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Hamilton - caste determination in stingless bees ?

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30 December 2003
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William L Hunt
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  1. Does anyone (Edser?) think that self-determination of caste is NOT occurring in the Melopine
    stingless bees?

    I will explain further. Self-determination of caste means there is an inherited genetic(allelic)
    difference between sister larva and because of this difference one follows a developmental pathway
    to become a sterile worker and the other to become a (potential) fertile queen. There should be an
    observable genetic difference in some alleles between the two. In all higher eusocial insects
    except one, the caste (worker/reproductive) of the larva is determined, not by the larva herself,
    but by the workers usually by the amount or type of food given the larva. The exception is the
    Melopine stingless bees. In Melopona all cells are the same size, provisioned with identical
    amounts of food, and after the egg is laid the cell is capped. There is no further contact with the
    developing larva until it emerges as a fully formed worker or queen. It would seem that self-
    determination of caste is occurring. Melopona is among those bees that reproduce only by hive
    swarming (fission) and only need a new queen when swarming (maybe once or twice a season). Other
    hive swarming bees produce very few new queens but in Melopona as many as 25% of females may emerge
    as queens. They are all killed immediately by the workers (unless the hive is in the process of
    swarming when it would keep one). What a waste! The inefficiency of the Melopona hive producing all
    these queens just to immediately kill them is striking. If the workers controlled/determined the
    larva caste, they would never overproduce unneeded new queens. Again it seems clear that self-
    determination of caste is occurring in Melopona. I should note that as far as I know no lab has
    looked or is currently looking for the genetic difference between worker and queen in Melopona. If
    self-determination of caste is occurring there must be a genetic difference and I am quite curious
    what it actually is. Possible allelic mechanisms have been proposed (Kerr 1950). I ask again if
    anyone thinks self-determination of caste is NOT occurring in the Melopine stingless bees?

    William L Hunt

  2. William L Hunt said:

    Does anyone (Edser?) think that self-determination of caste is NOT occurring in the Melopine
    stingless bees?


    Nothing you wrote seems to rule out maternal effects - i.e. it's the queen who's determining caste.
    You can then explain the excess females as bet-hedging, as their caste is determined some time
    before hatching.

    Just an alternative hypothesis.

    Bob

    Quoted message said:

    I will explain further. Self-determination of caste means there is an inherited genetic(allelic)
    difference between sister larva and because of this difference one follows a developmental
    pathway to become a sterile worker and the other to become a (potential) fertile queen. There
    should be an observable genetic difference in some alleles between the two. In all higher
    eusocial insects except one, the caste (worker/reproductive) of the larva is determined, not by
    the larva herself, but by the workers usually by the amount or type of food given the larva. The
    exception is the Melopine stingless bees. In Melopona all cells are the same size, provisioned
    with identical amounts of food, and after the egg is laid the cell is capped. There is no further
    contact with the developing larva until it emerges as a fully formed worker or queen. It would
    seem that self-determination of caste is occurring. Melopona is among those bees that reproduce
    only by hive swarming (fission) and only need a new queen when swarming (maybe once or twice a
    season). Other hive swarming bees produce very few new queens but in Melopona as many as 25% of
    females may emerge as queens. They are all killed immediately by the workers (unless the hive is
    in the process of swarming when it would keep one). What a waste! The inefficiency of the
    Melopona hive producing all these queens just to immediately kill them is striking. If the
    workers controlled/determined the larva caste, they would never overproduce unneeded new queens.
    Again it seems clear that self-determination of caste is occurring in Melopona. I should note
    that as far as I know no lab has looked or is currently looking for the genetic difference
    between worker and queen in Melopona. If self-determination of caste is occurring there must be a
    genetic difference and I am quite curious what it actually is. Possible allelic mechanisms have
    been proposed (Kerr 1950). I ask again if anyone thinks self-determination of caste is NOT
    occurring in the Melopine stingless bees?

    William L Hunt

    --
    Bob O'Hara

    Rolf Nevanlinna Institute
    P.O. Box 4 (Yliopistonkatu 5) FIN-00014 University of Helsinki Finland Telephone: +358-9-191 23743
    Mobile: +358 50 599 0540 Fax: +358-9-191 22 779 WWW: rni.helsinki.fi~boh

  3. "Anon." <[email hidden]> wrote in message news:<[email hidden]>...

    Quoted message said:
    William L Hunt said:

    Does anyone (Edser?) think that self-determination of caste is NOT occurring in the Melopine
    stingless bees?


    Nothing you wrote seems to rule out maternal effects - i.e. it's the queen who's determining
    caste. You can then explain the excess females as bet-hedging, as their caste is determined some
    time before hatching.

    Just an alternative hypothesis.

    Bob

    And, just to throw in yet another alternative, are we sure that the one queen that survives the
    slaughter is the winner of a lottery, or is she the winner of some kind of beauty contest? Workers
    unite - choose your own leaders!

    It would be nice though if you had found the exception that proves the rule. (Ouch, that stings!)

    Jim

    btw. I would be interested in seeing the allelic model that Kerr suggested. I tried to come up with
    one and failed miserably.

