Biological Markets: the role of partner choice in cooperation and mutualism
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Biological Markets: the role of partner choice in cooperation and mutualism
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Johnson (2016). Exploring social markets, partner debt, and mimetic currency in dolphins. Animal Behavior and Cognition, 3(4), 224-242

Johnson (2016). Exploring social markets, partner debt, and mimetic currency in dolphins. Animal Behavior and Cognition, 3(4), 224-242 | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
Abstract: Dynamic models of “Biological Markets” can provide a systematic and ecologically-valid approach to studying communication and social cognition in dolphins. These market models view interacting animals as traders engaged in a negotiation for social commodities, whose values vary with the state of their current market. Across the phyla, factors like the supply and demand of social resources can impact on investment and partner choice. Such models map well to the polyadic nature of typical social interactions, generating predictions based on configurations of participants, and enabling us to use behavioral observations to address issues of cognitive organization. Plus, by positing communicative signals as social currency, these models provide tools to discern which aspects of dolphins’ vocal and gestural repertoires impact on their social relationships. Adapting these models for dolphins highlights the premise that “partnerhood is good,” wherein both players gain when they partner. When considered over time, the gains and losses of valued partners may accumulate into “wealth” or “debt” for a particular player, altering its threshold for responding to its market’s odds. For example, “Partner Debt” could motivate a player to more readily take action to destabilize its current market. In dolphins, one type of social currency that bears further investigation is the use of vocal and/or gestural mimicry. Such mimesis may promote prosociality, cooperation, and even the coordination of third party information.

Keywords: Biological Markets, Dolphins, Social cognition, Communication, Imitation, Mimesis
Ronald Noë's insight:
This is a contribution to a workshop “Methods for Studying Communication and Social Cognition in Cetaceans”, organised by the author (Christine Johnson) and Denise Herzig during the 21st Biennial Conference of the Marine Mammal Society (San Francisco December 2015).

After an introduction to BMT with the help of some examples, mainly intra-specific markets in animals, the author delineates a detailed research program in dolphins based on market theory. Emphasis is placed on the value of partnerships as such in these animals that depend on cooperation and coordination in many aspects of life. Going from  “Who does what to whom?” to “Who does what to whom, while who else is present?” (p. 228) is perhaps the core element in the proposal to introduce more market in dolphin research.
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Batstone, Dutton, Wang, Yang & Frederickson (2016). The evolution of symbiont preference traits in the model legume Medicago truncatula. New Phytologist

Batstone, Dutton, Wang, Yang & Frederickson (2016). The evolution of symbiont preference traits in the model legume Medicago truncatula. New Phytologist | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
Abstract:
Many hosts preferentially associate with or reward better symbionts, but how these symbiont preference traits evolve is an open question. Legumes often form more nodules with or provide more resources to rhizobia that fix more nitrogen (N), but they also acquire N from soil via root foraging. It is unclear whether root responses to abiotically and symbiotically derived N evolve independently.

Here, we measured root foraging and both preferential allocation of root resources to and preferential association with an effective vs an ineffective N-fixing Ensifer meliloti strain in 35 inbred lines of the model legume Medicago truncatula.

We found that M. truncatula is an efficient root forager and forms more nodules with the effective rhizobium; root biomass increases with the number of effective, but not ineffective, nodules, indicating preferential allocation to roots harbouring effective rhizobia; root foraging is not genetically correlated with either preferential allocation or association; and selection favours plant genotypes that form more effective nodules. 

Root foraging and symbiont preference traits appear to be genetically uncoupled in M. truncatula. Rather than evolving to exclude ineffective partners, our results suggest that preference traits probably evolve to take better advantage of effective symbionts.

Keywords: cheating Ensifer meliloti genetic correlations Medicago truncatula mutualism stability partner choice root foraging sanctions
Ronald Noë's insight:
Two strains of rhizobia that differ in their efficiency of providing the plant with nitrogen (N), different regimes of N-fertilization, whole-root and split-root experiments and further typical experimental methods used in the analysis of the symbiosis between rhizobia and legumes were put to good use in this study. The authors aimed at answering a rather interesting question: do legumes select their microbial partners by excluding the bad ones (a process often called 'sanctioning'), or do they rather allocate more resources to the good ones? Their results firmly support the latter hypothesis. The paper further discusses some interesting chicken-or-egg-type problems in the evolution of partner choice mechanisms.
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Raihani & Barclay (2016). Exploring the trade-off between quality and fairness in human partner choice. Royal Society Open Science, 3(11)

Raihani & Barclay (2016). Exploring the trade-off between quality and fairness in human partner choice. Royal Society Open Science, 3(11) | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it

Partner choice is an important force underpinning cooperation in humans and other animals. Nevertheless, the mechanisms individuals use to evaluate and discriminate among partners who vary across different dimensions are poorly understood. Generally, individuals are expected to prefer partners who are both able and willing to invest in cooperation but how do individuals prioritize the ability over willingness to invest when these characteristics are opposed to one another? We used a modified Dictator Game to tackle this question. Choosers evaluated partners varying in quality (proxied by wealth) and fairness, in conditions when wealth was relatively stable or liable to change. When both partners were equally fair (or unfair), choosers typically preferred the richer partner. Nevertheless, when asked to choose between a rich-stingy and a poor-fair partner, choosers prioritized fairness over wealth—with this preference being particularly pronounced when wealth was unstable. The implications of these findings for real-world partner choice are discussed.


Keywords: partner choice biological market theory quality fairness

Ronald Noë's insight:
Experiments were conducted with subjects recruited via Amazon's Mechanical Turk in order to see whether people playing a modified Dictator game rather deal with 'poor but fair' dictators than with 'rich but stingy' ones. What is in fact tested is the relative importance of the willingness of a partner to offer a good deal (i.e. the partner has 'good intentions') as opposed to his ability to offer a good deal (i.e. the partner actually has the necessary resources).

