Evolutionary Conditions of Happiness

What it means to lead a happy life is among the most long-standing questions of humanity. This question is also inextricably connected to the question of human nature, an answer arguably has to take into account the hypotheses and results of evolutionary anthropology and psychology.

This paper provides a review of evolutionary conditions of happiness, understood as a descriptive notion, denoting a psychological state or condition of a subject that has a positive quality for her – in some non-evaluative sense of ‘positive’ (cf. Haybron, 2020, § 1). Thus, a state of happiness is positive from the perspective of the subject. Accordingly, a judgment that a subject is in this state has to take into account her own perspective, but does not have to imply any intersubjective judgment on whether the state is also ‘good for’ the subject in an evaluative sense.¹ That a state (psychological or otherwise) of a subject is ‘good for’ or ‘benefits’ her in an evaluative sense is denoted with the term ‘well-being’ (also ‘prudential value’, ‘flourishing’, or ‘welfare’). Thus, an ascription of well-being has to involve some kind of evaluative appraisal from an intersubjective perspective² (which a subject can also apply to herself).

While the terms ‘happiness’ and ‘well-being’ are not used consistently in the manner delineated here across or even within different disciplines,³ they are taken in the following to refer to different concepts employed in relation to different questions. Nonetheless, the concepts also seem to be closely interrelated. Happiness is often considered if not as a sufficient, then as a necessary condition for well-being (at least under normal circumstances): An overall state of a subject that is prudentially beneficial plausibly also has to include a favorable psychological condition as an aspect (Haybron, 2020, § 4.1). Accordingly, many accounts of well-being also comprise an account of happiness (Novak & Kiknadze, 2024).⁴ An example are ‘eudaimonic’ conceptualizations of happiness, which derive from the (evaluative) Aristotelean notion of eudaimonia, but consider only the psychological aspects of it (Delle Fave et al., 2011, p. 187).

Theoretical conceptions of happiness can be grouped, roughly, into ‘hedonistic’, ‘emotional state’, ‘life satisfaction’, and hybrid views (Haybron, 2020, § 2.1). According to hedonistic theories, happiness is constituted by a predominance of pleasant over non- or unpleasant psychological states. Emotional state theories conceive of happiness as an overall positive condition with regard to emotion⁵ or mood⁶ states. Life satisfaction views hold that happiness consists in a favorable attitude towards one’s life as a whole. Using the term ‘affect’ to stand for emotions, moods, and hedonic states taken together (a terminological convention adopted in the following), hedonistic and emotional state views may be grouped as ‘affect-based’ accounts of happiness (Haybron, 2020, § 2.3, cf. Intelisano et al., 2020, p. 164). Since life satisfaction judgments, in contrast, are typically not considered to be affects (Feldman, 2010, pp. 77–78), life satisfaction views can be opposed to the other two kinds of theories as not affect-based (for further terminological differentiations, cf. Haybron, 2020, notes 7 and 8, and Heathwood, 2022). Finally, there are hybrid accounts combining aspects of different views in a comprehensive conception of happiness. Empirical happiness constructs or indicators are also often of a ‘hybrid’ form. Here, ‘hedonic’ and ‘eudaimonic’ conceptions are usually distinguished (Ryan & Deci 2001, cf. Delle Fave et al., 2011, 2016),⁷ based on the degree to which affective or more cognitive (and possibly evaluative⁸) aspects (such as meaning, self-actualization, personal growth, and social connectedness) are included. Commonly cited examples are the constructs of Subjective Well-Being (Diener et al., 1985; Diener, 2000; Pavot & Diener, 2008) as a ‘hedonic’, and Psychological Well-Being (Ryff, 1989, 2014; Ryff & Singer, 2008)⁹ as a ‘eudaimonic’ conception. Recently, Intelisano et al. (2020) have proposed to transcend this binary opposition in favor of a gradual distinction based on the relative weight put on cognitive and affective aspects, respectively.

Why would a perspective integrating evolutionary anthropology and psychology be helpful for a deeper understanding of happiness? As Nettle and Scott-Phillips (2023) argue, there is a certain indispensability of the evolutionary perspective for psychology. The uncontentious commitment that ‘[p]sychological mechanisms can be usefully viewed as exhibiting some kind of functionality for the whole organism’ (Nettle & Scott-Phillips, 2023, p. 23) engenders an at least implicit appeal of evolutionary explanations, because any account of psychological mechanisms and states in terms of proximate functions ‘always leads to an invocation of the functionally organizing power of natural selection, either directly via evolved functions, or more indirectly via derived functions that depend on evolved functions’ (Nettle & Scott-Phillips, 2023, p. 30). The reference to evolution by natural selection as the ultimate basis of proximate functions¹⁰ may remain implicit, but this does not change the conclusion that an evolutionary perspective is, in a certain sense, indispensable.

