Looking in the Right Direction

Home , Ernst Mayr

www.landesbioscience.com RNA Biology11:3,1–6;March 2014;©2014LandesBioscience meet you.“ meet to moving decidedly is field the judge any Iam but if stands, to me: exciting …” handicaps, Despitethese subject. much of the by or penetrate engaged be can that of mind sort the Ihave think … Idon’t much biology, about Idon’t worse, very and know of by my most colleagues out mainstream of the as regarded are me.” with ter mat the to discuss My telephone youmanner. 3–9369, care if is a such in evolved and system dynamic acomplex indeed is cell the that fact for the appreciation an systems, of dynamic complex biologists carrying of card part on the some cognizance at least to see like Iwould about. I’m to what [sic] talking atune you are that know, [sic], and thereofore told interests, of been your have I to emerge. needs biology different very and anew and vision, its out of) (run lost has clearly biology 21st Molecular century. into the of biology teaching the you atsome point moving about point: to the on getting no time “ninja,” wasted sender the called mysteriously acomputer from Originating of my life. email important most Submitted: 03/21/2014; Accepted: 03/21/2014 http://dx.doi.org/10.4161/rna.28640 Correspondence to: Nigel Goldenfeld; Email: [email protected] class of complex dynamical systems. dynamical complex class of as anovel biology of understanding our for also but elsewhere, on life of origin the of understanding our for only not ramifications has which and genetics, population transcends ciated. appre widely so not Archaea!—are the not was discovery—it his of outcome principle as the saw he what and direction, tific this in scien driven was why he reasons the rRNA, of his studies on based developed he techniques this the discovery, through Archaea. as the known be to came which Life, of Domain third ticular, the of his discovery par in to microbiology, contributions his landmark through “You may not feel too much at home with biology as it now“You as biology much athome too not may with feel tremendously honest, to be and, frank was response Carl’s but tend to work physicist, on topicsthat atheoretical “I am that writing alacrity, with Iresponded Naturally, to to talk like I’d Sciences. Woese, Life over in Carl is “This the Ireceived 2002, 2pm 20, on FridaySeptember exactly At ese is known to the scientific community primarily primarily community scientific to the is known ese Carl Review I n thi W stitute for Universal Biology; Biology; Universal for stitute I n o s essay, I discuss his vision of evolution, one which which one evolution, of his discuss vision

Carl Woese and evolutionary biology evolutionary and Woese Carl W Looking in the right direction right the in Looking hi le it is well known how he made made he how known is it well le I n stitute for Genomic Biology, and Department of Physics; University of of University Physics; of Department and Biology, Genomic for stitute E a rth and and rth Nigel Goldenfeld RNA Biology - - - - ultimately leading to a new proposal for the classification of life classification for the proposal to anew leading ultimately accurately, defined) the lineages of organisms whose cellular cellular whose of organisms lineages the defined) accurately, (or more tracked only however, comparison sequence 16SrRNA of large; evolution writ dynamics the to mark choice a brilliant sequence molecular its made 16SrRNA, particular in ribosome, of the nature conserved highly The ribosome. the revelation: of biological instrument chosen of his limitation by anecessary it, disappointed with dissatisfied deeply he was accomplishment, remarkable this pride in manifest despite Carl’s and exaggerate, to hard is discoveries two of these significance the Although organisms. of associated view, acommunity another orism, in organ asingle view one in extreme representing and branches Archaeal/Eukaryotic the and Bacterial the between positioned to be now known (LUCA), ancestor common universal a last of existence the implies related is life all that finding the sized, empha view. Not frequently so mainstream the today is that and tripartite structure of life by Woese Fox 1977, of and in life structure tripartite and relatedness the of both discovery celebrated the in culminating work on rRNA, his into motion, through investigation this ting Evolution.” Cellular of actual period into the or so. i.e., backward years by a billion time in of evolution backward knowledge our to extend necessary be it will Ifeel (i.e., cells, simplest) prokaryotic evolution of the molecular biology, and evolution theory. evolution and biology, molecular microbiology, between relationship conceptual and historical the on article our in reproduced was of which 1969, extract alengthy June 24, dated Crick to Francis aletter in clarity with articulated been had of research program 1960s. the That during seriously about to think begun he had that of research aprogram on Earth, life of all history evolutionary of the understanding complete his to was wanted Carl what provide to biology. could tion fact, In quantifica that value great to the no stranger he was later. Thus, years 1950 in Yale from Biophysics three in aPhD College and Amherst from Physics and Mathematics in aBA with physicist, a as trained had himself Carl skills. math better with gist—one of biolo breed anew to help away create more than was cause to his physicist atheoretical enlisting but his systems, dynamical complex in roots with science aquantitative biology of making on agoal sights his set indeed had Carl exchanges. initial those in set was trajectory to note how future much of our remarkable it is of correspondence, fragments atthese back Looking wise. other phone on email the or via talked and day every nearly we met time, that During death. his until a decade more than

