mat]ial f ! ,google ,pr9t project47 ,,biology ,,by ,,edmund ,,bee*] ,,wilson ,,pr(essor ,,( ,,zoology ,,columbia ,,univ]sity ,new ,york ,,! ,,columbia ,,univ]sity ,,press #aijh ,,biology ,a ,,lecture ,,deliv]$ ,,at ,,columbia ,,univ]sity ,,9 ,,! ,,s]ies ,,on ,,sci5ce1 ,,philosophy ,,& ,,art ,,novemb] #bj1 #aijg ,,biology ,,by ,,edmund ,,bee*] ,,wilson ,,pr(essor ,,( ,,zoology ,,columbia ,,univ]sity ,new ,york ,,! ,,columbia ,,univ]sity ,,press #aijh ,,copyright1 #aijh1 by ,,! ,,columbia ,,univ]sity ,,press4 ,set up1 & publi%$ ,m>*1 #aijh4 ,,biology ,i mu/ at ! outset rem>k t among ! _m sci5ces t are occupi$ ) ! /udy ( ! liv+ _w !re is no "o t may prop]ly lay exclusive claim to ! "n ( ,biology4 ,! ^w does n1 9 fact1 d5ote any p>ticul> sci5ce b is a g5]ic t]m appli$ to a l>ge gr\p ( biological sci5ces all ( :i* alike are conc]n$ ) ! ph5om5a ( life4 ,to pres5t 9 a s+le address1 ev5 9 rudim5t>y outl9e1 ! specific results ( ^! sci5ces is obvi\sly an impossible task1 & "o t ,i h no int5tion ( attempt+4 ,i %all (f] no m ?an a k9d ( preface or introduction to ^? :o w speak aft] me on ! biological sci5ces ( physiology1 botany & zoology2 & ,i %all conf9e x to :at seem to me ! mo/ ess5tial & *>act]i/ic ( ! g5]al problems t[>ds :i* all l9es ( biological inquiry mu/ soon] or lat] conv]ge4 ,x is ! g5]al aim ( ! biological sci5ces to le>n some?+ ( ! ord] ( nature 9 ! liv+ _w4 ,p]haps x is n amiss to rem>k t ! biologi/ may n hope to solve ! ultimate problems ( life any m ?an ! *emi/ & physici/ may hope to p5etrate ! f9al my/]ies ( exi/5ce 9 ! non-liv+ _w4 ,:at he c d is to obs]ve1 comp>e & exp]im5t ) ph5om5a1 to resolve m complex ph5om5a into simpl] compon5ts1 & to ?is ext5t1 z he says1 to expla9 !m2 b he kn[s 9 advance t 8 explanations w nev] be 9 ! full s5se ( ! ^w f9al or complete4 ,inve/igation c d no m ?an pu% =w>d ! limits ( k4 ,! task ( ! biologi/ is a d\ble "o4 ,8 m imm$iate ef=t is to inquire into ! nature ( ! exi/+ organism1 to asc]ta9 9 :at measure ! complex ph5om5a ( life z !y n[ appe> are capable ( resolution into simpl] factors or compon5ts1 & to det]m9e z f> z he c :at is ! relation ( ^! factors to o!r natural ph5om5a4 ,x is (t5 practically conv5i5t to consid] ! organism z pres5t+ two diff]5t aspects--a /ructural or morphological "o1 & a functional or physiological--& biologi/s (t5 call !mselves accord+ly morphologi/s or physiologi/s4 ,morphological inve/igation has 9 ! pa/ l>gely foll[$ ! me?od ( obs]vation & comp>ison1 physiological inve/igation t ( exp]im5t2 b x is "o ( ! be/ signs ( progress t 9 rec5t ye>s ! fact has come cle>ly into view t morphology & physiology are really insep>able1 & 9 consequ5ce ! di/9ctions betwe5 !m1 9 respect bo? to subject matt] & to me?od1 h l>gely disappe>$ 9 a great] community ( aim4 ,morphology & physiology alike 7 pr(\ndly trans=m$ by ! introduction into biological /udies ( ! g5etic or hi/orical po9t ( view by ,d>w91 :o did m ?an any o!r to e/abli% ! fact1 suspect$ by _m e>li] naturali/s1 t exi/+ vital ph5om5a are ! outcome ( a def9ite process ( evolution2 & x 0 he :o fir/ fully br\ght home to u h[ defective & "o-sid$ is our view ( ! organism s long z we d n consid] x z a product ( ! pa/4 ,x is ! second & p]haps great] task ( ! biologi/ to /udy ! organism f ! hi/orical po9t ( view1 consid]+ x z ! product ( a cont9u\s process ( evolution t has be5 9 op]ation s9ce life began4 ,9 its wide/ scope ?is g5etic inquiry involves n only ! evolution ( high] =ms f l[] ones1 b also ! /ill l>g] "q ( ! primordial relation ( liv+ ?+s to ! non-liv+ _w4 ,"h is involv$ ! possibility s /rik+ly express$ _m ye>s ago by ,tyndall 9 t eloqu5t passage 9 ! ,belfa/ address1 :]e he decl>$ himself driv5 by an intellectual necessity to cross ! b\nd>y l9e ( ! exp]im5tal evid5ce & to disc]n 9 non-liv+ matt]1 z he said1 ! promise & pot5cy ( e =m & quality ( t]re/rial life4 ,?is intellectual necessity 0 creat$ by a conviction ( ! cont9uity & consi/5cy ( natural ph5om5a1 :i* is almo/ insep>able f ! sci5tific attitude t[>ds nature4 ,b ,tyndall's words /ood aft] all = a confession ( fai?1 n = a /atem5t ( fact2 & !y so>$ f> above ! t]ra firma ( ! actual evid5ce4 ,at ! pres5t day we too may f9d ourselves logically driv5 to ! view t liv+ ?