Ch. 3 - Cell Structure & Function in Bacteria & Archae (Ch. 4 -- 12 ed.) Flashcards

1
Q

Central Dogma of Molecular Bio

(pp. 176, 252)

A
  • Genetic material in the cell specifies the organization of the cell & its activity
  • Molecular processes (underlying genetic info flow) can be dividied into 3 stages:
  1. Replication - DNA is duplicated, producing two double helices
  2. Transcription - DNA participates in protein synthesis through an RNA intermediate (mRNA)
  • -> some genes contain info for other types of RNA: tRNA & rRNA
    3. Translation - sequence of amino acids in a polypeptide (making up a protein) is determined by the specific sequence of bases in mRNA

–> In Eukaryotes…each gene transcribed to give a single mRNA

–> in Prokaryotes…a signle mRNA may carry genetic info from several genes

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2
Q

Mimi Virus exception(s)?

A
  • Viruses contain only a single form of nucleic acid – either DNA or RNA

–> Mimi virus contains both RNA & DNA

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3
Q

Major Cell Morphologies

A

–> several morphologies are known among prokaryotes…

  • Coccus – spherical or ovoid
  • Bacillus (rod) – cylindrical shape
  • Spirilla – some rods twist into spiral shapes

–> several groups possess unusual shapes…

  • Spirochets – tightly coiled shape
  • Appendaged – possess extensions of their cells (hypha/stalks)
  • Filamentous bacteria – long, thin cells/chains
  • Vibrio – half-moon shaped

**Cell morphology is easily recognized…but generally a poor predictor of a cell’s other properties**

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4
Q

Describe Cell Size of Prokaryotes

  • Esherichia coli?
  • Thiomargarita namibiensis?
  • Mycoplasma pneumoniae
A
  • Prokaryotes vary in size from ~ 0.2 µm to ~ >700 µm (in diameter)
  • -> 1 µm (micrometers) = 10-6 m (meters)
  • E. coli = 2 µm
  • -> considered base volume for prokaryotic cells
  • -> E. coli vol. = 2 µm3 –> (EC vol. = 1)
  • T. namibiensis = 750 µm
  • -> T. namibiensis vol. = 200,000,000 µm3 –> (EC vol. = 108
  • M. pneumonia = 0.2 µm
  • -> M. pneuonmia vol. = 0.005 µm –> (EC vol = 2.5 x 10-3)
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5
Q

Significance of Small size of Prokaryotes

A
  • prokaryotic cells generally very small cells compared to eukaryotes
  • -> very large…not common
  • rate at which nutrients/waste products transport in/out of cell is generally **inversely proportional to cell size**
  • -> higher S/V (surface are/volume) ratio of smaller cells supports greater nutreint exchange per unit of cell volume…vica versa…
  • -> cell’s growth rate depends (among other important things) on the rate of nutrient exchange

**S/V = 3/r** (for spherical coccus..)

⇒ larger the cell…smaller the S/V ratio –> results in a lesser efficient exchange with environment

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6
Q

Structure of Viruses

A

Viruses come in numerous sizes & shapes…most smaller than prokaryotic cellsg

  • Acellular…thus “nonliving”
  • -> rely on host cells to provide E & materials needed for replicationg their genomes & synthesizing their proteins…**Obligate intracellular parasites**
  • Nucleic acids – contain either DNA or RNA.. (NOT both)
  • -> (exception)…Mimi Virus contains both DNA & RNA
  • Capsids – protein shell which surrounds nucelic acid of virion
  • -> made up of Capsomers (structural subunits)
  • -> Nucelocapsid – complete complex of nucelic acid & protein packaged in virion
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7
Q

Viriods

A
  • infectious RNA molecules that differ from viruses…lack capsids (protein coat)
  • consist of single-stranded (covalently-closed) circular RNA that forms a seemingly double-stranded structure by intra-strand base pairing
  • -> makes it stable to exist outide of host cell… (NOT an obligate intracellular parasite)
  • -> enters through wound in plant cells… (does not use a receptor to enter host like viruses)
  • **interferes with regulatory RNA in plants**…
  • -> viriods possess no protein-encoding genes…basically totally dependent on host function (for replication)
  • -> viroid replicated in host cell nucleus or chloroplast by plant RNA polymerases
  • NO animal diseases known
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8
Q