Carl had famously spent much of the following decade set decade following spent much of the famously had Carl to the leading of events course the to unravel ever we are “If So began a scientific partnership and friendship that lasted lasted that friendship and partnership ascientific began So 1 I l linois at Urbana-Champaign; Urbana, Urbana, Urbana-Champaign; at linois 1 I L U SA Review 2 and and 3,4 1 ------

©2012 Landes Bioscience. Do not distribute. 2 amino-acyl-tRNA-synthetases. of the history into the to dig collaborators and Carl led had this and phylogenies, HGT, for non-canonical and evidence showed clearly ribosome the and translation with closely associated those than other on genes based Phylogenies ref. 5). see (for areview, concept ameaningful aTree post-LUCA, was of that Life idea the doubt on casting thus of lineage, concept of the invalidation an as invoked being organism—was unrelated to another organism resolved. to be began problem LUCA also before life the issues, these understanding in Remarkably, (HGT). fer trans gene of horizontal impact evolutionary of the recognition increasing the from notion of Tree very arising itself, the of Life to were challenges there because mind, on his preoccupations were more urgent there but that found Isoon questions, related and by these animated extremely was Carl LUCA? before life about us it tell does What LUCA? is that singularity the about us tell this does What significant. surely is for example, hundred, of not order is one unity, number and that fact but the three, number to the significance special any not attaching Iwas Life? of Domains three only there why are was to him questions first One of my advance. aconceptual to make need he would track, on program original his to get that convinced was endeavor. Carl original Carl’s overshadowed of Life Domains three the ering Earth had apparently run into aroadblock. run apparently had Earth on of life history evolutionary Woese’s the to uncover program phylogeny. to 16SrRNA invisible be would thus and ribosomes, to today’s related as recognizable machinery translational have not would of life phylogeny. by 16SrRNA classes traced Earlier to be form advanced sufficiently in machinery ited translational exhib which organisms first of the but arepresentative ancestor common universal last not the just was LUCA of life. out all leaves it necessarily LUCA, before era but the in of course, viruses out leaves that era modern the In ribosomes. included structure because what interested him more was not whether evolution had evolution had not whether more was him interested what because of evolution. rate the HGT but from instead phylogeny arising not on the to focusing of receptive power HGT, was thus, and evolutionary the understood he well of evolution. Nevertheless, structure large-scale the appropriate for illuminating equally are genes not all meant: of lineage concept the of what derstanding misun afundamental showed genome phylogeny, they because whole from arising those as such of Life, Domains three to the challenges more extreme the with not impressed too was Carl 9). 8and refs. in articles related as well ref. 7as in discussion reviewer and paper the for example see perspective, a balanced from ries it (for summa enhances actually relatives, close among common more of it being or by virtue pattern, canonical not HGT disrupts or whether as debating—such still is community the which about issues many remain there conclusion, although this confirmed basically have analyses Subsequent thoughtful intact. still was ture struc large-scale Domains three but the disrupted certainly was pattern canonical that showed study synthetase The fluidity. any component with likely most the represented synthetases the and machinery, translation of the part conserved most the to be likely