What is going to be explored in the following is an explicit account of evolutionary conditions of happiness as a psychological state – in line with an approach which has recently been called ‘Positive Evolutionary Psychology’, and characterized as ‘the use of evolutionary principles to help guide people toward living richer lives’ (Geher & Wedberg, 2019, p. 2). While the term came into use only recently, the approach has been pursued for some time already, by a number of scholars, who have produced an array of publications in the last thirty-something years, but who seem to have taken only little notice of each other’s works. A review of the literature in this emerging field of research is the main aim of this paper, which may contribute to a stronger integration of the approach (cf. Geher et al., 2023, pp. 3–4, on this desideratum).

‘Evolution’ is meant in the following to refer not only to ‘natural’ (or biological), but also to ‘cultural’ evolution (cf. Chudek et al., 2016) – two processes that run in parallel (in different paces), but also intertwine in natural-cultural ‘co-evolution’. The resulting co-evolutionary perspective is not committed to uniformity or fixity of the human psyche, but opens ‘space for environment, context, culture, meaning or agency’ (Nettle & Scott-Phillips 2023, p. 21) - and especially, for cross-cultural variation and construction of how happiness is conceptualized and realized (cf. Miyamoto & Ryff, 2011, Uchida & Ogihara, 2012, Joshanloo, 2013, Ho et al., 2014, Delle Fave et al., 2011, 2016).

It is unlikely that evolutionary conditions of happiness can be elucidated through pointing out a specific function (or functions) of it, and then developing an account that basis. ‘Happiness’ is too broad a term to allow for any such simple reduction. Nonetheless, there seem to be more specific psychological states, which are ‘happiness-constituting’ (Haybron, 2008, p. 107) and for which functional characterizations can be given.¹¹ A paradigmatic example of psychological states which are widely considered to be happiness-(and unhappiness-)constituting are states that have a certain ‘positive’ or ‘negative’ affective valence as a central aspect – such as pleasure/displeasure, or positive/negative emotion, or positive/negative mood. In an overview of extant accounts of happiness in philosophy and psychology, Intelisano et al. (2020, pp. 189–191) point out that there are nearly no exclusively affective accounts of happiness, but they also find nearly no exclusively cognitive accounts, which would deny any relevance of affective states for happiness. In fact, the mere possibility of such a denial is given only in certain life satisfaction theories, which are not affect-based (if life satisfaction judgments are in fact understood as purely cognitive, cf. Feldman, 2010, pp. 84–85). Further, it is widely held that an actual denial would be counterintuitive (Haybron, 2020, §§ 2.2–2.3). Life satisfaction theorists have accepted this, but denied that positive life satisfaction judgments are even possible together with an unfavorable balance between negative and positive affective states (cf., e.g., Benditt, 1978). Similarly, empirical constructs for measuring happiness and well-being, such as the mentioned Subjective and Psychological Well-Being, PERMA (Peterson et al., 2005; Seligman, 2011) the Pemberton Happiness Index (Hervas & Vazquez, 2013), or the Flourish Index and Secure Flourish Indices (VanderWeele, 2017), usually include affective aspects. Given that they are apparently relevant to happiness, as well as amenable to an evolutionary-functional analysis, it seems useful to focus on positive and negative effective states for the purposes of the present review.

Defining ‘positive’ as opposed to ‘negative’ affective states, in a descriptive and non-evaluative sense of ‘positive’ and ‘negative’, is not easy (cf. Haybron, 2008, pp. 145–146). Since affective states in general seem to have multiple dimensions (cf. Drummond, 2020), ‘affective positivity’ in particular arguably also has to be a multifaceted phenomenon; the positivity or negativity of a given affective state is not constituted by simply one ‘affective dimension’ (the balance of pleasure over displeasure, say). Accordingly, for present purposes, I will adopt a pluralistic or multi-dimensional stance towards affective positivity/negativity. In correspondence to the ‘affect-based’ accounts of happiness, affective states may be considered to have a hedonic dimension, an emotional dimension, a mood dimension, or a combination of several of these dimensions. Another possibly relevant dimension concerns the quality of the experienced relationship to the world, or parts of it (persons, things, situations; as for example proposed in Rosa, 2019, cf. Höffken, 2024).

