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    The Peacock Problem
    By Gerhard Adam | May 24th 2011 04:37 PM | 19 comments | Print | E-mail | Track Comments
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    In a recent set of posts there was a discussion about evolutionary psychology and how it can be used to explain various behaviors.  However, one of the fundamental challenges raised is whether the references to biological phenomenon are, in fact, settled issues and whether they should serve as a basis for drawing additional conclusions.  As a result, I wanted to focus on one particular paragraph from one paper to illustrate the problem.
    "However, costly signaling theory is only beginning to influence thinking in psychology (Miller, 2000). The classic example of a costly signal is the peacock’s tail, whereby the quality of the tail—its size, color, luminosity, and symmetry—serves as an honest signal of the quality of the peacock’s genes to potential mates. A high-quality tail is costly to have because it takes much metabolic energy and esources to grow and maintain such a resplendent ornament, which is useless and even detrimental in other aspects of a peacock’s life; a high-quality tail is an honest signal of good genes because only those peacocks who are in good health and who have the traits required to survive and acquire abundant supplies of food can afford to waste their energy and resources to grow and maintain this showy and nutritionally costly ornament (Loyau, Saint Jalme, Cagniant,&Sorci, 2005; Møller&Petrie, 2002)."

    Blatant Benevolence and Conspicuous Consumption: When Romantic Motives Elicit Strategic Costly Signals
    Here we have a prime example of making a scientific assumption with little or no evidence for the actual facts.  Notice how the reference begins by citing a "classic example" despite the fact that there is nothing "classic" about it.  It simply suggests that this is a settled issue and therefore beyond questioning.

    The basis for this statement is rooted in the idea that the peacock's tail 1, because of its extravagance, is a costly feature that will negatively impact male fitness and consequently its display is an "honest signal" to females regarding the male's genetic superiority.  The first problem is whether this is actually "costly".

    To my knowledge there hasn't been a single study that actually relates peacock mortality to their tails.  In other words, if the tail is a handicap, then there must be some data which shows a higher vulnerability to predation than for those without it.  In the quote it is referred to as being metabolically "costly", although this is a value judgement and has no merit.  Unless it can be demonstrated to materially impede the peacock's ability to survive, by incurring a higher caloric requirement and hence cost, it is a meaningless statement.  More problematic, is the point that peacocks routinely molt and do not carry this tail with them for well over half their lives.  

    The point here isn't that the peacock's tail isn't the product of sexual selection, but rather it is that the interpretation of its costs and benefits is entirely extrapolated without data, often based on little more than our own perceptions.  One thing that is ultimately true about sexual selection, is that for any trait to manifest in a population, it must be more beneficial than any counter-selection that could incur which impacts fitness.  Therefore, we need to rethink the "cost" of the peacock's tail since it clearly has little apparent impact on the ability for peacocks to survive year to year.  There is a paper that attempts to correlate the number of tail feathers to age, but it simply isn't known how this relates to sexual selection in any meaningful way.
    "Such a choosy female would therefore mate with a peacock who has demonstrated his genetic quality by avoiding death by predation, disease and environmental extremes.  Whether peahens can and do assess numbers of ocelli remains to be demonstrated."
    http://www.gbwf.org/pheasants/peacock-train.pdf
    This also creates a secondary problem because it indicates that the tail cannot be a reliable indicator of the peacock's ability to survive predators, since it only grows during the mating season and does not reflect the actual history of the animal.  It is also problematic that the peacock will fan his tail when threatened, since this indicates exactly the opposite behavior that one would expect from an animal with such a "liability".