+s fir/ arose z a product ( non-liv+ matt]4 ,we mu/ fully recognize ! extraord9>y progress t has be5 made by ! *emi/ 9 ! artificial syn!sis ( comp\nds =m]ly kn[n only z ! direct products ( liv+ protoplasm4 ,b x mu/ also be admitt$ t we are /ill :olly )\t evid5ce ( ! orig9 ( any liv+ ?+1 at any p]iod ( ! e>?'s hi/ory1 save f "s o!r liv+ ?+2 & aft] m ?an two c5turies ,r$i's aphorism omne vivum e vivo reta9s to-day its full =ce4 ,x is my impression !re=e t ! "t has n yet come :5 hypo!ses reg>d+ a diff]5t orig9 ( life c be consid]$ z practically useful4 ,if ,i h ! tem]ity to ask y\r att5tion to ! fundam5tal problem t[>ds :i* all l9es ( biological inquiry soon] or lat] lead u x is n ) ! delusion t ,i c contribute any?+ new to ! prolong$ discussions & controv]sies to :i* x has giv5 rise4 ,i desire only to indicate 9 :at way x affects ! practical ef=ts ( biologi/s to ga9 a bett] und]/&+ ( ! liv+ organism1 :e!r reg>d$ z a gr\p ( exi/+ ph5om5a or z a product ( ! evolution>y process2 & ,i %all speak ( x1 n 9 any ab/ract or speculative way1 b f ! /&po9t ( ! work+ naturali/4 ,! problem ( :i* ,i speak is t ( organic me*anism & its relation to t ( organic adaptation4 ,h[ 9 g5]al are ! ph5om5a ( life relat$ to ^? ( ! non-liv+ _w8 ,h[ f> c we pr(itably employ ! hypo!sis t ! liv+ body is ess5tially an automaton or ma*9e1 a configuration ( mat]ial p>ticles1 :i*1 l an eng9e or a piece ( clockwork1 owes its mode ( op]ation to its physical & *emical con/ruction8 ,x is n op5 to d\bt t ! liv+ body is a ma*9e4 ,x is a complex *emical eng9e t applies ! en]gy ( ! food-/uffs to ! p]=mance ( ! "w ( life4 ,b is x some?+ m ?an a ma*9e8 ,if we may imag9e ! physico-*emical analysis ( ! body to be c>ri$ ?r\gh to ! v end1 may we expect to f9d at la/ an unkn[n some?+ t transc5ds su* analysis & is nei!r a =m ( physical en]gy nor any?+ giv5 9 ! physical or *emical configuration ( ! body8 ,%all we f9d any?+ correspond+ to ! usual popul> conception--:i* 0 also along ! view ( physiologi/s--t ! body is animat$ by a specific vital pr9ciple1 or vital =ce1 a dom9at+ ar*u t exi/s only 9 ! realm ( organic nature8 ,if su* a pr9ciple exi/s1 !n ! me*ani/ic hypo!sis fails & ! fundam5tal problem ( biology becomes a problem sui g5]is4 ,9 its be>+ on man's place 9 nature ?is "q is "o ( ! mo/ mom5t\s ) :i* natural sci5ce has to deal1 & x has occupi$ ! att5tion ( ?9k+ m5 9 e age4 ,i _c trace its hi/ory1 b x w be wor? our :ile to place side by side ! words ( ?ree ( ! great lead]s ( mod]n sci5tific & philosophic ?\ght4 ,! say+ has be5 attribut$ to ,desc>tes1 ,give me matt] & ,i w con/ruct ! _w--mean+ by ?is ! liv+ _w z well z ! non-liv+2 b ,desc>tes specifically except$ ! human m9d4 ,i d n "k :e!r ! great ,fr5* philosoph] actually us$ ^! p>ticul> words1 b !y express ! ess5ce ( ! me*ani/ic hypo!sis t he adopt$4 ,kant utt]ly repudiat$ su* a conception 9 ! foll[+ well kn[n passage3 ,x is q c]ta9 t we _c become adequately acqua9t$ ) organiz$ creatures & _! hidd5 pot5tialities by means ( ! m]ely me*anical pr9ciples ( nature1 mu* less c we expla9 !m2 & ?is is s c]ta9 t we may boldly ass]t t x is absurd = man ev5 to make su* an attempt or to hope t a ,newton may "o day arise :o w make ! production ( a blade ( grass compreh5sible to u accord+ to natural laws t h n be5 ord]$ by design4 ,su* an insight we mu/ absolutely d5y to man4 ,/ill1 9 ano!r place ,kant admitt$ t ! facts ( comp>ative anatomy give u a ray ( hope1 h[ev] fa9t1 t some?+ may be accompli%$ by ! aid ( ! pr9ciple ( ! me*anism ( nature1 )\t :i* !re c be no sci5ce 9 g5]al4 ,x is int]e/+ to turn f ?is to ! bold & aggressive ass]tion ( ,huxley3 ,liv+ matt] diff]s f o!r matt] 9 degree & n 9 k9d1 ! microcosm repeats ! macrocosm2 & "o *a9 ( causation connects ! nebul\s orig9 ( suns & planet>y sy/ems ) ! protoplasmic f\ndations ( life & organization4 ,d n expect me to decide :]e su* le>n$ doctors disagree2 b ,i w at ?is po9t v5ture on "o comm5t :i* may s\nd ! key-note ( ?is address4 ,p]haps we %all f9d t 9 ! long run & 9 ! l>ge s5se ,kant 0 "r2 b x is c]ta9 t to-day we "k v mu* m ab\t ! =mation ( ! liv+ body1 :e!r a blade ( grass or a man1 ?an did ! naturali/s ( ,kant's "t2 & = bett] or = worse ! human m9d seems to be s con/itut$ t x w cont9ue its ef=ts to expla9 su* matt]s1 h[ev] difficult !y may seem to be4 ,b ,i return to our m specific inquiry ) ! rem>k t ! hi/ory ( physiology 9 ! pa/ two hundr$ ye>s has be5 ! hi/ory ( a progressive re/riction ( ! notion ( a vital =ce or vital pr9ciple )9 n>r[] & n>r[] limits1 until at pres5t x may seem to _m physiologi/s t no room = x rema9s )9 ! limits ( our biological philosophy4 ,"o aft] ano!r ! vital activities h be5 %[n to be 9 great] or less degree explicable or compreh5sible consid]$ z physico-*emical op]ations ( v>i\s degrees ( complexity4 ,e physiologi/ w ma9ta9 t we _c "n "o ( ^! activities1 n ev5 ?