Prions

A
  • thought to be small replicating polypeptides
  • -> contain neither RNA or DNA
  • Mechanism of Prion misfolding:
  • -> neuronal cells produce normal form of prion protein (PrPC)
  • -> Pathogenic form (PrPSc) catalyzes refolding of normal prions into abnormal form… (PrPC into PrPSc)
  • PrPSc is protease resistant & insoluble…aggregates in host neural cells (brain cells)
  • infectious in animals only
  • -> Mad Cow Disease in cattle
  • -> Creutzfeldt-Jakob disease (CJD) in humans
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9
Q

General structure of Phospholipid Bilayer

A
  • main constituent of cytoplasmic membrane
  • -> separates cytoplasm from environment
  • -> highly selective permeability barrier
  • -> if broken…integrity of cell is destroyed
  • possess both hydrophobic (fatty acids) & hydrophilic (glycerol-phosphate) components
  • -> hydrocarbon chains (of fatty acid) points inward
  • -> glycerol-phosphates point outward toward environment & cytoplasm
  • 6-8 nm wide
  • viewed from electron microscope…appears as 2 light-colored lines separated by a darker area
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10
Q

Describe in-depthly composition of Phospholipids
–> chemical make-up…

A
  • consists of hydrophobic fatty acids (hydrocarbon chains) *esterified* to glycerol
  • -> **removal of H20 between alcohol group (from glycerol) & caroxyl group (from 2 fatty acids OR 1 phosphoric acid)**
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11
Q

Cytoplasmic membrane Proteins
–> 2 types…

A
  1. Peripheal Proteins (extrinsic proteins)
  • not embedded…but still associated with membrane surface
  • some easily removed from changes in environment (pH, ionic strenght, etc…)
  • typically on surface of membrane
  • some may be bound to integral proteins…important cellular processes (energy metabolism, transport, etc…)
  • -> some are *Lipoproteins*… proteins containing lipid tail that can anchor integral proteins
  1. Integral Proteins (intrinsic proteins)
  • firmly embedded in membrane
  • many span entire membrane but not all…possessing external & internal surfaces
  • -> must be *ampipathic* (both hydrophilic & hydrophobic regions)…hydrophobic inside membrabe & hydrophilic internal/external surfaces of membrane

**Cytoplasmic proteins are arranged in clusters** (NOT being distributed evenly)

  • -> allows grouping of proteins that interact or that have similar function
  • -> Lipid bilayer varies in thickness (from 6 - 8 nm) to accommodate various patches of proteins
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12
Q

Cytoplasmic Membrane Function

A
  1. Permeability Barrier
  • prevents leakage (which would distruct integrity of cell)
  • functions as a gateway for transport of nutrients into and out of cell
  1. Protein Anchor
    - site of many proteins involveed in transport, bioenergetics, signaling, & chemotaxis
  2. Energy Conservation
  • site of generation & use of proton motive force
  • -> *Proton Motor Force* – membrance possesses a charge separation (in which protons & hydroxyl ions are separate across surface)… results in form of energy
  • -> responsible for driving many energy-requiring functions in cell
  • -> includes some forms of transport, motility, and biosynthesis of ATP
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13
Q

Proton Motive Force

A
  • factor that gives cytoplasmic membrane ability to function as major site of E conservation in cell
  • **membrane possess an energetically charged form in which protons (H+) are separated from hydroxyl ions (OH-) across its surface**
  • -> charge separation is a form of E…analogous to potential E
  • -> responsible for driving many E-requiring functions in cell…some forms of transport, motility, biosynthesis of ATP
  • bc of charged membrane…charged molecules (even as small as a proton H+) canNOT passivle transport across membrane
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14
Q

Sterols

A
  • Eukaryotes possess **Sterols** in their membranes (while almost all prokaryotes do NOT)
  • -> (exceptions… 2 prokaryotes that do possess Sterols – Mycoplasmas)
  • -> Mycoplasmas lack cell walls…require Sterols to stabilize their membrane
  • Steroles make up anywhere from 5% - 25% of total lipids in eukaryotic membranes
  • Sterols are rigid planar molecules & the association of these with the membrane serves to *stabilize* its structure making it less flexible
  • -> Fatty Acids are more flexible
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15
Q

