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Prokaryotic and eukaryotic cells are similar in a number of ways including
Chemically similar – contain macromolecules: Nucleic acids, proteins, lipids and polysaccharides’
Similar metabolic reactions to metabolize food, synthesis proteins and nucleic acids and store energy
Contain a membrane, cytoplasm, DNA and ribosomes
Prokaryotic and eukaryotic cells differ in a number of ways (Table 4.2) including
Prokaryotic cells
DNA is usually in the form of a single circular dsDNA chromosome and not enclosed in a membrane
DNA is not associated with histones; other proteins are associated with DNA
They lack membrane bound organelles
Usually divide by binary fission
Eukaryotic cells
DNA is usually in the form of multiple linear dsDNA chromosomes found in a membrane bound nucleus
The DNA is consistently associated with chromosomal proteins called histones and with nonhistone proteins
They may have a number of membrane bound organelles, including endoplasmic reticulum,
Golgi complex, lysosomes, vacuoles, mitochondria and chloroplasts
Cell division usually involves mitosis
Prokaryotic cells: Archaea (member of Archaeal domain = archaeon) and Bacteria (member of Bacterial domain = bacterium)
Prokaryotic cell structures include the following (Fig. 4.6; Note: Underlined structures are found in all prokaryotic cells):
Plasma membrane
Cytoplasm
Nucleoid region
Ribosomes
Cell wall (and periplasmic space)
Flagellum
Pili and Fimbriae
Inclusions
Gas vacuole
Capsule and slime layers
Endospores
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Eukaryotic cells: Protozoa, Fungi and Algae
Eukaryotic cell structures include the following (Fig. 4.22; Note: not all cells possess all of these structures at all times):
Plasma membrane
Cell wall
True nucleus - membrane bound nucleus
Ribosomes
Membrane bound organelles
chloroplast
mitochondrion
endoplasmic reticulum
vacuoles
golgi apparatus
lysosomes
peroxisomes
Cytoskeleton
Flagellum/Cilium
A. Prokaryotic Cells
Two most common cell shapes
coccus ( pl.
cocci) e.g.,
bacillus ( pl.
bacilli) e.g.,
Other shapes include:
spirillum ( pl . spirilli) e.g.,
Mycelial – e.g., Actinomycetes
Stalked – e.g., Caulobacter, Hypomicrobium
Plates
Star shape
Some prokaryotes are variable in shape and lack a single characteristic form = pleomorphic e.g.,
B. Eukaryotic Cells
Highly variable: cell shapes vary in shape from sphere and cylinders to very irregular nerve cells.
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C. Cell Size
Cells come in a variety of sizes & shapes
Size limit is set by the logistics required to carry out metabolism
The smallest cells are nanobacteria (diameter of 0.05 to 0.2 µm).
Mycoplasmas = 0.2 µm in diameter
What factors constrain the lower size limit of cells?
Most bacterial cells are 10 times larger than mycoplasmas (1 to 10 µm in diameter)
Eukaryotic cells are typically 10 times larger than bacteria (10 - 100 µm in diameter).
What factors constrain the upper size limit of cells?
Generally prokaryotic cells are smaller than eukaryotic cells. But there are exceptions. e.g., Epulopiscium fishelsoni size up to 80 x 600 µm
Thiomargarita namibiensis size up to 750 µm in diameter
Nanochlorum eukaryotum
1 to 2 µm in diameter
Generally cells are microscopic. But there are exceptions
Loligo - Atlantic squid has neurons with axon diameters as large as 1.0 mm.
Ostrich egg
Every cell is surrounded by a plasma membrane (also known as a cytoplasmic or cell membrane)
Encloses the cytoplasm
This is an important feature distinguishing archaea from bacteria and eukaryotes
Membranes are selectively permeable barriers - Why?
What is the function of the plasma membrane?
Fluid mosaic model (Fig 4.14)
S.J. Singer and G. Nicolson (1972)
Dynamic structure
bifacial quality – sidedness
5 – 10 nm thick
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1. Membrane components i. Lipids
backbone or basic fabric
often as a lipid bilayer ( amphipathic) hydrophobic core hydrophilic exterior surfaces ii. Proteins
integral (70 to 80 % of the membrane proteins)
peripheral (20 to 30% of the membrane proteins)
Functions o Transport – import and export o Enzymes o Receptors o Intracellular junctions o Cell to cell recognition o Metabolic processes iii. Carbohydrates
Usually associated with the outer surface of the membrane
2. Differences between bacterial, eukaryotic and archaeal membranes i. Bacterial membranes
Like eukaryotes, most of the membrane lipids are phospholipids
unlike eukaryotes bacteria lack sterols such cholesterol but contain sterol like compounds called hopanoids. The role of hopanoids is likely similar to steroids – stabilized membranes.
