In this scene, as in many other situations in nature, we see a bizarre combination of altruistic and selfish behavior. The calling food cry of a seagull is a typical example of altruism. The seagull does not gain any benefit from this cry. The winnings go to the other seagulls: they get a chance to dine. The second part of the scene is a fight. Here, of course, we see only pure selfishness on the part of all participants.
The answer is in Hamilton's rule. Seagulls on the White Sea feed mainly on schooling fish, such as herring. If a seagull notices 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. Magnitude IN- the winnings of those who arrive at the cry will be quite large: they will have lunch. Since the fish are schooling, you may have to wait a long time for the next school. Magnitude R(relatedness) is also likely to be high, because gulls nest in colonies, often returning to the same place after wintering, and therefore, most likely, its relatives - parents, children, brothers and nephews - nest next to this gull.
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 advantageous to shout, and in the second, to remain silent. But such calculations require brains. And the brain, as we know, is an expensive organ. Selection, as a rule, tries to save on brains. Besides, brains are heavy. Seagulls need to fly, not solve algebraic problems. Therefore, the bird cannot figure out when it is beneficial for it to call its companions and when it is not, and its behavior turns out to be illogical. Not always, but only when there is a lack of fish.
The evolution of altruism has gone especially far among hymenopteran insects: ants, bees, wasps, bumblebees. In social Hymenoptera, most females forego 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 had to do with the peculiarities of gender inheritance. In Hymenoptera, females have a double set of chromosomes, and 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 share 50% of their genes (identical by origin). Magnitude R in Hamilton's formula is equal to 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, like other animals, have only 50% of their genes in common. So it turns out that, all other things being equal, it is more profitable for female Hymenoptera to raise sisters than daughters.
The mechanism of sex inheritance in Hymenoptera. The female is diploid, that is, she has a double set of chromosomes (2n). She can lay an unfertilized egg with a single set of chromosomes (p), 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 her chromosomes from her mother and half from her father. The male receives half of her chromosomes from his mother, but 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 (from the sister’s point of view) or 50% from the brother’s point of view. However, working females raise brothers too (although they do not like them). We will not go into this rather complex theoretical area, especially since the primates that interest us are not haplodiploid. But social Hymenoptera have (or had in the evolutionary past) another important property that sharply increases the likelihood of the development of altruism under the influence of kin selection. This property is monogamy.
The offspring of monogamous diploid parents share on average 50% of their genes ( R= 0.5). The offspring of a female mating with many males have an average R tends to 0.25 (if there are a lot of males). For kin selection, this is a very serious difference. At R= 0.5, any trifle is enough to tip the scales in favor of sisters and brothers. At R= 0.25 your children are definitely more expensive. It is very important that monogamy is characteristic of termites - the second order of insects in which eusociality is widespread, without any haplodiploidy. Termites work not only females, but also males (they are diploid, like their sisters).
As we remember, monogamy was probably characteristic of ancient hominids. This could become a powerful incentive for the development, under the influence of kin selection, of brotherly (and sisterly) mutual assistance, intrafamily cooperation and altruism. And also, of course, fatherly love, and at the same time the devotion of children to both parents, and not just the mother. Perhaps kin selection was able to support this entire range of altruistic feelings in our ancestors precisely because they were - at least partly - monogamous.
This slide shows definitions, I won’t dwell on them, I think everyone more or less understands 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 life problems are much easier to solve through joint efforts than alone. Why then has the biosphere never turned into a kingdom 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 the levels at which we consider evolution. At the gene level, evolution is based on competition between 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, there cannot be. Gene is always selfish. Now, if such a “good” allele suddenly appears, which, to its detriment, allows another competing allele to reproduce, then this “good” allele will be automatically forced out of the gene pool and will simply disappear. Therefore, there is no altruism at the gene level. But if we shift our gaze from the level of genes to the level of organisms, then 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 resides. Why? Because a gene, or more precisely an allele, a variant of a gene, is not a single object. It is present in the gene pool in the form of many identical copies. But an organism is a single object, and it carries within itself 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 its other copies, which are contained in other organisms. But here I must make a reservation; biologists are sometimes reproached for using metaphors such 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 understands nothing and strives for nothing. When biologists say “it is beneficial for the gene”, “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 the gene had brains and desires, it would change in the same way as it changes automatically under the influence of selection. I hope this is clear to everyone. It may be beneficial for a gene to sacrifice several copies of itself in order to give an 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. Important contributions to this matter were made by Ronald Fisher, John Haldane, and William Hamilton.
Creators of the theory of kin selection
And the theory they built is called the Kin Selection Theory. Its essence was expressed figuratively 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 alarmed, 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 promotes the altruistic behavior of the organism, will be supported by selection, that is, it will spread in the gene pool of the population if this inequality is satisfied:
gV > C
Where r- the degree of genetic relationship between the one who makes the sacrifice and the one who accepts the sacrifice. This degree of genetic relatedness is the likelihood that the person you are sacrificing yourself for has the same allele of the same gene as you. For example, this altruism gene. Let's say, if some allele is in me and I have a sibling, then, roughly speaking, there is a ½ probability that he has the same allele. If, say, a cousin, then it will be 1/8. IN(Benefit) is the reproductive advantage received by the recipient of the altruistic act, that is, the one for whom you sacrifice yourself. A WITH(Cost) is the “price” of an altruistic act, that is, the reproductive damage caused by the donor to himself. This can be measured in the number of, say, descendants born or unborn by you.
Haldane said “I would give my life for two brothers”, here we must modify a little more, if we sacrifice ourselves not for the sake of one individual, but for the sake of several, then we can also add n at the beginning:
nrB > C
n- this is the number of those accepting the sacrifice. Here two brothers are shown, n = 2, r=0.5, IN- 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 IN 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 for bash”, “a big deal”. For three brothers it will be profitable. Gene, not you. Now we can understand the behavior of those same seagulls. This food cry is a calling, why have seagulls developed such an instinct - to scream and call 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 doesn’t have anything will lose. Magnitude WITH– the price of an altruistic act is likely to be low. IN– the winnings of those who fly to the cry will be quite large, they will have lunch. Since, again, the fish are schooling, you may have to wait a long time for the next school. That is, the gain is quite tangible. r- kinship. The relatedness is also most likely quite high, because they nest in colonies, often returning to the same place after wintering, and therefore, most likely, various of its relatives nest next to this gull: parents, children, brothers, nephews, etc. .d. AND n- the number of seagulls that will hear, fly in and have lunch is also quite high. So she screams. And why doesn’t she share her prey? She doesn’t give back what she’s already grabbed - because here WITH She’s already doing much more, she’s actually left without lunch. AND n less. By giving her prey to another gull, she will feed one, and not the whole flock. So the inequality is not fulfilled, so such an instinct has not been developed. Of course, the most beneficial thing for a seagull would be to learn to distinguish a situation when there is a lot of food and enough for everyone, and then call. And when there is little food, eat in silence. 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 or unprofitable, and this illogical behavior results.
Hymenoptera - a group in which the evolution of altruism has gone particularly far
In general, Hamilton's rule has remarkable predictive and explanatory power. For example, in which group of animals the evolution of altruism 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 generally refuse to reproduce and raise their sisters. Working females do not reproduce, but help their mother raise her sisters. Why specifically Hymenoptera, why is this so common in this order of insects? Hamilton suggested that the whole point here was the peculiarities of gender inheritance. 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. Magnitude r in Hamilton's formula is equal to ½, and in Hymenoptera sisters have 75% of common genes. Because each sister receives from her father not half of his chromosomes, as is usual in other animals, but she receives the entire paternal genome. And all sisters receive this complete paternal genome, the same one. Because of this, they share 75% of their genes. It turns out that for female Hymenoptera, a sister is a closer relative than a natural daughter. And therefore, other things being equal, it is more profitable for them to force their mother to give birth to more and more new sisters and raise them, than to give birth to their own daughters. But in reality, everything is somewhat more complicated here, because there are also brothers who, on the contrary, turn out (brother and sister) to be more distantly related than those of ordinary animals. I will not go into these subtleties, but in this situation, where sisters are closer to each other than mother and daughter, there is apparently enough of it in the order 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, it is very widespread. Now one of the interesting areas in microbiology is the experimental study of the evolution of bacteria, “evolution in vitro.”
Evolution of “altruists” and “deceivers” in vitro: experiments with the bacterium Pseudomonas fluorescens
Altruists and deceivers in Myxococcus xanthus bacteria
Honest yeast and cheating 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 labors. Theoretically, this should lead to the complete displacement of altruists by egoists. But in reality, the number of altruists does not fall below a certain level. We began to investigate why. It turned out that the fact is that the altruism of yeast, upon closer examination, is not entirely selfless. They really help everyone around them, they 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 thanks to this little trick, given the 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 it is hardly possible to build a serious complex cooperative system on such small tricks. Another great trick of this kind is called the Simpson Paradox. The essence of this paradox is that, subject to 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's Paradox in action. There was a population where there were half altruists and egoists. 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 daughter 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 there were initially more altruists, in general, grow faster. Altruists still help others. As a result, in total, the percentage of altruists increases, despite the fact that in each individual population it decreased. Recently it was possible to experimentally show that this is not just a theory, but that such a mechanism can actually work in microbes. True, apparently, for this to happen, rather rare conditions must be met, but this is not yet entirely clear. But there is also a trick for maintaining the level of goodness in the world. It's time to move from microbes to multicellular ones. The appearance of multicellular organisms in general and animals in particular was the greatest triumph of the evolution of altruism. In a multicellular organism, most cells are altruists who have given up their own reproduction for the sake of the common good. Animals, compared to microbes, have new opportunities for the development of cooperation based on complex behavior and learning. But, unfortunately, the same opportunities appeared for the deceivers, and the evolutionary arms race continued at a new level. And again, neither the altruists nor the deceivers received a decisive advantage.
