Tag Archives: evolution

The Evolution of Cooperation: Thankfulness and Inspiration as Two Routes

Cooperation or Altruism or Prosociality is the tendency to help others, even at a cost to oneself. Naive conceptions of evolution, make us think that cooperation or altruism cannot evolve because the genes are selfish and only care about perpetuating themselves. However, the selfish gene view of evolution does not preclude organisms to become altruistic if for example they share genes ; one of the mechanisms for the same is kin selection.  

Another mechanism that can give rise to cooperation is direct reciprocal altruism. In colloquial language, it is akin to you scratch my back and I’ll scratch yours.  You help someone in their time of need and expect that the person will help you back in your time of need. Consider our hunter/ gatherer ancestors; if one of them made a big kill and shared with a con-specific, it becomes more likely that the peer will share with him when good fortune shines on the peer.

A necessary condition for reciprocal altruism to evolve is the ability to punish if someone doesn’t reciprocate or at least to not keep trusting that freeloader in future. Both punishment as well as not indulging with that person again or repaying a past benefactor, depends on the ability to remember the person whom who we have helped or who has helped us. This ability will typically involve remembering faces. Another ability that will come handy is the ability to gauge intentions- whether the person who helped did it on purpose or accidentally.

Yet another mechanism that gives rise to cooperation is indirect reciprocity. I help you and you help someone else. While in direct reciprocity, a pair or dyad helps each other over time or helps now with the hope/ expectation of a payoff in the future; in indirect reciprocity, one helps a stranger just because one has been helped by someone. Indirect reciprocity works by creating a culture of altruism, where helping becomes the norm and people accumulate reputations.

Now, direct reciprocity can work by encouraging feelings of gratitude in a beneficiary. These feelings of thankfulness and indebtedness towards the benefactor , act together to ensure that the person receiving favors, returns them.

Indirect reciprocity works similarly, by promoting feelings of elevation or inspiration in the general public who may be recipient or even just witnessing a virtuous act of kindness, morality etc. . Once such feelings are aroused, the person witnessing such acts become more likely to act prosocially.

The above may seem speculative, but there is solid evidence around the evolution of direct reciprocity as well as indirect reciprocity. and so too is there strong linkage between direct reciprocity and gratitude. The link between indirect reciprocity and elevation is also hinted.

For example, consider the following article by Sui et al that shows that trait gratitude and trait elevation have different neural correlates. From the abstract:

We demonstrated that trait gratitude was positively correlated with gray matter volume (GMV) in the left cerebellum extending to fusiform gyrus, and also the right middle occipital gyrus (MOG) extending to posterior superior temporal sulcus (pSTS) and temporoparietal junction (TPJ), while trait elevation was negatively correlated with GMV in the left dorsolateral prefrontal cortex. While controlling each other, all the regions still held significant, except the right MOG and pSTS/TPJ. The results indicate that there are distinct neuroanatomical correlates for proneness to gratitude and elevation, while the evidence is mixed that pSTS/TPJ may be the common correlates for them.

The authors discuss the implications of above: gratitude  is associated with more GMV in fusiform gyrus, an area important for remembering faces, and is thus the emotion associated with direct reciprocity where remembering faces is important. Also, both gratitude/ elevation are associated with  pSTS and TPJ, areas important for attributing and inferring intentions to others. Elevation further is negatively associated with GMV  in left DLPFC, a region that involves thinking in a utilitarian manner rather than in intuitive manner about moral issues. As per dual process theories of morality one can either take a intuitive, de-ontological stand where a moral act is moral because it is the right thing to do or a duty to uphold; or the second pathway consists of more deliberate, utilitarian reasoning whereby one looks at acting such that happiness/utility is maximized for maximum number of people. Trolley problems are famous examples of such utilitarian reasoning battling with moral intuitions. 

 To me the differential neural correlates of gratitude and elevation, as well their different manifestations at the emotion and behavioral level, strongly suggest that they are associated with different types of altruism- direct and indirect respectively.  

What Good (and Bad) are Positive Emotions?

