12. Gen´etica evolutiva

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Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas

12. Gen´ etica evolutiva

Fundamentos de Gen´ etica Grado en Bioqu´ımica Universidad de Granada

Prof. ´ Angel Mart´ın Alganza ([email protected]) Departamento de Gen´ etica

Gen´ etica, Universidad de Granada 12. Gen´ etica evolutiva

(2)

Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas

12. Gen´ etica evolutiva

1 Dimensiones del proceso evolutivo Anag´ enesis y cladog´ enesis

Microevoluci´ on y macroevoluci´ on

2 La variaci´ on gen´ etica y la selecci´ on natural

La variaci´ on gen´ etica es el ((sustrato)) de la evoluci´on La selecci´ on natural es el ((motor)) de la evoluci´on

3 El proceso de especiaci´ on Concepto biol´ ogico de especie Modos de especiaci´ on

Mecanismos de aislamiento reproductivo Modelo general de especiaci´ on

4 Evoluci´ on molecular

Reconstrucci´ on de filogenias moleculares Relojes moleculares

5 Teor´ıas evolutivas

La selecci´ on en acci´ on

(3)

Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas AnaClado MicroMacro

12. Gen´ etica evolutiva

1 Dimensiones del proceso evolutivo Anag´ enesis y cladog´ enesis

Microevoluci´ on y macroevoluci´ on

2 La variaci´ on gen´ etica y la selecci´ on natural

La variaci´ on gen´ etica es el ((sustrato)) de la evoluci´on La selecci´ on natural es el ((motor)) de la evoluci´on

3 El proceso de especiaci´ on Concepto biol´ ogico de especie Modos de especiaci´ on

Mecanismos de aislamiento reproductivo Modelo general de especiaci´ on

4 Evoluci´ on molecular

Reconstrucci´ on de filogenias moleculares Relojes moleculares

5 Teor´ıas evolutivas La selecci´ on en acci´ on

Gen´ etica, Universidad de Granada 12. Gen´ etica evolutiva

(4)

Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas AnaClado MicroMacro

Dimensiones del proceso evolutivo

El proceso evolutivo tiene dos dimensiones:

Anag´ enesis o evoluci´ on fil´ etica: dentro de una l´ınea evolutiva a lo largo del tiempo; debida a los procesos evolutivos

Cladog´ enesis o diversificaci´ on: cuando una

l´ınea filogen´ etica se diferencia

en dos (especiaci´ on y aparici´ on

de taxones superioes)

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Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas AnaClado MicroMacro

Estasis, anag´ enesis y cladog´ enesis

Gen´ etica, Universidad de Granada 12. Gen´ etica evolutiva

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Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas AnaClado MicroMacro

Microevoluci´ on versus macroevoluci´ on

Los procesos o mecanismos evolutivos son los responsables de los cambios que se observan a lo largo de la evoluci´ on

Seg´ un la escala o el nivel en que se analicen los procesos evolutivos, pueden clasificarse en:

Microevoluci´ on Conjunto de procesos evolutivos que ocurren dentro de una especie (peque˜ na escala)

Macroevoluci´ on Conjunto de procesos evolutivos que ocurren

por encima del nivel de especie (gran escala)

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Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas Variaci´ on Selecci´ on

12. Gen´ etica evolutiva

1 Dimensiones del proceso evolutivo Anag´ enesis y cladog´ enesis

Microevoluci´ on y macroevoluci´ on

2 La variaci´ on gen´ etica y la selecci´ on natural

La variaci´ on gen´ etica es el ((sustrato)) de la evoluci´on La selecci´ on natural es el ((motor)) de la evoluci´on

3 El proceso de especiaci´ on Concepto biol´ ogico de especie Modos de especiaci´ on

Mecanismos de aislamiento reproductivo Modelo general de especiaci´ on

4 Evoluci´ on molecular

Reconstrucci´ on de filogenias moleculares Relojes moleculares

5 Teor´ıas evolutivas La selecci´ on en acci´ on

Gen´ etica, Universidad de Granada 12. Gen´ etica evolutiva

(8)

Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas Variaci´ on Selecci´ on

La variaci´ on gen´ etica es el ((sustrato)) de la evoluci´on

La variaci´ on gen´ etica

Producida por mutaci´ on y recombinaci´ on Es el sustrato de la evoluci´ on

M´ etodos de cuantificaci´ on

Grado de polimorfismo (proporci´ on de loci polim´ orficos) Heterocigosidad (frecuencia media de heterocigotos) T´ ecnicas para detectar variaci´ on gen´ etica

Isoenzimas

RFLPs

VNTRs

Secuenciaci´ on

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Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas Variaci´ on Selecci´ on

La selecci´ on natural es el ((motor)) de la evoluci´on

La selecci´ on natural

Es el proceso evolutivo m´ as ((importante))

