 All right. Hey everybody, thanks very much for tuning in and watching the talk. I really appreciate the the invitation even even unfortunately, virtually, but it's nice to be able to just be able to talk with you guys. It's been a fantastic triptych. I'm sorry I couldn't make the discussion session last Friday. But I really have a high standard to live up to being in in the third position here is actually not the most desirable spot because the other talks have been so great so I'll hopefully live up to that live up to that high standard. If you don't know who I am. Let me just briefly introduce myself. I'm Charles Pence. I'm a philosopher and historian of biology for the last three years. I've been working here in Belgium at the University Catholic the Louvain few if you know the the Leuven schools. There's the there's K you live in the Dutch speaking institution. We are the Francophone institution. You say Louvain, both a little bit outside of Brussels. That's, that's enough about me. What am I going to, what am I going to talk to you about today so titled greatly exaggerated rumors of the death of biometry helping myself to some Mark Twain. But I think the way that I want to approach this talk is I want to start by developing as a foil, a sort of classic history of the field and I'm going to narrow down to a particular period I'm going to be really focusing in on like 1905 to 1920 or so. According to this classic history, there's really nothing interesting going on. Maybe there's one argument that's kind of exciting I'll talk about that in a second. But there's really not much going on between 1905 and 1920. And what I want to do is try to make trouble for this classic history I think there is actually some really cool stuff that's happening over this over this period. I want to show you what that cool stuff is is I want to turn to a more philosophy of science focused question what does it mean to make evolution, a chancey or statistical theory. And that's something I think I want to argue on which there is a lot of movement over precisely this period, in a way that merits I think more careful reconstruction and thought from sort of HPSE type approaches to to the development of evolution. And then I want to close by convincing you that this matters for anything but like my personal preoccupation with these collection of humans by showing you that there are real connections between the kind of work that I want to talk about over the course of this, this little chunk of time, and the modern synthesis and so if if these kinds of ideas if this kind of philosophical approach, sort of makes it into the early days of the modern synthesis, then it by extension, makes it into the early sort of contemporary evolutionary evolution so far as you're sort of in the line of transmission from from the history of biology straight through to today's to today's work. And so the take home that I really want to argue for is that this isn't a story of rupture and revolution, which is one of the things we learn from the classic history there's a real continuity of a really interesting philosophical approach. This underlies the sort of birth of the use of chance and the methods of statistics in evolutionary theory. Possible shameless plug I'm not sure if this actually counts as a shameless plug so this does come from a book project that is working titled pompous parade of arithmetic it's a quote from WFR Weldon reporting an insult that he received that at a scientific level if his work on statistics and evolution was a pompous parade of arithmetic. I say possible shameless plug because for the last like eight months this manuscript has been finished and sort of shuffling its way through manuscript purgatory trying to find a publisher. If you want a copy feel free to email me I'm it's not doing anybody any good sitting on my hard drive gathering virtual dust. So feel very free to drop me a line if if this kind of work interests you and you'd like to know more about the approach I'll say a little something more about some other connections to stuff that's going on in the book project to kind of situate this piece but this is one of the pieces this is sort of chapter five ish of that of that manuscript project. Okay, so there's a classic history that we get from people like will provide perhaps most prominently on the on the history of science side, but also from practitioners practitioners like the geneticists start event or punnett who make the same kind of make the same kinds of claims. We start with Darwin Darwin zone theory doesn't use chance or statistics very much at all the story here is a more complicated than that I have a whole chapter on it, Andre area is about to have an entire book on it. So the story here is is weird and and complex, but let's just let's just say for the sake of argument for the moment for the purposes of the classic history. Darwin's theory is pretty much non mathematical right there's not a formula to be found over the course of the origin of species there's not much to be found in the way of formulas in anything that Darwin writes. He quite simply it seems isn't very much attracted to the mathematical approach to the sciences. That changes right after Darwin with the advent of folks like these two gentlemen right here this is Carl Pearson on the left and Francis Galton on the right so operating sort of at Galtons impetus. We have a little school that develops the biometrical school Carl Pearson and wfr Weldon really pick up the banner from Galton as Galton steps aside just basically spent the last several decades of his life doing nothing that work is great and all according to the classic history, but after the