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Charles Darwin
1809-1882
- Theory shaped by several different fields of study and experiences: geology, economics, Beagle
- Proposed that species evolve over time through natural selection.
Alfred Russel Wallace
Naturalist in the Amazon and Southeast Asia ~ 1848
• Collected beetles
- Reached similar conclusions as Darwin
Natural Selection leads to...
Adaptations
Evidence of Evolutionary Change
• Fossil record
• Biogeography
• Observations of natural and artificial selection
• Homologies
• Anatomical
• Developmental
• Molecular
Evidence of Evolutionary Change - Fossil Records
- Fossils are preserved remains or traces of ancient organisms.
- Depth of the fossil can help inform its age
- Fossil record is incomplete, but shows gradual evolutionary change (e.g., horses)
Transitional Form
Intermediate between ancestral form and descendant
Evidence of Evolutionary Change - Biogeography
- Spatial record of evolution
It shows that organisms in similar environments but different locations evolve independently, while nearby regions tend to have related species.
- Similar species on continents that are far apart suggest past physical connection
Evidence of Evolutionary Change - Direct Observations (Natural
and Artificial Selection)
Selective breeding/ artificial selection
Breeders choose parents, over time can lead to large changes in morphology
• Made possible by genetic variation
Evidence of Evolutionary Change - Homologies: Anatomical
Refers to similarities in the physical structures of different organisms that arise from a common ancestor, even if those structures serve different functions today.
A classic example is the forelimbs of vertebrates:
- Human arm (grasping)
- Whale flipper (swimming)
- Bat wing (flying)
Evidence of Evolutionary Change - Homologies: Developmental
Refers to similarities between structures in different organisms that arise from the same embryonic origin or developmental pathway.
- Species that differ as adults often bear striking similarities during embryonic stages
- Presence of gill ridges in human embryos indicates that humans evolved from an aquatic animal with gill slits
- Human embryos have long bony tails
Evidence of Evolutionary Change - Homologies: Molecular
- The idea that DNA, RNA, or protein sequences are similar because they were inherited from a common ancestor.
- All living species use DNA to store information
- Sequences of closely related species tend to be more similar to each other than to distantly related species
Convergent Evolution
2 species from different lineages show similar
characteristics because they occupy similar
environments
The Mechanism of Natural Selection
1. Variation exists Individuals in a population are not identical—they differ in traits like size, color, behavior, or physiology. These differences often come from genetic variation (mutations, recombination).
2. Traits are heritable Some of these differences can be passed from parents to offspring through genes.
3. Overproduction and competition Organisms tend to produce more offspring than can survive. Limited resources (food, space, mates) create competition.
4. Differential survival and reproduction Individuals with traits better suited to their environment are more likely to survive and reproduce. This is often called “fitness.”
5. Accumulation of favorable traits Over generations, beneficial traits become more common in the population, because the individuals carrying them leave more offspring.
Descent with modification
A concept from the theory of evolution by natural selection stating that species change over time, with new species arising from pre-existing ones while retaining some ancestral traits.
Homology
Similarity between traits due to shared ancestry, even if the traits serve different functions.
Homologous structure
A physical feature in different species that has a common evolutionary origin but may have different functions (e.g., vertebrate forelimbs).
Analogous structure
Structures that serve similar functions but evolved independently, not from a common ancestor (e.g., wings of birds and insects).
Convergent evolution
The process where unrelated species independently evolve similar traits due to similar environmental pressures.
Fossil
Preserved remains, impressions, or traces of ancient organisms, providing evidence of past life and evolution.
Vestigial structure
A reduced or nonfunctional feature that was functional in an organism's ancestors (e.g., human tailbone).
Evolutionary tree
A diagram (also called a phylogenetic tree) showing relationships among species based on common ancestry and divergence over time.
Biogeography
The study of the geographic distribution of species and ecosystems, helping explain how evolution and Earth's history shape where organisms live.
Artificial selection
The intentional breeding of organisms by humans to enhance desired traits (e.g., dog breeding).
