R I N C I PL ES O F BR EED I N G

A T R E A T I SE O N T H R EM M A T O L O G Y
OR

T HE P R I N C I PLES AND PR A CT I C ES I N V O LV ED I N T HE

E C O N O M I C I M R O V EM EN T O F D O M EST I
P
G AT ED A N I M A LS A N D P LA N T S

E . D AV EN POR T, M A G R . .
,
LL D . .

P no r ns so u op T n nnu u A rO L oc v
'
m T H! U m v n ns r r v or [n arro w
D EA N or m : CO L a n or Ac nic
-
ru n n
-
D i a na o u o r r u n A G R ICU L TU R A L E x n nm m ‘
r S T A T ION

WIT H A PPE N D I X
BY

H L R I E I Z PH D
' ‘

. . .
.
,

ASS l S l ANT
‘ '
Pno n sso a o r MA Tu u ‘
A rlcs in run U m v n n s r rv op I L L m o rs

G INN C O M PA N Y
B OS TON N E W Y OR K C HI CA G O L O N D ON
EN T B R B D AT ST A T I ON E R S
'
HA L L

Co m m en t , 1 90 7

BY E U G E N E D A V E N POR T
A LL R IG H TS R ES E R V E D

fl tb cn c u n D r e“
G I NN CO M P AN
Y PR O
PR I E T O R S B OS TO N U S A
. . .
 P R E FA C E

T wo c la s ses o f people hav e been in m ind in the prepa rat io n o f

th is t ext v iz t he s t u d ent o f agr ic u lt u re in the c ollege and experi
,
.

m ent s t a t io n and t he pra ct ical br eed er u po n t he farm . Bo th n eed

t o kn o w a ll that ev o lu t io n ha s t o t each o f m eth od s that m ay b e
em plo y ed in s t ill f u rth er adapt ing t o o u r n eed s s u ch an im a ls and

plant s a s ha v e been d om es t ica t ed becau se o f th eir va lu able natu ral

qua lit ies .

Th e g en era l p u rpo s e has been fir s t o f all t o d efin e t he pr o blem s

in v o lv ed in a n im a l and plant im pr o v em ent ; to free t he s u bj ect
fro m t h e prej ud ic e and t rad it io n that hav e alway s befogged it ;
t o br in g t o t he s t u dy what ev er fa ct s a re f u lly kn o wn t o b io lo gica l
s c ie nc e ; t o r ec o gn ize and d efin e s o m ewhat c lear ly t he pr esent

lim itat io n s o f kn owledge and t o ind icat e as well as m ay b e t he

,

d irect io n s fr om wh ich f u rth er and m u ch n eed ed light is m os t

-

lik el y t o c o m e Las t o f all an d m o re th a n a ll it ha s b een t he

.
,

pu rpo se to enc o u rage and if po ss ib le ind u c e m o re exact m eth od s

, ,

o f s t u dy and o f p ract ic e than ha v e h ith ert o ch a ract er ized th is

b ra nch o f agr ic u lt u ra l sc ienc e .

I t is yet too ea r ly to pr epare an id ea l treat ise u po n th is m o s t

intricat e s u bj ect and n o o n e is m o re c o n s c io u s than t he a u th o r
,

o f t h e m any d efic ienc ies and s h o rtc om in g s o f th is a tt em pt . S ome

effo rt h o wev er is s u r ely n eed ed at th is t im e to c lear t he at m o s
, ,

p h e r e t o,g i v e t h e s t u d ent o f agr ic u lt u r e at leas t a ra t io na l po int

of V iew an d t o bring h im int o c om rad es h ip with th o s e who a re

,

ea rn es t ly s t u dy ing b io lo g ica l pr o b lem s a n d thr o u gh wh o s e effo rt s

th es e v exed q u est ion s are s u re soo n er or lat er to find a solu t ion .

Th is t o geth er with t he pr es s ing n eed o f a t ext in his o wn c las s

,

roo m is t he au th o r s o n ly warra nt for t he pres ent v olu m e

,
’
.

N o n ew th eo r ies o f ev o lu t io n ar e pro po s ed . T he ch ief o bj ect

has been to d is t ingu is h what is k n own fro m what is m erely t rad i
tio nal ; to giv e a s m u ch a s pos sible with in t he lim it s o f a v a ilable
,

spac e o f t he b es t es tab lis h ed fact s b ea r ing u po n th is su bj ect

, to
v i R
P EFA C E

call att ent io n to approv ed m ethod s o f s t u dy and to ind ica t e lin es ,

o f res earch m o s t lik ely to f u rn is h va lu ab le info rm at io n in t he n o t

d istant f u t u re .

I t is n ec essary t o introd u c e a co n sid erab le am o u nt o f m ath e

ma t ic al work in t he lat er chapt er s N o exc u se is o ffered fo r th is
.

introd u ct io n and it is earn es t ly d es ired that the read er g iv e

,

s pec ia l att ent io n t o th is po rt io n o f t he t ext wh eth er ea sy o r d iffi

,

c ult of followi ng beca u se it is by th is road t hat many n ew pr inc i

,

ples will arriv e an d that m any of o u r f u t u re Operat io n s m u s t b e

o rd er ed ; fo r n o th ing is c lea rer than that til e s ucc essf u l b reed er

qf t/ze f u tu r e will be a b ookkeeper a nd a s ta ti stic ia n . F or t he

co n v en ienc e of t he n on m ath em at ical read er gen eral fo rm u lae are
-

placed in foo tn ot es and some of t he m o re ab stract m att er is

,

placed in the fo r m o f an append ix fo r t he ben efit o f t he m o re

m ath em at ica lly inc lin ed .

T he wr it er has tau ght th is s u bj ect fo r fift een year s and is fu lly

aware of the pedagogic d iffi c u lt ies in v olv ed as well as of the
lim itat io n s of kn o wled ge He has tried m any d ifferent o u t lin es
.

and m any d ifferent m ethod s of presentat io n and has ch osen the ,

o n e h ere em p loy ed becau se in experienc e it seem s the m os t

fav o rable for the presentat io n o f the s u bj ect m att er in v o lv ed and -

at t he sa m e t im e for pu tt ing the st u d ent in a frame of m ind

fav orable for the u nd ert ak ing o f ec on om ic breed ing o pera t io n s
and fo r t he rec ept io n of n ew tru th s as th ey s hall b e d isc ov er ed .

V ariat io n rath er than h ered ity has b een ch o sen for t he in it ial
and lea d ing th ou ght becau se bett er calc ulat ed as exper ience has ,

s h o wn t o a fford a fa v or ab le o u t lo o k and t o d ev elo p s u ch c o n

,

c ept io n s of ev olu t io n as are m o s t u s ef u l lat er o n .

T he ev olu t io n is t who m ight chanc e to s can th es e pages wo u ld

b e s tru ck by the ab s enc e o f so m e of t he card inal feat u res o f
ev olu t io n as he wo u ld a ls o n o t e t he exc eed ing pr o m in enc e giv en
,

t o c ertain o th er q u es t io n s of s eem ing m in o r im po rtanc e Herein .

exis t s the d ifferenc e bet ween thr emm at o logy and ev olu t io n and ,

th is v ery matt er has giv en the au th o r m ore d iffic u lty th an all

o th er s v iz t o r ear ra nge v alu es an d to d et erm in e pr o per r elat io n s
,
.

o f o ld q u es t io n s in a n ew fi eld .

W e m u s t d isc u s s the ca u ses of v ariat io n ev en th ou gh we are

told by the best s t u d ent s t hat su ch att em pt s are prem at u re A .
 PR E FA C E v fi

m in o r m a t t e r in ev olu t ion c u rio u s rath er t han o th erwise it is a

, ,

v i tal o n e i n thr emm at o logy and we m us t d is c u s s t he s u bj ec t t he

,

b es t we a r e ab le if o n ly t o lea rn how litt le we rea lly kn ow abou t

,

it an d t o p o in t att ent io n in the r ight d irect io n .

N o a t t em pt has been mad e t o inc lu d e ex ha u s t iv e referenc es .

On t he o t h er han d th ey are c o n fi n ed for t he mo s t part to a few

,

s t a ndard b o o k s m o f acc e s an d t o s av e t im e t he ref erenc es

y s ,

are m o s t l y t o d efi ni t e pa ges A gen eral and m ore ext end ed lis t

.

follo ws t h e su m m ary of n ear ly ev ery chapt er enabling the s t u d ent ,

to p u r s u e that part ic u lar s u bj ect f u rth er if d es ired b u t th ere is

n o att em pt at a c o m plet e b ib lio graphy I t was h o ped that if t he
.

lis t o f r e ferenc es c o u ld b e k ept sm all the s t ud ent an d the br eed er

wo u ld b e t he m o re lik ely t o prov id e th em selv es with s tandard
li t er at u r e bea ring on t he s u bj ect I hav e m ad e t he frees t u se o f
.

s tan d ar d a u th o r s gi v ing f u ll cr ed it in all ca ses g en er ally in the

, ,

fo rm o f reference to t ext an d page Th is c ou rs e ha s been d ictat ed

.

by th e d es ire to f u rn is h the s t u d ent with reliable fact s rath er

than a ser ies o f acad em ic d is c u s sio n s u po n d is pu t ed su bj ect s .

I d esire t o ackno wledge t he v ery great serv ic es o f D r R iet z .

,

t o wh om I a m ind ebt ed for m u ch as s is tanc e in t he m o re s tat is t ica l

po rt io n s and for the preparat io n of the append ix es pec ially
,

d irec t ed to the m ath emat ical s t ud ent n o t a s a t ext b u t as an ,

in t r od u ct io n t o fu rth er s t u dy in th is s pec ial pha se o f s c ienc e .

I am also ind ebt ed t o m any o f m y c o lab orers in t he U n iv er

s it o f I llin ois an d elsewh er e as well a s to n u m er ou s breed er s in
y ,

t h is and o ther s tat es who by th eir as s is tanc e hav e co ntr ib u t ed

,

m u ch t o any s u cc ess wh ich th is v o lu m e may m eet .

I ts po s sible m erit s th erefo re I m us t s hare with o th ers ; it s

, ,

d efec t s and s h o rtcom in gs are my own .

E D AV E N PO R T .

U I
N V ER S YIT or I LL I N OI S
U BA N A
R
 CON TEN TS

I N T R O D U C T I ON
PART I V A R I A TI O N
C H A PT E R

I . V A I A TI N
R O IN GE N ALER

I . V a ria t io n U n iv ers a l am o ng Li v in g B ein gs

II . V a ria b i itl y th e Bas is fo r I m p ro v em en t
III . N a t u re of V a ria b i it l y
IV . Mea n in g o f t he T e rm C h a rac t er

V . D o m in a n t an d La t en t C h a rac t ers

VI . T h e U n it of V a ria b i itl y
VI I . D is tin c t io n s as t o Ki d n s of V a ria t io n s
R eferenc es

II . M O R P HO LO G I C A L V A I ATI
R ON

R eferen c es

III . S U B ST A N TI V E V A R I ATI O N
M ER I S I IC ’ ‘
V A I A TI N
R O

1 . S y mm t ye r

II . Meris t ic V a ria t io n In L in ea r S eries

III . Meris t ic V a ria t io n an d Bi l a t era l S y mm t y e r

IV . y
S mm et r y in V a ria b l e Pa rt s
V
Meris t ic V a ri a t io n in R a d ia S eries
. l
V I I mpo rta n c e o f M eris tic V a ria t io n
.

R eferen c es

V . FU NC TI O N A L V A I ATI N
R O

I . V aria t io n in t he D egree O f A c t iv it o f N o rm a F u n c t io n s b e y l
d d l
tween D ifferen t I n iv i u a s o f th e S a m e S p ec ies
II . V a ria t io n in t he D egree o f A c t iv i t Of N o rm a Fu n c t io n s wit iny l h
t he S a m e I n di id
v u al

I II . Mo d ific a t io n of N o rm a l F u n c t io n s b y E x t er n a l or Ot h er I n flu
en c es

IV . N o rm a l F u n c t io n s ex erc is e d un d er l
A b n o rm a C o n itio n s d
R eferen c es

VI . M u T AT I O N s
I . D is t in c tio n b etween M u ta tio n an d d
O r in a r yV a ria t io n

II . Examp l es o f M u ta t io n
X CON T E NTS

C H A PT E R PA G E

III . E xp erimen ts of D e V ries

IV . A m eric a n E xperien c es
V . E c o n o m ic S ign ific a n c e of M u ta t io n s
VI . B io l o g ic a l S ign ific a n c e o f M u t a t io n s

R eferen c es

PAR T —
I I C A U S ES OF V A R I A T I ON
I NT RO DU C T O N I
T H E M EC HA N I M S OF L M N T A N D D I FF E R E N TI ATI N
D EV E OP E O

I . Pro to p las m th e Phy i l B i f Lif

s ca as s O e

II . T he C e ll th e U n it of S tr u c t u re
I II . Mec h a n is m of C ell D iv is io n ( M i t o s is
)
IV . C ell D iv is io n wit h an d wit h o ut D ifferen t ia tio n
V . P hy s io lo gi c a l U n its
R eferen c es

VI II IN T N A L C A U
. ER S ES OF V A I A TI N
R O

I . I N TE R N A L I N F L U E N CE S A F F E C TI N G PR I JI A R I L Y TH E I N
D I VI D U A L
I . C ell D i v is io n
II . d
B is ex u al R ep ro u c t io n a F u n a m en ta C a u s e o f V a ria t io n d l
III . d
Ma t u ra t io n a n d the R e u c tio n o f t he C ro m o s o m es a C au s e h
Of V a ria t io n
IV . B u d V aria t io n
V . In fl u en c e o f t he C o n it io nd of t he G erm u po n D l
ev e o p m en t

VI . X en ia, or l
Fert i iza t io n of t he E n d os
p erm
VI I . l
T e ego n y
VII I . I n t ra U t e rin e I n
-
fl u en c es

IX . R ev ers io n an d A ta v is m
X . d d l Ch
I n iv i ua a ra c t ers d ep en d en t u p o n S ex
II . I N TE R N A L I N F L U E N CE S A F F E C TI N G TH E RA CE AS A

XI . R ela t iv e Fert ilit y , or G en et ic S e l ec t io n

XI I . Phy i l s o o gic a lS l e ec t io n

X III . S l ti
e ec ve D ea th R a te ; L o n gev i t y
X IV . B th m i
a c In fl u en c es

XV . Phy i l s o o gic a l U n it s
X VI . G erm in al S elec tio n
R eferen c es

E xT E R N A L I N LU N
F E C ES A s A
C U S ES OF V A ATI O N RI

I . G en era l Eff ec t of Lo c a it l y u po n Pla n t an d A n im a l D ev elo p

men t
II . In fl u enc e o f Fo o d u
po n V a ri ab ilit y
III . E ffec t Of Mo is t u re u p o n D ev e l pmo en t

IV . Effec t Of C o n tac t u po n Pro to p l mi

as c A c tiv it y
V . Eff ec t of G ra v it y u po n Mat ter
L iv in g ; G eo t ro p is m
VI . Effec t of Lig t h u po n Li v in g Ma tt er
 C O NTENTS x i

