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its original source was the edge my interest in evolution started in the summer of 1947 when i spent six weeks in the painted desert with a paleontologist named sam welles who had a group of students there officially in a summer course but we spent most of the time swinging picks and shovels digging fossils as part of welles research project he was a specialist in triassic amphibians evenings were spent sitting around the campfire talking about things like evolution for the first time in my life people real biologists real scholars were willing to sit and listen to my opinions i was twenty one years old i certainly became interested in many aspects of evolution then and shortly after that i signed up at the university of california at berkeley for a course in evolution with ledyard stebbins who at the time and for quite a while thereafter was the world s primary expert in evolution with respect to things botanical stebbins course introduced me to theodosius dobzhansky s genetics and the origin of species stebbins was great but dobzhansky s book was what got me interested in natural selection as a process at the university of chicago my job was strictly teaching i was in their early entrant undergraduate program taught freshmen and sophomores biology they had a great books approach we read darwin mendel and others also i attended seminars by people such as alfred emerson the termite specialist and recognized authority on things evolutionary i found his ideas absolutely unacceptable that motivated me to do something if it was biology emerson was discussing i would be better off selling insurance i remember especially his lecture on the role of death in evolution he was all in favor of death and said that the reason we grow old and die is to make room for successors so that they can have a chance this seemed so totally impossible given that evolution proceeds by natural selection there was absolutely no logical way you could reconcile his ideas with darwinism even though he claimed to be a darwinist this initiated my first theoretical obsession the evolution of senescence the decline in adaptive performance with age you can t run as fast at sixty as you could at thirty on the way home that evening talking about the problem with my wife i independently came up with an idea that peter medawar is chiefly responsible for and published in 1952 although he may have published something that foreshadowed it in the 1940s and that is that the effectiveness of selection in maintaining adaptation is essentially the product of reproductive value and survival the survival factor is easier to appreciate if you re more likely to be alive at thirty than at sixty then selection will be more effective at maintaining adaptation at thirty than at sixty at an age you d be extremely unlikely to survive to such as one hundred years old adaptation would be a lost cause and selection wouldn t be concerned with it as the effectiveness of selection declines the effectiveness of its products declines this explains the rising mortality rate that comes with age it seemed to me at the time and still does that this is an inevitable conclusion arising from just the simple fact of mortality if there s any possibility of dying at any age then you re less likely to be alive at a later age than you are at an earlier age another one of alfred emerson s ideas was that evolution is much more concerned with cooperation than with competition it seemed to me to be very much the other way around and that there was something very special about the social insects which accounted for their extreme cooperativeness that special thing was their kinship high levels of kinship within the colony this was the focus of a theoretical paper i published in 1957 it was a model of natural selection between families now i think that s a silly way to do it but at the time i wasn t smart enough to think of the kin selection idea which was some years later worked out by william d hamilton in extreme models this kind of selection can lead to things like forgoing reproduction if in so doing you can for example more than double the reproduction of a full sib the full sib is half as good as you are genetically that is from the standpoint of getting your genes into future generations in the social insects of course sisters may have a three quarter relationship because if they share a father then all the genes they get from the father are exactly the same these early experiences kindled an interest that has never gone away and resulted in adaptation and natural selection my first book length publication on this and related matters by then i had worked on the problem of senescence and on cooperation between relatives but i had a long list of other problems that interested me at that time group selection was not explicit v c wynne edwards big book on group selection animal dispersion in relation to social behaviour came out in 1962 but i discovered it only after i was largely finished with adaptation and natural selection i submitted the manuscript in late 1963 and it referred to wynne edwards work but i brought it in as a late revision of the manuscript there was some group selection modeling prior to that and explicit use of group selection ideas by alfred emerson and a h sturtevant in paper published in 1938 in 1945 sewall wright presented a group selection model in a book review of george simpson s tempo and mode in evolution but the group selection model wasn t easy to find if you didn t know about it already mostly the group selection idea was necessary to the way people were thinking about adaptation although and i find this extremely strange they didn t realize it they kept talking about things being for the good of the species if it s for the good of something and it s to arise by natural selection it has to be produced by the natural selection of those somethings in other words one species survives as another one goes extinct the basis of wynne edwards work on group selection was that you can t have things that work for the good of the group unless you have selection at the level of groups what he was doing was looking for selection at the level of local breeding populations and whether they could be called separate species wasn t particularly relevant to most people s satisfaction wynne edwards has been proved wrong not that there s no selection at levels higher than the individual or the family but simply that his particular formulation isn t likely to be a very strong force in evolution it s now generally conceded that the phenomena he was explaining by this mode of thought are much better explained by other processes by selection at lower levels selection among individuals for instance any reproductive restraint anytime it looks as if individuals aren t reproducing at the maximum possible rate is explainable simply on the basis of an individual optimal resource allocation model you don t kill yourself trying to do something today if working at it a little bit more easily will enable you to try again tomorrow maybe you don t do it at all today if conditions will be much better tomorrow this kind of thinking explains the fact for instance that birds do not necessarily lay as many eggs in a breeding season as they demonstrably might the allocation of their resources will be much more effective for reproduction with a lower level expenditure on eggs which will enable them later to spend more on