Horizontal gene transfer—the transmission of genes from one from of genes transmission transfer—the gene Horizontal The spectacular success of molecular phylogeny in uncov phylogeny in of molecular success spectacular The until around 2000, 2000, around until of Species Origin Darwin’s not read did Carl 6 Carl had chosen the rRNA as as rRNA the chosen had Carl RNA Biology ------as a whole can behave in some sort of synchrony to control the to control the of synchrony some sort in behave awhole can as behavior, system but the switching stochastic a complicated have might individually nodes The of proteins. levels expression gene and factors transcription be might links the and genes are nodes the for example, case, latter the In or regulation. gene metabolism as such of a system complex a model constitute that by links nected con nodes of anetwork: idea the is paradigm Atypical selves. them parts the than more is important which parts those between relationship it the is and identity their lose parts individual the effects. of collective question more specific but notion of related complexity, to the defined vaguely the from away turned quickly discussions our because no exception, was case This intuition. and imagination his up for it made with than but he more mathematics, biology, and physics, structural istry, biochem in knowledge technical he lacked that say often would Carl feeling. this why he orhad even detail in to him meant this what say he couldn’t ofmode evolution, but really pressed, when and tempo the to do with something had dynamics systems plex com that sure was Carl dynamics. and of complexity issue on the work on of phylogeny, his context the in but to focus here Iwant phase of life, which he called the “progenote.” the he called which of life, phase a on such discussion conceptual Fox, initiated with also paper magnificent another in had, Archaea of the discovery the as year same the in and genes, before to life alot of given thought already had theory. Carl of the of validity regime the beyond manifestly was such as were genes world there before biological tion, the when, by construc process, evolutionary of the explanation full a as considered be possibly can genetics how population to ask is inadequacy seeming this One to phrase way process. evolutionary of the aspect important an somehow missing was century, 20th of the half first the during synthesis modern the from emerged had of evolution, to say, which is picture the That he felt that issue. to the pertinent be would systems of dynamical complex some understanding that intuition the had Carl because issue, on this centered Carl with discussions My early years? one billion atmost be could of what frame atime in machinery translational modern essentially an attaining and abiotic earth an from starting much, so evolution achieved have How Fox: could with results early the from right Carl puzzled had that something echoed Elsewhere, I have described how Carl thought about this entity this about thought how Carl described Ihave Elsewhere, ref. 14). see enterprise, of this account computer (for astimulating modern world’s of the first construction to the devoted life of his automata work on cellular of theoretical asummary inspiration as citing process, translational of the for accuracy requirements to the ity complex and size the attributed Carl machinery. cellular other any more than complex, perhaps and large is apparatus lational trans the that observation simple on incisive the ment he based argu an cell, evolution of the the shaping force primary the be to relationship genotype–phenotype of the emergence the sidered con Carl not emerged. yet had phenotype and genotype between and Mode of Evolution Mode and G.G. Simpson’s was on him impression most the Tempo made had work that the of fact, evolution. In speed but the occurred,

Complex systems must be strongly interacting, so much so that that much so so interacting, strongly be must systems Complex The progenote was a phase of life in which the distinction distinction the which in of life aphase progenote was The 12 conducted by John Von conducted Neumann , 10 because its analysis of the fossil record record fossil of the analysis its because 11 13 during the phase phase the during V olume 11 I ssue 3 15 ------