Ha m : PA C E

VII . I n flu en c e of Tempera t u re u po n L iv in g M a t ter

V I II . Efi ec t of C h em ic a l Ag en ts u po n Pro to p l as m ic A c t iv it y
IX . Effec t of S a lin e S l o u t io n u
po n D ev elo p m en t in A q u a t ic
A n ima l s

X . In fl u en c e Of U se an d D is u s e u po n D ev e l o p m en t

XI . E xt ern a l I n flu en c es as C a us es of V a ria t io n in T pe y
R eferen c es

X . RE L TI
A VE ST A B I L IT Y AN D IN TA I L T
S B I Y OR LIV I N G M A TT E R
I . Ev i d en c e fro m S ta b il it y f T y p
o e

II . Ev id en c e fro m M u t b il it y
a f Sp i O ec es

I II . Ev id en c e fro m R ev ers io n and A ta v is m

IV . Ev id en c e f ro m D is a p p ea ra n c e of Pa rt s
V . Ev id en c e fro m t he D i rec t A c t io n of t h e E n v iro n m en t
VI . Ev id en c e l
f ro m A c c im a t iza tio n
VII . Ev id fro m R eg en era tio n
en c e

VIII . I n tern al Fac to rs in R egen era t io n

IX . Ev i d en c e f ro m G ra ft in g
X . Ev id en c e fro m th e O rigin o f N ew C e s a n d T iss u es ll
XI . Ev id en c e l
f ro m D ev e o pm en t an d D ifferen t ia tio n
R eferen c es

PART III T R A N S M S S ON I I
TR A N S M I SS I O N OF M O D I F I C ATI O N S D U E TO EXTE R N A L IN LU N F E C ES

I . I n t ro d u c to r y
II . Ev i d
en c e f ro m t he N a t u re of V a ria t io n
II I . Ev id
en c e fro m Mu t ila tio n s
IV . Ev id en c e f ro m Fo o d S pp l y u

V . Ev id
en c e f ro m A c c l im ti t i
a za on

VI . Ev id
en c e fro m Hab it an d I n s t in c t
VI I . Ev id en c e fro m U s e an d D is u s e
V II I . Ev id fro m D is a ppea rin g O rga n s
en c e

IX . V a ria t io n s d u e t o C a u s es n o t affec tin g t he G erm a re no t

T rans m it te d
R eferen c es

AND V A I A I L IT
R B Y

T pe y
V aria b ilit y , D ev ia tio n f ro m T p e
or y
Prac t ic a l Hin ts o n the T a in g an d G ro u pI n g k o fM eas u rem en ts

P ro b a b l e Erro r
C o m pa ra tiv e T yp e a n d V a ria b i it l y fOD iff r eren t C h a ra c t ers in
the S am e Po p u a t io n l
Effec t of Se l ec t io n u p o n y
T p e a n d V a ria b i it l y
d
I n irec t E ffec ts Of Se l y
ec tio n u p o n T p e an d V a riab i it l y
d
S t u ies in T p e y
O f t h e S a m e V a rie
a n d V a ria b i it l y ty fo Com
ra is e d d
u n er D ifferen t C o n it io n s a s t o F ert i it d l y
R ef eren c es
x ii C O N TE NTS

C H A RT E R PA G E

XI I I . COR R E L ATI N O

I . M ea n in g Of C o r re l a t io n

II . Ca l l
c u a t io n o f C o effi c ien ts of C o rre l a t io n

III . The C o rre l a t io n Ta b l e

IV . The C o rrela t io n C o e fli c ien t

V . T h e R egres s io n C o efli c ien t
VI . d d
S t u ies in S p ee R ec o r s o f T ro t ters d
R eferenc es

X IV . H E R E D IT Y
IHo w C h a ra c te rs b eh a v e in T ra n s m is s io n
.

I I S ta tis t ic a l M et h o ds O f S t u dy o f Hered ity

.

I II T he R egres s io n Ta b e l
l k Off p i
.

IV . k
Li e Pa ren ts b eget U n i e s r ng a n d, c o n v ers e ly , Lik e

Offs p rin g may b e b ego t te n b y U n i e Pa ren ts lk

V . R egres s io n I n gen era l, t h e O ff s p rin g is M o re M e io c re
. d
h h
t a n t he Pa ren t s ; t a t is , W a t ev er t h e I a ren t age t he
’
h ,

O ffs p rin g ex h ib it s a S t ro n g T en de y t nc o r eg res s t o wa r d th e

Mea n O f t he R a c e
V I T he Mea s u re O f Hered it y
.

V I I The M ea n o f t he O ffs p rin g

. no t n ec es sa ril y the Same as t he

Mean O f t he Pa ren tage

VI I I . E x t rem es of a R ac e r ela t iv e ly L es s Pod r u c t iv e t h an th e
Mea n s
IX . Pro g res s io n . Pa ren ts in gen era l p ro d uce a F e w I n iv i d d l ua s

M o re h
E x t rem e t a n t he R a c e
X . T h e E xc ep tio n a l I n iv i d d l i ua a r s es eit h er fro m M e io c rit d y
or f ro m t h e E xc e
p t i l P t ona a ren

XI . F ra t ern a l V a r i b il it y — Off p i g
a , s r n of S a me Pa ren t s no t

I d en t ic a l

X II . C h a rac ters t en d to co mb in e in D efin ite M a t h em a t Ic a l

Pro po rt io n s
X I II . Men d e l ’
s La w O f H b r i y d s

X IV . The La w of A n c es t ra l H ere it d y
XV . L im it to t he Re d u c t io n O f V a r ia b i it l y
X VI . Po wer Of S elec t io n to p erm a n en tly m d ify T y p
o es by t he

E s tab is l h m en t o f B ree s d
X VI I . d
B ree in g T ru e , or S ta b i l it y of a Ch a ra c t er es ta b lis h d
e by
S elec t io n
X VIII . D u ra t io n of V a riet ies , B ree d s , a nd Fa m il y St ra in s

R eferen c es

XV . P R EPOT E N CY

I . D a ta fro m th e T ro t t in g R ec o r d s illu s tra t in g Prep o t en c y

II . Prepo ten c yi n S ex
III . I n flu en c e of A ge on Prep o ten c y
IV . In fl u en c e o f C o n s t it u t io n a V igo r l u
p on Prep o t en c y
R eferenc es
 CO NT ENT S x iii

PA R T —
IV P R A C T I C A L P R O B L E MS
CH A P T E R PA G E

XV I . SE L ECT I N O

I . I d ea ls S e ec t io n
In l
II . l
His to ric a K n o w e ge o f t he B ree N ec es s a r ld d y
III . G en era l Pri n c ip es in v o v e I n S e ec t io nl l d l
IV . R a tio n al S elec tio n

XV I I . S Y ST E MS OP B R E ED I N G

I . P u rpo s es in B ree in g d
II G ra in g d .

Hy b ri dizin g
.

I II . C ross in g or

IV . Li n e B ree in g d
V . I n b ree in g d
VI . B reedin g f ro m t he Bes t
R eferen c es

XV I I I . T II E D ET E R I I I N A T I O N or S ex
I . T h eo ries

II . In fl u en c e o f N u t ri t io n
II I . In fl u en c e o f l
Fert i iza t io n
IV . S ex in Mamm a ls
V . T he A c c ess o r C y h ro m o s o me an d D et erm m a t I o n
R eferen c es

XI X . PLA NT B R E ED I N G

I . A d v a n ta
ges a nd Lim it a t io n s
II . So i l an d l
C u t u re C o n it io n s d
I II . S y s t ems o f P l a n tin g

R eferen c es

A N I MA L B R EED I N G

I . A d v a n ta ges a n d D is a d v a n ta
ges

II . Fewer C h a rac t ers fo r S e l ec t io n

II I . Fas io n h
IV . S h o w R in g C o n s eq u en c es
V . Tes t in g of S ires an d D ams
VI . Wea t h erin g a Perio d Of D ep ress io n an d d
Pres e r v in g t he Her
VII . R ec o r d s .

VIII . D is po sa l of S u rp l us F em a l es

IX . A Ma r k et fo r S ires
X . C o mm u n it B ree in g y d
XI . T he Y o u n g B ree er d
R eferen c es

XXI . D EV E LO P M E N T
 T HE PR I N C I PL ES O F B R EE D I N G

T H R EM M A T O L O G Y

I N T R O D U CT I ON

T he m ai n t it le Of t he pres ent v olu m e was ch osen becau se self

explan at o ry th o u gh it les s acc u rat ely expres ses t h e s c o pe Of t he
,

s u bj ect than d oes t he s u b t it le wh ich is o n ly beg inn ing t o c o m e

’

-
,

int o g en eral u se .

T hr em m at o logy from t he Greek t/zrem ma a th ing bred is a

, , ,

t erm pro po sed by R ay L anca s t er to c o v er t he pr inc iples and

1

pract ic es c o nc ern ed in t he im prov ement o f d o mes t icat ed an imals

and plant s .

T h e t erm is broad er than t he princ ip les Of breed ing beca u se

it i nc lu d es d ev elo pm ent a s well as reprod u ct io n I t is d is t inct .

fro m ev olu t io n in gen eral wh ich att em pt s to explai n the princ i

,

ples and fo rces co nn ect ed with t he o r igin and d ev elo pm ent o f all
fo rm s of life b u t with ou t t he slight es t referenc e t o ec o n o mic
c o n s id erat io n s I n ev olu t io n a pro to zo On is as im po rt a nt as a
.

pg i ,
a hyd ra O f as m u ch S ign ificanc e as a h o r s e and t h e m o s t
,

pes t ifero u s weed as m u ch an Obj ect o f int eres t as eith er c o rn

o r wh ea t .

T hrem m at olo gy lim it s it s elf t o th o s e S pec ies and v ar iet ies wh o se

n a t u ra l q u a lit ies m ad e th em u s efu l t o m a n in t h e b eg inn ing and ,

it as k s an d s eek s t o an s wer th is o n e q u es t io n Ho w can th ey b e ,

m ad e s t ill m o re u sefu l and b ett er adapt ed t o t he p u rp o ses Of an

a d v anc ing c iv ilizat io n ? I n th is s t u dy fo r m s Of life that a r e Of n o
e c o n o m ic v alu e a r e Of n o s pec ia l c o nc ern exc ept a s th eir c o n s id er

a t io n ma thr w l ight p n d m t icat d f o rm s T h i latt er i

y o u o o e s e s s
.

O ft en t he ca se fo r it is d o u bt les s tr u e that t h e s am e pr inc ip les

,

a pply t o all s pec ies ec o n o m ic o r o th er wis e b eca u s e n o n e wer e

, ,

1
S E y l p di B i t
ee nc i
c o e i t h d it i
a r XX IV 8 4
an n c a, n n e on , , I.
2 I NT R O D U CT I O N

s pec ially creat ed fo r ma n any m o re than for any o th er an im al .

I t is a gen era l b io lo g ic a l tr u th th a t ev eryth ing liv es u nt o it s elf ,

— n o t wh ere it ch o s es b u t wh ere it can ; n o t u po n what it likes

bes t b u t u po n what it c a n get .

I t may s eem t o t he s t u d ent th a t an u nd u e am o u nt o f att ent io n

is giv en to v ar ia t io n and that a d is pr o p o rt io nat e a m o u nt Of s pac e is
d ev ot ed to that s u bj ect I n that ev ent I h av e to say th a t v ar iat io n
.

is n o t t he ant ith es is o f h ered ity b u t rath er it s c o n s tant an d in sep

arable att endant and t ha t t he fact s Of v ar ia t io n c o n st it u t e t he
,

bes t po rt ion Of that s t ock Of info r m at io n with wh ich t he s t u d ent

m u s t b ec o m e po ss es sed b efo re h e is r eady to s t u dy t he pr inc ip les
in v olv ed in t ho s e gen era liza t io n s u po n wh ich pract ica l o perat io n s
m ay b e s afely ba sed .

On e is pa inf u lly aware t oo Of t he n ec es s ity o f ranging far and

, ,

wid e fo r fact s and t h e s t ud ent c a nn o t fa il t o feel Oftt im es that

,

t he s u bj ect m att er in hand is far r em o v ed fro m a gr ic u lt u r e Wh en

-
.

th is is t he ca s e it is beca u se we are forced to tak e wh at is av ail

a b le and m ak e t he m o s t Of it U nfo rt u nat ely t he wo rk er s in
.

s tr ict ly agr ic u lt u ra l fi eld s ar e all t oo few and t he r eliab le data

d eplo rab ly m eager th o u gh so m e o r iginal an d I tr u s t v alu ab le

,

m att er h as been r ec en t ly add ed to o u r s t ock o f kn o wledge .

Wh ile the sa me princ iples d o u bt les s apply in threm m at ology

as in ev o lu t io n y
,
e t im p o rtant d is t inct io n s a re t o b e O b ser v ed .

Fir s t th er e is ev ery r ea so n t o S I IppOs e that ev en f u nda m enta l

,

la ws app ly t o d ifferent s p ec ies in d iff erent way s Fo r ex a m ple

.
,

I nd ian c o rn seem s pa rt ic u lar ly s en s it iv e t o c lo s e br eed ing wh erea s ,

wh ea t is alm o s t exc lu s iv ely in br ed a n d ha s b een S O inbred fo r

u nkn o wn gen era t io n s A ga in t he c irc u m s t a nc es Of t he ca s e
.
,

Oft en intr o du c e int o t he pro b lem c erta in ec o n o m ic c o n s id erat io n s

n o t r es t ing u po n g en eral ev o lu t io n Fo r exa m ple m an cann o t

.
,

affo rd t he co u nt les s ages a n d u nt old gen erat io n s wh ich are

“

acc o rd ed n a t u r e fo r acc o m plis h ing res u lt s I n pract ical breed ing .

Operat io n s s u b s tant ia l r es u lt s m u s t fo llo w at o nc e and exh ib it a

h igh d egree Of s u cc es s wit hin the period o f a lifet im e o r th ey will

b e d is card ed a s v alu eles s .

E xper ienc e s h o ws tha t t he p u r po s es s tandard s and m eth od s , ,

Of a s u cc es s f u l breed er are s eld o m hand ed d o wn fr o m on e m a n

to an o th er ev en to h is o wn so n
,
E v en if that c o u ld b e d o n e it
.
,
 I NT R O D UCT I O N
3

wo u l d c o n s t it u t e no exc ept ion to the ru le tha t a man m u s t real

ize t h e fru it o f his o wn la b o rs and in his o wn gen erat io n for ,

br eed ing is a b u s in ess and mu s t b e m ad e to pay The breed er .

m u s t th erefo re wo rk fa s t er than nat u re and threm m at o logy can

,

n o t m ak e u se of a leis u rely o perat ing ev o lu t io nary pr inc iple u n les s

it s a ct io n can b e accelerat ed o r it s c u m u lat iv e effect s exaggerat ed .

