The Origin of Life On Earth

Foundations to Chemistry - adapted from "Chemistry, Matter and the Universe" Page 1 of 2
26. Origin of Life on Earth -- Jump to --
Introduction
Primates are naturally curious, and this curiosity is most highly But as we gradually come to understand our fellow creatures and
developed in Homo sapiens. to realize our biological kinship, the question has broadened to the
more comprehensive one:"Where did life come from?"
The question "Where did we come from?" has been one of the
most compelling quandaries for as long as man has been able to Two possibilities exist, special creation or spontaneous
frame enquiries. In one guise or another, this question has been at generation.
the root of most religions.
Special creation has long been the purview of theologians. For
http://www.chem.ox.ac.uk/vrchemistry/chapter26/page01.htm 2006/12/27
As long as animals and the rest of Earth's creatures were many centuries, the rational view was considered to be that of
considered only automata, as Descartes characterized them, or as spontaneous generation
subordinate creatures placed here for our express benefit, the
question of origins was narrowly confined to man.
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Introduction
Every practical observer of the world around him knew that life develops
spontaneously from nonliving matter by the action of heat, light, moisture, and
(after it was discovered) electricity. Maggots come from decaying meat, and
lice from sweat-soaked clothing. Beetles develop from rotting wood, and
horseflies from transmuted manure.
It is difficult to put forward so thoroughly eroded an idea as spontaneous

generation today without arousing smiles from the listeners. If ever a generally
accepted idea was revealed by careful experiments to be nothing but old wives'
tales, spontaneous generation was.
Francisco Redi demonstrated more than 300 years ago that meat, shielded
from egg-laying flies by cloth, never developed maggots. Others following him
showed that nutrient broths that are boiled and then kept isolated from airborne
contamination never produce microorganisms.
Spontaneous generation died hard; its proponents claimed that the life forces
were delicate and were destroyed by boiling. The early experiments were
crude, and failed just often enough to keep the controversy alive.
This reluctance to abandon spontaneous generation was not an example of the

obstinacy of the superstitious, but was the stubbornness of those who
considered themselves defenders of the rational approach, and the only
alternative to divine whimsy.
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Introduction
The defenders were wrong. Louis Pasteur sealed the fate of
spontaneous generation in a series of careful experiments, in
1861. He demonstrated clearly that microorganisms are
carried in the air, and that they grow in previously sterilized
broths only when the broths are contaminated by air or similar
sources.
"All Life from Life" became one of the fixed and immutable
points of biological dogma. This led to a dilemma that has
been expressed as the chicken-and-egg paradox.
Which came first, the chicken or the egg? If all eggs come
only from chickens. and if all chickens come only from eggs,
then there must once have been either a first chicken or a first
egg. This demanded a Creator, a celestial clockmaker who at
least set the entire machinery of life in motion before stepping
back to let things take their "natural" course thereafter.
The operations of life and the mechanisms of life hence were

areas of fruitful research, but the origin of life was not a
legitimate subject for scientific investigation. Pasteur
apparently had disproved the only theory of the origin of life
that was subject to scientific testing.
While Pasteur was tamping the last dirt over the grave of
spontaneous generation, another extraordinarily important
idea was developing in biology - one that would not have its
impact on chemistry for nearly a century. This was the theory
of evolution, as proposed by Charles Darwin, Alfred Wallace,
and the very able propagandist, Thomas Huxley.
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Introduction
Reproduction in all living creatures is never perfect. Variations This is the key to the chicken-and-egg paradox. If we trace the
show up in the offspring, which give them different efficiencies in evolution of chickens and eggs back far enough, we will not find a
meeting the challenges of any given environment. first Egg. Instead, we will realize slowly that we are not looking at
chickens any more, but at feathered reptiles.
The environment exerts a selective action on the population of
offspring: The best adapted survive in the greatest numbers to Tracing the line back further, we will see amphibia, bony fish,
breed and produce new offspring. Thus the traits that encourage cartilaginous fish, and invertebrates. The trail, if pursued long
survival in any environment are preserved. enough, leads back to one-celled life.
As adaptation to a given environment improves, and as But where did this one-celled life come from? Is a bacterium-and-
environments gradually change on the planet, the organisms spore paradox any less frustrating than the chicken-and-egg?
themselves change, adapt, and evolve.
Unless we suffer from mental fatigue or atrophied curiosity along
the way, we must eventually ask "Where did the earliest one-
celled life come from?" With such simple organisms, the problem
becomes as much chemical as biological.
Page 4 of 36
Introduction
Reproduction in all living creatures is never perfect. Variations This is the key to the chicken-and-egg paradox. If we trace the
show up in the offspring, which give them different efficiencies in evolution of chickens and eggs back far enough, we will not find a
meeting the challenges of any given environment. first Egg. Instead, we will realize slowly that we are not looking at
chickens any more, but at feathered reptiles.
The environment exerts a selective action on the population of
offspring: The best adapted survive in the greatest numbers to Tracing the line back further, we will see amphibia, bony fish,
breed and produce new offspring. Thus the traits that encourage cartilaginous fish, and invertebrates. The trail, if pursued long
survival in any environment are preserved. enough, leads back to one-celled life.
As adaptation to a given environment improves, and as But where did this one-celled life come from? Is a bacterium-and-
environments gradually change on the planet, the organisms spore paradox any less frustrating than the chicken-and-egg?
themselves change, adapt, and evolve.
Unless we suffer from mental fatigue or atrophied curiosity along
the way, we must eventually ask "Where did the earliest one-
celled life come from?" With such simple organisms, the problem
becomes as much chemical as biological.
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Introduction
The question of the origin of life was studiously ignored by the This chapter is concerned with the reawakening of the concept of
scientific community for three quarters of a century after Pasteur, spontaneous generation in a new, restricted, and scientifically
with two isolated exceptions: A. I. Oparin in Russia, and J. B. S. verifiable form.
Haldane in England.
We do not claim now that it happens all the time; Pasteur took
The very finality of Pasteur's experiments had made chemical care of that. What we do believe is that the spontaneous
inquiry into the development of life from nonliving chemicals not generation of life happened once on this planet, and that it then
really respectable. destroyed the conditions under which it could happen again.
The picture is "The Second Day of Creation" a 1925 woodcut by

the Dutch artist M.C. Escher
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The Common Biochemical Heritage of Life