There is a seminal article by Barbara Fredrickson titled ‘What good are positive emotions?’ which introduces the broaden-and-build theory of positive emotions. As per this theory, while negative emotions are associated with specific action tendencies, positive emotions broaden the thought-action repertoire available at that moment and help build physical, social and intellectual resources over the long run.

To take an example, joy is associated with creativity, and more loose associations, etc, all involving a move away from rigidity and fixedness to flexibility and fluency in thinking ;  joy is also associated with rough-and-tumble play in many species, including humans, and this play although apparently without any purpose, prepares the young ones for avoiding predators later on. Thus, being in a state of joy over time prepares/ enables one to take care of an important evolutionary problem – that of avoiding predators.

However, there is a downside to extreme continued state of joy or euphoria- Mania that is marked by energy (needed for play), paranoia (predator avoidance and suspiciousness kicking in) and other characteristics like racing thoughts and verbal diarrhea (creativity and loose associations gone haywire). Some of the features of this syndrome (like paranoia) can only be understood when one applies the evolutionary lens to the phenomena.

I have listed elsewhere the major adaptive problems faced by all humans in the EEA, and will like to use as an example the first two problems: Avoiding predators and finding food (by hunting/ gathering).

One may surmise that not being able to avoid predator and getting eaten partly or wholly is painful and distressing and that these emotions play a great part in ensuring survival in the short term.  In the long term however if one has to avoid predator one needs to rehearse in peace time to be prepared for the show when one encounters the predator next. Joy signals such a peaceful state where one can indulge in rough and tumble play and practice predator avoidance tactics and strategies.

Now how is the state of joy induced? Once the organism is not in a state of pain or in a state that can lead to more pain, it can focus on hunting/ gathering to get food and become string and heal. Eating such food leads to pleasure/joy and the cycle repeats.

….joy leads to play leads to predator avoidance leads to less or no pain leads to hunting/gathering leads to eating food leads to pleasure joy leads to ….

Thus the positive signal of Joy/pleasure moves the organism towards food (other consummatory rewards)  while the negative signal of pain/ distress makes the organism avoid predator and move away from such in danger states.

Normally, these work in perfect balance, but if pain/distress takes over one loses the drive to hint/ gather and may suffer from depression; if pleasure/ joy takes over one is too focused on avoiding (imaginary) predators that one may become manic (paranoid type).

The natural question arises, does this analysis apply to other emotions too? Below I try to list the cycles for each emotion (two emotions form a pair and work in conjunction)  along with the adaptive problems they solved in EEA.

  1. Predator Avoidance: failure of predator avoidance leads to pain/distress. When there is no pain/ distress (and one has healed) one hunts/ gathers.
  2. Hunting/ gathering food: eating food leads to pleasure/joy. Joy leads to rough and tumble play which preparers one to avoid predators in future.
  3. Nature’s fury (unpredictability/ novelty ) avoidance: failure of not being able to predict weather or encountering a new phenomenon etc lead to fear/ guilt. When there was no fear or guilt and conditions were stable one explored the world.
  4. Exploration of the world/ environment: Encountering novel phenomenon leads to interest in such phenomenon. Interests leads to learning maybe via nighttime dreams where one subconsciously elaborates the cognitive maps and explores the dark corners including nature’s fury and tries to understand them so as to avoid them in future. The main function involving fear/ interest here is learning about the good and the bad stimuli in the environment.  When fear takes over it manifests as phobias, when interest takes over it manifest as obsession and compulsion (rituals to avoid unpredictability?).
  5. Misunderstanding avoidance (theory of mind) : failure of this leads to misunderstanding among the dyad and leads to anger/ aggression etc. When there is no anger / aggression one indulges in communicating with the other and build bridges of understanding.
  6. Communicating with others: When one feels listened to and understood, one feels love and compassion for the other. Love leads to social play (say flirting) that ensures that you have a better grasp on what the other person is thinking. The love/ anger cycle is responsible for ensuring proper bonding and communication between con-specifics. When anger takes over you have antisocial/ conduct disorders; when love takes over you may have dependence/ addiction.
  7. Incest Avoidance: failure of incest avoidance led to disgust. When the ancestral human was disgust free he mastered the art of remembering faces so as to distinguish kin from non kin.
  8. Face recognition: When one recognizes a familiar face/ pattern, one is in awe or get elevated.  Being in Awe one experiments with morality and builds character which leads to incest avoidance.  this system of disgust/ Awe has been co-opted for morality. When disgust takes over you may have dissociation.
  9. Wrong mate avoidance (mate selection): Selecting the right mate is very important. Failure to do so leads to envy/jealousy. When one is free of envy/  jealousy one can take care of the marriage- by investing in kids and relatives.
  10. Helping children and kin (Parental investment):  When one is helping one’s children and kin, one feels the contentment of merely being able to serve them. Contentment leads to savoring where one enjoys the time together with the same mate; the actions you take cements the relation and leads to less chances of mate selection being bad.  This cycle is about family systems.
  11. Cheating avoiding (cheater detection): In altruistic and social beings, detecting cheaters is very important. Failure to detect them timely leads to feelings of dissatisfaction/ vengeance. When such feelings are lacking one can make new friends and alliance partners.
  12. Making new friend and alliances: When one makes new friends and alliance partners one feels gratitude towards them.  Gratitude leads to reciprocal plays like repeated prisoner’s dilemma, where you learn not to get waylaid by cheaters and can avoid them in future.