Y e t a ga in m an ca nn o t a ffo rd t he im men s e n u m bers and t he wh ole

,

sa le d es t ru ct io n that charact erize nat u re s m eth od s of wo rk ing

’

C h a nges . A n im als and ev en plant s c os t m o n ey and a relat iv ely,

la r g e pro po rt io n m u s t meet t he c o nd it io n s o r t he ent erpr ise m u s t

,

b e a band o n ed .

B u s in es s c o n s id erat io n s th erefo re set ard u o u s lim itat io n s u po n

t h r em m at ology in respect t o bo th t ime and n u m ber s fro m wh ich
ev o lu t io n in gen era l is ent ir ely fr ee T he breed ing b u s in es s ha s
.

it s Own part ic u lar pro blem s so me o f t he mo s t im po rtant Of wh ich

,

u nfo rt u nat ely t he kn o wn fact s o f ev o lu t io n a re lea s t ab le t o

an s wer . The pro fitab le s t u dy Of th is su bj ect will h o wev er b e , ,

a s s is t ed by a c lear s tat em ent Of th es e pr o b lem s .

The prob l ems of t h e b reeder C ertain q u es t io n s s tand c lea rly

.

o u t in the m ind s o f pract ica l br eed er s and th o u gh an att em pt

,

t o an s wer th em s er ia t im wo uld n o t b e t he bes t m eth od Of s t u dy ,

and th ou gh so me of th em cann o t b e an swered with certainty

in t he present s tat e Of kn o wledge yet n o th ing is Of m o re c o n
,

s eq u enc e at t he o u t s et than that t he s t u d ent get a c lea r id ea o f

t he pr o b lem s n eed ing so lu t io n and to ward s wh o s e s olu t io n the

s t u dy o f thr emm at o lo y is d irect ed T h ey e s u b s tant ia lly a s
g a
. r

follo ws :
T O what ext ent are t he chara c terist ic s of an ind iv id u a l at m at u r
ity d u e to it s anc es try ( h er ed ity ) and to what ext ent a re th ey
,

d u e t o t he c o nd it io n s Of life (en v ir o n m ent ) su ch a s fo od c lim at e

, , ,

ex erc is e and gen eral care d u r ing d ev elo p m ent

,

A re the infl u enc es Of t he c o nd it io n s Of life lim it ed to t he ind i

v id u a l o r ar e th ey in c erta in in s tanc es and t o s o m e ext ent c a rr ied

Ov er u po n t he Offs pr ing ? that is a re t he effect s Of en v ir o n m ent

,

inh erit ed
Can va riat io n s b e d ir ect ly co ntrolled to any ext ent what ev er ,

or o n ly ind irect ly thro u gh select io n and by S pec ia l car e d u r ing

d ev elo pm ent
 I NT R O D U CT I O N
4

Ho w effect iv e is s ele ct io n in c o ntrolling v ar iat io n that is are

,

co ngen ital v ar iat io n s d ue ent irely t o parent age o r a re th er e back

o f t he par entage c erta in inh er ent and c o n s t it u t io na l t end enc ies

t hat largely fix t he gen eral d irect io n of v a r iat ion s ind epend ent
,

Of s elect io n

D o es im pro v em ent c o n sis t in ra is ing t he s tandard ab solu t ely o r

o n ly in ra is ing t he gen eral a v erag e by elim in at ing t he les s d es ir

ab le ? that is d oes breed ing im pro v e u po n t he bes t o r d oes it

,

on l br ing t h e gen era l m a s s n earer to t h e u pper lev el

y
T O what ext ent is ev o lu t io n a grad u a l pro c es s and t o what
ext ent m ay pr o fo u nd ad v a nc es appear s u dd en ly a s in S po rt s ?
,

and is t he on e c las s o f im pro v em ent any m o re perm an ent or reli

ab le than the o th er
D O all po s sib le v alu es Of a v a r iable ch a ract er a pp ea r o r are
c ertain v alu es s eld o m o r n ev er pres ent ed ? that is to say is ,

v ariat io n a lway s c o nt in u o u s o r is it s o m et imes d is c o nt in u o u s ,

mak ing c erta in th ings im p o ss ible bec a u s e t he pro per v aria

t io n s o r c o m binat io n s d o n o t appear o n which select io n may
b e ba s ed ?
What v ar ia t io n s are m o s t lik ely t o a p pear in s u cc es s iv e gen er
a t io n s Of any giv en br eed v ar iety o r typ e ?
, ,

A r e v ar iat io n s c o rrelat ed that is d o th ey t end at a ll to m o v e

,

t ogeth er s ugges t ing r elat ion s Of cau se and eff ect

,

What are t he pro per standard s for select io n How m u ch s hall

b e giv en t o u t ility a n d ho w m u ch t o appearanc e ?
T o what ext ent is ind iv id u al exc ellenc e a s afe g u id e t o breed
in g p o wer s ?
To wh at ext ent is t he Off s pr in g lik e t he im m ed ia t e parent and
t o wha t ext ent d oes it r esem b le m o r e rem o t e anc es t o r s

What is t he rela t iv e infl u enc e o f S ire and dam with r es pec t to

tra n sm iss io n o f charact ers ?
T O wh at ext en t d o t he c o nd it io n Of t he m ale at t he t ime Of
s er v ic e and t he c a re o f t h e f ema le d u r ing pr egnancy infl u enc e

t he O ffs pr ing ?

Wh at are t he r eal danger s from Clo se breed ing if an y and are

, ,

th ey c ertain or o n ly probable
Ho w can the ad v antages Of c lo se o r lin e breed ing b e realized
with o u t enc o u nt er ing it s d a nger s
 PA R T I V A R I A T I ON

C H A PT E R I

V AR IATI ON IN GE NERAL

SECTI O N I V A R I ATI O N U N I V ER SA L AM O N G LI V I N G
BE I N G S

T he m os t b fact abou t liv ing beings is th eir v ariability

O v iou s .

Not o nly d o s pec ies d iffer from ea ch o th er by m any and wid ely
d iffer ent cha ra ct er s b u t ind iv id u a ls with in t he S pec ies are d ist in
,

g u is h e d by d if ferenc es rea d ily d is c e rn ib le at leas t by t he t ra in ed

,

o bser v er T he gen era l d ifferenc es bet ween h o rs es and catt le for

.
,

exam ple a re s pec ific and d is t inct and th er efo re s trik ing ev en t o
,

t he cas u al o b s erv er ; b u t to t he tra in ed eye all h o r s es a re n ot

alik e and so it is that d ifferences are d et ect ed with in the s pec ies
,
.

Two ind iv id u al s m ay b e rec o gn ized as po sses s ing the sa m e char

ac ter s and th erefo r e r elat ed by d es c ent b u t in v ar ia b ly th ese
,

charact er s d iffer in d egree or in th eir pro po rt ion s o n e to ano th er .

T wo an ima ls m ay b e Of t he s ame o r Of d ifferent co lo r s b u t in ,

eith er ev ent t he pa rt s a re d ifferent ly pr o po rt io n ed The leg Of .

on e is lo nger larger o r m o r e croo k ed than that o f t he o th er

, , .

The bo n es com po s ing t he two are n o t Of eq ua l or ev en o f p ro ,

po rt io nal lengt h s Two co ws o f the sam e breed d iffer m arv el

,
.

ou s l in t he a m o u nt o f m ilk th ey c a n y ield in a y ear and s o me

y ,

are kn o wn to pro d u c e th ree t imes as m u ch b u tt er fat as o th ers

fro m t he sa m e a mo u nt Of the sa m e k ind o f feed A ga in so me1

.
,

milk is rich i n fat ( 6 o r ev en 8 per c ent ) wh ile o th er is poo r

( p
2 e r c ent o r ev en les s ) S o me. h o r ses beca u s e o f th ei r c o n
,

forma t io n trav el mo re eas ily o r m o re ra p id ly than o th ers and

, ,

so me a re mo re i nt elligent o r mo re end u ri ng o r mo re d oc ile .

1 l
S ee data fro m A gric u lt u ra E xperi m en t S ta t io n , U n iv ers it y of ll
I in o is .
8 VA R IAT I O N

So me d ogs (bloo dh ou nd s ) trail m arv elo u s ly well ; o th er s ( grey

h o u nd s ) s carc ely a t all S o m e h en s lay m o re eggs than o th ers
.
,

and of d iff erent c olo r and s ize .

S o m e an i m a ls are hard feed ers wh ile o th ers lay o n fl esh ,

read ily So m e beef is c o arse in its gra in ; o t her is fin e and ten

.

d er . So me is well marb led with fat ; o th er is n o t S o m et im es .

t he fla v o r is d elicate again it is rank and Often it is in s ip id , .

N O t wo trees a re alik e in th eir gro wth o r branch ing hab it ,

th o u gh similar with in the sa me v ariety and t he wid es t d iff erenc e ,

is o ft en f o u nd in lea v es fro m t he s a m e tree .

Differen ces ex t en d t o mi n u t e particu lar s an d in cl u de al l c h arac

t er s . t d ent S h o u ld early fo rm a c lea r c o nc ept io n Of t he
T he s u

fact that d ifferenc es extend to all characters h owev er in s ign ifi

c ant Or m in u t e Besid es he s h ou ld u nd ers tand t hat t hey inc lu d e
.
,

f u nct ion as well as s tru ct u re and that n o t o n ly ext erna l anatomy

,

and c o nfo r m at io n a re in v olv ed b u t int ernal o rgan s and th ei r

act iv ity as well and n o great er mis tak e c a n b e m ad e than to
,

d efine ev olu t io n as a s t u dy in m o rph olo gy .

”

I f we S O d efi n e t he wo rd v ari at io n a s to c o v er any change

in d etail Of s t ru ct u re o r f u nct io n wh ic h o u r fac u lt ies enable u s

t o d et ect th en we m ay s ay that v ariat io n ext end s t o a ll charac
,

t ers int ernal o r ext erna l s tr u ct u ra l o r f u nct io na l and if t he

, , ,

s t u dy la in t he r ealm o f eth ic s ec o n o m ic s ph ilo so phy o r r eli

y , , ,

gio n we Sh o u ld add mat erial o r im mat erial

, ,
.

T he i nd iv id u a l is th erefo re S O d is t inct ly a u n it that it s id en

t ity is at o nc e rec ogn ized and the princ ip le is c o nc ed ed that no
t wo a re alik e and that v ariat io n is u n iv ers al .

Limit ation of v ariab ilit y T he exc ept io n t o th e u n iv ers ality

.

Of v ari ab ility is in t he r ea lm o f n o n liv in g m att er The s pec i fic

-
.

g rav ity and o th er propert ies Of iro n go ld s o d iu m o r ch lo ri n , , ,

a re c o n s tant and t heir rela t io n s and c o m b in ing po wers with

,

o th er ch em ica l s u b s tanc es a re u nd er id ent i cal c o nd it io n s in v a ri

, ,

able and th eref o re well kn o wn .

Oxygen hyd ro gen and ev en ni tro gen and carb o n c o m b in e

, , ,

al way s in d efin it e pro p o rt io n s and th o u gh th eir c o m b inat io n s

,

a re exc eed ing ly n u m er ou s et w h e n t he c nd i t io n a e kn w n

y o s r o

the exact c o m b in at io n can b e f o ret old m o reo v er t he pro pert ies

o f th is c o m b inat io n will n o t o n ly b e d efin it e b u t th ey will b e

id ent ical with th os e of all o t her S im ilar c o m po u nd s I n th is .

 V A R I AT I O N I N G E N ER A L 9

way sod i u m C h lo rid for exa m p le ha s a lway s

, ,
d efin ite and well
known pro pe rt ies n o t s u bj ect to v ar iability .

I t is t o b e n o t ed of c o u rse that the pro pert ies o f the c o m ~

pou nd N a C l a re t o tally d ifferent fro m t he pro pert ies Of t he ele

ment s that c o m pose i t N a and C l T h ey a re n ev erth eles s d is tinc t
,
.

and in v a riab le as well as n ew Th is d is t inct io n bet ween the v aria

.

b ility o f liv ing m att er and t he c o n s tancy Of n o n liv ing matt er -

sh o u ld b e bo rn e in m ind lat er o n wh en d is c u ss ing s o me Of t he

cau ses Of v ariatio n .

SECT I O N I I —V A R I ABI LITY T H E BA SIS FO R

I M P R O V E M ENT

I m pro v ement is po ss ible o n ly wh ere v ariab ility ex is t s The .

co m po u nd NaC l being c o n s ta nt it wo u ld b e im po s s ib le to p ro
,

d u c e an im prov ed v ariety of sod iu m C h lo rid b ecau se t he c o m ,

po u nd is a lway s t he s a me and cann o t b e had with o th er than its

s tanda rd and i n v ariab le pro pert ies I m pro v ement in th is c o m .

mod ity is lim ited th erefo re to its m echan ica l fo r m and cann o t
, ,

ext end t o it s c o n s t it u t io n .

L iv ing m at ter u p o n t he o th er hand wh ile po s s es s ed o f d efi

, ,

n it e pro pert ies d oes n o t exh ibit thes e pro pert ies a lway s in t he
,

sam e d egree and o b s er v at io n and ex per ienc e ha v e b o th s h o wn

,

that pro fo u nd changes may b e mad e in eith er the fo r m o r the

cons t it u t io n o f bo th p lant s and an imals by the s imp le m eth od Of
j u d ic iou s c o m b inat io n s of d es irable d ev iat io n s .

I f th ere were n o v ariab ility and if liv i ng m att er were as c on

,

s tant in it s pro pert ies as is no n liv ing m att er th en we s h o u ld b e

-
,

c ertain of what we already hav e b u t n o im pro v ement wo u ld b e

,

po s sible A s it is wit h v ariab ility ev erywh ere liv ing o rgan is ms

.
, ,

a re bo th cap a b le Of im pro v em ent and liab le to d egen erat io n fo r ,

bo th a re the lo gical c o n seq u enc e Of v ariability Man m u s t th ere .

fo re wo rk for what he po ssesses in t he way Of an imals and

plant s and they will serv e him well o r ill acc o rd ing to his k n o wl
,

edge and s k ill in d ealing wi th th eir v ariat io n s .

A cc o rd ingly he canno t kno w too m u ch ab o u t t he v ariat io n s that

a re lik ely t o o cc u r —th eir nat u re th ei r ext ent and t he ca u s es
, , ,

that c o ntro l th ei r appea ranc e and d et er mine th eir perm anency

IO VA R I AT I O N

He cann o t kno w to o m u ch abo u t life and it s v ic is s it u d es abo u t ,

li v ing th ings and what th ey d o .

A n an ima l is b o rn int o t he wo rld I t s en erg ies a re fi r s t

.

d ev o t ed to n u trit io n and gro wth I t b u ild s it s o wn mach in e

.

and b u ild s it q u ick ly o u t o f m at er ials ly ing c lo se at hand I n .

goo d t ime it is fin is h ed and a ll i ts en ergies a re at a m ax imu m .

