# Genetics and Eugenics in Frank Herbert's Dune

Discussion of fictional eugenics program in the SF Dune-verse and how it contradicts contemporary known human genetics but suggests heavy agricultural science and Mendelian inspiration to Frank Herbert's worldview.
criticism, transhumanism, genetics, SF
5 May 201812 Jan 2019 finished certainty: highly likely

Frank Herbert’s SF Dune series features as a central mechanic a multi-millennium human eugenics breeding program by the Bene Gesserit, which produces the main character, Paul Atreides, with precognitive powers. The breeding program is described as oddly slow and ineffective and requiring roles for incest and inbreeding at some points, which contradict most proposed human eugenics methods. I describe the two main historical paradigms of complex trait genetics, the Fisherian infinitesimal model and the Mendelian monogenic model, the former of which is heavily used in human behavioral genetics and the latter of which is heavily used in agricultural breeding for novel traits, and argue that Herbert (incorrectly but understandably) believed the latter applied to most human traits, perhaps related to his longstanding autodidactic interest in plants & insects & farming, and this unstated but implicit intellectual background shaped Dune and resolves the anomalies.

One of the odder things in the 1965 SF novel , among its many exuberant ideas, is the role of genetics. Genes are constantly being invoked as a central theme, cause, and goal, molding and being molded by all main characters, but to a reader familiar with human genetics, especially modern molecular genetics, or historical eugenics, the role of genes makes no sense.

This is odd because (1920–1986), while definitely a soft SF author who will never be mistaken for , was nevertheless a widely-read autodidact and great enthusiast for science and technology, turning his farm home into a demonstration project for fuel efficiency & home automation among other things, and writing a home computer manual (Without Me You’re Nothing: The Essential Guide to Home Computers, 1980). Dune famously began in a magazine article on ecological research in Oregon on dune reclamation, and Herbert did extensive background research to develop the world—eg he says “I did a year at the Library of Congress. I did about six years on the whole book (Dune). I leaned on Muslim and Arab history very heavily. I did an extensive study of Arab history. I also used the Library of the British Museum. I’ve lived in the desert. I was doing other things during those six years. Don’t get the idea that was all I did. But I did the research over a six-year period (from 1959 to 1965).”, which includes obscure historical episodes like the Caucasus resistance to Russian imperialism. The Bene Gesserit philosophy draws on , the Mentats are based on idiot savants, the stillsuits are high-tech Bedouin robes, the are old ideas for powered flight, the melange spice bears many resemblances to psychedelics like LSD, the ancestral memory recalls Jung’s speculations about a & western occultic “”, “Dune Tarot” is based on Tarot, the “” accidentally wound up having not one but two references (to Herbert’s anti-development activist friend & to of )—even the bizarre speech by Leto II in with the comparison of war to orgasms as justification for Leto II’s all-female army had a basis (Herbert cites the obscure book The Sexual Cycle of Human Warfare, Walter 19501). Herbert’s use of genetics in Dune is also far from a one-off, as themes of selective breeding/cloning/genetic engineering show up in many of his other SF books as well, like or or or or or (for discussions of them, a good source is Touponce 1988, Frank Herbert). So where does the all-pervasive role of genetics come from, even if only loosely? It certainly doesn’t come from a standard behavioral genetics textbook or from standard research findings on the genetics of intelligence (the sort of topic which one might expect to inspire a fictional eugenics breeding program for intelligence and ESP).

# The Bene Gesserit breeding program

‘“Ever sift sand through a screen?” she asked…“We Bene Gesserit sift people to find the humans.”’

RM Gaius Helen Mohiam, Dune

“Grow up, humans!…That’s their [the BG’s] dream. Start acting like adults and not like angry children in a schoolyard.”

RM Murbella,

To summarize the genetics and breeding program as described in the Dune series: for thousands of years (indeed, “across thousands of generations”), stemming from, Herbert implies in comments2, American and the USSR ‘New Soviet Man’ origins, the sisterhood has been running a persistent highly organized breeding program focused on the sisterhood’s members and the Great Houses of the human Empire, using arranged marriages and political subterfuge as necessary to create the desired crossings in the overall pedigree, aimed at increased intelligence, motor skill, social skills & manipulation, self-discipline, and most of all, the creation of the ‘’, a man capable of highly powerful long-range accurate precognitive ESP of the future and also accessing the ‘ancestral memories’ (the lifelong memories of all their forebears) of both their male & female lineage without losing their sanity or being ‘possessed’ by ancestral personalities.3 Precognition is already possible to a limited extent, and the female half of the ancestral memories are available to the most elite Bene Gesserit (the ‘Reverend Mothers’), but the KH apparently would possess it in far greater degree. It is unclear to what extent these are due to the previous successes of the breeding program. All of these traits are necessary to create a humanity which increases its capabilities without limit, growing into its ‘adulthood’, and becoming capable of long-term planning and survival in an open-ended universe without any certainties or hard rules, without falling into crutches like the destructive “hero myth” and making true democracy a possibility4. (This emphasis on self-mastery and increased capabilities leading to liberation ’s Second Apocalypse fantasy series, which draws heavily on Herbert, but where self-mastery of psychology & biology ultimately leads not to human flourishing but to nihilism, hedonism, damnation, war against the gods, and death.)