Altruism in social insects is far from selfless
In many species of Hymenoptera, workers sometimes show selfishness by laying their own eggs. In Hymenoptera, as we said, males are born by virgin birth, parthenogenetically, from haploid unfertilized eggs. The workers of some wasps try to lay such unfertilized eggs and hatch their own sons. This is the most profitable strategy, as I mentioned, the most profitable thing for a female Hymenoptera is to raise sisters and native sons. This is what they are trying to do. But this is not liked by other workers, for whom it is beneficial to lay their eggs, but it is not beneficial for their sisters to do this, so they destroy the eggs laid by their sisters. It turns out to be a kind of morality police. And special studies have shown that the degree of altruism in colonies of such wasps seems to depend not so much on the degree of relationship 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 a family, of which only one, the oldest, lays eggs. The rest take care of the larvae. When the queen dies, the next oldest wasp takes her place. That is, they strictly adhere to the principle of seniority. At the same time, helper wasps, which do not yet reproduce on their own, vary greatly in the degree of their work enthusiasm. Some work hard without sparing themselves, while others sit back in the nest and relax. And so, as it turned out, their work enthusiasm depends on how great the chances of a given wasp for the royal throne are. How great are her chances of leaving her own offspring, starting her own family. If these chances are small, as with low-ranking wasps, 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 tendency to altruism is stronger among those who have nothing to lose. Is it possible to create a society in which altruism will be supported without violence, and where there will be no deceivers? Neither wasps nor humans have yet succeeded in this, 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 prevent the emergence of cheaters is to reduce the genetic diversity of individuals in the system to zero, so that everyone is genetically identical. Then the symbionts simply will not be able to compete with each other over which of them will grab a larger piece of the common pie. That is, the symbionts will be able to, but the genes that reside 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: heredity, variability and selection, one of the components is excluded, namely variability. That is why evolution never managed to create a normal multicellular organism from genetically diverse cells, but managed to create it from clones, descendants of a single cell. There is such an interesting phenomenon as insect agriculture.
Some ants and some termites grow mushrooms, “domesticated” mushrooms, in special gardens in their nests. In such a situation, it is 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, the easiest way for the host to ensure the genetic identity of its symbionts is to pass them on by inheritance. Moreover, only one of the sexes should do this: either males or females. This is exactly how leaf-cutter ants pass on their mushroom crops from generation to generation. When symbionts are transmitted vertically, they take with them a small amount of seed material, this mushroom material, before establishing a new anthill. And this leads to the fact that genetic diversity, due to constant bottlenecks in fungal numbers, is constantly maintained at a very low level. But, however, there are also symbiotic systems with horizontal transmission of symbionts, that is, for example, each host collects symbionts for itself in the external environment. In such systems, the symbionts of each host will be genetically heterogeneous, they retain the ability for selfish evolution, and therefore deceivers appear among them every now and then. And here nothing can be done. Deceivers appear, for example, many deceiver strains are known among symbiotic luminous bacteria, which are symbionts of fish and squid. They work as lanterns 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 the symbionts, and therefore the hosts have to fight the deceivers with 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 only notices immediate 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 might be like an earthly paradise. But evolution is blind, and therefore cooperation develops only where one or another set of special circumstances helps to curb deceivers or prevent their emergence. If in a certain species of animals 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 take away resources from their group mates. This part of the resources spent on intra-group squabbles is called the “selfish effort” of a given 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 a group, that is, nested two-level competition is obtained. And the more energy individuals spend on intragroup struggle, the less it remains for intergroup competition and the smaller the “common pie” of the group turns out - the total amount of resources obtained by the group. Research into this model has shown that competition between groups should be the strongest incentive for the development of intragroup cooperation. This model seems to apply to human society as well. Nothing unites a team more than joint opposition to other teams, many external enemies; Clearly, this is a prerequisite for the existence of totalitarian empires and the most reliable means of uniting the population into an altruistic anthill. But before we apply 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 in bees and bacteria, because one can immediately confidently assume that the answer lies in genes, and not in upbringing or cultural traditions. And research in recent years has 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) in 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 recorded in our gene pool. It is clear that all people have a certain genetic basis for altruism. The question is 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 do 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 influence a person’s moral qualities have also been identified. I don’t have time to talk in detail about these genes, but the general conclusion is clear: altruism in people, even today, can still develop under the influence of biological mechanisms. And therefore evolutionary ethics is quite applicable to us.
Reciprocal (mutual) altruism
In animals, altruism is usually directed either towards relatives, or, another option, it can be based on the principle: you - for me, I - for you. This phenomenon is called reciprocal or reciprocal altruism. It is found in animals intelligent enough to choose reliable partners and punish deceivers, because systems based on mutual altruism are extremely vulnerable and generally cannot exist without effective means of combating deceivers. The ideal of reciprocal altruism is the so-called “Golden Rule of Ethics”: do unto others as you would have them do unto you. But truly unselfish care for non-relatives is rare in nature; perhaps humans are almost the only species in which such behavior has received some development. But recently an interesting theory was proposed, according to which altruism in people developed under the influence of frequent intergroup conflicts. I have already said that models show that intergroup hostility promotes the development of ingroup altruism. According to this theory, altruism among our ancestors was initially aimed only at members of their own group. Naturally, in such a situation, researchers, even using mathematical models, showed that it seemed that altruism could develop immediately only in combination with parochialism. Parochialism refers to devotion to one's own and hostility to strangers. And it turns out that our opposite qualities, such as, on the one hand: kindness, altruism, on the other hand: belligerence, hatred of strangers, of 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 brought any benefit to their owners. But to test this theory, facts are needed, which they are now trying to obtain - in particular, with the help of various psychological experiments. For example, it turned out that most three- or four-year-old children usually behave like selfish people, but by the age of 7-8 they already have a clearly expressed willingness 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 dividing candy, both in their own and in others' favor. That is, it no longer looks like selfless altruism, but like a desire for equality, egalitarianism, this is some form of struggle against deceivers, in fact, perhaps. And the proportion of such lovers of justice among children grows very quickly 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 towards strangers.
Altruism among “insiders” and hostility towards outsiders: 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 if they lost in an intergroup conflict and gained much more if they won. For example, in case of victory, they could take captives; in case of defeat, they most likely lost their lives. And in many cases women only faced the danger of changing their husbands. And therefore, it is not surprising that in men both intra-group cooperation and hostility towards outsiders are more pronounced. 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, our ancestors could have formed the foundations of morality. Such reasoning cannot do without intergroup wars. Accordingly, we know that intergroup competition can promote intragroup altruism, but several conditions must be met for this to happen. In particular, intergroup enmity among our ancestors must have been quite acute and bloody. Was this really so? Recently, archaeologist Samuel Bowles, one of the authors of this theory of the coupled evolution of altruism and hostility towards strangers, tried to assess whether the tribes of our ancestors were at odds with each other enough for natural selection to ensure the development of intragroup altruism.
Are intergroup wars the cause of altruism?
Extensive archaeological data from the Old Stone Age and the Paleolithic were analyzed, and the conclusion was that conflicts in the Paleolithic were generally very bloody. Between 5 and 30% of all deaths were violent, apparently usually occurring in intergroup conflicts. This is, in fact, a colossal number. Up to 30% of violent deaths. This seems completely counterintuitive and hard to believe, but it is 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 20th century, taking into account 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 20th century. And calculations show that this level of bloodshed is more than enough for natural selection to contribute to maintaining high levels of intragroup altruism in hunter-gatherer populations. Moreover, this should happen even in cases where within each group selection favors exclusively egoists. But this condition most likely was not met, because selflessness and military exploits most likely increased the reputation and, consequently, the reproductive success of people in primitive groups.
Indirect reciprocity
This mechanism for maintaining altruism through reputation enhancement 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 doesn't only work in humans; Surprisingly, it is also found in animals, and a wonderful example is such social, social birds, Arabian gray blackbirds. They live in colonies and raise their chicks together. They have sentries that sit in the treetops and watch for predators. It is customary for them to feed each other and help each other in this way. Males help females take care of the chicks, in general, this is a social way of life. And it turned out that among these blackbirds, 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 perform a good deed. And only a high-ranking male can serve as a sentry. That is, altruistic acts acquire symbolic meaning. They begin to serve as status signs, to demonstrate and maintain their own status. Reputation has been 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 incriminating information about unreliable members of society, which helps to unite the team and punish deceivers. With this I am already approaching 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 talk about all the interesting research in this area.
Some ideas not included in the report
This slide lists in abstract form some points that were not included in the lecture. For example, it has been 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, problems that are difficult to solve or guess. But problems can be presented in different contexts. Maybe about Masha and Petya and how many apples they have. Or you can find a different setting for this problem. And it turned out that if the surroundings are connected with exposing a deceiver, with exposing a violator of some kind of social order, then people are reliably more successful in solving such problems. That is, if it’s not about Masha, Petya and apples, but about the fact that someone deceived someone, stole someone, some kind of deception - the problem 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 in order to properly punish that scoundrel. This is also a manifestation of altruism. A person sacrifices himself for the sake of the public good, so to speak. Or at least what he considers to be a public good. Then there are more interesting arguments, work on the emotional regulation of the processes of forming 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, first of all, the departments associated with emotions are activated in our brain. And there are also very interesting results obtained on people who have certain parts of the brain 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 experience feelings of guilt, sympathy, empathy - while all other functions of the intellect are fully preserved. There are various other neurobiologically interesting things. There is also a whole such branch - evolutionary religious studies, where the evolutionary roots of religions and the possible role of religious beliefs in strengthening and strengthening this parochial altruism are examined. In particular, the function of rituals, joint religious rites, as some special studies show, may be to prevent the emergence of deceivers and 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: 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 thereby justified, that it is good and correct.
Conclusion
When we do evolutionary ethics, we are talking about the morality that emerged from biological evolution during the hunter-gatherer stage. It is clear that with the development of civilization the situation changes - what was good and highly moral for a hunter-gatherer is not necessarily good and highly moral for a modern city dweller. Fortunately, evolution also gave man reason, and, for example, evolutionary ethics warns us that we actually have an innate tendency to divide people into “strangers” and “us.” And to feel disgust, hostility, and enmity towards “strangers”. 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.
Discussion of the lecture
Boris Dolgin: Thanks a lot. It seems that this topic would be good for some kind of 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, it seems to me, much tougher in distinguishing where there is an evidential judgment, and where there are interpretations and constructions on top of these judgments, which is what the presented part of the supposedly natural science sins with. It seems that there are some strange sags 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 people, but for social animals. And somewhere in the argument, the line was not very clearly drawn between what could be considered directly proven, what kind of experiment was carried out, what it 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 mostly told the conclusions of some article in thesis form, just in one phrase. Conclusion after conclusion. Naturally, I simply did not have time to discuss the degree of reliability of certain conclusions. There is a separate discussion for each phrase about how reliable it is.
Novel: The question is next. You connected a huge percentage of deaths in the Paleolithic and, as a consequence, the development of altruism. Can we conclude that in the twentieth 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 an advantage was given to those individuals who knew how to cooperate with their own, with members of their tribe, and were even ready to sacrifice themselves for the sake of their own. For the sake of my little tribe. And how to tie this to modern wars, to modern society is a rather difficult task, and there is no serious data here, there is no direct connection, because now social and cultural evolution plays a much larger 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 happening, 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. These are, as it were, slightly different things from different areas.