I t s eem s lik e a s tab le th ing that m u s t liv e f o rev er B u t repro .

d u ctio n o cc u rs s ec u r ing a s u cc ess io n Of its k ind On e a ft er

,
.

ano th er o f it s fac u lt ies fail and its c o nd it io n is aga in red u c ed t o

,

that Of bare ex is tenc e with y o u thf u l rec u perat iv e po wers go n e

,

f o rev er B y and by so m e V ita l f u nct io n fa ils T h en life go es

. .

o u t ; the o rga n is m br eak s d o wn and ret u rn s it s elem ent s to t he

in o rgan ic wo rld S u ch is t he brief h is t o ry o f a b it o f m a tt er

.

t em po rarily end o ed ith lif e

w w — fleet in g as a breath ; any
,

s erv ic e it m ay rend er u s m u s t b e ca u ght i n t he p as s i ng .

S ECTI O N I I I — N AT U R E OF VA R I A B I LITY

T he xact nat u re Of v ariability is a m o s t Ob s c u re s u bj ect

e ,

and o n e that canno t b e f u lly c o m preh end ed in the pres ent

s tat e o f kn o wledge Wh eth er t he d is t inct io n s bet ween liv ing
.

and n o n liv ing m att er will alway s rem a in as m ark ed as th ey

-

n o w s eem to b e o n ly f u t u re d is c o v eries will d et er m in e

,
We .

hav e as yet o n ly t o u ch ed t he fringe o f th is great s u bj ect b u t ,

en o u gh is kn o wn t o enab le u s to b egi n t o pen etrat e s o m e o f i t s

m y s t e ries .

A t lea s t two gen era l pr inc ip les m ay b e la id d o wn in t he

pres ent s tat e o f kn o wledge with o u t m u ch chanc e Of erro r
I
. T hat a ll cha racters o f p lant and an i m a l life wh eth er s t ru c ,

t u ra l o r f u nct io nal a re exc eed ingly v ariable

, .

2
. T hat o rd ina ry v ariat io n is t he res u lt Of a change in the
relat io n s bet ween a n u m ber Of as soc iat ed characters thro u gh the
d ev iat io n Of o n e o r mo re o f the m em bers and n o t t he in trod u c ,

t io n Of an abso lu t ely n ew charact er .

We s peak loo sely Of introd u c ing n ew cha ract ers b u t in

“
,

tr u th im pro v ement c o n s is t s n o t in t he intro d u ct io n Of ab solu t ely

,

n ew cha ract ers b u t in t he int en s ify ing o f d es i rab le Old o n es and

,

t he s u b o rd in a t io n Of th o se that a re u nd es irab le Fo r exam p le .

,
 VA R IAT IO N IN G EN E R A L 1 1

wh en we u nd ertak e to im pro v e the q u ality of woo l we lim it o u r

attem pt s t o t he s heep with wh ich woo l bearing is a nat u ra l
,

charact e r Wh eth er the h o rse or the ho g c o u ld b e mad e to gro w

.

woo l is a q u es t io n I f the hen c o u ld b e m ad e to pro d u c e m ilk

.

o r t he c o w to gro w feath er s that wo u ld b e the intro d u ct io n o f

,

a n ew cha ract er in the s trict es t sen se o f t he t erm .

Bu t n o th ing s im ilar t o th is has ev er been acc o m p lis h ed by m a n .

T he part ic u lar gro u p o f ch a ract er s th a t c o n s t it u t es a g iv en s pec ies

a ppear s t o b e s tra n gely fi x ed and im prov em ent s eem s t o c o n
,

s is t in c hangi ng t he r elat io n s o f th es e c haract er s a m o ng th em

selv es rath er than in t he intr od u ct io n o f n ew m em ber s Ho w .

t h is pa rt ic u lar gr o u p ing ar o se o r igin ally and ho w a n ew m em ber

( charact er ) m ight b e intr od u c ed a r e q u es t io n s fo r t he s t u d ent o f

gen era l ev o lu t io n Th ey are q u es t io n s m o reo v er u po n wh ich t he

.
, ,

present s tate of kn owledge S h ed s litt le light and s o far as is , ,

known t he s t udy o f the pract ical br eed er is lim it ed to m eth od s

,

of d ealing with gr o u p s o f charact er s a lr eady a s s o c iat ed and c o n

st it u t in g well m ark ed typ es and fo r m s

-
.

S ECT I O N I V — M EA N I N G O F T H E T ER M “
C HA R A CT E R

T h is is a
m u ch a b u s ed t erm loo s ely u sed in a v ar iety o f m ea n
-
,

ings . Fo r exa m ple when an ind iv id ua l d iffer s s light ly fro m

,

ano th er we say he has d iffer ent charact er is t ic s Wh at we r eally .

mean is s im ply that his chara ct er s d iffer in th eir d ev elo pm ent ,

no t that he ha s d iff er ent chara ct er s His bo n e is n o t s o ro u nd

.

or h is h ock s o cro o k ed t he cro p s a re n o t s o f u ll o r t he m ilk s o

rich ; t he eye of t he po tat o is n o t s o s u nk en o r t he c o lo r o f t he
fr uit so h igh in o n e spec im en as c o m pared with an o th er and ,

we say loosely that the cha ract ers a re d iffer ent .

N o w t he tr u th is t he C haract ers are n o t d ifferent in k ind b u t

on ly in d egr ee and pro po rt io n W e sa y o f o n e h o r se that he ha s
.

S peed and of an o th er that he has n o t s p eed T he fact is that

.

th ey bo th hav e so m e s peed b u t o n ly on e has en ou gh to attract

,

att ent io n and b e wo rthy of rem ark T his u s e of t erm s u n fo rt u

.
,

nat e as it m ay b e is pro bably too co m m o n to b e chan ged ind eed

, ,

t he m ere u s e o f t erm s is o f less im po rtan c e t han a c lear c o m pr e

hen s io n of t he fact s .
1 2 V A R I ATI ON
T he t erm charact er is em plo yed in th is t ext to d es ignat e o n e
o f th o se d eta ils o f fo r m o r f u nct io n wh ich t a k en t ogeth er c o n s t i , ,

t u te a well m ark ed gro u p o f an im als or p la nt s m o re or les s c lo sely

-

relat ed by d esc ent and t his is t he o nly sen se in wh ich the t erm
,

o u ght to b e u s ed T hus t he c o lo r charact er s o f t he h o r se ar e

.

black bay brown gray etc b u t n o t red gr een o r b lu e alth o u gh

, , , ,
.
, , , ,

these charact ers are n ot u nkn own to the animal wo rld being ,

co mm on with b ird s .

U sed in th is s en se a “
charact er belo ngs pr imarily to t he
,

race or gro u p o f wh ich t he ind iv id u al is a m em ber I t is th ere .

fo re n o t pec u lia r to any part ic u lar ind iv id u a l and is in n o sen se

perso nal property Th u s n ot o n ly the c olo r o f t he coat b u t the
.

fo rm of the body the pec u liar f u nct io n o f any o f its orga n s as in

, ,

m ilk prod u ct io n o r t he s ecret io n of po iso n s any S p ec ia l m en tal ,

att it ud e or int ellect u al fu nct ion or ev en a part ic u lar croo k o f

'

lim b o r s pec ial body m ark ing o f any k in d that ru n s c o m m o n ly

throu gh the gro u ps is properly s po k en o f as a rac ial charact er
, .

T h o se charact er s d o n ot c o m e and go b u t o n the c o ntrary th ey ,

r em ain with t he rac e ind efin it ely T he ind iv id u al h o r se for .

,

exa m ple will b e m ark ed by o n e o r po ss ib ly m o re o f th e c o lo r

—
,

C haract ers o f his k ind — b ack h t

l ,
w i e b a
y et c
,
b u t he will , , .
,

n o t b e m ark ed with charact er s n o t o f h is k ind as red green , , ,

etc . F r o m th is we s ee that t he ind iv id u al is n o t so goo d a

u n it fo r s t u dy a s is t he g ro u p t o wh ic h he b elo ng s and t h e ra c ia l

c haract ers that c o m p o se it .

N o w t he per so na lity Of the ind iv id u al s o s tro ngly im press es

u s that we in s t inct iv ely r egard him as an a ct u al u n it and we ,

s peak lo os ely o f h is cha ract er s as if th ey were p er s o nal pr o per ty

pec u liar to th is ind iv id u al alo n e wh erea s he po ssesses no th ing

,

that is n o t c o mm o n to his rac e His d iffer enc es are in d egr ee

.
,

n o t in k ind .

What we mea n to d esignat e in the ind iv id u al is the pa rt icu lar

c o m b inat io n of rac ial Charact ers that m ak e U p his perso n ality ,

kn o wing perfect ly well that the charact ers o f all ind iv id u als with in
t he rac e are rac ial charact er s and n o o th er and that ev ery ind i ,

v id u a l that m a ev er aris e by d es c ent will b e limit ed as t o h is

y
d etails to some co m binat io n o f t he charact ers of his ra ce N o w .

t h e charact ers o f any rac e a re S O m a ny th eir d ev iat io n s ar e s o ,

 V A R I A T I ON
1 4

th is we c o nc lu d e that t he a ppa rent m ak e u p o f t he ind iv id u a l -

d epend s u po n the parti cu lar charact ers that happen t o b e

s tro nges t that is
,
m o s t h igh ly d ev elo ped in his ca s e b u t that
, ,

he is i n act u a l po s s es s io n and m ay tran s m it any and a ll the

characters o f t he rac e to wh ich he belong s b u t n o o th er , .

W e n o w arriv e at t he d is t inct io n be t ween d o m inant and lat ent

charact ers wh ich is as fo llo ws :T h o se charact ers that a re pro m
,

i n en t in any i nd iv id u al a re s a id to b e d o m in a nt with h im b ec a u s e
well d ev elo p ed and p la in ly ev id ent and a ll o th er rac ia l charac
,

t ers a re s a id to b e latent bec a u s e n o t ev id ent a lth o u gh th ey a re ,

k n o wn to b e pres ent fro m the fa ct that th ey a re tran s m itt ed t o

t he o ffs pr ing o ft en bec o m ing t he d o m in a nt ch a ract ers i n f u t u re
,

gen erat io n s .

The t er m la t ent “
S h o u ld n o t c o n v ey the im pres s io n o f
h idd en o r lu rk ing charact ers b u t rath er u nd ev elo ped po s s ib il
,

it ies o f the ra c e with in t he ind iv id u a l in q u es t io n With th is .

c o nc ept io n t he s t u d ent will b e s a v ed m u ch menta l c o nf u s io n

wh en d ea ling with h ered ity and r ev ers io n .

Elemen tary ch ar ac t ers I n a b io lo g ica l s en s e t he u lt im at e

.

u n it o f v ariab ility th eref o re m u s t b e s o m eth ing les s than t he

, ,

rac ia l charact ers wh ich we h a v e been d is c u s s ing beca u s e th ey ,

th em selv es a re c o m plex rath er than s im ple We s peak o f t he .

leg o f a h o rs e o r t he q u ality o f m ilk a s a wh o le E v en if we .

n a rro w the po int to the c o nfo r m at io n o f the h o ck o r the pro po r

t io n o f fat we yet hav e ch a ract ers c lear ly m ad e u p o f p a rt s
, .

T he h o ck is an exc eed i ng ly c o m p lex s t ru ct u re and s ev en po s , ,

s ib l
y n i n e o r t en fa t s and o ils a re fo u nd in t he fat o f m ilk .

A s alm o s t u n lim it ed c o lo r eff ect s a re m ad e u p by few p ri

m ar ies in d iff erent pro p o rt io n s and a s all o rd inary m at eria ls a re
,

m ad e u p o f a few ch em ica l elem ent s in d ifferent c o m b inat io n s ,

so in a ll pro bab ility if we c o u ld m ak e t he u lt im at e a n a ly s is we

s h o u ld fi n d that a ll th es e charact ers a re m ad e u p o f d efin it e liv

i n g u n i t s that we m ay call fo r want o f a bett er nam e elem entary

, , ,

charact ers .

T h ese elem ent a ry charact ers ha v e rec eiv ed m any and v ario u s
names T h ey a re t he s t i rp o f Galt o n t he bio ph o rs o f Weis m ann
'

.
, ,

and t he phys io logical u n it s o f b iolo gis t s generally I n general .

th ey a re the sma lles t c o nc eiv ab le liv in g u n it s c o mparab le wit h ,

 VA R I A TION I N G ENE R A L 1 5
the mole c u le
ch em ical c o m po u nd s S u ch elementa ry cha rac
Of .

ters a re s u p p o s ed no t t o b e v ariable exc ept a s th ey effect d if

feront c o m b inat io n s a mo ng th em s elv es .

SECT I O N V I — T HE U N IT OF V A R I A B I L I TY

The n it o f v ariat io n is th erefo re n o t t he ind iv id u al b u t the

u

rac ial cha ra ct ers that c o n s t it u te t he partic u lar gro u p a n d that ,

run d o wn t he lin e o f d es c ent lik e t he s trand s o f a ro pe and o u t

o f wh ich ind iv id u als a re m ad e u p — s o m e with o n e c o m b inat io n

, ,

o th er s with an o th e r aft er the fas h io n o f t hread s in a fabric

, ,

form ing patt erns h ere and th ere n ow o f o n e d es ign n o w of ,

ano th er as th ey wand er apparent ly aim les sly here and th ere

,
.
‘

I t is ev id ent ho wev er that the act u a l bas is o f cha ract er d ev ia

, ,

tio n is somet imes exc eed ingly c o m p lex Milk s ecr et io n fo r .

,

exa m p le wh ile lim it ed t o c erta in an im a ls and c o nfin ed t o t he

,

female s ex is pro per ly rec o gn ized as a d is t inct charact er ; yet

,

its s u cc es s f u l f u nct io n ing d epend s u po n a v ariety o f c o n s id era

t io n s ,
— t he gen eral h ea lth o f t h e b od y t he n er v o u s t em pera ,

tu re o f t he ind iv id u al t he po wer of t he s t o mach to pro v id e large

,

q u ant i t ies o f pr e pa re d f oo d t he ab ili ty o f t he k idn ey s t o d o

,

t heir wo rk and the po wer o f ev ery o rgan in t he bo dy to d is

,

charge its f u nct io n su cc es s f u lly and f u lly u nd er h eav y s t ra in s .

A ll th es e a re a s im po rtant to s u cc es s f u l m ilk prod u ct io n as

are large and act iv e m ilk g land s a nd an acc id ent at any po int ,

will ca u s e d ev iat io n i n m ilk y ield eith er as t o q u ant ity o r q u ality

o r bo th .

D ev iat io n in c o lo r o n t he o th er hand m ay b e d u e t o pres enc e

, ,

o r ab s enc e o f p ig m ent wh ich m ay b e regard ed as a ch em ica l

,

su bs tanc e secret ed at a s ing le p o int I n th is and in s im ilar .

cas es t he act u al bas is o f d ev iat io n is S im ple and read ily d et ected .

F ro m th is it will appear that t he u lt im at e s eat o f v ar iat io n ,

wh os e flu ct u at io n s a re res p o n s ib le fo r c haract er d ev iat io n s m ay ,

b e exc eed ing ly d iffic u lt if n o t im po s s ib le to id ent ify .