    Another point made is that because of this "cost", the peacock generates an "honest signal" regarding genetic health.  
    "Moller and Petrie took blood samples from male Blue Peafowl (Pavo cristatus) and recorded the numbers of B- and T-cells, and also measured the peacocks' tails and counted the number of eye spots. They discovered that the condition and length of the peacock's tail was related to the production of B-cells, and the size of the eye spots to T-cell production."
    http://people.eku.edu/ritchisong/birdcirculatory.html
    http://www.springerlink.com/content/j705122208485740/
    So, it appears that the tail may signal a healthy immune system, although it says nothing about the overall genetic quality nor survival capability of the peacock itself.  This certainly suggests that some information is being conveyed by the tail which is of importance during the mating rituals and acts as a "signaling" mechanism.  However, this does not support the claims made in the original quote which indicate "good genes" and the ability to acquire abundant supplies of food.  

    One final problem remains.
    "Combined with previous results, our findings indicate that the peacock's tail (1) is not the universal target of female choice, (2) shows small variance among males across populations and (3) based on current physiological knowledge, does not appear to reliably reflect the male condition."
    http://www.sciencedirect.com/science/article/pii/S0003347207005301
    In this study, it doesn't appear that the size of the tail actually influences female breeding behavior, at least not alone.
    "A seven-year study of feral Indian peahens and peacocks in Japan [by Mariko Takahashi et al.] titled “Peahens Do Not Prefer Peacocks with More Elaborate Tails” notes that these peahens disregard plumage in making their mating choices. The study further observes that variations in male plumage do not reflect anything in particular about the male’s physiological condition. Most strikingly, this 2008 study finds that a high level of female hormones dulls a bird’s coloration, suggesting there has been selection on females with dull plumage. These scientists propose that both sexes started brightly colored. Females evolved dull coloration, which increases survival chances for this ground-nesting bird, since there is little male involvement in parenting and since the female is vulnerable to predation while incubating eggs on the ground. (The scientists also reviewed the literature on peacocks, found it contradictory in terms of supporting or refuting sexual-selection theory except in the sexual-selection field, and suggested that researchers take care to publish negative as well as positive results.)"
    http://www.theamericanscholar.org/the-peacock-problem-by-long/
    In the end, we are left with a "classic example" which turns out to not be classic at all.  In fact, the truth is that much of what we use to explain such behaviors is based on our own quaint notions of how we think this works, instead of building on detailed knowledge that can truly be used as a reliable indicator of biological behavior.  Darwin's theory was successful because it was simple and general enough to provide true insight, in addition to being nuanced enough to allow for varied interpretations.  There needs to be a much more cautious approach in explaining biological phenomenon, lest we engage in building ever flimsier hypothesis on little more than "just-so" stories.
    ---------------------------------
    Note that the peacock's trail is actually a "train" of covert tail feathers that comes off the back of the animal. However, for our purposes we will use the term "tail".

    Comments

    Stellare
    Great debunking.

    And this particular phrase made me smile -
    ...relates peacock mortality to their tails....

    I do not know exactly why. Maybe it is a 'classic example' of silliness. On my part. :-)
    Bente Lilja Bye is the author of Lilja - A bouquet of stories about the Earth
    Gerhard Adam
    Thank you and I'm glad it made you smile :)
    Stellare
    Actually it made me do more than smile. It also made me rise from my chair, grab my iphone and take pictures of - you guessed it - peacock 'tails' :-)

    I posted the image with a link to your article on my FB. Hopefully you can see the image from this link:

    So it made me smile AND move. :-)
    Bente Lilja Bye is the author of Lilja - A bouquet of stories about the Earth
    Gerhard Adam
    Well, then many more thanks and I will resist commenting on how these tails move :)
    Bonny Bonobo alias Brat
    Interesting article Gerhard. We can't make these false assumptions from quaint notions and we don't even know why the male peacock evolved such a spectacular tail and what the costs were for him.
    In fact, the truth is that much of what we use to explain such behaviors is based on our own quaint notions of how we think this works, instead of building on detailed knowledge that can truly be used as a reliable indicator of biological behavior. 
    I'm not sure how we can even build this detailed knowledge as the evolution occurred in the past so we can't ever 100% recreate and test the original environment that favoured spectacular male tail selection. It would be interesting to know how long ago the male peacock first evolved this spectacular tail and what conditions and  predators were around at the time though. 