Responsable de la mayor parte de los fenotipos por adaptaci´ on Act´ ua a niveles inferiores (genes, c´ elulas, individuos)

Tiene efectos en los niveles superiores (reforzamiento MARs)

Gen´ etica, Universidad de Granada 12. Gen´ etica evolutiva

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Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas Especie Modos MARs Modelo

12. Gen´ etica evolutiva

1 Dimensiones del proceso evolutivo Anag´ enesis y cladog´ enesis

Microevoluci´ on y macroevoluci´ on

2 La variaci´ on gen´ etica y la selecci´ on natural

La variaci´ on gen´ etica es el ((sustrato)) de la evoluci´on La selecci´ on natural es el ((motor)) de la evoluci´on

3 El proceso de especiaci´ on Concepto biol´ ogico de especie Modos de especiaci´ on

Mecanismos de aislamiento reproductivo Modelo general de especiaci´ on

4 Evoluci´ on molecular

Reconstrucci´ on de filogenias moleculares Relojes moleculares

5 Teor´ıas evolutivas

La selecci´ on en acci´ on

(11)

Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas Especie Modos MARs Modelo

Concepto biol´ ogico de especie

Grupo de poblaciones de individuos de la misma especie que se pueden cruzar real o potencialmente y que en la naturaleza est´ an reproductivamente aisladas de todos los dem´ as grupos.

Gen´ etica, Universidad de Granada 12. Gen´ etica evolutiva

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Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas Especie Modos MARs Modelo

Modos de especiaci´ on

Alop´ atrica Divergencia evolutiva de poblaciones separadas Vicariante Barrera de separaci´ on entre poblaciones Perip´ atrica Aislamiento de una peque˜ na poblaci´ on Parap´ atrica Divergencia evolutiva de poblaciones adyacentes

Simp´ atrica Divergencia evolutiva en el seno de una poblaci´ on Instant´ anea Por hibridaci´ on y/o poliploid´ıa Cromos´ omica Por aparici´ on de reordenaciones

Gradual Producida por selecci´ on disruptiva

(13)

Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas Especie Modos MARs Modelo

Mecanismos de aislamiento reproductivo

Gen´ etica, Universidad de Granada 12. Gen´ etica evolutiva

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Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas Especie Modos MARs Modelo

Especiaci´ on incipiente en poblaciones de D. pseudoobscura

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Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas Especie Modos MARs Modelo

Modelo general de especiaci´ on

Etapa I (reversible)

1. Interrupci´ on del flujo g´ enico

2. Acumulaci´ on de diferencias gen´ eticas (mecanismos evolutivos) 3. Aparici´ on de potenciales MARs postcig´ oticos

Etapa II (irreversible)

1. Restablecimiento del flujo g´ enico

2. Aparici´ on de MARs precig´ oticos por reforzamiento del aislamiento reproductivo por parte de la selecci´ on natural

Gen´ etica, Universidad de Granada 12. Gen´ etica evolutiva

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Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas Filogenias RM

12. Gen´ etica evolutiva

1 Dimensiones del proceso evolutivo Anag´ enesis y cladog´ enesis

Microevoluci´ on y macroevoluci´ on

2 La variaci´ on gen´ etica y la selecci´ on natural

La variaci´ on gen´ etica es el ((sustrato)) de la evoluci´on La selecci´ on natural es el ((motor)) de la evoluci´on

3 El proceso de especiaci´ on Concepto biol´ ogico de especie Modos de especiaci´ on

Mecanismos de aislamiento reproductivo Modelo general de especiaci´ on

4 Evoluci´ on molecular

Reconstrucci´ on de filogenias moleculares Relojes moleculares

5 Teor´ıas evolutivas

La selecci´ on en acci´ on

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Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas Filogenias RM

Evoluci´ on de la vida en La Tierra

From Teaching About Evolution and the Nature of Science, 1998, The National Academies Press, https://www.nap.edu/read/5787/

Gen´ etica, Universidad de Granada 12. Gen´ etica evolutiva

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Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas Filogenias RM

Filogenia construida usando la secuencia del citocromo c

(19)

Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas Filogenias RM

Reconstrucci´ on filogen´ etica mediante UPGMA

Emparejamiento no ponderado utilizando medias aritm´ eticas

Gen´ etica, Universidad de Granada 12. Gen´ etica evolutiva

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Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas Filogenias RM

Reloj molecular

del gen que codifica para la hematoglutinina del virus de la gripe A

(21)

Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas Filogenias RM

Relaciones evolutivas

de mitocondrias, α-proteobacterias, cianobacterias y cloroplastos

Gen´ etica, Universidad de Granada 12. Gen´ etica evolutiva

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0.4

Candidate Phyla Radiation

Microgenomates Parcubacteria

Eukaryotes

Archaea Bacteria

DPANN

Opisthokonta

Amoebozoa Chromalveolata Archaeplastida Excavata RBX1

WOR1 Cyanobacteria

Melainabacteria

PVC superphylum

TACK

Major lineage lacking isolated representative:

Major lineages with isolated representative: italics Dojkabacteria WS6

Peregrinibacteria Gracilibacteria BD1-5, GN02Absconditabacteria SR1

Katanobacteria WWE3

Berkelbacteria SM2F11

CPR1 CPR3 Nomurabacteria Kaiserbacteria

Adlerbacteria Campbellbacteria

Wirthbacteria Chloroflexi

Armatimonadetes

GiovannonibacteriaWolfebacteria Jorgensenbacteria

Azambacteria Yanofskybacteria

Moranbacteria Magasanikbacteria Uhrbacteria Falkowbacteria

Saccharibacteria

Woesebacteria Amesbacteria Shapirobacteria Collierbacteria Pacebacteria Beckwithbacteria Roizmanbacteria Gottesmanbacteria Levybacteria Daviesbacteria Curtissbacteria

Nanoarchaeota Woesearchaeota Pacearchaeota Nanohaloarchaeota

Micrarchaeota

Altiarchaeales Aenigmarchaeota

Diapherotrites

Z7ME43 Loki.

Thaumarchaeota Archaeoglobi Methanomicrobia

Halobacteria

Thermoplasmata Methanococci Spirochaetes

Firmicutes (Tenericutes)

Bacteroidetes Chlorobi

Gammaproteobacteria Alphaproteobacteria

Betaproteobacteria Actinobacteria

Planctomycetes Chlamydiae, Lentisphaerae, Verrucomicrobia

Omnitrophica AminicentantesRokubacteriaNC10

Elusimicrobia Poribacteria

Ignavibacteria

Dadabacteria

TM6 Atribacteria Gemmatimonadetes

Cloacimonetes Fibrobacteres

Nitrospirae Latescibacteria TA06

Caldithrix Marinimicrobia

WOR-3 Zixibacteria

Synergistetes Fusobacteria Aquificae Calescamantes

Deinococcus-Therm.

Caldiserica Dictyoglomi

Deltaprotebacteria(Thermodesulfobacteria)

Epsilonproteobacteria DeferribacteresChrysiogenetes Tectomicrobia, ModulibacteriaNitrospinae

Acidobacteria

Zetaproteo.

Thermotogae

Acidithiobacillia

Parvarchaeota Hydrogenedentes NKB19

Thor.

BRC1

Thermococci Methanobacteria Hadesarchaea Methanopyri

Aigarch.

Crenarch.

YNPFFA Korarch.

Bathyarc.

Figure 1 | A current view of the tree of life, encompassing the total diversity represented by sequenced genomes. The tree includes 92 named bacterial phyla, 26 archaeal phyla and allﬁve of the Eukaryotic supergroups. Major lineages are assigned arbitrary colours and named, with well-characterized lineage names, in italics. Lineages lacking an isolated representative are highlighted with non-italicized names and red dots. For details on taxon sampling and tree inference, see Methods. The names Tenericutes and Thermodesulfobacteria are bracketed to indicate that these lineages branch within the Firmicutes and the Deltaproteobacteria, respectively. Eukaryotic supergroups are noted, but not otherwise delineated due to the low resolution of these lineages. The CPR phyla are assigned a single colour as they are composed entirely of organisms without isolated representatives, and are still in the process of deﬁnition at lower taxonomic levels. The complete ribosomal protein tree is available in rectangular format with full bootstrap values as Supplementary Fig. 1 and in Newick format in Supplementary Dataset 2.

LETTERS

NATURE MICROBIOLOGYDOI: 10.1038/NMICROBIOL.2016.48

NATURE MICROBIOLOGY| VOL 1 | MAY 2016 |www.nature.com/naturemicrobiology 2

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0.4

Candidate Phyla Radiation

Microgenomates Parcubacteria

Eukaryotes

Archaea Bacteria

DPANN

Opisthokonta

Amoebozoa Chromalveolata Archaeplastida Excavata RBX1

WOR1 Cyanobacteria

Melainabacteria

PVC superphylum

TACK

Major lineage lacking isolated representative:

Major lineages with isolated representative: italics Dojkabacteria WS6

Peregrinibacteria Gracilibacteria BD1-5, GN02Absconditabacteria SR1

Katanobacteria WWE3

Berkelbacteria SM2F11

CPR1 CPR3 Nomurabacteria Kaiserbacteria

Adlerbacteria Campbellbacteria

Wirthbacteria Chloroflexi

Armatimonadetes

GiovannonibacteriaWolfebacteria Jorgensenbacteria

Azambacteria Yanofskybacteria

Moranbacteria Magasanikbacteria Uhrbacteria Falkowbacteria

Saccharibacteria

Woesebacteria Amesbacteria Shapirobacteria Collierbacteria Pacebacteria Beckwithbacteria Roizmanbacteria Gottesmanbacteria Levybacteria Daviesbacteria Curtissbacteria

Nanoarchaeota Woesearchaeota Pacearchaeota Nanohaloarchaeota

Micrarchaeota

Altiarchaeales Aenigmarchaeota

Diapherotrites

Z7ME43 Loki.

