In this scene, as in many other situations in wildlife, we see a bizarre combination of altruistic and selfish behavior. The inviting food cry of a seagull is a typical example of altruism. The seagull does not benefit from this cry. The other gulls win: they get a chance to dine. The second part of the scene is the fight. Here, of course, we see only pure selfishness on the part of all the participants.

The answer lies in Hamilton's rule. Seagulls in the White Sea feed mainly on schooling fish, such as herring. If a seagull has noticed one fish, then most likely there are many others nearby: there is enough for everyone. This means that the value With- the price of an altruistic act - will be low on average. Value AT- the payoff of those who fly to the scream will be quite large: they will have lunch. Since the fish are schooling, the next school may have to wait a long time. Value R(relatedness) is also likely to be high, because gulls nest in colonies, often return to the same place after wintering, and therefore, most likely, its relatives nest next to this gull - parents, children, brothers and nephews.

Of course, the most beneficial thing for a seagull (more precisely, for its genes) would be to learn to distinguish between a situation when there is a lot of food and enough for everyone, and when there is little food. In the first case, it is beneficial to shout, and in the second, to keep quiet. But such calculations require brains. And the brain, as we know, is an expensive organ. Selection, as a rule, tries to save on brains. Plus, the brain is heavy. Seagulls need to fly, not solve algebraic problems. Therefore, the bird cannot figure out when it is beneficial for it to call the companions, and when it is not, and its behavior turns out to be illogical. Not always, but only with a lack of fish.

The evolution of altruism has gone especially far in hymenoptera insects: ants, bees, wasps, bumblebees. In social Hymenoptera, most females give up their own reproduction in order to feed their sisters. This is the highest manifestation of altruism. Such animals are called eusocial, that is, "truly social". But why hymenoptera?

Hamilton suggested that this was due to the peculiarities of the inheritance of sex. In Hymenoptera, females have a double set of chromosomes, while males have a single set. Because of this, a paradoxical situation arises: sisters turn out to be closer relatives than mother and daughter. In most animals, sisters have 50% common (identical in origin) genes. Value R in Hamilton's formula is 1/2. In Hymenoptera, sisters share 75% of their genes ( R= 3/4), because each sister receives from her father not half of his chromosomes, but the entire genome. Mother and daughter in Hymenoptera have, like in other animals, only 50% of common genes. So it turns out that, other things being equal, it is more profitable for female Hymenoptera to raise sisters than daughters.

Mechanism of sex inheritance in Hymenoptera. The female is diploid, that is, it has a double set of chromosomes (2n). She can lay an unfertilized egg with a single set of chromosomes (n), from which a haploid male will hatch. If the egg is fertilized, then its chromosome set will be double, and a female will hatch from it. The female receives half of the chromosomes from the mother, half from the father. The male receives half of her chromosomes from the mother, but he does not have a father. This mechanism of sex inheritance is called haplodiploid.

In reality, everything is somewhat more complicated. In addition to sisters, there are also drone brothers who share only 25% of their genes with their sisters (when viewed from the side of the sister) or 50% (from the point of view of the brother). However, working females also raise brothers (although they do not like them). We will not go into this rather complex theoretical area, especially since the primates we are interested in are not haplodiploids. But social hymenoptera have (or had in the evolutionary past) another important property that dramatically increases the likelihood of developing altruism under the influence of kin selection. This property is monogamy.

The offspring of monogamous diploid parents have an average of 50% of common genes ( R= 0.5). In the offspring of a female mating with many males, the average value R tends to 0.25 (if there are enough males). For kin selection, this is a very serious difference. At R= 0.5, any trifle is enough to tip the balance in favor of siblings. At R= 0.25 their children are definitely more expensive. It is very important that termites are monogamous, the second order of insects in which eusociality is widespread, and without any haplodiploidy. Not only females work for termites, but also males (they are diploid, like their sisters).

As we remember, monogamy was probably characteristic of the ancient hominids. This could become a powerful stimulus for the development under the influence of kin selection of fraternal (and sisterly) mutual assistance, intra-family cooperation and altruism. And, of course, paternal love, and at the same time the devotion of children to both parents, and not just the mother. It is possible that kin selection was able to support this range of altruistic feelings in our ancestors precisely because they were - at least in part - monogamous.

This slide shows the definitions, I will not dwell on them, I think everyone is more or less clear what altruism is - both in ethics and in biology. We are faced with two main questions: first, on the one hand, it is clear that many of life's tasks are much easier to solve through joint efforts than alone. Why, then, did the biosphere never turn into a realm of universal love, friendship and mutual assistance? This is the first question. And the second question is the opposite: how can altruistic behavior develop in the course of evolution if evolution is based on the selfish mechanism of natural selection. If the fittest always survives, then what kind of altruism can we talk about?! But this is an extremely primitive and incorrect understanding of evolution. The error here is due to the confusion of levels at which we consider evolution. At the level of genes, evolution is based on the competition of different variants or alleles of the same gene for dominance in the gene pool of a population. And at this genetic level there is no altruism and, in principle, cannot be. Gene is always selfish. Now, if such a “good” allele suddenly appears, which, to the detriment of itself, allows another competing allele to multiply, then this “good” allele will be automatically ousted from the gene pool and simply disappear. Therefore, at the level of genes, there is no altruism. But if we shift our gaze from the level of genes to the level of organisms, the picture will be different. Because the interests of a gene do not always coincide with the interests of the organism in which this gene sits. Why? Because a gene, or rather an allele, a variant of a gene is not a single entity. It is present in the gene pool in the form of many identical copies. And an organism is a single entity, and it carries only, roughly speaking, one or two copies of this allele. And sometimes it is beneficial for a selfish gene to sacrifice one or two copies of itself in order to provide an advantage to other copies of itself that are contained in other organisms. But here I must make a reservation, biologists are sometimes reproached for using such metaphors as "the gene benefits", "the gene wants", "the gene strives". I hope you understand that a gene doesn't really want anything, it doesn't have any desires, a gene is just a piece of a DNA molecule. Of course, he does not understand anything and does not strive for anything. When biologists say “the gene benefits”, “the gene wants”, “the gene strives”, they mean that under the influence of selection the gene changes as if it wanted to increase the efficiency of its reproduction in the gene pool of the population. That is, if a gene had brains and desires, it would change in the same way that it changes automatically under the influence of selection. I hope this is clear to everyone. It can be beneficial for a gene to sacrifice a few copies of itself to give advantage to other copies, and due to this, altruistic sacrificial behavior can develop in organisms. For the first time, biologists began to approach this idea quite a long time ago, back in the 30s of the twentieth century, this idea began to be expressed and developed. An important contribution to this matter was made by Ronald Fisher, John Haldane, William Hamilton.

Creators of kin selection theory

And the theory they built is called "Kind Selection Theory". Its essence was figuratively expressed by Haldane, who once said: "I would give my life for two brothers or eight cousins." What he meant by this can be understood from the following formula.

Hamilton's rule:

I ask you not to be afraid, this will be only one formula in the lecture and there will be no more. This is a very simple formula. This is called "Hamilton's Rule". The altruism gene, that is, the allele that contributes to the altruistic behavior of the organism, will be supported by selection, that is, it will be distributed in the gene pool of the population, if this inequality is true:

gV > C

where r- the degree of genetic relatedness of the one who makes the sacrifice and the one who accepts the sacrifice. This degree of genetic relationship is the probability that the one for whom you sacrifice yourself has the same allele of the same gene that you have. For example, this gene of altruism. Let's say if some allele is sitting in me and I have a sibling, then, roughly speaking, the probability is ½ that it has the same allele. If, say, a cousin, then it will be 1/8. AT(Benefit) is a reproductive advantage received by the addressee of an altruistic act, that is, by those for whom you sacrifice yourself. BUT With(Cost) is the "price" of the altruistic act, that is, the reproductive damage inflicted by the donor on himself. This can be measured in terms of, say, the number of children born or not born by you.

Haldane said "I would give my life for two brothers", here we have to modify a little more, if we sacrifice ourselves not for the sake of one individual, but for the sake of several, then we can add n at the beginning:

nrB > C

n is the number of those who accept the sacrifice. Here are two brothers, n = 2, r=0.5, AT- this can be substituted for any number, say the number of children produced by each person. With- this is your damage, you sacrifice yourself, that is, you do not give birth to these children, well, for example, if AT and With= 2, then in this case, these values ​​will be equal, that is, if you give your life for two brothers, then it’s like “bash on bash”, “wash on soap”. It will be profitable for three brothers. Gena, not for you. Now we can understand the behavior of those same seagulls. This food call is inviting, why do seagulls develop such an instinct to scream and call for others when they see something edible? Look, these seagulls in our White Sea feed mainly on schooling fish: herring, stickleback - and if a seagull notices one fish, then most likely there are many, many others nearby, and there is enough for everyone, that is, it does not will lose. Value With– the price of an altruistic act is likely to be low. AT- the winnings of those who fly to the scream will be quite large, they will have lunch. Since, again, the fish are schooling, it may take a long time to wait for the next flock. That is, the gain is quite tangible. r- kinship. The relationship is also, most likely, quite high, because they nest in colonies, often return to the same place after wintering, and therefore, most likely, its various relatives nest next to this gull: parents, children, brothers, nephews, etc. .d. And n- the number of seagulls that will hear, fly in and dine is also quite high. Here she is screaming. And why does she not share her prey, what she has already grabbed does not give - because here With already much more turns out, she really remains without lunch. And n smaller. By giving her prey to another gull, she will feed one, not a whole flock. So the inequality is not fulfilled, therefore such an instinct has not been developed. Of course, it would be most beneficial for the seagull to learn to distinguish between a situation where there is a lot of food and enough for everyone, and then call. And when food is scarce, eat silently. But for this you need - what? Brain. And this is a very “expensive” organ, selection usually saves on brains. Birds need to fly, they need to lighten their body weight, and not solve all sorts of algebraic problems. Therefore, the bird cannot figure out in which case it is profitable - it is not profitable, and such illogical behavior is obtained.

Hymenoptera - a group in which the evolution of altruism has gone especially far

In general, Hamilton's rule has remarkable predictive and explanatory power. For example, in which group of animals the evolution of altruism has led to the most significant consequences. Apparently, these are Hymenoptera insects - ants, bees, wasps, bumblebees. In these insects, several times, apparently more than a dozen times, the so-called eusociality arose, that is, a social way of life in which most individuals refuse to reproduce at all and raise their sisters. Working females do not breed, but help their mother raise sisters. Why exactly Hymenoptera, why is it so common in this order of insects? Hamilton suggested that the whole point here is in the characteristics of the inheritance of sex. In Hymenoptera, females have a double set of chromosomes like most animals, but males have a single set of chromosomes, males develop from unfertilized eggs in Hymenoptera - parthenogenetically. Because of this, a paradoxical situation arises - sisters turn out to be closer relatives than mother and daughter. In most animals, sisters share 50% of their genes. Value r in Hamilton's formula is ½, and in Hymenoptera sisters have 75% of common genes. Because each sister receives from her father not half of his chromosomes, as usual in other animals, but she receives the entire paternal genome as a whole. And this complete paternal genome is received by all sisters, one and the same. Because of this, they share 75% of their genes. It turns out that the female Hymenoptera sister is a closer relative than her own daughter. And therefore, other things being equal, it is more profitable for them to force their mother to give birth to more and more sisters and raise them than to give birth to their own daughters. But in reality, everything is somewhat more complicated here, because there are still brothers who, on the contrary, turn out to be (brother and sister) more distant relatives than ordinary animals. I will not go into these subtleties, but in this situation, where sisters are closer to each other than mother and daughter, it appears that there is enough of her in the order of Hymenoptera for such altruistic systems to arise repeatedly. But besides kin selection, there are other mechanisms that help or, conversely, hinder the evolution of altruism. Let's look at specific examples and start with bacteria. Bacteria also have altruism, very widespread. Now one of the interesting directions in microbiology is the experimental study of the evolution of bacteria, "evolution in a test tube."

The evolution of "altruists" and "deceivers" in vitro: experiments with the bacterium Pseudomonas fluorescens

Altruists and deceivers in the bacterium Myxococcus xanthus

Honest yeast and fake yeast can live together

In yeast populations, some cells behave like altruists - they produce an enzyme that breaks down sucrose into easily digestible monosaccharides: glucose and fructose, while other individuals are selfish yeasts, they do not secrete this enzyme, but use what is produced by altruists. Enjoy the fruits of other people's work. Theoretically, this should have led to the complete displacement of altruists by egoists. But in reality, the number of altruists does not fall below a certain level. They began to investigate why. It turned out that the altruism of yeast, on closer examination, is not entirely unselfish. They really help everyone around them, release the enzyme into the external environment, but they still take 1% of the glucose produced for themselves immediately, as if bypassing the “common boiler”. And due to this little trick, with a low frequency of occurrence of altruists, it turns out to be more profitable to be an altruist than an egoist. Hence the peaceful coexistence of these two varieties of yeast in one population. However, it is clear that on such small tricks it is hardly possible to build a serious complex cooperative system. Another great trick of this kind is called the Simpson Paradox. The essence of this paradox is that under a certain set of conditions, the frequency of occurrence of altruists in a group of populations will increase, despite the fact that within each individual population this frequency is steadily decreasing.

Simpson's paradox

This slide shows a hypothetical example of the Simpson Paradox in action. There was a population where there were altruists and egoists in half. It has divided into small populations, where the ratio of altruists and egoists varies widely, this is the key point. There needs to be so much variability in these small daughter populations. To do this, these child populations must be very, very small, preferably only a few individuals. Then each daughter population grows, in each population the proportion of altruists decreases, in each of the three populations the proportion of altruists decreases, but those populations where initially there were more altruists, in general, grow faster. Altruists still help others. As a result, at the output, in total, the percentage of altruists increases, despite the fact that in each individual population it decreased. Recently it was possible to show experimentally that this is not only a theory, but that such a mechanism can actually work in microbes. True, apparently, rather rare conditions must be met for this to happen, but this is not yet entirely clear. But there is also such a trick to maintain the level of goodness in the world. It's time to move from microbes to multicellular. The advent of multicellular organisms in general, and animals in particular, was a major triumph in the evolution of altruism. In a multicellular organism, most of the cells are altruists who have given up their own reproduction for the sake of the common good. Animals, compared with microbes, have new opportunities for the development of cooperation based on complex behavior and learning. But, unfortunately, the same opportunities appeared in the deceivers, and the evolutionary arms race continued at a new level. Again, neither the altruists nor the deceivers gained a decisive advantage.