Boris Dolgin: The question contained a very important, albeit slightly strangely expressed idea: would you like to try measuring altruism? That is, introduce some kind of unit, somehow try to isolate it from behavior? If you use this category all the time, you might want to instrumentalize it. The question, as I understand it, was how do you measure altruism over a period? Your answer: other factors are playing a big role now. And here, I hope, most of us will completely agree with you. But then what to do with “altruism”? Why do you even need this category? What are you doing with her?
Alexander Markov: In biology, altruism is always nothing more than a kind of metaphor, an image. And some researchers don’t like to use this word at all; they replace it with all sorts of euphemisms. For example, in my opinion, those authors who worked on yeast, one yeast secretes an enzyme, helps others, another yeast does not secrete an enzyme. To call this one yeast an altruist and the other an egoist - perhaps some authors believe that it is not necessary. Call it something else. Each specific situation means something different. The person has some special psychological tests. It's a multifaceted thing. And in the case of yeast, they simply measure: 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 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 each specific case has its own idea. In general, this is a common concept - sacrificing one’s own reproductive interests in order to increase the reproductive success of another. Although, of course, I understand that this is all quite vague, but people are interested in knowing where their moral instinct came from. And that's why I think it's useful to talk about these things.
Dmitry Gutov: The train of thought is interesting, maybe this is not your specialization, but if we were to radically summarize, then it would be necessary to extend this concept to the inorganic world, that is, maybe physicists are doing 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 are striving for something. Specifically, they strive to increase the efficiency of their reproduction. As if. Therefore, you can use such 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 purpose do, say, inorganic objects have if we begin to apply the concepts of altruism and egoism to them? For living beings, altruism is sacrificing one's own goal to help another achieve that goal. This is understandable.
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 expansion more deeply.
Boris Dolgin: Let's say, for crystals?
Alexander Markov: First of all, I never thought about this topic. Secondly, at first glance I don’t 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 altruism genes. How can the fact that these genes for modified vasopressin and oxytocin receptors be associated with the functions of these hormones be related?
Alexander Markov: So you mean the genes that a person has?
Olga: Yes.
Alexander Markov: As is your custom, I can talk about them for an hour?
Boris Dolgin: Approach wisely. There are still people who clearly want to ask questions, but at the same time try to answer somehow.
Alexander Markov: This is a very interesting topic. Simply a wonderful topic.
Boris Dolgin: You can send to work.
Alexander Markov: Oxytocin and vasopressin are neuropeptides, small protein molecules that are secreted by certain neurons of the brain, hypothalamus, and they serve as signaling substances. In general, there are a lot of signaling 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'm going over in my head what to choose for the story now. Let's say there is such a wonderful object - American voles, in which in one genus there are species that are monogamous, that is, they form strong mating pairs, the male is actively involved in caring for the offspring, and there is a lifelong attachment between the male and female. There are polygamous species, where there are no such stable relationships between males and females, and males do not participate in caring for the 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 the receptor is a protein that responds to this vasopressin - and, accordingly, the neuron is excited. And it turned out that by changing the operation of the gene for this very vasopressin receptor, even a male from a polygamous species can be forced to become a faithful husband, that is, so that he can form a strong attachment, a lifelong love for one female. Until recently, they did not know whether humans have the same problem. It turned out that there is one after all. Of course, we have the same vasopressin receptor gene, we began to look at variability, polymorphism in this gene, and whether polymorphism in this gene correlates with any aspects of personality. And it turned out that yes, it correlates. For men who have one of the variants of this vasopressin receptor gene, firstly, the emergence of a romantic relationship with a girl is half as likely to lead to marriage as for all other men. And if they do get married, they are more likely to be unhappy in their family life. And the wives of such men are almost always dissatisfied with family relationships. And the same gene as in voles affects marital fidelity and marital affection. Here it is difficult to doubt that a person has a genetic basis for such things, say, as love between spouses. In addition, the variability of oxytocin and vasopressin receptor genes turned out to correlate with such qualities as kindness and generosity. This is tested, for example, in various economic games. And a lot of different experiments are being carried out. They put oxytocin in people's noses and see how their behavior changes. This has a great effect on men. So that, for example, they begin to understand the interlocutor’s facial expression better, look into the eyes more often, and so on. That is, it is clearly absolutely clear that kindness, sensitivity - all this very much depends on the oxytocin-vasopressin system.
Olga: Variations in receptors that lead to, say, changes in the population of voles, do they bind to the hormone better or worse?
Alexander Markov: There is a level of expression, I’ll try to remember now. In one case there are simply more of these receptors, the gene expression is higher, and in the other there is less. But, to be honest, I don’t remember which one, I’ll have to look.
Alexander: Please tell me, 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, some bacteria “go astray”? And what are the criteria for moving from one to the other? Or did they remain as they were born?
Alexander Markov: It is very difficult to register such cases in which a bacterium “re-educates” during its lifetime and changes its behavior; even if they exist, it is not clear how.
Alexander: What if we take it higher?
Alexander Markov: That is, a mutation occurs - and then the behavior changes. But this will already happen in the next generation.
Alexander: What if we take other organisms rather than bacteria?
Boris Dolgin: That is, at what level, as I understand the question, does variability in behavior arise - within the life of the same organism? Did I understand the question correctly?
Alexander: In particular, yes, if it is not clear how to record this process in bacteria, then what about others?
Alexander Markov: And animals can, of course, modify their behavior depending on circumstances. But again, always following Hamilton's formula. I spoke about wasps: as the wasp's chances for the royal throne grow, it works less and less and shifts this work more and more to others. That is, the degree of altruism in her behavior decreases, because she understands that she needs to take care of herself, otherwise her wings will fray and she will die.
Question from the audience: That is, she is loosening her waist, preparing to become a uterus?
Alexander Markov: Yes.
Valeria: If there are bacteria that are representatives of two types: altruists and egoists, it turns out to be a kind of consumer society. If there is a tendency towards education, that is, towards an increase in altruists, then everyone ends up with the same genes, this kind of communism results, and there will be no incentive for any progress if everyone is the same, then what if this really happens to human society? Will there be any desire for a transition of world dominance in Asia, if there is such a thing? They are known to be prone to repetition. The Chinese - they copy some inventions.
Boris Dolgin: 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 must be a counterbalance to good, some kind of evil. Will there be ballast?
Alexander Markov: Balancing selection is likely to be at work here. That is, these are frequency-dependent things: the more altruists there are, the more profitable it is to be an egoist among them. If almost everyone is altruistic, and I alone am an egoist, can you imagine, everyone will help me. Very profitable. And in this situation, selfish people begin to quickly multiply and infect this population. Then there are a lot of egoists, no one helps anyone anymore. Only a few altruists work there, in their garden, 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 entire system will die. This is called in evolutionary ethics the “Tragedy of the Common Grazing.” This is a situation when a village has a common pasture, everyone grazes their own sheep there and there is overgrazing, the pasture is depleted. It is necessary to reduce the number of sheep being grazed, but every peasant thinks: let the neighbor remove his own, and I will still graze mine. And everyone is only interested in tending as many of their sheep as possible. This ends with the pasture being completely destroyed and all the peasants dying of hunger. But even when they are already dying of hunger, half have already died, the most profitable strategy for every peasant until the very end is to still 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 tricks, for example, statistical paradoxes or the fact that the altruist still takes for himself, bypassing the common pot, a certain balance is established. That is, given 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 of a hereditary, genetic nature. That's all?
Boris Dolgin: In general, this is not a trivial thesis at all.
Svetlana: And let’s say, from the simplest to children, everything. We don't go any further. And so it’s interesting, what about today, a person, an individual, a group? Right now, today, as we are, countries. What should we call altruism and egoism in this sense?
Boris Dolgin: This question should be asked to psychologists. Thank you.
Svetlana: But the point is that we say: it’s interesting to look at the evolutionary origins. And for what? We live now, today, among people - and just understand: are altruism and selfishness of a genetic nature?
Boris Dolgin: You can leave no comments, 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 its reputation, does the individual weigh the risk of death?
Alexander Markov: Of course, not every time, in general this is quite a rarity, that is, indirect reciprocity is a mechanism of reputation. In humans it is well developed, in birds, and perhaps a little in some higher primates. Of course, these are very smart animals, they have 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 about their interests and preserving their lives.
Zukhra: Once again I want to return to children and psychology, since you talked about it. Caring for talented children, and for me altruism is a moral talent. Are there any tests that measure altruism in children? You talked about such experiments with children, can you elaborate? Do they exist or not?
Alexander Markov: Yes. Lots of different things.
Zukhra: Can their talent be measured?
Boris Dolgin: Sorry, for now we are not talking about talent, but about altruism.
Zukhra: About altruism - yes, but for me this is the highest talent.
Maria Kondratova: You brought up a rather interesting topic - gender differences in altruism - when you talked about these patterns associated with Paleolithic evolution. In connection with the different evolutionary strategies of females and males, is it even possible to talk about differences in altruism: 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 polymorphism that correlates with different behavior within one sex, but are there any correlations between the sexes in vasopressin-oxytocin receptors that determine altruism?
Alexander Markov: Somehow, in people it is usually specific to one of the sexes - the influence of these genes, and the influence of these neuropeptides themselves is different on men and women. Correlation between genders? I don't remember anything specific in this regard.
Boris Dolgin: That is, you have partially outlined the correlation between gender and this factor. As I understand it, the question was a continuation of this topic. Are there any other signs of sex differences? I wouldn’t talk about gender, of course, because gender is social sex.
Alexander Markov: Are there any other 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, frankly, don’t know.
Boris Dolgin: There are works by Geodakyan, but in my opinion they are not substantiated in any way.
Alexander Markov: Yes, these are controversial things. Therefore, it’s difficult to answer.
Konstantin Ivanovich: I would like to say that altruism and civilization are the number of charitable societies and the resources that circulate in these charitable societies. Is it interesting to compare, say, America, Russia, China, Sweden, Germany?
Alexander Markov: Not everything is so simple either.
Question from the audience: Do bacteria have such societies?
Alexander Markov: Charity?
Question from the audience: Yes.
Alexander Markov: In a sense, when they secrete some socially useful substance.
Dmitry Ivanov: Do you agree with the theory of the selfish gene, that it makes sense to consider natural selection not at groups, not even at individuals, but at the level of genes. What exactly is each gene interested in continuing, copying itself as an elementary replicator that has such an opportunity?
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 theory of kin selection is a gene-centric approach.
Dmitry Ivanov: Thus, the gene for altruism... makes it difficult to survive. That is, he can manifest himself only in social societies, that is, only in society?
Alexander Markov: Naturally, if you have no society, if you live alone in a big forest, then what is altruism if there is no one to show it to? This is understandable.
Dmitry Ivanov: There is great competition for resources in society, that is, we have a primitive society where different groups compete with each other. There is a welfare society where everyone is supposedly altruistic and helps each other. Is it possible to be an altruist in such a society?