Thaumarchaeota Archaeoglobi Methanomicrobia

Halobacteria

Thermoplasmata Methanococci Spirochaetes

Firmicutes (Tenericutes)

Bacteroidetes Chlorobi

Gammaproteobacteria Alphaproteobacteria

Betaproteobacteria Actinobacteria

Planctomycetes Chlamydiae, Lentisphaerae, Verrucomicrobia

Omnitrophica AminicentantesRokubacteriaNC10

Elusimicrobia Poribacteria

Ignavibacteria

Dadabacteria

TM6 Atribacteria Gemmatimonadetes

Cloacimonetes Fibrobacteres

Nitrospirae Latescibacteria TA06

Caldithrix Marinimicrobia

WOR-3 Zixibacteria

Synergistetes Fusobacteria Aquificae Calescamantes

Deinococcus-Therm.

Caldiserica Dictyoglomi

Deltaprotebacteria(Thermodesulfobacteria)

Epsilonproteobacteria DeferribacteresChrysiogenetes Tectomicrobia, ModulibacteriaNitrospinae

Acidobacteria

Zetaproteo.

Thermotogae

Acidithiobacillia

Parvarchaeota Hydrogenedentes NKB19

Thor.

BRC1

Thermococci Methanobacteria Hadesarchaea Methanopyri

Aigarch.

Crenarch.

YNPFFA Korarch.

Bathyarc.

Figure 1 | A current view of the tree of life, encompassing the total diversity represented by sequenced genomes. The tree includes 92 named bacterial phyla, 26 archaeal phyla and allﬁve of the Eukaryotic supergroups. Major lineages are assigned arbitrary colours and named, with well-characterized lineage names, in italics. Lineages lacking an isolated representative are highlighted with non-italicized names and red dots. For details on taxon sampling and tree inference, see Methods. The names Tenericutes and Thermodesulfobacteria are bracketed to indicate that these lineages branch within the Firmicutes and the Deltaproteobacteria, respectively. Eukaryotic supergroups are noted, but not otherwise delineated due to the low resolution of these lineages. The CPR phyla are assigned a single colour as they are composed entirely of organisms without isolated representatives, and are still in the process of deﬁnition at lower taxonomic levels. The complete ribosomal protein tree is available in rectangular format with full bootstrap values as Supplementary Fig. 1 and in Newick format in Supplementary Dataset 2.

LETTERS

0.2

Korarchaeota Diapherotrites

Nanohaloarchaeota Unclassified archaea

Pacearchaeota Woesearchaeota, Nanoarchaeota

Woesearchaeota Altiarchaeales Z7ME43

Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciArchaeoglobi, Methanomicrobia, Halobacteria Aciduliprofundum, Thermoplasmata Uncultured Thermoplasmata

Thermoplasmata

Opisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa

Crenarchaeota Crenarchaeota Thorarchaeota Lokiarchaeota

YNPFFA Thaumarchaeota Thaumarchaeota Cyanobacteria, Melainabacteria

Dojkabacteria WS6 CPR3

Katanobacteria WWE3 Katanobacteria WWE3

Microgenomates Roizmanbacteria Microgenomates Roizmanbacteria Microgenomates Microgenomates Curtissbacteria

Microgenomates Daviesbacteria

Microgenomates Levybacteria Microgenomates Woesebacteria Microgenomates Amesbacteria

Microgenomates Shapirobacteria Microgenomates Beckwithbacteria, Pacebacteria, Collierbacteria

Microgenomates Gottesmanbacteria KAZAN

CPR2, Saccharibacteria TM7 Berkelbacteria

Berkelbacteria Berkelbacteria Berkelbacteria CPR Uncultured unclassified bacteria

Peregrinibacteria Peregrinibacteria

Absconditabacteria SR1 Gracilibacteria BD1-5 / GNO2

SM2F11

ParcubacteriaParcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, Magasanikbacteria Parcubacteria

Parcubacteria Parcubacteria

Parcubacteria

Parcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, Kaiserbacteria Parcubacteria

Parcubacteria Moranbacteria Parcubacteria Parcubacteria Yanofskybacteria Deinococcus-Thermus

Aquificae, Calescamantes EM19 Caldiserica, Dictyoglomi Thermotogae

Omnitrophica Omnitrophica

Spirochaetes Spirochaetes Hydrogenedentes NKB19

Deltaproteobacteria Epsilonproteobacteria TM6 Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres

Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot.