Adaptation
A heritable trait that increases an organism's ability to survive and reproduce in a specific environment.
Why variation is common and necessary for natural selection
Variation exists in populations because of:
Mutations (new genetic changes)
Genetic recombination during meiosis
Gene flow between populations
This variation is essential for natural selection because:
Selection can only act on existing differences
Without variation, all individuals would respond the same to environmental pressures → no evolution
Using the Hardy-Weinberg equilibrium
Key equations:
Allele frequencies: p + q = 1 (p = dominant allele, q = recessive allele)
Genotype frequencies: p^2 + 2pq + q^2 = 1
p^2 = homozygous dominant
2pq = heterozygous
q^2 = homozygous recessive
Five conditions for Hardy-Weinberg equilibrium
No mutations
Random mating
No natural selection
Extremely large population size (no drift)
No gene flow (no migration)
Four patterns of natural selection
- Directional selection
- Stabilizing selection
- Disruptive selection
- Balancing selection
Directional selection
Favors one extreme phenotype
Example: larger beak size becoming more common
Stabilizing selection
Favors intermediate traits
Reduces variation (average individuals do best)
Disruptive selection
Favors both extremes over the intermediate
Can lead to speciation
Balancing selection
Maintains multiple alleles in a population
Often due to heterozygote advantage or changing environments
Sexual selection
A type of natural selection focused on mating success rather than survival.
Types:
Intrasexual selection
Competition within the same sex (e.g., male vs male fighting)
Intersexual selection
One sex chooses mates (often female choice based on traits)
Genetic drift
- Random change in allele frequencies, especially in small populations.
Key points:
Not based on fitness (pure chance)
Can reduce genetic variation
Strongest in small populations
Examples:
Bottleneck effect: sudden population reduction
Founder effect: small group starts a new population
Biological Species Concept (BSC)
Species are groups of actually or potentially interbreeding populations that are reproductively isolated from others.
Limitation: doesn't work for asexual organisms or fossils
Morphological Species Concept
Species are defined by physical traits (appearance).
Limitation: subjective; different species can look similar (cryptic species)
Phylogenetic Species Concept
Species are the smallest group sharing a common ancestor (based on DNA/evolutionary history).
Limitation: requires detailed data; can over-split species
Why reproductive isolation is necessary
- Prevents gene flow between populations.
Without it:
Populations would keep mixing genetically
Differences would not accumulate
With isolation:
Genetic differences build up over time
Eventually leads to new species (speciation)
Eight mechanisms of reproductive isolation (Prezygotic)
Habitat isolation – live in different environments
Temporal isolation – breed at different times
Behavioral isolation – different mating behaviors
Mechanical isolation – incompatible reproductive structures
Gametic isolation – sperm and egg cannot fuse
Eight mechanisms of reproductive isolation (Postzygotic)
Reduced hybrid viability – offspring don’t survive well
Reduced hybrid fertility – offspring are sterile (e.g., mule)
Hybrid breakdown – offspring viable but weak in later generations
Sympatric speciation (same location)
Occurs without geographic separation.
Common mechanisms:
Polyploidy (extra chromosome sets, especially in plants)
Habitat differentiation within the same area
Sexual selection (mate preferences split populations)
Disruptive selection
Allopatric speciation (most common)
Happens when populations are geographically separated (mountains, rivers, distance).
Why it's most common:
Physical barriers easily stop gene flow
Populations evolve independently under different conditions
Speciation
Formation of new species
Reproductive isolation
Barriers preventing interbreeding
Biological species concept
Defines species by ability to interbreed
Allopatric speciation
Speciation due to geographic separation
Sympatric speciation
Speciation without geographic separation
Hybrid zone
Area where two species meet and interbreed
Polyploidy
Extra sets of chromosomes (common in plants)
Hybrid
Offspring of two different species
Intraspecific
Within the same species
Interspecific
Between different species
Adaptive radiation
Rapid diversification into many species from a common ancestor (often when new niches open)
How eukaryotic cells evolved from prokaryotic cells
The endosymbiotic theory.