Altruism in social insects is far from selfless

In many species of Hymenoptera, workers sometimes become selfish by laying their own eggs. In Hymenoptera, as we have said, males are born by immaculate conception, parthenogenetically, from haploid unfertilized eggs. Worker individuals in some wasps try to lay such unfertilized eggs and breed their own sons. This is the most profitable strategy, as I mentioned, for a female Hymenoptera, the most profitable business is to raise sisters and native sons. This is what they are trying to do. But this is not to the liking of the other workers, who benefit from laying their own eggs but not from the sisters, so they destroy the eggs laid by their sisters. It turns out such a kind of morality police. And special studies have shown that the degree of altruism in the colonies of such wasps depends not so much on the degree of kinship between individuals, but on the severity of such police measures, on the effectiveness of the destruction of illegally laid eggs. That is, apparently, the cooperative system created by kin selection, even in Hymenoptera, will still be destroyed by deceivers if it fails to develop additional means of combating egoism.

Another example showing that the altruism of social insects is far from the ideal of selflessness. There are wasps that have several adult females in the family, of which only one, the oldest, lays eggs. The rest take care of the larvae. When the queen dies, the next most senior wasp takes her place. That is, they strictly observe the principle of seniority. At the same time, helper wasps, which do not yet breed themselves, differ greatly in the degree of their labor enthusiasm. Some work without sparing themselves, while others sit in the nest, rest. And now, as it turned out, their labor enthusiasm depends on how great the chances of this wasp for the royal throne are. How great are her chances to leave her own offspring, to start her own family. If these chances are not great, like those of low-ranking wasps, the last in line for the royal throne, then the wasp is working actively. And if the assistant has a high rank, then she tries to take care of herself and work less. This behavior of wasps is also well explained by Hamilton's rule. It must be taken into account that the value With- the price of altruistic behavior - varies depending on the circumstances. In this case, from the chances of the royal throne. That is, the propensity for altruism is stronger in those who have nothing to lose. Is it possible to create a society in which altruism will be maintained without violence, and at the same time there will be no deceivers? Neither wasps nor humans have succeeded so far, but some cooperative systems that exist in nature indicate that it is possible to prevent the appearance of deceivers in some cases. One way to keep cheaters out is to reduce the genetic diversity of the individuals in the system to zero so that everyone is genetically identical. Then the symbionts simply won't be able to compete with each other for which one of them grabs a bigger piece of the common pie. That is, symbionts can, but the genes that sit in them will not be able to compete: they are all the same. That is, if all symbionts are genetically identical, then selfish evolution within the system becomes impossible. Because from the minimum set of conditions that are necessary for evolution, and this is the Darwinian triad of heredity, variability and selection, one of the components, namely variability, is excluded. That is why evolution never managed to create a normal multicellular organism from genetically heterogeneous cells, but managed to create it from clones, descendants of a single cell. There is such an interesting phenomenon as insect agriculture.

Some ants, some termites grow mushrooms, "domesticated" mushrooms, in special gardens in their nests. In such a situation, it is just very important to ensure the genetic homogeneity of the symbionts so that deceivers do not begin to appear among them, among mushrooms, in this case. When a cooperative system, as in the case of insect agriculture, consists of a large multicellular host, in this case an insect, and small symbionts, then the easiest way for the host to ensure the genetic identity of its symbionts is to pass them down. And only one of the sexes should do this: either males or females. This is exactly how leaf-cutting ants pass their mushroom cultures from generation to generation. In the vertical transfer of symbionts, they take a small amount of seed, this mushroom, with them, before establishing a new anthill. And this leads to the fact that genetic diversity, due to constant bottlenecks in the number of fungi, is constantly maintained at a very low level. But, however, there are also symbiotic systems with a horizontal transfer of symbionts, that is, for example, each host collects symbionts for himself in the external environment. In such systems, symbionts in each host will be genetically heterogeneous, they retain the ability to selfish evolution, and therefore deceivers appear among them every now and then. And here nothing can be done. Tricksters appear, for example, many trickster strains are known among symbiotic luminous bacteria that are symbionts of fish and squid. They work as flashlights for fish and squid, symbiotic bacteria. But there are deceivers who live there but do not shine. There are deceivers among nitrogen-fixing nodule bacteria, plant symbionts. There are deceivers among mycorrhizal fungi, among unicellular algae zooxanthellae - these are symbionts of corals. In all these cases, evolution failed to ensure the genetic homogeneity of symbionts, and therefore the hosts have to deal with deceivers by some other methods, and most often simply tolerate their presence, relying on certain mechanisms that ensure a balance in the number of deceivers and honest cooperators. All this is not so effective, but, unfortunately, selection notices only momentary benefits, it cannot look ahead and is not at all interested in long-term prospects, so this is how it turns out. In general, if it were not for the problem of deceivers, then our planet, perhaps, would look like an earthly paradise. But evolution is blind, and so cooperation develops only where one or another set of special circumstances helps to curb or prevent deceivers. If in some kind of animal cooperation has already developed so much that the species has switched to a social way of life, then more interesting and more complex things begin, competition begins not only between individuals, but also between groups of individuals.

Intergroup competition promotes intragroup cooperation

What this leads to is shown, for example, by this model developed by American ethologists, they called it the “Nested Tug of War Model”. In this model, each individual selfishly spends part of the resources to increase his share of the “social pie”. They are trying to sort of take away resources from their comrades in the group. This part of the resources spent on intra-group squabbles is called the "selfish effort" of this individual, and a typical example of such internal squabbles is when social wasps prevent each other from laying eggs, but at the same time try to lay their own. That is, there is competition within the group between individuals, but there is also competition between groups. And it is built on the same principles as between individuals within the group, that is, it turns out to be a nested two-level competition. And the more energy individuals spend on intra-group struggle, the less it remains for inter-group competition and the less the “common pie” of the group turns out - the total amount of resources obtained by the group. The study of this model showed that competition between groups should be the strongest stimulus for the development of intragroup cooperation. This model seems to apply to human society as well. Nothing unites a team like a joint confrontation with other teams, a multitude of external enemies; Clearly, this is a prerequisite for the existence of totalitarian empires and the most reliable means of rallying the population into an altruistic anthill. But before applying any biological evolutionary models to humans, we must make sure that human morality is at least partly genetic in nature. It is easier to study the evolution of altruism on bees and bacteria, because one can immediately confidently assume that the key lies in the genes, and not in upbringing and not in cultural traditions. And studies of recent years have shown that the moral qualities of people are largely determined not only by upbringing, but also by genes.

Kindness, altruism and other "socially useful" qualities of people are partly hereditary (genetic) nature

Moreover, the available methods allow us to evaluate only the tip of the iceberg, only those hereditary traits of our behavior for which modern people still have variability, that is, which have not yet been fixed in our gene pool. It is clear that all people have some genetic basis of altruism. The question is, in what phase is the evolution of altruism in modern humanity. Either the genetic stage has already ended, or the evolution of altruism continues at the gene level. Special studies, based, in particular, on twin analysis, have shown that such traits as a tendency to good deeds, gullibility, gratitude - all this is subject to hereditary variability in modern people. Hereditary, that is, genetic variability. This is a very serious conclusion. It means that the biological evolution of altruism in humans may not be complete yet. Some specific genes that affect the moral qualities of a person have also been identified. I don't have time to talk about these genes in detail, but the general conclusion is clear: altruism in humans, even today, can still develop under the influence of biological mechanisms. And so evolutionary ethics is quite applicable to us.

Reciprocal (mutual) altruism

In animals, altruism is usually directed either towards relatives, or, alternatively, it can be based on the principle: you - to me, I - to you. This phenomenon is called reciprocal or reciprocal altruism. It occurs in animals intelligent enough to choose reliable partners and punish deceivers, because systems based on mutual altruism are highly vulnerable and cannot exist at all without effective means of combating deceivers. The ideal of reciprocal altruism is the so-called "Golden Rule of Ethics": treat others the way you want to be treated. And truly disinterested concern for non-relatives is rare in nature, perhaps the person is almost the only species in which such behavior has received some development. But recently an interesting theory has been proposed, according to which altruism in humans developed under the influence of frequent intergroup conflicts. I have already said that models show that intergroup hostility contributes to the development of intragroup altruism. According to this theory, altruism in our ancestors was originally directed only at members of their group. Naturally, in such a situation, the researchers, even with the help of mathematical models, showed that it seems that altruism could develop immediately only in combination with parochialism. Parochialism refers to loyalty to one's own and hostility to strangers. And it turns out that our opposite properties, such as, on the one hand: kindness, altruism, on the other hand: militancy, hatred for strangers, for everyone who is not with us, who is not like us - these opposite qualities of ours developed in a single complex, and neither one nor the other of these traits individually did not bring any benefit to their owners. But to test this theory, facts are needed, which are now being tried to be obtained - in particular, with the help of various psychological experiments. For example, it turned out that the majority of three- or four-year-old children usually behave like egoists, but by the age of 7-8 they already have a clearly expressed readiness to help their neighbor. And special tests have shown that most often altruistic behavior in children is based not on a disinterested desire to help, but on the desire for equality and justice.

For example, children tend to reject dishonest, unequal options for sharing sweets, both in their favor and in someone else's favor. That is, it no longer looks like disinterested altruism, but a desire for equality, egalitarianism, this is some form of struggle against deceivers, in fact, maybe. And the proportion of such lovers of justice among children is growing very rapidly with age. The results of various psychological experiments, in general, are in good agreement with the theory of the joint development of altruism and hostility to strangers.

Altruism among “friends” and hostility towards strangers: two sides of the same coin

It turned out that altruism and parochialism develop in children almost simultaneously, and both properties are more pronounced in boys than in girls. This is easy to explain from an evolutionary point of view, because in the conditions of primitive life, male warriors lost much more in case of defeat in intergroup conflict and gained much more in case of victory. For example, in case of victory, they could take captives; in case of defeat, they most likely lost their lives. And women in many cases were only in danger of changing their husbands. And therefore, it is not surprising that men have more pronounced intra-group cooperation and hostility towards strangers. The idea of ​​a connection between the evolution of altruism in humans and intergroup conflicts was expressed by Darwin.

As we know, this is a quote from his book, where he sets out his views on how, in the course of evolution, the foundations of morality could have been formed in our ancestors. Such arguments cannot do without intergroup wars. Accordingly, we know that intergroup competition can promote the development of intragroup altruism, but several conditions must be met for this to happen. Here, in particular, intergroup enmity should have been quite sharp and bloody among our ancestors. Was it really so? Recently, the archaeologist Samuel Bowles, one of the authors of this theory of the coupled evolution of altruism and hostility to strangers, tried to assess whether the tribes of our ancestors were sufficiently hostile among themselves for natural selection to develop in-group altruism.

Intergroup wars - the cause of altruism?

Extensive archaeological data on the Old Stone Age, on the Paleolithic were analyzed, and the conclusion was drawn such that the conflicts in the Paleolithic in general were very bloody. Between 5 and 30% of all deaths were violent, apparently usually due to intergroup conflicts. This is actually a huge number. Up to 30% of violent deaths. This seems completely counter-intuitive and hard to believe, but it's a fact. This is not only Bowles, and our researchers believed and came to the same conclusions that the level of bloodshed in the Stone Age was much higher than even in the twentieth century, taking into account the two world wars - per capita, of course. That is, in the Stone Age you were much more likely to die at the hands of a murderer or an enemy from another tribe than - even taking into account two world wars - in the twentieth century. And, calculations show that this degree of bloodshed is more than enough for natural selection to help maintain a high level of intragroup altruism in hunter-gatherer populations. Moreover, this should occur even in those cases when within each group the selection favors exclusively egoists. But this condition, most likely, was not observed, because selflessness and military exploits, most likely, increased the reputation and, consequently, the reproductive success of people in primitive collectives.

Indirect reciprocity. (indirect reciprocity)

This mechanism for maintaining altruism through reputation improvement is called indirect reciprocity, that is, you perform an altruistic act, sacrifice yourself - this increases your reputation in the eyes of your fellow tribesmen, and you have reproductive success, leaving more descendants. This mechanism does not only work in humans; Surprisingly, it also occurs in animals, and a wonderful example is such social, public birds, Arabian gray thrushes. They live in colonies and raise chicks together. They have sentries that sit in the tops of the trees and watch for predators. It is customary for them to feed each other, to help each other in this way. Males help females take care of the chicks, in general, such a social way of life. And it turned out that among these thrushes, only high-ranking males have the right to feed other males. If a low-ranking male tries to feed his older relative, he will most likely receive a thrashing - this is a violation of subordination. That is, these social birds compete for the right to do a good deed. And only a high-ranking male can also be sentry. That is, altruistic acts acquire a symbolic meaning. They begin to serve as status signs, serve to demonstrate and maintain their own status. Reputation was very important for people at all times.

There was even such a hypothesis, there is such a hypothesis, that one of the incentives for the development of speech was the need to gossip. Gossip - what is it? This is the oldest means of disseminating compromising information about unreliable members of society, which contributes to team building and punishment of deceivers. With this, I'm already nearing the end. I must say that this topic is very large and is now actively developing, and in one lecture it is absolutely impossible to tell about all the interesting research in this area.

Some ideas not included in the report

Here on this slide are listed in abstract form some points that were not included in the lecture. For example, it is shown that people have innate psychological properties, predispositions, aimed at effectively identifying deceivers. Such very beautiful experiments were carried out. There are some tests developed by psychologists a long time ago that are very difficult for a person to pass, puzzles that are difficult to solve, to guess. But problems can be submitted in different contexts. You can talk about Masha and Petya and how many apples someone has. And you can find another entourage for this problem. And it turned out that if the entourage is associated with the exposure of a deceiver, with the exposure of a violator of some social order, then people are significantly more successful in solving such problems. That is, if not about Masha, Petya and apples, but about the fact that someone deceived someone, stole, some kind of deception - the task is solved better than in various other frames. "Costly punishment" is a widespread phenomenon, also a manifestation of altruism - people are ready to make sacrifices in order to effectively punish deceivers. That is, I am ready to sacrifice my own interests, if only to properly punish that scoundrel. This is also a manifestation of altruism. A person sacrifices himself for the sake of the public, so to speak, good. Or at least what he considers a public good. Then there are more interesting arguments, works on the emotional regulation of the processes of formation of moral judgments, there are very interesting neurobiological works that show that, firstly, moral judgments in people are made mainly through emotions. When we solve some moral dilemmas, in our brain, first of all, the departments associated with emotions are excited. And there are also very interesting results obtained on people in whom certain parts of the brain are disabled, as a result of a stroke, for example, and how this affects their morality. For example, a part of the brain has been identified, damage to which leads to the fact that a person loses the ability to feel guilt and sympathy, empathy - while all other functions of the intellect are fully preserved. There are various other neurobiological interesting things. There is another such branch - evolutionary religious studies, where the evolutionary roots of religions and the possible role of religious beliefs in strengthening, strengthening this parochial altruism are analyzed. In particular, the function of rituals, shared religious rites, as shown by some special studies, may be to prevent the appearance of deceivers and to strengthen parochial altruism. In general, this is such a young rapidly developing area. In conclusion, I want to emphasize that it is very important to remember that if we say that this or that aspect of our behavior, our morality, has an evolutionary explanation, has evolutionary roots, this does not mean at all that this behavior is thus justified, that it is good and correct.