We have no fossilized citric acid cycle or glycolytic enzymes for Other bacteria such as the Clostridia, which produce botulism in
study, and never shall have, so the starting point for foods and gangrene in wounds, do not respire at all. They obtain
understanding chemical evolution must be the reactions that occur all of their energy by anaerobic fermentation (glycolysis), giving off
in present-day organisms. as waste products lactate, acetate, ethanol, butyrate, propionate,
or other small organic molecules.
Eucaryotes all obtain energy by oxygen-using respiration; and
those that are photosynthetic obtain reducing power from water They all are compulsory or obligate anaerobes, for whom free
and release oxygen. This metabolic uniformity is missing in the oxygen gas is lethal. (This is why botulism only develops in sealed
older procaryotes. but imperfectly sterilized cans of food, and why aerating a wound
helps to prevent gangrene.)
Some bacteria do respire with O2, but others can use nitrate as an
oxidant if O2 is not available. Since the same enzymes are The ability to respire and oxidize foods is a special talent not
involved, and O2 always is their first preference, nitrate respiration possessed by all life, but glycolysis is universal. Glycolysis
probably is a relatively recent special adaptation that is of little accompanied by the storage of energy as ATP appears to be the,
interest in tracing the evolution of life. irreducible minimum for life.
Desuifovibrio sewage bacteria respire and extract energy from

their foods by using sulfate as an oxidizing agent, emitting the H2S Those organisms that go no farther than glycolysis cannot tolerate
that contributes to sewage stench. Different enzymes are used in the presence of gaseous O2. In contrast, with few exceptions,
the electron-transport chain of sulfate respiration, and this appears those bacteria that can live in the presence of O2, also have
to be a genuinely independent solution to the problem of getting learned to use it for respiration. It is too good a source of extra
more energy from foods by combining them with an oxidizing energy to neglect.
agent.
These facts suggest that life began as fermenting one-celled
Sulfate is not as good an oxidant as O2, but it is acceptable. organisms, at a time when no free oxygen was present in the
Sulfate-respiring bacteria are strict anaerobes, which are poisoned atmosphere.
by the mere presence of O2. They are restricted to life in rotting
sewage and other microenvironments that are reducing in
character. They may be living fossil remains of an era when the
planet had little or no free atmospheric oxygen.
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The eurcaryotic pattern of two-center photosynthesis yielding ATP and
NADPH, in which water is broken down and O2 is released, is followed also
by blue-green algae but not by bacteria.
Purple nonsulfur bacteria avoid the need for a reducing agent by running the
same electrons around again and again in cyclic photophosphorylation, but
then must depend on a citric acid cycle for production of reducing power in
the form of NADH (not NADPH).
They also possess a respiratory chain and, if grown aerobically in the dark,
can obtain energy from glycolysis, the citric acid cycle, and O2, respiration
just as we do. They are unique among bacteria in combining photosynthesis
with respiration, and appear to be a metabolic halfway house on the road to
blue-green algae and eucaryotes.
The purple and green sulfur bacteria are less versatile, and depend on
noncyclic photophosphorylation for production of both ATP and NADH. This
demands a source of reducing equivalents; lacking Photocenter II, they use
H2S or H2, both of which are intrinsically stronger reducing agents than H20.
These sulfur bacteria are compulsory anaerobes that are poisoned by an

oxygen atmosphere.
Once again the biochemical evidence suggests that early life arose under
conditions where free oxygen was absent, but where hydrogen and
hydrogen sulfide might be found.
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The chemoautotrophs are a special class of bacteria
that can synthesize carbohydrates like the
photosynthetic bacteria do, but which obtain the energy Chemoautotrophs and the inorganic materials from
for doing so from inorganic reactions rather than from
which they obtain their energy
the sun.
o Nitrifying bacteria Nitrosomonas, Nitrococcus - Ammonia

Some of them obtain energy by oxidizing ammonia to
nitrite or nitrate, others convert H2S to elemental sulfur,
thiosulfate, or sulfate, and still others oxidize Fe (II) to o Nitrifying bacteria Nitrobacter, Nitrococcus - Nitrite
Fe (III).
One might think that chemosynthesis, which uses o Thiobacillus - Hydrogen Sulfide, Sulfur, Sulfate
inorganic reactions for energy, is an older mechanism
than photosynthesis.
o Ferrobacillus, Gallionella - Ferrous Iron Salts

This is unlikely, since all chemoautotrophic bacteria
have well-developed respiratory chains and use O2 as
an oxidant. It is more likely that these chemoautotrophs
are specially adapted forms, which found an alternative
means to solar radiation to power their carbohydrate
syntheses. They can be neglected in a search for the
origin of life.
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Conditions for the Appearance of Life

The most important conclusion from a comparison of how bacteria Organic compounds themselves are unstable in an O2
and higher forms of life obtain energy is that all living creatures atmosphere, and are auto-oxidizable. Organic matter today is
have a common fermentative metabolism, which suggests a constantly produced anew by the action of living organisms. If all
common evolutionary origin. The various types of respiration and life were to end tomorrow, O2 would begin to reclaim the organic
photo- or chemosynthesis that have been added to glycolysis, do matter on our planet, and the process would stop only when no
not obscure this basic unity. more free oxygen remained.
Oxidation with O2, yields vastly more energy than fermentation It is inconceivable that large quantities of organic substances
alone, and had oxygen been present at the time when life evolved, could have remained unoxidized long enough for life to evolve
it surely would have been used. In that case, O2 respiration would from them, if they were constantly exposed to 02 in the
be as common to all life as fermentation is. atmosphere.
However, this is not the case, which leads us to a second If the original atmosphere was reducing, why is it oxidizing today?
conclusion: Life arose from less complex, nonliving chemical
systems at a time when the atmosphere was reducing in One source of O2, is the photodissociation of water vapor by
character, not oxidizing. ultraviolet light in the upper atmosphere, followed by the loss of
light hydrogen atoms from the Earth's gravitational field. This
Other evidence points to the same conclusion. alone could lead to an oxygen concentration of around 0.1% of the
present-day level.
The atmospheres of the other planets generally are reducing, as
we shall see later in the section on geological evidence. The main source of oxygen in the atmosphere today is green-plant
photosynthesis, and this probably is what turned the planetary
Old mineral beds on this planet suggest that they were laid down atmosphere from reducing to oxidizing.
in contact with a reducing atmosphere.
Life evolved under reducing conditions, where organic molecules
would be stable for long periods of time; but this same life was
responsible later for changing the original atmosphere to its
present-day composition.
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The Oparin-Haldane Theory of the Origin of Life
The first scientist after Pasteur to address himself seriously to The English biologist J. B. S. Haldane began thinking
questions about the origin of life was the Russian biologist A. I. independently along the same general lines, although he never
Oparin. read Oparin's writings.
He presented his ideas in a paper before the Botanical Society of In an eight-page article in the "Rationalist Annual" for 1929,
Moscow in 1922. They were published two years later, not in a Haldane published a complete synopsis of a theory of the origin of
scientific journal, but as a monograph. life
The paper sank into obscurity and had no effect on his The ideas of these two men were simple, elegant, and almost
contemporaries. It was not translated into English until 1967. Only identical.
when Oparin expanded this pioneering article into a full-length
book in 1936, and this book was translated from the Russian, did
his ideas begin to attract attention outside his homeland..
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According to their theory, life evolved in the oceans during a This Oparin-Haldane theory was a remarkably complete blueprint
period when the atmosphere was reducing - containing H2, H2O, for the ideas still held today. It was especially remarkable because
NH3, CH4, and CO2, but no free O2. in 1929 virtually none of the biochemical details of the previous
chapters were known.
Organic compounds were synthesized nonbiologically by
ultraviolet light energy, which in the absence of an ozone shield None of the chemistry of glycolysis, respiration, or photosynthesis
would penetrate the upper layers of the ocean. was understood, aside from the overall reactions.
Without free O2 to oxidize them, these organic molecules would Enzymes were a mystery, and were not even thought to be
be stable, and would accumulate in a warm, dilute broth that has proteins. The nature of the genetic machinery was unknown -
been nicknamed "Haldane soup." scientists were as likely to choose proteins as they were nucleic
acids for the carriers of genetic information.
The first living organism would be little more than a few chemical
reactions wrapped up in a film or membrane to keep them from The Oparin-Haldane theory was an accurate extrapolation far
being diluted and destroyed. These organelles would absorb beyond the limits of chemical knowledge of the time, which
chemicals, grow, divide, and obtain energy by fermenting the undoubtedly contributed to its general neglect.
available organic molecules around them.
It is to the credit of both men that much of what we have learned
Photosynthesis would arise eventually as an alternative energy since then has been a filling in of the blanks in their proposals.
source when natural foods ran short. The oxygen released by
photosynthesis would have the side effect of screening out the
ultraviolet radiation with an ozone layer in the upper atmosphere,
and eventually would turn the atmosphere from reducing to
oxidizing. Free oxygen would lead to the evolution of respiration
and to modern eucaryotic metabolism.
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What hard evidence do we have today for a theory of the origin of life?
The first area is comparative biochemistry along the lines we have been
following. The more we learn, the more the Oparin-Haldane ideas make sense.
In addition, we have geological evidence from the Earth's own history,

