Group, individual, or gene level
selection: Which has the best explanatory power for our ‘moral’ emotions?
Mark Sloan 12-27-2014
Charles Darwin
proposed group selection to explain how “altruistic” behaviors toward others
and self-sacrifice for the group, which could easily be taken to be
counterexamples to his theory of evolution, might instead be examples of its
explanatory power. He observed that, all else being equal, groups that were
more cooperative would out compete less cooperative groups. Selection for more cooperative
groups selects for biology that better motivates cooperative behavior (Darwin,
1871) and, though Darwin did not mention it, cultural norms that do the same. Thus,
we arrive at the simple and appealing notion that people’s experience of
compassion and loyalty and cultural norms such as “Do to others as you would
have them do to you” (or variations with the same function in increasing
cooperation) are arguably near inevitable products of blind evolution of beings
with sufficiently complex brains. And no supernatural deity is required to
explain human, or any other species’, “goodness”.
Mark Borrello (2005)
described group theory’s subsequent rise and then fall from respectability in
the 1960’s (mainly due to a theoretical objection and some bad science from a chief
advocate) and its rise again, starting in the 1980’s and now called multilevel
selection, which is how I will refer to it here where appropriate.
Steven Pinker and
twenty three other respected evolutionists provided an accessible snapshot (Pinker
2012) of the lively state of the unresolved debate on group selection’s proper
role in evolutionary science. The diversity of pro and con opinions expressed
makes it appear the debate about group selection is not close to over.
However,
Pinker’s focus on “altruistic” and “self-sacrifice for the group” behaviors as
“what must be explained” was a poor choice for resolving the relative
explanatory power of group versus individual and gene level selection
perspectives. First, as they are commonly understood, these behaviors can have
many explanations which make them insensitive tests for differentiating between
hypotheses. Also, these explanations included cultural norms and forces, which
divert attention to a secondary issue, group selection’s role in cultural evolution.
Finally, Pinker appeared to be taking “altruism” to mean something very close
to “biological altruism” (behavior by an individual that increases the lifetime
fitness of others while decreasing the lifetime fitness of the actor). If
what-must-be-explained is biological altruism, his argument is almost won
before he starts. Without the requirement to explain cooperation behavior, the
most credible candidate is kin altruism, the obvious choice from the gene level
selection perspective.
To avoid these
and other problems, future group level selection discussions might be made more
fruitful by choosing human cooperation as what-must-be-explained. Also, making
certain stipulations will clarify the discussions and avoid other above described
problems. I propose the following:
1) Multilevel
selection’s explanatory power for biology will be evaluated separately from
culture, requiring what-must-be-explained to be chosen to insure only biology
is needed to explain it.
2) Multilevel,
individual, and gene level selection are different perspectives on (or
bookkeeping systems for) the same phenomena. In theory, they might all have the
same explanatory power.
3) To be a
useful part of an evolutionist’s toolkit, multilevel selection’s perspective must
show better explanatory power in practice, meaning ability to more readily
produce correct explanations of specific biology.
4) A hypothesis’
“in practice” explanatory power may be inferior to its “in theory” explanatory
power.
5) Multilevel
selection’s “in practice” explanatory power will ideally be evaluated for
multiple phenomena, not just one. Requiring explanation of multiple phenomena
increases discriminatory power between alternate hypotheses.
I expect these
stipulations would be acceptable to many in the field. If they are not
acceptable, perhaps discussions could clarify what will be stipulated and any
dissenting opinions.
The biology
underlying what makes human beings the amazingly cooperative social animals we
are, and arguably the heart of what makes us ‘human’, provides two candidates
for what-must-be-explained.
The first candidate is our moral emotions triggered by our moral sense that, in part, motivate unselfishness. For reasons described below, I will here take these to be compassion, loyalty, gratitude, anger, disgust, contempt, shame, guilt, and ‘elevation’. (‘Elevation’ is an emotion encompassing feelings of satisfaction and pride.) Note that the existence of the emotions triggered by our moral sense is not culturally dependent. That is, the existence of these emotions does not vary from culture to culture. Only the circumstances when they are triggered and their intensity with which they are triggered are culturally and individually dependent.
