Can we behave altruistically?


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IT is evident that if we hope to mitigate the effects of the environmental catastrophe that is already upon us, we must cooperate with each other. In other words, we will need to take actions that are for the collective good. Inevitably, that will mean acting selflessly, which means acting in a manner that may leave at least some of us (the actor included) worse off than before. Actions that are directed at the well-being of others to the detriment of one’s own interests are said to be altruistic and the behaviour is referred to as altruism.1

The specie to which we belong, Homo sapiens, is the product of a long evolutionary history. Our physical traits have been shaped by evolution. In addition, through the intermediary of evolution, our behavioural patterns too display responses to the experiences of our non-human ancestors. To state it in the most general terms, human culture has biological roots.2 One aim of this essay is to point out that similarities to the form of behaviour known as altruism exist in other organisms; in fact, their study has engaged biologists for a great many years. By seeing how the issue of altruism in non-human animals has been analyzed, we may obtain insights that throw light on the link between the evolutionary roots of human behaviour and environmental degradation. A second aim is to ask, does behaving in the manner required for maintaining a high quality of environment come naturally?

It is widely acknowledged that our environment is deteriorating. Also, it is generally agreed that the reason for this is the behaviour of us humans. Vikram Soni’s book Naturally gives vivid examples of how humans degrade the quality of the places they inhabit, the food they eat and the very climate of the world.3 It shows the price that is being paid, already, for human misbehaviour. It also draws attention to a number of reasons why we behave as we do. Among them are short-sightedness and a lack of attention to long-term consequences. Both lead to actions that offer immediate benefits to some but jeopardize the future of the many. Soni draws the conclusion that those of us who belong to this generation must cooperate with others if the planet that we bequeath to our descendants is to be at least as liveable as the one we inherited from our ancestors. My view is that what some of us are called upon to do is more difficult than mere cooperation. We need to perform actions that not only help others, but also harm our own interests.4 However, the instinctive tendency to keep one’s individual interests foremost inevitably comes in the way of doing something for the good of the group. That is especially true when doing good to others entails a cost and ‘being good to oneself’ does not.


The essential point was brought out starkly by Garrett Hardin in a classic essay titled ‘The Tragedy of the Commons’.5 It is central to the arguments developed in ‘Naturally’ and remains worth reading even today. A more homely version of Hardin’s thesis comes to us as a well known folktale involving the emperor Akbar and his adviser Birbal. When Akbar began boasting that his subjects followed the highest ethical standards of behaviour, Birbal countered that not one of them would act honestly, if it was to their advantage to be dishonest, and there was a good chance of escaping detection. Akbar objected angrily. To settle the issue, Birbal designed a test. A lake bed was dug and Akbar ordered that each citizen pour one glass of milk into it at night and create something wonderful – a lake of milk. Everyone agreed to obey the command.

To the emperor’s shock, instead of the expected lake of milk, there was a lake of water to be seen the next morning. Birbal’s explanation was simple. Each citizen had reasoned identically: ‘Since everyone else is likely to contribute milk anyway, it won’t be noticed if I alone substitute water for milk and save myself some money. After all, even if most of us contribute milk, suppose someone else happens to pour a glass of water, it will not be noticed. Rather, the person who does so will both get away with it and gain financially relative to me.’ Akbar duly acknowledged defeat. We do not know whether he also drew the conclusion that Hardin did some 400 years later: a social system in which it is possible to gain the benefits of others’ cooperation without incurring the cost of cooperating oneself, is inherently unstable. In the somewhat unfortunate terminology that has become part of contemporary biology, one would say that whereas Akbar expected his subjects to behave altruistically, they knew that cheating, and behaving selfishly, was the rational and superior option.


