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another cell who’s DNA had been removed and

produced a functioning living cell


Origins of Life
we can also build virus particles by hand and they act like
Why is there such a diversity of living forms yet all natural virus particles
life shows remarkable similarities:
but we cannot yet build a cell completely from
1. genetic code = DNA scratch
!same basic code for all forms of life ……………………………………………………
Previous ideas on the Origin of Life
2. common chemicals & metabolic pathways
eg. 20 amino acids used by all life
Until mid 1800’s it was generally thought that life
3. all made of cells could arise by spontaneous generation (600BC-
!cell structures are amazingly similar 1800’s)
only two basic forms:
procaryotes and eucaryotes
=Nonliving matter had capacity within itself to
because of so many shared traits biologists have turn into certain types of living organisms
assumed that all live had a common ancestor eg: moist soil ! toads, snakes, mice
manure ! flies
in 2010 a new modeling study of life at a chemical level added
fruits ! molds
significant additional support to this idea
believed there was some “vital force” in the elements
Darwin provided a mechanism that showed how all that could transform matter
life could be interrelated
Late 1600’s: Redi performed experiments that dealt
! But, how did it all start? 1st major blow to this theory

Later: Spallanzani -- sealed flasks


today it is impossible to simply take all the ingredients
of life, in the proper proportions, mix them Pasteur: Final downfall of the idea
showed it included microrganisms
together and get life
BUT: This left biologists with no plausible hypothesis
can’t even do it for a single cell for the Origin of Life until fairly recently
but we have recently (2010) succeeded in constructing the ……………………………………………………
complete DNA of a bacterial cell and inserting it in
Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 1 Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 2

since Darwins time we have learned how to very Major Events in the Origin of Life:
accurately date rocks & fossils by radioactive
decay 1. Suitable Environment
Decay System Half Life Useful Timespan
Rubidium! Strontium
Lutetium ! Hafnium
1.42X10-11 yr-1
1.94x10-11 yr-1
48.8 BY
35.7±1.2BY
2. Formation of Basic Building Blocks
Uranium! Lead 1.55 X10-10 yr-1 4.47BY
Potassium ! Argon 0.581 X10-10 yr-1 1.93BY
14
C 5568±30 yr3 3. Metabolism & the Formation of Large
Polymers
from the Fossil record we have learned:
4. Compartmentalization; ie. Cells
!the early earth was very different from today
5. Operating Instructions
!all forms of life did not appear at the same time
These are probably not sequential events:
!life progressed from small, simple forms to
more complex and larger forms most or all probably occurred at the same time
eg. interstate system: none ! fully developed

not: first roads, then gas stations, state police


system, then dealerships, etc

Our thoughts on what the first cell were like can only
be based on what cells are like today, but…
it is possible the first cells were considerably less complex and
efficient ! perhaps very little like cells around today

Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 3 Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 4
1. Suitable Environment within about 150 MY of earth’s formation, it
collided with a smaller planet to form the
4.54 BY: Earth formed moon

the sun is a second or third generation star it remelted the earth and reset the timescale of
life’s origin
even today, a star explodes each second with a
brightness greater than a galaxy oldest known piece of earth’s crust is 4.4 BY old

! all matter on earth has been through 1 or 2 as earth cooled, water condensed to form rivers,
previous cycles of “stellar alchemy” lakes and oceans

sun and planets form from previous supernova by 3.9 or 4 BY:


explosion producing cloud of dust and gas
intensive bombardment slowed
heavy bombardments by comets and meteorites
(size of Ohio) – each impact would have crust cooled and solidified
caused any liquid water to boil away
completely cooled enough for liquid water to collect in basins
when the moon formed by a grazing impact of a mars-sized
body it would have completely destroyed any life that
intense storms
might have arisen
atmospheric water condensed to form oceans
even today 40,000 tons of space debris fall to earth each
year, mainly as dust most of this water came from volcanic activity

surface temperatures up to 1000-3000º C up to 1/3rd of water on the earth (and in us) may have
come from comets
no solid ground
even today 10 M small comets hit earth each yr
no liquid water (no oceans, lakes) ! each comet = wt of 60 compact cars together
only steam from geysers, volcanoes
if this is same as rate throughout earths history it would
equal the volume of today’s oceans

Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 5 Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 6

earth’s early atmosphere was derived mainly


from volcanoes: if life took much longer to appear; eg. several billion
years, it would imply that the origin of life is an
contained: H2O, CO2, N2, CO, H2, H2S,
HCl, HCN (reducing atmosphere)
extremely unlikely event and may have implications
for finding life on other planets

