Extreme Environments and Extremophiles: Surviving the Unthinkable

 
E
X
T
R
E
M
O
P
H
I
L
E
S
:
 
T
h
e
r
m
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p
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i
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&
A
c
i
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o
p
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s
 
E
X
T
R
E
M
O
P
H
I
L
E
S
 
Organisms found living in extreme harsh
 
environments.
W
o
r
d
 
o
r
i
g
i
n
a
t
e
d
 
f
r
o
m
 
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-
 
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Definition: Extremophiles are organisms that have been
discovered on  earth that survive in environments that
were once thought  not to be able to sustain life.
 
M
o
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,
 
h
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s
,
t
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s
,
 
a
l
k
a
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,
 
p
s
y
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,
h
a
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,
 
b
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b
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e
n
d
o
l
i
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h
s
.
 
A
d
a
p
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a
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s
 
E
x
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t
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t
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.
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a
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t
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r
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e
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o
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,
 
p
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t
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a
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w
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i
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a
c
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a
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e
 
o
t
h
e
r
 
 
e
n
z
y
m
e
s
.
Important 
in 
industries because 
of this
property.
 
E
X
T
R
E
M
E
 
C
O
N
D
I
T
I
O
N
S
 
T
e
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s
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e
n
R
a
d
i
a
t
i
o
n
P
r
e
s
s
u
r
e
 
Response 
to Environmental
 
factors
 
T
Y
P
E
S
 
P
s
y
c
h
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p
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s
T
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m
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r
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i
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-
1
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t
o
 
1
5
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C
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k
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c
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.
H
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(
9
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b
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g
l
a
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r
s
,
 
s
n
o
w
f
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d
s
 
e
t
c
Metabolism is quite normal at colder
 
temperatures.
C
e
l
l
 
m
e
m
b
r
a
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e
s
-
h
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h
 
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p
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t
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c
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d
 
D
N
A
Some of them 
cause 
spoilage in refrigerated food
 
materials.
E
g
:
 
A
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r
o
b
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s
p
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,
 
P
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s
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,
 
H
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,
 
P
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d
o
m
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a
s
,
 
S
p
h
i
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g
o
m
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a
s
 
F
I
R
M
I
C
U
T
E
S
 
G
r
a
m
 
p
o
s
i
t
i
v
e
,
 
s
p
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e
 
f
o
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b
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e
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c
o
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d
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s
.
T
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a
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e
 
b
a
c
t
e
r
i
a
 
(
e
u
b
a
c
t
e
r
i
a
)
.
Plays an important role in 
the 
spoilage 
of 
beer, 
wine
and
 
cider.
E
g
:
 
H
e
l
i
c
o
b
a
c
t
e
r
i
u
m
 
s
p
p
,
 
M
y
c
o
p
l
a
s
m
a
,
C
l
o
s
t
r
i
d
i
u
m
 
s
p
p
.
 
M
a
n
y
 
m
e
m
b
e
r
s
 
o
f
 
t
h
e
 
F
a
m
i
l
y
 
F
i
r
m
i
c
u
t
e
s
 
a
r
e
 
a
l
s
o
t
h
e
r
m
o
p
h
i
l
e
s
.
E
g
:
 
B
a
c
i
l
l
u
s
 
s
t
e
a
r
o
t
h
e
r
m
o
p
h
i
l
u
s
R
e
c
e
n
t
l
y
,
 
a
 
D
N
A
 
p
o
l
y
m
e
r
a
s
e
d
e
r
i
v
e
d
 
f
r
o
m
 
t
h
e
s
e
b
a
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r
i
a
,
 
B
s
t
 
p
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l
y
m
e
r
a
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e
 
h
a
s
 
b
e
c
o
m
e
 
i
m
p
o
r
t
a
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t
 
i
n
b
i
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t
e
c
h
n
o
l
o
g
y
.
B
s
t
 
p
o
l
y
m
e
r
a
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-
 
h
e
l
i
c
a
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e
 
l
i
k
e
 
a
c
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v
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y
 
(
m
a
k
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t
a
b
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t
o
 
u
n
w
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d
 
D
N
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s
t
r
a
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d
s
.
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p
t
i
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f
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a
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t
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i
s
 
6
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6
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0
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a
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d
 
g
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t
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a
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A
 
t
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s
 
a
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s
,
 
b
e
t
w
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n
 
4
5
 
a
n
d
 
1
2
2
 
°
C
 
(
1
1
3
 
a
n
d
 
2
5
2
 
°
F
)
.
 