Some bacteria may have extensive in folding of the plasma membrane to increase membrane surface area for the purpose of greater metabolic activity
Glycerol diesters (phospholipids) o Glycerol bonded to phosphate (negatively charged) and two fatty acids o Linkage between fatty acids and glycerol is an ester linkage
Ester linkage
O
||
R-O-C-(CH
2
) n
CH
3
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Environmental conditions affect fatty acid composition of membranes o Fatty acids are generally unbranched and 16 to 18 carbons long e.g., palmitic acid CH
3
(CH
2
)
14
COOH stearic acid - 18 carbons o Fatty acids may be saturated or unsaturated
Membrane composition also varies with species and these differences can be used to identify bacteria
(Fatty acid methyl ester analysis - FAME ). ii. Eukaryotic membranes o generally the same structure as bacterial membranes including phospholipids but differs in the major lipids: phospholipids, sphingolipids and cholesterol. o Microdomains that differ in protein and lipid composition may be found – lipid rafts – span membrane and appear to be involved in a variety of cellular processes (e.g., signal transduction and cell movement)
Sterols o compounds consisting of carbon skeleton of 4 interconnected rings o targets for polyene antibiotics - damage cell membranes o Mycoplasmas acquire sterols from eukaryotic cells iii. Archaeal membranes
Many are thought to be “Extremophiles”
Membranes are distinctive
Phospholipids are not the main structural components
Have branched chain hydrocarbons attached to glycerol by ether linkages
Glycerol diethers
Ether linkage R-C-O-C-R
Bilayers o glycerol bound to branched hydrocarbons (e.g., phytanyl) by ether linkage o glycerol diethers
Monolayers o diglycerol tetraether o often found in extreme thermophiles
Phosphate, sulfur and sugar containing groups are attached to the third carbon of glycerol resulting in a polar lipids that are the predominant lipids (70 – 93%) in the membrane
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Nonpolar lipids (squalene derivatives) make up the rest of the membranes
3. The movement of materials across membranes
Review this material on your own – it is largely review from Biol 1010. You should be familiar with the following
Passive Transport
Simple diffusion
Facilitated diffusion o Transporter proteins
Osmosis o Aquaporin o Osmotic pressure
Active Transport o Group translocation
1. Bacterial cell walls
Functions
Define cell shape
Protection from osmotic shock (turgor pressure) and toxic substances
Point of anchorage for flagella
May contribute to pathogenicity
Most bacteria have cell walls but there are exceptions – e.g., mycoplasmas
Forms a strong, protective layer that is relatively porous, elastic and somewhat stretchable
Gram positive and Gram negative (Fig. 4.13)
Can be differentiated through the Gram stain - an important diagnostic tool.
Gram negative
Gram positive
Gram variable i. Peptidoglycan (murein)
Components a. Polysaccharide
1,4 linkages between N-acetylglucosamine (NAG) and N-acetylmuramic acid (NAM) monomers
(Fig 4.12).
Note: N-acetylmuramic acid is found only in Bacteria
Polymers 10 to 65 monomers long
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...NAM – NAG - NAM - NAG - NAM - NAG... b. Peptide chains
Tetrapeptide chains linked to M residues - composed of unusual amino acids (D- amino acids) – Why?
