EVOLUTION'S ARROW

The direction of evolution and the future of humanity

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(new) The most recent and refined version of the evolutionary worldview that was first presented in Evolutionís Arrow can be found in the 34 page document The Evolutionary Manifesto which is here

Chapter 4.     Barriers to Cooperation       

 

If cooperation is so good, why isn’t it universal? If cooperative organisations are better in purely evolutionary terms, if they are able to out-compete non-cooperators in evolutionary struggles, why are we surrounded by so many examples of animals that aren’t cooperating? Why haven’t the potential benefits of cooperation that we discussed in Chapter 3 already driven the evolution of cooperation amongst all living things?

We humans are obviously only partial cooperators. It is true that human economies can be spectacularly cooperative with an extraordinary specialisation and division of labour. It is also clear that internally, within our own bodies, we are cooperative all the way down. But anyone who tries to convince us that humans are always cooperators has a hard job in front of them.

The 20th century has seen millions killed in the largest wars in human history. Hundreds of millions of people in the world are chronically underfed, even though there are sufficient resources to feed us all. Millions die each year as a result[1]. Thousands of millions will live and die without having any chance to obtain the knowledge and skills needed to participate effectively in a modern economy. They will not share in the understanding of the world and of ourselves that has been made possible by the growth of human knowledge. They will live and die without fulfilling the potential for personal development and self-awareness that is possible at this time in the evolution of humanity. And individuals, corporations and countries continue to pollute and degrade our environment even though they know they are damaging the lives of others, now and into the future.

On a smaller scale, most of us do not feel that we are cooperators. There are strict limits to what we are prepared to give up for the common good. We know that tens of thousands die each year from starvation. But most of us will not sacrifice much to save any of them. We do not do for others what we would have them do for us.

We are very choosy about whom we will cooperate with: we are more comfortable cooperating with people who are likely to return any favours and to share the benefits of cooperation with us. So we cooperate more with family, friends and people we know we can trust. We are far more wary about cooperating with strangers. Before we do so, we want to be sure the strangers are trustworthy, or are put in a position where they will not take more than their fair share of the benefits of cooperation. For example, where the cooperation is a business venture we might insist on an enforceable contract, or that the investments and profits are not solely under the control of the stranger.

The other animals about us cooperate even less than we do. Like us, each of them is a cooperative organisation of smaller-scale living processes. But very few animals cooperate with each other. There is little cooperative organisation amongst the members of most species of earthworms, snails, crabs, prawns, spiders, insects, fish, frogs, snakes, lizards, birds or mammals. There is some cooperation between parents and offspring, but little beyond this. And in the few cases where cooperation is more extensive, it is nowhere as complex as the cooperative division of labour found within the bodies of individuals and within human society. Even ant societies have differentiated into at most four different types of ants.

If cooperation is so good, why is cooperation between animals so scarce and underdeveloped? There is no controversy amongst evolutionists about the answer. Cooperation does not evolve easily. There is a fundamental barrier to the evolution of cooperation. Why is this so?

In most circumstances, the only features that natural selection will produce in animals are those that benefit the individual. No matter how much a feature benefits the group or the species, it will not evolve unless it also benefits the individual. So if a feature causes an animal to help others without any benefit to itself, the feature will die out. And it will die out even if the benefits provided to others far outweigh the costs to the cooperator[2].

The reason for this is simple. The only way the gene that causes an individual to be cooperative can survive in a population is by reproducing successfully. For this to happen, the gene must cause individuals who carry it to have greater numbers of successful offspring than individuals that do not carry it. If the cooperator gene does this, the numbers of individuals in the population that carry the gene will increase, until eventually all will be cooperators. If it does not do this, non-cooperator genes will do better, and cooperator genes will die out. 

So what will happen to a cooperator gene that causes its carriers to use resources to help others without benefit to themselves? How will it do in competition with non-cooperators who use the same resources to help themselves rather than to help others? There is no doubt that the cooperator’s ability to survive and reproduce will be inferior to non-cooperators, and the cooperator gene will die out.

The more the cooperator sacrifices its own interests to help others, the worse it will do and the quicker it will die out. And cooperator genes will die out no matter how superior the advantages of cooperation. In fact, the more efficient and effective cooperation is at providing extra benefits to others, the worse the cooperator will do. This is because some of the others it helps are likely to be non-cooperators. And the more the cooperator helps its competitors, the worse it will do in comparison.

Evolutionary struggles tend to be won by genes that cause individuals to put their own interests ahead of the interests of others. The useful effects that an individual has on others will not improve its own competitive ability. Only the effects it has on itself will do this. This essentially is the ‘selfish gene’ perspective that has been argued so persuasively by writers such as Richard Dawkins[3] and George Williams[4].

