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Re: Life on other planets, a real possibility ? on Thu Jul 15, 2010 4:13 am
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In the distant past, some early eucaryans (perhaps already endowed
with a nucleus, but still very small and lacking other organelles) may have
routinely ingested other prokaryotic cells for food. This in itself was a major
advance over the prokaryotes, for it necessitated the evolution of an outer
cell wall that could devour, or phagocytize, other cell material—in other
words, it made predation possible. Some of these ingested prokaryotic cells,
however, were not rapidly digested and hence destroyed within the host cell.
Instead, they may have lived there for some time. (Alternatively, these organelles
may have invaded the host cell, rather than being captured by it;
perhaps they bored in and established parasitic colonies within the larger eukaryotic
cell environments.) Eventually, the host cell came to benefit from
this association in some way: Prokaryotes, being very efficient chemical factories,
may have performed services the host could not carry out for itself,
such as energy transformation or even energy acquisition, and metabolic
functions. The organelles known as mitochondria (which are involved in energy
formation and transformation), plastids (the sites of chlorophyll), and
perhaps even flagella (which are used for locomotion) may have evolved in
this fashion.
The prime evidence for this hypothesis comes from DNA. Mitochondria
and plastids contain their own strands of DNA, which are closer in structure
to prokaryotic than to eukaryotic DNA. Mitochondria may have been
free-living bacteria that were capable of oxidizing simple carbohydrates into
CO2 and water and liberating energy in the process. There are living bacteria
today, such as forms known as purple nonsulfur bacteria, that may be close
to the ancestral mitochondrial form. When incorporated into the host cell,
these “guests” eventually lose their cell walls and become part of the host.
With the addition of the cell organelles, our eukaryote approaches or attains
a level of organization that would be familiar to us.
We can now sketch the evolutionary steps necessary to arrive at the eukaryotic
grade of organization. We start with a cell membrane enclosing
DNA—a simple bag of protoplasm and DNA—and then evolve the ability to
phagocytize (or engulf material), evolve a cytoskeleton (which allows us,
among other things, to get larger), evolve aerobic respiration, and then bring
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into our much larger bag various organelles: mitochondria, the nucleus, ribosomes,
and so forth.
This last is among the most interesting and controversial aspects of the
evolution of the eukaryotic cell type. Few scenarios have been proposed that
make evolutionary and adaptive sense, but one intriguing possibility has been
described by Dr. Joseph Kirschvink of Cal Tech, who has summarized the
problems faced by the evolving eukaryote as follows:
The problems for the eukaryotic host cell are that:
The host must be large enough to engulf other bacteria.
The host cell must be capable of phagocytosis, so that the
invaders are put into a membrane-bound vacuole (a small
space within the cell), leading to the characteristic double
membrane of the mitochondria and chloroplasts.
The cell should have at least a rudimentary cytoskeleton.
The host cell should offer a better, more controlled environment
for the symbionts, so that natural selection would
favor the association.
The only known bacterium that meets all of these constraints
is called Magnetobacter, discovered in Germany, which
dwarfs most other protists (in size). Each cell of this bacterium
makes several thousand organelles called magnetosomes, which
are tiny crystals of the mineral magnetite (Fe3O4) encased in a
membrane bubble—a bubble that forms by phagocytosis. These
magnetosomes are held in place in chain-like structures that keep
each crystal aligned properly; this can only be done if an intracellular
mechanical support structure such as the cytoskeleton exists.
Magnetobacter has the ability to keep itself in the optimal environment
by swimming along the magnetic field lines generated by the
Earth’s magnetic field. This ability makes it an attractive partner
for symbiosis, as many organisms spend a great deal of their metabolic
energy staying in the correct environment.
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This scenario for the evolution of the eukaryotic cell has two
major implications for the timing of higher life on Earth. First, the
most probable path for the evolution of magnetotaxis (the ability
to align along magnetic fields) and magnetite biomineralization
(the formation of minerals by living organisms) is a result of natural
selection for iron storage. Anaerobic microbes do not need iron
storage mechanisms, as ferrous iron is freely available in solution.
But in oxygen-rich environments the iron rusts out the [ferrous]
form, which drops out of solution. Hence, magnetotaxis is unlikely
to evolve in an anaerobic world, which on Earth ended
about 2.5 to 2 billion years ago. The oldest magnetofossil—the
fossil remains of bacterial magnetosomes—date to about 2 billion
years ago. Second, magnetotaxis requires the presence of a strong
planetary magnetic field. On Earth, a strong early field probably
decayed after 3.5 billion years ago, only to reach its present level
after nucleation of the inner core about 2.8 billion years ago.
Kirschvink has thus postulated a novel scenario—and perhaps the most
plausible scenario—for the formation of the eukaryotic cell: a pathway necessitating
the presence of magnetite and a strong planetary magnetic field.
As we shall see in a later chapter, not all planets maintain magnetic fields. If
this pathway is the only way to large eukaryotic cells (a hypothesis that still
awaits verification), then we have another requirement we must impose on
planets that aspire to host animal life—a magnetic field.
E N V I R O N M E N T A L CO N D I T I O N S L E A D I N G
TO T H E E V O L U T I O N O F E U K A R Y O T E S
What environmental conditions led to the evolution of the forerunners of animal
life? New discoveries of the 1980s and 1990s have given us a much
clearer view of the early Earth during the great evolutionary transitions we
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saw in the last chapter. The Earth’s earliest life seems to have formed during
or soon after cessation of the heavy comet bombardment. By about 3.8 billion
years ago that heavy cosmic bombardment ended, and by 3.5 billion
years ago we find the first fossilized evidence of life.
The region that has yielded Earth’s oldest fossils found to date is known
to Australians as the North Pole, because even in this isolated continent, it is
uniquely remote and inhospitable. The rocks in this region belong to a unit of
interbedded sedimentary and volcanic rocks known as the Warrawoona Series.
Geologists have deduced that the deposits were consolidated in a shallow
sea over 3.5 billion years ago. There is the evidence of storm layers and evidence
as well that on occasion, a hot sun evaporated small pools of seawater
into brine deposits. But it is not these structures that have created so much excitement
about the Warrawoona rocks. This ancient bit of Australia holds the
world’s oldest stromatolites, low mounds of lime and laminated sediment that
have been interpreted as the remains of microbial mats—in other words, life.
Stromatolites (the “stone mattresses” we mentioned earlier as an anomaly,
multicellular prokaryotes) are the most conspicuous fossils and the most
commonly preserved evidence of life for more than 3 billion years of Earth
history; they provide our best record of early life. They have been found on
every continent in rocks half a billion years old and older. Today, they are
found in only one type of environment on Earth, in quiet, briny tropical waters.
Such environments are refuges from algal grazers; stromatolites can no
longer exist on most of our planet’s surface because they would quickly be
eaten. The photosynthesizing bacteria termed cyanobacteria are modern
equivalents of these ancient deposits.
The presence of stromatolites is a sure clue that by 3.5 billion years ago,
life on this planet had left its earliest, probably hydrothermal or deep-earth environments
and diversified onto the surface of the planet. For a billion years the
prokaryotes were masters of the world, but life was still scattered. According to
the fossil record, it was not until about 2.5 billion years ago that the organisms
that produced stromatolites had released sufficient quantities of oxygen to form
sedimentary deposits known as banded-iron formations. Prior to the appearance
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of common stromatolites, there was no dissolved oxygen in the sea, no gaseous
oxygen in the atmosphere, and hence no possibility of mineral oxidation. With
the appearance of oxygen, however, large volumes of iron that had been dissolved
in seawater precipitated out as it oxidized into iron oxides—rust, in
other words. Today, there still exist at least 600 trillion tons of such iron oxides
deposited before 2.5 billion years ago in these banded-iron formations.
The time interval commencing about 2.5 billion years ago is marked by
a change in the very tectonic nature of the planet Earth—its rate of mountain
building and continental drift. By this time, the heat production from radioactive
elements locked in Earth’s rocks had diminished, for some of the
radioactive elements decayed rapidly early in Earth’s history. This material
was like a finite amount of fuel within the interior of the planet, and as it was
used up, heat flow declined. It turns out that the processes of continental drift
and mountain building are by-products of heat rising from within Earth, and
as the amount of heat decreased over time, so did these two activities. There
is also some evidence that around this time, a major pulse of land formation
occurred, allowing larger continental land masses to form. As the new continents
formed, many shallow-water habitats were created, and these proved
favorable environments for the growth of photosynthesizing bacteria. We
can speculate that from about 4 billion to about 2.5 billion years ago, there
were few large continents, but numerous volcanic island chains dotted the
world. After 2.5 billion years ago, continental land masses began to form, and
volcanism on a global scale lessened.
With this increase of habitat, ever more stromatolites grew and flourished.
This in turn relentlessly pumped ever more oxygen into the sea. As
long as there was dissolved iron in the seawater, all of the liberated oxygen
was quickly locked up in the banded-iron formations. By about 1.8 billion
years ago, however, this reservoir of dissolved iron material was used up. We
know this because after that time, no more banded-iron formations were laid
down. This changeover left an indelible mark on the sedimentary record of
Earth, for as the sea became saturated with oxygen, the time of banded-iron
formations ended forever—or at least until some far-distant future when our
planet may again no longer have oxygen. With nowhere else to go, oxygen
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began to emerge into our planet’s atmosphere and, in so doing, probably gave
life its first impetus toward animal life.
T H E OX Y G E N R E V O L U T I O N
It is probably impossible for us to conceive how entirely alien to ours this
world truly was. Yet the strange microbial world of 2 billion years ago may be
the norm in the Universe for those planets that harbor life. Traces of it exist
still, here on Earth, in the bacterial froths and pond scum that persist across
our planet, and perhaps nowhere more prolifically than in the rotting garbage
dumps and landfills created by our own species—places where huge, visible
colonies of rapidly growing bacteria exist still. But the rainbow slick of the
oozing swamp is the exception in a world where the eucaryans are so much
more in evidence than the prokaryotic forms. What would that 2-billionyear-
old world look like? The best description we know of was penned by
two scientists who have journeyed back to this world, in their imaginations,
many times. We owe the following image of the ancient Proterozoic era (the
formal name for the time interval of 2.5 to 0.5 billion years ago) to Lynn Margulis
and Dorion Sagan, in their 1986 book Microcosmos:
To a casual observer, the early Proterozoic world would have
looked largely flat and damp, an alien yet familiar landscape, with
volcanoes smoking in the background and shallow, brilliantly colored
pools abounding and mysterious greenish and brownish
patches of scum floating on the waters, stuck to the banks of rivers,
tainting the damp soils like fine molds. A ruddy sheen would coat
the stench-filled waters. Shrunk to microscopic perspective, a fantastic
landscape of bobbing purple, aquamarine, red, and yellow
spheres would come into view. Inside the violet spheres of Thiocapsa,
suspended yellow globules of sulfur would emit bubbles of
skunky gas. Colonies of ensheathed viscous organisms would
stretch to the horizon. One end stuck to rocks, the other ends of
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some bacteria would insinuate themselves inside tiny cracks and
begin to penetrate the rock itself. Long skinny filaments would
leave the pack of their brethren, gliding by slowly, searching for a
better place in the sun. Squiggling bacterial whips shaped like
corkscrews or fusilli pasta would dart by. Multicellular filaments
and tacky, textilelike crowds of bacterial cells would wave with the
currents, coating pebbles with brilliant shades of red, pink, yellow
and green. Showers of spheres, blown by breezes, would splash
and crash against the vast frontier of low-lying mud and waters.
This prokaryotic world was creating what has been called the Oxygen
Revolution. The initiation of an oxygenated atmosphere was one of the most
significant of all biologically mediated events on Earth. Prokaryotic bacteria,
using only sunlight, water, and carbon dioxide, ultimately transformed the
planet by generating an ever-increasing volume of atmospheric oxygen. This
outpouring of oxygen created both biotic opportunity and biotic crises. Many
of Earth’s primitive organisms were metabolically incapable of dealing with
abundant oxygen. For most of the archaeans, the oxygen boom of about 2 billion
years ago was an environmental disaster, driving some species into airless
habitats, such as lake and stagnant ocean bottoms, sediments, and dead organisms.
Others were incapable of such migration and simply died out. For yet
other creatures, however, the profound change in atmospheric conditions created
new opportunities. Some prokaryotic cells began to exploit the enormous
power of oxygen metabolism to break down food sources into carbon dioxide
and water. This new metabolic pathway yielded far more energy than any of
the anaerobic pathways. Organisms that adopted it soon began to take over
the world. The most efficient of these were members of the domain Eucarya,
which, more than 2 billion years ago, evolved true eukaryotic cell machinery.
