EES 227: Paleobiology
Spring 2004
return to
syllabus
Lecture 6: February 12 2004:
Darwin
Day
Reading:
Web resources for this lecture:
Lecture Notes: Evolutionary
Theory.
Before Darwin:
A popular view of nature was
'Natural
Theology' (e.g., Linnaeus
), as contrasted with 'revealed
theology' where insight in the nature of the deity is arrived at
without observations. Natural Theology
was very popular in the 1700s, and implies:
- God made the world and all its
creatures
- One can learn about God by studying
Nature, his creation
Natural theology:
- No extinction has occurred: an
omnipotent benevolent God would not allow His creatures to go
extinct ('Divine Benevolence'): Many European explorers were
looking for organisms that we now consider extinct (e.g.,
President Jefferson argued that many large animals could be alive
in the American West).
- God had created a complete and perfect
nature ('Fullness of Nature')
- All of nature formed a 'Great
Chain of Being' linking animals
to man to angels to God. God, representing perfection in the
highest degree, was at the top of the chain, which in its intirety
represents all degree of perfection or lack thereof from the
highest and fullest to the lowest and least, with 'nothingess' at
the bottom of the chain. In images, this concept survived long
after acceptance of the fact of evolution in the 'amoeba-to-man'
type of figures (see text book 'Tree of Life' figure). Note that
in 'amoeba to man' figures there really is no place for the
protostomes, a large branch of organisms not in line between
amoeba and man (deuterostomes).
But from the 1700s on: decreasing believe
in Bible as literally true
- Voyages of exploration: many species;
how would they have arrived where they are from Mount Ararat?
Especially the species that do not live between Ararat and where
they do live (e.g., kangaroos) ?
- Fossils: many of organisms that were
extinct (not known from present world); record too complex for
just one flood.
- Age of the Earth under discussion, but
accepted as being more than a few thousand years
Lamarck (1744-1829):
- Species not fixed
- Did believe in 'Great Chain of Being',
'Ladder of creation': lower beings were continuously created from
mud ( they 'climb the ladder' until they reach the level of
angels.
- Lamarck did NOT see life as a branching
tree, although he did not see species as fixed
entities.
NOTE: Pasteur
lived 1822-1895; the argument on the theory of 'spontaneous
generation' (origin of life from
non-life) ran from 1668 through 1859 when Pasteur did his winning
experiment. For insatnce, a seventeenth century recipe for the
spontaneous production of mice required placing sweaty underwear and
husks of wheat in an open-mouthed jar, then waiting for about 21
days, during which time it was alleged that the sweat from the
underwear would penetrate the husks of wheat, changing them into
mice.
Debate started seriously in 1668 by
Francesco Redi (no maggots on meat covered with fine netting, in
closed jar). But invention of microscopy seemingly supported
spontaneous generation, revealing a whole new world of organisms that
appeared to arise spontaneously. It was quickly learned that to
create "animalcules," as the organisms were called, you needed only
to place hay in water and wait a few days before examining your new
creations under the microscope.
Lamarckianism (or 'adaptation on
demand'); what we now call Lamarckianism was really only a very small
part of Lamarck's book.
- Changes in the environment
=>
- changes in the physical needs of
organisms =>
- changes in behavior of organisms
=>
- specific organs are used
more/less/differently =>
- changes in the structure of these organs
=>
- such changes are heritable
Note that organisms
themselves 'strive to change'. Darwinian ideas and modern dogma of
biology: efforts by parents not inherited by their
offspring.
Charles Darwin, 'The Origin of Species' (1859);
[1809-1872]
"I had two distinct objectives in view;
- firstly to show that species had not
been separately created, and
- secondly that natural
selection had been the chief agent of change."
Things that were NOT Darwin's
idea:
- Species have not been created
separately, but are related (discussion > 100 years, e.g.,
Darwin's grandfather)
- No divine creator (e.g.,
Buffon)
- The word 'evolution': Darwin used
'descent with modification'
Components of
Darwinism as Darwin knew it:
- Evolution occurred
- Common descent: all life on Earth has a
single ancestor
- Multiplication of species: number of
species increased during earth history
- Gradualism: evolution occurs
gradually
- Natural selection is main
cause
Natural selection:
- All individuals of a species are
slightly different
- These differences are
heritable
- Not all young of a species survive into
adulthood and reproduce
- The individuals who are best adapted to
their environment leave most offspring
- Individuals who leave most offspring
transfer their heritable characteristics into the next
generation
What Darwin did not know about:
- How character traits were passed from
parents to offspring
- Inheritance would be 'blended', i.e.,
that offspring would have mixed properties from father and
mother
- Mendel (contemporary of Darwin):
inheritance 'discrete' (not 'mixed')
Neodarwinism: 'The Modern
Synthesis':
- Darwinism (1859)
- Mendelian inheritance (1865
-1900)
- Julian Huxley: Modern Synthesis
(1920s-1950s)
- Population genetics (Fisher, Haldane,
Sewall Wright): mathematical models
- Genetics: Dobzhansky (fruit
flies)
- Population field studies: Mayr
(birds)
- Paleontology: Simpson
Afterwards modified: structure of genetic
material (DNA-RNA) 1953 (Watson & Crick)
Separation of body cells (somatic cells) and
reproductive cells (germ cells): Weisman's
barrier'. Later this was redefied as
the 'central dogma of biology': DNA => RNA => protein
(transcription)
Natural Selection and
Genes
- Natural selection does not (and can not)
affect the genotype of an organism
- Natural selection acts upon the whole
organism (called phenotype, i.e., how the organism looks
like)
- Some characters are defined almost
completely by the genetic make up of an organism, others by
environmental conditions
What is Evolution?