L-alanine
D-glutamic acid
L-lysine (Gram positive) or diaminopimelic acid (DAP; Gram negative)
D-alanine
There are either peptide interbridges (G-G-G-G-G: Gram positive) or direct peptide linkages (Gram negative) between tetrapeptides
Results in a strong multilayer sheet or sacculus
Autolysins - enzymes used by the bacteria to recycle, reshape or restructure cell wall
Lysozyme
Hydrolyes the
1,4 linkages between M and G monomers
Sources of lysozyme
predators
tears
saliva
chicken egg white
spheroplast - partial removal of cell wall
protoplast - complete removal of cell wall
Penicillin
Prevents formation of peptides cross linkages between tetrapeptides
Penicillin binds to transpeptidase
Not effective against bacteria lacking cell walls such as mycoplasmas ii. Gram positive cell wall
Up to 20% of the organism
peptidoglycan represents 50-90% of this structure
relatively thick (ca. 40 nm – ranges from 20 to 80 nm)
a thick peptidoglycan layer is more resistant to desiccation
Polysaccharides such as teichoic acids (e.g., glycerophosphate or ribitol phosphate residues – negatively charged) or teichuronic acids are bonded to the peptidoglycan or plasma membrane lipids
Putative function - bind to cations and regulate movement into and out of the cell
- Prevent extensive cell wall lysis
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Responsible for cell wall’s antigenicity e.g., Bacilli, Staphylococci, Streptococci iii. Gram negative cell envelope
more complex than Gram positive cell wall
Cell wall
thin layer of peptidoglycan (ca. 2 to 7 nm)
1 - 10 % of the cell wall
lipoproteins instead of teichoic acids
Outer membrane - lipid bilayer
phospholipids
proteins - e.g., porins
lipoproteins – anchored to peptidoglycan
The outer membrane may be linked to the plasma membrane in a number of places
lipopolysaccharides (LPS)
composed of i) lipid A, ii) the core polysaccharide and iii) the O polysaccharide side chain.
main structural component of outer half of outer membrane believed to aid in creating a permeability barrier as well as contributes to negative charge of the cell surface.
Protects pathogenic bacteria from host defenses
The LPS (Lipid A componenet in particular) is frequently toxic to animals and known as endotoxin (i.e., because it is still attached to cell).
O polysaccharide is an antigen that is used to distinguish species of bacteria
The outer membrane is more permeable than the plasma membrane to most small molecules due to the presence of porin proteins
Less permeable than the plasma membrane to hydrophobic and amphipathic molecules - makes cells less susceptible to certain antibiotics.
Keeps periplasmic enzymes from diffusing away.
Periplasmic space (30 - 70 nm wide)
Gel-like in consistency due to abundance of proteins
Import region where number of chemical reactions occur:
oxidation-reduction reactions
osmotic regulation
solute transport
hydrolysis
protein secretion
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2. Archaeal cell wall
No peptidoglycan, in particular lacking in N-acetylmuramic acid
May be composed of
Pseudopeptidoglycan – N-acetylglucosamine and N-acetyltalosaminuronic acid linked by
1,3 linkages
Protein or glycoproteins - most common form of cell wall
Polysaccharide
NOTE: Some Bacteria and Archaea do not have cell walls e.g., mycoplasma
Thermoplasma
3. Eukaryotic cell wall
Like Bacteria and Archaea, not all eukaryotes have cell walls
Algal cell wall
Polysaccharides are the major components e.g., cellulose
May contain high concentrations of calcium or silicon - diatoms
Fungi
Many contain chitin - polysaccharide consisting of N-acetylglucosamine monomers.
Composition is used in classification schemes e.g., primitive Chytridiomycetes fungal cell walls lack chitin but contain cellulose
Protists
Many protists have a pellicle for support – rigid layer of components beneath the plasma membrane.
Pellicle is composed of protein
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Prokaryotes
Prokaryotes have a variety of layers outside the cell wall
Functions
protection
ingestion
dehydration
loss of nutrients
attachment
pathogenicity
Form thick or thin, rigid or flexible layers depending upon composition
Variable composition – glycoproteins and polysaccharides
1. Capsule
Rigid – tight matrix that excludes particles
protein or polysaccharide
2. Slime layer
loosely bound layer – easily deformed
3. Extracellular Polymeric Substance (EPS)
glycocalyx that helps cells bind to surfaces and other cells – formation of biofilm
Glycocalyx – term that collectively refers to both capsule and slime layer
3. Surface layer (S-layer)
nearly all bacteria and Archaea
crystalline protein layer
unknown function – may function as permeability or protective barrier
Deoxyribonucleic acid (DNA) - macromolecule consisting of nucleotide monomers
Backbone = 5'...-Phosphate-Sugar-Phosphate-Sugar -Phosphate-Sugar-…3'
Nucleotide = nucleoside plus phosphate
Nucleoside = deoxyribose + nitogenous base
Purines – adenine & guanine
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Pyrimidines - cytosine & thymine
Review structure of DNA – Chapter 2
1. Bacterial and Archaeal DNA
These organisms do not have a nucleus – the bulk of the genetic material is found in the nucleoid region
Chromosome
single double stranded DNA molecule in the form of a covalently closed circular chromosome – also known at the genophore
Exceptions: Borrelia burgdorferi and some Streptomyces spp. have a linear chromosome;
Rhodobacter sphaeroides has two circular chromosomes
usually only one chromosome and it may be present as multiple copies in rapidly growing cells
DNA arranged into supercoiled domains that are stabilized with structural proteins
In some Archaea – DNA is extensively complexed with proteins that closely resemble histone proteins of eukaryotic organisms
Plasmids
autonomously replicating units that usually contain only a few genes (< 30; 1 – 5% of the size of the chromosome). Most the bacterial and archaeal genomes sequenced contain plasmids
Usually covalently closed circles (CCC) but may be linear
May contain one to many plasmids
Range in size from several kbp to Mbp
Examples of plasmid coded factors (Table 3.3)
1.