Many thermophiles 
are 
archaea. Thermophilic 
eubacteria 
are 
suggested 
to  
have been
among the 
earliest
 
bacteria.
 
"Thermophile" 
is derived from 
the Greek: 
(
thermotita
), meaning 
heat,  
and Greek:
(
philia
),
 
love.
 
Thermophiles 
are classified into 
obligate 
and 
facultative 
thermophiles
:  
Obligate
thermophiles (also called extreme thermophiles) require 
such high  temperatures for
growth,
 
whereas 
facultative thermophiles 
(also 
called moderate thermophiles) 
can  
thrive at high
temperatures, but also at lower temperatures (below
 
50°C).
 
H
y
pert
h
ermophil
e
s
 
are
 
pa
r
t
i
c
ul
arly
 
ex
t
r
eme
 
t
h
ermophi
l
es
 
for
 
which
 
the
optimal 
temperatures are above
 
80°C.
 
Their 
membranes 
and 
proteins are unusually 
stable at these 
extremely 
high
temperatures.
 
 
 
Thus,      
 
many      
 
important
 
biotechnological processes use
thermophilic enzymes because of their ability to withstand intense
 
heat.
 
Many of the hyperthermophiles Archea require 
elemental sulfur for
 
growth.
 
Some 
are 
anaerobes 
that 
use the 
sulfur 
instead 
of oxygen 
as 
an 
electron  
acceptor
during cellular
 
respiration.
 
Some 
are 
lithotrophs 
that 
oxidize sulfur 
to 
sulfuric acid 
as an 
energy 
source,
thus 
requiring 
the microorganism 
to be adapted to 
very low pH 
(i.e., 
it is  
an
acidophile 
as 
well 
as
 
thermophile).
 
These 
organisms 
are 
inhabitants 
of hot, 
sulfur-rich environments 
usually
associated with 
volcanism, 
such 
as 
hot 
springs, 
geysers, and
 
fumaroles.
 
T
H
E
R
M
O
P
H
I
L
E
S
Temperature 
loving
 
organisms.
M
o
s
t
 
m
e
m
b
e
r
s
 
a
r
e
 
A
r
c
h
a
e
G
r
o
w
s
 
i
n
 
a
 
t
e
m
p
e
r
a
t
u
r
e
 
r
a
n
g
e
 
o
f
 
5
5
-
1
1
3
0
C
M
o
s
t
l
y
f
o
u
n
d
i
n
g
e
o
t
h
e
r
m
a
l
l
y
h
e
a
t
e
d
r
e
g
i
o
n
s
o
n
e
a
r
t
h
v
i
z
.
,
 
h
o
t
 
s
p
r
i
n
g
s
,
 
h
y
d
r
o
t
h
e
r
m
a
l
 
v
e
n
t
s
 
e
t
c
.
As 
they need 
extreme 
temperature, its 
very 
hard 
to
 
study
them under laboratory
 
conditions.
 
hea
t
 
by
 
A
l
s
o
 
that
 
some
 
mem
bers
 
can
 
pro
du
ce
themselves 
(compost 
and garbage
 
landfills).
E
g
 
:
 
C
y
a
n
i
d
i
u
m
 
c
a
l
d
a
r
i
u
m
,
 
C
h
a
e
t
o
m
i
u
m
t
h
e
r
m
o
p
h
i
l
e
 
D
e
i
n
o
c
o
c
c
u
s
-
t
h
e
r
m
u
s
 
i
s
 
a
 
s
m
a
l
l
 
g
r
o
u
p
 
o
f
 
e
u
b
a
c
t
e
r
i
a
w
h
i
c
h
 
c
a
n
 
t
h
r
i
v
e
 
e
n
v
i
r
o
n
m
e
n
t
a
l
 
h
a
z
a
r
d
s
.
Stains 
Gram 
positive (thick cell wall) but possesses
 
an
outer membrane, similar 
to the Gram 
negative cell
 
wall.
Several thermophilic bacteria comes under 
this
 
group.
I
t
 
i
s
 
t
h
e
 
s
o
u
r
c
e
 
o
f
 
h
e
a
t
 
r
e
s
i
s
t
a
n
t
 
e
n
z
y
m
e
-
 
t
a
q
p
o
l
y
m
e
r
a
s
e
,
 
w
h
i
c
h
 
i
s
 
w
e
l
l
 
u
s
e
d
 
i
n
 
P
C
R
.
T
h
e
 
e
n
z
y
m
e
 
i
s
 
i
s
o
l
a
t
e
d
 
f
r
o
m
 
T
h
e
r
m
u
s
 
a
q
u
a
t
i
c
u
s
.
 