©2012 Landes Bioscience. Do not distribute. www.landesbioscience.com by the abstraction of a genetic code, evolved in specificity and and specificity in evolved code, of agenetic abstraction by the represented as machinery, translational resulting the that was however asurprise was What arise. could this that anticipated already we had because not asurprise, was This effect. network collective of the of evolution, because dynamics the accelerated HGT Emergent apparent. became simple quickly conclusion very it To were. as a not put bywould hand be in astonishment, our and calculation the from dynamically emerge actually would of HGT, ingredient added efficacy HGT the whose but an with or (2) by mutation alone, (1) adapting ways: two in ronments, envi niche to occupy trying and competing organisms, of early dynamics the evolution.” explore we would is, That “Darwinian vertical purely with evolution of translation clear: very was model control the For separately. us, ingredients of the contributions dynamical the isolate we could dynamics, the in included dients ingre different with models minimal two by comparing that so a theorist’shypothesis), version of a null like (something model acontrol to have we needed that meant evolution of translation of the models minimal with working that idea the work was to our Central Wisconsin. and atHarvard career distinguished appropriately gone on to an has who me, with working student graduate abrilliant time atthat Vetsigian, of Kalin ticipation par the with accomplished was but this not trivial, project was acontribution! make perhaps could physics theoretical that away was Here at last checked. robustness the and analysis, or other by simulation enumerated be could outcomes the tems, sys of dynamical mathematics the in formulated be could anism amech if that it meant because thing, agood to me was Generic literature. the were in evolution that early about stories” so “just the but all not ours, were, just arguments these how all generic me it struck matters, other and these about more we discussed The code? genetic to auniversal lead and compete themselves would of organisms, cohorts competing many across distributed codes, of genetic set initial an by which mechanism reinforcing self- autocatalytic, an be there Could recipientby the organism. expressed be can gene transferred the that so code, genetic versal auni is there of HGT. much unless not accomplish HGT does presence the in evolve would machinery translational how the we wondered particular, In effect). body single point mutation (a to opposed as effect) by collective HGT (a dominated was tion evolu whose populations in arise would behavior emergent what such a mechanism could be significant. be could amechanism such of impact evolutionary the workers, by earlier recognized As gene. transferred the is that by alink connected recipientand microbes, donor the being nodes the effect, a network is also transfer gene life. 7yof Biology,his Genomic past he where for worked the for Institute up at the we set library interdisciplinary the through lent book “Sync” book lent Strogatz’s of Steven excel copies adozen half went bought out and of phenomenon he eventually that type by this struck so was Carl Actually, of systems. complex phenomenon one hallmarks is of the emergent this and to inputs, response and node’seach dynamics of details specific much on too depend the not necessarily does whole of system the behavior collective The for example. cycle cell The task of turning these early ideas into a concrete research research into aconcrete ideas early these of turning task The Returning to the problem at hand, we realized that horizontal horizontal that we realized problem to the athand, Returning 16 to give to friends and to distribute to students to students to distribute and to friends to give

