Thanks, Eric. That puts it nice and succinctly. That said, I take the points about how best to characterize "fitness" and will adjust my draft accordingly (and I had some chuckles over the lighter responses too). I'll revise it and re-send it sometime over the next few days (it might be old news by then, but at least it motivates me to keep working on it!). I've just seen Vladimir Burachynsky's post, and will respond to that momentarily too.
Hugh ----- Original Message ----- From: ERIC P. CHARLES To: Nicholas Thompson Cc: Hugh Trenchard ; Friam@redfish.com Sent: Monday, March 29, 2010 10:13 AM Subject: Re: [FRIAM] Sperm pelotons; article in Nature But Nick, Hugh's point is that we DO NOT need trait-group selection to explain the clustering sperm. We merely need sperm to swim in the same direction, AND have a variety of abilities. Given that alone, Hugh thinks he can prove, sperm will cluster based on their swimming abilities (which he calls 'fitness'). Thus I (captial 'I') declare that the real empirical question is whether or not sperm-in-clusters are more genetically similar than Hugh's model would predict. Only if THAT were true, would we conclude that group selection was involved, as the authors of the Nature article have claimed. That is, the authors of the Nature article have a flawed notion of what would happen by chance if sperm were swimming along without 'relatedness' detectors, and hence they have a flawed 'null hypothesis', and hence they have a flawed statistical test. (This is all in the same sense that Schank's models have convincingly demonstrated that the results of so-called 'menstrual synchrony' research are exactly what you would expect due to chance. Those who think they showed 'menstrual synchrony' just have a flawed notion of what happens by chance.) Eric On Mon, Mar 29, 2010 12:30 PM, "Nicholas Thompson" <nickthomp...@earthlink.net> wrote: Hugh, I yield to no man in my ignorance of subject we are talking about. However, two points: The term "genefur" is one I use to remind myself (and anyone who happens to be listening) that the common expression, "a gene for", (as in "a gene for blue eyes" or "a gene for prostate cancer" is deeply problematic. I should probably say something with more words, such as, "a gene for peletonizing, whatever the hell that might mean." Although we know that the path from a trait in parents to the same trait in an offspring is much more tortured than a Dawkinsian argument requires, and that the material basis for parent-offspring is not as "atomic" as the expression "a gene for" implies, we continue to need a term for a unit of inheritance and "genefur" is a quietly ironic way to speak of units of inheritance while acknowledging that that sort of speech is silly. As I understand this discussion it has a lot to do with the group/individual selection argument. Think of it this way. Think of a bike race containing 20 riders from 5 teams. Let it be the case that the winning TEAM takes down all the prize money but that it is shared unequally by members of the team, with half taken by the winning rider, a quarter by the second rider, and the an eighth by the 3rd rider, and the balance by the fourth, etc. Now we have set up a conflict between group level and individual level success. My comments on fitness are only to remind us that "fitness" in a Darwinian conversation means winning the race by any means. In your terms, "fitness" means using your resources to produce the maximum output. Call these "fitnessD" and "fitnessT". One could be "fitT" all by oneself on a stationary bike. However, as the scene in Breaking Away demonstrates, there are lots of way to be "fitD" without being "FitT". I wish we could engage David Sloan Wilson in this discussion, but he is too damned busy running around the world being famous and talking about the evolution of religion. Gawd I hate when that happens. Nick Nicholas S. Thompson Emeritus Professor of Psychology and Ethology, Clark University (nthomp...@clarku.edu) http://home.earthlink.net/~nickthompson/naturaldesigns/ http://www.cusf.org [City University of Santa Fe] ----- Original Message ----- From: Hugh Trenchard To: ERIC P. CHARLES;Nicholas Thompson Cc: friam@redfish.com Sent: 3/29/2010 9:42:09 AM Subject: Re: [FRIAM] Sperm pelotons; article in Nature Thanks Eric for taking the time to look through my post. For Nick's last post, I am not entirely sure what a "genefur" is, although it sounds like it is a reference to an inherent genetic trait, as you also discuss. Yes, I agree it will help my argument if I hone in more closely on what I mean by fitness, and I will add some description to clarify this. My useage relates to inherent physical fitness in terms of maximum power output capacity. That too needs fine-tuning because I refer to "maximum sustainable output", which is not the same as absolute maximum power output, and I would need to outline more carefully what this means. Regardless, I think there are ways of testing for the actual power-output capacities of individual sperm - I have seen references in the literature to testing procedures for this. Because I know very little about genetics, for my part I would be treading dangerously to begin describing the process in a gene-related sense (and I would not want to get into discussion about chromosomes), but to address the issue you raise (if I understand it correctly), it would be necessary to measure the power output of the sperm of individual male mice to determine the range of their output capacities and/or the sperms' average output. This is no doubt not easy, but I imagine there would be some sampling size that would provide an accurate