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Geology and the Fossil Record
Much of what we know about life's origins has come from the fossil record,
and the study of how the earth has changed over time.
The Fossil Record
Sedimentary rock forms as layers of minerals settle out of water. Dead
organisms are buried in these sediments, so the oldest fossils are found in
the oldest (which are often the deepest) sediments. But how do we tell
the age of the fossils?
Relative Dating
Because more recent sediments cover older ones,
the strata (layers) of sedimentary rock present a record of organisms
that have existed through time.
- Strata at different locations can be matched by the presence of
similar organisms known as index fossils.
- animals with hard body parts make the best fossils:
- shells (the best index fossils, as they are numerous and widespread)
- bones (less numerous)
- This method does not allow a fossil to be dated in years, but only set into a relative time scale of existence compared with other fossils.
- The Geological Time Scale was devised by
geologists studying the fossils in
the various strata at different locations.
Absolute Dating
Unlike relative dating, absolute dating of fossils involves establishing a discrete age (in years) to the fossils under study. This is generally done via radiometric dating in which the half life of a radioactive isotope is used to calculate how long the substances in the fossil have been decaying. Half life is the time it takes for 50% of the original radioactive sample to decay to its stable form. It is not
affected by temperature, pressure, or other environmental variables.
Hence, radiometric dating is a reliable way to gauge the age of fossils. Examples:
Carbon Dating
- Devised by W.F. Libby and his colleagues at the University of Chicago, carbon dating is used to estimate the age of relatively "young" objects, from a few hundred years old to about 50,000 thousand years old.
- In the atmosphere, free neutrons generated by cosmic radiation constantly react with atmospheric nitrogen to form Carbon-14:
147N + 10n --> 146C + 11H
- Hence, atmospheric carbon exists as both stable 12C and as
radioactive 14C in the form of CO2.
- The relative concentration in the atmosphere of of 14CO2 to 12CO2 is about 1 : 10,000,000,000,000.
- Living organisms constantly take in CO2, and so have the same 14C:12C ratio as the atmosphere while they are alive.
- Once they die, however, they stop taking up atmospheric CO2, and the 14C in their tissues begins to decay back into stable 14N.
- Because 14C decays at a constant, known rate, a measurement of the proportion of 14C to 12C in an organic object will yield that object's age.
- The half life of 14C is 5730 years, a relatively short
time. Hence, this isotope is useful primarily for dating relatively recent
fossils.
- Here's how to calculate carbon-14 decay.
- But if you're feeling lazy, you can just use this cute Carbon Dating Calculator to figure out the age of your fossil.
- Because of 14C's relatively short half life, paleontologists use isotopes with longer half lives to date older fossils.
Potassium-Argon Dating
- Radioactive parent isotope 40K decays to daughter isotopes 40Ar and 40Ca over a half-life of 1.26x109 years.
- Because 40Ca is the most common form of calcium, increase in its abundance due to 40K decay is less useful as a measurement of a sample's age.
- The 40Ar is much more rare. Hence, increases in its abundance are easier to measure, making it more useful for radiometric studies.
Uranium-Lead Dating
- Uranium-lead is one of the oldest and most refined radiometric dating methods.
- Uranium-lead dating couples the decay of 238U to 206Pb (half-life = 4.47 billion years) with the decay of 235U to 207Pb (half-life = 704 million years) to construct a geochronometer with a range of 1 million - over 4.5 billion years with accuracy to 1 - 0.1%.
- 238U does not occur in live organisms, but is found in
igneous (volcanic) rock.
- A paleontologist can
date fossils found in close proximity to igneous rocks by measuring their
concentration of 206Pb.
Amino Acid Racemization
- Isomers of amino acids (L- and D-, which stand for "levo" and
"dextro") also can be used to date fossils.
- Since living things
make only L-amino acids, they don't contain detectable amounts of the D
isomer while alive.
- Once they die, the L-form slowly converts to the
D isomer (a process termed racemization).
- Measurement of the proportions of L- and D- amino
acids in a fossil can yield a age for relatively recent (tens of thousands of
years) fossils.
- PROBLEM: Racemization is affected by climate, so it's reliable only in areas
(e.g., the tropics) where the temperature and other aspects of climate
have not changed much since the fossils of interest were deposited
there.
Biogeography
The continents did not always exist as they are today. They once were
present as a single land mass called Pangaea. This split into northern
(Laurasia) and southern (Gondwanaland) land masses that then underwent
further fragmentation. This process is known as continental drift. You can see the
physical manifestation of abutment of the continental plates in some areas, such as
the San Andreas Fault.