The BG decline to use technological methods like AI or genetic engineering in their breeding program. Shortly before Dune, the BG believe that they are finally within a generation or two of success, and plan a final cross: , daughter of , will have a daughter with , and the daughter will then be with Baron Harkonnen’s nephew, , and this rather inbred & incestuous grandson will likely be the KH. Jessica, however, has a son (), who unexpectedly becomes the KH and leads a rebellion against the Harkonnen, being pitted against Feyd-Rautha & Baron Harkonnen, both of whom are killed. In response to the impending disaster, the BG “preserve the [Harkonnen] bloodline” by a last-ditch mating of a BG sister () with Feyd-Rautha, and count on Lady Jessica’s second child, a daughter (), to the other half.

In , it is revealed the Bene Gesserit have a counterpart, the , who specialize in biological and genetic engineering (especially cloning); they casually reveal that they have “dabbled in various pure essences”, including a KH of their own, to study natural extremes (“nature often produces creations as deadly as ours”) in scientific conditions but they found him uncontrollable & he committed suicide. In , Paul Atreides’s two fraternal twins, and , are at the center of political intrigue in the now-headless Empire, and both of them are potential KHs, driving the BG to desperate moves to retrieve their genes somehow, with their preferred solution being a brother-sister mating5, to lock in the KH genes into an offspring despite the political scandal and absolute condemnation of their traditional Fremen supporters, and the risk of recessive genes complicating it. They reject that option, and Leto II instead merges himself with the sandworms to become a nigh-immortal worm-human hybrid, Alia ultimately is driven insane and commits suicide, and the BG breeding program is taken over by the transformed God-Emperor Leto II. In , Leto II takes over the BG breeding program6 for additional millennia as part of his grand plan to ensure the permanent survival of the human race against the possibility of prescient machines/artificial intelligences being developed & turning against humanity; the Golden Path breeding program continues until the birth of a very distant Atreides descendant, Siona, who is invisible to prescience (a biological version of “no-ships”/“”, so perhaps one should dub Siona as having a ‘no-gene’). By this point the breeding program has successfully developed humans to the extent that the last ghola of Duncan Idaho, famed martial artist, can be defeated by even an old man;7 and by the end of the series, being a Mentat, an achievement of only the most intelligent & well-trained in Dune and something remarkable in Paul Atreides, has become commonplace (and the final mysterious enemy scoffs at Mentats–“Dime a dozen, they are.”) Siona’s actions precipitate the death of Leto II, the collapse of the human Empire, and the exodus of countless billions of humans in a diaspora across the multiverse where they will adapt and evolve to local conditions and change in unpredictable ways with the development of brand new genes. (“Think of the uncounted genes out there! Think of the potential talents floating free in universes where they might be lost forever!”)

The remaining 2 books (, ) deal with genetics mostly in the form of “” clones, only somewhat touching on the ‘uncounted genes’ manifested in new kinds of human capabilities like the “futars” & “Honored Matres”; the final book, Chapterhouse: Dune, ends with the protagonists escaping from a mysterious enemy (who appears in the guise of kindly old ‘farmers’ on a farm discussing how to best breed their crops) along with the last Bene Tleilax who is carrying an archive of cells taken from all the major characters of the series. It is implied that they will all be cloned as gholas, with their multifarious talents preserved intact, to be used in the continued development of humanity towards the goal of adulthood.

(The Brian Herbert/Kevin J. Anderson novels apparently go into considerable detail about the origins of the BG and goals, based on the ever-expanding secret material Frank Herbert left behind, but as there is no credible third-party evidence that they are drawing on authentic material, and their novels are too lousy to read, I will ignore them here although it’s possible at least their sequels to Chapterhouse: Dune, /, are based on some degree of authentic material which could be reverse-engineered with sufficiently careful reading.8 It would also be worth remarking on Destination: Void’s use of clones, but I don’t recall it sufficiently well, and I haven’t read yet.)

# A human/animal genetics evaluation

Given this, the breeding program in Dune makes no sense. (For a good review of quantitative genetics animal breeding methods, see , Gianola & Rosa 2015; for what Frank Herbert would’ve had access to at the time, see the dean of American animal breeding, ’s enormously influential textbook, Animal Breeding Plans.)