\ght1 t is n c>ri$ on by a physical me*anism4 ,he w ma9ta9 fur!r t 9 mo/ cases ! vital actions are n m]ely accompani$ by physico-*emical op]ations b actually consi/ ( !m2 & he may g s f> z def9itely to ma9ta9 t we h no evid5ce t life itself c be reg>d$ z any?+ m ?an _! sum total4 ,he is able to br+ =w>d cog5t evid5ce t all modes ( vital activity are c>ri$ on by means ( en]gy t is set free 9 protoplasm or its products by means ( def9ite *emical processes collectively kn[n z metabolism4 ,:5 ! matt] is r$uc$ to its l[e/ t]ms1 life1 z ?us view$1 seems to h its root 9 *emical *ange2 & we c und]/& h[ an em95t ,g]man physiologi/ (f]s u a def9ition or *>act]ization ( life t runs3 ,! life-process consi/s 9 ! metabolism ( proteids4 ,i ask y\r p>ticul> att5tion to ?is def9ition s9ce ,i n[ wi% to contra/ ) x ano!r & v diff]5t "o4 ,i %all introduce x to y\r att5tion by ask+ a v simple "q4 ,we may admit t dige/ion1 = example1 is a purely *emical op]ation1 & "o t may be exactly imitat$ outside ! liv+ body 9 a glass flask4 ,my "q is1 h[ does x come to pass t an animal has a /oma*8--&1 pursu+ ! inquiry1 h[ does x happ5 t ! human /oma* is practically incapable ( dige/+ cellulose1 :ile ! /oma*s ( "s l[] animals1 su* z ! goat1 readily dige/ ?is sub/ance8 ,! e>li] naturali/s1 su* z ,l9naeus1 ,cuvi] or ,agassiz1 7 ready ) a reply :i* seem$ s simple1 adequate & f9al t ! plodd+ mod]n naturali/ _c repress a feel+ ( envy4 ,9 _! view plants & animals are made z !y 7 orig9ally creat$1 ea* accord+ to its k9d4 ,! biologi/ ( to-day views ! matt] diff]5tly2 & ,i %all give 8 answ] 9 ! =m 9 :i* ,i n[ & !n make x to a /ud5t :o may *ance to ask :y an insect has six legs & a spid] eight1 or :y a yell[bird is yell[ & a bluebird blue4 ,! answ] is3 ,= ! same reason t ! elephant has a trunk4 ,i tru/ t a c]ta9 rugg$ p$agogical virtue 9 ?is reply may atone = its lack ( elegance4 ,! elephant has a trunk1 z ! insect has six legs1 = ! reason t su* is ! specific nature ( ! animal2 & we may ass]t ) a degree ( probability t am\nts to practical c]ta9ty t ?is specific nature is ! outcome ( a def9ite evolution>y process1 ! nature & causes ( :i* x is our trem5d\s task to det]m9e to su* ext5t z we may be able4 ,b ?is does n yet t\* ! mo/ ess5tial side ( ! problem4 ,:at is mo/ significant is t ! clumsy1 %ort-neck$ elephant has be5 end[$--by nature1 z we say--) precisely su* an organ1 ! trunk1 z he ne$s to comp5sate = 8 lack ( flexibility & agility 9 o!r respects4 ,if we are ask$ :y ! elephant has a trunk1 we mu/ answ] because ! animal ne$s x4 ,b does su* a reply 9 itself expla9 ! fact8 ,evid5tly n4 ,! "q :i* sci5ce mu/ seek to answ]1 is h[ came ! elephant to h a trunk2 & we d n prop]ly answ] x by say+ t x has develop$ 9 ! c\rse ( evolution4 ,x has be5 well said t ev5 ! mo/ complete k ( ! g5ealogy ( plants & animals w\ld give u no m ?an an ance/ral portrait-gall]y4 ,we mu/ det]m9e ! causes & conditions t h coop]at$ to produce ?is p>ticul> result if our answ] is to con/itute a true sci5tific explanation4 ,& evid5tly he :o adopts ! ma*9e-!ory z a g5]al int]pretation ( vital ph5om5a mu/ make cle> to u h[ ! ma*9e 0 built be=e we c admit ! validity ( 8 !ory1 ev5 9 a s+le case4 ,our app>5tly simple "q z to :y ! animal has a /oma* has ?us reveal$ to u ! full magnitude ( ! task ) :i* ! me*ani/ is confront$2 & x has br\ght u to t "p ( our problem t is conc]n$ ) ! nature & orig9 ( organic adaptations4 ,)\t t>ry+ to attempt a def9ition ( adaptation ,i w only emphasize ! fact t _m ( ! great naturali/s1 f ,ari/otle onw>d1 h recogniz$ ! purposeful or design-l quality ( vital ph5om5a z _! mo/ ess5tial & fundam5tal *>act]i/ic4 ,h]b]t ,sp5c] def9$ life z ! cont9u\s adju/m5t ( int]nal relations to ext]nal relations4 ,x is "o ( ! be/ t has be5 giv51 ?