They have shown that there is a fundamental barrier to the evolution of cooperation amongst animals that can be overcome only in limited circumstances. Even though cooperation can provide significant advantages, the genetic evolutionary mechanism is not very effective at exploiting them. The ‘selfish gene’ perspective can successfully explain why we see only limited cooperation amongst animals.

The conclusions of the ‘selfish gene’ perspective are an important part of my demonstration that the potential benefits of cooperation can drive progressive evolution indefinitely. If evolution could rapidly exploit all the benefits of cooperation amongst living processes, there would still have been a period of progress towards increasing cooperation, but it would have been short-lived. The potential benefits of cooperation would soon have been exhausted. The potential would not have been on-going in the way needed to drive indefinite progress. Paradoxically, the reason why cooperation continues to be so important in the evolution of life is that it does not evolve easily.

But as we discussed briefly in Chapter 3, the barrier also presents a direct threat to progressive evolution. Evolution will not progress if the barrier to the evolution of cooperation is insurmountable. This threat is even greater once we recognise that the barrier to the evolution of cooperation does not apply only to the animals we see around us. It is not just genetic evolution that is restricted by the barrier. The ‘selfish gene’ perspective can easily be extended to show that a similar barrier applies to the evolution of cooperation amongst all other living processes. It applies to cooperation amongst human organisations, individual humans, single cells, and molecular processes[5].

As is the case for the animals around us, evolutionary struggles amongst other living processes will be won by individuals that put their own interests first and foremost. Those that sacrifice their own interests for the interests of others will die out. The advantages of cooperation can be exploited only where evolution finds a way around this barrier. As we shall see in detail later, cooperation emerges only where evolution discovers how to build cooperative organisations out of self-interested components.

A number of examples will illustrate how the barrier to the evolution of cooperation applies to these other living processes.

Consider human corporations that sell their products in a highly competitive market. The corporations that are less efficient will tend to be out-competed, and will go out of business or be taken over. What will happen if the most efficient way corporations can make their products also pollutes the environment? Any corporation that cooperates with the community by reducing pollution will be less efficient and will go out of business. The corporations that pursue only their own interests will be more competitive, and will pollute. No matter how good the intentions of a corporation, it cannot stop polluting if it is to survive. And it does not matter whether it is run by responsible people who genuinely care about the environment. All they will achieve if they steer the corporation towards environmental responsibility is to send it broke.

The so-called “free rider problem” undermines cooperation in many human activities[6]. For example, it prevents businesses in an industry from cooperating together to train sufficient employees for the industry. If an industry is to be successful and to expand, enough workers must be trained in the general skills needed in that industry. However, businesses that make the investment needed to train employees can have their trained employees poached by other companies. This will often happen before the businesses have got a good return on their investment in training.

Free rider companies will rely on other businesses to train the skilled employees they need. Free riders will end up in front because they can get trained employees without paying the high costs of training. In contrast, companies that train can end up having paid for training without being able to hold onto trained employees. To remain competitive, more and more companies have to reduce their general training as much as they can, and join the free riders. Businesses that train for the good of the industry will be at a competitive disadvantage. As a result, the industry as a whole trains insufficient workers, and increasing numbers of businesses suffer shortages of skilled employees. And there is nothing any individual business can do about it if it is to remain competitive.

Free riding also undermines the ability of employees to band together to bargain with their employers for higher wages and better conditions. Any improvements won by the bargaining will apply to all employees. So free-riding employees will benefit even though they do not lose wages in strike action, and do not risk retaliation from their employer. Again, the result is that the free riders win out, and cooperation is undermined.

Free riding occurs wherever individuals, whether they are molecular processes, cells, animals, humans or organisations of humans, can get the benefits of cooperation without contributing to its costs. Free riders will always end up ahead of cooperators who use energy or resources by cooperating. Wherever free riding is possible, it undermines cooperation.

Cells provide further examples. Cells that put their own success first will out-compete cooperative cells that sacrifice their own interests. Consider a number of cells that are teamed up to form a group. To take full advantage of the benefits of cooperation, the cells must develop a cooperative division of labour, with different cells specialising in different functions. However, once a cell significantly changes its structure and function to become an efficient specialist, it will no longer be able to reproduce to form a new organisation of cells. If a cell is to start a new group of cells, it must be able to differentiate to form all the types of cells in the new group. A cell that is already highly differentiated will not be able to do this. Only a cell that is non-specialised can start a new group and produce all the specialised cells.