The authors find that fairness has higher priority in human partner choice than the ability to offer high payoffs. This conclusion fits well with other studies, even though in this one the subjects played in the noisy world of MTurk (according to the authors own judgement) and for rather low payoffs.
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Loverdo & Viciana (2016). Cultural transmission and biological markets. bioRXiv preprint

Loverdo & Viciana (2016). Cultural transmission and biological markets. bioRXiv preprint | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
Abstract: Active cultural transmission of fitness-enhancing behavior can be seen as a costly strategy, one whose evolutionary stability poses a Darwinian puzzle. In this article, we offer a biological market model of cultural transmission that substitutes or complements existing kin-selection based theories for the evolution of cultural capacities. We explicitly formulate how a biological market can account for the evolution of deference and prestige-related phenomena, as well as how it can affect the dynamics of cumulative culture. We show that, under certain conditions, teaching evolves even when innovations are not sufficiently opaque and can be acquired by emulators via inadvertent transmission. Furthermore, teaching in a biological market is a precondition for enhanced individual learning abilities.

Keywords: Social learning, comparative advantage, teaching, deference, cumulative culture, partner choice
Ronald Noë's insight:
The authors assume, in many cases rightly so I think, that the teaching of a skill is a commodity that can be exchanged against another commodity. The authors assume the latter commodity to consist of "deference" and "prestige enhancement". I could also imagine payment for learning opportunities that a bit more straightforward, but that doesn't really change the basic argument: teachers and learners can be seen as two classes of traders in a biological market. The authors offer a plausible argument that this simple assumption improves our understanding of the evolution of teaching and cultural accumulation of skills considerably.

A quote that illustrates the spirit of the paper: "In this article, we analyze conditions of the evolution of cultural transmission capacities in a biological market model. Originally proposed by behavioral ecologists Ronald Noë and Peter Hammerstein, biological markets arise when associations between biological individuals are sufficiently uncoerced that competition occurs not so much by force or its threat, as due to a need to offer more of what the choosing party “demands”. The idea of biological markets thus sheds light on certain selective mechanisms, namely market effects in which “members of one class can “force” members of another class to evolve traits that would have a negative effect on fitness in the absence of 77 the cooperative interaction” (Noë and Hammerstein, 1994, p. 2)."

The authors forgot to actually list our paper in the references, but I guess the readers of these pages know where to find it.The paper can use a few more improvements here and there, but this is only preliminary version apparently.
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Heintz, Karabegovic & Molnar (2016). The co-evolution of honesty and strategic vigilance. Frontiers in Psychology, 7(1503)

Heintz, Karabegovic & Molnar (2016). The co-evolution of honesty and strategic vigilance. Frontiers in Psychology, 7(1503) | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it

Abstract: We hypothesize that when honesty is not motivated by selfish goals, it reveals social preferences that have evolved for convincing strategically vigilant partners that one is a person worth cooperating with. In particular, we explain how the patterns of dishonest behavior observed in recent experiments can be motivated by preferences for social and self-esteem. These preferences have evolved because they are adaptive in an environment where it is advantageous to be selected as a partner by others and where these others are strategically vigilant: they efficiently evaluate the expected benefit of cooperating with specific partners and attend to their intentions. We specify the adaptive value of strategic vigilance and preferences for social and self esteem. We argue that evolved preferences for social and self-esteem are satisfied by applying mechanisms of strategic vigilance to one’s own behavior. We further argue that such cognitive processes obviate the need for the evolution of preferences for fairness and social norm compliance.

Ronald Noë's insight:
This is a carefully written and thought-provoking review/opinion piece about the role the choice of cooperative partners might have played in the evolution of human morality, honesty and prosociality.

The authors' line of thought is roughly as follows:
1) People chose among potential partners for cooperation in a way that is ultimately beneficial to themselves.
2) The fact that people choose partners induces competition among individuals trying to outbid others in making the impression to be a good (or the best) partner
3) People do everything to obtain a good reputation as a cooperation partner
4) The possibility that people fake their partner quality selects for (cognitive) mechanisms to tell fake apart from the real thing. This makes people sensitive, for example, for the intentions of a cooperating individual rather than the outcome of the cooperation
5) The best way of monitoring the impression one makes on others is by monitoring the impression one has of self. Hence, people like seeing themselves as god and trustworthy partners.

Heintz et al clearly build on the work of other authors, notably Barclay and Baumard, that connected "partner choice theory" with the evolution of prosociality etc. They don't necessarily agree with these authors, however, and notably contrast their evolution of 'strategic vigilance' with explanations Baumard and colleagues offer for the evolution of a sense of fairness.

Especially interesting is the distinction they make in the Conclusions-section between the explanatory power of their brand of partner choice theory and of cultural group theory.
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Dunayer & Berman (2016). Biological markets: theory, interpretation, and proximate perspectives. A response to Sánchez-Amaro and Amici (2015). Animal Behaviour, 121, 131-136

Dunayer & Berman (2016). Biological markets: theory, interpretation, and proximate perspectives. A response to Sánchez-Amaro and Amici (2015). Animal Behaviour, 121, 131-136 | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
In their review of the applicability of biological markets theory (BMT) to primate exchanges, Sanchez-Amaro and Amici (2015) make numerous valid and important points about theoretical and methodological problems that have so far limited the application of BMT to primates. While we agree with many points of their excellent review, there are a few areas that we feel would benefit from further discussion.

Keywords: Biological Markets Theory cooperation time frame of exchange primates stress supply and demand
Ronald Noë's insight:
This is a second answer to Sanchez-Amaro's and Amici's (2015) question" Are primates out of the market?" after Kaburu & Newton-Fisher's (2016) first reaction.

Here Erica Dunayer and Carol Berman discuss the pros and cons of both BMT and the paper they target in a very thorough and thought-provoking way. I especially liked their insights in the notorious "time-frame discussion" that has haunted the primate cooperation literature for much too long now.

I was also very happy with their balanced approach to the question whether or not 'stress' is just a proximate mechanism driving adjustments to changes in supply and demand or an alternative explanation for the observed phenomena by itself.