\gh ,i am n sure t ,pr(essor ,brooks has n improv$ ^u x :5 he says t life is response to ! ord] ( nature4 ,?is seems a long way f ! def9ition ( ,v]worn1 h]et(ore cit$1 z ! metabolism ( proteids4 ,to ?is ,brooks opposes ! tell+ epigram3 ,! ess5ce ( life is n protoplasm b purpose4 ,)\t attempt+ adequately to illu/rate ! nature ( organic adaptations1 ,i w direct y\r att5tion to :at seems to me "o ( _! mo/ /rik+ features reg>d$ f ! me*ani/ic position4 ,?is is ! fact t adaptations s (t5 run c\nt] to direct or obvi\s me*anical conditions4 ,nature is cramm$ ) devices to protect & ma9ta9 ! organism aga9/ ! /ress ( ! environm5t4 ,"s ( ^! are giv5 9 ! obvi\s /ructure ( ! organism1 su* z ! t5drils by means ( :i* ! climb+ plant su/a9s itself aga9/ ! action ( gravity or ! w9ds1 ! protective %ell ( ! snail1 ! protective colors & %apes ( animals1 & ! l4 ,any /ructural feature t is useful because ( its con/ruction is a /ructural adaptation2 & :5 su* adaptations are giv5 ! me*ani/ has = ! mo/ "p a relatively easy task 9 8 int]pretation4 ,he has a f> m difficult knot to dis5tangle 9 ! case ( ! s-call$ functional adaptations1 :]e ! organism modifies its activities 7& (t5 also its /ructure7 9 response to *ang$ conditions4 ,! nature ( ^! ph5om5a may be illu/rat$ by a few examples s *os5 z to =m a progressive s]ies4 ,if a spot on ! sk9 be rubb$ = "s "t ! fir/ result is a direct & obvi\sly me*anical "o2 ! sk9 is worn away4 ,b if ! rubb+ be cont9u$ long en\gh1 & is n too sev]e1 an indirect effect is produc$ t is precisely ! opposite ( ! initial direct "o2 ! sk9 is replac$1 becomes ?ick] ?an be=e1 & a callus is produc$ t protects ! spot f fur!r injury4 ,! heal+ ( a w\nd involves a simil> action4 ,aga91 remove "o kidney or "o lung & ! rema9+ "o w 9 "t enl>ge to assume1 z f> z x is able1 ! functions ( bo?4 ,if ! leg ( a salam&] or a lob/] be amputat$1 ! w\nd n only heals b a new leg is reg5]at$ 9 place ( t :i* has be5 lo/4 ,if a flatworm be cut 9 two1 ! front piece gr[s out a new tail1 ! h9d piece a new head1 & two p]fect worms result4 ,f9ally1 x has be5 f\nd 9 c]ta9 cases1 includ+ animals z highly organiz$ z salam&]s1 t if ! egg be sep>at$ into two p>ts at an e>ly p]iod ( developm5t ea* "p develops into a p]fect embryo animal ( half ! usual size1 & a pair ( tw9s results4 ,9 ea* ( ^! cases ! a/oni%+ fact is t a me*anical injury sets up 9 ! organism a complicat$ adaptive response 9 ! =m ( op]ations :i* 9 ! end c\nt]act ! initial me*anical effect4 ,x is no d\bt true t some:at simil> self-adju/m5ts or responses may be said to take place 9 c]ta9 non-liv+ me*anical sy/ems1 su* z ! sp9n+ top or ! gyroscope2 b ^? t occur 9 ! liv+ body are ( su* g5]al occurr5ce1 ( su* complexity & v>iety1 & ( s design-l a quality1 t !y may fairly be reg>d$ z among ! mo/ *>act]i/ic ( ! vital activities4 ,x is precisely ?is *>act]i/ic ( _m vital ph5om5a t r5d]s _! accurate analysis s difficult & complex a task2 & x is l>gely = ?is reason t ! biological sci5ces1 z a :ole1 /ill /& f> beh9d ! physical sci5ces1 bo? 9 precision & 9 complet5ess ( analysis4 ,:at is ! actual work+ attitude ( naturali/s t[>ds ! g5]al problem t ,i h endeavor$ to outl9e8 ,x w\ld be a piece ( presumption = me to speak = ! body ( work+ biologi/s1 & ,i w !re=e speak = only "o ( !m4 ,x is my own conviction t :atev] be ! difficulties t ! me*ani/ic hypo!sis has to face1 x has e/abli%$ itself z ! mo/ useful work+ hypo!sis t we c at pres5t employ4 ,i d n mean to ass]t t x is adequate1 or ev5 true4 ,i believe only t we %\ld make use ( x z a work+ program1 because ! hi/ory ( biological rese>* proves x to h be5 a m effective & fruitful means ( advanc+ k ?an ! vitali/ic hypo!sis4 ,we %\ld !re=e cont9ue to employ x = ?is purpose until x is cle>ly %[n to be unt5able4 ,:e!r we mu/ 9 ! end adopt x w dep5d on :e!r x proves ! simple/ hypo!sis 9 ! l>ge s5se1 ! "o mo/ 9 h>mony ) our k ( nature 9 g5]al4 ,if su* is ! outcome1 we %all be b\nd by a deeply ly+ in/9ct t is almo/ a law ( our intellectual be+ to accept x1 z we h accept$ ! ,cop]nican sy/em r ?an ! ,ptolemaic4 ,i believe ,i am "r 9 say+ t ! attitude ,i h indicat$ z a m or less p]sonal "o is also t ( ! body ( work+ biologi/s1 ?