G
r
a
n
d
 
P
r
i
s
m
a
t
i
c
 
S
p
r
i
n
g
 
a
n
d
 
M
i
d
w
a
y
 
G
e
y
s
e
r
 
B
a
s
i
n
-
Y
e
l
l
o
w
s
t
o
n
e
 
N
a
t
i
o
n
a
l
 
P
a
r
k
,
 
U
S
A
S
o
u
r
c
e
:
 
I
n
t
e
r
n
e
t
 
C
L
A
S
S
I
F
I
C
A
T
I
O
N
 
O
F
 
T
H
E
R
M
O
P
H
I
L
E
S
 
1
.
 
O
b
l
i
g
a
t
e
 
t
h
e
r
m
o
p
h
i
l
e
s
Also known 
as 
extreme
 
thermophiles.
T
e
m
p
e
r
a
t
u
r
e
 
r
a
n
g
e
 
i
s
 
8
0
-
1
2
2
0
C
.
Membranes and proteins are unusually stable 
at
these extreme
 
temperatures.
For this reason, 
most 
biological processes utilize
thermophilic enzymes because 
of 
their ability 
to
withstand intense
 
heat.
Many 
of this 
group can 
resist 
radiation
 
too.
E
g
:
 
M
e
t
h
a
n
o
p
y
r
u
s
 
k
a
n
d
l
e
r
i
,
 
c
a
n
 
s
u
r
v
i
v
e
 
a
n
d
r
e
p
r
o
d
u
c
e
 
a
t
 
1
2
2
0
C
,
 
S
u
l
f
o
l
o
b
u
s
 
s
p
p
 
,
 
P
y
r
o
c
o
c
c
u
s
s
p
p
,
 
P
y
r
o
d
i
c
t
i
u
m
 
s
p
p
 
(
o
p
t
i
m
u
m
 
o
f
 
1
1
3
0
C
)
 
M
o
s
t
 
o
f
 
t
h
e
 
m
e
m
b
e
r
s
 
r
e
q
u
i
r
e
 
e
l
e
m
e
n
t
a
l
 
s
u
l
f
u
r
 
f
o
r
g
r
o
w
t
h
.
A
n
a
e
r
o
b
i
c
 
m
e
m
b
e
r
s
 
u
s
e
 
s
u
l
f
u
r
 
a
s
 
e
l
e
c
t
r
o
n
 
a
c
c
e
p
t
o
r
instead of oxygen in cellular
 
respiration.
S
o
m
e
 
a
r
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Rare 
group of 
organisms 
that 
can live both 
in 
higher
temperature 
and normal 
temperature 
are referred 
to
as facultative thermophiles.
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Acidophiles 
or 
acidophilic organisms 
are 
those that thrive under 
highly 
acidic
conditions (usually 
at pH 2.0 or 
below). These 
organisms 
can 
be 
found 
in 
different
branches 
of 
the tree 
of 
life, including Archaea, Bacteria, and also
 
Eukaryotes.
 
Archaea
Sulfolobes, an order in the Crenarchaeota branch 
of
 
Archaea
Thermoplasmatales, an order in the Euryarchaeota branch of
 
Archaea
Acidianus
 
brierleyi,
 
A.
 
infernus
,
 
facultatively
 
anaerobic
 
thermoacidophilic
archaebacteria
Haloarchaeum 
acidiphilum
, acidophilic 
member 
of 
the
 
Halobacteriacaeae
Metallosphaera 
sedula
,
 
thermoacidophilic
 
Bacteria
Acidobacterium, a phylum 
of
 
Bacteria
Acidithiobacillales, an order 
of 
Proteobacteria e.g. 
A.ferrooxidans, 
A.
 
thiooxidans
Thiobacillus 
prosperus, 
T. 
acidophilus, 
T. 
organovorus, 
T.
 
cuprinus
Acetobacter
 
aceti
,
 
a
 
bacterium
 
that
 
produces
 
acetic
 
acid
 
(vinegar)
 
from
 
the
oxidation 
of
 
ethanol.
Alicyclobacillus
, a genus 
of 
bacteria that 
can 
contaminate fruit
 
juices.
]
 
A
C
I
D
O
P
H
I
L
E
S
 
Microorganisms 
that 
lives 
in 
highly
 
acidic
environments are called as
 
acidophiles.
The 
pH range is
 
1-5.
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.
Mechanism 
of 
action is 
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they
 
have
 
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proton
pump machinery 
to 
eliminate protons 
from the
cytoplasm 
of the 
cell 
to 
maintain low
 
pH.
E
g
:
 
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b
u
s
 
Eukaryotes
Mucor
 
racemosus
Urotricha
Dunaliella
 
acidophila
Philodina
 
roseola
 
Acidophiles 
are 
acid-loving microbes. Most 
natural 
environments 
on the 
earth 
are
essentially neutral, 
having pH values between 
five and
 
nine.
 