feeding the young and later still next year having another breeding season there s a great conceptual deficiency in my earlier work one that i shared with just about everybody else who was working at the time i failed to realize what a tremendous problem the existence and prevalence of sexual reproduction is i got interested in that in the early seventies and i published a book in 1975 titled sex and evolution there are a lot of complications that i didn t appreciate at the time but john maynard smith and bill hamilton and many others have advanced our understanding tremendously in the last twenty years richard dawkins went in the right direction when he made the distinction between replicators and vehicles david hull s substitution of the term interactor for vehicle is a good idea but that s a minor terminological matter dawkins didn t go nearly far enough in making that distinction because he defines a replicator in a way that makes it a physical entity duplicating itself in a reproductive process this is fine but the important distinction lies at a still more basic level he was misled by the fact that genes are always identified with dna evolutionary biologists have failed to realize that they work with two more or less incommensurable domains that of information and that of matter i address this problem in my 1992 book natural selection domains levels and challenges these two domains will never be brought together in any kind of the sense usually implied by the term reductionism you can speak of galaxies and particles of dust in the same terms because they both have mass and charge and length and width you can t do that with information and matter information doesn t have mass or charge or length in millimeters likewise matter doesn t have bytes you can t measure so much gold in so many bytes it doesn t have redundancy or fidelity or any of the other descriptors we apply to information this dearth of shared descriptors makes matter and information two separate domains of existence which have to be discussed separately in their own terms the gene is a package of information not an object the pattern of base pairs in a dna molecule specifies the gene but the dna molecule is the medium it s not the message maintaining this distinction between the medium and the message is absolutely indispensable to clarity of thought about evolution just the fact that fifteen years ago i started using a computer may have had something to do with my ideas here the constant process of transferring information from one physical medium to another and then being able to recover that same information in the original medium brings home the separability of information and matter in biology when you re talking about things like genes and genotypes and gene pools you re talking about information not physical objective reality they re patterns i was also influenced by dawkins meme concept which refers to cultural information that influences people s behavior memes unlike genes don t have a single archival kind of medium consider the book don quixote a stack of paper with ink marks on the pages but you could put it on a cd or a tape and turn it into sound waves for blind people no matter what medium it s in it s always the same book the same information this is true of everything else in the cultural realm it can be recorded in many different media but it s the same meme no matter what medium it s recorded in in cultural evolution obviously the idea of a coffee cup or a table is something that persists the coffee cups and tables don t persist they recur as a result of the persistence of the information that tells people how to make coffee cups and tables it s the same way in biology hands and feet and noses and so on don t persist they recur as a result of genetic instructions for making hands and feet and noses it s the information that lasts and evolves obviously it s because of the physical manifestations of the information that we know about the information dawkins has had trouble in convincing people and this stems from his thinking of the gene as an object of emphasizing the importance of replication rather than of proliferation of information until you ve made the distinction between information and matter discussions of levels of selection will be muddled comparing a gene with an individual for instance in discussions of levels of selection is inappropriate if by individual you mean a material object and by gene you mean a package of information it should be gene and genotype you have to look at levels of selection in both of these domains and realize what you re doing comparisons of levels of selection should be within the same domain having made the domain distinction you then go to levels and you find that in the two domains the levels do not correspond exactly as a general rule if we restrict our attention to sexually reproducing populations there are only two possible levels of selection in the informational or what i call the codical domain the gene and the gene pool selection can operate on alternative genes within a population selection can act on alternative gene pools in a biota both of these are evolutionary factors that can produce interesting effects in the material domain on the other hand selection can operate at the level of alternative individuals in the usual sense of individual or on groups of individuals such things as insect colonies or families whether they form elaborate colonies or not these temporary groupings of individuals give rise to what the biologist david sloan wilson calls trait group selection and also to selection between alternative populations that s the physical basis for selection between gene pools but the physical levels of selection below that level for instance between competing colonies of the same species of social insects don t have a corresponding level in the codical domain the events in the competition between insect colonies are recorded at the level of a gene there are no sufficiently persistent genetic differences among colonies for effective selection in the codical domain i believe david wilson agrees with that he s interested in selection among the interactors in the material domain the main messages of my 1966 book are now generally accepted this would have been the case whether i wrote that book or not the ideas would have prevailed by today because people like hamilton dawkins robert trivers and others were doing work at the same time more or less and if there hadn t been a single book in the mid sixties to deal with the idea of levels of selection i think one of those people probably would have written it dawkins book the selfish gene is very much a case in point it advanced things a lot further than mine did my lasting contribution will be for a clarification of the problems of the two domains and the levels of natural selection i ll also be known as one of the people who first became interested in explaining why there is such a thing as sexual reproduction and why it s so widespread in the future breakthroughs in evolutionary biology may come in the field of paleontology fieldwork now going on will be recognized several decades from now as having provided extremely important information people i ve never heard of are out there digging looking for pollen grains in lake sediments or dusting off trilobites from paleozoic shales other important insights will come from people working in traditionally unrelated fields for instance on things like con...