1 Evolutionary motivations towards pro-social moral norms
Descriptive ethics is the study of why and how humans behave individually and collectively in ways that give meaning to the terms âethicalâ or âmoralâ. The motivation to behave in these ways has within descriptive ethics a more naturalistic, not to say pragmatic, nature. A motivation to cooperate and to behave altruistically is at the heart of resolving the dilemmas of social life. If cooperative and altruistic behaviour runs counter to the evolutionary interests of the individual, we must consider the likelihood of such behaviours occurring. Research on evolution offers both evidence and models of the selection by these types of behaviour, both in individuals and in groups. Understanding the relationship between individuals and their interaction within the broader situation offers an indispensable insight into why cooperative and moral behaviours are selected in the complexity of human social groups. These social decisions are the most telling in nature, for they are existential. Evolutionary theory is concerned with fitness through the process of natural selection (Darwin 1859). It therefore follows that if ethical behaviours are subject to evolutionary processes, these must be understood by the descriptive ethicist. Evolutionary theory is concerned with why genes and humans (in their very different ways) are selected. Much of the explanation of processes that influence the development of ethics and moral behaviour, be it in neurology, psychology or theories of group behaviour, do not answer the question why; they in fact answer the question of how. To confuse the two can lead to, and in many cases has (El Mouden 2011; West et al. 2007) led to, the assertion that the reason why humans behave in more or less moral ways is due to the nature of the brain, personal values and group norms, among other evolved phenomena. Essential though these processes are to the understanding of ethical behaviour, they can and should only be used in a manner consistent with the ultimate motivation of evolutionary theory.
This chapter introduces some of the arguments and research that bear on descriptive ethics and some of the recent debates within evolutionary biology. It specifically aims at highlighting evolutionary theory and evidence that can be applied to human social groups and the relationship between the situation and the propensity to behave cooperatively and altruistically, both of which are essential to pro-sociality. From this basis, how these processes are manifest in humans and groups may be explored in a systematic and grounded manner. Only then will it be possible to offer a description of the ethics within a group with any claim to ultimate causality.
Although the ground covered in this chapter is substantial, it is necessary for two reasons: first there is an increasing literature on the misconception and resulting confusion within the evolutionary literature (not to mention social sciences) of some of its key concepts (Dawkins 1979; Scott-Phillips et al. 2011; West et al. 2007). For that reason, the rationale for the models and points of analysis must be explained, themes that will be expanded in later chapters. Finally, introducing an evolutionary dimension into any discussion of the ethics of moral norms invites the criticism of âreductionismâ. We will argue that this is precisely a consequence of the confusion between âhowâ and âwhyâ, and in fact current evolutionary biology and anthropology offer a coherent thread of motivation that provides unique insights into the motivations we may expect to be present in the resolutions of the dilemmas of social life.
Recognising the evolutionary value of social interdependence
That social behaviour exists does not require discussion. The underlying motivations for selection on the basis of group behaviour for social behaviour, however, are more contested. Why is it that altruistic behaviour occurs, and what factors affect the intra-group and inter-group selection?
The dominant explanation for altruistic behaviour is that of inclusive fitness theory (Hamilton 1964). Some recent alternatives will be highlighted later in the areas in which they seek to add substantially new insights on the evolution of social behaviour within and between groups. The theory against which these alternatives are compared will be that of inclusive fitness, which is the most established. The discussion of social behaviour from an evolutionary perspective offers clarity of definition and an argument that will facilitate later discussions within the psychological social sciences.
The mechanisms of selection between individuals, groups and context (ecosystem) are debated; however, the classification of behaviour within interactions may be common to each.
An evolutionary case for altruistic behaviour is conventionally based on the âinclusive fitnessâ that results from it (Hamilton 1964). Inclusive fitness captures both direct fitness, that is to the individual, and indirect fitness, that of the most similar gene pool. This can be represented in the simplified formula:
- where c is the reproductive cost to the actor of altruistic behaviour
- r is the likelihood over the mean of the recipient sharing genes with the actor
- b is the reproductive benefit to the recipient of the altruistic act
A social behaviour within evolutionary theory is that behaviour that has an outcome on both the actor (who acts) and the recipient (who is acted on) of selfishness, mutual benefit, altruism and spite (Hamilton 1964; West et al. 2007), and each behavioural strategy is subject to natural selection.
