Archaeal Membranes – Ether linkages in phospholipids of Archaea – Bacteria and Eukarya that have ester linkages in phospholipids – Can exist as lipid monolayers, bilayers, or mixture (Figure 3.7d and e) © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 10 10/5/2013 Figure 3.7d,e Membrane structure in Archaea may be bilayer or monolayer (or a mix of both) Out Glycerophosphates Phytanyl Membrane protein In Out Lipid bilayer Biphytanyl In Lipid monolayer © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 3.4 Functions of the Cytoplasmic Membrane (Figure 3.8) • Permeability Barrier – Polar and charged molecules must be transported – Transport proteins accumulate solutes against the concentration gradient • Protein Anchor – Holds transport proteins in place • Energy Conservation © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 11 10/5/2013 Figure 3.8 The major functions of the cytoplasmic membrane. Permeability barrier: Protein anchor: Energy conservation: Prevents leakage and functions as a gateway for transport of nutrients into, and wastes out of, the cell Site of many proteins that participate in transport, bioenergetics, and chemotaxis Site of generation and use of the proton motive force Although structurally weak, the cytoplasmic membrane has many important cellular functions. © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 3.5 Transport and Transport Systems • Carrier-Mediated Transport Systems (Figure 3.9) – Show saturation effect – Highly specific © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 12 10/5/2013 Rate of solute entry Figure 3.9 Transport versus diffusion. Transporter saturated with substrate Transport In transport, the uptake rate shows saturation at relatively low external concentrations Simple diffusion External concentration of solute © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 3.5 Transport and Transport Systems • Three transport events are possible: uniport, symport, and antiport (Figure 3.11) – Uniporters transport in one direction across the membrane – Symporters function as co-transporters – Antiporters transport a molecule across the membrane while simultaneously transporting another molecule in the opposite direction © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 13 10/5/2013 Figure 3.11 Structure of membrane-spanning transporters and types of transport events Out In Uniporter © 2012 Pearson Education, Inc. Antiporter Symporter Membranespanning transporters are made of 12 αhelices (each shown here as a cylinder) that aggregate to form a channel through the membrane. Shown here are three different transport events; for antiporters and symporters, the cotransported substance is shown in yellow. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI III. Cell Walls of Prokaryotes • 3.6 The Cell Wall of Bacteria: Peptidoglycan • 3.7 The Outer Membrane • 3.8 Cell Walls of Archaea © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 14 10/5/2013 3.6 The Cell Wall of Bacteria: Peptidoglycan Peptidoglycan (Figure 3.16) – Rigid layer that provides strength to cell wall – Polysaccharide composed of • • • • N-acetylglucosamine and N-acetylmuramic acid Amino acids Lysine or diaminopimelic acid (DAP) Cross-linked differently in gram-negative bacteria and gram-positive bacteria (Figure 3.17) © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI Figure 3.16 Cell walls of Bacteria. (a, b) Schematic diagrams of gram-positive and gram-negative cell walls. © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 15 10/5/2013 3.6 The Cell Wall of Bacteria: Peptidoglycan • Gram-Positive Cell Walls (Figure 3.18) – Can contain up to 90% peptidoglycan – Common to have teichoic acids (acidic substances) embedded in the cell wall • Lipoteichoic acids: teichoic acids covalently bound to membrane lipids © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 3.6 The Cell Wall of Bacteria: Peptidoglycan • Prokaryotes That Lack Cell Walls – Mycoplasmas • Group of pathogenic bacteria – Thermoplasma • Species of Archaea © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 16 10/5/2013 3.7 The Outer Membrane • Total cell wall contains ~10% peptidoglycan (Figure 3.20a) • Most of cell wall composed of outer membrane (lipopolysaccharide [LPS] layer) • Structural differences between cell walls of gram-positive and gram-negative Bacteria are responsible for differences in the Gram stain reaction © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI Figure 3.20a The gram-negative cell wall. Arrangement of lipopolysaccharide, lipid A, phospholipid, porins, and lipoprotein in the outer membrane O-polysaccharide Core polysaccharide Lipid A Protein Out Lipopolysaccharide (LPS) Porin Cell wall 8 nm Outer membrane Periplasm Peptidoglycan Phospholipid Lipoprotein Cytoplasmic membrane © 2012 Pearson Education, Inc. In Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 17 10/5/2013 3.8 Cell Walls of Archaea • No peptidoglycan • Typically no outer membrane • Pseudomurein – Polysaccharide similar to peptidoglycan Composed of N-acetylglucosamine and Nacetyltalosaminuronic acid – Found in cell walls of certain methanogenic Archaea • Cell walls of some Archaea lack pseudomurein © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI IV. Other Cell Surface Structures and Inclusions • • • • 3.9 Cell Surface Structures 3.10 Cell Inclusions 3.11 Gas Vesicles 3.12 Endospores © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 18 10/5/2013 3.9 Cell Surface Structures • Capsules and Slime Layers – Polysaccharide layers (Figure 3.23) • May be thick or thin, rigid or flexible – Assist in attachment to surfaces – Protect against phagocytosis – Resist desiccation © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI Figure 3.23 Bacterial capsules. Capsules of Acinetobacter species observed by phase-contrast microscopy after negative staining of cells with India ink. India ink does not penetrate the capsule, so the capsule appears as a light area surrounding the cell, which appears black. Transmission electron micrograph of a thin section of cells of Rhodobacter capsulatus with capsules (arrows) clearly evident; cells are about 0.9 µm wide. Cell Capsule Transmission electron micrograph of Rhizobium trifolii stained with ruthenium red to reveal the capsule. The cell is about 0.7 µm wide. © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 19 10/5/2013 3.9 Cell Surface Structures • Fimbriae – Filamentous protein structures (Figure 3.24) – Enable organisms to stick to surfaces or form pellicles (film) © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI Figure 3.24 Fimbriae Flagella Fimbriae Electron micrograph of a dividing cell of Salmonella typhi, showing flagella and fimbriae. A single cell is about 0.9 µm wide. © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 20 10/5/2013 3.9 Cell Surface Structures • Pili – – – – Filamentous protein structures (Figure 3.25) Typically longer than fimbriae Assist in surface attachment Facilitate genetic exchange between cells (conjugation) – Type IV pili involved in motility © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI Figure 3.25 Pili Viruscovered pilus The pilus on an Escherichia coli cell that is undergoing conjugation (a form of genetic transfer) with a second cell is better resolved because viruses have adhered to it. The cells are about 0.8 m wide. © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 21 10/5/2013 3.10 Cell Inclusions • Carbon storage polymers – Poly--hydroxybutyric acid (PHB): lipid (Figure 3.26) – Glycogen: glucose polymer • Polyphosphates: accumulations of inorganic phosphate (Figure 3.27) • Sulfur globules: composed of elemental sulfur • Magnetosomes: magnetic storage inclusions (Figure 3.28) Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI © 2012 Pearson Education, Inc. Figure 3.26 Poly-β-hydroxyalkanoates. Polyhydroxyalkanoate Electron micrograph of a thin section of cells of a bacterium containing granules of PHA. Nile red–stained cells of a PHA-containing bacterium. © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 22 10/5/2013 3.11 Gas Vesicles • Gas Vesicles – Confer buoyancy in planktonic cells (Figure 3.29) – Spindle-shaped, gas-filled structures made of protein (Figure 3.30) – Gas vesicle impermeable to water – Molecular Structure of Gas Vesicles • Gas vesicles are composed of two proteins: GvpA and GvpC • Function by decreasing cell density © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI Figure 3.29 Buoyant cyanobacteria. Flotation of gas-vesiculate cyanobacteria that formed a bloom in a freshwater lake, Lake Mendota, Madison, Wisconsin (USA). © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 23 10/5/2013 3.12 Endospores • Endospores – Highly differentiated cells resistant to heat, harsh chemicals, and radiation (Figure 3.32) – “Dormant” stage of bacterial life cycle (Figure 3.33) – Ideal for dispersal via wind, water, or animal gut – Only present in some gram-positive bacteria © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI Figure 3.32 The bacterial endospore. Terminal spores Subterminal spores Central spores Phase-contrast photomicrographs illustrating endospore morphologies and intracellular locations in different species of endospore-forming bacteria. Endospores appear bright by phase-contrast microscopy. © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 