R-plasmids - antibiotic resistance genes
2.
Bacteriocins
3.
Conjugal factors
4.
Metabolic factors - substrate utilization or fixation
5.
Virulence factors - toxin production
2. Eukaryotic DNA
Most possess a nucleus that contains the bulk of the genetic material
Nucleus contains a number of linear chromosomes
Chromosomes composed of DNA complexed with protein
chromatin
DNA associated with Histones in structural subunits called nucleosomes
Chloroplasts and mitochondria also contain small circular genomes
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Sites of protein synthesis
Approximately 20 - 25 nm in diameter
Large numbers may be present in cells (>10,000 in bacterial cells and more in eukaryotic cells)
Number varies depending on the level of protein synthesis going on in the cell.
Ribosome
Bacteria and
Archaea
70 S*
Eukarya
80S
Large subunit
Small subunit
50 S
23S rRNA and 5S rRNA
Large number of proteins
e.g., 34 proteins in E. coli
60S
25 - 28S rRNA and 5.8S rRNA
Large number of proteins
* Svedberg units (S)
Eukaryotic organisms have 70S ribosomes in their mitochondria and chloroplasts
Implications in the endosymbiotic theory
Practical implications of ribosome structural differences
Antibiotic treatment chloramphenicol tetracycline kanamycin erythromycin streptomycin diptheria toxin anisomycin
30S
16S rRNA
Large number of proteins
e.g., 21 proteins in E. coli
40S
18S rRNA
Large number of proteins all bind 70S bacterial ribosomes and disrupt protein synthesis bind 70S archaeal and 80S eukaryotic ribosomes and disrupts protein synthesis
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Prokaryotic Cells
For many years it was thought that prokaryotic cells lacked cytoskeletal elements. Recently homologues have been discovered for all three elements of the eukaryotic cytoskeleton e.g., FtsZ – tubulin homologue; widely observed in Bacteria and Archaea
MreB – actin homologue; Many rod shaped cells
Crescentin – intermediate filament proteins; Caulobacter
Eukaryotic Cells
Have a well developed three dimensional network of fibrous proteins (microtubules, microfilaments and intermediate filaments
Microtubules and microfilaments are very dynamic structures – can be quickly disassembled and assembled elsewhere
Functions
Support
Maintenance of cell shape
Cell movement - e.g., muscle cell contraction, amoeboid movement, cilia
Cell division (mitosis, cytokinesis and meiosis)
Cell wall deposition
Provides spatial organization and movement of organelles and cytosolic enzymes
Regulation of biochemical activities in the cell – mechanical signaling
Cytoskeleton components a) Microtubules
thickest cytoskeletal elements - hollow rods - 25 nm
found in the cytoplasm of all eukaryotes
composed to and tubulin dimers
readily assembled and disassembled
grow by addition of subunits to ends
centrosome - microtubule organizing centre (MOC) important in cell division
a pair of centioles (9 sets of microtubule triplets) often found in the centrosome region of animal cells. Replicate during cell division. May function in cell division but not necessary as they are generally not found in plant cells.