including evidence for a primitive reducing atmosphere and fossil remains of
primitive microorganisms. These fossils allow us to assign dates to the various
biochemical steps in the origin of life.
Finally, laboratory demonstrations, point out the feasibility of the nonbiological

synthesis of the molecules of life, and of the formation of simple, organized
chemical systems.
These considerations cannot prove the theories about what actually happened
billions of years ago, but they can give them plausibility.
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The Geological Evidence

What geological evidence is there to suggest that the Earth We can learn much from old sedimentary rocks about the
originally may have had a reducing atmosphere? conditions under which they were deposited.
The elemental composition of the universe, given in Chapter 8, Rocks that crystallized in the interior and then were thrust to the
supports this idea with an overwhelming predominance of surface have little to tell us about the atmosphere of the time. In
hydrogen. contrast, sedimentary rocks deposited by the weathering away of
older minerals during a long contact with the atmosphere preserve
The atmospheres of other planets, especially the larger ones a record of that atmosphere.
whose gravitational fields would have prevented loss of their early
atmospheres, are composed primarily of H2, He, CH, CO, CO2, If the atmosphere were oxidizing, then the sediments would be at
N2, NH3, and H20, with no free oxygen. least partially oxidized; if reducing, then the sediments would
remain reduced. Present-day sands are mainly quartz and other
The Earth as a whole is built mainly from metal silicates. These forms of SiO2. Most other minerals in the rocks that weathered to
silicates are 90 % oxygen by volume, but this oxygen is locked up make sand have been oxidized. Their oxidized metal cations have
in the mineral framework. been leached out, ultimately to be redeposited elsewhere as clay
minerals.
Iron in minerals can serve as a barometer of the state of oxidation
of its surroundings. The familiar red and orange of oxidized Fe(III) The result is that sedimentary rocks deposited during oxidizing
compounds from sands are only skin deep. conditions are of three main types: silicate sands, clay minerals,
and carbonate deposits of biological origin (from shells of marine
A short distance below the surface, these colors give way to the life). The sedimentary rocks laid down during the past 500 million
green and black of reduced Fe (II) compounds. The oxidized years and more are of this type. All indicate that oxidizing
minerals are a thin surface layer that is exposed to an O2- conditions existed during their original weathering period.
containing atmosphere that is anomalous among the planets.
Life has "rusted" the surface of our planet, but has had little effect
on its interior.
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The Geological Evidence

Weathering proceeds differently under a reducing atmosphere. These deposits are thought to have been laid down under
Quartz is still a major component of sedimentary material. The reducing conditions, although the evidence is not as conclusive.
other minerals that contain metals in lower oxidation states are These banded iron beds are 1.8 to 2.5 billion years old. In
less soluble and do not leach out completely. contrast, "red beds" of fully oxidized hematite ore (Fe203) are
never dated earlier than 1.4 billion years ago.
Among the reduced iron minerals in such sands one finds pyrite
(Fe(II)S), siderite (Fe(II)CO3), and magnetite (Fe3O4, or Fe(II)
O.Fe2(III)O3). Other metal oxides and sulfides in low oxidation The conclusion from these iron deposits is that the atmosphere
states are common. was predominately reducing prior to 1.8 billion years ago, has
been oxidizing for the past 1.4 billion years, and underwent a
Ancient Precambrian sediments containing sands with such period of gradual transition in the time between.
reduced minerals have been found in Canada, Brazil, and South
Africa. These deposits have been studied intensively because the This is not to imply that oxygen suddenly appeared in the
reduced materials often include elemental gold and uranium ore. atmosphere 1.8 billion years ago, or that water-using
photosynthesis was invented only then.
Geologists have concluded that these are the remains of ancient
sedimentary beds laid down under reducing atmospheric Photosynthesis on a small scale probably appeared nearly a
conditions. Radioisotopic methods have dated these various beds billion years earlier.
from 1.8 billion to 3.0 billion years old. The Earth is 4.5 billion
years old, so for the first 2.5 billion years of its existence, it had a It is difficult to calculate how fast O2 would accumulate in the
reducing atmosphere like the other planets. atmosphere, or how great the O2 concentration would have to be
before it could begin to influence the oxidation state of iron
Additional evidence comes from banded iron ore deposits with minerals in sediments. All we can say is that by the time this had
mixed oxidation states, found in Minnesota, Finland, Russia, begun, the O2 concentration must have been appreciable.
South Africa, India, and Australia.
So the stage was set for an Oparin-Haldane type of evolution of
life in a reducing atmosphere.
But did the actors really appear on cue?
For this, we must turn to the fossil evidence.
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Precambrian Fossils
Only a few years ago, the expression "Precambrian
fossils" would have been considered almost a
contradiction in terms.
At the beginning of the Cambrian era, 600 million years

ago, there was a veritable explosion in the fossil record.
Every major branch of modern animal life was present
at the start of the Cambrian era except vertebrates.
Geologists even define and identify the beginning of the
Cambrian era by this sudden increase in volume of the
fossil record.
Prior to 600 million years ago the fossil record quickly

