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    Inclusive Fitness Is Group Selection
    By Gerhard Adam | October 14th 2012 08:00 AM | 36 comments | Print | E-mail | Track Comments
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    Despite the intent in biology to eliminate group selection, it invariably turns up as the only reasonable explanation for the cohesion of species and the behavior of large groups of animals.

    Additionally, the struggle to explain altruism using kin selection and inclusive fitness is perpetually haunted by the requirement that the entire premise hinges on the existence of an actual "altruism" gene.  In other words, Hamilton's rule and inclusive fitness are meaningless if there isn't a genetic component to cooperation and altruism.

    In that respect, can we be reasonably confident that genes are involved?  If we simply consider how cooperation works and examine it purely from a behavioral [not cognitive] approach, we can see cooperation/altruism (1) occurs in organisms that don't even possess a nervous system.  Therefore concepts like cooperation and altruism, are not created by the existence of cognitive processes.  However, this fails to address whether such behavior is actually genetic.

    To approach this problem we need to determine whether cooperation/altruism is a trait that is actually selectable.  If so, what is the heritability of such cooperation/altruism?  

    Even the cells of a multi-celled organism require a high level of cooperation and altruism to form the representative creature.  As such, one must postulate a means by which such a characteristic is passed on from generation to generation.  One can also easily imagine that as cooperation becomes a staple of the group, then those individuals that lack such a trait or characteristic will tend to either be expelled or attempt to compete independently.  Given the lower total energy costs of cooperation, it follows that the advantage cooperation provides ensures that cooperative groups tend to thrive.

    This certainly wouldn't be true for all organisms, since the total energy costs of fitness would vary, so that there might be many creatures for which there is no intrinsic advantage to belonging to a group.  Similarly, other adaptations turn fitness into a pure numbers game, where by simply creating tremendously large numbers of viable offspring, sheer probability assures that some number will likely survive.

    In these cases, then cooperation may actually be a liability and consequently one wouldn't expect to see it persist in a population.

    However, what is more interesting is that the arguments in favor of inclusive fitness require groups.  It makes no sense to talk of altruism among different animal species, just as it makes no sense to consider cooperation as evolutionarily stable between separate species.

    That this may occur isn't relevant, since it conveys neither advantage nor disadvantage on the practitioner unless such encounters produce a conflict of interest [which may often occur when altruistic or cooperative behavior changes due to one of the creatures reaching sexual maturity].

    In effect, the gene-centric view in biology was a direct attempt to stop the persistence of group selection theories that suggested that evolution occurred "for the good of the group".  Specifically, no one could offer an explanation for how traits that were carried by individuals could possibly translate into group-level selection, so the idea was dismissed as mere wishful thinking.

    Yet, we find that on examination, inclusive fitness is precisely the theory necessary to explain group selection.  While many may balk at such a conclusion, one must remember that the essence of the altruistic explanation is that the individual does NOT benefit.  Therefore the relevant question is to ask who does benefit?  Previously the simple answer was; the genes.  Yet, this is unsatisfactory since it is obvious that 23,000 + genes are not actively competing for propagation.  In addition, we know that survival of any species is enhanced with greater genetic diversity, not less.  Therefore the problem that evolution "solved" was how to maintain a faithful enough transfer of information to ensure sufficient uniformity so that reproduction could be maintained, but not so uniform as to risk the liabilities of amplifying negative traits.

    Certainly the argument has been made that blood-related kin can benefit by passing on such genes, in kin selection theory.  Yet, this is inadequate because overwhelming almost no species is capable of recognizing kin (2), and other than the eusocial insects, acts of altruism and cooperation still occur with regularity among unrelated individuals.  In addition, many offspring have no siblings [related kin] so this fails to explain why social behavior should have developed.

    Inclusive fitness extends the arguments of kin selection, by claiming that individuals within the same group will then benefit by such behaviors.  Yet, isn't this precisely what group selection claims?

    Doesn't this render the altruistic act of an unrelated individual being made precisely for "the good of the group"?  Moreover, it is well known that without a group [sufficient number of breeding pairs] extinction is all but inevitable.  

    Part of the problem is that Hamilton's equation is often tossed around as "evidence" of inclusive fitness explaining how an altruistic gene can persist in a population (3).

    r * b > c

    Where r (relatedness) times b (benefit) is greater than the c (cost).  So, the benefits must be greater than the cost where b is adjusted by the probability (r) of the relative carrying the genes [i.e. the degree of relatedness].

    Yet, on examination, this begs the question, since there is NO instance of where there can ever be a benefit greater than simply not making the sacrifice.  In other words, if the altruist doesn't perform the act, then their probability of passing on their traits is 100%, compared to some lesser probability if a relative were to reproduce.

    Using Haldane's famous quote about saving two drowning brothers, the statement is problematic.  In the first place, both brothers would need to be drowning to balance the equation, which is already somewhat more unlikely.  However, there is no argument that can be made for saving the brothers that doesn't produce a greater benefit to the initial actor than simply standing on the shore.  In other words, the altruistic act invariably falls short of propagating the desired genes regardless of the degree of relatedness.

    More importantly, one doesn't observe altruistic behavior that fits into Haldane's quote.  An altruist will likely risk sacrificing themselves for considerably less lofty results, so the postulate granting such a gene becoming pervasive based only on individual genetic success is unsatisfactory.

    The only way to make sense of this situation is if the benefit [fitness benefit] is assigned to the group rather than the individual.  In that situation, if the group survives, then it will provide benefit for considerably more members than the singular act addresses, and therefore any altruist's cost will always make sense within that context. 

    On the other hand, if the cost is zero, then ALL actions are justified since any action, regardless of how small, is beneficial.  This is also an argument that makes sense if one views the cost as merely a risk-taking venture, where the fitness cost is not assured, but merely a probability.  In those cases, then any success will automatically produce a benefit with the cost being zero.

    What makes this important is that inclusive fitness, as it is currently defined, does not adequately account for those situations in which it is not beneficial to incur a cost.  Despite popular belief, it is often better for a parent to sacrifice their children than to risk themselves.  Since the parent can produce more children, but the children may not survive without the parent, to behave altruistically places both at risk, and has a slim probability of success.  Therefore, Hamilton's Rule is necessarily restricted to those instances of where there is no dependency between the individuals if one is not capable of reproducing.