Table 1.1 The range of altruistic behaviours permitted in inclusive fitness optimisation | Effect on Recipient | |
| + | â |
Effect on Actor | | |
+ | Mutual Benefit | Selfishness |
â | Altruism | Spite |
An important attribute of this classification is that its categories are intentional. This is clearly relevant when considering human groups, in which situation may significantly change the intentionality of behaviour.
Not every interaction needs to be cooperative to support individual or collective fitness. Inclusive fitness predicts that cooperative and altruistic behaviours are driven by the costs and benefits and the degree of relative genetic similarity between the parties. However, (see Table 1.1) altruistic behaviour may also have the effect of spite in the case of negative relatedness with the recipient in question, a result that may have implications for the attributes of groups,1 but also of behaviours that are locally non-costly to both parties yet perceived as cooperative to the fitness of the group as a whole.
The theory of inclusive fitness enables behaviours that are not exclusively concerned with personal reproductive outcome to be explained in evolutionary terms. To be clear, as misuse of this term is widespread (Grafen 1984), inclusive fitness captures the reproductive success of the individual (direct fitness) with the harm or benefit their behaviour has on other members multiplied in each case by their degree of relatedness (indirect fitness). Hamiltonâs equation is concerned wholly with the hereditary costs and benefits of a given behaviour. The motivation towards balance between direct and indirect fitness behaviours (individual or group reproductive success), however, is not unqualified. Hamilton himself drew explicit attention to this fact: âAltruistic or selfish acts are only possible when a suitable social object is available. In this sense behaviours are conditional from the startâ (Hamilton 1987: 420). This social object refers to the attributes of the environment in which the relatedness group functions. The implication then of Hamiltonâs theory is that the external environment will affect the evolutionary value of an altruistic behaviour between individuals.
Non-kin altruism
The theory of inclusive fitness has, since almost its first appearance, been associated with kin selection; that is, one is altruistic to those who share common genes. Hamilton himself argued (Hamilton 1987: 420) that inclusive fitness was not fully explained by kin selection, and this confusion was due to an overly restrictive definition of the motivation to discount direct fitness costs in a social interaction.
In the original formulation of the Hamilton equation, ârâ was defined as immediate shared genetic heredity. This however did not explain altruism within heterogeneous groups, and certainly not in human social structures in which altruism is common among unrelated individuals. In fact, as Bell and colleagues observe, âthe scale of costliness of human altruism is extraordinaryâ (Bell et al. 2009) and certainly beyond the expectation of genetic similarity.
In order to render inclusive fitness more explanatory of these observed attributes of both genetic and human social structures, an adaptation has occurred in the definition of the concept of ârâ, which has resulted in the use of the term ârelatednessâ rather than relationship.
Hamiltonâs relatedness captures the relative rather than absolute genetic similarity between an actor and a recipient: âa statistical measure of genetic similarity regardless of its sourceâ (Pepper 2000). This represents a process of gradually broadening the domain of indirect fitness. It can now be fitness optimal to be altruistic to anyone who is more genetically similar to oneself than is the average for the population, that is, not limited to the group with which you interact; for example, those within a group or organisation. The implication is that inclusive fitness does not imply a specific type of behaviour; indeed, even altruism may be harmful to a recipient if it improves inclusive fitness for the individual and, by extension, the relatedness group to which the actor belongs.
In the strictest sense, inclusive fitness has difficulty in explaining cooperation and even altruism between species and non-related kin. It cannot, for example, explain inter-species symbiosis, or reciprocal altruism (Trivers 1971) which has, in many iterated prisonerâs dilemma games, been found to yield cooperation between non-relatives (Axelrod 1986). Reciprocal altruism introduces a model of survival of altruistic genes through behaviour that is conditional on that of the other player. The most famous example of this is the tit-for-tat strategy. In this a player cooperates and then mirrors what the other player did in the previous round. This strategy minimises exploitation while exploiting cooperation, and is therefore fit over time without relatedness.
Hamilton, too, does not directly account for non-additivity between generations â that is, genetic mutations (like eye colour) that are not always passed directly from parent to child. Generational non-additivity does not fall within our central concerns, so we may simply observe that there is no guarantee that altruistic genes will be transmitted from parents to children.