In hindsight I am glad I decided not to react to Sanchez-Amaro & Amici myself. I couldn't have done a better job than these authors and Kaburu & Newton-Fisher together - and I would have been seen as highly biased anyway.

Just for the record: I was not the anonymous reviewer of this paper and saw it for the first time today. Makes my day!


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Norscia & Palagi (authors/editors) 2016 The Missing Lemur Link Cambridge UP

Norscia & Palagi (authors/editors) 2016 The Missing Lemur Link Cambridge UP | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
A comparative study of lemurs in the context of shared ancestral links with both humans and primates.
Ronald Noë's insight:
All I can say about this book is that, judging from the index, it has quite a few pages with something on markets. Probably not surprising in the light of earlier work by the authors: Norscia, I., Antonacci, D., & Palagi, E. (2009). Mating first, mating more: biological market fluctuation in a wild prosimian. PLoS ONE, 4(3), e4679
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Sánchez-Amaro & Amici (2016). Markets carefully interpreted: a reply to Kaburu and Newton-Fisher (2016). Animal Behaviour, 119, e7-e13

Sánchez-Amaro & Amici (2016). Markets carefully interpreted: a reply to Kaburu and Newton-Fisher (2016). Animal Behaviour, 119, e7-e13 | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
In our recently published essay (Sánchez-Amaro & Amici, 2015) we aimed to discuss the literature on primate biological markets (BMs) by drawing attention to the problems that, in our view, affect most primate studies endorsing the existence of BMs. In this way, we aimed not only to warn of conclusions based on dubious methodological approaches, but also to provide some possible new avenues to more efficiently test biological market theory (BMT) in primates. Finally, we hoped to stimulate debate with experts in BMT, including primatologists working in the field, to critically discuss the points we raised in our essay and find new ways to collaboratively improve empirical work on primate BMs.
Ronald Noë's insight:
... and the answer (see earlier scoop of this paper for my comment)
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Macfarlan (2016). Social evolution: The force of the market. Current Biology, 26(16), R756-R758

Macfarlan (2016). Social evolution: The force of the market. Current Biology, 26(16), R756-R758 | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
Abstract: Biological market forces shape patterns of cooperation typical of small-scale human societies that are organized by division of labor based on age and gender. Labor specialization promotes trade, while supply and demand affect the amount individuals exchange for commodities.
Ronald Noë's insight:
This is a 'Dispatch' about the paper by Jaeggi et al (2016 Current Biology, 26(16), 2180-2187 - mentioned a few scoops further down) in which Shane Macfarlan explains very well how the exchanges of goods and services in horticulturalists such as the Tsimane from Bolivia can be seen as a transition between 'biological' markets without binding contracts, common currencies etc. and 'modern' human markets with all kinds of institutions that stabilise the market.
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Jaeggi, Hooper, Beheim, Bret, Kaplan & Gurven (2016). Reciprocal exchange patterned by market forces helps explain cooperation in a small-scale society. Current Biology

Jaeggi, Hooper, Beheim, Bret, Kaplan & Gurven (2016). Reciprocal exchange patterned by market forces helps explain cooperation in a small-scale society. Current Biology | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
Abstract: Social organisms sometimes depend on help from reciprocating partners to solve adaptive problems [1], and individual cooperation strategies should aim to offer high supply commodities at low cost to the donor in exchange for high-demand commodities with large return benefits [2, 3]. Although such market dynamics have been documented in some animals [4–7], naturalistic studies of human cooperation are often limited by focusing on single commodities [8]. We analyzed cooperation in five domains (meat sharing, produce sharing, field labor, childcare, and sick care) among 2,161 household dyads of Tsimane’ horticulturalists, using Bayesian multilevel models and information-theoretic model comparison. Across domains, the best-fit models included kinship and residential proximity, exchanges in kind and across domains, measures of supply and demand and their interactions with exchange, and household-specific exchange slopes. In these best models, giving, receiving, and reciprocating were to some extent shaped by market forces, and reciprocal exchange across domains had a strong partial effect on cooperation independent of more exogenous factors like kinship and proximity. Our results support the view that reciprocal exchange can provide a reliable solution to adaptive problems [8–11]. Although individual strategies patterned by market forces may generate gains from trade in any species [3], humans’ slow life history and skill-intensive foraging niche favor specialization and create interdependence [12, 13], thus stabilizing cooperation and fostering divisions of labor even in informal economies [14, 15].

Keywords: reciprocity kin selection biological market theory trade division of labor Tsimane
Ronald Noë's insight:
When I started thinking about biological markets (> 25 years ago), I assumed that anthropologists, sociologists and economists would have studied, described and analysed all kinds of human markets from the very basic bargaining in pre-agricultural, pre-industrial populations up to modern global stock markets. I guess my biologist brain expected others to see markets as the result of a continuous evolutionary process and describe the transitions from simple and old forms to complex and recent ones as a continuum. This turned out to be a mistake: anthropologists were more likely to look at cooperative exchanges among the people they studied in terms of kinship, reciprocal altruism and geographical distances than as exchanges in a market context. 

This paper fills the gap very nicely with an impressive Bayesian multilevel analysis of the exchange pattern of 5 commodities among the Tsimane horticulturalists of Bolivia. It turns out that adding supply and demand effects to the mix considerably increases the explanatory power of the models.

Two little moans nevertheless:
It seems odd that biological market theory is mentioned many times throughout the paper without ever referring to its origins.

