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    In Mendel 's Mirror

    Philosophica Refectons on Bioogy

    PHILIP KITCHER

    OXOUNIVERSTY PRESS

    2003

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    30 n d's

    35 Fr eampes see K Wessels, Tu I ntton n Dvlopmnt (Menl ark,Cal Benjamn, 1977) espeally apters 7 3 5 and Dnald Ede An Intoutonto Dvlopmntl Boloy (Lndn Blake, 197) espeally apter 3

    3 I atempt t sw w te same perspetve an be rutully appl ed t ter eamples n lp Kter, Eplanatry Unatn, setns 3 and 4 Kter, Abun Sn(Cambrdge, Mass M ress 19 ) apter Kter, Darwn's evement (aper3 ts bk

    37 Te etreme versn redutnsm s deended by Rard Dawkns n Th SlhGn (ew Yr Ord Unversty ress, 197) and Th Extn Phnoyp (San Frans Freeman, 9) Fr an eellent rtque, see Ell tt Sber and Rard C. Lewntn, rtat, Cause, and Gen Seletn, Phloophy of Sn 49 ( 9) 1 571 Mreambtus rms ant redutn sm wt respet t evutn ary tery are advaned nS. J. Guld, Is a ew and General ery Evlutn Emergng? Ploboloy (19)1 19 3 Eldredge and J. Crara, Phylont Ptt n th Evoluton Po(New Yrk Clumba Unversty ress, 19) and Steven M Stanley, Movoluton (SanFrans Freeman, 1 979) a ss early sue sme (but nt all ) later antredutnst temes s Est Mayrs Anml Sp n Evoluton (Cambrdge, Mass. arvard Unversty ress, 1 93) espeally apter 1

    3 Guld's Ontony n Phylony (Cambrdge, Mass Harvard Unversty ress,1 977) prvdes stral l lumnatn bt areas debate abut redutnsm Cntem

    prary antredutnst arguments abt embrylgy are epressed by Wessels, Tu Int-ton, and Ede, Dvlopmntl B oloy See as G. Oster and . lber, Evlutn andBuratn Devepmental rgrams, Evoluton 3 (19) 444459

    39 Oster and lber, Evlutn and B uatn, 44 Te dagram n p. 45 prvdes an equally stragtrward aunt ter antredutnst pstn

    2

    The Hegemony of MolecuarBiology ( 1999)

    Dic Lewoti has poal ha oe iuec e o cotepoa phiosoph ofiolog ha a ohe livig ioogistpal ecause of his illiat a wie-agig coiutios to geeics a evoluioa heo pal ecause of he

    wah a iess he h as exee o philosophes a especial ecause ofhis ow ajo p hiosophica cotiuios Fo eal two ecaes ow wiigs have ee sustaial iee o Dics isights a Ive fou self ght-ig o he sae sie i a of the sae attes But hee have ee ipotaieeces A Dic has oe oe his ow oppositio o vaious popula oc-ies especiall i socioioog a othe fos of geetic eeiis has eeoe aica ha ow hee I have accepte the gou ules of a paila eepise a ague that he alege cocusios ot folow Dic has oewae o sweep awa he eepise as isguie. I eec weve eplae the elatioship ewee eal tweiehcetu Biish socialiss a hei oe evolutioa coutepas i cotieal Euope Kei Haie to Dics Lei. I cathi of o ete wa to hoo his egac to phiosoph of iolog ha o pla it

    agai.Athough his picipal coce aou he cotepoa pacice of olecula

    iolog has cetee o ieas aout geetic causatio Lewoti has a oaeiees i euig what he sees as a Caesia saeg of expaaio issecio. ! e opposig he ieolog" ha we ca stu all the aue eaig he wol up ito iepeet pats a ae coeig oscuatistholis Lewoi cotiues: The pole is to costuct a hi view oeha sees he eie wol eithe as a iissolule whole o with the eualicoect u cue oia view tha a eve level he wol is ae upof its a pieces tha ca e isolae a that have popeties tha ca esuie i isoaio. I effec Lewoi was o esist the hegeo of olecu-

    a ioog wihout apsig io sicis o o I I wha foows I shal toaicuae a aieucioist view tha sees oecula stuies as a ipoatpa of ut ot he whoe of coepoa ioog suspec hat his view wilassig olecua iolog a oe ipoa ole ha ha wh ich Lewoi woulfavo.