17-22 We began to wonder We began RNA Biology ------had been guessed by Woese in a remarkable paper, by Woese aremarkable in guessed been had “optimality” genome. This of pointthe mutations indeed and translation in of errors effects the mitigating in so) nearly very (or optimal some sense in was that to acode precision, leading account tRNA abundance tRNA account into to take refinements later it (and because quite remarkable, the preceding one of Ardell and Sella and one of Ardell preceding the and model our of all, First code. genetic evolution ofof the the example of what has been presciently called “universal biology.” “universal called presciently been has of what example an is and protocols, innovation-sharing we termed to what generic is that something is process This complexification. organismal and refinement of code process of aco-evolutionary attractors dynamical were the characteristics these but because molecules, of properties or chemical tuning not due to fine optimal, nearly and universal both be would code genetic workour that showed Fourth, dynamically. interpreted progenote, finally mysterious HGT: rampant Carl’s one with of life, state ofresidue acollective but the organism not to be asingle identified was LUCA Third, atLUCA. of Life Domains of three emergence the lution and evo to vertical transition the in not properly understood still mechanisms through culminated that process growth nential expo an in novelty genetic generating state collective evolving arapidly with LUCA, before were different tempo and mode the both yes, of mode evolution: and tempo the with cupation it.”— (1969). F. letter unpublished money to finance Crick enough attract well might project acombined Such disappointing. prove results evolutionary the if even results to give bound is which on function, of sequence of change effects the study gramme—to pro enlarged up aslightly you draw Isuggest reason, For this sequences. the in frozen still is evidence enough whether gamble a rather its as money for it, toespecially get difficult be well may wrote: Crick to Carl. anathema now been to have know I that something function, and of sequence astudy with hedged it be that suggested and program, evolutionary the about doubts he expressed to my surprise; supportive, not fully was response Crick’s of life. history evolutionary out to map the sequences lar molecu intention to use his he announced which in to Crick, to me recently. letter to Carl’s Previously, clear Ireferred became only for this reasons the and wrong, was argument Crick’s that out of showing However, gotakick Carl intoto it detail. go in place not is the this one, complex and and interesting an was Crick with accident.” relationship Carl’s “frozen memorable term his through code evolved of an dismissal to Crick’s arguments the canonical degeneracy structure). degeneracy canonical the retaining (although table code the in acids amino mutation of the per one through canonical the from differed that codes genetic of synthetic sampling Monte Carlo using others, later and Hurst, and by Haig quantified and rediscovered been subsequently had optimal nearly be would itself code canonical modern the that idea The for life. complex necessary machinery deterministic highly day present to of the proteins distributions statistical duce able to pro being from machinery of atranslational refinement gradual of the scenario adynamical to describe first the was

That this arose from HGT but not vertical evolution was to us to us evolution was HGT from but not vertical arose this That Second, our work finally answered Carl’s overriding preoc overriding Carl’s answered work our finally Second, one but it important an one and agood projectis the “I think 31 ) solved in one stroke the key facets facets key the one in stroke ) solved 24-30 32-34 were concrete counter were concrete 23 which also also which 35 3 ------

©2012 Landes Bioscience. Do not distribute. he pointed out that Carl’s heritage as a physicist had influenced influenced had a physicist as heritage Carl’s he out pointed that arguments, his order In to buttress classification. one biological in meaningful only the being as distinction prokaryote–eukaryote of the concept of the refutation concomitant the and Domains, three Carl’s He challenged emerged. that of life view centric microbe- the from recoiled Mayr apparent, became program of Carl’s results However, the of as Sciences. Academy National to the Carl one nominated who the been had indeed and porter of Sciences. Academy National of the Proceedings the and of Nature pages the in undertaken of titans battle true Mayr—a Ernst with mish skir famous his in issue ascientific became that something cist, aphysi as self-identification enduring his is well-appreciated less 4 a physicist wondering within the world the of biology. within wondering a physicist as long journey of his avindication some sense in fulfilling, truly found Carl that something was theory systems dynamical and mathematics using of power reasoning the that Ibelieve forseen. been had that not something evolution was for code mechanism dynamical of the inevitability remarkable the by Carl, anticipated work were long of this roots the that Ifeel Although results.