- subduction zone - area where the edge of one plate is being forced
underneath the edge of another one. Deep trenches form in these
regions.
- seafloor spreading zone - area where volcanic activity causes the
ocean floor to split and spread apart. Oceanic ridges form in these
areas.
Living organisms rode the continents as they broke up, and their
previously contiguous populations underwent allopatric speciation.
Phylogenies are linked to continental drift, with those land masses that
first broke away still having the most primitive organisms (not counting
human introductions of exotic species).
An excellent overview of the study of plate tectonics is once again provided by the University of California at Berkeley Museum of Paleontology.
The Origin of Life
Tracking the history of fossils and the changing continents tells us
something about how life has changed over time, but the big question still
remains...
How did life originate? We can't go back and watch. But many have
contributed pieces of the puzzle, which has yet to be completed.
In 1924, Russian biologist Aleksandr (Alexander) Oparin published a paper entitled
The Origin of Life, in which he suggested that chemical reactions in
the primitive oceans could have eventually "created" life. The work was never translated from Russian, it had little
impact at the time.
In 1929, British biologist J.B.S. Haldane published
similar ideas, but still, they were still essentially ignored.
In 1936 Oparin published his ideas--first described in his paper--as a
book entitled The Origin of Life. This was translated into several
languages (including English), was widely read...and the race was on!
The Primordial Ooze
Earth was formed 4.5 - 5 billion years ago.
Earliest life on earth appeared between 3.5 - 4 billion years ago.
The oldest known fossils are stromatolites--sedimentary rock with
striations very similar to those made even now by extant cyanobacteria.

What was Primordial Earth like?
The early atmosphere was composed mostly of
- carbon dioxide
- water vapor (for perhaps its first billion years of its existence,
earth was too hot for water to remain in liquid form)
- hydrogen
- nitrogen
- ammonia
- hydrogen sulfide
- carbon monoxide
- methane (the simplest organic molecule)
(NOTE:
The latter six are Greenhouse Gases, responsible
for the Greenhouse Effect.
Don't confuse the Greenhouse Effect (Our Friend) with the anthropogenic
Global Warming (Not such a Good Pal), which has been caused by the quick
release of vast quantities of CO2 previously trapped in fossils and
non-fossilized organic matter.)
Eventually, sufficient cooling allowed pockets of liquid water
to persist, and over the millenia, the oceans were formed. Still, there
was no oxygen in the atmosphere.
The only organisms were obligate anaerobes, whose only metabolic
pathway was fermentation.
The Origin of the Oxidizing Atmosphere: Chlorophyll a Appears
There was no oxygen in the Earth's atmosphere until photosynthetic organisms appeared.
- chlorophyll a (the most primitive form of chlorophyll) first appeared
about three billion years ago, in the early ancestors of the
cyanobacteria.
- about one billion years ago, the first eukaryotic autotrophs
appeared. These were early forms of green algae which looked very much
like an extant species (Chlamydomonas).
- It took about 1.2 billion years of photosynthesis by these organisms
to oxidize all the iron in the earth's crust that was exposed to the
atmosphere. Once that was done, oxygen could begin to enter the atmosphere.
- by the beginning of the Cambrian (0.6 billion years ago), oxygen was present
at about 1% of PAL (PAL = "Present Atmospheric Level") due to the
photolysis of water by the unicellular autotrophs (cyanobacteria and green
algae)
- This was enough oxygen to result in two changes:
1. fermentation gradually gave way to cellular respiration as the major metabolic pathway.
(obligate anaerobes are fried by oxygen)
2. the generation of a protective ozone (O3) layer in the upper
atmosphere allowed the upper levels of the ocean to be colonized by small
organisms, now shielded from deadly ultraviolet (UV)
and gamma radiation.
- By 0.4 billion years ago, when land plants were present, oxygen was about
10% PAL, and it just kept creeping up. Today's atmosphere is about 21% oxygen.
How did the first cells appear?
1953 - Stanley Miller (then a graduate student) and Harold
Urey (then his major professor) at the University of Chicago built an apparatus
that simulated the conditions of the primordial oceans.
Analysis of the reaction solution yielded...
- amino acids (glycine, alanine, methylalanine, glutamate,sarcosine)
- formic acid
- acetic acid
- urea
Analysis of gas portion yielded
- hydrogen cyanide
- formaldehyde
- other organic volatiles
Altering proportions of gases gave similar results,
as long as there was no free oxygen, which would oxidize everything. (Remember: there was no oxygen in the primordial atmosphere.)