Further, it is plausible that affective positivity and negativity are not simply mutually exclusive ends on a continuum, but that they can also merge in mixed or ambivalent affective states with both positive and negative valence. This proposal of ‘Second Wave’ Positive Psychology (Lomas & Ivtzan, 2016, Lomas, 2017, cf. Delle Fave et al., 2023) has already been acknowledged and taken up in Evolutionary Positive Psychology (Nesse, 2004; Dornisch, 2022, p. 170).

Similarly, it is acknowledged that happiness-constituting affective states do not only have a biological basis, but are also culturally formed. For example, based on the distinction between evolved ‘primary emotions’ and developmentally/culturally shaped ‘secondary emotions’ (Ekman et al., 1987, Dornisch, 2022, pp. 196 − 170) suggests that we can understand affective valence on the positive-negative continuum as ‘primary’ (as well as affective arousal on the low activation-high activation continuum). On this basis, ‘secondary’ processes pertaining to how the primary states ‘are labeled, valued, and attended to are driven by sociolinguistic learning and cultural environment, even more so for emotions characterized by physiologically ambiguous or complex combinations’ (Dornisch, 2022, pp. 170, cf. Feldman Barrett, 2006, Matsumoto & Hwang, 2012).

In the following, the mentioned definitional questions concerning affective positivity/negativity are going to be set aside. Rather, the review of the extant literature in ‘Positive Evolutionary Psychology’ is going to be based on a certain requirement implied by any conception of happiness accepting positive/negative affect as happiness-constituting. Namely, if positive and negative affect are relevant for happiness, then their relative prevalence at a given time (point or interval) in a subject’s life should be considered as relevant to the subject’s happiness at that time. Thus, whatever a full account of happiness would consist in, a defining aspect¹² of it is a specific favorable balance between certain positive and certain negative affects. It can be expected that such a favorable balance would be experienced by subjects as desirable and in this sense ‘optimal’ (including the possibility of the optimum lying on an interval rather than a point, as well as different subjects having different optima). We may call this the ‘Affective States Assumption’.

Plausibly, the prevalence of positive in comparison to negative affect may not fall under a certain threshold for the overall affective balance to be favorable. It is a challenging task to make such a threshold or the favorable balance itself measurable in some precise quantitative form (in some kind of ‘positivity ratio’, cf. Larsen & Prizmic, 2008). Moreover, relevant for happiness is arguably also the temporal profile of the occurrence of positive, neutral, and negative affective states (cf. Klausen, 2019), as well as the intensity of arousal in the respective states (as modelled in the ‘circumplex map of emotion’, which is created via crossing the two dimensions valence and arousal and circularly plotting different emotional or affective states, cf. Kuppens et al., 2013). Lastly, there seems to be a dialectic ‘complementarity’ between positive and negative affect in the constitution of happiness, as suggested in Second Wave Positive Psychology (Lomas & Ivtzan, 2017, p. 1775) and through cross-cultural comparisons of conceptions of happiness, especially in an Asian context (Miyamoto & Ryff, 2011; Uchida & Ogihara, 2012). But a precise account of none of these aspects of happiness-constitution is required for the present purposes. Rather, the ‘Affective States Assumption’ is intended to express the in-principle importance of affective states for happiness (however their respective ‘positivity’ and ‘negativity’ is explicated in detail). The relevant affective states may of course also interact with cognitive, or partly cognitive, states such as judgements about life satisfaction or meaning in life, as well as harmony (Delle Fave et al., 2023, Uchida & Ogihara, 2012), attunement (Haybron, 2008), or ‘resonance’ (Rosa, 2019) with the surrounding world.