    Quoted message said:
    Quoted message said:

    I will explain further. Self-determination of caste means there is an inherited
    genetic(allelic) difference between sister larva and because of this difference one follows a
    developmental pathway to become a sterile worker and the other to become a (potential) fertile
    queen. There should be an observable genetic difference in some alleles between the two. In all
    higher eusocial insects except one, the caste (worker/reproductive) of the larva is determined,
    not by the larva herself, but by the workers usually by the amount or type of food given the
    larva. The exception is the Melopine stingless bees. In Melopona all cells are the same size,
    provisioned with identical amounts of food, and after the egg is laid the cell is capped. There
    is no further contact with the developing larva until it emerges as a fully formed worker or
    queen. It would seem that self-determination of caste is occurring. Melopona is among those
    bees that reproduce only by hive swarming (fission) and only need a new queen when swarming
    (maybe once or twice a season). Other hive swarming bees produce very few new queens but in
    Melopona as many as 25% of females may emerge as queens. They are all killed immediately by the
    workers (unless the hive is in the process of swarming when it would keep one). What a waste!
    The inefficiency of the Melopona hive producing all these queens just to immediately kill them
    is striking. If the workers controlled/determined the larva caste, they would never overproduce
    unneeded new queens. Again it seems clear that self-determination of caste is occurring in
    Melopona. I should note that as far as I know no lab has looked or is currently looking for the
    genetic difference between worker and queen in Melopona. If self-determination of caste is
    occurring there must be a genetic difference and I am quite curious what it actually is.
    Possible allelic mechanisms have been proposed (Kerr 1950). I ask again if anyone thinks self-
    determination of caste is NOT occurring in the Melopine stingless bees?

    William L Hunt

    --
    Bob O'Hara

    Rolf Nevanlinna Institute
    P.O. Box 4 (Yliopistonkatu 5) FIN-00014 University of Helsinki Finland Telephone: +358-9-191 23743
    Mobile: +358 50 599 0540 Fax: +358-9-191 22 779 WWW: rni.helsinki.fi~boh

  4. Anon. said:
    William L Hunt said:

    Does anyone (Edser?) think that self-determination of caste is NOT occurring in the Melopine
    stingless bees?


    Nothing you wrote seems to rule out maternal effects - i.e. it's the queen who's determining caste.
    You can then explain the excess females as bet-hedging, as their caste is determined some time
    before hatching.

    Just an alternative hypothesis.

    Bob


    Well, both the queen and the workers should have the same interest here in maximizing the hive
    efficiency by producing only queens as needed with maybe just a little "bet-hedging". Other hive
    swarming species do produce a small excess over what is needed ("bet-hedging"😉 but nothing like the
    excess seen in Melipona. I made this post hoping to get a response from John Edser. From his logic
    he reasons that a sterility gene(s) cannot be selected. Self-determination of caste requires that
    this is, in fact, happening. Maybe I should have made this clearer to provoke a response from
    Edser. I can picture Melapine bees buzzing Edser and telling him to take a look at our larva. They
    are determining their own caste fate and choosing sterility 75+ percent of the time. They have been
    doing this for millions of years. Surely sterility is being selected. This is not mutualism. The
    larva are acting in their own self-interest, the only reason they all don't become queens is that
    there must be some "choosing" to be workers to form the swarm that is required to found a hive. A
    queen cannot found a hive by herself; no workers and everyone dies. The question might be how does
    one predict what this optimal percentage is? These prediction formula have been derived by various
    methods and they all will have a mention of the name Hamiltion and relatedness r values. In fact,
    the formula is just: percentage of larva becoming queens P=(1 - Rf)/(1 + Rm) where Rf is
    relatedness of larva to sister and Rm is relatedness of larva to male. Melopines are always single
    mated, so Rf= .75 and Rm ranges from .25 to .75 depending on species. So the predicted P% is 14% -
    20%. For a paper deriving this (from the same site given by Wirt Atmar): "Caste fate conflict in
    swarm-founding social Hymenoptera: an inclusive fitness analysis".
    www.shef.ac.uk/uni/projects/taplab/pdf/wrbjeb2003.pdf

    I think Edser might grind his teeth just reading the title of this paper -).

    In any case, I think some lab will eventually find the allelic mechanism and that should end any
    questions of whether it really is self-determination that is happening. William L Hunt

    ... [snip] ...

  5. On Sat, 20 Dec 2003 22:05:05 +0000 (UTC), [email hidden]

    (Jim Menegay) said:

    "Anon." <[email hidden]> wrote in message
    news:<[email hidden]>...

    Quoted message said:
    William L Hunt said:

    Does anyone (Edser?) think that self-determination of caste is NOT occurring in the Melopine
    stingless bees?


    Nothing you wrote seems to rule out maternal effects - i.e. it's the queen who's determining
    caste. You can then explain the excess females as bet-hedging, as their caste is determined some
    time before hatching.

    Just an alternative hypothesis.

    Bob

    And, just to throw in yet another alternative, are we sure that the one queen that survives the
    slaughter is the winner of a lottery, or is she the winner of some kind of beauty contest? Workers
    unite - choose your own leaders!

    It would be nice though if you had found the exception that proves the rule. (Ouch, that stings!)

    Jim

    btw. I would be interested in seeing the allelic model that Kerr suggested. I tried to come up with
    one and failed miserably.


    The paper is Kerr,W.E. 1950 "Genetic determination of castes in the genus Melipona". Genetics The
    model is quite simple and similiar to how many bees determine

    Melopona might be close to 25% so Kerr used a 2 loci 2 allele model. Aa Bb, if the female is
    heterozygotic at both loci she develops into a queen, if homozygotic at either loci she developes
    into a sterile worker. Remember males are all hapoid. Work it out. A Female Aa Bb mating with any
    haploid male will produce 25% Aa Bb females. With a 3 loci model you would have 12.5% and a 4 loci
    6.25%. If you have two loci on the same chromosome and have some degree of linkage (under
    selection?), you could produce any intermediate percentage value. The actual allelic mechanism is
    not known and the mechanism may limit these percentage so they can not reach a predicted optimal
    value. William L Hunt

    ... [snip] ...

  6. [email hidden] (William L Hunt) wrote in message news:<[email hidden]>...

    Quoted message said:
    (Jim Menegay) said:
    Quoted message said:

    William L Hunt wrote:
    > Does anyone (Edser?) think that self-determination of caste is NOT occurring in the Melopine
    > stingless bees?

    Quoted message said:
    Quoted message said:

    btw. I would be interested in seeing the allelic model that Kerr suggested. I tried to come up
    with one and failed miserably.