So cells that help the group by specialising give up their chance of contributing to future generations. They will be unable to found new groups. As a result, they will not appear in the next generation, and will die out. They will be out-competed by less specialised cells that contribute less to the group but retain the chance of establishing new groups. No matter how beneficial to the group, specialisation and differentiation will not evolve because it is not in the interests of individual cells. The great American evolutionary theorist Leo Buss has argued convincingly that a strong division of labour was not possible within multicellular organisms until this barrier to cooperation was overcome[7].

Another example involves the mitochondria, the organelles inside modern cells that release and convert energy for the use of the cell. It is now widely accepted that mitochondria are the descendants of free-living bacteria that took up residence within cells[8]. To a certain extent, the first mitochondrial bacteria to live inside cells had interests in common with their hosts: the bacteria could do useful things the cell could not, so they could make the cell more efficient. The cell provided food and a protected environment for the bacteria. And if the bacteria helped the cell to do better, the bacteria did better as well.

But the very close cooperative relationship that exists between mitochondria and modern cells has not been achieved by either sacrificing their interests to the other. In fact, there is clear evidence that mitochondria which do not put their interests before those of the cell will be out-competed by mitochondria that do. And this is true even if the self-interest actually damages the interests of the cell[9].

Consider the following example: in most multicellular organisms, an individual obtains all its mitochondria through the egg cell from its mother[10]. The sperm cell from the father usually does not contain any mitochondria. So, from the point of view of mitochondria, the production of male offspring is a waste of time and resources. The mitochondria that a female passes to a son will not be passed to his offspring. They will die with the son. In contrast, the more daughters that a female produces, the more mitochondria it will pass to the next generation. If a female puts all its energy into producing daughters, its mitochondria will do better, and will increase in numbers in the population.

Of course, it is not in the best interests of the organism itself to produce only daughters. Its genes will generally do better if it produces an equal number of sons and daughters. But the interests of the organism has not stopped mitochondria in a number of species from manipulating their host organism to produce a higher proportion of females. For example, in some hermaphrodite plants, the genes of the mitochondria operate to inhibit the development by the plant of male pollen, and instead enhance the production of the female seeds that pass mitochondria to the next generation[11]. These mitochondria will out-compete any mitochondria that cooperate with the plant and do not pursue their own interests at the expense of the plant.

Self-interest is also the best strategy for molecular processes that reproduce and compete. We will consider the case of autocatalytic sets formed of protein molecules. Such sets are thought to be very important in the origin of life because they are probably one of the simplest ways in which protein molecules can get themselves replicated[12].

The basic members of an autocatalytic set are protein molecules that are able to catalyse each other’s production. That is, they are able to speed up reactions that might not otherwise occur at a significant rate, and these reactions produce protein molecules that are also members of the set. Catalysts can do this by collecting together smaller molecules and holding them in positions in which they are likely to react. In this way they can organise reactions that would be extremely improbable if they occurred only by the chance meeting of the smaller molecules.

The set as a whole becomes autocatalytic and self-replicating when the formation of every member is catalysed by some other member of the set. Every member of the set gets reproduced by the catalytic activity of other members of the set. Once a set of protein catalysts links up in this way, it gains the ability to reproduce itself and its members indefinitely through time. This would be the case even though none of the members could survive indefinitely alone.

The particularly exciting thing about autocatalytic sets is that they form very easily. American biologist Stuart Kauffmann has shown that once you get a mixture of proteins that contain enough different types of proteins, it is likely that they will form a self-replicating autocatalytic set[13]. The chances are that the mixture will contain a set in which each component is catalysed by another component. These different proteins do not have to be specially selected. Even if they are chosen at random, if there are enough of them they are likely to include an autocatalytic set.

So once you get enough different proteins together, self-replication will arise easily. And the formation of these mixtures of protein should not be a rare and improbable event on a planet such as ours. On this basis, Kauffmann argues very persuasively that the fundamental step in the origin of life is easily taken, and is highly probable on this and other suitable planets.

Once a number of these sets form in a particular location, they will compete with each other for the resources needed to build the members of the set. The most competitive sets will win out. However, within the set itself, cooperative molecules that contribute most to the competitiveness of the set as a whole will not necessarily survive. They will be out-competed by molecules that are better at looking after their own interests, even where this damages the interests of the set as a whole[14].

For example, consider a cooperative catalyst that greatly improves the effectiveness of the set as a whole. It does this by catalysing the production of a molecule used in the construction of many other components of the set. But it will not survive in the set unless its own production is catalysed, no matter how useful it is to the set as a whole. And it will be out-competed by a similar molecule that uses its catalytic ability to further its own interests instead of the set’s. Self-interested molecules do not catalyse molecules that are widely used in the set, but of no use to themselves. Instead they catalyse the production of molecules that in turn catalyse the reproduction of the self-interested molecules themselves.

