Posts tagged evolution
Altruism, put simply, is helping others or cooperating with others, even if it is costly to self. Of course, something like that cannot evolve, unless there are benefits too, associated with such acts of apparent selflessness.
Cooperation, to start with, can evolve based on three forms of reciprocity: direct, indirect and network. All are based on the fact that there re repeated interactions between group of people- dyads, triads or many more. Reciprocity can typically be measured in the lab using the repeated Dictator/ Trust game.
Direct reciprocity was proposed by Robert Trivers as a mechanism for the evolution of cooperation. If there are repeated encounters between the same two players in an evolutionary game in which each of them can choose either to “cooperate” or “defect”, then a strategy of mutual cooperation may be favoured even if it pays each player, in the short term, to defect when the other cooperates.
Here, in direct reciprocity A trusts/helps B and hopes that when time comes B will reciprocate/help A. The top-of-the-mind factor is whether or not to trust somebody and whether or not to reciprocate someone’s trust. Trust and exploitation may be relevant issues here. In the Dictator/ Trust game this trust/exploitation manifests as the amount that is split and given to the other person vs kept with oneself.
In the standard framework of indirect reciprocity, there are randomly chosen pairwise encounters between members of a population; the same two individuals need not meet again. One individual acts as donor, the other as recipient. The donor can decide whether or not to cooperate. The interaction is observed by a subset of the population who might inform others. Reputation allows evolution of cooperation by indirect reciprocity.
This is partially correct that reputation for being trustworthy helps in indirect reciprocity; however that is only true for the downstream version; for the upstream version feelings of gratitude/happiness/awe/elevation in persons receiving the help/ witnessing the act also lead to more pro-social behavior by those receiving help/ witnessing. Thus feelings of gratitude/ awe/elevation mediate this kind of upstream indirect reciprocity. See below for upstream and downstream variants.
Individual acts of indirect reciprocity may be classified as “upstream” or “downstream”:
Upstream reciprocity occurs when an act of altruism causes the recipient to perform a later act of altruism in the benefit of a third party. In other words: A helps B, which then motivates B to help C.
Downstream reciprocity occurs when the performer of an act of altruism is more likely to be the recipient of a later act of altruism. In other words: A helps B, making it more likely that C will later help A.
Before touching upon network reciprocity, I will take a quick detour about kin selection. I believe kin selection or inclusive fitness is also a type of reciprocity (that between related individuals sharing genes) and may be rechristened genetic reciprocity. After all if A is likely to help B because they share x % of genes, the reverse is equally true and applicable. And of course this is mediated by emotional attachment to the kid/kin.
As per one definition of kin selection:
A biological theory stating that a gene that causes an organism to exhibit behavior detrimental to its survival will increase in frequency in a population if that behavior benefits the organism’s relatives, which will pass the gene on to subsequent generations.
If I slightly change words form above definition, I can now define a neighbor selection process as a cultural theory stating that a meme that causes an organism to exhibit behavior detrimental to its survival will increase in frequency in a population if that behavior benefits the organism’s neighbors, which will pass the meme on to subsequent neighbors.
We are now ready to look at network reciprocity:
Real populations are not well mixed, but have spatial structures or social networks which imply that some individuals interact more often than others. One approach of capturing this effect is evolutionary graph theory, in which individuals occupy the vertices of a graph. The edges determine who interacts with whom. If a cooperator pays a cost, c, for each neighbor to receive a benefit, b, and defectors have no costs, and their neighbors receive no benefits, network reciprocity can favor cooperation.
Basically, what I understand from the above is that if you help your neighbors sometimes such that the cost is not too high but benefits to neighbors are high and if cost to benefit compares favorably with average number of neighbors/ neighborly interactions you have, then in the long run you will benefit and this form of cooperation can evolve. To me the effects are mediated by the number of neighbors or sociability of a person.
Of course, even if you have all these mechanisms in place, cooperation may not evolve, as you may have free-riders. One important mechanism that has evolved to keep the free-riders in check is that of punishment. And once punishment is part of the picture you don’t even need repeated interactions, one-off games may be sufficient. I call this phenomenon Direct Punishment. One way it has been measured is with the Ultimatum game.
In the Ultimatum game, the second player can inflict costly punishment on first player by refusing to accept the division; this costly punishment is dyadic in nature and the aggression/hostility/vengefulness of the second player ensures that cooperation in even one-off encounters happens.
