Chapter 1 EVOLUTIONARY BIOLOGY

Summary of Chapter 1: EVOLUTION, D. Futuyma, 2

nd
Ed., 2!.
WHAT IS EVOLUTION?
The "ord e#o$ut%on &ome' from the Lat%n "ord evolvere, "h%&h mean' to unfo$d or unro$$.
(To re#ea$ the h%dden potent%a$%t%e'.
(It ha' &ome to mean )&han*e+.
Biological evolution refer' to the &han*e %n the &hara&ter%'t%&' of *roup' of or*an%'m' o#er many
*enerat%on', o#er t%me.
The de#e$opment or ontogeny of an %nd%#%dua$ or*an%'m %' not &on'%dered e#o$ut%on.
Ind%#%dua$ or*an%'m' do not e#o$#e.
,opu$at%on' of or*an%'m' e#o$#e- they under*o descent with modification.
Chan*e' that are pa''ed from one *enerat%on to the ne.t are &on'%dered e#o$ut%onary- the'e are
*enet%& &han*e'.
The re'ur*en&e of ant%/%ot%& re'%'tant d%'ea'e' %' an e.amp$e of natura$ 'e$e&t%on.
BEFORE ARWIN
Dar"%n0' theory of /%o$o*%&a$ e#o$ut%on re#o$ut%on%1ed the pre#a%$%n* "or$d#%e" %n the 2e't that
had e.%'ted from the t%me of ,$ato and 3r%'tot$e.
!lato" 425(645 7. C. 8ree9 ,h%$o'opher.
)Idea+ or )form+ %' rea$ and e#eryth%n* e$'e %' a ref$e&t%on of th%' rea$%ty. The e''en&e or %dea %'
%mperfe&t$y %m%tated /y %t' earth$y repre'entat%on'. The rea$ "or$d %' the "or$d of %dea', and "hat
"e 'ee on earth %' on$y an %m%tat%on of tho'e %dea'. The e''en&e or form of a 'tru&ture &ou$d /e
under'tood from %t' fun&t%on, '%n&e the fun&t%on d%&tated the form. Spe&%e' /e&ame the %n%t%a$
mo$d for a$$ $ater rep$%&ate' of that 'pe&%e'. 3n %nd%#%dua$ or*an%'m ha' %mperfe&t%on'. Var%at%on'
/et"een or*an%'m' of the 'ame 'pe&%e' are a&&%denta$ %mperfe&t%on'.
Theory of e''ent%a$%'m.
A#i$totle" 654(622 7. C. 8ree9 ph%$o'opher.
3r%'tot$e de#e$oped the ,$ato0' &on&ept of %mmuta/$e e''en&e %nto the not%on that 'pe&%e' ha#e
f%.ed propert%e'. 3r%'tot$e propo'ed that the $a't 'ta*e of de#e$opment, the adu$t form, e.p$a%n'
the &han*e' that o&&ur %n the %mmature form' :te$eo$o*%&a$ e.p$anat%on;- the ad#an&ed 'ta*e'
%nf$uen&e the ear$%er 'ta*e'. <e a$'o propo'ed that a$$ $%#%n* form' are $%n9ed %n a pro*re''%on
from %mperfe&t to mo't perfe&t. <e &a$$ed th%' the Scale o% Natu#e :S&a$a Naturae;.
Chr%'t%an thou*ht e$a/orated on ,$aton%& and 3r%'tote$%an ph%$o'ophy that e.%'ten&e %' *ood and
the 8od0' /ene#o$en&e %' &omp$ete, therefore, <e mu't ha#e /e'to"ed e.%'ten&e on e#ery
&reature *%#%n* ea&h %t' o"n e''en&e.
S%n&e order %' 'uper%or to d%'order, 8od0' &reat%on mu't fo$$o" a p$an, e'pe&%a$$y a *radat%on from
%mperfe&t to perfe&t.
Inan%mate o/=e&t' to p$ant' to %n#erte/rate' up to #erte/rate', human, and f%na$$y, an*e$'.
S&a$a Naturae or S&a$e of Nature or the 8reat Cha%n of 7e%n*.