    The paper is Kerr,W.E. 1950 "Genetic determination of castes in the genus Melipona". Genetics The
    model is quite simple and similiar to how many bees determine

    Quoted message said:

    Melopona might be close to 25% so Kerr used a 2 loci 2 allele model. Aa Bb, if the female is
    heterozygotic at both loci she develops into a queen, if homozygotic at either loci she developes
    into a sterile worker. Remember males are all hapoid. Work it out. A Female Aa Bb mating with any
    haploid male will produce 25% Aa Bb females. With a 3 loci model you would have 12.5% and a 4 loci
    6.25%. If you have two loci on the same chromosome and have some degree of linkage (under
    selection?), you could produce any intermediate percentage value. The actual allelic mechanism is
    not known and the mechanism may limit these percentage so they can not reach a predicted optimal
    value. William L Hunt

    Thx. That is a cute mechanism, but if it applies, there doesn't seem to be any relevance to
    Edser/Hamilton. No way to cheat. Each allele A,a,B,b in a haploid gamete is just as likely to make
    you fertile as any other. It is a crapshoot.

    But maybe I am missing why you think this example is significant. I'm also curious why Kerr thought
    there are differences between Melopines and honeybees in which castes are closely related to which
    other castes. Are the Melopines polyandrous (if that is the right word)?

  7. Bob said:
    William L Hunt said:

    Does anyone (Edser?) think that self-determination of caste is NOT occurring in the Melopine
    stingless bees?


    Nothing you wrote seems to rule out maternal effects - i.e. it's the queen who's determining caste.
    You can then explain the excess females as bet-hedging, as their caste is determined some time
    before hatching.

    Just an alternative hypothesis.

    A more recent, very brief reference to Kerr's thoughts on Melopine caste determination can
    be found at:

    chemecol.orgp 60.html

    An even more detailed, although short summary of the differences between the caste determination
    mechanisms associated with Melopina vs. the other social bees can be found at:

    shef.ac.uktwenseleers.html

    Wirt Atmar

  8. [email hidden] (William L Hunt) wrote in news:bs78kk$2a88$1
    @darwin.ediacara.org:

    Quoted message said:
    Anon. said:
    William L Hunt said:

    Does anyone (Edser?) think that self-determination of caste is NOT occurring in the Melopine
    stingless bees?

    Quoted message said:
    Quoted message said:

    Nothing you wrote seems to rule out maternal effects - i.e. it's the queen who's determining
    caste. You can then explain the excess females as bet-hedging, as their caste is determined some
    time before hatching.

    Quoted message said:
    Quoted message said:

    Just an alternative hypothesis.

    Quoted message said:

    Well, both the queen and the workers should have the same interest here in maximizing the hive
    efficiency by producing only queens as needed with maybe just a little "bet-hedging". Other hive
    swarming species do produce a small excess over what is needed ("bet-hedging"😉 but nothing like
    the excess seen in Melipona. I made this post hoping to get a response from John Edser. From his
    logic he reasons that a sterility gene(s) cannot be selected. Self-determination of caste
    requires that this is, in fact, happening. Maybe I should have made this clearer to provoke a
    response from Edser. I can picture Melapine bees buzzing Edser and telling him to take a look at
    our larva. They are determining their own caste fate and choosing sterility 75+ percent of the
    time. They have been doing this for millions of years. Surely sterility is being selected. This
    is not mutualism. The larva are acting in their own self-interest, the only reason they all don't
    become queens is that there must be some "choosing" to be workers to form the swarm that is
    required to found a hive. A queen cannot found a hive by herself; no workers and everyone dies.
    The question might be how does one predict what this optimal percentage is? These prediction
    formula have been derived by various methods and they all will have a mention of the name
    Hamiltion and relatedness r values. In fact, the formula is just: percentage of larva becoming
    queens P=(1 - Rf)/(1 + Rm) where Rf is relatedness of larva to sister and Rm is relatedness of
    larva to male. Melopines are always single mated, so Rf= .75 and Rm ranges from .25 to .75
    depending on species. So the predicted P% is 14% - 20%.


    (snip)

    Quoted message said:

    In any case, I think some lab will eventually find the allelic mechanism and that should end any
    questions of whether it really is self-determination that is happening.

    To be charitable, John Edser may not have seen this particular follow - I often find that stuff on
    newsgroups disappears from some ISP's and/or newsreaders. To be uncharitable, Edser doesn't
    particularly like to respond to real research - it often fails to correspond to what must be true
    according to his logic.

    But my question is, how did this behavior evolve in the first place? Granted that self-determination
    is happening, how would you start to select it? In most of the examples of kin helping that I have
    seen, the involved entities still retain their fertility. So you can see how they can sort of "slip
    into" kin selection. In the case of the bees, it would seem there would never be an initial
    advantage to choosing sterility, even though there might be an advantage in choosing not to
    reproduce. It would seem that the self-determination would have to be a trait derived from a
    previous "standard" bee. The way I can see this working is if it was frequently true that hives were
    started with a small number of bees and that the original queen frequently died.

    If on the other hand Melapine bees are the ancestral condition and other bees the "derived"
    condition, I would think there would have to be some additional cost in being a queen, so that
    choosing sterility would be similar to menopause in humans. But I thought that bee queens live
    longer, not shorter, than sterile workes. Of course, as I know little about bees and nothing about
    Melapine bees, I would not be surprised to find that I am way off base on the above.

    Yours,

    Bill Morse

  9. William Morse said:

    [email hidden] (William L Hunt) wrote in news:bs78kk$2a88$1 @darwin.ediacara.org:

    Quoted message said:
    Anon. said:

    William L Hunt wrote:
    > Does anyone (Edser?) think that self-determination of caste is NOT occurring in the Melopine
    > stingless bees?

    Quoted message said:
    Quoted message said:

    Nothing you wrote seems to rule out maternal effects - i.e. it's the queen who's determining
    caste. You can then explain the excess females as bet-hedging, as their caste is determined some
    time before hatching.

    Quoted message said:
    Quoted message said:

    Just an alternative hypothesis.