\gh !re are "s conspicu\s exceptions4 ,9 endeavor+ to illu/rate h[ ?is "q actually affects rese>* ,i w (f] two illu/rative cases1 "o ( :i* may indicate ! fruitfulness ( ! me*ani/ic conception 9 ! analysis ( complex & app>5tly my/]i\s ph5om5a1 ! o!r ! nature ( ! difficulties t h 9 rec5t ye>s l$ to attempts to re-e/abli% ! vitali/ic view4 ,! fir/ example is giv5 by ! s-call$ law or pr9ciple ( ,m5del 9 h]$ity4 ,! pr9ciple reveal$ by ,m5del's wond]ful discov]y is n %[n 9 all ! ph5om5a ( h]$ity & is probably ( m or less limit$ application4 ,x possesses h[ev] a pr(\nd significance because x gives almo/ a demon/ration t a def9ite1 & p]haps a relatively simple1 me*anism mu/ lie beh9d ! ph5om5a ( h]$ity 9 g5]al4 ,h]$it>y *>act]s t con=m to ?is law und]go comb9ations1 disassociations & recomb9ations :i* 9 c]ta9 way sugge/ ^? t take place 9 *emical reactions2 & l ! latt] !y con=m to def9ite quantitative rules t are capable ( ari?metical =mulation4 ,?is analogy mu/ n be press$ too f>2 = *emical reactions are individually def9ite & fix$1 :ile ^? ( ! h]$it>y *>act]s involve a =tuit\s elem5t ( su* a nature t ! num]ical result is n fix$ or con/ant 9 ! individual case b foll[s ! law ( probability 9 ! aggregate ( individuals4 ,nev]!less1 x is possible1 & has already become ! cu/om1 to designate ! h]$it>y organization by symbols or =mulas t resemble ^? ( ! *emi/ 9 t !y imply ! quantitative results ( h]$ity t foll[ ! union ( comp\nds ( kn[n composition4 ,quantitative pr$iction--n precisely accurate1 b 9 accordance ) ! law ( probability--has ?us become possible to ! biological exp]im5t] on h]$ity4 ,i w give "o example ( su* a pr$iction made by ,pr(essor ,cun 9 exp]im5t+ on ! h]$ity ( color 9 mice 7see ! foll[+ table74 ,! exp]im5t ext5d$ ?r\gh ?ree g5]ations4 ,( ! f\r gr&p>5ts ?ree 7 pure :ite alb9os1 id5tical 9 outw>d appe>ance1 b ( diff]5t h]$it>y capacity1 :ile ! f\r? 0 a pure black m\se4 ,! fir/ pair ( gr&p>5ts consi/$ ( an alb9o ( gray ance/ry1 ,,ag1 & "o ( black ance/ry1 ,,ab4 ,! second pair consi/$ ( an alb9o ( yell[ ance/ry1 ,,ay1 & a black m\se1 ,,cb4 ,! result ( ! fir/ union1 ,,ag x ,,ab is to produce aga9 pure :ite mice ( ! composition ,,agab4 ,! second union1 ,,ay x ,,cb is to produce mice t appe> pure yell[1 & h ! =mula ,,aycb4 ,:at1 n[1 w be ! result ( unit+ ! two =ms ?us produc$--i4e4 ,,agab ,,aycb8 ,cun's pr$iction 0 t !y %\ld yield eight diff]5t k9ds ( mice1 ( :i* f\r %\ld be :ite1 two yell[1 "o black & "o gray4 ,! actual aggregate result ( su* unions1 repeat$ly p]=m$1 comp>$ ) ! !oretic expectation1 is %[n 9 ! =ego+ table4 ,z w be se51 ! correspond5ce1 ?\gh close1 is n absolutely exact1 yet is ne> en\gh to prove ! validity ( ! pr9ciple on :i* ! pr$iction 0 bas$1 & we may be c]ta9 t _h a mu* l>g] numb] ( ^! mice be5 re>$ ! correspond5ce w\ld h be5 /ill clos]4 ,i h purposely select$ a some:at complicat$ example1 & "t w n admit ( a full explanation ( ! mann] 9 :i* ?is p>ticul> result 0 rea*$4 ,i w h[ev] attempt to give an indication ( ! g5]al ,m5delian pr9ciple by means ( :i* pr$ictions ( ?is k9d are made4 ,?is pr9ciple appe>s 9 its simple/ =m 9 ! behavior ( two contra/+ *>act]s ( ! same g5]al type--= in/ance two colors1 su* z gray & :ite 9 mice4 ,if two animals1 :i* %[ respectively two su* *>act]s are br$ toge!r1 only "o ( ! *>act]s 7kn[n z ! dom9ant7 appe>s 9 ! (fspr+1 :ile ! o!r 7kn[n z ! recessive7 disappe>s f view4 ,9 ! next g5]ation1 obta9$ by bre$+ ^! hybrids toge!r1 bo? *>act]s appe> sep>ately & 9 a def9ite ratio1 !re be+ 9 ! long run ?ree individuals t %[ ! dom9ant "* to "o t %[s ! recessive4 ,?us1 9 ! case ( gray & :ite mice1 ! fir/ cross is always gray1 :ile ! next g5]ation includes ?ree grays to "o :ite4 ,?is is ! fundam5tal ,m5delian ratio = a s+le pair ( *>act]s2 & f x may readily be d$uc$ ! m complicat$ comb9ations t appe> :5 two or m pairs ( *>act]s are consid]$ toge!r4 ,su* comb9ations appe> 9 def9ite s]ies1 ! nature ( :i* may be work$ out by a simple me?od ( b9omial expansion4 ,by ! use ( ?is pr9ciple a/oni%+ly accurate num]ical pr$ictions may be made1 ev5 ( r complex comb9ations2 & fur!rmore1 new comb9ations may be1 & h be51 artificially produc$1 ! numb]1 "* & h]$it>y capacity ( :i* are kn[n 9 advance4 ,! fundam5tal ratio = a s+le pair ( *>act]s is expla9$ by a v simple assumption4 ,:5 a dom9ant & a recessive "* are associat$ 9 a hybrid1 ! two mu/ und]go 9 "s s5se a disjunction or sep>ation 9 ! =mation ( ! g]m-cells ( ! hybrid4 ,?is takes place 9 a q def9ite way1 exactly half ! g]m-cells 9 ea* sex receiv+ ! pot5tiality ( ! dom9ant "*1 ! o!r half ! pot5tiality ( ! recessive4 ,?is is r\ghly express$ by say+ t ! g]m-cells are no long] hybrid1 l ! body 9 :i* !y arise1 b be> "o "* or ! o!r2 & al?