Granted that the Affective States Assumption does in fact hold, if we understand relevant factors of meeting the ‘favorable balance’, we have also gained understanding of how a necessary aspect of happiness may be realized – or missed. Positive Evolutionary Psychology, as seen in light of the Affective States Assumption,¹³ proceeds by understanding those relevant factors in explicitly evolutionary terms. The evolutionary perspective can be applied to the Affective States Assumption by inferring, from our knowledge of the human evolutionary development, hypotheses regarding which circumstances of life are correlated to a favorable balance between positive and negative affective states for human beings.¹⁴

Human evolution, natural and cultural, is the process of the development and transmission of certain traits and behaviors in individuals or groups. These traits come into being with a certain degree of randomness – by way of genetic mutation in the case of natural evolution and by way of spontaneous ideas or deliberate design, which may happen more or less randomly or intentionally, in the case of cultural evolution. In the course of evolution, some individuals and groups survive and thrive thanks to their traits and subsequently pass on these traits and behaviors through genetic inheritance or through enculturation. Other individuals and groups do not have the ‘fit’ traits to survive and thrive, and their traits and behaviors vanish with them.

Natural evolution (cf. Millstein, 2024) is (at least with regard to certain traits and timescales, cf. Wilkins & Godfrey-Smith, 2009) mainly driven by the processes of natural selection, sexual selection, and kin selection. These selection processes lead to the propagation of certain traits via genetic variation and inheritance. The evolutionary success of the bearers of these traits is measured by their ‘fitness’ or ‘adaptiveness’ in relation to the inanimate environment and to the behavior of other organisms in the environment, such as predators or prey, but also conspecifics (in terms of cooperation/competition and sexual selection).

Since all traits of an organism always come as an evolutionary ‘package deal’, the overall adaptiveness of this set of traits includes trade-offs between them. Especially, there may be competition between adaptiveness in terms of natural and sexual selection. For example, sexual selection may lead to the evolution of traits and behaviors that are costly with regards to natural selection since such ‘handicaps’ signal adaptive attributes in an ‘honest’ fashion (such as, for example, the peacock’s tail). These costly signals often do so exactly by being an obstruction to the chances of survival (Zahavi & Zahavi, 1997). This can lead to runaway evolutionary processes, a kind of overbidding through which the signal becomes ever more pronounced (and costly in terms of survival).

The expression ‘cultural evolution’ stands for the notion that our cultural traits and behaviors ‘evolve’ in a manner that is analogous to biological evolution – while ‘selection’, ‘inheritance’, and ‘fitness/adaptiveness’ mean something similar, yet different, in the cultural compared to the biological sphere. Accordingly, the selective processes of cultural evolution go beyond or sometimes even against those of biological evolution.¹⁵

To begin with, cultural transmission works differently than genetic inheritance. Transmission in cultural evolution is instantiated by social learning from other conspecifics. This goes beyond the actualization of innate, genetically encoded developmental trajectories. Further, social learning is opposed to individual or ‘asocial learning’ from one’s own experience of and interaction with the environment, without interference of others (Henrich, 2020, pp. 12–13, p. 370, note 3).

This affords an understanding of what is meant with ‘culture’ in the context of cultural evolution: it is anything – ‘practices, techniques, heuristics, tools, motivations, values, and beliefs’ (Henrich, 2016, p. 3) – that opens or constrains possibilities for certain behaviors and actions, and that can be transmitted through social learning. Accordingly, a ‘cultural species’ – with homo sapiens as the paradigm case – ‘is one that has evolved to socially transmit complex behavior-shaping information’ (Chudek et al., 2016, p. 750).

Further, the selection processes driving cultural evolution operate in a different fashion than in biological evolution. What is selected are not biological traits (phenotypes, behavioral dispositions, and the genes underlying them), but cultural ‘behavior-shaping information’, such as acquired practices, beliefs, norms, and values. For these, the selection consists in sui generis in-group and between-group dynamics (while such ‘group selection’ is usually rejected for biological evolution; Chudek et al., 2016, p. 762, cf. Wilson, 2016, pp. 335–336).

The analogy between biological and cultural evolution also includes the possibility of costly signaling and runaway selection in the cultural case (Chudek et al., 2016, p. 761). Furthermore, as with the trade-off between traits within biological evolution, there are trade-offs between biological and cultural traits and behaviors. Under the condition that it does not jeopardize its biological basis by making the ‘carrier population’ die out, the evolution of cultural traits may even run counter to biological evolution, since the genes or the biological phenotype of its bearers are not decisive for the ‘reproductive success’ of cultural traits. Individuals who are highly successful in passing on cultural behaviors and norms may still fail to leave copies of their genes, up to the point of having no reproductive success at all.