- An ancestral prokaryotic cell engulfed smaller bacteria
- Instead of digesting them, they formed a symbiotic relationship
- These internalized bacteria evolved into organelles:
- Mitochondria (from aerobic bacteria)
- Chloroplasts (from photosynthetic bacteria, in plants/algae)
Biases of the fossil record
Hard parts preserve better (bones, shells > soft tissue)
Certain environments favor fossilization (sediments, low oxygen)
Recent fossils are more common than older ones
Geological processes (erosion, heat) destroy fossils
What we know (and don't) about the origin of life (Best Understood)
- Early Earth had simple molecules (water, methane, ammonia)
- Organic molecules can form naturally (e.g., Miller-Urey type experiments)
- RNA may have been an early genetic system (RNA world hypothesis)
What we know (and don't) about the origin of life (Less Certain)
Exact pathway from nonliving chemistry → first living cells
First self-replicating system
Exact environment where life began (deep-sea vents vs shallow pools)
Timeline of life on Earth
Earth formed about 4.6 billion years ago.
Prokaryotes: ~3.5–3.8 billion years ago
Eukaryotes: ~2.0–2.5 billion years ago
Multicellular eukaryotes: ~1.0 billion years ago
Animals: ~600 million years ago
Land plants: ~470 million years ago
Humans (Homo sapiens): ~300,000 years ago
Changes in Earth's environment and their effects
Oxygen revolution (~2.4 billion years ago)
- Photosynthetic microbes released oxygen
- Caused mass extinction of anaerobic organisms
- Enabled evolution of aerobic life
Climate changes (ice ages, warming periods)
- Alter habitats and drive natural selection
Mass extinctions (e.g., asteroid impacts)
- Wipe out species → open niches → adaptive radiations
Estimating fossil age (radiometric dating)
Basic idea:
Unstable isotopes decay at a known rate (half-life)
Measure ratio of parent → daughter isotopes
Example:
If half-life = 1 million years
50% parent left → 1 million years old
25% parent left → 2 million years old
Common isotopes:
Carbon-14 (recent fossils)
Uranium-238 (very old rocks)
Early development and evolutionary relationships
Studying embryos helps reveal shared ancestry.
- Different species often show similar early stages
- Controlled by conserved developmental genes (e.g., Hox genes)
Interpreting and drawing phylogenetic trees
How to read one:
- Root = common ancestor
- Branches = lineages through time
- Nodes = common ancestors where lineages split
- Sister taxa = closest relatives (share a recent node)
Drawing one (basic steps):
Identify shared traits (derived characteristics)
Group organisms by shared derived traits
Place the most ancestral traits near the root
Add branches where new traits appear
Trees can...
change
Phylogenetic trees are hypotheses, not final truths.
New DNA evidence or fossil discoveries can change relationships
Advances in molecular biology often lead to revisions
Monophyletic group (clade)
Includes a common ancestor and all descendants
Why monophyletic is preferred:
- Accurately reflects evolutionary history
- Matches patterns of descent in evolution
Paraphyletic group
- Includes a common ancestor but not all descendants
Polyphyletic group
- Includes organisms without their common ancestor
Principle of parsimony
The simplest explanation (fewest evolutionary changes) is most likely correct.
When choosing between trees:
- Prefer the one requiring the fewest mutations or trait changes
How neutral mutations spread
- Changes in DNA that don’t affect fitness.
They spread through populations via genetic drift:
- Random chance determines whether they increase or disappear
- Especially important in small populations
- Can eventually become common or fixed without providing an advantage
Using molecular clocks
A molecular clock uses the rate of DNA mutations to estimate how long ago two species diverged.
- Mutations accumulate at roughly constant rates in some genes
- By comparing DNA differences, scientists estimate time since a common ancestor
Basic idea:
- More genetic differences → longer time since divergence
- Calibrated using fossils or known dates
Complications in phylogenetic trees
(HGT): Horizontal gene transfer is when genes move between unrelated organisms (common in bacteria).