rediscovery of Gregor Mendel's work. We basically toss it right we said all of that aside we the biometrical school is sort of falls into ill repute. Not in large in no small part because Carl Pearson is just an abrasive abrasive human being. And so we have to wait for the maturation of these kinds of methods of statistics or uses of concepts of chance to be sort of reinvented from the whole cloth by people like are a Fisher. I'm not so right JBS halting the sort of classic architects of the modern synthesis, but on the classic history. It's a reinvention story, right they are. They are starting over from scratch these these synthesis architect authors, there's really anything like a continuity here. And so this is can be framed as a sort of impact of a Mendelian Revolution, right you could say that there was this approach that was being developed slowly surely to doing evolutionary science over the course of the 1880s, 1890s 19 odds. Mendel lands like a bombshell in the middle of all of this, and that sort of resets the story. A brief aside that I should that I should make right here that I think is very important to underline. Of course, another thing that's very important that's happening during all of this history is that all of the people that I just mentioned save Darwin are the architects of eugenics as a movement. And I want to be very clear here, I am not a historian of eugenics eugenics, the history of eugenics is a very strong, a very well represented discipline there are really phenomenal people working in it, I'd be happy if if you're interested during the period to get some references about some of the authors that have really shaped my view about how to understand that side of the thought of these figures I'd be happy to share them. But I'm not going to get into that today that sort of touching the history of eugenics tends to be a morass and this isn't a eugenics book I don't I don't want this to be another sort of worse copy of the work of someone like Pauline who has resumed or who's done phenomenal work on the history of eugenics already. There's no need for me to write that book again badly. So with that with that very important, very important caveat. What's the problem with this history why don't I like it why do I feel that an important goal of this book is sort of revising revising this story. The whole answer to that question in some sense is the whole book so I go through the work of Darwin the work of Galton the entire trajectory of the Biometrical School, the bit today on the period between modern synthesis and I close with some exegesis pretty careful exegesis of our Fisher, the ways in which he builds on the the ingredients that came before but let me just point out two problems to weird, weird features that got me started thinking about how we might want to tell another story why telling another story might be a good idea. First of all, the following question was the triumph of Mendelism really that straightforward. Is it really the case that Mendel's paper is rediscovered in late 1900 widely disseminated over the course of 1901 is really the case that everyone drops everything and becomes Mendelians over the course of like 18 months and very later you hear this as a sort of very quick way of describing what what went on. This isn't true at all. The story is in fact much more complicated than this and I just want to point to two different ways of trying to think about that. First is a sort of analysis by by Kim it's a it's really a sociology of science analysis. The goal is to try to talk about how consensus forms in a community. What Kim really underlines that I think is absolutely brilliant there's there's problems with this book but I think it's a I think it's still required reading for anybody who's interested in in picking up this this period of the history of biology. What Kim really underlines is there's a huge role that's being played not just by these major figures, but by a whole host of smaller figures some of whom I'll talk about today. Work to sort of move the needle on public opinion about biometry and Mendelism, and that's a much slower, much murkier, more complicated process than we might have given it credit for and a second source that really makes this point well as well as as an article of Rachel Ankeny so you really cast doubt on whether one of the classic figures here that we talk about having been converted to Mendelism as someone who started working with the biometricians and becomes a full a full fledged Mendelian very quickly after Rachel I think very compellingly that this conversion metaphor just doesn't make sense the sociology the experiences of the science of the scientists what caused them to change their minds, how, why, when is way more complicated. These personal stories deserve to be analyzed in detail and when we do that, we're going to get a much more sort of reticulated confusing history that I think is worth spending our time trying to understand. So that's the first kind of question that got me made me begin to wonder whether or not there was another story that could get told here. The second question has to do with biometry itself. And so on the classic history right there's all this work that goes on between a basically roughly 1880 and 1905. That's just useless. Right. There's a 25 year false start. Sturtevant says this very, very clearly. The personal quarrel here is talking about between William Bateson and WFR Weldon certainly delayed the utilization of powerful methods of statistics and much of genetics and so there's this sort of wistful nostalgia, like if only we had been able to recognize that we could just all get along. If only these people