T&e !#inci'le o% !lenitu(e
The $e*a&y of ,$ato to European th%n9%n* a/out the natura$ "or$d ha' /een e.pre''ed a' the ,r%n&%p$e of ,$en%tude
:Lo#e=oy, 1!6>- ?o$fe, 1!5@;. Th%' "a' the /e$%ef that a$$ po''%/$e 9%nd' of th%n*' e.%'t %n the "or$d a$ready and noth%n*
more &an /e &reated. 3r%'tot$eA' $e*a&y "a' more &omp$%&ated /ut ha' /een 'ummar%1ed /y ?o$fe a' the &on&ept of
&ont%nu%ty and *radat%on /et"een ad=a&ent 9%nd' of /e%n* "hen h%erar&h%&a$$y arran*ed :1!5@, p. 6;. To*ether, the
,r%n&%p$e of ,$en%tude and the 8reat Cha%n of 7e%n* $ed to the /e$%ef, from med%e#a$ t%me', that a &ont%nuou' &ha%n
e.tended from the %nan%mate "or$d of non($%#%n* matter, 'u&h a' earth and 'tone', throu*h the an%mate "or$d of p$ant',
1oophyte', and the $o"e't form' of an%ma$ $%fe, up"ard' to the Buadruped' and e#entua$$y throu*h Can to the rea$m' of
an*e$' and f%na$$y to the Chr%'t%an 8od. Th%' /e$%ef a$'o enta%$ed the #%e" that =u't a' noth%n* ne" &ou$d /e &reated-
ne%ther &ou$d anyth%n* /e e.term%nated, '%n&e th%' "ou$d &ountera&t the "%$$ of 8od :?o$fe, 1!5@, p. 1;.
http:DD"e/.e/'&oho't.&omDeho'tDdeta%$E'%dF>d6!562&(@e6&(4eG4(565e(&&G5e&1G/1/
H4'e''%onm*r11I#%dF1Ih%dF1@I/dataFJnNpd8U!K2h#&6Lt/8$2KSK1M2!"KT11aN?$Od/
FaphI3NF!>12>G>2
A#i$totle" t&e Scale o% Natu#e" an( )o(e#n attitu(e$ to ani)al$ * In t&e +o)'any o% Ani)al$
So&%a$ ?e'ear&h, Fa$$, 1!!@ /y Ju$%et C$utton(7ro&9
Linnaeu$" +a#olu$, 1GG(G5, S"ed%'h /otan%'t and ta.onom%'t,
L%nnaeu' %' &on'%dered the founder of the -ino)ial $y$te) o% no)enclatu#e and the or%*%nator
of modern '&%ent%f%& &$a''%f%&at%on of p$ant' and an%ma$'. <e u'ed the 'pe&%e' a' the /a'%& un%t %n
/u%$d%n* h%' 'y'tem up"ard to $ar*er &ate*or%e'. <e *rouped re$ated 'pe&%e' %nto *enera, and
re$ated *enera %nto order', and 'o on. In h%' m%nd, th%' 'pe&%e' had /een re$ated %n the m%nd of the
Creator.
Hutton" .a)e$, 1G2>(!G, S&ott%'h *eo$o*%'t.
<utton formu$ated &ontro#er'%a$ theor%e' of the or%*%n of the earth %n 1G5@. <e "a' of the op%n%on
that the earth mu't /e #ery o$d.
Uni%o#)ita#iani$)/ the do&tr%ne that pa't *eo$o*%&a$ &han*e' %n the earth "ere /rou*ht
a/out /y the 'ame &au'e' a' tho'e no" ta9%n* p$a&e. It 'tre''ed the '$o"ne'' and
*radua$ne'' of rate' of &han*e.
Lyell" Si# +&a#le$, 1G!G(15G@, En*$%'h *eo$o*%'t.
Lye$$ ar*ued %n h%' /oo9 that, at the t%me, pre'ent$y o/'er#a/$e *eo$o*%&a$ pro&e''e' "ere
adeBuate to e.p$a%n *eo$o*%&a$ h%'tory. <e thou*ht the a&t%on of the ra%n, 'ea, #o$&anoe' and
earthBua9e' e.p$a%ned the *eo$o*%&a$ h%'tory of more an&%ent t%me'. Lye$$ &on&$u'%#e$y 'ho"ed
that the earth "a' #ery o$d and had &han*ed %t' form '$o"$y, ma%n$y from &ond%t%on' 'u&h a'
ero'%on. Lye$$ "a' a/$e to date the a*e' of ro&9' /y u'%n* fo''%$' em/edded %n the 'tone a' t%me
%nd%&ator'. Lye$$ he$ped "%n a&&eptan&e of Jame' <uttonA' theory of uni%o#)ita#iani$) and of
Char$e' Dar"%nA' theory of e#o$ut%on.
La)a#c0" .ean Ba'ti$te !ie##e Antoine (e 1onet" +&evalie# (e" 1G44(152!.
Lamar&9 "a' /orn %n 7a1ent%n($e(,et%t, Fran&e. Sent to a Je'u%t '&hoo$ %n 3m%en'.
<e re&e%#ed a &$a''%&a$ edu&at%on unt%$ 1G@!.
