The Genetic Basis of Altruism and Cooperative Behaviour in Human Societies

 

Muhammad Muzammal 1*, Abida Bibi 1, Maria Shafiq 2, Hunza Malik 1, Shafiqua Istiaq 1,

Sabeeha Asad 1, Harris Khan 1, Harmain Saba 3,

Umar Raoon 1, Muhammad Ismail 1, Nazia Farid Burki 1, Hamna Batool Hashmi 1, Hadia Gul 4 and Muzammil Ahmad Khan 1

1 Gomal Centre of Biochemistry and Biotechnology, Gomal University, Dera Ismail Khan, Pakistan

2 Gomal Medical College, Dera Ismail Khan, Pakistan

3 Government Degree College no 1, Dera Ismail Khan, Pakistan

4 Institute of Biological Sciences, Gomal University, Dera Ismail Khan, Pakistan

* Corresponding author

ABSTRACT

Altruism and cooperative behaviour are selected characteristics of societies that range from private to public ones. Though such commu- nal sacrificing seems to go as far as being not in congruence with the principle of natural selection, evolutionary science offers some expla- nations about how altruism persists. Theories like kinship selection, reciprocal altruism, and group selection suggest that prosocial behav- iours develop because they have had evolutionary benefits in terms of survival of genes and group cohesion. Advancement in genetics and

Recommended citation: Muzammal M. et al. The Genetic Basis of Altruism and Cooperative Behaviour in Human Societies. Social Evolution & History, Vol. 24 No. 2, September 2025, pp. 103–117. DOI: 10.30884/seh/2025.02.05.

© 2025 ‘Uchitel’ Publishing House

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neuroscience enables showing the biological basis by which altruism has specific genes and neurochemical pathways’ effects on empathy, trust, and co-operation. Studying oxytocin, dopamine, serotonin, and the MAOA gene bring the genetics of altruistic behaviour into focus, and epigenetic research shows how environmental factors shape altru- istic tendencies. However, altruism is not only a product of biology; but also cultural, social, and economic factors go along in reinforcing or curtailing cooperative behaviour, and altruism within societies is well maintained by moral systems, social norms, and institutional frame- works, but at the same time it is increasingly threatened with new types of challenges – like economic inequality, anonymity on the digital platform, and global collective action problems – confronted in the mo- dern era. The essay concentrates on the genetic, evolutionary, and cul- tural foundations of altruism, emphasizing how biological predisposi- tions interact with environmental influences to mold human coopera- tion. Such an understanding will enable us to develop strategies that could enhance cooperation toward the response to some of these con- temporary global dilemmas.

Keywords: genetic, evolutionary, cultural, foster.

• INTRODUCTION

Altruism and cooperation form important categories of human social life, dictating interpersonal relations between individuals, families, and whole societies. These can range from the smallest acts, like shar- ing food or helping a stranger, to some of the largest-scale humanitari- an efforts. Humans build a life of prosocial acts in ways that benefit others, sometimes at their expense. In this regard, one question be- comes interesting: if natural selection favors traits underlying individ- ual survival and reproductive success, why is it so often that humans act against such biological inclinations? (Alexander 2017)

The answer to why we have developed altruism has historically interested biologists, psychologists, and anthropologists. If one accepts classic Darwinism, it should be clear that Darwin stated that individu- als acted in a way that is beneficial for them. There is a trade-off be- tween costs and benefits. Darwin realized that in social species, indi- viduals cooperate with each other, mothers risk protecting their off- spring, etc. More recent frameworks of evolutionary theory have sug- gested that cooperative behaviour can be an adaptive benefit in certain instances. Kin selection, reciprocal altruism, and group selection theo- ries support the notion that cooperation and altruism have survived evolution because they promote the survival of genes on an individual

level, as well as the family, social group, and population levels (Api- cella et al. 2012).

After the introduction of genetics and neuroscience, some re- searchers have ventured into the biological basis of prosocial behav- iour. Genetics of the twins, neurotransmitters, and hormonal influ- ences have proven that the propensities towards altruism, in very gen- eral terms, have been mediated by particular genetic and neurobiologi- cal factors in a person. Thus, genes regulating the synthesis and release of oxytocin, dopamine, and serotonin play a crucial role in determining how humans’ bond with others, develop empathy, and cooperate. One of the many discoveries of recent epigenetic studies is that environmen- tal factors, what one experiences in their youth, the surrounding so- cialization mechanisms, and cultural training, influence the expression of genes correlated with altruistic tendencies (Bowles 2006; Butov- skaya et al. 2020a, 2022; Yadav et al. 2024).