hadn't been such obnoxious characters on a personal level obsessed with fighting with each other. Now we would have been able to develop the modern synthesis maybe 2020 or 30 years earlier than it in fact arrived. And so there's this. There's this sense that that biometry was a waste of time. And so that that there's a lot of reasons personally that that that doesn't sit right with me not least, it seems as though, of course, it's possible that an entire branch of science like that could be ruled out as a waste of time that feels like a sort of reductive, much too quick way to try to understand the history of what's going on here. Secondly, something I've some things I've worked on in the past, well before I picked up the book project, the turnaround this question of their sophisticated philosophy of science that's taking place in the community from the 1880s to the to the late 1900s, really cool stuff they're thinking very very reflect a self reflectively about what it means to be introducing math to be introducing chance to be introducing randomness to be introducing statistics into do into the doing of evolution because they knew that this wasn't a thing that we had needed before classic morphological approaches to biology simply hadn't been mathematized yet. I think in the Q&A if you're interested about the ways in which Mendel is implicated here, there's actually there's some really neat things we could things we can threads we can pull on. But in general there's there's a really that they're thinking really interesting thoughts, but if it's all abandoned anyway why should I care about it, right why should I spend time trying to analyze, other than again because I'm like personally obsessed with what some people were doing because I find their work, their evolutionary theorizing really interesting. Why bother right why bother spending time. And I think what I want to do today is sort of bring out this philosophical work is trying to help you see what philosophy of science was going on in this period at this time at least briefly to briefly, but then show you how you see it being preserved in the community of practicing biologists straight through to the early days of the modern synthesis so far from being a sort of 20 year false start that it was thrown away by the following by the following work of the biological community. In fact, the kinds of issues that the biometricians were interested in raising are important to the community they're followed they're there they're picked up upon they are in the air to use a classic but very weasel history, history phrase. They're in the air during this period. And so there is sense in trying to understand how they were developed how they were preserved why they were preserved when they weren't preserved. Why weren't they preserved etc etc. This is a story that deserves to be told. So let's start by looking at what the late biometricians were doing. So this is wfr Weldon. I love showing this photo because he looks like he's dropped straight out of a Western movie from, you know, the spaghetti Western era or something. History does get one thing pretty much right. And that is, after Weldon dies, young, unexpectedly of an illness in 1906, April of 1906, the traditional biometrical school for all that there's a sort of dedicated cluster of people as Pearson Weldon but there's also loads of laboratory assistance calculators etc etc there was a whole whole structure here it span to universities. Weldon was at Oxford and Pearson was at University College London. So there was a there was a real institutional sort of force to this. It all goes away. It all it all it all disappears. And that's pretty much that's pretty much right. And a little bit when I talk about this this sort of the period between the death of Weldon and the early works of the early works of Fisher but for now let me just leave that there. So what I want to do then is look at the view of sort of late biometry the view in particular that Weldon was developing in his manuscripts and his notebooks over the course of 1905 and 1906. He was resembling to work on someone who worked themselves to death he quite literally he quite literally killed himself with with overwork. So here's my reconstruction. It takes two chapters of book to really make the pitch for this to really tease out, not only how this, how this approach comes out in the in the biometrical school but also the biometrical school itself of course changes radically as not only technical methods are developed more data becomes available but also the philosophical sort of underpinnings, the approach to thinking about why statistics and chance are interesting for evolution. That whole approach is radically modified over the course of the few decades of the biometrical school. I'm only going to be able to give you a tiny window into this I'm sort of giving you the finished product I'm giving you the theory that I think Weldon was trying to build when he died. And that's on the basis in particular of an entire book manuscript well, and a fragmentary book manuscript but a significant chunk of fragmentary book manuscript that he left behind at his death he was actually in the process of writing a textbook of his thoughts on everything from and very interesting. This is entirely unnecessary aside but I'm going to go here anyway. Interesting as well for thoughts about what was left in or left out of the modern synthesis Weldon's textbook includes reasoning thoughts about cellular biology thoughts about development thoughts about natural selection thoughts about statistics and mathematics for Weldon this is all a sort of unified picture. As regards evolution though evolution