Boris Dolgin: What kind of societies are there?
Dmitry Ivanov: If we talk hypothetically. Is this the kind of society we want? It turns out that in such a society these same deceivers can spread until the number of altruists again reaches a critically small level and fierce competition for all resources begins again. Logical?
Alexander Markov: What's the question? I don't quite understand.
Dmitry Ivanov: The question is the spread of those very genes of altruism in the human environment.
Boris Dolgin: Do you think that a stable social situation is possible where this gene wins? Did I understand the question correctly?
Dmitry Ivanov: Yes, that this can only be done through education and the development of culture, and not through natural selection?
Alexander Markov: The altruism that arises through education and cultural development faces exactly the same problems. Just like among unconscious beings, some bacteria, in this situation it is advantageous to be an altruist, but in this situation it is disadvantageous to be an altruist. It’s the same in human society - even if we assume that there is no genetic variability for these characteristics, that altruism or selfishness of a person depends only on upbringing. Let's say. All the same, in one situation it will be beneficial to behave altruistically, and in another - selfishly. Let’s say, the more altruists there are, the more profitable it is, the more tempting it is to start behaving like an egoist. Since people are intelligent creatures and actively adapt throughout 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 beneficial for everyone to behave well. The ideal of reciprocal altruism is something we should probably strive for. The golden rule of ethics, it is not by chance that it is 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 them to treat you?
Alexander Markov: Yes.
Dmitry Ivanov: Another small question about children. How was the influence of culture distinguished from the influence of genes in experiments with children? That is, the influence of upbringing received from parents? The fact that he wants to share with others not because his mom and dad raised him that way?
Alexander Markov: But in this experience - no way. In this experiment, genes were not touched; behavior was simply studied and how it changes with age. How it changes with age, how the percentage of certain behavior patterns changes. Unaccountable altruism, desire for equality, and so on. In this particular study, no genes were touched.
Grigory Chudnovsky: If possible, a short discussion and in this sense not a question - if you consider it necessary to comment. The Hamilton equation that you displayed on the screen, both in single and multiple versions, is an exact proportionality that is not regulated by the brain, but by other mechanisms in simple organisms and communities. The exact proportion of what I am willing to lose by transferring to someone else what is important to him. And there it is clear that there is a limit, that even in this inequality there is a limit to what to transmit. That is, some kind of quantum that can be transferred to preserve the life of the organism. What got me interested in this inequality is this limiting 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 ended your religious lecture, including fragments that you did not expand, it is indicated that all liturgical practices and ceremonies so expensive are, as it were, a form of altruism, as I understand . It seems to me that a form of mental suppression, the more expensive and complex the procedure, the more selfish.
Boris Dolgin: This is a little different.
Grigory Chudnovsky: Yes, this is a little different. But I’m just coming to this topic now, that for example, a donation is given based on calculation - this is altruism, right? Give a coin to the poor. But they are calculated, because the rich will become poor if he gives to everyone who asks. This is also the first question, where are the boundaries between altruism, which brings benefits both socially. Thank you.
Alexander Markov: To avoid confusion, I will first answer the first question: where is the border? Everything is just written here, there is no special additional essence here. This is the whole border, it is here, this is the inequality. That's if rB>C, the altruism gene 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 - it's called - the murder of siblings. Some birds are able to feed only one chick, but lay two eggs, just in case. The first chick has hatched; If he is alive by the time the second one hatches, he will peck the second chick - or throw it out. This is their norm of life. Because, in this case, obviously, for them the price of saving their brother’s life turned out to be much higher than this matter. That is, if we consider an altruistic act as not killing a brother. That is, everything depends on the relationship between these variables. That's all. And no mysticism. And I’ve already conveniently forgotten the second question about religion. 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? I think it sounded completely different in your lecture?
Alexander Markov: Not a manifestation of altruism, but you said they can suppress the psyche?
Grigory Chudnovsky: Yes, this is probably why they are created.
Alexander Markov: But there is no contradiction here. It may very well be that the manifestation of parochial altruism, that is, selfless devotion to one’s own, the readiness to die for one’s faith, for one’s co-religionists, can be facilitated by the suppression of the psyche.
Alexander Nikitin: It seems to me that this model: talking about human society is fundamentally unsuitable, because man is fundamentally different from the animal and biological world. He has consciousness, he has goals and objectives, besides reproducing, he also has creative ones. Therefore, an example can also be illustrated according to this model of 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 and that's all. And these people for some reason, due to some forces, set themselves, perhaps, other goals. To write poetry like Pushkin - and from the point of view of primitive Darwinists - they are simply deceivers. And this black and white model, it seems to me, is fundamentally unsuitable.
Alexander Markov: When studying complex objects, you always need to take into account a bunch of everything, a bunch of specifics of all kinds. Naturally, you can apply some methodological approaches correctly to an object and you can apply it incorrectly. It is clear that no one is going to take it head-on - and to any situation: someone is digging, someone is writing poetry - no one applies this formula like that, naturally. 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 didn’t expect to hear something new for myself, I’m very grateful. I listened to a lecture about this back in the school years of 66-67. Whatever you call human exceptionalism, I will give an example that it is not so. 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 Efroimson's ideas, they are simply proven in a way that is little understood, and there is no question associated with it. And immediately the question is the most interesting for me. Still, the egoistic gene - what is meant, and is this whole elegant theory of altruism and egoism 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 “Bilingualism” " more. If everyone is so politically correct, then how can we explain Anglo-Saxon national egoism, and this is an extremely painful issue now, for our world. And lastly, were there any searches and studies of “altruism genes” before Vladimir Pavlovich Efroimson? What is important, I was faced with the fact that many journalists do not even know about the phenomenon that this gene of altruism has circled the globe several times.
Alexander Markov: Last time I was asked two questions in a row, and now you asked four. I would still prefer it to be one question at a time. The first question was: what is a selfish gene - this needs to be read in a separate lecture. There is a book by Dawkins, “The Selfish Gene,” where this is popularly stated. I based my entire lecture on this model. I’m not ready to formulate this in a nutshell right now.
Boris Dolgin: Thank you. The next question was: how much of a role did Efroimson play in the development of the concept?
Alexander Markov: Darwin himself began to think about this topic. He already made the first hints of the theory, then Fischer developed this topic, then Haldane - this was the beginning of the twentieth century. So all these ideas have been developing for quite some time.
Boris Dolgin: The third question, I think, was: would you like to apply this to “media viruses”?
Alexander Markov: To the memes, right? As you probably know, Dawkins wrote about the possibility of drawing an analogy between genes and units of information of cultural inheritance, which may also behave in part like genes. They are also selected, mutated, and spread. Let's say, jokes, some popular pictures, songs, melodies, some sayings, little words, things like that - they also spread partly like genes, like viruses in a population. But can the concepts of altruism and egoism be applied to them? I think it will be a little difficult, because with genes, why does this happen? I said that a gene cannot be altruistic. An altruistic gene would be a genetic variant that sacrifices its own spread to help another competing genetic variant spread. What will happen to such an altruistic gene - it will simply disappear automatically, it will be supplanted. Therefore this cannot be. Altruism arises due to the fact that the interests of genes and the organisms in which these genes reside do not coincide. An organism can be altruistic. Gen - can't. What is the analogue of an organism for a meme? I don't quite understand this, this theory is not very developed.
Boris Dolgin: Well, maybe a tradition?
Alexander Markov: The gene complex does what it creates, builds an organism from a fertilized egg. And what does the complex of memes do?
Boris Dolgin: I am opposed to this metaphor, but if we proceed from it, then it is tradition.
Alexander Markov: It's difficult, you have to think about it.
Evgeniy Teslenko: Thank you very much for the lecture. To be honest, I felt a little scared. Because, if we extend the scientific logical trend, the question arises: with the modern developments of genetic engineering, it is quite possible for the emergence of certain theories, and then practices of correcting the human essence with a great good desire to increase altruists, for example, to reduce egoists.
Boris Dolgin: A continuation of eugenics?
Evgeniy Teslenko: Yes, yes, absolutely right, we are returning to the same eugenics, returning to more rational forms of the structure of humanity, and so on. How do you feel about this, especially since both scientific and technical developments have already come quite close. Why is it your lecture that makes this trend scary? Because it seems that 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 this perfectly at the beginning, that the words, terms, metaphors that are labeled are altruistic. Well, what kind of altruists are these, what kind of egoists are they? Maybe it makes sense for scientists who are engaged in fundamental research to be very careful about such metaphors? Because they cause temptation. What do you think about this?
Alexander Markov: What temptation?
Evgeniy Teslenko: The temptation to use and correct the entrusted people in the right direction.
Boris Dolgin: Do social genetic engineering?
Evgeniy Teslenko: Not social, but technical genetic engineering
Alexander Markov: The temptation here is not because of metaphors. When it comes to people, altruistic, egoistic behavior is no longer metaphors, but is already quite what was originally called that way. If we see that changes in a gene affect the tendency to do good deeds, we are talking about good deeds, and not about the secretion of some 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 understood the question correctly.
Alexander Markov: This is, of course, a complex question. Humanity will have to face this. Of course, it now seems to us that it is impossible to genetically modify a person so that he becomes kinder. But this seems unethical. Let's start from the other end, what 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 its cells, and we will fix it, and then your child will most likely be born healthy and normal. Well, of course, the parents will agree to this. Depriving parents of the opportunity to make such a choice is also wrong. What if there is no genetic disease? But the geneticists 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 his family life, he has a bad 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 built into his brain, and then he will be happy in his family life. So choose, fellow parents. This is a more complex question. Yes, I don't dare 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 broad 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 can be happy in family life?
Sergey Kapustin: I have two questions. The first is an explanation of agriculture among ants. Why do they need the mushrooms to be genetically homogeneous? To prevent them from becoming poisonous, for example? Were they still edible?
Alexander Markov: These mushrooms behave as if altruistic towards the ants. The mushroom has different options. In principle, if these mushrooms were engaged in selfish evolution, inside these anthills or termite mounds, then deceiving mushrooms would necessarily appear there, which would only exploit the ants, but would feed them poorly or would not feed these ants at all. Well, for example, the fungi that live in termite mounds produce two types of fruiting bodies: small, round fruiting bodies to feed the termites, and large, stalked fruiting bodies that grow through the mound and disperse the spores. That is, small fruiting bodies are, roughly speaking, altruism for the termites that feed, grow, and spud them. And large fruiting bodies - it’s like selfishness - the mushroom makes for itself. Accordingly, what will happen if a mutant mushroom appears that spends more energy on the production of large fruiting bodies and less energy on the production of small fruiting bodies? If these fungi are allowed to compete calmly and evolve inside the termite mound, the egoist will win, the fungus that produces the most large fruiting bodies will win, and the termites will remain “with their noses.” They will have less food. So that this does not happen, so that there is no such competition between different strains of mushrooms with different numbers of such and such fruiting bodies, for this they need to be genetically identical. Then their evolution will not work.