Although some of the above is speculative, I as usual am very much excited by the above framework- its makes a number of empirical predictions that can be easily tested. What disorders do you foresee from the excess of both positive and negative emotions? Do let me know via the comments!

Evolution of the 4 Emotion Dimensions

PAD is a popular dimensional theory of emotions, whereby all emotions can be classified on three dimensions: Pleasure (Pleasant- Unpleasant), Arousal (Ready-Relaxed), and Dominance (Control- Lack of control). To this model has been added a fourth dimension called Predictability (Ambiguous- Certain) (please see my earlier post for why this fourth dimension is relevant).

Manga emotions

Manga emotions (Photo credit: Wikipedia)

As an example, anger and fear are both unpleasant emotions, but angry person is in control (has high dominance) while a fearful person is not in control of the situation.

Similarly, both contentment and excitement are pleasant emotions, but the former is low arousal and the latter high arousal.

Thus, emotions differ on four basic dimensions. I’ll address each of these dimensions below:

Pleasure (pleasant – unpleasant). This is similar to pleasure-pain polarity as highlighted by Millon and works at the Affective level in the ABCD model. The pleasure polarity addresses the physiological needs (Maslow’s hierarchy, see here) for maintaining body, while the pain polarity ensures that we stay out of harm’s way and take care of our safety needs. If one were to measure well-being related to this dimension, the appropriate measure would be something like PANAS, a difference between your positive affect and negative affect. In a nutshell, this is characterized by in-the-moment feelings and if your needs are met here, you live a happy life in the hedonic sense. The existential challenge would be body-givennenss and what to do given the body- a potential answer being – survive and protect body integrity.

This dimension, related to feelings, may have evolved to help our bodies/genes survive. If something leads to unpleasant emotions, avoid it; if it leads to pleasant emotions, indulge and approach!


Arousal (Ready-Relaxed): This is similar to the active-passive polarity as highlighted by Millon and works at the Behavioral level of ABCD model. The active polarity, which is related to being excited/ inspired, addresses the self-actualization needs of Maslow’s, while the passive polarity, which may be related to tranquility/ calmness/ meditation etc is related to transcendence needs.  If one were to measure well-being related to this dimension, the appropriate measure would be something like Ryff’s Psychological well-being measuring things like psychological growth etc.  This dimension may be related to living authentically in this world and experiencing life to the fullest. The existential challenge relevant here would be how to cope with a meaningless/ absurd world. The answer may lie in living life fully and experiencing it deeply.

This emphasis on experience is related to the ‘experience’ component of consciousness (recall that consciousness is made up of two parts- experience and agency).  This dimension of emotion, related to energy/ experience, may have evolved to give the emotion a vibrant and vivid tone.  That vibrancy may be required if after gaining a  mind, we can retain it i.e. remain sane.