NC10, Rokubacteria, Aminicenantes, Acidobacteria Planctomycetes Chlamydiae Lentisphaerae

Verrucomicrobia Verrucomicrobia RBX-1 WOR-1

Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes

Uncultured bacteria (CP RIF32)

Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres

Cloacamonetes Atribacteria (OP9) BRC1, Poribacteria Latescibacteria WS3

Gemmatimonadetes, WOR-3, TA06

Elusimicrobia Uncultured bacteria

Uncultured bacteria (CP RIF1) Aigarchaeota, Cand. Caldiarchaeum subterraneum Unclassified archaea

Parcubacteria

Candidate Phyla Radiation Cyanobacteria, Melainabacteria

Deinococcus-Thermus Aquificae, Calescamantes EM19 Caldiserica, Dictyoglomiqq ,, ThermotogaeAifi C

Omnitrophica Omnitrophicapp

Spirochaetes Spirochaetes S iht Hydrogenedentes NKB19 Deltaproteobacteria

H d dtN

Epsilonproteobacteria b

TM6 Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres

Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot.

NC10, Rokubacteria, Aminicenantes, Acidobacteria D fb D fb

, , p,

Planctomycetespp Chlamydiaey Lentisphaerae ChC a yd

Verrucomicrobia Verrucomicrobiapp RBX-1 WOR-1

Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes

Uncultured bacteria (CP RIF32), y, ygg

Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres

Cloacamonetes Atribacteria (OP9) BRC1, Poribacteria( ) Latescibacteria WS3

Gemmatimonadetes, WOR-3, TA06b M

Elusimicrobia Uncultured bacteria

Uncultured bacteria (CP RIF1)

O h

Dojkabacteria WS6 CPR3

Katanobacteria WWE3 Katanobacteria WWE3

Microgenomates Roizmanbacteria Microgenomates Roizmanbacteria MicrogenomatesMicrogenomates Curtissbacteriagg

Microgenomates Daviesbacteriagg

Microgenomates Levybacteria Microgenomates Woesebacteria Microgenomates Amesbacteria

Mi tLbt i

Microgenomates Shapirobacteria

Mi tW b

Mi t

Microgenomates Beckwithbacteria, Pacebacteria, Collierbacteria

Mi Shi b i

Microgenomates Gottesmanbacteria

tR i bt i

g y

KAZAN CPR2, Saccharibacteria TM7 Berkelbacteria

Berkelbacteria Berkelbacteria Berkelbacteria CPR Uncultured unclassified bacteria

Peregrinibacteria Peregrinibacteria

Absconditabacteria SR1 Gracilibacteria BD1-5 / GNO2

SM2F11

Parcubacteriate aParcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, Magasanikbacteria Parcubacteria

Parcubacteria Parcubacteria

Parcubacteria AbscAbs

Parcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, ggCampbellbacteria, Adlerbacteria, Kaiserbacteria Parcubacteria

Parcubacteria Moranbacteria Parcubacteria Parcubacteria Yanofskybacteria

P b i

Candidate Phyla Radiation Diapherotrites

Nanohaloarchaeota Unclassified archaea

Pacearchaeota Woesearchaeota, Nanoarchaeota

Woesearchaeota Altiarchaeales Z7ME43

Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, Thermococci E43

Archaeoglobi, Methanomicrobia, Halobacteria,, ,, ,, Aciduliprofundum, Thermoplasmatagg Uncultured Thermoplasmatapp ,,

Thermoplasmatap Unclassified archaea

Korarchaeota

,

Crenarchaeota Crenarchaeota Thorarchaeota Lokiarchaeota

YNPFFA Thaumarchaeota Thaumarchaeota

b lb

Aigarchaeota, FFA Cand. Caldiarchaeum subterraneum

C bt iM l ibt i

Opisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa

Th h ,

Th ht Eukaryotes

Bacteria Archaea

Katanobacteria WWE3

Bootstrap ≥ 85%

85% > Bootstrap ≥ 50%

Woesearchaeota, Nanoarchaeota

Figure 2 | A reformatted view of the tree in Fig. 1 in which each major lineage represents the same amount of evolutionary distance. The threshold for groups (coloured wedges) was an average branch length of <0.65 substitutions per site. Notably, some well-accepted phyla become single groups and others are split into multiple distinct groups. We undertook this analysis to provide perspective on the structure of the tree, and do not propose the resulting groups to have special taxonomic status. The massive scale of diversity in the CPR and the large fraction of major lineages that lack isolated representatives (red dots) are apparent from this analysis. Bootstrap support values are indicated by circles on nodes—black for support of 85% and above, grey for support from 50 to 84%. The complete ribosomal protein tree is available in rectangular format with full bootstrap values as Supplementary Fig. 1 and in Newick format in Supplementary Dataset 2.

