The Immorality of Morality
Morality and the Dopamine Reward System
A dopamine neuron in the brain's reward system
The recent revelations in Scott McClellan's book, What Happened, has provoked two very different and predictable responses from conservatives and liberals. The empathetic liberals are predictably outraged that a president could risk the lives of troops under false pretenses, while the conservatives view the whole matter as a justifiable and perfectly moral attempt to improve national security.
This adds fuel to the viewpoint that the study of morality runs right through the hearts and minds of the religious conservatives and secular liberals. Although this viewpoint originated on this web-site, it is now making its way into the more conventional world of moral psychology, starting with Jonathan Haidt's discussion on liberal and conservative morality in The New Synthesis in Moral Psychology (2007).
The richness and diversity of morality is a byproduct of the many domains of processing within the brain: the reward system; the empathy system; the harm avoidance system; quantitative and probability estimation systems; the linguistic system; the genetic distance assessment subsystems; memory systems; the sexual arousal and response systems; the body and spatial mapping systems; motor-mirror neuron systems; and last but not least, the olfactory-gustatory system.
These "systems" are diffuse neural networks that overlap to a great degree, that is, a single neuron, like the one in the above picture, may be reused as a conductor of information across many of the aforementioned domains. As such, morality, like any simple colloquial term used for aggregate cognitive traits, is highly problematical when it comes to consistency in neuroimaging results.
Morality is often perceived to be consistent and immutable, following some sort of divine plan on how one should relate with others. But morality has a dark side, and one that has arisen from the bowels of Hamiltonian inclusive fitness. We can lend a hand to a fallen stranger, and at the same time, quietly let children starve to death on the other side of the world. We are simultaneously moral and immoral, and have only one thing to thank for this perpetual state of hypocrisy: evolution.
The Evolutionary Game of Morality
The evolutionary pressure for the development of morality can be laid at the feet of organism complexity and sexual reproduction, which should be expected, given the substantial bias of morality in regulating reproductive behaviors. Sexual reproduction creates genetic diversity and rapidly reduces the percentage of genes shared between parent and child. Further, organism complexity, a co-product of sexual reproduction, requires longer maturation periods and a correspondingly greater parental investment in offspring.
This has transformed the elevated self-interest of asexually reproducing organisms into the wider social-interest of sexually reproducing organisms. This evolutionary game surfaces as empathy and morality. The fundamental essence of this game is exposed by how much it varies with genetic distance. On average, moral and altruistic behaviors increase with decreasing genetic distance, and are highest within nuclear families, where genetic distance is lowest.
But how did evolution execute the moral paradigm founded upon Hamiltonian inclusive fitness? The brain has indeed been under extreme selective pressure for a spectrum of behaviors that have been recklessly categorized as "moral", but no domain of neural processing has been as targeted by evolution as the human dopamine reward system.
Reward, Religiosity, and the Brain
The relationship between religiosity and the brain's dopaminergic reward system is a strong one, as evident in the many references to heavenly rewards in religious canon. Indeed, Previc proposed a theory (2004) involving the evolution of the dopamine system and religiosity, which ingeniously integrates the spatial mapping and orientation systems of the brain (see God, Dopamine, and 3-Dimensional Space).
Further, we suspect that this rapid evolution of the human dopaminergic system has resulted in the transference of earthly reward-seeking into heavenly reward-seeking: one of the most remarkable events in the evolution of the religiosity and morality. Previc also hinted at this: "religious experience and practice in humans are a consequence of the expansion of the dopamine-rich ventral brain systems that, in other mammalian species, mediate behavioral interactions with the most distant portions of the surrounding 3-D space". Rewards are now sought in the dopaminergic system's orientation towards distant upper space, that is, "heaven".
Our initial proposal, in 2005, that the religious conservatives were more organized around reward-seeking was based on a very Darwinian argument: the religious conservatives produce more children and have a greater reproductive burden. The diverse conservative and liberal opinions on global warming are intimately connected to the greater conservative tendency for higher reproductive rates and the required elevation in reward-seeking, and therefore less deference to environmental problems. Ironically, the religious are playing quite the Darwinian game of reproductive advantage (see God, meet Darwin).
