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Modelling and the fall and rise of the handicap principle

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Abstract

The story of the fall and rise of Zahavi’s handicap principle is one of a battle between models. Early attempts at formal modeling produced negative results and, unsurprisingly, scepticism about the principle. A major change came in 1990 with Grafen’s production of coherent models of a handicap mechanism of honest signalling. This paper’s first claim is that acceptance of the principle, and its dissemination into other disciplines, has been driven principally by that, and subsequent modeling, rather than by empirical results. Secondly, there is a vast literature on biological signalling but few studies that make all of the observations necessary to diagnose the handicap mechanism. My final claim is that many of the applications of “costly signalling theory” in other disciplines are conceptually confused. Misinterpretations of what is meant by “costly signalling” are common. Demonstrating that a signal is costly is insufficient and is not always necessary in order to prove that, and explain why, a signal is honest. In addition to the biological modelling of signals, there is an economic literature on the same subject. The two run in parallel in the sense that they have had little mutual interaction. Additionally, it is the biological modelling that has been picked up, and often misapplied, by other disciplines.

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Notes

  1. This is taken from a review of Zahavi and Zahavi’s (1997) The Handicap Principle. Pomiankowski & Iwasa do not actually address the first of the questions that they pose, that is, why the change? They say something on the second along lines similar to my section six, which is that hard evidence of examples is difficult to find.

  2. Arnqvist and Rowe’s Sexual Conflict (2005) does have some history in it but that appears in their chapter that discusses runaway versus indicator (handicap) models.

  3. It provides no information in the statistical sense (Skyrms 2010).

  4. It should be noted that this is the case where individuals are in a “game” in which their interests conflict. As discussed in section five, where interests coincide perfectly, honest signals can be cheap. For now it suffices to note that, when it comes to potential instances of handicap signals, we are dealing with conflicts of interest.

  5. Here and in much of what follows I refer to signallers as “males” and receivers as “females” purely for ease of exposition. Many examples of animal communication do follow this pattern but they need not.

  6. This is a typical and reasonable assumption in population genetic modelling.

  7. See Web of Science counts for Zahavi (1975) and Spence (1973).

  8. Møller (1994) demonstrates that tail size correlates with offspring viability, suggesting that it is a reliable signal of quality. Saunders (2009) points out some methodological problems with this study. Møller and de Lope use survival as a proxy for lifetime reproductive fitness. It is true that the two are not perfectly correlated but use of a proxy is very often unavoidable. He also comments that they do not compare swallows with the same original tail length as should be done to examine costs at the same point on the cost curve.

  9. Searcy and Nowicki (2005) make the same point.

  10. Although this case is striking it has been noted that, compared to its prominence in the literature, there is surprisingly little empirical support, Homo sapiens excepted, for the reciprocal altruism/cooperation models stemming from Robert Trivers’ (1971) early work (Hammerstein 2003). Joan Silk goes so far as to argue that there is slim evidence in humans of the reciprocity of tit-for-tat models (Silk 2003).

  11. Web of Science 24/8/10, 28/5/10 and 30/8/10 respectively.

  12. Web of Science 30/8/10.

  13. Gintis and Bowles are economists who buck the trend but the paper is in The Journal of Theoretical Biology.

  14. My discussion of evidence of the handicap mechanism in action can be fitted into current philosophical debates on modelling in biology and economics. In the case of costly apologies, we find a model purporting to share a common structure with the handicap model but which, in fact, makes different substantive assumptions (Kuorikoski and Lehtinen 2009) about costs and benefits. In other cases, we find an unwarranted inference from observation of the result of the model (honest signalling), to the fact that it must be this model that explains the result. Put another way, to the truth of the model’s substantive assumptions, such as that interests conflict. As Sugden (2009) argues, biologists are typically silent about precisely in what sense their model is similar to, or well-represents, the real world. This is a problem even if we have good reason to think that the substantive assumptions hold. I am arguing that there are “costly signalling” cases where it is clear that the model is not a good representation because even that initial hurdle is not cleared; at least one substantive assumption does not hold, often that there are differential costs. As Kuorikoski and Lehtinen put it, “it is the truthlikeness of the substantial assumptions that ultimately carries the epistemic weight in a model” (2009, 127).

  15. Koziel et al.’s (2010) paper on tattoos as “signals of biological quality” suffers from the same currency ambiguity as the blood donation case.

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Acknowledgments

Thanks to audiences at The British Society for the Philosophy of Science conference, Dublin 2010 and at the University of Bristol for helpful discussions and to two anonymous referees for very useful comments. Research funded by the Arts and Humanities Research Council project “Evolution, Cooperation and Rationality”. Grant AH/F017502/1.

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  1. Department of Philosophy, University of Bristol, Bristol, UK

    Jonathan Grose

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Grose, J. Modelling and the fall and rise of the handicap principle. Biol Philos 26, 677–696 (2011). https://doi.org/10.1007/s10539-011-9275-1

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