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
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Eric Charles

  Professional Student and
  Assistant Professor of Psychology
  Penn State University
  Altoona, PA 16601


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