While they follow analogous (and sometimes cross-cutting) patterns, there is an important difference between natural and cultural evolution: Natural evolution is much slower. Thus, in the short term, cultural evolution unfolds within the confines of natural evolution, while in the long run, it can have feedback effects on natural evolution (Chudek et al., 2016, pp. 763–764). So, cultural and natural evolution influence one another, and indeed, they cannot be told apart from a certain point onwards in the evolution of our species. We are an ‘evolved cultural species’ (Chudek et al., 2016, p. 750), it is our nature to have culture. This process of natural-cultural co-evolution may be illustrated through the major developments that will become relevant to the discussion of evolutionary conditions for positive and negative affect.

Beginning at least 5 million years ago, after leaving the trees in the rainforest as their preferred habitat, hominins faced new environmental conditions on the African savanna: better opportunities for social learning, more diverse and fluctuating environments, and increased danger from predators. These environmental pressures and opportunities created an evolutionary pathway with mutually reinforcing factors, including upright walk (and free hands), brain growth, intensified tool use, diversified diet (especially big game hunting), use of fire (especially for cooking food), increased cooperation in larger groups, and pair-bonding strategies in mating (Henrich, 2016, pp. 298–312).

Cultural evolution plausibly has had a decisive influence on our nature as social beings (apart from other factors such as kin selection, reciprocity, and mutuality in in-group cooperation). Analogies to great apes make it plausible that the social life of chimpanzee-like human ancestors was initially characterized by routine behavior of competition and dominance, with only occasional incidents of cooperation. But at some point, the described co-evolutionary processes made social cooperation an ecological success factor to the degree that prosocial norms, increasing in-group cooperation and harmony were selected for (Richerson et al., 2016; Henrich, 2016, p. 167).¹⁶ Such norms harness the genetically encoded social instincts and can have feedback effects on the biological evolution of the human mind, catalyzing a process of ‘self-domestication’ (Wrangham, 2019) and further entrenching a norm psychology geared towards the internalization of cooperative behavior (Henrich, 2016, Chap. 11).

Especially, as the most adaptive form of social group organization, a so-called ‘reverse dominance hierarchy’ (Boehm, 2001) was culturally selected for in the ecological circumstances of our ancestors’ African habitat (the so-called ‘environment of evolutionary adaptedness’). This concept describes social constellations in which the group dominates individuals striving for status at the expense of others. In line with this hypothesis, extant hunter-gatherers have even been characterized in ethnography as being ‘fiercely egalitarian’ (Lee, 1979, p. 24), they defend the equal distribution of access to resources and prestige with conscious effort and vigor (including unparalleled gender equality; cf. Kelly, 2013, pp. 243–248).

The hunter-gatherer lifestyle underlying egalitarianism was (and is, for the still-remaining populations) typically nomadic and characterized by an ‘immediate-return’ subsistence system in which food is consumed shortly after procurement (Woodburn, 1980, 1982). Accumulation of material resources (in the form of useful or valued goods, or food stocks) was practically impossible. The correlated mode of social resource distribution is wide and tendentially equal sharing, a kind of ‘insurance system’ to even out resource fluctuations (Kelly, 2013, Chap. 6).

Moreover, the nomadic lifestyle rendered ‘fission and fusion’ the default mechanism to deal with social tensions: When insurmountable conflicts arise in a group, some of the members choose to leave and join other bands to which they have ties (Couzin & Laidre, 2009).

Apart from the group an individual is staying with at the moment, the larger social structure she is embedded in is the ‘extensive kinship network’. Such networks are established along biological relatedness and other social ties, such as fictive kinship, and span multiple smaller groups of flexible composition (Henrich, 2020, pp. 71–72).

Finally, the cultural-evolutionary adaptation package of immediate-return hunter-gatherers includes a strong emphasis on personal autonomy (Gardner, 1991). This fits with the egalitarian ethos and the necessity to self-sufficiently navigate an ‘open landscape’ – both in the physical and the social sense. In effect, the only populations on earth valuing individual autonomy as high as persons with a ‘Western’ cultural background are immediate-return hunter-gatherers (Henrich, 2020, p. 224).

All in all, the deeply-rooted drive for dominance and competition, together with the shift to a reverse dominance hierarchy has endowed us with an ambivalent evolutionary heritage. This shows most markedly with the transition to a ‘delayed-return’ system (in which a considerable time lies between the efforts to attain resources and their consumption, cf. Woodburn, 1980, 1982) and a sedentary lifestyle, which spread around the world with the advent of agriculture.