Later experiments with the apparatus yielded...
- sugars
- lipids
- thymine, guanine, adenine, cytosine and uracil (which are the
components of...)
Meanwhile, back in Russia, Oparin was working with
another aspect of the origin of early life.
Recall how shaking oil and water together can create
a tiny interface layer of bubbles! Other substances can do this,
too.
Oparin mixed gelatin (a protein) and gum arabic (a
polysaccharide) and stirred them with a motion mimicking that
of the early seas.
The results: tiny, stable, globular structures. He called these coacervates.
Adding additional substances to the reaction mixture yields different
results...
- Add lipids, and these tend to coat the coacervates
in a membrane-like sheet, adding stability.
- Add enzymes and substrates, and these are
incorporated into the inside of the coacervates and begin to function normally
A coacervate with working enzyme
systems inside can be called a protobiont.
A protobiont has some of the properties of life, depending on what was
added to the mix, but not all. To be considered truly alive, a thing must have...
1. organized structure (anatomy)
2. chemical reactions coordinated to perform vital functions (metabolism)
3. ability to maintain constant internal environment (homeostasis)
4. reaction to stimulus (internal and external)
5. growth and development
6. ability to adapt to environmental changes
7. ability to reproduce
Early Genetic Material
It is generally believed that RNA was the original
genetic material
- it is simpler to make from raw materials
- retroviruses exist that use RNA as original template,
and sequester the host cell's machinery to perform reverse transcription
(RNA-->DNA)
- self-splicing RNA (ribozyme) can snip and splice
some introns out of mRNA before translation, and may have functioned as
early enzymes
However, comparatively stable DNA might have conferred a tremendous selective
advantage over RNA. Almost all known living organisms use it as their
permanent genetic blueprint.
Of course, there are some who remain unconvinced.
Origins of Life and Metabolism: Prokaryotes
The term "prokaryote" is more descriptive than phylogenetic, as the lack of
a characteristic (in this case, a membrane-bounded nucleus) isn't a very
good basis for classification. What is the meaning of
the word "prokaryote?"
pro =
karyon =
What is the meaning of "eukaryote"?
eu =
karyon =
PROKARYOTES...
They're everywhere.
Formerly, all prokaryotes were lumped into "Kingdom Monera."
Now, they are classified into distinct lineages:
- Domain Bacteria ("true" bacteria)
- Bacteroides
strict anaerobes, these are a major component of
normal intestinal flora in many vertebrates. Bacteroides
fragilis is the most common species. These bacteria are
beneficial as long as they stay in the gut: they help digest food,
provide essential nutrients, and help keep harmful bacteria in check
(via competition). They can cause abscesses when they are introduced
into other tissues, however. This can be problematic, as they readily
evolve resistance to antibiotics.
-
Cyanobacteria (Blue-green bacteria)
These are the earliest photoautotrophs. Using the photosynthetic pigments
chlorophyll a, phycoerythrin and phycocyanin, they capture sunlight and store
its energy in the bonds of sugar via photosynthesis.
- Flavobacteria
Relatively new group, this one is still being
divided into subtaxa
- Gram-positive Bacteria
These stain darkly with Gram Stain due
to the special anatomy of the cell wall, rich in peptidoglycan.
- Green Nonsulfur Bacteria
Once classified as "gliding bacteria," these form flexible filaments that
allow them to move with a gliding motion. They are anoxygenic phototrophs
with bacteriochlorophylls c or d, and small quantities of chlorophyll
a.
- Green Sulfur
Bacteria
Apparently closely related to the Bacteroides group, these bacteria may
be rod-shaped, spherical, or spiral. They may form stalks with which to
attach to substrate. Photosynthetic, they use bacteriochlorophylls c, d,
or e, and have small quantities of chlorophyll a.
- Purple Sulfur
Bacteria
Anoxygenic phototrophs
with bacteriochlorophylls c or d, and small quantities of chlorophyll
a. These are colorful, and may express purple, yellow, orange, and red
pigments. They make spectacular sludge.
- Spirochaetes
These are large, spiral-shaped bacteria. Some species are zoonotic pathogens (i.e.,
they can be transmitted between animal species, including humans).
Various species of spirochaetes are responsible for such well-known
diseases as Lyme Disease, syphilis, leptospirosis, and other nasty
illnesses. Other species are normal symbionts in the intestines of
ruminant animals such as cattle.
