Research questions based on Ratnieks et al. 2006 (Clara B. Jones)

Research questions for group-living vertebrates based on Ratnieks et al. 2006*, **

Introduction

Inclusive fitness theory explanations for both cooperation and conflict

What do we mean by conflict resolution?
1. "Potential conflict is any difference in the reproductive optima of individuals or groups within a society."
2. Individuals have three options...to be workers, reproductives, or dependents (e.g., "social parasites") or some combination of these serially or concurrently. Categories can be analyzed by class or rank, by sex, by age, by race or ethnicity, etc.
3. For vertebrates, including, humans, how do we estimate "fitness optima?"

Kinship, coercion, and constraint
"...low relatedness makes wasteful conflicts more likely because the cost of conflict falls upon more distant kin."

Sex allocation
...differential investment in males and/or females & differential costs/benefits...

Conflict-resolution
1. "Coercion and constraint may help to resolve conflict if one party has complete power over sex allocation." ...etc., e.g., over access to limiting resources...
2. "Mixed power seems likely in many but not all situations." ...e.g., "division of task"..."division of labor"...n.b. DoL more efficient than DoT; also note, male & female interests may not coincide since males are expected to be time-minimizers, females, energy-maximizers...
3. Under some conditions, is it more beneficial for an individual to assume the costs of conflict rather than to resolve conflict? ...i.e., When are costs of conflict [over reproduction &/or limiting resources (convertible to offspring] tolerated or preferred relative to benefits of conflict-resolution?
4. Importance of information, especially, competition for information [see EO Wilson 1971 on "communication" in societies]...Humans: language, who shares information with whom, etc...
5. Investigate "policing"*** as a form of conflict-resolution [mitigation of selfishness], mitigation of conflict [e.g., force, coercion, constraint, persuasion, etc. (see French & Raven's "bases of power")]...
6. What factors lower incentives for individual selfishness [& subsequent "tragedy of the commons"]
7. High "r" lowers the proportion of reproductives in a population...
See "reproductive skew" literature, e.g., R Hager & CB Jones 2009 CUP...

Conflicts among totipotent individuals
1. "Totipotency greatly increases the potential for conflict...."
2. If optimal family and/or group size can be estimated, excess individuals are likely to increase potential for conflict...

Discussion & Conclusions
1. Class conflicts relatively easy to resolve [because dominants "police" subordinates?]...
2. Conflict easily resolved when one party "powerless"...
3. Should the incorporation of steriles or other non-reproductives in groups reduce conflict?...relative to what [e.g., "r"]...
4. How do humans escape control in groups? [see French & Raven's "bases of power"]
5. What role does genetics play in conflict-resolution [e.g., genetic conflict]...

*Ratnieks FIW, Foster KR, Wenseleers T (2006) Conflict resolution in insect societies. Annu. Rev. Entomol. 51: 581-608.****
**Sentences in quotation marks are quotations from paper; other statements, comments, or questions are my interpolations relative to group-living vertebrates, including, humans.
***New paper: "'Enforcement' is central to evolution of cooperation"...i.e., &, also, to evolution of repression of selfishness... https://www.nature.com/articles/s41559-019-0907-1