Thus there are two difficulties with being an altruist. The first is that if you perform an altruistic act, you pay a cost but derive no benefit – indeed the benefit accrues to someone else. The second difficulty is that altruism is open to exploitation. If you act selfishly, you automatically gain an advantage from altruistic acts that are directed at you. That being the case, one is forced to ask: What might it take for people to sacrifice their personal interest in order to further the interest of the larger group to which they belong? It is unlikely that there is a general answer. Fortunately, it is possible to take a look at the question as it applies to non-human animals and ask whether analogous explanations might hold in the case of humans.6 If nothing else, the exercise might point to considerations that should be kept in mind while formulating hypotheses that apply to humans. Besides, it might help us rule out hypotheses that appear attractive at first sight.


There are several explanations for animal altruism. However, they come with associated difficulties. Experience shows that for understanding any issue concerning living organisms, an evolutionary perspective is helpful.7 But there is an important distinction between humans and other creatures concerning what constitutes a satisfactory evolutionary explanation.8 When it is applied to a microbe, plant or animal, an evolutionary explanation for some trait (e.g., the streamlined body of a fish)9 tries to make a plausible case for three testable assertions pertaining to that trait: (a) the trait differs from one individual to the next; (b) the trait is heritable, meaning that children resemble parents in respect of the trait; and (c) differences in the trait translate to differences in longevity, or number of offspring, or both. The differences that are relevant for (c) are referred to as differences in fitness. In popular terminology, (a) to (c) deal with fitness and explain what is meant by ‘survival of the fittest’.


But an evolutionary explanation for a trait exhibited in human beings may depend on factors that, even though analogous to fitness, are not the same. The factors lead to successful propagation of the trait; however, the success may be measured in terms of something other than survivorship of the individual or the number of children he or she has. Behavioural traits that lead to the accumulation of wealth is an obvious instance. They exemplify traits that spread because their possession is associated with prestige, or because they convey a sense of well-being.10

The simplest form of cooperative behaviour is one in which helping others overlaps with self-help. Aesop’s fable of the bundle of sticks is a good example of ‘united we stand, divided we fall’.11 Amoebae belonging to the group known as the cellular slime moulds provide another instance. These amoebae come together and form cooperative groups when they run out of food.12 They are thought to do so because an amoeba that joins others has some chance of survival, whereas one that remains solitary has none. Analogously, as Aesop did, one can think of circumstances in which each human in a group helps himself or herself by cooperating with others.

The existence of cooperation in animals, including apparent altruism, has long intrigued biologists. It seems that altruistic behaviour runs counter to a basic tenet of Darwinian evolutionary theory, which is that for a trait to get established in a population, it must provide a reproductive advantage to those who possess the trait. By definition, an altruistic trait would appear not to satisfy this requirement. The most famous example is the behaviour of worker insects in ant, bee, wasp and termite societies. The workers, as their name indicates, work for the community – the hive or nest. However, in many cases they do not reproduce. In fact, by virtue of being sterile, often they cannot reproduce. It is someone else who benefits from their altruism – the queen, who lays eggs that give rise to the next generation of adults.13


Are there ways of getting around the difficulty? Altruism, in the sense used here (see note 1), appears to lower the biological fitness of the altruistic individual. Therefore, if it exists, it must be because either the decrease in fitness is only apparent, not real, or because its spread has largely to do with factors other than biological fitness. Interestingly, both hypotheses have been advanced to account for altruism in humans. Hints of the first sort of explanation go back all the way to the work of the 14th century Arab philosopher Ibn Khaldun.14 For him, related-ness, or common descent, lay behind ‘Asabiyah’ – cooperativeness.


The importance of relatedness was alluded to by Darwin himself and fleshed out later by a number of evolutionary theorists, prominently R.A. Fisher, J.B.S. Haldane and W.D. Hamilton. It has acquired a certain popularity, or perhaps notoriety, as the theory of the ‘Selfish Gene’.15 Its essence is that what looks like altruism at the level of the individual may be selfishness at the level of the gene.16 J.B.S. Haldane offered the same explanation many years ago and, in his inimitable style, outlined the underlying genetic calculus.17 In brief, if there is a genetic basis for altruistic behaviour,18 the behaviour can spread when it is more likely to benefit relatives, who have a high chance of carrying the same altruism-promoting gene as the altruist, than the average individual in the group.19