Sergey Kapustin: And the second question, we summarize various arguments about memo viruses, such cultural phenomena. How do you feel about this idea: in principle, evolution is the spread and reproduction of genetic information. The gene, as a carrier of information, “has a goal” to replicate itself. Do other media appear in the human environment, in any natural environment, at the information level? That is, a person is a carrier of information in another non-genetic form, an idea, a statement 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 information equivalent of reproduction.
Boris Dolgin: Unfortunately, the question is not entirely clear. Or do you understand?
Alexander Markov: No, unfortunately, I didn’t understand either.
Boris Dolgin: Undoubtedly, people usually want 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, replication of information in a different form - not genetic. Somehow similar to the process of evolution... Question: isn’t altruism, for example, in humans and altruism in nature in general some kind of first step towards moving away from the genetic, that is, they sacrifice their genetic in favor of alternative replication, for example, the idea of altruism .
Boris Dolgin: How do you imagine the mechanism for scientific testing of this hypothesis?
Sergey Kapustin: This is probably hard.
Alexander Markov: This is just an interesting opinion.
Maria Kondratova: Since we know people who sacrifice their lives and their reproductive capabilities for the sake of implementing some ideas, apparently this makes sense for a person to be. That's not what my question is about. I really liked that you included in your report the point that genetic, evolutionary description does not mean justification. Because, unfortunately, this is very often replaced. If there is something in our nature, then it must be so; this is the most common, trivial judgment, but the question that arises is not biological, but more general: what, in this case, can be a justification at the present moment, when religious authority is no longer a justification, human nature, scientific description is a description, but also not a justification, and what, in this case, can be a justification?
Boris Dolgin: Maybe your value system? For you - yours, for Alexander - his.
Maria Kondratova: Then the concept of altruism as a common good, as something specifically common, is lost.
Boris Dolgin: But the system of values is more or less common for some communities. But the idea of the common good is still nothing more than part of this value system.
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 and what is bad for a person now.
Reply from the audience: Therefore, there was no need to talk about people today!
Alexander Markov: I don't agree.
Boris Dolgin: From the very beginning we were going to talk about people, 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 money in, I'm afraid, is not entirely relevant to a biologist.
Irina: Do you have any information based on everything you told us, like the abstracts of previous large materials? What are the prospects?
Boris Dolgin: In other words: what areas of biology do you find most interesting, and where would you advise investing money or what to pay attention to?
Alexander Markov: There is such 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 - brain research. 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, feelings, and so on are formed.
Reply from the audience: This is good?
Alexander Markov: Man knows himself.
Victor: The position that we heard, the work has both practical and any other significance. They are all posted on the website - are your provisions written on the website? Is the entire lecture there?
Boris Dolgin: I’ll 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 said, has been hanging on my website for about five months (evolbiol.ru/altruism.htm). I went to one conference, 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 audience: There was such a three-volume work, published before the revolution, “The Nature of Love.” It examines in great detail, from bacteria to humans, the evolutionary process from the point of view of altruism and all other categories.
Boris Dolgin: Before the revolution of 1917?
Question from the audience: Certainly. So, please tell me, did you rely on this work to some extent?
Question from the audience: Bailey.
Alexander Markov: No, I don't know him.
Which, under certain conditions, reduce the chances of individuals to reproduce, can spread in a population when the amount of contribution to the reproduction of other individuals is greater than the cost of help. In this case, this individual thus produces more copies of its genes than by spending all its 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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Excerpt describing Hamilton's Rule
The postilion set 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 très bien. - He kissed the tips of his fingers. - Et tout a fait francaise. [Well, my dear, your little princess is very sweet! Very sweet and perfect Frenchwoman.]
Hippolytus snorted and laughed.
“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 pretends to be a sovereign person.]
Contrary to popular misconception among laypeople, modern evolutionary biology successfully explains the origins 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 in recent years, which shed 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 is invading the “forbidden” territory where philosophers, theologians and humanists have until now reigned supreme. A central issue in evolutionary ethics is 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 one’s own chances of successful reproduction. This definition essentially differs little from the definitions of altruism adopted in ethics, taking into account the fact that the action of natural selection in the general case is aimed specifically at increasing reproductive success. This allows us to speak metaphorically about it as the main “goal” in which evolving organisms are “interested” in achieving. Of course, we are only talking about the fact that 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 could consciously influence their own evolution, then the direction of evolutionary change would be exactly what is observed in reality. It is in this, partly metaphorical, sense that such concepts 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 through joint efforts than alone. Cooperation, that is, joint problem solving, usually implying some degree of altruism on the part of cooperators, could be an ideal solution to most problems for many organisms. Why, then, has the biosphere never turned into a kingdom 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 if the driving force of evolution is the mechanism of natural selection, which at its core seems to be purely selfish? A primitive, simplified understanding of the mechanisms of evolution can lead to the completely 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 “the struggle for existence” and especially “survival of the fittest.” If the fittest always survive, what kind of altruism can we talk about?
The error in such reasoning lies in the confusion of 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 gene level. Therefore, it is from the genetic level that consideration should begin. Here, the basis of evolution is 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, there cannot be. Gene is always selfish. If an “altruistic” allele appears, which, to its detriment, allows another allele to reproduce, such an “altruist” will be automatically forced out of the gene pool and disappear.
Relatedselection
However, if we move our view 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 meaning that evolutionary biologists attach to the concept of “interest”). The discrepancy of interests stems from the discrepancy in 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 contrary, is a single entity, each cell of which usually carries only one or two of these copies. In many situations, it is beneficial for a selfish gene to sacrifice one or two copies of itself in order to provide an advantage to the remaining copies that are contained in other organisms.
Biologists began to approach this idea already in the 30s of the 20th century. Important contributions to understanding the evolution of altruism were made by R. Fisher (Fisher 1930), J. Haldane (Haldane 1955) and W. Hamilton (Hamilton 1964). The theory they developed is called the theory of kin selection. Its essence was figuratively expressed by Haldane in the famous 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”). The altruism gene (more precisely, the allele promoting altruistic behavior) will be supported by selection and spread in the population if:
rB > C,
Where r – the degree of genetic relationship between the “donor” and the “receiver of the sacrifice” (the probability that the latter’s genome contains the same “altruism allele” depends on it); B – reproductive advantage received by the recipient of the altruistic act; C – reproductive damage caused by the “sacrificer” to himself. Reproductive advantage or disadvantage can be measured, in part, by the number of offspring produced (or not produced). 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 accepting 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 nrB > C, the altruism allele 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 forces its carriers (organisms) to behave altruistically, but thereby the allele looks after its “selfish interests.” An allele sacrifices several copies of itself to give an advantage to other copies contained in the bodies of closely related organisms. Natural selection is the process of automatically weighing the sum of the gains and losses for an allele (for all its copies together!), and if the gains outweigh it, the allele spreads.
Hamilton's rule has remarkable explanatory and predictive power. In particular, it allows us to explain the repeated occurrence of eusociality in insects of the order Hymenoptera(Hymenoptera). In eusocial Hymenoptera (ants, bees, wasps, bumblebees), most females forego their own reproduction in order to help the mother raise other daughters. Apparently, an important factor promoting the development of eusociality in this particular 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 relatedness between sisters and between mothers and daughters is the same (50% of common genes, the value r in Hamilton's formula is equal to 1/2). In Hymenoptera, siblings share 75% of their genes (r = 3/4), because each sister receives from their father not a randomly selected half of his chromosomes, but the entire genome. Mother and daughter in Hymenoptera, like other animals, have only 50% of their genes in common. Therefore, in order to effectively transmit their genes to the next generations, female Hymenoptera, all other things being equal, are more profitable 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 ensures an abnormally high level of genetic relatedness between individuals in the colony (Hughes etal. 2008).
Kin selection appears to underlie many cases 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 look at these mechanisms using specific 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, given the necessary minimum conditions, is capable of quickly evolving before the eyes of researchers, mastering new niches and developing original adaptations.
In order for a social system to develop beyond the very first steps, it needs to develop a mechanism to combat deceivers. Such mechanisms are sometimes actually developed. This often leads to an evolutionary “arms race”: deceivers improve methods of deception, and cooperators improve methods of identifying deceivers, fighting them, or trying to prevent the very appearance of deceivers.
The ability to defend against deceivers may appear as a result of single mutations
Let's look at another example involving bacteria. Myxococcus xanthus. These microbes are characterized by complex collective behavior. Sometimes they gather in large clusters and organize a collective “hunt” for other microbes. “Hunters” release toxins that kill “prey” and then absorb organic substances released during the breakdown 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 actions of millions of individual bacteria, of which only a part receives direct benefit, 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 subsequent generations. The rest act as “building material”, doomed to die without leaving offspring.
In this experiment, the altruists were never able to develop a defense against deceivers. Something else happened: the deceivers themselves experienced a mutation, as a result of which the bacteria restored the lost ability to independently form fruiting bodies and at the same time received an additional advantage (!). These mutant bacteria were protected from parasites, that is, from their direct ancestors - deceiving bacteria. Thus, a single mutation turned deceivers into altruists, protected from deception. The mutation occurred in one of the regulatory genes that influence the behavior of bacteria. The specific molecular mechanism for this effect has not yet been elucidated (Fiegna etal. 2006).
The problem of deceivers is also familiar to more complex single-celled organisms, such as social amoebas. Dictyoste-lium. Like many bacteria, these amoebas, when there is a lack of food, gather into large multicellular aggregates (pseudoplasmodia), from which fruiting bodies are then formed. Those amoebas whose cells go to build the stem of the fruiting body sacrifice themselves for the sake of their comrades, who get a chance to turn into spores and continue the race (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 beyond 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 in the body).
As already mentioned, in order to survive, social organisms need to protect themselves from parasites. Experiments conducted on amoebas showed that the likelihood of resistance developing as a result of random mutations in this organism is also quite high, as in myxococci (Khare etal. 2009). 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, the deceivers occupy the best places in the fruiting body and turn into spores, leaving honest amoebas to build the stem of the fruiting body alone. As a result, among the resulting disputes, disputes of deceivers predominate.
During the experiment, honest amoebas had an artificially increased 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 reproduce. After this, selection for resistance to parasites began, and the parasites themselves were used as a selecting agent. Amoebas from thousands of mutant strains were mixed in equal proportions and combined with deceptive amoebas. The mixed population was kept under conditions of lack of food, forcing them to form fruiting bodies. Then the resulting spores were collected and the amoebas were removed from them. Naturally, deceivers predominated among them, but the experimenters killed all the deceivers with an antibiotic (a gene for resistance to this antibiotic was previously inserted into the genome of honest amoebas). The result was a mixture of mutant amoebas, but from the thousand original strains, it was now dominated by those who were able to resist the deceivers better than others. These amoebas were again mixed with the dupes and again forced to form fruiting bodies.