    As a result, this also turns the altruism attributed to eusocial insects on its head, since there is no benefit in an insect sacrificing itself for another sterile insect.  Therefore the only context that makes sense is group selection.  The sacrifice is beneficial because it conveys benefit to the group.

    In fact, even if we were to argue from the gene-centric position that somehow the objective of the gene is to be propagated into future generations, there is little benefit in a gene supporting an individual versus supporting the group.  There is far greater benefit (to the gene) to be conserved within the group, rather than struggling to be passed on to a single individual. 

    If anything, the gene-centric view, might offer some insight into how novel genes are introduced into a population, but it contributes nothing to the notion of conserved genes already present within a population.  After all, if they are already pervasive throughout the population, then what's the point in evolving altruism, since every member of the group already carries the gene?

    Instead, we have to consider that this explanation is insufficient for anything except genes that haven't gone to fixation within a population. 

    In short, the most likely explanation is a combination of all these ideas, to varying degrees, so that at individual levels we find cooperation producing economies of scale that render it beneficial, coupled with altruistic behaviors that promote the survival of the group.  Once group survival is assured, then competition between individuals within groups becomes possible.   In many cases, the development of cooperate strategies can "bypass" phenotypic developments by providing a group level benefit that is beyond the individuals themselves.   It is clear that the gene is insufficient to account for all the traits and criteria for successful survival of individuals, therefore one must consider that the solution exists within the myriad number of groups that constitute life.

    The greater the benefit of cooperation, the greater the dependency of individuals to group survival.  The individual cells of a multi-celled organism have completely become committed to "group" survival, while other organisms may experience varying degrees of such a dependency.  So, while the gene is certainly a crucial element for conveying information into the next generation at the individual level, it is not the only or necessarily the most significant factor shaping the next generation in an evolving population.

    Additional reading:
    http://www.world-science.net/exclusives/060529_altruism.htm
    http://plato.stanford.edu/entries/altruism-biological/#2
    http://www.genetics.org/content/176/3/1375.full

    ======================================
    (1) Admittedly one of the difficulties occurs in defining "altruism" itself.  The conventional definition is "any act that increases the fitness of an organism while decreasing the fitness of the actor".  A portion of this post is to refute whether such a definition is actually accurate.

    (2) Interestingly enough, there is some suggestion that while organisms may not recognize kin, their microbiota may be capable of such recognition, or at least recognition of similar microbes.

    (3) In a previous article I noted that altruism didn't appear likely to be related specifically to fitness in humans anyway. 

    However, there is a more fundamental problem with the idea of an altruistic gene.  How does an altruist become the beneficiary of an altruist?  The concept is contradictory, since the nature of the general view of altruism is that all members are self-sacrificing.  Therefore, the logic conclusion of a population filled with altruists, is a suicidal group.

    It is for this reason that I argue that altruism is a risk-taking behavior with the beneficiary being the individual, but primarily the group at large.  This ensures that the "altruist" is not merely a self-sacrificing individual [since such individuals are destructive to the group], but rather someone that can take a risk versus a huge benefit.

    Comments

    I agree that the gene-centered argument is not the whole story. That leaves me with a lack of understanding of how a multi-celled creature, say a mammal, organizes itself to be altruistic. (This supposes that one has a firm definition of the term in mind.)

    There are so many different cell types contained in a mammal that it boggles my mind trying to see how these cells communicate the need to cooperate. Do they communicate, or have they done so in the past? It seems unlikely that there is direct communication as we know it, but there must be some mechanism for the cooperation. Do you have any thoughts along this line?

    Great article, BTW.

    Gerhard Adam
    Cells receive many signals and communicate with each other for a variety of purposes, not the least of which is when they have to do something.  This communication is purely through chemical means, but it represents a quite sophisticated network of information and responses.

    http://www.biotechprimerblog.com/2010/08/how-do-cells-communicate.html

    Actually Wikipedia also has some good examples
    http://en.wikipedia.org/wiki/Cell_signaling

    Of course, cell communication isn't just confined to multi-celled organisms, although in those cases then there clearly has to be a more well-defined set of functions and roles.  I'm sure you've heard of quorum sensing in bacteria, which is also a form of communication, so we recognize that all cells are capable of communicating with each other.

    Personally I think you could also argue that even viruses are capable of communicating, since they also must recognize and respond to the chemical signals employed in the cell, but also that they have the means to enter a cell, hijack the reproductive machinery, subvert the DNA to their own purposes and replicate themselves. 
    Mundus vult decipi
    Yes, I have heard of quorum sensing. I hadn't made the communication connection though.

    Thanks for the links. I've some study to do.

    Gerhard Adam
    That leaves me with a lack of understanding of how a multi-celled creature, say a mammal, organizes itself to be altruistic. (This supposes that one has a firm definition of the term in mind.)
    I certainly agree that the definition is often an issue.  But if we consider that altruism exists at all levels, then it is something that can be intrinsic in individual cells, as readily as it can be in a large multi-celled organism.  However, the question is ... "what" is being altruistic?

    In the case of individual cells, apoptosis [Programmed Cell Death] is a specific mechanism where the cell essentially "commits suicide", in response to some directive.  Is this altruistic or merely self-sacrifice?  In my view, there is a difference.

    Part of the difference is that apoptosis occurs when a cell has no purpose or can no longer fulfill its obligations.  For example a nerve cell that fails to find a point to connect to will be directed to "kill itself".  So, it's not purely altruistic in the sense that it is a viable entity that has sacrificed itself so that another may survive.  These cells are all in the same boat, since their survival is linked to the survival of the organism they represent.

    In the case of the eusocial insects the picture changes a little, but ultimately the entity being served is the nest.  As I mentioned in the article, since none of the ants are viable by themselves, then sacrificing themselves for each other makes no sense.  So, again, this is more of a set of actions where the benefit is directed to the group.  In that respect, eusocial insects can be treated as a distributed type of "multi-celled" organism where each ant represents a kind of cell or system.

