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Early vertebrate with 4 eyes
Early fossils thought the extra eyes were pineal organs, but melanin evidence suggests the pineal eyes likely functioned as camera-type eyes
Metazoans
Multicellular heterotrophs that can be motile (at least as larvae)
Endoderm
Forms linings of the gut and lung
Mesoderm
Forms muscles, bones, cartilage, heart, kidneys, and notochord
Ectoderm
Forms nervous system, epidermis, and sensory organs
Bilaterians
Animals with two sides
Protostomia
~90% of bilaterian phyla; “mouth first” development
Deuterostomia
“Mouth second” development
Blastopore
Opening that forms first during development; becomes the anus in deuterostomes
“We all start as an anus”
In deuterostomes, the blastopore becomes the anus first
Embryogenesis
Development from fertilization to hatching
Basal deuterostome body plan
Shown by sea urchin and tunicate development
Sea urchin cleavage type
Radial holoblastic cleavage
Vegetal pole/plate (sea urchin)
Region where mesenchymal cells detach and where invagination begins
Micromeres (sea urchin)
Undergo epithelial-mesenchymal transition and migrate into the blastocoel
Sea urchin gastrulation
Vegetal cells invaginate to form the gut which elongates toward the animal pole
Tunicate cleavage type
Bilateral holoblastic cleavage
Tunicate early cleavage pattern
Vegetal half divides before the animal half
Tunicate gastrulation
Involution of endoderm and spreading over of ectoderm
Tunicate neurulation
Notochord forms and extends down a tail bud
Vertebrates
Allow study of development of notochord and vertebrae
Xenopus blastula division
Animal side divides faster; divisions become asynchronous between animal and vegetal sides
Xenopus gastrulation
Vegetal middle region invaginates, forming the blastopore lip and blastopore (anus forms first)
Xenopus neurulation
Strong head-to-tail neurulation with notochord extension and neural tube formation
Xenopus tailbud stage
Whole body elongates; tadpole stage ~37, adult stage ~66
Zebrafish cleavage
Discoidal cleavage
Zebrafish blastula formation
Cleavage happens at the top; yolk forms the bottom
Zebrafish gastrulation
Involution occurs atop yolk and cells move down sides of yolk
Primary neurulation
Most vertebrates use primary neurulation (like Xenopus)
Secondary neurulation
Zebrafish use secondary neurulation (some vertebrates use it only in the tail)
Amniotes
Group where development involves importance of the amnion
Avian gastrulation structure
Primitive streak
Primary hypoblast formation
Hypoblast islands form primary hypoblast with congregation around Koller’s sickle
Secondary hypoblast formation
Primary hypoblast connects to posterior marginal zone (PMZ) to form secondary hypoblast cells
Primitive streak function
Cells migrate through the streak to form mesoderm and endoderm
Primitive streak equivalent to blastopore lip
The streak is equivalent to the blastopore lip
Primitive groove equivalent to blastopore
The groove is equivalent to the blastopore
Primitive groove timing
Appears ~18–20 hours after egg is laid
Notochord and somite formation timing (birds)
~20–22 hours after egg is laid
Primitive streak regression timing
~23–25 hours after egg is laid
Somites
Paired structures forming vertebral column, ribs, occipital bone, skeletal muscle, cartilage, tendons, and skin
Why primitive streak exists
Blastopore is circular but streak defines body axes and separates embryo from yolk
Animals with a primitive streak
Birds, reptiles, and mammals
Haeckel’s law
“Ontogeny recapitulates phylogeny”
Problem with Haeckel’s embryo drawings
Accidental printing of the exact same woodcut three times
What was missing in Haeckel embryo comparisons
Yolks or placenta
Issue with drawings vs photographs
Bias and Missing structures may be less obvious in drawings
Why Haeckel was wrong
Biogenetic law (embryos pass through adult ancestral stages) is not true
Von Baer’s Laws of Divergence
General traits appear early; specialized traits appear later
Embryological diversity
Shown across fish, amphibians, reptiles, birds, and mammals
Phylogeny (phylogenetic tree)
Branching diagram showing evolutionary relationships among species
Tips/taxa (taxon singular)
The endpoints of a phylogeny representing species/groups
Nodes
Branch points on a phylogeny
MRCA
Most recent common ancestor
Closest relative (on a tree)
Taxon that shares the most recent MRCA
Phylogenetic systematics
Classifying organisms based on evolutionary relationships
Clade
A monophyletic group including an ancestor and all descendants
Monophyletic
Includes an ancestor and all its descendants
Paraphyletic
Does not include all descendants (not a clade)
Molecular scaffolding
Combine molecular data for extant taxa with morphology to place fossils on tree
Extinct taxa on phylogenies
Can be shown with daggers (†)
Chronogram
Phylogeny where branch lengths represent time since divergence
Divergence times estimated using
Fossil ages, molecular clocks, and geological events
Molecular clock concept
Genetic differences accumulate at a known rate to estimate divergence time
Character states on phylogenies
Traits can be mapped on tips, nodes, and branches
Tick marks on branches
Show when a character state change likely occurred
Ancestral vs derived states
Ancestral = original state; derived = changed state
Homology
Trait similarity due to shared ancestry
Homoplasy
Trait similarity not due to shared ancestry (convergent evolution)
Apomorphy
A trait that has changed from its ancestral form
Plesiomorphy
An ancestral trait
Synapomorphy
A shared derived trait inferred to be in the MRCA of a group
Misconception (most evolved taxon)
Tip order does not mean “most evolved/advanced/derived”
Reality (phylogeny rotation)
Tip order can rotate; only branching pattern matters
Misconception (basal = primitive)
First branching taxon is not the ancestor or “most primitive”
Reality (sister taxa)
Early-branching taxa are sister to the rest; no extant taxon is ancestral
Misconception (species branch from species)
Extant species do not evolve from other extant species
Correct view of evolution
Humans did not evolve from chimps; humans and chimps share a common ancestor
Human classification summary
Animal → bilaterian → deuterostome → vertebrate → jawed vertebrate → bony fish → tetrapod → amniote → mammal → primate → Homo sapiens
Heterochrony
Changes in timing of gene expression or development
Heterotopy
Changes in location/position of gene expression
Heterometry
Changes in the degree/amount of gene expression