indication of the overall output range. And certainly one would want clearly to correspond average sperm outputs and ranges with the genetic descriptions of the various mice tested, but this could be done according to a replication of the Fisher and Hoesktra procedures. It would also be necessary to determine percentages! of ! energy savings that occur when sperm are coupled (if this does in fact occur). My model assumes that there is a difference in the average power output of individual males' sperm, whether related or unrelated or of the same species or not - a difference sufficiently significant to demonstrate that sorting occurs according to fitness (in the power-output sense) and not according to some mechanism for identifying the genetic relatedness of the sperm, as the authors of the Nature article appear to suggest. The fact that sperm aggregate indicates coupling and energy savings, which is why (in my view) the peloton model applies. In terms of chance, it seems to me Fisher and Hoekstra have taken a lot of care to establish that there is sorting beyond chance, but implicitly ascribe that sorting to some sensory/perceptual capacity of the sperm to identify related sperm. My model begins with their proven result that there is sorting beyond chance, and asks whether there is some sorting mechanism involved other than an unidentified mechanism to perceive the location of related sperm, which is intuitively problematic because (it seems) sperm do not have a sufficiently developed sensory system (i.e. eyes, ears, or other) to do this. My model provides a simpler explanation for the sorting process than the Hoekstra & Fisher explanation, because, in my model, sorting occurs according to self-organized energetic principles, and not according to a perceptual/sensory mechanism, as apparently implied by the authors. I can see how a basic computer simulation would be helpful as a starting point for making predictions according to my model, which I see is really my next step. Does anyone know how/where one could apply for some funding to resource such a simulation? I could develop it myself (and have developed at least one simulation, but it really needs to be worked through again), but it would happen a whole lot faster if I could engage someone more adept at computer modelling than me. ----- Original Message ----- From: ERIC P. CHARLES To: Nicholas Thompson Cc: Hugh Trenchard ; friam@redfish.com Sent: Saturday, March 27, 2010 2:54 PM Subject: Re: [FRIAM] Sperm pelotons; article in Nature Hugh, Very interesting model! One of my doctoral adviser's, Jeffrey Schank has demonstrated repeatedly that scientists are very bad at predicting what 'chance' looks like when trying to do experiments involving synchrony. This seems one of those situations, and the only way around it is modeling. Nick's sarcasm aside, he has a point, and it has to do with some of the flavor text surrounding your model (for geeks of the wrong variety to know what flavor text is, see: http://en.wikipedia.org/wiki/Flavor_text). If I can take a shot at identifying the problem: Rather than looking at 'fitness' as if it were a unified trait, you have created a model that needs some mutli-stage selection language (the better term escapes me at the moment). The reality is that what makes a 'fit' sperm is not necessarily what makes a 'fit' orga! nism. To fix the flavor text of your model, you would need to explicitly recogni! ze that (if the sperm sort, then) the sperm are going to sort based on a similarity in the genes that 'build' the sperm. Their sorting will be completely independent of all the other genes, or of any role that the sperm-building genes might later play as body-building genes. Ignoring chromosomal linkages (which you shouldn't), two sperm could be identical on all the genes important for building sperm, but completely different in terms of all other genes. Your model would thus al! low a much clearer test of the prediction that sperm identify each other in some way. It does so because it provides a vastly improved predicted relatedness due to chance. GIVEN: We would expect sperm to cluster along the race track based on the similarity of certain, specifiable genes. MODEL: If we know the genes important for building sperm, we can model the expected relatedness of sperms within a cluster. IF: Sperm are implementing some weird sort of kin selection mechanism - THEN: we wo! uld expect the relatedness to be significantly larger that what our mo del predicts. Any help? Eric On Sat, Mar 27, 2010 01:36 PM, "Nicholas Thompson" <nickthomp...@earthlink.net> wrote: Hugh, Even if it has nothing to do with sperm it is a nifty model. There is an idea lurking here that i dont know whether it plays a covertrole in your thinking or not, but what about the fate of a "genefur"peletonizing. My email program is misbehaving and my computer is about to crash so I wontsay more, now. Nick Nicholas S. ThompsonEmeritus Professor of Psychology and Ethology, Clark University (nthomp...@clarku.edu)http://home.earthlink.net/~nickthompson/naturaldesigns/http://www.cusf.org [City University of Santa Fe]> [Original Message]> From: Hugh Trenchard <htrench...@shaw.ca>> To: <nickthomp...@earthlink.net>; The Friday Morning AppliedComplexityCoffee Group <friam@redfish.com>>! Date: 3/27/2010 10:54:41 AM> Subject: Re: [FRIAM] Sperm pelotons; article in Nature>> Thanks for taking a peek at my post. Great que! stions, and they help me to > see how/where my descriptions can be clarified.