Functions
compression resisting function
cell motility (flagella and cilia)
chromosome movement
organelles movement
cell shape
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Tortora et al – Chap 4 b) Microfilaments
thinnest cytoskeletal element - 7 nm in diameter
composed of protein called actin
readily assembled and disassembled
Functions
tension bearing function
muscle contraction (myosin motors molecules-burn ATP)
cytoplasmic streaming (myosin motors molecules-burn ATP)
cell motility (pseudopodia)
cell division
cell shape – three dimensional network just inside the plasma membrane c) Intermediate filaments
8 to 12 nm in diameter
diverse class composed of different protein subunits including keratins
more permanent structures not readily assembled and disassembled like microtubules and microfilaments
Functions
tension bearing function
maintenance of cell and organelle shape e.g. nuclear lamina
fixing positions of certain organelles
Bacterial and Archaeal Movement
1. Flagellum ( pl.
flagella)
one to many flagella
up to 60 cell lengths/s
Atrichous – no flagella
Monotrichous - single polar flagellum
Amphitrichous - single flagellum at each pole
Lophotrichous - polar tuft of flagella
Peritrichous - multiple flagella disgributed over the entire cell
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Parts of a prokaryotic flagellum (Fig 4.8) i. filament - whiplike extension that rotates - helical in shape – 15 to 20 µm long composed of flagellin – highly conserved in bacteria ii. curved hook - single protein - connects filament to motor iii. Basal apparatus or motor composed of many proteins (~30) central rod
a series of rings embedded in the cell wall, plasma membrane and outer membrane (Gram negative cells). This structure is around 20 nm in diameter
Mot proteins drive the flagellar motor - energy comes from proton motive force (about 1000
H
+
/rotation)
Fli proteins act as a switch
> 40 genes ( fla, fli, flg ) required for synthesis and motility (structural, export of components and timing of synthesis)
2. Gliding
cells glide along a surface.
Mechanism is unknown but there are several models a) excreted slime adheres to surface pulls cells along - cyanobacteria b) movement of surface proteins - Flavobacterium
3. Gas Vesicles
confer buoyancy on cells and allow to move up and down in a water column
composed of two types of protein (97% of the gas vacuole is composed of GvpA (
-sheets). The remainder is made of GvpC (
-helix) that acts like a cross linker between the GvpA molecules
4. Behavioural Responses to Stimuli
In a heterogenous environment prokaryotes are capable of movement ( taxis; pl. taxes ) towards or away from stimuli (light, heat, chemicals and electricity)
Chemotaxis
movement towards or away from a chemical stimulus (chemoattractant or chemorepellent).
Respond to temporal gradient
Respond to very low levels of some materials – 10 -8 M for some sugars
Types of Taxes phototaxis - light stimulus scotophobotaxis - entering darkness has a negative effect on a cell geotaxis - gravitational stimuli magnetotaxis - magnetic stimuli aerotaxis – oxygen stimulus
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Bacteria lack spatial sensing capabilities- they are too small to sense a gradient along the cell length.
Consequently they respond in a temporal fashion.
1. Periodically sample environment e.g., chemoreceptors – sensory proteins
2. Process information through a signal transduction pathway
3. Control of direction of flagella rotation
Movement is through a series of a) alternating runs (straight line movement- counterclockwise rotation of flagella) and tumbles (reversal of direction of flagella rotation) b) The direction of the next run is random; however, if there is a gradient of a chemical attractant or repellent present, the random movements become biased
If the organisms sense an increase in attractant concentration then this results in longer runs and tumbles are less frequent. This is also the case if the cell senses a decrease in repellant concentration
Eukaryote Movement
Cilia and flagella are the most prominent structures for movement
Cilia (sing. - cilium) and Flagella (singl. - flagellum) are composed of microtubules
Cilia and flagella are 0.25 µm in diameter
A plasma membrane encases a core of microtubules arranged in the 9 + 2 pattern (Fig 4.23): and outer ring of 9 evenly spaced doublets around 2 two single microtubules
radial spokes and cross-linking proteins connect the microtubules;
dynein arms (named after the protein composing these structures) connect microtubules
anchored to cell by a basal body - identical in structure to centrioles
chemical energy required for movement
Comparison of Flagella and Cilia
Cilia
Flagella
Length
2 - 20 µm
10 - 200 µm
Number large number
(100-1000’s)
1 to several
Movement back and forth - rowing undulating or snakelike
(sperm cell)
The eukaryotic flagellum is much different in structure than prokaryotic flagella ( Another difference between Prokaryotes and Eukaryotes!)