shows that taxonomic trees are more balanced than evolutionary trajectories as measured by 16SrRNA by sequences. measured as trajectories evolutionary than balanced more are trees taxonomic that shows NCB the from constructed trees Red, atree. of node each for size subtree Figure 1. Carl’s lonely journey is well-known of course, well-known is lonelyCarl’s journey by these thrilled very was Carl that to say It no exaggeration is Scaling properties of phylogenetic and taxonomic trees, as described in the text. The cumulative subtree size is plotted as a function of the the of afunction as isplotted size subtree cumulative The text. the in described as trees, taxonomic and phylogenetic of properties Scaling 37-39 Paradoxically, Mayr had been Carl’s strong sup strong Carl’s been had Mayr Paradoxically, 36 but perhaps but perhaps RNA Biology - - - I database. Blue, trees constructed from the Greengenes database. The figure figure The database. Greengenes the from constructed trees Blue, database. al. et by Herrada proposed were simultaneously which this, quantify to some ways up with came and trees phylogenetic branching of properties topological the about thinking I started to do this, order In itself. process evolutionary of the dynamics statistical the of phylogeny, in laws someorder understanding toin get scaling to look for possible it became accumulated, genomes as and trees, of phylogenetic properties statistical the about curious I became 2000s. late the during it developed as of metagenomics science new Woese’s blast. call last we might one that that, about anecdote another to end with want and conversation, this to acontribution of making However, privilege the Ihad passing. Mayr’s with ended debate the of course and argument, scientific abeautiful is rebuttal Carl’s of life. classification Domains three the of proposing taxonomy, mistake the not he made would have of practice the through identified features of organismal tance impor of the appreciation an with abiologist, been had Carl If trajectory. evolutionary by their lineages classifying lution and of evo course the for tracing techniques of choice molecular his the number of subtaxa diversifying from the node). The second second The node). the from diversifying of subtaxa number the (ie. size subtree the A: call we will first The tree. a phylogenetic 40

For various reasons, Carl and I had become interested in the the in interested become Ihad and Carl reasons, For various The basic idea is to measure two quantities on each node of of node on each quantities two to measure is idea basic The V olume 11 I ssue 3 - -

©2012 Landes Bioscience. Do not distribute. www.landesbioscience.com his unpublished PhD thesis. PhD unpublished his in detail great in question this explored has who Jeraldo, Patricio student, outstanding by another extended and confirmed a result z~1.4, practice In 2. 1and between be zcan exponent the where A~z log C/log relationship: by law apower related were essentially pose of classification is to facilitate information retrieval. information to facilitate is of classification pose pur the that Mayr’s from view diversity,” arises aprinciple that of degree and size of equal possible, as far as are, rank categorical atagiven taxa the if facilitated greatly is of information retrieval tion is: are C and A related? Herrada et al. et Herrada Arelated? Cand are is: tion ques The value. possible on largest its Ctakes branch, a single from time the all splitting branches with subtree, comb-like ric, asymmet most for the whereas value, possible smallest the C is trees, balanced equally For symmetric, some sense. in subtree the of shape the measures quantity This nodes. subtree the of all sizes branch of the sum the C: size branch cumulative the call we will board attached to a SUN workstation—the famed “ninja” famed no workstation—the to aSUN attached board key at agrubby pecking he is Later bench. alab really desk, the up on are feet His of design. ergonomic forerunner zero-budget and aprimitive as arms to the taped towels of washroom pad athick one with the chair, ancient his in indecorously lounging 2002, September in sessions early those during Hall at Morrill scientist. Atrue That’s he who emerged. was. that understanding scientific the was to him important Mayr, was but and what Crick of both ghosts the vanquished had Carl not gloat. did tion—he classifica human vs. classification views—nature’s of his cation Figure lineage. evolutionary a common shared that together toorganisms group is of classification purpose the not acollector: evolutionist, of an that was view, of course, Principle of Balance!” Mayr’s Principle of Balance of Principle Mayr’s Balance!” of Principle you’ve “Ah,” Mayr’s “I think discovered he said, fascinating: and immediate was reaction His z. exponent of the values to different corresponding look like would trees the of what out acaricature drawing for him, interpretation the and calculation the explained I and shortly He arrived interesting. something to see lab into the to come him asked and up Carl, Icalled plotted, was analysis Patricio’s as soon As Why? trees. phylogenetic than more balanced were somehow trees taxonomic the that suggested to unity close zbeing exponent of the value the because puzzling especially was their statistical scaling properties, as shown in Figure in shown as properties, scaling statistical their in reflected is this and different, very are trees taxonomic and trees of phylogenetic structure topological the words, other unity. In to close law, apower is very zis exponent but the also result the that discovered Patricio genomes. sequenced of fully set the than rather database, NCBI taxonomic the from constructed are trees if tion-based perspective. Mayr’s and onclassifica biology perspective evolutionary Carl’s between difference vast the reflects and processes, evolutionary the of dynamics natural the to 1.4 close exponent indicates ent scaling adiffer exhibit trees phylogenetic the that fact The classification. into taxonomic put by hand in been apparently has that Balance Principle of the reflecting of Mayr’s to classification, fact approach arti an is to unity zclose exponent with law ascaling exhibited taxonomy that fact the viewpoints: two these tension between the When I think of Carl Woese, the scientist, he is in his office office his in he is scientist, Woese, the of Carl Ithink When Although Carl considered this analysis as some sort of vindi some sort as analysis this considered Carl Although