1 Th e un it f i b il ity m t
O v ar a t b f us d with th
no it f l t i th
e c o n us e e un o s e ec on e

l tt
a er, o f i t h i d i id
c o u rs e, s l W
e n v t ua p . t h i h t
e c an n o b t m t
s e a ra e s c a ra c ers , u us

t k
a e him as h i f e s, b t or f w b t w m t d
e t er o r or f ll y
o rs e li i g th t
u e us o so u rea z n a

eac h f hi p t h t h
O s se a ra e c id
a rac ers t it y f it w
as a n th t th
en i of i s O n , so a e un t o v ar

a b il it y i f
s wi th i t h
ar n y it f l ti
e n ec ess ar un o s e ec on.
I 6 V A R I AT I O N

I t has been d that t he u lt imat e u n it o f o rgan ized

as s u m e

beings is the c ell Th is is tru e in a s t ru ct u ral s en s e o n ly fo r

.
,

th ere is po s it iv e ev id enc e that t he c ell is its elf m ad e u p o f

v ario u s and d is t inct elem ent s capab le o f s o m ewhat i nd epend ent

act io n in bo th c ell div isio n and gro wth The c o nt ent o f a c ell .

is n o t to b e regard ed as a m as s o f a m o rph ou s pro t o p las m t o b e

halv ed o r q u art ered by chanc e b u t o n the c o ntrary it is an ,

o rga n ized b o d y with a d is t inct d i fferenc e bet ween the n u c leu s

with it s d efin it e n u m ber o f chro m o so m es ( t he s u ppo s ed s ea t o f

the phy s io lo gical u n i t s that g iv e cha ract er t o i t s act iv it ies ) and
its s u rro u nd ing cyt o p las m o r foo d m at erial .

A ga in a wh ole gro u p o f s im ilar c ells m ay c o n s t it u t e a s pec ial

,

o rgan ( liv er k idn ey o r h eart ) d is charging a h igh ly s pec ialized

, , ,

f u nct io n q u it e d ifferent fro m that o f any o th er po rt io n of t he

body Th is c olo ny o f m any c ells d ischarging t he s a m e f u nct io n
.

appears to mo v e t o geth er th u s c o ns t it u t ing a k ind o f f u nctio nal

,

u n i t larger than and q u it e d is t inct fro m t he u ltim at e phy s io

lo g ical u nit s that m u s t res id e with in t he c ell .

Corr elat ed v ariation S t ill a ga in it is fo u nd in p ract ic e that

.
,

oc cas io n ally wh o le gro u p s Of charact er s s eem to b e s o c o rrelat ed

as to mo v e t o geth er s o that ha v i ng f o u nd o n e we m ay reas o n

,

ab ly expect t o d isc o v er the o th er F a m ilia r exam p les o f th is .

a re f o u nd in n ear ly a ll cases o f r ev e rs io n F o r in s tanc e a wh it e .

,

calf amo ng D ev o n catt le will alm os t c ertain ly S h o w black o r

bro wn po int s (ea rs n ose and legs ) wh ile a whi te S ho rt ho m will
, , ,

not The o n e is a c ase o f rev ers io n t o t he anc es t ra l c o lo r o f t he

breed — t he wild catt le Of B rit ain ; the o th er is s im ply t he
.

appea ra n c e o f on e of the n o rmal c o lo r cha ract ers o f t he rac e .

I n i ncreas e o f n u m b er s o f part s th ere is so m e t end ency t o repeat

a who le gro u p as in cases of d o u ble hand
,
1
.
”

T he sa me tend ency for m any d is t inct cha ract ers to mo v e

togeth er in gro u ps is fou nd in cas es of so called S po rt s in “ -
,
”

wh ich we in s t inct iv ely rec o gn ize so meth in g mo re than o rd inary

v a riat io n .

Th ese i n s ta nc es o f gro u ping of cha ra ct ers n o rm a lly in d epen d

en t in s u ch a way as aft erward t o mo v e t o geth er is t o b e d is

t in g u is hed fro m su ch v ariat io n as is in v o lv ed in extrem e m ilk

S d M i ti V i t i
1 ee u n er er s c ar a on .
 VA R IAT I O N IN G E N E RA L I 7

prod u ct io n in wh ich res u lt s a re to b e as cribed rath er to fo rt u i

,

tous v a ria t io ns a mo ng ind epend ent u n it s than to anyth ing lik e a

lin k ing t o ge th er o f t he s eparat e charact er s in v o lv ed .

The u l ti mate u nit T he s t u d ent m u s t m a inta in a c lea r V is io n

.

as t o d is t i nct i o n s o f th is so rt F ro m t he b io lo gica l s ta ndpo int

.

we acc ept as t he u ni t o f v ar iab ility the u lt im at e phy s io lo g ical

u n it s that m u s t res id e wi th i n t he c ell ; fo r p u rpo s es o f ev eryday

us e h o wev er we as s u m e as a u n i t that d eta il o f fo r m o r f u nct io n

, ,

that is im po rtant to t he breed er u nd ers tand ing perf ectly well ,

that c o n s id ered phy s iolo gic ally i t d o u bt les s c o n s is t s o f a n u m

, ,

b er o f u lt imat e u n it s wh ich lik e t he element s in a ch em ica l c o m

,

po u nd are ent irely capable of o th er and d is t inct c o m b inat io ns

, .

To r epeat we may as s u m e eith er o n e o f th es e c o nc ept io n s as

,

the u n it fo r s t u dy acc o rd ing to t he p u rpo se in hand I n t he .

text t he wo rd charact er will b e u sed to d eno t e that d etail Of

( rac i a l) s t ru ct u re o r f u nct io n w h ich i s i m p o rtant t o t he far m er
and to th is s t u dy The cha ract er will b e c o n s id ered as the
.

pract ical u n it o f v ariat io n kn o wing perf ect ly well that the u lt i

,

mat e u n it Of v ariab ility lies m u ch fart her back in t he c o n s t it u t io n

of the pr o t o p las m it s elf Wh en th is is in mind t he t erm s ele
.

menta ry charact er phy s io lo gical u nit o r so me eq u iv alent

“
” ”

, ,

term will b e u sed .

What ev er the S it u at ion t he s t u d ent m u s t n ot c on sid er t he

,

in d iv id u a l as the u n it o f v ar iab ility o r he will c o m e t o grief bo th

in s t u dy an d in pract ic e n o r s h o u ld he bec o m e c o nf u sed by
,

those oc cas ion al an d r em arkab le c o rrelat io n s o f charact ers that

s ugges t a u n it o f v ar iab ility u nd u ly large .

SEC TI O N V I I —D I STI N CT I O NS A S TO K I N D S OF
V A R I AT I O NS

I n t he c rit ical s t u dy of v ar iat io n it is n ec essary to o bserv e

certain d is t inct io n s that are o ft en ov erloo k ed res ult ing in m o re ,

or les s c o nf u s io n as t o what is r ea lly in v o lv ed in t he t erm

v ariat io n
“ ”
.

V ariati on q u anti tativ e or q u al ita t iv e T he fi rs t q u es t io n that .

sh o u ld ar is e in t he m ind o f t he s t u d ent t o u ch ing any d ev iat io n is

th is : I s t he d ifferenc e o n e in d egr ee merely (q u ant itat iv e) o r

 I 8 V A R I A TI O N

is it a d ifferenc e in k in d ( q u a litat iv e) Fo r exam ple on e h o rse

?
,

is exact ly lik e a n o th er o n ly la rger : t he d ifferenc e is q u ant i

,

t at iv e. A n o th er is n o larger b u t he c a n dra w m o re and has

,

great er end u ranc e : t he d ifferenc e is q u a lita t iv e On e c ow giv es .

m o re m ilk than a n o th er : t he d ifferenc e is q u ant itat iv e ; b u t a

th ird g iv es bett er m ilk a n d t he d ifferenc e is q u a litat iv e On e
,
.

apple is larger than a n o th er o f t he sa m e v ariety ( q u ant itat iv e

v ar iat io n b u t an o th er is d ifferent in t ext u r e and fla v o r q a li
) , ( u

ta t iv e v a riat io n ) .

\Vh en th erefo re in t he s t u dy o f a rac ia l charact er a s repre

, ,

s ent ed in t he s a m e o r in d ifferent ind iv id u a ls it is fo u nd t o ha v e

v ar ied t he fi r s t q u es t io n t o a s k and a n s wer is th is : I s t he d ev i

,

at io n o n e in k ind or m erely in a m o u nt I s it q u alitat iv e o r m erely

q ua n t it a t iv e ? I S t he ch ange to b e r egard ed a s o n e in nat u re or
o n ly in d eg r ee I f t he s t u d ent will carry th ese d is t inct io n s a lway s
in m ind h e will a v o id m u ch n eed less c o nf u s io n
,
.

V ariati on con tin u ou s an d disc on tin u ou s I t is n o t t o b e a s s u med

.

that v ar iat io n s d iffer fro m o n e an o th er by infin it es ima l increm ent s .

T he d ifferences m ay b e infin it es im a l (c o nt in u o u s v ar iat io n ) o r th ey

m ay b e d is cret e
“
( d is c o nt in u o u s v ar iat io n
) .

D a r win s u ppo s ed and it is c o m m o n ly a s s u m ed th a t v ar iat io n

, ,

is by nat u re c o nt in u o u s and that n ew fo rm s o r iginat e by t he

,

g rad u a l a cc u m u la t io n o f in s en s ib le d iff er enc es thr o u gh t he agency

o f lo ng c o nt in u ed s elect io n
-
T h is m ean s that if a ll t he ind iv id u als
.

that ev er liv ed c o u ld b e a s s em b led and s o a s so rt ed as to bring

n eares t t o geth er tho s e th a t a re n eares t alik e it wo u ld th en b e ,

fo u n d th a t th ey wo u ld grad e int o o n e an oth er by im percept ible

d iffer enc es and that a n y gaps that m ight o cc u r wo u ld b e d u e t o
,

t he effect o f s elec t io n in b lo tt ing o u t int erm ed iat e fo rm s .

N o w th is is a ha s ty a s s u m p t io n ind icat ing in th es e day s b u t

,

a s u p erfic ia l acq u a intanc e with t he m ann er o f v ar iat io n We .

ca nn ot as s u m e that a ll po s s ib le v alu es in v ar iable charact er s ar e

pres ent ed ; ind eed we kn o w v ery well that in many cases all
,

po s s ible v a lu es a re n o t pres ent ed and that som e int er m ed ia t e

,

fo rm s n ev er ar ise Fo r exam ple p each es o ft en giv e r ise t o

.
,

n ecta rin es b u t th ere is a ga p bet ween t he t wo that apparent ly is

,

n ev er fi lled D ar win called an o cc u rrenc e o f th is k ind a s po rt

.
,
”

a s if it wer e a n in s tanc e in wh ich a ll o rd inary la ws wer e s et a s id e ,

 VA RIATI O N I N G ENERA L 19

wh ereas it on ly s h ows that the va riat ion s of the pea ch are o ft en

disc on t in u o u s , with wid e ga p s represen t in g S pa c es n ot fi lled by
va riat ion .

To get a f u ll u nd er s tan d ing of th is gr ou n d the s t u d ent m u s t

form a c lea r co ncept io n of t he d is t inct io n bet ween c o nt in u ity an d
d isc o nt in uity d in th is c o nn ect ion
as u se .

A ma n gr ows fr om ch ildh ood to m at u rity I n d o in g s o he .

pas ses throu gh all possible weight s an d height s bet ween th ose of
infancy and m at u r ity W e cann o t r epresent all th es e v alu es by
.

any of ou r u nit s of weight or m easu remen t s becau se all n um bers

are by na t u re d isc o nt in u o u s The on ly meas u re of c o nt in u ity is
.

a lin e b ec ause a lin e c u rv ed o r s traight represent s all v alues

, , ,

sen sib le an d in sen s ible bet ween it s t wo extrem es W e can th u s .

plot co nt in u ity b u t we cann ot m eas u re it except by c u tt ing it

,

int o sect io n s and m eas u ring it at s tat ed po in t s as if it were d is

cont in u ou s ign oring the int er v als
,
.

C hanges o f t em perat u re are c o nt in u ou s as are th os e of h u m id ity , ,

illu m ina t ion and all gro wth in t he s en se o f ext en s io n in s ize

, ,

wh eth er pla nt or an im al A ll mo t io n wh eth er r egu lar o r irregu lar

.
, ,

is c o nt in u ou s beca u se a ll int erv en ing S paces are inc lu d ed .

D isc o nt in u ity o n t he o th er ha nd im plies v acan t S pac es n o t

, ,

represent ed by valu es The good s inger goes abru pt ly from on e

.

not e to t he n ext giv ing a d isc o nt in u ou s series of t o n es while t he

, ,

un s k illed v ocalis t S lid es u p or d o wn t he s c ale giv ing r is e to a ,

co nt in u ou s series o f t o n es in his effort to fi n d the pro per n o t e .

Th e latt er is n ot m u s ic bec a u se t he ear is n ot pleas ant ly a ffect ed

by th is co nfu sed j u m ble of so u nd wav es ar ising from the int er
med iat e t on es G ood m u s ic co n s ist s of a series of t on es n o t
.

fl owing int o on e an o th er b u t c u t s harply o ff and cas t int o a

d is co nt in u ou s series s trik ing the ca r at int erv als with sou nd
,

wav es that fi t with m ath emat ical prec is ion .

A ll n u m ber is by n at u re d is c o nt in u o u s By fract ion s we att em pt.

to bridge t he s pa c e b et ween c o nt igu ou s u n it s as bet ween I and ,

2 ; b u t h owev er s ma ll t he fract io n th er e is et a S pac e and a

y , ,

sen s ib le a nd m ea s u rab le o n e bet ween t he fract io n and t he n ext

,

u n it 1 9 9 9 9 is n o t 2 n o r will it ev er bec o m e 2 th is s id e o f
1 3 65 6
.
,

in fin ity by t he add it io n of any n u m ber o f n in es t o t he n u m erat o r

and c iph er s t o t he d en o m inat o r I t will c o n s tant ly appro ach 2
.
,
 0 V A R I AT I O N

b u t will n ev er reach it beca u s e d efin it e n u m ber is d is c o nt in u ou s

,
.

T h is is why we can n ev er acc u rat ely m eas u re c o nt in u ity exc ept

by a lin e and th is is why we cann o t expres s in n u m ber s t he
,

g r o w th o f an an im a l o r p lant exc ept appro x im at ely in t er m s o f ,

d isco nt in u ity .

A ll ch em ic a l c o m po u nd s ar e m ad e u p o f elem ent s in d efin it e

pro po rt io n T h ey are th erefo re d isc o nt in u o u s W e hav e H20
. .

and 14 202 b u t n o int er med iat e is po ss ible

,
th is again fo r n u m er ,

ica l rea so n s Plant s and an im a ls gen era lly are d im o rph ic t he o n e

.
,

fo rm being m ale t he oth er fem ale : th is is d isco nt in u ity S o me

,
.

s pec ies are tr im o rp h ic o r ev en po ly m o rph ic T he ant is eith er .

m ale fem a le wo rk er or so ld ier an d th o u gh th ey all belo ng to

, , , ,

t he s am e S pec ies th ere are n o c o nn ec t ing

link s in th is d is co nt in u o u s ch a in .