A continuous polygenic trait responds quickly to selection, and in discussing eugenics, even the most pessimistic estimates by R.A. Fisher of how many generations it might take to drastically increase average human intelligence or almost entirely eliminate a nasty recessive might be 20 generations—certainly not “thousands of generations”. This would hold true of other traits one might select for, and selecting for many traits simultaneously would increase the number of generations relatively modestly. Since there are few or no rare variants fostering extremely high intelligence or other desirable traits, all of the necessary variants exist already in the human gene pool and merely need to be increased or decreased in frequency, which can be done rapidly without waiting centuries (or tens of millennia) for “hopeful mutants”. Due to the CLT, for a highly polygenic additive trait, the starting population mean may be extremely distant from the end result of a selective breeding program (no will ever have a puppy as big as the average ), so it would be incorrect for the Bene Tleilax to claim that the baseline human population could occasionally throw up extremes anywhere as extreme as their ultimate ‘essences’.

If anyone did such a program, it would be self-defeating to restrict the program to a few aristocrats because the families would regress to the mean and would be constantly diluted by intermarriage from the general population (especially over millennia) and such a restriction risks various ills from a small effective population as it greatly increases the risk of bad luck and exaggerates the effects of ; it would be best to do it on as egalitarian a basis as possible, and if it has to be limited to a certain size, select anyone with high trait values with little regard for lineage.

Should a high genetic level be reached and maintained, it would then not be especially important to mate this person with that person, since it is only the average which matters, and a misaimed marriage simply means a slight reduction in selection efficiency; nor would there be anything particularly special about a brother-sister inbreeding, other than incurring & an increased risk of birth defects/genetic diseases (which is not good but tolerable as long as it is not repeated, in some eras such a mating was quite common, and even now is common & preferred in many societies). Likewise, it would not be terribly important to carry the exact cells of particular noted figures to clone them rather than, say, a sibling or several more distant relatives.

So is it all just complete nonsense?

The Mendelian monogenic paradigm, with heavy emphasis on various nonlinear or interaction rather than simple additive effects, remains highly influential in scientific research. (Indeed, perhaps far more than it should. It seems to me that the disappointed hopes of rapidly finding most disease-causing genetic variants after the Human Genome Project rested on quasi-Mendelian beliefs and disregarding the evidence of high additive & polygenicity of most human traits including diseases, and were responsible for the candidate-gene debacle where almost all candidate-gene hits were shown by later GWASes to be false positives. Indeed, so opposed to the standard behavioral genetics paradigm were many researchers & commentators that they used the initial GWAS null findings, indicating that the “” was due to polygenicity with many small effects and thus requiring sample sizes close to n>100,000, as reductio ad absurdum which disproved the entire enterprise; , of course, those sample sizes were reached and the hits have kept coming ever since.

Why is this the case? Perhaps because it suffers the dual problem of being offensively theoretically simple yet practically difficult to deal with; early Mendelians (eg ) complained of the difficulties of understanding Galton, , or Fisher’s mathematics, while applying the statistics at all must have been enormously painful in an era where even mechanical calculators were not always available, and the implications are that for some things like GWASes hundreds of thousands or millions of samples were required, all in the service of a theory whose intellectual charm & subtlety are difficult to appreciate, and which seems prima facie false to anyone familiar with the intricate endlessly complex pathways and feedback loops of real biological systems. And yet, ‘it moves’, for all the sophistication and of Mendelian theories reveling in epistasis and dominance. It can be easy to read small n data in Mendelian ways, assuming away anomalies as measurement error and the usual ‘crud factor’ of scientific research—a striking recent example is the 60-year-long mistaken belief that response is a single Mendelian autosomal dominant trait based on ’s 34 cats which turns out to be when studied more rigorously with n~210. In addition, the monogenic approach is indisputably successful in describing many dramatic genetic diseases. And, of course, the eugenics implications for humans of Mendelian-style genetics are much less, in exactly the ways Herbert inadvertently illustrates. So perhaps we should not be too hard on researchers who naively expected to find a few dozen genetic variants which could account for most differences in intelligence or health, and which could be found looking under the lamp post using easy samples like n=100.)

In particular, as extended by , it is heavily used in animal and plant breeding in creating new strains of plants with a specific desired trait, often crossed in from another varietal or even species. In those scenarios, where one is trying not to exaggerate existing traits but to copy an entire novel trait—resistance to a particular pesticide or insect, perhaps, or salt resistance, or a coat color—there may be more than one genetic variant at work, you may need a whole gene copied over from the other organism, perhaps several of them working in concert, acting epistatically, a “gene complex” as Wright dubbed them. Epistasis makes breeding difficult because the new set of genes might be broken up immediately by the recombination. If there are, say, 3 new genes brought over into an organism and it has some offspring with an unmodified organism, each offspring will have, say, odds of inheriting the full set of 3; so of 8 offspring, perhaps 7 will not have the desired trait because only the 8th managed to get all 3 simultaneously. Only 1 or 2 is no good. These are not good odds and complicate things (if you can only see the result when all 3 genes are inherited, how do you know whether there were 0, 1, or 2 in the ones without the trait?). As Lush puts it in his 1943 textbook:

Selection for epistatic effects is somewhat like building a sand pile on the seashore exposed to each incoming wave. It is easy to build a little pile between waves, but each wave which rolls over it tends to flatten out the pile. When building is stopped, some traces remain after the first wave and perhaps even a few after the second and third, but soon practically all traces of the pile are leveled away. If building continues between waves, the pile can be built a little higher before the second and third waves than it was built before the first wave but soon a size is approached which can just be maintained, the building between waves being just enough to repair the leveling action of the preceding wave.