\gh 9 a te*nical s5se ?is is probably n precisely accurate1 x w suffici5tly answ] our purpose4 ,if1 n[1 x be assum$ t f]tilization takes place =tuit\sly--t is t union is equally probable betwe5 g]m-cells be>+ ! same "* & ^? be>+ opposite *>act]s1--! obs]v$ num]ical ratio 9 ! foll[+ g5]ation foll[s accord+ to ! law ( probability4 ,?us is expla9$ bo? ! =tuit\s elem5t t diff]5tiates ^! cases f exact *emical comb9ations1 & ! def9ite num]ical relations t appe> 9 ! aggregate ( individuals4 ,gr&p>5ts ,,ag 7:ite7 ,,ab 7:ite7 ,,ay 7:ite7 ,,cb 7black7 --------- ----------- ,p>5ts ,,agab 7:ite7 ,,aycb 7yell[7 ---------------------- ,obs]v$ ,calculat$ ,,agay ,,abay 7,:ite7 #ha #gf ,,agab ,(fspr+ ---------------,,abab ,,agcy 7,yell[7 #cd #ch ,,abcy ,,abcb 7,black7 #bj #ai ,,agcb 7,gray7 #af #ai ---- ---- #aea #aeb ,n[1 ! po9t t ,i desire to emphasize is t "o or two v simple me*ani/ic assumptions give a lum9\sly cle> explanation ( ! behavior ( ! h]$it>y *>act]s accord+ to ,m5del's law1 & at "o /roke br+ ord] out ( ! *aos 9 :i* facts ( ?is k9d at fir/ sight seem to be4 ,n less significant is ! fact t direct microscopical inve/igation is actually reveal+ 9 ! g]m-cells a physical me*anism t seems adequate to expla9 ! disjunction ( *>act]s on :i* ,m5del's law dep5ds2 & ?is me*anism probably gives u also at lea/ a key to ! long /&+ riddle ( ! det]m9ation & h]$ity ( sex4 ,^! ph5om5a are !re=e becom+ intelligible f ! me*ani/ic po9t ( view4 ,f any o!r !y appe> z an insoluble enigma4 ,:5 su* progress z ?is is be+ made1 h we n a "r to believe t we are employ+ a useful work+ hypo!sis8 ,b let u n[ turn to a second example t w illu/rate a class ( ph5om5a :i* h ?us f> almo/ :olly elud$ all attempts to expla9 !m4 ,! "o t ,i select is at pres5t "o ( ! mo/ enigmatical cases kn[n1 namely1 ! reg5]ation ( ! l5s ( ! eye 9 ! tadpoles ( salam&]s4 ,if ! l5s be remov$ f ! eye ( a "y tadpole1 ! animal proce$s to manufacture a new "o to take its place1 & ! eye becomes z p]fect z be=e4 ,t su* a process %\ld take place at all is rem>kable en\gh2 b f a te*nical po9t ( view ?is is n ! extraord9>y feature ( ! case4 ,:at fills ! embryologi/ ) a/oni%m5t is ! fact t ! new l5s is n =m$ 9 ! same way or f ! same mat]ial z ! old "o4 ,9 ! normal developm5t ( ! tadpole f ! egg1 z 9 all o!r v]tebrate animals1 ! l5s is =m$ f ! out] sk9 or ectod]m ( ! head4 ,9 ! replacem5t ( ! l5s aft] removal x arises f ! cells ( ! iris1 :i* =m ! edge ( ! optic cup1 & ?is orig9ates 9 ! embryo n f ! out] sk9 b z an outgr[? f ! bra94 ,z f> z we c see1 nei!r ! animal itself nor any ( its ance/ors c h _h exp]i5ce ( su* a process4 ,h[1 !n1 c su* a p[] h be5 acquir$1 & h[ does x inh]e 9 ! /ructure ( ! organism8 ,if ! process ( repair be due to "s k9d ( intellig5t action1 z "s naturali/s h suppos$1 :y %\ld n ! high] animals & man possess a simil> useful capacity8 ,to ^! que/ions biology c at pres5t give no reply4 ,9 ! face ( su* a case ! me*ani/ mu/ simply confess himself = ! "t be+ br\ght to a /&/ill2 & !re are "s able naturali/s :o h 9 rec5t ye>s argu$ t by ! v nature ( ! case su* ph5om5a are incapable ( a rational explanation along ! l9es ( a physico-*emical or me*ani/ic analysis4 ,^! writ]s h urg$1 accord+ly1 t we mu/ po/ulate 9 ! liv+ organism "s =m ( controll+ or regulat+ ag5cy :i* does n lie 9 its physico-*emical configuration & is n a =m ( physical en]gy--some?+ t may be ak9 to a =m ( intellig5ce 7consci\s or unconsci\s71 & to :i* ! physical en]gies are 9 "s fa%ion subject4 ,to ?is suppos$ factor 9 ! vital processes h be5 appli$ su* t]ms z ! entele*y 7f ,ari/otle71 or ! psy*oid2 & "s writ]s h ev5 employ$ ! ^w s\l 9 ?is s5se--?