Conclusion

When we deal with evolutionary ethics, we are talking about the morality that was formed as a result of biological evolution at the stage of hunter-gatherers. It is clear that with the development of civilization the situation is changing - what was good and highly moral for a hunter-gatherer is not necessarily good and highly moral for a modern city dweller. Fortunately, evolution has also given man a mind, and, for example, evolutionary ethics warns us that we have, indeed, an innate tendency to divide people into “strangers” and “ours”. And to "strangers" to feel disgust and hostility, enmity. And we, as rational beings at the current stage of cultural and social development, must understand and overcome such things. All. Thank you for your attention.

Lecture discussion

Boris Dolgin: Thank you very much. It seems that this topic would be good for some big public discussion, perhaps with representatives not so much of the humanities as of the social sciences. Social science is now beginning to look much stricter, as it seems to me, much tougher in distinguishing where there is a demonstrative judgment, and where there are interpretations and constructions on top of these judgments, which is the sin of the stated part of supposedly natural science. It seems that there are some strange sagging in place of the statement about the genetic nature of the inheritance of altruism, although this is clearly not the only possible interpretation of the data presented - not even for humans, but for social animals. And somewhere in the argument, the line was not very clearly drawn between what can be considered directly proven, what kind of experiment was carried out, what he could, in general, prove - and for what statement. And that, in turn, is not a completely verifiable interpretation of the results.

Alexander Markov: Naturally, I basically told the conclusions of some article in a thesis form, just in one sentence. Conclusion after conclusion. Naturally, I simply did not have time to discuss the degree of reliability of certain conclusions. There is a separate conversation for each phrase, how reliable it is.

Novel: The question is next. You linked the huge percentage of deaths in the Paleolithic and the consequent development of altruism. Can we conclude that in the 20th century, at the beginning and in the middle, the level of altruism was extremely low, which is why there was a huge number of deaths?

Alexander Markov: This could be an evolutionary factor that acted for a long time, which directed selection in such a way that those individuals who, with their own, with members of their tribe, were able to cooperate and were even ready to sacrifice themselves for their own, received an advantage. For your little tribe. And how to tie this to modern wars, to modern society - this is a rather difficult task, and there are no serious data here, there is no direct connection, because now social and cultural evolution plays a much greater role in the changes that are happening to humanity . The development of our knowledge, our culture, science, and not at all biological evolution, which, of course, is going on, but it is going very slowly. And such episodes as the 20th century are nothing for evolution, nonsense. Less than 10-50 thousand years - there is nothing to talk about. It's like a little something from different areas.

Boris Dolgin: The question had a very important, albeit somewhat strangely expressed, thought: do you want to try measuring altruism? That is, to introduce some kind of unit, somehow try to isolate it from behavior? If you use this category all the time, I would like to instrument it. The question, as I understand it, was how do you measure altruism for a certain period? Your answer: other factors are playing a big role now. And here, I hope, most of us will fully agree with you. But then what to do with "altruism"? Why do you need this category at all? What are you doing with her?

Alexander Markov: In biology, altruism is always nothing more than a metaphor, an image. And some researchers do not like to use this word at all, they replace it with all sorts of euphemisms. For example, in my opinion, those authors who dealt with yeast, one yeast secretes the enzyme, helps others, the other yeast does not secrete the enzyme. To call this one yeast an altruist, and another an egoist, perhaps, some authors believe that it is not necessary. Call it something else. Each situation has something different in mind. A person has some special psychological tests. This is a multifaceted thing. And in the case of yeast, they simply measure it: it releases an enzyme, it does not release an enzyme. Artificial systems of altruists - egoists from microbes are being created. Now genetic engineers are conducting experiments, artificially creating altruistic bacteria that secrete some kind of socially useful product, and selfish bacteria that do not secrete this product. And they look at how they will interact with each other, who will displace whom and how such a system will behave. That is, if these are not people, but bacteria, then in each case there is a different idea. In general, this is a general concept - sacrificing one's own reproductive interests in order to increase the reproductive success of another. Although, of course, I understand this is all rather vague, people are interested in knowing where their moral instinct came from. And so, I think, it is useful to talk about such things.

Dmitry Gutov: The train of thought is interesting, maybe this is not your specialization, but if we were to radically sum up the results, then it was necessary to extend this concept to the inorganic world, that is, maybe physicists do this?

Alexander Markov: But I don’t quite understand this, because this metaphor of teleology, purposefulness, is applicable to living beings. Because, as I said, natural selection works in such a way that genes and organisms change as if they want something and aspire to something. Specifically, they seek to increase the efficiency of their reproduction. As if. So you can use these metaphors. They “want” this, but all this, of course, is in quotation marks. It's all automatic. That is, they have a goal - to leave as many descendants as possible. What is the purpose of, say, inorganic objects if we begin to apply the concepts of altruism and egoism to them? For living beings, altruism is sacrificing one's goal in order to help another achieve that goal. This is clear.

Alexander Markov: Yes, although the goal - in biology is also only our metaphor. In fact, there is no goal in biology either.

Dmitry Gutov: That is, you do not see the possibility of logical propagation more deeply.

Boris Dolgin: Let's say crystals?

Alexander Markov: First of all, I never thought about this topic. Secondly, at first glance, I do not see how.

Dmitry Gutov: In any case, the train of thought, if we go to bacteria, requires continuation, of course.

Olga: I have a more biological question. Please tell me a little more about genes for altruism. How can the fact that these genes for mutated vasopressin and oxytocin receptors be related to the functions of these hormones be related.

Alexander Markov: So you mean the genes that a person has?

Olga: Yes.

Alexander Markov: How is it customary for you, I can talk about them for an hour?

Boris Dolgin: Approach wisely. There are others who clearly want to ask questions, but at the same time, somehow try to answer.

Alexander Markov: This is a very interesting topic. Just a great topic.

Boris Dolgin: You can send to work.

Alexander Markov: Oxytocin and vasopressin are such neuropeptides, such small protein molecules that are secreted by certain neurons of the brain, hypothalamus and they serve as signal substances, there are a lot of signal substances in the nervous system, but these oxytocin and vasopressin are specialized mainly to regulate social and sexual relations, relationships between individuals. Moreover, this is a very ancient signaling system. All animals have these neuropeptides, and in all animals they do just that - they regulate social relations and relationships between individuals. I scroll in my head what to choose now for the story. For example, there is such a wonderful object - American voles, which in one genus have monogamous species, that is, they form strong marriage pairs, the male is actively involved in caring for offspring, and there is attachment between the male and the female for life. There are polygamous species, where there are no such stable relationships between males and females, and males do not participate in caring for offspring. It turned out that the difference in behavior between these species depends to the greatest extent on the variability of the vasopressin receptor gene. Receptors are proteins that sit on the surface of neurons and respond to something, in this case vasopressin. Vasopressin is a signaling substance, and a receptor is a protein that reacts to this vasopressin - and, accordingly, the neuron is excited. And it turned out that by changing the work of the gene of this very vasopressin receptor, even a male from a polygamous species can be forced to become a faithful husband, that is, so that he forms a strong attachment, a lifelong love for one female. Until recently, they did not know if a person has the same thing. It turned out that it still exists. Of course, we have the same vasopressin receptor gene, so we started looking at the variability, polymorphism in this gene, and whether the polymorphism in this gene correlates with any aspects of personality. And it turned out that yes, it correlates. In men who have one of the variants of this vasopressin receptor gene, firstly, the occurrence of a romantic relationship with a girl leads to marriage half as often as in all other men. And if they do marry, they are more likely to be unhappy in family life. And the wives of such men are almost always dissatisfied with family relationships. And the gene is the same as in voles, it affects marital fidelity, marital affection. Here it is already difficult to doubt that a person has the genetic basis of such things, say, as love between spouses. In addition, the variability of the genes for oxytocin and vasopressin receptors turned out to correlate with such qualities as kindness and generosity. This is checked, for example, in various economic games. And there are a lot of experiments going on. They put oxytocin in their noses and watch their behavior change. It works very well for men. So that, for example, they begin to better understand the facial expression of the interlocutor, look into the eyes more often, and so on. That is, it is clearly absolutely perfect that kindness, sensitivity - all this very much depends on the oxytocin-vasopressin system.

Olga: Variations in receptors that lead to, say, changes in the vole population, do they bind better to the hormone or worse?

Alexander Markov: There's a level of expression, now I'll try to remember. In one case, there are simply more of these receptors, the gene expression is higher, and in the other, less. But in which, to be honest, I don’t remember, I’ll have to look.

Alexander: Tell me, please, if we return to bacteria, are altruism and selfishness permanent characteristics of individuals, or are they temporary and cases of re-education are known - or, conversely, do some bacteria “go astray”? And what are the criteria for transition from one to another? Or were they born and left?

Alexander Markov: It is very difficult to register such cases that a bacterium “re-educates” during its lifetime, changes its behavior, even if they exist, it is not clear how.

Alexander: What if you take it higher?

Alexander Markov: That is, a mutation occurs - and then the behavior changes. But that will be in the next generation.

Alexander: And if we take not bacteria, but other organisms?

Boris Dolgin: That is, at what level does the variability of behavior arise, as I understand the question, within the framework of the life of one and the same organism? Did I understand the question correctly?

Alexander: In particular, yes, if it is not clear how to fix this process in bacteria, then what about others?

Alexander Markov: And animals can, of course, modify their behavior depending on the circumstances. But again, always following Hamilton's formula. I was talking about the wasp: as the wasp's chances for kingship grow, she works less and less and more and more shifts this matter to others. That is, the degree of altruism in her behavior is reduced, because she understands that she ought to take care of herself, otherwise the wings fray, you will still die.

Question from the floor: That is, she dissolves her waist, is preparing to become a uterus?

Alexander Markov: Yes.

Valeria: If there are bacteria representatives of two types of altruists and egoists, it turns out such a kind of consumer society. If there is a tendency to education, that is, to an increase in altruists, then everyone gets the same genes, it turns out such a kind of communism, and there will be no incentive for any progress if everyone is the same, then what if this really happens with human society? Will there be any aspiration for the transition of world domination in Asia, if there is such a thing? As you know, they are prone to repetition. The Chinese - they also copy some inventions.

Boris Dolgin: And what does this have to do with the question of evolutionary ethics?

Valeria: Is a society of altruists possible in the world? What will happen if there are altruists instead of egoists? Because, I think that there is some kind of world symmetry, and there should be a counterbalance to good, some kind of evil. Will there be ballast?

Alexander Markov: Here, most likely, balancing selection will work. That is, these are frequency-dependent things: the more altruists are around, the more profitable it is to be an egoist among them. If almost all are altruists, and I alone am an egoist, can you imagine, everyone will help me. Very profitable. And in this situation, egoists begin to multiply rapidly, to infect this population. Then there are a lot of egoists, no one helps anyone anymore. Only altruists work there, in their garden, isolated, and everyone walks around and asks for help. In this situation, when there are very few altruists left, one of two things will happen: either the altruists will finally die, and then the whole system will die. This is called in evolutionary ethics the Tragedy of Common Grazing. This is a situation when the village has a common pasture, everyone grazes their sheep there and overgrazing there, the pasture is depleted. It is necessary to reduce the number of sheep grazing, but every peasant thinks: let the neighbor take away his own, and I will graze mine anyway. And everyone is only interested in grazing as many of their sheep as possible. It ends with the fact that the pasture is finally destroyed, and all the peasants die of hunger. But even when they are already dying of hunger, already half have died, still the most profitable strategy for every peasant to the very end is to graze as many of his sheep as possible on the last blades of grass. In this situation, everything dies. But often, thanks to all sorts of different tricks, for example, statistical paradoxes or the fact that the altruist takes anyway, bypassing the common boiler, a certain balance is established. That is, with a certain number of egoists, it turns out to be more profitable to be an altruist than an egoist. Also, of course, intergroup enmity is a very powerful means of preserving intragroup altruism.

Svetlana: It seems to me that the lecture is quite long and somewhat interesting, but you are banal: kindness, altruism and other socially useful qualities of people are partly hereditary, genetic in nature. And all?

Boris Dolgin: In general, this is not a banal thesis at all.

Svetlana: And let's say, from the simplest to children, everything. We don't go any further. And it’s so interesting, but how is it today, a person, an individual, a group? Right now, today, such as we are, countries. How to call altruism and egoism in this sense?

Boris Dolgin: This question should be asked to psychologists. Thank you.

Svetlana: But the fact is that we say: it is interesting to see the evolutionary origins. And for what? We live now, today, among people - and just understand: does altruism and egoism have a genetic nature?

Boris Dolgin: You can leave without comment, or you can try to answer.

Alexander Markov: I'd rather leave it without comment.

Vladimir: If everything is more or less clear with Hamilton's formula, then I have a question about indirect reciprocity: every time an individual has a chance to perform any action that affects the reputation, does the individual measure the risk of death?

Alexander Markov: Of course, not every time, in general, this is a rather large rarity, that is, indirect reciprocity is a reputation mechanism. In humans, it is well developed, in birds, and perhaps a little bit in some higher primates. Of course, such very smart animals, they have a very complex behavior, which depends on a lot of different factors, and, of course, they will behave differently in different situations. Of course, they usually remember their interests and the preservation of their lives.

Zuhra: Once again I want to return to children, psychology, since you have already spoken about it. Caring for talented children, and for me altruism is a moral talent. Are there any tests that serve to measure altruism in children? You talked about such experiments with children, can you elaborate? Exist or not?

Alexander Markov: Yes. Lots of stuff.

Zuhra: Can their talent be measured?

Boris Dolgin: Forgive me, so far we are not talking about talent, but about altruism.