Or as would put it, directly considering human geniuses (, 1978):

Truly exceptional individuals, weak or strong, are, by definition, to be found at the extremes of statistical curves…Since each individual produced by the sexual process contains a unique set of genes, very exceptional combinations of genes are unlikely to appear twice even within the same family. So if genius is to any extent hereditary, it winks on and off through the gene pool in a way that would be difficult to measure or predict. Like Sisyphus rolling his boulder up to the top of the hill only to have it tumble down again, the human gene pool creates hereditary genius in many ways in many places only to have it come apart in the next generation.11

To investigate or select, one must carry out time-consuming and difficult breeding of multiple generations, various of related organisms, and so on, and one can eventually deduce all of the relevant parameters and introduce everything enough times.

But… what you can do is, once you have managed the cross, create a line of organisms which breeds true for the trait by extensive inbreeding or cloning, “line-breeding”. (The use of inbreeding for developing new lines is generally attributed to early English breeder 12; breeders then avoided inbreeding as ‘incest’ and worried about inbreeding depression, engaging in extensive cross-breeding of varieties, which while avoiding both of those problems, drastically slows down progress and obscures heritability and inevitably muddles any sharp distinctions.) If you get say 16 offspring from that organism, and take the two sibling organisms with the trait, both of which you know have the full gene complex of 3 variants, and you mate them, then all of their offspring will express the trait because the 3 variants have been fixated within that line. (This sort of incestuous inbreeding approach would also help with purging harmful recessives: because they are so related, offspring will often have two copies of a harmful recessive and it will immediately cause ill health or death, rather than continue floating around the general population.) Or you could clone them and ensure the genes (and thus trait) is preserved that way—cloning is especially common in plants, and many famous plant varieties are propagated clonally because their characteristics would be lost if they were propagated sexually. ( are a famous example: wild apples exhibit tremendous variety but typically all taste bad and are useful mostly for making ; the supermarket apple varieties all stem from single discovered on farms to be unusually tasty, and then are propagated clonally for commercial sale. The commemorates the legendary discovery of the popular green apple underneath Maria Ann Smith’s kitchen window, although it may also have been found growing in a pile of discarded French crab-apples, while was accidentally discovered after a hurricane knocked down its surrounding normal apple trees; apple varieties can be discovered when individual limbs of trees mutate, creating , see ///. Basically, apples work in real life the way the X-Men work in fiction.) Then you can use that strain directly, or employ it in various other breeding programs while perpetuating the line indefinitely. All of this is common in, for example, plant cultivars or in the special mice & rat breeds (Green 1966) used in lab work. Or if a desired mutation suddenly pops up in an individual, perhaps encouraged by the use of mutagenesis like , the mutation will be lost if it is not bred heavily, possibly with relatives. There can be additional advantages to inbreeding selection (see chapter 23/30 of the draft Walsh & Lynch textbook on selection), especially in situations in which past generations can be re-bred, such as with saved seeds or in the case of Dune, gholas repeatedly cloned from old cells.

# Bene Gesserit as Farmers

So—this alternate paradigm can neatly explain all of the oddities of the Dune breeding program! The reason it is so odd is because Herbert was drawing on the obsolete Mendelian interpretations which were heavy on epistasis and de novo mutations, as opposed to the more plausibly relevant biometric Fisherian paradigm of highly polygenic additive traits with selection on standing variation. Herbert was throughout his life interested in agriculture & genetics, as demonstrated by his demonstration home farm project and the repeated use of agricultural themes in his works (eg Hellstrom’s Hive, where a group of humans develops into eusocial insects, or The Green Brain, where human extermination of insects has catastrophically destabilized global agriculture & provoked evolution of intelligent insects).