\gh ?is te*nical & limit$ use ( ! ^w %\ld n be conf\nd$ ) ! m usual & g5]al "o ) :i* we are famili>4 ,views ( ?is k9d repres5t a return1 9 "s measure1 to e>li] vitali/ic conceptions1 b diff] f ! latt] 9 t !y are an outcome ( def9ite & exact exp]im5tal "w4 ,!y are !re=e (t5 spok5 ( collectively z neo-vitalism4 ,x is n my purpose to ent] ^u a detail$ critique ( ?is doctr9e4 ,to me x seems n to be sci5ce1 b ei!r a k9d ( metaphysics or an act ( fai?4 ,i mu/ own to complete inability to see h[ our sci5tific und]/&+ ( ! matt] is 9 any way advanc$ by apply+ su* names z entele*y or psy*oid to ! unkn[n factors ( ! vital activities4 ,!y are words t h be5 writt5 into c]ta9 spaces t are o!rwise blank 9 our record ( k1 & z f> z ,i c see no m ?an ?is4 ,x is my impression t we %all d bett] z inve/igators ( natural ph5om5a frankly to admit t !y /& = matt]s t we d n yet und]/&1 & cont9ue our ef=ts to make !m kn[n4 ,& h we any o!r way ( do+ ?is ?an by obs]vation1 exp]im5t1 comp>ison & ! resolution ( m complex ph5om5a into simpl] compon5ts8 ,i say aga91 ) all possible emphasis1 t ! me*ani/ic hypo!sis or ma*9e-!ory ( liv+ be+s is n fully e/abli%$1 t x may n be adequate or ev5 true2 yet ,i c only believe t until e o!r possibility has realty be5 exhau/$ sci5tific biologi/s %\ld hold fa/ to ! work+ program t has creat$ ! sci5ces ( biology4 ,! vitali/ic hypo!sis may be held1 & is held1 z a matt] ( fai?2 b we _c call x sci5ce )\t misuse ( ! ^w4 ,:5 we turn1 f9ally1 to ! g5etic or hi/orical "p ( our task1 we f9d ourselves confront$ ) precisely ! same g5]al problem z 9 case ( ! exi/+ organism4 ,biological inve/igators h long s9ce ceas$ to reg>d ! fact ( organic evolution z op5 to s]i\s discussion4 ,! transmutation ( species is n an hypo!sis or assumption1 x is a fact accurately obs]v$ 9 our laboratories2 & ! !ory ( evolution is only que/ion$ 9 ! same v g5]al way 9 :i* all ! great g5]alizations ( sci5ce are held op5 to modification z k advances4 ,b x is a v l>ge "q :at has caus$ & det]m9$ evolution4 ,"h1 too1 ! fundam5tal problem is1 h[ f> ! process may be me*anically explicable or compreh5sible1 h[ f> x is susceptible ( =mulation 9 physico-*emical or me*ani/ic t]ms4 ,! mo/ ess5tial "p ( ?is problem relates to ! orig9 ( organic adaptations1 ! production ( ! fit4 ,) ,kant1 ,cuvi] & ,l9naeus believ$ ?is problem sci5tifically insoluble4 ,lam>ck attempt$ to f9d a solution 9 8 !ory ( ! inh]itance ( ! effects ( use1 disuse & o!r acquir$ *>act]s2 b 8 !ory 0 insecurely bas$ & also begg$ ! "q1 s9ce ! p[] ( adaptation ?r\gh :i* use1 disuse & ! l produce _! effects is precisely t :i* mu/ be expla9$4 ,d>w9 believ$ he _h f\nd a p>tial solution 9 8 !ory ( natural selection1 & he 0 hail$ by ,haeckel z ! biological ,newton :o _h set at naught ! obit] dictum ( ,kant4 ,b ,d>w9 himself did n consid] natural selection z an adequate explanation1 s9ce he call$ to its aid ! subsidi>y hypo!ses ( sexual selection & ! inh]itance ( acquir$ *>act]s4 ,if ,i correctly judge1 ! fir/ ( ^! hypo!ses mu/ be consid]$ z ( limit$ application if x is n s]i\sly discr$it$1 :ile ! second c at be/ receive ! ,scot* v]dict1 n prov54 ,9 any case1 natural selection mu/ fight its own battles4 ,latt] day biologi/s h come to see cle>ly t ! inadequacy ( natural selection lies 9 its failure to expla9 ! orig9 ( ! fit2 & ,d>w9 himself recogniz$ cle>ly en\gh t x is n an orig9ative or creative pr9ciple4 ,x is only a condition ( survival1 & h5ce a condition ( progress4 ,b :e!r we conceive ) ,d>w9 t selection has act$ ma9ly ^u slight individual v>iations1 or ) ,de,vries t x has op]at$ ) l>g] & m /able mutations1 any adequate g5]al !ory ( evolution mu/ expla9 ! orig9 ( ! fit4 ,n[1 "u ! !ory ( natural selection1 pure & simple1 adaptation or fitness has a m]ely casual or accid5tal "*4 ,9 itself ! fit has no m significance ?an ! unfit4 ,x is only "o out ( _m possibilities ( *ange1 & evolution by natural selection resolves itself into a s]ies ( lucky accid5ts4 ,= ,agassiz or ,cuvi] ! fit is t :i* 0 design$ to fit4 ,= natural selection1 pure & simple1 ! fit is t :i* happ5s to fit4 ,i1 = "o1 am unable to f9d a logical flaw 9 ?is conception ( ! fit2 & p]haps we may be =c$ to accept x z suffici5t4 ,b ,i believe t naturali/s d n yet re/ cont5t ) x4 ,d>w9 himself 0 repeat$ly br\ght to a /&/ill1 n m]ely by specific difficulties 9 ! application ( 8 !ory1 b also by a c]ta9 in/9ctive