The humpback whale Megaptera novaeangliae

The peacock, Pavo cristatus

Section 1: What is evolutionary biology?

Evolutionary biologists ask many kinds of questions:

Observations in nature: Medicine:

Why do whales have lungs? Where do new pathogens come from?
Why are peacock tails so fancy? Why do bacteria so often become resistant to
Why do snakes have no legs? antibiotics?
Why do we have an appendix, wisdom teeth?
Why do we age and die?
Genetics
Are human races different at the genetic level? How and why?
Are humans still evolving and changing genetically? Is this change rapid?
Why do some species (such as salamanders) have 10x the genome size of humans?

Sociology, Psychology, Anthropology

Why does every human (other than monozygotic twins) appear to be unique?
Why do humans differ across the globe? How did these differences develop?
Why do certain things universally taste good (sugar) or bad (soap)?
Why are there behavioral differences between men and women?
Section 1: What is evolutionary biology?

Proximate versus ultimate causation:

two approaches to ask causal questions in biology:

•  Proximate questions ask what is the immediate cause

of a trait, behavior, or process:
–  Proximate causes are observable and include the molecular,
physiological, and chemical mechanisms that shape
characteristics of organisms in nature.

•  Ultimate questions ask how a trait, behavior, or process

came to be over time:
–  Ultimate causes are the historical, evolutionary forces that shape
characteristics of organisms in nature.

•  Evolutionary biology focuses on Ultimate Questions

–  Examples: DNA replication structure, plant-pollinator interactions
•  http://www.ted.com/talks/jonathan_drori_the_beautiful_tricks_of_flowers.html
•  http://www.ted.com/talks/drew_berry_animations_of_unseeable_biology.html
Section 1: What is evolutionary biology?

•  What is known and what is not known: In evolutionary

biology (like most modern sciences) the unknown greatly
outweighs the known. In this class we will emphasize:

–  The processes that are used to by evolutionary biologists

to understand nature.

–  The hypotheses that biologists have made about the

origins and diversification of life.

All along the way we emphasize what is known and

what remains unknown.
Lecture Outline

2.    An  example  of  evolu+on  in  ac+on  

Section 2: Evolution in action

Pathogenic disease is important

•  Most people that have ever lived have died from microbial
infections.

•  Some biologists have estimated that 25% of all people that

have ever lived have died of one pathogen in particular
(Plasmodium, Apicomplexa)

•  In human pathogens the evolution of antibiotic

resistance represents a vexing problem for medicine
and is also a key example of rapid evolutionary change.
Section 2: Evolution in action

A tuberculosis ward at a U.S. Army base hospital in France during World War I

• Previous to 1940 most patients suffered from bacterial

infections. Deaths were caused by tuberculosis, pneumonia,
meningitis, typhoid fever, syphilis, cholera.
• After 1950’s: Most U.S. patients die of cancer, heart disease.
Section 2: Evolution in action

• Discovery  of  anRbioRcs  in  the  late  1920’s  and  development  of  
anRbioRc  therapies  in  the  1940s-­‐1950s  lead  to  the  near  conquest  
of  many  bacterial  diseases.  

• More  recently,  many  bacterial  strains  have  evolved  that  are  

resistant  to  some  of  our  most  potent  anRbioRcs.  

• Currently,  hospitals  are  witnessing  an  explosion  of  evoluRonary  

change  in  bacterial  resistance.  
• Example  Staphylococcus  aureus  (staph.)  ,  Neisseria  
gonorrhea,  Vibrio  cholera,  Clostridium  difficile  have  all  
evolved  resistance  to  mulRple  drugs.  
• SUPERBUG:  MRSA,  Methicillin  Resistant  Staphylococcus  
aureus  is  becoming  prevalent  in  hospital  seings  and  in  the  
community.  
Section 2: Evolution in action

Moraxella (middle-ear) infections in Finland

Section 2: Evolution in action

Evolu+onary  studies  oHen  begin  with  an  observa+on,  such  as  a  

paIern,  and  follow  with  ques+ons,  hypotheses,  and  experiments.  