LETTERS

(24)

Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas Filogenias RM

Relaciones evolutivas y simbi´ oticas

By Maulucioni y Dorid´ı - Own work, CC BY-SA 3.0,

htps://commons.wikimedia.org/w/index.php?curid=25888693

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Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas La selecci´ on en acci´ on

12. Gen´ etica evolutiva

1 Dimensiones del proceso evolutivo Anag´ enesis y cladog´ enesis

Microevoluci´ on y macroevoluci´ on

2 La variaci´ on gen´ etica y la selecci´ on natural

La variaci´ on gen´ etica es el ((sustrato)) de la evoluci´on La selecci´ on natural es el ((motor)) de la evoluci´on

3 El proceso de especiaci´ on Concepto biol´ ogico de especie Modos de especiaci´ on

Mecanismos de aislamiento reproductivo Modelo general de especiaci´ on

4 Evoluci´ on molecular

Reconstrucci´ on de filogenias moleculares Relojes moleculares

5 Teor´ıas evolutivas La selecci´ on en acci´ on

Gen´ etica, Universidad de Granada 12. Gen´ etica evolutiva

(26)

Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas La selecci´ on en acci´ on

Teor´ıas evolutivas

Herencia de los caracteres adquiridos (Lamark) Evoluci´ on por selecci´ on natural (Darwin y Wallace) Teor´ıa sint´ etica de la evoluci´ on (Fisher, Haldane, Wright) Microevoluci´ on Cambios gen´ eticos graduales en poblaciones Macroevoluci´ on Sucesos de especiaci´ on

Teor´ıa neutralista de la evoluci´ on molecular (Kimura) Teor´ıa del equilibrio puntuado (Eldredge y Gould) Gradualismo extremo (Dawkins)

Teor´ıa sint´ etica extendida de la evoluci´ on (Pigliucci, Noble)

(27)

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Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas La selecci´ on en acci´ on

Lo que puede hacer la selecci´ on (artificial)

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Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas La selecci´ on en acci´ on

La selecci´ on natural en acci´ on

Gen´ etica, Universidad de Granada 12. Gen´ etica evolutiva

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Dimensiones VarSel Especiaci´ om EvolMol Teor´ıas La selecci´ on en acci´ on

La selecci´ on sexual en acci´ on

(31)

Nothing in Biology Makes Sense Except in the Light of Evolution

THEODOSIUS DOBZHANSKY

As

RECENTLY AS 1966, sheik Abd el Aziz bin Baz asked the king of Saudi Arabia to suppress a heresy that was spreading in his land. Wrote the sheik:

"The Holy Koran, the Prophet's teachings, the ma- jority of Islamic scientists, and the actual facts all prove that the sun is running in its orbit ... and that the earth is fixed and stable, spread out by God for his mankind. ... Anyone who professed otherwise would utter a charge of falsehood toward God, the Koran, and the Prophet."

The good sheik evidently holds the Copernican theory to be a "mere theory," not a "fact." In this he is technically correct. A theory can be verified by a mass of facts, but it becomes a proven theory, not a fact. The sheik was perhaps unaware that the Space Age had begun before he asked the king to suppress the Copernican heresy. The sphericity of the earth had been seen by astronauts, and even by many earth-bound people on their television screens.

Perhaps the sheik could retort that those who ven- ture beyond the confines of God's earth suffer hal- lucinations, and that the earth is really flat.

Parts of the Copernican world model, such as the

contention that the earth rotates around the sun, and not vice versa, have not been verified by direct observations even to the extent the sphericity of the earth has been. Yet scientists accept the model as an accurate representation of reality. Why? Because it makes sense of a multitude of facts which are other- wise meaningless or extravagant. To nonspecialists most of these facts are unfamiliar. Why then do we accept the "mere theory" that the earth is a sphere revolving around a spherical sun? Are we simply submitting to authority? Not quite: we know that those who took time to study the evidence found it convincing.

The good sheik is probably ignorant of the evi- dence. Even more likely, he is so hopelessly biased that no amount of evidence would impress him. Any- way, it would be sheer waste of time to attempt to convince him. The Koran and the Bible do not con- tradict Copernicus, nor does Copernicus contradict them. It is ludicrous to mistake the Bible and the Koran for primers of natural science. They treat of matters even more important: the meaning of man and his relations to God. They are written in poetic symbols that were understandable to people of the age when they were written, as well as to peoples of all other ages. The king of Arabia did not comply with the sheik's demand. He knew that some people fear enlightenment, because enlightenment threatens their vested interests. Education is not to be used to promote obscurantism.

The earth is not the geometric center of the uni- verse, although it may be its spiritual center. It is a mere speck of dust in cosmic spaces. Contrary to Bishop Ussher's calculations, the world did not ap- pear in approximately its present state in 4004 B.C.