The second
“cooperation biology” candidate is the cross-cultural universal set (and
therefore likely biological, but not certainly biological) of categories of circumstances
which can trigger our moral sense to make one of its near instantaneous
judgments (and in turn trigger our ‘moral’ emotions). The most commonly
referred to version of Moral Foundations Theory identifies five cross-culturally
universal categories: care/harm, fairness/cheating, loyalty/betrayal,
authority/subversion, sanctity/degradation (Graham 2012).
Another
candidate for what-must-be-explained is the evolution of cancers with their
intrinsic complexities of simultaneous cooperation and competition at the
cancer’s gene, cell, and cell strain levels in the hostile environment of the
patient’s immune system. Due to the specialized knowledge required, such group
selection debates would likely be most effective among experts.
However, it may be that such a debate is not needed. Researchers in the field use group selection perspectives as standard practice (Merlo, 2006). It seems unlikely that any debate among those experts would conclude using the group selection perspective is not useful in actual practice.
However, it may be that such a debate is not needed. Researchers in the field use group selection perspectives as standard practice (Merlo, 2006). It seems unlikely that any debate among those experts would conclude using the group selection perspective is not useful in actual practice.
There are
doubtless other candidate biological phenomena that could be useful choices for
what-must-be-explained.
The following
is an example of how the character of group selection discussions could change
if what-must-be-explained was human cooperation and the above stipulations were
accepted. In the interest of providing a brief example, I will only compare the
“in practice” explanatory power of multilevel and gene selection perspectives for
our ‘moral’ emotions, the emotions triggered by our moral sense.
Drawing on
many sources, Johnathon Haidt argued moral emotions include compassion,
gratitude, anger, disgust, contempt, shame, embarrassment, guilt, and
‘elevation’ (Haidt 2003). I largely agree, but add ‘loyalty’, meaning the
emotion that motivates “loyal” behavior, since unselfishness motivated by
loyalty is a central part of morality. I prefer the word indignation to anger
because “indignation” better captures an emotion that is intrinsically about
right and wrong. Also, I would omit the emotion “embarrassment” since
embarrassment can be about matters having nothing to do with right and wrong,
and shame and guilt adequately convey this category of moral emotions. For
these reasons, I will use explanatory power for compassion, loyalty, gratitude,
anger, disgust, contempt, shame, guilt, and ‘elevation’ as what-must-be-explained.
If our moral
emotions are components of cooperation strategies, what necessary functions are
they fulfilling?
Tit-for-tat2
is the simplest highly effective cooperation strategy known in game theory. It
seems almost inevitable that, if evolution did encode cooperation strategies in
our moral sense, something like tit-for-tat would be one of those strategies.
Tit-for-tat has two necessary behavioral components: 1) cooperation by
unselfishly risking exploitation in any first interaction and also if the other
individual cooperated in the previous interaction and 2) retaliation if the
other individual attempted to exploit unselfishness in the previous
interaction.
However, this
simple tit-for-tat strategy is not always the best, and what is better varies
with circumstances. Perhaps sometimes there are ways to predict who is likely
to be a good cooperator and who is likely to exploit you – for instance from
gossip about other’s interactions with the individual. Perhaps ‘forgiveness’ of
an instance of exploitation could avoid cooperation destroying endless cycles
of retaliation.
How might
evolution most efficiently encode motivation for people to develop more
effective variations of tit-for-tat and even switch between them in the many
different circumstances encountered over a lifetime, and sometimes even over
the course of a single day – as we know people readily do? If a mutation
produced a psychological reward for achieving cooperation, then that mutation
could be selected for because it motivates the individual to find those
strategies that are more effective variations of tit-for-tat and to switch
between them according to circumstances.
So if
evolution did encode cooperation strategies in our moral sense, the simplest implementation
that would be effective in diverse circumstances would have three necessary
components: 1) motivation for unselfish behaviors, such as helping actions,
which can initiate cooperation but risk exploitation, 2) motivation for
retaliation against exploiters, and 3) psychological rewards for achieving cooperation.