Zuhra: About altruism - yes, but for me it is the highest talent.

Maria Kondratova: You brought up a rather interesting topic - gender differences in altruism - when you talked about these models associated with the evolution of the Paleolithic. In connection with the different evolutionary strategies of the female and male sex, is it possible to talk about differences in altruisms: male and female, are there any studies on this topic? And to the question about the polymorphism of these genes. You say that there is a polymorphism that correlates with different behavior within the same sex, but are there any correlations between the sexes in vasopressin-oxytocin receptors that determine altruism?

Alexander Markov: Somehow, in humans, this is usually specific to one of the sexes - the influence of these genes, and the influence of these neuropeptides themselves, is different for men and women. Correlation between the sexes? I don't remember anything specific about this.

Boris Dolgin: That is, you partially indicated the correlation between gender and this factor. I understand that the question was in continuation of this topic. Are there any other gender differences? Of course, I would not talk about gender, because gender is a social gender.

Alexander Markov: So there are some gender differences?

Boris Dolgin: Yes, in relation to this very altruism.

Alexander Markov: I don’t know, probably, psychologists are actively studying this, I just, to be honest, I don’t know.

Boris Dolgin: There are works by Geodakyan, but they, in my opinion, are not substantiated in any way.

Alexander Markov: Yes, these are debatable things. Therefore, it is difficult to answer.

Konstantin Ivanovich: I would like to say that altruism, civilization is the number of charitable societies and the resources that rotate in these charitable societies. Is it interesting to compare, say, America, Russia, China, Sweden, Germany?

Alexander Markov: Not everything is so clear either.

Question from the floor: Do bacteria have such societies?

Alexander Markov: Charity?

Question from the floor: Yes.

Alexander Markov: In a sense, when they secrete some kind of socially useful substance.

Dmitry Ivanov: Do you agree with the theory of the selfish gene, that it makes sense to consider natural selection not on groups, not even on individuals, but at the level of genes. That it is precisely each gene that is interested in continuing, copying itself as an elementary replicator that has this ability?

Alexander Markov: If you heard the beginning of the lecture, you probably noticed that I build everything on this gene-centric approach. Of course, I admit it just works. It just is. The kin selection theory is a gene-centric approach.

Dmitry Ivanov: Thus, the gene for altruism... is difficult to survive. That is, it can manifest itself only in social societies, that is, only in society?

Alexander Markov: Naturally, if you do not have a society, if you live alone in a big forest, then what kind of altruism, if there is no one to show it to? This is clear.

Dmitry Ivanov: There is a lot of competition for resources in society, that is, we have a primitive society where various groups compete with each other. There is such a society of general welfare, where all supposedly altruists and help each other. Is it possible to be an altruist in such a society?

Boris Dolgin: What are such societies?

Dmitry Ivanov: If you think hypothetically. Do we want such a society? It turns out that in such a society these very deceivers can spread until the number of altruists again reaches a critically small level and fierce competition for all resources begins again. Is it logical?

Alexander Markov: And what is the question? I did not quite understand.

Dmitry Ivanov: The issue is the distribution of those genes of altruism in the human environment.

Boris Dolgin: Do you think that a stable social situation is possible when this gene wins? Did I understand the question correctly?

Dmitry Ivanov: Yes, that it is possible to do this only through education and through the development of culture, and not through natural selection?

Alexander Markov: The altruism that arises through the upbringing and development of culture faces exactly the same problems. As among unconscious beings, some bacteria, in this situation it is beneficial to be an altruist - in this situation it is not beneficial to be an altruist. It is the same in human society - even if we assume that there is no genetic variability in these traits, that altruism or selfishness of a person depends only on education. Let's say. All the same, in one situation it will be beneficial to behave altruistically, and in another - selfishly. For example, the more altruists around, the more profitable, the more tempting to take it and start behaving like an egoist. Since people are sentient beings, actively adapting during their lives, changing their behavior, the same problems arise.

Dmitry Ivanov: It turns out that this is the so-called reasonable egoism?

Alexander Markov: The ideal is, of course, when it is personally profitable for everyone to behave well. The ideal of reciprocal altruism is what we should probably strive for. The golden rule of ethics, it is not accidentally called the “golden rule”, people have long understood that it is on this basis that one must live.

Dmitry Ivanov: Treat others the way you want to be treated?

Alexander Markov: Yes.

Dmitry Ivanov: Another little question about children. How did the influence of culture differ from the influence of genes in the experience with children? That is, the influence of upbringing received from parents? The fact that he wants to share with others is not because his mom and dad raised him that way?

Alexander Markov: But in this experience, no way. In this experiment, the genes were not touched, they simply studied the behavior, how it changes with age. How it changes with age, how the percentage of certain patterns of behavior changes. Unaccountable altruism, the desire for equality and so on. In this particular study, the genes were not touched.

Grigory Chudnovsky: If possible, a short discussion and in this sense is not a question - if you consider it necessary to comment. Hamilton's equation, which you highlighted on the screen, both in single version and in multiple version, is an exact proportion, which is regulated not by the brain, but by other mechanisms in simple organisms and communities. The exact proportion of what I am willing to lose by giving someone else that important thing he needs. And there you can see that there is a limit, that even in this inequality there is a limit to what to transmit. That is, some quantum that can be transferred to save the life of the organism. I was interested in this inequality in this limit state. To what extent has it been studied? That is, some experiments, explanations, where a clear boundary is given. And the last thing to this question, for example, in civilized societies, where you finished your religious lecture, including fragments that you didn’t expand, it points out that any liturgical practice and ceremonies that are so expensive are, as it were, a form of altruism, as I understand it. . It seems to me that a form of psychic suppression, the more expensive and complex the procedure, the more selfish.

Boris Dolgin: This is a little different.

Grigory Chudnovsky: Yes, it's a little different. But I'm just now on this topic, that for example, a donation is calculated - this is altruism, right? Give a coin to the poor. But they are calculated, because the rich will become poor if he distributes to everyone who asks. This is in addition to the first question, where are the boundaries between altruism, which is good both socially. Thank you.

Alexander Markov: In order not to get confused, I will first answer the first question: where is the border? It's just that everything is written here, there is no special additional essence here. That's the whole border, it's here, that's the inequality. Now if rB>C, the gene of altruism will spread. Note that if rB<C, then the gene of selfishness will spread. This rule is retroactive. If your With much more than yours rB, then you will not save your own brother, but will gnaw at his throat, as a result of the action of natural selection automatically. This is observed, for example, in the chicks of many birds. Siblicide - this is called - the murder of siblings. Some birds are able to feed only one chick, but lay, just in case, two eggs. The first chick has hatched; if he is alive by the time the second hatches, then he will peck the second chick - or throw it away. This is the norm for them. Because, in this case, obviously, their price of saving the life of their brother turned out to be much higher than this case. That is, if we consider an altruistic act as a non-murder of a brother. That is, it all depends on the ratio of these variables. And that's all. And no mysticism. And the second question about religion, I already safely forgot. There was something interesting there, and I wanted to say something.

Boris Dolgin: The second question is, do you think that religious practices are a manifestation of altruism? In my opinion, in your lecture it sounded completely different?

Alexander Markov: Not a manifestation of altruism, but you said, can suppress the psyche?

Grigory Chudnovsky: Yes, that's what they're made for.

Alexander Markov: And there is no contradiction here. It may very well be that the suppression of the psyche can just contribute to the manifestation of parochial altruism, that is, selfless devotion to one's own, the readiness to die for one's faith, for one's fellow believers.

Alexander Nikitin: It seems to me that this model: talking about human society is fundamentally unsuitable, because a person is fundamentally different from the animal and biological world. He has consciousness, he has goals and tasks, besides multiplying, still creative. Therefore, an example can also be illustrated, according to this model, altruists and egoists. But according to this model, all people who set themselves some kind of high goal, unlike primitive altruists, fall into the category of deceivers. Because those altruists do not understand what their task is. They wanted them to just dig in the ground with a shovel next to them, that's all. And these people set themselves for some reason, by virtue of some forces, higher, maybe other goals. To write poetry like Pushkin - but from the point of view of primitive Darwinists - these are just deceivers. And this black and white model, it seems to me, is fundamentally unsuitable.

Alexander Markov: When complex objects are being studied, you always need to take into account a bunch of everything, a bunch of specifics of everyone. Naturally, some methodological approaches can be applied correctly to an object and can be applied incorrectly. It is clear that no one is going to take it in the forehead - and to any situation: someone digs, someone writes poetry - no one applies this formula like that, of course. It is clear that everything is much more complicated. This is a general saying, in life everything is more complicated than in your model. This is a universal refutation of any scientific research in biology.

Lev Moskovkin: I did not expect to hear something new for myself, very grateful. I listened to a lecture about this back in the academic years of 66-67. Whatever you call human exclusivity, I will give an example that it is not. This seems obvious. And I will never agree with the very common thesis about the slowness of human evolution, but this is not the topic of today's lecture. Geodakyan's ideas are absolutely conclusive. Unlike the ideas of Efroimson, they are simply proved in such a way that they are little understood, and the question is not connected with this. And immediately the most interesting question for me. After all, a selfish gene - what is meant, and is all this elegant theory of altruism and selfishness applicable to media viruses that circulate in the public infosphere, Dawkins, if I'm not mistaken, called them memes, and there was an excellent lecture, by the way, in Bilingual " more. If everyone is so politically correct, then how to explain then the Anglo-Saxon national egoism, and this is an extremely painful issue now, for our world. And the last thing - were there any searches and studies of "genes of altruism" before Vladimir Pavlovich Efroimson? What is important, I have come across the fact that many journalists are not even aware of the phenomenon that this gene of altruism circled the globe several times.

Alexander Markov: Last time I was asked two questions in a row, and now you have asked four. I would still prefer it to be on one issue. Here the first question was: what is a selfish gene - it is necessary to read a separate lecture. There is a book by Dawkins, The Selfish Gene, which popularizes this. I built my entire lecture on this model. I'm not ready to put it into words right now.

Boris Dolgin: Thank you. The next question was: to what extent did Efroimson play a role in the development of the concept?

Alexander Markov: Darwin himself began to think about this topic. He already built the first hints of the theory, then Fisher developed this topic, then Haldane - this was the beginning of the 20th century. So all these ideas have been developing for a long time.

Boris Dolgin: The third question, I think, was: do you want to apply this to "media viruses"?

Alexander Markov: For memes, right? As you probably know, Dawkins wrote about the possibility of drawing an analogy between genes and information units of cultural inheritance, perhaps, which also behave somewhat like genes. They are also selected, mutated, distributed. Say, jokes, some popular pictures, songs, melodies, some sayings, buzzwords, such things - they also spread in part the same way as genes, like viruses in a population. But is it possible to apply the concept of altruism and egoism to them? I think it will be difficult, because with genes, why does it work like that? I said that a gene cannot be altruistic. An altruistic gene is what it is - it would be a genetic variant that would sacrifice its own propagation to help another competing genetic variant to spread. What will happen to such an altruistic gene - it will simply disappear automatically, it will be forced out. Therefore, this cannot be. Altruism arises from the fact that the interests of genes and the organisms in which these genes sit do not coincide. The body can be altruistic. Gene can't. And what is the equivalent of an organism for a meme? I do not quite understand this, this theory is not very well developed.

Boris Dolgin: Well, maybe a tradition?

Alexander Markov: The complex of genes is engaged in what it creates, builds an organism from a fertilized egg. And the meme complex, what does it do?

Boris Dolgin: I am against this metaphor, but if we proceed from it, then it is a tradition.

Alexander Markov: It's hard, you have to think.

Evgeny Teslenko: Thanks a lot for the lecture. To be honest, I got a little scared. Since, if we extend the scientific logical trend, then the question arises: with the modern developments of genetic engineering, it is quite possible that some theories will appear, and then practices of correcting the human essence with a great wish to increase altruists, for example, to reduce egoists.

Boris Dolgin: In continuation of eugenics?

Evgeny Teslenko: Yes, yes, absolutely right, we are returning to the same eugenics, returning to more rational forms of the structure of mankind, and so on. How do you feel about this, especially since both scientific and technical developments have already come close enough. Why is it your lecture that makes this trend scary? Because it seems - yes, scientific and technological progress cannot be stopped, there will still be research. But why can they be good or bad? Why do they invade the realm of morality? Because, and you yourself showed it perfectly at the beginning, that the words, terms, metaphors that are hung up are an altruist. Well, what kind of altruists are these, what kind of egoists are there? Maybe it makes sense for scientists who are engaged in fundamental research to approach such metaphors very carefully? Because they are tempting. What do you think about this?

Alexander Markov: What temptation?

Evgeny Teslenko: The temptation to use and correct entrusted people in the right direction.

Boris Dolgin: Do social genetic engineering?

Evgeny Teslenko: Not social, but technical genetic engineering

Alexander Markov: Here the temptation is not because of the metaphors. When it comes to people, then altruistic, egoistic behavior is no longer a metaphor, but already quite what was originally called that. If we see that changes in a certain gene affect the tendency to do good deeds, we are talking about good deeds, and not about the secretion of some kind of enzyme by yeast.

Reply from the audience: In fundamental science, the phrase "good deeds" is very strange.

Alexander Markov: Of course, formal definitions are given there. It's just long and boring. Naturally, they are in the articles.

Boris Dolgin: The question was whether you are afraid of the social consequences of the activities of this scientific direction. If I understand the question correctly.

Alexander Markov: This is, of course, a difficult question. Humanity will have to face this. Of course, it seems to us now that it is impossible to genetically modify a person so that he becomes more kind. But this seems unethical. Let's start from the other end, and if we are talking about hereditary diseases? For example, parents are told: you will have a child with a severe hereditary disease.

Boris Dolgin: With a certain percentage of probability?

Alexander Markov: Maybe with a certain percentage of probability, if before conception, or when there is already an embryo. We can do gene therapy. We can inject viruses into it, and the necessary genes will be inserted into the cells, and we will fix it, and then your child will most likely be born healthy and normal. Well, of course, parents will agree to this. Depriving parents of the opportunity to make such a choice is also wrong. And if there is no genetic disease? But the genetics of the future simply tell parents: your child has such an allele of the vasopressin receptor gene that he will almost certainly be unhappy in family life, he has an unsuccessful option, he cannot feel sympathy, he will not have a good family (with such and such a probability) . We can insert viruses into him now, genetically modify him, the necessary genes will be inserted into his brain, and then he will be happy in family life. Take your pick, fellow parents. This is a more difficult question. Yes, I do not undertake to decide.