Haldane went on to point out that altruism could have an entirely different basis, one that was independent of genetic relatedness. Suppose the individuals of a species lived in groups,20 and groups came into conflict with one another. Then, argued Haldane, as Darwin had done much before him, a group in which the proportion of altruists was high, could be more successful as a group than one in which the proportion of altruists was low. By definition, within each group the altruists would do worse than the rest; they would decrease in proportion. But when it came to competition between groups, the presence of altruists could boost the reproductive advantage of groups that contained altruists so much, that – counter-intuitively – it could lead to an increase in the proportion of altruists overall.21 In effect, Darwin and Haldane postulated that the benefit of altruism (accruing to the group through inter-group competition) could more than compensate for the cost of altruism (imposed by inter-individual competition within the group). It turns out that stringent conditions must be met for the compensation to work, which is one reason why many think that group selection, as the process is called, is an unlikely route for the evolution of altruism.


We have looked at two ways in which altruism can evolve, either via genetic relatedness (kinship) or via group selection. There is a third way, and it comes closest to what one might call old-fashioned Darwinian natural selection. In it, the reproductive interest of the individual – and not of the genes of the individual or the group – is paramount. One means through which individual self-interest could be consistent with altruism was proposed by R.L. Trivers. It goes by the name of reciprocal altruism or, in ordinary language, ‘you scratch my back and I scratch yours’.22 An altruistic act performed by X towards Y may be reciprocated later by an altruistic act by Y towards X; on average, neither gains nor loses relative to the other.


Are hypotheses for altruism based on biological evolution applicable to humans? There is a problem with extending explanations for altruistic behaviour from other animals to humans. The problem is that when it comes to human behaviour, the complexity of the human brain, and the correlated complexity of human social organization, forces one to move beyond arguments that depend solely on reproductive interest.23 The evolution of the brain has led to a state of affairs in which it is possible to maintain patterns of behaviour that would have been impossible in most of our animal ancestors.

Let us consider the most obvious consequences. The evolution of complex brains can lead to learned behaviour becoming vastly more important than instinct. Unlike a trait with a strong innate component, learning is not restricted to being passed from parents to offspring; it can be transmitted between any two individuals within the same or different generations. This makes non-biological cultural evolution a force in itself.24 Further, the criteria that determine whether a trait will spread or not need no longer rest on reproductive success. Rather, other considerations become relevant; for example, a sense of well-being.25 Economic self-interest can supplant reproductive fitness as a force for change or, depending on circumstances, as a force for stability. Traits that ipso facto run counter to reproductive fitness can spread: birth control and the use of hallucinogenic drugs, for instance.


Such considerations imply that the problem of altruism in humans has to be viewed on a different footing from that in other animals. The obvious question suggests itself: What (non-inherited) cultural factors might foster altruistic behaviour? An optimistic answer is that the human brain permits us to plan for a future that benefits the group as a whole and to be prudent in our dealings with the environment. Examples abound from traditional cultures, in which the physical and living environments participate in one’s day-to-day living.26 The displayed behaviour would be altruism of the sort that most ethical systems promote; it would involve the suppression of individual self-interest when it collides with group interest.

The question posed in the previous paragraph permits a pessimistic answer too. That answer follows from the evolution of a sophisticated technological culture, which too is a fallout of the evolution of the brain.27 Technological skill, when combined with long-range planning, can permit equitable social life with a dependence on the environment that is sustainable. In an inequitable system, it can also lead to one group of people dominating another and compelling the latter to behave altruistically towards the former.28 An imbalance in strength and a capacity for deceit, both of them products of biological evolution, are major factors behind the persistence of successful slave-making in ants (see note 27). From the available evidence, it seems that a similar outcome, deriving from similar causes, could also result from purely cultural (non-biological) evolution. The examples are many, and they come from many different cultures and different periods. It is of interest to direct a glance at the situation in India.


