Obrázek1 kill_bill surikata2
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sault-rifle-simulator.jpg Výsledek obrázku pro trump putin image

natural selection essentially a competitive process Þ
cooperation between organisms is one of nature´s most peculiar features
social insects, humans
mutualism
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http://pensci.files.wordpress.com/2012/01/honeybee2.jpg
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713/crocodile-plover.jpg http://nd04.jxs.cz/034/547/5359f3a9e5_74617442_o2.jpg
http://scribbler.cz/wp-content/uploads/2014/01/Magnetick%C3%A1-termiti%C5%A1t%C4%9B-4.jpg
http://www.xterm-sumitomo.com.au/sites/default/files/images/termite_hq.gif

http://upload.wikimedia.org/wikipedia/commons/thumb/8/81/Dicty_Life_Cycle_H01.svg/533px-Dicty_Life_
Cycle_H01.svg.png
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jpg http://www.clemson.edu/extension/hgic/graphics/2354/slime_mold.jpg
Eg.: slime molds
https://encrypted-tbn1.gstatic.com/images?q=tbn:ANd9GcQ5tYEWNU3f-mEv25f5vqIc1vlHGhk2XHihic65r-dUSPU
Lynkhnw
http://islandnature.ca/wp-content/uploads/2010/03/yellow_slime_mold.jpg
http://greatecology.com/wp-content/uploads/2012/12/Chocolate-Tube-Slime-Mold.jpg
http://sciencegeekgirl.com/files/2008/10/picture1_1.jpg
http://blogs.discovermagazine.com/loom/files/2011/10/slime-mold-5.jpg

How can, in spite of conflict between organisms,
cooperation evolve?
Charles Darwin: struggle for life
but also cooperation between a cow and her calf (cooperation between
relatives))
Neodarwinism: evolution in populations, selection affects individuals
´ till the 1960s, this assumption rather implicit (cf. Wright´s „interdemic
selection“)
Darwin, Wallace, Konrad Lorenz etc.: „benefit of species“, „survival
of species“....

http://www.ruraltech.org/video/2011/ITC_Workshop/19_Morishima/flash_files/lgslide_25_25.png
http://upload.wikimedia.org/wikipedia/commons/f/f1/Herd_of_sheep.JPG
William Forster Lloyd (1833) ® Garrett Hardin (1968):
Tragedy of the commons
adding 1 sheep to the herd Þ direct benefit for the owner
´ costs (drop of pasture) shared by the whole group
 Þ if people behave with respect to their benefit independently and
rationally, eventually the sources are necessarilly depleted
Solution =  voluntary restriction by herders ®
Why should such behaviour be favoured by selection?

1962 – Vero Copner Wynne-Edwards:
Animal Dispersion in Relation to Social Behaviour
flocking, dispersion, restriction of reproduction, altruism
cooperation explained as the selection of whole groups rather than
individual selection (in extreme form „adaptation for species´ survival“)
V. C. Wynne-Edwards
cheater
http://www.britishbirds.co.uk/wp-content/uploads/article_files/V90/V90_N06/V90_N06_P209_T50.jpg
cheater

File:John Maynard Smith.jpg
reaction:
1964: William D. Hamilton,
          John Maynard Smith
1966: George C. Williams
1976: Richard Dawkins
richard-dawkins File:W D Hamilton.jpg Image:George C. Williams.jpg
příbuzenský výběr
kin selection
důležité jsou geny
genes are important!

GROUP SELECTION
V.C. Wynne-Edwards:
 dispersion in order to avoid depletion of sources
 production of fewer offspring than potentially possible
alarm calls, fish shoals
http://i.huffpost.com/gen/1359251/images/o-FISH-SCHOOLS-facebook.jpg http://i.imgur.com/PFPhmSD.jpg
http://www.wascgroup.com/wp-content/uploads/2013/02/Grey-Nurse-Sharks-Swimming-through-Fish-School-
940x627.jpg http://www.duskyswondersite.com/wp-content/uploads/2014/11/School-of-Fish.jpg

„stotting“
Individual advantage!
http://upload.wikimedia.org/wikipedia/commons/4/47/Springbok_pronk.jpg
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http://medias.photodeck.com/00f4e78c-55de-11e3-80b6-e7323dcc0875/016030jj_xlarge.jpg
http://s.hswstatic.com/gif/gazelle-body-language-1.jpg
Thomson´s gazelle, springbok, mule deer, pronghorn etc.
https://www.youtube.com/watch?v=qr5Sru8gGSk
https://www.youtube.com/watch?v=jMIiB9DnRXg

Turdoides
guards of the Arabian babbler (Turdoides squamiceps)
and meerkats (Suricata suricatta)
surikata1
Suricata suricatta
T. squamiceps
sentinel
alpha male
Individual advantage!