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    32 In ndl s iro

    I

    Athogh the ea of the hegeony of oear oogy i oen peente yphioopher n te of the notion of ntertheoreti reton a ore oonforaton n io ogy on wo ephaze two thee

    (H 1 ) A organ are opoe of oee(H2) Rea (rgoro, opete) expanaton of th e properte of ng thng

    trae thoe propetie to interation aong oee (fe i to eexpane at the oear ee")

    (H 1) i a ti he ea eate ente on ( H2)roponent of (H2 ) enage a eforate ioogy in wh ih t he propetie of

    ogani ae ere n a angage that aow for appaton of ioheiaprnipe to ere ioogia oneqene The rt ojeton i that the eniage eration ae notanae eae we ant proe the approprate angage The eon that een f we ha h eraton, they wo not awaye expanatoy One ery oio way to poe the rt to ak how we o eer

    hope to proe a iohea expiaton of h noton a speces pedao anecossem Bt the e an e o re hapy poe f we fo on what ee toe a h oe poing ae for the hegeonit to wit genet

    one two tateent fo a a geneti

    ( G ) Han eng wh o ae ho ozygo fo the ik ing aee expeenee at ow ee of oxygen

    (G2 ) Gene on ierent hoooe or fienty fa apart on the aehroooe aort n epenenty

    Hegeonit an point to (G ) a a patia e t a age, (G2) repe-ent tota fae

    Sine the ate 1940, oogit hae known how the heogoin traniean tranate fro the ikng aee er fo that tanate fo the noaaee gnoring opexite of eeopent, they an teat eythroyte a a ontaining heogon an ing pinipe of hetry, they an then how that,ner onition of ow oxygen, the tant heogon wo ten to p nway that poe the haraterti rig eent that ge ikee anea itnae O ne t eognze that thee eent wo ten to ok narrow apiare we hae an expanaton fo (G ) Athogh that expanation int fy o-eaea that wee gnore the eeopenta poe entey an haetaken a ery aoee ew of the petinent phyioogyt a tat

    The eniage expanation wo tat wth the eriation of the nora an

    tant eqene of ano ai o the peation of NA eqene anthe genet oe Of ore, to ahiee that we ony nee the eqene peaton of patia aee By ontat, when we tn to (G), the eforation inoheia ter wo eqie a peiation of the genea popety beng agene hat , what nee e i opeton of the open entene

    x a gene if an ony if x

    T oy of oar Biooy 3 3

    Now rey we know somehng aot how to opete th A n iportant neeay onton on gene that they e egent of NA o RNA; t of oethere are ot of egent of DNA an RNA (ot of the n fat) that are notgene The tak th to ent th e propery that itingihe the ght egentof nei a fro the wong one

    Thee i an portant ontrant on oing thi a ontaint that oeteneognze f the pinipe of heitry are to e epoye in eringthe eforate iooga onon, then we nee a haatezation ofthe petinent entitiein th ntane genethat wi eh wth tanarway of rawng hea oneqene hat eant that the haraterzatonwi hae to e sucua ientng gene in ter of ther ontitent oeeHene a propoa to pe gene a ntiona entitie fo exape thoene a egent that are tanie an tranate to proe poypeptie,wont ee the hegeont trn ( ho note that th propoa i ao inaeqate eae t wongy exe egent that happen to oe ther regatoryegion)

    No ttra peiation of the genera notion of a gene renty aai-

    ae hat not eae the pojet of nng one wont e iportant to onteporay oea oogy On the ontrary, a ae of eqene ata po in inetigato hntng for gene wo weoe a yteat etho o f earhingthe ong tng of A, , G a n T he et they an o i to pk ot OpenReaing Frae (ORF)eatiey ong trethe one y tat an topoonteatng thee a anate an then hekng to ee f they an oeoeponng RNA