The oldest known fossils of an organism that appears to have attained
the eukaryotic grade of organization have been found in banded-iron deposits
located in Michigan. The fossils themselves are about 1 millimeter in
diameter and are found in chains as much as 90 millimeters long. The organism,
then, is far too large to be a single-celled prokaryote or even a singleHow
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celled eukaryote. This creature, which has been named Grypania, is preserved
as coiled films of carbon on smooth sedimentary rock bedding planes, the
places where sedimentary beds split apart. Its 1992 discovery indicates that
the evolution of the first eukaryotic cell occurred during the banded-iron formation
process, when there was still little free oxygen in the sea and probably
none in the atmosphere. These early eukaryotes may have been vanishingly
rare, for other eukaryotes do not occur in the fossil record for 500
million years after this first appearance, but with this form, a beachhead in
life’s advance had been established.
For the period between 2 and 1 billion years ago (see Figure 5.2), few notable
achievements of life are recorded as fossils in the rocks. The first common
appearance of eukaryotes begins about 1.6 billion years ago, when microscopic
fossils called acritarchs begin to appear in the geological record. These are
spherical fossils with relatively thick, organic cell walls. They are interpreted to
be the remains of planktonic algae, forms that used photosynthesis and lived in
the shallow waters of the world’s oceans. Other life forms evolved as well, but
as is also true of most living protists, such as the amoeba and the paramecium,
their lack of skeletons renders them invisible in the fossil record. With a proliferation
of plant-like forms, new varieties of predatory protists surely evolved.
Whole armadas of single-celled, floating pastures and the somewhat larger
and more mobile grazers on these fields of plankton lived and died in this
seemingly endless epoch of geological time. The open ocean would have had
little life, but the coastal regions richer in nutrients would have been awash
with life—microscopic life. It was the Age of Protists, the Age of the Small.
We have now reached 1 billion years ago, in our march through evolutionary
time. Finally, the tempo of evolutionary development increased, if we
are correctly interpreting the fossil record, for there is a burgeoning in the
number of eukaryotic species found in the rock record at this time. Some of
these new forms include the first red and green algae, forms still crucial and
varied in marine ecosystems. This diversification of eukaryotic species, including
protozoans and plants, set the stage for the evolution of larger, multicellular
forms and may have been triggered by the evolution of important
new morphologies within the eukaryotic cell.
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E V O L U T I O N I N E U K A R Y O T I C
FORM AND F U N C T I O N
Four biological innovations may have been especially significant in paving
the way for the emergence of larger animals: (1) the development of sexual
cycles; (2) new methods of shuffling information coded along the chromosomes
(through the nascent ability to excise and relocate entire gene sequences);
(3) new methods of communicating between cells via substances
called protein kinases, and (4) the development of a new type of intracellular
skeleton, called a cytoskeleton, that allowed eukaryotic cells to increase
enormously in size. These innovations greatly enhanced the ability of cells to
Archaean Proterozoic Phanerozoic
Paleoproterozoic Mesoproterozoic Neoproterozoic
Grypania megafossils
Undoubted multicellular algae
Chuaria-Tawuia assemblage
Longfengshania
Worm-like megafossils
Ediacara-type fossils
Simple trace fossils
Cloudina
Skeletal fossils
Complex trace fossils
Trilobites
Chengjiang fauna
2.5 2.0 1.5 1.0 0.5 0 Ga
Varanger ice age
?
Figure 5.2 Early multicellular fossils. Broken bars indicate uncertain time ranges.
E A R L Y MU L T I C E L L U L A R F O S S I L S
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evolve new morphologies in response to natural selection and their ability to
band together into multicellular creatures.
We can now better categorize what we term “advanced” life: eukaryotic
multicellular organisms. There are, of course, many types of multicellular organisms,
including a considerable number of prokaryotic forms. In most cases
these multicellular prokaryotes are composed of only two cell types. Cellular
slime molds are multicellular, as are some cyanobacteria. In a way, however,
these forms are evolutionary dead ends. They have existed on Earth for several
billion years and are highly conservative in an evolutionary sense. It is
multicellular creatures of the other category that became so important in the
history of life. We refer here to true metazoans.
The jump from single-celled organisms to organisms of multiple cells
requires numerous evolutionary steps. The jump from single-celled organisms
to metazoan animals, where a high degree of intercellular cooperation in organization
exists, involves even more. In their recent book Cells, Embryos and
Evolution, biologists John Gerhart and Marc Kirschner discuss this evolutionary
accomplishment. The first step, they argue, seems almost paradoxical: It
was not some new structure gained that allowed this transition, but an important
structure lost. Long ago in our planet’s past, some organism of the eukaryotic
lineage made a brave (or lucky) morphological change—it shed its
external cell wall. Why this occurred is still unclear, but the net effect was
far-reaching. A tough outer coating protects most unicellular creatures from
their surrounding environment. At the same time, however, it isolates these
cells from other members of their own kind. By divesting themselves of this
outer wall, individual cells could begin exchanging living material—and
information—with one another. The naked cells could adhere to each other,
crawl over each other, and communicate. These were the first steps in the formation
of a tissue, which is an aggregation of cells united for mutual benefit.
Larger animals require highly integrated systems of cells that can accomplish
the myriad functions necessary for all life. Respiration, feeding,
reproduction, the elimination of waste material, information reception,
locomotion—all require the integration of many cells acting in concert. Each
of these functions ultimately requires one or many types of tissues.
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Among tissue types, the outer wall of any organism (the epithelium) is
of utmost importance. The epithelium must protect the organism from the
rigors of the external environment but at the same time allow adsorption of
critical gas and, sometimes, nutrients. The evolution of the epithelium was a
decisive first step in the evolution of metazoans.
Which group of unicellular creatures first achieved this breakthrough?
The most primitive and enigmatic of larger eukaryotic metazoans are
sponges. These curious creatures seem to bridge the gap between singlecelled
eukaryotes or even colonial protozoa and the highly integrated invertebrate
metazoan phyla. Sponges have several cell types that perform specialized
tasks, but there is a very low level of organism-wide organization.
There is no gut or body cavity specialized for processing food, nor is there
any nervous system. Yet the sponges may be an important clue to the identity
of our actual metazoan ancestors.
The stem, or ancestral, metazoan probably had a larger number of cell
types than sponges (perhaps 10 to 15 rather than the 3 to 5 individual cell
types found in sponges). There was probably a body cavity of some sort segregated
into two cell layers: an outer ectoderm and an inward-facing endoderm.
This two-tissue plan seems to have been an evolutionary dead end, and
it wasn’t until a third layer—the mesoderm—was added that animals with
real internal complexity formed. Eventually, a small worm-like shape with
three tissue layers evolved, a creature with a gut running through the long
axis of the body and a separate space known as a coelom to serve as an internal
hydrostatic skeleton. With this tiny organism (the first may have been
less than a millimeter long), the evolutionary stage was set for the emergence
of animals on planet Earth.
T H E TWO DI V E R S I F I C A T I O N S
O F A N I M A L P H Y L A
With the advent of this form—what evolutionary biologists call the “roundish
flatworm”—a body plan was in place that could be modified to shape all the
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categories of metazoan life, the major body plans that we call phyla. The
phyla living today include the arthropods; the mollusks; the echinoderms;
our own group, the chordates; and about 25 more. These are the complex
metazoans that we hope to find—but that may be very rare—on other planets.
These are animals. They appeared relatively late in the history of life on
Earth. One of the great novel insights of the 1990s was our realizing that
their origin and their subsequent diversification and rise to abundance were
two separate events, not one, as had been believed since the time of Charles
Darwin.
Fossils of macroscopic animals (those visible to the unaided eye) first
appear in abundance less than 600 million years ago, during the “Cambrian
Explosion,” a diversification event resulting in the rapid formation of thousands
of new species; we will describe it in more detail in the next chapter.
Yet the appearance of abundant animal fossils at this time actually marks the
second of the two diversification events that led to the proliferation of larger
animals on the planet. As we will show, fossils of such complex animals as
trilobites and mollusks—common members of the Cambrian Explosion—are
advanced descendants of a much earlier, diversification event that took place
between 1 billion and 600 million years ago. Yet there is no fossil record of
this first diversification—paleontologists have been stymied by an almost
compete lack of fossils in strata older than 600 million years, when this initial
event must have taken place. Our understanding of the initial diversification
of animals comes not from paleontology but from an entirely different line of
investigation: genetics. Geneticists have arrived at answers about the “when”
of the first diversification event by examining the genetic code of living animals
via a technique called ribosomal RNA analysis.
Gene sequences are simply strings of base pairs lined up along the double
helix of a DNA molecule. As we saw earlier, if a DNA molecule is likened
to a twisted ladder, the base pairs can be considered the steps of the ladder,
and it is the sequence of the steps that is used in this type of analysis. Genes
are simply instructions for protein formation coded by the sequence of nucleotides
on the DNA ladder. There are only four types of nucleotides, but
they provide the genetic code that is the basis for all Earth life. All organisms
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share more genes with their ancestors than with nonrelated species. By comparing
the genes from various organisms, it is possible to produce a model of
evolutionary history (an evolutionary tree, as it were) with the branches of
the tree showing which species gave rise to which other species. Yet according
to many geneticists, such an analysis not only tells us how the branching
occurred, it can also tell us when.
In 1996 G. Wray, J. Levinton, and L. Shapiro published a paper claiming,
on the basis of results obtained by using this genetic technique, that the
first event—the earliest divergence of animals—occurred 1.2 billion years
ago. This result drew a collective gasp from the paleontological fraternity: It
seemed much too ancient. The fundamental assumption of the Wray et al.
paper is that gene sequences evolve with sufficient regularity that a sort of
molecular “clock” can be used to date the divergence of various groups. The
reasoning behind the molecular clock technique is that changes in the genetic
code—evolution, in other words—occur at a rather constant rate. The
more distinct two DNA sequences are, the longer it has been since they diverged
from a common ancestor. Other scientists, however, dispute that
changes in gene frequency occur at a constant rate, and therefore they do not
believe in the molecular clock. It is these molecular clock data that led the
Wray group to their conclusion. This finding was a bombshell. If animals
evolved this early, why did they not appear in the fossil record until less than
600 million years ago? What were they doing for such a long time?
The Wray group’s findings were extremely controversial not only because
they contradicted long-held paleontological dogma but also because
they provoked criticism among other geneticists. There is fierce debate
among geneticists about the reliability of the molecular clock technique. The
Wray study itself, yielded both minimum and maximum figures for the earliest
divergence. One group of genes suggested that the fundamental splitting
of the phylum made up of annelids (worms) from the phylum of chordates
(our phylum) occurred only 773 million years ago, whereas a second group of
genes (in the same organisms) suggested 1621 million years ago—a very wide
spread indeed! These results give us minimum and maximum ages for the diHow
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vergence. Even with the minimum figure, however, there were (according to
the molecular data, anyway) recognizable chordates and annelids 700 million
years ago—yet there is no trace of their presence in the fossil record. Where
were they? Or were they not there at all? Could it be that no rocks of this age
survive or that no fossils from the interval of about 1 billion to less than 600
million years ago were preserved? This seems to be stretching things, as suggested
by British paleontologist Simon Conway Morris:
Appeals to gaps in the rock record and pervasive metamorphism
of the sediments are not going to work: if there were large metazoans
capable of either fossilization or leaving traces, they had an
uncanny knack of avoiding areas of high preservation potential.
Since the original, tantalizing analysis by the Wray group, other geneticists
have reconsidered the basic data. Most concede that the 1.2-billion-year
figure is too old. (However, a report published in Science magazine in late 1998
by a team headed by Adolf Seilacher of Yale University announced the discovery
of billion-year-old trace fossils (worm-tracks) possibly derived from small,
worm-like organisms. Critics of this finding suggest that the marks in question
could just as easily have been produced by inorganic actions, and even if these
trace fossils turn out to have been produced by organisms, the question remains:
Why are no further such fossils found for hundreds of millions of years?)
Let’s say, then, that divergence occurred less than a billion years ago. We must
still account for a significant period of time with animals but without fossils. Paleontologists
have long believed that only a single major diversification event
occurred—the event coincident with the appearance of fossils, the so-called
Cambrian Explosion that began about 550 million years ago. Now this evolutionary
event is seen as a follow-up to the much earlier first event.
The answer to this seeming conundrum is that the animals were indeed
present, but they were so small as to be essentially invisible in the fossil record.
A recent and spectacular discovery of microscopic fossil animal embryos
seems to confirm this view. Using newly developed techniques of searching
for tiny (but complex) animals in minerals called phosphates, paleontologist
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Andy Knoll and his colleagues have uncovered a suite of tiny but beautifully
preserved fossils interpreted to be the embryos of 570-million-year-old
triploblasts—animals with three body layers, like most of those found today.
These fossils tell us that the ancestors of the modern phyla were indeed present
at least 50 million years before we find any conventional fossil record of
them. The combination of genetic information and new discoveries from the
fossil record now give us a robust view of the rise of animals: They did not
exist 1 billion years ago, and perhaps not 750 million years ago. Animals are
indeed very late arrivals on the stage of life on Earth.
Thanks to these new discoveries and interpretations, the question of
“when” has been answered to most people’s satisfaction: The emergence of animals
was a two-stage event. The initial stage seems to have occurred less (and
perhaps much less) than the billion years ago proposed by Wray and his colleagues.
