mitos, monstrous y dioses - eva.fcien.udelar.edu.uy
TRANSCRIPT
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Origen de los eucariotas: Mitos, Monstrous y Dioses
29 Junio, 2021
Curso Genómica Evolutiva
Fernando Alvarez
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Last Universal Common Ancestor
LUCA
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(FUCA)
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Archeabacteria
(Archeabacteria)
Woese’s group showed that prokaryotes are composed of two very distantly related groups, which they named Eubacteria and Archaebacteria, now synonymous with Bacteria and Archaea
(Eubacteria)
Phylogenetic structure of the prokaryotic domain: The primary kingdoms Carl R. Woese and George E. Fox PNAS November 1, 1977 74 (11)
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Rooting the tree
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Poniéndole la raíz al árbol
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Surgimiento de los eucariotas Woese’s group showed that prokaryotes are composed of two very distantly related groups, which they named Eubacteria and Archaebacteria, now synonymous with Bacteria and Archaea
LUCA
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Surgimiento de los eucariotas Árbol filogenético de Carl Woese et al. basado en rRNA. La ubicación de la raíz se realizó usando una duplicación muy antigua, entre el factor de elongación TU (EF-Tu) y el factor de elongación G (EF-G) 1990
LUCA
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Procariotas Eucariotas
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Surgimiento de los eucariotas
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Eucariotas mosaicos genómicos
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Surgimiento de los eucariotas
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Surgimiento de los eucariotas, entrada y adaptación de las mitocondrias (domesticación o esclavización)
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great oxidation event [g.o.e.]),
Aparición de las cianobaterias 14
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OoL (Origin of Life)=LUCA
OoL
OoL
FECA
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Hipótesis del Eocito
Hipótesis alternativas sobre el árbol de la vida
LECA
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Transición de FECA a LECA
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Eucariotas:un mosaico de bacterias y archeas
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La hipótesis del chronocyte (1984) y las ESPs (2002)
E=A+B+C Eukaryote=Archea+Bacteria+Chronocyte
Inclusión de Giardia garantiza diversidad por cosnidrealro “early diverging”
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• Speculations Sci Technol. 1984;7(2):77-81. 19
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Painting by Peter Paul Rubens of Cronus devouring one of his children
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La hipótesis del chronocyte (1984) y las ESPs (2002)
E=A+B+C Eukaryote=Archea+Bacteria+Chronocyte
Inclusión de Giardia garantiza diversidad por cosnidrealro “early diverging”
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Cromatina, interacciones proteínas-ADN
Poro nuclear
Metabolismo ARN
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FECA y LECA
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Cuando surgen las mitocondrias?
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Phylogeny of Eukarya
Euglenozoa
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Archezoa:
Eucariotas ancestrales (aun existentes) surgidos antes de la incorporación de las mitocondrias If the mitochondrion, like the plastid, originated by an endosymbiotic event, then it was also possible that some amitochondrial eukaryotes diverged prior to this event. This means that LECA no tendría mitocondrias.
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Fitting the expectations! This phylogenetic evidence fulfilled the most basic prediction
of the Archezoa hypothesis for these groups: if they
predate the origin of the mitochondrion, then they must
branch earlier than mitochondrion-containing eukaryotes in
phylogenetic trees.
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Se cae la hipótesis de archezoa
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More substantial evidence came from a direct search of the Entamoeba genome for molecular relics of the mitochondrial symbiont. Most of the many hundreds of mitochondrial protein-coding genes are encoded in the nucleus and targeted to the organelle after they are translated in the cytoplasm.
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Val-tRNA-S trees obtained with the ML method of protein phylogeny (JTT-F model) 40
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Hydogenosomas de parabasalia
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Características de LECA Inferidas a partir de la diversidad de eucariotas
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Surgimiento de los eucariotas
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Mirando en dirección de las Archeas: la hipótesis del ecocito de Lake
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Dos topologías alternativas usando rRNA
Eocyte
Lake 1984, Lake et al 1988 Topología de Woese
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Hipotesis del eocito (Lake 1984)
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Hipotesis del eocito (Lake 1984)
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Los eucariotas presentan membranas de tipo bacteriano
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Incremento de densidad filogenetica:TACK superfilum
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TACK superphylum, a clade originally comprising the archaeal phyla Thaumarchaeota, Aigarchaeota, Crenarchaeota and Korarchaeota23. In support of this relationship, comparative genomics analyses have revealed several eukaryotic signature proteins (ESPs)24 in TACK lineages
(nanoarchaea or ultra-small archaea)
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Filogenia con 16S
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Filogenia usando rRNA 16s
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Enviromental data handling
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Amplificado No amplificado
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Filogenia usando 36 genes concatenados
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Composicion de genes (usando blastp de los ensamblajes)
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Miscellaneus Crenacheota Group
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The endosomal sorting complexes required for transport (ESCRT) machinery is made up of cytosolic protein complexes, known as ESCRT-0, ESCRT-I, ESCRT-II, and ESCRT-III.
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Distribución de proteínas ESPs
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Distribución de proteínas ESPs
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La densidad filogenética acorrala a los eucariotas
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Filogenia rRNA 16S
ASGARD
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SAGs
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Son buenos estos ensamblajes?
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SAGs=Single cell Amplified Genomes
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Distribución de proteínas ESPs
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Contexto filogenético del origen de los eucariotas
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Reconstrucción de una historia intrincada
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Isolation of an archaeon at the prokaryote-eukaryote interface
Hiroyuki Imachi et al, BioXchi
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We propose an “Entangle-Engulf-Enslave (E3) model” for eukaryogenesis through archaea-alphaproteobacteria symbiosis mediated by the physical complexities and metabolic dependency of the hosting archaeon.
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