After six such cycles, representatives of only one of the thousand original strains remained in the population of mutant amoebas. These amoebas were reliably protected from deceivers as a result of the mutation that occurred in them. Moreover, they protected themselves not 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 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 mutant amoebas, and different genes were mutated and different mechanisms of resistance emerged. Some resistant strains themselves became deceivers in relation to the “wild” amoebas, 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, subject to 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 original population there were equal numbers of altruists and egoists. Then the population 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 ensured by simple chance). As each individual subpopulation grows, altruists lose out (their share decreases). However, those subpopulations that initially had more altruists grow faster due to the fact that they have at their disposal more of the “socially useful product” produced by altruists. As a result, if you add together the increased subpopulations, it turns out that the “global” percentage of altruists has increased. The fundamental possibility of such a mechanism for maintaining the number of altruists was assumed by Haldane and Hamilton, but experimental evidence of the effectiveness of Simpson’s paradox was obtained only recently (Chuang et al. 2009). The main difficulty was that in each specific 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 whether Simpson's Paradox alone could make altruists thrive, a model system of two strains of genetically modified E. coli was created. The gene for an enzyme that synthesizes the signal substance N-acyl-homoserine lactone, used by some microbes for chemical communication, was added to the genome of the first of the two strains (“altruists”). In addition, a gene for an enzyme that provides resistance to the antibiotic chloramphenicol was added to the genome of both strains. A promoter was attached to this gene, activating the gene only if the above-mentioned signal 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 altruists is necessary for both strains for successful growth in the presence of the antibiotic. The benefit received by both strains from the signal 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 did not receive additional benefits from their altruism.
In a medium with the addition of an antibiotic, pure cultures of egoists, as expected, grew worse than pure cultures of altruists (since in the absence of a signal substance, the gene for protection against the antibiotic in egoists remained turned off). However, they began to grow better than altruists if either living altruists or a purified signaling substance were added to the environment. Altruists in a mixed culture grew more slowly because they had to spend resources on the synthesis of a signaling substance. Having verified that the model system worked as expected, the researchers began simulating Simpson's Paradox.
To do this, they placed mixtures of two cultures in different proportions in 12 test tubes with a medium containing an antibiotic, waited 12 hours, and then measured the number of bacteria and the percentage of altruists in each test tube. It turned out that in all test tubes the percentage of altruists decreased significantly. Thus, altruists in all cases lost in competition to egoists. However, the size of those populations where there were initially more altruists grew significantly more 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 noticeably: Simpson’s paradox “worked” successfully.
However, in nature, no one will deliberately mix altruists with egoists in different proportions and place 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 severe population decline followed by its subsequent recovery. 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 by simple chance there may be an increased percentage of altruists among them. The population formed by this founding group will grow quickly, while other populations founded by groups of microbes dominated by egoists will grow slowly. As a result, Simpson's paradox will ensure an increase in the “global” share of altruists in the totality 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 into test tubes in portions of different sizes with an approximately known number of microbes in each portion. The size of the portions turned out to be the main factor on which the further fate of the altruists depended. As would be expected, when portions were large, Simpson's paradox did not occur. In a large portion, that is, in a large sample from the original culture, the ratio of altruists and egoists, according to the laws of statistics, cannot differ much from the original one. The populations founded by these samples grow at approximately the same rate, and altruists are the losers not only in each individual population, but in all populations as a whole.
However, if each portion contained only a few bacteria, then among these portions there were sure to be those in which altruists predominated. Such founding groups gave rise to rapidly growing colonies, and due to this, the overall percentage of altruists in the aggregate of all populations increased. In the specific conditions of this experiment, for the Simpson effect to manifest itself, it is necessary that the average number of microbes in the founder group be no more than 10. The authors also showed that, after repeating this sequence of actions several times (dilute the culture, settle in small groups in test tubes, grow, connect populations in one, dilution again, etc.), you can achieve an arbitrarily high percentage of altruists in a culture.
Another condition necessary for the spread of “altruism genes” in the model system was identified: mixed populations should not be allowed to grow for too long. Dilution and dispersal must be carried out before populations reach a stable population level, populating the entire culture medium in a test tube, because then differences in population levels between populations are smoothed out and Simpson's paradox cannot occur (Chuang etal. 2009).
Thus, natural selection, under certain conditions, can ensure the development of altruism even when in each individual population it favors egoists and condemns altruists to gradual extinction. However, the range of conditions under which Simpson's paradox can operate is quite narrow, and therefore its role in nature is probably small.
Altruists and deceivers among social animals
The greatest triumph of the evolution of altruism was the emergence of true multicellular organisms, including animals. Animals, compared to microbes, have new opportunities for the development of cooperation and altruism, based on complex behavior and learning. But the same new opportunities opened up for deceivers. Deceivers learned to deceive cooperators more cunningly, and they, for their part, began to develop new methods of identifying deceivers and combating them. The evolutionary “arms race” continued at a new level, and again neither altruists nor deceivers received a decisive advantage.
One of the important innovations in this endless war was the possibility of physical (and not just chemical) punishment of deceivers. This phenomenon occurs, in particular, in social insects. Working individuals of Hymenoptera usually do not reproduce, 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. Let us recall that in Hymenoptera males develop from unfertilized eggs. Due to the peculiarities of sex inheritance, the most profitable strategy for female Hymenoptera is to raise other people's daughters (their sisters) and their 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 serve as a kind of “morality police.”
Recently, German entomologists tried to test which of two factors is more important for maintaining altruism in insect society: voluntary adherence to the principle of “reasonable selfishness,” i.e., kin selection in its purest form (1), or “police surveillance” (2) (Wenseleers, Ratnieks 2006). For this purpose, data on 10 species of social Hymenoptera were processed. It turned out that the stricter the “morality police,” the less often workers commit acts of selfishness by laying their own eggs. We also tested the influence of the degree of relatedness between workers in a nest on altruistic behavior. The degree of relatedness between them in reality is often lower than the ideal 75%, since the queen can mate with several different males. It turned out that the lower the degree of kinship between sister workers, the stronger the “police surveillance”, and the less often workers behave selfishly. This corresponds to the second hypothesis (about the leading role of police measures). With a low degree of relatedness 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 profitable, but, as can be seen from the results obtained, effective “police supervision” clearly outweighs the selfish aspirations of working individuals (Wenseleers, Ratnieks 2006).
The peculiarities of sex inheritance in Hymenoptera played an important role in the development of altruistic behavior and sociality, however, in many modern species, altruism is maintained 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 as those observed in Hymenoptera families, will still be destroyed by deceivers if it fails to develop additional means of combating selfishness.
This pattern may also 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 ultimately everyone benefits from this. However, in many cases it is still beneficial for each individual to act selfishly, pursuing selfish interests to the detriment of the collective. And to effectively combat egoism, one has to use violent methods.
Let's consider another example showing that the altruism of social insects is far from the ideal of selflessness. Wasps Liostenogasterflavolineata They 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 oldest 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 to get food for the larvae, which is associated with wear on the wings and the risk of being caught by a predator. With the promotion of an assistant to the rank of queen, her life expectancy increases sharply (Field et al. 2006).
In this species, as in many others, helper wasps vary greatly in the degree of “work enthusiasm”. Some, without sparing themselves, spend up to 90% of their time searching for food, while others prefer to sit in a safe nest and fly out for food much less often. At first glance, these differences are difficult to explain from the standpoint of the theory of kin selection, since the degree of work enthusiasm of the 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 of becoming a queen and leaving her own offspring. If these chances are small (as for low-ranking young wasps, the last in the “queue” for the royal throne), then it makes sense to work more actively in order to pass on their genes to future generations, at least 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, the wasp occupying the second place in the hierarchy was removed (i.e., the first in seniority after the queen), and from another family of the same size, a low-ranking young wasp was removed. After this, they monitored the behavior of the wasp, which before the experiment occupied third place in the hierarchy. In the first nest, after the removal of the senior assistant, this wasp increased its rank, moving from third place to second; in the second, it remained in third place. The size of both families remained the same. It turned out that in the first case the wasp begins to work approximately half as much. In the second case, when a low-ranking assistant was removed from the nest, wasp number three continued to work as much as before (Field etal. 2006).
These results indicate that the amount of "altruistic effort" in wasps is indeed regulated depending on a given wasp's chances of its own reproductive success. In other words, the tendency towards altruism is stronger among those who have nothing to lose. The appearance of such behavior during 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 emergence of cheaters
Is it possible to create a social system where altruism will be supported without violence and where there will be no deceivers and egoists? Neither wasps nor people have succeeded in this yet. 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 the cooperative system to zero. This eliminates the possibility of competition between genetically different varieties of symbionts over 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 from the minimum set of conditions necessary for evolution - the Darwinian triad of “heredity, variability, selection” - one of the components, namely variability, is excluded. As a result, the evolutionary interests of 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 has never succeeded, despite repeated “attempts,” to create a full-fledged multicellular organism from genetically heterogeneous cells. All true 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 the symbionts is to transmit them vertically, that is, 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-cutter ants also pass on their crops from generation to generation. With vertical transmission, 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, the symbionts of each host are genetically heterogeneous, they retain the ability for selfish evolution, and therefore deceivers appear among them every now and then. For example, strains of deceivers are known among luminous bacteria (symbionts of fish and squid), nitrogen-fixing bacteria-rhizobia (symbionts of plants), mycorrhizal fungi, and zooxanthellae (symbionts of corals). In all these cases, evolution failed to ensure the genetic homogeneity of the symbionts, and the hosts have to fight the deceivers using 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, Simpson's paradox or balancing selection, which is based on the fact that sometimes being a deceiver is beneficial 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 short-term benefits and is completely indifferent to long-term evolutionary prospects.
In order for a mechanism to develop that ensures the genetic homogeneity of symbionts, this mechanism must provide an immediate benefit, otherwise selection will not support it. The benefit that we have talked about so far - depriving symbionts of the opportunity to evolve into deceivers - belongs to the category of “distant prospects” and therefore cannot work as an evolutionary factor at the microevolutionary level. But if some species is so lucky that the vertical transmission of symbionts will be associated with immediate benefits for it and therefore will be secured by selection, this can provide its distant descendants with triumphant success.