That year, h%' father d%ed, and Lamar&9 entered the m%$%tary and /e*an to 'tudy p$ant'. In
1G>5 he $eft m%$%tary 'er#%&e and 'tud%ed med%&%ne %n ,ar%' for four year', dur%n* "h%&h t%me
he /e&ame %ntere'ted %n meteoro$o*y, &hem%'try, and 'he$$ &o$$e&t%n*.
Lamar&9 "a' e$e&ted to the 3&ademy of S&%en&e' and /e&ame an a''o&%ate /otan%'t %n 1G56,
<%' mo't '%*n%f%&ant "or9 "a' done "hen he /e*an to "or9 at the Jard%n du ?o% :P%n*A'
8arden; %n 1G55.
2hen the *arden "a' reor*an%1ed %n 1G!6, Lamar&9A' %dea' he$ped to frame the 'tru&ture of
the ne" Cu'eum of Natura$ <%'tory.
<e /e&ame profe''or of %n'e&t' and "orm' %n the Cu'eum of Natura$ <%'tory, a d%#%'%on he
named %n#erte/rate 1oo$o*y.
Lamar&9A' theoret%&a$ o/'er#at%on' on e#o$ut%on, referred to %n the ear$y 1!th &entury a'
tran'form%'m or tran'mutat%on, pre&eded h%' e.ten'%#e o/'er#at%ona$ "or9 on %n#erte/rate'.
Lamar&9 a&&epted the #%e" that an%ma$' %n nature "ere arran*ed on one &ont%nuou' Q'&a$a
naturaeQ.
F%r't /%o$o*%'t to a&t%#e$y propo'e e#o$ut%on.
Lamar&9 d%ed "%th $%tt$e '&%ent%f%& re&o*n%t%on of h%' "or9 or h%' %dea'. Not unt%$ the 'e&ond
ha$f of the 1!th &entury "ere Lamar&9A' %dea' 'er%ou'$y &on'%dered a*a%n.
LA1AR+2IS1
Lamar&9 propo'ed the pr%n&%p$e of the in&e#itance o% ac3ui#e( c&a#acte#i$tic$.
Lamar&9 /a'ed h%' theory of e#o$ut%on %n part on h%' 'tudy of the fo''%$' of mar%ne %n#erte/rate',
"a' that 'pe&%e' do &han*e o#er t%me.
<e /e$%e#ed, furthermore, that an%ma$' e#o$#e /e&au'e unfa#ora/$e &ond%t%on' produ&e need' that
an%ma$' try to 'at%'fy.
Spe&%e' not on$y /e&ome e.t%n&t /ut a$'o under*o a *radua$ mod%f%&at%on throu*h t%me.
Or*an%'m' ha#e an inne# 'e#%ecting '#inci'le that R
&ou$d 'en'e the need' of the en#%ronment
re'pond /y de#e$op%n* adaptat%on'R
mo't$y from '%mp$e to &omp$e..
The%r en#%ronment doe' not pa''%#e$y a$ter or*an%'m'.
3 &han*e %n the en#%ronment &au'e' &han*e' %n the need' of or*an%'m' $%#%n* %n that
en#%ronment, "h%&h %n turn &au'e' &han*e' %n the%r /eha#%or.
T"o )ec&ani$)$ are %n#o$#ed %n e#o$ut%on:
,r%n&%p$e of u'e and d%'u'e.
Inher%tan&e of a&Bu%red &hara&ter'.
Lamar&9 #%e"ed e#o$ut%on a' a pro&e'' of %n&rea'%n* &omp$e.%ty and Qperfe&t%on,Q not dr%#en /y
&han&e.
QNature, %n produ&%n* %n 'u&&e''%on e#ery 'pe&%e' of an%ma$, and /e*%nn%n* "%th the $ea't
perfe&t or '%mp$e't to end her "or9 "%th the mo't perfe&t, ha' *radua$$y &omp$%&ated the%r
'tru&ture.Q Philosophie zoologique.
2h%$e the me&han%'m of Lamar&9%an e#o$ut%on %' Bu%te d%fferent from that propo'ed /y Dar"%n,
the pred%&ted re'u$t %' the 'ame: adapt%#e &han*e %n $%nea*e', u$t%mate$y dr%#en /y en#%ronmenta$
&han*e, o#er $on* per%od' of t%me.
Lamar&9 d%d not /e$%e#e %n e.t%n&t%on: for h%m, 'pe&%e' that d%'appeared d%d 'o /e&au'e they
e#o$#ed %nto d%fferent 'pe&%e'.