Oxford Concise Science
Dictionary: Evolution: "The gradual process by which the
present diversity of plant and animal life arose from the earliest
and most primitive organisms, which is believed to have been
continuing for the past 3000 million years." NOTE:
definition requires 'gradual': not accepted by everyone.
New Webster Encyclopedic Dictionary of the
English Language: "Evolution: ...that
theory which sees in the history of all things, organic and
inorganic, a development from simplicity to complexity, a gradual
advance from a simple or rudimentary condition to one that is more
complex and of a higher character". NOTE: definition
requires 'progress'; would exclude evolution of parasites that e.g.,
lack eyes from ancestors that have them
" Biological evolution ... is change in
the properties of populations of organisms that transcend the
lifetime of a single individual. The ontogeny of an individual is not
considered evolution; individual organisms do not evolve. The changes
in populations that are considered evolutionary are those that are
inheritable via the genetic material from one generation to the next.
Biological evolution may be slight or substantial; it embraces
everything from slight changes in the proportion of different alleles
within a population (such as those determining blood types) to the
successive alterations that led from the earliest proto-organism to
snails, bees, giraffes, and dandelions." (Douglas
Futuyma)
Definition MUST contain:
- Living organisms (thus excluding the
origin of life)
- Evolution works on populations, not on
individuals
- Heritable changes must be
involved.
Some Terms:
- Gene: hereditary unit that can be passed
on unaltered for many generations.
- Gene pool: the set of all genes in a
species or population.
- Allelles: variants of a specific gene
(e.g., eye color) within the gene pool
Organisms with a specific allele are doing
better than those with a different allele. The
allele whose possessor does better will become more and more common
in the population, because the organisms which carry the favorable
allele are leaving more offspring. Alleles that make their
possessor less able to reproduce than other organisms of the same
species will disappear from the total population.
What happens during evolution?
- Genes mutate
- Natural selection works on
individuals
- Populations evolve
Evolution: more definitions:
'Evolution is a process that results in
heritable changes in a population of organisms, which changes are
spread over many generations'. Note that this definition lacks
such terms as 'gradual', or 'to more complex forms'!
This contrasts with a much more limited
definition (Neodarwinian) of evolution as seen, e.g., in some high
school text books:
"In fact, evolution can be precisely
defined as any change in the frequency of alleles within a gene pool
from one generation to the next."
Many scientists, however, including many
paleontologists and embryologists would disagree with this
definition: "Evolution is change in gene frequencies"
- Irreversibility? such changes might be
reversed in another generation, not lead to formation of a new
species and reproductive isolation. (see the story of the
peppered
moth)
- Is generation of whole new groups of
organisms (e.g., mammals) just adding up of many small changes?
How do these changes in gene frequencies become transferred into
formation of reproductive isolation? How do we add new species
rather than transform a whole species into another?
- Is microevolution the same as
macroevolution?
What some people do not like about strict
Neodarwinianism (as e.g., argued for by Richard
Dawkins) and specifically the
statement that evolution is a change in gene frequency are the
following aspects:
- Reductionism: genes say all there is to
say about an organisms
- Panselectionism: natural selection of
individuals is the only cause of evolution.
- Extrapolationism: small scale evolution
is the same as large scale evolution.
- Gradualism: all evolutionary change is
gradual.
Why does evolution occur? What drives
evolution? What is the limiting factor on the rate of
evolution?
- New mutation may occur which helps its
possessors to leave more offspring than organisms without
it.
- Gene frequencies change because the
environment keeps changing.
- Gene frequencies change because the
organisms keep change in response to co-existing
species.
What drives evolution?
- Interaction with other organisms
(internal to the biota): "Red Queen hypothesis"
- Interaction with the environment
(external to the biota): good/bad luck
- But can biota and environment be
strictly separated??
Arguments against strict Neodarwinism from
within biology/genetics:
- Inheritance of acquired characters
revisited: Some processes change genes, changes inherited:
'jumping genes' in corn; lateral gene transfer (human-operated);
'reverse transcription'; human gemetically modified food; activity
of retroviruses.
- Neutralism: in panselectionism, EVERY
change in genes is positive or negative, however little, but never
neutral. BUT there are 64 possible different codons (sets of 3
bases in DNA and RNA) which code only for 20 proteins. Some
mutations thus appear to be selectively neutral.
- Structural and regulatory genes:
structural genes code for proteins. Many genes apparently do not
code for anything ('Junk DNA'). Regulatory genes: switch on and
off. Homeotic genes: major pattern of body plan (back/front; where
appendages occur).
- 'Hopeful monsters': mutations in
regulatory genes causes 'new species' in one step. Thoroughly
discredited in the past (Goldschmidt); now people look into parts
of Goldschmidt's hypothesis again.
Arguments against strict Neodarwinism from
within Paleontology.
- Paleontological
problem: the time dimension. Only morphological species can be
recognized. How do such species change over time?? IF species
change gradually over time, paleontologist can NOT really
distinguish species easily: they grade imperceptibly into each
other. One could only separate species if the record if really
incomplete: then a 'new species' seems to occur at each gap in the
record.
- In fact, paleontologists DO in
most cases easily recognize species over time. Over long times,
species do not appear to change significantly
in morphology, then there is rapid change, possibly splitting into
more species. Long periods of little change are called stasis,
short period of rapid change are called punctuations. Theory that
calls for alternation of periods of stasis and punctuations is
called punctuated equilibria (Eldredge & Gould, mid
1970s).
- If most species indeed originate by
allopatric
speciation, then this is the
pattern that we expect to see: new species originate in a small
region only.
Punctuated
Equilibrium:
- New species develop mostly by splitting,
not by evolving species
- New species develop rapidly (compared to
total 'species life')