Altruistic behaviour is evolved genetically, but is by no means naturally obligatory. Different factors – social, economic, and techno- logical – can have diverse influences on human behaviour, some con- tributing to prosociality, others fostering selfishness or competition. For example, economic inequality, ambiguous identity on the internet, and large collective action dilemmas such as climate change challenge altruism in modern society. Understanding the genetic basis of altruism and cooperation is of more than scientific interest. It is increasingly required in practical terms as the world faces problems of such as scale and complexity that the levels of cooperation required will be unprec- edented. Such research can illuminate the biological and cultural foun- dations of prosocial behaviour and, hence, move toward a more com- passionate and cooperative world (Krebs 2011; Darwin 2023). This review will discuss the evolution and adaptive advantages of altruism, genetics and neurobiology of cooperation, cultural and environmental influences, and the major challenges facing altruism in contemporary society. Through this lens, we may gain a better understanding of hu- man cooperation and how to encourage pro-social behaviour for the benefit of individual and societal welfare (Bowles 2006; Clutton- Brock 2009).

• THE EVOLUTIONARY FOUNDATIONS OF ALTRUISM AND COOPERATION

The evolutionary roots of altruism have been proposed to explain why organisms, including humans, sometimes cooperate instead of com-

peting with one another. This behaviour is understandable and forms part of the most important theories regarding prosocial-human behav- iour persistence. Key theories are shown in Table 1.

Table 1

Key Theories Explaining Altruism and Cooperation

Theory	Proponent(s)	Core Principle	Examples in Nature & Human Society
Kin Selection	William D. Hamilton (1964)	Altruism is directed toward genetically related individuals to enhance shared gene survival	Parental care, family support, eusocial in- sects (bees, ants)
Reciprocal Altruism	Robert Trivers (1971)	Individuals help non- kin with the expecta- tion of future recipro- cation	Food sharing in pri- mates, cooperative hunting, business transactions
Group Selec- tion	Darwin, E.O. Wilson	Groups with coopera- tive individuals outper- form less cooperative groups	Religious communi- ties, cooperative mili- tary units, collective farming
Strong Reci- procity	Fehr & Fisch- bacher (2003)	Humans enforce coop- eration by punishing cheaters, even at a personal cost	Legal systems, whis- tleblowing, social ostracization of rule- breakers
Cultural Evolution	Richerson & Boyd (2005)	Altruism is reinforced by learned norms and cultural traditions	Charitable organiza- tions, ethical systems, moral education

Note: Trivers 1971; Fehr et al. 2002a; Lieberman et al. 2007; Leigh 2010; Birch and Okasha 2015; Birch and Heyes 2021.

Kin Selection and Hamilton’s Rule

One of the earliest explanations for altruism, which is still widely ac- cepted, is kin selection, which was proposed by the British evolution- ary biologist William Hamilton in the 1960. Hamilton’s rule states that altruistic acts evolve under conditions when:

rB > C

Where:

• r stands for the genetic relatedness between individuals,
• B is the reproductive benefit to be realized by the recipient, and
• C is the cost borne by the altruist at the very instance.

This can explain why people are more likely to help close kin than those who are unrelated. Those relatives share incredibly high propor- tions of genes, so the survival and reproduction of relatives indirectly benefit from the altruistic actions of their genetic lineage (Emlen 2001).

Some natural examples include eusocial insect species such as bees and ants, where sterile worker castes renounce their own repro- ductive potential to serve their queen and colony of genetically related individuals. Among mammals, including humans, there are investments in children who can be too expensive even for parents because they preserve their genetic inputs through avenues of offspring (Abid et al. 2022; Ali et al. 2022; Hussain et al. 2022; J. Alsalman et al. 2022a; N. Alhashem et al. 2022). Kin selection among humans is found in fami- ly support, inheritance, and caregiving activities (Hussain et al. 2022). For example, people are generally more willing to perform more for their children, siblings, or even close relatives than other people, which indicates a cross-cultural and cross-generational access pattern (The age of empathy: nature’s lessons for a kinder society, 2010).

• RECIPROCAL ALTRUISM: COOPERATION BEYOND KIN

Kin selection explains why individuals help their relatives, but not cooperation among unrelated individuals. According to Robert Triv- ers’ reciprocal altruism theory of 1971, cooperation can evolve when individuals are helped with the expectation that something will be re- turned in the future (Trivers 1971; Muzammal et al. 2019; Gul et al. 2021; J. Alsalman et al. 2022b; Khan et al. 2022; Mohaini et al. 2022a; Muzammal et al. 2022; Ahmad et al. 2023; Ayaz et al. 2023).