- Thermatogales
- Domain Archaea (archaebacteria)
- Not yet subdivided into phylogenetic groups, but they can be
classified on the basis of their metabolic oddities:
- Their metabolisms don't necessarily imply evolutionary relatedness.
Stay tuned for more information as new research reveals where they came
from.

Note that the phylogenetic tree shows "Domain Bacteria" as branching from
the universal ancestor first, and sister taxa Archaea and Eukarya branching off
"next".
Remember that the entire tree can be swiveled at any node, and it would be
just as correct to have Archaea and Eukarya at the left of the diagram.
This might be less confusing to some, as the very first fossil organisms
are nearly indistinguishable from archaebacteria.
Both Archaea and Bacteria are considered to be "prokaryotic"--pre-nuclear.
They lack membrane bounded organelles or nuclei, though many do have
internal membrane systems.
Archaean-like prokaryotes were the first inhabitants of earth,
and spent the first 2 billion years after their appearance alone on earth.
A huge diversity of Bacteria exists today
Bacteria are both primitive and highly successful.
(Remember: primitive does not equal inferior)
The divergence of prokaryotic taxa happened so long
ago that it may not be possible to determine true evolutionary
relationships.
Prokaryotic Structure and Function
Prokaryotes may be unicellular, aggregate or colonial.
Some of the more derived species may have colonies with a division of
labor among cells (analogous to multicellularity in "higher" organisms).
Bacteria may be classified by their shape:
- round (cocci)
- rod-shaped (bacilli)
- helix (spirillae and spirochaetes - superficially similar
only)
- A clustering bacteria may be given the prefix "staphyl-" (as in
Staphylococcus spp.
- A link-forming bacteria may be given the prefix "strept-" (as in
Streptococcus spp.
...but Bbacterial shape may not reflect phylogenetic relationships, as there may be
convergence in shapes (e.g., spirilla & spirochetes), as well as a variety
of shapes in closely related bacteria. But shape is useful for
grouping and identifying on a gross level.
Bacteria range in size from 1-5 micrometers--much smaller than most
eukaryotic cells (100-1000 micrometers).
Bacterial genome consists of a single, circular chromosome of
double-stranded DNA. This can be very large, and is organized in the
nucleoid region of the cell.
The average bacterium has about 1000 genes on this nucleoid c'some.
a plasmid is a small, circular piece of DNA, and it may contain only
a few genes. This replicates autonomously, and is not considered part of
the bacterium's own genome. However, it may confer phenotypic traits on the bacteria containing it.
Genes on plasmids may confer antibiotic resistance and other mutant characters on bacteria that have them. (For example, some
bacterial species are not toxic unless they have a plasmid allowing them
to manufacture specific toxins (e.g., Clostridium spp.)
Another diagnostic character is the nature of the cell
wall, which is present in most bacteria. (Exception: mycoplasmas, all of
which are intracellular parasites) Two major types of cell wall can be
distinguished with
Gram Staining.
- The major difference is the amount and location of a
mucopolysaccharide known as peptidoglycan.
- Peptidoglycan forms a thick,
rigid layer in both Gram positive (G+) and Gram negative (G-) cells.
It composed of an
overlapping lattice of two sugars crosslinked by amino acid "bridges".
The exact molecular composition of peptidoglycan layers is species specific.
- The two sugars are N-acetyl glucosamine (NAG)
and N-acetyl muramic acid (NAM). NAM is found ONLY in
bacterial cell walls--nowhere else.
- Gram Positive: stain dark with Gram method
- G+ have a very thick external layer of peptidoglycan
- Gram Negative: do not stain with Gram method (appear pink from
safranin counterstain)
- G- have a thin layer of peptidoglycan sandwiched between two plasma
membranes (inner and outer)
- Gram staining properties are linked to pathogenicity, with G-
often being more dangerous than G+.
- Because the peptidoglycan layer of G- is protected by a plasma
membrane, it is less susceptible to attack by antibiotics which
interfere with the formation of peptidoglycan amino acid "bridge" bonds (e.g., penicillins)
- A pilus is a surface extension from a bacterial cell that may act as a
bridge between bacteria (for exchange of genetic material), or a means of
attachment to a substrate or a host cell.
- Many bacteria are motile (they can move), and the means of
locomotion is another way to identify them.
- flagellum (composed of unique protein known as flagellin)
- Note the difference between the prokaryotic and eukaryotic flagella:

- gliding (on a secreted slime trail)
- taxis means "movement. Bacteria exhibit various forms of
positive or negative taxis, depending on species and specific
environmental conditions. (e.g. phototaxis, chemotaxis, etc.)