shrinks to almost nothing. Jellyfish and other softbodied
marine life from 900 million to 600 million years ago are
known from deposits in Australia and a few other
places.
Much of the trouble is that these creatures are soft-

bodied. Jellyfish do not preserve well in the fossil record
in comparison with shellfish. Part of the Cambrian
explosion is not a sudden burst of life, but a sudden
increase in the use of hard, protective materials such as
shells and body armor.
No matter what the reason, most paleontologists only a

few years ago considered that little was to be learned
from the fossil record prior to this Cambrian population
explosion.
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Precambrian Fossils
A change came about when we learned how, and where, to look for fossil
microorganisms. Elso Barghoorn and his associates have studied polished thin
sections of silica-rich cherts from the Gunflint region of northern Minnesota and
southern Canada.
With the aid of optical and electron microscopes they have found a rich
collection of fossil bacteria, blue-green algae, fungi, and other microorganisms
with unknown present-day relatives. Two examples are shown on the previous
page.
Their association with banded iron ore formations means that these cherts
probably were laid down under reducing conditions, and radioisotopic methods
date them at 1.8 billion to 2.1 billion years old.
The Gunflint cherts also were found to contain pristane and phytane,
diagrammed on the right. These are organic compounds that can occur as
breakdown products of chlorophyll, and have been regarded as possible
evidence for photosynthesis.
Other microfossil deposits around 2.7 billion years old, from Australia,
Rhodesia, and South Africa, contain what appear to be fossil remains of
bacteria and blue-green algae.
The oldest sediments with true microfossils are the Fig Tree cherts from the
Transvaal, and the Onverwacht sediments from Swaziland, both in South
Africa. The Fig Tree cherts, which are 3.1 billion years old, contain fossil
bacteria that are spheroids resembling blue-green algae, filamentous organic
structures, and complex hydrocarbons including pristane and phytane.
The Onverwacht sediments are more than 3.2 billion years old, and are carbon-
rich cherts containing spheroids and filaments that possibly are of biological
origin.
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Precambrian Fossils
For unambiguous evidence of photosynthesis one must return to It is clear that organisms resembling bacteria and blue-green
no more than 1.6 billion years ago, to limestone deposits identical algae were in existence 3 billion years ago, and is probably true
to those produced today in hot springs by blue-green algae. that some of these organisms were photosynthetic and oxygen-
liberating.
These deposits, called stromatolites, are scattered widely over the
world. Some in Rhodesia are as much as 2.7 billion years old. The Well over a billion years may have been required for
1.6-billion-year-old stromatolites in the western Sahara are photosynthetic life to pour so much O2 into the atmosphere that its
unusual in that they contain alternating layers of CaCO3 and Fe character was changed. By 1.6 billion years ago, oxygen-emitting
(OH)3 as if they were laid down by colonies of photosynthetic photosynthesis and oxygen-using respiration were in full swing.
blue-green algae and O2-respiring iron-containing bacteria.
It is encouraging that the date for the Sahara stromatolites falls
The oxygen released by the algae would be used by the bacteria, right in the middle of the atmospheric transition period predicted
which would not then be dependent on significant amounts of from the oxidation states of iron deposits.
atmospheric oxygen.
What is remarkable is that the South African rocks from the
It is likely that such mutual aid, or symbiosis, was common in this Transvaal and Swaziland tell us that less than 1.5 billion years
era, with respirers living next to and using the oxygen from elapsed from the condesation of the Earth to the evolution of life at
photosynthesizers, just as bacteria live in mixed colonies in the bacterial level.
sewage and swamps today, with one species being dependent on
the waste products of another species for its food or raw As an indication of how difficult the next step - the development of
materials. eucaryotes - was, this second step required fully as much time as
the creation of the planet and evolution of bacteria.
It is not necessary to assume that oxygen respiration had to wait
for the complete conversion of the atmosphere to oxidizing
conditions before it could develop.
Page 17 of 36
Precambrian Fossils
Page 18 of 36
Precambrian Fossils
The first fossil evidence of cells with nuclei and internal structure like
eucaryotes comes from dolomite rock from Beck Springs, California.
These rocks are 1.4 billion to 1.2 billion years old (shown on previous page).
From this time on, the evidence is increasingly solid. The changeover of the
atmosphere to oxidizing conditions, the development of enough O2-respiring
procaryotes to show up plentifully in the fragmentary fossil record, and the
development of eucaryotic cells, all apparently took place 1.8 to 1.3 billion years
ago.
As an interesting sidelight to this chronology, one can compare the amino acid
sequences from a protein that is present in many forms of life to obtain a rough
measure of how distantly related these forms are, and how long ago their
ancestors diverged.
The sequences of respiratory cytochrome c from more than 67 eucaryotic

species have been compared, including vertebrates of all kinds, insects,
microorganisms, and higher plants.
Examination of the rates at which the cytochromes change in different lines of

descent suggests that plants and animals diverged approximately 1.2 billion
years ago, in excellent agreement with the fossil evidence for early eucaryotes.
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The Laboratory Evidence

One aspect of the Oparin-Haldane theory that has been neglected The results of a typical run beginning with H2, H2O, NH3, and CH4
so far is the very beginning of life. are recorded in the graph on the next page.
Is it reasonable that the organic compounds necessary as the Ammonia disappeared steadily during the experiment. During the
precursors of life would have been synthesized naturally first 25 hours of boiling and refluxing, most of the ammonia and
and abiotically in a reducing atmosphere? methane was being converted to HCN and aldehydes, with a slow
synthesis of amino acids.
Where would the energy have come from?
During the next 100 hours, HCN and aldehydes reached a steady
The first such simulation experiments were attempted by Harold state, being used in further reactions as rapidly as they were
Urey and his graduate student, Stanley Miller, in 1953. In 1952, made. The main products from these compounds were amino
Urey had published, in his book The Planets, a survey of the acids.
atmospheric chemistry of the planets, and had pointed out the
consistently reducing character of their atmospheres. They probably were the result of a Strecker synthesis , in which
ammonium cyanide reacts with aldehydes to make amino acid
Miller decided to see if biological molecules could be produced in nitriles, and these nitriles hydrolyze In water to amino acids.
a mixture of such reducing gases by a spark discharge, as an
analog of lightning. After 125 hours, as the supplies of ammonia and methane were
depleted, HCN and aldehyde concentrations began to decrease.
His experimental setup, shown on the next page, consisted of a The amino acid concentration leveled off as more of the simple
completely closed system, with gases flowing past a spark amino acids were incorporated into short peptides.
discharge; the condensed gases were recirculated by boiling. The
gases tried were mixtures of methane, ammonia, water, hydrogen,
and other reduced molecules.
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http://www.chem.ox.ac.uk/vrchemistry/chapter26/Page21.htm 2006/12/27
Page 21 of 36