Acidophiles thrive in the 
rare 
habitats 
having a pH below
 
five.
 
 
Highly acidic environments can 
result 
naturally from geochemical activities 
(such
as 
the 
production 
of 
sulfurous gases in hydrothermal vents and some 
hot 
springs)
and from 
the 
metabolic activities 
of 
certain acidophiles
 
themselves.
 
Acidophiles 
are also found 
in the debris left over 
from 
coal
 
mining.
 
Interestingly, 
acid-loving extremophiles cannot tolerate great acidity inside 
their
cells, 
where 
it would 
destroy such 
important molecules as
 
DNA.
 
They 
survive 
by keeping the 
acid out. 
But the 
defensive molecules 
that 
provide  
this
protection, 
as 
well 
as 
others that come into 
contact with 
the environment, 
must  be
able to operate in extreme 
acidity. 
Indeed, extremozymes 
that 
are able to 
work  
at 
a
pH 
below one--more acidic 
than 
even vinegar 
or 
stomach fluids--have been
isolated from 
the 
cell wall and underlying cell 
membrane of 
some
 
acidophiles.
 
 
M
o
s
t
 
a
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d
o
p
h
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l
e
 
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v
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e
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e
x
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m
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m
p
 
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s
 
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t
 
o
f
 
t
h
e
 
i
n
t
r
a
c
e
l
l
u
l
a
r
 
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p
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i
n
 
o
r
d
e
r
t
o
 
k
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p
 
 
t
h
e
 
c
y
t
o
p
l
a
s
m
 
a
t
 
o
r
 
n
e
a
r
 
n
e
u
t
r
a
l
 
p
H
.
 
Therefore, intracellular 
proteins do not 
need to 
develop 
acid 
stability
through 
evolution. 
However, 
other 
acidophiles, 
such 
as 
Acetobacter aceti
,
have an 
acidified 
cytoplasm which 
forces nearly all proteins 
in 
the
genome to evolve 
ACID
 
stability.
 
For 
this 
reason, 
Acetobacter aceti 
has become 
a 
valuable resource
 
for
understanding the 
mechanisms by 
which proteins can attain acid
 
stability.
 
Studies of 
proteins adapted 
to 
low pH have revealed 
a 
few 
general
mechanisms 
by which proteins can achieve acid
 
stability.
 
prot
e
in
 
f
r
om
 
Sulpho
l
ob
u
s
 
a
c
ido
c
alda
r
ius
),
 
t
h
ere
 
is
 
an
 
 
In
 
m
ost
 
a
c
i
d
 
stab
l
e
 
pr
o
t
e
ins
 
(such
 
as
 
pepsin
 
a
nd
  
the
soxF
         
over
 
abundance of 
acidic 
residues which 
minimizes low pH destabilization
induced by a buildup of positive
 
charge.
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Extremophiles are organisms that thrive in extreme conditions such as high temperatures, acidity, salinity, and more. They have unique adaptations like extremozymes and can withstand harsh environments where life was once deemed impossible. Types include psychrophiles and Firmicutes, showcasing remarkable abilities to survive and even play important roles in various industries.

  • Extremophiles
  • Extreme Environments
  • Adaptations
  • Firmicutes
  • Psychrophiles

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  1. EXTREMOPHILES: Thermophiles & Acidophiles

  2. EXTREMOPHILES Organisms found living in extreme harsh environments. Word originated from Greek- Extremus + Philiawhich means extreme loving Definition: Extremophiles are organisms that have been discovered on earth that survive in environments that were once thought not to be able to sustain life. Most members of this group comes under the domain Archae. These include thermophiles, hyperthermophiles, thermoacidophiles, alkaliphiles, psychrophiles, halophiles, barophiles, radiation resistant bacteria and endoliths.

  3. Adaptations Extremozymes- specialized enzymes that are highly stable. Can tolerate extremes of temperature, pH, salinity that would inactivate other enzymes. Important in industries because of this property.