[todays atm: 78% N2, 20% O2, 4% H2O, 0.03% CO2] some believe that the spontaneous origin of life
would be impossible:
CHONPCa all major atoms were available
the odds of life arising on the early earth are about the
there remained lots of energy sources on the early same as the odds of a tornado going through a
earth: junkyard and randomly assembling a fully functional
jumbo jet, 747

!lots of volcanic activity unfortunately, there is no rock left on the surface


!severe lightning storms of the earth older than 3.85 BY due to earth’s
!unshielded solar radiation: xrays and UV active geology
(no ozone layer yet)
!more radioactive materials ! everything older has been lost to weathering and
continental drift
most agree there was a suitable environment some suggest that the best place to look for signs for the
for life by 3.9 BY ago origin of life are on the moon

How quickly did life appear after this? some estimate there are thousands of tons of earth
rock on the moon

if the origin of life occurred relatively quickly after if we can find rocks older than 3.5 BY we might be
conditions were suitable, it would indicate that life able to find fossils or signs of life in them
did indeed begin very simply and that natural laws
earliest true fossils appear 3.5 BY
are conducive to its origin given the suitable
conditions and materials;
oldest “trace fossils” = 3.85 BY, Greenland
! the origin of life is inevitable given the proper
conditions
Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 7 Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 8
by 3.8 BY rocks show high concentrations of 2. Formation of Basic Building Blocks: small
carbon; indications are that life was thriving organic molecules
then
~4.0-3.9 BY ago there were ideal conditions for
!life must have begun 4.0 to 3.8 BY ago chemical reactions that could produce small
organic molecules:
geological evidence also points to the fact that
sugars, amino acids, nucleotides, fatty acids, etc
this earliest indications of life were due to
photosynthetic life Miller and Urey (1953) and others later modeled the
composition of the early ocean & atmosphere and
but photosynthysis is an ‘advanced’ form of used electrical sparks to mimic energy
energy procurement so life must have
appeared BEFORE 3.8 BY produced:
all 20 amino acids
As soon as there was a suitable environment, several sugars
life showed up lipids
purines and pyrimidines
short chains of nucleotides
ATP
short chains of nucleotides

Miller and others (early 1950’s) could produce building


blocks of virtually all small organic molecules
given early atmosphere and energy

these same reactions could also have happened at


deep ocean vents on early earth

but generally cannot occur in the presence of O2

Also, comets and meteorites that helped to form the


early earth brought organic molecules
eg. amino acids are fairly common in these objects
Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 9 Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 10

to form polypeptides
3. Metabolism and the Formation of large
some believe, given enough time, such reactions were
complex, interacting inevitable on early earth

even a single living cell consist of 1000’s of highly also, HCN was common then and is commonly
organized chemical reactions, all occurring at the used as a catalyst in chemistry labs
same time
over millions of years large polymers could
all these reactions together are called metabolism conceivably be constructed

in living cells today, each of these reactions requires a another question is how any reactions got organized
specific enzyme (= organic catalyst) into something we could call metabolism

! but enzymes are proteins (large polymers); modern cells today have 10’s of 1000’s of
proteins weren’t around yet chemical reactions occurring each second

without enzymes, need high temperatures or some all of these reactions are interlinked and
kind of catalyst to do these reactions coordinated into numerous reaction pathways

there were many high energy sources in these scientists today believe that all these complex
early days: pathways could have originated from the most
basic set of reactions that occur in every cell
!lots of volcanic activity
! the energy producing pathway
! many more deep ocean thermal vents
the simplest such pathway may include only a
!horrendous thunder & lightning few dozen interacting molecules

!lots of UV radiation (earth had no ozone from this set of chemicals all other reactions
shield) could be derived

!meteorite impacts produce enough energy


Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 11 Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 12
4. Compartmentalization; ie. Cells
the more organic matter present the greater
the internal chemical environment must be amount of foam
isolated from external environment
the early ocean contained lots of organic
large organic molecules must be enclosed and matter and produced abundant ocean
contained or at least collected together in one foam
place
this foam is somewhat similar to the cell
must have high concentrations of interacting membranes of living cells (lipid bilayer
organic molecules to approximate with proteins)
metabolism in living organisms
it is permeable ! things can move in and
several possible scenarios have been proposed: out

a. clay particles these bubbles can grow and fuse with


other bubbles
the chemicals in clay allow it to absorb and
hold onto many kinds of organic molecules over time we would get a kind of chemical
natural selection
areas on the sea floor with high
!those droplets that were most stable and best able to
concentrations of clay could absorb and accumulate organic molecules would grow and split
concentrate these organic chemicals into
functional groupings !other droplets would fall apart or fail to grow and divide