Basically instead of trusting and helping B, A starts by exploiting B and B retaliates by punishing A at cost to oneself.
Of course one can then surmise that there can be a phenomena of indirect punishment. This again may happen in two ways:
- Indirect punishment upstream: A is exploitative in nature; A exploits B; B punishes A, who then feels guilt/ gets reformed and stops exploiting C or even starts helping C.
- Indirect punishment downstream: A is exploitative in nature: A exploits B, B punishes A; B gets a reputation for being tough/competent and stops getting exploited by others say C or C may now even help B.
The Indirect reciprocity effects can be seen in Public goods/ trust game.
I will now take a detour and introduce the HEXACO model of personality which set me thinking about this in the first place.
HEXACO is an alternate personality model that is based on the same principles as the Big Five/FFM; i.e. it uses factor analysis of lexical terms in various languages to arrive at major personality traits.
The six factors are generally named Honesty-Humility (H), Emotionality (E), Extraversion (X), Agreeableness (A), Conscientiousness (C), and Openness to Experience (O). The personality-descriptive adjectives that typically belong to these six groups are as follows:
Honesty-Humility (H): sincere, honest, faithful, loyal, modest/unassuming versus sly, deceitful, greedy, pretentious, hypocritical, boastful, pompous
Emotionality (E): emotional, oversensitive, sentimental, fearful, anxious, vulnerable versus brave, tough, independent, self-assured, stable
Extraversion (X): outgoing, lively, extraverted, sociable, talkative, cheerful, active versus shy, passive, withdrawn, introverted, quiet, reserved
Agreeableness (A): patient, tolerant, peaceful, mild, agreeable, lenient, gentle versus ill-tempered, quarrelsome, stubborn, choleric
Conscientiousness (C): organized, disciplined, diligent, careful, thorough, precise versus sloppy, negligent, reckless, lazy, irresponsible, absent-minded
Openness to Experience (O): intellectual, creative, unconventional, innovative, ironic versus shallow, unimaginative, conventional
The factor H is a new factor not present in Big Five/FFM. The E though looking similar to N of big Five, is conceptually different; it no longer contains anger/hostility which are instead present in HEXACO A. similalrly there are important differences between HEXACO A and Big Five A. the other 3, C, O and X (extarversion) are similarly conceptualized and defined in both systems and have same loadings when tested together.
Ashton and Lee, the proponents of the HEXACO model, have themselves related evolution of altruism to these traits [pdf] and I am building on their work.
Basically as per them,
To begin, we have proposed that the Honesty- Humility and Agreeableness factors represent two complementary aspects of the construct of reciprocal altruism (Trivers, 1971). Honesty-Humility represents the tendency to be fair and genuine in dealing with others, in the sense of cooperating with others even when one might exploit them without suffering retaliation. Agreeableness represents the tendency to be forgiving and tolerant of others, in the sense of cooperating with others even when one might be suffering exploitation by them. (For a discussion of two broadly similar, although not identical, constructs, see Perugini, Gallucci, Presaghi, & Ercolani, 2003.) Presumably, high levels of Honesty- Humility are associated with decreased opportunities for personal gains from the exploitation of others but also with decreased risks of losses from withdrawal of cooperation by others. In a similar manner, high levels of Agreeableness are associated with increased opportunities for personal gains from long-run reciprocal cooperation with others, as well as with increased risks of losses from exploitation by others. (Note that we use the term altruism in terms of a dimension of altruistic versus antagonistic tendency, which involves both a willingness to help or provide benefits to others and an unwillingness to harm or impose costs on others.)In addition, we have proposed that Emotionality represents tendencies relevant to the construct of kin altruism (Hamilton, 1964), including not only empathic concern and emotional attachment toward close others (who tend to be one’s kin) but also the harm-avoidant and help-seeking behaviors that are associated with investment in kin (see also Lee & Ashton, 2004). Presumably, high levels of Emotionality are associated with increased likelihood of personal and kin survival, as well as with decreased opportunities for gains that are often associated with risks to personal and kin survival.
According to Indian Aesthetics , as outlined in Bharat Muni’s Natyashatra, there are eight Rasas or impacts that an aesthetic piece of work like dance form/ literary piece, tries to achieve/invoke in the beholder/reader. These can be loosely thought of as eight different kinds of aesthetic experiences.