Implicitly, as is often done, 'reciprocity' (i.e. reciprocal altruism) is seen as THE alternative to kinship as an explanation for cooperation and thus any model describing non-kin based cooperation is automatically pigeon-holed as 'an extension of reciprocity'. This ignores a dichotomy that I consider at least as fundamental: the one between partner control and partner choice models.
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Noë, R. (2016) How do biological markets compare to the markets of economics? - Munich Personal RePEc Archive

Noë, R. (2016) How do biological markets compare to the markets of economics? - Munich Personal RePEc Archive | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
Abstract: After an introduction to biological markets written for non-biologists, I explore whether and to what extent natural markets, i.e. markets on which non-human traders exchange goods and services with members belonging to their own or to other species, can be compared to human ‘economic’ markets, i.e. the markets analysed by economists. Biological Market Theory (BMT) borrows jargon and ideas from economics, but was at least as much inspired by sexual selection theory, a collection of models of ‘mating markets’, including human mating markets. Here I ask two main questions: (1) Is there more than only a superficial resemblance between both types of markets? (2) Can the analysis of one yield insights about the other? First, I consider the different forms of human trading and markets and propose some biological ones to which these can best be compared, e.g. companies trading goods in markets shaped by ‘comparative advantage’ to underground nutrient exchange markets between plants and rhizobial bacteria and mycorrhizal fungi; job and retail markets with pollination, seed dispersal and protection markets between plants and insects; ‘embedded markets’ with grooming markets in non-human primates and so forth. Then I look at some phenomena that are considered to be exclusive to human markets, such as common currencies and binding contracts, and ask whether these are indeed that exclusive. Finally I look at the common ground: negotiations that take place on several types of markets, natural or not; the honesty of advertisements, which is recognised as a major problem for both human and non-human clients; the biological equivalent of the market – firm dichotomy and the importance of the costs of partner choice, which are known to economists as ‘transaction costs’ and to sexual selection theoreticians as ‘search costs’. I conclude that there are several good reasons to have a closer look at those properties that set human and biological markets apart, but certainly also at those features that make them comparable to each other.

Keywords: cooperation, mutualism, trade, biological markets, mating markets, embedded markets, sexual selection, partner choice, currency, binding contract, negotiation, bargaining, honest advertisement, transaction costs, search costs, market – firm dichotomy
Ronald Noë's insight:
Pushing my own stuff this time. This is a revised and updated version of a working paper I wrote for the workshop series ‘The Market and Marketization’, organized by Uskali Mäki and Pekka Mäkelä, Academy of Finland Centre of Excellence in the Philosophy of the Social Sciences & Department of Political and Economic Studies, University of Helsinki, Finland.

This very inspiring workshop with philosophers of economics, economists, sociologists, lawyers and so forth made me think a bit more about the similarities and differences between biological markets and the markets we took as inspiration: those with human traders studied by economists, but also anthropologists, sociologists etc.

The paper is written in the form of a book chapter, but since it may take a while till it sees the light in that form, I went the unorthodox way, for a biologist at least, to deposit it on a web-based archive that is used (a lot) by economists.
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Fehrler & Przepiorka (2016). Choosing a partner for social exchange: Charitable giving as a signal of trustworthiness. Journal of Economic Behavior & Organization

Fehrler & Przepiorka (2016). Choosing a partner for social exchange: Charitable giving as a signal of trustworthiness. Journal of Economic Behavior & Organization | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
Abstract: People benefit from being perceived as trustworthy. Examples include sellers trying to attract buyers, or candidates in elections trying to attract voters. In a laboratory experiment using exchange games, in which the trustor can choose the trustee, we study whether trustees can signal their trustworthiness by giving to charity. Our results show that donors are indeed perceived as more trustworthy and they are selected significantly more often as interaction partners. As a consequence of this sorting pattern, relative payoffs to donors and non-donors differ substantially with and without partner choice. However, we do not find donors to be significantly more trustworthy than non-donors. Our findings suggest that publicly observable generosity, such as investments in corporate social responsibility or donations to charity during a political campaign, can induce perceptions of trustworthiness and trust.
Ronald Noë's insight:
Not really about markets in the sense of measuring the effect of changes in supply & demand ratios by experimentally changing the number of potential partners, but an interesting experiment about the effect of charitable donations on the trustworthiness and reputation of donors to charitable causes when subjects have an option to choose their partners among donors and non-donors.
  
The authors appropriately stress the lack of attention in experimental economics for partner choice:  "Despite the important role partner choice plays in many settings, studies of social preferences have so far mainly focused on situations with random matching." This is notably odd in the context of economics, since partner choice in its many disguises (consumer preferences, employment etc.) to my mind is and remains the main motor of markets.

A result that I found especially interesting is that a minority of subjects did NOT choose to give to a donor when given the choice between a donor and a non-donor. The authors explain this as follows: "A possible explanation for this might be that these subjects feel morally obliged to give more to certain interaction partners and so avoid these partners when given the choice." Well .... if someone would show that for non-human animals then I'll eat all the hats I have!
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Ravindra Krishnamurty (June 3, 2016) A Peep Into the Underground Biological Market - The Permaculture Research Institute

Ravindra Krishnamurty (June 3, 2016) A Peep Into the Underground Biological Market - The Permaculture Research Institute | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
Symbiotic relationship between plants and mychorrhizal fungi has been in existence for over 400 million years. Fungi cannot survive on their own as they are incapable of producing their own food. Hence, they latch on to the plants roots to get essential food. Plants share a part of the carbohydrates produced during photosynthesis and in return, fungi supply phosphates and other minerals, by mining them from the soil. These natural …
Ronald Noë's insight:
For what it is worth .... (to be honest I have no idea what this site stands for, but perhaps this is of some use to someone).
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Wannagat (2016). Study of the evolution of symbiosis at the metabolic level using models from game theory and economics. PhD, Université de Lyon

Wannagat (2016). Study of the evolution of symbiosis at the metabolic level using models from game theory and economics. PhD, Université de Lyon | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
Summary: Symbiosis, a term that brings all types of species interaction under one banner, is defined as a close association of different species living together. Species interactions that comprise microorganisms are of particular interest for agriculture, health, and environmental issues. All kinds of species interactions such as mutualism, commensalism, and competition, are omnipresent in nature and occur often at the metabolic level. Organisms release metabolites to the environment which are then taken up by other individuals of the same or of different species. In this thesis, we study how species interactions shape the environment. We examine the questions of (i) what are the minimal nutrient requirements to sustain growth, and (ii) which metabolites can be exchanged between an organism and its environment. Enumerating all minimal stoichiometric precursor sets, and all minimal sets of exchanged metabolites, using metabolic network models, provide a better insight into species interactions. In a spatially homogeneous environment, the metabolites that are released to such an environment are shared by all individuals. The problem that then arises is how cheaters, individuals that profit from the released metabolites without contributing to the public good, can be prevented from the population. This and other configurations were already modeled with approaches from game theory and economics. We examine how the concepts of minimal stoichiometric precursor sets and minimal sets of exchanged compounds can be introduced into such models.