Lamar&9A' Philosophie zoologique ment%on' the *reat #ar%ety of an%ma$ and p$ant form' produ&ed
under human &u$t%#at%on :Lamar&9 e#en ant%&%pated Dar"%n %n ment%on%n* fanta%$ p%*eon'S;- the
pre'en&e of #e't%*%a$, non(fun&t%ona$ 'tru&ture' %n many an%ma$'- and the pre'en&e of em/ryon%&
'tru&ture' that ha#e no &ounterpart %n the adu$t.
Lamar&9 /e$%e#ed that the Earth "a' #ery o$d.
+&a#le$ a#4in" 15!(1552.
Char$e' ?o/ert Dar"%n "a' /orn %n Shre"'/ury, En*$and %n 15!. The 'on of an em%nent
$o&a$ do&tor, Dr ?o/ert Dar"%n, Char$e' "a' /orn %nto a mode't$y "ea$thy fam%$y.
<e "a' the *rand'on of Era'mu' Dar"%n, En*$%'h natura$%'t, and promoter of the %dea of
e#o$ut%on.
Dar"%n 'tud%ed med%&%ne at Ed%n/ur*h Un%#er'%ty from 152@ to 152G.
<e tran'ferred to Chr%'tA' Co$$e*e, Cam/r%d*e %n 1525 "%th the %ntent%on of /e&om%n* a
m%n%'ter %n the Chur&h of En*$and.
<e "a' a med%o&re 'tudent and d%d not $%9e &$a''%&a$ edu&at%on. <%' %ntere't' "ere %n natura$
h%'tory, /otany, *eo$o*y, &o$$e&t%n* and hunt%n*.
In 1561, "%th the he$p of h%' /otany profe''or John <en'$o" and h%' un&$e Jo'%ah
2ed*e"ood, he too9 the po't of natura$%'t on /oard the H,1,S, Beagle on a '&%ent%f%&
m%''%on to South 3mer%&a.
7y the t%me of h%' return %n 156>, he had /e&ome an author%ty on many form' of $%fe.
7et"een 1542 and 1544 he de#e$oped h%' t&eo#y o% natu#al $election, a$thou*h he d%d not
announ&e h%' "or9 unt%$ 15@5.
In 15@! he pu/$%'hed a &on'%dera/$y e.panded #er'%on of h%' re'ear&he' %n the &ontro#er'%a$
Origin of Species by Means of Natural Selection.
<e pu/$%'hed The Descent of Man %n 15G1.
THE VOYAGE OF THE BEAGLE
<.C.S. 7ea*$e, under the &ommand of Capta%n ?o/ert F%t1roy, $eft for ,ata*on%a, South
3mer%&a, %n 1561 "%th Char$e' Dar"%n on /oard.
3t th%' t%me Char$e' d%d not /e$%e#e %n e#o$ut%on, %n&$ud%n* h%' *randfather0' theory, a' any
e#%den&e pre'ented 'o far &ou$d not &on#%n&e h%m.
The 7ea*$e rea&hed South 3mer%&a %n 1562 and Char$e' too9 &are to o/'er#e the f$ora and
fauna.
Char$e' "ent then to 7ueno' 3r%e' "here he 'a" fo''%$' of more an&%ent an%ma$', %n&$ud%n* a
ma'todon.
<e e.per%en&ed a #%o$ent earthBua9e %n Ch%$e that ra%'ed the $and %n 'ome p$a&e' /et"een 2 and
1 feet.
Dar"%n "a' mo't %ntere'ted %n the p$ant' and an%ma$' on the 8a$Tpa*o', a *roup of 1> $ar*e
%'$and' :and many 'ma$$er; off the &oa't of E&uador.
8%ant torto%'e' %nha/%t e#ery one of the %'$and', "h%&h *a#e the %'$and &ha%n %t' name,
from the Span%'h 8a$Tpa*o', mean%n* torto%'e.
Ea&h %'$and ha' %t' o"n type of torto%'e, d%'t%n*u%'ha/$e /y the 'hape and pattern of
%t' 'he$$.
Dar"%n "a' a'ton%'hed that the %'$ander' fe$t that th%' "a' due to the d%fferen&e %n
en#%ronment on ea&h %'$and.
Dar"%n a$'o o/'er#ed the f%n&he', "h%&h #ar%ed %n '%1e and 'hape from %'$and to
%'$and.
The%r /ea9' a$'o #ar%ed depend%n* on "h%&h food they ate and 'ome e#en had e.tra
$on* ton*ue' for *ra//%n* &erta%n type' of food' 'u&h a' %n'e&t', nut' or 'eed'.
I'$and an%ma$' and p$ant' "ere d%fferent to tho'e on the ma%n$and /ut a re$at%on'h%p &ou$d /e 'een.