  4. EXTREME CONDITIONS Temperature pH Salinity Nutritional scarcities Absence of oxygen Radiation Pressure

  5. Response to Environmental factors

  6. TYPES Psychrophiles Temperatue range is -15 to 150C Also known as cryophiles. Have an optimum temperature of 150C orlower Isolated from Arctic and Antarctic habitats (90% ofthe ocean is 50C or colder) Also found in ice bergs, glaciers, snowfieldsetc Metabolism is quite normal at colder temperatures. Cell membranes-high levels of fatty acids whichremain fluid at colder temperatures. Proteinaceous antifreeze mechanism to protect thecell and DNA Some of them cause spoilage in refrigerated food materials. Eg: Arthrobacter spp, Psychrobacter spp,Halomonas spp, Pseudomonas, Sphingomonas

  7. FIRMICUTES Gram positive, spore forming bacterial family that can survive desiccation and can survive extreme conditions. This group also is an example for extremophilic true bacteria (eubacteria). Plays an important role in the spoilage of beer, wine and cider. Eg: Helicobacterium spp, Mycoplasma, Clostridium spp.

  8. Many members of the Family Firmicutes are also thermophiles. Eg: Bacillus stearothermophilus Recently, a DNApolymerase derived from these bacteria, Bst polymerase has become important in biotechnology. Bst polymerase- helicase like activity (making it able to unwind DNAstrands. Optimum functional temperature is 60-650C and get inactivated at temperatures above 800C

  9. A thermophile is an organism a type of extremophile that thrives at relatively high temperatures, between 45 and 122 C (113 and 252 F). Many thermophiles are archaea. Thermophilic eubacteria are suggested to have been among the earliest bacteria. "Thermophile" is derived from the Greek: (thermotita), meaning heat, (philia), love. and Greek: Thermophiles are classified into obligate and facultative thermophiles: thermophiles (also called extreme thermophiles) require such high temperatures for growth, Obligate whereas facultative thermophiles (also called moderate thermophiles) can thrive at high temperatures, but also at lower temperatures (below50 C). Hyperthermophiles are particularly extreme thermophiles for which the optimal temperatures are above 80 C. Their membranes and proteins are unusually stable at these extremely high temperatures.

  10. Thus, thermophilic enzymes because of their ability to withstand intenseheat. many important biotechnological processes use Many of the hyperthermophiles Archea require elemental sulfur forgrowth. Some are anaerobes that use the sulfur instead of oxygen as an electron acceptor during cellular respiration. Some are lithotrophs that oxidize sulfur to sulfuric acid as an energy source, thus requiring the microorganism to be adapted to very low pH (i.e., it is an acidophile as well as thermophile). These organisms are inhabitants of hot, sulfur-rich environments usually associated with volcanism, such as hot springs, geysers, and fumaroles.

  11. THERMOPHILES Temperature loving organisms. Most members are Archae Grows in a temperature range of 55-1130C Mostly found in geothermally heated regions on earth viz., hot springs, hydrothermal vents etc. As they need extreme temperature, its very hard to study them under laboratory conditions. Also themselves (compost and garbage landfills). Eg : Cyanidium caldarium, Chaetomium thermophile that some members can produce heat by

  12. Deinococcus-thermus is a small group of eubacteria which can thrive environmental hazards. Stains Gram positive (thick cell wall) but possesses an outer membrane, similar to the Gram negative cell wall. Several thermophilic bacteria comes under this group. It is the source of heat resistant enzyme- taq polymerase, which is well used in PCR. The enzyme is isolated from Thermus aquaticus.

  13. Grand Prismatic Spring and Midway GeyserBasin- Yellowstone National Park, USA Source: Internet

  14. CLASSIFICATION OF THERMOPHILES 1. Obligate thermophiles Also known as extreme thermophiles. Temperature range is 80-1220C. Membranes and proteins are unusually stable at these extreme temperatures. For this reason, most biological processes utilize thermophilic enzymes because of their ability to withstand intense heat. Many of this group can resist radiation too. Eg: Methanopyrus kandleri, can survive and reproduce at 1220C, Sulfolobus spp , Pyrococcus spp, Pyrodictium spp (optimum of 1130C)

  15. Most of the members require elemental sulfur for growth. Anaerobic members use sulfur as electron acceptor instead of oxygen in cellular respiration. Some are lithotrophs that oxidizes sulfur to sulfururic acid as an energy source. Such organisms require a very low pH and hence known as thermoacidophiles. Inhabits regions associated with volcanic eruption viz; hot, sulfur rich, acidic regions such as hot springs, natural geysers, fumaroles etc .