    Fthe, a the intiae of genoe eoe ore eent the poiitieof pt gene, oerapping gene, trnate eqene tat are ti aoate wthregatory egion, eqene that hae ot ther regatory regon, eeegene, an o foth ake any trtra an fntona riteria, wth ORF ope

    with fntona RNA a the entra ntane an wth perphea exapeette y onention that oete ary fro ty to tyThe t toe with (G2) th that the reqe oiene ienta-

    tion arent aaae now arge that een i f they wee, a eration of (G2)fo prnipe of hetry wont e expanatory

    egn nrety wth a otationa tory n 1 1 , John Athnot, a phyian,pointe ot that the preio eightytwo yea n onon were a ae" that i,

    in eah of thee yea there wa a peponeane of ae irth n ononhing h ning n the Phosophca Tansacons of he Roa SoceArthnot aate the poaiity that th oe y han e an n ing thatpoaity to e inte, he hake p the phenoenon to ine roieneLet agne wo ear harate who ty to ge a eer expanaton

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    34 In endels Mio

    Fr he Ma Mehan MM) H gng prinipe ha fe i o eexpaine a he moear ee" an he p h o work n oeing an ex-panaion. More exay he poie a repe for an expanaion amiing haeae of h gnoane of ron of eai, e an go frher. The repe na foow

    Start wth the rt brth of 1 6 Go bak to the o pulatory at that began the preg-

    nany reultng n th bth Gve a moleular haraterzaton of the rumtane that preeded fetlzaton From th haraterzaton derve a onluonabout the perm that wa norporated nto the zygote. Contnue wh the moleula aount of the oure of the pregnany and brth. You have now explanedthe ex of the rt nfant of 1 6

    Contnue n the ame fahon wth the eond brth the thrd brth and oon When you are done add up the total fo both exe You now have a om-plete expanation of why 16 wa a male yea

    Repeat the ame poedure fo ubequent year untl you reah 179 Stop.You now have a omplete explanaton for why all year are mae

    Aay yo on.To ee why, oner or nex haraer, he Sen e SexRao Theor (S ST).

    She poee from R. A. Fiher nig h ao he eoon of ex ao

    In pee wthout peal ondton of matng nludng Homo pen) f the exrato at exual maturty depat fom 1 1 there wll be a eetve advantage to atendeny to produe membe of the underrepreented ex (th wll how up nterm of nreaed number of expeted grand offprng. In huma populatonthat are ufently large, we hould thu expet the ex rato at exual matuty toapproxmate 1 : (the more loey the ager the populaton even n the even-teenth entury London had a lage populaton.

    If one ex more vulnerable than the other to motalty between brth andexual matuty then th at ex wll have to be produed n greater number f theex rato at exual matuty to be 1 1 In human beng male are more vulnerable to prepubertal death Thu the brth ex rato kewed toward male.

    I aim ha he SST wo gie a eer expanaon han he MM een if heae o aay eier he ea ar of he reaon i ha he SST aonhow ha Arhno aa ae no ke. The ignan pon fo or prpoe ha we on nee he mae of aena moea mnae we wan o eehow a regary n nare par of a roa genera paern.

    A NoSoMa Mehani ( NSM M) wo ee he poin an moi hi po-on Reognzng he fa ha he e expanaon of he phenomenon oeng hogh he moea ea he mgh ak wheher here are ieren faeof h aon ha moear reearh migh minae Inee here are. SSTe wh year are mae or more exay key o e mae for age popa-on B ha eae open how ao popaion of

    Hm apienn her

    way o rogh erim . NS MM w propoe a ion of expanaory aor.Aer ha hown he hae o he exanaon hyoog an ee ino hemehanm o Ybae eraion ae Ybearng erm aer? are agna on-ion moe ie o he oage o Ybeang erm? are here poymorphm in

    he egemony o olecula Biology 35

    hman popaon?) eang eenay o a mo ear neranng of he mompoan proee. I ern o he gnane of hi pon eow