Predictability (Ambiguous- Certain): This is similar to the broad-narrow polarity as highlighted by Millon, and works at the cognitive level of ABCD model. The broad polarity, which is ambiguous and amorphous, is related to the Aesthetic and beauty needs of Maslow, while the narrow polarity to Knowledge and understanding needs. If one were to measure well-being related to this perhaps Satisfaction with Life survey might work. This dimension is related to attracting mates (both beauty and brains are attractive) and perhaps reproduction. The existential challenge here may be Death and the answer may be transcending death by leaving progeny. Cognitive abilities allow one to reflect on one’s own death and this leads to obsession with procreation. Intelligence (and beauty) associated with this dimension may be a result of sexual selection.

This dimension, associated with intelligence, may have evolved to help out bodies/ genes replicate. To figure if a mate is the best possible mate, and to attract / coax it, one may need intelligence and beauty.


Dominance (Control/ Lack of control): This is similar to self-other polarity, related to Drive level of ABCD.  The Self polarity is associated with self-esteem needs while the Other polarity with Belonging needs. Another way to conceptualize the same polarity is on interpersonal dimension of competence and warmth. If one were to measure well-being related to this dimension, the appropriate measure would be something like Key’s Scoial well-being measuring things like social trust. This dimension is related to controlling/ influencing others either via power or via love. The existential given here is Isolation and the solution is domination and control through exercising one’s ego. Agency/ ego/self may be important here. the issue whether we have control or nor makes this a part of moral domain too.

This dimension, associated with ego, may have evolved to spread the memes associated with the ego far and wide.


In essence, while Feelings and Intelligence are more closely related to evolution (survival and reproduction) of our physical bodies, Agency and Experience are more closely related to the evolution of our minds.

The Evolution of Altruism And Its Relationship to Personality

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.

Evolutionary theory mandates that there be some genetic payoffs in terms of either inclusive fitness or future benefits to self (reciprocity) for any kind of altruism/ cooperation to evolve.

I recommend reading Wikipedia articles on reciprocity, kin selection, and evolution of cooperation if they are not familiar to you or you need a refresher.

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:

  1. 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.
  2. 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.
To me, this looks like equating direct reciprocity with H and direct punishment with A.  With this in mind I now list the personality trait (HEXACO)/ evolution of altruism linkages.
1. Kin Selection/ Genetic Reciprocity : mediated by emotional attachments etc and related to Emotioanlity.
2. Neighbor Selection / Network Reciprocity: mediated by sociability and related to sociability aspects of Extrarversion.
3. Direct Reciprocity : mediated by trusting others and being honest and related to Honesty-humility.
4. Direct Punishment:  mediated by punishing others and being aggressive/ hostile when needed and related to Agreeablness.
5. Indirect Reciprocity upstream:  mediated by feelings of gratitude/ awe/ elevation and possibly related to Openness to Experience
6. Indirect Punishment upstream: mediated by feelings of guilt/responsibility and related to Conscientiousness.
 7. Indirect Reciprocity downstream: mediated by signaling and being trustworthy; need a new personality dimension for this.
8. Indirect Punishment downstream: mediated by signaling and being competent/ tough and possibly related to Surgency dimension of Extarversion.
I am more and more convinced, looking at the above model that we evolved to be cooperative/ altruistic rather than otherwise.

The Eight Rasas (aesthetic essences)

Guru N?ty?c?rya M?ni M?dhava Ch?ky?r performin...

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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. .

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The four mechanisms of evolution

Darwin's finches or Galapagos finches. Darwin,...

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Regular readers of the blog will know of my enthusiasm for evolutionary and developmental theories especially the eight stage evo-devo theory.

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.


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The Four Fundamental Causes

Bust of Aristotle. Marble, Roman copy after a ...

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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.

Niko Tinbergen , a great ethologist, asked four questions or explanations about any biological phenomenon, inspired by the Aristotle’s 4 causes. . From the Wikipedia page:


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.


