Genetics and Eugenics in Frank Herbert's Dune-verse

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, insight-porn
2018-05-052019-01-12 finished certainty: highly likely importance: 3

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,

“I think we are the best equipped sur­vival ani­mal that this planet has ever pro­duced. I don’t depend just on ratio­nal­i­ty, I depend on the need to sur­vive, on the urge to sur­vive, on the desire to sur­vive as a species. This is behind a lot of what I write. It pleases me to think that 20,000 years in the future, 20 mil­lion years in the future, there will be human beings around enjoy­ing life the way I enjoy life.”

Frank Her­bert, part 5 of his 1983 pre- inter­view

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 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

To quickly sum­ma­rize some rel­e­vant human & ani­mal genet­ics: the behav­ioral genet­ics par­a­digm traces back to the British “bio­met­ric” school of genet­ics which began with & dis­cussing kinds of “blend­ing inher­i­tance” for the very grad­ual process of evo­lu­tion, lead­ing to sub­tle changes com­pound­ing for eons, which Dar­win set out in Ori­gin of Species; Gal­ton’s inves­ti­ga­tions ulti­mately led to the cen­tral role of the and and and in math­e­mat­i­cally mod­el­ing the inher­i­tance of con­tin­u­ous traits like height, which even­tu­ally was per­fected by infin­i­tes­i­mal mod­el, where the con­ti­nu­ity of traits like height (as opposed to sim­ple dis­crete Mendelian traits which shift between differ­ent phe­no­types under the influ­ence of one or a few genes) is due to the trait being caused by the sim­ple addi­tive sum of the aver­age effect from thou­sands or tens of thou­sands of genetic vari­ants. This could even account for the count­less binary or dis­crete traits which clearly had genetic influ­ence and ran in fam­i­lies but failed to fol­low any kind of Mendelian pat­tern what­so­ev­er, such as alco­holism or schiz­o­phre­nia, by the where a thresh­old is defined and the phe­no­type man­i­fests if the sum of all genet­ics vari­ants & envi­ron­men­tal influ­ences (a nor­mal­ly-dis­trib­uted vari­able) passes a cer­tain crit­i­cal total. This par­a­digm matches data from twin stud­ies, adop­tion stud­ies, pedi­gree and fam­ily stud­ies, and has enjoyed immense suc­cess in recent decades with the advent of genome sequenc­ing. (And we could also use it to explain other fic­tional genetic sce­nar­ios, like 9 or the 3 pony races in .10) Intelligence/IQ in par­tic­u­lar fits this par­a­digm well and the con­sen­sus is that it is highly poly­genic, addi­tive, much of the rel­e­vant genetic vari­ants are com­mon ones, and while rare vari­ants & de novo muta­tions are usu­ally respon­si­ble for cases of severe retar­da­tion, most below-av­er­age intel­li­gence is sim­ply the lower end of a con­tin­u­um, and there are few or no rare vari­ants which cause extremely high intel­li­gence or which offer a large boost in intel­li­gence.

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, 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.

Alternative paradigms

So is it all just non­sense?