The evolutionary imperative of conditionally helping altruists
Inclusive fitness in its simple form is left with a series of explanatory challenges. It does not explain altruism between relatively distinct non-kin, as we have illustrated earlier in the case of reciprocal altruism. Its explanatory weakness stems from a conflation of two distinct processes by which inclusive fitness is attained by large-brained animals. When accounted for, these lead to a dramatic alternative understanding of how altruism survives in social groups that is also found in human social groups.
Queller (1985) shows through a coherent elaboration of the Hamilton equation that it is a combination of genotype and phenotype that achieves positive assortment (that is, breeding with more genetically similar types) and therefore the survival of altruistic genes. Queller does not provide an alternative to Hamilton, but places phenotype as a fundamental means of how altruism is transmitted over time.
In Quellerâs analysis including the role of phenotype, the average genetic type of the population plays little or no part. In fact, he showed that it was phenotype that played a greater influence on the survival of the gene, as the gene requires sufficient âhelpingâ behaviour from non-related actors to achieve related mating. Altruists will survive if they receive more fitness benefits than costs that they absorb (Fletcher and Zwick 2006). This costâbenefit evaluation of behaviour may be efficiently enhanced by punishment and policing (Lehmann and Keller 2006) on the part of the group.
This seemingly counter-intuitive result stems from the observation that an altruistic allele (mutation) is not shared throughout the same genetic group. It is therefore the central motivation of the gene to reproduce itself through assortment (mating) with a similar altruist or those having the helping phenotype (and therefore not necessarily âkinâ). In order that the altruist survives and the benefit to the group is maintained, it is in the interest of other members of the group that the altruist be compensated for the costs of their altruism to the extent to which they value the survival of altruistic behaviour. Helping behaviours in others ensures that the altruistic actor does not become extinct and, with it, the allele. Phenotypic helpers are then central to the survival of the altruistic gene. They become suitable mating partners for non-altruists, as they have a fitness effect on their gene.
In short, altruism can survive through positive assortment, by phenotypic transmission rather than by genetic relatedness alone.
In an extreme case, Hamiltonâs r would be zero, and therefore the aggregate nature of the r term would lead to flawed expectations of altruistic behaviour exclusively derived from genetic rather than phenotypic transmission.
Two major implications follow from Quellerâs work that are of importance for our discussion:
- i) Altruistic behaviour survives through the conditional reciprocated helping behaviour from others rather than a direct contribution to the related gene pool. The conditions in which helping is valid are contingent on the situation.
- ii) The direct fitness benefit (B) of an action is a consequence of the behaviour (phenotypes) of the population rather than the genotype of the altruist alone.
The central analytical focus is therefore shifted to the situational attributes that ensure that altruistic behaviour is net positive to the altruist and as such the group. Combining the claims of both theories, we have four conditions for cooperation and altruism (helping) as an evolutionary phenomenon (Lehmann and Keller 2006):
- i) The direct cost of helping is less than the benefit received. Helping conditionally increases the direct fitness of both.
- ii) Helping behaviour will increase the likelihood of receiving helping behaviour in the future. As noted earlier, this likelihood is situationally contingent.
- iii) Altruistic helping occurs subject to genetic distinctiveness (i.e. kin selection).
- iv) Altruistic behaviour occurs subject to phenotypic distinctiveness, that is, recognising individuals who have known behavioural profiles.
Using Quellerâs introduction of phenotype, we may now specifically consider the effect of demography on the situations specifically conducive to helping behaviours in the phenomena of âgroup formation and extinction, migration between groups, reproduction, offspring dispersal, and deathâ (Pepper 2000, p. 7). As demography changes, so, too, does the value of helping behaviour to the individual and the group. If helping is less favourable to the group, social dilemmas faced locally may put altruists at risk by favouring more or less likely helping behaviours in response to them, and therefore risk becoming extinct in the group. As a response, âhelpersâ may adopt conditional behaviour in repeated interactions between non-relatives (Fletcher and Doebeli 2006) in order to maintain a viable population by changing the structural nature of the social dilemmas being encountered. In repeated interactions, then, a behavioural (cultural) phenotype will enable helping to remain viable in changing conditions.
The evolutionary rationale for altruistic behaviour does not change because of Quellerâs work. It does, however, expand the means by which demography and notably culture or phenotype support, albeit conditionally, helping behaviour. The means of transmission are quite different between genes directly and phenotype. As the role of phenotype provides the most significant explanation of the survival of altruistic genes (following Queller), it is to the understanding o...