    Now ak o GZ) We an enage a onerpar o MM raey rying ohow how gory hema ea ye he nepen en aomen of gene pro-e ha he gene are on een hromoome or are fen far apar onhe ame hromoome). B here no reaon o hnk ha hee eo wo eany more mnaing han MM. Fo hee ao a onerpar o SST, whoe

    expanaion goe a foowConder the followng knd of proe a poe fo png and eption)There are ome ba entte whh ome n pa For eah par there a orrepondene relaton between the part of one member of the par and the part ofthe othe member. At the rst stage of the process, the entities are placed in anen. Whle they are n the a rena they an exhange egment o that the partof one member of a par are eplaed by the orrepondng part of the othermember, and conversely. Aer exactly one round of exchanges, one and only one

    member of each pair is drawn from the arena and placed in the winner's box.In any PSpoe the hane that mall egment that belong to member

    of derent pairs or that are sufciently far apart on members of the same par wil

    be found in the winner's box are ndependent of one another. (G2) holds because

    the distbution of chromosomes to gametes a t meiosis is a PS-process.

    Th, I m a f expanaon of (GZ) an expanaon ha pren enieyfrom he ha gene are mae of. Uneranng he proa regaeha goern he ranmion of gene a maer of eeng ha ranmon i a Sproe an rreean wheher he gene ae mae of ne a or of wheee.

    The onon we ogh o raw ha ome mporan iooga regar-ie anno e apre n he angage of moear oogyo, moe iy, na moear ooga angage ha rer ef o rra noion an hahee regarie are fy expane who grnng o moear ea Annghene Hegemon (EH ) ogh o appreae he poin, reognzng he nee

    o aor fnona onep an am nong hoe onep from raonaarea of ioogy. EH w n, howeer, ha here are mporan moear ieao he fnonay haraerze regare eon onernng he meh-anm of Yae ferizaon or he moear n epnnng of he parng ofhomoogo homoome a meo Tha pon I arge ae, i ore. If EH am o, h oweer hee may e a frher propoa Ahogh gh for mo-ea oogy o aor fnona nigh from he aia area of oogyhe negaon n hee area are nneeary; from now on moearoogy i a he new oogy we nee. I now wan o gge ha we ogh o eh hegemon yeanng.

    IVn 917 Ary Wenworh Thomon pbhe a remakae book. ike TomSopar ay homana, homon eane or he mahemai o he

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    36 In endels Mio

    anmate wo n eent yea mathematal ologt hae egn to eazehompon' pogam an the elt l gget a e w of eeopmental ologythat oth agn an mpotant pae to moea te an eepen the halenge to the heg emony of moela oogy

    Fo peent ppoe we'l only nee to one the mot elementay pat ofa few majo appoahe an want to emphaze that the mple moe lee ae elaoate n mh moe te eon eg n wth the e of Ldemeyer sstems-ytem to h aateze the gowth of plant A strg OLsystem a tpe V P whee V a oaay an nta tng an P aet of poton le A devemeta seqece n an Oytem eqene oftng whoe t meme the ntal tng an h that the + 1 t meme otane fom the th meme y applyng al the poton le that an eappe to th e th tng So fo example one th e ytem

    1: aPI a abP : a ba

    g : b a P4 b a

    Wthn th ytem we an otan the folowng eelopmenta eqene

    aabbaaaababbabaabaa

    h fomam an e e to moe the eelopment of a mtel la lamentfon n the legeen atea Aabaea catea (the a an epeent ffeent type of el an the fxe how the poay; ee (ge 21 .6

    n genea ytem moe the eelopment of pant y ppong that theeae elementay ologal poee that ae apple eely to etan k n oftte nttely n a gowng plant a patla kn of tte ge way toa eent kn of tte an the poe of epaement epeente y apoton le Note that th teat the eeopment of plant n an extemeyatat way penng fom the et al of the ype of poee nole h the gowth of two qte eent plant o e epeente y the ame ytemf n the one ntane a poton le ale fo the eplaement of a pata

    kn of noa el wth a anh an n the othe t pee that a ey eenttype of e ho e one y a pe geometa te of etan knof e.

    o a e atat teatment o e n eeopment we an tn to themathemat of on eqaton npe y wo of Aan ng a nme of