****Topics/questions based on titles of literature cited:
--evolution of male and female traits
--causes & consequences of informational constraint
--conflict over class/SES determination
--status-allocation to offspring
--Does it "pay' kin to favor relatives who are "losers" or inferior reproductives and/or competitors?
--n.b. Charnov EL (1978) Evolution of eusocial behavior: offspring choice or parental parasitism? JTB 75: 451-465.
--n.b. Crespi BJ, Ragsdale JE (2000) A skew model for the evolution of sociality via manipulation: why it is better to be feared than loved. Proc R. Soc. London B Biol. Sci. 267: 821-828.
--facultative policing
--conflict over paternity [n.b. causes & consequences of multiple-mating by females]
--n.b. Keller L (1997) Indiscriminate altruism: unduly nice parents and siblings. TREE 12: 99-103.
--genes regulating complex social behavior
--ancestral states of complex sociality
--sex-ratio determination in groups & populations [e.g., conflict over sex-ratio determination]
--causes & consequences of sexual deception [e.g., in some mammal species, males are indistinguishable from adult females for some time before expressing secondary sexual characteristics (see, e.g., mantled howler monkeys, Alouatta palliata)]
--selfish tactics & strategies that promote sociality [e.g., "social cohesion"]
--implications: "heirs & spares"
--mechanisms of conflict-management & conflict-resolution in eusocial taxa
--n.b. Ratnieks FLW, Wenseleers T (2005) Policing in insect societies. Science 307: 54-56.
--alternative reproductive strategies and conflict-resolution/conflict-management
--n.b. Sachs JL, Mueller UG, Wilcox TP, Bull JJ (2004) The evolution of cooperation. Q. Rev. Biol. 79: 135-160.
--policing x age, sex, class/SES, race, etc.
--Is multiple-mating by females rare in vertebrates?
--causes & effects of totipotency; ubiquitous in vertebrates?
--genetic basis of sterility
--mechanisms of kin discrimination
--What traits differentiate members of classes [within & between]?
--relevance of behavioral flexibility & phenotypic plasticity to conflict-management & conflict-resolution [also, flexibility of group structures]

When to exhibit Social Behavior [Cooperation, Altruism]? Clara B. Jones [Graph, Legend]

Fig. 1. When should an Actor or a Recipient exhibit Social* Behavior [Cooperation, Altruism]? Clara B. Jones, 3/13/2019

1. This [idealized] graph**** depicts & predicts when an [idealized] organism--an Actor or a Recipient--should give/exhibit Cooperation or Altruism, defined, formally, as those interactions during which the Recipient benefits [in relative reproductive units].
2. X-axis: Time [T], Energy [E] investment in Social Behavior [reflects organism's T/E budget for a given interaction].
3. Y-axis: Costs [C, in reproductive units to Actor or Recipient] or Benefits [B, in reproductive units to Actor or Recipient] to Inclusive Fitness** [IF= selfish reproductive units (usually, & clasically,  offspring, though, offspring might be delayed, such as when transaction in wealth or other goods--see "Altruism" below) + reproductive units of kin]. r, coefficient of relatedness, is incorporated in measure/estimate of IF [how to measure?].
4. C rise or reach asymptote over T. Costs, for Actor or Recipient, or both, may prevent Cooperation or Altruism between kin when, for example, Cooperation or Altruism increases Competition [for limiting resources such as food, mates, burrows], increasing [for ego or ego's offspring] likelihoods of  dispersal, "competitive exclusion" [demand for ~identical limiting resources], death, etc. Cooperation and Altruism are expected, by definition, to be responses for the avoidance or neutralization of Competition.
5. B rise, then decrease over T.
6. "x" represents the optimal point at which an Actor or a Recipient should exhibit Cooperation [both Actor and Recipient gain reproductive units] or Altruism. [Actor loses, Recipient gains reproductive units]. "x" is the extension to the X-axis of the vertical, black line connecting the points with the greatest separation between the C and B curves--where B > C. "x" is the idealized point maximizing inclusive fitness benefits.
7. The shaded area represents the zone around "x" for which Actor's benefits from Social Behavior are highest. Ideally, the broken vertical lines extend vertically & downward from those points on either side of "x" on the B curve at which the B curve first begins to descend. The shaded area represents the zone of maximal benefit to Inclusive Fitness of A or B, in this case, maximum benefit from an act of Cooperation or Altruism ["IF maximizing"].
8. Actor & Recipient curves and values will usually be asymmetric, by definition [possibly, even, for clones (e.g., identical twins)]. In other words, zones of maximal benefit from social behavior should differ for Actor and Recipient. Overlaying Actor's & Recipient's curves will determine whether there is overlap between Actor & Recipient zones of greatest benefit to ego from Cooperation or Altruism. If there is overlap, the zone of mutual benefit is defined, so that Actor & Recipient should exhibit Cooperation [+, +] within these parameters [of T, E]. If there is no overlap, then conflict arises [between Actor and Recipient &, possibly, kin of one or both].
9. Similarly, an Actor should exhibit Altruism [-, +] to a Recipient when the long-term B of such action[s] outweigh the short-term C [determined by hard-wired Hamiltonian algorithm?].
10. If B from Social Behavior are minimal or non-existent [e.g., low T/E "budget;" little or no overlap between Actor-Recipient zones of greatest benefit], then interactants should adopt an alternate behavior [Alternative Reproductive Behavior--tactic or strategy], such as fight***, avoidance, flight, force, coercion, persuasion, manipulation, exploitation, coexistence--or some combination of these.
11. Group-living organisms experience relatively ongoing "decisions" about how to behave relative to conspecifics; however, it is important to keep in mind that "fitness budgets" change over T and that behavior is condition- [context-] dependent.
12. Ceteris paribus, and on average, curves, B & C, as well as, shapes of zones of maximum benefits for Males [time-minimizers] and Females [energy-maximizers] should differ. Curves, also, are expected to be Age-, Class-, & Role-dependent.