    Is the male peacock tail still evolving and if so is it getting more or less long and/or spectacular in the general peacock population? You mention experiments that tested whether the peahen was even impressed by the spectacular male tail and found that they weren't and others that found no correlation between the health of the male peacock and how spectacular his tail is :-
    Combined with previous results, our findings indicate that the peacock's tail (1) is not the universal target of female choice, (2) shows small variance among males across populations and (3) based on current physiological knowledge, does not appear to reliably reflect the male condition."
    I can't help coming up with a couple more quaint notions as to why the male peacock evolved a large spectacular tail covered in what looks like big staring eyes. Maybe it was simply to try to browbeat the peahen into submission? Maybe also peahens are shy creatures who don't enjoy the limelight, and a male peacock's tail immediately makes her feel as though she is being stared at by a large audience, so she is more likely to mate with him just to bring on the final curtain? 

    Alternatively maybe peahens who had recently mated with peacocks with large spectacular  tails had a better chance of survival than peahens who had mated with males with less spectacular tails because when she was preparing her nest after mating, or tending to her chicks, any predator that happened along at the time was distracted by his peacock tail away from her, because the peacock also displays his tail when he's feeling aggressive towards predators? So maybe more male peacocks with splendid tails were eaten by predators allowing the nearby peahen mate and her chicks a better chance of survival? Anyway, I know that is not the point of the article but it is interesting to speculate as to why the peacock did evolve such a spectacular tail isn't it? 

    The point of your article I think, is that evolutionary psychologists should not make false claims and references to biological phenomenon, as if they are in fact settled issues that should serve as a basis for drawing additional conclusions, and I have to agree with you. It must be frustrating for evolutionary psychologists though, not to be able to make these false claims and conclusions sound like settled issues when they are trying to argue their case. 

    The Diagnostic and Statistical Manual of Mental Disorders or DSM IV is another example of an evolving source of often false claims and references which are treated by psychologists as if they are facts and settled issues, that should not be used as a basis to draw additional conclusions from but as we all know they often are. At best the DSM is also just an attempt by psychologists to build upon detailed knowledge to create a reliable indicator of behaviour that they should acknowledge can also never be 100% correct.
    Make love not war
    Gerhard Adam
    I'm not sure how we can even build this detailed knowledge as the evolution occurred in the past so we can't ever 100% recreate and test the original environment that favoured spectacular male tail selection.
    I don't think anyone knows the level of detail we'll eventually discover, but we can't make claims and then use them to develop other claims.  If we don't know, then we simply don't know.

    My point is that we can't discuss altruism as a trait (and extrapolate it into mate selection) when we haven't clearly established how, when, and why altruism occurs in biology.  The units of selection is still not a reconciled issue (i.e. selfish genes, group selection, etc.).  These are fundamental elements that need to be examined to see if actual principles can be established.  Once that occurs, then we might be able to use those to build up additional knowledge about how things work or may have evolved (not necessarily with irrefutable certainty).

    However, when even the basics are wrong or questionable, then what's the point in elaborating on even more sophisticated principles and referencing 100 papers that have no better proof beyond comparable levels of speculation.

    People like to invoke game theory, but game theory is not proof of anything.  It is simply a mathematical model that demonstrates that a particular strategy may work in a particular way, but it provides us no information about how life actually reacts.  The "selfish gene" may be a useful viewpoint in some cases, but genes aren't the complete description of the living organism.   Cooperation is one of the fundamental aspects of life on this planet and yet virtually nothing is known about it (beyond examples in select cases).   We don't even have an explanation for why the original single celled organisms reproduced in the first place.  After all, if you're in a resource rich environment, then what is the need to reproduce and create competitors?  Why not simply maximize your growth and continue on indefinitely with each organism unique?  No reproduction; no selection. 

    The point is that we make assumptions about all these things, but actually "know" very little.