in the narrow sense. Let me let me try to let me try to reconstruct this. Well then was in search of a statistical theory, which could let us understand the action of natural selection across generations at the population level, and which could be harmonized with Mendelian inheritance, at least as a special case so back all four of the things in blue there I don't want to not just kind of believe that. So, statistics, why statistics, why were the methods of statistics so important. So a lot of things I could say here, but the thing I want to at least want to start by underlining is, there's too many counter examples to Mendelism that these scientists are viewing the people like Pearson and Weldon are viewing in the data that they have available to them about heredity and inheritance so here's an image from a 1902 paper that Weldon offers. These are this is actually a photographic plate it's reproduced at great expense in color in a journal. There's all kinds of correspondence about how they're spending a jillians of pounds on on getting this done. Well then wants to underline here we could just look at the top row right. Remember that if you're if your mental peas are either green or yellow. And so you're supposed to very easily be able to differentiate peas into the green category and the yellow category and Weldon says well hang on. Here's a bunch of peas and by the way he mentions in the journal article. I'll mail you my extras I have a lot of extras if anybody wants more peas just send me a letter and until I run out I'll send them to you for free. Here's a here's a spectrum ranging from really green to really yellow but through yellow we green and green yellow and all the colors in between. Well then reads a well then looks at a photograph like this and says there's just no way to think about character traits that doesn't involve statistical distribution. Everything comes in degrees, practically. There are a few exceptions things like eye color we know that but it to first order the case that matters is just is evenly distributed uniformly distributed statistically distributed characters, not necessarily even your uniformly strike that statistically distributed characters. Perhaps as a special case, real mentalism might happen in some characters things like I things like the color of eyes, but that's going to be this that's going to be the exception not the rule. Natural selection is always important, in part because the biometricians consider themselves to be sort of arch gradualist Darwinians. And so they're really interested in explaining how the mechanism of natural selection could be taken to work. I think part of the reason for this that that comes out over the over the course of of my book is is that Galton had made a mess of this in his work God's, God's approach to natural selection in particular is just it's it's confused and confusing it was confusing at the time to his to his colleagues. And so the biometricians really saw okay we gonna we need to make sense if we're really going to say that we're Darwinians we really need to make sense figure out how to make sense of, of natural selection. Next up ever a population level so evolution is always a distributional population phenomenon for the biometricians on the one hand this comes sort of naturally as a consequence of the fact that they're focused on distribution traits statistically distributed that that sort of follows follows directly, but also they saw here another weakness of the Mendelian approach that is to say you're too preoccupied with deriving stuff the biometricians would say from a single mating of two parents and what offspring even if in a Mendelian case you are indeed you're breeding many parents of course Mendel. Evidence is some sort of proto statistics in the way that he approaches the populations of peas. Setting aside a detailed mental analysis which is absolutely a different talk. The biometricians are still going to say you're worried about a sort of one generation mating event. And you're worried about a one generation mating event from one particular set of parents, you're not thinking about the populations a whole, and that's a that's a blind spot. But, and this I think it's very important because this is something that the classic history gets entirely wrong after about 1903 or 1904. There's no opposition to Mendelian inheritance as something that happens. Sometimes, not all the time, not perhaps even frequently, but mentalism they think is a special case of what's happening overall in the transmission of of characters. It's important to remember right for those of you who who have worked in later periods in the in the history of biology. At this point this is still Mendelism with complete dominance it's Mendelism with two alleles usually the rest of the complications to that come later so actually think it's not weird at all to look at that and say, yeah okay I mean maybe this is maybe this happens, but it's not going to happen that frequently. And so, well done well done writes to Pearson and you know three and says it's easy to say Mendelism does not happen for that it just takes right the right kind of statistical distribution you just have to show. You just have to show a non Mendelian ratio arising in the transmission of a character from from parent from parent generation to an offspring generation. It does happen it's harder every day. Right so how do we actually understand what's going on here if we're going to reject Mendelism as sort of a universal foundation for the inheritance and transmission characters. What are we going to build in its place. Okay, so that's the kind of theory that well done wanted and of course I'm