  16. HABITATS OFEXTREMOPHILES Hot spring situated in Atlanta, USA Courtsey: http://www.idahohotsprings.com/destinations/atlanta/atlanta_hot_ springs_01.jpg

  17. Castle Geyser, Yellowstone National Park, USA Courtesy: http://upload.wikimedia.org/wikipedia/commons/4/49/Steam_Phase_eru ption_of_Castle_geyser_with_double_rainbow.jpg

  18. Black smoker at a mid-ocean ridge hydrothermal vent

  19. Thermoacidophiles Requires both high temperature and highly acidic environment for optimum growth. Preferred temperature range is 70-800C and have anoptimum pH range of 2-3. All the organisms discovered belongs to the DomainArchae, so far. They can thrive in acidous and sulfur richenvironments. Instead of cell wall, possesses a unique membrane composed of tetraether lipoglycan, which gives the unusual stability for the bacteria. Eg: Thermoplasma acidophilum and T.volcanium

  20. Facultative Thermophiles Rare group of organisms that can live both in higher temperature and normal temperature are referred to as facultative thermophiles. These organisms can live at 200C, and have an optimum of 500C. Maximum temperature that they can survive is 600C. Eg: Bacillus flavothermus

  21. Acidophiles or acidophilic organisms are those that thrive under highly acidic conditions (usually at pH 2.0 or below). These organisms can be found in different branches of the tree of life, includingArchaea, Bacteria, and also Eukaryotes. Archaea Sulfolobes, an order in the Crenarchaeota branch of Archaea Thermoplasmatales, an order in the Euryarchaeota branch of Archaea Acidianus brierleyi, A. infernus, facultatively anaerobic thermoacidophilic archaebacteria Haloarchaeum acidiphilum, acidophilic member of the Halobacteriacaeae Metallosphaera sedula, thermoacidophilic Bacteria Acidobacterium, a phylum of Bacteria Acidithiobacillales, an order of Proteobacteria e.g. A.ferrooxidans, A. thiooxidans Thiobacillus prosperus, T. acidophilus, T. organovorus, T. cuprinus Acetobacter aceti, a bacterium that produces acetic acid (vinegar) from the oxidation of ethanol. Alicyclobacillus, a genus of bacteria that can contaminate fruit juices. ]

  22. ACIDOPHILES Microorganisms that lives in highly acidic environments are called as acidophiles. The pH range is 1-5. Some members that mainly found in the drainage of coal mines are able to oxidize sulfur into sulfuric acid. Mechanism of action is that they have a proton pump machinery to eliminate protons from the cytoplasm of the cell to maintain low pH. Eg: Pyrodictium, Picrophilus, Ferroplasma, Sulfolobus

  23. Eukaryotes Mucor racemosus Urotricha Dunaliella acidophila Philodina roseola Acidophiles are acid-loving microbes. Most natural environments on the earth are essentially neutral, having pH values between five and nine. Acidophiles thrive in the rare habitats having a pH below five. Highly acidic environments can result naturally from geochemical activities (such as the production of sulfurous gases in hydrothermal vents and some hot springs) and from the metabolic activities of certain acidophiles themselves. Acidophiles are also found in the debris left over from coalmining. Interestingly, acid-loving extremophiles cannot tolerate great acidity inside their cells, where it would destroy such important molecules as DNA. They survive by keeping the acid out. But the defensive molecules that provide this protection, as well as others that come into contact with the environment, must be able to operate in extreme acidity. Indeed, extremozymes that are able to work at a pH below one--more acidic than even vinegar or stomach fluids--have been isolated from the cell wall and underlying cell membrane of some acidophiles.

  24. Most mechanisms to pump protons out of the intracellular space in order to keep the cytoplasm at or near neutral pH. acidophile organisms have evolved extremely efficient Therefore, intracellular proteins do not need to develop acid stability through evolution. However, other acidophiles, such as Acetobacter aceti, have an acidified cytoplasm which forces nearly all proteins in the genome to evolveACIDstability. For this reason, Acetobacter aceti has become a valuable resource for understanding the mechanisms by which proteins can attain acid stability. Studies of proteins adapted to low pH have revealed a few general mechanisms by which proteins can achieve acid stability. In soxF most acid stable proteins (such as pepsin and the over protein from Sulpholobus abundance of acidic residues which minimizes low pH destabilization induced by a buildup of positive charge. acidocaldarius), there is an

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