Lec 9 Flashcards
Developmental abnormalities are widespread
Observations of development found abnormalities are widespread
Homeotic transformations
Certain mutations in insects and vertebrates result in one body part replacing another or duplicating - these are callled homeotic transformations
Homeotic transformations seem to be most common in parts of the body that were either repeated (appendages, ribs, and so on), segmented, or both
What can homeotic transformation tell us about evolutionary change?
Evolutionary Development (Evo-devo_
Structural variation in body shape and form depends in part on when and where certain genes are expressed
This field is a fusion of developmental and evolutionary biology
If we can understand this process, we can understand the origin and evolution of morphological variation and innovation - major transitions in the evolution of life
History of Evo-devo
Ontogeny: Development of an individual over its lifetime
Pre-Darwinian naturalists noticed that the development of an individual started with “simple” traits and moved on to complex traits later in development
Meckel-Serres Law (early 1800s): Embryos display characteristics of embryos from species that came before them in the scala naturae - the classification of life forms from the “highest” to the “lowest”
von Baer’s law (1828): General traits appear early in development, and more specific traits that separate species appear later
Von Baer: Characteristics that unite species appear _______
Early
Stage 1 embryos are most similar to each other
Stage 4 embryos are least similar to one another
Haeckel’s Theory of Recapitulation
Ernst Haeckel expanded on the Meckel-Serres law with his Biogenetic Law: Ontogeny is a precise and compressed recapitulation of phylogeny
First theory to tie development to evolutionary theory
Put development in terms of phylogeny
Thought that evolution produced novelty by “tacking” new structures on to the terminal part of development of an ancestor
-This is NOT true: Evolution acts on ALL stages of an organism’s life, including the embryological stages. Development, like all traits, is thus a fusion of phylogenetic history and ongoing adaptive change
Evo-devo and the Modern Synthesis
Experimental work on evolutionary genetics in the 1930s and 40s demonstrated that genes not only code for physical traits, but also control the rate and timing of development
-When different structures appear during embryogenesis
Heterochrony: The time in the developmental process at which a trait is first expressed, relative to when that trait is first expressed in the ancestor
-Puts developmental stages (when things appear) into an evolutionary lens
Types of heterochrony
2 categories: Changes that affect time of onset of reproductive traits, and changes that affect timing and appearance of somatic traits
Heterochrony: Recapitulation via acceleration
A trait that appears late in development in an ancestral species, but earlier in development in the descendant species (AKA peramorphosis or overdevelopment). Genetic change can lead to: Somatic trait appearing earlier (acceleration)
Appearance of somatic = accelerated
Appearance of Reproductive = unchanged
Heterochrony: Recapitulation via hypermorphosis
Genetic change can lead to : Reproductive trait appearing later (hypermorphosis)
Somatic - unchanged
Reproductive - retarded
Heterochrony: Paedomorphosis via progenesis
A trait that appears EARLY in ancestors but LATER in descendants
Reproductive trait appearing earlier (progenesis)
Somatic - unchanged
Reproductive - accelerated
Heterochrony: Paedomorphosis via neoteny
Somatic trait appearing later (neoteny)
Somatic - retarded
Reproductive - unchanged
Recapitulation
A trait appears EARLIER in descendent species
Paedomorphosis
Trait appears later in descendent
Concept of heterochrony was a significant step forward in our understanding of the evolution of development
Incorporates evolutionary history by comparing ancestral and descendent species
Focus on genetic change
Recognizes that traits associated with reproduction are fundamentally different from somatic traits
The concept of heterochrony allows us to:
a) Think about development within an evolutionary framework
b) Compare the time at which traits appear during development in ancestors and descendants
c) Construct a phylogeny base don when traits appear during development
d) A and B
d) A and B
Best studied example of heterochrony occurs in neotenic axolotls
Salamanders spend juvenile stage in the water
They then lose juvenile traits and move onto the land
The axolotl never loses its juvenile traits (gills, flattened tail) and stays in the water
Reproductive traits appear at same time as ancestors, but somatic traits never appear at all
What causes neoteny?
Most salamanders have a spike in thyroid hormone associated with metamorphosis, but axolotls don’t
Adding thyroid hormone to water makes juvenile axolotles metamorphose into terrestrial-like forms
Mechanism may be reduced expression of TH regulatory genes in axolotls
Why would neoteny be favored by natural selection?
Maybe staying in the water is safer
Some other salamander species can stay neotenous under certain environmental conditions (“facultative neoteny”)
Facultative = under certain conditions
Axolotls are OBLIGATE neonates
Maybe staying in the water is safer
Some other salamander species can stay neotenous under certain environmental conditions (“facultative neoteny”)
This occurs more frequently when there are few predators in ponds, water levels are stable, and there is little competition
Supports the idea that neoteny is favored when these conditions are met
What observation might support the hypothesis that neoteny occurs because it’s safer to stay in the water?
Facultatively neotenous species stay in the water when the water level is stable and there are few predators in the ponds
Axolotls
Native to Mexico
When Spanish settled in 1521, drained lakes = first step to pushing axolotls to extinctions
Neotenic: Reach adulthood without undergoing metamorphosis
Inject iodine in lab to stimulate metamorphosis
Live around 15 years in the wild
Able to regenerate PERFECTLY; can regenerate limbs, spinal chord, jaw, and skin with NO scarring
1000% more resistant to cancer than any other animal
Cells become pluripotent after limb cut off; can then be used to differentiate as needed to regrow limb
Can also place removed limb from one axolotl onto another and it will grow that into another limb
How do multicellular creatures differentiate into so many different forms?
Every multicellular organism develops from a single cell
Except for sperm and eggs, every cell in the body of a multicellular creature contains the same set of genes
yet skin cells function very differently than do the cells in muscles, cells in the liver, and so on
Regulation, expression and switches
Very early in development each cell in an embryo is totipotent - it could develop into any kind of cell
We talked about how some cells give up the ability to be reproductive in favor of performing other tasks
Which type of cell they become depends on how their genes are regulated and expressed, and the environment surrounding the cell
Homeotic genes
Tells genes what to become
Genes that play key role in development and construction of the phenotype
Encode proteins that switch other genes on and off in a specific sequence
This affects cell size, shape, division,a dn positioning within an organism’s body plan
Act as a map for where structures should develop
In plants the most important homeotic genes are called the MADS-box genes
In animals the key homeotic genes are the HOM (inverts) or Hox (vertebrates) genes