The proposal that the religious conservatives are more inclined towards reward-seeking is also justified on the grounds that liberals have a higher tendency towards depressive disorders, which greatly reduce the tendency for reward-seeking. And finally, conservatives rate themselves as more competitive than liberals, another indicator of reward-seeking.
Taking a Ride on the VTA
The connection between religiosity and reward runs through the dopamine system, and in particular, the mesolimbic and mesocortical dopamine systems. The mesolimbic dopamine projections, part of Previc's "action-extrapersonal" pathways, and central to Previc's dopaminergic theory of religiosity, consists of the brain's primary reward circuitry: the ventral tegmental area (VTA), which is connected with dopaminergic afferent and efferent fibers to the nucleus accumbens (NA), among other regions. Orgasmic rush involves strong activation of the ventral tegmental area, and is also a primary target of drugs such as cocaine and heroin.
As an area of the brain involved in both orgasmicity and religiosity, the VTA provides an interesting bridge between God and sex, and Andrew Newberg's view that "the neurological machinery of transcendence may have arisen from the neural circuitry that evolved for mating and sexual experience". We have previously noted an improvement in orgasmicity among the very religious during intercourse (see God, meet Darwin), which is consistent with common dopaminergic substrates between sexual performance and religiosity. However, religiosity involves a much wider spectrum of neurology than that associated with sexual experience.
Previc, in The role of extrapersonal brain systems in religious experience, states: "in terms of their neuroanatomy, dreams, hallucinations, and religious experiences may all be mediated by ventromedial (cortico-limbic) pathways extending from the medial temporal lobe to the anterior cingulate and prefrontal cortex". This, by proxy, implicates the brain's key reward region, the VTA, which maintains an unusual array of connectivity with Previc's aforementioned structures.
The VTA is one of the regions of the brain that receive diverse inputs from other brain regions, and is one of the crucial areas for information processing. These areas typically employ dopaminergic transmission, as does the VTA, increasing the probability of switching between sources of input competing for control. The lateral septum, amygdala, accumbens, lateral habenula, entorhinal cortex, and frontal cortex are other examples of these special integrative regions in the brain.
Evolution has preserved the VTA to be remarkably intact, as there is "astonishing general similarity between the VTA nuclei of mammals ranging from rodents to man" (Oades, 1987). (Much of the discussion here is based on Oades' work). This certainly implies a "modular" nature to the VTA, that is, it is highly specialized for reward processing.
However, one interesting difference in the VTA between rats and men involves the number of neurons, and in particular, the number of dopamine neurons. These increase dramatically with phylogenetic progression (German, 1983), which is interesting in the light of the dopaminergic theories of the evolution of human intelligence (Previc, 1999). Indeed, there is a greater dopaminergic influence in the cortices as one moves up the evolutionary ladder.
The VTA maintains the large majority of connections within the same hemisphere, that is, the left VTA maintains the majority of connectivity within the left hemisphere, which is a very common attribute of neural organization, and the backbone of the poorly conceived concept of "hemisphericity". The VTA dopaminergically innervates different layers within the ipsilateral cerebral cortex, whereas serotonin or noradrenaline predominate in other, earlier cortical layers--a very interesting evolutionary occurrence.
Many forebrain structures show reciprocal connections with the VTA, as the brain seems to be weaving a very tangled web when it comes to reward-processing. Most of the connecting structures of the VTA are feedback loops, with the exception of the septo-hippocampal complex. The VTA is reciprocally connected to both the phylogenetically newer mesocortical system, as well as the mesolimbic systems.
The moral and religious significance of the VTA lies in its connectivity to the socially "hot" regions of the brain: the ventral prefrontal, orbitofrontal, dorsolateral, cingulate, and temporal cortices. These regions participate in social interactions and social attitudes to a great degree, and the fact that the brain's reward system is closely linked with the neurology modulating social behavior is more than a coincidence.
Indeed, the integration of the reward system with social behavior has been under heavy evolutionary pressure, with strong implications on human morality and behavioral inhibition in general.
One of the more interesting aspects of morality lies in its contextual inconsistency. As Joshua Greene (2003) noted, people generally believe it is immoral to abandon a bleeding stranger by the side of a road to preserve one's leather car seats, and at the same time, believe it is moral to spend money on luxury goods instead of using it to provide a better life for impoverished people.