Boris Dolgin: Yes, but still, we will clarify that, as you said today, for a modern person, the moment of culture, the social moment turns out to be at least no less significant.

Alexander Markov: Naturally.

Boris Dolgin: That is, there is always hope for re-education (in the broadest sense).

Alexander Markov: In such cases, when such a sharp effect as with these alleles of the vasopressin receptor is, of course ... Well, how do you raise a boy? I have three sons, how will you raise him so that he is happy in family life?

Sergey Kapustin: I have two questions. The first is an explanation about ant farming. Why do they need mushrooms to be genetically homogeneous? So that they don't become poisonous, for example? Were they edible?

Alexander Markov: These mushrooms behave, as it were, altruistically towards ants. The mushroom has different options. In principle, if these mushrooms were engaged in selfish evolution, inside these anthills or termite mounds, then deceiver mushrooms would necessarily appear there, which would only exploit the ants, but feed them poorly or not feed these ants at all. Well, for example, mushrooms that live in termite mounds form two types of fruiting bodies: small, round fruiting bodies to feed termites, and large, stalked fruiting bodies that grow through the termite mound and disperse spores. That is, small fruiting bodies are, roughly speaking, altruism for termites that feed, grow, and spud them. And large fruiting bodies are like selfishness - a mushroom makes for itself. Accordingly, what will happen if a mutant fungus appears that uses more energy for the production of large fruiting bodies and less energy for the production of small fruiting bodies? If these fungi are allowed to compete calmly, to evolve inside the termite mound, the egoist will win, the fungus that will give more large fruiting bodies will win, and the termites will remain “with a nose”. They will have less food. So that this does not happen, that there is no such competition between different strains of fungi with a different number of such and such fruiting bodies, for this it is necessary that they be genetically identical. Then they won't evolve.

Sergey Kapustin: And the second question, we summarize different arguments about memo-viruses, such cultural phenomena. How do you feel about this idea: in principle, evolution is the distribution and reproduction of genetic information. The gene as a carrier of information "has a purpose" to replicate itself. Do not appear in the human environment, in any natural environment, at the informational level, other carriers? That is, a person is a carrier of information in a different non-genetic form, ideas, statements as one of the possible options, he tries to disseminate this information no longer in a genetic form, but in a cultural form, for example. And thus, some behavior that may seem altruistic for genetic reproduction may not be altruistic at all as the informational equivalent of reproduction.

Boris Dolgin: Unfortunately, the question is not entirely clear. Or do you understand?

Alexander Markov: No, unfortunately I don't understand either.

Boris Dolgin: Undoubtedly, people usually tend to spread their ideas. But what is your question?

Sergey Kapustin: Is there any analogy here, any research on the fact that there is reproduction of genes, replication of genes, there is replication of information in a different form - not genetic. Somehow similar to the process of evolution ... The question is: is altruism, for example, in humans and altruism in general in nature, some first step towards getting away from the genetic, that is, they sacrifice their genetic in favor of alternative replication, for example, the idea of ​​altruism .

Boris Dolgin: And how do you imagine the mechanism of scientific verification of this hypothesis?

Sergey Kapustin: This is hard, I guess.

Alexander Markov: It's just such an interesting opinion.

Maria Kondratova: Since we know people who sacrifice their lives and their reproductive capabilities for the sake of some ideas, apparently, this makes sense for a person. My question is not about that. I really liked that you included a point in your report that a genetic, evolutionary description does not mean justification. Because, unfortunately, it is very often replaced. If there is something in our nature, then it must be so, this is the most common, trivial judgment, but such a question arises not biological, but more general: what, then, can be a justification for today, when religious authority is no longer a justification, human nature, a scientific description is a description, but also not a justification, and what, then, can be a justification?

Boris Dolgin: Maybe your value system? For you - yours, for Alexander - his.

Maria Kondratova: Then the concept of altruism is lost as a common good, as something specifically common.

Boris Dolgin: But the system of values ​​is more or less common for some communities. And the idea of ​​the common good is still nothing more than a part of this system of values.

Alexander Markov: But this question, of course, is not for biologists. It seems to me that biology should not, biology can explain why we have such or such instincts, innate inclinations, but it is not our business to decide what is good for a person now and what is bad.

Reply from the audience: Therefore, it was not necessary to talk about people today!

Alexander Markov: Disagree.

Boris Dolgin: We were going to talk about people from the very beginning, this is also in the topic of the lecture. So we knew what we were getting into.

Irina: Thank you for a very interesting lecture. I wanted to ask you, as a biologist, in what direction is biology going to develop, what will the money be invested in?

Boris Dolgin: What they invest in, I'm afraid, this is not entirely for a biologist.

Irina: Do you have any information on the basis of everything that you told us, as the abstracts of the previous large materials. What are the prospects?

Boris Dolgin: In other words: what areas of biology seem most interesting to you, and where would you advise to invest money or what to pay attention to?

Alexander Markov: There is an opinion - not mine, but I want to believe that it will be so, that just as the 20th century is sometimes called the century of genetics, the 21st century, perhaps, will be the century of neurobiology - the study of the brain. And, indeed, there are very encouraging results in this direction in understanding the mechanisms of the brain of animals, including humans. Maybe by the end of the 21st century we will understand how it all clicks for us, how thoughts are formed, feelings, and so on.

Reply from the audience: This is good?

Alexander Markov: Man knows himself.

Victor: The position that we have heard, the work has both practical and any other significance. They are all displayed on the site - your provisions, everything is written on the site? Is there a full lecture?

Boris Dolgin: I will give a partial answer right away, and Alexander can answer for his part. The transcript of this lecture will be posted along with the video on the Polit.ru website. And now Alexander, apparently, will talk about other forms in which you can get acquainted with the provisions of the report.

Alexander Markov: Actually, this report in an expanded form, twice as long as I told you, has been hanging on my website for five months (evolbiol.ru/altruism.htm). I went to one conference, I reported it there, and then I posted almost everything on the Internet. A significant part of what I just said is already on the Internet, on my website. Website "Problems of evolution" www.evolbiol.ru.

Question from the floor: There was such a three-volume work, published even before the revolution, - "The Nature of Love." It examines the evolutionary process in great detail, from bacteria to man, from the point of view of both altruism and all other categories.

Boris Dolgin: Before the revolution of 1917?

Question from the floor: Certainly. So tell me, please, did you rely on this work to some extent?

Question from the floor: Bailey.

Alexander Markov: No, I don't know him.

Contrary to popular misconception among lay people, modern evolutionary biology is successful in explaining the origin of morality and altruistic behavior. Cooperation, mutual assistance and self-sacrifice are not unique to humans: they are found in many animals and even microorganisms. As in human society, the altruism of some individuals creates an ideal breeding ground for the selfishness of others. The article discusses the results of experimental and theoretical studies of recent years, shedding light on the evolution of cooperation and altruism in bacteria, unicellular eukaryotes and animals, including humans.

Evolutionary ethics is a relatively young area of ​​biological research, moving along which biology invades the "forbidden" territory, where philosophers, theologians and the humanities have so far reigned supreme. The central question of evolutionary ethics is the question of the origin and evolution of cooperation and altruistic behavior.

“Altruism” in biology is understood as behavior that leads to an increase in the fitness (reproductive success) of other individuals to the detriment of their own chances of successful reproduction. This definition essentially differs little from the definitions of altruism accepted in ethics, given the fact that the action of natural selection in the general case is aimed precisely at increasing reproductive success. This allows us to speak metaphorically about it as the main "goal" in the achievement of which evolving organisms are "interested". Of course, we are only talking about the fact that the changes automatically undergone by organisms under the influence of natural selection, as a rule, lead to increased reproductive success. In other words, if organisms had a conscious goal of maximizing their reproductive success, and if they could consciously influence their own evolution, then the direction of evolutionary changes would be exactly what is observed in reality. It is in this somewhat metaphorical sense that concepts such as "goal" and "interest" are used in evolutionary biology.

Biologists studying the origins of cooperation and altruism face two main questions. It is obvious that almost all vital tasks facing organisms are easier to solve by joint efforts than alone. Cooperation, i.e., joint problem solving, usually involving some degree of altruism on the part of the cooperators, could be the ideal solution to most problems for many organisms. Why, then, did the biosphere never turn into a realm of universal friendship and mutual assistance?

The second question is the opposite of the first. How can cooperation and altruism arise in the course of evolution at all, if the driving force behind evolution is the mechanism of natural selection, which at its core, it would seem, is purely egoistic? A primitive, simplified understanding of the mechanisms of evolution can lead to an absolutely wrong conclusion that the very idea of ​​altruism is incompatible with evolution. This is facilitated by such, in my opinion, not very successful metaphors as “struggle for existence” and especially “survival of the fittest”. If the fittest always survives, what kind of altruism can we talk about?

The error of such reasoning is to confuse the levels at which we consider evolution. It can be considered at the level of genes, individuals, groups, populations, species, communities. But all evolutionary changes are recorded (remembered) only at the level of genes. Therefore, it is from the genetic level that consideration should begin. Here, evolution is based on the competition of different variants (alleles) of the same gene for dominance in the gene pool of the population. At this level, there is no altruism and, in principle, cannot be. Gene is always selfish. If an “altruistic” allele appears, which, to its detriment, allows another allele to multiply, such an “altruist” will be automatically ousted from the gene pool and disappear.

Relatedselection

However, if we look from the level of competing alleles to the level of competing individuals, the picture will be different, because the interests of the gene do not always coincide with the interests of the organism (see above about the metaphorical sense that evolutionary biologists put in the concept of "interest"). The discrepancy between interests stems from the discrepancy between the material nature of these objects. An allele is not a single object: it is present in the gene pool in the form of many copies. An organism, on the other hand, is a single entity, each cell of which carries, as a rule, only one or two of these copies. In many situations, it is advantageous for a selfish gene to sacrifice one or two copies of itself in order to provide an advantage to the rest of the copies that are contained in other organisms.

Biologists began to approach this idea already in the 30s of the XX century. R. Fisher (Fisher 1930), J. Haldane (Haldane 1955) and W. Hamilton (Hamilton 1964) have made important contributions to understanding the evolution of altruism. The theory they built is called kin selection theory. Its essence was figuratively expressed by Haldane in the well-known aphorism: "I would give my life for two brothers or eight cousins." What he meant by this can be understood from the following formula (known as "Hamilton's rule"). A gene for altruism (more precisely, an allele that promotes altruistic behavior) will be supported by selection and spread in a population if:

rB > C,

where r – the degree of genetic relationship between the “donor” and the “acceptor” (it determines the probability that the genome of the latter has the same “altruism allele”); B - reproductive advantage received by the addressee of the altruistic act; C - reproductive damage caused by the "donor" to himself. Reproductive advantage or disadvantage can be measured, in particular, by the number of produced (or not produced) offspring. Taking into account the fact that not one, but many individuals can benefit from an act of altruism, the formula can be modified as follows: nrB > C, where n - the number of those who accept the sacrifice.

It should be emphasized that Hamilton's rule does not introduce additional entities and is not based on any special assumptions. It follows logically from the basic facts and models of population genetics. If a nrB > C, the allele of altruism will purely automatically, without any external guiding forces, increase its frequency in the gene pool of the population.

From the point of view of the allele itself, there is no altruism in this, but only pure selfishness. In fact, this allele makes its carriers (organisms) behave altruistically, but in this way the allele watches over its “selfish interests”. An allele sacrifices several copies of itself to give advantage to other copies contained in the bodies of closely related organisms. Natural selection is an automatic weighing of the sum of gains and losses for an allele (for all its copies together!), and if the gains outweigh the allele, it spreads.

Hamilton's rule has remarkable explanatory and predictive power. In particular, it makes it possible to explain the repeated occurrence of eusociality in insects of the order Hymenoptera(hymenoptera). In eusocial hymenoptera (ants, bees, wasps, bumblebees), most females give up their own breeding to help the mother raise other daughters. Apparently, an important factor contributing to the development of eusociality in this order is the haplodiploid mechanism of sex inheritance. In Hymenoptera, females have a double set of chromosomes and develop from fertilized eggs. Males are haploid (have a single set of chromosomes) and develop from unfertilized eggs. Because of this, a paradoxical situation arises: sisters turn out to be closer relatives than mother and daughter. In most animals, the degree of relationship between sisters and between mothers and daughters is the same (50% of common genes, the value r in Hamilton's formula is 1/2). In Hymenoptera, siblings share 75% of their genes (r = 3/4), because each sister receives from her father not a randomly selected half of his chromosomes, but the entire genome. Mother and daughter in Hymenoptera have, like in other animals, only 50% of common genes. Therefore, for the effective transfer of their genes to the next generations, it is more profitable for female Hymenoptera, other things being equal, to raise sisters than daughters. Another factor in the development of eusociality in insects, not only in hymenoptera, but also in termites, is monogamy, which provides an abnormally high level of genetic relationship between individuals in the colony (Hughes etal. 2008).

Kin selection seems to underlie many instances of altruism in nature. However, in addition to kin selection, there are a number of mechanisms, some of which help, while others, on the contrary, hinder the evolution of altruism. Let's consider these mechanisms on concrete examples.

Altruists and deceivers among bacteria

Experimental study of the evolution of bacteria (“evolution in vitro”) is one of the promising areas of modern microbiology. Interesting results were obtained on bacteria Pseudomonas fluorescens, which, under the necessary minimum conditions, is capable of rapidly evolving in front of researchers, mastering new niches and developing original adaptations.

In order for the social system to be able to develop beyond the very first steps, it needs to develop a mechanism to combat deceivers. Such mechanisms are sometimes actually worked out. Often this leads to an evolutionary "arms race": deceivers improve their methods of deception, and cooperators improve ways to identify deceivers, fight them, or try to prevent deceivers from appearing.

The ability to defend against deceivers can come from single mutations

Consider another example related to bacteria Myxococcus xanthus. These microbes are characterized by complex collective behavior. Sometimes they gather in large clusters and arrange a collective "hunt" for other microbes. "Hunters" secrete toxins that kill "prey", and then absorb organic substances released during the decay of dead cells.

With a lack of food, myxococci form fruiting bodies, in which some of the bacteria turn into spores. In the form of spores, they can survive times of famine. The fruiting body is formed from many individual bacterial cells. The creation of such a complex multicellular structure requires the coordinated action of millions of individual bacteria, of which only a part directly benefits, while the rest sacrifice themselves for the common good. The fact is that only some of the participants in the collective action can turn into disputes and pass on their genes to the next generations. The rest act as "building materials", doomed to die without leaving offspring.