not giving you the details about his proposal for it that would take another different 45 minute talk. Absolutely you can ask me in the Q&A I could talk about it for hours. So I'm not giving you the details what I'm giving you though is this this philosophical orientation right. This is what the biometricians thought it meant to say that we would have built a good approach a good theory that took account of the phenomena that they were important. These are the reasons for which they thought it was necessary, not just not just useful, but absolutely essential to make evolution of chancee statistical process. Okay, well done dies well done well done dies what happens after well done's death, the classic history has this is a period that sort of devoid of theoretical import. Maybe Provine talks about this there's a debate in here about continuous versus discontinuous natural selection so this is the sort of mutationist saltationist debate does evolution proceed by big steps or by small steps. That's, that's an interesting debate I could I could say something about it if people want me to but it's not really I don't, I don't see it as being as all encompassing as is often taken to be to be the case, historically, by by historic historiographically, I should say. Now, it is true. And this is why I think the big the popular history the the classic history became a classic. It is true of the two major players that everything dies and they quit talking to one another so this is Carl Pearson in I believe 1911. Now, Pearson basically moves on to only work on eugenics. He keeps getting funding for the eugenics laboratory he becomes the director of the Gaulton eugenics lab after Gaulton's death at at university college and so he basically transitions into full time full time eugenic and statistical work as a supplement. He doesn't like fighting about biology alone. He liked doing it when Weldon was around he actually writes letters to this effect. Now he likes doing it if he had Weldon with him. But he doesn't feel like he has the right kind of facility with biological details with biological knowledge the facts of the case to really be able to sort of stand in a debate at the Oxford Museum and fight with William Bates and he just doesn't like it. He just stops doing it. For his part William Bates and shifts into sort of full blown condescending dismissal toward anything having to do with mathematics. So Bates and puts in a book as early I think is 1909 something something to the effect of you know it's it's quite nice that I don't even have to waste my time talking to you about all of the nonsense that went around went along under the turn under the under the name of biology because nobody likes it anymore. So base so as far as these these big towering figures are concerned I think the classic history in some sense has it right right Bates and goes on to being a sort of incremental Mendelian developing his Mendelian perspective Pearson just leaves the field essentially stops there's an asterisk and a caveat here. I owe I owe a more nuanced reading of Pearson post 1906 to Mike but all to give a great talk about this at the last ish yeah that was the last ish. Yeah, but but but long story short, it still is I think generally true that these these towering figures are done. And so what I want to do now is look at the look at some of these people that we don't normally talk about and the argument that I want to make first I want to pick up some people who I think were combining some of these threads, not all of them not all five of these sort of desiderata for the final theory of evolution that the biometricians had been advancing, but that people were sort of keeping each of those threads alive over this kind of over this kind of interregnum period as I've as I've as I've called it. So first, as a tradition of using precisely the same kinds of statistical methods that biometry that Pearson had developed that Pearson had developed well than it picked up. Without grounding in any kind of statistical view of inheritance so not saying that these will sort of directly straightforwardly describe how characters are transmitted from parents offspring but still that the methods of statistics. Contra, for example, that quote from start event that I showed you earlier are being used everywhere so here's William Castle working with his I believe former graduate student Hansford Hansford McCurdy. It started William Castles a very important Harvard mammalian geneticist he's often described without further ado has just sort of oh yeah well he's a one of the he's an American Mendelian. But here's his major report from this period from this interregnum period with McCurdy on rodent coat genetics. And he's using Pearson Indian statistics so of course he says you know, I don't mind just giving a kind of informal gloss on frequency tables, but for those who place confidence in the more precise methods of statistical analysis devised by Pearson and others, it may be more satisfactory to treat the tables which have been constructed for the various groups of individuals as correlation tables and derive from them the constants which measure the variability of parents and children respectively in the several groups, and the degree of correlation between the two. So he's doing exactly the same kind of data analysis that Pearson would have done if you'd given Pearson these these these sets of rodent coat data. There's not any sense in in in anything as a slight dismissiveness just before this quote. There's not really a sense of kind of warfare going on here. There's really just yeah easy if you if you like these tools. There's a hint under there of course if you're