*After Hamilton (1964), "Social Behavior" is defined as Cooperation [both Actor and Recipient gain reproductive units] or Altruism [Actor forfeits reproductive units; Recipient gains reproductive units]
**The term "inclusive fitness" can be confusing since, technically, "fitness" is the frequency of an allele in a population averaged across the reproductive success of all individuals in the population bearing the allele, leading one to inquire how important it is to measure individual variation of traits [since individual variation will be subsumed in the measure of an allele's "fitness"].
***As pointed out by Geoff Parker [1974], aggression should not usually be a first resort because the C of damaging behavior generally outweigh the B.
****The same graphical/conceptual framework could be used for Coexistence if axes modified so that X-axis= savings in T, E to Mean fitness of allele 1 [or 2]; Y-axis= B, C from T, E savings to fitness [Mean frequency] of allele 1 [or 2] in a population. In the case of Coexistence, a Community Ecology process, the population would be escaping Competitive Exclusion. [See, also, my 2014 Springer Brief, Ch 2, for discussion of Coexistence in relation to Hamilton's Rule.]

Acknowledgments: Thanks to my son, Luke [M. Luke Jones] for stimulating discussion and for drawing the figure. Further, via e-mail, I asked Andrew Bourke [Univ. of E. Anglia] to comment on this blogpost. On 4/11/2019, I received the following reply [n.b.: my original blogpost above, including, the graph, have NOT been revised subsequent to Bourke's comments]:

1.     Perhaps the X axis needs clarifying a little – after all, time and energy seem to have most relevance in this context insofar as they are proxies for fitness, so then there would be some relationship between the metrics on the X axis and on the Y axis, and how these interact might then require consideration.

2.     It would be good to have concrete, empirical examples of when the scheme in the figure needs invoking, i.e. showing how considering the time/energy investment dimension in the manner advocated in the figure enlarges our understanding of the occurrence of social behaviours.  Personally I find it hard to think of things in the abstract all the time, so even some hypothetical biological examples/applications might be useful.

3.     I think your footnote 8 makes a good point, i.e. that actor and recipient interests need not overlap even for altruistic and cooperative interactions.  This point was explored theoretically in a fairly old paper by Yamamura and Higashi (1992)*****.  Their work does not seem to have been followed up empirically to any great extent, perhaps because the usual assumption that one of the parties has most control over the interaction (in which case conflict is minimised – e.g. an insect larva may often have little choice but to accept the level of care it’s given) holds.

References
Hamilton WD (1964) The genetical evolution of social behaviour. J Theor Biol 7: 1-52.
Parker, G (1974) Assessment strategy and the evolution of fighting behavior. J Theor Biol 47: 223-243.
*****Yamamura N, Higashi M (1992) An evolutionary theory of conflict-resolution between relatives: altruism, manipulation, compromise. Evolution 46: 1236-1239.

http://www.kurims.kyoto-u.ac.jp/~kyodo/kokyuroku/contents/pdf/0827-14.pdf