We can conclude confidently that the barrier to the evolution of cooperation applies to all living processes. The circumstances that cause it are universal. Individuals who use resources to help others without benefit to themselves will be out-competed. They will be disadvantaged compared with those who use the resources for their own benefit. And the barrier applies no matter what the evolutionary mechanisms that adapt and evolve individuals. In the examples we have considered, the barrier has applied whether the evolutionary mechanisms are those that adapt corporations, individual humans, other multicellular organisms, single cells, or autocatalytic sets.

This barrier makes it difficult for evolutionary mechanisms to take advantage of the potential benefits of cooperation. It makes it difficult, but not impossible. If the barrier completely prevented the evolution of cooperation, evolution would not progress. But as we shall see in the next three Chapters, the barrier is not insurmountable. Evolution can exploit the advantages of cooperation by finding ways to make cooperation pay for the individuals who cooperate. To evolve, cooperation must be organised so that it is in the interests of individuals.

So if evolution is to progress, it must meet this central challenge: it must discover ways of building cooperative organisations out of self-interested components—it must learn how to make it in the interests of individuals to cooperate.

As we have seen, evolution has had great success at this already. It has discovered how to build complex cooperative organisations of molecular processes and highly cooperative organisations of cells. Evolution on earth has been progressively producing cooperative organisations of wider and wider scale.

But so far evolution has had only limited success in building cooperative organisations of greater scale than multicellular organisms. To date, human society is as far as it has progressed. However, as we have seen, human organisation is nowhere near as spectacularly cooperative as the components of cells and as the cells within animals. The potential for beneficial cooperation amongst humans and other organisms on this planet is far from exhausted, and will continue to drive progressive evolution. We are only a step along this way. We are evolutionary work-in-progress. And as we shall see in detail, evolution’s next great challenge on earth is our challenge. To progress, evolution must discover ways to expand and improve cooperative organisation on this planet. Whether this occurs in the near future depends on us. The only evolutionary mechanisms on earth that are capable of discovering these improvements operate through our minds and our social systems.

In the next Chapter we will begin to look at how past evolution has found ways to overcome the barrier to the evolution of cooperation. We will see how evolution has repeatedly discovered ways to build cooperative organisations of living processes out of self-interested components. In later Chapters we will use this understanding to see how humanity can build larger-scale cooperative societies that can further exploit the advantages of cooperation to achieve future evolutionary success.

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[1].       Loftas, T. Ed. (1995) Dimensions of need: an atlas of food and agriculture. Rome, Italy: Food and Agriculture Organization of the United Nations.

[2].       Williams, G. C. (1966) Adaptation and Natural Selection. Princeton: Princeton University Press.

[3].       Dawkins, R. (1976) The Selfish Gene. New York: Oxford University Press.

[4].       Williams: Adaptation and natural selection. op. cit.

[5].       Maynard Smith, J., and Szathmary, E. (1995) The Major Transitions in Evolution. Oxford: W. H. Freeman; and Stewart, J. E. (1995) Metaevolution. Journal of Social and Evolutionary Systems 18: 113-147.

[6].       Olson, M. (1965) The Logic of Collective Action. Cambridge, Mass: Harvard University Press.

[7].       Buss, L. W. (1987) The Evolution of Individuality. Princeton: Princeton University Press.

[8].       Margulis, L. (1981) Symbiosis in cell evolution. San Francisco: W. H. Freeman.

[9].       Frank, S. A. (1997) Models of symbiosis. American Naturalist 150: S80-S99.

[10].     Eberhard, W. G. (1980) Evolutionary consequences of intracellular organelle competition. Quarterly Review of Biology 55: 231-249.

[11].     Hanson, M. R. (1991) Plant mitochondrial mutations and male sterility. Annual Review of Genetics 25: 461-486.

[12].     Farmer, J. D., Kauffman, S. A., and N. H. Packard (1986) Autocatalytic replication of polymers. Physica D, 22: 50-67.

[13].     Kauffman, S. A. (1993) The Origins of Order: Self-organisation and selection in evolution. New York: Oxford University Press; and Kauffman, S. A. (1995) At home in the universe: The search for laws of self-organisation and complexity. New York: Oxford University Press.

[14].     Bagley, R. J. and Farmer, J. D., (1991) Spontaneous Emergence of a Metabolism. In: Artificial Life II. (Langton, C. et al, eds.) New York: Addison and Wesley; and Maynard Smith, J. (1979) Hypercycles and the origin of life. Nature, Lond. 280: 445-446.

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