Agriculture changed the cultural fitness calculation both within and between groups, effectively uprooting the core aspects of immediate-return hunter-gatherer culture: The dependence on crop cycles and farming grounds favored ‘intensive kinship’ institutions organized around bloodlines, clans, kindreds, and religious congregations (instead of extensive social networks with fluctuating group composition). This shift stabilized group composition and cohesion as well as long-term mutual commitments among the members (Henrich, 2020, pp. 96–112). Intensive kin-based institutions are typically associated with steeper hierarchies and less personal autonomy. In addition, agriculture allowed dominance behavior, to which especially men are disposed due to their great ape animal nature, to spread (von Hippel, 2018, Chap. 3). As a matter of economic necessity, agriculture comes with stockpiling and territoriality. This opened the pathway to increasing inequality in power and resource distribution (including pronounced gender differences) through the (now-expedient) accumulation of resources, prestige-based leadership, or simply more organized forms of violence and coercion, and control over labor (Mattison et al., 2016).

The agrarian conditions of life favored these ‘cultural adaptations’ both through both within-group and between-group selection. Within groups, the individuals complying with dominance (as dominators, but also as subordinates, or fellow travelers) were in the long run more successful in reproducing it than individuals opposing it. In comparison between groups, more hierarchical ones were more efficient in the organization of agricultural production and military action than less hierarchical ones (especially, the hunter-gatherer communities coexisting with agriculturalists), thus effectively crowding them out (Mattison et al., 2016, pp. 193–195).

Clans and kindreds, together with intensifying functional specialization, laid the basis for the development of chiefdoms and premodern states (Henrich, 2020, pp. 112–120). The transition to modern states, impersonal markets, and industrialized mass societies, in turn, as has been suggested by Henrich (2020), was facilitated by the dismantling of intensive kin-based institutions and norms. The effect was, at core, a shift from ‘interpersonal’ (mainly kin-based) to ‘impersonal prosociality’. In a gradual process spanning hundreds of years, clan structures were largely replaced by more and more nuclearized families. The socialization in more nuclearized families (instead of intensive kinship networks) fueled a self-reinforcing cultural-evolutionary process with the development of a psychological profile characterized by the explicit or implicit adherence to individualism, analytic thinking, anonymous norms and principles, and openness to relative strangers as well as to ‘voluntary associations’ (which, unlike kinship institutions, individuals can join voluntarily; examples are monasteries, charter towns, guilds, universities, and companies).

Evolutionary thinking is always a complex inference to the best explanation, with all the concomitant uncertainties. We will probably never know with certainty which evolutionary developments ultimately account for what portion of the success of our species and of particular cultures. But there have to be some success factors, otherwise homo sapiens would not be around anymore and certain cultures would not have gained the dominant role they have.

What is also certain, though, is that whatever the crucial success factors were and are, their being success factors in evolutionary terms ultimately lies solely in their contribution to the reproduction of certain traits, not in the fact that the bearers of these traits have a favorable balance between positive and negative affect.

Concerning the relationship of natural evolution to positive and negative affective states, a basic question is why such states evolved in the first place. The answer would surely be complex (cf. Dornisch, 2022, for an overview of theoretical and empirical approaches), and an important caveat in this regard is that affective states never serve only one specific function (Nesse, 2004, pp. 1337–1338).

Nonetheless, many evolved capacities – and especially the capacity to experience certain affective states under certain circumstances – do not simply have one specific evolved function, but also more complex and derivative adaptive effects. A theory that proceeds on basis of this insight is Broaden-and-Build Theory (Fredrickson, 2008), which conceives of positive affect, such as joy, gratitude, contentment, interest, hope, pride, amusement, inspiration, and love (Fredrickson, 2008, pp. 3–6), not simply as being part of a reward/incentivizing mechanism. Thus, while negative affective states serve a short-term purpose in motivating the subject to face imminent challenges (through fight or flight, say), or to work towards a change of the situation if things are not going well (in fitness-terms), positive affective states have developed more long-term ‘side’ effects. Positive affect allows for a broadened awareness when occurring, opening possibilities for discovery of new knowledge, personal relationships, and skills – in short: of new fitness-relevant resources. Resources built through positive emotions also increase the chances of experiencing subsequent positive emotions, with further broaden-and-build-benefits, thus creating an upward spiral toward improved fitness (Fredrickson, 2013, pp. 15–17). So, the importance of positive emotions is hypothesized to have amplified, over and above its basic incentivizing function, through a self-reinforcing (‘runaway’) evolutionary process.