    (r-) r8 br( . ) l8 bl

    e Hegeoy o Moea Bioogy 37

    F R 1 Devepmet me (Anben cenul) multed ug DOytem Frm . Ldemyer d rkewz, The Alorihmic Beuy of Plns (NewYrk Sprger, 1 99

    ologt hae exploe the pol ty that ao kn of patten o e geneate thogh atatonh to" ytem Ba to th appoah the thoghtthat a el m ght tat to poe geate onentaton of a patla molelethat th molele ol then ffe nto ajaent el wth the ataton anon pomptng the poton of an nhto n the ong ae an entete of el eahe a nfom teay tate. Mh moe nteetng the poly that a loal epate fom nfomty ge e to a tale paen

    One agan ' fo on the ey mplet ytem We magne an nteatonamong wo molee the atato (whoe onentaton a an the nh to(whoe onentaton b . he nh to ame to e mh moe aplythan the atato an ao to eah eqlm amot ntantaneoy he onentaton ae goene y the eental eqaton

    /t = aZ -a'b/t = aZ - b

    t not ha to ee that thee a teay tate at a b 1 t pole to geneate apaen howee f one el n an aay ha a ghty neae atato onen

    taton. Beae of the ampton that the nh to e aply t epon tothe aeage onentaton of the atato an th eman taly ontant.Hene the atato wl ontne to neae ( ne y the t eqaton the tmeeate of t onentaton wll e pote) One the neae eomefenty age thee l e an eet on the aeage ent to poe

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    38 edes iror

    a

    ctor

    Inhibitor

    d

    -- ._./

    c

    -_en ='

    G R 22 Pattrn formaton by autocataly and ongrang bton a Ractonchm A actator catay t ow producto ad that of t hghly dffug atagotth nbtor. b Stag pattr formato ar a local prturbaton Computrmulato n a nar array of cll. A omognou dtrbuton of bot ubtanc untab. A mnut ocal ncra of t actato - grow urtr ut a tady tat rachd wch lfactaton an th urroundng clou of nbtor a balacd[S22] rm . Mnart he Alothmc Beauty of Sea Shell Nw York Sprgr998)

    h bor o so he rocess e hs oba a seady sae wh a locally h gh co-cerao of acvaor ad reavely eevaed evels of hbor esewhere (seege 2.2).8

    Sose he ha h e growh of seashels s a rocess wh ch coceraosof acvao olecles ad of hb o olecles are govered by a coed se ofaral dereal eaos ha allow fo orva seady saes If he deecebewee hese coceraos s assocaed wh geao ( or ossby wh d f-ferealy dreced cel growh ad dvs o he s ossle o desad howaes of varos ds eerge Has Mehard has exlored a wde rage ofgrowh rocesses showg how he aers fod a dverse cass of seshells cabe geeraed fro arclar ses of eao. Hs a ayss w hle les s absrac hahe deayerPrsewcz eae of las coes o esc d fo heoeclar deals. Two shells g h resl fro he sae growh rocessaccre-o of ew aeral a he argad gh cofo o he sae se of der-eal eaos eve hogh he olecles ha lay he roles of acvaor adhbor ae dee he wo cases. I gh eve r o ha he oesace he relaosh bewee he oleces odces a geed aerwhle he oher ha reaoh yel ds a aer of reef (rdges ad valeys ohe shell srface).

    My las exale ches frher he dreco of dshed absracoMehards ae o d a geeral se of odes for shel aer rages orewdey ha a edeavor of aes Mrray o exla he dversty of aalacoa aers ollowg g Mrray cosders a reaco dso syse gov-eed by he followg eaos

    he egemony o olecla Biology 39

    I GU R E 23 Ect of body urfac ca on t patal pattrn formd by th ractondffuon mchanm Mammalan Coat Pattrn "ow th Lopar ot t Spot wthparamtr alu 1 5 , K 0 1 2 5 P 1 3 a 103 b 77 tady tat Us = 23 V 24)d = 7. Doman dmno rlat drctly to y a Y< 0 b Y 0 5 ; c y= 25 ; d y=250; Y= 250; f Y= 3000; g Y 5000 From Muray Mathematcal Boloy wYork: Sprngr 19 89)