In this experience, the altruists failed to develop protection against deceivers. Something else happened: the deceivers themselves underwent a mutation, as a result of which the bacteria restored their lost ability to independently form fruiting bodies and at the same time gained an additional advantage (!). These mutant bacteria turned out to be protected from freeloaders, that is, from their direct ancestors - deceiver bacteria. Thus, a single mutation turned deceivers into altruists, protected from deception. The mutation occurred in one of the regulatory genes that affect the behavior of bacteria. The specific molecular mechanism of this effect has not yet been elucidated (Fiegna etal. 2006).

The trickster problem is also familiar to more complex single-celled organisms such as social amoeba. Dictyoste-lium. Like many bacteria, these amoeba, when there is a lack of food, gather into large multicellular aggregates (pseudoplasmodia), from which fruiting bodies are then formed. Those amoeba whose cells go to build the stem of the fruiting body sacrifice themselves for the sake of comrades, who get a chance to turn into spores and continue the genus (Kessin 2000).

It seems that the evolution of social bacteria and protozoa repeatedly began to move towards the formation of a multicellular organism, but for some reason things did not go further than plasmodia and rather simply arranged fruiting bodies. All truly complex multicellular organisms are formed in a different way - not from many individual cells with different genomes, but from the descendants of a single cell (which guarantees the genetic identity of all cells of the body).

As already mentioned, in order to survive, social organisms need to defend themselves from freeloaders. Experiments conducted on amoebas have shown that the likelihood of developing resistance as a result of random mutations in this organism is also quite high, as in myxococci (Khare etal. 2009). The experiments were carried out with two strains of dictyostelium - "honest" and deceivers. With a lack of food, they form chimeric (mixed) fruiting bodies. At the same time, deceivers occupy the best places in the fruiting body and turn into spores, leaving honest amoeba alone to build the stem of the fruiting body. As a result, disputes of deceivers predominate among the resulting disputes.

During the experiment, honest amoebas artificially increased the rate of mutation. Then, from the many resulting mutants, a thousand individuals with different mutations were selected and each of them was given the opportunity to multiply. After that, selection for resistance to freeloaders began, and freeloaders themselves were used as a selecting agent. Amoebas from a thousand mutant strains were mixed in equal proportions and combined with deceit amoebas. The mixed population was kept under conditions of lack of food, forcing the formation of fruiting bodies. Then the resulting spores were collected and amoebas were removed from them. Naturally, deceivers prevailed among them, but the experimenters killed all deceivers with an antibiotic (the gene for resistance to this antibiotic was previously inserted into the genome of honest amoebas). The result was a mixture of mutant amoebas, but of the thousands of original strains, it was now dominated by those who were best able to resist the deceivers. These amoebas were again mixed with deceivers and again forced to form fruiting bodies.

After six such cycles, only one of the thousand original strains remained in the mutant amoeba population. These amoebas were reliably protected from deceivers as a result of a mutation that occurred in them. Moreover, they did not defend themselves from any deceivers, but only from those with whom they had to compete in the experiment. Moreover, it turned out that these mutant amoebas protect not only themselves from deception, but also other strains of honest amoebas, if they are mixed. It is clear that the mutual assistance of honest strains opens up additional opportunities for combating deceivers.

These experiments were repeated many times, and each time resistance arose in one or another strain of amoeba-mutants, and different genes were mutated and different mechanisms of resistance arose. Some resistant strains themselves became deceivers in relation to "wild" amoeba, while others remained honest (Khare etal. 2009).

"Peaceful coexistence" of altruists and egoists

Another trick of this kind is called Simpson's paradox. Its essence is that under a certain set of conditions, the frequency of occurrence of altruists in a group of populations will increase, despite the fact that within each individual population this frequency is steadily decreasing. Let's say that in the initial population there were equally altruists and egoists. Then the population was divided into many very small subpopulations, in which the ratio of altruists and egoists varies greatly (with a sufficiently small size of subpopulations, the high variability of this ratio is provided by simple chance). In the course of the growth of each individual subpopulation, altruists are the losers (their share is reduced). However, those subpopulations that initially had more altruists grow faster due to the fact that they have at their disposal more of the “public good” produced by altruists. As a result, if you add together the grown subpopulations, it turns out that the "global" percentage of altruists has grown. The fundamental possibility of such a mechanism for maintaining the number of altruists was assumed by Haldane and Hamilton, however, it was only recently possible to obtain experimental evidence of the effectiveness of Simpson's paradox (Chuang et al. 2009). The main difficulty was that in each case, when we see the spread of “altruism genes” in a population, it is very difficult to prove that some other, unknown to us, benefits associated with altruism in a given species of organisms are not involved.

To find out if Simpson's paradox alone could make altruists thrive, a model system was created of two strains of genetically engineered E. coli. The genome of the first of the two strains (“altruists”) was supplemented with the gene for an enzyme synthesizing the N-acyl-homoserine-lactone signaling substance used by some microbes for chemical communication. In addition, the gene for an enzyme providing resistance to the antibiotic chloramphenicol was added to the genome of both strains. A promoter was attached to this gene, which activates the work of the gene only if the above-mentioned signaling substance enters the cell from the outside. Egoists differed from altruists in the absence of a gene necessary for the synthesis of a signaling substance.

Thus, the signaling substance secreted by the altruists is necessary for both strains for successful growth in the presence of the antibiotic. The benefit that both strains get from the signaling substance is the same, but only altruists spend resources on its production. Since both strains were created artificially and had no evolutionary history, the experimenters knew for sure that there were no “secret tricks” in the relationship between altruists and egoists in their model, and altruists do not receive additional benefits from their altruism.

In the medium with the addition of the antibiotic, pure cultures of egoists, as expected, grew worse than pure cultures of altruists (because in the absence of a signaling agent, the gene for protection against antibiotics in egoists remained turned off). However, they began to grow better than altruists if either live altruists or a purified signaling agent were added to the medium. Altruists in a mixed culture grew more slowly because they had to spend resources on the synthesis of a signaling substance. After confirming that the model system worked as expected, the researchers set about modeling Simpson's paradox.

To do this, they placed mixtures of two cultures in different proportions in 12 tubes with an antibiotic-containing medium, waited 12 hours, and then measured the number of bacteria and the percentage of altruists in each tube. It turned out that in all test tubes the percentage of altruists decreased significantly. Thus, altruists in all cases lost in competition with egoists. However, the size of those populations where initially there were more altruists grew much stronger than those where egoists predominated. When the authors summed up the numbers of microbes in all 12 test tubes, it turned out that the overall percentage of altruists increased markedly: Simpson's paradox successfully "worked".

However, in nature, no one will deliberately mix altruists with egoists in different proportions and put them in test tubes. What natural process can serve as an analogue of such a procedure? Apparently, this role can be played by "bottlenecks" - periods of a strong reduction in the population size with its subsequent restoration. This can occur, for example, when new substrates are colonized by a very small number of “founder” microbes. If the number of founders is small, then, due to mere chance, there may be an increased percentage of altruists among them. The population formed by this group of founders will grow rapidly, while other populations founded by selfish-dominated groups of microbes will grow slowly. As a result, Simpson's paradox will ensure the growth of the "global" share of altruists in the aggregate of all populations.

To prove the effectiveness of this mechanism, the authors mixed altruists with egoists in equal proportions, greatly diluted the resulting culture and began to sow it in test tubes in portions of various sizes with an approximately known number of microbes in each portion. Portion size turned out to be the main factor on which the further fate of altruists depended. As you might expect, when the portions were large, Simpson's paradox did not manifest itself. In a large portion, i.e., in a large sample from the initial culture, the ratio of altruists and egoists, according to the laws of statistics, cannot differ greatly from the initial one. Populations based on these samples grow at approximately the same rate, and altruists lose not only in each individual population, but in all populations as a whole.

However, if there were only a few bacteria in each portion, then among these portions there were necessarily those in which altruists predominated. Such founder groups gave rise to rapidly growing colonies, and due to this, the overall percentage of altruists in the aggregate of all populations increased. Under the specific conditions of this experiment, for the manifestation of the Simpson effect, it is necessary that the average number of microbes in the group of founders be no more than 10. The authors also showed that by repeating this sequence of actions several times (dilution of the culture, settling in small groups in test tubes, growth, connection of populations in one, again dilution, etc.), it is possible to achieve an arbitrarily high percentage of altruists in a culture.

Another condition was identified that is necessary for the spread of "altruism genes" in the model system: mixed populations should not be allowed to grow for too long. Dilution and settlement should be carried out before the populations reach a stable population level, populating the entire nutrient medium in the test tube, because then the differences in abundance between populations are smoothed out and Simpson's paradox cannot manifest itself (Chuang etal. 2009).

Thus, under certain conditions, natural selection can ensure the development of altruism even when it favors egoists in each individual population, and dooms altruists to gradual extinction. However, the range of conditions under which Simpson's paradox can operate is rather narrow, and therefore its role in nature is probably small.

Altruists and deceivers among social animals

The biggest triumph of the evolution of altruism was the emergence of true multicellular organisms, including animals. Animals, compared with microbes, have new opportunities for the development of cooperation and altruism, based on complex behavior and learning. But the same new possibilities opened up to the deceivers. The deceivers learned to deceive the co-operators more cunningly, and the co-operators, for their part, began to develop new methods for identifying the deceivers and fighting them. The evolutionary "arms race" continued at a new level, and again neither altruists nor deceivers gained a decisive advantage.

One of the important innovations in this endless war was the possibility of physical (not just chemical) punishment of deceivers. This phenomenon occurs, in particular, in social insects. Hymenopteran workers usually do not breed, devoting themselves to caring for the queen's offspring. The development of altruism in Hymenoptera is associated with kin selection (see above). However, in many species of Hymenoptera, workers are physiologically quite capable of reproduction, and sometimes they actually show "selfishness" by laying their own unfertilized eggs. Recall that in Hymenoptera, males develop from unfertilized eggs. Due to the nature of sex inheritance for female Hymenoptera, the most profitable strategy is to raise other people's daughters (your sisters) and your own sons. This is exactly how worker wasps of many species try to behave. However, these "unauthorized" eggs laid by workers are often destroyed by other workers, who thus act as a kind of "moral police".

Recently, German entomologists have tried to test which of the two factors is more important for maintaining altruism in an insect society: voluntary adherence to the principle of "reasonable selfishness", i.e. pure kin selection (1), or "police surveillance" (2) (Wenseleers, Ratnieks 2006). For this, data on 10 species of social Hymenoptera were processed. It turned out that the stricter the "moral police", the less often workers commit acts of selfishness, laying their own eggs. We also tested the influence of the degree of kinship between workers in the nest on altruistic behavior. The degree of relatedness between them is often less than the ideal 75% in reality, since a queen may mate with several different males. It turned out that the lower the degree of relationship between sister workers, the stronger the "police surveillance", and the less often the workers behave selfishly. This corresponds to the second hypothesis (about the leading role of police measures). With a low degree of relationship between workers, it becomes more profitable for them to destroy the eggs of other workers. A low degree of relatedness also makes selfish behavior more beneficial, but as can be seen from the results obtained, effective “policing” clearly outweighs the selfish aspirations of workers (Wenseleers and Ratnieks 2006).

Features of sex inheritance in Hymenoptera played an important role in the development of altruistic behavior and sociality, however, in many modern species, altruism is supported mainly not by the indirect “genetic benefit” received by workers from such behavior, but by strict “police control”. Apparently, the cooperative system created by kin selection, even under such "ideal" conditions that are observed in families of Hymenoptera, will still be destroyed by deceivers if it fails to develop additional means of combating egoism.

This pattern may be true for human society, although it is difficult to verify experimentally. Social life is impossible without altruism (the individual must sacrifice his interests for the sake of society), and in the end everyone benefits from this. However, in many cases it is still beneficial for each individual person to act selfishly, pursuing selfish interests to the detriment of the team. And in order to effectively combat egoism, one has to use violent methods.

Let us consider one more example showing that the altruism of social insects is far from the ideal of unselfishness. wasps Liostenogasterflavolineata live in families, including from 1 to 10 adult females, of which only one - the oldest - lays eggs, and the rest take care of the larvae. When the queen dies, the next most senior wasp takes her place. Outwardly, the helpers are no different from the queen, but they lead a much more difficult and dangerous life: if the queen almost never leaves the nest, then the helpers have to fly for food for the larvae, which is associated with wear and tear of the wings and the risk of being caught by a predator. With the transition of a helper to the rank of queen, her life expectancy increases dramatically (Field et al. 2006).

In this species, as in many others, helper wasps vary greatly in the degree of "labor enthusiasm." Some, not sparing themselves, spend up to 90% of the time looking for food, while others prefer to sit in a safe nest and fly out for food an order of magnitude less often. At first glance, these differences are difficult to explain from the point of view of the theory of kin selection, since the degree of labor enthusiasm of helpers does not depend on the degree of their relationship with the queen and the larvae they care for. However, as it turned out, each assistant strictly doses altruism, depending on how great her chances are to become a queen and leave her own offspring. If these chances are small (like those of low-ranking young wasps, the last in the “line” for the royal throne), then it makes sense to work more actively in order to pass on their genes to the next generations, even through other people's children. If the assistant has a high rank, it is more profitable for her to take care and take less risks.

This conclusion is based on the results of elegant experiments. From one family, a wasp that occupies the second place in the hierarchy (i.e., the first in seniority after the queen) was removed, and a low-ranking young wasp was removed from another family of the same size. After that, the behavior of the wasp, which, prior to the start of the experiment, occupied the third place in the hierarchy, was monitored. In the first nest, this wasp, after the removal of the senior assistant, increased its rank, moving from third place to second, and in the second nest, it remained in third place. The size of both families remained the same. It turned out that in the first case, the wasp starts to work about half as much. In the second case, when a low-ranking helper was removed from the nest, wasp number three continued to work as long as before (Field etal. 2006).

These results show that the amount of "altruistic effort" in wasps is indeed regulated by the wasp's chances of its own reproductive success. In other words, the propensity for altruism is stronger in those who have nothing to lose. The appearance of such behavior in the course of evolution is well explained by Hamilton's rule, if we take into account the fact that the quantity c, i.e., the price of altruistic behavior varies depending on the circumstances, including the chances for the "royal throne".