2hat "a' e#en 'tran*er thou*h, to Dar"%n, "a' the fa&t that or*an%'m' on d%fferent %'$and'
#ar%ed, /ut 't%$$ 'eemed re$ated.
ALFRE RUSSEL WALLA+E" 1526(1!16.
5 January 1526: 3$fred ?u''e$ 2a$$a&e /orn at U'9, Conmouth'h%re.
m%dD$ate 156G: Jo%n' the e$de't /rother 2%$$%am %n 7edford'h%re to $earn the 'ur#ey%n* trade.
2@ 3pr%$ 1545: 2a$$a&e and 7ate' $ea#e En*$and for 3ma1on%an South 3mer%&a to /e*%n a
natura$ h%'tory &o$$e&t%n* e.ped%t%on.
Car&h 15@4: Lea#e' En*$and for the Far Ea't to /e*%n a natura$ h%'tory &o$$e&t%n* e.ped%t%on.
2 3pr%$ 15@4 to 2 Fe/ruary 15>2: Co$$e&t%n* e.ped%t%on %n the Ca$ay 3r&h%pe$a*o.
Fe/ruary 15@5: 2r%te' 'On the Tendency of Varieties to Depart Indefinitely From the
Original Type' and 'end' %t off to Char$e' Dar"%n for &omment.
1 Ju$y 15@5: 2a$$a&e and Dar"%nA' "r%t%n*' on natura$ 'e$e&t%on are pre'ented at a meet%n* of
the L%nnaean So&%ety.
No#em/er 15@!: )On the oological !eography of the "alay #rchipelago$% the paper
de'&r%/%n* 2a$$a&e0' L%ne %' read /efore the L%nnaean So&%ety- Dar"%nA' )On the Or%*%n of
Spe&%e'+ %' pu/$%'hed.
1 3pr%$ 15>2: ?eturn' to En*$%'h 'o%$.
G No#em/er 1!16: D%e' at O$d Or&hard.
QIn Fe/ruary of 15@5, "h%$e 'uffer%n* from an atta&9 of ma$ar%a %n the Co$u&&a' :%t %' not fu$$y &erta%n "h%&h %'$and he
"a' a&tua$$y on, thou*h e%ther 8%$o$o or Ternate 'eem' the $%9e$y &and%date;, 2a$$a&e 'udden$y, and rather
une.pe&ted$y, &onne&ted the %dea' of Thoma' Ca$thu' on the $%m%t' to popu$at%on *ro"th to a me&han%'m that m%*ht
%n'ure $on*(term or*an%& &han*e. Th%' "a' the &on&ept of the Q'ur#%#a$ of the f%tte't,Q %n "h%&h tho'e %nd%#%dua$
or*an%'m' that are /e't adapted to the%r $o&a$ 'urround%n*' are 'een to ha#e a /etter &han&e of 'ur#%#%n*, and thu' of
d%fferent%a$$y pa''%n* a$on* the%r tra%t' to pro*eny. E.&%ted o#er h%' d%'&o#ery, 2a$$a&e penned an e''ay on the 'u/=e&t
a' 'oon a' he "a' "e$$ enou*h to do 'o, and 'ent %t off to Dar"%n. <e had /e*un a &orre'ponden&e "%th Dar"%n t"o
year' ear$%er and 9ne" that he "a' *enera$$y %ntere'ted %n Qthe 'pe&%e' Bue't%onQ- perhap' Dar"%n "ou$d /e 9%nd
enou*h to /r%n* the "or9, t%t$ed AOn the Tenden&y of Var%et%e' to Depart Indef%n%te$y from the Or%*%na$ Type,A
to the attent%on of Lye$$E Q
http:DD"""."9u.eduDU'm%th&hD%nde.1.htm
ARWIN5S EVOLUTINARY THEORY
Dar"%n0' e#o$ut%onary theory ha' t4o t&e$e$:
6, e$cent 4it& )o(i%ication: 3$$ $%#%n* and e.t%n&t 'pe&%e' ha#e de'&ended "%thout %nterrupt%on
from one or 'e#era$ or%*%na$ form' of $%fe. The'e ne" 'pe&%e' a&&umu$ated &han*e' o#er *reat
e.pan'e of t%me and no" $oo9 d%fferent from one another.
7, Natu#al Selection: U'efu$ #ar%at%on "%$$ a$$o" tho'e or*an%'m' that po''e'' them to 'ur#%#e
and reprodu&e, and pa'' the'e *ood #ar%at%on' to the%r off'pr%n*.
ARWIN8S THEORY OF NATURAL SELE+TION,
Fa&t O1 ( 2%thout &on'tra%nt', popu$at%on' "%$$ *ro" e.ponent%a$$y, produ&%n* an e#er more
rap%d$y *ro"%n* num/er of or*an%'m'.