D im o rph is m with ou t res pect to s ex oc

c u rs in m any beet les and is exc eed ingly ,

m ar k ed in t he co mm on ea r wig as s h o wn1
,

in t he acc o m pa ny ing d iagram .

A T h e s hap e o f th is c u rv e s h o ws c lea r ly
Fm . L D im p h i m ill or
that h er e are t wo d is t inct fo rm s o f m ale
s us
.

t ra t ed t w ty p
: f th
o a large an d a s m all o n e liv ing t ogeth er
es O e ,

co mm mon ie 3 i and aris ing n at u ra lly o u t o f t he c om mon

ov s

m o re mm co th A on an
m a ss yet s h o wing alm o s t n o int erm ed iat es
’

d t h t yp i m
.
,
an e p e s o re ro

n o unc e d b t th , u S t u d ent s o f breed i ng fa mI lIar W It h t he

e re are

a lm tos i t m di t
"O n e
o ld er types o f Her efo rd will r ecall that t he
e a eS

A ft B er “ a l es
breed was almo s t d imo rph ic in that t wo d is
t inct types t end ed to appear with S ing ular perv ersen es s ref u sing ,

eith er t o blend o r t o u nd ergo m od ificat io n T h ere was t he old .

,

large s o lid bo d ied th ick m ea t ed d eep r ibbed type id eal exc ept
,
-
,
-
,
-
,

a s t o lat en es s o f m at u r ity ; th en th er e wa s a lso t he po ny b u ilt -

type —
,
S h o rt in t he bar r el and lac king in d epth beh in d th o u gh ,

well pro po rt io n ed in fro nt .

T he S h o rth o rn s a re a lmo s t po ly m o rph ic in po s ses s ing n ot o n e

b u t a v ariety o f types ea ch s ta nd ing o u t with extrem e d is t inct
,

n es s and n o t read ily m erged .

T he m o re t he m att er is exam in ed t he m o re it will b e s een tha t

s tra nge and u nacc o u ntab le gap s a re fo u nd ev ery wh er e M an y a .

B 1 M t i l f th
a t es o n , St dy f V i t i
a er a s 6—
pp 3 4 or e u o ar a on, . 2 .
 2 VA R I ATI ON

T he t d ent m u s t n ot th erefo re as su m e t he po ss ibility o f int er

s u

med iat e gr ad at io n s and in s en s ib le d ifferenc es wh en d ea ling with

biologica l ph en om ena Many of th ese d ifferences are es sent ially
.

and n ec es s arily d is c o nt in u o u s I t rem ain s to d isc ov er wh ich

.

these a re and t o d iscov er t he bea r ing o f d is co nt in u ity u po n t he

resu lt s to b e accom plis h ed by s elect ion .

I f all v aria t io n s were c o nt in u o u s we m ight h o pe to b e ab le

th eoret ically to acc om plis h any d es ired r es u lt and sec u re an y
d es ired s h ad e of d ifference by select io n ; b u t if n o t th en th ere ,

will r em ain n o tab le gap s that c a nno t b e fi lled T he nat u ral c o rol .

lary of all th is is that we can acc o m plis h by select io n alm o s t any

d es ired S had e of res u lt with th ose v ar ia t io n s wh ich are by nat u re
co nt in u ou s b u t that with th ose v ariat io n s wh ich are by nat u re
,

d is co nt in u ou s ou r effo rt s in th is r es pect will b e lim ited

, .

Di st in ct ion s ar i sin g f r om th e n atu re of th e cha ract er s in v ol v ed .

Hav ing d et er m in ed wh eth er t he d ev iat io n is q u ant itat iv e o r q u ali

t at iv e c o nt in u o u s o r d is c o nt in u o u s we n ext inq u ir e int o t he r ea l
, ,

nat u re o f t he v ariatio n as it a ffect s t he o rga n is m Man ifes t ly .

th is d epend s u po n t he charact er or charact ers in v olv ed .

Th o s e c o nc ern ed wi th fo rm will in th eir d ev iat io n s g iv e r is e to

, ,

m o rph o logical d iffer enc es On t he o th er hand d ev iat io n in char

.
,

a c t ers d is t inct ly f u nct io nal will giv e r is e t o d iffer enc es in o rga n ic

act iv ity with ou t rega rd t o fo r m .

A cc o rd ingly fo u r d is t inct ly d iffer ent k ind s o f v aria t io n are

recogn ized
I . M orpho log ica l relat ing t o d ifferenc es in fo rm o r size By
,
.

nat u re th ey are alway s q ua nt itat iv e b u t m a y b e eith er c ont in u o u s

,

o r d is c o nt in u o u s .

2 . S u bs ta n t i v e relat ing to d ifferenc es in q u a lity o f t he s t ru e

,

t u re as d is t inct from m ere fo rm or s ize By d efin it io n th ey are .

alway s q ua litat iv e and gen erally if n o t alway s c o nt in u ou s , ,

.

3 . Al er l
z s ti
‘

c ,
r elat ing t o d ev ia t io n s in pa t t ern es p ec ia lly as t o ,

repeat ed part s a s in extra fi nger s and t o es d o u bling o f petals

, , ,

s t o o ling o f ra in et V ar iat i n f th i k ind ith r q ant i

g c ,
o s. o s a r e e e u

t at iv e o r q ua litat iv e gen era lly t he fo r m er b u t a re o f n ec es s ity

, ,

d is c ont in uo u s .

4 Fu nc t iona l r elat ing to d ev iat io n s in t he n o rm al act iv ity of

.
,

t he v ario u s o rgan s and p a rt s o f t he body o r t he p lant s u ch a s ,

 V A R I AT I O N I N G EN ER AL 2
3

mus c ular ac t iv ity gland u lar secret io n s etc Th ey are eith er

, ,
.

quan t itat iv e o r qua lita t iv e c o nt in u ou s or d is co nt in u o u s th ou gh

, ,

rarely the latt e r .

A c lear u nd er s tand ing o f th ese d is t inct io n s is n ec essary to an

int elligent s t u dy o f t he nat u re and c a u ses of v ariat io n A cc o rd .

in gly en o u gh att ent io n will b e giv en t o ea ch to acq u ai nt t he

st ud ent with t h e way in wh ich v ar iat io n b ehav es part ly for its ,

o wn s a k e and pa rt ly a s prepa rat io n for c a ref u l inq u iry int o

meth od s of d ea ling with d ev iat io n s in th ose plant s and an imals

that we hav e d om es t icated and appropriat ed to o u r u se and ,

wh ich we wou ld see s t il l bett er adapt ed to t he p u rpos es o f m an .

S u m ma r y V ariability is t he u n iv er sa l r u le am o ng liv ing

.

beings L it erally n o two are alik e The d ifferences ext end to

. .

all c ha ra ct er s and t o t he m o s t m in u t e part ic u lar s N o n liv ing .

-

com po u nd s exis t in d efin it e pro po rt io n s an d th eir q u alit ies ar e ,

con s tan t n ot v ariab le V ar iability is the on ly bas is for im prov

,
.

ment N o im prov m ent is po s s ib le in t he s t rict es t sen se o f t he

.
,

t erm with res pect to in orga n ic c om po u nd s b u t liv i ng m att er being

, ,

va riab le is ca pab le o f change and th er efo r e o f im pro v em ent .

V ariat io n c o n s ist s no t in the introd u ct io n o f n ew c haract ers

b u t in d ifferent pr o po rt io n s o r r elat io n s a m o ng the o ld o n es .

A ll charact ers are rac ial and all ind iv id u als act u a lly po s ses s all
,

the cha ra ct er s of t he ra c e and n o n e o th er Th is is s h o wn by t he .

chara ct er s that are tran sm itt ed to the offspr ing .

The u nit o f v ar iab ility is in n o s en se th e ind iv id u a l th ou gh he ,

mu s t b e acc ept ed as the u n it for s elect ion The rea l u n it of d ev ia .

t ion is the rac ial cha racter b u t back of that in a biologi cal sen se
, , ,

lie t he elem entary charact er s o r phy s io lo gi cal u n it s wh ose v a ri ,

o u s c o m b inat io n s c o n s t it u t e ra c ial ch a ract er s .

V a riat io n is bo th q u ant itat iv e and q u al itat iv e bo th c o nt in u o u s ,

an d d isc o nt in uo u s and th ese d is t inct io n s s h o u ld b e c lea rly in

,

mind at all t im es .

S PE C I A L E x E R C I S E s

Prepa re a lis t o f v ari a ti o n s th at a re ( ) I q u a n t i t a tiv e,

( ) 2 q u a l i t a t iv e,

( 3) m o r p ho logi c al , (4) s u b s t an ti v e,
( 5) meri s tic , 6
( ) fu n c t io n al, ( 7) con

t in u o u s , 8
( ) d is c o n ti n u o us .
24 VA R IAT I O N

I I
A DD T ON A L REF ERE N CES

A N I M A LS AN D PL A N TS U N DER DO M E S TI C AT I O N . By C ha rles D a r wi n .

2 v o ls .

D I S C O N TI N U O U S V A R I A TI O N .
( A n ex a m p le .
) By E . R . S a u n d ers . Pro
c eed in gs of t he R o y al S o c iet y ,X I I 1 1 —2 5
L , .

OR I G I N OF S PE C I E S BY MEANS O F N A T U R A L S E L E CT I O N . By C ha r les

D a rwin . I v ol .

T HEO R Y O F OR G AN IC V A RI ATI ON . By H . S . W illia m s . S c ien c e, V I ,

7 3 84
-

T Y PE , H o w F I x E D .
( O n g en eti c en ergy o f o r a n is m s
g I S v ari ab ili t
y a n d .

n ot perm a n en c y th e n o rm al l w o f o rg n i c li fe )
a a ? B y H S W illia ms . . .

S c ien c e, V II , 72 I —7 2 9 .

V A R I AT I O N A ND SO ME PH E N O M EN A CON NEC TED W IT II R E P R O D U C T I O N

AND S EX . By A d am S ed gwi c k
S c ien c e, X I , 8 8 1 8 94 , 92 3 - 93 0.
-
.

V A R I AT I O N D I S C O N TI N U O U S ( S tud o f a rec en t v a riety o f fla t fis h ) B y

. y .

H C B u m p u s S c ien c e , V I I , 1 9 7
. . . .

V A R I AT I O N I N PL A N T S ( A s tu d o f the po rtio n s o f a lea f o n w ic h c hlo

. y h
ro ph yll is fo u n d E x peri m en t S ta tio n R ec o rd , X I , 4 2 3 I I
) . .

V A R I AT I O N IN TRI L LI U M G R A N D I PL O R U M .
( A rec o rd of the v a ria ti o ns
Ob serv ed in 1 85 c a s es
) By H W B ri tc her M ain e E x per imen t
—96
. . . .

S ta tio n B u lleti n N o 8 6 , pp . . I 69 I a lso E x peri men t S ta t io n R ec o rd ,

X I V , 6 34 .
 C HA PT E R I I

MORPHOLOGI CAL V ARIA TI ON

Morph olo g ic a l ar iat ion has referenc e to d ifferenc es in fo rm I f

v .

two or m o r e in d iv id ua ls po s sess t he sa m e s tr u ct u ra l charact er s and

if they hav e a ll attain ed t he sa m e relat iv e d ev elo pm ent th en t he ,

different ind iv id u als will d iffer o n ly in s ize b u t if t he charact er s

hav e n ot at t a in ed pr opo rt io nal d ev elo p m ent in t he d ifferent in d i
v id uals,
th en we S hall not e d ifferenc es in fo rm ind epend ent o f
mere s ize T h is is t he S im ples t of all fo r m s o f v ariat io n and is
.

t he o n e ch iefly in t he m ind o f Old er b io lo gi st s ev en lead ing to ,

t he m is tak e of s u ppo s ing that ev olu t io n is ess ent ially a s t u dy in

m o rph ology .

The ca u se of m o rph ologica l d ifferenc es may lie in extrem ely

fav orable or u nfav orab le c o nd it io n s of life es pec ially as rega rd s
,

food and c lim at e a ffect in g d ifferent charact ers d ifferent ly o r

, ,

th ey m ay arise from int ern al and con s t it u t io nal cau ses as in ,

giant s an d d wa rfs o r in s u ch extrem e d ifferenc es as in the m ul

,

berry leav es s ho wn in Fig 2 . .

I n s tan c es o f m o rph o lo gica l v ar iat io n a re so c o m m o n and s o

eas ily n o t ed as to s carc ely req u ir e m ent io n T wo app les ar e .

exact ly alik e exc ept that o n e is larger than t he o th er I t is a .

clea r ca se of m o rph ological v ar iat io n I n th is in s tanc e th ere is

.

n o d iffer enc e in t he charact er s o f t he t wo ind iv id u a ls exc ept that

cell d iv isio n and gr o wth hav e proc eed ed farth er in o n e cas e than
in t he oth er A s id e fro m th is th ey ar e id ent ical I f t wo s ta lk s
. .

of c o rn or if a n u m ber o f p igs S h eep c o ws or h o r s es a re exact ly

, , ,

alike except as to S ize th en th eir d ifferenc es are q u ant itat iv e

,

onl y an d t he effect is m o rph o logica l m erely

, .

A ga in t wo h o rses a re o f t he sa m e breed — that is po s s es s t he

, , ,

sa me cha ra c t er s — b u t th ei r chara ct er s a re n o t eq u a lly that is

, , ,

p r o po rt io nat ely d ev e,
lo ped I n o n e. t h e leg is lo ng er t he h o ck ,

S ho rt er o r t he fac e wid er b et ween t he ey es

, T h es e d ifferenc es
.

25
FI G . 2. Mo rph o logica l v a ri a tio n d diffe
illu s tra te : ren t y
fo rms o f m u lb err
lea v es p ic k edf ro m t h e sa m e t ree o n th e sa me d ay
 M O R P HO LO G I C A L V A R I AT I ON 2 7
are all q u an t i t a t iv e and mo rph o logica l b u t th ey infl u enc e fo rm ,

ra th er than S iz e as a who le beca u s e th ei r d ev elo p m ent is n o t ,

propo rt io n a l o n e part with ano th er .

V ar iati on sel dom simple I n s tanc es o f t he abo v e k ind are .

,

howev er e xtrem ely rare V ariat io n is so c o mmo n that o th er

,
.

d ifferenc es generally acc o m pany t hose o f f o r m T he t wo apples .

may d iffer in c o lo r fla v o r o r t ext u re a s well as in s ize in which

, , ,

cas e s u b s tant iv e v ariat io n has also o cc u rred On e of t he h o rses .

may ha v e an extra rib o n e o f t he p igs a so lid h oo f or o n e o f

, ,

the S h eep m o re fi ber s o f woo l t o t he s q u are inch in wh ich c as e ,

meris t ic v a riat io n is pres ent T he p u ls e o f o n e o f t he h o rs es .

may b e fas t er than t hat o f t he o th er o r t he m ilk o f o n e c o w ,

may b e rich er in fat s h o wing f u nct io nal d ev iat io n

,
.