>> On the paradox part - that is one of the really interesting features of a > peloton: the energy savings effect of drafting narrows the range offitness > between the strongest and weakest riders. In contrast, think of a packof > runners of varying fitness levels. There is negligible drafting effect - > there is some, esp if running into a headwind, but overall it's smallenough > that it can be ignored for this illustration. Say there are 50 runners,all > separated incrementally by 1% difference in fitness; say they run acouple > of miles. If they all start off slowly at say the max speed of theslowest > runner, they can all run in a big group, separated only by enoughdistance > between them to keep them from kicking and elbowing each other. As they > pick up speed, the gr! oup thins into a line and are separatedincrementally &! gt; by d istances that correspond to their differences in fitness. In thespace > of two miles, they all finish individually in a single long lineaccording > to their fitness, and it can be predicted accurately where runners will > finish if you know their starting levels of fitness.>> This is not the case with a peloton. For example at 25mph, riders cansave > at least 25% by drafting (approx savings 1%/mph) - all theriders who are > within 25% fitness of the fastest rider can ride together even at the max > speed of the strongest rider. So their fitness levels are effectively > narrowed, and they can all finish together as a group (ie. globallycoupled > by finishing within drafting range of each other), and so theparadox. Part > of the paradox is also that, while fitness levels are effectivelynarrowed > by drafting, it means, conversely, that a broader range of fitn! ess levels > can ride together in a group, which maybe isn't something that is clearfrom > my initial post (though it is certainly implied). Also, thereare other > important things going on in a peloton which precede the sorting ofriders > into groups, some of which I see I do need to clarify to make my model > clearer.>> Of these, particularly important are 1) the occurrence of pelotonrotations, > and 2) points of instability when riders are forced into positionswhere > they do not have optimal drafting advantage. Below a certain output > threshold, when all drafting riders in a group are sufficiently below max > output, riders have sufficient energy to shift relative positions withinthe > peloton, and in this particular phase, a self-organized rotationalpattern > forms whereby riders advance up the peripheries and riders are forced > backward down! the middle of the peloton. However, instabilities in pace > oc cur along the way, caused by such things as course obstacles, hills(when > lower speeds reduce drafting advantage, but when output may be at leastas > high), cross-winds, narrowing of the course, or short anaerobic burstsamong > riders at the front - all of which cause splits (i.e. PDR>1 atthese > points). In a competitive situation, instabilities occur frequently > causing temporary splits at various places in the peloton, but these are > often closed when the cause of the instability has ceased. Sorting thus > occurs according to some combination of peloton rotations in whichstronger > riders are able to get to the front and the continual splits in thepeloton > at points of instability and reintegrations. I would need to develop the > model some more to show this as an equation (though I touch on abasic > version of it in my Appendix).>> For sperm, I! don't know what the initial state of the aggregates are when > they begin their travels, but I am assuming (perhaps quiteincorrectly), > that there is some initial phase in which they are mixed (such ascyclists > on a starting line), and then they begin to sort as they increasespeed. > During the process, they aggregate like cyclists because a broader rangeof > fitness levels can aggregate together (causing an effective narrowingof > fitness). As in a peloton, there are instabilities that allow for > continuous re-adjustments to the relative positions of all the sperm, and > over time they begin to sort into groups where each have fitness levels > closer to the average. This is my hypothesis, at least.>> On the second last question, there would be an advantage to sperm amongthe > first pulse aggregation if all the pulsed aggregations do not mix first,but &a! mp;a! mp;g! t; the principles apply to each aggregation. However, I don't! know wh ether > there is some other process of mixing first among all the pulses of sperm > aggregations before they begin traveling (I imagine I could find theanswer > in the literature), in which case there could easily be a sperm in, say, > the second pulse, which could end up impregnating the egg.>> I don't know about the kamikaze sperm - I'll leave that one for now! ButI > do remember that scene from the movie as clear as day!>> In any event, my aim is really to ask the question - are there energeticand > coupling principles that allow sperm to end up in groups which otherwise > appear to have occurred because genetically related sperm can somehow > identify each other? I am really only suggesting the existence of some > dynamics of the sperm aggregations that could be studied for, which don't > yet appear to have been addressed.>> Hugh>============================================================ FRIAM Applied Complexity Group listserv Meets Fridays 9a-11:30 at cafe at St. John's College lectures, archives, unsubscribe, maps at http://www.friam.org Eric Charles Professional Student and Assistant Professor of Psychology Penn State University Altoona, PA 16601
============================================================ FRIAM Applied Complexity Group listserv Meets Fridays 9a-11:30 at cafe at St. John's College lectures, archives, unsubscribe, maps at http://www.friam.org