But even recalibrated, the Wray group’s finding has given us yet another
tantalizing insight into the potential incidence of animal life in the Universe.
The Wray work confirms that there were indeed two “explosions.” The
first was the actual differentiation of the various body plans; the second was the
differentiation and evolution, in these various phyla, of species large and abundant
enough to enter the fossil record. The geneticists can show that genes of
annelid worms and genes of chordates were differentiating hundreds of millions
of years before the emergence of these creatures as large entities that
could appear in the fossil record. This leads us to ask a crucial question: Even if
they evolve, do animals necessarily, or inherently, go on to diversify, enlarge, and survive?
Does the second flowering of animal life—the Cambrian Explosion event so
long known to geologists—inevitably follow the first diversification, or is it yet
another threshold of possibility that may be (but is not necessarily) attained?
Perhaps on some worlds in the Universe, animals diversify but never attain
larger size and greater numbers in some Cambrian Explosion equivalent. This
particular insight was first expressed by paleontologist Simon Conway Morris:
We need to discuss to what extent metazoan history was implicit a
billion years ago, at least in outline, as opposed to what was inevitable
500 million years later at the onset of the Cambrian exHow
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plosion. Even if metazoans have a deep history, which paleontologically
remains cryptic, the actual organisms would have been of
millimeter size and perhaps without the potential for macroscopic
size and complex ecology. . . . Wray et al. may have been correct
in tracing the gunpowder back as far into the mists of the Neoproterozoic
(the late Precambrian time period of a billion years
ago), but the keg itself still looks as if it blew up in the Cambrian.
In other words, it seems that the development of animals was a two-step process,
with step two—the Cambrian Explosion—not necessarily being an outcome
predetermined by the initial differentiation of the animal phyla.
Over and over the same question arises: Why did it take so long for animals
to emerge on planet Earth? Was it due to external environmental factors,
such as the lack of oxygen for so long in the history of this planet, or to
biological factors, such as the absence of key morphological or physiological
innovations?
T H E E V O L U T I O N O F A N I M A L S :
B I O L O G I C A L B R E A K T H R O U G H O R
E N V I R O N M E N T A L S T I M U L U S ?
Complex animals surely cannot appear on any planet without following some
evolutionary pathway from simpler, single-celled organisms. The change
from single-celled microbes to multicellular creatures must be the common
route on any planet, and even if the molecules of life are different from world
to world, the pathway from simple to complex may be universal. Because of
this, the example of how animals evolved on our Earth may be of the utmost
importance in understanding the frequency with which animals occur on
other planets.
If we are to understand how animals evolved from single-celled ancestors,
we must first understand the environments where these monumental
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Million
years Traditional view Wray et al. view Compromise view
300
400
500
600
700
800
900
1000
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1300
1400
Perm
Dev
Sil
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Cam
Carb
Mollusca
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Echinodermata
Arthropoda
Agnatha
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Mollusca
Annelida
Echinodermata
Arthropoda
Agnatha
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Mollusca
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Proterozoic
Riphean Vendian
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Million
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Figure 5.3 Differing views of metazoan phylogeny. Most paleontologists follow the “traditional view”
(left), accepting the fossil record as a fairly reliable indicator of original events. Molecular clocks are interpreted
by Wray et al. (center) as indicating very deep origins for the principal metazoan phyla. The recognition
that some molecular clocks run much faster than others suggests a “compromise view” (right), which implies
that our search strategy for the first metazoans should be concentrated in the interval from about 750
million years onward. Perm, Permian; Carb, Carboniferous; Dev, Devonian; Sil, Silurian; Ord, Ordovician;
Cam, Cambrian.
evolutionary advances were made. We know well the “when” of this change—
it took place during a 500-million-year interval from 1 billion to 550 million
years ago. The second event, the Cambrian Explosion of between 550 and 500
million years ago, included the morphological diversification of the phyla into
subdivisions based on body plans, as well as the appearance, within the various
phyla, of species with skeletons and large size (see Figure 5.3).
During this interval of time, Earth went though major environmental
changes, among them ice ages of unprecedented severity, rapid continental
movements, and drastic changes in ocean chemistry. We are thus left with
perplexing questions: Did the environmental changes of this interval (which
are described in more detail below) somehow trigger the diversification of anHow
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imals? Or would the rise of animals have occurred even in the absence of
these profound environmental changes? These questions, which of course are
central to understanding the evolution of life on our planet, have great relevance
to understanding the frequency of animal life on other planets as well.
Does animal life always (or even commonly) evolve, once a suitable ancestor
appears? Or does there need to be an additional trigger of some sort, a sequence
of environmental steps? We might compare this whole process to
baking a cake. Say the ingredients for the batter were all assembled and
mixed by 1 billion years ago. Does the cake need to be cooked for a given
time at a highly restricted temperature in order to rise? Or will any amount of
cooking at any temperature accomplish the task just as well? Or will our cake
be completed without any cooking at all? (That is, does simply assembling
the ingredients into a batter ensure success?)
The beginning of this fecund period in Earth history is marked by the
appearance not of new types of animals, but of plants. Around 1 billion years
ago, many types of algae begin to appear in the fossil record, including the
green and red algae still so prominent on Earth today. These were not the ancestors
of animals, of course, but their appearance was the opening salvo of
an evolutionary assault that was the most significant up to that time. It was
followed, hundreds of millions of years later, first by the initial diversification
of animal phyla and then (after more hundreds of millions of years) by the
Cambrian Explosion of animal life.
What were the environmental events of this interval of time 1 billion to
600 million years ago? By this period, land masses approaching the size of
today’s continents had formed, and the total area of land on the planet may
not have been significantly different from what we see in the present day.
The land, however, was not a tranquil place. The period was one of significant
mountain building and continental drift. It was also marked by episodes
of continental glaciation unmatched in severity since that time. Did these
events have anything to do with the diversification of animals? One school of
thought says yes. Work by Martin Brasier and others suggests that rapid
changes in sea level, and especially the formation of broad, shallow seas
within the new continents, would have opened up many new habitats very
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hospitable in terms of temperature and nutrients. This, in turn, may have
stimulated the diversification of animals and plants. There are dissenters, notably
James Valentine, who cautions, “the link between plate tectonics . . .
and the origins and radiation of animals remains to be demonstrated.” But, as
Harvard paleobiologist Andy Knoll points out, there is another way in which
the new and active tectonic events could have influenced the initial radiation
of animals that occurred during this time. In 1995 Knoll noted that “tectonic
processes could have influenced one or more of the great radiations (of animals)
. . . through their participation in the biogeochemical cycles that regulate
Earth’s surface environments.”
Examples of such effects include the role of hydrothermal influences on
ocean chemistry. The hydrothermal vents, as we saw in Chapter 1, are submarine
regions where great volumes of hot and chemically distinctive water
are mixed with seawater. The amount of this volcanically derived water entering
the oceans fluctuated during the interval of 1 billion to 550 million
years ago, and these fluctuations had marked effects on the chemistry of the
seawater, on the composition of the atmosphere, and on climate. The tectonics
events also affected the rate of burial and exhumation of organic carbon
in sediments. Oxygen and carbon dioxide values shifted, and as they did
so, major changes in the temperature and oxygenation of the planet ensued.
Yet another environmental stimulus may also have contributed to the
initial animal diversification. Changes in ocean chemistry caused by increased
tectonic activity beginning a billion years ago facilitated the evolution of
skeletons. This period is marked by the appearance of rocks called phosphorites.
Some authors credit these rocks with bringing about an increase in the
fertility of the oceans at this time, which may in turn have helped trigger the
sudden appearance of many diverse animals beginning about 600 million years
ago. Phosphorus is much more concentrated in living things than in the environment,
so it is a limiting nutrient. The sudden presence of abundant sources
of this element could have acted as a veritable fertilizer for growth.
Knoll has discussed all of these disparate factors and has proposed three
alternatives. First, it may be that the complex physical events and the equally
complex series of biological events that occurred from 1 billion to 550 milHow
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111
lion years ago are simply coincidental—they had nothing to do with one another.
If this is true, the great biological diversification must be attributed
solely to biological innovations (such as the ability of cells to bind together,
build an outer cell wall, and evolve internal cooperation between contained
cells) that were unrelated to concurrent environment changes.
The second alternative is that evolution was indeed facilitated by
changes in the physical environment. The most important of these changes
may have been in levels of oxygen. The first appearance of larger metazoans,
the ediacarans, about 600 million years ago occurred immediately after a sudden
increase in atmospheric oxygen (evidence for this comes from stable isotopes).
Thus it may be that the initial animal diversification of around 700
million years ago was itself a response to the oxygen level reaching some critical
threshold.
The third alternative is that the biological revolutions themselves somehow
triggered some of the physical events—just the opposite of alternative two!
In this scenario, the common use of calcium carbonate shells by newly evolved
animals changed the way calcium was distributed in the oceans. Similarly, organisms
may have favored the formation of phosphorus, not the other way
around: The presence of many organisms may have changed the physical chemistry
of the ocean environment, boosting the formation of this mineral type.
Knoll leans toward the last alternative. He stresses that the first major
evolutionary radiation among protists and algae (about 1 billion years ago)
may have occurred because of the first evolution of sexual reproduction. The
invention of sex, rather than an environmental trigger, stoked the fires of diversification.
But Knoll also acknowledges the central role of oxygenation in
the evolution of larger animals. Without oxygen, larger animals could never
have evolved, and oxygenation during this interval was facilitated by tectonic
processes—specifically, the role of changes in sea level and erosion of continents
in complex geochemical cycles. For a variety of physiological reasons,
oxygen is a key to the appearance of larger animals; the metabolism of animals
requires oxygen.
Indeed, we may well ask whether oxygenation, and hence the rise of animals,
would ever have occurred on a world where there were no continents to
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erode. Perhaps “water worlds” are ultimately inimical to animal life. But there
may have been even more sudden and catastrophic changes than those listed
by Knoll—most important, dramatic changes in planetary temperature. Evidence
uncovered in the late 1990s has led to a radical new concept: that Earth
almost completely froze over at least twice in its history—once 2.5 billion
years ago and a second time (perhaps repeatedly) during the interval from
about 800 to some 600 million years ago. These times of intense global cold,
when even the oceans were covered with ice, are known as Snowball Earth.
Their biological significance is explored in the next chapter.
Snowball
Earth
“Let it snow, let it snow, let it snow.”
—Christmas song
It is hard to hide our genes completely.
—Philip Kitchner, The Lives to Come, 1996
Spring is universally associated with birth, growth, and fertility. It is a
time of warmth and renewal after the frigid lifelessness of winter. And so
it would seem that the emergence of animals long ago on Earth should
have resulted from a protracted period of warm and fertile, spring-like conditions.
But new information uncovered by several insightful scientists suggests
that the birth of animal life on Earth was initiated not by a time of warmth
but, rather, by the most fearful winter ever to grip the planet. If this phenomenon,
known as Snowball Earth, turns out to be linked to the origins of animal
life, what will it mean for the possibility of animal life on other planets?
As we noted earlier, a majority of astrobiologists believe that the temperature
of early Earth from the time of the first life, about 3.8 billion years
ago, until the origin of eukaryotic cells, about 2.5 billion years ago, was
high—probably too hot for the existence of animal life. (Yet there are others
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who suggest that Earth may have undergone a “cool start,” because the sun at
that time was giving off much less energy than now). Both camps agree that
the planet’s atmosphere was almost devoid of oxygen. Those who believe in
a “hot start” suggest that gradually, as greenhouse gases in the atmosphere
were reduced in volume, the temperatures declined. But Earth may have
cooled too much (or, if you are of the “cool start” persuasion, failed to warm
up enough), at least in the short term. There is evidence of as many as four
major episodes of glaciation on a scale far exceeding anything before or
since—times of cold and ice that make the last ice age, the Pleistocene epoch
of 2.5 million to 10,000 years ago, seem but a brief cold snap.
The first known Snowball Earth episode began about 2.45 billion years
ago, and a second protracted siege of several such events occurred between
800 and 600 million years ago. These two dates are of great interest, because
they are also the times of the two most signal events in biological history
since life’s first appearance here: Around 2.5 billion years ago the first eukaryotic
cells appeared, and the fossil record reveals that about 550 million
years ago, diverse and abundant animal life blossomed, in the event known as
the Cambrian Explosion, the subject of the next chapter. Perhaps it is just coincidental
that these two spectacular and far-reaching biological events occurred
immediately after the two most severe episodes of glaciation and ice
cover in Earth history. But according to a controversial new theory, both may
have been triggered by the Snowball Earth episodes.
I M P R I S O N E D I N I C E
Continental glaciations leave evidence of their former presence: a characteristic
topography on the landscape, grooves and scratches caused as the passing
glaciers ground over hard rock, and (perhaps most important) telltale sedimentary
deposits called tillites. The latter are deposits of angular rock
fragments, which were carried and then left by moving glaciers. The recently
concluded ice ages of 2.5 million to 12,000 years ago left many such deposits
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in both the Northern and the Southern hemispheres. Such tillite deposits are
also found in much older rocks. Thick tillite deposits have been recovered
from two different intervals in Precambrian Earth history: around 2.4 billion
years ago and during the interval from about 800 to 650 million years ago.