b. iron pyrite around hydrothermal vents !the “environment” would “select” for some over others

could also do that


5. Operating Instructions
c. ocean foam
to understand the origin of life we must explain
how these collections of organic chemicals
organic matter particularly fats and oils tend
to form hollow spheres
Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 13 Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 14

could become organized into living self one way to get an idea of the likelihood of life
autonomous, reproducing cells appearing is to consider how ordered life is
compared to nonliving matter
cells need instructions to operate propery
how many “instructions” do cells need to function
! need to control metabolism properly
! need to be able to copy and pass on these
instructions to new cells we can view the genetic code as “information storage”
of data bytes
today the DNA in chromosomes contain all the
each byte is 1 unit of information (1 byte = 8 bits)
instructions for life
gigabytes (billions of bytes)
Shared Genes
= the amount of information stored in typical eucaryote
the simplest life today are the procaryotes cells today

eg. DNA in human cells contains of 1 gig of RAM in each


there are two major kinds of procaryotes; the archea cell (~21,000 genes)
and the eubacteria
comparable to information levels today’s computers

they most likely developed from the same ancestor megabytes (millions of bytes)

by looking at genes shared between them we can get = the amount of information stored in typical procaryote
cells (~4000 genes)
an idea of what their common ancestor was like
comparable to the info storage in old Pentium and
60 genes are shared between the two groups Power PC computers

kilobytes
Information from how these genes work suggest that
the earliest cells might have lived in volcanic hot =the amount of information stored in the simplest
springs procaryote cells capable of independent existence

eg. the simplest organism capable of independent existence


Life as Information Storage has 60 Kilobytes of information, or ~200 genes

Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 15 Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 16
comparable to the first desktop computers on the
market
RNA uses that code to make either
eg. even (nonliving) viruses with 2% as much genetic
info as E coli have ~15 kilobytes enzymes for metabolism
or
eg. one typical gene contains ~ 300 bytes new copies of DNA for reproduction
some biologists are trying to determine the minimum
number of genes that any living organism needs enzymes metabolism

they genetically engineer bacteria by sequentially removing


DNA RNA
new DNA reproduction
genes, one at a time, to determine which genes, and the
minimum number of genes needed for life
A simpler genetic code?
so far the most likely estimate is that any living
cell needs at least 151 genes for life or ~45 metabolism is the result of the activity of 10’s of
kilobytes of information 1000’s of different enzymes

biologists have succeeded in randomly producing very enzymes are some of the largest, most complex
small DNA molecules of about 50 nucleotides organic molecules in a cell
from a mixture of simple building blocks
computer analysis indicates that today’s enzymes
! much less than that needed for even 1 gene
could have originated from a very few, much
based on natural laws, the early earth could have simpler enzyme molecules constructed with a
randomly accumulated no more than ~25 bytes much simpler genetic code than the one in cells
of information by pure chance today

if life on earth had a spontaneous origin there must A simpler molecule than DNA?
have been some simpler intermediate state
many believe that for life to appear some simpler
In every cell today: genetic material had to be there 1st

DNA stores the genetic code ! probably RNA appeared 1st


Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 17 Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 18

new (2010) studies indicate that RNA molecules could


1. RNA is a much simpler molecule than DNA actually have formed easily under conditions on
the early earth
2. RNA carries the same kind of genetic
information as does DNA the packaging of RNA “genes” and their enzyme
products within some kind of container would
3. RNA is able to replicate itself have been a significant milestone in the origin of
life:
4. all life requires RNA to make DNA
would now have had all basic requirements
5. all life requires RNA to make proteins for life together and interacting
(enzymes)
!these simple cells could then “evolve” as units
6. RNA can act directly as enzyme or
catalyst
!favorable interactions could have been selected,
7. some viruses have only RNA as genetic preserved and perpetuated
material and are able to function
! Natural Selection could occur
8. all components of RNA: sugar, phosphate and
N-bases could have been formed under RNA “life” could have arisen ~ 4BY ago and lasted for
primitive earth conditions ~ 200MY