    In larger animals, we also have to consider other means of information transfer; behavioral, cultural, etc. that can have an influence on how an organism reacts.  So that social animals will have some portion of their behaviors determined by what they are taught and the demands of the environment they live in [environment, in this case could include the group].

    So, as you can see, it's hard to define exactly what the "altruistic" entity is and who the beneficiary is. 

    Personally, I think the focus on altruism as a singular trait is ultimately wrong.  It treats it as if it were some unique behavior that isn't connected to anything else, but this doesn't really explain much and it makes no sense.  There has to be some evolutionary benefit to such a trait, which is why I argue that it is for support of the group.  Yet, why should the group be important?  The group is important because it represents an economy of scale, so that less energy is expended by each individual to survival, thereby enhancing the overall fitness of the individuals in such a group.  At this point, the group becomes the means by which individuals succeed.

    It seems that in such a scenario, that risky-taking behaviors that may benefit more individuals can be selected for, without turning everyone into a self-sacrificing maniac.  Too much of a risk taker; the odds are your fitness will be impacted.  Judicious risk-taking can be beneficial, because the individual is more prudent, thereby ensuring their own survival as well as enhancing that of the group.

    Anyway ... I think I'm starting to ramble, but you get the idea.
    Mundus vult decipi
    Human altruism is a different critter. Here my definition of altruism differs from that which could be assigned to insects or flora of any type. Humans regularly give goods or services to other humans and animals, sometimes separated by thousands of miles, with no expectation of benefit. No benefit of either survival or reproduction.

    The costs of giving is usually small, but the expectations would seem to be non-existent. I don't think this is a case of heritage either. In some cases one sibling will give and another could care less. It seems like a cognitive process were one recognizes a need and does what is needed.

    Gerhard Adam
    Yes, but how would you separate these acts between altruistic versus merely cooperative?  In humans it becomes easier to consider by asking what motivates someone to be a fireman, police officer?  How about those fighting in the military?

    These are clearly altruistic acts, where fitness can be impacted.  However, I think you would agree that none of these individuals is specifically "self-sacrificing".
    Mundus vult decipi
    Way too many degrees of freedom here!

    I'm not trying to be argumentative, just trying to understand my, and your, interpretations. The various definitions of altruism leave many aspects of the term up for grabs.

    Agreed that the fireman, police officers, and military members do have altruistic tendencies. However, they also expect to gain from their employment.

    In the case of someone donating $50 to the Haiti relief agency, there is no expectation of gain for the individual who gave. In many (most?) cases the donating person can't be sure that the altruistic act of giving actually impacted the person or group that the giver intended.

    There is a certain satisfaction in the, supposedly, selfless act of giving that could be interpreted as an expected gain. And, that may be the answer. It's all in our head.

    Which brings full circle. How does human altruism originate, and how is it propagated?

    Gerhard Adam
    I don't interpret your comments as argumentative, so feel free to comment however you see fit.
    The various definitions of altruism leave many aspects of the term up for grabs.
    I certainly agree, which is why I've tried to define within the context of taking a risk without presuming self-sacrifice.  In other words, the typical biological definition requires that someone win and someone lose.  I'm arguing that the "loser" doesn't start out with that intent, so it becomes a calculated risk, rather than simply a sacrifice.
    Agreed that the fireman, police officers, and military members do have altruistic tendencies. However, they also expect to gain from their employment.
    Yes, this is often mentioned as a counter-argument, but there are certainly many more ways to gain employment, and certainly better advantages in other employment.  Certainly the fact that it is employment conveys some benefit back, but it is hardly sufficient compared to the risk involved.   Again, it would be different if just anyone off the street could do that job, and especially in the case of the military, there may be additional motives that can be exploited to advance themselves in life later [i.e. education, etc.].  However, none of this is enticing enough to motivate the majority of people to pursue these careers, so I'm not inclined to assign much motivation to the economic incentives.  In short, I don't believe the financial benefits are sufficient to induce altruism.
    In the case of someone donating $50 to the Haiti relief agency, there is no expectation of gain for the individual who gave. In many (most?) cases the donating person can't be sure that the altruistic act of giving actually impacted the person or group that the giver intended.
    Well, there may be a gain in terms of respect, reputation, or simply self-satisfaction of having contributed.  We don't need to be sure that it impacted the person or group, we simply have to believe that it does.  I'm sure you would agree that if someone made a donation and found that the funds were being funneled to some other purpose, they would be furious, because they would feel that their "altruism" had been taken advantage of.
    There is a certain satisfaction in the, supposedly, selfless act of giving that could be interpreted as an expected gain. And, that may be the answer. It's all in our head.
    To a lesser extent, some of that may be true, but in human society we tend to assess people on the values they hold and display.  So, unless you're being extremely secretive about it, such acts aren't merely self-satisfactory, but they build on a reputation for the individual within the group.  This can effectively become another means of competing within the group for status.  It doesn't make it a selfish act, nor does that even have to be the motivation, although most people would feel that their reputation and how others perceive them is an extremely important attribute of living in human society. 

    I also think too much is made of "selflessness" as if the only alternative is a selfish self-centered attitude.  Self-interest is not the same as selfishness.  I've used the example before, but if I make myself a sandwich because I'm hungry, I'm not being selfish by not making you one.  I'm merely pursuing my own hunger.  If I were to actively deny you food despite your also being hungry, then that would be selfish.  In other words, it requires energy be expended in order to be selfish.  One has to make an effort to deny others.

    If we were two castaways stuck on an island, we wouldn't even have to like each other to recognize the benefits of cooperating to acquire food, build shelter, etc.  As a result, we would tend to help each other out, with the expectation that we might need a favor in return.  So, by building on such a cooperative effort [with our respective self-interest] we are achieving a cooperative society.  This doesn't have to be a "My Little Pony" scenario.  It simply has to work to keep the peace and ensure that effort isn't being expended against the other party, when we can gain an economy of scale by cooperating.