It appears nonsensical if you aren’t familiar with animal/plant breeding, and stuff like brother-sister mating looks arbitrary or a weird sexual fetish, but such sibling mating is an established tactic for fixating a specific set of multiple genes which all depend on each other to work at all (epistasis) and common in breeding. Hence, the reason the program took so many thousands of years is that they were waiting for de novos to pop up and were fragilely assembling multiple gene complexes of epistatic variants. The BG had to maintain an elaborate program to hunt the hints of various recessive/epistatic patterns in their pedigrees (“We Bene Gesserit sift people to find the humans”), gradually concentrating sets of useful genes in prized aristocratic lineages, and occasional premature deaths or political errors could let slip a ball of yarn they had spent generations gradually winding. And there were no Siona no-genes floating around the gene pool for Leto II to select on, he had to wait patiently for it to pop up naturally in Siona. This is also why the Bene Tleilax could observe “essences” in the wild13 or offhandedly create their own Kwisatz Haderach: the BG & Leto II refuse to use anything but natural procreation, but the BT can edit and clone freely, which massively speeds things up. Finally, this is why unusual genetics jargon from the Mendelian tradition (like Leto II describing Duncan Idaho as “a first-generation cross”) pops up occasionally in the novels.

This is not to say that this is a correct view of human genetics, or that Frank Herbert necessarily had an explicit idea of epistasis and de novo mutations in mind while plotting out Dune, but it shows that in his wide reading (especially related to his hobby of farming), he could easily have picked up on existing research and methods, and these influences were woven into his novels.

# Appendix

## Race in My Little Pony

(For background on , .)

Another fictional universe with genetic mechanisms is My Little Pony: Friendship Is Magic, where there are 3 pony races which are heritable. One outlier family which has all 3 races represented challenges simple Mendelian interpretations of MLP races. I review 4 attempts to reconcile the outlier with Mendelian mechanisms, and propose another interpretation, drawing on polygenic mechanisms, treating race as a polytomous liability threshold trait, which is flexible enough to explain all observations in-universe (at least for the first few seasons of MLP).

In My Little Pony: Friendship Is Magic, there are 3 apparently separate & specialized pony races: “Earth ponies” (physically strong & naturally gifted at agriculture), the Pegasi (fliers who magically manipulate the weather), and Unicorns (magic specialists—the fourth type of pony, the all-female “Alicorns”, are extremely rare and are usually explained as being magical); they are not separate species and can interbreed, but can be considered breeds/races: almost all depicted families are homogenous & have offspring of the same race, they are clearly distinct, distantly related (eg Pinkie Pie & the Apples/Pears), typically reproduce within their own race, and tend to cluster geographically (with the noted exception of Ponyville).

All main characters’ families are, as far as shown, the same race as their daughter. The one exception is the Cake Family of bakers, where the two Earth ponies Mr. and Mrs. Cake have two children in season 2 episode 13 (“Baby Cakes”), a pair of fraternal twins, who are discordant—Pegasi and Unicorn. When queried by the confused main characters, Mr Cake appeals to a vague genetic explanation, claiming “My great-great-great-great grandfather was a unicorn, and Cup Cake’s great aunt’s second cousin twice removed was a Pegasus. That makes sense, right?”

The puzzle here is how can we explain that almost all families except the Cakes are homogenous in pony race, but it is still possible for one (or two) occasional discordant offspring?

A polytomous is sufficiently flexible as to explain all of these patterns by simply setting thresholds appropriately and expecting assortative mating. For example, there could be a single race trait which is polytomous with 2 thresholds, Pegasi|Earth|Unicorn, and the thresholds are set sufficiently extreme as to ensure most families of 2–3 children (MLP families tending to the very small) are homogenous, such as at −2SD and +2SD. Then ensures that most Pegasi families have a ‘race trait’ mean somewhere well below −2SD, Earth families have a mean ~0, Unicorn families have a mean trait >2, etc. Then, if Mr Cake is unusually close the 2SD threshold and Mrs Cake is unusually close to the Pegasus threshold, the two fraternal twins could inherit differentially from their parents and wind up being pushed across different thresholds. (Siblings/fraternal twins may both inherit 50% from each parent, but they will inherit different 50%s at random due to the randomization of , and their relatedness to each other can easily be ±5%, so in a particularly extreme case perhaps that would be enough.) Alternately, it could be determined by multiple binary liability threshold traits, one for each race, and the expressed race is simply the maximum of the 3 trait values; in which case similar logic holds for the two twins. The mane problem, you might say, with the liability threshold model is that it is, if anything, too flexible and so it’s hard to provide good evidence picking it out with just the MLP evidence. (In real world genetics, one could accumulate evidence for a liability threshold model by examining ‘risk’ in increasingly distant relatives and whether it drops off with genetic distance as it should, looking for continuous measurements reflecting an increased genetic trait value like depressed IQ or schizophrenia symptoms in relatives of schizophrenics, or using molecular genetic methods like or to see if a binary trait is highly polygenic, etc, but none of that is possible here.)