- Makes species appear more closely related than they are
- Creates a “web” of relationships instead of a simple tree
Convergent evolution: Convergent evolution occurs when unrelated species evolve similar traits.
- Leads to analogous structures
- Can mislead scientists into grouping species incorrectly
Binomial nomenclature & taxonomy
Binomial nomenclature
- Developed by Carl Linnaeus:
- Each species has a two-part name:
Genus (capitalized)
Species (lowercase)
Example: Homo sapiens
Taxonomic hierarchy (broad → specific):
Domain
Kingdom
Phylum
Class
Order
Family
Genus
Species
Phylogeny
Evolutionary history of a species or group
Systematics
Study of classifying organisms and their relationships
Taxon
Any named group (e.g., species, genus, family)
Sister taxa
Two groups that share an immediate common ancestor
Analogy
Similarity due to convergent evolution, not shared ancestry
Homology
Similarity due to shared ancestry
Clade
A monophyletic group (ancestor + all descendants)
Monophyletic
Includes common ancestor and all descendants
Paraphyletic
Includes ancestor but not all descendants
Polyphyletic
Excludes common ancestor
Shared ancestral character
Trait inherited from a distant ancestor (not unique to a group)
Shared derived character (synapomorphy)
Trait unique to a group, used to define clades
Bacteria
Prokaryotic (no nucleus)
Cell walls contain peptidoglycan
Very diverse metabolically
Archaea
Prokaryotic
Cell walls lack peptidoglycan
Often live in extreme environments
Biochemically more similar to eukaryotes
Archaea and Eukarya are more closely related to each other than either is to Bacteria
Eukarya
Have nucleus and membrane-bound organelles
Include animals, plants, fungi, protists
Ecological roles of prokaryotes
Decomposers: break down dead organic matter, recycle nutrients
Producers: some perform photosynthesis (e.g., cyanobacteria)
Nitrogen fixers: convert N₂ → ammonia (usable by plants)
Pathogens: cause disease
Symbiosis:
Mutualistic (both benefit)
Commensal (one benefits, other unaffected)
Parasitic (one benefits, one harmed)
Horizontal gene transfer (HGT)
Transformation: uptake of free DNA from environment
Transduction: DNA transferred by viruses
Conjugation: direct transfer via cell-to-cell contact
Importance:
Rapid spread of traits (e.g., antibiotic resistance)
Contributes to genetic diversity
Played a role in early evolution and possibly in the origin of eukaryotes (e.g., gene exchange before organelles stabilized)
Bacterial Shapes
Coccus (spherical)
Bacillus (rod-shaped)
Spirillum (spiral)
Bacterial Cell wall types:
Gram-positive: thick peptidoglycan
Gram-negative: thin peptidoglycan + outer membrane
Bacterial Modes of Nutrition
Photoautotroph: light energy + CO₂
Photoheterotroph: light energy + organic carbon
Chemoautotroph: chemical energy + CO₂
Chemoheterotroph: chemical energy + organic carbon
Bacterial Oxygen Use:
Aerobic: require oxygen
Anaerobic: do not use oxygen (may be harmed by it)
Facultative anaerobes: can switch
Halophile
Thrives in high salt
Extremophile
Lives in extreme conditions
Thermophile
Thrives in high temperatures
Methanogen
Produces methane (archaea)
Nitrogen fixation
Conversion of N₂ to usable ammonia
Aerobic
Requires oxygen
Anaerobic
Does not require oxygen
Unifying characteristics of protists
Protists are grouped mainly by what they are not:
- Eukaryotic (have nucleus and organelles)
- Mostly unicellular (some multicellular or colonial)
- Do not fit into animal, plant, or fungi kingdoms
They are incredibly diverse in:
- Nutrition (autotrophs, heterotrophs, mixotrophs)
- Habitat (aquatic, moist environments)
- Structure and movement