The unusual aspect of these features is that they are recovered from virtually
all latitudinal regions of the globe, which shows that the glaciations extended
to near equatorial latitudes (in contrast to the more recent glaciations, which
extended from the poles to mid-latitudes). It may be that no region then on
Earth escaped the glaciation. So much of the planet was covered by ice in
these two Precambrian ice ages that in 1992, Dr. Joseph Kirschvink of Cal
Tech dubbed them “Snowball Earth” events. Far different from the later ice
ages, they were times when Earth teetered dangerously close to becoming
too cold for any life. The Snowball Earth theory received a boost in August
1998 with Harvard geologist Paul Hoffman’s publication, in Science, of new
evidence that ice extended to near equatorial latitudes in the late Precambrian,
about 700 million years ago.
The more recent glaciations, those that occurred since skeletons
evolved about 550 million years ago, affected only land regions; except for an
increase in icebergs, or at most ice cover near the continents, the oceans remained
open. Such may not have been the case in the Precambrian glaciations.
During these two “Snowball Earth” episodes, all of the oceans may have
been covered with ice to considerable depths. And although the deeper regions
of the seas remained liquid, thick icebergs, or pack ice to depths of 500
to 1500 meters, may have covered the ocean. Earth would have been cold indeed.
Average surface temperatures on the planet would have varied between
20°C and 50°C.
These extremely cold temperatures would have had an enormous influence
on the surface of our planet. For example, continental weathering would
have slowed or even stopped. In the interior of continents, the covering of ice
would eventually ablate (evaporate) away, just as it does in the dry valleys of
Antarctica today, leaving behind a sterile rock surface. Dust from these regions
would be blown out to sea, making the pack-ice cover of the oceans
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brown from terrigenous material. From space, Earth would have looked white
and brown—the white being the ice covers on the oceans, the brown the denuded
land areas.
The presence of the pack ice covering the sea would act as a lid on a
pot. Normally, much free exchange occurs across the vast interface of ocean
and atmosphere. Water evaporates from the sea into the air and then rains
back into the sea. If the sea were covered with ice, however, the ocean and
the atmosphere would become “decoupled.” Chemical changes in the ocean
would be separated from the atmosphere by the kilometer-thick lid of ice on
the ocean surface. Very drastic chemical change could—and according to
Kirschvink and others, did—occur within the sea itself.
Even with the icy cover, volcanism would have continued both on the
land’s surface and along the mid-ocean volcanic ridges at the bottoms of the
world’s oceans. At such sites today (see Chapter 1), great volumes of metalrich
fluids gush forth from these submarine volcanoes. In a covered ocean,
this material would have become toxic, producing what are known as reducing
conditions. The oceans would have begun to accumulate with metal ions,
mainly iron and manganese. For as long as 30 million years, the glaciers and
ice never relaxed their frigid grip on the planet’s surface.
All of this global cold would surely have adversely affected life in the
shallow-water regions of all the world’s oceans. The biosphere became restricted
to a narrow belt around the equator and to deep-sea hot springs and
hydrothermal vent settings. Perhaps some life also survived in occasional
Yellowstone-like hydrothermal systems.
Astronomers once thought that a previously warm world’s descent into
such an “icehouse” or “snowball” would be irreversible. Their reasoning was
that as a planet gets more and more thickly coated by ice, the fraction of sunlight
reflected back into space increases and solar heating of the surface declines.
On Earth today, sunlight is adsorbed by the darker land and seas but
is reflected into space by cloud cover. A planet completely covered with ice
would reflect most sunlight into space, causing the planet to become ever
cooler. Yet it is clear that Earth was able to escape from the deep freeze—not
once but several times. The means of that escape was through the volcanic
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emissions of greenhouse gases such as carbon dioxide into the atmosphere,
producing a “greenhouse effect.”
E S C A P E
As we saw in Chapter 2, a planet’s average temperature is greatly affected by
the volumes of greenhouse gases in its atmosphere. Much of this gas enters a
planet’s atmosphere from actively erupting volcanoes. Although there are
abundant volcanic eruptions in the sea as well, most of the carbon dioxide
from these events does not make its way into the atmosphere. Cold seawater
can hold large amounts of dissolved carbon dioxide, and below 700 meters,
CO2 will settle to the bottom of the ocean as it reaches saturation in the
water. At the time of Snowball Earth, enough CO2 would eventually reach
the atmosphere to melt back the sea ice and, in so doing, expose the metalrich
waters of the sea to the atmosphere. The time necessary for this “meltback”
has been estimated by Hoffman and his group to be between 4 and 30
million years. With the ice melted back from the sea, and temperatures again
warming, Earth would have undergone spectacular changes. Here is how
Kirschvink has described these events:
Escape from this “icehouse” condition was only accomplished by
the buildup of volcanic gases, particularly carbon dioxide, mostly
from undersea volcanic activity. Deglaciation during the end of
these glacial events must have been spectacular, with nearly 30
million years of carbon dioxide, ferrous iron, and long buried nutrients
suddenly being exposed to fresh air and sunlight. Hundreds
of meters of carbonate rock are preserved capping the glacial sediments,
at all latitudes, on all continents, as a direct result of wild
photosynthetic activity. For a brief time, the Earth’s oceans would
have been as green as Irish clover, and the sudden oxygen spikes
may have sparked early animal evolution.
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The most important source of biological productivity in the oceans of
today derives from the growth of phytoplankton, the single-celled plants that
are the pastures of the sea. The growth of these plants, so important for producing
oxygen, is limited by the availability of nutrients and iron. If iron is
dropped into the oceans of today, a great bloom of phytoplankton results.
Such was probably the case soon after the end of the first Snowball Earth
event. As the ice-covered seas began to melt, the fine iron- and magnesiumrich
dust coating the surface of the sea ice would have acted as a fertilizer,
tremendously stimulating growth of the blue-green “algae” (really photosynthesizing
bacteria known as cyanobacteria). Enormous populations of
cyanobacteria would have clotted the surface regions of the liberated seas, releasing
huge volumes of oxygen as a consequence of their photosynthetic activity.
This sudden appearance of so much life, after the millions of years of
cold and dearth of life, would have been a great revolution, and it probably
stimulated new evolutionary changes.
These events would have had profound geological as well as biological
ramifications. The sudden rush of oxygen into the sea and air would
have caused the iron- and manganese-rich oceans to precipitate out iron
and manganese oxides. In a previous chapter we saw how banded-iron deposits
began to accumulate about 2.5 billion years ago. Kirschvink and his
group argue that the appearance of banded-iron deposition occurred soon
after the first Snowball Earth ended. Not only iron deposits but magnesiumrich
deposits as well were immediate results of the end of the first Snowball
Earth event. Evidence of this is seen in South Africa, where the world’s
largest land-based deposit of manganese minerals has been dated at 2.4 billion
years of age and sits just above sedimentary deposits that were laid
down during the 2.5-billion-year-old Snowball Earth episode. Like the
banded-iron formations, these manganese-rich deposits appear to be a direct
consequence of the oxygen bloom that occurred when the planetary
snowball melted.
The cessation of the 2.5-billion-year-old Snowball Earth thus appears
to have resulted in a rise in the amount of oxygen both dissolved in the sea
and free in the atmosphere. Probably for the first time in Earth’s history, the
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sunlit portions of the sea became too oxygen-rich to allow iron to exist in solution
in seawater. Kirschvink and his colleagues argue that this dramatic
change in the chemistry of the sea would have exerted intense evolutionary
pressure on life on Earth, then no more advanced than prokaryotic bacteria.
Oxygen, indispensable to the survival of animals, was at that time a poison to
perhaps the majority of life forms. Having evolved in environments with little
or no oxygen, most life experienced the sudden appearance of the chemically
reactive element as a global disaster—but for the rest it was a powerful
evolutionary spur. There were but two choices facing life on Earth in that
long-ago time: Adapt through evolution, or die.
All organisms in the sea had to adapt in two major ways. First, they had
to evolve enzymes capable of mitigating the ravages of dissolved molecular
oxygen and chemicals called hydroxyl radicals. (We humans are still trying to
do this. Our ingestion of antioxidants such as vitamin E and vitamin C is an
attempt to reduce the ravaging effects that dissolved oxygen and “free radicals”
have on living cells.) Second, with the banded-iron formations’ precipitation
out from seawater, living cells no longer inhabited a solution rich in
iron. After having been surrounded by high-iron solution since the first formation
of life, proteins within cells had to be reengineered for life in an environment
low in iron.
Recent DNA sequencing has shown that several enzymes found in archaeans
and eukaryotes are left over from this event of 2.5 billion years ago.
No such enzymes occurred in the older bacteria. The implications of this are
profound: Kirschvink and his colleagues are proposing no less than complete
rejection of the Tree of Life models we examined at the end of Chapter 3,
which suggest that the three great domains (Archaea, Bacteria, and Eucarya)
all arose soon after life’s first evolution at least 3.8 billion years ago. The new
study has not only uprooted this tree; it has burned it. If the Kirschvink group
is correct, two of the three domains—Archaea and Eucarya—arose only after
the 2.5-billion-year-old Snowball Earth and are thus much younger than the
bacteria. Soon after this, in rocks about 2.1 billion years of age, we find a
record of the oldest organelle-bearing eucaryan—the creatures known as
Grypania, which we mentioned in Chapter 3.
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This new version of the Tree of Life is a revolutionary scientific discovery,
and if true, it will utterly reshape our understanding of life’s evolutionary
path. The Snowball Earth events can be seen as biologically important in two
ways. First, the inception of the Snowball produced what may have been the
largest “mass extinction” (the subject of Chapter
in our planet’s history.
The persistence of globally freezing temperatures, the isolation of the ocean
from sunlight, the change in the precipitation patterns on Earth, and the removal
of all water from the surfaces of continents would have removed the
majority of surface habitats then available for microorganisms. In only a few
places could microorganisms have survived: in the deep earth, around hot
springs, and in hydrothermal deposits. Second, Earth’s release from this icy
prison after 30 million years brought about a new catastrophe: from cold to
hot, from oxygen-free to oxygen-rich. Again, organisms had to adapt rapidly.
It is this legacy that we may be seeing in the DNA of all living organisms;
those that survived all bear witness in their DNA to this dual catastrophe—
first cold, then warmth and oxygen. Life on the early Earth went through an
icy bottleneck, and it came out the other side radically changed.
The Snowball Earth of 2.5 billion years ago may have given our planet
eucaryans and the eukaryotic cell necessary for animal life. The second series
of Snowballs (there were several in rapid succession) may have bequeathed
our planet an even more interesting biological legacy—animal life as we
know it.
T H E S E C O N D GL O B A L GL A C I A T I O N
As we saw in Chapter 5, by the next round of Snowball Earth events, those
spanning the time interval from 800 to 600 million years ago, animal life was
present on Earth, but it was newly formed. Either simultaneously with or
soon after the appearance of the new animal phyla, Earth was once more
locked into a global icehouse. Once again, there must have been a period of
mass extinction, as the warm planet froze and the heat-loving organisms of
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Earth had to retreat to oases of heat, such as around volcanoes and hydrothermal
vents, or die. Yet the very severity of these events may have benefited
the newly arisen animals. The great stress inflicted by environmental
conditions imposed by the Snowball events would have stimulated inordinately
rapid evolution among the newly evolved animals. It would also have
caused the isolation of various populations, because the small populations of
life huddled around the undersea volcanoes would have been cut off from any
exchange of genes with other animal groups. This very isolation may have
been largely responsible for the diversity of phyla that emerged at the other
end of these crises, for when the final Snowball Earth event ended, about 600
million years (or less) ago, an entirely new group of creatures was ready to
take over the planet. This is the interval when animal life began to diversify
dramatically, in an event known as the Cambrian Explosion, the subject of
the next chapter.
Would this have happened if the glaciations had not occurred?
Kirschvink and Hoffman suggest that there is a causal link between the cessation
of these major glaciations and the emergence of animals. Hoffman has
noted, “Without these ice events, it is possible there wouldn’t be any animals
or higher plants.” He believes that the melting of the ice at the end of these
ice ages boosted biological productivity—and in the process stimulated evolutionary
activity. This idea has yet to be confirmed, but it remains a tantalizing
possibility.
Both of the two great episodes of Snowball Earth nearly ended life on
Earth, as we know it. But each, ultimately, may have been crucial in stimulating
the great biological breakthroughs necessary for animal life: the evolution
of the eukaryotic cell and then the diversification of animal phyla. This leads
us to ask whether Snowball Earth events are necessary to produce animal life as
diverse as that seen on Earth today.
The end of the last Snowball Earth event brought the time interval
known as the Precambrian to a close. Soon thereafter, abundant skeletons of
larger animals began to fill the sea, in the Cambrian Explosion. If the two
groups of scientists led by Joseph Kirschvink and Paul Hoffman are correct
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about Snowball Earth, a good case can be made that life on Earth is to some
extent due to these events.