Fa&t O2 ( In 'p%te of th%' pred%&t%on, the num/er' of %nd%#%dua$' %n a popu$at%on rema%n' near
eBu%$%/r%um, f$u&tuat%n* a/o#e and /e$o" 'ome mean #a$ue.
Fa&t O6 ( ?e'our&e' are $%m%ted.

+onclu$ion: From the'e three fa&t', Dar"%n &on&$uded that there was a struggle for e&istence.
Dar"%n &om/%ned th%' "%th t"o add%t%ona$ fa&t':

Fa&t O4 ( Ind%#%dua$' are un%Bue. There %' %nd%#%dua$ #ar%at%on. Th%' &ame from o/'er#%n* an%ma$
/reed%n*.

Fa&t O@ ( Cu&h, /ut not a$$, of the %nd%#%dua$ #ar%at%on %' her%ta/$e. Th%' o/'er#at%on a$'o &ame
from an%ma$ /reeder'. Some of the o/'er#ed #ar%at%on %' en#%ronmenta$, 'ome %' *enet%&.
+onclu$ion: The'e fa&t' $ed Dar"%n to the &on&$u'%on that 'ome %nd%#%dua$' are /etter eBu%pped
to 'ur#%#e and reprodu&e :Natura$ Se$e&t%on; %n the%r 'tru**$e for e.%'ten&e.
Throu*h many *enerat%on' of t%me, e#o$ut%on %' the re'u$t. Dar"%n u'ed 9(e$cent 4it&
)o(i%ication,9
Cayr :1!52; 'tated that Dar"%n0' Theory of E#o$ut%on %n&$uded f%#e theor%e':
1. Evolution a' 'u&h: $%nea*e' of or*an%'m' &han*e o#er t%me.
2. +o))on (e$cent: a$$ $%fe &an /e repre'ented a' a '%n*$e fam%$y tree "here there %'
&ommon an&e'tor to a$$ 'pe&%e'.
6. G#a(uali$): *radua$ &han*e' produ&ed ne" 'pe&%e'- th%' %' %n &ontra't "%th $altation,
the hypothe'%' that propo'e' that e#o$ut%on o&&ur' /y 'udden $eap'.
4. !o'ulation c&ange: e#o$ut%on o&&ur' due to the proport%on of %nd%#%dua$' %n the
popu$at%on that %nher%t a *%#en &hara&ter%'t%&.
@. Natu#al $election: 'ur#%#a$ and reprodu&t%on.
3 /%* *ap %n Dar"%n0' theory "a' the $a&9 of an e.p$anat%on a/out the or%*%n of the #ar%at%on'
found %n a popu$at%on: "here do the'e #ar%at%on' &ome fromE
Dar"%n0' theory "a' a )va#iational t&eo#y+ %n "h%&h popu$at%on' &han*e due to 'ur#%#a$
reprodu&t%on and 'e$e&t%on of &erta%n *ene'.
Lamar&90' theory "a' a )t#an$%o#)ational t&eo#y+ %n "h%&h the %nd%#%dua$ &han*e', %'
tran'formed, and then pa''e' tho'e tra%t' to the off'pr%n*.
Blen(ing In&e#itance.
Th%' "a' the pre#a%$%n* hypothe'%' a/out the %nher%tan&e of tra%t'.
Caterna$ and paterna$ &ontr%/ut%on' /$end %n the off'pr%n*.
3' a re'u$t, a ne" adaptat%on "%$$ /e d%$uted %n 'u&&e''%#e *enerat%on' and e#entua$$y
d%'appear.
Therefore, #ar%at%on 'hou$d de&rea'e and not %n&rea'e.
Dar"%n ne#er 9ne" that 8re*or Cende$ had 'o$#ed the pro/$em of %nher%tan&e %n a paper
pu/$%'hed %n 15>@.
EVOLUTIONARY THEORIES AFTER ARWIN
3fter Dar"%n0' theory "a' propo'ed %n the 15@' there "a' per%od of 'u/'tant%a$ &ontro#er'y and
'e#era$ theor%e' "ere propo'ed, "h%&h %n&$uded neo(Lamar&9%'m, ortho*enet%& and mutat%on%'t'
theor%e'.
3 *reat dea$ of %nformat%on "a' a&&umu$ated after the 15G due to the "or9 done %n
pa$eonto$o*y, &omparat%#e morpho$o*y and &omparat%#e em/ryo$o*y
Neo*La)a#c0i$): 'e#era$ theor%e' /a'ed on the %dea of the %nher%tan&e of a&Bu%red
&hara&ter%'t%&'
O#t&ogenetic t&eo#y: e#o$ut%on pro&eed' to"ard a predeterm%ned *oa$ and doe' not
reBu%re natura$ 'e$e&t%on. No me&han%'m "a' e#er propo'ed for th%' theory.