    I'm hoping to put an article(s) out here that will discuss some of those problems more specifically by addressing some areas like "definitions of life", "cooperation" and "altruism". 
    Gerhard Adam
    The Diagnostic and Statistical Manual of Mental Disorders or DSM IV is another example of an evolving source of often false claims and references which are treated by psychologists as if they are facts and settled issues...
    I understand and agree, but I would argue that there is one major difference at work here.  Psychologists in the field have to treat actual patients and they don't have the luxury of gaining decades worth of research before commencing treatment.  As a result, mistakes will occur, but invariably it is a case of working with the best suppositions (even if incomplete) that can be mustered (along with the "common-sense" of the therapist).  It's not an excuse for bad science or bad medicine, but it is in the application that it becomes necessary to operate with incomplete knowledge.

    Evolutionary psychology is under no such pressure and therefore has no excuse for engaging in such speculation.
    Bonny Bonobo alias Brat
    We don't even have an explanation for why the original single celled organisms reproduced in the first place.  After all, if you're in a resource rich environment, then what is the need to reproduce and create competitors?  Why not simply maximize your growth and continue on indefinitely with each organism unique?  No reproduction; no selection.  
    I seem to remember from my schoolgirl biology that the ratio of the surface area to the protoplasm in a cell means that eventually as the cell grows it is forced to divide as osmosis alone cannot remove the toxins and by-products of metabolism that the cell produces efficiently enough and if it didn't divide and it kept growing it would die. I suppose that any single cells that stopped growing and therefore didn't need to divide would have eventually died without reproducing so they are no longer around.


    Make love not war
    Gerhard Adam
    I understand, but why not "evolve" a more efficient method to remove toxins rather than the machinery for reproduction?  I realize that there will be reasonable explanations, and it could be as simple as "those that divided" competed better than those that didn't; no specific purpose beyond an accidental trait.

    Some species have some degree of "immortality" so it isn't clear that cells that didn't divide would be subject to death.
    http://en.wikipedia.org/wiki/Biological_immortality

    After all, what difference does it make?  If you die you're still dead even if you reproduce, so what's the benefit?   I know we can argue about the benefits to populations and species, but my point is to address this before such groupings existed (i.e. the first cell that divided).
    Interesting.

    >>. . . serves as an honest signal of the quality of the peacock’s genes to potential mates. <<
    So, what would a dishonest signal look like? And, would the hen be able to tell the difference? If she could not tell the difference, what would the researchers of the 'classic example' make of the whole deal? (no need to answer these)

    We are left to wonder if the author of the quoted article is serious, or just seriously confused.

    Looking forward to you writing more like this one.

    Steve Davis
    Well done Gerhard. You made the point I was about to make, that the tail might well be signalling, but the "costly" adjective is pure conjecture. And where does it leave the "costly" signalling of all the brightly coloured parrots, who according to this reasoning should have been eliminated long ago.
    Mark-n-PRMantis
    I understand, but why not "evolve" a more efficient method to remove toxins rather than the machinery for reproduction?  I realize that there will be reasonable explanations, and it could be as simple as "those that divided" competed better than those that didn't; no specific purpose beyond an accidental trait.

    Well, if Natural Selection is the rule, or model, then "'those that divided' competed better" wouldn't only be the result of an accidental trait. One cell - one HUGE cell - cannot accommodate inevitable environmental change (probably). The 'accidents' have a shot since, especially early on perhaps, the replications didn't always go so well. 

    Also, isn't the general paradigm that self-replication preceded environmental barrier membrane and internal metabolism?

    Also, as Ms. Barratt pointed out, gravity, which probably existed back then, would have had something to say about the giant monster cell before some sort of nice, strong frame could evolve. Lobsters can be gigantic Arthropods because their bodies don't collapse when they first molt and the proteins aren't linked up yet. 

    And if you got too big, you could get eaten by a... oh - nevermind.
    Gerhard Adam
    Those are all good points and I understand completely what the issues are.  However, we don't need to have arbitrarily large cells and those attendant problems to consider the fundamental question.  There's no question that the points you raise all provide positive benefits to the cells in question, but we end up coming full circle, being that the act of division conveys no advantage to the original cell.