24 10/5/2013 Figure 3.33 The life cycle of an endospore-forming bacterium. Vegetative cell Developing spore Sporulating cell Mature spore The phase-contrast photomicrographs are of cells of Clostridium pascui. A cell is about 0.8 m wide. © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 3.12 Endospores • Endospore Structure (Figure 3.35) – – – – Structurally complex Contains dipicolinic acid Enriched in Ca2+ Core contains small acidsoluble proteins (SASPs) © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 25 10/5/2013 3.12 Endospores • The Sporulation Process – Complex series of events (Figure 3.37) – Genetically directed Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI © 2012 Pearson Education, Inc. Figure 3.37 Stages in endospore formation. Coat Growth Spore coat, Ca2 uptake, SASPs, dipicolinic acid Maturation, cell lysis Free endospore Stage VI, VII Germination Stage V Cortex Sporulation stages Vegetative cycle Cell division Cell wall Cytoplasmic membrane Asymmetric cell division; commitment to sporulation, Stage I Cortex formation Stage IV Prespore Septum Engulfment Mother cell Stage II Stage III Stages are defined from genetic and microscopic analyses of sporulation in Bacillus subtilis, the model organism for studies of sporulation. © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 26 10/5/2013 V. Microbial Locomotion • 3.13 Flagella and Motility • 3.14 Gliding Motility • 3.15 Microbial Taxes © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 3.13 Flagella and Motility • Flagellum (pl. flagella): structure that assists in swimming – Different arrangements: peritrichous, polar, lophotrichous – Helical in shape Animation: Flagella Arrangement © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 27 10/5/2013 3.13 Flagella and Motility • Flagella increase or decrease rotational speed in relation to strength of the proton motive force • Differences in swimming motions – Peritrichously flagellated cells move slowly in a straight line – Polarly flagellated cells move more rapidly and typically spin around © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 3.14 Gliding Motility • Gliding Motility – – – – – Flagella-independent motility Slower and smoother than swimming Movement typically occurs along long axis of cell Requires surface contact Mechanisms • Excretion of polysaccharide slime • Type IV pili • Gliding-specific proteins © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 28 10/5/2013 3.15 Microbial Taxes • Taxis: directed movement in response to chemical or physical gradients – – – – – Chemotaxis: response to chemicals Phototaxis: response to light Aerotaxis: response to oxygen Osmotaxis: response to ionic strength Hydrotaxis: response to water © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 3.15 Microbial Taxes • Chemotaxis – Best studied in E. coli – Bacteria respond to temporal, not spatial, difference in chemical concentration – “Run and tumble” behavior (Figure 3.47) – Attractants and receptors sensed by chemoreceptors © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 29 10/5/2013 Figure 3.47 Chemotaxis in a peritrichously flagellated bacterium such as Escherichia coli. Tumble Attractant Tumble Run Run No attractant present: Random movement. In the absence of a chemical attractant the cell swims randomly in runs, changing direction during tumbles. © 2012 Pearson Education, Inc. Attractant present: Directed movement In the presence of an attractant runs become biased, and the cell moves up the gradient of the attractant. The attractant gradient is depicted in green, with the highest concentration where the color is most intense. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 3.15 Microbial Taxes Measuring Chemotaxis (Figure 3.48) Measured by inserting a capillary tube containing an attractant or a repellent in a medium of motile bacteria Can also be seen under a microscope © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. Microbiology – Prof. BARIŞ ÇALLI 30 10/5/2013 Figure 3.48 Measuring chemotaxis using a capillary tube assay. (a) Insertion of the capillary into a bacterial suspension. As the capillary is inserted, a gradient of the chemical begins to form. (b) Control capillary contains a salt solution that is neither an attractant nor a repellent. Cell concentration inside the capillary becomes the same as that outside. Time course showing cell (c) Accumulation of bacteria in a capillary numbers in capillaries containing an attractant. containing various chemicals. (d) Repulsion of bacteria by a repellent. © 2012 Pearson Education, Inc. Marmara University – Enve303 Env. Eng. 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