A n d s o it is in pract ic e that t wo o r mo re fo rm s o f v ariat io n

may b e and lik ely will b e fo u nd pres ent in the s a m e ind iv id u a l .

Bu t h o wev er that m ay b e a ll d i fferenc es in f o rm o r S ize a re ,

rega rd ed as mo rph o lo gic al n o m att er what o th er d ifferenc es ,

may b e f o u nd and it is im po rtant that the s t u d ent early fo rm

,

t he hab i t o f d is t ingu is h ing c learly b et ween t he d ifferent k ind s

of v ari at io n pres ent ev en in t he s a m e i nd iv id u a l

,
.

Th e limi ts of size Ev ery s pec ies has a gen eral a v erage o f

.

s ize t o wh ich m os t ind iv id u a ls c lo s e ly appro x im at e A few ho w .

,

ev er ( giant s ),
great ly exc eed th is s ize and o th ers (d warf s) fa ll ,

far s h o rt of it A ll inv es t igat o r s agree u po n the c o nc lu s io n that

.

th is d iff erenc e in S ize is d u e t o t he n u m b er and n o t t he s ize

o f ind iv id u a l c ells ; in o th er wo rd s s ize is d ep end e nt u p o n t he ,

energy o f c ell d iv is io n T h is en ergy is exc eed ingly act iv e in

1
.

you th gra d u ally d ecrea s ing to zero at m at u rity exc ept as t o

, ,

c ertain part s ( repro d u ct iv e o rgan s s k in and so met im es the , ,

t eeth and h o rn s ) I n m o s t s pec ies acc id ent to a part will s t im

.

n la t e c ell d iv is io n lead ing t o a m o re o r less s u cc es s f u l r egen era

,

tio n F o r the m o s t part h owev er c ell d iv is io n d oes n o t pro c eed

.
, ,

rap id ly aft er m at u ri ty and t he lim i t o f it s act iv ity is in gen era l

,

t he limi t o f gro wth G iant s th erefo re repres ent exc es s iv e c ell

.

d iv isio n ab ov e the n o rm al and d wa rfs arres t ed d ev elo p m ent

, , ,

an abn o rm ally early c es s at io n o f c ell d iv isio n .

1 W ils o n , T he C e l l l
in D ev e o p m en t , pp 3.88 394
-
.
28 VA R I AT I ON

T he ca u ses inv o lv ed in th is abno rm al behav io r o f t he c ell in

d iv is io n a re exc eed ingly o bs c u re T hey a re c ertainly s o met im es
.

c o nnect ed with food and care in early lif e and no d o u bt th ey ,

a re o ft en c o n s t i t u t io na l Ev ery s t ock m an kno ws a b o u t t he

'

s t u nt ed pig ,
calf o r c olt and that it so met imes b u t rarely
, , , ,

rec o v ers ; t hat is t o say cell d iv is io n o nc e ch ecked d oes n o t eas ily

,

ret u rn to the no rm al rate T he t rai t howev e r eas ily bec o m es

.
, ,

c o n s t it u tio nal and hered ita ry fo r wh ole fa m ilies ( s t rai ns ) b ec o me

,

u nd ers ized and o th ers as m u ch a b o v e t he m ed iu m .

En ergy of gr owt h n ot t o b e con fu sed wi th b odi ly or fu n c

ti onal act i v i ty . W hile arger an imal o f his kind wheth er
t he l ,

it b e t he ind iv id u al o r t he s trai n represent s t he grea t er energy

,

o f c ell d iv is io n in bo dy b u ild ing it by n o m ean s f o llo ws that

,

t he bo dy when b u ilt will po s ses s a great er d egr ee o f act iv i ty

than will its no rm al o r s maller neighbo r ; ind eed the o ppo s ite is ,

lik ely to b e t ru e beca u se the larger bo dy wo rk s at great er d is

,

ad v anta ge hav ing greater in ert ia to o v erc o me and m o re dea d

,

weight t o carry ab ou t T h is is em inent ly tru e of all ani mals

.

wh o se serv ic e i n v o lv es t ran s po rt ing t he bo dy fro m plac e t o

plac e as amo ng h o r ses I t is m an ifes t ly im po s s ib le fo r a h eav y
,
.

h o rse to eq u al a light o ne in s peed with ou t t he expend itu re o f

far mo re po wer in d o ing it T h is is no t o n ly becau se of t he
.

ext ra weight b u t b eca u s e o f m echan ical d is ad v antages as well .

I n act iv it ies no t in v o lv ing m o t io n th is d ifferenc e in s ize wi th in ,

reas o nab le lim it s d oes n o t ex is t ; s m all men fo r exam p le a re

, , ,

d ou bt les s n o mo re and n o les s int ellect u a l than a re large o nes .

I mportance of morph ologi cal v ar iat ion N ext to th ose o f c o lo r .

,

d ifferenc es in s ize a re t he m os t n o t ic eab le of all v ariatio ns ; b u t

th ey a re by no mean s the mo s t S ignificant and their im po rtanc e ,

is lik ely to b e great ly ov eres t im at ed Exc ept in a few in s tanc es

.
,

as wi th d raft h o rs es m ere S ize is o f far les s c o n s eq u enc e than

,

is c o m mo n ly s u ppo s ed .

Generally s peak ing it is s o me q u ality o th er than b ulk that

,

d et er mines v alu e and it will b e fo rt u nate fo r breed ing wh en t he

,

po p u lar not io n that t he b igges t is the b es t s hall hav e passed

a way The larges t apple is no t t he bes t fo r eat ing n o r t he
.
,

larges t b u ll t he bes t fo r breed ing pu rpo ses Ho wev er th is d oes .

,

no t free the s t u d ent and breed er fr o m c o n s id erat io n s o f s ize ,

 C H A PT E R I I I

S U BS TA NTIVE VAR IA TI ON

S u b s tant iv e ariat io n has referenc e to d ifferenc es in q u ality

v

as d is t inct fro m f o rm o r S ize I t rega rd s t he c o m po s it io n o r .

m ak e u p o f t he bo dy o r it s m em b ers and refers to t he c o n s t itu

-
,

t io n o r inh erent nat u re of t he o rga n is m

, , .

Ev eryb o dy rec o gn izes d ifferenc es in m u s c les wh eth er fi rm ,

a n d s tro ng o r s o ft flabby and weak

,
W e d is t ingu is h t he bo ne
, ,
.

o f a h o r s e as d en s e o r as s o ft po ro u s and s po n gy T he h o rn , ,
.

o f t he h oo f is hard and t o u gh o r s o ft and S h elly .

Meat is fi n e in g ra in and h igh in fla v o r o r c o ars e in gra in and

lack ing in q u ality 1
I t is ei th er j u icy and r ich o r dry and tas t e
.

les s T he ga m y fla v o r o f wild m eat bo th o f m a m m a ls and b i rd s

.
, ,

is es p ec ia lly ta s t ef u l t o t he h u nt s m an and wh eth er d u e to breed ,

in g o r t o feed i t is c erta in ly charact eris t ic o f wild life ev ery wh ere

, .

N o t wo c u t s o f m eat a re a lik e wh eth er wild o r ta me and th es e , ,

d ifferenc es a re s o pro n o u nc ed a s to b e c o m mo nl y rec o gn ized ;

ind eed lang u age abo u nd s in adj ect iv es d es cr ipt iv e o f d ifferenc es
,

in q u a lity o f f o o d s t u ff s .

A t o n e t ime m ilk was s o ld o n t he q u ant itat iv e ba s is o n ly b u t ,

n o w t he per c ent o f fat is t he ba s is o f v a lu e T h e int elligenc e o f .

a n an im a l o r o f a m a n d ep end s les s u p o n t h e s ize a n d weight o f

t he bra in th a n u p o n t h e q u a lity o f t h e bra in m a tt er and t he

d epth of t he co n v o lu t io n s .

On e apple is s weet an oth er is s o u r a n d s t ill an o th er is in s ipid

, , .

On e fr u it is h igh ly fla v o red a n o th er is ta s t eles s T he s u gar o f , .

beet s of can e and of m aple is t he sam e ; b u t t he t wo fo rm er are

, ,

S im p ly s weet wh ile t he latt er is a cc o m pan ied by a h igh ly v o lat ile

,

eth er th a t add s a p ec u liar ly d elic io u s a ro m a

2
.

Pig f d h
1 s ily
e ttea v d m l m k on p k
co t l y fl
o n -s ee
g d wi t h ea a e 3 or s ro n a v o re

co tt d il S G i d l y j
o n -s ee o . l f Am
ee r n Ch m i l S i ty V l XX VI I
e , ou r n a o er t ca n e ca oc e , o . .

I l t d by K d i
9 so a e f th M i h ig e A g i lt
z e, o l C ll g f m mp l f
e c an r cu u ra o e e, ro sa es ur

i h d b y th
n s e th e au or .
 V A R I AT I ON 3 1

The c o nt ent of beet s varies fro m 4 or 5 per cent to o v er

s u ga r

2 0 per c ent I t is a lso exceed in gly v ar iab le in can e Wh eat is

. .

richer in pr o t ein than is c om b u t bo t h are v ariable and co rn , ,

has been bred with a pro t ein c o nt ent h igh er than that o f wh eat 1
.

Plant s d iffer in t heir ab ility to with s tand fro s t a s d o an ima ls to

res is t d is ea se A s ing le s talk o f c o rn m ay rem ain fres h and green
.

wh en all it s n eighbo rs ha v e been k illed C erta in ind iv id u als seem .

imm u n e t o pa rt ic u lar d iseas es and appear t o b e a ble to res is t ,

infect io n ind efi n it ely .

C olo r in gen era l is bas ed u po n d efin it e ch em ica l c o n st it u ent s

o r u po n t he charact er o f t he s u rfac es pr esent ed fo r r efra ct io n o i

light .I n eith er cas e it is a m att er o f inh erent q u a lity an d is

su bs tant iv e .

I mportance of su b stan ti v e v ariation T he s ign ificanc e o f s u b .

s ta n t iv e d iffer enc es d ep end s u po n t he in s tanc e S peak ing gen .

erall
y th,
es e d ifferenc es a re o f h igh v alu e T h ey a re u s u a lly .
,

th ou gh n ot in v ariably co rr elat ed with effic iency and in su ch cases

, ,

th ey po sses s a u t ilitar ian interest .

A d en se bo n e is bett er than a so ft and fibro u s o n e E v ery .

body prefer s a good a pple to a po o r o n e ; we hav e a d ec id ed pr ef

erenc e for c erta in ar o m as ; t he j u icy h igh ly fl av o red s t ea k is ,

bett er tha n t he dry t ou gh and tas t eles s o n e , ,

.

Colo r is a u t ility charact er a m o ng fl o wer s W e b u y th em fo r .

their colo r th eir fo r m and th eir od o r T h ey ha v e n o o th er v alu e

, ,
.
,

and of th ese charact er s th eir c olo ring is o f t he m o s t im po rt anc e .

Colo r h o wev er is in gen eral t he m os t d ec ept iv e o f all charac

, ,

t ers — d ec ept iv e beca u se it is str ik in g and bec a u se we great ly

,

prefer c ertain c olo r effect s ov er o th er s ev en th o u gh n o t c o rrelat ed ,

wit h u t ility W e carr y th is prefer enc e b ey o n d r ea s o n

. A red .

apple will sell fo r m ore tha n o n e of any o t her colo r ; yet we b u y

an apple n o t to loo k at b u t to ea t ; and n o o n e ha s s h own a c o r
relat io n bet ween c o lo r and q u ality in fr u it .

Hor ses with wh it e s k in a re pro v erb ially s u bj ect to c ert a in

d isea ses Fo r th is and o th er r ea s o n s c olor ha s n o litt le s ign ifi
.

cance in hor ses b u t a mo ng catt le it has pract ically n o m ean ing

,

what ev er ; an d yet ho w d ec id ed ly d o c o lo r m ark ings fig u r e in

1 Th l w t p t i
e o es t t di d i th b d i g p im t t th
ro e n c o n en s c o v ere n e r ee n ex er en s a e

U i
n v ers it y f I ll i i o p t d t ( 9 7) i 6
no s 3 p
u t o d h h igh
a e t i l 0 s . I e r c en , an t e es s
3 2 V AR IAT I ON

m any c r card s for p u re bred a n im als ! No w t he fact s are that

s o e -

a c o w is k ept for what s he can do and th ere is n oth ing inh erent
,

in mer e c o lo r that is ind ica t iv e o f h er ability to c o n v ert feed int o

eit her m ilk o r m ea t . S he is th erefo r e n eith er bett er n o r wo r s e
fo r her colo r exc ept a s it is an ind ex o f b lood lin es wh en S he is
t o b e u sed as a breed er .

S o s tr ik ing are c o lo r d ifferenc es h o wev er and s o d is t inct are

, ,

our prefer enc es t hat we in s t inct iv ely fo llo w o ur pr ej u d ic es in th is

,

r es pect q u it e regard less o f m o re im po rtant co n sid era t io n s u nt il

, ,

t he wh ole fabr ic o f br eed ing is int er wo v en with fancy po int s

“

in t he s ha pe o f co lo r m ar kings that gr eat ly c o nf u se t he breed er

in his att em pt s to s elect ind iv id u als for breed ing p u rpos es .

Tod ay t he m aj ority o f pr ize winn ing S h o rth o rn s are eith er

-

roan o r pu re wh it e Twenty fi v e year s ago n o breed er wou ld

.
-

hav e dared to S h ow a roan an im al and if a wh it e ca lf had been

,

dro pped in his h erd he wo u ld hav e d es troyed it at o nc e and k ept

t he m att er as secr et a s po s sib le so s tro ng was t he r ed c o lo r craze
,

following t he A mer ican wo r sh ip o f Bat es catt le T h is craze .

,

wh ic h was a lway s gro u nd les s c o s t t he breed and th eir ad m irer s

,

clea r ly a nd ch eck ed by a d ecad e o r m o re it s pr o gr es s u p ward .

Pro ba b ly o f all s u b s tant iv e v ar iat io n s c olo r is exc ept ing am o ng

,

fl o wer s and o rn am ental plant s of the leas t c o n sequ ence to man ;

,

et t he pr ej u d ic e is with u s and t h e br eed er wh o expect s t o sell

y ,

his prod u ct m u s t r eck o n wit h it He S h o u ld d o it int elligent ly

.
,

h owev er realizing f u lly that ind iv id uals v ary great ly in inh erent
,

q uality qu it e ind epend ent ly of c o lo r .

I n gen era l it m ay b e sa id that s u b s t a nt iv e d iff er ences th ou gh ,

n ot so ea s ily d et ect ed ,
ar e yet o f far m o re S ign ificanc e t o t he

farm er t han are those of eith er f o rm or S ize .

 C HA PT E R I V

MER I STI C V AR I ATI ON

Meris t ic t io n has r eference to a d ev iat io n in t he n u m ber

v a r ia

or arrangem ent o f repea t ed part s in v o lv ed in t he p lan o r patt ern

upo n wh ich any pa rt ic u lar o rgan ism is b u ilt .