competent to do that to handle the mathematics in these tools. Here's what here's what they would tell you. Here for William Johansson, whose treatise on evolution includes extensive methodological discussions of the use of statistics to analyze genetic data he lobbies in the introduction for more teaching of statistics to biologists. So really knowing really evaluating a theoretical proposal in evolution he writes is only possible however if the methods that have been or at least should have been followed in the research or given special consideration. An essential aspect of these methods is their mathematical character and can be described as applied mathematics familiarity with these methods is absolutely necessary for a real understanding from many hereditary questions. And so there's several chapters is a big chunk of you have Johansson's work dedicated to grinding out the statistical method. Of course, and just to underline the kind of personal disagreeableness here so Johansson at one point writes to Pearson and says hey, we should, you know, probably collaborate because I think your stuff is really good. And Pearson basically says I don't want to talk to you go away. Please leave me alone. So this, you know, Pearson is Pearson is a problematic figure. So next up, let's look at early approaches to mathematical inheritance. So trying to mathematicalize a sort of a Mendelian approach to inheritance that don't involve statistics. So this is Herbert Spencer Jennings a Johns Hopkins geneticist who pioneered some of these early some of these early mathematical methods that are that are at work in what I've kind of this sort of American school of doing this kind of longhand mathematics. They're grinding them out as algebraic recurrence equations. So this is a rule take the series of results as finished products and make an independent study of them endeavoring by processes of trial to fit them to some series or to some formula. It's here that there is scope for ingenuity, a given series of results may resist for weeks, the discovery of the law that unites them after a law or regular series subtane that fits the first five or six generations the laws applied to give the results for three or four generations These results are then tested by the actual detailed working out so actually drawing up symbolically like the gametes that would be in the population actually mating them out longhand to try to make sure that your formula gave you the answer that you wanted. And this is what Jennings did Jennings spent two or three years doing basically just doing this from like 1914 and 1916. It gives these awesomely unreadable papers. Where he just picks different ways that you might imagine a breeding experiment different patterns of mating, assortative or non assortative different initial populations, etc, etc, and crunches out these recurrence equations and then derives. Over here on the right you see one thing that he's interested in of course thinking about breeding experiments. He's interested in the steady state. So what are the long term steady state ratios between the different the different phenotypes and how many generations would it take for you to get there so here you've got me so you get to 74% homozygous dominant after 72 generations right that could be useful if you were trying to I don't know if cows or something and you wanted to know how long it might take you to get to 74% homozygous dominant in your cow pasture or what have you. So, cool as far as it goes. There's there's there's more. There's more to say about the story Rainer Robbins of Michigan mathematician who interestingly enough would go on to run the academic investment firm Tia who might hold your retirement depending on what university you work at is the first to start doing other mathematical induction based proofs here it seems pretty clear Jennings doesn't know how to sort of handle mathematical induction which is kind of the first a proof method that would seem like if you were looking at this from a perspective of contemporary mathematics you would think that's what you would be doing. But they're still painful. This is hard to do with algebraic tools. Edward Murray East. He worked also primarily at Harvard with the same departments castle East is important for my story here in being one of the many biologists to re derive the work of the Swedish botanist Nilsson a la, who demonstrated that large characters of independently assorting Mendelian characters could produce the appearance of statistical distribution so this is how to get a sort of statistically distributed character without moving to the population level without abandoning the kind of standard structure of idealism. So you're right here when one considers the difficulty of distinguishing the zygote setting various genetic formula even when dominance is comparatively perfect. You might expect the population of offspring individuals with almost continuous quantitative variation if dominance is imperfect or absent. This gives a clue to a Mendelian interpretation of the inheritance hitherto known as blended so he's talking here about imperfect dominance. And that's where about if there's a large number of Mendelian factors that will also produce the appearance of statistical distribution but this was clearly something that was very important to him. Emerson sorry East spends quite a bit of time thinking about how those kinds of characters might arise in part because he sees them. Emerson in East 1913 is a report on corn. He sees them in his experimental subjects so for exactly the same reasons that people like Pearson and Weldon were interested in. And last and perhaps most importantly this is a