In India, the current perilous state of the environment has a lot to do with implicitly sanctioned measures for improving the living conditions of the better off relative to that of the poor.29 What are the numbers and relative proportions of the two classes? Let us begin by looking at an estimate of how many fall under the category of middle class. That much used label may represent people who are, relative to the vast bulk of the population, doing quite well indeed. Credit Suisse defines the Indian middle class in terms of wealth per adult within the band 13,700-1,37,000 US dollars, or ~Rs 9-90 lakh. Its data show that, ‘Only 3% of adults have wealth above the middle class minimum, but the group accounts for 64% of total household wealth’, implying that ‘members of the middle class are not "in the middle". Rather, they are toward the top of the distribution and there are relatively few people above them.’30 Based on those figures, and using an average household size of 4, at least 75 million Indians qualify for membership of the middle and upper classes,31 leaving an astonishing ~1125 million on the other side of the ledger. The proportions work out to 6% and 94%. Even allowing for the fact that this is a rough calculation, the numbers are staggering.


Figures like these contain lessons relevant for altruism in humans. A highly skewed distribution of wealth can arise and be maintained only by the enforced altruism of the many towards the few.32 If the degradation of the environment is a consequence of the prevailing state of affairs, its improvement is unlikely to come about without redressing the existing inequalities in knowledge, power and influence, which simultaneously feed into, and are sustained by, the distribution of wealth. That requires taking active steps to reverse the existing pattern of altruism. In matters affecting the way we live, it is unlikely that a policy of non-intervention and ‘nature knows best’ will succeed on its own.


If conditions are such that the exploitation of others favours the short-term interests of some, minor disparities that arise accidentally can get reinforced and lead to the establishment of major inequalities. The inequalities can be between individuals or, when individuals live in groups, among groups. The upshot is enforced altruism: the less powerful have no choice but to act in ways that benefit the interests of others more than of themselves. In a sense, the outcome is natural. But that does not qualify it to be thought of as inevitable or desirable, let alone ethically justifiable. (The practice of slave-making in ants is not something that gives humans a licence to behave similarly.)

How can the situation be redressed? When it comes to degraded lands or polluted water bodies, a philosophy of non-interference and ‘leave it to nature’ may have a good chance of succeeding. The reasons for thinking so have been brought out lucidly in ‘Naturally’. However, if we demand that fairness must be a component of success, it is unlikely to work with us humans – unless we base our behaviour on considerations that go beyond short-term individual advantage.

To sum up, many human problems appear intractable in the absence of actions that benefit others without heed to the actor’s self-interest. Indeed, the actions may need to be detrimental to self-interest. Behaviour of this sort is said to be altruistic. The existence of apparent altruism in microorganisms and non-human animals has puzzled biologists, because it seems to go against the expectations of Darwinian evolutionary theory. Nevertheless, careful analysis shows that altruism in animals can be accounted for in a number of different ways. A common element in the explanations is that costs and benefits are estimated in the currency of fitness, i.e. in terms of the number of surviving offspring that an individual leaves behind.


This essay asks what lessons might be drawn from such explanations when it comes to humans, and points out that criteria other than biological fitness have to be considered. Enforced altruism can and does occur among humans. Going by the Indian example, it can be sustained by an oligarchic system in which wealth, power and influence are restricted to a minority. The issue of altruistic behaviour that is ethically acceptable raises exceptional problems. Behavioural modes that were moulded largely by biological evolution must give way to behaviour based on ethics and the realization of long-term, and therefore impersonal, good. The transformation is unlikely to emerge by itself. It will require active, interventionist steps.

* Dedicated to Pushpa Mittra Bhargava, with respect and affection.


1. Here and in what follows, we use the definition ‘Altruism is when we act to promote someone else’s welfare, even at a risk or cost to ourselves.’ (http://greatergood.berkeley. edu/topic/altruism/definition).

2. It is important to stress that this by no means implies that biological evolution provides the sole, or even the most important, explanatory framework for understanding human culture. See V. Nanjundiah, ‘Culture and Evolution’, in History of Science, Philosophy and Culture in Indian Civilization, Vol. XIV, Part I: Nature and Culture’, R. Narasimha and S. Menon (eds.), Centre for Studies in Civilizations, New Delhi, 2011, pp. 343-356.