    ala {(u, ) u

    la g(u ) + dz(u, ) = a u h(u, )g(u ) (b- )- h(u )h(u, ) u+u+Ku

    Here u a d are oeca coceraos a, b, d ad are desoless ara-eers d beg he ao of dffso coefces ad a scalg aaee ( varesas he aea of he sface o wh ch h e aer s beg ad dow) . Mray showsha whe d > 1 ocesses coforg o hese eaos ca gve rse o saalyhoogeeos aers. heher sch a aer occrs ad wha fo aesdeeds o he vae of A hs vae creases he c haracer of he gea

    o ae chages fro fo o bcolored o bloched o sred o soed(see gre 2 3 ) .s ow ossble o arve a a ceve heoe Asse ha aa la coa

    ags are geeaed fo reacos aog checals ha sas he gvesyse of eqaos or a gve vale of d > 1 , rovded ha alows fo boh

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    0 In endel s Mio

    pe an poe paern heell e a hrehol y* ch ha fo y *, herelan paen will e poe an for y < Y* he paen will e pe Theae of y fo an ana oy will alway e geaer han he ale for ha analal (oe are alway gger in aea han al) Hence i can' happen ha heale for he a le aoe he hreho an he ale fo he oy eow hehehol n oher wor we hae he heore"

    Alhogh hee can e poe ana w h rpe ail here c an e rpeana wh poe ail

    Mray oel of aaian coa paern h expan a eglary we n nnare

    Coner now hree iffeen propoal for eeach n eeopenal ologyThe he oigina hegeon poiion gge ha e of oganceelopen ae e pre y arng wh a coplee neranng of hegenec connng wh an negaon of he way n which ieren geneae aciae an ppree an on hi a explong he oleclar ae ofcellar ieeniaon n igh of he coneaon aie n eaer econhegeon ay concee he n ee for ppleenng he oo p" analye

    wh fnconal concep rawn fo claica phyoogy (an ohe raional -ciplne) ee no reaon for frher fncona anaye ha o no aen o heoleclar ea The exaple e choen fo he aheacal y of ee-open are nene o how ha h conceion oo iie A h an oeenlighene appoach wol iew he aheaical an oleclar poga aworking n ane

    A he o concree leel of aheaica analyi heori ay ry oforlae eena eqaion ha goern he neacon of olecle whoeiene hey on know eekng n h way o nean a general paern ofeeopen in oe grop of ogana in Mrray eaen of aalancoa paern wh prey re ao po an rpe. The reearch hen poehe proe of yng o n he hypohecal olecle an qie poiy of

    elng he oel o accooae he coplexe ha eergeAoe h a ee of analyi epreene y Men ha work on eahell

    where he ephai on a faly of relae oe Hee heori aep ocoe oe geneal reglariie coniing n confot o a faly of e oferenal eqaon. Fo h enepre o ccee wll e poran o howhow paiclar phenoena in paricar ogan are goene y parclarebe of he faly an n coneqence o ppeen he aheacaleail wh he ienicaon of he pernen olecle

    en or e arac i a tye o f analy ha foce on foral feare of gowhwho efeence o a pecc inerpeaion of he ol ogca procee epreeney foa ranforaon an who peccaon o eqaon ha ae o be a-e. The y of pan growh n er of inenayer ye alow or aophyologca eang" of he procon e nepeaon ha gh e genn e o acocopc pan phyology n e o oe aheacaly cha-acee poce o n e o olecar neacon eay o ee how herecol be a nee eence o abac accon be a he o oral eel

    The egemony o olecula Biology I

    n cla ao ye wh neeae leel of aheaical oogy haeenally are nanae n eale ie of heeogeneo olece n eyren organi.

    f he hegeon lkely ake con n he lo of neranng hrogheron n eal wh conco an falre o repreen geneal reglare haare poan o gowh an fo" he aheacal analy can ealy loe ochwih he iological reale Hegeonic glng ao he eae wh whch one

    can ake p pleang aheacal oe nfai n o eay haa pon The lleele pce of heoring ao eelopen ha haeecoene nee oleclar ae. To a ha o recogne ha heqeon ao nial ech an ha exce olecar o log (paly ofcore ecae hey hae powerfl ool for areng he ) are poran ohfor he conraon of he oe aac oe an fo he ncoerng ofconran on oellng. The ake o hink ha hee ae he lypoan qeon ha once we hae C R here no frher nee of clacawhoe organ" ology. DAcy Thopon ion hol e negae wh heacheeen an progra of conepoary oleclar oogy o generae alileele y of eelopen