Hopanoids

A
  • similar to Sterols…rigid, planar molecules which strengthen & stabilize membrane making it less flexible
  • ***present in many Bacteria***
  • -> NOT present in Archaea
  • widely distributed example = *Diplotene (C30)*
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16
Q

Achaeal Membranes

A
  • Archaea lipids are chemically unique
  • -> possess ***Ether*** linkages between L-glycerol & their hydrophobic side chains…
  • -> …Bacteria & Eukarya possess *Ester* linkages between D-glycerol & fatty acid hydrocarbon chains
  • Archaea lipids lack fatty acids…
  • -> … possess side chains of repeating units of parent 5-carbon hydrocarbon **Isoprene** [(c) of image @ bottom]

***it was 16S rRNA sequence differences that led Woese to discover Archaea as a new domain…distinct from Bacteria & Eukarya***

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17
Q

Major Lipids of Archaea

A

(both still possess repeating units of Isoprene side chains)

  • ***Glycerol diethers***
  • -> possess 20-C side chains… (4 linked Isoprene units)
  • -> …*Phytanyl* – 20-C side chains
  • ***Diglycerol Tetraethers***
  • -> possess 40-C side chains… (8 linked Isoprene units)
  • -> …*Biphytanyl* – 40-C side chains
  • -> 2 phytanyl from each glycerol molecule are covalently bonded together – Tetraether actually called Di-biphytanyl diglycerol tetraether
  • (in Tetraether lipids)…
  • -> within a membrane, this structure yields a lipid ***monolayer***
  • Lipid Monolayers are resistant to peeling apart
  • -> monolayer membranes widespread among hyperthermophilic Archaea, prokaryotes…prevents cell lysis
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18
Q

Necessary Properties of Transport Proteins

A
  1. transport systems demonstrate **Saturation Effect**
  • -> carrier proteins are saturable sometime even at low [solute]
  • -> rate of uptake becomes maximal @ substrate saturation & addition of more solute does not increase the rate
  1. has high specificity
  • -> many carrier protons react only with a single molecule
  • -> others show affinities for closely related class of molecules (such as sugars; amino acids)
  1. biosynthesis of transport proteins is typically regulated by cell
  • -> function of both nutrients present in environment & their concentrations
  • -> particular nutrient may need to be transported by one transporter when at high [given nutrient]….and by a different (higher-affinity) transporter when at low [given nutrient]
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19
Q

Rate of Solute Entry

A

**Y axis : Si
**X axis: Time

  • Initial velocities of transport: /\Si/ /\T
  • velocities meased when Si is initially 0 & S0 is varied & rate is measured @ early times
  • each line represents a different initial [solute] on outside, S0
  • eventually *max velocity* is achieved
  • -> arrow line would be same for S4, S5, S6, etc…
  • -> Vmax of transport reached
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20
Q

Types of Passive Transport

A

**do not utilize source of E**
–> solute travels down conc. gradient…
…from [high] to [low]
- **Net transport internally when [S]o > [S]i**
–> if [S]o = [S]i then v = 0

  1. Simple Diffusion
    - non-saturable
  2. Facilitated Diffusion
  • saturation @ high [solute]
  • -> carrier proteins possess specificity for solute
21
Q

Types of Active Transport

A

**use E to accumulate solute against conc. gradient**
–> transports from [low] to [high]

  1. Simple Active
  • utilize Ion gradients/*Proton Motice Force*
  • consists of only a membrane-spanning integral protein
  1. ATP-binding Cassette (ABC) System
  • E from ATP
  • consists of 3 components: 1) substrate-binding protein; 2) membrane-integrated transport; 3) ATP-hydrolyzing protein

–> 2 ATP → 2 ADP + 2 Pi

  1. Group Translocation
  • E from chemical modification of transported substance (phosphorylation) driven by Phosphoenopyruvate
  • involved series of proteins in transport event
22
Q

Membrane Spanning Proteins & their structural similarities which correspond to their ability to act as Transporter membranes

–> 3 classes of transport events?