A Mendelian model is a little harder to come up with but there are at least 4 attempts. HelloNurse suggests a monogenic model with 3 alleles each causing a different race, Earth pony dominance, and epistasis or environmental randomization (ie low penetrance):

Mendelian ponies could have a single gene with 3 alleles, E/U/P, with EE, EU and EP earth, UU and PP respectively unicorn and pegasus and UP either unicorn or pegasus at random (i.e. depending on other genes). If the two parents are EU and EP they are earth but offspring of all three types is possible. An even simpler model: one gene with two alleles, earth and fancy, earth dominant, fancy ponies become unicorns or pegasi depending on rainbow intensity or other non-genetic factors.

Grim-S-Morrison gives a 2 gene model, with a dominant/recessive Earth/Unicorn effect and a dominant/recessive Pegasus/Unicorn, giving a Punnet table (reproducing in Markdown):

Grim S Morrison’s 2-gene model using Earth pony dominance; hypothetical for Mr & Mrs. Cake
EuPu (Earth parent)
EuPu (Earth parent) EP Eu uP UU
EP (Earth) EEPP (Earth) EEPu (Earth) EuPP (Earth) EuPu (Earth)
Eu (Earth) EEPu (Earth) EEuu (Earth) EuPu (Earth) Euuu (Earth)
uP (Earth) EuPP (Earth) EuPu (Earth) uuPP (Pegasus) uuPu (Pegasus)
uu (Earth) EuPu (Earth) Euuu (Earth) uuPu (Pegasus) uuuu (Unicorn)

This admittedly has some problems explaining why Pegasi and Unicorns appear to be quite common in Equestria; it would require an extreme caste system to keep Pegasi & Unicorn families highly homogenous & a large fraction of the population, rather than being submerged in Earth ponies and mostly popping up in Earth pony families, which is so rare that multiple adult ponies could be quite surprised at any discordance. And of course, if two Pegasi ever have Earth pony offspring (rather than Pegasi or Unicorn) or two Unicorns are ever shown as having non-Unicorn offspring, the model would be immediately falsified.

Finally, Tad Stone & Soryu Aleksi take as a starting point that Earth ponies are much more common, and Unicorns rare; they don’t try to explain to explain Alicorns. They also go with a blocking method:

Going from here, the ponies would have two genes, a Unicorn and a Pegasus gene. Both genes have a dominant allele that blocks the development of wings or horns respectively, and a recessive one to develop the features:

• P: Pegasus blocker
• p: Pegasus
• U: Unicorn blocker
• u: Unicorn

An Earth pony is a pony where both genes contain at least one blocking allele…The possibilities then are as follows:

Tad Stone & Soryu Aleksi 2-gene model, racial types
Race Possible allele combinations
Earth pony PPUU, PpUU, PPUu, PpUu
Pegasus ppUU, ppUu, ppuu
Unicorn PPuu, Ppuu

This would make Mr and Mrs Cake’s children possible if both parents are PpUu:

 PU Pu pU pu PU PPUU (Earth) PPUu (Earth) PpUU (Earth) PpUu (Earth) Pu PPUu (Earth) PPuu (Unicorn) PpUu (Earth) Ppuu (Unicorn) pU PpUU (Earth) PpUu (Earth) ppUU (Pegasus) ppUu (Pegasus) pu PpUu (Earth) Ppuu (Unicorn) ppUu (Pegasus) ppuu (Pegasus)

Of course this would mean that Mr and Mrs Cake each have a (possibly very distant) Pegasus and a Unicorn ancestor. However, in the series only a Pegasus cousin for Mrs Cake and a Unicorn for Mr Cake are named. With only these two as non-Earth pony ancestors it would not be possible following my own theory, but in all likelihood they don’t know all their ancestors. Also a pretty complicated combination of genes would be needed in that case and we can’t describe such a thing without knowing much, much more.

This explanation works nicely and violates the minimum of elements.

There are some further theories which I think are inconsistent with the evidence:

• CocoaNutCakery suggests a more complicated epistasis, explaining the non-observed combinations as being fatal/producing nonviable embryos, and explaining the Earth pony-like magic of Fluttershy (a Pegasus) as being due to additional magic resistance & explaining Alicorns similarly; this has the serious defect of requiring Unicorn & Pegasi families to have much lower fertility or severe infant mortality rates (which are not supported in canon and are rather against the spirit of the show).
• theaceofspadez’s “The Genetics of the Pony: The Nelson Theory for Pony Inheritance” founders on attempting to explain too much and making Alicorns far too frequent, and also postulating “Alicorn Earth/Pegasus/Unicorn” variants too.
• Tacticalrainboom’s environmental determination theory argues that it’s not genetic at all but environmental (eg Pegasi are caused by exposure to high altitude & weather magic), but this is quite a stretch as it can’t explain why Earth ponies colonizing new places have Earth pony offspring (eg. the entire Apple clan from many different locations throughout Equestria are all Earth ponies), what environment Manehattan & Ponyville have in common causing Earth ponies but which the capital city of Canterlot does not to cause mostly Unicorns, or why, if deviations from the local norm are due to random environmental fluctuations, both Cake twins would be discordant instead of being the same type (presumably any random environmental fluctuation affecting Mrs. Cake ought to affect the fraternal twin fetuses simultaneously & equally).