From an evolutionary perspective, this inconsistency follows the rules of dopaminergic processing. Exposure to a bleeding stranger has a neurologically different impact than exposure to the more abstract concept of impoverished people, whom may be at least partially to blame for their impoverishment. We propose that this moral dichotomy may indeed involve the reward system, more specifically, the dopaminergic phenomenon of unanticipated loss.
The China Quake: human altruism is elevated by the dopaminergic phenomenon of unanticipated loss
Dopamine neurons in the VTA and its nextdoor neighbor, the substantial nigra (SN), are more activated in the presence of unanticipated rewards than with expected rewards. But this interesting anomaly also occurs with unanticipated losses (Shepard, 2006). In the VTA and SN, dopamine neurons cease firing when a fully expected reward is withheld, and also in the case of punishment.
This appears to be under the control of a midline structure, close to the thalamus, called the lateral habenula (LN). Electrical stimulation of this area will inhibit the spontaneous activity in the VTA and SN by as much as 90%. The LN is another region of the brain that receives a wide variety of inputs, and may be functioning to inhibit the reward experience of the VTA and SN, thereby functioning as a conduit for the enhanced moralistic response to unanticipated loss.
Humans indeed seem to be more motivated to provide help in those cases of unanticipated loss, that is, in cases of natural disasters or unexpected circumstances. Poverty is anticipated, and coincidentally, does not generate the same level of dopaminergic inhibition and its corresponding moral and altruistic response. Therefore, the strange moral acceptance of spending extra money on leather seats as other people experience the stress of poverty has a distinctive dopaminergic root.
While our proposal that the dopaminergic phenomenon of unanticipated loss has an unusually strong influence on moral attitudes, it is one of many neurological phenomenon that collectively paint the fuzzy landscape of morality. What if, in Greene's dichotomy, the impoverished people were the children of the person spending the money on luxury goods? Most people would now consider this to be unacceptable moral conduct, and there is even child-support legislation to regulate it.
It Takes a Village
What evolutionary game is being played by liberals and conservatives?
Hillary Clinton's proposal that "it takes a village to raise a child", and the conservative response, "it takes a family", highlights the genetic bandwidth variations in their respective social attitudes. All of our cognitive data on conservatives indicate a general bias towards altruistic behaviors over a smaller range of genes than liberals. These two political cohorts are playing two divergent sides in the same evolutionary game, with the conservatives favoring smaller, genetically similar social groups, while the liberals favor larger, genetically diverse social groups.
This evolutionary game is survival of the gene, with the conservatives banking on force of numbers to provide their genes an advantage in future generations. That is, if gene A from one parent is combined with gene A from another parent, then the offspring carry two copies of gene A. The liberals are banking on the value of diversity, with gene A from one parent combining with gene B from another parent, and hoping that the heterozygous condition of both genes A and B will have an advantage over the homozygous condition in future generations.
This is evident in the greater propensity of liberals to interbreed across racial barriers than conservatives. We have previously proposed that the coefficient of inbreeding, F, is higher in conservative populations than in more liberal populations, although this difference is generally quite small, except in isolated religious groups.
Conservatives and liberals are, for the most part, unaware of this evolutionary game. So how are they playing it so successfully? We have long proposed that conservatives are more under the influence of the dopamine system in their cognitive styles, while the liberals are more likely to reflect the cognitive style of the noradrenergic (and serotonergic) systems.
The dopamine system is asymmetrically distributed in the left hemisphere, while the noradrenergic and serotonergic systems are asymmetrically distributed in the right, which is the reason we refer to "left-brained" conservatism and "right-brained" liberalism. Indeed, there is evidence that if these two hemispheres are separated by commissurotomy, they create two simultaneous and opposite political orientations in the same person! In one sense, we might all be conservative and liberal, with one mode being favored while the other is suppressed by the phenomenon of hemisphericity, or the way the brain resolves conflicts between competing neural regions in the brain.
And once again, the dopamine system also seems to be behind the conservative and liberal variations in the tolerance for genetic distance.
The Origin of Political Species: How did us become us?