A
  • contain 12 alpha-helix domains that weave back & forth forming channels which solutes enter cell
    –> transport event involves protein *conformational change* upon solute binding…
    …this shuttles solute across membrane

3 clases of transport events:

1) Uniporter
- proteins that transport a molecule unidirectionally across membrane

2) Antiporter
- proteins that transport a molecules across membrane while simultaneously transporting a second molecule in *opposite* direction

3) Symporter
- proteins that transpot a molecule across membrane while simultaneously transporting a second molecule in *same* direction
- -> frequents a proton (H+)
- -> function as *cotransporters*

23
Q

Phosphotransferase System

A
  • best studied model of *group translocation*
    –> type of active transport – requires E…
    …energy from PEP (Phosphoenol-pyruvate)
    –> results in phophorylation (chemically modification) of substance being transported
  • **Phosphotransferase Sys. transports the sugars glucose, mannose, & fructose in E. coli**
  • consists of a family of proteins…5 of which are necessary for given sugar
  • (before transport of sugar) these proteins are alternately phosphoylated & then dephosphorylated in cascading fashion … until the actual transporter (Enzyme IIC) phosphorylates the sugar
  • Enz I; HPr; Enz IIa are all cytoplasmic proteins
  • Enz IIb lies on the inner surface & Enz IIc is an integral protein
  • Enz I & HPr are non-specific components
  • all Enz II proteins are specific components…
  • -> specific Enz II exists for each sugar (glucose, fructose, & mannose)

***phosphorylation of Glucose to Glucose-6-P is 1st step in its intracellular metabolism – Glycolysis***

24
Q

The ABC System

A

ATP-binding Cassette
–> necessary for uptake of organic compounds (sugars, amino acids) & inorganic compounds (sulfate & phosphate) & trace metals

**Gram- (-) bacteria possess Periplasmic region**

  • -> Periplasm – lies between cytoplasm & peptidoglycan
  • -> periplasm contains many different periplasmic-binding proteins
  • Gram- (+) bacteria also have ABC systems
  • -> substrate-binding protein anchored to external surface of cytoplasm

ABC System:

  • ATP hydrolyzing protein (Kinase) provides E
  • 3 components
    1) Periplasmic (substrate) - binding protein
    –> located in periplasm
    –> typically high substrate affinity
    2) membrane-spanning transporter protein
    3) ATP-hydrolyzing protein (Kinase)
    2 ATP → 2 ADP + Pi
25
Protein Transport
- cell to function properly...proteins transported through cytoplasmic membrane or inserted into membrane - -\> possible by \*\*Translocases\*\* - many bacterial enzymes function as exoenzymes - Preproteins -- presecretory proteins which synthesize transported proteins - -\> preproteins possess a signal peptide at amino terminus...recognized by Sec System - -\> Chaperon -- proteins that bind to signal peptides & delay protein folding (keeping it in necessary conformation for transport) - -\> Signal Peptidase -- remove signal peptide & protein folds into shape - key example -- \*Sec (secretory) System\* - -\> both exports/inserts into membrane - consists of 7 proteins: 1) Sec Y, E, G -- transmembrane transporter proteins 2) Sec A -- ATP hydrolyzing enzyme (energy) 3) Sec B -- prevents folding of protein in cytoplasm 4) Sec D, F -- assists in translocation - -\> also consumes E from Proton Motive Force
26
Function of Cell Wall of Bacteria
\*bc of activities of transport systems...cytoplasm of bacterial cells maintains a high [dissolved solutes]\* - protects cell from Osmotic Pressure & prevents Cell Lysis - -\> bacteria tend to live in environments typically hypotonic to cell...thus H20 tends to move into cell - -\> this causes significant Osmotic Pressure...would cause cell lysis if it were not for rigid cell wall - confers shape & regidity to cell
27
Gram + vs Gram - Bacteria cell walls
- \*\*Peptidoglycan is rigid layer primarily responsible for strength of cell wall\*\* - Gram + : - -\> 1 thicker layer of peptidoglycan - -\> Gram stain test results in Purple stained cells - -\> B. subtilis - -\> S. aureus - Gram - : - -\> 2 layers -- thin peptidoglycan + LPS - -\> Gram stain test results in Pink stained cells - -\> E. coli
28
Composition of Peptidoglycan
- rigid layer or sheet - -\> thick in Gram + - -\> thin in Gram - - composed of 2 sugar derivatives: N-acetylglucosamine (G) & N-acetylmuramic acid (M) - -\> always connected in Beta-1,4 glycosidic bond - also composed of small group of amino acids: L-alanine & D-alanine, D-glutamic acid; either Lysine (gram +) or Diaminopimelic acid (DAP) (gram -)
29
DAP vs Lysine
- 2 of numerous amino acids found in peptidoglycan cell wall of bacteria - identical in chemical structure besides terminal end - Gm (-): - -\> possess DAP (Diaminopimelic Acid) - -\> E. coli - -\> - COOH - Gm (+): - -\> possess Lysine - -\> B. subtilis - -\> S. aureus - -\> - H
30
Peptidoglycan variations: Cross-links & interbridge connections
\*\*Glycan backbone portion constant among all species of Bacteria\*\* - -\> G-M-G-M (N-acetylglucosamine & N-acetylmuramic acid) - -\> beta 1,4 Glycosidic bond - Tetrapeptide variation only in 1 amino acid in \*\*\*position 3\*\*\* -- DAP/Lysine - Gm (-): - -\> amino group (- NH2) of DAP cross-linked to carboxy group (- COOH) of terminal D-alanine via \*\*peptide bond\*\* - -\> NO interbridge - Gm (+): - -\> cross-linkage occurs via \*\*peptide interbridge\*\* - -\> kinds & numbers of amino acids in interbridge vary from organism to organism - -\> interbridge in S. aurerus...consists of Glycine pentapeptide bridge
31
Peptidoglycan Sheet
- Glycosidic bonds confer strength in X direction - Peptide bonds confer strength in Y direction
32
Diversity of Peptidoglycan --\> Defining characteristics...?
- only present in species of Bacteria - sugar N-acetylmuramic acid (M) & amino acid Diaminopimelic acid (DAP) never found in cell walls of Archaea or Eukarya - most Gm (+) possess Lysine instead of DAP - -\> 3rd molecule distal from M - (unusual feature) presence of 2 amino acids that are D-configuration (D-alanine & D-glutamic acid) - greatest diversity in peptide crosslinks & interbridge - -\> Gm (+) -- Lysine -- cross-linkage @ peptide interbridge bond (Lysine & varying amino acid bridge of adjacent glycan chain) - -\> Gm (-) -- DAP -- cross-linkage @ peptide bonds (DAP & D-alanine of adjacent glycan chain) - Gm (+) possess more cross-links - Gm (-) possess cross-links that are "longer"
33
Teichoic Acids
- acid substances embedded in cell wall - -\> of Gm (+) Bacteria - bc of (-) charge of Teichoic acids...partially responsible for (-) charge on Gm (+) cell surface as whole - encompasses all cell wall, cytoplasmic membrane, & capsular polymers containing \*polyalcohols\* - -\> Polyalcohols -- glycerophosphate; ribitol phosphate residues - (in wall) covalently bound to N-acetylmuramic acid via \*\*Phosphate Esters\*\* - -\> usually possess D-alanine & other sugars attached - \*\*Lipoteichoic acids\*\* -- teichoic acids covalently linked to membrane lipids (Lipoproteins)
34
Lysozyme
- Peptodoglycan can be destroyed by certain agents - -\> causes weakening of cell & possibly eventual cell lysis - -\> such as \*\*Lysozyme\*\* - -\> Lysozyme -- protein that breaks the B-1,4-glycosidic bond between N-acetlymuramic acid & N-acetyl glucosamine - -\> Lysozyme function as major line of defense against bacterial infections via this mechanism
35
Protoplasts