many traces of the ancient RNA-dominated world RNA is not very stable, so cells that began to
remain in our genome today store information in a much more stable but
similar molecule, DNA, became much more
eg. ALL life today contains a particular sequence that codes for successful
an RNA enzyme that plays a key role on the synthesis of
proteins
once DNA appeared, natural selection would have
eg. at least 100 genes can be traced back to this “universal greatly favored it over RNA
common ancestor to ALL life
eventually ALL the RNA only cells would have
gone extinct, leaving no trace
Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 19 Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 20
= autotrophs (=”self feeders”)
What were the first cells:
other organisms must consume preformed organic
1. 1st cells were Prokaryotes molecules to break down for energy
= heterotrophs ( ~”feed on others”)
the earliest fossils we find are prokaryotes
of the two, autotrophs require considerably more
the only fossils we find for the first 2 Billion cellular “machinery” to make their own
years of life are prokaryotes sugars more genes are required; more
complex
!small, simple, inefficient metabolism, little
internal structure 3.9 BY ago the oceans were thick with 100’s
Millions of years of accumulated organic
2. simpler cell membrane chemicals

protein analysis indicates that the earliest cell There were plenty of organic molecules after 100’s
membranes were much simpler and “leakier” of millions of years of earth history
than modern cell membranes
life didn’t require primary producers
allowed much greater exchange of genes, ! the planet was awash in “food”
nutrients, enzymes, proteins and products
! First cells were simpler heterotrophs
3. First cells were heterotrophs
! able to capitalize on this abundance of
all life requires nutrients and energy nutrients

all life produces the energy they need by breaking 4. First cells produced energy anaerobically
down sugars and other organic molecules
= respiration there was no free oxygen (O2) on the early earth,
either in atmosphere or in oceans
some organisms are able to make their own
sugars the breakdown of sugars to produce energy
Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 21 Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 22

= respiration
= fermentation
respiration can occur with or without O2
fermenters excrete acids, alcohols, etc.
! with O2 the process is very efficient
(chemicals that contain less energy than the food
but more complex ingested)

requires additional pathways and vs aerobic respiration that produces CO2 & H2O
enzymes, ie. more genes
= aerobic respiration 5. 1st cells used RNA as genetic instructions

! without O2 some energy can be made RNA molecular evidence supports the idea that for
the 1st billion years of life RNA was the
it is a much simpler process, fewer primary genetic material
enzymes required, fewer genes
the genes that work with DNA are much younger
= anaerobic respiration

but nowhere near as efficient

( 2 units of energy vs 34 units of energy per


sugar molecule)

!first cells were anaerobic heterotrophs who


broke down sugars without the use of O2 gas

they took organic molecules, extracted


energy from them and produced organic
molecules as waste products

! this is the simplest form of energy


production
Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 23 Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 24
Where did life begin? found in bacteria in such habitats today seem
to be precursors to other genes
A. Shallow warm water
C. some other planet or solar system
on soft clay sediments
=panspermia; ie. seeded from space
habitat closest to suspected energy source needed
for “prelife” events 100’s to 1000’s of meteorites and comets
brought organic molecules from space which
some of earliest fossils are stromatolites: banded could have jump started the process
domes of sedimentary rock and bacterial mats
some suggest bacteria may have traveled
these form today in shallow salt marshes and from other places (Mars meteorite-now
warm lagoons disproven)

consist of colonies of bacteria and cyanobacteria bacteria, esp spores, can survive very harsh
in jellylike secretions interspersed with conditions
sediment layers eg. some bacteria (Microbispora) that were on the shuttle
Columbia, survived the fiery reentry
produces a banding pattern
but: but only traveled about 1/5th the speed of a
meteorite
B. Deep Ocean Thermal Vents
still survived extreme heat and impact
less exposed seafloor and
that was 6 times faster than anything tested before

molecular biology supports idea that earliest life ! just defers the problem
thrived in hot conditions and may have used
sulfur compounds – common conditions in
vent communities

structure of proteins and genes resembles those


Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 25 Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 26

LUCA (Last Universal Common Ancestor) found in all other kingdoms today were
already present in bacteria
for the first billion years or so of earths history there
with the exception of only a few exotic chemicals
lived a single kind of organism in the earth’s
oceans = LUCA eg, essential oils and hallucinogens of flowering
plants
the ocean was a global genetic swap shop for 100’s of
eg. animal venoms
millions of years
!photosynthetic bacteria were producing oxygen
! free exchange; no competition gas; building up in the atmosphere

probably all the cells in the world’s oceans shared the earth’s modern atmosphere and surface
genes, proteins, nutrients, enzymes, metabolic were largely established by bacteria
pathways
it was only when some of the cells evolved ways to
none lived independently from the others produce everything they needed within their own
cell did life begin to diversify into the major
they were all interdependent on each other domains that exist today:
archaea
by ~1.5 BY ago most “biochemical evolution” had bacteria
been accomplished: eukaryotes

these early prokaryotes, working together, could


assemble and disassemble all molecules of
modern life; virtually all metabolic pathways:
photosynthesis
fermentation
aerobic respiration