Many similar experiments followed, by Miller and others, using both electrical
discharge and ultraviolet light. The compositions of the gas mixtures were
varied, and included H2S, CO, and CO2. Almost any starting mixture containing
compounds of both nitrogen and carbon led to amino acids, as long as free
oxygen was absent.
It seems that the spontaneous formation of amino acids by lightning and

ultraviolet radiation would have been virtually inevitable on Earth in a reducing
atmosphere, but impossible in an oxidizing atmosphere.
The first products in these experiments usually were hydrogen cyanide (HCN),
cyanogen (NC-CN), cyanoacetylene (H-C=C-C=N), formaldehyde (HCHO),
acetaldehyde (CH3CHO), and propionaldehyde (CH3CH2CHO). These
products then reacted to form various nitriles (R-CN), which subsequently
hydrolyzed in aqueous solution:
The results were mixtures of formic, acetic, propionic, lactic, succinic, and other
organic acids; glycine, alanine, aspartic and glutamic acids, and other
biological and nonbiological amino acids; urea, methylurea, and various other
small molecules.
None of these artificially synthesized molecules showed optical activity; they all
were equal mixtures of D and L isomers. As has been mentioned previously,
the optical activity that biological molecules exhibit today is a result of choices
by enzymes in living organisms.
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Even heat was effective in bringing about prebiotic
reactions. At elevated temperatures aqueous solutions
of formaldehyde (HCHO) and hydroxylamine (HO-NH2)
yielded amino acids and short polymers.
Solutions of HCN kept at 90°C for several days yielded

adenine, as shown at the left. Perhaps the fact that
adenine is simply a pentamer of HCN and is so easily
formed abiotically is the reason it is used in the energy
storage ATP molecules, in preference to guanine,
cytosine, thymine, or uracil.
Comparable experiments showed possible synthetic

pathways for purines, pyramidines, and carbohydrates.
Given ultraviolet radiation, electrical discharge, and
other energy sources on the primitive Earth, and a
reducing atmosphere, one might expect the inevitable
appearance of amino acids, purines, pyramidines,
ribose and deoxyribose, and even nucleosides and
nucleotides.
At least the building blocks of life would have been

available on the primitive Earth. A molecule of Adenine
The "Haldane soup" was real.
Page 23 of 36
The Problems of Organized Cells

So far we have glibly bypassed a massive problem. How do we Nevertheless, we do have this record to study from the protocells
get from the Haldane soup to even the simplest fermenting of the Fig Tree deposits to the first eucaryotes. For periods earlier
bacterium? than this, we have nothing at all.
A long, long step exists between amino acids, sugars, and We know how the planet began, and how this first phase in the
nucleosides, and simple cells of the Fig Tree type, and this is the evolution of life ended. The gap between must be filled by
step about which we know least. imagination tempered by the results of laboratory experiments.
From the short time span involved from the formation of the planet The chemical problems to be overcome are many.
to the development of these simple photocells, it can be argued
that the problem must be simpler than we think. How were polymers of proteins, nucleic acids, and lipids formed in
an aqueous environment, when polymer formation requires the
An equal time span occurred from protocells to eucaryotes, and removal of water and is thermodynamically nonspontaneous?
we think we have a good idea as to how this change came about.
The difficulty is that we have visible evidence for this latter How were the first reacting systems isolated from their
process, in microfossils and living survivors, but no such evidence surroundings to avoid a lethal dilution and cannibalism by other
has survived for the evolution of protocells. competing systems?
This is a continual problem in evolutionary history, the erasure of How were the chemical reactions of a protocell integrated into a
the older record. Survival on this planet is based on efficiency, and coherent and efficient "metabolism" that would increase its
there are no museums of unsuccessful types. chances for survival?
Even among the bacteria, we do not have samples of all of the And finally, having achieved all these things, how did the
ancestral chemistries, only those that enabled their possessors to successful protocell find a way of preserving its gains and passing
get along in odd corners where their more "advanced" eucaryotic them on?
competitors could not survive.
These are the next questions that we must try to answer.
We should not view the present-day bacteria as representative of
the ancestors of the main stream of development, but rather as
the "oddballs."
Page 24 of 36
Polymers and Microspheres

The first problem beyond the stage of Haldane soup is imagining how protein
polymers could form in dilute aqueous solution, when polymerization is a
dehydration, or water-removing, process. Equilibrium strongly favors cleavage,
not polymerization.
Sidney Fox has found that dry amino acids, heated to 160-210°C, will form
polymers of molecular weights up to 300,000, provided that aspartic and
glutamic acids are included in the mixture.
The sequences of these "thermal proteinoids" are not completely random, but
show some internal order. These polymers display a limited catalytic activity,
probably resulting from their charged side chains of acidic and basic amino
acids. They catalyze the decomposition of glucose reasonably well.
It is important not to read too much into this catalytic activity, since even
protons and platinum atoms are catalysts. It would be surprising if a polymer
with such a mixture of side chains was not catalytic for some reaction.
However, some such weakly catalytic polypeptides, with or without metal ions,
probably were the ancestors of the much more efficient and selective enzyme
catalysts of today.
These thermal proteinoids have another interesting property. If a hot proteinoid

mixture is washed with water or salt solution, microspheres of a fairly uniform
20,000-Å diameter are formed, as in the photograph to the right.
These are small globules of proteinoid polymer solution, enclosed by a

semipermeable proteinoid film with some of the physical properties of simple
cell membranes.
Microspheres shrink and swell in salt solutions of different concentrations. They
will grow at the expense of dissolved proteinoid material, and have been
observed to bud like yeast cells to produce "daughter" microspheres. They can
be induced to fission by MgCl2 or by a pH change. The enclosing film is a
double layer resembling those found in soap films and artificial and natural
membranes.
Page 25 of 36
Polymers and Microspheres

Fox hypothesizes that proteinoid material first polymerized on hot, Various mechanisms have been suggested by which amino acids
dry volcanic cinder cones, and then was leached into the oceans could be induced to polymerize spontaneously, even in an
by rain to form microspheres, which then could have become the aqueous environment.
early segregated chemical systems that eventually led to
protocells. These mechanisms usually involve making intermediate
molecules with high free energy, and using this free energy to
This cinder cone hypothesis, while ingenious, is not accepted by bring about the joining of amino acids during the polymerization
many scientists as more than an interesting idea. process.
What has been demonstrated is that membrane formation, Such mechanisms are analogous to the priming of molecules with
swelling, budding, and division all can occur by physical chemical phosphate groups or coenzymes, which we have seen in
forces, and are not necessarily tied in with living organisms. glycolysis and the citric acid cycle.
A weakness of the theory is the requirement of dry heat for While the problem of natural formation of protein chains by abiotic
polymerization. It is hard to imagine such high concentrations of means has not been solved completely, we tend to think of it as
dry amino acids occurring on the early planet. solvable.
It is difficult to make roast beef out of Haldane soup. No matter how an early proteinoid polymer might have been
formed, Fox's microsphere experiments, divorced from their
volcanic cinder cone hypothesis, remain relevant as a possible
way of producing isolated, enclosed regions of aqueous solution
for further evolution.
Page 26 of 36
Coacervate Drops and "Protobionts"
Oparin in Russia has had similar goals to Fox's, namely, to see If materials of smaller molecular weights are added to a solution of
how isolated and bounded regions of a solution could arise coacervate drops, they will distribute themselves unequally
naturally, as potential centers for the development of life. between drops and bulk solution, depending on their relative
solubilities in the two polymer phases.
If a concentrated solution of polypeptides, nucleic acids,
polysaccharides, or almost any polymer is gently shaken, it will Coacervates tend to concentrate some molecules in their interior,
separate into two phases of different polymer concentrations. an ability that the most rudimentary of protocells would need.
Concentrated droplets will form in a more dilute solution.
This behavior of coacervates shows how early protocells could