As is evident I like all things eight, as I believe they have an underlying eight fold evo-devo stage structure behind them. In this case in visual art from each of the rasa is associated with a predominant color that signifies that rasa. For eg Shringar or Love/beauty/erotic rasa has as its color, color green.
Now, I have also blogged previously about color terms and how they may have evolved in an eight step model. In brief, I believe that Black came first , white next, then Red followed; after which followed yellow, blue and green and then other terms like orange, brown and grey followed. This I believe is also tied to the way our color vision and sensitivity would have grown/evolved.
Taking cue from that color term evolution model , I belive the eight rasas evolved in the same order as the color used to denote them, and that nicely fits with other aspects of the eight stage theory too, like the eight stage emotion development.
So, in essence, please find listed below the eight Rasaas in the correct eight stage order:
1. Bahyanak Rasa: color Black : emotion fear.
2. Hasya Rasa: color white: emotion joy/laughter
3. Rudra Rasa: color red : emotion anger
4. Adbhuta Rasa: color yellow , emotion wonder
5. Bibhitsa Rasa: color blue, emotion disgust
6. Shringar rasa: color green; emotion erotic/love
7. Veera rasa : color orange; emotion heroism
8. Karuna rasa: color grey; emotion compassion/ sadness.
Contrast this with the basic emotions list I have come up earlier viz.
1. Fear – Bahyanak
2. Joy/happiness – Hasya
3. Anger – Raudra
4. Sadness – Karuna
5. Disgust – Bibhitsa
6. Surprise – shringar
7. Contempt – Veera
8. Interest (anticipation) – Adbhuta
With just one switch between the Adbhuta (wonder/interest) and Karuna (sadness/ compassion), the eight rasas fit both the color term evolution and basic emotions evolution. That surely means we are at something and moving in the right direction as different routes are leading to the same destination. .
I have also advocated the four primary problems faced by all creatures undergoing evolution, as delineated by Theodore Millon– the problems of Existence; Adaptation; Replication and Abstraction which lead to polarities of pain/pleasure; active/passive; self/other and broad/narrow at each of the stages/domains/ solutions.
However, when we pause to look at what the mechanism of evolution actually is, we clearly note that there are a few prerequisites for evolution to take place and unless all the four mechanisms/ preconditions are present it is unlikely that the creatures will evolve. I have been having this in the back of my mind for quite some time especially as I have been ruminating on the BVSR (blind variation and selective retention) theory of Donald Campbell as applied to creativity.
I was recently reading ‘Driven’ and in that book too a lot of emphasis is placed on the V-S-R (Variation, Selection, Retention) mechanism of evolution. I think this popular portrayal of evolution misses another important ingredient required for evolution that of Struggle due to limited resources and excess fecundity. If the problem of limited resources and excess fecundity was not there, probably there would be no pressure to evolve.
Thus I would like to frame the four evolutionary prerequisites/ mechanisms as Struggle-Retention- Variation-Selection or S_R_V_S. To elaborate:
1. Struggle: This is driven by the fact of limited resources and overproduction due to excess fecundity. Here two strategies, leading to 2 polarities, work; one can either compete for resources or one can cooperate and exist symbiotically. Also, one can either have a r-strategy (low investment in many) of reproduction or A K-strategy (high investment in few). In the eight stage model, the organisms at first 2 levels would be primarily constrained by this evolutionary mechanisms.
2. Retention: There must exist some mechanism by which the traits that confer survival/reproductive or selection advantage can be retained over time in the same individual and over generations in the same species. Again the mechanism of stable trait over time as well as over generations may be in conflict with each other and may lead to a polarity. Creatures at stage 3 and 4 of 8 stage evo-devo theory would likely face issues regarding stability and retention of traits; retaining in individual the same trait is an active process; while retaining in generations is more passively driven.
3. Variation. There must exist some mechanism that causes minor changes in the stable traits such that variation may lead to deleterious or beneficial effect over the individual having that variation. This is classically implemented using mutations and sexual-recombinations. While mutations confer (dis)advantages at an individual level; recombinations take that to the next level by affecting offsprings variability. The creatures at stages 5 and 6 of evo-devo stages are grappling with these problems of adequate variation in self and other.