!all the unique and unusual metabolic pathways

Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 27 Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 28
Major Events in the History of Life
2. Aerobic Respiration (?2.0 by)
3.85 BY
indirect evidence that life may have existed 3. Eukaryotes (2.9 by)

3.5 BY for the first billion years or so of earths history there


1st true fossils: very simple cells = prokaryotes lived a single kind of organism in the earth’s
oceans
11 different species have been found

Once the first living cells arose, 3.8 BY ago, Natural the ocean was a global genetic swap shop for 100’s of
Selection easily explains the appearance of all millions of years
other species
by ~1.5 BY ago most “biochemical evolution” had
Chemical evolution became biological evolution been accomplished:
_____________________________________
!prokaryotes could assemble and disassemble all
In 1800’s, Challenger Expedition – deep sea collections
molecules of modern life; virtually all
! thought they had found the first life forms: metabolic pathways:
Bathybius haekelii – “primordial slime” photosynthesis
fermentation
! turned out to be precipitate formed when sediment aerobic respiration
was mixed with alcohol
!all the unique and unusual metabolic pathways
Once life arose, what were some of the major events
found in all other kingdoms today were
that occurred from then until now?
already present in bacteria
Major events in History of Life (fossils): with the exception of only a few exotic chemicals

eg, essential oils and hallucinogens of flowering


1. Photosynthesis (~3.5 by) plants

around 2.9 BY ago oxygen began to eg. snake venoms


accumulate in the earths atmosphere
Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 29 Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 30

!the earth’s modern atmosphere and surface


were largely established by bacteria 4. Multicellular Forms (~0.8 by)
first animals
first fungi
!bacteria invented asexual and a simple form of first plants
sexual reproduction

many of the bacteria alive today seem to have


changed very little from those earliest ones:
eg. stromatolites are still formed today

eg. bg bacteria seem to have changed little

even today bacteria are the most tenaceous beings


known

they grow and divide quickly


! life cycle in minutes not days or years

bacteria are essentially immortal


! they don’t age as complex cells do.

bacteria can mutate & exchange genes quickly


and freely to adapt quickly to changing
conditions

about 2.9 BY ago the original group of


prokaryotes split into 3 major “domains”
of life that we have today

bacteria
archaea
eukaryotes
Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 31 Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 32
What is life?
History of the Earth in a Single Year
days yrs
b. p. “Living things are systems that tend to respond to changes
(1) Jan 1, 12:00am - earth formed (4.6 BY) in their environment, and inside themselves, in such a way as
(63) May 5: - 1st Cells (3.8 BY) to promote their own continuation.”
-Witzany
(135) May 16: - O2 producing photosynthesis (2.9 BY)
“Life is the representation, the presencing of past chemistries,
(206) July 26: - aerobic respiration (~2.0 BY) a past environment of the early Earth that, because of life,
(222) Aug 11: - origin of eukaryotes (1.8 BY) remains on the modern Earth.”
-Margulis & Sagan
(301) Oct 28: - multicellularity-1st animal fossils (800 MY)
“It is the watery, membrane-bound encapsulation of
(320) Nov 16: - O2 levels near 20%; Cambrian Explosion (570 MY) spacetime.”
-Margulis & Sagan
(324) Nov 20: - earliest vertebrate (520 MY)
st “Life is a potentially self-perpetuating open system of linked
(329) Nov 24: - 1 plant fossils (460 MY)
organic reactions, catalyzed simultaneously and almost
(335) Nov 30: - 1st land vertebrate fossils (380 MY) isothermally by complex chemicals that are themselves
produced by the open system.”
(345) Dec 10: - Earth’s greatest mass extinction (250 MY)
-author unknown

(353) Dec 18: - 1st fossil bird (150 MY)


“Life is a nexus of increasing sensitivity and complexity in a
(355) Dec 21: - 1st fossils of flowering plants (125 MY) universe of parent matter that seems stupid and unfeeling
in comparison.”
(359) Dec 25: - end of dinosaurs; beginning of age
-Margulis & Sagan
of mammals;1st primate fossils (65 MY)

(363) Dec 28: - spread of grasses “A network of inferior negative feedbacks subordinated
to a superior positive feedback.”
(364) Dec 30, 9:00pm: - 1st human fossils -Korzeniewski

Dec 30, 11:24pm: - 1st Homo sapiens fossils “Preserving the past, making a difference between past
and present, life binds time, expanding complexity and
creating new problems for itself.”
-Margulis & Sagan

“Life is the state of being of proteins.”


-Engels

Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 33 Life, Biodiversity, History: Origins of Life, Ziser, Lecture Notes, 2015.1 34

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