Keywords: symbyosis; metabolism; metabolic network modeling; enumeration; minimal stoichiometric precursor sets; minimal stoichiometric factories; minimal sets of exchanged compounds; game theory; economics
Ronald Noë's insight:
This French PhD-thesis (in English) is not the kind of document I would read cover to cover. Specialists, notably theoreticians, will probably find some of interest here. There are good descriptions of 'comparative advantage' and Walrasian theory in connection to symbiosis. I wouldn't mind a better connection to the biological market literature (and more careful citation :) )
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Kafashan, Sparks, Rotella & Barclay (2016). Why heroism exists. Evolutionary perspectives on extreme helping. In S. T. Allison, G. R. Goethals & R. M. Kramer (Eds.), Handbook of heroism and heroic...

Kafashan, Sparks, Rotella & Barclay (2016). Why heroism exists. Evolutionary perspectives on extreme helping.  In S. T. Allison, G. R. Goethals & R. M. Kramer (Eds.), Handbook of heroism and heroic... | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
First 2 par. On June 17, 2014, thirteen-year-old Robert Pritchard Junior rushed into a burning mobile home to rescue a six-year-old girl from being engulfed by the flames. Almost a year earlier, Christopher Ihle saved an eighty-four-year-old male and his seventy-eight-year-old wife from being struck by a train. A year prior to this incident,Kyle Hardman attempted to save three men and two children from drowning in the Mississippi River. Pritchard Junior, Ihle, and Hardman are recipients of the prestigious Carnegie Medal. These heroes were awarded their medals for voluntarily and knowingly risking their lives to attempt to save others. Why would anyone do such a thing?Why would one incur a cost—such as risk their life—to benefit others?Why would anyone be a hero? From an evolutionary perspective, incurring any such costs appears puzzling at first glance. Natural selection favors traits that increase the propagation of one’s genetic material into future generations, and it ruthlessly eliminates any costly traits that provide no net reproductive advantage. This logic prompts the following crucial questions: do heroes receive any benefits that counteract the costs of their actions (thus providing a net selective advantage for heroic traits), and if so, what is the nature of those benefits?
Ronald Noë's insight:
For all those interested in the type of (human) biological markets driven by what Gilbert Roberts called "competitive altruism' : the idea that one shows off - one way or the other - to improve one's status as a partner.
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Hellmann (2016). Neighbor effects: The influence of colony-level social structure on within-group dynamics in a social fish. PhD-thesis. Ohio State University.

Hellmann (2016). Neighbor effects: The influence of colony-level social structure on within-group dynamics in a social fish. PhD-thesis. Ohio State University. | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
Abstract (1st section): In nearly all species, social interactions are highly influential in determining an individual’s success within its environment. In humans, positive social interactions are correlated with increased job success, better health, and reduced stress; in other species, social interactions are critical determinants of disease spread, predation avoidance, and learning. Behavioral ecologists have long been interested in strategic interactions among group members; however, most studies of these do not consider how interactions within a group may change due to the presence of or interactions with other groups. This is problematic, as it is difficult to understand why individuals would cooperate or resolve conflict within their group without understanding the extent to which neighboring groups provide opportunities for individuals to leave their current group. Similarly, it is difficult to understand the extent to which individuals pursue reproductive opportunities within the group without understanding how neighboring groups offer additional opportunities for current and future reproduction. Theoretical models have long suggested that neighboring groups strongly influence individual behavior and reproductive success, but there have been few empirical experiments testing these theoretical predictions. My dissertation research explores how neighboring groups alter within-group social and reproductive dynamics in Neolamprologus pulcher, a cooperatively breeding fish. These fish live in colonies of 2-200 groups, each with a dominant breeding pair and 1-20 subordinates that help maintain the territory and care for offspring.
Ronald Noë's insight:
Quite a few references to biological markets in this thesis apparently, but I didn't manage to read all 202 pages (yet).

This is a cooperatively breeding species and the question is whether dominant breeders have leverage over their helpers by playing them off against each other and by threat of eviction from the territory and/or aggression.
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Who pays first? Nutrient exchange dynamics in the arbuscular mycorrhizal symbiosis BBSRC Project 12 - Helgason / Field - Postgraduate study, The University of York

Who pays first? Nutrient exchange dynamics in the arbuscular mycorrhizal symbiosis BBSRC Project 12 - Helgason / Field - Postgraduate study, The University of York | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
Project description: Arbuscular mycorrhizal fungi (AMF) are soil-dwelling fungi that form mutualistic symbioses with the vast majority of extant land plants. These symbiotic associations are a key link between plants and soils, as they occupy both niches simultaneously. As obligate biotrophs, AMF are unable to obtain organic carbon other than as photosynthates derived from their host plant through the bidirectional exchange for fungal-acquired soil nutrients.

There is a heated debate about the nature of the cross-talk between plant hosts and their fungi, specifically regarding how carbon-for-nutrient exchange is regulated between symbiotic partners.

The ‘biological market’ hypothesis suggests that symbionts engaged in mutualisms exchange goods in the same way as commodities are traded in human economic markets. Under this framework, AMF partner selection and evolutionary stability is underpinned by the proportional bidirectional exchange of photosynthates for nutrients, where the most ‘cooperative’ partners are ‘rewarded’ with resources, while so-called ‘uncooperative partners’ face nutritional ‘sanctions’.

