Leactures 8-11 Flashcards
What definition did Reverend William Paley give for adaptations in 1802 book ‘Natural Theology’
Organisms appear contrived as it towards an end and that they show relation in parts
- organisms did not need to be perfect—same way broken watch still appears designed
Example and explanation of self-organisation
- self organisation is where order or patterns occur spontaneously in systems
- rapid cooling of molten basalt 50-60 mya—hexagonal columns
- also happens in biology
What are some of the important extensions to Darwin’s theory?
x 3
- Hamilton and inclusive fitness
- Fisher and Price with the fact other processes may decrease fitness
- Grafen formalised link with apparent design and mathematics of allele frequency change
Hamilton’s and concept of inclusive fitness
- social evolution, idea that actions have consequences for members of same population, rather than just their own reproductive success
- includes biological altruism and spite
- personal fitness=own reproductive rate
- inclusive fitness=direct + indirect fitness
What does eusocial mean?
The structuring of populations where the majority of the organisms are sterile
- e.g naked mole rates or ants
Fisher and Price
Fisher = rate of increase in fitness of any organism at any time is equal to its genetic variance in fitness at that time
- price explained this so that it was true- by stating he didnt mean total evolutionary change but only on allele frequency change due to NS where environments remain constant
- then natural selection always acts to increase fitness but that other processes may decrease it
(like drift)
Grafen
connect understanding of mathematics of allele frequency change and then the idea of apparent purpose/design
- HOW- correspondences between allele frequency changes and an optimisation programme
- that natural selection when isolated as a component of evolutionary change is an optimising process leading to organisms that appear designed as if to maximise inclusive fitness
- also- all genes with same pattern of inheritance tend to correspond to same optimisation programme
What is an optimisation programme?
pretends that organisms are agents employing strategies to maximise inclusive fitness
Active research is on marginal cases where unclear if a trait has an adaptive function
- give troglobite example
- troglobite is an animal that only lives in caves
- many convergent features
- no eyes and lack of pigment
- regressive evolution—all came from a species with both eyes and pigment
2 ideas
1. directional (so has adaptive purpose)
2. Drift (due to relaxed selection) - eye loss = directional
- pigmentation = drift
- Protas et al in Maxican cave tetra using QTL mapping found this
What is a fitness landscape?
AKA adaptive topography or surfaces of selective value
- metaphor introduced by Wright in 1932
- visualising evolution
- fitness valleys with global optimum and local optima
2 types of fitness landscape
- organism level
- vertical axis is individual fitness
- other axes represent either genotype of phenotype
(only useful is fitnesses are not frequency dependent) - population level
- vertical is population mean fitness
- other can be population mean phenotype
Example of individual-level phenotypic landscape
Brodie (1992)
- garter snakes
- colour pattern and escape behaviour
- Juvenile snakes better is longitudinal stripes and straight line escape or blotched colours and multiple reversals during escape
- intermediates didnt have as high a fitness
What is the shifting balance theory?
Wright - used to explain how evolve from local optimum to peak
- large natural populations frequently subdivided
Phase 1 - small Ne enables cross fitness valleys with aid of genetic drift
Phase 2 - natural selection pushes subpop to new, sometimes higher fitness peaks
Phase 3 - subpops at higher peaks send out more migrants so gene flow to other pop
Problems with shifting balance theory of evolution
- phase 1 requires small Ne and low migration but then 2 needs large Ne and 3 needs high migration rate
- shown can work but only if limited range of conditions
- others same solving non-problem as natural population rarely stuck at suboptimal peaks as landscapes change over time and high dimensionality of peak-world fitness means that ridges are likely to be common and dont need to cross fitness valley
Experiments are difficult for shifting balance theory, two main experiments give conflicting evidence
- Weber 1996 - Drosophila fight speed
2. Lenski and Travisano 1994 - E.coli
Weber experiment 1996
wind tunnel automatically select for faster flies - 9 million flies (over all) - 100 generations - top 4.5% of speed roughly 2 increased to 170cm per second
- flies showed little to no decline in fitness compared to control lines and no decrease in flying speed when selection relaxed so no fitness valley
Lenski and Travisano 1994
E.coli in new environment for 10,000 generations
- regular intervals fitness of 9 replicate lines compared to ancestral strain (kept frozen)
- all populations increased in fitness initially but some later plateaued at suboptimal levels-stuck at local fitness peaks?
- though a few experimental errors like maybe bottleneck built into experimental design? as all from single clone
What is the neutral theory of molecular evolution?
Kimura in 1968
the vast majority of molecular evolution was due to drift
- so most substitutions and polymorphisms were effectively neural
- do not deny widespread action of negative selection
What observations do the neural theories aim to explain?
x 6
- substitution rate was higher than expected
- roughly constant in different lineages
- unreleased to rate of phenotypic evolution
- highest in less important proteins and parts of proteins
- molecular diversity much higher than expected
- diversity higher in larger populations
How to test neutral theory
- examine distributions of fitness effects
- in labs by introducing mutations or by bioinformatics methods
What is speciation?
The formation of two or more species from a single ancestral population
What 5 traits do populations need to fulfil to be species?
- Unique phenotypic character found in no other population
- Multivariate distributions of quantitive trait value that does not overlap with distributions found in other population
- Monophyly (most recent common ancestor of population has no descendants outside of population
- Distinct ecological niche
- Not hybridising with other populations
(in practice not all of these species criteria are filled)