    All of us recognize our participation in such activities, and we don't think much of it, until the time comes when we need a favor or help and the other individual refuses.  It is at that point that we recognize that the cooperative "agreement" has been compromised.  Again, we judge such an individual on those actions and it will have ramifications on their reputation in the group.  In a sufficiently large group, such individuals become outcasts.
    Mundus vult decipi
    As is usually the case, I've increased my understanding of small segments of the larger question while discovering that deeper questions remain.

    I'll have to internalize all of this so that I can assess the deeper areas. Then, perhaps, I can ask a more intelligent question. Once again, thanks for your efforts.

    Gerhard Adam
    Frank, I always enjoy hearing from you.
    Mundus vult decipi
    vongehr
    if the altruist doesn't perform the act, then their probability of passing on their traits is 100%, ... both brothers would need to be drowning to balance the equation
    Though I agree with a lot in this article, it seems that either there are assumptions you do not tell us about or your probability calculus is off. Intuitions about statistics are usually wrong, and inclusive fitness is backed up by numerical simulations that focus on genes. The need to have both brothers drown is a limit case in parameter space. The equations only bear out if looking at the expectation values of returns over several generations and in competition with different genes that try to invade the gene pool.
    The only way to make sense of this situation is if the benefit [fitness benefit] is assigned to the group rather than the individual.
    Or to genetic traits' predominance in the gene pool. Gene centrism avoids mysticism about the mechanisms. If group selection equals inclusive fitness, I go with the term "inclusive fitness". Group selection probably combines several such mechanisms.
    eusocial insects on its head, since there is no benefit in an insect sacrificing itself for another sterile insect.
    Sounds good, but questionable once you look at the details of how similar the genes of the queen and the sisters are and do the math. Time for numerical simulation.
    Steve Davis
    "...genes that try to invade the gene pool."
    Followed by "Gene centrism avoids mysticism about mechanisms."
    The first sentence disproves the second.
    Gene centrism is all about mysticism, as you demonstrated.
    vongehr
    You have lately nothing left but ultra short quibs that illustrate your ignorance. "try to" is the usual convenient shortcut for what otherwise would be a very long sentence. Everybody who has just a basic introduction to simple biological evolution knows that it does not mean that genes "try" or "want" anything. Fine, I should have obviously formulated differently here, because the "try to" is not even necessary; I am sorry for having let exactness slip here for a second. But your turning this into a claim of my support for mystical mechanisms is just silly. You are simply still pissed off about that our argument way back made people abandon your own esoteric views on "life" that refuse to properly define terms.
    Steve Davis
    Oh my, we are prickly today!
    "try to" is the usual convenient shortcut for what otherwise would be a very long sentence.
    So enlighten us; give us the very long sentence.
    But your turning this into a claim of my support for mystical mechanisms is just silly.
    But I did not claim that, did I?
    I said gene-centrism is all about mysticism, and you demonstrated that by your statement. You were sucked in. Alpha Meme? More like meme victim.
    vongehr
    Not biting Steve. Toss the troll bait someplace else. You know what I mean, or if not, you refuse to. You have read more about genes than I ever will. If you mean to say something useful, perhaps that in your opinion/experience, gene based descriptions and related shortcuts like "the gene/animal/system wants to ..." are doing more harm than good, just say so and best provide alternatives. Else, I do no longer "discuss" with people who simply refuse to know or pretend to not know to save face.

    Numerical simulations based on gene models have shown that cooperation can be an evolutionary stable strategy. Read: No mystic ingredients. If you can do better and protect the population against invading defectors via other mechanisms, do it. Anybody can do such simulations nowadays - your home computer is plenty fast enough. Good luck.
    Steve Davis
    "Toss the troll bait someplace else."
    Now, why would I want to keep fishing when I just got the catch of the day!
    This is not the first time you've presented gene-centrism as a concept with scientific credibility. One mistake can be overlooked, two indicates a problem.
    Gene-centrism is the science world's version of a cult.
    It's a scientific cult because its proponents tolerate no dissent. (Just ask E O Wilson or Mary Midgley.)
    The name itself should have told you it's a cult.
    Genes are the centre of what, exactly?
    The universe? They wish!
    Of evolution?
    Don't make me laugh.
    You've been sucked in by a meme.
    There's nothing in biology that we can claim to be central, except life itself. And you're so unsure of yourself there that you have told us, from memory, "everyone can define it as they like as long as they are explicit"! What a cop-out.


     
    Gerhard Adam
    I would agree about how the statistics works out, but that also depends on how accurately the model reflects actual behavior.  Part of my point is that it isn't just a simple numerical simulation, but also depends on the genes being able to influence the behavior to achieve this objective/result.  I'm not assigning "intent" to the genes, but nevertheless some process must recognize the "benefit" of such behavior.

    From this, I have to question why an altruistic individual would ever sacrifice themselves, if the logic is that doing so will preserve the altruistic genes in their brothers.  In the first place, such a guarantee isn't possible [if it were it would render the act meaningless since the gene is already pervasive].  Therefore, there is no situation of where rescuing a brother is more advantageous that merely preserving yourself [where the gene is assured].  However, such an assurance also means that the gene will not have fulfilled it's objective [of being altruistic].

    Moreover, how does an altruistic drowning brother justify another altruistic brother risking his life to save him?  It appears to be a circular argument.  This becomes especially problematic if they both drown.
    Sounds good, but questionable once you look at the details of how similar the genes of the queen and the sisters are and do the math. Time for numerical simulation.
    Again, I don't disagree, but the question remains that even in this situation the queen isn't actually viable any longer;  only the nest is.  So, the only way the genetic sacrifice is useful is if the entire existing nest is preserved giving rise to the possibility of future queens migrating to form other nests.  If all the ants merely sacrificed themselves for the existing queen, the entire nest dies. 