• RECIPROCAL ALTRUISM KEY PRINCIPLES:
• They should meet each other at least many times in future to form a possible reciprocity.
• It is necessary to identify cheaters and punish them for accept- ing without returning favors (Trivers 1971; Emlen 2001; Leigh 2010; Gardner et al. 2011; Bourke 2014).
• Cooperation must have a positive and greater payoff over the long-term than the cost incurred through cooperation over the short-haul.

For example, human and animal behaviours include:

• Consider chimpanzees observed to groom non-relatives and later return that by sharing food,

• In hunter-gather societies, food sharing is an activity where everyone contributes to the very group during a time of plenty and then expects help from that group during times of short- age.
  • It has spread its roots into the most modern economies, and in principle, it is considered a positive factor in most modern economic theories: trust and mutuality in economic transac- tions (Birch and Okasha 2015).
• GROUP SELECTION AND CULTURAL EVOLUTION

Co-operation benefits not only individuals but also society as a whole. The theory of group selection suggests that cooperative and altruistic groups are more likely to outcompete less cooperative groups. Geog- raphers were previously discontent over the idea of group selection, but have gradually come to accept the fact that cultural evolution in- deed plays a significant role in promoting prosocial behaviour (Cohen 2013; Author 2013).

• Examples of cultural reinforcement of cooperation:
• Religious and moral systems encourage altruism and often frame prosocial behaviour as a moral duty. Many religions promote charity, kindness, and community service.
• Legal and social institutions enforce cooperation through laws, contracts, and norms that discourage selfish behaviour (e.g., tax- funded welfare programs, anti-fraud regulations).
• Nationalism and collective identity foster cooperation on a large scale, as people tend to act altruistically toward those they per- ceive as part of their ‘in-group.’
• THE GENETIC BASIS OF ALTRUISTIC AND COOPERATIVE BEHAVIOUR

Genetics and neuroscience have advanced to the point where they re- veal that altruism is not simply acquired behaviour, but rather a bio- logical phenomenon. Various genetic, hormonal, and neural mecha- nisms have been linked to prosocial tendencies in individuals.

• Twin Studies and Heritability of Altruism

Twin studies have been conducted by behavioural geneticists to esti- mate the heritability of altruism and cooperative behaviour. Identical

twins have a greater resemblance in their prosocial behaviours than fraternal twins, and the latter share only half of their genetic material. These studies have put forward evidence that genetics accounts for approximately 30 % to 50 % of altruistic tendencies while environ- mental and cultural factors account for the rest (West et al. 2007).

• Key Genes Associated with Altruism and Cooperation

Some gene candidates have been implicated in prosocial behaviour:

• Oxytocin and Vasopressin Receptor Genes (OXTR & AVPR1A)

Oxytocin, the so-called ‘love hormone,’ determines how much social bonding, trust, and empathy humans have. Variations in the OXTR gene are linked to differences in generosity and emotional intelligence. In addition, some studies have shown that carriers of the OXTR rs53576 GG variant display altruism significantly more frequently, both to- wards friends and strangers (Butovskaya et al. 2020b). The AVPR1A gene is responsible for regulating the vasopressin system which is re- sponsible for social bonding. The vasopressin system is responsible for social bonding and cooperativeness, especially among men (Nowak et al. 2010; Muzammal et al. 2021; Stonerook 2021; Ahmad et al. 2022).

• Dopamine and serotonin pathways

Prosocial behaviour is reinforced by the dopamine system through the attachment of positive emotions and rewards to such acts. Therefore, any generous act will result in the release of dopamine in the brain and hence cause a pleasurable feeling. Higher serotonin levels are associ- ated with patience, fairness, and cooperative decision-making. The sero- tonin system regulates mood and impulse control (Smortchkova 2017; Mohaini et al. 2022b; Ahmed et al. 2024).

• MAOA (‘Warrior Gene’)

Gene MAOA is thought to influence aggression and social behaviour. Some variations in the gene are associated with increased aggression and reduced empathy, but environmental factors still play a huge role in their expression (Golya 2005).

• EPIGENETICS: THE ENVIRONMENT SLIGHTLY AF- FECTS GENETIC EXPRESSION

Epigenetics indicates that life experiences can affect gene expression without changing DNA sequences. Some examples include childhood experiences: children raised in a nurturing environment develop strong- er prosocial tendencies, while those raised in neglectful conditions show low levels of trust and empathy (Ramsay 2005). Stress and adversity: stress can inhibit oxytocin production, which can impede bonding to social networks. The genetic and neurobiological basis of altruism is shown in Table 2.