The runaway selection for positive affective states suggested by Broaden-and-Build theory is plausibly further amplified in the context of cultural evolution. While the subjective components of affective states appear as mere ‘psychological carrots and sticks’ from the vantage point of natural evolution, and not ‘as being good or bad, per se’ (Hill et al., 2013, p. 876), from the perspective of the experiencing individual, they are exactly this. Accordingly, Kováč (2012, p. 298) suggests that the conscious orientation towards affective states has long been the primary driver of cultural evolution (in a runaway process, as seen from the lens of natural evolution) – which is thus to a large extent an ‘emotional evolution’.

Moreover, already on a more basic level, it may be expected with Broaden-and-Build theory that natural evolution has already given us, on average, the maximum inclination towards a favorable balance between positive and negative affective states that could possibly be afforded without inhibiting their incentivizing function. Namely, as long as an inclination to such a favorable balance is not as strong as preventing its bearer from adaptive behavior, it is of help for realizing such behavior. A favorable affective balance seems to be correlated with optimism and openness to novelty (Fredrickson, 2008, pp. 455–462), which is helpful for acquiring fitness-relevant resources. Accordingly, we may hypothesize that in the absence of adverse conditions, humans tend to be in a state of ‘default contentment’ (Grinde, 2012, p. 7), of low-arousal positive affect.

But in general, the role of affective states as ‘psychological carrots and sticks’ (Hill et al., 2013, p. 876) makes them, their existential importance for happiness notwithstanding, only means to an end (Nesse, 2004, pp. 1335–1336; Grinde, 2016, p. 177; von Hippel, 2018, p. 217). For example, humans have evolutionarily deeply ingrained concerns with relative social status and status competition (‘positional bias’; Hill & Buss, 2008, pp. 64–66). These have been a potent driver of unhappiness under pre-historical, historical, and modern conditions – although they play out in different forms in the respective contexts. Another challenge for the pursuit of happiness may lie in the structure of the motivational states themselves, as Nettle (2006) points out. Nettle distinguishes between ‘liking’ – the actual enjoyment of positive affect (which he relates to the opioid system in the brain) – and ‘wanting’ – the desire for whatever is expected to bring about positive affect (which he relates to the dopamine system, cf. also Esch, 2022). The challenge is that it may well be adaptive for humans to expect positive affect of certain things (such as high social status, for example), even if these fail to deliver the expected degree of positive affect. In short, ‘evolution hasn’t set us up for the attainment of happiness, merely its pursuit’ (Nettle, 2006, p. 166) – making the relentless pursuit of happiness ‘mirages’ a predictable phenomenon (Nettle, 2006, Chap. 5).

Similar to natural evolution, cultural evolution as a selection process is indifferent, and often detrimental, to the quality of life of its subjects. The transition to agriculture, for instance, created disadvantageous conditions for health, such as field work, crowded living, and a one-sided diet, and initiated severe social stratification and gender inequality (von Hippel, 2018, Chap. 3).

The conclusion that may be drawn from the considerations so far is that if happiness grows out of the selective process of (natural and cultural) evolution, then it does so only as ‘an accidental by-product’ (Grinde, 2016, p. 177). However, this mere ‘by-product’ is exactly what humans, as conscious beings, are able to set for themselves as a goal. In this vein, Grinde holds:

We can extend Grinde’s appeal also to cultural evolutionary objectives. For example, just as we may choose to ‘give a damn’ about the pressures exerted by the mechanics of sexual selection, we may choose to do so with regard to inherited cultural norms (although neither the one nor the other may be easy).

Dealing with evolutionary objectives in pursuit of happiness (or, with the Affective States Assumption, a favorable balance of positive and negative affect as a necessary condition for happiness) presupposes knowledge of these evolutionary objectives and factors. Only on this basis is it possible to consciously engage with evolutionary influences. This may be considered the central idea in the approach of ‘Positive Evolutionary Psychology’.

The review of Positive Evolutionary Psychology provided here may serve to illustrate that an account of the evolutionary conditions of (positive or negative) affective states can deepen our understanding of human happiness.

Thus, we may attain, if not an exhaustive account of what constitutes happiness (which may go beyond a favorable balance between positive and negative affect), then at least useful advice concerning central aspects of happiness and important roots of unhappiness (Nesse, 2004, p. 1337).