4. Selection: There must exist some selection criteria based on which the struggling creatures having stable but slightly varying traits can be selected for or against. While Natural selection (stage 7) employs the three methods of directional, stabilizing and disruptive selection; the stage 8 deploys a qualitatively different method of sexual selection where the criteria for selection may be arbitrarily driven by choices of the other gender conspesifics. While the Natural selection criteria is broad, the sexual selection criteria can be said to be narrow. Another way to look at this is that the selection crteria is either to survive (natural selction) or to reproduce (sexual slection) and those who are sucessful can very well move from one level/species to another (speciation due to sexual selection).
To me this is further corrobrating evidence of the eight stage evo-devo theory and ABCD model being on the right track.
This post is an ambitious attempt to link the four causes delineated by Aristotle, to the four questions asked by Tinbergen to the four types of explanations that can be made for any human/animal ability.
First a bit of a background.
Aristotle had listed four causes – Material, Efficient, Formal and Final causes. From the Wikipedia:
Aristotle held that there were four kinds of causes:
- A thing’s material cause is the material of which it consists. (For a table, that might be wood; for a statue, that might be bronze or marble.)
- A thing’s formal cause is its form, i.e. the arrangement of that matter.
- A thing’s efficient or moving cause is “the primary source of the change or rest.” An efficient cause of x can be present even if x is never actually produced and so should not be confused with a sufficient cause.(Aristotle argues that, for a table, this would be the art of table-making, which is the principle guiding its creation.)
- A thing’s final cause is its aim or purpose. That for the sake of which a thing is what it is. (For a seed, it might be an adult plant. For a sailboat, it might be sailing. For a ball at the top of a ramp, it might be coming to rest at the bottom.)
While studying these causes scholars have stressed on two sorts of distinctions, the first is the concept of actuality and potentiality– while some causes like the final cause (teeos or entelchiea) and formal cause (eidos or energiea) are actual; the material cause is deemed to be poetntial.
The other concept is that of Hylomorphism, or the distinction between matter and form. I believe that the four causes can be broken down along the two dimensions of potentiality/actuality and matter/form as below to lead to a 2×2 matrix of causes.
However, I like to arrange it slightly differently. I retain the ultimate/proximate distinction, but supplement it with Accidental/ Teleological distinction. Accidental to me means phylogeny due to random genetic drift and genetic baggage; and physiologic mechanisms like hormones etc kicked in accidentally by random interaction with immediate environment; on the other hand the process of adaptation and developmental unfolding to an adult form (ontogeny) appear purposeful and pseudo teleological. Thus my arrangement:
Which leads to how I became re- interested in these four causes recently- it was while reading an article by Dean Simonton on ability , wherein he conjectured that studying both the generic and specific factors that affect performance as well as acquisition of ability is problematic and non-fruitful, that I remembered about these and saw how fruitful it may be to conceptualize ability in just so many terms. Hence my conceptualization:
In general for ability I think the following four factors are applicable and all four have strong influences on ability; I have named them talent, hard work, grit and optimism etc elsewhere. In general for any trait like intelligence, emotionality etc I think the 2×2 factor matrix is relevant and worth keeping in mind.
That brings me eventually to my own levels of explanations for any phenomenon. I believe evo-devo explanations as well as psycho-social factors have equal weight-age while explaining say behavior. while some of these explanations are mechanistic/deterministic other are of non-deterministic or chaotic origin. Similarly, while some are governed by factors internal to the organism others are mostly affected by extrinsic factors.
I hope the above conceptualization makes sense. It is inline with my eight stage model and four domains model where Evo is stage 1&2; social- stage 3&4, devo- stage 5&6 and psycho stage 7&8.
I started the emotion and personality series with a focus on the eight stage model and how that informs us about both personality and emotions. I ended up changing tracks and pursuing Millon’s evolutionary stages and polarities and extending it to the ABCD of four broad psychological domains. Avid readers will notice that both my eight stage model and the Millon’s stages/polarities are based on evolutionary considerations and thus there should be neat synthesis involving the two. this post is an attempt to do so under the framework of the four basic domains of psychology : the ABCD model of psychology.
To recap: all psychology basically can be broken into study of :
- Affects or emotions
- Behaviors or social factors
- Dynamics or drives and motivations and
- Cognition or learning, memory etc.