• Do  anRbioRcs  cause  mutaRons  in  bacteria  that  make  them  

resistant  to  those  anRbioRcs?  
• Do  anRbioRc  resistance  mutaRons  occur  even  without  
exposure  to  the  anRbioRcs?  
• Are  mutaRons  present  in  populaRons  that  have  not  been  
exposed?  
• Can  we  prevent  the  evoluRon  of  anRbioRc  resistance?  
• Will  use  of  lower/higher  anRbioRc  doses  prevent  
resistance?  
• Will  use  of  mulRple  anRbioRc  drugs  prevent  resistance?  
• Does  the  faithful  use  of  an  anRbioRc  prescripRon  prevent  
the  evoluRon  of  drug  resistance?    
Section 2: Evolution in action

The  principles  of  evolu+on  have  been  used  to  address  some  but  
not  all  of  these  ques+ons.    

Key  results:  
1  Bacterial  mutants  that  are  resistant  to  anRbioRcs  occur  in  low  
frequency  in  most  natural  populaRons  of  bacteria.  
2  The  use  of  anRbioRcs  intensely  selects  for  these  resistant  
mutants.  
3  The  increased  usage  and  diversity  of  anRbioRcs  is  selecRng  
for  more  and  more  mulRple  drug  resistant  bacteria.  
4  The  evoluRon  of  anRbioRc  resistance  in  bacteria  is  extremely  
rapid.  We  can  observe  the  evoluRon  occurring  over  years  or  
even  months!  
3.    Early  history  of  evolu+onary  biology  
Section 3: Early history of evolutionary biology

Plato 424-348BC and Aristotle 384-322 BC

The Greek philosophy of Essentialism dominated biology

for many centuries

• The main philosophy of Essentialism was the ‘eidos’:

• All organisms have a transcendental ideal form.
• Each species has an immutable essence, and
imperfections only exist as accidents.

• Because of the philosophy of eidos, variation was often

ignored by naturalists

A great deal of evolutionary biology is the study of

variation and its causes
Section 3: Early history of evolutionary biology

Another key theme of Essentialism was the great chain of being,

the Scala naturae (a scheme for organizing species)

• A gradation from the inanimate, to “lower” life forms, to

plants and invertebrates, and up to the “higher” forms of life

• A core ideal of essentialism is that creation followed an

ordered plan

• Order is better than disorder and the creator made a

detailed, ordered design

• Since the creator made all plants and animals with perfect
form, no changes were expected in species over time.
Section 3: Early history of evolutionary biology

The Scala naturae

• The language that the

Scala naturae inspires
still pervades
evolutionary biology

We must be very careful

not to describe life forms
as ‘higher’ or ‘lower’
Section 3: Early history of evolutionary biology

Carolus Linnaeus (Sweden 1707-1778)

His life goal was to catalogue nature

and make manifest the plan of creation

He established the framework of

modern classification Systema
Naturae (1735)

He classified species into genera,

families, classes: all classification was
intended to relate life in its propinquity
(closeness) to the creator.

He was only 28 when he published this

book that realized worldwide acclaim!
Systema Naturae (1735)!
Section 3: Early history of evolutionary biology

James Hutton (Scotland 1726-1797) &

Charles Lyell (Scotland 1797-1875)

Principle of Uniformitarianism (idea

that greatly influenced Darwin)

“The present is the key to the past”

Observations of geology can and

should be explained by causes and
mechanisms that we now observe
Section 3: Early history of evolutionary biology

Lamarck (1744-1829 France)

First coherent model of evolution

Species are constantly being generated

(spontaneous generation) and thus some are
older and more developed than others
Section 3: Early history of evolutionary biology

Each species originates via spontaneous generation (Aristotle)

from non-living matter starting at the bottom of the chain of
being. Spontaneous generation was finally put to rest by
Pasteur in 1862.

