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The evolution of complexity A primer on evolution and selection 1.Selection is on Phenotypes not genotypes 2. Most mutations are neutral because of the degeneracy of genetic information. The wobble position of the 3rd codon often leads to mutations that have no phenotype. 3. Evolution occurs on multiple (all) genes simultaneously – but at different rates (tempos). 4. There is a potential for lateral gene transfer – especially in prokaytotes. 5. Sexual recombination greatly increased genetic variability – because genetic information is contained in discrete units (genes) . Molecular “clocks” The basic Poisson function for a mutation at any point in a codon dN/dt (I) = e- kt (kt)I/ I ! The evolution of Eukaryotes The acquisition of mitochondria and plasids – driving forces and evolutionary consequences Massive lateral gene transfers Modes of Evolution 1. Horizontal versus lateral gene transfer 2. Selection versus neutral rhizaria te s ell a Di no fla g es s Cili at oph y te s Parabas alid Diplomonad s Core Jakobids on ads Ox ym la op Archaea sm e a/f rr * ia Euryarchaea Me th an ot he Me t rm h an oba us ct er ium Me th an opy rus m a ia er ct a b b ia ) icro do i m ( GNSB Ac rruc o ac te ria B r u lf u Ve N on-s a Gre en b ac te ri a c te ria - proteo -p rote ob ? mitocho ndrion - prot eob ac te ria -pr - p oteobac teria rot Spir eo ba Ch och ae t c te es la m ria yd C ia Ch y to p h lo Ac ro a ga s bi tin (C ob (G FB r a ee ct Gr De e n ou ri a in Su p) oc † l fu oc rB cu ac s/ Th te ri a er ,G m us sb ) r te ac ob an Cy s st id pla er Th sm te s my ce ROOT as a an os Me th m u irill osp 1 h an t e ne M Am la op excavates la sm My c op Firmicutes rc in a /t he rm Halophile s P yro c oc c u s Archaeoglobus oco cc us Pyrobaculum/thermoproteus of ilum The rm Na no a rc ha eot a De s u lfuro Sulfolo Ae ropy c rum/py ro oc c us b us dic tium c ha e ot a The rmo t og Hy drog en a o bac te ri a Ko ra r ids ra s e ba s c A d Amo ampf ii lk h a V Euglena Tryp Le i a nos ome sh m s ani a P la nc to Ce na rch ae um nts Op 11 Crenarchaea het ero ko a te s y te s ce ms s oph y m ia to Ch ry o D Phaeop hy tes O Ch ry s ophy te s lids u th n ri y b phy te s Cry pt o ia s porid i Mic ro ng Fu h ist p o La ex pl dis cic r i s tat e s ts on k o om ic Ap Ha p t am oe bo zo a d Re ee nA lga Gla e uc Ce o p llul hyt ar S Lob o eA s lim lga P las m e M e Am e odia l S lim o ld s oe b e Mo as lds Ch oa n ofla ge lla te s Anima ls E ug l y p hid A moe ba e Foraminifera ia s la r mona d Ce rc o te s dio y Ra ph e nio ga Al a ch Gr rar G n ts P la a e g nd La Al en re s nt pla alveo lates lo Ch Eukaryotes Bacteria Secondary symbionts • The evolution of modern eukaryotic phytoplankton • Drivers for secondary symbiosis – • retention of fixed N in the