    My "thought experiment" in pushing this point is that it strongly suggests that "benefit" can only be accrued to a group, rather than the individual.  The original cell invariably dies, so the act of division automatically promotes "group/species" level advantage.   Similarly, natural selection cannot benefit an individual; only its offspring and the population at large.

    This may actually be more philosophical than biological, but it comes down to the fact that reproduction conveys no advantage to the parent, except within the context of that particular "group" having a greater competitive advantage over those that don't reproduce.  From here we can easily extend this idea to those that reproduce faster.  In that sense, reproduction only provides a benefit "in principle" (namely that my copies have a chance of survival).

    In short, what I'm getting at, is that reproduction requires that available resources must be divided up among a greater number of organisms.  However, if this greater number are copies of the original, then it becomes an initial act of "cooperation" to gain an economy of scale that is unavailable to a singular organism.

    In this one act, it would require that an organism have the ability to recognize its own kind (i.e. it doesn't divide and immediately fight to consume its "offspring" as food).  This creates a level of "cooperation" where the copies are all intended to work towards the same objective (i.e. survival of the group, rather than the individual - although each individual must look out for itself).  Once again, if this didn't occur, then the act of reproduction would be detrimental to the original.

    Anyway, that's part of what I'm trying to explore with that question.

    kerrjac
    Nice article. It's a neat example of how superficial assumptions and values might bias science.

    The metaphor of a 'costly tail' reminds me about our past discussions of consumer behavior.

    Often in research, I find that there are 2 overlapping ways to interpret results. One is to use research to reinforce your beliefs, eg, I found just what I was looking for, and so I should study it more. The other is to use your findings to adjust your perspective. I try to constantly remind myself to take the latter route. A few times you do find what you're looking for, but rarely, it seems, do you find it *exactly*.
    Gerhard Adam
    Thanks for the comment.  I agree it is easy to to be influenced by your own bias'.  I don't really have a problem with that, but it reinforces how important it is for opposing sides to also present evidence.  I've often found that that is the most reasonable way to get "facts" reconsidered and possibly even lead to an alternate solution/perspective.
    vongehr
    Hey - I somehow totally missed this article two months back. Interesting Mariko's study that finds little correlation between his tail and her liking.
    "Classical" means that it is the old and well known, always put forth example, so yes, the peacock tail is certainly the classical example. Without a doubt the tail is more costly than no such tail - maintenance and growing it in the first place - I am not sure why one would possibly think otherwise. That tail has no solar cells.
    A non-classical example would be the poor Chinese guys constantly flashing expensive cigarettes and phones. Costly? Of course. Do Chinese girls actually go for it? No study exists. "Honest" I guess would mean it is actually a Nokia and not just the label, ha ha.
    Steve Davis
    Sascha, you said, "Without a doubt the tail is more costly than no such tail - maintenance and growing it in the first place - I am not sure why one would possibly think otherwise."
    By that reasoning, singing in the shower is costly. Any expenditure of energy by that reasoning, is costly.
    That's obviously not the case.
    In biology, for an act to be costly it must reduce the organism's capacity to survive.
    In Australia we have a bird called the Wagtail. It has a fan-shaped tail that is quite large for the size of the bird, and it waves this tail around at all times when not in flight.
    Not only is this a huge expenditure of energy over the course of a day, but it would also attract the attention of predators; birds of prey being numerous in its habitat.
    But clearly its survival is not affected.
    Just as the peacock's tail does not affect survival.
     
    vongehr
    "Costly" in biology = decrease in survival during reproductive age
    Hmm - If this is the accepted definition in this area, I guess one has to avoid that term when for example describing the high cost, eh, pardon, energy expenditure of certain expensive (in terms of energy, not cost) survival enhancing features. Thanks for the reply.
    costly = getting eaten by tigers.