Most people, at some point in their lives, practice a form of communism known as the family. Within these genetically close social organizations, resources are produced based on ability and distributed based on need, which is the definition of Marxian communism. This form of communism is not even noticed by the minions of anti-communists that practice it daily. But extend the genetic bandwidth of this familial communism, and they begin to notice. The breakdown of communal behaviors with increasing genetic distance rears its ugly head in larger social groups.
This leads to the fundamental question on the origin of the political species: how are humans drawing the line between their altruistic and indifferent (or even spiteful) behaviors towards others? Who is us, and who is them? The information we've collected pertaining to the social behavior of conservatives and liberals points to a very obvious difference: conservative social behavior is heavily organized around reproduction.
Conservatives are more likely to be in a heterosexual relationship that supports reproduction. Further, their relationships last longer, and support a higher-yield model of reproduction (see Sociosexuality). This leads us to the proposal that the conservative "high reproductive yield" lifestyle is anchored by the functioning of the dopamine reward system and the associated neuropeptides known as the relationship "bonding" molecules: oxytocin and vasopressin. This may also be the reason that the conservatives report higher empathy levels for infants versus adults (soon to be reported on this web-site).
Dopamine, oxytocin, and vasopressin have been linked to parent-offspring bonding (Hansen, 1993, Febo, 2005, Bielsky, 2004) and monogamous relationships. Strong social bonds have only been observed in species where receptors for oxytocin and vasopressin are highly-expressed in the dopamine reward system. Depletion of dopamine in the aforementioned VTA blocks the development of maternal behaviors (Hansen, 1991). Social recognition also increases with the administration of vasopressin in male rats (Engelmann, 1996).
Given the dopaminergic emphasis of the left hemisphere, and the general dopaminergic orientation of conservatism, we suspect that the dopaminergic-oxytocin-vasopressin (DOV) bonding mechanism is providing the impetus for the conservative's propensity to prefer smaller social groups, as they indeed seem to exhibit enhanced social orientation to the people they are directly exposed to. Conservatives are more prone to organize their behavior around their immediate social contacts (see Conservatives, Liberals, and Social Networks), and this may be due to the greater influence of the DOV.
Further, pair-bonding seems to increase competitiveness in many species. Among prairie voles, once a monogamous pair has staked out a territory, intruder voles of either sex are expelled from the nest (Getz, 1993). Monogamous species, like the prairie vole, exhibit enrichment of oxytocin or vasopressin receptors in the mesolimbic dopamine pathways (Young, 2001).
In general, maternal behavior, pair bonding, and infant attachment recruit the mesolimbic dopamine pathways, which include the ventral tegmental area and nucleus accumbens (Insel, 2004). Animals, on average, are more competitive during peaks in sexual activity, and are more organized towards defense after birth. In Wistar rats, oxytocin triggers maternal aggression towards conspecifics (Bosch, 2005).
The phylogenetically ancient neurology involved in maternal defense and infantile bonding is a very good place to start when backtracking to the neurology that facilitates human xenophobia and the wide divergence in moral and altruistic behaviors associated with genetic distance. It is certainly interesting that reproductive output is correlated with higher levels of racial prejudice and elevated responses to threats from outgroups.
Resolving genetic distance crosses many domains of sensory processing, with the evolutionarily ancient olfactory system getting the first crack at the identification of reproductively relevant stimuli. The olfactory system is bimodal: a main system that supports the identification of food, predators, and prey; and, an accessory system that supports social recognition of reproductively relevant stimuli, which includes mates, relatives, and offspring.
This accessory system is anchored by the accessory olfactory bulb, and the genes that encode olfactory receptors combine to make about 4% of the mammalian genome. This is an indication not only of how important olfaction has been in the processing of the wide away of chemical signals regarding both food and danger, but also in the evolution of social behavior and the Hamiltonian identification of us versus them. It is interesting that conservatives tend to report greater olfactory efficiency than liberals, and this tendency varies with the strength of conservatism.