41 It is interesting to ask what happens happens what to ask It interesting is

1 dramatically captures 40 found that C and A A Cand that found 38 states that “the “the that states

1 . The result result . The 42 Carl’s Carl’s RNA Biology ------Jeraldo in the work briefly summarized in Figure in summarized work briefly the in Jeraldo That was Carl’s “Rosebud” moment. “Rosebud” Carl’s was That youth. his in horses riding remembering moment he when was that as happy away. so him far saw Inever are eyes his and smile abeatific wears face His once. Carl out, was rider, it so turns and Zippy. to my daughter She’s ahorse- talking house of his kitchen today.” again mind your “Woese, you destroyed have he went evening: homeas the in to himself saying he recalls reminiscences, personal his In of life. history the he deduced which from catalogs molecular the create to experiments dull mind-numbingly performing wilderness, the 10 the yin through gothim which it that is of course and dinary, simply extraor was to focus ability Carl’s never-ending. seemed sometimes loop that feedback perfect—a it was honed until and again gone to be over and small too was no detail And he write! could But, man, speaker. articulate not an public he was in that avisitor, know or introduce you’ll a lecture give Carl heard ever you If over commas. now, one. But right good arguing we are it’sWe avery even paper, that agood perhaps to know be going to write. a year us taken already paper. It code has genetic of our version final over the going department physics the in at my desk people. other did than way different avery in atlife he looked that fact the betrayed but it always taste; by good unfettered sometimes thus, humor was, His person. other of the mind the in put himself to unable he was that meant this and of personality, type Asperger’s asub-clinical had Carl He He didn’t jested. jokes. tell between. in bawdy. even Very sometimes little and rambunctious or off-scale witty and subtle, subdued, encountered: ever Ihave of humor that sense idiosyncratic most and cleverest the perhaps He had worked. we while not fun to have no reason saw Carl hand. his with wall the beating and barking, aseal like asound atajoke, raucously and loudly IGB. the laughing He in is offices our between hallway pretentions. without aman is this Clearly think. I what to know He wants papers. and books in read Ican what me telling in no interest He has to talk. gently, he starts then and closing eyes the head, of the lowering aslight with acknowledges He he who is. Iknow that know him to let gratitude, and respect of expression but genuine acareful Imake lesson. biology first my Ireceive that day the to be is of it. This years 3.8billion last the Or atleast on Earth. of life history the ously, unraveled Carl room, 25 yprevi this In bench. lab his facing wall the adorns flag American an and Davis of Miles poster agiant is him Behind times. of the Moore’s a decade’s is worth behind Law less—that NNA13AA91A issued through the Science Mission Directorate. Mission Science the through issued NNA13AA91A No. Agreement Cooperative under Institute Astrobiology NASA the through Administration Space and Aeronautics National by upon the work supported based is material This text. the in

It is a pleasure to acknowledge the collaboration of Patricio collaboration the to acknowledge It a pleasure is were disclosed. of interest conflicts No potential When I think of Carl Woese, the man, he is standing in the the in standing he is man, Woese, the of Carl Ithink When Woese,sitting my we are collaborator, of Carl I think When the in Woese, standing he is Carl of my friend Ithink When Disclosure of Potential Conflicts of Interest Conflicts ofPotential Disclosure Acknowledgments

1 and described described and 5 - -

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