3. V. Soni, Naturally. HarperCollins India, 2015.

4. Here ‘our’ refers particularly to members of what are called the upper and middle classes.

5. Garrett Hardin, ‘The Tragedy of the Commons’, Science (162)3859, 1968, pp. 1243-1248; freely accessible at http://science.

6. It deserves pointing out that plants too display traits that – at least formally – are reminiscent of animal behaviour; see R.U. Shaanker and K.N.Ganeshaiah, ‘Conflict Between Parent and Offspring in Plants: Predictions, Processes and Evolutionary Consequences’, Current Science 72(12), 1997, pp. 932-939. Pertinently, they also exhibit something like altruistic behaviour that is based on genetic relatedness; see C-C Wu, P.K. Diggl and W.E. Friedman, Proceedings of the National Academy of Sciences 110(6), 2013, pp. 2217-2222. The present discussion is restricted to animals.

7. Theodosius Dobzhansky, a major contributor to the post-Darwinian elaborations of evolutionary theory, said: ‘Nothing in biology makes sense except in the light of evolution.’ Altruism is no exception, except that the underlying issues are not restricted to biology. They involve psychology, economics, sociology and, centrally, history. Biological evolution plays a role in all of them, but the extent to which it does is debatable. Of course, one can take the extreme view that all the other fields are offshoots of biology, but that would be a sterile view. It reflects crude reductionism and suffers from serious problems. For a perceptive analysis of likely pitfalls see P.W. Anderson, ‘More is Different’, Science 177(4047), August 1972, pp. 393-396; www.

8. By ‘evolutionary explanation’ I mean the explanation for organic evolution that was developed independently and publicly announced in 1858 by C. Darwin and A.R. Wallace. It is also known as natural selection. So far it has been the most successful explanation for evolutionary phenomena. However, it is by no means the only way in which evolution can take place. See the last part of V. Nanjundiah, ‘The Origin of Species After 150 Years’, Resonance 14(2), February 2009, pp. 124-153,; describe/article/reso/014/02/0124-0153

9. To repeat, we mean an explanation for the evolution of the trait by natural selection.

10. And thereby subvert the lesson learnt through evolution, because normally a sense of well-being, or the absence of stress, means that one is doing well. Addictive drugs are one agency of subversion. Behaving in some way only because it is fashionable can be another.

11. See The maxim attributed to Benjamin Franklin, to which politicians and other public figures appear to pay heed (‘We must all hang together, or assuredly we shall all hang separately’), offers another example (see http://www. 71.htm).

12. See =vjRPla0BONA

13. The full story is too intricate to be discussed here. Termite colonies also have a ‘king’. The restriction of reproduction to a small number of individuals is known as reproductive division of labour. The phenomenon is widespread. The naked mole rat, a mammal like us, exhibits highly social behaviour that includes reproductive division of labour ( Also see, R. Gadagkar, Survival Strategies: Cooperation and Conflict in Animal Societies. Universities Press, 2001.

14. Not that he expressed his thoughts in a manner that would be viewed as constituting an acceptable explanation today. See A. Gierer, ‘Ibn Khaldun on Solidarity (‘Asabiyah’) – Modern Science on Cooperativeness and Empathy: A Comparison’, Philosophia Naturalis 38, 2001, pp. 91-104. See html#c2973

15. R. Dawkins, The Selfish Gene. Oxford University Press, 1976. Dawkins has softened his earlier views, if only slightly; see the third edition of the book published in 2006.

16. Obviously the word ‘selfish’ cannot be applied to a gene in the way it can to humans. Dawkins, who popularized the term ‘selfish gene’, seems to have been trying to describe the genetic basis of cellular and organismal phenomena as if in a Panchatantra story, with participants who had goals and employed various strategies to achieve those goals. As he explained later, he meant to say that if we look at genes in an anthropomorphic way, the properties that they exhibit in certain situations resemble what we would expect from a sentient individual who behaves selfishly. Thus ‘selfish’ is simply a convenient, if (in hindsight) injudicious, shorthand description of certain properties. No more should be read into it than into the descriptors ‘colour’ or ‘charm’ used by theoretical physicists for quarks. The tendency to use it in the context of a gene is regrettable, but the mistake may be too far gone to be redressed.