This is exactly
what we find in our moral emotions. 1) Compassion, loyalty, and gratitude
motivate unselfish behaviors that initiate or maintain cooperation. 2)
Indignation, disgust, and contempt motivate behaviors that punish other’s
exploitation of unselfishness; shame and guilt internally punish our own
selfishness. 3) The emotional experience of ‘elevation’ rewards our own
successes at cooperation, motivates imitating other’s successes, and encourages
staying in cooperative groups and seeking to increase cooperation within those
groups.
In contrast,
consider the gene level selection perspective’s most common explanation of
unselfish behavior toward non-kin (and moral behavior in general): that unselfishness
toward non-kin is best explained as a product of kin altruism and cultural
influences (Pinker 2012). How well does that explanation explain the existence
of our moral emotions?
Kin altruism
readily explains the existence of compassion and loyalty toward close kin and the
reward of ‘elevation’ for cooperation with close kin. While speculative, the
argument that these emotions are triggered toward non-kin due to a kind of error
in kin detection is possible.
But what about
the other six: gratitude, Indignation, disgust, contempt, shame and guilt? How
are these explained by kin altruism? They appear to have nothing to do with kin
altruism. I expect clever people can find arguments that all these emotions can
be explained by kin altruism. But inventive speculations about how they might
be explained as products of kin altruism fall far short on the “best
explanation scale” compared to multilevel level perspective’s robust explanation.
Based on
relative explanatory power, we can conclude that the majority of our moral emotions
are elements of reciprocity cooperation strategies, not kin altruism.
Does this failure
to explain our moral emotions mean that gene level selection cannot, in theory,
explain all gene determined biology? Again, no, it simply means that the gene
level perspective can, in practice, incline investigators to produce false
explanations of biological adaptations.
Multilevel
selection robustly explains the set of emotions that are motivated by our moral
sense. Gene level selection’s common claim about unselfishness (and moral
behavior in general) toward non-kin being kin altruism misapplied cannot directly
explain two thirds of the emotions motivated by our moral sense. Based on its
poor explanatory power, this common claim, which flows naturally from the gene
level selection perspective, appears false. Gene level selection’s perspective appears,
in practice, to have led Pinker and other gene-level-only advocates astray.
In practice, it appears that the
multilevel selection perspective produces correct explanations of our moral
emotions more reliably than gene level selection. Therefore, the multilevel
selection perspective deserves an important place in the evolutionist’s tool
kit.
Notes:
1.
Unselfishness (including “altruism” and “self-sacrifice for the group”) is here
defined as accepting a cost in order to intentionally benefit others without
consideration of possible future benefits.
2. Tit-for-tat is a highly effective, but
extremely simple, cooperation strategy in game theory (Axelrod, 1984). An agent
using this strategy will first cooperate, then subsequently replicate an
opponent's previous action. If the opponent previously was cooperative, the
agent is cooperative. If the opponent previously tried to exploit the agent,
the agent will retaliate by attempting to exploit the opponent.
References:
Axelrod, David, (1984). The Evolution of
Cooperation.
Borrello, Mark E., (2005). The rise, fall
and resurrection of group selection.
Darwin, Charles, (1871). The Descent of
Man.
Grahama, Jesse, Haidt, J., et al. (2012).
Moral Foundations Theory: The Pragmatic Validity of Moral Pluralism. Available at http://ssrn.com/abstract=2184440
Haidt, J. (2003). The moral emotions. In
R. J. Davidson, K. R. Scherer, & H. H. Goldsmith (Eds.), Handbook of
affective sciences. Oxford: Oxford University Press. (pp. 852-870).
Merlo, Lauren M.F., Pepper, John W., et
al. (2006). Cancer as an evolutionary and ecological process.
Pinker, Steven, et al. (2012).The False Allure of Group Selection - An Edge Original Essay with Replies.
http://edge.org/conversation/the-false-allure-of-group-selection
Mark Sloan’s personal interest is in
showing how understanding morality as an evolutionary adaptation can be
culturally useful. As associate editor of This View of Life, Morality section, his
goal is to promote a cross-disciplinary view of the science of morality and how
that science could be culturally useful. He has degrees in engineering and
physics and has had a career in the aerospace industry.
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by Mark Sloan – All Rights Reserved – No Reproduction or Publication for
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