1utationi$t t&eo#y: *enet%&%'t' o/'er#ed that mutat%on' ar%'e and produ&e ne" #ar%at%on'
%n the popu$at%on, therefore, natura$ 'e$e&t%on "a' not ne&e''ary for e#o$ut%on to o&&ur-
the mutant' repre'ent ne" 'pe&%e'.
THE EVOLUTIONARY SYNTHESIS
Th%' 'ynthe'%' %n&orporate' the %nformat%on o/ta%ned /y *enet%&%'t', 'y'temat%'t' and
pa$eonto$o*%'t' %nto the theory of e#o$ut%on propo'ed /y Dar"%n.
Cutat%on' and natura$ 'e$e&t%on together &au'e adapt%#e e#o$ut%on.
Cutat%on' are the ra" mater%a$ of natura$ 'e$e&t%on.
Cutat%on', re&om/%nat%on, natura$ 'e$e&t%on and other pro&e''e' operat%n* "%th%n 'pe&%e' a&&ount
for the or%*%n of ne" 'pe&%e' and for the ma=or, $on*(term feature' of e#o$ut%on :ma&roe#o$ut%on;.
Fun(a)ental !#inci'le$ o% Evolution
1. The phenotype %' d%fferent form the *enotype- phenotyp%& d%fferen&e' may /e due to *enet%&'
or en#%ronmenta$ fa&tor'.
2. En#%ronmenta$ effe&t' do not affe&t *ene' pa''ed on to the ne.t *enerat%on- a&Bu%red
&hara&ter%'t%&' are not %nher%ted.
6. 8ene' reta%n the%r %dent%ty and do not /$end.
4. 8ene' mutate u'ua$$y at a '$o" rate.
@. E#o$ut%onary &han*e o&&ur' %n the popu$at%on.
>. The rate of mutat%on %' too '$o" for mutat%on a$one to &au'e e#o$ut%onary &han*e, the 'h%ft
from one *enotype to another. Chan*e %n *enotype proport%on' %n a popu$at%on &an /e &au'ed
/y *enet%& dr%ft :random f$u&tuat%on'; and /y 'ur#%#a$ of the f%tte't :nonrandom- natura$
'e$e&t%on;.
G. Natura$ 'e$e&t%on &an, under &erta%n &%r&um'tan&e', /r%n* a/out 'u/'tant%a$ e#o$ut%onary
&han*e %n a rea$%'t%& amount of t%me.
5. Natura$ 'e$e&t%on &an &han*e the freBuen&y of a$$e$e' that may re&om/%ne "%th other *ene'
that affe&t the 'ame tra%t, and *%#e r%'e to ne" phenotype'.
!. Natura$ popu$at%on' are #ar%a/$e and &an &han*e rap%d$y "hen en#%ronmenta$ &ond%t%on'
&han*e.
1. ,opu$at%on' %n d%fferent *eo*raph%& re*%on' d%ffer %n &hara&ter%'t%&' that ha#e a *enet%& /a'%'.
11. The num/er of d%fferent *ene' form' the /a'%' for d%fferen&e' /et"een popu$at%on' of the
'ame 'pe&%e' and /et"een 'pe&%e'. Th%' 'upport' the hypothe'%' that 'pe&%e' e#o$#ed %n a
'er%e' of 'ma$$ 'tep'.
12. D%fferen&e' /et"een *eo*raph%& popu$at%on' of a 'pe&%e' are often adapt%#e and are the
&on'eBuen&e of natura$ 'e$e&t%on.
16. ,henotype a$one doe' not def%ne a 'pe&%e'- there are d%fferent phenotype' "%th%n an
%nter/reed%n* popu$at%on. D%fferent 'pe&%e' repre'ent d%fferent *ene poo$' and do not
e.&han*e *ene' "%th other 'pe&%e'.
14. S'eciation %' the or%*%n of t"o or more 'pe&%e' from a &ommon '%n*$e an&e'tor. Spe&%at%on
o&&ur' /y the *enet%& d%fferent%at%on of a *eo*raph%&a$$y 'e*re*ated popu$at%on.
1@. There are many *radat%on' of phenotyp%& d%fferen&e' %n $%#%n* or*an%'m' a''%*ned to 'pe&%e'
*rouped under the 'ame *enu', *enera to fam%$%e', et&. <%*her ta.a ar%'e /y the 'eBuent%a$
a&&umu$at%on of 'ma$$ d%fferen&e' rather /y the 'udden appearan&e of a ne" )type+.