These coacervate drops are typically 20 gm (200,000 A) in have achieved internal compositions that were different from their
diameter, and may contain 5% to 50% polymer, depending on surroundings, and could have developed a certain amount of
how they are formed. They have a skin or membrane around chemical independence.
them, which is visible in a microscope.
Coacervate drops, like microspheres, are the result of physico-

chemical forces and have no direct connection with life.
Page 27 of 36

Even more interesting are coacervates prepared with enzymes inside. These
can absorb substrate molecules from solution, catalyze chemical reactions, and
let the products diffuse out.
If coacervates containing the enzyme phosphorylase are prepared, and

glucose-1-phosphate is added to the bulk solution, the primed glucose
molecules will diffuse into the coacervate droplets and be polymerized there
into a starch polymer.
If the coacervates also contain the enzyme amylase, then the starch produced
by the first enzyme is chopped back to disaccharide molecules of maltose,
which diffuse into the bulk solution again. Coacervates with these two enzymes
are miniature factories for turning glucose-1-phosphate into maltose, using the
energy of the phosphate bond in the starting molecules.
In another experiment, coacervates were prepared that contained

mitochondrial NADH dehydrogenase, the flavoprotein enzyme found at the
beginning of the respiratory chain. These droplets could absorb NADH and a
reducible dye from the solution, reduce the dye molecules, and release dye
and NAD+ back into solution.
In the most spectacular model experiment of all, coacervate drops containing

chlorophyll were allowed to absorb ascorbic acid and an oxidized dye that
could not be reduced spontaneously by ascorbate alone.
When the droplets were kept in the dark, nothing happened; but when they
were illuminated by light the dye became reduced. In a very close parallel to
the single-center photosynthesis of bacteria, chlorophyll molecules absorbed
light energy, and used their excited electrons to reduce the dye molecules.
Ascorbate merely played the role of H2S in the electron-transfer chain by

providing the electrons required to restore the electron deficit in the chlorophyll
molecules. Reduction occurred with a net increase in free energy, the
increased energy coming from the absorbed light.
Page 28 of 36

All of these, of course, are models and nothing more. They show This would have been the era of chemical evolution, where the
what could have happened, and what is not impossible. criterion for success would be the ability to find or synthesize the
chemicals necessary for the continuance of the droplet, and the
Oparin has suggested an evolutionary scheme for protocells or ability to prevent its own materials from being cannibalized for use
"protobionts" along the lines suggested by his coacervate in a neighboring system.
experiments. He proposes that in lakes or ponds with appreciable
concentrations of polymerized material, coacervate droplets would The development of an efficient outer membrane that could exert
be formed naturally by wave action, as diagrammed on the control on what came into and left the protobiont would be a
preceding page for a lipidlike material.In general, the composition strong aid to survival, as would an active-transport system that
of these droplets would differ from that of the bulk solution. could concentrate certain substances inside the membrane.
These "microenvironments" in time could develop into enclosed The ability to carry out reactions quickly, and to grow to such a
systems of chemical reactions that absorb high-energy size that the protobiont droplet would fall apart into many
compounds from their surroundings (like the glucose-1-phosphate independent daughter droplets, also would be advantageous to
experiment) to perform protective reactions or other necessary the survival of one particular kind of protobiont. Enzymes or their
syntheses. simpler catalytic precursors therefore would confer a great
survival advantage on a droplet.
The absorption of light for synthetic purposes, as in the
chlorophyllascorbate experiment, might have occurred first at this The second stage in the development of a living cell would be
prebiological stage. In this limited sense, "photosynthesis" might characterized by the ability to transfer to daughter fragments
have preceded life. during division, not samples of all catalytic substances, but the
instructions for making more of these pre-enzymes from simpler
The experiments of Fox and Oparin with microspheres and molecules.
coacervates suggest a model for how living organisms might have
developed. This would mark the beginning of hereditary information storage,
and of evolution by genetic variation and natural selection. It is a
The first stage along the road to life would have been stable, self- convenient point at which to draw a boundary between prelife and
maintaining, enclosed chemical systems such as these - perhaps life. This original information-storage system would have been far
growing and propagating by simple fission or division into smaller simpler than today's DNARNA-protein machinery, but all traces of
droplets that had the same chemical abilities and growth potential. it would have been erased as later improvements took over.
Control of reactions in these protobionts would be effected by

weak natural catalysts, also made by the protobiont and passed
along to each daughter fragment during fission.
Page 29 of 36
The Drama of Life

We can bring together everything that has been said in this chapter about the
evolution of life, and some of the earlier remarks about the formation of the
planet, into a fourteen-point scenario, a script for one element of the cosmic
drama as seen from our planet.
Although the scenario format enables us to describe events in simple

declarative sentences without a constant repetition of "probably" and "most
likely," bear in mind that this is at best a theory, a consistent set of hypotheses
to account for what happened. Some of the statements are on very firm ground,
but others are outlines for future research, to which chemists will make a major
contribution.
1. The universe began roughly 20 billion years ago, and our galaxy, the Milky
Way, approximately 13 billion to 15 billion years ago. Our sun is a second-
generation star in this galaxy, formed from the heavy-element-rich debris from
earlier stars. As the new star formed by gradual accretion of matter from a dust
cloud, local nodes of material in the plane of rotation of the flattened dust cloud
began to build up into protoplanets, moving around the sun. One of these
aggregates became our Earth.
2. The young Earth was too small to retain whatever original atmosphere it may
have had, and what remained was an airless ball of rock, made up mainly of
elements, such as silicon, oxygen, and metals, that could form nonvolatile
compounds. Heat from compression and from natural radioactive decay caused
the interior of the planet to become fluid, leading to the stratification by density
that exists today: iron-nickel core, olivine mantle, and a crust of lighter silicates
and other minerals.
Page 30 of 36
The Drama of Life