Lamarck’s  theory  of  organic  progression  

Section 3: Early history of evolutionary biology

Lamarck (1744-1829 France)

Lamarckian evolution

Inheritance of acquired characteristics

Species vary because each has different
needs and strongly exercised organs
attract more ‘nervous fluids’

Famous example: A giraffe’s necks grows

over time to reach foliage in high trees

First cogent theory of evolution, but

famously wrong
Section 3: Early history of evolutionary biology

• Charles  Robert  Darwin  (1809-­‐1882)  

• Darwin  was  young  when  he  came  

up  with  some  of  his  most  earth  
changing  ideas.  
• Son  of  a  physician,  started  studying  
medicine  at  Edinburgh  (1830’s  
surgery  was  scary!  No  anesthesia!)  
• Dropped  out  of  medicine,  switched  
to  clerical  studies  (clergy)  
• Became  friendly  with  naturalists  in  
the  faculty  and  these  interacRons  
changed  him  forever  
Section 3: Early history of evolutionary biology

• Darwin invited to serve as a naturalist

on the H.M.S. Beagle by Robert
Fitzroy, a captain of a British ship sent
to chart the waters off the coast of
South America (12/27/1831-10/2/1836)

• Darwin collected specimens and

made biological, geological
observations

• Most of these specimens and his

journals @ Cambridge U. Library/
Museum
Section 3: Early history of evolutionary biology

• Darwin’s 5 year journey on the H.M.S. Beagle

• Charted the waters off the coast of South
America (he is 22 years old at outset)
Section 3: Early history of evolutionary biology

Several  events  occurred  soon  aHer  

Darwin’s  long  journey  that  had  
important  impacts  on  his  theory  of  
evolu+on  (1837-­‐1838)  

1.  World  famous  ornithologist:  John  

Gould’s  observaRon  upon  return:    

Mockingbirds  from  neighboring  

islands  are  different  enough  to  be  
species  

Triggered  idea  in  Darwin’s  mind  that  

different  species  evolved  from  
common  ancestors  
Section 3: Early history of evolutionary biology

2.  WriRng’s  of  Thomas  Malthus  

(Bri+sh  1766-­‐1834)  economist:  
Essay  on  the  principle  of  popula8on  

Journal  entry:  September  1838  (age  

29)  Darwin  found  himself  reading  
Malthus  "for  amusement”    

Key  point:  Human  growth  is  faster  than  

food  producRon,  famine  will  be  result  

Led  to  Darwin’s  idea  of  the  “struggle  

for  existence…”  
Section 3: Early history of evolutionary biology

Darwin’s idea of the “struggle for

existence”

Natural selection
Individuals with superior features
survive and reproduce more
successfully than individuals with
inferior features

Evolution
If these differences are inherited then
the average character of a species
can be altered over time
Section 3: Early history of evolutionary biology

Darwin’s first tree (Late

1837)

Less than two years after

the journey of the Beagle
is complete ---the main
idea of evolution was set
in Darwin’s mind
Section 3: Early history of evolutionary biology

• Darwin spent the next 22+ years

amassing evidence to support his
theories of evolution and natural
selection without publishing any of his
controversial ideas… although he wrote
dated assays that described each of his
theories in detail

• 1858, date unknown, Darwin received a

manuscript from Alfred Russel Wallace
(British 1823-1913) that described a very
similar theory of natural selection.

• While Darwin had been living a

comfortable life in England, Wallace had
been spending all his time collecting
specimens in Southeast Asia (Research
in Amazon/Malay archipelago)
Section 3: Early history of evolutionary biology

Darwin presented both of their

ideas together at a Royal
Society Meeting in London and
quickly wrote an abstract of his
book on evolution and natural
selection that he had intended
(490 pages long)

“The origin of species by

means of natural selection or
the preservation of favoured
races in the struggle for life”
Published 11/24/1859
 Darwin’s  Evolu+onary  Theory  –  Five  big  ideas:  

1  Evolu+on  –  CharacterisRcs  of  the  lineages  of  organisms  change  

over  Rme  (Darwin  wasn’t  the  first  to  have  this  idea  but  was  the  
first  to  amass  all  the  evidence).  