92
In the distant past, some early eucaryans (perhaps already endowed
with a nucleus, but still very small and lacking other organelles) may have
routinely ingested other prokaryotic cells for food. This in itself was a major
advance over the prokaryotes, for it necessitated the evolution of an outer
cell wall that could devour, or phagocytize, other cell material—in other
words, it made predation possible. Some of these ingested prokaryotic cells,
however, were not rapidly digested and hence destroyed within the host cell.
Instead, they may have lived there for some time. (Alternatively, these organelles
may have invaded the host cell, rather than being captured by it;
perhaps they bored in and established parasitic colonies within the larger eukaryotic
cell environments.) Eventually, the host cell came to benefit from
this association in some way: Prokaryotes, being very efficient chemical factories,
may have performed services the host could not carry out for itself,
such as energy transformation or even energy acquisition, and metabolic
functions. The organelles known as mitochondria (which are involved in energy
formation and transformation), plastids (the sites of chlorophyll), and
perhaps even flagella (which are used for locomotion) may have evolved in
this fashion.
The prime evidence for this hypothesis comes from DNA. Mitochondria
and plastids contain their own strands of DNA, which are closer in structure
to prokaryotic than to eukaryotic DNA. Mitochondria may have been
free-living bacteria that were capable of oxidizing simple carbohydrates into
CO2 and water and liberating energy in the process. There are living bacteria
today, such as forms known as purple nonsulfur bacteria, that may be close
to the ancestral mitochondrial form. When incorporated into the host cell,
these “guests” eventually lose their cell walls and become part of the host.
With the addition of the cell organelles, our eukaryote approaches or attains
a level of organization that would be familiar to us.
We can now sketch the evolutionary steps necessary to arrive at the eukaryotic
grade of organization. We start with a cell membrane enclosing
DNA—a simple bag of protoplasm and DNA—and then evolve the ability to
phagocytize (or engulf material), evolve a cytoskeleton (which allows us,
among other things, to get larger), evolve aerobic respiration, and then bring
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93
into our much larger bag various organelles: mitochondria, the nucleus, ribosomes,
and so forth.
This last is among the most interesting and controversial aspects of the
evolution of the eukaryotic cell type. Few scenarios have been proposed that
make evolutionary and adaptive sense, but one intriguing possibility has been
described by Dr. Joseph Kirschvink of Cal Tech, who has summarized the
problems faced by the evolving eukaryote as follows:
The problems for the eukaryotic host cell are that:
The host must be large enough to engulf other bacteria.
The host cell must be capable of phagocytosis, so that the
invaders are put into a membrane-bound vacuole (a small
space within the cell), leading to the characteristic double
membrane of the mitochondria and chloroplasts.
The cell should have at least a rudimentary cytoskeleton.
The host cell should offer a better, more controlled environment
for the symbionts, so that natural selection would
favor the association.
The only known bacterium that meets all of these constraints
is called Magnetobacter, discovered in Germany, which
dwarfs most other protists (in size). Each cell of this bacterium
makes several thousand organelles called magnetosomes, which
are tiny crystals of the mineral magnetite (Fe3O4) encased in a
membrane bubble—a bubble that forms by phagocytosis. These
magnetosomes are held in place in chain-like structures that keep
each crystal aligned properly; this can only be done if an intracellular
mechanical support structure such as the cytoskeleton exists.
Magnetobacter has the ability to keep itself in the optimal environment
by swimming along the magnetic field lines generated by the
Earth’s magnetic field. This ability makes it an attractive partner
for symbiosis, as many organisms spend a great deal of their metabolic
energy staying in the correct environment.
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This scenario for the evolution of the eukaryotic cell has two
major implications for the timing of higher life on Earth. First, the
most probable path for the evolution of magnetotaxis (the ability
to align along magnetic fields) and magnetite biomineralization
(the formation of minerals by living organisms) is a result of natural
selection for iron storage. Anaerobic microbes do not need iron
storage mechanisms, as ferrous iron is freely available in solution.
But in oxygen-rich environments the iron rusts out the [ferrous]
form, which drops out of solution. Hence, magnetotaxis is unlikely
to evolve in an anaerobic world, which on Earth ended
about 2.5 to 2 billion years ago. The oldest magnetofossil—the
fossil remains of bacterial magnetosomes—date to about 2 billion
years ago. Second, magnetotaxis requires the presence of a strong
planetary magnetic field. On Earth, a strong early field probably
decayed after 3.5 billion years ago, only to reach its present level
after nucleation of the inner core about 2.8 billion years ago.
Kirschvink has thus postulated a novel scenario—and perhaps the most
plausible scenario—for the formation of the eukaryotic cell: a pathway necessitating
the presence of magnetite and a strong planetary magnetic field.
As we shall see in a later chapter, not all planets maintain magnetic fields. If
this pathway is the only way to large eukaryotic cells (a hypothesis that still
awaits verification), then we have another requirement we must impose on
planets that aspire to host animal life—a magnetic field.
E N V I R O N M E N T A L CO N D I T I O N S L E A D I N G
TO T H E E V O L U T I O N O F E U K A R Y O T E S
What environmental conditions led to the evolution of the forerunners of animal
life? New discoveries of the 1980s and 1990s have given us a much
clearer view of the early Earth during the great evolutionary transitions we
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saw in the last chapter. The Earth’s earliest life seems to have formed during
or soon after cessation of the heavy comet bombardment. By about 3.8 billion
years ago that heavy cosmic bombardment ended, and by 3.5 billion
years ago we find the first fossilized evidence of life.
The region that has yielded Earth’s oldest fossils found to date is known
to Australians as the North Pole, because even in this isolated continent, it is
uniquely remote and inhospitable. The rocks in this region belong to a unit of
interbedded sedimentary and volcanic rocks known as the Warrawoona Series.
Geologists have deduced that the deposits were consolidated in a shallow
sea over 3.5 billion years ago. There is the evidence of storm layers and evidence
as well that on occasion, a hot sun evaporated small pools of seawater
into brine deposits. But it is not these structures that have created so much excitement
about the Warrawoona rocks. This ancient bit of Australia holds the
world’s oldest stromatolites, low mounds of lime and laminated sediment that
have been interpreted as the remains of microbial mats—in other words, life.
Stromatolites (the “stone mattresses” we mentioned earlier as an anomaly,
multicellular prokaryotes) are the most conspicuous fossils and the most
commonly preserved evidence of life for more than 3 billion years of Earth
history; they provide our best record of early life. They have been found on
every continent in rocks half a billion years old and older. Today, they are
found in only one type of environment on Earth, in quiet, briny tropical waters.
Such environments are refuges from algal grazers; stromatolites can no
longer exist on most of our planet’s surface because they would quickly be
eaten. The photosynthesizing bacteria termed cyanobacteria are modern
equivalents of these ancient deposits.
The presence of stromatolites is a sure clue that by 3.5 billion years ago,
life on this planet had left its earliest, probably hydrothermal or deep-earth environments
and diversified onto the surface of the planet. For a billion years the
prokaryotes were masters of the world, but life was still scattered. According to
the fossil record, it was not until about 2.5 billion years ago that the organisms
that produced stromatolites had released sufficient quantities of oxygen to form
sedimentary deposits known as banded-iron formations. Prior to the appearance
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of common stromatolites, there was no dissolved oxygen in the sea, no gaseous
oxygen in the atmosphere, and hence no possibility of mineral oxidation. With
the appearance of oxygen, however, large volumes of iron that had been dissolved
in seawater precipitated out as it oxidized into iron oxides—rust, in
other words. Today, there still exist at least 600 trillion tons of such iron oxides
deposited before 2.5 billion years ago in these banded-iron formations.
The time interval commencing about 2.5 billion years ago is marked by
a change in the very tectonic nature of the planet Earth—its rate of mountain
building and continental drift. By this time, the heat production from radioactive
elements locked in Earth’s rocks had diminished, for some of the
radioactive elements decayed rapidly early in Earth’s history. This material
was like a finite amount of fuel within the interior of the planet, and as it was
used up, heat flow declined. It turns out that the processes of continental drift
and mountain building are by-products of heat rising from within Earth, and
as the amount of heat decreased over time, so did these two activities. There
is also some evidence that around this time, a major pulse of land formation
occurred, allowing larger continental land masses to form. As the new continents
formed, many shallow-water habitats were created, and these proved
favorable environments for the growth of photosynthesizing bacteria. We
can speculate that from about 4 billion to about 2.5 billion years ago, there
were few large continents, but numerous volcanic island chains dotted the
world. After 2.5 billion years ago, continental land masses began to form, and
volcanism on a global scale lessened.
With this increase of habitat, ever more stromatolites grew and flourished.
This in turn relentlessly pumped ever more oxygen into the sea. As
long as there was dissolved iron in the seawater, all of the liberated oxygen
was quickly locked up in the banded-iron formations. By about 1.8 billion
years ago, however, this reservoir of dissolved iron material was used up. We
know this because after that time, no more banded-iron formations were laid
down. This changeover left an indelible mark on the sedimentary record of
Earth, for as the sea became saturated with oxygen, the time of banded-iron
formations ended forever—or at least until some far-distant future when our
planet may again no longer have oxygen. With nowhere else to go, oxygen
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began to emerge into our planet’s atmosphere and, in so doing, probably gave
life its first impetus toward animal life.
T H E OX Y G E N R E V O L U T I O N
It is probably impossible for us to conceive how entirely alien to ours this
world truly was. Yet the strange microbial world of 2 billion years ago may be
the norm in the Universe for those planets that harbor life. Traces of it exist
still, here on Earth, in the bacterial froths and pond scum that persist across
our planet, and perhaps nowhere more prolifically than in the rotting garbage
dumps and landfills created by our own species—places where huge, visible
colonies of rapidly growing bacteria exist still. But the rainbow slick of the
oozing swamp is the exception in a world where the eucaryans are so much
more in evidence than the prokaryotic forms. What would that 2-billionyear-
old world look like? The best description we know of was penned by
two scientists who have journeyed back to this world, in their imaginations,
many times. We owe the following image of the ancient Proterozoic era (the
formal name for the time interval of 2.5 to 0.5 billion years ago) to Lynn Margulis
and Dorion Sagan, in their 1986 book Microcosmos:
To a casual observer, the early Proterozoic world would have
looked largely flat and damp, an alien yet familiar landscape, with
volcanoes smoking in the background and shallow, brilliantly colored
pools abounding and mysterious greenish and brownish
patches of scum floating on the waters, stuck to the banks of rivers,
tainting the damp soils like fine molds. A ruddy sheen would coat
the stench-filled waters. Shrunk to microscopic perspective, a fantastic
landscape of bobbing purple, aquamarine, red, and yellow
spheres would come into view. Inside the violet spheres of Thiocapsa,
suspended yellow globules of sulfur would emit bubbles of
skunky gas. Colonies of ensheathed viscous organisms would
stretch to the horizon. One end stuck to rocks, the other ends of
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some bacteria would insinuate themselves inside tiny cracks and
begin to penetrate the rock itself. Long skinny filaments would
leave the pack of their brethren, gliding by slowly, searching for a
better place in the sun. Squiggling bacterial whips shaped like
corkscrews or fusilli pasta would dart by. Multicellular filaments
and tacky, textilelike crowds of bacterial cells would wave with the
currents, coating pebbles with brilliant shades of red, pink, yellow
and green. Showers of spheres, blown by breezes, would splash
and crash against the vast frontier of low-lying mud and waters.
This prokaryotic world was creating what has been called the Oxygen
Revolution. The initiation of an oxygenated atmosphere was one of the most
significant of all biologically mediated events on Earth. Prokaryotic bacteria,
using only sunlight, water, and carbon dioxide, ultimately transformed the
planet by generating an ever-increasing volume of atmospheric oxygen. This
outpouring of oxygen created both biotic opportunity and biotic crises. Many
of Earth’s primitive organisms were metabolically incapable of dealing with
abundant oxygen. For most of the archaeans, the oxygen boom of about 2 billion
years ago was an environmental disaster, driving some species into airless
habitats, such as lake and stagnant ocean bottoms, sediments, and dead organisms.
Others were incapable of such migration and simply died out. For yet
other creatures, however, the profound change in atmospheric conditions created
new opportunities. Some prokaryotic cells began to exploit the enormous
power of oxygen metabolism to break down food sources into carbon dioxide
and water. This new metabolic pathway yielded far more energy than any of
the anaerobic pathways. Organisms that adopted it soon began to take over
the world. The most efficient of these were members of the domain Eucarya,
which, more than 2 billion years ago, evolved true eukaryotic cell machinery.
The oldest known fossils of an organism that appears to have attained
the eukaryotic grade of organization have been found in banded-iron deposits
located in Michigan. The fossils themselves are about 1 millimeter in
diameter and are found in chains as much as 90 millimeters long. The organism,
then, is far too large to be a single-celled prokaryote or even a singleHow
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celled eukaryote. This creature, which has been named Grypania, is preserved
as coiled films of carbon on smooth sedimentary rock bedding planes, the
places where sedimentary beds split apart. Its 1992 discovery indicates that
the evolution of the first eukaryotic cell occurred during the banded-iron formation
process, when there was still little free oxygen in the sea and probably
none in the atmosphere. These early eukaryotes may have been vanishingly
rare, for other eukaryotes do not occur in the fossil record for 500
million years after this first appearance, but with this form, a beachhead in
life’s advance had been established.