17. ‘Let us suppose that you carry a rare gene which affects your behaviour so that you jump into a river and save a child, but you have one chance in ten of being drowned, while I do not possess the gene, and stand on the bank and watch the child drown. If the child is your own child or your brother or sister, there is an even chance that the child will also have the gene, so five such genes will be saved in children for one lost in an adult. If you save a grandchild or nephew the advantage is only two and a half to one.’ J.B.S. Haldane, ‘Population Genetics’, New Biology 18, 1955, 34-51, p. 44.

18. That is, other things being equal, a difference in genetic make-up can be correlated reliably with a difference in behaviours that differ in degrees of altruism.

19. The group may or may not comprise the entire population.

20. Many different factors can favour group formation or the association of individuals, for example, the manner in which food sources are distributed in space.

21. The effect would be indirect – it would require taking the entire population into account. There is an implicit ecological assumption here: the individuals who constitute the population live in groups that exist separately, compete with each other and mix only intermittently. A recent experiment with bacteria has shown that this works best when group sizes are small – more precisely, when the size distribution is Poissonian. See J.S. Chuang, O. Rivoire and S. Leibler, ‘Simpson’s Paradox in a Synthetic Microbial System’, Science 323, pp. 272-275.

22. R.L. Trivers, ‘The Evolution of Reciprocal Altruism’, The Quarterly Review of Biology 46(1), 1971, pp. 35-57. See

23. Human behaviour is influenced by both biological inheritance (‘genes’), learning and, in a broader sense, culture. This leads to a system of dual inheritance, in which traits can be passed on as correlates of genetic transmission, or as elements of culture that can vary in the absence of genetic variation, or via an interplay of the two. Also, culture is not restricted to humans. See https://en.wikipedia. org/wiki/Dual_inheritance_theory and J.T. Bonner, The Evolution of Culture in Animals. Princeton University Press, 1983.

24. That is not influenced by biology, except in the trivial sense that it involves living organisms.

25. It has been said that the selfish gene is gradually giving way to the hedonistic neuron.

26. See M. Gadgil, ‘Cultural Evolution of Ecological Prudence’, Landscape Planning 12, 1985, pp. 285-299; available in http://wgbis.

27. This is not to say that enforced altruism is impossible in other animals, only that it is rare. Slavery in ants provides a famous example. The enslaved workers belong to another species, are ‘kidnapped’ by the slave makers when they are eggs or larvae, and are fooled into thinking that they are working for members of their own species. See The enslaved ants are not always passive; slave revolts do occur.

28. One could cite any number of instances from human societies. The history of human slavery and colonialism are glaring examples. The ability to manage access to information should be counted among the necessary technological skills.

29. This is not to say that other factors, for instance population pressure, play no significant role. Of course environmental degradation can be caused by the way of life led by a large number of poor people. But that begs the question why their number is large. Mahmood Mamdani drew the conclusion many years ago that in Third World countries, people are poor not because they have too many children; they have too many children because they are poor. (The Myth of Population Control, Monthly Review Press, New York, 1972.)

30. The number of adults in India is ~792 million, so the category would consist of ~25 million adults, or (my guess) ~75-100 million if one includes other family members (2015 Global Wealth Databook;

31. Rough estimate: 792 million adults x~3.5% (Table 3-1) plus twice as many children ~75 million. Because the lower boundary is set at 13,700 US dollars, this may overestimate the size of the middle-cum-upper classes.

32. With reference to the impressive number of 55 (presumably dollar) billionaires we have, a recent news item states: ‘The fact that most billionaires gained wealth because of their access to natural resources such as land or government contracts raised disturbing questions about the nature of India’s growth process.’ In the same article, the Governor of the Reserve Bank is reported to have asked – rhetorically, it would appear – how India could not be called an oligarchy of the rich. See ‘Why Worry About Inequality in India?’, Economic Express, 15 November 2015; 2221473700.html