2  Common  Descent-­‐  All  life,  living  or  exRnct  has  descended  

without  interrupRon  from  one  or  a  few  original  forms.  
•  Species  diverge  from  common  ancestors,  are  at  first  very  
similar  and  accumulate  differences  over  Rme  and  can  
become  radically  different  
•  Different  from  Lamarck’s  ideas  (species  have  common  
origins)  
 Darwin’s  Evolu+onary  Theory  –  Five  big  ideas:  

3.  Gradualism    
• Even  radical  differences  between  organisms  have  evolved  from  
incremental  changes  over  Rme  
• We  will  also  discuss  different  hypotheses  that  radical  change  
can  occur  without  intermediate  steps  

4.  Popula+onal  changes  
• EvoluRon  occurs  by  changes  in  the  proporRons  of  individuals  in  
populaRons  
• This  simple  idea  was  a  radical  departure  from  other  ideas  of  
evoluRon  
 Darwin’s  Evolu+onary  Theory  –  Five  big  ideas:  

5.  Natural  selec+on  (independently  conceived  by  Wallace)  

• Changes  in  the  proporRons  of  individuals  in  a  populaRon  are  

caused  by  differences  in  their  ability  to  survive  and  reproduce  

• Non-­‐random,  differenRal  survival  and/or  reproducRon  of  

organisms  that  vary  in  one  or  more  heritable  characterisRcs.    

• Natural  selecRon  can  result  in  the  evoluRon  of  adapta+ons,  

features  that  make  an  organism  appear  “designed”  to  fit  it’s  
environment  
The  big  gap  in  Darwin’s  theory  –  What  is  the  
source  of  varia+on?  

• Dominant  theory  of  inheritance  is  “blending”  –  

leads  to  reduced  variaRon  over  Rme.  

• Darwin  did  not  realize  that  Gregor  Mendel  

(Austria  1822-­‐1884)  had  solved  this  problem  in  
1865,  with  his  theory  of  Par+culate  inheritance.  

• Stated  that  traits  were  inherited  by  parRcles  that  

remained  unaltered  over  Rme,  as  opposed  to  by  
‘fluids’  that  mixed  anew  in  each  generaRon-­‐  
MutaRons.    
• Mendel’s  papers  weren’t  rediscovered    unRl  
~1900  
Section 3: Early history of evolutionary biology

The theory of evolution was almost universally accepted

and inspired much research

• …but Natural Selection was not accepted by most

biologists for more than 60 years

• Many mutationists held that all change over time is

caused by mutations and that new species form when
major genetic reorganizations occur “Hopeful monsters”

• Although Darwin almost singularly formulated the original

theories of evolution, much of what we will learn has been
discovered much more recently:
Section 3: Early history of evolutionary biology

Major events in evolutionary biology

1920s-1930s The Modern Synthesis

• Darwin’s ideas of evolution and natural selection were reconciled with
genetics
• R.A. Fisher, J. B. S. Haldane and Sewell Wright formulated the field of
population genetics which showed that mutation and natural selection
together cause adaptive evolution, and that mutation is not an alternative, but
the raw material for natural selection

• 1950’s Discovery of DNA structure (1953 Watson & Crick) kicks off the field of
Evolutionary genetics

• 1960’s Evolutionary theory expands into ecology, animal behavior, reproductive

biology (Evolutionary biology is young and much of what we will learn has only
been examined in the last 50 years)

• 1990’s-2000’s Molecular Revolution – Sequencing of genes and genomes

opens up a whole new realm of biological study.
• We could also have a molecular evolution course that focused only on information
that has been learned in the last 20 years.