host cell (extreme oligotrophy) • plastid could obtain protection from predation (armor) Euglenophyta Chlorophyta Bacillariophyta Chlorella Odontella ntcA odpA odpB petJ preA rpl28 trpA trpG trxA chlN cpcA cpcB cpcG dfr glnB gltB hisH infC nblA Nephroselmis Cryptophyta cemA cpeB ftrB ilvB ilvH infB minD pbsA psaK rne tsf Euglena ftsW ndhA-I ndhK psaM Guillardia thiG bas1 Cyanidium pgmA rpl9 rps1 syfB syh upp Secondary endosymbiosis clpP ftsW psbM rbcLg rpl22 accD bas1 cpeB infB minD pbsA psbX rps20 accA accB accD apcA apcB apcD apcE apcF argB carA Mesostigma Porphyra hisH minD ndhJ odpB rpl33 rps15 rps16 Glaucocystophyta accD cysA ccsA cysT cemA ftsW chlB infA chlL minD chlN ndhA-I clpP ndhK Secondary endosymbiosis rps16 rpl21 Cyanophora chlB chlL cpeA dsbD fabH fdx moeB atpI rne cemA rpl23 minD rpl32 odpB cystA cystT infA ndhA-K rps15 > 90% of genes lost Primary endosymbiosis Ancestral photosynthetic prokaryote bioY crtE groES hemA mntA mntB nadA rbcSg accA accB argB bas1 carA clpC cpcG cpeA cpeB dnaB dfr dsbD fabH fdx ftrB ftsH glnB gltB ilvB ilvH infB infC moeB nblA ntcA odpA pbsA petJ pgmA psaD psaK psaL rbcLr rbcSr rpl4 rpl9 rpl13 rpl24 rpl27 rpl29 rpl31 rps1 secA syfB syh thiG trpA trxA tsf upp acpP apcA apcB apcD apcE apcF atpD atpG cpcA cpcB dnaK groEL hisH petF petM preA psaE psaF psbV psbW psbX rbcR rpl1 rpl3 rpl6 rpl11 rpl18 rpl28 rpl34 rpl35 rps5 rps6 rps10 rps13 rps17 rps20 secY trpG psbM petA petD petL psaI rne rpl19 rpoA Marchantia cysA, cysT rpl21, ndhA-K Pinus cysA, cysT, rpl21 chlB,L,N, psaM Nicotiana acc D Oryza chlI ftsW minD odpB rne rpl5 rpl12 rpl19 rps9 tufA Zea Ancestral Rhodophyta genes? minD [minE] ilvB (pbsA) (cpeB) ilvH psaK ftrB (infB) tsf Cryptophyta Guillardia [accD] [odpB] cemA [(rne)] Euglenophyta Euglena atpI rpl23 rpl32 Bacillariophyta Odontella thiG (bas1) accA accB argB carA cpcG (cpeA) dfr (dsbD) (fabH) (fdx) glnB gltB infC moeB nblA ntcA odpA petJ (pgmA) rpl9 (rps1) Rhodophyta (syfB) (syh) trpA trxA (upp) cbbX clpC dnaB ftsH psaD psaL rbcSr rpl4 rpl13 rpl24 rpl27 rpl29 rpl31 secA acpA/P atpD atpG dnaK groEL petF petM psaE psaF psbV psbW (psbX) rbcR apcA apcB apcD apcE rpl1 rpl3 rpl6 rpl11 rpl18 rpl21 rpl34 rpl35 rps5 rps6 rps10 rps13 rps17 (rps20) secY atpA atpB atpE atpF atpH chlI petB petG psaA psaB psaC psaJ [psaM] psbA psbB ccsA petA petD petL psaI rpl19 [rpl33] [rps16] rpoA psbC psbD psbE psbF psbH psbI psbJ psbK psbL psbN psbT rbcLg/r rpl2 rpl5 rpl12 rpl14 rpl16 rpl20 [rpl22] rpl36 rps2 rps3 rps4 rps7 rps8 rps9 rps11 rps12 rps14 rps18 rps19 rpoB rpoC1 rpoC2 tufA [rdpO] cysA cysT infA [I-CvuI] [ndhA] [ndhB] [ndhC] [ndhD] [ndhE] [ndhF] [ndhG] [ndhH] [ndhI] [ndhJ] [ndhK] [rps15] chlB (chlL) chlN apcF cpcA preA cpcB rpl28 (hisH) trpG Glaucocystophyceae Cyanophora bioY crte groES hemA clpP [ftsW] psbM mntA mntB nadA rbcSg Chlorophyta