    Quoted message said:

    Well, both the queen and the workers should have the same interest here in maximizing the hive
    efficiency by producing only queens as needed with maybe just a little "bet-hedging". Other hive
    swarming species do produce a small excess over what is needed ("bet-hedging"😉 but nothing like
    the excess seen in Melipona. I made this post hoping to get a response from John Edser. From his
    logic he reasons that a sterility gene(s) cannot be selected. Self-determination of caste
    requires that this is, in fact, happening. Maybe I should have made this clearer to provoke a
    response from Edser. I can picture Melapine bees buzzing Edser and telling him to take a look at
    our larva. They are determining their own caste fate and choosing sterility 75+ percent of the
    time. They have been doing this for millions of years. Surely sterility is being selected. This
    is not mutualism. The larva are acting in their own self-interest, the only reason they all
    don't become queens is that there must be some "choosing" to be workers to form the swarm that
    is required to found a hive. A queen cannot found a hive by herself; no workers and everyone
    dies. The question might be how does one predict what this optimal percentage is? These
    prediction formula have been derived by various methods and they all will have a mention of the
    name Hamiltion and relatedness r values. In fact, the formula is just: percentage of larva
    becoming queens P=(1 - Rf)/(1 + Rm) where Rf is relatedness of larva to sister and Rm is
    relatedness of larva to male. Melopines are always single mated, so Rf= .75 and Rm ranges from
    .25 to .75 depending on species. So the predicted P% is 14% - 20%.


    (snip)

    Quoted message said:

    In any case, I think some lab will eventually find the allelic mechanism and that should end any
    questions of whether it really is self-determination that is happening.

    To be charitable, John Edser may not have seen this particular follow - I often find that stuff on
    newsgroups disappears from some ISP's and/or newsreaders. To be uncharitable, Edser doesn't
    particularly like to respond to real research - it often fails to correspond to what must be true
    according to his logic.

    But my question is, how did this behavior evolve in the first place? Granted that self-
    determination is happening, how would you start to select it? In most of the examples of kin
    helping that I have seen, the involved entities still retain their fertility. So you can see how
    they can sort of "slip into" kin selection.


    In looking at a route to sociality in bees one should not look at the advanced eusocials especially
    these swarming bees that have completely replace individual queen dispersal as means of
    replication. Look at subsocial bees and you see more what you expect where they sort of "slip into"
    kin selection. I should note that I was a little loose in my use of the term "sterile". In some
    eusocial species worker have no ovaries and are truly sterile, but in the Melipine bees I think in
    all species workers have ovaries and in some species in fact the workers make almost all the males.
    In others the queen produces all the males, and only in these species should one even think of the
    workers as being "effectively" sterile.

    Quoted message said:

    In the case of the bees, it would seem there would never be an initial advantage to choosing
    sterility, even though there might be an advantage in choosing not to reproduce. It would seem
    that the self-determination would have to be a trait derived from a previous "standard" bee. The
    way I can see this working is if it was frequently true that hives were started with a small
    number of bees and that the original queen frequently died.

    If on the other hand Melapine bees are the ancestral condition and other bees the "derived"
    condition, I would think there would have to be some additional cost in being a queen, so that
    choosing sterility would be similar to menopause in humans. But I thought that bee queens live
    longer, not shorter, than sterile workes. Of course, as I know little about bees and nothing about
    Melapine bees, I would not be surprised to find that I am way off base on the above.


    Well, queens live longer than workers sometimes a lot longer. Melapine and Trigonine bees are very
    similiar and closely related. In the Trigonines workers control caste fate by providing a much
    greater food mass to a female larva they have choosen to be a queen. Which method of determining
    caste fate was present in the ancestral line before the split is not known. My view would be that
    it most probably was Trigonine method. There is a genera in the Trigonines where occasionally larva
    cheat by breaking through the cell wall to also eat the neighbor's food mass and in doing so become
    what are called "dwarf queens". They have not been observed to become accepted as true queens but
    one would think they must even if very infrequently else why would the behavior persist. It seems
    similiar in some way to the Melipine "self-determination", at least some Trigonine larva are trying
    to determine their caste fate even if they have not been observed to be successful. William L Hunt

    Quoted message said:

    Yours,

    Bill Morse

  10. Quoted message said:

    WH:- Well, both the queen and the workers should have the same interest here in maximizing the
    hive efficiency by producing only queens as needed with maybe just a little "bet-hedging". Other
    hive swarming species do produce a small excess over what is needed ("bet-hedging"😉 but nothing
    like the excess seen in Melipona. I made this post hoping to get a response from John Edser. From
    his logic he reasons that a sterility gene(s) cannot be selected.

    JE:- WH has misrepresented my argument. I did not respond originally because I was sure that this
    was the case. Now WH have written down what WH thought JE was saying JE can now arue that it was not
    what JE was actually saying.

    I argued that eusocial sterility cannot be selected using Hamilton's rule where the altruistic genes
    are selected _for_ within the body of a _sterile_ form. A sterile eusocial form has _zero_ fitness
    so nothing within it CAN be selected _for_. Thus the only possible argument that makes sense is that
    genes for sterility are only selected _for_ within the body of their _fertile_ parent,
    i.e. from whence they came. Thus the only possible use a sterile form has is to serve the parent as
    modular extension of the parental soma. This argument requires _zero_ organism fitness altruism
    (OFA) of the _parent_, who alone, had any fitness to give away in the first place. All that is
    happening here is that the parents that increase their absolute fitness as _one_ Darwinian
    total of the number of FERTILE forms they reproduce within the _same_ population, by allowing
    SOME of their reproductives to remain sterile were selected _for_. Genes were subselected to
    pass this on. Sterile casts are an _investment_ cost for the parent and not an altruistic
    sacrifice of sterile forms, who only have zero Darwinian fitness to give away.