Theoretical arguments against group selection:
altruism = behaviour increasing recipient´s fitness and,
      at the same time, decreasing donor´s fitness
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infiltration of selfish individual
spread of „selfish“ allele in the population
Wynne-Edwards´ population of altruists
fixation of „selfish“ allele
lemmings

Low heritability and longer generation time of the group relative to
heritability and generation time of individuals Þ changes at the
individual level much faster
  Þ infiltration of selfish individuals, extinction of the altruistic
  population
skenovat0002

rapid alternations of extinction and re-creation of demes
Eg.: fig wasps (Agaonidae)
virtually no migration:

   c … cost for an individual
   (b – c) … benefit for the group

island model:
Conditions for group selection:
Conclusion: interdemic (group) selection will be stronger than intrademic
(individual) selection only if the group benefit relative to the individual
cost is higher than the average number of migrants per generation.

Wade
Michael Wade (1977): group selection experiment in the red flour
beetle (Tribolium castaneum)
But in nature the role of group selection probably minimal
fast-growing group
slow-growing group
individual selection

KIN SELECTION
http://www.voxeurop.eu/files/Bertrams-greece.jpg
http://kgov.com/files/images/science/JBS-Haldane-RSR.jpg
If I rescued my two brothers from drowning in the river, it would be the same as to rescue myself!
http://flexpeditions.com/wp-content/uploads/2013/03/WWR-rescue-jump.jpg

William Hamilton (1964):
Hymenoptera: haplo-diploid system of sex determination:
  females 2N, males N
  Þ relationship:
  worker – worker = ¾
  queen – descendants = ½
  worker – drone = ¼
inclusive fitness = fitness of an individual and his/her relatives
altruism between relatives = kin altruism

http://upload.wikimedia.org/wikipedia/commons/thumb/3/3f/Coefficient_of_relatedness.png/1024px-Coef
ficient_of_relatedness.png
Degree of
relationship
Relationship
Coefficient of
relationship (r)
0
identical twins; clones
100%[4]
1
parent-offspring[5]
50% (2−1)
2
full siblings
50% (2−2+2−2)
2
3/4 siblings or sibling-cousins
37.5% (2−2+2⋅2−4)
2
grandparent-grandchild
25% (2−2)
2
half siblings
25% (2−2)
3
aunt/uncle-nephew/niece
25% (2⋅2−3)
4
double first cousins
25% (2−3+2−3)
3
great grandparent-great grandchild
12.5% (2−3)
4
first cousins
12.5% (2⋅2−4)
6
quadruple second cousins
12.5% (8⋅2−6)
6
triple second cousins
9.38% (6⋅2−6)
4
half-first cousins
6.25% (2−4)
5
first cousins once removed
6.25% (2⋅2−5)
6
double second cousins
6.25% (4⋅2−6)
6
second cousins
3.13% (2−6+2−6)
8
third cousins
0.78% (2⋅2−8)
10
fourth cousins
0.20% (2⋅2−10)[6]
Soubor:Koeficientpribuznosti.png
coefficient of relationship:

dependence on degree of relationship between donor and recipient
(= on probability they share genes)
Hamilton´s rule:
rb > c
r = relationship; b = benefit; c = cost
relation between relationship and group selection:

Eusociality:
Hymenoptera
Isoptera (termites)
mammals: naked mole-rat (Heterocephalus glaber),
Fucomys mole-rats (Bathyergidae)
Florida scrub jay (Aphelocoma coerulescens)
(Florida): c = 7%, b = 14%
GLABER
H. glaber
Fukomys sp.
http://www.xterm-sumitomo.com.au/sites/default/files/images/termite_hq.gif

INTRAGENOMIC CONFLICT
conflict between individuals within populations
conflict between relatives (siblings, mother – descendant)
conflict between males and females (sexual selection)
cooperation and conflict at the genomic level:
George Williams:
body mortal ´ genes (almost) immortal
„gene view“
Image:George C. Williams.jpg

Richard Dawkins:

the term selfish gene (book The Selfish Gene, 1976):
body only as a vehicle for spreading replicators (genes)
which cannot spread on their own
therefore selection affects genes rather than the whole organism
genes must cooperate (the eight analogy)
BUT! the term „selfish“ must be understood as a metaphor!
sometimes some genetic element behaves „unfair“
 ® ultraselfish DNA

Aa
A
a
50%
50%
Gregor Mendel
Law of segregation

Intragenomic conflict results in higher frequency of some
   genomic elements in the next generation
Aa
A
a
95%
5%
segregation (transmission) distortion
Gregor Mendel
?!
drive
drag

Intragenomic conflict may have many forms, eg.:

Interference
  = prevention of transmission of an alternative allele
Gonotaxis
  = preferential transmission to germinal lineage
Overreplication
  eg. transposons
MEIOTIC DRIVE
D
driver
interference: normal allele eliminated from transmission to the next generation
D
D
gonotaxis: mutant allele gets to ovum whereas normal allele gets to the polar body
ovum
polar body