An optimistic conclusion to be drawn is that if we can avoid the pitfalls of unhappiness, then happiness may be a quite expectable outcome. In the absence of adverse conditions, default contentment is to be expected, it does not have to (and possibly cannot) be intentionally produced. A pessimistic conclusion is that there are so many pitfalls, many of which are, ironically, created by supernormally amplified and phenotypically indulgent promises of happiness - pursuit of those may lead to a particularly unfavorable imbalance of negative and positive affect (leading to an evolutionary version of the ‘paradox of happiness’, cf. Nesse, 2004, p. 1343). Moreover, results from evolutionary anthropology and psychology suggest that concerns with status, wealth, and consumption have strong evolutionary underpinnings, while at the same time leading to cognitive overload, addictive dynamics, and estrangement from deeply ingrained needs for belonging and cooperation.

Beyond these conclusions, there are still open questions with regard to the conceptual basis and possible practical impact of the approach described here. One open question concerns the relative importance of the sketched dimensions of affective positivity/negativity, which is contentious (cf. Haybron, 2008). This conceptual challenge affects Positive Evolutionary Psychology just as much as other approaches in happiness research. But it seems that Positive Evolutionary Psychology can make a contribution in this regard: Similar to the predictions on evolutionary ‘imperatives’ concerning sources of positive/negative affect, and complementing ‘proximate’-functional accounts, we may devise evolutionary hypotheses on the adaptive relevance of the different dimensions of positivity/negativity, and consequently make predictions about their respective relevance to the overall affective condition.

But so far, there seem to have been few efforts to ground Positive Evolutionary Psychology on a clearer conception of positive and negative affect and their respective contributions to happiness. There is not even an explicit distinction between the questions of well-being and happiness, as differentiated here, in the Positive Evolutionary Psychology literature.²³Another question concerns the interplay of affect and cognition as happiness-constituting factors. Giving an account of cognition in parallel to the one provided here for affect from an evolutionary perspective can be expected to be significantly more complex, taking into account the pronounced developmental plasticity and individual as well as cultural variation with respect to cognition. Especially relevant in this respect are the different cultural ‘styles’ in conceptualizing happiness, such as more or less pronounced ‘dialectical emotional styles’ (Miyamoto & Ryff, 2011) or diverging ‘interpersonal’ and ‘individualistic’ conceptions of happiness (Uchida & Ogihara, 2012). This opens a vast field for cultural-evolutionary research. For example, Uchida and Ogihara (2012, pp. 357–358) hypothesize that a cultural setting characterized by an agricultural subsistence style and low social relationship-mobility favors an interpersonal conception of happiness.

When drawing practical conclusions from the described enabling or inhibiting factors for realizing a favorable balance between positive and negative affective states (as a necessary condition for happiness, according to the Affective States Assumption) in modern societies, Positive Evolutionary Psychology has mainly focused on the level of individual, as opposed to collective, circumstances of life (Positive Evolutionary Psychology has pursued a more ‘individualist’ than ‘contextualist’ approach to happiness, cf. Haybron, 2008, pp. 263–264).

Nonetheless, widening the focus of the evolutionary perspective on happiness from the individual to the collective level seems to be a realistic future possibility. This may yield important implications not only for the pursuit of happiness, but also, and relatedly, for pressing societal and political questions concerning the transition to more equitable resource distributions and more sustainable ways of life. Whatever an answer to the current social, political, and ecological challenges may consist in, it will likely have to include a foregoing of excessive consumption and status competition by privileged classes and individuals (both on a global scale and within societies). But argumentation in favor of this cause is often obstructed by attitude-behavior gaps and rationalizations of cognitive dissonance.

Evolutionary reasoning may both help us to better understand and to counter such challenges: An awareness of natural and cultural human evolution can elucidate reasons why the hedonic lures of a modern world as well as status concerns and conspicuous consumption make it so hard to promote sufficient and sustainable lifestyles. But instead of leading into determinism (since it is part of our nature to be susceptible to hedonic objectives and status concerns), this approach invites us to actively engage with evolutionary pressures and path dependencies – and to go beyond them, where necessary (a capacity that is part of our nature as well). Thus, understanding ourselves as evolved beings may support a conscious distancing from tendencies for excessive status and pleasure seeking based in natural and cultural evolution, unfolding new motivational potentials and complementing more normatively inclined appeals. Overall, a broader dissemination of knowledge about the evolutionary influences on our behavior may itself become a factor in cultural evolution – potentially, and hopefully, moving humanity towards greater ecological and social sustainability.