    v

    rern in conclon o ewonn concern ao he hegeony of oleclarology expree n h cqe of he Drea of he H an Genoe " Becaeof ch of he propagana ha ha rone he Han Genoe ojec(HG) ha ecoe a yol of he hegeony of olecla ology. n parewonn cqe foce on eren qeon fo hoe ha hae concernee hee H e ee qe correcly ha he a on wh ch he HG wa aerieconi of a o e of clai ao he caal role of gene an ao heexience of a oya roa o a re ole clar e cn e of enoo power There

    no reaon o eee ha when wee appe an eqence he h an genoewe wll neran who we ae." Silarly erely knowing he eqence of oof han gene n going o ell er ch recly gen he fclie of heproenfolng prole he ncerane of eho of racng proein fncono gnorance of eelopenal pahway an o forh The ieae io-ecal pho of he HG wl l e an enoroly enhance aly o ge genece an oh ewonin an hae o ao wheher h i lkely o e ocallyenecal en f n pncple poie o apply he new ean of eng opooe he wefare of cen (a hae arge a oe lengh) econgepengly clear ha he neee afegar ae no lkely o e n place y hee he echnologe oo he akeplace

    B wan o ingh h a of he HG a par of a ocoeical agenafro i oe n coneporay ologca eearch ewonn crqe an heknre reak of hoan an phloophe of oogy 5 coney he eage hahe HG ologcally ge ehe ecae he a of eqence aa wlgenerae ele o becae nexcaby enwne wh a econc

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    2 I Mendel's irror

    research program. These reactions reect a disposition to accept te propagandafor the Hep at face value. t is quite right to point out that there is nothingbiologicaly special about te genome of our own species and to question theegemonist suggestion tat we can proceed from knowledge of sequence data toknowledge of genes and th ence to all manner of bioogi cal un derstandings. Yet teresearch actually conducted under the auspices of the Hep fully absorbs thesepoints

    From a bioogica point of view, the most important work being conducted witHep funding ( or the parallel resea rch carried ou t wit private support, most notabythat of Craig Venter and his colleagues) consists in negrained mapping andsequencing of nonuman organisms, from bacteria to yeast, to nematode worms,and stil in progress, ie s Te fruits of this research are ikel y to make any numberof researc projects in physiology and developmenta bioogy enormously easier incoming decades (as wel a s paving the way for evolutionary ins igts obtained fromcomparison of te genomes of cosely related species) Specicaly, molecularbiologists working on Caenorhabditis elegans Drosophila melanogaster, andDictyostelium discoideum already envisage the possibil ity of identiing major deve-opmenta pathways, possibly pathways tat ave been ighy conserved in the evo

    lutionary process Tere's no automatic route to picking out such pathways but teability to discover wich genes are activated in which cells whic will ow fromcomplete genome sequencing) is ikely to oer important clues.

    There shoud be no ilusi ons that this molecular work can proceed by ignor-ing macrolevel studies of deveopment and physiology. On te contrary the fullexploitation of te sequence data generated by the genome project wil require justte kinds of nctiona studiesincuding matematical modeingthat I aveephasized trougout tis essay.16 Critics of te Hep may be correct in thinkingthat the current balance of research in biology has tipped too far toward this par-ticular moecuar endeavor that it is not the only proect of bioogica vaue It iswrong, owever, to overstate te claim by taking the project to be devoid of bio-logical signicance and to accuse it of commitment to te hegemonist manifesto.

    Provided we have a ric enough repertoire of visions, the dream of say) te fruit-y genome is a dream worth aving.