- a Gm (+) bacterium that has lost its cell wall - -\> did not under cell lyses - -\> destruction of peptidoglycan caused from agents such as Lysozyme - Lyses occurs.... ...in dilute solution bc cell wall is structurally very weak - Lyses does NOT occur... ...in solutation containing an \*\*isotonic\*\* concentration of a solute (such as Sucrose) ...H20 does NOT enter cell ⇒ \*\*\*Protoplasts only possess cytoplasmic membrane & internal constituents\*\*\* --\> devoid of cell wall; no residual wall left
36
Spheroplasts
- Gm (-) bacteria produce Spheroplasts when treated with Lysozyme - (similar to protoplasts)...but possess residual wall attached
37
Examples of Cells that lack Cell Walls
- Mycoplasmas - -\> group of pathogenic bacteria that cause variety of infectious diseases in humans & other animals - -\> like protoplasts & live in osmotically protected environment - -\> Sterols add strength to membrane - -\> ...Lysozyme found in animal secreations... - Thermoplasmas - -\> species of Archaea that naturally lack cell wall - -\> unusually tough cytoplasmic membranes
38
Pseudomurein
- found in some Archaea cell wall - -\> Methanogenic Archaea - constructed of a polysaccharide very similar to Peptidoglycan - backbone composed of alternating repeats of N-AcGlu & N-acetyltalosaminuronic acid (T) - -\> Glycosidic bonds are B-1,3 (instead B-1,4) - -\> Lysozyme-insensitive - possess Lysine as 3rd position on Glycan Chain - possess only L steroisomer - some archae have S layers
39
Penicillin
- type of Beta-lactam ring antibiotic - potent inhibitors of cell wall synthesis - -\> disrupts formation of cross-links - -\> bind to Transpeptidase enzymes - Archae naturally resistant to Penicillin (& Lysozyme) - Bacteria affected by both Penicillin & Lysozyme
40
S layer
- most common cell wall type among Archaea - Paracrystalline surface layer (or S layer) - -\> showed ordered appearance under microscope - -\> typically hexagonal symmetry - consists of protein or glycoprotein - provides some protection from osmotic lysis - -\> many organisms possess both S layer & peptidoglycan -- (S layer being the outermost) - selective sieve - -\> not alloweing large molecules in (such as viruses) - may retain proteins near surface
41
Lipopolysaccharide layer (LPS)
- Gm (-) bacteria possess outer LPS in addition to peptidoglycan - contains polysaccharides & phospholipids... - -\> ... linked in outer membrane to form lipopolysaccharide structures
42
Structure of LPS
- 3 components: 1) O-specific polysaccharides - often branched - most external facing component 2) Core polysaccharides 3) Lipid portion (Lipid A) - most internal portion of LPS - is endotoxic (toxicity to animals) - embedded in phospholipid portion - -\> lipoproteins present on inner side of LPS anchor outer membrane to peptidoglycan - considered lipid bilayer but distinct from cytoplasmic membrane
43
Endotoxin
- LPS layer [Gm (-)] bacteria commonly toxic to animals - toxic properties associated with Lipid A - -\> Innermost portion of LPS - produce fever, decrease B.P., activation of inflammation, coagulation of blood presence in blood can lead to \*\*septicemia\*\*
44
LPS Permeability & Porins
- LPS not permeable to large molecules (proteins) - -\> major function -- prevent proteins from diffusing - O-specific polysaccharides prevent hydrophobic molecules from diffusing pass LPS - -\> hydrophobic antibiotics not effective to Gm (-) - Lipid A phspholipid layer prevent hydrophilic small molecules from diffusing pass LPS - \*\*Porins\*\* -- proteins present in LPS - -\> function as channels for entrance/exit of hydrophilic low-molecular-weight substances - -\> H20 filled channels - -\> several different porins exist...both specific & nonspecific - consists of 3 identical subunits
45
Capsules vs. Slime Layers
both may be thick/think; rigid/flexible depending on chemical makeup & hydration 1. Capsules - material organized in tight matrix (like an envelope around cell) - exclused small particles -- such as india ink 2. Slime layers - material more easily deformed; organized in loose matrix - will not exclude particles; more difficult to see
46
Functions of Capsules
- attachment of microbes to solid surfaces - -\> pathogens typically first bind to surface components on human tissues - -\> nonpathogenic often bind to surfaces in nature -- forming Biofilm ( -- thick layer of cells) - encapsulated pathogenic bacteria more difficult for phagocytic cells to engulf & destroy - polysaccharide layers bind water & may make cells resistant to desiccation (drying)
47
Fimbriae & Pili
- both filamentous structures composed of protein that extend from surface of cell - -\> not typically involved in motility even though similar to flagella - allow cells to stick to surfaces - -\> including animal tissue - Fimbriae are considerably shorter than flagella & are more numerous - Pili are similar to fimbriae but typically longer & only 1 or a few present on cell surface
48
Functions of Pili
pili often receptors for certain types of viruses... --\> can best be seen under electron microscope when they have become virus-coated pilus - \*Congugation\* -- facilitation of genetic transfer between prokaryotic cells - -\> F+ (donar) x F- (recipient) ⇒ F- becomes F+ - -\> F factor (Plamid) transferred while replicated - \*Adhesion of pathogens\* to specific host tissue - \*Cell Twitching\* -- unusual form of cell motility