new you'll you'll start out as an assistant of Pearson's but he offers us an early attempt at a synthesis between biometry and Mendelism really actually picking up all five of those desiderata you'll has the same goals for an evolutionary theory that the late biometricians did. Again, I could spend an entire talk laying out you'll approach but let me just pick out a few quotes. So first he's reinterpreting Mendel as offering a sort of mechanism for the kinds of statistical and distributional facts that were being studied by biometry. The law of Mendel and his successors he writes lies not in discovering a phenomenon inconsistent with that law, the law of ancestral heredity one of the core mathematical pieces of biometry, but in showing the process consistent with it though neither suggested nor postulated by it might actually occur right so this is something that really could happen that could give rise to Mendelian results. And this should be what a synthesis, a new theory should offer us right away to show how we can get from the statistical constants describing a population to the causal influences at work on the population at issue right so what's required from the physical theory of heredity he says a bit later on is that it should assign a meaning to the variations in the constants that do occur, enabling one given the law of ancestral heredity for an organ to state the relative influences there on of the different agencies concern selection in all forms circumstance and so forth. Okay, so much for some particular instances of particular people keeping some of these biomedical claims in the air. That's fine as far as it goes for a few for a few people in particular but would people have really known about this work so that was the second the next question I wanted to tackle here. How would anybody were people actually reading this or were they being exposed to these ideas perhaps in a different way. I think the answer is yes. So, the way I want to show you that is to look at the textbook tradition so I did a deep dive on on three of the major topics that were at play in exactly these relevant years Robert Keith locks recent progress in the study of variation heredity and evolution that's a 1906 book. J Arthur Thompson's heredity from 1908 and Ed Edwin as good riches, the evolution of living organisms. And so I'm going to generalize for you because you know time time ticks by but but I want to bring out three big features. First, there's no evidence of open warfare between these traditions so here's a quote from good rich. These variations can often be accurately measured and the statistical study of variation begun by Katelyn Galton carried on by WFR Weldon K Pearson and others has yielded many important results. This is right after talking about some kinds, perhaps not all kinds but some kinds of biological variation there's nothing of the kind of condescending scorn of William Bates and anywhere in any of these three, these three textbooks. There's a general belief that we want a synthesis a synthesis would be good. So here's JR Thompson who writes developing a new vocabulary for picking up evolutionary questions is all the more justifiable so we cannot doubt that all the ordinary phenomena of a piece. Many of the ordinary modes will be embraced eventually in one general formula, probably some modification of Galton's law of ancestral inheritance and that others will be embraced in Mendelian formula. So, there's a little bit of a little bit of an equivocation here it's not super clear that this is what he means Thompson is not a good writer this book is actually painful. But let me assure you that what he means here is that we want a theory that can treat these two classes of phenomena as deriving from some kind of common foundation. And that's a thing that all three of these textbook authors make really clear over the course of their their discussion. There's caution against excess, and that's true. There's a to the extent that there's a sort of negative, negative side to this here's a quote from locks book he says you know some students of biometry however would go very much further than this. This is the best position that their own form of study is the only method by which any real advance in our understanding of the processes of evolution can be brought about. This opinion is based upon the assumption of which proof is wanting that new species have arisen exclusively through the accumulation by natural selection of variations of a strictly indefinite fluctuating or normal kind. Notably here he frames this though as sort of saying yeah the jury's out on whether or not gradual variation is actually produced all species. But perhaps with Bateson, Mendelism is showing us a way that larger variations could take place and perhaps create a species sort of from whole cloth much quicker. Okay, that's, that's what I want to do in terms of showing you that that the biomedical view stays very much in play over this period. Now I want to do even a little bit more precise digging and try to argue that it does actually show up in the architects of the modern synthesis so it gives us some common ground that played a real role in the construction of the modern synthesis. So Locke's book was really important. E.B. Wilson uses it to lecture at Columbia, it is by some anecdotal accounts the book that convinces H.J. Muller to study genetics. He's in the, he's in the lectures he's reading the book and he goes crazy. The rest of the Morgan group as well including Sturdivant Bridges definitely talk about it we have a we have a we have a testimony to that effect. He reads it during, reads it during graduate school. These are all figures