For the period between 2 and 1 billion years ago (see Figure 5.2), few notable
achievements of life are recorded as fossils in the rocks. The first common
appearance of eukaryotes begins about 1.6 billion years ago, when microscopic
fossils called acritarchs begin to appear in the geological record. These are
spherical fossils with relatively thick, organic cell walls. They are interpreted to
be the remains of planktonic algae, forms that used photosynthesis and lived in
the shallow waters of the world’s oceans. Other life forms evolved as well, but
as is also true of most living protists, such as the amoeba and the paramecium,
their lack of skeletons renders them invisible in the fossil record. With a proliferation
of plant-like forms, new varieties of predatory protists surely evolved.
Whole armadas of single-celled, floating pastures and the somewhat larger
and more mobile grazers on these fields of plankton lived and died in this
seemingly endless epoch of geological time. The open ocean would have had
little life, but the coastal regions richer in nutrients would have been awash
with life—microscopic life. It was the Age of Protists, the Age of the Small.
We have now reached 1 billion years ago, in our march through evolutionary
time. Finally, the tempo of evolutionary development increased, if we
are correctly interpreting the fossil record, for there is a burgeoning in the
number of eukaryotic species found in the rock record at this time. Some of
these new forms include the first red and green algae, forms still crucial and
varied in marine ecosystems. This diversification of eukaryotic species, including
protozoans and plants, set the stage for the evolution of larger, multicellular
forms and may have been triggered by the evolution of important
new morphologies within the eukaryotic cell.
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E V O L U T I O N I N E U K A R Y O T I C
FORM AND F U N C T I O N
Four biological innovations may have been especially significant in paving
the way for the emergence of larger animals: (1) the development of sexual
cycles; (2) new methods of shuffling information coded along the chromosomes
(through the nascent ability to excise and relocate entire gene sequences);
(3) new methods of communicating between cells via substances
called protein kinases, and (4) the development of a new type of intracellular
skeleton, called a cytoskeleton, that allowed eukaryotic cells to increase
enormously in size. These innovations greatly enhanced the ability of cells to
Archaean Proterozoic Phanerozoic
Paleoproterozoic Mesoproterozoic Neoproterozoic
Grypania megafossils
Undoubted multicellular algae
Chuaria-Tawuia assemblage
Longfengshania
Worm-like megafossils
Ediacara-type fossils
Simple trace fossils
Cloudina
Skeletal fossils
Complex trace fossils
Trilobites
Chengjiang fauna
2.5 2.0 1.5 1.0 0.5 0 Ga
Varanger ice age
?
Figure 5.2 Early multicellular fossils. Broken bars indicate uncertain time ranges.
E A R L Y MU L T I C E L L U L A R F O S S I L S
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evolve new morphologies in response to natural selection and their ability to
band together into multicellular creatures.
We can now better categorize what we term “advanced” life: eukaryotic
multicellular organisms. There are, of course, many types of multicellular organisms,
including a considerable number of prokaryotic forms. In most cases
these multicellular prokaryotes are composed of only two cell types. Cellular
slime molds are multicellular, as are some cyanobacteria. In a way, however,
these forms are evolutionary dead ends. They have existed on Earth for several
billion years and are highly conservative in an evolutionary sense. It is
multicellular creatures of the other category that became so important in the
history of life. We refer here to true metazoans.
The jump from single-celled organisms to organisms of multiple cells
requires numerous evolutionary steps. The jump from single-celled organisms
to metazoan animals, where a high degree of intercellular cooperation in organization
exists, involves even more. In their recent book Cells, Embryos and
Evolution, biologists John Gerhart and Marc Kirschner discuss this evolutionary
accomplishment. The first step, they argue, seems almost paradoxical: It
was not some new structure gained that allowed this transition, but an important
structure lost. Long ago in our planet’s past, some organism of the eukaryotic
lineage made a brave (or lucky) morphological change—it shed its
external cell wall. Why this occurred is still unclear, but the net effect was
far-reaching. A tough outer coating protects most unicellular creatures from
their surrounding environment. At the same time, however, it isolates these
cells from other members of their own kind. By divesting themselves of this
outer wall, individual cells could begin exchanging living material—and
information—with one another. The naked cells could adhere to each other,
crawl over each other, and communicate. These were the first steps in the formation
of a tissue, which is an aggregation of cells united for mutual benefit.
Larger animals require highly integrated systems of cells that can accomplish
the myriad functions necessary for all life. Respiration, feeding,
reproduction, the elimination of waste material, information reception,
locomotion—all require the integration of many cells acting in concert. Each
of these functions ultimately requires one or many types of tissues.
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Among tissue types, the outer wall of any organism (the epithelium) is
of utmost importance. The epithelium must protect the organism from the
rigors of the external environment but at the same time allow adsorption of
critical gas and, sometimes, nutrients. The evolution of the epithelium was a
decisive first step in the evolution of metazoans.
Which group of unicellular creatures first achieved this breakthrough?
The most primitive and enigmatic of larger eukaryotic metazoans are
sponges. These curious creatures seem to bridge the gap between singlecelled
eukaryotes or even colonial protozoa and the highly integrated invertebrate
metazoan phyla. Sponges have several cell types that perform specialized
tasks, but there is a very low level of organism-wide organization.
There is no gut or body cavity specialized for processing food, nor is there
any nervous system. Yet the sponges may be an important clue to the identity
of our actual metazoan ancestors.
The stem, or ancestral, metazoan probably had a larger number of cell
types than sponges (perhaps 10 to 15 rather than the 3 to 5 individual cell
types found in sponges). There was probably a body cavity of some sort segregated
into two cell layers: an outer ectoderm and an inward-facing endoderm.
This two-tissue plan seems to have been an evolutionary dead end, and
it wasn’t until a third layer—the mesoderm—was added that animals with
real internal complexity formed. Eventually, a small worm-like shape with
three tissue layers evolved, a creature with a gut running through the long
axis of the body and a separate space known as a coelom to serve as an internal
hydrostatic skeleton. With this tiny organism (the first may have been
less than a millimeter long), the evolutionary stage was set for the emergence
of animals on planet Earth.
T H E TWO DI V E R S I F I C A T I O N S
O F A N I M A L P H Y L A
With the advent of this form—what evolutionary biologists call the “roundish
flatworm”—a body plan was in place that could be modified to shape all the
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categories of metazoan life, the major body plans that we call phyla. The
phyla living today include the arthropods; the mollusks; the echinoderms;
our own group, the chordates; and about 25 more. These are the complex
metazoans that we hope to find—but that may be very rare—on other planets.
These are animals. They appeared relatively late in the history of life on
Earth. One of the great novel insights of the 1990s was our realizing that
their origin and their subsequent diversification and rise to abundance were
two separate events, not one, as had been believed since the time of Charles
Darwin.
Fossils of macroscopic animals (those visible to the unaided eye) first
appear in abundance less than 600 million years ago, during the “Cambrian
Explosion,” a diversification event resulting in the rapid formation of thousands
of new species; we will describe it in more detail in the next chapter.
Yet the appearance of abundant animal fossils at this time actually marks the
second of the two diversification events that led to the proliferation of larger
animals on the planet. As we will show, fossils of such complex animals as
trilobites and mollusks—common members of the Cambrian Explosion—are
advanced descendants of a much earlier, diversification event that took place
between 1 billion and 600 million years ago. Yet there is no fossil record of
this first diversification—paleontologists have been stymied by an almost
compete lack of fossils in strata older than 600 million years, when this initial
event must have taken place. Our understanding of the initial diversification
of animals comes not from paleontology but from an entirely different line of
investigation: genetics. Geneticists have arrived at answers about the “when”
of the first diversification event by examining the genetic code of living animals
via a technique called ribosomal RNA analysis.
Gene sequences are simply strings of base pairs lined up along the double
helix of a DNA molecule. As we saw earlier, if a DNA molecule is likened
to a twisted ladder, the base pairs can be considered the steps of the ladder,
and it is the sequence of the steps that is used in this type of analysis. Genes
are simply instructions for protein formation coded by the sequence of nucleotides
on the DNA ladder. There are only four types of nucleotides, but
they provide the genetic code that is the basis for all Earth life. All organisms
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share more genes with their ancestors than with nonrelated species. By comparing
the genes from various organisms, it is possible to produce a model of
evolutionary history (an evolutionary tree, as it were) with the branches of
the tree showing which species gave rise to which other species. Yet according
to many geneticists, such an analysis not only tells us how the branching
occurred, it can also tell us when.
In 1996 G. Wray, J. Levinton, and L. Shapiro published a paper claiming,
on the basis of results obtained by using this genetic technique, that the
first event—the earliest divergence of animals—occurred 1.2 billion years
ago. This result drew a collective gasp from the paleontological fraternity: It
seemed much too ancient. The fundamental assumption of the Wray et al.
paper is that gene sequences evolve with sufficient regularity that a sort of
molecular “clock” can be used to date the divergence of various groups. The
reasoning behind the molecular clock technique is that changes in the genetic
code—evolution, in other words—occur at a rather constant rate. The
more distinct two DNA sequences are, the longer it has been since they diverged
from a common ancestor. Other scientists, however, dispute that
changes in gene frequency occur at a constant rate, and therefore they do not
believe in the molecular clock. It is these molecular clock data that led the
Wray group to their conclusion. This finding was a bombshell. If animals
evolved this early, why did they not appear in the fossil record until less than
600 million years ago? What were they doing for such a long time?
The Wray group’s findings were extremely controversial not only because
they contradicted long-held paleontological dogma but also because
they provoked criticism among other geneticists. There is fierce debate
among geneticists about the reliability of the molecular clock technique. The
Wray study itself, yielded both minimum and maximum figures for the earliest
divergence. One group of genes suggested that the fundamental splitting
of the phylum made up of annelids (worms) from the phylum of chordates
(our phylum) occurred only 773 million years ago, whereas a second group of
genes (in the same organisms) suggested 1621 million years ago—a very wide
spread indeed! These results give us minimum and maximum ages for the diHow
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vergence. Even with the minimum figure, however, there were (according to
the molecular data, anyway) recognizable chordates and annelids 700 million
years ago—yet there is no trace of their presence in the fossil record. Where
were they? Or were they not there at all? Could it be that no rocks of this age
survive or that no fossils from the interval of about 1 billion to less than 600
million years ago were preserved? This seems to be stretching things, as suggested
by British paleontologist Simon Conway Morris:
Appeals to gaps in the rock record and pervasive metamorphism
of the sediments are not going to work: if there were large metazoans
capable of either fossilization or leaving traces, they had an
uncanny knack of avoiding areas of high preservation potential.
Since the original, tantalizing analysis by the Wray group, other geneticists
have reconsidered the basic data. Most concede that the 1.2-billion-year
figure is too old. (However, a report published in Science magazine in late 1998
by a team headed by Adolf Seilacher of Yale University announced the discovery
of billion-year-old trace fossils (worm-tracks) possibly derived from small,
worm-like organisms. Critics of this finding suggest that the marks in question
could just as easily have been produced by inorganic actions, and even if these
trace fossils turn out to have been produced by organisms, the question remains:
Why are no further such fossils found for hundreds of millions of years?)
Let’s say, then, that divergence occurred less than a billion years ago. We must
still account for a significant period of time with animals but without fossils. Paleontologists
have long believed that only a single major diversification event
occurred—the event coincident with the appearance of fossils, the so-called
Cambrian Explosion that began about 550 million years ago. Now this evolutionary
event is seen as a follow-up to the much earlier first event.
The answer to this seeming conundrum is that the animals were indeed
present, but they were so small as to be essentially invisible in the fossil record.
A recent and spectacular discovery of microscopic fossil animal embryos
seems to confirm this view. Using newly developed techniques of searching
for tiny (but complex) animals in minerals called phosphates, paleontologist
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Andy Knoll and his colleagues have uncovered a suite of tiny but beautifully
preserved fossils interpreted to be the embryos of 570-million-year-old
triploblasts—animals with three body layers, like most of those found today.
These fossils tell us that the ancestors of the modern phyla were indeed present
at least 50 million years before we find any conventional fossil record of
them. The combination of genetic information and new discoveries from the
fossil record now give us a robust view of the rise of animals: They did not
exist 1 billion years ago, and perhaps not 750 million years ago. Animals are
indeed very late arrivals on the stage of life on Earth.
Thanks to these new discoveries and interpretations, the question of
“when” has been answered to most people’s satisfaction: The emergence of animals
was a two-stage event. The initial stage seems to have occurred less (and
perhaps much less) than the billion years ago proposed by Wray and his colleagues.
But even recalibrated, the Wray group’s finding has given us yet another
tantalizing insight into the potential incidence of animal life in the Universe.