Table 2

Genetic and Neurobiological Basis of Altruism

Gene/Hormone	Function	Impact on Altruism & Cooperation	Empirical Findings
OXTR (Oxytocin Receptor Gene)	Regulates oxyto- cin, the ‘bonding hormone’	Enhances trust, empathy, and so- cial bonding	Variations in OXTR influence generosity and emotional intelli- gence
AVPR1A (Vaso- pressin Receptor Gene)	Regulates vaso- pressin, involved in social behaviour	Affects bonding, particularly in males	Associated with pair bonding in mammals and cooperation in humans
Dopamine (D4R, DRD4 gene)	Reward system neurotransmitter	Reinforces proso- cial behaviours through pleasure and reward	Higher dopamine activity linked to generosity and fair decision-making
Serotonin (5-HTTLPR gene)	Regulates mood and impulse con- trol	Promotes patience, fairness, and co- operative behav- iour	Increased seroto- nin associated with prosocial choices in eco- nomic games
MAOA (‘Warrior Gene’)	Regulates aggres- sion and emotional control	Certain variants linked to reduced empathy and in- creased aggression	Low MAOA linked to antisocial behaviour, espe- cially under stress

Evolutionary benefits of altruism are shown in Table 3.

Evolutionary Benefits of Altruism

<br>

Table 3

Benefit	Mechanism	Example
Enhanced Sur- vival of Kin	Kin selection ensures shared genetic material is passed on	Parents invest in children, siblings support each other
Reputation & Social Status	Altruistic individuals are trusted and respected, increas- ing social capital	Philanthropy enhances public image; generous leaders gain loyalty
Reciprocal Ben- efits	Helping others increases chances of future support	Business partnerships, alli- ances in warfare
Group Success & Stability	Cooperative groups outper- form selfish groups in compe- tition	Nation-building, teamwork in corporations
Reduction of Conflict	Altruism fosters social har- mony and reduces aggression	Mediation in disputes, legal systems promoting fairness

• OBSTACLES TO ALTRUISM IN TODAY’S WORLD

However, the modern world presents a series of challenges to altruism in general, despite its evolutionary and genetic underpinnings.

• Economic inequalities and social fragmentation

Enormous disparities in wealth can erode trust and cooperation within societies, leading to less prosocial behaviour. Studies have shown that people in highly unequal societies are less likely to give or support charitable causes (Golya 2005; Ramsay 2005; Smortchkova 2017).

• The Digital Era and Online Behaviour

Online social media creates a phenomenon known as ‘moral outrage amplification,’ in which people participate in performative versions of altruism, such as virtue signaling, instead of cooperating to achieve meaningful actions (Fehr et al. 2002b; Birch and Heyes 2021). Digital anonymity tends to diffuse responsibility, thus creating more hostile interactions and less cooperation online (Santos and Pacheco 2011a).

• Global Collective Action Problems

Climate change, pandemics, and international conflicts all require the parable of large-scale cooperation, yet many people and nations stand in the way and choose self-interest, which frustratingly interferes with

the optimality of joint solutions (Seyfarth and Cheney 1988; Fehr et al. 2002b; Santos and Pacheco 2011a, 2011b; Author 2013; Claidie et al. 2014; A natural history of human morality 2016; Tomasello 2018). Challenges to altruism in modern society are shown in Table 4.

Table 4

Challenges to Altruism in Modern Society

Challenge	Cause	Impact on Altruism	Possible Solutions
Economic Inequality	Wealth disparities reduce trust and cooperation	Less charitable giving, social fragmentation	Progressive taxa- tion, social safety nets
Digital Anonymity	Online interactions lack accountability	Increased trolling, cyberbullying, reduced empathy	Stronger regula- tions on social media behaviour, digital identity verification
Global Collective Action Problems	Climate change, pandemics require large-scale coop- eration	Nations act in self- interest, leading to failures in global solutions	International trea- ties, incentives for cooperation
Cultural & Politi- cal Polarization	Ideological divi- sions weaken so- cial cohesion	Reduced willing- ness to help out- groups	Promoting dia- logue, education on shared values
Declining Face-to- Face Interaction	Virtual communi- cation replaces real-world interac- tions	Reduced emotion- al connection, lower empathy	Community- building initia- tives, encouraging in-person en- gagement

• CONCLUSION

The genetic basis of altruism and cooperation is a topic of great fasci- nation and complexity, invoking a tangled interplay between biologi- cal inheritance and environmental factors. Genes provide the founda- tion for prosocial behaviour, while cultural evolution, social norms, and individual experiences determine its expression. Cooperation is at higher demand than ever in today’s world, with pressing solutions re- quired to rectify economic inequality, digital isolation, and the chal- lenges of collective action on a global scale. With an understanding of the biological and cultural roots of altruism, we will be able to make significant progress toward a more cooperative and compassionate

world. Figure showing the Altruism and Cooperative behaviour in Human Societies.

Fig. Representation of Altruism and Cooperative behaviour in Human Societies

Note: Aoki 2017; BioRender 2023.

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