    A eusocial reproductive increases the survival of the fertile parents it is just a Darwinian slave
    of, allowing a possible increase in such a parents absolute Darwinian fitness and thus the sterile
    genes that code ofr it. Thus genes for eusociality are selected for within the parent and not
    within the sterile form. It does not matter if the parent turns them on or off, or the sterile
    reproductive does.

    I can't imagine why such a simple argument has escaped Neo Darwinists who seem hell bent on finding
    non existent OFA everywhere in nature. I can only conclude that OFA is just a product of a massive
    "political correctness" bias.

    Quoted message said:

    WH:- Self-determination of caste requires that this is, in fact, happening. Maybe I should have
    made this clearer to provoke a response from Edser. I can picture Melapine bees buzzing Edser and
    telling him to take a look at our larva. They are determining their own caste fate and choosing
    sterility 75+ percent of the time. They have been doing this for millions of years. Surely
    sterility is being selected. This is not mutualism.

    JE:- Yes "sterility is being selected" _for_ but the sterile genes are only being selected _for_
    within the soma of a _fertile_ parent. This is "not mutualism" between parents and sterile casts
    because sterile casts only have zero fitness to be mutual with. Mutualism is only possible between
    FERTILE PARENTS. One Day Neo Darwinians might realise that only FERTILE forms are valid units of
    selection (UOS) and thus, valid units of fitness. (UOF). While they keep counting sterile forms as
    valid UOS their UOF totals are hopelessly wrong.

    Respectfully Yours,

    John Edser Independent Researcher

    PO Box 266 Church Pt NSW 2105 Australia

    [email hidden]

  11. On Tue, 27 Jan 2004 06:40:28 +0000 (UTC), "John Edser"
    <[email hidden]> wrote:

    ... [snip] ...

    Quoted message said:
    Quoted message said:

    WH:- Self-determination of caste requires that this is, in fact, happening. Maybe I should have
    made this clearer to provoke a response from Edser. I can picture Melapine bees buzzing Edser and
    telling him to take a look at our larva. They are determining their own caste fate and choosing
    sterility 75+ percent of the time. They have been doing this for millions of years. Surely
    sterility is being selected. This is not mutualism.

    JE:- Yes "sterility is being selected" _for_ but the sterile genes are only being selected _for_
    within the soma of a _fertile_ parent. This is "not mutualism" between parents and sterile casts
    because sterile casts only have zero fitness to be mutual with. Mutualism is only possible between
    FERTILE PARENTS. One Day Neo Darwinians might realise that only FERTILE forms are valid units of
    selection (UOS) and thus, valid units of fitness. (UOF). While they keep counting sterile forms as
    valid UOS their UOF totals are hopelessly wrong.

    WH:- Just to be sure I am have no misunderstanding at all could you answer my original question with
    a "Yes' or "No". The question was: Do you think it is even possible that "self-determination of
    caste" is occurring in the Melapine bees?

    In all other eusocial insects the selection for sterility is clearly occurring through the fertile
    queens (as you say it must). The only question is how it is happening in Melapine bees. "Self-
    determination of caste" means a larva becomes a worker or queen depending upon its own "sterility"
    alleles. William L Hunt

    Quoted message said:


    Respectfully Yours,

    John Edser Independent Researcher

    PO Box 266 Church Pt NSW 2105 Australia

    [email hidden]

  12. Quoted message said:
    Quoted message said:

    WH:- Self-determination of caste requires that this is, in fact, happening. Maybe I should have
    made this clearer to provoke a response from Edser. I can picture Melapine bees buzzing Edser and
    telling him to take a look at our larva. They are determining their own caste fate and choosing
    sterility 75+ percent of the time. They have been doing this for millions of years. Surely
    sterility is being selected. This is not mutualism.

    Quoted message said:

    JE:- Yes "sterility is being selected" _for_ but the sterile genes are only being selected _for_
    within the soma of a _fertile_ parent. This is "not mutualism" between parents and sterile casts
    because sterile casts only have zero fitness to be mutual with. Mutualism is only possible between
    FERTILE PARENTS. One Day Neo Darwinians might realise that only FERTILE forms are valid units of
    selection (UOS) and thus, valid units of fitness. (UOF). While they keep counting sterile forms as
    valid UOS their UOF totals are hopelessly wrong.

    WH:- Just to be sure I am have no misunderstanding at all could you answer my original question with
    a "Yes' or "No". The question was: Do you think it is even possible that "self-determination of
    caste" is occurring in the Melapine bees?

    JE:- Yes.

    WH:- In all other eusocial insects the selection for sterility is clearly occurring through the
    fertile queens (as you say it must). The only question is how it is happening in Melapine bees.
    "Self-determination of caste" means a larva becomes a worker or queen depending upon its own
    "sterility" alleles".

    JE:- I can't see it makes any difference. These alleles remain selected for within the fertile form
    of Melapine bees. Being born a worker or a queen and not made into one by a pheromone just fixes the
    sterility genes. This does not refute the proposition that the genes that code for either caste can
    only be selected for within a fertile parent.

    If the castes are now fixed by inherited genes then no pheromone is required. Thus the normal
    restriction for an almost totally enclosed nest environment to disperse a pheromone is now removed.
    I would suggest that this form of eusociality may have evolved from the more common queen induced
    pheromone system that requires enclosure. Once such a system has evolved then the assimilation of
    caste forms can remove the encumbrance of a pheromone allowing eusociality to occur with almost no
    enclosure of the nest. I have no idea if Melapine bee nests are fully enclosed or not, but I would
    predict they would be less enclosed.

    Respectfully Yours,

    John Edser Independent Researcher

    PO Box 266 Church Pt NSW 2105 Australia

    [email hidden]

  13. "John Edser" <[email hidden]> wrote in message news:<[email hidden]>...

    Quoted message said:
    Quoted message said:
    Quoted message said:

    WH:- Self-determination of caste requires that this is, in fact, happening. Maybe I should have
    made this clearer to provoke a response from Edser. I can picture Melapine bees buzzing Edser
    and telling him to take a look at our larva. They are determining their own caste fate and
    choosing sterility 75+ percent of the time. They have been doing this for millions of years.
    Surely sterility is being selected. This is not mutualism.