This accessory olfactory bulb is connected to vomeronasal organ in earlier species, and supported the detection of molecules that have a social significance, such as gender. However, in humans, this organ is presumed to be vestigial, although the evidence to support this notion is equivocal. Much of its original functionality has been assumed by the main olfactory system. While it appears that the evolution of the human brain has aggressively targeted the elimination of the vomeronasal organ, as it grows until the 30th week of gestation and then recedes, its value in human evolution seems to be the ancilliary neurology that has been hijacked by the other sensory domains, such as vision, hearing, and the main olfactory system.
Perceptual Imprints and Political Behavior
While olfaction is the only sensory system plugged directly into the cortex, its usefulness in resolving genetic distance in humans is limited, due primarily to high degree of genetic complexity. However, the visual and auditory systems would assume proportionately more control, and adapt the rather curious neurological phenomenon known as imprinting via perceptual templates (see Insel's discussion).
Greylag geese following their mother, Konrad Lorenz
Imprinting resolves the problem of attachment of offspring towards parents, and therefore must occur very soon after birth. This process results in some unusual behaviors, such as precocial birds adopting the first moving object (either animate or inanimate) they encounter as their mother. This was documented by Lorenz (as described by Insel), who became the mother of greylag geese reared by him from the time of hatchling. Further, when they reached sexual maturity, the geese courted him in preference to conspecifics.
Another common example of imprinting involves cats, which are unlikely to tolerate human contact unless exposed within the first nine weeks after birth. Somehow, animals have an innate "object-class" template, and may also employ "motion" and "sound" templates, which are only sensitive during a specific time window. Once an object fulfills the template requirements (during the time window), the animal is imprinted. The animal is then more likely to be approach this imprinted object, and its subsequent behavior is under proportionately greater control by that object.
Filial imprinting supports attraction towards parents and conspecifics, while sexual imprinting supports attraction to potential reproductive partners. The functioning, relationship, and timing involved between these two types of imprinting, at least in humans, is not well understood.
It is interesting to note that "morphological studies [in chicks] have shown that imprinting is correlated with an increase in the length of the postsynaptic density of spine synapses in the IMHV [intermediate and medial hyperstriatum ventrale] but only in the left hemisphere" (Insel). This result, as the regular readers of this web-site will immediately see, has potential political implications.
First, the left-hemisphere has been implicated in both religiosity and political conservatism, and by proxy, therefore implicated with a reduction in genetic distance in affiliative behavior and mate selection. Whether or not the human left-hemisphere plays a disproportionate role in imprinting is still speculative, but the left-hemisphere's dopaminergic bias in approach and social bonding certainly points to this conclusion.
Therefore, according to our proposal, imprinting has a greater influence over conservative behavior in general. Thus, the definition of us would be more closely aligned to the qualities associated with those objects encountered during the imprinting window, which presumably would be parents and relatives, that is, those with lower genetic distance. Altruistic behavior would therefore be more likely to focus on this more genetically-limited version of us.
They'll Name a City After Us
The adaptation of the dopaminergic reward system in moralistic and altruistic behavior has evolved under the invisible hand of Hamiltonian inclusive fitness. It has also led to a number of contextual inconsistencies that have puzzled morality researchers such as Joshua Greene.
One of these inconsistencies is a by-product of the way the brain stores memories associated with the phenomenon of unexpected loss. Interestingly, the dopaminergic system is more activated when gains or losses are unexpected, as this process facilitates memory storage for future reference. Expected rewards and losses do not activate the dopaminergic system in the same magnitude, even though they may be greater than unexpected rewards and losses.
Thus, humans do not respond with the same altruistic or moralistic attitudes when the issue involves expected loss. However, unexpected losses, such as those involving natural disasters, induce greater altruistic and moralistic responses.
Another politically active feature of the dopaminergic system is how it supports the resolution of genetic distance. In general, a more active dopaminergic system results in a lower genetic spectrum from which one selects mates, performs altruism, or applies moralistic attitudes. This supports the curious coexistence of extremes of moral and immoral behaviors associated with warfare, such as killing and self-sacrifice.
The dopamine system provides the most intriguing and diverse system of checks and balances for human behavior, and one that responds the most quickly to intergroup conflict, which is why the conservatives are so effective at intergroup competition. Correspondingly, it also provides for the widest range of moral and immoral behavior across genetic distance, and further, reduces the genetic bandwidth in the politically and evolutionarily important definition of us.
Charles Brack, June 2008
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