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emotion and personality: take 5

Scared child
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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 :

  1. Affects or emotions
  2. Behaviors or  social factors
  3. Dynamics or drives and motivations and
  4. 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:

  1. Affect/emotions: Evolutionary explanations as most emotions are evolved mechanisms. Biological context is relevant. Genes drive this.
  2. Behaviors/social functioning: Social/situational explanations dependent on interactions with other people/cultural effects. Cultural context is more relevant. Environment drives this.
  3. 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.
  4. 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:

  1. 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.
  2. Behavior maps to the problem of Adaptation and the polarity of Active and Passive mapped respectively to the ‘RAGE’ and ‘PANIC’ systems of panksepp.
  3. Dynamics/Drive maps to the problem of Replication and the Polarities of Self and Other mapped respectively to ‘LUST’ and ‘CARE’ of Panksepp.
  4. 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.

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Altruism as a result of sexual selection


First published in Fun, Nov 1872. Original cap...
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There is a new article in BPS, that has found some evidence for the fact that altruism may have evolved by the process of sexual selection.

There are many mechanisms that underlie exactly how and why sexual selection takes place- one is the ‘handicap’ /’costly honest signal‘ theory according to which a trait that is actually disadvantageous or a handicap for the host evolves to signal exactly that fact- that despite this handicap I am able to function well and must therefore be of better genetic quality; the most common example being the evolution of peacocks tail which is a handicap and makes the male peacock carrying a big tail more vulnerable for predation, but also is attractive to females and preferred by them.  Another theory is that of ‘runaway selection’ i.e . a trait may evolve in a direction due to some genetic drift and the preference for it may also evolve in tandem such that there is  a slight leaning or preference towards that trait. Now, in a competition, those, typically males, who have that trait will be selected by the females and their progeny will have an advantage as they are more likely to display that trait and be favored by subsequent generations; thus an arbitrary trait may get fixed by this runaway selection where all members of the species want to be part of the new fashion/club in town. I know I am drawing very loose analogy, but just to give an idea. Nakedness or loss of hairs in humans is predicted to have followed this pattern.

Of course sexual selection also differs on whether it is largely intrasexual, driven by competition (selection pressure) between males for eg  having big antlers to defeat and subjugate another male; or is driven by mate preferences and has intersexual selection dynamics- like the peacocks tail.

What the authors of this paper hypothesized was that sexual selection is behind the evolution of altruism or selfless concern with non-kin and that this being the case and this sexual selection dynamics being driven by inetrsexual dynamics, there are bound to be genetic underpinnings to both the trait altruism as measured in males/females and the mate preferences for altruistic trait in both females as well as males. The reason they didn’t anticipate any differences in male sand females was that parental investment, as per them, is roughly equal in case of humans and so both males and females exert equally strong sexual selection pressure son each others traits and ‘choose’ their mates equally.

However, in this study they looked only at females and their genetic basis for altruistic traits as well as mate preference for altruism. The altruistic traits and mate preferential weer measured using self-report instruments.  the genetic components underlying these were estimated using classical twin studies paradigm where correlation between mono-zygotic twins and di-zygotic twins are compared to estimate the genetic contribution. They also calculated the phenotypic correlation between mate preference for altruism and altruistic trait in individuals and tried to calculate how much of this correlation again was genetic in nature or in other words was a result of  mating between those who had the trait and those who preferred the trait. . They hypothesized that in the ancestral environmental this type of mating for trait altruism would have taken place and thus these would be genetically correlated.

This is exactly what they found; they found that both altruistic personality and mate preference for altruism had genetic components and that they both co-varied and that covariance again had genetic component.  the pare itself is full text open access and is written very well, so go ahead and read it yourself. this is an important paper that has come timely when the whole kin-selection paradigm for evolution of eusociallity is being challenged by E o Wilson and team and provides a fresh and alternative perspective of why altruism may have evolved.

Here is a tit-bit from the discussion:

We believe that the sexual selection hypothesis for the evolution of human altruistic traits should now be considered alongside other more established theory (Bshary & Bergmu¨ller, 2008; Lehmann & Keller, 2006), particularly as there is the possibility that multiple mechanisms might underlie a complex behaviour such as altruism. Empirical testing of contrasting theories might even be possible. For example, reciprocal altruism (Trivers, 1971) does not strictly predict the genetic correlation between MPAT and ‘altruistic personality’ found here as ongoing reciprocation towards others would not necessarily result in such a selective process. Indirect reciprocity (Leimar & Hammerstein, 2001) concerns reputation directed towards all other group members while the sexual selection hypothesis focuses solely on altruistic displays that can be evaluated by potential mates (Phillips et al., 2008). A study that examined ‘costly signalling’ of altruistic behaviour through personal donation to a children’s charity found a significant effect on male behaviour when witnessed by a female observer while no such effect was found when male participants were observed by same sex others (Iredale et al., 2008), a finding that could be seen to be at odds with indirect reciprocity. Additional studies could further elucidate the effects of altruistic reputation when directed towards same sex others as opposed to potential mates, thus testing the relative claims of indirect reciprocity against the sexual selection hypothesis.