    Quoted message said:

    JE:- Yes "sterility is being selected" _for_ but the sterile genes are only being selected _for_
    within the soma of a _fertile_ parent. This is "not mutualism" between parents and sterile casts
    because sterile casts only have zero fitness to be mutual with. Mutualism is only possible
    between FERTILE PARENTS. One Day Neo Darwinians might realise that only FERTILE forms are valid
    units of selection (UOS) and thus, valid units of fitness. (UOF). While they keep counting
    sterile forms as valid UOS their UOF totals are hopelessly wrong.

    WH:- Just to be sure I am have no misunderstanding at all could you answer my original question
    with a "Yes' or "No". The question was: Do you think it is even possible that "self-determination
    of caste" is occurring in the Melapine bees?

    JE:- Yes.

    WH:- In all other eusocial insects the selection for sterility is clearly occurring through the
    fertile queens (as you say it must). The only question is how it is happening in Melapine bees.
    "Self-determination of caste" means a larva becomes a worker or queen depending upon its own
    "sterility" alleles".

    JE:- I can't see it makes any difference. These alleles remain selected for within the fertile
    form of Melapine bees. Being born a worker or a queen and not made into one by a pheromone just
    fixes the sterility genes. This does not refute the proposition that the genes that code for
    either caste can only be selected for within a fertile parent.

    If the castes are now fixed by inherited genes then no pheromone is required. Thus the normal
    restriction for an almost totally enclosed nest environment to disperse a pheromone is now
    removed. I would suggest that this form of eusociality may have evolved from the more common queen
    induced pheromone system that requires enclosure. Once such a system has evolved then the
    assimilation of caste forms can remove the encumbrance of a pheromone allowing eusociality to
    occur with almost no enclosure of the nest. I have no idea if Melapine bee nests are fully
    enclosed or not, but I would predict they would be less enclosed.

    Respectfully Yours,

    John Edser Independent Researcher

    According to one interpretation of what John has written in the past, "self-determination" of
    ANYTHING is impossible until the self has

    on behalf of the parents. Since in most species, the bulk of development

    as taking place on behalf of the parent. By these rules, genetically caused infertility is always
    parentally determined. It is logically impossible for it to be self-determined. (This does not apply
    to early menopause - that is self-determined, I guess).

    There are some other curious aspects to this. What a Hamiltonian would call parent-child conflict
    must be viewed as a conflict between the two parents in John's accounting. As a consequence, we
    might expect the children of strictly monogamous parents to be better behaved. :-}

  14. Quoted message said:

    WH:- Just to be sure I am have no misunderstanding at all could you answer my original question
    with a "Yes' or "No". The question was: Do you think it is even possible that "self-determination
    of caste" is occurring in the Melapine bees?

    Quoted message said:

    JE:- Yes.

    Quoted message said:

    WH:- In all other eusocial insects the selection for sterility is clearly occurring through the
    fertile queens (as you say it must). The only question is how it is happening in Melapine bees.
    "Self-determination of caste" means a larva becomes a worker or queen depending upon its own
    "sterility" alleles".

    Quoted message said:

    JE:- I can't see it makes any difference. These alleles remain selected for within the fertile
    form of Melapine bees. Being born a worker or a queen and not made into one by a pheromone just
    fixes the sterility genes. This does not refute the proposition that the genes that code for
    either caste can only be selected for within a fertile parent.

    If the castes are now fixed by inherited genes then no pheromone is required. Thus the normal
    restriction for an almost totally enclosed nest environment to disperse a pheromone is now
    removed. I would suggest that this form of eusociality may have evolved from the more common queen
    induced pheromone system that requires enclosure. Once such a system has evolved then the
    assimilation of caste forms can remove the encumbrance of a pheromone allowing eusociality to
    occur with almost no enclosure of the nest. I have no idea if Melapine bee nests are fully
    enclosed or not, but I would predict they would be less enclosed.

    WH:- According to one interpretation of what John has written in the past, "self-determination" of
    ANYTHING is impossible until the self has

    JE:- It is just another "chicken or the egg" problem. What came first a fertile or infertile form?
    The answer is NEITHER because they are both part of the same unit of selection. The artificial
    division of a unit of selection into parts is a necessary act of analysis but analysis on its own
    cannot provide a correct synthesis.

    WH:- Immature forms are seen by John as acting on behalf of the parents.

    JE:- What else could they possibly do?

    WH:- Since in most species, the bulk of development

    must be viewed as taking place on behalf of the parent.

    JE:- Absolutely. This may produce bizarre syndromes. Maybe cot death in infants is a way genes in
    children kill off excess immature forms. Parents can serially reproduce. Thus they may benefit by
    the removal of excess infertile forms that they wish to get rid of now and replace later on.

    WH:- By these rules, genetically caused infertility is always parentally determined. It is logically
    impossible for it to be self-determined. (This does not apply to early menopause - that is self-
    determined, I guess).

    JE:- It is impossible for the egg to be selected without a chicken and it is impossible for the
    chicken to be selected without an egg.

    WH:- There are some other curious aspects to this. What a Hamiltonian would call parent-child
    conflict must be viewed as a conflict between the two parents in John's accounting. As a
    consequence, we might expect the children of strictly monogamous parents to be better behaved. :-}

    JE:- Please elaborate.

    I would appreciate information re: the nest enclose of stingless bees. Are their nests less enclosed
    than pheromone induced eusocials?

    Respectfully Yours,

    John Edser Independent Researcher

    PO Box 266 Church Pt NSW 2105 Australia

    [email hidden]

  15. John Edser said:

    JE:- I would appreciate information re: the nest enclose of stingless bees. Are their nests less
    enclosed than pheromone induced eusocials?