    It's no different than the behavior of individual cells in your body.  They are already committed.  There is no separate future for them, regardless of what they do.  Their only option is to preserve the body [i.e. group] in the hopes that some of the germ cell line can propagate. 
    Mundus vult decipi
    vongehr
    I cannot follow you.
    it isn't just a simple numerical simulation, but also depends on the genes being able to influence the behavior
    Is this going back to anti-racism that denies behavior being influenced by genes? Again, if we deny gradualism leading up to speciation, we may be nice humanists, but we effectively deny evolution! Behavior is influenced by genes, otherwise different behaviors cannot evolve. Other primates have no schools that you can blame for teaching anything that makes the young already attack females in order to prepare later rapes. Once you admit that genes are involved, you start the maths with b and c parameters to model gene pools, and the models work rather well, even predictive.
    In the first place, such a guarantee isn't possible
    There is never guarantees; there are probabilities.
    how does an altruistic drowning brother justify another altruistic brother risking his life to save him?
    There are no justifications. Lower animals react. They either do jump in to help or they don't. If they both drown, c is high. If the brother helping gets them both out, c is low. Do the population maths, that is all.
    Gerhard Adam
    If they both drown, c is high. If the brother helping gets them both out, c is low. Do the population maths, that is all.
    ... and that's the problem I have with it.  C isn't just high, it is 100%.  Similarly if the both survive, then c = 0.  However if c = 0 then the act is not considered altruistic, since [by definition] an altruistic act must confer a fitness cost to the actor.  In my view this is a rigged "game" by only examining or allowing those cases where there is a cost, then the equations are skewed.

    This is why I proposed that altruism is actually a risk-taking behavior where the benefit [i.e. both surviving] far outweighs any other consideration because the cost could be zero.  Otherwise it seems that the entire concept is skewed towards self-sacrifice, which doesn't make any sense to me.

    Even more important, is that brother saving the drowning brother doesn't actually confer a fitness benefit either, other than ensuring that his brother survives with the possibility of reproducing.  If the brother is already older and isn't going to have more children, then any benefit would be zero. 

    So, by definition, if someone isn't going to have any more children, there is no fitness cost to anything they do, which should render older adults 100% altruistic.  That's what would be predicted.
    Mundus vult decipi
    Thor Russell
    "So, by definition, if someone isn't going to have any more children, there is no fitness cost to anything they do, which should render older adults 100% altruistic.  That's what would be predicted."Not sure what you mean by altruistic in the case. If they are not going to have any more children, then it would make sense for them to invest all their efforts into the survival of their existing children/grandchildren/related offspring etc.  The only question is how. Saving a brother would still have a fitness benefit if that brother is looking after your children. Also helping your brother look after his children also advances some of your genes even if that brother is not going to reproduce either. Not saving someone you have only a very low chance to save thus preserving yourself for later would also be expected.


    Do you accept that if you are to look at the gene pool several generations hence, saving several of your brothers children could be very similar to saving one of your own, (in terms of what genes of yours would be around in it) and if so then why distinguish the two situations from an evolutionary gene pool/group makeup perspective. I don't see why the benefit has to far outweigh the risk. It could make sense for someone to take a 5-10% chance on their life to save their brother both where their brother was looking after your children or where you were looking after your brothers.
    Thor Russell
    Gerhard Adam
    I don't disagree with what you're saying which is precisely why I object to the definition of altruism as it is stated.
    In evolutionary biology, an organism is said to behave altruistically when its behaviour benefits other organisms, at a cost to itself. The costs and benefits are measured in terms of reproductive fitness, or expected number of offspring. So by behaving altruistically, an organism reduces the number of offspring it is likely to produce itself, but boosts the number that other organisms are likely to produce. This biological notion of altruism is not identical to the everyday concept.
    http://plato.stanford.edu/entries/altruism-biological/
    Altruistic behaviour involves a ‘loss’, ‘cost’, ‘risk’ or ‘disadvantage’ to the personal fitness of the altruist, as measured by its production of offspring, while conferring a ‘gain’, ‘benefit’, or ‘advantage’ to one or more other individuals in the population. An altruist gains no direct benefit from its actions. It is implicit in Hamilton’s treatment that the altruist has lower personal fitness as a result of its altruistic behaviour than if it had not behaved altruistically.
    http://www.gnxp.com/wp/2011/03/01/defining-biological-altruism/
    ... and you could find many more.

    I'm proposing that it is a risk-taking behavior, in which case your point about the probability of success or failure applies.  However, you can see from these definitions, there is no probability of a cost, it is defined as REQUIRING a cost.  More explicitly that cost must translate directly into fitness.
     I don't see why the benefit has to far outweigh the risk.
    Not risk; cost.  It does so because that's what Hamilton's rule demands  r * B > C, where r is the relatedness, B = benefit [in terms of fitness], and C = cost [in terms of fitness].  So the only probability in this relationship is the probability associated with the relatedness.

    You can also see by this definition, that it is very difficulty to argue about how personal fitness would be reduced [especially among humans] short of death.  Again, this is why I'm arguing that it makes little sense, because that requires self-sacrificing behavior, and [in my view] most such behaviors do not intend to sacrifice themselves, but rather are willing to incur the risk [whether it be high or low probability of success].  In such a situation, the benefits might be huge while the costs may be negligible or zero.

    Again, in my opinion, that fits better with what we actually see behaviorally.  Even some of the common examples, such as bird issuing a warning cry, presumes that the bird intends to sacrifice itself, rather than simply arguing that the bird may well undertake the risk without any intent of dying.  Success helps more than one individual.  Failure is what incurs the cost, but with the act the costs may be minimized instead of distributed over a larger portion of the population.