The Wray work confirms that there were indeed two “explosions.” The
first was the actual differentiation of the various body plans; the second was the
differentiation and evolution, in these various phyla, of species large and abundant
enough to enter the fossil record. The geneticists can show that genes of
annelid worms and genes of chordates were differentiating hundreds of millions
of years before the emergence of these creatures as large entities that
could appear in the fossil record. This leads us to ask a crucial question: Even if
they evolve, do animals necessarily, or inherently, go on to diversify, enlarge, and survive?
Does the second flowering of animal life—the Cambrian Explosion event so
long known to geologists—inevitably follow the first diversification, or is it yet
another threshold of possibility that may be (but is not necessarily) attained?
Perhaps on some worlds in the Universe, animals diversify but never attain
larger size and greater numbers in some Cambrian Explosion equivalent. This
particular insight was first expressed by paleontologist Simon Conway Morris:
We need to discuss to what extent metazoan history was implicit a
billion years ago, at least in outline, as opposed to what was inevitable
500 million years later at the onset of the Cambrian exHow
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plosion. Even if metazoans have a deep history, which paleontologically
remains cryptic, the actual organisms would have been of
millimeter size and perhaps without the potential for macroscopic
size and complex ecology. . . . Wray et al. may have been correct
in tracing the gunpowder back as far into the mists of the Neoproterozoic
(the late Precambrian time period of a billion years
ago), but the keg itself still looks as if it blew up in the Cambrian.
In other words, it seems that the development of animals was a two-step process,
with step two—the Cambrian Explosion—not necessarily being an outcome
predetermined by the initial differentiation of the animal phyla.
Over and over the same question arises: Why did it take so long for animals
to emerge on planet Earth? Was it due to external environmental factors,
such as the lack of oxygen for so long in the history of this planet, or to
biological factors, such as the absence of key morphological or physiological
innovations?
T H E E V O L U T I O N O F A N I M A L S :
B I O L O G I C A L B R E A K T H R O U G H O R
E N V I R O N M E N T A L S T I M U L U S ?
Complex animals surely cannot appear on any planet without following some
evolutionary pathway from simpler, single-celled organisms. The change
from single-celled microbes to multicellular creatures must be the common
route on any planet, and even if the molecules of life are different from world
to world, the pathway from simple to complex may be universal. Because of
this, the example of how animals evolved on our Earth may be of the utmost
importance in understanding the frequency with which animals occur on
other planets.
If we are to understand how animals evolved from single-celled ancestors,
we must first understand the environments where these monumental
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Million
years Traditional view Wray et al. view Compromise view
300
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Perm
Dev
Sil
Ord
Cam
Carb
Mollusca
Annelida
Echinodermata
Arthropoda
Agnatha
Gnathostomata
Mollusca
Annelida
Echinodermata
Arthropoda
Agnatha
Gnathostomata
Mollusca
Annelida
Echinodermata
Arthropoda
Agnatha
Gnathostomata
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Proterozoic
Riphean Vendian
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years
Figure 5.3 Differing views of metazoan phylogeny. Most paleontologists follow the “traditional view”
(left), accepting the fossil record as a fairly reliable indicator of original events. Molecular clocks are interpreted
by Wray et al. (center) as indicating very deep origins for the principal metazoan phyla. The recognition
that some molecular clocks run much faster than others suggests a “compromise view” (right), which implies
that our search strategy for the first metazoans should be concentrated in the interval from about 750
million years onward. Perm, Permian; Carb, Carboniferous; Dev, Devonian; Sil, Silurian; Ord, Ordovician;
Cam, Cambrian.
evolutionary advances were made. We know well the “when” of this change—
it took place during a 500-million-year interval from 1 billion to 550 million
years ago. The second event, the Cambrian Explosion of between 550 and 500
million years ago, included the morphological diversification of the phyla into
subdivisions based on body plans, as well as the appearance, within the various
phyla, of species with skeletons and large size (see Figure 5.3).
During this interval of time, Earth went though major environmental
changes, among them ice ages of unprecedented severity, rapid continental
movements, and drastic changes in ocean chemistry. We are thus left with
perplexing questions: Did the environmental changes of this interval (which
are described in more detail below) somehow trigger the diversification of anHow
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imals? Or would the rise of animals have occurred even in the absence of
these profound environmental changes? These questions, which of course are
central to understanding the evolution of life on our planet, have great relevance
to understanding the frequency of animal life on other planets as well.
Does animal life always (or even commonly) evolve, once a suitable ancestor
appears? Or does there need to be an additional trigger of some sort, a sequence
of environmental steps? We might compare this whole process to
baking a cake. Say the ingredients for the batter were all assembled and
mixed by 1 billion years ago. Does the cake need to be cooked for a given
time at a highly restricted temperature in order to rise? Or will any amount of
cooking at any temperature accomplish the task just as well? Or will our cake
be completed without any cooking at all? (That is, does simply assembling
the ingredients into a batter ensure success?)
The beginning of this fecund period in Earth history is marked by the
appearance not of new types of animals, but of plants. Around 1 billion years
ago, many types of algae begin to appear in the fossil record, including the
green and red algae still so prominent on Earth today. These were not the ancestors
of animals, of course, but their appearance was the opening salvo of
an evolutionary assault that was the most significant up to that time. It was
followed, hundreds of millions of years later, first by the initial diversification
of animal phyla and then (after more hundreds of millions of years) by the
Cambrian Explosion of animal life.
What were the environmental events of this interval of time 1 billion to
600 million years ago? By this period, land masses approaching the size of
today’s continents had formed, and the total area of land on the planet may
not have been significantly different from what we see in the present day.
The land, however, was not a tranquil place. The period was one of significant
mountain building and continental drift. It was also marked by episodes
of continental glaciation unmatched in severity since that time. Did these
events have anything to do with the diversification of animals? One school of
thought says yes. Work by Martin Brasier and others suggests that rapid
changes in sea level, and especially the formation of broad, shallow seas
within the new continents, would have opened up many new habitats very
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hospitable in terms of temperature and nutrients. This, in turn, may have
stimulated the diversification of animals and plants. There are dissenters, notably
James Valentine, who cautions, “the link between plate tectonics . . .
and the origins and radiation of animals remains to be demonstrated.” But, as
Harvard paleobiologist Andy Knoll points out, there is another way in which
the new and active tectonic events could have influenced the initial radiation
of animals that occurred during this time. In 1995 Knoll noted that “tectonic
processes could have influenced one or more of the great radiations (of animals)
. . . through their participation in the biogeochemical cycles that regulate
Earth’s surface environments.”
Examples of such effects include the role of hydrothermal influences on
ocean chemistry. The hydrothermal vents, as we saw in Chapter 1, are submarine
regions where great volumes of hot and chemically distinctive water
are mixed with seawater. The amount of this volcanically derived water entering
the oceans fluctuated during the interval of 1 billion to 550 million
years ago, and these fluctuations had marked effects on the chemistry of the
seawater, on the composition of the atmosphere, and on climate. The tectonics
events also affected the rate of burial and exhumation of organic carbon
in sediments. Oxygen and carbon dioxide values shifted, and as they did
so, major changes in the temperature and oxygenation of the planet ensued.
Yet another environmental stimulus may also have contributed to the
initial animal diversification. Changes in ocean chemistry caused by increased
tectonic activity beginning a billion years ago facilitated the evolution of
skeletons. This period is marked by the appearance of rocks called phosphorites.
Some authors credit these rocks with bringing about an increase in the
fertility of the oceans at this time, which may in turn have helped trigger the
sudden appearance of many diverse animals beginning about 600 million years
ago. Phosphorus is much more concentrated in living things than in the environment,
so it is a limiting nutrient. The sudden presence of abundant sources
of this element could have acted as a veritable fertilizer for growth.
Knoll has discussed all of these disparate factors and has proposed three
alternatives. First, it may be that the complex physical events and the equally
complex series of biological events that occurred from 1 billion to 550 milHow
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lion years ago are simply coincidental—they had nothing to do with one another.
If this is true, the great biological diversification must be attributed
solely to biological innovations (such as the ability of cells to bind together,
build an outer cell wall, and evolve internal cooperation between contained
cells) that were unrelated to concurrent environment changes.
The second alternative is that evolution was indeed facilitated by
changes in the physical environment. The most important of these changes
may have been in levels of oxygen. The first appearance of larger metazoans,
the ediacarans, about 600 million years ago occurred immediately after a sudden
increase in atmospheric oxygen (evidence for this comes from stable isotopes).
Thus it may be that the initial animal diversification of around 700
million years ago was itself a response to the oxygen level reaching some critical
threshold.
The third alternative is that the biological revolutions themselves somehow
triggered some of the physical events—just the opposite of alternative two!
In this scenario, the common use of calcium carbonate shells by newly evolved
animals changed the way calcium was distributed in the oceans. Similarly, organisms
may have favored the formation of phosphorus, not the other way
around: The presence of many organisms may have changed the physical chemistry
of the ocean environment, boosting the formation of this mineral type.
Knoll leans toward the last alternative. He stresses that the first major
evolutionary radiation among protists and algae (about 1 billion years ago)
may have occurred because of the first evolution of sexual reproduction. The
invention of sex, rather than an environmental trigger, stoked the fires of diversification.
But Knoll also acknowledges the central role of oxygenation in
the evolution of larger animals. Without oxygen, larger animals could never
have evolved, and oxygenation during this interval was facilitated by tectonic
processes—specifically, the role of changes in sea level and erosion of continents
in complex geochemical cycles. For a variety of physiological reasons,
oxygen is a key to the appearance of larger animals; the metabolism of animals
requires oxygen.
Indeed, we may well ask whether oxygenation, and hence the rise of animals,
would ever have occurred on a world where there were no continents to
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erode. Perhaps “water worlds” are ultimately inimical to animal life. But there
may have been even more sudden and catastrophic changes than those listed
by Knoll—most important, dramatic changes in planetary temperature. Evidence
uncovered in the late 1990s has led to a radical new concept: that Earth
almost completely froze over at least twice in its history—once 2.5 billion
years ago and a second time (perhaps repeatedly) during the interval from
about 800 to some 600 million years ago. These times of intense global cold,
when even the oceans were covered with ice, are known as Snowball Earth.
Their biological significance is explored in the next chapter.
Snowball
Earth
“Let it snow, let it snow, let it snow.”
—Christmas song
It is hard to hide our genes completely.
—Philip Kitchner, The Lives to Come, 1996
Spring is universally associated with birth, growth, and fertility. It is a
time of warmth and renewal after the frigid lifelessness of winter. And so
it would seem that the emergence of animals long ago on Earth should
have resulted from a protracted period of warm and fertile, spring-like conditions.
But new information uncovered by several insightful scientists suggests
that the birth of animal life on Earth was initiated not by a time of warmth
but, rather, by the most fearful winter ever to grip the planet. If this phenomenon,
known as Snowball Earth, turns out to be linked to the origins of animal
life, what will it mean for the possibility of animal life on other planets?
As we noted earlier, a majority of astrobiologists believe that the temperature
of early Earth from the time of the first life, about 3.8 billion years
ago, until the origin of eukaryotic cells, about 2.5 billion years ago, was
high—probably too hot for the existence of animal life. (Yet there are others
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who suggest that Earth may have undergone a “cool start,” because the sun at
that time was giving off much less energy than now). Both camps agree that
the planet’s atmosphere was almost devoid of oxygen. Those who believe in
a “hot start” suggest that gradually, as greenhouse gases in the atmosphere
were reduced in volume, the temperatures declined. But Earth may have
cooled too much (or, if you are of the “cool start” persuasion, failed to warm
up enough), at least in the short term. There is evidence of as many as four
major episodes of glaciation on a scale far exceeding anything before or
since—times of cold and ice that make the last ice age, the Pleistocene epoch
of 2.5 million to 10,000 years ago, seem but a brief cold snap.
The first known Snowball Earth episode began about 2.45 billion years
ago, and a second protracted siege of several such events occurred between
800 and 600 million years ago. These two dates are of great interest, because
they are also the times of the two most signal events in biological history
since life’s first appearance here: Around 2.5 billion years ago the first eukaryotic
cells appeared, and the fossil record reveals that about 550 million
years ago, diverse and abundant animal life blossomed, in the event known as
the Cambrian Explosion, the subject of the next chapter. Perhaps it is just coincidental
that these two spectacular and far-reaching biological events occurred
immediately after the two most severe episodes of glaciation and ice
cover in Earth history. But according to a controversial new theory, both may
have been triggered by the Snowball Earth episodes.
I M P R I S O N E D I N I C E
Continental glaciations leave evidence of their former presence: a characteristic
topography on the landscape, grooves and scratches caused as the passing
glaciers ground over hard rock, and (perhaps most important) telltale sedimentary
deposits called tillites. The latter are deposits of angular rock
fragments, which were carried and then left by moving glaciers. The recently
concluded ice ages of 2.5 million to 12,000 years ago left many such deposits
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in both the Northern and the Southern hemispheres. Such tillite deposits are
also found in much older rocks. Thick tillite deposits have been recovered
from two different intervals in Precambrian Earth history: around 2.4 billion
years ago and during the interval from about 800 to 650 million years ago.