Termite subfamily Macrotermitinae those who have mastered effective “agriculture”—growing mushrooms—still seemed to be the exception to the rule. The transmission of symbionts (domesticated mushroom crops) is not vertical, but horizontal, but deceiver mushrooms are completely absent from 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 subfamily of mushroom-growing termites includes 10 genera and about 330 species, which play an important role in the circulation of substances and the functioning of tropical communities of the Old World. Unlike mushrooms grown by leaf-cutter ants, mushrooms “domesticated” by termites have already lost the ability to exist independently. They grow only in termite mounds on specially arranged beds made from plant material passed through the intestines of termites.
Having founded a new colony, termites collect fungal spores in the surrounding area. Termitomyces and sow their plantations with them. Naturally, the initial seed material turns out to be genetically very heterogeneous. Fungi 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 serve for the reproduction of fungi inside the termite mound. Termites feed on the nodules, and the spores pass through their intestines intact and are used to seed new plantations.
Fungi also need to take care of getting into new termite mounds. Conidia usually do not spread beyond the termite mound. For this purpose, sexual spores (basidiospores) are used. They are formed in fruiting bodies of a different type - large ones that grow outward through the walls of the termite mound. Small haploid mycelia grow from basidiospores brought by termites to a new nest. Cells of different haploid mycelia merge and turn into dikaryons - cells with two haploid nuclei. From them grow large dikaryotic mycelia 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.
Thus, fungi produce small fruiting bodies mainly for termites (altruism), and large ones mainly for themselves (selfishness). The trickster fungi's strategy could be, for example, to produce more large fruiting bodies and spend fewer resources feeding termites. But among the mushrooms Termitomyces there are no deceivers, and until now it was not known why. This mystery was only recently solved. It turned out that only one strain of fungi is grown in each termite mound. At the same time, different strains are cultivated in different termite mounds. Consequently, termites prevent the appearance of deceivers in the usual way - through 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 strains of fungi during dense sowing, combined with the fact that the reproduction of fungi inside a termite mound is completely controlled by termites. U Termitomyces 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 arises 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 after several cycles of “reseeding” carried out by termites.
Positive feedback is based on the fact that processes of dikaryotic mycelia can grow together 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 increased yields in monoculture and the displacement of “minorities”.
Apparently the wild ancestor of fungi Termitomyces turned out to be a successful candidate for “domestication” precisely because it was inclined to form monocultures when sowed densely. The increased yield of monocultures could become the “momentary advantage” that allowed selection to support and develop this tendency in the early stages of the formation of the symbiosis. In the long-term (macroevolutionary) perspective, it turned out to be decisive, because it saved the mushroom-growing termites from the threat of the appearance of deceiver mushrooms. Ultimately, this ensured the symbiotic system's evolutionary success ( Ibid. ).
During the transition of people from hunting and gathering to food production (Neolithic Revolution), the problem of selecting candidates for domestication, apparently, was also extremely acute. A good symbiont is very rare, and in many regions there simply were no suitable species of animals and plants. Where they were most numerous, human civilization began to develop at the greatest speed (Diamond 1997).
The examples considered suggest that if not for the problem of deceivers, generated by evolution’s lack of foresight and concern for the “good of the species” (not 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 set of specific circumstances helps to curb deceivers or prevent their emergence. There are not many good “engineering solutions” to deal with the problem of cheaters. Evolution has repeatedly “stumbled upon” each of them in its wanderings through the space of the possible.
Intergroup competition promotes intragroup cooperation
If in a certain animal species cooperation has already developed so much that the species has switched to a social way of life, then additional mechanisms may come into play to further strengthen intragroup cooperation. In social animals, an individual, as a rule, can reproduce successfully only by being a member of a successful group. Moreover, 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, Hölldobler 2007). The purpose 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 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 a given individual. The share that ultimately goes to each individual depends on the ratio of his own egoistic efforts and the sum of the egoistic efforts of the remaining members of the group. Something similar is observed in social insects when they carry out “mutual supervision” - they prevent each other from laying eggs, while trying to lay their own (see above).
The model also builds relationships between groups on the same principles. This creates a nested, two-level tug-of-war. The more energy individuals spend on intra-group struggle, the less it remains for inter-group “pulling” and the smaller the “common pie” of the group turns out to be.
The study of this model using game theory showed that it explains empirically observed patterns well. The model confirmed that intragroup cooperation should increase with increasing intragroup relatedness (which is fully consistent with the theory of kin selection). But the model also showed that cooperation can occur even in the absence of any relatedness between group members. 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 brings a team together more than joint opposition to other groups; 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 certain models developed within the framework of evolutionary ethics to humans, 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. Using 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 answer lies in the genes that determine behavior, and not in upbringing, culture, traditions, etc. With primates , especially with humans, is more complicated: 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 memes such as moral norms, rules behavior in society, etc.) (Dawkins 1976).
Research in recent years has shown that the moral qualities of people are largely determined by genes, and not just by upbringing. Available methods allow us to evaluate only the tip of the iceberg - those hereditary traits for which variability has remained in modern people and which have not yet been recorded in our gene pool. Many of the alleles that ensured the growth of altruism in our ancestors were fixed long ago, that is, they reached one hundred percent frequency. All people have them, and therefore methods such as twin and comparative genetic analysis can no longer identify 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 thereby created a “nutrient medium” for the spread and evolution of memes. Any healthy person, with appropriate upbringing, is capable of learning to behave more or less “cooperatively” and “altruistically.” This means that everyone has a certain genetic basis for altruism (the corresponding genes are firmly fixed in the human population). However, until recently there was very little experimental data on the basis of which one can judge what phase the evolution of altruism in modern humanity is in: has the “genetic” stage already ended, so that only the socio-cultural aspects of this evolution are relevant today, or The evolution of altruism continues at the gene level.
In the first case, we should expect that the hereditary variability of people in terms of traits associated with altruism is very small or completely absent, and the behavioral, 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 explained by genes. In part, because the role of external factors in the development of the human personality is too obvious to deny. The question is posed as follows: Do individual genetic differences have any influence on the observed variability of people in the degree of cooperation, altruism and mutual trust?
In search of an answer to this question, twin analysis is used, in particular. Using special tests, they determine the degree of altruism (or, for example, qualities such as gullibility and gratitude) in many pairs of identical and fraternal twins, and then compare the similarity of the results between different pairs. If identical twins are more similar to each other on a given 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 among people in the degree of gullibility and gratitude are at least 10–20% genetically determined (Cesarini etal. 2008).
Specific genes are also identified that influence 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, it turned out that in humans, peronasal administration of oxytocin increases trustingness and generosity (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 may influence people's tendency to engage in altruistic behavior. Recently, a connection was discovered between some allelic variants of the oxytocin receptor gene ( OXTR) and the tendency of people to show selfless altruism. The oxytocin receptor is a protein produced by some brain cells and is responsible for their sensitivity to oxytocin. Similar properties were also found in the vasopressin receptor gene ( AVPR1a). The regulatory regions of these genes contain so-called single nucleotide polymorphisms. These are nucleotides that can be different 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 a lower, and others a greater, propensity for altruism (Israel etal. 2009). Such facts indicate that altruism in people, even today, can still develop under the influence of biological mechanisms, and not just socio-cultural factors.
Altruism, parochialism and the desire for 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 is found in animals intelligent enough to choose reliable partners, monitor their reputation and punish deceivers, because systems based on mutual altruism are extremely vulnerable and generally cannot exist without effective means of combating deceivers.
Truly selfless care for nonrelatives is rare in nature (Warneken and Tomasello 2006). Perhaps humans are 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 “friend” for us does not always coincide with the concept of “relative.”
Recently, an interesting theory was proposed according to which altruism in humans developed under the influence of frequent intergroup conflicts (Choi and Bowles 2007). According to this theory, altruism among our ancestors was aimed mainly at members of the “own” group. Using mathematical models it was shown that altruism could only develop in combination with parochialism (hostility towards strangers)(!). In conditions of constant wars with neighbors, the combination of intragroup altruism with parochialism provides the greatest chances for the successful reproduction of an individual. Consequently, such seemingly opposite human properties as kindness and belligerence may have developed in a single complex. Neither of these traits alone would benefit their owners.
To test this theory, facts are needed, which can be obtained, in particular, through psychological experiments. Oddly enough, we still know very little about how the formation of altruism and parochialism occurs during the development of children. Recently, the gap has begun to be filled thanks to special experimental studies (Fehr etal. 2008).
Among children, there are about 5% of good-natured people, selfless altruists who always take care of others, and the proportion of such children does not change with age. There are “bad guys” 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 divide everything equally; the proportion of such children grows rapidly with age.
The results obtained also agree well 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 mental properties is generally repeated during 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. The main participants in intergroup conflicts and wars have always been men. 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 “preserving justice” at their expense. As for women, if the group lost in an intergroup conflict, their chances of successful reproduction did not decrease as much as for men. For women, the consequences of such a defeat could be limited to only a change of sexual partner, while men could die or be left without wives. In case of victory, women also clearly won 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 genes. This phenomenon is known as the “Baldwin effect” and has nothing to do with the Lamarckian inheritance of acquired characteristics (Dennett 2003).
Are intergroup wars the cause of altruism?
The idea that the origins of human morality should be sought in the instincts that developed among our ancestors in connection with the social way of life was expressed by Charles Darwin (1896); He also came up with the idea of a 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 (and the concept of “reproductive success” also includes the transmission 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 survivors have a reduced chance of raising healthy and numerous offspring. For example, during intergroup conflicts among chimpanzees, groups that lose in the fight with neighbors gradually lose both their members and territory, i.e., access to food resources.
Secondly, intergroup enmity among our ancestors must have been quite acute and bloody. This is much more difficult to prove.
Thirdly, the average degree of genetic relatedness between fellow 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 towards strangers, tried to assess whether the tribes of our ancestors were at odds with each other strongly enough and whether the degree of relatedness within the group was high enough for natural selection to 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 assessed by the level of mortality in wars; 2) 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 kinship within a group exceeds kinship between warring groups; 4) on the size of the group.
To understand the range of these four parameters in groups of primitive people, Bowles drew on extensive archaeological data. He concluded that conflicts in the Paleolithic were quite bloody: between 5 and 30% of all deaths apparently occurred in intergroup conflicts. In the book by A.P. Nazaretyan “Anthropology of violence and 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 based on data from archaeology, genetics and ethnography. As a result, there remains only one value that is almost impossible to assess directly - the degree to which the group’s military successes depend on the presence of altruists (heroes, brave men) in it. Calculations have shown that even at the lowest values of this value, natural selection in hunter-gatherer populations should help maintain a very high level of intragroup altruism. A “very high” level in this case corresponds to values of the order of 0.02–0.03. In other words, "gene for altruism» will be distributedVpopulations, if the chances of survivalAndleave offspringatcarrier of such a gene on 2–3 % below, howatselfish fellow tribesman. It may seem, What 2–3 % – not a very high level of self-sacrifice. However, in reality this is a significant amount. Bowles provides two illustrative calculations.