For such a rewards/sanctions-based mechanism to exist, each symbiont must detect variation in benefit received, and be in a position to withhold nutrients until maximum reward for their investment is achieved. However, this model does not account for the persistence of AMF across a wide variety of plant life-histories, in particular in plants exhibiting heterotrophy where ‘cheating’ is prevalent. Evidence from lab and field experiments suggests bidirectional exchange of carbon-for-nutrients between plants and fungi is not always proportional; that the symbiotic nutrient exchange is an emergent property of a source-sink driven relationship. Critically, while much data has been published that purports to support one or other of these hypotheses, all of them could be interpreted either way because they lack information on timing.

This PhD project will investigate these competing hypotheses by studying the timing of carbon-for-nutrient exchange between AMF and their plant hosts using a powerful combination of molecular methods such as gene expression (York) and physiological measurements carbon for nutrient exchange using tracers and metabolomics analyses (Leeds). Working within two groups with complementary expertise, you will have the opportunity to shed light on this important symbiosis, while developing research and data skills that are important in a wide range of areas.
Ronald Noë's insight:
Not something I normally put on this site, but I thought this PhD project description looks interesting in itself. I look forward to see these conflicting ideas being tested against each other.

And who knows ... perhaps a good candidate stumbles upon this page and applies.
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Wild & Cojocaru (2016). Runaway competition: A correction and extension of results for a model of competitive helping. PLoS ONE, 11(10), e0164188

Wild & Cojocaru (2016). Runaway competition: A correction and extension of results for a model of competitive helping. PLoS ONE, 11(10), e0164188 | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it

We investigate and generalize an existing model of competitive helping within a biological market, first introduced for a population of competing individuals in which one individual provides help to all others; the rest compete for the help available from this individual by providing help themselves. Our generalized model comprises two strategies in which each individual of a specific type provides the same amount of help as all other individuals of that type. Each individual’s fitness function is dependent on this level of help, the cost of providing the help, and the fact that help is proportionally reciprocated by other individuals. Competitive helping occurs when individuals actively try to help more than other individuals. To assess the emergence of equilibrium help strategies as adopted by proportions of the population, we examine the competition over available help within two settings: replicator dynamics and agent-based numerical simulations. To move one step further in our generalization, we use the agent-based model to study the N-person competitive helping game, where all individuals in the population are heterogeneous with respect to help provided. Our results show that helping does not increase indefinitely with the population size, as concluded previously, and while there are some instances of an increase in help provided as a result of competition, this competition can be detrimental to all individuals and in most cases, one type simply gives up (thus evolving to a “no help” strategy). The degree to which an individual’s help is reciprocated by the others in the population has strong implications in the long-term behaviour of equilibrium help levels of types of individuals (and of individuals themselves); these equilibrium help levels diverge from existing conjectures in current literature. Lastly, small amounts of passively provided (costless) help results in runaway competition among all individuals.
Ronald Noë's insight:
I am not going to pretend having followed the math in this paper and having understood the details, but this is an extension (and partial contradiction) of Barclay, P. (2011). Competitive helping increases with the size of biological markets and invades defection. Journal of Theoretical Biology, 281(1), 47-55
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Grillo, Stinchcombe & Heath (2016). Nitrogen addition does not influence pre-infection partner choice in the legume–rhizobium symbiosis. American Journal of Botany

Grillo, Stinchcombe & Heath (2016). Nitrogen addition does not influence pre-infection partner choice in the legume–rhizobium symbiosis. American Journal of Botany | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
PREMISE OF THE STUDY: Resource mutualisms such as the symbiosis between legumes and nitrogen-fixing rhizobia are context dependent and are sensitive to various aspects of the environment, including nitrogen (N) addition. Mutualist hosts such as legumes are also thought to use mechanisms such as partner choice to discriminate among potential symbionts that vary in partner quality (fitness benefits conferred to hosts) and thus impose selection on rhizobium populations. Together, context dependency and partner choice might help explain why the legume–rhizobium mutualism responds evolutionarily to N addition, since plant-mediated selection that shifts in response to N might be expected to favor different rhizobium strains in different N environments.

METHODS: We test for the influence of context dependency on partner choice in the model legume, Medicago truncatula, using a factorial experiments with three plant families across three N levels with a mixed inoculation of three rhizobia strains.

KEY RESULTS: Neither the relative frequencies of rhizobium strains occupying host nodules, nor the size of those nodules, differed in response to N level. 

CONCLUSIONS: Despite the lack of context dependence, plant genotypes respond very differently to mixed populations of rhizobia, suggesting that these traits are genetically variable and thus could evolve in response to longer-term increases in N.
Ronald Noë's insight:
(Note added 31 Oct 2016)
I have now read this paper, but I can't say I can really follow the reasoning. This is not doubt for a large part due to my lack of knowledge of legume - rhizobia mutualisms, but there also could be some more fundamental misunderstandings. The story hinges on the difference between 'sanctioning' (the plant allocating less resources to the nodule formed by a certain bacterial strain) and 'partner choice' (the plant preferentially forming nodules with certain strains on the basis of information obtained through 'signalling'.

I have two puzzles:
(1) Isn't sanctioning not just another form of partner choice? If I buy my bread from 2 bakeries and at some point don't like the bread of one of them and stop buying there - didn't I exert partner choice? The difference some may see is that the legume invested resources first in one case and only communicated before making a choice in the other. However, communicating is not free, neither sending a signal nor reading one is, so the difference is at best a matter of the height of the initial investment.
(2) Why is partner choice not considered both ways? Can the rhizobia for some reason not choose between different legume individuals? Or, over evolutionary time, between different legume genotypes?