- External to the cell wall, some species have a gel capsule that is
often protective against predators (or a host's immune system).
Bacterial Reproduction
Bacteria may reproduce asexually:
...or sexually
Bacterial cells can be grown in culture on appropriate nutrient media
(usually agar with broth added). A bacterial colony on an agar plate is
called a lawn, and various species have characteristic lawn phenotypes.
This can be very useful in the study of bacterial genetics!
Some species can form environmentally resistant structures called
endospores, which is little more than the chromosome surrounded by a thick
wall. It's nearly impossible to kill an endospore, so pathogens that can
form them are particularly pernicious!
Competition exists at even the microscopic level. Antibiotics are
substances that inhibit the growth of prokaryotic cells. They are
manufactured not only by plants and fungi, but also by some bacteria.
Metabolic Diversity of Prokaryotes
Three basic types of organisms re: oxygen tolerance/metabolism
- oblicate anaerobe - can do only fermentation; killed by oxygen
- obligate aerobe - operates primarily on aerobic metabolism;
cannot survive long without oxygen.
- facultative anaerobe - can do either aerobic
respiration or fermentation, depending on environmental conditions.
In the Krebs Cycle, the terminal electron acceptor can be
- oxygen (in aerobes)
- nitrate or nitrite (in denitrifying bacteria, which
return nitrogen gas to the atmosphere)
- sulfate (in sulfur bacteria)
Four Main Categories of Prokaryotic Energy Transduction
- Photoautotrophs
- photosynthetic
- use CO2 as carbon source
- use light as Energy source
- cyanobacteria are the most common in this group
- Chemoautotrophs
- chemosynthetic
- use CO2 as carbon source
- use inorganic compound oxidation as Energy source (e.g.,
H2, ammonia or iron ions)
- many archaebacteria fall into this category
- Photoheterotrophs
- photosynthetic
- use organic molecules as a carbon source
- use light as energy source
- relatively few bacteria in this group
- Chemoheterotrophs
- heterotrophic
- obtain energy from organic compounds
- vast majority of bacteria are in this category, along with most
protists, animals, fungi and some plants.
- a saprobe is a chemoheterotroph that breaks down decaying organic matter for energy.
- a parasite is a chemoheterotroph that uses the organic
molecules of living tissue for energy
How did it all begin?
Early hypotheses about the origin of bacterial metabolism suggested that the earliest cells used ATP from the "primordial soup".
Problem: it's not likely there was enough ATP out there to fuel those newly made cells. ATP is highly
unstable, and won't remain in solution for long.
More plausible is the idea that CO2 was the first Carbon
source, and that early cells had plasma-membrane anchored enzymes that
could oxidize inorganic compounds to make the energy needed to drive
synthesis of carbon compounds.
Ecological Importance of Prokaryotes
Along with fungi, they are the biosphere's main decomposers.
Many are symbiotic
- parasitic (pathogens, opportunistic and not)
- mutualistic
- commensal
The Importance of Nitrogen Metabolism
Let's have a look at the Nitrogen Cycle 
And here are some of our pals who perform this marvel:

Nitrogen-fixing bacteria in the roots of a
leguminous plant
Nitrogen fixation - conversion of atmospheric nitrogen
to nitrogen compounds (ammonia, nitrite or nitrate) that are usable
by plants
Denitrification - nitrate-->nitrite-->ammonia-->N2
Nitrosomonas spp. - converts
ammonia to nitrite
Pseudomonas spp. - converts
nitrite or nitrate into N2 (denitrification)
(Note: Pseudomonas (e.g., aeruginosa and cepacia) are
often opportunistic pathogens.
A pathogen is a disease-causing agent.
Although Pseudomonas are very common in the environment, they usually do not cause disease.
However, in an immunocompromised animal, they can proliferate,
creating a very difficult-to-treat infection. Many strains are
resistant to a wide variety of antibiotics.) Hence, they are opportunistic pathogens.
Prokaryotes as Pathogens: Koch's Postulates
Everyone knows that certain bacteria can cause disease in plants and animals.
In order for a microorganism to be declared the cause of a particular
disease, however, it must meet the following criteria (Koch's Postulates):
- it must be isolated from a diseased individual
- it must be grown in pure culture from that sample
- it must cause the disease in a healthy individual when introduced
from that culture
- it must be isolated from the newly infected individual
Some pathogens don't meet the postulates, so clinicians must be a
little bit flexible when trying to determine how to treat difficult-to-culture pathogens.
How do bacteria cause diseases?