3. Volcanic outgassing from the interior created a new atmosphere for the
planet, made up of reduced compounds: H2, CH4, N2, NH3, H2O, and H2S. In
the absence of today's high-altitude ozone shield, ultraviolet radiation from the
sun penetrated all the way to the surface of the Earth. This radiation, lightning
discharges, volcanic heat, and natural radioactivity all provided energy sources
for the spontaneous reaction of the atmospheric gases to form more complex
molecules: HCN and aldehydes, nitriles, organic acids and bases, simple
carbohydrates and amino acids. These were leached into the oceans by rain,
where they slowly built up a thin "Haldane soup", which was stable for long
periods of time in a reducing atmosphere.
4. Life evolved from this soup, perhaps through intermediate stages of localized
but nonreproducing chemical systems, protected by simple barrier membranes.
Catalytic proteins, or crude enzymes, developed from random polymers of
amino acids, sometimes in association with metal ions and organic molecules.
Energy for chemical syntheses was provided by the breakdown of
polyphosphates, or later by molecules such as ATP, both formed originally by
nonbiological means. As competition depleted the natural supply of many
necessary substances in their surroundings, the more successful "protobionts"
developed the ability to synthesize these substances from more plentiful
molecules. Those primitive chemical systems that also developed a machinery
for duplicating all of this chemistry in daughter systems, crossed the threshold
of what we would define as life. The primitive information-transfer system need
have borne little resemblance to the elaborate DNA-RNA-ribosome system of
today, but the function would have been the same.
5. As the natural supply of polyphosphates and ATP ran short, some protocells
evolved glycolysis as a means of degrading organic molecules and saving the
energy as homemade ATP. This pattern of metabolism became so
advantageous that only those organisms that possessed it survived to the
present. Glycolysis and gluconeogenesis developed, with the necessary
enzymes floating freely in the cell fluid. The stage of the fermenting bacteria
was reached. Even with this metabolic capability, the amount of life on the
planet was strictly limited by the available supply of nonbiologically produced
organic molecules for use as energy sources.
Page 31 of 36
The Drama of Life

6. In the face of constant competition for the limited amounts of organic matter,
certain bacteria (if we may now call them that) found ways to enhance their
survival by using metalloporphyrins and similar delocalized ring molecules to
absorb solar energy. Perhaps at first the absorbed energy was used only as
heat to accelerate all reactions uniformly. Later, this electronic excitation of
chlorophyll molecules was coupled to the production of ATP and NADH. Two of
the best reducing agents that were available, H2 and H2S, were used to supply
reducing equivalents for making NADH. Carbohydrates were synthesized from
this ATP, NADH, and atmospheric CO2 by taking some of the reactions of
gluconeogenesis and turning them into the Calvin cycle. The stage of the
present-day green and purple sulfur bacteria had been reached.
7. Sulfate, although not a substance that would have been present in quantity
on the primitive Earth, was given off as a waste product from bacterial
photosynthesis. The ancestors of Desulfovibrio developed the ability to squeeze
a little more energy out of their foods by oxidizing them with this sulfate.
Colonies of green sulfur bacteria and sulfate-respiring bacteria could have
existed in close symbiotic association, as they sometimes do today, passing
oxidized and reduced sulfur compounds back and forth and drawing their
common support from the sun. Respiration had been invented, although not the
kind that was to dominate the planet in later years.
8. The slow development of a citric acid cycle as an alternate source of NADH

gradually liberated the purple sulfur bacteria from their dependence on H2S and
noncyclic photosynthesis. The ancestors of the purple nonsulfur bacteria arose,
which were dependent mainly on cyclic photophosphorylation for ATP energy.
Page 32 of 36
The Drama of Life

9. Close relatives of these photosynthetic bacteria found a way, via a second
chlorophyll photocenter, to absorb two photons of light where one had been
absorbed before, and to use the extra energy to make an acceptable reducing
agent out of H2O. Instead of abandoning a scarce reducing agent, H2S, they
managed to trade it for a much more plentiful one, H2O. In these ancestors of
the blue-green algae, green-plant photosynthesis was born. This step may have
been reached as early as 3 billion years ago.
10. Oxygen began to accumulate locally around these photosynthetic

organisms. They and the purple nonsulfur bacteria learned to use O2 with
NADH from their citric acid cycle to obtain much more energy than ever before.
The sequence of glycolysis-citric acid cycle-respiration, familiar in eucaryotes
today, was complete. As today, blue-green algae and purple nonsulfur bacteria
made relatively little use of respiration, depending mainly on photosynthesis for
ATP energy, but the facility was there. Oxygen respiratior need not have
required more than local concentrations of free O2, just as the earlier sulfate
respiration would have required only local concentrations of sulfate around
green and purple sulfur bacteria.
11. The great efficiency of water-splitting photosynthesis led to an explosion of

life on the planet, and this may be why we see fossil remains for the first time in
the Fig Tree cherts. With oxygen respiration still of minor importance, excess
O2 gradually accumulated in the atmosphere, changing it slowly from reducing
to oxidizing. This development had three important consequences for the future
evolution of life. An ozone shield in the upper atmosphere blocked off the
shorter ultraviolet wavelengths, thereby ending one source of nonbiological
synthesis of organic molecules as possible foods for living organisms. Free
oxygen in the atmosphere hastened the destruction of those organic molecules
that already had been synthesized, with the result that for all time to come,
organic compounds would be associated almost entirely with living organisms.
Lastly, with the lethal ultraviolet radiation screened out, life could come up from
the lower depths to inhabit the upper ten meters of the seas and, eventually, the
land itself.
Page 33 of 36
The Drama of Life

12. With the increasing oxygen content of the atmosphere, respiration became
more important. Oxygen-respiring bacteria evolved from purple nonsulfur
photosynthetic bacteria by the loss of photosynthetic ability. This explanation of
the origin of respiration would account for the remarkable similarity of the
electron-transport chains of photosynthesis and O2. respiration, and their great
difference from the processes involved in sulfate respiration in Desulfovibrio. It
also would explain the near identity in molecular structure of cytochrome c in
respiring eucaryotes and in respiring and photosynthetic bacteria.
13. Eucaryotes developed from procaryotes by a symbiotic relationship