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Glycocalyx – discussed already
Pili and/or Fimbriae
hair like proteinaceous projections used as attachment structures fimbriae
Slender tubes (3 – 10 nm in diameter) composed of helically arranged protein subunits
function in attachment and twitching movement
more numerous than pili pili
generally longer and thicker than fimbriae but fewer in number
attachment between mating bacterial cells
consist of phosphate-carbohydrate-protein (single type of pilin)
F pilus - important structure involved in bacterial mating (conjugation) which is carried exclusively by the donor cell
Storage of energy or structural compounds
Often form in response to excess or imbalance in nutrients that are available
Usually bounded by a thin non-unit membrane examples
phosphate – polyphosphate – metachromacit granules
sulfur
glycogen or starch
poly-
-hydroxyalkanoates (e.g., poly-
-hydroxybutyric acid - PHB) are common carbon reserves produced by Bacteria and Archaea e.g., B. thuringienisis
YPD mediium – very rich medium – PHB
NA medium – spore formation
Functions
• reproduction
• dispersal - fungi and some actinomycetes produces spores that are readily dispersed
• survival under adverse conditions
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1. Bacterial endospore
Differentiated cells
Formed within the cell – refractile body
Dormant structures that are highly resistant to heat, desiccation, UV irradiation, chemical disinfectant – Not a reproductive structure
Found commonly in soil
Contains little water
Complex multilayer spore coat contains peptidoglycan and calcium dipicolinate (required for heat reistance) in its core
Endospore producing bacteria - at least 16 genera, including
Bacillus
Clostridium
Sporosarcina
Oscillospira
Desulfotomaculum i) Endospore Structure
Exosporium
thin delicate proteinaceous covering
Spore coat
several protein layers
Impermeable and responsible for spores resistance to chemicals
Cortex
up to half the spore volume
peptidoglycan (less cross linked than vegetative cell)
Core or spore wall
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Spore core
plasma membrane, cytoplasm, nucleoid
energy supply
phosphoglycerate
contains Ca
2+
complexed with dipicolinic acid
10 – 30 % water content of vegetative cells – water moves freely in and out of endospores
heat resistance of endospore is determined by state and amount of water in core
pH is one unit lower than vegetative cells
contains small acid-soluble spore proteins (SASP) which bind DNA and protect it from damage due to UV, heat and desiccation. May also serve as a carbon and energy source during germination
Ca 2+ dipicolinic acid complexes and SASP form a gel in cytoplasm – gel like material excludes water from the endospore protoplasm
Low metabolic activity, no macromolecule synthesis and few or no mRNA molecules
The core also contains DNA repair enzymes that repair the the DNA once the spore germinates ii Sporulation (sporogenesis)
a very complex inducible process and is controlled by > 200 sporulation specific genes.
triggered by environmental conditions which are unfavourable for vegetative growth. In Bacillus species
C, N or P limitation.
a multistage process (seven stages have been described for Bacilli) that can last up to 8 hours or more. Vegetative cell or mother cell becomes compartmentalized forming a spore compartment through invagination of the plasma membrane (sporangium).
Endospores dormant for many years
If conditions become favourable for vegetative growth then the endospore can rapidly convert back to a vegetative cell.
Three stages i) Activation
prepares endospore for germination
reversible process
treatments like heating conditions endospore to germinate when placed in a nutrient solution ii) Germination
breaking of an endospore’s dormant state, germinants - such as glucose and certain amino acids may induce gemination
characterized by
loss of refractility
loss of resistance to heat and chemicals
loss of Ca dipicolinate and cortex components, and degradation of SASPs
and increased ability to be stained by dyes iii) Outgrowth
develops into a vegetative cell
uptake of water
swelling
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macromolecule synthesis
vegetative growth until encounters environmental triggers again
Review cell structures and concepts including
Endomembrane system and all of its components
Mitochondria
Chloroplast
Endosymbiotic theory – eukaryotic cells arose from larger cells engulfing smaller cells
Mitochondria and chloroplasts
contain DNA – closed covalently circular form
contain their own ribosomes – 70 S
protein synthesis in these organelles is affected by antibiotics such as streptomycin that also kill or inhibit bacteria
rRNA sequences of mtDNA and cpDNA are more closely related to bacteria rRNA
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