Tim Phillips1, Eamonn Ferguson2, & Fruhling Rijsdijk (2010). A link between altruism and sexual selection: Genetic influence on altruistic behaviour and mate preference towards it British Journal of Psychology DOI: 10.1348/000712610X493494

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Life History theory and eight stage evo-devo model


Hoverflies (Simosyrphus grandicornis) mating i...
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I’ve touched upon life history theory earlier, in an oblique fashion, while discussing evolutionary perspectives on personality.

Life History theory posits that an individual’s life efforts can be subsumed under two headings- somatic life efforts
and reproductive life efforts. The latter relates to selection due to being able to successfully replicate one-self; the former relates to the ability of an organism to survive and thus act as a vehicle for genes that can be replicated at a later date.  To elaborate more on the life history theory I quote:

Life History Theory is a mid-level theory from evolutionary biology that describes the strategic allocation of bioenergetic and material resources among different components of fitness (e.g., calories and nutrients devoted to growth vs. reproduction). Somatic Effort anchors one end of the first dimension of this trade-off whereas Reproductive Effort anchors the other. Somatic Effort refers to resources devoted to continued survival of the individual organism whereas Reproductive Effort refers to resources devoted to production of new organisms as vehicles for survival of the individual’s genes. The second dimension of this trade-oV further partitions Reproductive Effort. Mating Effort anchors one end of this continuum whereas Parental Effort and Nepotistic Effort jointly anchor the other. Mating Effort refers to resources devoted to obtaining and retaining sexual partners whereas Parental/Nepotistic Effort refers to resources devoted to enhancing the survival of existing offspring and other genetic relatives. Thus, a life-history strategy allocates an individual’s bioenergetic and material resources among the competing demands of survival and reproduction.

I break the somatic effort into five different functions:

  1. Survival– each organism needs to survive and maintain its body integrity
  2. Growth – each organism needs to consume energy and grow
  3. Maintenance– each organism needs to maintain the body it accumulates as a result of previous efforts
  4. Development– there is developmental unfolding and stages where different needs are met at different times
  5. Differentiation/specialization– the organism makes efforts to create an environmental niche for itself

I agree with the authors that reproductive effort can be broken into 3 parts:

  1. Mating effort effort to find and retain a mate.
  2. Parental effort– efforts devoted to nurture offspring
  3. Nepotistic efforts– efforts devoted to helping close genetic kins.

The first three stages/efforts are related to energy balance while the next two are timing related. Finally the final three are efforts proper, thus completing the eight stage model.

The life history theory says that one has limited energy, time and efforts and needs to invest these limited resources wisely- between current and future reproduction; between quantity and quality of offspring; and between self and offspring.

Let me elaborate.

Current and future reproductive trade-offs can be conceptualized as a giant trade-off between somatic effort and reproductive effort. If I reproduce now, I do not invest in my growth, maintenance, survival etc and thus have lowered future chances of being able to survive and reproduce.  On the other hand if I do not reproduce now, I can invest the excess energy in somatic efforts- granting me future reproductive benefits, but incurring the opportunity cost of not reproducing now.

Quality and quantity of offspring trade-off can be conceptualized as how best to allocate resources, once I have decided to reproduce,  amongst a litter- the size of litter (no. of offsprings), the size of individual baby at birth and the quality or survivability of offspring at birth. All these are orthogonal to each other and involve trade-offs.