The genetic identity of cooperators prevents the appearance of cheaters

Is it possible to create a social order where altruism will be maintained without violence and at the same time there will be no deceivers and egoists? Neither wasps nor humans have succeeded so far. But some cooperative symbiotic systems that exist in nature indicate that in principle it is possible to prevent the very appearance of deceivers. To do this, it is necessary to reduce the genetic diversity of individuals in a cooperative system to zero. This excludes the possibility of competition between genetically different varieties of symbionts for which of them will exploit common resources more efficiently (grab a larger piece of the common pie). If all symbionts are genetically identical, selfish evolution within the system becomes impossible, because one of the components, namely variability, is excluded from the minimum set of conditions necessary for evolution - the Darwinian triad of "heredity, variability, selection". As a result, the evolutionary interests of the twin symbionts are automatically identified with the interests of the entire system. In this case, selection ceases to act at the level of individual symbionts and begins to act at the level of entire symbiotic systems.

That is why evolution never succeeded, despite repeated “attempts”, to create a full-fledged multicellular organism from genetically heterogeneous cells. All real multicellular organisms are formed from clones - descendants of a single cell.

If the cooperative system consists of a large multicellular "host" and small "symbionts", then the easiest way for the host to ensure the genetic identity of symbionts is to pass them vertically, i.e., by inheritance, and only one of the sexes should do this - either males, or females. This is how, for example, mitochondria are transmitted in all eukaryotes - strictly through the maternal line, and the mitochondria themselves reproduce clonally. Leaf-cutting ants also pass on their crops from generation to generation. With vertical transfer, the genetic diversity of symbionts is automatically maintained at a level close to zero due to genetic drift and bottlenecks.

There are, however, also symbiotic systems with horizontal transfer of symbionts. In such systems, symbionts in each host are genetically heterogeneous, they retain the ability to selfish evolution, and therefore deceivers appear among them every now and then. For example, strains of deceivers are known among luminous bacteria (fish and squid symbionts), nitrogen-fixing rhizobia bacteria (plant symbionts), mycorrhizal fungi, and zooxanthellae (coral symbionts). In all these cases, evolution failed to ensure the genetic homogeneity of symbionts, and the hosts have to deal with deceivers by other methods, for example, immunological, or simply tolerate their presence, relying on certain mechanisms that ensure a balance in the number of deceivers and honest cooperators. For example, on Simpson's paradox or on balancing selection, which is based on the fact that sometimes it is beneficial to be a deceiver only as long as the number of deceivers is not too high - otherwise there will be no one to deceive. All this is not so effective, but natural selection notices only momentary benefits and is completely indifferent to distant evolutionary prospects.

In order for a mechanism to ensure the genetic homogeneity of symbionts to evolve, that mechanism must provide an immediate benefit, or selection will not support it. The benefit that we have been talking about so far - depriving symbionts of the opportunity to evolve into deceivers - belongs to the category of "remote prospects" and therefore cannot work as an evolutionary factor at the microevolutionary level. But if a species is so lucky that the vertical transmission of symbionts will be for it momentary benefits and therefore will be fixed by selection, this can ensure its distant descendants a triumphant success.

Subfamily termites Macrotermitae, those who have mastered effective "agriculture" - the cultivation of mushrooms - have so far seemed to be the exception to the rule. The transmission of symbionts (domesticated mushroom crops) is not vertical, but horizontal, however, trickster mushrooms are completely absent in their gardens (Aanen etal. 2009).

The symbiosis of termites with fungi arose once over 30 million years ago in equatorial Africa and turned out to be very successful. Currently, the termite mushroom subfamily includes 10 genera and about 330 species that play an important role in the cycle of substances and the functioning of tropical communities of the Old World. Unlike mushrooms grown by leaf-cutting ants, mushrooms “domesticated” by termites have already lost the ability to exist independently. They grow only in termite mounds on specially equipped beds from plant material passed through the intestines of termites.

After establishing a new colony, termites collect fungal spores in the vicinity Termitomyces and plant them in their plantations. Naturally, the initial inoculum turns out to be genetically very heterogeneous. Mushrooms form special small fruiting bodies (nodules) containing asexual spores (conidia) in the termite mound. These spores are called "asexual" because they are formed without meiosis, and their genome is identical to the genome of the parent mycelium. Conidia are used to reproduce fungi inside the termite mound. Termites feed on nodules, and the spores pass through their intestines intact and are used to seed new plantations.

Fungi also need to be taken care of in order to get into new termite mounds. Conidia usually do not spread beyond the termite mound. For this, sexual spores (basidiospores) are used. They form in fruiting bodies of a different type - large ones that grow outward through the walls of the mound. From basidiospores brought by termites to a new nest, small haploid mycelia grow. Cells of different haploid mycelia merge and turn into dikaryons - cells with two haploid nuclei. Large dikaryotic mycelia grow from them, capable of forming fruiting bodies. Nuclear fusion occurs only during the formation of basidiospores, immediately before meiosis. Conidia contain two haploid nuclei, like mycelial cells, and basidiospores contain one each.

Thus, fungi produce small fruiting bodies mainly for termites (altruism) and large ones mainly for themselves (selfishness). A trick fungus strategy could be, for example, to produce more large fruiting bodies and spend fewer resources feeding termites. But among mushrooms Termitomyces there are no cheaters, and it has not yet been known why. This mystery has only recently been solved. It turned out that only one strain of mushrooms is grown in each termite mound. At the same time, different strains are cultivated in different termite mounds. Therefore, termites prevent the appearance of deceivers in the usual way - with the help of monoculture breeding of symbionts. But how do they manage to create a monoculture from an initially heterogeneous crop? It turned out that everything is explained by the peculiarities of the relationship between fungal strains at dense sowing, combined with the fact that the reproduction of fungi inside the termite mound is completely controlled by termites. At Ter-mitomyces there is a positive correlation between the frequency of occurrence of a strain in a mixed culture and the efficiency of its asexual reproduction. In other words, genetically identical mycelia help each other—but not other mycelia—produce conidia (Aanen etal. 2009). As a result, a positive feedback occurs between the relative abundance of a strain in a mixed culture and the efficiency of its reproduction. This inevitably leads to the formation of a monoculture already after several cycles of "reseeding" carried out by termites.

Positive feedback is based on the fact that the processes of dikaryotic mycelia can fuse with each other, but only if these mycelia are genetically identical. The larger the mycelium, the more resources it can devote to the production of nodules and conidia. This contributes to the growth of yields in monoculture and the displacement of "minorities".

Apparently the wild ancestor of mushrooms Termitomyces turned out to be a good candidate for "domestication" precisely because he was inclined to form monocultures with dense sowing. The increased productivity of monocultures could become the very “momentary advantage” that allowed selection to support and develop this tendency in the early stages of the formation of symbiosis. In the long term (macroevolutionary) perspective, it proved to be decisive, because it saved the termites-mushroom growers from the threat of the emergence of trick fungi. Ultimately, this provided the symbiotic system with evolutionary success ( ibid. ).

During the transition of people from hunting and gathering to food production (the Neolithic Revolution), the problem of choosing candidates for domestication, apparently, was also extremely acute. A good symbiont is a rarity, and in many regions there were simply no suitable animal and plant species. Where there were the most of them, human civilization began to develop with the greatest speed (Diamond 1997).

The above examples suggest that if it were not for the problem of deceivers, generated by the lack of foresight in evolution and concern for the “good of the species” (rather than the gene), cooperation and altruism could become the dominant form of relationships between organisms on our planet. But evolution is blind, and therefore cooperation develops only where one or another combination of specific circumstances helps to curb deceivers or prevent their occurrence. There are not many good "engineering solutions" to deal with the problem of deceivers. Evolution repeatedly "stumbled" on each of them in its wanderings through the space of the possible.

Intergroup competition promotes intragroup cooperation

If in some animal species cooperation has already developed so much that the species has moved to a social way of life, then additional mechanisms may come into play that further strengthen intragroup cooperation. In social animals, an individual, as a rule, can only reproduce successfully by being a member of a successful group. In this case, competition usually exists not only between individuals within a group, but also between groups. What this leads to is shown by the nested tug-of-war model developed by American ethologists (Reeve and Hölldobler 2007). The aim of the study was to find an explanation for a number of quantitative patterns observed in the social structure of social insects. In the model, each individual selfishly spends a part of the "social pie" in order to increase his share of this pie. This part spent on intra-group competition is called the "selfish effort" of this individual. The share that each individual eventually gets depends on the ratio of his own egoistic efforts and the sum of the egoistic efforts of the other members of the group. Something similar is observed in social insects when they exercise "mutual supervision" - they prevent each other from laying eggs, while trying to lay their own (see above).

Relationships between groups are built on the same principles in the model. Thus, a nested, two-level "tug of war" is obtained. The more energy individuals spend on intra-group struggle, the less energy remains for inter-group “pulling” and the less the “common pie” of the group turns out.

The study of this model with the help of game theory showed that it explains well the empirically observed patterns. The model confirmed that intra-group cooperation should increase with the growth of intra-group kinship (which is fully consistent with the theory of kin selection). But the model also showed that cooperation can take place even in the absence of kinship between members of the group. This requires intense competition between groups. The main conclusion is that intergroup competition is one of the most important, and perhaps the most important factor stimulating the development of cooperation and altruism in social organisms (!) (Reeve, Hölldobler 2007).

Theoretically, this model can be applied not only to insects, but also to other social animals, and even to human society. The analogies are quite obvious. Nothing unites a team like a joint confrontation with other teams; a multitude of external enemies is a prerequisite for the sustainable existence of totalitarian empires and a reliable means of “rallying” the population into an altruistic anthill.

Genetic basis of altruism in humans

Before applying to man one or another model developed within the framework of evolutionary ethics, we must make sure that human morality is at least partly hereditary, genetic in nature, that it is subject to hereditary variability and therefore selection can act on it. On bees, bacteria and other social organisms that are not capable of cultural evolution, it is easier to study the formation of altruism, since one can immediately confidently assume that the key lies in the genes that determine behavior, and not in upbringing, culture, traditions, etc. With primates , especially with humans, it is more difficult: here, in addition to the usual biological evolution based on the selection of genes, it is also necessary to take into account social and cultural evolution based on the selection of ideas, or memes (in this case, we are talking about such memes as moral norms, rules behavior in society, etc.) (Dawkins 1976).

Recent studies have shown that the moral qualities of people are largely determined by genes, and not just upbringing. The available methods make it possible to evaluate only the tip of the iceberg - those hereditary traits for which variability has been preserved in modern people and which have not yet been fixed in our gene pool. Many of the alleles that ensured the growth of altruism in our ancestors have long been fixed, that is, they have reached one hundred percent frequency. All people have them, and therefore methods such as twin and comparative genetic analysis can no longer detect them.

It is clear that the ability for altruistic behavior is fundamentally embedded in our genes, because cooperation was necessary for our ancestors long before they mastered speech and thus created a “nutrient medium” for the spread and evolution of memes. Any healthy person with the appropriate upbringing is able to learn to behave more or less "cooperatively" and "altruistically." This means that everyone has a certain genetic basis of altruism (the corresponding genes are firmly fixed in the human population). However, until recently there were very few experimental data on the basis of which it is possible to judge in what phase the evolution of altruism is in modern humanity: has the “genetic” stage already ended, so that today only the socio-cultural aspects of this evolution are relevant, or the evolution of altruism continues at the level of genes.

In the first case, it should be expected that the hereditary variability of people in terms of traits associated with altruism is very small or completely absent, and the behavioral and moral and ethical differences between people that are so obvious to all of us are explained solely by upbringing, living conditions and various random circumstances. In the second case, we should expect that these differences are partly due to genes as well. In part, because the role of external factors in the development of the human personality is too obvious to be denied. The question is posed as follows: do individual genetic differences have any effect on the observed variability of people in the degree of cooperativity, altruism and mutual trust?

In search of an answer to this question, twin analysis is used, in particular. With the help of special tests, the degree of altruism (or, for example, such qualities as gullibility and gratitude) is determined in many pairs of identical and fraternal twins, and then the similarity of the results in different pairs is compared. If identical twins are more similar to each other in this trait than fraternal twins, this is a strong argument in favor of its genetic nature.

Such studies have shown that the tendency to act kindly, to be trusting, and to be grateful is largely genetic in nature. The differences observed in people in the degree of gullibility and gratitude are at least 10–20% genetically determined (Cesarini etal. 2008).

Specific genes are also identified that affect a person's personality, including his moral qualities (Zorina et al. 2002). In recent years, the effect of the neuropeptides oxytocin and vasopressin on the social behavior of animals and humans has been actively studied. In particular, nasal administration of oxytocin has been shown to increase trustfulness and generosity in humans (Donaldson and Young 2008). However, twin analysis shows that these character traits are partly hereditary. This suggested that certain alleles of genes associated with oxytocin and vasopressin can influence people's tendency to altruistic behavior. Recently, it was possible to find a link between some allelic variants of the oxytocin receptor gene ( OXTR) and people's tendency to display selfless altruism. The oxytocin receptor is a protein produced by some brain cells and is responsible for their susceptibility to oxytocin. Similar properties were also found in the vasopressin receptor gene ( AVPR1a). In the regulatory regions of these genes, there are so-called single nucleotide polymorphisms. These are nucleotides that can vary from person to person (most of the nucleotides in each gene are the same in all people). It turned out that some of the alleles of these genes provide less and others more propensity for altruism (Israel etal. 2009). Such facts suggest that altruism in humans, even today, can still develop under the influence of biological mechanisms, and not just socio-cultural factors.

Altruism, parochialism, and the pursuit of equality

In animals, altruism in most cases is either directed towards relatives (which is explained by the theory of kin selection), or is based on the principle “you give me - I give you”. This phenomenon is called "reciprocal or reciprocal altruism" (Trivers 1971). It occurs in animals intelligent enough to choose reliable partners, monitor their reputations, and punish deceivers, because systems based on mutual altruism are extremely vulnerable and cannot exist at all without effective means of combating deceivers.

Truly unselfish concern for non-relatives is rare in nature (Warneken and Tomasello 2006). Perhaps man is almost the only animal species in which such behavior has developed noticeably. However, people are much more willing to help “their own” than “strangers”, although the concept of “ours” for us does not always coincide with the concept of “relative”.

Recently, an interesting theory has been proposed, according to which altruism in humans developed under the influence of frequent intergroup conflicts (Choi, Bowles 2007). According to this theory, altruism among our ancestors was directed mainly towards members of "their" group. Using mathematical models, it was shown that altruism could develop only in combination with parochialism (hostility towards strangers)(!). In conditions of constant wars with neighbors, the combination of intra-group altruism with parochialism provides the greatest chances for successful reproduction of the individual. Consequently, such seemingly opposite human properties as kindness and militancy may have developed in a single complex. Neither one nor the other of these traits, taken separately, would benefit their possessors.