that will go on to play crucial roles in the development of later 20th century biology. Goodrich is a really interesting character at Oxford, a crucial player in the sort of Oxford biological scene for decades is a great chapter. Another hat tip to Greg Radick who found this there's a chapter on zoology in morals books science at Oxford 1914 and 1939 that really goes into detail about Goodrich's lab and what Goodrich is up to. This connects him to Edie Poulton and the Huxleys so these are other you know very important figures over the course of this period in early genetics. This is published for years by James Mark Baldwin so there's American connections here as well that are kind of cool. Goodrich's textbook becomes recognized as an absolute shining exemplar of doing good pedagogical popularization of evolutionary theory. Thompson's does as well. This is talked about extensively and very clearly in Bowler's science for all it's about popularization of science during this period. Haldane so there's Haldane on the left in before the war. Haldane's only formal training in biology comes from attending Goodrich's lectures at Oxford that sees otherwise a mathematician. So he's he's getting his evolution from Goodrich are a Fisher and here we have some amazing detective work from and the Edwards who crawl through the library to find the copy of locks book in the library of the ox of the college at which Fisher did his undergraduate studies. It turns out there's some there's some hen scratch in the corner next to one of the diagrams that definitely is Fisher's because Fisher had really weird handwriting. So Fisher studies Bayton's textbook he studies Pearson's articles but he also and he also studies locks textbook so we know that there's a real a real connection between these folks. These books are these books are mattering these articles are mattering this stuff is is is absolutely feeding into the intellectual environment at work around around this time. There's a pragmatist connection here I'm just I'm just tossing this in for for Trevor. Visado argues that the American side of this and especially including Jennings and East but but several others right. So one thing you might notice I talked a lot about Harvard and Hopkins I also could have talked about Chicago. These are all major institutions where pragmatism is being developed and all of these biologists later on in their careers become super interested in social and ethical issues. There's a really interesting question about about whether or not they're sort of absorbing pragmatism in the in the coffee room. I would I want to pick that up someday I didn't have a chance to do it for this for this book project yet. So in short let me give you a couple concluding thoughts I'm a little bit over time already so let me let me try to wrap up fast. The story is a lot more complex than we thought right if we look for the elements of a late biomedical worldview, they absolutely persist through this period straight through this period from from 1905 to 1920. They absolutely influence the architects of the modern synthesis we know that they were exposed to all of this writing that I've that I've talked about today. And so there's there's not a sense in which biometry has died biometry has has sort of been metabolized in interesting kinds of ways been transmitted in interesting kinds of ways. But that's not to say that it's been sort of scoured from the earth over the course of this period. I think this narrative can serve some really good HPSC type goals history and philosophy goals. On the one hand it can sort of deemphasize the revolution of the early synthesis right I think it's good to not saddle people like Fisher and right important as they were. I talk about their papers in the book I think we have to understand them, but the idea that they sort of recreate evolution from 1918 to 1925 or so. I think that story doesn't hold up right there's a very nice sort of classic gradualist intellectual transmission story unfolding over this period. And so that's that that story of continuity I think is is important I think it can help us produce more grounded sort of more plausible, especially from a sociology of science perspective more plausible stories about what actually happens here. And three, this is the most philosophical point again to double back to one of the things that started me on this whole journey. I think it helps us focus on what it meant during this period to bring chance and statistics to evolution for each of these different authors and these are choices these are active choices that are being made. They're choosing very directly to engage with parts of this tradition to leave other parts aside. There's reasons. Institutional reasons model organism related reasons theoretical philosophical especially for are a Fisher who's personal personal philosophy of science is crazily interesting. I think it helps them to pick up those ingredients and run with them in different ways and I think that nuanced historical philosophical story is is one that that I'm really happy to have been able to engage with and at least in a tiny degree bring out for you guys here today. So, with that, thank you very much this is normally the part where I would ask you for questions but I will see you all very soon in the in the live zoom Q&A. So on any of this as well feel very free to drop me an email as I said this is a sort of live active project. I'm already revising this chapter after writing this talk because I thought of some better things to add and phrase differently. So thanks again very much for the opportunity and I'm really looking forward to talking with you guys here soon. Bye bye.