There have been paradigms in psychology like affective psychology, behaviorism, psychodynamcis and cognitive psychology that have focused on one domain more than the other. One can even extrapolate the diffrent approaches and discplines that are releavnt to each domain as below:
- Affect/emotions: Evolutionary explanations as most emotions are evolved mechanisms. Biological context is relevant. Genes drive this.
- Behaviors/social functioning: Social/situational explanations dependent on interactions with other people/cultural effects. Cultural context is more relevant. Environment drives this.
- Dynamics/ drives/motivations: Developmental explanations of how life course and needs/drives interact. Ecological context is instrumental. Environment chooses genes here and only certain phenotypes expressed.
- Cognition: economic/information-processing explanations that explain decisions, perceptions etc. Constructed and created context is important here. Niches are build and genes choose environment.
The Affect and Dynamics can be clubbed together as Evo-devo explanations and refer to subjective phenomenon; while the Behavior and Cognition can be clubbed together as Socio-Economic explanations and refer to objective observable phenomenons. But anyway this is digressing a bit from the main topic.
Getting back to topic at hand, the four domains correspond to the four evolutionary stages of millon and each stage has two polarities and thus map to eight stage model as follows:
- Affect maps to problem of Existence and the polarity of pain and pleasure mapped respectively to say the ‘FEAR’ and ‘SEEKING’ system of Pankseep. These (the panksepp emotions systems) I have previously shown how they are mapped to the eights stage model.
- Behavior maps to the problem of Adaptation and the polarity of Active and Passive mapped respectively to the ‘RAGE’ and ‘PANIC’ systems of panksepp.
- Dynamics/Drive maps to the problem of Replication and the Polarities of Self and Other mapped respectively to ‘LUST’ and ‘CARE’ of Panksepp.
- Cognition maps to the problem of abstraction and the polarities of Broad/creative versus narrow/rigid and maps respectively to ‘PLAY’ and ‘SELF’ systems of Panksepp.
Seen form this angle, the eight stages are just the eight polarities manifesting one after the other in the developmental course.
Extending Millon’s evolutionary considerations,it behooves to remember that another way to look at his problems is to view the change sin phylogeny, ontogeny , function (ultimate) and causation (proximate) of any trait.
Phylogeny is dependent on historical environment and leads to the trait having its ultimate thematic value. This I argue is the problem of existence (of trait/individual) and manifests as the domain of Affect.
But a trait with a fixed value would be of no use. Around the ultimate thematic value there will be ultimate variation that charts the possible functional map of what that feature is supposed to do. The abstracted ancestral environment (EEA) is teh context in which function evolves. This is the problem of abstraction and manifests as domain of cognition,.
Apart form the ultimate thematic value, one needs to tune that value to the immediate ecological and developmental context. Ontogeny is dependent on such an ecological context and is a proximate thematic mechanism that leads to a particular stable thematic value of a trait. This is also the problem of Replication (r-K) and manifest as Dynamics.
However, even a fixed proximate thematic value of trait will be useless as situations keep changing. Causation is responding to immediate environment in appropriate and adaptive manner. This is the problem of adaptation and manifests as domain of behavior.
I can relate the above to Aristotle’s four causes, but will leave that for another day; time now to wrap up the personality part. Emotions we have seen can be easily subsumed under the ABCD domains of psychology. As Pankspess model has been related to cloninger personality traits, I’ll leave the case rested that personality can also be adequately explained using the ABCD construct.
Before signing off, I’ll hastily note that to me, these ABCD domains map to underlying neurotransmitter systems:
Affect (pain/pleasure/’FEAR’/’SEEKING”) is associated with Noradrenaline system (NE). the role of ?NE in FEAR is clear, but for SEEKING DA also seems to play a role.
Behavior(active/passive/’RAGE’/’PANIC’) is associated with Serotonin system (SE). The relationship of Serotonin with aggression and separation distress as well as behavioral manifestations like eating/sleeping cycle are well known. Known abnormalities in SE also cause OCD/Panic attack etc. Known abnormalities lead to fatigue, lethargy etc.
Dynamics (self/other/LUST/CARE) is associated with Dopamine (DA). Known abnormalities in DA cause motivational obscurity like paranoia/psychosis.
Cognition (creative/rigid/’PLAY'”/SELF) is asscoited with Acetylcholine (ACh) and known abnormalities in ACh here lead to Alzheimer etc.
That shall be all for now.