Interference
1. Autosomal
SD (segregation distorters) genes:
males Drosophila melanogaster
preferential transmission 95–99%
distorter and responder
spermatogenetic block in cells with
disabled allele
often emergence of modifiers
SD genes = „outlaw genes“
„Spore killers“ (sk genes):
Neurospora crassa

thaplotyp
t haplotype:
male house mouse
~ proximal third of Chromosome 17
preferential transmission 95–99%
4 paracentric inversions Þ recombination only 2%
responder + several distorters
t/t males sterile Þ more than 15 lethal genes

diverse genetic structure leads to different drive results:
both chromosomes normal Þ 1:1 segregation
highest transmission distortion when complete t haplotype and normal chromosome combination
respoder
distorters
Ó Vesmír 2006/12

TRD mechanism different from drosophila:
responder = Smok (fused gene)
regulation of gene cascade involved
in flagellum formation
Ó Vesmír 2006/12
distorter
responder

2. Maternal-effect killers
´
♀ M/+       ♂ +/+
M/+      +/+
Medea gene:
Maternal-Effect Dominant Embryonic Arrest
Tribolium castaneum
mother M/+
the gene eliminates all descendants who
do not possess it – the +/+ individuals die
in the second larval instar

3. Sex-biased inheritance
uniparentally inherited genes are interested just in reproduction of
the particular sex Þ sex ratio distortion
X chromosome drive Þ female-biased sex ratio Þ selection will favour
return to the original state
cytoplasmic male sterility (CMS)
in 5-10% populations of monoecious plants
mixed populations with sterile male plants
this sterility caused by mutant mitochondrial
genome
advantage when the plants with sterile male sex invest more to pollen
than to seeds Þ transmission of more mitochondria
chromosome%20X%20et%20Y

male
female
if mother has 1 son and 1 daughter number of copies of her mtDNA remains the same
´
´
´
´
´
mtDNA
CMS

similar effect is caused by Wolbachia
intracellular parasite of arthropods
killes males who do not possess Wolbachia
reduction of competition for sources –
   kin selection
besides killing males Wolbachia can have other phenotypic effects:
feminisation: infected males are developing as females or infertile
pseudofemales
parthenogenesis: eg. in Trichogramma wasps males rare (likely due to
wolbachias) ® wolbachias help females to reproduce
parthenogenetically, ie. without males
cytoplasmic incompatibility: inability of males with wolbachias to reproduce
with females which does not possess them or which have wolbachias
of other strain ® reproductive barier, speciation

Overreplication
Transposable elements (transposons)
incorporating of copies to other genome site
(Barbara McClintock: „jumping genes“ in maize)
usually not removed from genome
® molecular fossils
usually huge numbers
human: > half of genome
horizontal transfer, also between species
in some cases effect on gene regulation

1. DNA elements
„cut-and-paste“
enzyme transposase
Ac a Ds elements in maize (B. McClintock), mariner in animals,
P elements in Drosophila
2. Retroelements
„copy-and-paste“
through RNA stage, reverse transcription (reverse transcriptase)
template stays at the original place Þ increase of copy numbers
novel site
DNA
RNA
reverse transcription

Retroelements
LTR-retrotransposons: copia in D. melanogaster
retroposons: LINE – L1 in human: 17% of genome
SINE: short, do not code for own reverse transcriptase
Alu sequence in human – 12% of genome; B1, B2 in mouse
3. MITE (miniature inverted-repeat transposable elements)
 Stowaway, Tourist

gene effects can extend outside organisms –
  R. Dawkins: The Extended Phenotype
Eg.: cases of caddisfly larvae, spider webs
normal
mescaline
LSD
caffeine
marihuana (THC)
amphetamine
chloral hydrate

flukes: parasited individuals bulid thicker shells
Toxoplasma gondii: decrease of host´s reaction time
http://toxoplasmosisinpregnancy.net/wp-content/uploads/2013/06/01.jpg

similarly parasitic flukes:
eg. abdomen of parasited ant Cephalotes atratus turns red
so that resembles edible berry (other species change
ants´ behaviour ® they climb up a grass blade where
they are eaten by cattle or sheeps)
http://stockarch.com/files/13/01/grazing_sheep.jpg

ant Monomorium santschii: absence of workers
® invasion of foreign ant nests, „command“ to kill
own queen and to adopt the invader queen
http://2.bp.blogspot.com/-MSQJxNqsV78/UvZrZxq1bKI/AAAAAAAAEj4/8Yl2UfANcwk/s1600/ant.jpg

http://www.hellodaly.com/uploaded_images/ants-730589.jpg
Duke of Burgundy (Hamearis lucina) caterpillars:
on head an organ producing a narcotic nectar; another pair
of glands causing increased aggressiveness against
all organisms except the carerpillar itself ®
protection („bodyguard“), several days of ants´drug
addiction, ants do not leave the caterpillar
http://www.kolumbus.fi/~kr5298/lnel/lyc/Hamlucin.jpg
https://encrypted-tbn0.gstatic.com/images?q=tbn:ANd9GcTOpMWVbUsy5wjXj3Krh7ACvzXvNcpB3rOJ_ugSQ3lMvwp
ArC7u7g http://cdn.phys.org/newman/gfx/news/hires/2009/1-orphanarmyan.jpg Výsledek obrázku pro
Hamearis lucina larva image