    Mundus vult decipi
    Thor Russell
    OK I see your point. By the first definition even a mother stopping at 10 offspring could be considered altruistic. By not having the 11th child they are reducing the number of offspring they have but increasing the number of offspring their existing children will have by looking after them more. The definition already assumes that fitness is defined by direct offspring in the immediate generation rather than genes represented in several generations hence. If you take the second definition of fitness, then altruism would have to mean more like saving a completely unrelated stranger. If you include the expectation that the strange will do you a favor in the future, then altruism needs to mean saving a completely unrelated stranger who will definitely not return the favor.
    Thor Russell
    Gerhard Adam
    Yes, exactly my point, which is why I'm arguing that altruism makes sense if we consider it a risk-taking behavior whose beneficiary is the "group" since that increases the genetic diversity from which mates can be selected, and by ensuring the viability of the group, then "everyone" benefits [even if just indirectly].  In other words, if more individuals can benefit that it becomes easier to see how a large benefit could accrue from a relatively small [i.e. individual] sacrifice.  Of course, if no cost is actually incurred, then the entire venture is a success for everyone.
    Mundus vult decipi
    vongehr
    Why would the probability of drowning be 100%?
    Similarly if the both survive, then c = 0.
    If I fish my brother out of the water, I get a cold with P=99%, that I die from that cold: ~0.1%. You verify the overall model qualitatively [can cooperation arise at all in the (b, c) parameter space, are the predicted population oscillations between three mating strategies of lizards observed in nature?]. The quantitative values of b and c is what a working model then tells you if you put in empirical data (say minimum lizard ratios). c is not zero! And yes, c or IQ is defined/determined via the model/measurement combination, scientific knowledge is thus constructed - scientists who understand how science works know that this is precisely how it must be. Talk about what is "true life" or "the real cost" is nonsense.
    This is why I proposed that altruism is actually a risk-taking behavior where the benefit [i.e. both surviving] far outweighs any other consideration because the cost could be zero.
    So in fact your suggestion explicitly hangs on your intuition about probability calculus (must far outweight). Sorry to be harsh, but human intuition about probability is worth crap. Read up on Simpson paradox and base rate errors. That humans suck at probability is the reason they come up with ID. There is little alternative to actually doing the math.
    Even more important, ... benefit would be zero ... older adults 100% altruistic.  That's what would be predicted.
    Nope. You need to do the math. ;-)
    And complex systems are nonlinear. You change (b_i, c_i) just a tiny little near a threshold that you do not know about, and all of a sudden, bang, the systems behave counterintuitively. Some people call such arguments "mystical", and they would indeed be mere handwaving if one does not actually do the math. But that math has been done in evolutionary game theory.
    Gerhard Adam
    So in fact your suggestion explicitly hangs on your intuition about probability calculus (must far outweight).
    No, not at all.  Hamilton's relationship is quite explicit in defining the cost and benefits in terms of fitness [i.e. number of offspring].  Therefore if any act of risky behavior succeeds, such that the individual survives without any impact on their fitness [which is highly likely if they survive] then the cost is zero.

    This is why I indicated that older adults that will no longer have children have zero cost investment in the process.  Their fitness will not be altered by any act.  They've already had the maximum number of offspring they intended [or are capable of], so no matter what they do, they cannot incur a fitness cost. 

    I will check out your links, but the fuzzy part of Hamilton's rule is that the explicit assumption is that all creatures seek to maximize their fitness [number of offspring], yet there is no way to know what the maximum fitness is except in hindsight.  Death is an obvious limiting factor, but anything short of death that doesn't impact fitness would be a zero cost, and consequently not considered altruistic.  So we are stuck with the dilemma that the same identical act, in which the actor survives isn't counted, while that in which they die is considered altruistic and Hamilton's rule applied.  It seems that we are arbitrarily defining conditions so that the results are pre-determined rather than allowing for actual prediction based on real behaviors.  We simply don't allow success.
    Mundus vult decipi
    vongehr
    Hamilton's relationship is quite explicit in defining the cost and benefits in terms of fitness [i.e. number of offspring].  Therefore if any act of risky behavior succeeds, such that the individual survives without any impact on their fitness [which is highly likely if they survive] then the cost is zero.
    This is why I indicated that older adults that will no longer have children have zero cost
    I should be careful, since it could of course be correct that somehow, in certain popular literature for example (and I don't read books), this is how it is introduced, and if so, I agree it may be no good. The parameters should certainly not be like "number of possible offspring of a sterile dead man = 0", but expectation values along the lines of "average boost to the preponderance of shared genes in the gene pool". If the sterile old man drowns trying to fish out his younger brother, he can no longer help his grand kids with the homework, so c could be larger than 0 (although there will be particular instances where the grand kids do therefore not get a job and father twenty children raping around out of boredom). I suspect there are b and c as introduced with (too) simple examples in certain texts, and then there are b and c parameters in models.

    BTW, if you write
    This is why I indicated that older adults that will no longer have children have zero cost investment in the process.  Their fitness will not be altered by any act.  They've already had the maximum number of offspring they intended [or are capable of], so no matter what they do, they cannot incur a fitness cost.
    this seems to propose extremely high level evolved behavior, like that an old man once he is sterile, all of a sudden completely changes his behavior. The genes make the guy more or less caring for people who look and smell like him all along, and that this is due to genes is already doubted by many "all is nurture" freaks.

    Gerhard Adam
    ...like that an old man once he is sterile, all of a sudden completely changes his behavior.
    Which is exactly my point.  We don't such a stark change in behavior.  Instead we find that risky behavior would be fairly uniform around any particular individual's behavior.  Perhaps they're more reckless when they are young, but overall we don't see such a marked change in behavior simply because the "gene" recognizes that fitness is no longer an issue.

    So, in my opinion, if a gene can't tell, then it is hard to argue that it is responsible for the behavior in the first place.  It can't "know" whether the altruistic behavior is appropriate or not, any more than it can know that the individual I'm saving is my brother versus a dog.
    Mundus vult decipi
    vongehr
    Yes: Genes are not likely to evolve to make old people suddenly become reckless.
    [The system would have to carry a mechanism that only does something really special if the system becomes sterile! This means a cost expectation value <c_old-reckless>. Such does not invade the gene-pool, because <b_old-reckless> will not outweight the cost.]

    But NO: The second section is not supported by that. We become reckless once a relative is just insulted, while we hardly care if a stranger is killed across the street.
    [Genes "care", because <b_help-brother> is huge!]

    Of course genes "know" or "care" nothing. This is short for: They are vitally involved in ("central to") shaping behavior, and thus "naturally selected", meaning they invade the gene-pool to a certain ratio (perhaps ~ zero) given a particular environment, which is what then can be modeled with b and c parameters.