    Nests of Melipona stingless bees are well enclosed in a fashion similiar to other bees that
    reproduce by swarming. One difference might be the hive size which is 1000-2000 workers in
    Melapines and can sometimes be much larger in other swarming species. In the closely related
    Trigonine bees as in honey bees, female caste fate is nutritionally controlled by workers.
    Trigonine bees generally produce no excess queens. William L Hunt

    Quoted message said:

    Respectfully Yours,

    John Edser Independent Researcher

    PO Box 266 Church Pt NSW 2105 Australia

    [email hidden]

  16. "John Edser" <[email hidden]> wrote in message news:<[email hidden]>...
    John,
    You have attributed to WH something that I (JM) wrote. I have
    corrected this in what follows. Sorry, maybe I should have
    put my own "JM:-" in front of what I wrote.

    Quoted message said:

    JM:- According to one interpretation of what John has written in the past, "self-determination" of
    ANYTHING is impossible until the self has

    Quoted message said:


    JE:- It is just another "chicken or the egg" problem. What came first a fertile or infertile form?
    The answer is NEITHER because they are both part of the same unit of selection. The artificial
    division of a unit of selection into parts is a necessary act of analysis but analysis on its own
    cannot provide a correct synthesis.

    JM:- Immature forms are seen by John as acting on behalf of the parents.

    JE:- What else could they possibly do?

    JM:- Since in most species, the bulk of development

    Quoted message said:

    must be viewed as taking place on behalf of the parent.

    JE:- Absolutely. This may produce bizarre syndromes. Maybe cot death in infants is a way genes in
    children kill off excess immature forms. Parents can serially reproduce. Thus they may benefit by
    the removal of excess infertile forms that they wish to get rid of now and replace later on.

    JM:- By these rules, genetically caused infertility is always parentally determined. It is
    logically impossible for it to be self-determined. (This does not apply to early menopause - that
    is self-determined, I guess).

    JE:- It is impossible for the egg to be selected without a chicken and it is impossible for the
    chicken to be selected without an egg.

    JM:- I am happy to find that I have interpreted John right, so far.

    Quoted message said:

    JM:- There are some other curious aspects to this. What a Hamiltonian would call parent-child
    conflict must be viewed as a conflict between the two parents in John's accounting. As a
    consequence, we might expect the children of strictly monogamous parents to be better behaved. :-}

    JE:- Please elaborate.

    JM:- Trivers, I believe, has written about parent/child conflict - for example, the child does not
    appreciate being weaned. Trivers interprets this as a conflict of interest between child and mother.
    However, Edser believes

    maturity. Therefore, weaning conflict is a conflict between the interests of the mother and the
    interests of the father (in whose interests the child is also acting). The father wants this child
    to be well fed. The mother, on the other hand, thinks that the child is ready to start eating solid
    food, so that she can get pregnant again (perhaps with a different father). However, in a monogamous
    species, the interests of father and mother are the same, so the child is expected to align with
    those common interests.

    John's "accounting rule" that attributes the child's actions to the parents creates a more
    complicated conceptual structure than does Hamilton's method of accounting which has every organism
    (even immature and infertile ones) acting in its own interest. On the other hand, Hamilton's idea of
    "inclusive fitness" is much more complicated than John's. Hamilton needs his fitness to be
    inclusive, otherwise an infertile form would have no fitness.

    Those who are familiar with the philosophical foundations of economics and game theory may worry
    about how a child can be said to balance the interests of both parents, giving equal weight to each.
    In critiques of the ethical doctrine of utilitarianism, for example, theoretical economists object
    to the idea of "equal weight". Those objections may well apply in economics, where utility functions
    are freely chosen. The objections do not seem to apply in evolutionary theory, because Nature is
    assigning the utility function. (PS. If this paragraph makes no sense to you, don't worry about it.
    It is addressed only to people who understand the distinction between cardinal utility theory and
    ordinal utility theory.)

  17. Quoted message said:

    JE:- I would appreciate information re: nest enclosure of stingless bees. Are their nests less
    enclosed than pheromone induced eusocials?

    WH:- Nests of Melipona stingless bees are well enclosed in a fashion similiar to other bees that
    reproduce by swarming. One difference might be the hive size which is 1000-2000 workers in Melapines
    and can sometimes be much larger in other swarming species. In the closely related Trigonine bees as
    in honey bees, female caste fate is nutritionally controlled by workers. Trigonine bees generally
    produce no excess queens.

    JE:- Not surprisingly, many ways seem to have evolved to produce eusocial sterility. The common link
    seems to be nest enclosure to disperse a regulatory pheromone. Nest size would be limited by
    pheromone production and dispersion. Trigonine bees have found another way using pheromone laced
    food. From what you say, it appears Melapines can induce eusocial sterility without the need for any
    pheromone at all so they do not need an enclosed place to disperse any. Thus their nest size could
    be unlimited. However, such a system would be less flexible and unable to meet environmental
    changes. Melapine bees may be more recently evolved because non pheromone induction could have
    evolved from pheromone induction by genetic assimilation. Are 1000-2000 workers within Melapine
    nests a largee or smaller number compared to other species which employ a pheromone induced eusocial
    sterility?

    Many Thanks,

    John Edser Independent Researcher

    PO Box 266 Church Pt NSW 2105 Australia

    [email hidden]

  18. John Edser said:
    Quoted message said:

    JE:- I would appreciate information re: nest enclosure of stingless bees. Are their nests less
    enclosed than pheromone induced eusocials?

    WH:- Nests of Melipona stingless bees are well enclosed in a fashion similiar to other bees that
    reproduce by swarming. One difference might be the hive size which is 1000-2000 workers in
    Melapines and can sometimes be much larger in other swarming species. In the closely related
    Trigonine bees as in honey bees, female caste fate is nutritionally controlled by workers.
    Trigonine bees generally produce no excess queens.