But wait—there are more possible explanations! Could Mrs. Cake have been cheating on Mr. Cake or they did a threesome, since fraternal twins don’t have to be fertilized by sperm from the same father, and this is why he had shifty eyes/was vague? Were surrogacies and/or egg/sperm donors involved, because one or both of the Cakes are infertile? Or could the twins have been affected prenatally by Discord at the beginning of season 2? Or perhaps ponies are not even diploid (so all the Mendelian explanations are fundamentally wrong), but , in which case just about any kind of inheritance is possible?

That said, reviewing the theories, I think my polygenic threshold models or Tad Stone’s two-gene model fit the best overall.

1. Frank Herbert, “Listening to the Left Hand”, Harper’s (December 1973):

…We tend to react together with a remarkable degree of similarity across boundaries that are real only to individual cells, but remain transparent to the species. We tend to go psychotic together.

Touch one part and all respond.

The totality can learn.

This implies a nonverbal chemistry of species-wide communication whose workings remain largely unknown. It implies that much of our collective behavior may be preplanned for us in the form of mechanisms that override consciousness. Remember that we’re looking for patterns. The wild sexuality of combat troops has been remarked by observers throughout recorded history and has usually been passed off as a kind of boys-will-be-boys variation on the male mystique. Not until this century have we begun to question that item of consensus reality (read The Sexual Cycle of Human Warfare by N.I.M. Walter). One of the themes of my own science fiction novel, Dune, is war as a collective orgasm. The idea is coming under discussion in erudite journals such as the General Systems Yearbook. …

• Something like pheromones (external hormones) interacting between members of the human species to weld groups into collective-action organs. (How does a mob unite and hold itself together?)
• Isolation cues that separate groups into identifiable substructures, a system possibly influenced by diet. (Aside from accent and mannerism, how do members of the British upper class recognize each other?)

Walter’s book, as hard to get as it is (it has never been reprinted), occasionally surfaces in odd places. For example, was impressed enough to spend several pages describing it in his 2006 autobiography Confessions Of The Father Of The Neutron Bomb. Cohen, incidentally, gives some of the few biographical details about Walter I’ve seen:

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2. Dune, of course, takes you through the creation of the superhero, Paul Atreides, who becomes the Mahdi of the Fremen. Please focus on that word creation because many of the elements which go into the making of the superhero are laid out for you in this work. You have here a kind of distillation of an aristocratic bureaucracy, one of whose unmentioned ancestors is the Soviet experiment. You are taken through a history of many power instruments which have been tried and discarded (or adapted to new forms).

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3. It’s worth noting that the ancestral memories are probably part of the KH package and rely on prescience. The usual interpretation by readers that ancestral memories are encoded into DNA and passed on that way may be an over-reading of the mentions of DNA: encoding into DNA doesn’t make any sense because it would imply, among other things, that ancestral memories work only for direct ancestors, cannot recall anyone whose lineage went extinct, cannot recall anything in an ancestor’s life after fertilization of their children, cannot recall dying (except for gholas), ought to decay or mutate over many millennia, have limited capacity, etc, and cannot explain how the Duncan Idaho ghola eventually recalls memories from all his gholas, even ones whose cells were never used in the final ghola! So the connection of DNA to ancestral memories is probably more in the role of a ‘key’ or ‘index’ where relatedness opens up access across time.↩︎

4. The general theme throughout the Dune series of humans becoming ever more capable and flexible generalists is undeniable, but the ultimate culmination in democracy is a little doubtful; Herbert was not a fan of dictatorship or monarchies, despite those being about the only forms of government depicted, and it seems to be consistent with a few passages like the discussions of the Honored Matres, but the only source for democracy as the end-game is a late recollection in January 2009 by :

SR: Do you believe Frank had intentions of a “Dune 7”? Along the same lines, what do think of the claim of Brian Herbert to have found floppy disks containing the “complete” Dune 7 outline in a safety deposit box years after Frank’s death? If there were notes, do you believe Brian and Kevin J. Anderson used them faithfully in their new “Dune” books?

NS: Maybe. Frank kept going as long as the big money kept rolling in. Knowing Frank’s political philosophy, I once asked him how he could keep writing this royalist stuff. He told me he planned to end the series with a novel that would transition to a fictional universe of democratic rule. Never wrote it, of course. And Brian and Kevin certainly didn’t from any 7 notes.

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5. From Children of Dune:

“If … if Jessica has gone back to the Sisterhood completely …”

“That’d be very dangerous to us,” he said, completing the thought. “We carry the blood of their Kwisatz Haderach—their male Bene Gesserit.”