I was a bit ill at ease when I realised that the data of an experiment done in 2007 were used to test ideas that were partially formulated later. This gives the impression that some post hoc interpretation was going on.
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Jandér & Herre (2016). Host sanctions in Panamanian Ficus are likely based on selective resource allocation. American Journal of Botany

Jandér & Herre (2016). Host sanctions in Panamanian Ficus are likely based on selective resource allocation. American Journal of Botany | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
PREMISE OF THE STUDY: Fig trees and their pollinators, fig wasps, present a powerful model system for studying mutualism stability: both partners depend on each other for reproduction, cooperation levels can be manipulated, and the resulting field-based fitness quantified. Previous work has shown that fig trees can severely reduce the fitness of wasps that do not pollinate by aborting unpollinated figs or reducing the number and size of wasp offspring. Here we evaluated four hypotheses regarding the mechanism of sanctions in four Panamanian fig species. METHODS: We examined wasp and fig samples from field experiments with manipulated levels of pollination. KEY RESULTS: In unpollinated figs, the fig wall and the wasp offspring had a lower dry mass. Unpollinated figs had as many initiated wasp galls as pollinated figs but fewer galls that successfully produced live wasp offspring. Across three experimentally increasing levels of pollination, we found nonlinear increases in fig wall mass, the proportion of wasp galls that develop, and wasp mass. CONCLUSIONS: Our data did not support the hypotheses that lack of pollination prevents gall formation or that fertilized endosperm is required for wasp development. While our data are potentially consistent with the hypothesis that trees produce a wasp-specific toxin in response to lack of pollination, we found the hypothesis that sanctions are a consequence of trees allocating more resources to better-pollinated figs more parsimonious with the aggregate data. Our findings are completely analogous to the selective resource allocation to more beneficial tissues documented in other mutualistic systems.

Keywords: coevolution cooperation Ficus fig wasp Moraceae mutualism partner choice pollination resource allocation sanctions species interaction
Ronald Noë's insight:
When I see that a Discussion of a paper starts with a sentence such as “The data presented here in conjunction with already published studies suggest that the most likely mechanism underlying host sanctions in figs is relatively higher resource allocation to better pollinated figs.”, then I am inclined to expect a nice paper about partner choice at least, and perhaps biological markets. Unfortunately the authors only use the first term in their keywords and the latter not at all. They opted for the rather unfortunate term ‘sanctions’, even though they report a number of very nice experiments and observations in which they show rather neatly that fig trees allocate resources to those figs that contain the best pollinators among the wasps that lay eggs in their ovules.

There is nothing wrong with the term sanctions as such, but I would propose to use it only in the way suggested by its use in a human context: negative actions against bad mutualistic partners. Personally, I only use it in the case of big guys killing their small partners, e.g. plants with obligate nursery pollination (such as fig trees) selectively eliminating bad pollinators by abortion of fruits.

Apart from an overview of previously published and new data from these well-designed experiments, the authors also make another point that I find important: the mechanisms that are used by mutualists to implement partner choice don’t have to be the result of evolution from scratch or even a modification of existing processes; they can serve the actor's own needs for 100% and have partner choice as a side-effect. In this case the fig trees can invest in their well-developing seeds and, as a result, fatten up the little wasp larvae in the same fig too.
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Kaburu & Newton-Fisher (2016). Markets misinterpreted? A comment on Sánchez-Amaro and Amici (2015). Animal Behaviour, 119, e1-e5

Kaburu & Newton-Fisher (2016). Markets misinterpreted? A comment on Sánchez-Amaro and Amici (2015). Animal Behaviour, 119, e1-e5 | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
In a recent essay, Sanchez-Amaro and Amici (2015) reviewed evidence in support of biological market theory (BMT) in primates. Since the pioneering work by Noe (1990, 1992; No € e, van Schaik, € & van Hoof, 1991), and Barrett and colleagues (Barrett, Gaynor, & Henzi, 2002; Barrett, Henzi, Weingrill, Lycett, & Hill, 1999), several studies have looked for and found evidence of BMT in a variety of primate species, from lemurs (Norscia, Antonacci, & Palagi, 2009; Port, Clough, & Kappeler, 2009) to monkeys (Fruteau, Lemoine, Hellard, van Damme, & Noe, 2011; Gumert, € 2007; Tiddi, Aureli, & Schino, 2012) and apes (Kaburu & NewtonFisher, 2015a, 2015b; Koyama, Caws, & Aureli, 2012; NewtonFisher & Lee, 2011). With an increasingly large number of studies, a review such as the one by Sanchez-Amaro and Amici (2015)  would be warmly welcome as a timely summary of the evidence for BMT, and an indication of future directions. The authors identify four areas of interest and usefully highlight some potential issues with BMT, for example where free trading is compromised by extortion or the need for comparable methods across studies. However, while their aims may be laudable, we feel there are particular flaws in some of their arguments and some misrepresentation of cited literature that we would like to correct:
Ronald Noë's insight:
The official publication - 8 months after was accepted. (See earlier scoop about this paper for my comment)
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Akçay 2016 Population structure reduces the benefits from partner choice in mutualism BioRXiv

Akçay 2016 Population structure reduces the benefits from partner choice in mutualism BioRXiv | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it

Abstract: Mutualisms are key drivers of evolutionary and ecological processes. Understanding how different species can evolve to interact in mutually beneficial ways is an important goal of evolutionary theory, especially when the benefits require costly investments by the partners. For such costly investments to evolve, some sort of fitness feedback mechanism must exist that more than recoups the direct costs. Several such feedback mechanisms have been explored both theoretically and empirically, yet we know relatively little how they might act together, as they probably do in nature. In this paper, I model the joint action of three of the main mechanisms that can maintain symbiotic cooperation: partner choice by hosts, population structure amongst symbionts, and undirected rewards from hosts to symbionts. My results show that population structure reduces the benefit from partner choice to hosts. It may help or hinder beneficial symbionts and create positive or negative frequency dependence depending on the nature of host rewards to the symbiont. Strong population structure also makes it less likely that host choosiness and symbiont cooperation will be jointly maintained in a population. The intuition behind my results is that all else being equal, population structure reduces local variation available to the host to choose from. Thus, population structure is not always beneficial for the evolution of cooperation between species. My results also underscore the need to do full analyses of multiple mechanisms of social evolution to uncover their interactions to uncover the interactions between them.