or temp]am5tal dissatisfaction ) su* a g5]al conclusion z ! "o ,i h indicat$2 & _m able naturali/s feel ! same difficulty to-day4 ,:e!r ?is be ju/ifi$ or n1 x is und\bt$ly ! fact t few work+ naturali/s feel conv9c$ t ! problem ( organic evolution has be5 fully solv$4 ,"o ( ! que/ions ) :i* rese>* is s]i\sly engag$ is :e!r v>iations or mutations are indet]m9ate1 z ,d>w9 on ! :ole believ$1 or :e!r !y may be 9 great] or less degree det]m9ate1 proce$+ along def9ite l9es z if impell$ by a vis a t]go4 ,! !ory ( or?og5esis1 propos$ by ,naegeli & ,eim]1 makes ! latt] assumption2 & x has f\nd a consid]able numb] ( adh]5ts among rec5t biological inve/igators1 includ+ "s ( our own colleagues1 :o h made important contributions to ! inve/igation ( ?is fundam5tal "q4 ,x is too soon to v5ture a pr$iction z to ! ultimate result4 ,t evolution has be5 or?og5etic 9 ! case ( c]ta9 gr\ps1 seems to be well e/abli%$1 b _m difficulties /& 9 ! way ( its acceptance z a g5]al pr9ciple ( explanation4 ,! unc]ta9ty t /ill hangs ov] ?is "q & t ( ! h]$ity ( acquir$ *>act]s be>s witness to ! unsettl$ /ate ( op9ion reg>d+ ! :ole problem1 & to ! inadequacy ( ! attempts ?us f> made to f9d its consi/5t & adequate solution4 ,"h1 too1 accord+ly1 we f9d ourselves confront$ ) wide gaps 9 our k :i* op5 ! way to vitali/ic or transc5d5tal !ories ( developm5t4 ,i ?9k we %\ld resi/ ! temptation to seek su* refuge4 ,x is m ?an probable t !re are factors ( evolution /ill unkn[n4 ,we c b seek = !m4 ,no?+ is m c]ta9 ?an t life & ! evolution ( life are natural ph5om5a4 ,we mu/ approa* !m1 & z f> z ,i c see mu/ attempt to analyze !m1 by ! same me?ods t are employ$ 9 ! /udy ( o!r natural ph5om5a4 ,! /ud5t ( nature c d no m ?an /rive t[>ds ! tru?4 ,:5 he does n f9d ! :ole tru? !re is b "o gospel = 8 salvation--/ill to /rive t[>ds ! tru?4 ,he kn[s t ea* =w>d /ep on ! highway ( discov]y w br+ to view a new horizon ( regions /ill unkn[n4 ,x w be an ill day = sci5ce :5 x c f9d no m fields to conqu]4 ,& s1 if y ask :e!r ,i look to a day :5 we %all "k ! :ole tru? 9 reg>d to organic me*anism & organic evolution1 ,i answ]3 ,no6 ,b let u g =w>d4 ,,columbia ,,univ]sity ,,press ,a ,s]ies ( tw5ty-two lectures descriptive 9 unte*nical language ( ! a*ievem5ts 9 ,sci5ce1 ,philosophy & ,art1 & indicat+ ! pres5t /atus ( ^! subjects z concepts ( human k1 are be+ deliv]$ at ,columbia ,univ]sity1 dur+ ! academic ye> #aijg-aijh1 by v>i\s pr(essors *os5 to repres5t ! sev]al dep>tm5ts ( in/ruction4 ,,ma!matics1 by ,cassius ,jackson ,keys]1 ,adra9 ,pr(essor ( ,ma!matics4 ,,physics1 by ,erne/ ,fox ,ni*ols1 ,pr(essor ( ,exp]im5tal ,physics4 ,,*emi/ry1 by ,*>les ,f4 ,*&l]1 ,pr(essor ( ,*emi/ry4 ,,a/ronomy1 by ,h>old ,jacoby1 ,ru!rfurd ,pr(essor ( ,a/ronomy4 ,,geology1 by ,james ,furman ,kemp4 ,pr(essor ( ,geology4 ,,biology1 by ,edmund ,b4 ,wilson1 ,pr(essor ( ,zoology4 ,,physiology1 by ,fr$]ic ,s4 ,lee1 ,pr(essor ( ,physiology4 ,,botany1 by ,h]b]t ,maule ,ri*>ds1 ,pr(essor ( ,botany4 ,,zoology1 by ,h5ry ,e4 ,crampton1 ,pr(essor ( ,zoology4 ,,an?ropology1 by ,franz ,boas4 ,pr(essor ( ,an?ropology4 ,,ar*aeology1 by ,james ,rignall ,:eel]1 ,pr(essor ( ,greek ,ar*aeology & ,art4 ,,hi/ory1 by ,james ,h>vey ,rob9son1 ,pr(essor ( ,hi/ory4 ,,economics1 by ,h5ry ,rog]s ,seag]1 ,pr(essor ( ,political ,economy4 ,,politics1 by ,*>les ,a4 ,be>d1 ,adjunct ,pr(essor ( ,politics4 ,,jurisprud5ce1 by ,munroe ,smi?1 ,pr(essor ( ,roman ,law & ,comp>ative ,jurisprud5ce4 ,,sociology1 by ,frankl9 ,h5ry ,gidd+s1 ,pr(essor ( ,sociology4 ,,philosophy1 by ,ni*olas ,murray ,butl]4 ,presid5t ( ! ,univ]sity4 ,,psy*ology1 by ,rob]t ,s4 ,woodwor?1 ,adjunct ,pr(essor ( ,psy*ology4 ,,metaphysics1 by ,fr$]ick ,j4,e4 ,woodbridge1 ,johnsonian ,pr(essor ( ,philosophy4 ,,e?ics1 by ,john ,dewey1 ,pr(essor ( ,philosophy4 ,,philology1 by ,a4,v4,w4 ,jackson1 ,pr(essor ( ,indo-,iranian ,languages4 ,,lit]ature1 by ,h>ry ,?ur/on ,peck1 ,an?on ,pr(essor ( ! ,lat9 ,language & ,lit]ature4 ,^! lectures are publi%$ by ! ,columbia ,univ]sity ,press sep>ately 9 pamphlet =m1 at ! uni=m price ( tw5ty-five c5ts1 by mail tw5ty-eight c5ts4 ,ord]s w be tak5 = ! sep>ate pamphlets1 or = ! :ole s]ies4 ,address ,,! ,,columbia ,,univ]sity ,,press ,columbia ,univ]sity1 ,new ,york 99 99 99 99 99