The estimates of the age of the universe given by modern cosmologists are still only rough approxi- mations, which are revised (usually upward) as the methods of estimation are refined. Some cosmol- ogists take the universe to be about 10 billion years old; others suppose that it may have existed, and will continue to exist, eternally. The origin of life on earth is dated tentatively between 3 and 5 billion years ago; manlike beings appeared relatively quite recently, between 2 and 4 million years ago. The estimates of the age of the earth, of the duration of the geologic and paleontologic eras, and of the antiq- uity of man's ancestors are now based mainly on radiometric evidence-the proportions of isotopes of certain chemical elements in rocks suitable for such studies.

125 One of the world's leading geneticists, Theo-

dosius Dobzhansky is professor emeritus, Rockefeller University, and adjunct profes- sor of genetics, University of California, Davis 95616. Born in Russia, in 1900, he is a graduate of the University of Kiev and taught (with J. Philipchenko) at the Uni- versity of Leningrad before coming to the U.S., in 1927; thereafter he taught at Colum- bia University and the California Institute of Technology be- fore joining the Rockefeller faculty, in 1962. He has been president of the Genetics Society of America, the American Society of Naturalists, the Society for the Study of Evolution, the American Society of Zoologists, and the American Teil- hard de Chardin Association. Among his many honors are the National Medal of Science (1964) and the Gold Medal Award for Distinguished Achievement in Science (1969). He holds 18 honorary doctorates from universities in this country and abroad. Among his well-known books are The Biological Basis of Human Freedom (1956) and Mankind Evolving (1963). The present paper was presented at the 1972 NABT convention.

(32)

Nothing in Biology Makes Sense Except in the Light of Evolution

THEODOSIUS DOBZHANSKY

As

RECENTLY AS 1966, sheik Abd el Aziz bin Baz asked the king of Saudi Arabia to suppress a heresy that was spreading in his land. Wrote the sheik:

"The Holy Koran, the Prophet's teachings, the ma- jority of Islamic scientists, and the actual facts all prove that the sun is running in its orbit ... and that the earth is fixed and stable, spread out by God for his mankind. ... Anyone who professed otherwise would utter a charge of falsehood toward God, the Koran, and the Prophet."

The good sheik evidently holds the Copernican theory to be a "mere theory," not a "fact." In this he is technically correct. A theory can be verified by a mass of facts, but it becomes a proven theory, not a fact. The sheik was perhaps unaware that the Space Age had begun before he asked the king to suppress the Copernican heresy. The sphericity of the earth had been seen by astronauts, and even by many earth-bound people on their television screens.

Perhaps the sheik could retort that those who ven- ture beyond the confines of God's earth suffer hal- lucinations, and that the earth is really flat.

Parts of the Copernican world model, such as the

contention that the earth rotates around the sun, and not vice versa, have not been verified by direct observations even to the extent the sphericity of the earth has been. Yet scientists accept the model as an accurate representation of reality. Why? Because it makes sense of a multitude of facts which are other- wise meaningless or extravagant. To nonspecialists most of these facts are unfamiliar. Why then do we accept the "mere theory" that the earth is a sphere revolving around a spherical sun? Are we simply submitting to authority? Not quite: we know that those who took time to study the evidence found it convincing.

The good sheik is probably ignorant of the evi- dence. Even more likely, he is so hopelessly biased that no amount of evidence would impress him. Any- way, it would be sheer waste of time to attempt to convince him. The Koran and the Bible do not con- tradict Copernicus, nor does Copernicus contradict them. It is ludicrous to mistake the Bible and the Koran for primers of natural science. They treat of matters even more important: the meaning of man and his relations to God. They are written in poetic symbols that were understandable to people of the age when they were written, as well as to peoples of all other ages. The king of Arabia did not comply with the sheik's demand. He knew that some people fear enlightenment, because enlightenment threatens their vested interests. Education is not to be used to promote obscurantism.

The earth is not the geometric center of the uni- verse, although it may be its spiritual center. It is a mere speck of dust in cosmic spaces. Contrary to Bishop Ussher's calculations, the world did not ap- pear in approximately its present state in 4004 B.C.

The estimates of the age of the universe given by modern cosmologists are still only rough approxi- mations, which are revised (usually upward) as the methods of estimation are refined. Some cosmol- ogists take the universe to be about 10 billion years old; others suppose that it may have existed, and will continue to exist, eternally. The origin of life on earth is dated tentatively between 3 and 5 billion years ago; manlike beings appeared relatively quite recently, between 2 and 4 million years ago. The estimates of the age of the earth, of the duration of the geologic and paleontologic eras, and of the antiq- uity of man's ancestors are now based mainly on radiometric evidence-the proportions of isotopes of certain chemical elements in rocks suitable for such studies.