The final traditional trade-off is between self and offspring- the parent-offspring conflict. Given that I am reproducing some  offsprings now, what resources should I invest in them now,  such that my future reproductive costs are not sky-high and I can survive and reproduce later and invest in later offsprings too.  Here the two sexes have different incentives when a baby is gestating- for mother later offsprings will also be hers; for father later offsprings may not be his; thus father wants the baby to consume as many resources now as possible; while mother wants to spread resources more evenly- the genomic imprinting conflict theory.

Here it is instructive to pause and note the cost of reproductive (to mother)-

  1. future survival (death in childbirth – for mammals important)
  2. future energy capture (pregnancy, lactation and nursing resulting in less  loco mobility and foraging)
  3. future reproduction (lowered fecundity as a result of prior child birth)

The avid reader will note that the three trade-offs refrred to above relate to the three stages of reproductiuve effort – mating (now/future), parenting (one/many child) and nepotism (future related babies/unrelated babies)

There do exist of course tradeoff between somatic and reproductive efforts and it has been shown time and gain that they are inversely related; however more interesting is the fact that somatic efforts themselves are multi-dimensional; thus forgoing reproduction may not simply lead to longer lifespan; it may just result in more growth and heavier bodies.

Also important to note the concept of r and K strategies.

Let us now look at some of the life history variables and again present them is a stage fashion, with special focus on r-K strategy and how these have antagonistic and opposite effects on variables under consideration:

  1. Mortality rate (survival)- low in r , high in K
  2. Body size (growth) -low in r, high in K
  3. Metabolic activity related lifespan (maintenance)- low in r , high in K
  4. Age of sexual maturity (developmental) – early in r, later in K
  5. Age of first reproduction (differentiation/specialization)- early in r, later in K
  6. duration of gestation/pregnancy (mating) – lesser in r, greater in K
  7. no. and size of offspring in a litter (parenting)- more number and lesser size in r, smaller no. and bigger size in K
  8. no. of reproductive events (nepotism)- semelparous or one reproductive event only in many r; iteraparous or multiple reproductive events in most K

Another way of parsing the data is to link these to developmental and evolutionary tasks and personality traits, as I had done earlier:

  1. Foes: Survival
  2. Food: Growth
  3. Friends: Maintenance
  4. Maturity: Development
  5. Mate selection: specialization/differentiations (displays of creativity etc)
  6. Mate retention: mating long term for future reproductive benefits
  7. Parenting: parental efforts for ensuring well being of offsprings
  8. Nepotism /altruism: favoring kin/ non-kin to ensure inclusive fitness

The last three are generally referred to as dilemmas of parental investment- balancing reproduction, well being of offsprings and inclusive fitness via nepotism with somatic or self fitness.

Its also instructive to note that many ecological variables affect the somatic as well as reproductive effort distribution (the r and K strategy are under ecological constraints to an extent).

For ex, changes in ecology like Mortality hazard (foe related) , food availability (food related) , density of con-specifics (friends related) , shortage of food during critical developmental window( maturity related) or differential habitats leading to different genetic variants (niche related) all lead to changes in these life history variables sometime affecting growth, sometimes reproduction, sometime lifespan etc etc. For eg caloric restriction may lead to increased lifespan etc etc

The above life history analysis was applied to organisms, but the same can be extended to brains and neurons-the following processes are involved in neuronal life history (neural Darwinism)

  1. Neuronal survival
  2. Neuronal growth
  3. Neuronal maintenance
  4. Neuronal developmental and
  5. Neuronal specialization/differentiation.

In all of the above neurotropins or growth factors are instrumental and drive these processes. Whether or not this applies to neuronal level, but to organism level life history theory makes a lot of sense. What do you make of it and its fit with the 8 stage evolutionary theory?

Graf, M., Cellerino, A., & Englert, C. (2010). Gender Separation Increases Somatic Growth in Females but Does Not Affect Lifespan in Nothobranchius furzeri PLoS ONE, 5 (8) DOI: 10.1371/journal.pone.0011958
FIGUEREDO, A., VASQUEZ, G., BRUMBACH, B., SCHNEIDER, S., SEFCEK, J., TAL, I., HILL, D., WENNER, C., & JACOBS, W. (2006). Consilience and Life History Theory: From genes to brain to reproductive strategy Developmental Review, 26 (2), 243-275 DOI: 10.1016/j.dr.2006.02.002

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