The unusual aspect of these features is that they are recovered from virtually
all latitudinal regions of the globe, which shows that the glaciations extended
to near equatorial latitudes (in contrast to the more recent glaciations, which
extended from the poles to mid-latitudes). It may be that no region then on
Earth escaped the glaciation. So much of the planet was covered by ice in
these two Precambrian ice ages that in 1992, Dr. Joseph Kirschvink of Cal
Tech dubbed them “Snowball Earth” events. Far different from the later ice
ages, they were times when Earth teetered dangerously close to becoming
too cold for any life. The Snowball Earth theory received a boost in August
1998 with Harvard geologist Paul Hoffman’s publication, in Science, of new
evidence that ice extended to near equatorial latitudes in the late Precambrian,
about 700 million years ago.
The more recent glaciations, those that occurred since skeletons
evolved about 550 million years ago, affected only land regions; except for an
increase in icebergs, or at most ice cover near the continents, the oceans remained
open. Such may not have been the case in the Precambrian glaciations.
During these two “Snowball Earth” episodes, all of the oceans may have
been covered with ice to considerable depths. And although the deeper regions
of the seas remained liquid, thick icebergs, or pack ice to depths of 500
to 1500 meters, may have covered the ocean. Earth would have been cold indeed.
Average surface temperatures on the planet would have varied between
20°C and 50°C.
These extremely cold temperatures would have had an enormous influence
on the surface of our planet. For example, continental weathering would
have slowed or even stopped. In the interior of continents, the covering of ice
would eventually ablate (evaporate) away, just as it does in the dry valleys of
Antarctica today, leaving behind a sterile rock surface. Dust from these regions
would be blown out to sea, making the pack-ice cover of the oceans
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brown from terrigenous material. From space, Earth would have looked white
and brown—the white being the ice covers on the oceans, the brown the denuded
land areas.
The presence of the pack ice covering the sea would act as a lid on a
pot. Normally, much free exchange occurs across the vast interface of ocean
and atmosphere. Water evaporates from the sea into the air and then rains
back into the sea. If the sea were covered with ice, however, the ocean and
the atmosphere would become “decoupled.” Chemical changes in the ocean
would be separated from the atmosphere by the kilometer-thick lid of ice on
the ocean surface. Very drastic chemical change could—and according to
Kirschvink and others, did—occur within the sea itself.
Even with the icy cover, volcanism would have continued both on the
land’s surface and along the mid-ocean volcanic ridges at the bottoms of the
world’s oceans. At such sites today (see Chapter 1), great volumes of metalrich
fluids gush forth from these submarine volcanoes. In a covered ocean,
this material would have become toxic, producing what are known as reducing
conditions. The oceans would have begun to accumulate with metal ions,
mainly iron and manganese. For as long as 30 million years, the glaciers and
ice never relaxed their frigid grip on the planet’s surface.
All of this global cold would surely have adversely affected life in the
shallow-water regions of all the world’s oceans. The biosphere became restricted
to a narrow belt around the equator and to deep-sea hot springs and
hydrothermal vent settings. Perhaps some life also survived in occasional
Yellowstone-like hydrothermal systems.
Astronomers once thought that a previously warm world’s descent into
such an “icehouse” or “snowball” would be irreversible. Their reasoning was
that as a planet gets more and more thickly coated by ice, the fraction of sunlight
reflected back into space increases and solar heating of the surface declines.
On Earth today, sunlight is adsorbed by the darker land and seas but
is reflected into space by cloud cover. A planet completely covered with ice
would reflect most sunlight into space, causing the planet to become ever
cooler. Yet it is clear that Earth was able to escape from the deep freeze—not
once but several times. The means of that escape was through the volcanic
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emissions of greenhouse gases such as carbon dioxide into the atmosphere,
producing a “greenhouse effect.”
E S C A P E
As we saw in Chapter 2, a planet’s average temperature is greatly affected by
the volumes of greenhouse gases in its atmosphere. Much of this gas enters a
planet’s atmosphere from actively erupting volcanoes. Although there are
abundant volcanic eruptions in the sea as well, most of the carbon dioxide
from these events does not make its way into the atmosphere. Cold seawater
can hold large amounts of dissolved carbon dioxide, and below 700 meters,
CO2 will settle to the bottom of the ocean as it reaches saturation in the
water. At the time of Snowball Earth, enough CO2 would eventually reach
the atmosphere to melt back the sea ice and, in so doing, expose the metalrich
waters of the sea to the atmosphere. The time necessary for this “meltback”
has been estimated by Hoffman and his group to be between 4 and 30
million years. With the ice melted back from the sea, and temperatures again
warming, Earth would have undergone spectacular changes. Here is how
Kirschvink has described these events:
Escape from this “icehouse” condition was only accomplished by
the buildup of volcanic gases, particularly carbon dioxide, mostly
from undersea volcanic activity. Deglaciation during the end of
these glacial events must have been spectacular, with nearly 30
million years of carbon dioxide, ferrous iron, and long buried nutrients
suddenly being exposed to fresh air and sunlight. Hundreds
of meters of carbonate rock are preserved capping the glacial sediments,
at all latitudes, on all continents, as a direct result of wild
photosynthetic activity. For a brief time, the Earth’s oceans would
have been as green as Irish clover, and the sudden oxygen spikes
may have sparked early animal evolution.
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The most important source of biological productivity in the oceans of
today derives from the growth of phytoplankton, the single-celled plants that
are the pastures of the sea. The growth of these plants, so important for producing
oxygen, is limited by the availability of nutrients and iron. If iron is
dropped into the oceans of today, a great bloom of phytoplankton results.
Such was probably the case soon after the end of the first Snowball Earth
event. As the ice-covered seas began to melt, the fine iron- and magnesiumrich
dust coating the surface of the sea ice would have acted as a fertilizer,
tremendously stimulating growth of the blue-green “algae” (really photosynthesizing
bacteria known as cyanobacteria). Enormous populations of
cyanobacteria would have clotted the surface regions of the liberated seas, releasing
huge volumes of oxygen as a consequence of their photosynthetic activity.
This sudden appearance of so much life, after the millions of years of
cold and dearth of life, would have been a great revolution, and it probably
stimulated new evolutionary changes.
These events would have had profound geological as well as biological
ramifications. The sudden rush of oxygen into the sea and air would
have caused the iron- and manganese-rich oceans to precipitate out iron
and manganese oxides. In a previous chapter we saw how banded-iron deposits
began to accumulate about 2.5 billion years ago. Kirschvink and his
group argue that the appearance of banded-iron deposition occurred soon
after the first Snowball Earth ended. Not only iron deposits but magnesiumrich
deposits as well were immediate results of the end of the first Snowball
Earth event. Evidence of this is seen in South Africa, where the world’s
largest land-based deposit of manganese minerals has been dated at 2.4 billion
years of age and sits just above sedimentary deposits that were laid
down during the 2.5-billion-year-old Snowball Earth episode. Like the
banded-iron formations, these manganese-rich deposits appear to be a direct
consequence of the oxygen bloom that occurred when the planetary
snowball melted.
The cessation of the 2.5-billion-year-old Snowball Earth thus appears
to have resulted in a rise in the amount of oxygen both dissolved in the sea
and free in the atmosphere. Probably for the first time in Earth’s history, the
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sunlit portions of the sea became too oxygen-rich to allow iron to exist in solution
in seawater. Kirschvink and his colleagues argue that this dramatic
change in the chemistry of the sea would have exerted intense evolutionary
pressure on life on Earth, then no more advanced than prokaryotic bacteria.
Oxygen, indispensable to the survival of animals, was at that time a poison to
perhaps the majority of life forms. Having evolved in environments with little
or no oxygen, most life experienced the sudden appearance of the chemically
reactive element as a global disaster—but for the rest it was a powerful
evolutionary spur. There were but two choices facing life on Earth in that
long-ago time: Adapt through evolution, or die.
All organisms in the sea had to adapt in two major ways. First, they had
to evolve enzymes capable of mitigating the ravages of dissolved molecular
oxygen and chemicals called hydroxyl radicals. (We humans are still trying to
do this. Our ingestion of antioxidants such as vitamin E and vitamin C is an
attempt to reduce the ravaging effects that dissolved oxygen and “free radicals”
have on living cells.) Second, with the banded-iron formations’ precipitation
out from seawater, living cells no longer inhabited a solution rich in
iron. After having been surrounded by high-iron solution since the first formation
of life, proteins within cells had to be reengineered for life in an environment
low in iron.
Recent DNA sequencing has shown that several enzymes found in archaeans
and eukaryotes are left over from this event of 2.5 billion years ago.
No such enzymes occurred in the older bacteria. The implications of this are
profound: Kirschvink and his colleagues are proposing no less than complete
rejection of the Tree of Life models we examined at the end of Chapter 3,
which suggest that the three great domains (Archaea, Bacteria, and Eucarya)
all arose soon after life’s first evolution at least 3.8 billion years ago. The new
study has not only uprooted this tree; it has burned it. If the Kirschvink group
is correct, two of the three domains—Archaea and Eucarya—arose only after
the 2.5-billion-year-old Snowball Earth and are thus much younger than the
bacteria. Soon after this, in rocks about 2.1 billion years of age, we find a
record of the oldest organelle-bearing eucaryan—the creatures known as
Grypania, which we mentioned in Chapter 3.
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This new version of the Tree of Life is a revolutionary scientific discovery,
and if true, it will utterly reshape our understanding of life’s evolutionary
path. The Snowball Earth events can be seen as biologically important in two
ways. First, the inception of the Snowball produced what may have been the
largest “mass extinction” (the subject of Chapter
The persistence of globally freezing temperatures, the isolation of the ocean
from sunlight, the change in the precipitation patterns on Earth, and the removal
of all water from the surfaces of continents would have removed the
majority of surface habitats then available for microorganisms. In only a few
places could microorganisms have survived: in the deep earth, around hot
springs, and in hydrothermal deposits. Second, Earth’s release from this icy
prison after 30 million years brought about a new catastrophe: from cold to
hot, from oxygen-free to oxygen-rich. Again, organisms had to adapt rapidly.
It is this legacy that we may be seeing in the DNA of all living organisms;
those that survived all bear witness in their DNA to this dual catastrophe—
first cold, then warmth and oxygen. Life on the early Earth went through an
icy bottleneck, and it came out the other side radically changed.
The Snowball Earth of 2.5 billion years ago may have given our planet
eucaryans and the eukaryotic cell necessary for animal life. The second series
of Snowballs (there were several in rapid succession) may have bequeathed
our planet an even more interesting biological legacy—animal life as we
know it.
T H E S E C O N D GL O B A L GL A C I A T I O N
As we saw in Chapter 5, by the next round of Snowball Earth events, those
spanning the time interval from 800 to 600 million years ago, animal life was
present on Earth, but it was newly formed. Either simultaneously with or
soon after the appearance of the new animal phyla, Earth was once more
locked into a global icehouse. Once again, there must have been a period of
mass extinction, as the warm planet froze and the heat-loving organisms of
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Earth had to retreat to oases of heat, such as around volcanoes and hydrothermal
vents, or die. Yet the very severity of these events may have benefited
the newly arisen animals. The great stress inflicted by environmental
conditions imposed by the Snowball events would have stimulated inordinately
rapid evolution among the newly evolved animals. It would also have
caused the isolation of various populations, because the small populations of
life huddled around the undersea volcanoes would have been cut off from any
exchange of genes with other animal groups. This very isolation may have
been largely responsible for the diversity of phyla that emerged at the other
end of these crises, for when the final Snowball Earth event ended, about 600
million years (or less) ago, an entirely new group of creatures was ready to
take over the planet. This is the interval when animal life began to diversify
dramatically, in an event known as the Cambrian Explosion, the subject of
the next chapter.
Would this have happened if the glaciations had not occurred?
Kirschvink and Hoffman suggest that there is a causal link between the cessation
of these major glaciations and the emergence of animals. Hoffman has
noted, “Without these ice events, it is possible there wouldn’t be any animals
or higher plants.” He believes that the melting of the ice at the end of these
ice ages boosted biological productivity—and in the process stimulated evolutionary
activity. This idea has yet to be confirmed, but it remains a tantalizing
possibility.
Both of the two great episodes of Snowball Earth nearly ended life on
Earth, as we know it. But each, ultimately, may have been crucial in stimulating
the great biological breakthroughs necessary for animal life: the evolution
of the eukaryotic cell and then the diversification of animal phyla. This leads
us to ask whether Snowball Earth events are necessary to produce animal life as
diverse as that seen on Earth today.
The end of the last Snowball Earth event brought the time interval
known as the Precambrian to a close. Soon thereafter, abundant skeletons of
larger animals began to fill the sea, in the Cambrian Explosion. If the two
groups of scientists led by Joseph Kirschvink and Paul Hoffman are correct
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about Snowball Earth, a good case can be made that life on Earth is to some
extent due to these events.