Well, there is an alter­na­tive par­a­digm of genet­ics, which may be more famil­iar to most read­ers: the , trace­able to and his peas, but taken up with great enthu­si­asm by Amer­i­cans. In Mendelian genet­ics, the focus is over­whelm­ingly on sin­gle genetic vari­ants with large effects, which since they come in pairs can have sim­ple addi­tive dose-re­sponse effects (0/1/2 copies), non-ad­di­tive effects such as (one copy is enough to cause the trait), “reces­sive” (two copies are required), and lead to com­pli­cated inher­i­tance pat­terns where a trait may dis­ap­pear but then pop up many gen­er­a­tions lat­er, or where sev­eral genetic vari­ants may only have a par­tic­u­lar effect when all of them are present simul­ta­ne­ously (“”). Mendelian genet­ics applies well to a num­ber of rare human dis­eases, and a few odd­ball traits, but works par­tic­u­larly well in agri­cul­tural and sci­en­tific set­tings, where it can be demon­strated vividly and used to track muta­tions and inves­ti­gate their effects, among many other things. The devel­op­ment of Mendelian genet­ics thus lead to a noto­ri­ously bit­ter aca­d­e­mic dis­pute between the bio­me­tri­cians and the Mendelians, because nei­ther side was wrong: there clearly were Mendelian traits which were busily being exper­i­men­tally demon­strated in plants and flies and mice, but it was also clear that Mendelian approaches could­n’t account for traits like height. (For more back­ground, see Provine’s The Ori­gins of The­o­ret­i­cal Pop­u­la­tion Genet­ics, Gilham’s A Life of Sir Fran­cis Gal­ton, and Paul & Spencer 1995.) The feud was only par­tially resolved by R.A. Fish­er’s famous uni­fi­ca­tion demon­strat­ing that the con­tin­u­ous traits could be seen as sim­ply the sum of indefi­nitely many genetic vari­ants each of which acted in a Mendelian man­ner. (In par­tic­u­lar, Mendelian­ism was avidly adopted by Amer­i­can eugeni­cists, who pro­ceeded to inter­pret traits like low intel­li­gence or alco­holism or schiz­o­phre­nia as being sin­gle Mendelian genetic vari­ants, often reces­sive, rather than being part of a con­tin­uum in which suffer­ers merely have bad luck and some­what lower aver­age num­ber of favor­able vari­ants. While under a bio­met­rics par­a­digm, it would be about equally effec­tive to try to increase intel­li­gence by increas­ing the fer­til­ity of more intel­li­gent peo­ple ver­sus decreas­ing the fer­til­ity of lower intel­li­gence, under Amer­i­can Mendelian­ism run amok, all low intelligence/mental illness/disease ‘must’ be due to reces­sives as proven by slop­py, biased—or per­haps even fal­si­fied—­pedi­gree charts of arbi­trar­ily dichotomized traits, and as increas­ing the fer­til­ity of high­-trait peo­ple is largely futile then, in the absence of any kind of genetic test­ing coer­cive gov­ern­men­t-backed ster­il­iza­tion approaches then become the nat­ural approach—e­spe­cially for the strong lib­eral pro­gres­sive tra­di­tion in Amer­ica which was an advo­cate of gov­ern­ment inter­ven­tion to reshape soci­ety eg Pro­hi­bi­tion or min­i­mum wage.)

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, 1⁄23 = 1⁄8 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.

See Also


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 inde­pen­dent 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:

Tad Stone & Soryu Aleksi 2-gene mod­el, Pun­nett square
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:

    When I had fin­ished Wal­ter’s book, through the pub­lisher I wrote a long let­ter to him, telling him how impressed I was and ask­ing about his back­ground—cer­tain it was aca­d­e­mic, that he was a pro­fes­sor of inter­na­tional rela­tions, or phi­los­o­phy, or anthro­pol­ogy at some pres­ti­gious British uni­ver­si­ty. Some weeks lat­er, I received a reply thank­ing me for my kind words and telling me I was lit­er­ally the only one to write to him about his book. As for his back­ground, no he was­n’t a pro­fes­sor. He had grad­u­ated from Sand­hurst (the British West Point) and after a very long career retired as a colonel. He had been a com­bat infantry offi­cer, fight­ing mainly to pro­tect the Empire—in Africa, the Mid­dle East, Asia, wherever; and finally in World War I and II. Over all these years of fight­ing and see­ing war in the raw and coun­tries gripped in the throes of war, this sex­ual theme grad­u­ally dawned upon him. He vowed that when his career ended he would write a book about it. Which he did, but was unable to get a pub­lish­er, so he dipped into his mea­ger pen­sion and had 500 copies printed at his own expense, most of which never sold.

  2. “Frank Her­bert LP: the Ban­quet Scene”:

    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).

  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.

  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.”

  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.”

  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.↩︎