To test this theory, facts are needed, which can be obtained, in particular, with the help of psychological experiments. Ironically, we still know very little about how altruism and parochialism develop in the course of child development. Recently, the gap has begun to be filled thanks to special experimental studies (Fehr etal. 2008).

Among children there are about 5% of kind people, selfless altruists who always take care of others, and the proportion of such children does not change with age. There are “harm” who try to take everything from others and give nothing to anyone. Their number decreases with age. And there are “lovers of justice” who try to share everything equally, the proportion of such children is growing rapidly with age.

The results obtained are also in good agreement with the theory of the joint development of altruism and parochialism under the influence of intense intergroup competition. It is possible that the evolutionary history of these properties of the psyche in general terms is repeated in the course of the development of children. It turned out that altruism and parochialism develop in children more or less simultaneously - at the age of 5–7 years. Moreover, both properties are more pronounced in boys than in girls ( ibid. ). This is easy to explain from an evolutionary point of view. Men have always been the main participants in intergroup conflicts and wars. In the conditions of primitive life, male warriors are personally interested in ensuring that not only themselves, but also other men of the tribe are in good physical shape: there was no point in “keeping justice” at their expense. As for women, if a group was defeated in an intergroup conflict, their chances of successful reproduction were not reduced as much as for men. For women, the consequences of such a defeat could be limited to a change of sexual partner, while men could die or be left without wives. In the event of a victory, women also won clearly less than men, who could, for example, capture captives.

Of course, these properties of the child's psyche depend not only on genes, but also on upbringing, that is, they are a product of both biological and cultural evolution. But this does not make the results less interesting. After all, the laws and driving forces of biological and cultural evolution are largely similar, and the processes themselves can smoothly flow into each other (Grinin et al. 2008). For example, a new behavioral trait may first be passed down from generation to generation through learning and imitation, and then gradually become fixed in the genes. This phenomenon is known as the "Baldwin effect" and has nothing to do with the Lamarckian inheritance of acquired traits (Dennett 2003).

Intergroup wars - the cause of altruism?

The idea that the origins of human morality should be sought in the instincts that our ancestors developed in connection with the social way of life was expressed by Charles Darwin (1896); he also owns the idea of ​​the connection between the evolution of altruism and intergroup conflicts. As noted above, mathematical models show that intense intergroup competition can promote the development of intragroup altruism. To do this, several conditions must be met, of which three are the most important.

First, the reproductive success of an individual must depend on the prosperity of the group (moreover, the concept of "reproductive success" includes the transfer of one's genes to offspring through relatives whom the individual helped to survive and who have many genes in common with him). There is no doubt that this condition was fulfilled in the collectives of our ancestors. If a group loses an intergroup conflict, some of its members die, and the chances of surviving to raise healthy and numerous offspring are reduced. For example, in the course of intergroup conflicts among chimpanzees, groups that lose in the fight against their neighbors gradually lose both their members and territory, i.e., access to food resources.

Secondly, the intergroup enmity among our ancestors should have been quite sharp and bloody. Proving this is much more difficult.

Thirdly, the average degree of genetic relationship between tribesmen should be significantly higher than between groups. Otherwise, natural selection will not be able to support sacrificial behavior (assuming that altruism does not give the individual any indirect benefits - neither through increased reputation, nor through the gratitude of fellow tribesmen).

S. Bowles, one of the authors of the theory of the coupled evolution of altruism and hostility to strangers, tried to assess whether the tribes of our ancestors were strong enough at enmity with each other and whether the degree of kinship within the group was high enough so that natural selection could ensure the development of intragroup altruism (Bowles 2009) . Bowles showed that the level of development of altruism depends on four parameters: 1) on the intensity of intergroup conflicts, which can be estimated from the death rate in wars; 2) on the extent to which an increase in the proportion of altruists (for example, brave warriors who are ready to die for their tribe) increases the likelihood of victory in an intergroup conflict; 3) on how much the kinship within the group exceeds the kinship between the warring groups; 4) on the size of the group.

In order to understand the range of these four parameters in the groups of primitive people, Bowles drew on extensive archaeological data. He concluded that the conflicts in the Paleolithic were very bloody: from 5 to 30% of all deaths, apparently, were due to intergroup conflicts. In the book by A.P. Nazaretyan “The Anthropology of Violence and the Culture of Self-Organization. Essays on evolutionary-historical psychology” (2008) collected anthropological data indicating a very high level of violent mortality in archaic societies. The size of human groups in the Paleolithic and the degree of kinship in them can also be estimated from the data of archeology, genetics and ethnography. As a result, only one value remains, which is almost impossible to assess directly - the degree of dependence of the military successes of the group on the presence of altruists (heroes, brave men) in it. Calculations have shown that even at the lowest values ​​of this quantity, natural selection in hunter-gatherer populations should help maintain a very high level of intra-group altruism. The “very high” level in this case corresponds to values ​​of the order of 0.02–0.03. In other words, altruism gene» will spreadinpopulations, if the chances of survivalandleave offspringatcarrier of such a gene 2–3 % below, howatselfish fellow tribesman. It could seem, what 2–3 % – not a very high level of self-sacrifice. However, this is actually a significant amount.. Bowles gives two illustrative calculations.

Let the initial frequency of occurrence of this allele in the population be 90%. If the reproductive success of carriers of this allele is 3% lower than that of carriers of other alleles, then after 150 generations the frequency of occurrence of the "harmful" allele will decrease from 90 to 10%. Thus, from the point of view of natural selection, a three percent reduction in fitness is a very expensive price. Now let's try to look at the same value (3%) from a "military" point of view. Altruism in war is manifested in the fact that warriors attack enemies without sparing their lives, while egoists hide behind their backs. Calculations showed that in order for the degree of altruism to be equal to 0.03, military mortality among altruists should be over 20% (taking into account the real frequency and bloodshed of Paleolithic wars), i.e., whenever a tribe collides with neighbors for life , and to death, every fifth altruist must sacrifice his life for the sake of a common victory. Admittedly, this is not such a low level of heroism (Bowles 2009). This model is applicable to aspects and cultural factors of altruism transmitted through training and education.

Thus, the level of inter-group aggression among primitive hunter-gatherers was quite sufficient for the “genes of altruism” to spread among people. This mechanism would work even if within each group the selection favored exclusively egoists. But this condition, most likely, was not always observed. Selflessness and military exploits could increase the reputation, popularity and, consequently, the reproductive success of people in primitive collectives.

The mentioned mechanism of maintaining altruism by improving the reputation of the one who performs the altruistic act is called "indirect reciprocity" (Alexander 1987). It works not only in humans, but also in some animals. For example, in Arabian gray thrushes Turdoides squamiceps only high-ranking males have the right to feed their relatives. These social birds compete for the right to do a "good deed" (to sit over the nests as a "sentinel", to help care for the chicks, to feed a comrade). Altruistic acts have acquired a partly symbolic meaning for them and serve to demonstrate and maintain their own status (Zahavi 1990). Reputation issues are extremely important in any human team. According to one authoritative hypothesis, an important stimulus for the development of speech in our ancestors was the need to gossip. Gossip, within the framework of this hypothesis, is considered as the oldest means of disseminating compromising information about "unreliable" members of society, which contributes to team building and punishment of deceivers (Dunbar 1998).

It is impossible to cover all areas of research related to the evolution of altruism in one review. In particular, the following remained outside the scope of this article: 1) works devoted to the study of innate psychological predispositions found in humans to effectively identify deceivers; 2) the phenomenon of "expensive punishment" ( costly punishment), which manifests itself in the fact that people are ready to make sacrifices for the effective punishment of deceivers (this can also be considered a form of altruism, because a person sacrifices his interests for what he considers to be a public good or justice); 3) study of the system of emotional regulation of the formation of moral judgments (according to the results of the latest neurobiological studies, it is the brain regions associated with emotions that play a key role in solving moral dilemmas; the emotion of disgust was probably “recruited” in the course of evolution to form a hostile attitude towards strangers) ; 4) the study of the role of religion, "expensive" rituals and religious rites as a means of strengthening parochial altruism (see: Markov 2009), etc.

In conclusion, it is necessary to consider briefly the question of what ethical conclusions can be drawn from the data of evolutionary ethics, and which should never be drawn. If one or another aspect of our behavior, emotions and morality follows from evolutionary patterns (has an evolutionary explanation), this does not mean at all that this behavior has thus received an evolutionary “justification”, that it is good and correct. For example, hostility to strangers and wars with foreigners have been an integral part of our evolutionary history and even, perhaps, a necessary condition for the development of the foundations of our morality, propensity for cooperation and altruism. But the fact that historically our altruism was directed only at “our own”, and our ancestors felt disgust and enmity towards strangers, does not mean that this is the model of morality that we should imitate today. Evolutionary ethics explain, but do not justify, our innate tendencies. At present, the development of moral and ethical standards is determined by cultural and social evolution to an immeasurably greater extent than by biological evolution, which is much slower, and therefore its influence on changes in moral Zeitgeist(“spirit of the times”) on short time intervals (on the scale of decades and centuries) is negligible. Fortunately, in addition to archaic instincts and emotions, evolution also gave man reason, and therefore we can and must rise above our biological roots, timely revising the outdated ethical framework that evolution imposed on our ancestors. Far from all the emotional and behavioral stereotypes that contributed to the spread of Stone Age hunter genes are optimal for a modern civilized person. In particular, evolutionary ethics warns us that we have an innate tendency to divide people into friends and foes, and to feel disgust and hostility towards strangers. We, as rational beings, must understand and overcome this.

Literature

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Darwin, Ch. 1896. The origin of man and sexual selection/ per. I. Sechenov. SPb.: Ed. O. N. Popova.

Zorina, Z. A., Poletaeva, I. I., Reznikova, Zh. I. 2002. Fundamentals of ethology and genetics of behavior. M.: Higher school.

Markov, A. V. 2009. Religion: a useful adaptation, a by-product of evolution, or a "brain virus"? Historical psychology and sociology of history 2(1): 45–56.

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“... we are facing two main questions. On the one hand, it is clear that many life tasks are easier to solve by joint efforts than alone.

Why, then, did the biosphere never turn into a realm of universal friendship and mutual assistance? This is the first question.

The second question is the opposite of the first. How can altruism arise in the course of evolution at all, if the driving force of evolution is natural selection - a process that, at first glance, seems to be absolutely selfish?

The whole point is that this “first look” is wrong.

The mistake here is to confuse the levels at which we consider evolution.

Evolution can be considered at different levels: genes, individuals, groups, populations, ecosystems, the entire biosphere. Each level has its own patterns and rules.

At the level of genes, evolution is based on the competition of different variants (alleles) of the same gene for dominance in the gene pool of a population. At the gene level, there is no altruism and cannot be. Gene is always selfish. If a “good” allele appears, which, to its detriment, allows another allele to multiply, then this altruistic allele will be forced out of the gene pool and simply disappear.

But if we shift our view from the level of genes to the level of organisms, the picture will be different. Because the interests of the gene do not always coincide with the interests of the organism. A gene, or, more precisely, an allele, is not a single object; it is present in the gene pool in the form of many identical copies. The "interest" of all these copies is the same. After all, they are just molecules, and they are absolutely identical. And they, and us, and natural selection do not care at all which of the identical molecules will multiply and which will not. Only the total is important: how many copies of the allele were and how many became.

An organism, on the other hand, is a single entity, and, to put it simply, only one or two copies of the allele of interest to us can be present in its genome.

Sometimes it is beneficial for a selfish gene to sacrifice one or two copies of itself in order to provide an advantage to the rest of its copies, which are contained in other organisms. Biologists began to approach this idea already in the 30s of the last century. An important contribution to understanding the evolution of altruism was made by Ronald Fisher, John Haldane and William Hamilton.

The theory they built is called kin selection theory. Its essence is expressed figuratively Haldane who once said, "I would give my life for two brothers or eight cousins." What he meant by this can be understood from the formula that entered science under the name "Hamilton's rule."

Here is the formula. An “altruistic gene” (more precisely, an allele that promotes altruistic behavior) will be supported by selection and will spread in the population if

RB > C

where R is the degree of genetic relationship between the donor and the “receiver” (in fact, kinship is not important in itself, but only as a factor that determines the likelihood that the “receiver” has the same altruism allele as the donor); B - reproductive advantage received by the addressee of the altruistic act; C - reproductive damage caused by the "donor" to himself. Reproductive gain or loss can be measured, for example, by the number of offspring left or not left.

Taking into account the fact that not one, but many individuals can benefit from an act of altruism, the formula can be modified as follows:

NRB > C,

where N is the number of those accepting the sacrifice.

Note that Hamilton's rule not introduces no additional entities, requires no special assumptions, and does not even need experimental verification. It is purely logically derived from the definitions of R, B, C, and N, just as geometrical theorems are derived from axioms. If NRB > C, the "altruism allele" will quite automatically increase its frequency in the population's gene pool."

Markov A.V. , human evolution. Monkeys, neurons and the soul. In 2 books. Book two, M., "Ast"; Corpus, 2013, p. 298-300.

Which, under certain conditions, reduce the chances of individuals to reproduce, can spread in a population when the value of the contribution to reproduction of other individuals is greater than the price of help. In this case, this individual thus produces more copies of its genes than by spending all the resources on its own reproduction.

The rule was formulated by the British biologist W. Hamilton in

see also

Sources

• Hamilton W. D. (1963) The evolution of altruistic behavior. American Naturalist 97:354-356

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An excerpt characterizing Hamilton's Rule

The postilion moved off, and the carriage rattled its wheels. Prince Hippolyte laughed abruptly, standing on the porch and waiting for the viscount, whom he promised to take home.

“Eh bien, mon cher, votre petite princesse est tres bien, tres bien,” said the viscount, getting into the carriage with Hippolyte. - Mais tres bien. He kissed the tips of his fingers. – Et tout a fait francaise. [Well, my dear, your little princess is very cute! Very nice and perfect French.]
Hippolyte laughed with a snort.
“Et savez vous que vous etes terrible avec votre petit air innocent,” continued the viscount. - Je plains le pauvre Mariei, ce petit officier, qui se donne des airs de prince regnant.. [Do you know, you are a terrible person, despite your innocent appearance. I feel sorry for the poor husband, this officer who poses as a possessive person.]