Ronald Noë's insight:
Haven't read this yet, but seems relevant to biological market theory too
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Sheskin, Lambert & Baumard (2016). Biological markets explain human ultrasociality. Behavioral and Brain Sciences, 39, e113

Sheskin, Lambert & Baumard (2016). Biological markets explain human ultrasociality. Behavioral and Brain Sciences, 39, e113 | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
The evidence Gowdy & Krall (G&K) provide is more consistent with a biological markets explanation of human ultrasociality than a group selection explanation. Specifically, large-scale societies provide a better biological market for cooperation than do small-scale societies, allowing individuals to increase their fitness. Importantly, many of the quality-of-life costs G&K discuss (e.g., patriarchy) are not fitness costs.
Ronald Noë's insight:
A defence of the BMT approach in a comment on a rather puzzling group-selection based target paper in Behavioural and Brain Sciences (Gowdy, J., & Krall, L. (2016). The economic origins of ultrasociality. Behavioral and Brain Sciences, 39, e92)
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Schott, Valdebenito, Bustos, Gomez-Porras, Sharma & Dreyer (2016). Cooperation through competition - Dynamics and microeconomics of a minimal nutrient trade system in arbuscular mycorrhizal symbios...

Schott, Valdebenito, Bustos, Gomez-Porras, Sharma & Dreyer (2016). Cooperation through competition - Dynamics and microeconomics of a minimal nutrient trade system in arbuscular mycorrhizal symbios... | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
In arbuscular mycorrhizal (AM) symbiosis, fungi and plants exchange nutrients (sugars and phosphate, for instance) for reciprocal benefit. Until now it is not clear how this nutrient exchange system works. Here, we used computational cell biology to simulate the dynamics of a network of proton pumps and proton-coupled transporters that are upregulated during AM formation. We show that this minimal network is sufficient to describe accurately and realistically the nutrient trade system. By applying basic principles of microeconomics, we link the biophysics of transmembrane nutrient transport with the ecology of organismic interactions and straightforwardly explain macroscopic scenarios of the relations between plant and AM fungus. This computational cell biology study allows drawing far reaching hypotheses about the mechanism and the regulation of nutrient exchange and proposes that the ‘cooperation’ between plant and fungus can be in fact the result of a competition between both for the same resources in the tiny periarbuscular space. The minimal model presented here may serve as benchmark to evaluate in future the performance of more complex models of AM nutrient exchange. As a first step towards this goal, we included SWEET sugar transporters in the model and show that their co-occurrence with proton-coupled sugar transporters results in a futile carbon cycle at the plant plasma membrane proposing that two different pathways for the same substrate should not be active at the same time.
Ronald Noë's insight:
This is a rather remarkable and refreshing approach to the question of the mechanisms behind the exchange of nutrients (phosphorus and carbon notably) between plants and arbuscular mycorrhizal fungi. The authors apply computational cell biology to construct in silico models of the C - P exchange at the common plant/fungus membranes. They arrive at the remarkable conclusion that the system could be relatively simple, simpler at least than many empiricists working on the AM symbiosis generally assume. Obviously the model needs empirical testing, but the results seem to fit existing data well. More interesting here: the model supports viewing plants and AM fungi as traders on a market that cooperate by each maximising their own net gain. In order to show this the authors discuss some basic micro-economic principles and engage in a bit of game theory. The fit with basic market characteristics even inspires them to adapt a quote from Adam Smith in their conclusions.

I am not well placed to judge the technical details of both the model and the biological mechanisms involved, but if even if the assumptions and assertions of this paper only hold halfway, it could prove to be a rather interesting new way of tackling the problem of the evolution and stability of this both economically important and scientifically intriguing inter-specific mutualism.
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Wubs, Bshary & Lehmann (2016). Coevolution between positive reciprocity, punishment, and partner switching in repeated interactions. Proceedings of the Royal Society of London B: Biological Science...

Wubs, Bshary & Lehmann (2016). Coevolution between positive reciprocity, punishment, and partner switching in repeated interactions. Proceedings of the Royal Society of London B: Biological Science... | Biological Markets: the role of partner choice in cooperation and mutualism | Scoop.it
Cooperation based on mutual investments can occur between unrelated individuals when they are engaged in repeated interactions. Individuals then need to use a conditional strategy to deter their interaction partners from defecting. Responding to defection such that the future payoff of a defector is reduced relative to cooperating with it is called a partner control mechanism. Three main partner control mechanisms are (i) to switch from cooperation to defection when being defected (‘positive reciprocity’), (ii) to actively reduce the payoff of a defecting partner (‘punishment’), or (iii) to stop interacting and switch partner (‘partner switching’). However, such mechanisms to stabilize cooperation are often studied in isolation from each other. In order to better understand the conditions under which each partner control mechanism tends to be favoured by selection, we here analyse by way of individual-based simulations the coevolution between positive reciprocity, punishment, and partner switching. We show that random interactions in an unstructured population and a high number of rounds increase the likelihood that selection favours partner switching. In contrast, interactions localized in small groups (without genetic structure) increase the likelihood that selection favours punishment and/or positive reciprocity. This study thus highlights the importance of comparing different control mechanisms for cooperation under different conditions.
Ronald Noë's insight:
This paper is of marginal interest as far as BMT is concerned, since what is meant by 'partner switching' here is switching in its pure form: partner desertion and forming a new (dyadic) partnership with a randomly chosen partner. This lacks the crucial element of partner choice on the basis of characteristics of the partner, which - notably due to the selective effect of partner choice - is the 'raison d'être' of BMT.

The authors make one interesting point that applies to biological markets in general: partner switching (and partner choice too) are less beneficial the smaller the group of partners is from which one can choose. This makes of course (common) sense, but is nevertheless often overlooked when BMT is tested in species living in small groups in which potential cooperative partners are recruited within the group only.

The way in which the authors define partner choice and partner switching, which makes sense to me too, made clear to me that we need a 3rd term to describe switching partners in order to swap the present partner for a more profitable one. This then may force the abandoned partner to some switching too, of course, but the abandoned one has a choice between no partner and any partnership that generates a net gain. This applies obviously only to cases in which the number of partners is fixed and notably to dyadic forms of cooperation.

"Advantageous partner switching" would be an option, but  it is a bit clunky and I welcome better ideas.

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