between a nonrespiring host, respiring bacteria that were the ancestors of
mitochondria, and photosynthesizing blue-green algae that degenerated with
time into chloroplasts. This step probably was complete about 1.6 billion years
ago, judging from the Beck Springs fossil deposits.
14. In the interval between the development of the first eucaryotes and the
beginning of the Cambrian era, plants and animals diverged, soft-bodied
multicelled organisms developed, and most of the evolutionary lines arose that
later would lead to the major classes of living organisms. We move solidly from
chemical evolution and prehistory into the known fossil record.
This is the picture of life on Earth that we have been able to develop so far.
Whether life on other planets would have the same chemistry is a question we
cannot answer. We would assume it to be carbonbased and water-mediated,
but whether nucleic acids are inevitable as genetic records, and proteins as
structural materials and catalysts, is more than we can predict. The real
understanding of the limits of chemical systems and their organization into living
creatures has yet to begin.
Page 34 of 36
Questions
1. In what sense has spontaneous creation been discredited as an 9. What geological evidence is there for a reducing atmosphere on
explanation for the origin of life on Earth, and in what sense is it the early Earth? Approximately when did the transition to oxidizing
still the most acceptable theory? conditions occur, judging from the geological record?
2. If spontaneous creation is rejected as the origin of life, what 10. What are chemoautotrophs, and why are they unlikely to be
other explanation could there be? examples of a very primitive and ancient metabolism?
3. Why has the evolution of life at one moment on this planet 11. How does the presence of atmospheric O2 interfere with the
made it unlikely that life ever will evolve independently on Earth spontaneous synthesis of organic molecules by the nonbiological
again? What was wrong with the medieval picture of spontaneous process that must have occurred during the evolution of life?
generation?
12. How does the presence of atmospheric O2, make it
4. What portion of the energy-extracting machinery of eucaryotes improbable that organic molecules would evolve into living
is a common heritage of all forms of life? Give an example of organisms a second time, if all life on Earth were to be quickly
bacteria that depends solely on this type of energy production. extinguished?
5. What types of respiration are encountered in bacteria, in 13. What is the main source of organic compounds on the Earth
addition to respiration with O2? Which type probably is related to today?
O2 respiration, and which is quite different in evolutionary history?
14. How can organic compounds exist in an atmosphere of
6. There is no a priori reason why nearly all bacteria that do not oxygen, if oxidation of all of these compounds is
make use of molecular O2 as an oxidant should be poisoned by thermodynamically spontaneous?
the presence of O2, but this state of affairs is at least
understandable in terms of how bacterial metabolism evolved. 15. What is "Haldane soup," and how is it relevant to the problem
Why is it reasonable that the nonuse of O2 and the intolerance of of the origin of life?
its presence should go together?
16. How is it now believed that the organic molecules of "Haldane
7. How do bacterial and green-plant photosynthesis differ? Which soup" were formed?
more closely resembles the photosynthesis of blue-green algae?
8. What suggestions are there from bacterial metabolism that life

evolved in an O2-free, reducing environment?
Page 35 of 36
Questions
17. Why would the fossil record tend to underestimate the 25. How did Oparin's experiments with coacervate drops mimic
proportions of jellyfish to clams living at any one time? How is this some of the metabolic activities of living organisms?
relevant to the issue of a shortage of Precambrian fossils?
26. What advantage does water-decomposing photosynthesis
18. What are pristane and phytane, and why is their presence have for the organism that possesses it, in comparison with H2S-
considered as evidence for photosynthesis? using photosynthesis, if water is such a poor reducing agent
compared with H2S?
19. What is the oldest fossil evidence for bacterialike organisms?
Where are these fossils found, and how old are they? 27. What radical change in the environment of nonphotosynthetic
organisms was brought about by water-decomposing
20. What is the oldest fossil evidence for eucaryotes? Where were photosynthetic organisms? How did this lead in time to a much
they found, and how old is the deposit? more efficient means of extracting free energy from organic
molecules?
21. What are stromatolites? What organisms are believed to build
them? How does the presence of stromatolites constitute
evidence of a tentative sort for photosynthesis? How old are the
oldest stromatolite formations?
22. What arguments led Stanley Miller to choose hydrogen,

methane, ammonia, and water as components of his trial
"primitive atmosphere"? What energy source was used to bring
about chemical change? What natural phenomenon would this
correspond to? What compounds were formed in the course of the
reaction?
23. What is the Strecker synthesis, and how does it lead to the
formation of amino acids? What determines the nature of the
amino acid side chain, R-?
24. What are microspheres and coacervate drops, and what
relevance do they have to the problem of the evolution of life?

Why is a relative isolation from the surroundings advantageous to
any living organism?
Page 36 of 36

The Origin of Life On Earth

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Foundations to Chemistry - adapted from "Chemistry, Matter and the Universe" Page 1 of 2

26. Origin of Life on Earth -- Jump to --

26. Origin of Life on Earth -- Jump to --

It is difficult to put forward so thoroughly eroded an idea as spontaneous

This reluctance to abandon spontaneous generation was not an example of the

26. Origin of Life on Earth -- Jump to --

The operations of life and the mechanisms of life hence were

26. Origin of Life on Earth -- Jump to --

26. Origin of Life on Earth -- Jump to --

26. Origin of Life on Earth -- Jump to --

The picture is "The Second Day of Creation" a 1925 woodcut by

26. Origin of Life on Earth -- Jump to --

The Common Biochemical Heritage of Life

Desuifovibrio sewage bacteria respire and extract energy from

26. Origin of Life on Earth -- Jump to --

The Common Biochemical Heritage of Life

These sulfur bacteria are compulsory anaerobes that are poisoned by an

26. Origin of Life on Earth -- Jump to --

The Common Biochemical Heritage of Life

o Nitrifying bacteria Nitrosomonas, Nitrococcus - Ammonia

o Ferrobacillus, Gallionella - Ferrous Iron Salts

26. Origin of Life on Earth -- Jump to --

Conditions for the Appearance of Life

26. Origin of Life on Earth -- Jump to --

The Oparin-Haldane Theory of the Origin of Life

26. Origin of Life on Earth -- Jump to --

The Oparin-Haldane Theory of the Origin of Life

26. Origin of Life on Earth -- Jump to --

The Oparin-Haldane Theory of the Origin of Life

In addition, we have geological evidence from the Earth's own history,

Finally, laboratory demonstrations, point out the feasibility of the nonbiological

26. Origin of Life on Earth -- Jump to --

The Geological Evidence

26. Origin of Life on Earth -- Jump to --

The Geological Evidence

But did the actors really appear on cue?

For this, we must turn to the fossil evidence.

26. Origin of Life on Earth -- Jump to --

At the beginning of the Cambrian era, 600 million years

Prior to 600 million years ago the fossil record quickly

Much of the trouble is that these creatures are soft-

No matter what the reason, most paleontologists only a

26. Origin of Life on Earth -- Jump to --

26. Origin of Life on Earth -- Jump to --

26. Origin of Life on Earth -- Jump to --

26. Origin of Life on Earth -- Jump to --

The sequences of respiratory cytochrome c from more than 67 eucaryotic

Examination of the rates at which the cytochromes change in different lines of

26. Origin of Life on Earth -- Jump to --

The Laboratory Evidence

26. Origin of Life on Earth -- Jump to --

The Laboratory Evidence

26. Origin of Life on Earth -- Jump to --

The Laboratory Evidence

It seems that the spontaneous formation of amino acids by lightning and

26. Origin of Life on Earth -- Jump to --

The Laboratory Evidence

Solutions of HCN kept at 90°C for several days yielded

Comparable experiments showed possible synthetic

At least the building blocks of life would have been

26. Origin of Life on Earth -- Jump to --

The Problems of Organized Cells