Let the initial frequency of occurrence of a given allele in a 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 to pay. 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 selfish people hide behind their backs. Calculations have shown that in order for the degree of altruism to be equal to 0.03, the military mortality rate among altruists must be over 20% (taking into account the real frequency and bloodshed of Paleolithic wars), i.e., every time a tribe faces its neighbors for dear life , and in death, every fifth altruist must sacrifice his life for the sake of common victory. Admittedly, this is not such a low level of heroism (Bowles 2009). This model is applicable to the aspects and cultural factors of altruism, transmitted through training and education.
Thus, the level of intergroup aggression among primitive hunter-gatherers was quite sufficient for the “altruism genes” to spread among people. This mechanism would work even if within each group selection favored exclusively egoists. But this condition, most likely, was not always observed. Selflessness and military exploits could enhance the reputation, popularity, and therefore reproductive success of people in primitive groups.
This mechanism of maintaining altruism through 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 blackbirds Turdoides squamiceps only high-ranking males have the right to feed their relatives. These social birds compete for the right to perform a “good deed” (sit over the nests as a “sentinel”, help care for the chicks, feed a friend). Altruistic acts have acquired partly symbolic meaning among them and serve to demonstrate and maintain their own status (Zahavi 1990). Reputation issues are extremely important in any human group. According to one authoritative hypothesis, an important stimulus for the development of speech among our ancestors was the need to gossip. Gossip within the framework of this hypothesis is considered as the oldest means of disseminating incriminating information about “unreliable” members of society, which contributes to team unity and punishment of deceivers (Dunbar 1998).
It is impossible to cover all areas of research related to the evolution of altruism in one review. Outside the scope of this article were, in particular: 1) works devoted to the study of innate psychological predispositions discovered in humans to effectively identify deceivers; 2) the phenomenon of “costly punishment” ( costly punishment), which manifests itself in the fact that people are willing to make sacrifices in order to effectively punish cheaters (this can also be considered a form of altruism, because a person sacrifices his interests for the sake of what he considers to be the 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 research, it is the parts of the brain associated with emotions that play a key role in solving moral dilemmas; the emotion of disgust was probably “recruited” during evolution to form a hostile attitude towards strangers) ; 4) studying the role of religion, “expensive” rituals and religious rites as a means of enhancing parochial altruism (see: Markov 2009), etc.
In conclusion, it is necessary to briefly consider 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 laws (has an evolutionary explanation), this does not mean that this behavior has thereby received an evolutionary “justification”, that it is good and correct. For example, hostility to strangers and wars with foreigners were 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 aimed only at “our own people”, 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 explains, but does not justify, our innate tendencies. Currently, the development of moral and ethical standards is determined by cultural and social evolution to an immeasurably greater extent than biological evolution, which proceeds much more slowly, and therefore its influence on changes in moral zeitgeist(“spirit of the times”) over short time periods (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, promptly revising the outdated ethical framework that evolution imposed on our ancestors. Not all emotional and behavioral stereotypes that contributed to the spread of the genes of Stone Age hunters are optimal for modern civilized people. In particular, evolutionary ethics warns us that we have an innate tendency to divide people into friends and strangers and to experience disgust and hostility towards strangers. We, as intelligent beings, must understand and overcome this.
Literature
Grinin, L. E., Markov, A. V., Korotaev, A. V. 2008. Macroevolution in wildlife and society. M.: LKI/URSS.
Darwin, Ch. 1896. Human origins and sexual selection/ lane I. Sechenov. SPb.: Publishing house. 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: Beneficial Adaptation, Evolutionary Byproduct, or “Brain Virus”? Historical psychology and sociology of history 2(1): 45–56.
Nazaretyan, A. P. 2008. Anthropology of violence and culture of self-organization. Essays on evolutionary-historical psychology. 2nd ed. M.: LKI/URSS.
Aanen, D. K., de Fine Licht, H. H., Debets, A. J. M., Kerstes, N. A. G., Hoekstra, R. F., Boomsma, J. J. 2009. High Symbiont Relatedness Stabilizes Mutualistic Cooperation in Fungus-Growing Termites. Science 326: 1103–1106.
Alexander, R. D. 1987. The biology of moral systems. N. Y.: Aldine De Gruyter.
Bowles, S. 2009. Did Warfare Among Ancestral Hunter-Gatherers Affect the Evolution of Human Social Behaviors? Science 324: 1293–1298.
Cesarini, D., Dawes, C. T., Fowler, J. F., Johannesson, M., Lichtenstein, P., Wallace, B. 2008. Heritability of Cooperative Behavior in the Trust Game. 105(10): 3721–3726.
Choi, J. K., Bowles, S. 2007. The coevolution of parochial altruism and war. Science 318: 636–640.
Chuang, J. S., Rivoire, O., Leibler, S. 2009. Simpson's Paradox in a Synthetic Microbial System. Science 323: 272–275.
Dawkins, R. 1976. The Selfish Gene. Oxford: Oxford University Press.
Dennett, D. 2003. The Baldwin Effect, a Crane, not a Skyhook. In Weber, B. H., Depew, D. J., Evolution and learning: The Baldwin Effect Reconsidered. Cambridge, MA: MIT Press, p. 69–106.
Diamond, J. 1997. Guns, Germs, and Steel: The Fates of Human Societies. N. Y. Norton & Company.
Donaldson, Z. R., Young, L. J. 2008. Oxytocin, Vasopressin, and the Neurogenetics of Sociality. Science 322: 900–904.
Dunbar, R. 1998. Grooming, gossip, and the evolution of language. Cambridge, Ma: Harvard University Press.
Fehr, E., Bernhard, H., Rockenbach, B. 2008. Egalitarianism in Young Children. Nature 454: 1079–1083.
Fiegna, F., Yu, Y.-T. N., Kadam, S. V., Velicer, G. J. 2006. Evolution of an Obligate Social Cheater to a Superior Cooperator. Nature 441: 310–314.
Field, J., Cronin, A., Bridge, C. 2006. Future Fitness and Helping in Social Queues. Nature 441: 214–217.
Fisher, R. A. 1930. The Genetical Theory of Natural Selection. Oxford: Clarendon Press.
Gore, J., Youk, H., van Oudenaarden, A. 2009. Snowdrift Game Dynamics and Facultative Cheating in Yeast. Nature 459: 253–256.
Haldane, J.B.S. 1955. Population Genetics. New Biology 18: 34–51.
Hamilton, W. D. 1964. The Genetic Evolution of Social Behavior. Journal of Theoretical Biology 7(1): 1–52.
Hughes, W. O. H., Oldroyd, B. P., Beekman, M., Ratnieks, F. L. W. 2008. Ancestral Monogamy Shows Kin Selection is Key to the Evolution of Eusociality. Science 320: 1213–1216.
Israel, S., Lerer, E., Shalev, I., Uzefovsky, F., Riebold, M. et al. 2009. The Oxytocin Receptor (OXTR) Contributes to Prosocial Fund Allocations in the Dictator Game and the Social Value Orientations Task. Public Library of Science ONE 4(5): e5535.
Kessin, R. H. 2000. Cooperation can be dangerous. Nature 408: 917–919.
Khare, A., Santorelli, L. A., Strassmann, J. E., Queller, D. C., Kuspa, A., Shaulsky, G. 2009. Cheater-resistance is not futile. Nature 461: 980–982.
Maynard Smith, J. 1982. Evolution and the Theory of Games. Cambridge: Cambridge University Press.
Rainey, P.B. 2007. Unity from conflict. Nature 446: 616.
Reeve, H. K.,Holldobler, B. 2007. The emergence of a superorganism through Intergroup Competition. Proceedings of the National Academy of Sciences USA 104(23): 9736–9740.
Stoner, D.S., Weissman, I.L. 1996. Somatic and Germ Cell Parasitism in a Colonial Ascidian: Possible Role for a Highly Polymorphic Allorecognition System. Proceedings of the National Academy of Sciences USA 93(26): 15254–15259.
Trivers, R.L. 1971. The Evolution of Reciprocal Altruism. Quarterly Review of Biology 46: 35–37.
Warneken, F., Tomasello, M. 2006. Altruistic Helping in Human Infants and Young Chimpanzees. Science 311: 1301–1303.
Wenseleers,T.,Ratnieks, F.L.W. 2006. Enforced Altruism in Insect Societies. Nature 442: 50.
Zahavi, A. 1990. Arabian babblers: The Quest for Social Status in a Cooperative Breeder. In Stacey, P. B., Koenig, W. D. (eds.), Cooperative Breeding in Birds: Long-term Studies of Ecology and Behavior. Cambridge: Cambridge University Press, p. 103–130.
“...we are faced with two main questions. On the one hand, it is clear that many life problems are easier to solve through joint efforts than alone.
Why, then, has the biosphere never turned into a kingdom 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 if the driving force of evolution is natural selection - a process that, at first glance, seems absolutely selfish?
The thing is that this “first glance” is wrong.
The mistake here is the confusion of 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 gene level, evolution is based on competition between different variants (alleles) of the same gene for dominance in the gene pool of a population. At the genetic level there is no altruism and there cannot be. Gene is always selfish. If a “good” allele appears, which, to its detriment, allows another allele to reproduce, then this altruistic allele will be forced out of the gene pool and simply disappear.
But if we shift our gaze from the level of genes to the level of organisms, then 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 we, and natural selection are completely indifferent to which of the identical molecules will multiply and which will not. Only the total result is important: how many copies of the allele there were and how many there were.
An organism, on the contrary, is a single object, and its genome may contain, to put it simply, only one or two copies of the allele of interest to us.
Sometimes it is beneficial for a selfish gene to sacrifice one or two copies of itself in order to provide an advantage to its remaining copies, which are contained in other organisms. Biologists began to approach this idea already in the 30s of the last century. Important contributions to understanding the evolution of altruism have been made by Ronald Fisher, John Haldane And William Hamilton.
The theory they developed is called the theory of kin selection. Its essence was 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.”
This is the formula. The “altruism gene” (more precisely, the allele promoting altruistic behavior) will be supported by selection and spread in the population if
RB > C,
where R is the degree of genetic relatedness between the donor and the “receiver” (in fact, relatedness 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 is the reproductive advantage received by the recipient of the altruistic act; C - reproductive damage caused by the “sacrificer” to himself. Reproductive gain or loss can be measured, for example, by the number of offspring left or not.
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, does not require special assumptions and does not even need experimental verification. It is purely logically deduced from the definitions of the quantities R, B, C and N - in the same way as geometric theorems are deduced from axioms. If NRB > C, the “altruism allele” will completely 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.