125 One of the world's leading geneticists, Theo-

dosius Dobzhansky is professor emeritus, Rockefeller University, and adjunct profes- sor of genetics, University of California, Davis 95616. Born in Russia, in 1900, he is a graduate of the University of Kiev and taught (with J. Philipchenko) at the Uni- versity of Leningrad before coming to the U.S., in 1927; thereafter he taught at Colum- bia University and the California Institute of Technology be- fore joining the Rockefeller faculty, in 1962. He has been president of the Genetics Society of America, the American Society of Naturalists, the Society for the Study of Evolution, the American Society of Zoologists, and the American Teil- hard de Chardin Association. Among his many honors are the National Medal of Science (1964) and the Gold Medal Award for Distinguished Achievement in Science (1969). He holds 18 honorary doctorates from universities in this country and abroad. Among his well-known books are The Biological Basis of Human Freedom (1956) and Mankind Evolving (1963). The present paper was presented at the 1972 NABT convention.

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Erika Aguirre-Planter y Valeria Souza Erika Aguirre-Planter y Valeria Souza Lab. de Evolución Molecular y Experimental, Depto. de Ecología Evolutiva, Instituto de Ecología, Universidad Nacional Autónoma de México. Ciudad Universitaria, C.P. 04510, Deleg. Coyoacán, México, D.F. E-mail: *[email protected]

Nota: Artículo recibido el 01 de febrero de 2013 y aceptado el 11 de marzo de 2013.

ARTÍCULO DE REVISIÓN

R RE S U M E NE S U M E NSS EE

La teoría de la genética de poblaciones surgió hace más de 80 años y nos permite explicar los patrones de variación genética dentro y entre las poblaciones que forman a las especies en términos de las fuerzas evolutivas. Este programa de investigación generó las preguntas que se han abordado empíricamente mediante marcadores moleculares desde hace medio siglo. Una pregunta fundamental ha sido hasta dónde un conjunto reducido de loci es o no representativo del efecto de las fuerzas evolutivas, sobre todo el genoma de una especie. Esto ha llevado al desarrollo creciente de aproximaciones que permitan conocer de manera representativa los niveles de variación genética en las poblaciones naturales, dando origen a la genómica de poblaciones. En años recientes, las técnicas de secuenciación masiva, llamadas Next generation sequencing, o next-gen, han permitido obtener datos de grandes secciones del genoma de diferentes especies, sin que sea un requisito conocer marcadores previos. Así, al comparar los genomas de muchos individuos de diferentes poblaciones, tenemos acceso al archivo de su historia evolutiva, que nos habla del complejo y dinámico balance en el tiempo entre la selección natural y las otras fuerzas evolutivas de carácter neutral, como la deriva y el flujo génico. La existencia de enormes cantidades de información ha requerido el desarrollo de nuevas herramientas estadísticas y bioinformáticas para su análisis.

Diversas disciplinas se han visto beneficiadas de estos desarrollos. Para la biología evolutiva se abre la posibilidad de estudiar de manera más precisa y clara los patrones adaptativos de la variación. Tener genomas anotados y loci bien mapeados es relevante y arduo, pero el desarrollo técnico hace que lo anterior sea cada vez más plausible, y el reto será ser capaces de plantear preguntas adecuadas para hacer inferencias del mar de información disponible. El uso de una perspectiva evolutiva y de genética de poblaciones, enriquecerá a la genómica, de la misma manera que los datos genómicos nos ayudarán a avanzar en el desarrollo del programa iniciado por Theodosius Dobzhansky a mediados del siglo pasado.

Palabras Clave: Adaptación, genética de poblaciones, maíz, next generation sequencing, selección natural, teosinte.

ABBSTRACTA

The theory of population genetics originated over 80 years ago and allowed to explain, in terms of the evolutionary forces, the patterns of genetic variation within and between the populations that conform species. This research program generated the questions that have been empirically analyzed with the use of molecular markers for the last 50 years. A fundamental question within population genetics is if a reduced number of genes are representative of the evolutionary forces that affect the total genome of a species. This question has led to the development of molecular methods that allow the study of large sections of the genome in natural populations, giving rise to the field of population genomics. In recent years, techniques that are able to sequence DNA massively, usually called "Next generation sequencing" or "next-gen", are helping us to obtain genome wide data in many species, without needing previous molecular information. Comparing the genomes of many individuals from different populations, now we have access to an archive of their evolutionary history that narrates the complex and dynamic balance in time between natural selection and other evolutionary forces, such as genetic drift and gene flow, which act mainly in neutral regions of the genomes. The amount of information that is being produced has required the development of new statistical and bioinformatics tools for their analyses. Diverse disciplines have profited from these new developments. In particular in evolutionary biology it is now possible to study in a more precise way the adaptive patterns of variation. The annotation of genomes and the mapping of traits are important and complicated, but recent technical developments are making these goals easier, and thus the future challenge will be in asking the right questions to make relevant inferences from the sea of information these new methods generate. The evolutionary and population genetics perspective will enrich genomics, in the same way that the genomic data will help us advance in the development of the program initiated by Theodosius Dobzhansky several decades ago.

Key Words: Adaptation, population genetics, maize, natural selection, next generation sequencing, teosinte.