“They won’t abandon that search,” she said, “but they may abandon us. Our grandmother could be the instrument.”

“There’s another way,” he said.

“Yes—the two of us . . . mated. But they know what recessives might complicate that pairing.”

“It’s a gamble they must’ve discussed.”

“And with our grandmother, at that. I don’t like that way.” “Nor I.”

“Still, it’s not the first time a royal line has tried to . . .”

“It repels me,” he said, shuddering. She felt the movement, fell silent. “Power”, he said.

And in that strange alchemy of their similarities she knew where his thoughts had been. “The power of the Kwisatz Haderach must fail,” she agreed. “Used in their way,” he said…“You are not Osiris.” Ghanima reminded him. “Nor will I try to be.”

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6. GEoD: Leto II says “The Sisterhood has never forgiven me for taking their breeding program away from them” in discussing motives the BG would have for acting against him.↩︎

7. In God Emperor of Dune, Leto II’s 118 year old majordomo, Moneo, initially disapproves of the use of original Duncan Idaho gholas, noting that “The Duncans are slower and less alert than anyone in your Guard.” Leto II points out that using Idaho “gives me access to a first-generation cross between an older human form and the current products of my breeding program. Siona is 21 generations from such a cross.” Duncan later attacks Moneo and is easily defeated:

“You just haven’t the guts to pay the price he’s asking”, Moneo said.

In one blurred motion, Idaho whipped his knife from its sheath and lunged at Moneo. As fast as he moved, Moneo moved faster—sidestepping, tripping Idaho and propelling him face-down onto the floor. Idaho scrambled forward, rolled and started to leap to his feet, then hesitated, realizing that he had actually tried to attack an Atreides. Moneo was Atreides. Shock held Idaho immobile. Moneo stood unmoving, looking down at him. There was an odd look of sadness on the majordomo’s face.

“If you’re going to kill me, Duncan, you’d best do it in the back by stealth”, Moneo said. “You might succeed that way.”

Idaho levered himself to one knee, put a foot flat on the floor, but remained there still clutching his knife. Moneo had moved so quickly and with such grace, so . . . so casually! Idaho cleared his throat. “How did you. . .”

“He has been breeding us for a long time, Duncan, strengthening many things in us. He has bred us for speed, for intelligence, for self-restraint, for sensitivity. You’re. . . you’re just an older model.”

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8. It’s worth noting that Frank Herbert originally worked as a journalist and did not have a track record of developing things for long periods or taking extensive notes, aside from the original Dune trilogy, where he notes in an Omni essay (“Dune Genesis”) that “I conceived of a long novel, the whole trilogy as one book about the messianic convulsions that periodically overtake us.”, and in the preface to Heretics of Dune that “Parts of Dune Messiah and Children of Dune were written before Dune was completed. They fleshed out more in the writing, but the essential story remained intact.”↩︎

9. Note that Ramagopalan et al 2007 suggest HP magic is a few-gene model driven by dominance, which is an improvement over the simplistic (to the point of simple-mindedness) recessive monogenic magic model of , but—Klenotiz 2012’s exotic monogenic dominant model invoking -like dynamics notwithstanding—the evidence they give is equally or even more consistent with magic being a continuous trait (with perhaps a threshold), driven by polygenic additive variants.↩︎

10. The pony races are a little difficult to explain under a simple Mendelian paradigm because of an Earth pony couple who have Pegasus and unicorn children. For details, see the appendix.↩︎

11. For some more on this idea of genius, see Johnson & Bouchard 2014, ; Simonton 1999, ; Lykken et al 1992, ; Lykken 2008, “The Genetics of Genius”; Lykken 2006, ; Lykken 1982, ; Simonton 2005, .↩︎

12. For more background, see .↩︎

13. If the extremes of nobility or sadism or KH-ness the Bene Tleilax are interested in are driven by de novo mutations or occasional freak aggregations of complexes, this will happen once in a while observing on a scale of centuries & interstellar-sized human populations, in the same way that albinism and other bizarre traits can pop up at random, either by a chance mutation or by a chance aggregation of recessives etc. Indeed, it has been noted that the current human population is large enough that any mutation in the human genome not quickly fatal can probably be observed somewhere in the current population, and the absence of any carrier of a particular mutation is a strong indicator that it would be a lethal mutation (because everyone with it died).

In contrast, if the extremes in question were continuous polygenic additive traits where the extreme was something like 10 or 20 standard deviations from the norm, waiting for an extreme to appear naturally would require waiting the age of the universe (ie never); this is because random fluctuations are vanishingly unlikely to drive any individual that far away from the mean. So, for example, it is quite common to observe dwarfism or giantism due to rare diseases or genetic disorders or mutations of large effect, but one will never observe a 3-meter-tall man caused by just common genetic variants at their current mean frequencies.↩︎