    JE:- Not surprisingly, many ways seem to have evolved to produce eusocial sterility. The common
    link seems to be nest enclosure to disperse a regulatory pheromone. Nest size would be limited
    by pheromone production and dispersion. Trigonine bees have found another way using pheromone
    laced food.


    WH:- As to "pheromone laced food", I don't think so. As in most eusocials, workers are using
    "nutritional control", the amount of food given a larva, as the direct means of controlling caste
    fate. Workers are receiving pheromone signals to guide their behavior but the direct cause of
    sterility is the amount of food a larva receives not a pheromone in the food. In sub-social bees you
    may find direct inducement of sterility via queen pheromones but not in these higher eusocials.
    Somewhat outside this thread but I will note that one should not look at higher eusocials to
    understand how sterility in social insects evolved. One should look at closely related non-social
    and sub-social species to best understand how this transition occurred.

    Quoted message said:

    From what you say, it appears Melapines can induce eusocial sterility without the need for any
    pheromone at all so they do not need an enclosed place to disperse any. Thus their nest size could
    be unlimited.


    WH:- The key pheromone system for determining hive size is not the pheromone signal to make a worker
    or queen but rather the pheromone signal for swarming. This is probably the same in all the swarming
    bees (Melapine, Trigorine and honeybee) and this, at least in part, would be the absense or reduced
    pheromone of the active egg-laying queen. As hive size/density increases each worker receives less
    of this queen pheromone until, at some point, swarming behavior is triggered. After swarming hive
    size is reduced by 1/2 and the process repeats. William L Hunt

    Quoted message said:

    However, such a system would be less flexible and unable to meet environmental changes. Melapine
    bees may be more recently evolved because non pheromone induction could have evolved from pheromone
    induction by genetic assimilation. Are 1000-2000 workers within Melapine nests a largee or smaller
    number compared to other species which employ a pheromone induced eusocial sterility?

    Many Thanks,

    John Edser Independent Researcher

    PO Box 266 Church Pt NSW 2105 Australia

    [email hidden]

  19. Quoted message said:
    Quoted message said:

    JE:- I would appreciate information re: nest enclosure of stingless bees. Are their nests less
    enclosed than pheromone induced eusocials?

    Quoted message said:

    WH:- Nests of Melipona stingless bees are well enclosed in a fashion similiar to other bees that
    reproduce by swarming. One difference might be the hive size which is 1000-2000 workers in
    Melapines and can sometimes be much larger in other swarming species. In the closely related
    Trigonine bees as in honey bees, female caste fate is nutritionally controlled by workers.
    Trigonine bees generally produce no excess queens.

    Quoted message said:

    JE:- Not surprisingly, many ways seem to have evolved to produce eusocial sterility. The common
    link seems to be nest enclosure to disperse a regulatory pheromone. Nest size would be limited
    by pheromone production and dispersion. Trigonine bees have found another way using pheromone
    laced food.

    WH:- As to "pheromone laced food", I don't think so. As in most eusocials, workers are using
    "nutritional control", the amount of food given a larva, as the direct means of controlling
    caste fate.

    JE:- Does this mean that these eusocial lava can be maintained permanently sterile by just limiting
    nutrition? Is this based on calorific and/or protein limitation or micro nutrient limitation? I
    doubt if all eusocials could be maintained permanently sterile by just nutrition so its seems
    reasonable to assume that nutrition has been selected to act as a switch?

    WH;- Workers are receiving pheromone signals to guide their behavior but the direct cause of
    sterility is the amount of food a larva receives not a pheromone in the food. In sub-social bees you
    may find direct inducement of sterility via queen pheromones but not in these higher eusocials.

    JE:- That appears to be in agreement with what I am supposing: non pheromone control of sterility
    evolved from pheromone control.

    WH;- Somewhat outside this thread but I will note that one should not look at higher eusocials to
    understand how sterility in social insects evolved. One should look at closely related non-social
    and sub-social species to best understand how this transition occurred.

    JE:- I agree. Non social and sub social forms, as they exist today, could indicate how eusocial
    forms may have evolved. If eusociality could have evolved without an enclosed area needed to
    disperse a controlling pheromone I predict that eusociality may have become much more common and
    may have included humans. However, if selection must firstly evolve a pheromone stage requiring an
    enclosed nest then it should be rare, as it appears to be today. Raising some of your own
    offspring to become slaves for your own survival is a remarkably good strategy and is allied to
    the mass reproduction of forms with little parental care so that most of them die. Clearly, saving
    some of these immature forms to act as sterile casts instead of letting them all die is a gain for
    the parent.

    Naked mole rats seem to be the only mammal eusocial. Maybe in the future they might assimilate the
    eusociality sterility phenotype so that no pheromone is required and evolve a non enclosed eusocial
    mammal species. Humans use machines as eusocials. Instead of turning all resources into offspring we
    manufacture sterile casts called “machines” that do the same job as sterile eusocials. If the net
    benefit is a rise in absolute parental fitness, eusocial sterility or machine building, can evolve.

    Quoted message said:

    JE:- From what you say, it appears Melapines can induce eusocial sterility without the need for any
    pheromone at all so they do not need an enclosed place to disperse any. Thus their nest size could
    be unlimited.

    WH:- The key pheromone system for determining hive size is not the pheromone signal to make a worker
    or queen but rather the pheromone signal for swarming. This is probably the same in all the swarming
    bees (Melapine, Trigorine and honeybee) and this, at least in part, would be the absense or reduced
    pheromone of the active egg-laying queen. As hive size/density increases each worker receives less
    of this queen pheromone until, at some point, swarming behavior is triggered. After swarming hive
    size is reduced by 1/2 and the process repeats.

    JE:- The pheromone signal for swarming breaks the hive up so that it determines the size of the
    hive? Does this avoid the possibility of two queens within one nest when the nest becomes too large
    to distribute the sterility pheromone?

    Regards,

    John Edser Independent Researcher

    PO Box 266 Church Pt NSW 2105 Australia

    [email hidden]

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