1>. The fo''%$ re&ord ha' many *ap' amon* Bu%te d%fferent 9%nd' of or*an%'m', /ut %t a$'o
%n&$ude' e.amp$e' of *radat%on from an&e'tra$ form' to ne"er one.
EVOLUTIONARY BIOLOGY SIN+E THE SYNTHESIS
S%n&e the 1!@' ad#an&e' %n *enet%&' and mo$e&u$ar /%o$o*y ha#e re#o$ut%on%1ed the 'tudy of
e#o$ut%on.
In the '%.t%e', e#o$ut%onary theory ha' e.panded %nto e&o$o*y, an%ma$ /eha#%or and reprodu&t%#e
/%o$o*y.
1olecula# evolution: ana$y'%' of the pro&e''e' and h%'tory of &han*e %n *ene'.
Neut#al Hi$to#y o% 1olecula# Evolution: a hypothe'%' that ho$d' that the e#o$ut%on of
DN3 'eBuen&e' o&&ur' /y *enet%& dr%ft rather than /y natura$ 'e$e&t%on.
Evolutiona#y evelo')ental Biology: 'tud%e' ho" de#e$opmenta$ pro&e''e' /oth e#o$#ed and
&on'tra%nt' e#o$ut%on: Evo*evo for 'hort.
Tr%e' to under'tand and determ%ne the an&e'tra$ re$at%on'h%p' of or*an%'m' and the
pro&e''e' %n#o$#ed %n de#e$opment.
Evolutiona#y Geno)ic$: dea$' "%th the #ar%at%on and e#o$ut%on of mu$t%p$e *ene' and ent%re
*enome'.
!HILOSO!HI+AL I1!LI+ATIONS
The theory of e#o$ut%on &han*ed the e''ent%a$%'m of ,$ato and 3r%'tot$e and p$a&ed #ar%at%on %n %t'
p$a&e.
E#ery &hara&ter%'t%& of a 'pe&%e' &an #ary and rad%&a$$y 'o %f *%#en enou*h t%me.
The theory of e#o$ut%on &han*ed the 'tat%& &on&ept of the "or$d for one of &han*e.
Chan*e and not 'ta'%' %' the natura$ order.
?andom purpo'e$e'' #ar%at%on a&ted on /y /$%nd, purpo'e$e'' natura$ 'e$e&t%on pro#%ded a
re#o$ut%onary ne" 9%nd of an'"er to a$mo't a$$ Bue't%on' that /e*%n "%th )2hyE+.
The 'tru&ture' of or*an%'m' ha#e a fun&t%on, not a purpo'e.
ETHI+S" RELIGION AN EVOLUTION
Creat%on%'t mo#ement: $%tera$ read%n* of the &reat%on 'tor%e' found %n the 7%/$e.
The 'uppo'%t%on that natura$ %' *ood %' &a$$ed /y ph%$o'opher' the Natura$%'t%& Fa$$a&y.
There are no eth%&a$ or mora$ pr%n&%p$e' %n '&%en&e.
S&%en&e doe' not deny or aff%rm the e.%'ten&e of a Creator.
EVOLUTION
3 &y'ot&e$i$ %' an edu&ated *ue''- an %nformed &on=e&ture of "hat may /e true.
3 $cienti%ic t&eo#y %' a mature, &oherent /ody of %nter&onne&ted 'tatement', /a'ed on rea'on%n*
and e#%den&e that e.p$a%n a #ar%ety of o/'er#at%on'.
E#o$ut%on %' a fa&t that %' e.p$a%ned /y the e#o$ut%onary theory.
De'&ent "%th mod%f%&at%on' %' 'upported /y e#%den&e pro#%ded /y pa$eonto$o*y, *eo*raph%&
d%'tr%/ut%on' of 'pe&%e', &omparat%#e anatomy, em/ryo$o*y, *enet%&', /%o&hem%'try and mo$e&u$ar
/%o$o*y.
<o" mod%f%&at%on' o&&ur and ho" an&e'tor' *%#e r%'e to d%#er'e de'&endant' &on't%tute' the
theory of e#o$ut%on.
3 /ody of %dea' a/out the &au'e' of e#o$ut%on, %n&$ud%n* mutat%on, re&om/%nat%on, *ene f$o",
%'o$at%on, random *enet%& dr%ft, the many form' of natura$ 'e$e&t%on, and other fa&tor', &on't%tute
our &urrent theory of e#o$ut%on.
L%9e a$$ theor%e' %n '&%en&e, %t %' %n&omp$ete.