    Notes

    1 Ths opposton s evdet n many of the contrbutons to The Dialectical Biologist(Cambridge Mass.: Harvard University Press 1986), a voume of essays some of which areby Lewontn alone some by his colleague Rchard Levns and some tha are jointly wtenThe attack on genetic determinism also permeates this volume as wel as surfacing in

    Not in Our Genes (writte with Leon Kamin and Steven Rose [New Yor: Pantheon1984]) and the more recent Bio logy as Ideology( ew York: Harper, 1992). I discuss the vai-

    ees of Lewontn's crtque of genetic determn sm n Ph ilip Ktcher, "Battng the Undead:How nd How ot) to Resst Genetc Determnsm n a etschrit for Dck Lewontin thatesay erte as chater 1 of ths book should be seen as a cop ao pece to my eortshere

    2 ewot Bioogy a Ideoogy 1 5

    The Hegemony of Molecular Bioogy 43

    3 This problem afects the suggestion made n a provocative essay by C . KennethWaters "Genes Made Molecular Philosophy of Science 6 (1994) : 163185

    4 In fact coemporary molecular bolog s permeated by langage that cant berepaced wth an austere physcochemca dom Consder the standard account of transcrp ton One talks of R A polymerases assocat ng th D A. The suggestion of courses that the R A polymerases come close bt ho close s clos enough ? Well thats gongto depend on the conformaton of the DA ad theres o genera sructural creron. neffect molecular bologsts here and esewhere quey take over functonal concepts. Thsmoved Sylva Cu lp and me to suggest hat moecular b olo us out not to be reducbleto molecuar bolog see Culp and Ktcher Theor Structue and Theo Change Coteporary Moecular Bology Britih J oual for the Philosophy of Science, 40 [1989] 459-483

    5 DArcy Thompso On Growth and Fo Cambrdge: Cambrdge Unversty Press19 17 ); an abrdged verson edted by the dstngushed developmental bologst John TylerBonner was publshed by Cambrdge Unversty Press 1961 Thompsons wsh for amathematcal account of development and morpholog s echoed in severa speeches by theherone of Soppard's Arcadia.

    6 Ths example s used as the rst smplest llustraton by Przemyslaw Prusnkewczand Arstd ndenmayer n The Algorihmic Beauy of Plants ew York Sprnger 1990)The example comes from p 5 I hav e slghtly moded the noaton) A usnkewcz and

    Lndenmayer note systems are related to Chomskyan grammars Subsequent examplesreveal the possbtes of far more complex relatons between symbos and bologcal enttes partcularly through processes that draw out shapes dependen on he symbos) contextdependence threedmensionalty probablsc systems and so fort The resultan systemscan smulate the groh of owers ad rees generate the bonacc sprals found n sunowers and model compound leaves among other achevements

    7 Speccall a paper Alan Turng "The Chem ca Bas s of Morphogenesis Philosophical Transactions of the Royal Socie B 23 7 ( 19 52) 3772 Ths essay was wrten shorlybefore Turngs tragc sucde. t s terestng to ponder whether f he had lved the paceof work n mathematcal developmental bolo would have accelerated producng a verderent dstrbuton of work n the current e ld

    8 Ths exemely eementar example s om Hans Menhardt The Algorithmic BeauyoSea Shells New York: Sprnger 1998) As Menh ardt shows much more complex sets ofpartal derental equatons gve rse to a wde varety of patterns ncluding the elaboratebranching and meshwork found n some she ls The case n the ext s he foundaton of asystem that wil y iel d regular str pes.

    9 fact Murray has a ve broad pogram of tryg to uderstand patte formatonbut Il only consider one aspect of it here. The dscusson s drawn om chapter 1 5 of J. DMurray Mathematical Biology New York Sprnger 1989), altough the essentals werealready gve n Murrays "O Patte Formaton Mechansms for Lepdoptera WngPattes and Mammalan Coat Markngs Philosophical Transactions o the Royal SocietyB 29 5 ( 196 1 ) 473496 Murray provded an accessbl e overview n How the Leopad GesIts Spots," Scientifc American 2 5 8 ( 1 988) : 80-87.

    0 Murray provdes a lucd dscsson of these equatons n chapter 14 ofMathemat-ical Biology He notes there that the equatons descrbe the chemca knetcs of a substrate

    nhbon system whch has been studied expermentaly. Real nstanations of the systemare thus kown.1 1 As n other parts of scence technques molecular bology have a lfe of ther

    own sometmes nspng people to pursue queston becase they can be addressed For anllumnatng stdy of the ways n wch struments and expermental skills po sess an nerta

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