    Gerhard: YOU NEED TO DO THE MATH! I am concerned that your arguments become effectively ever more anti-Darwinistic. The refusal of genes shaping behavior and of gradualism leading up to speciation implies God actively interfering! Lefty pseudo progressives selling out to PC establishment consensus do not grasp or do not care that this backfires. If you do the science properly, all that anti-racism, anti-sexism, anti-fringe physics (e.g. bashing v>c_light math like just again happened on Science2.0) is strictly anti-science.
    Fred Phillips
    Traveling tomorrow from Seoul to Daejeon, and I've printed out this entire thread to re-read carefully on the train. At first reading, though, it seems to add real clarity to the issue. No small feat, as the argument has engaged heavy thinkers from Daniel Dennett to Steven Jay Gould. Bravo, Gerhard!
    Gerhard Adam
    Thank you, that's a hell of a nice thing to say.  Let's hope it survives your re-reading :)

    In any case, if you have time I would appreciate any criticisms. 
    Mundus vult decipi
    Thor Russell
    OK some more thoughts of mine.
    Relating to an inclusive fitness like situation without a group:

    Consider a population of animals on a vast area that stretches further than individual members will shift in their lifetime. Lets say these animals do not live in groups, they are like say cats/tigers. However offspring will often end up in adjacent areas with overlapping  territories and come into contact with each other. So there is no meaningful group to speak of, but interaction nonetheless.
    Now consider sometimes these creatures get into a situation (a predator etc) where if they are not helped they will die, but if they are helped by another member of their species there is a 95% chance they both live, 5% they both die. Lets consider the behavior where one member helps another (helper vs nonhelper). Now to avoid being accused of gene centrism etc lets say that this helping behavior is inherited, but not from genes. If either parent has this behavior the offspring are all taught it, keep it for life, and teach their offspring. 
    So, will this behavior spread through the population?

    To answer this question, the best way I can see is through modelling, without group concepts being much help. 
    Lets say a helper spreads into a group of non-helpers. They will be at a disadvantage before breeding because they will risk themselves for no benefit. However if they do breed, then their offspring will have an advantage because they will help each other (assuming they don't immediately disperse the same way). Their proximity to each other will make it likely that a helper helps another one. The helping behavior will spread out because of this (and also because of the extreme inheritance characteristic where if either parent helps, all offspring do). Now if you change the percentages say from 5/95 live/die to 50/50 then the behavior will likely die out. So how can you determine the actual results without simulation and some kind of (behavior centric) concept? 
    If you then change things to make helper/nonhelper genetic, then sure the simulation will change because inheritance will be different but won't the principle be the same. 

    Inclusive fitness in general
    I don't have a problem with inclusive fitness because it just seems like a general case of a parent helping its offspring. I don't see why you need to get poorly defined "altruism" involved. As I noted earlier if you want to know whether an individual is successful, just counting offspring in one generation isn't the best way (they may fail to reproduce), some measure of their genes passed on after 5-10 generations makes more sense. In this context, a mother looking after her offspring is just helping individuals with many similar genes, the same individual looking after a cousin is a similar situation.  It follows that making a small effort that is a big help to a cousin is similar to making a moderate effort that is a moderate help to immediate offspring in terms of genes that will be present many generations hence.  If this helping behavior sometimes goes "wrong" and an unrelated creature is helped, then if that happens seldom, and there is little risk to the individual then it will hardly be selected against and needs no further explanation. Also then a gene that makes individuals help others that look or smell like their litter mates would  spread through the population too. It would spread better than just naive helper behavior because there would be more chance that the individual being helped was themselves a helper.
    I don't see why you need the group concept here. However where you do have clear groups you may be quite right and the pure inclusive fitness effect is almost completely swamped by all the other things that go on. If this was the case it wouldn't make inclusive fitness wrong, just not useful most of the time. Inclusive fitness could help groups get established, with other concepts taking over after that when the group has formed.











    Thor Russell
    Gerhard Adam
    Not a problem, and this is precisely the type of discussion that typically occurs to explore how an altruistic gene can become pervasive in a population.

    I don't have a problem with it.

    Here's the problem I do have.  In order for this scenario to work, it is only applicable during the time that such a gene is actually selectable.  After all, if it is pervasive in the population, then there's nothing to pass on.  Everyone already has it.

    Therefore such a gene would have to have high heritability [i.e. heavily influenced by genes] in order for this discussion to be meaningful.  If heritability is near zero, then there is no selection criteria and the environment is almost exclusively responsible for an variability among individuals.

    So, in effect we can say that this is merely a historical discussion to explain how an altruistic gene may have moved through a group, and since it has low heritability, then there is nothing further to explain.  It is pervasive and it is evolutionarily stable, so that there is no ability for selfish behavior to invade.

    In that situation, then talking about saving drowning brothers, or rationalizing any act, is irrelevant since we all fundamentally possess the altruism gene.  We could certainly argue that we [as humans] modify it through culture, learning, etc. but it is pretty well settled.

    However, the question remains.  Why would such a strategy develop in the first place?  You essentially pointed it out in your initial scenario.  It makes no sense to discuss altruism in the absence of a group, of some type.  Therefore when your example animals help each other out, they are ensuring the viability of the group, in the event that they may need assistance in the future.  Hence, by supporting the group, that trait will tend to get passed down because those that possess it improve their odds of surviving, over those that don't. 

    BTW ... don't worry about gene-centrism in this case.  My problem with it is when the assertion is made that all this occurs because of an "objective" the gene seeks.  In other words, I don't buy the argument that the gene seeks greater representation.  I expect that greater genetic representation is the byproduct of successful traits being passed on.  

    I don't have an argument with doing simulations.  Again, my problem stems from the unnecessarily rigid definition of altruism.  The fact that it relies on the fitness metric, which seems arbitrary [since how does one agree on maximum fitness?].  In addition, it becomes completely arbitrary if an animal has a choice in its fitness [or limitations - consider bears, elephants].

    By requiring such absolute measures, the relationship always holds, because we've defined all the exceptions out of it.  We don't count those that don't die, so therefore they weren't altruistic, so therefore they aren't part of the model.  The problem isn't in making the numbers work, it's trying to answer the daft questions like ... how could altruism spread amongst selfish individuals.

    It's obvious ... because it was more beneficial, but you'll never see that if you always assume that those that engaged in it are dead.

    If we accept the idea that this is merely risk-taking behavior, then we can see that the benefits can become significant very rapidly [and the original actor doesn't have to die nor sacrifice their own fitness].  Then we can predict that such behavior has such immense rewards it solidifies cooperative behavior, in those animals that gain an economy of scale.  The economy of scale is also important, because we can predict that the largest predators [for example] would tend to not cooperate because they don't gain any energy benefit from sharing a kill [especially if they likely need the whole thing for themselves].  So for them to cooperate, they would need to hunt more often to satisfy every member of the group.  Whereas you take an animal like a wolf, where a group can take down a large animal with a singular effort, then they all benefit from that one foray into hunting.

    These are the kinds of economies of scale I'm referring to, so if cooperation provides such large benefits, then altruism becomes self evidence as risk-taking behavior to preserve the group. 
    Mundus vult decipi
    Thor Russell
    OK if a gene has spread through the population, then yes inclusive fitness has nothing more to add. I see it as helping to ask what genes will spread. However I expect many such behaviors to have high heritability, after all there are many genes that make parents look after offspring. Parents in some species do it a lot, others not at all and its genetic to a large extent. The question to me is what would keep those genes from causing behavior to just help their own offspring rather than individuals generally? Well looking after offspring would obviously be more beneficial to passing on your genes, but there would be constant chance for "spillover" where the genes didn't distinguish completely. So there would be highly heritable genes that would also cause cousins to be helped (as well as total strangers sometimes) as well as immediate children. Such "altruism" could likely evolve then as soon as looking after children did.  So yes all species that look after their young would have a gene that sometimes causes behavior we call "altruism". 
    I am not sure then why you ask "why would it develop in the first place?". Genes aren't specific enough to stop such "spillover" where a parent will help a child in need even though its not their own. I still don't know why you need a group because I deliberately had a situation where it didn't make sense to talk of one. There was only helper vs non-helper behavior but no group. You can define all the helpers as being part of a group but I don't really see the significance of that. "inclusive" implies something like that anyway. In my helpers example, before helpers appeared, the "group" was massive and already surviving/viable, a plain with animals in all directions, but no boundaries you could put around them and really no "group". The helpers aren't ensuring the viability of the group (the unlimited animals stretching in all directions), we are just asking how/if the helper behavior will spread. If helper behavior spread to all individuals it could perhaps increase the population density per area (if predation rather than food was limiting), but it wouldn't necessarily affect the species survival. 

    Regarding "gene centrism" I am not sure that academic biologists really think of it as the objective of a gene, perhaps thats a popularization that has lead to misunderstandings. 
    " I expect that greater genetic representation is the byproduct of successful traits being passed on"
    Whats the difference? Genes recording successful behaviors or causing them seems pretty much the same to me.


    I agree that for groups, economies of scale etc need to be considered and inclusive fitness could very well take a back seat to other considerations and not be helpful or give mislead/wrong answers.
    Thor Russell
    vongehr
    Regarding "gene centrism" I am not sure that academic biologists really think of it as the objective of a gene, perhaps thats a popularization that has lead to misunderstandings.
    Perhaps some do, and Gerhard and Steve etc. seem to charge so, but it does not matter. You see it best in the historical context of that if you do not strongly focus on DNA molecules and their chemically completely described interactions, you have the still huge crowd of religious people simply not grasping that evolution needs no mystic ingredients. Gene centrism means "look, there is a mechanism that one can understand reductively". It is overpopularized of course by "skeptics" and mediocre "new atheists" and their simplistic bashing of higher level descriptions like group selection etc, but "gene centrism" never meant to say that the universe has been created by racist little DNA molecules that consciously favor Romney over Obama.
    vongehr
    It is pervasive and it is evolutionarily stable, so that there is no ability for selfish behavior to invade.
    That contrary behavior is kept out consistently is nothing but the fact of being "evolutionarily stable"! You are wrong to imply that the game is all over once equilibrium is reached. Equilibrium is where the game happens to stay while it is consistently played. This is how the equilibrium adapts to environmental change; this is how evolution goes on to gradually transform the systems (red queen).
    In that situation, then talking about saving drowning brothers, or rationalizing any act, is irrelevant since we all fundamentally possess the altruism gene.
    No we don't! We put huge numbers of psychopaths into jail for example.
    However, the question remains.  Why would such a strategy develop in the first place?
    You started this reply saying that this is precisely no longer the question. Now it all of a sudden is again.
    A couple of days of reading have revealed a different view of my world. The drapes opened slightly, or the shade partially raised, show me a glimpse of a whole world of study of which I was unaware. You have my eternal gratitude for the links in the article.

    First, as for the communication in a cellular environment: I actually was aware of the communication, I just wasn’t aware of the differences in my head. After I adjusted the polarization of my internal binoculars, it was clear that certain cells were in communication with others. My problem was that I generally conceive of communication having a feedback loop. Moreover, it may be that there are feedback loops that aren’t apparent to us mere humans.

    Second, I was wrong to introduce the singular case of human altruism into this discussion. I now understand that there are so many variables in what we call altruism, in every day life, which it will likely take many more generations before the issue is centered on a generally accepted definition. On the other hand, it may never happen.

    Next, I find that the terms altruism, cooperation, and mutualism, are at times used to mean the same thing. This finding serves to confuse my efforts at understanding even more.

    As you no doubt understood at the outset of my correspondence, I have very little knowledge of, or understandings of, the studies by evolutionary biologists, or evolutionary behaviorists. I find that I am much more comfortable believing that I understand some cellular biology. I believe that I can understand the binary function of DNA methylation as well as the way that canalization preserves a phenotype. I have no such belief for the supposed ‘altruism gene’.

    As a matter of no particular interest, I believe in neither an altruism gene nor a smoking gene.

    What I am starting to believe is that we humans, with our outsized egos and impressive intelligence, are no more than the vehicles that our cells use to propagate themselves.

    Finally, I completely agree with your summation that, given your definition, altruism serves the group and not individuals. Thank you.