A plant o r an an im a l is n o t an am orph ou s lu m p of liv ing

matt er On t he c o ntrary it is mad e u p o f part s each o f wh ich has
.
, ,

a kind o f id ent ity of its own m any of wh ich are s imilar and all
, ,

of wh ich ar e d efin it ely r elat ed an d p lac ed in so m e s o rt of o rd er ly

arrangem ent .

R efl ect io n d is c lo ses t he fact that each o rga n is m is d ev elo ped

u po n a s pec ific p lan es s ent ially d iff er ent fr o m that o f any o th er
, ,

and that with m ost o rgan ism s the patt ern is c om po sed of a d efi
n it e n u m ber o f s im ilar part s m o r e o r les s r epeat ed .

T h u s t he ch ick en has t wo legs and t he h o r s e has fo u r legs

, ,

that are m ore o r less alike T he fl ower ha s many petals t he

.
,

corn pla n t many leav es ; t he s p inal c olu m n is com po sed of v ert ebra
v ery m u ch alik e and o rgan is m s gen era lly po s ses s m any part s
,

tha t m o re or les s c losely resem ble o n e an o th er .

F ro m th is it appea r s that t he ind iv id u al an im al or plant is

n ot a u n it in res pect t o fo r m b u t rath er t hat it is m ad e u p o f
,

many u n it s som e of wh ich are pr act ica l d u p licat es T h u s t he

, .

id ea of mu lt iple pa rt s in o rd er ly arra n gem ent ( meri s m ) c o m es at

onc e int o t he s t u dy and v ar iat io n in t he n u m ber o r chara ct er o f
,

th ese repeat ed part s ( m eri s t ic v ariat io n ) is a broad and com pli

cated s u bj ec t a ffo rd ing c o n s id erable in s ight int o t he nat u re of

v a r iat io n
. A cc ord ingly it is pro fi table to p u r s u e it at c o n s id erable
len gt h n ot so m u ch fo r t he m at er ia l in v olv ed wh ich c o n s is t s
, ,

largely o f abn o rm alit ies o f n o pract ical int er est or v alu e b u t ,

beca u se n o oth er phase of v a riat ion a ffo rd s so m u ch inform at io n

u po n t he r ea l nat u re o f liv i ng m att er and it s adh erenc e t o o r

dev ia t io n from a d efin it e plan

 V A R I AT I O N
34

SEC TI O N I SY M M ET R Y

Th e c entral tho u ght in a ll meris t ic s t u d ies is sym m etry by ,

wh ich is m ea nt t hat o p po s it e s id es o f an o rga n is m po s s es s part s

t ha t a re eith er id ent ica l o r at lea s t S im ilar Th u s the petals u po n .

o n e s id e o f a fl o wer a r e in m o s t ca s es lik e th o s e u po n t he o ppo

s it e s id e and t h e b lo s s o m is m ad e u o f a n u m ber o f s im ila r part s

, p
v e ry m u ch a lik e and s ev eral t im es repea t ed
1
.

A m o ng h ig h er an im a ls h owev er Oppo s it e S id es are s im ilar b u t , ,

n o t id ent ica l an d h ere a r is es t he d is t inct io n bet ween rad ia l s m

, y
m et ry and bilat eral s y m m etry .

Ra dial symm etr y and radial serie s By th is is m ea nt that k ind .

o f patt ern in wh ich t he s epa ra t e pa rt s ar e id ent ic a l an d e a c h part

ca pable o f replac ing any o th er in the s er ies Comm o n exam ples .

a re t he peta ls of m o s t fl o wer s ( legu m in o u s and t he lik e exc ept ed ) ,

t he lea v es and la t era l S h oo t s o f plant s t he ca p s u les o f m a ny seed s , ,

s u ch as t he a pple o range et c t he ro ws o f c o rn U po n t he c o b
, ,
.
, ,

t he pa rt s o f t he s ea u rch in the s tarfi s h and t h e R a d iolar ia , ,

gen era lly

I n a ll c ases o f th is s o rt t he ind iv id u al part s c o u ld each replac e
any o th er pa rt o f the ser ies T he patt er n is th erefo re s po k en o f .

a s o n e o f rad ia l s y m m etry and t he part s as m em ber s o f a rad ia l ,

s eries .

Bilat eral sym metry A m o ng h igh er an ima ls a d ifferent sym

.

metry is o b serv a b le Wh ile each s id e has it s c o u nt er par t u po n

.

t he o th er yet th ere is a d is t inct io n bet ween t he r ight an d t he

,

left s id es and bet ween t he d o rs a l and t he v entral s u rfac es In .

s u ch c a ses t he pa rt s wh ile s im ilar ar e n o t a lik e n o r c o u ld th ey ,

r eplac e each oth er .

l Sy m m etry i s well -
tri
g }
: u n iv er s a l . A ll o rgan is ms a ris e t h ro u g h div is ion in
c ell

o ne o r m o re p a n es a n d l , so me d e
g ree o f s ymm t y e r is t o b e ex pec t ed fro m t h e

h h
m a n n e r i n w ic g ro wt ht k a es p la c e .

Bu t s ym m t y ie r d t m l ti ll l t t e A pp d g
s n o t c o nfine o u i t ce u a r s ru c u r s . en a es c on s s

in g fo i s n
g l ll
e ce f q t ly
s a re y mm t i
re l l y p l
u en d d m y
s g i m w h i
e r ca h ac e , an an o r an s s, c

a re s n i gl ll d
e ce d th
e f an mi p i d
ere o re i t m t k l t
c ro s c ow i t h c , as a o s , s ec re e a s e e on

re g l
u ar m k
ari g ny m m t i
s as s l h f t d
e r ca
q it b t
as if l o a r ro s an u e as ea u u .

A ll t h i i s i
s c u r o us th th l b l t th t d t f th m m t l gy wh
ra er an v a u a e o e s u en o re a o o o

is i t
n e res et d i m l t i ll l
n b i g ; y t it ll th w l igh t
u ce u ar e n t h m th d
s f lif
e a ro s on e e o o e,

an d w e a re a b l t l y d w th p i ip l t h t ym m t y i
e o a o n e t ly t h
r nc t l
e a s e r s no on e n a u ra

c o ro ll y f d
ar o l pm t b y
ev e o ll d i i i b t t h t it i l
en ce g e l p i
v s on,ip le i u a s a so a n era r nc n

l iv in g m a t t er .
 M ER I ST I C V A R I AT I O N 35

The right han d and ar m are mad e u po n the sa me plan as t he

left b u t c o u ld n o t replac e th em beca u s e th ey wo u ld n o t fit ; t he
,

on e is t he r ev er s e o f t he oth er .

R efl ect ed in t he m irr o r t he r ight hand seems to b e the left ,

b ut it is an illu s io n fo r t he right S id e is t he n ega t iv e or o pt ical

,

ima ge o f t he left with a ll its element s rev er sed Henc e a part .

u po n t he o n e S id e c o u ld n o t rep lac e t he c o rres po nd ing part u p o n

the o th er I t is it s c o u nt erpa rt n o t it s d u plicat e as a m o ng leav es

.
,

and petals T h us we arr iv e at t he d is t inct io n bet ween the c om

.

p le x i t y o f b ilat e ra l s y m m etry and t he s i m p l ic ity o f r a d i a l s y m m e try .

I t is als o s ign ificant that b ilat eral s y m m et ry is charact er is t ic o f

h igh er an im al life and rad ia l sy m m etry o f lower an ima ls and plant s
,
.

Dorsal an d v entr al su r f aces T he f u ndam ental fact at t he .

bott o m o f b ilat eral sy mm etry is t he d is t inct io n bet ween d o rsal

and v entral su rfaces n ec es s itat ing d ifferences in the q u adrant s
,

that ar e fo rc ed t o work in Oppos it io n to ea ch oth er 1

.

I nd ia n c o rn an d t he gra s s fam ily gen erally ar e as d is t inct ly

bilat era l as is the h o rse o r m an yet th er e is n o th o ught o f d o rsal ,

and v entra l d iff er enc es and h enc e n o d is t inct io n bet ween r ight ,

an d left .

Fo r exa m ple let t he lea v es of a c o rn p la nt and t he arm s o f a

,

man ext end eas t and wes t T h en the n o rth and so u th S id es o f .

the c o rn p la n t will b e alik e N o t so with t he m a n . I n t he o n e .

d ir ect io n ( we will say the so u th ) will b e his s pinal c olu m n and

the gen era l fra m ewo rk o f t he bo dy ; in t he o t h er ( to t he n o rth )
will b e h is fac e his n os e his ey es and a ll t he act iv e ext erna l
, , ,

par t s Mo reov er his hand s are m ad e t o Oppose each oth er and to

.

work t o geth er with th is ( the v entra l) s id e of t he b ody T h ere is .

th erefo re b ilat eral s ym m et ry in o n e d irect io n b u t n o t in the oth er .

W ith t he c o rn plant the ca se is d ifferent I t d oes n ot m ov e .

fr om p lace to place and it present s it s pla in s id es ind ifferent ly

,

t o t he wo r ld A cco rd ingly no d is t inct io n s s im ilar t o d o rsa l and

.

v entra l are po ss ible

2
.

1 Th w d e pp or it i “
o i h os d on t i t h s ere u s e f t g i m
no b t
n e s en s e o
“
an a on s u

as p l d ace
pp i t o d os w k i e an
g w it h T h
or t h n mb i pp d t as, t h e u s o os e o e

o th er m mb e f th h ers O d th b y m k i g w k i g it
e an , ere a n a or n un .

2 N f th
o ne o d i ti ti es e h ld b s nc f d with h m l g
o ns s ou p t
e c o n use o o o ous ar s or

wit h l g p t
ana o o us h l d t h id b
a r s , no r S f
ou d d e ea s e c o n o un e .

H m l g o o o
p t b
o us l g t t
ar s w i d i i
e ond l t
o do th y n v h ua s, no o n e. a n e a re s u c as

be a r c o rres
p di g t on t l
n l t i t
s ru c u ra t h i p re at i g ons i m gg
o t i g
e r res ec v e o r an s s, su es n
 6 V A R I ATI O N
3

B i lat eral s ymm et ry n ot c om pl et e. C u r iou s ly en ou gh no t a ll t he

pa rt s follo w t he sa me pla n as to b ilat era lism and sy m m etry W hat .

ha s been s a id r efer s to pa ired o rgan s s tand ing o n Oppo s it e S id es

o f the body a s hand s a r m s leg s ey es c a rs et c
, , , , , ,
.

Many o rga n s n o t pa ired present c u rio u s fact s t o t he ev olu

t io n is t The n o s e for exa m p le ha s n o c o u nt erpart b u t it s tand s
.
, , ,

o n t he m ed ian lin e and has a b ila t era l s y mm etry of it s o wn ,

being m ad e u p o f r ight and left halv es The liv er and t he h eart .

,

h o wev er wh ile c o ns is t ing o f right and left halv es are u n pair ed

, ,

o rgan s p lac ed n o t o n t he m ed ia n lin e b u t t he o n e u po n t he right

, , ,

t he o th er u p o n t he left Ea ch has a b ilat eral s y m m etry of it s .

o wn with d is t inct r ight and left S id es et bo t h a r e u n sy m m etr ic

, y ,

a lly plac ed .

The s t om ach o n the oth er hand is an u npaired o rga n ly ing

, ,

u n sy m m etr ica lly a cr o s s t he body and it s o wn b ilat era lis m is n o t ,

bet ween right and left b u t f r om f r on t to ba ck The k idn ey s pre .

s ent t he an o m a lo u s ph en o m en a Of paired o rgan s with a b ilat eral

is m of th eir o wn b u t at r ight an g les t o that o f t he body being ,

also from fro nt to back .

Lon gi tu din al sym metr y an d lin ear seri es I nasm u ch a s all .

g r o w th is by c ell d iv is io n we m ight expect lo n it u d inal s y m m etry

g ,

as well as lat era l s y m metry O wing h o wev er to t he d efin it e .

, ,

relat ion s of bo th an imals and plant s to the ext ernal wo rld it is ,

n o t m u ch d ev elo ped an d th er e is b u t a s u gges t io n o f lo ngit u d ina l

,

sy m m etry t o b e fo u nd in eith er p lant or an im a l fo r m s .

Mo st plant s are bo th geo tr op ic and helio tro p ic ; that is on e ,

part goes d o wn int o t he earth in r es po n se to grav ity and t he

o th er u p ward t o ward t he light and aga in s t grav ity Th is m ak es .

c o mmo n d es c en t . T h us t he leg of a m a n is h o mo lo go u s wit h h

t at o f a h o rs e o r a
b ir d , b ec a u s e o f s t ru c t u ra l res em b a n c es l . I n th e sa m e s en s e his a rm is h o m o lo

go u s wit h t he fo re leg o f t he h o rs e o r t he win g of a d

b ir .

A n alo go u s p a rts a re s u c h as s erv e t h e s a me

p p
u r o s e d iff in eren t o rga n is m s

h h
t oug s tr u c t u ra ll y d i t i s nc t . T h us th e fl ipp e r of t he wh l wh i h a e, c is m o ifi e
a d d
h d i
an , l
s an a o go u s to th e fin of a fish , a n d t he
gi ll of a fi s h is a n a lo go u s t o t h e
lun g l
ma m m a b ec a u s e it s erv es t h e s a m e p u rpo s e, t o u g t ere is n o s t ru c
of a , h h h
t u ra l rela tio n b etween th e two .

h l
T he o m o lo g u e o r t he a n a o g u e o f a p a rt is t erefo re t o b e fo u n in a n o t er h d h
d d l d
in i v i u a a n d o f a ifferen t S p ec ies S m met r o n t h e c o n t ra r wit it s c o ro la r .

y y y , y ,
h l
l
o f m u t ip le p a rt s refe rs t o in i v i u a s t a en s in
,
g d d l
a n d t o t he in t errela tio n s o f k ly
h
t eir pa rts .
 8 V A R LAT I O N
3

ho m re o t ic aria t io n s and h o m oeo is o f so m e so rt is a fr eq u ent

v ,
s

ac c o m pa n im ent o f m eris t ic v a ria t io n in lo n git ud in a l s eries T he .

b es t in s ta nc e o f th is is fo u nd in t he pet a ls o f fl o wers wh ich a re

rec og nized by b o t a n is t s a s m od ifi ed lea v es an d in s ta nces a re n ot ,

rar e in wh ich t he S pec im en p la in ly S h o ws t he v a r io u s t ra n s it io n

s tat es fro m lea f t o s epa l s epa l t o pet a l an d pet a l t o s ta m en

, ,
.

T his t end ency o f on e pa r t t o a s s u m e t he cha ra c t er an d d is

c har ge t he fun ct io n s o f a n eighbo ri ng pa rt is an im p ort a n t ph as e
o f v a r ia t io n thro wing m u ch light u po n t he gen era l s u bj ect o f
,

d ev elo p m ent .

\V ith th is introd u ct io n t he s t u d ent is prepar ed fo r t he s t u dy o f

m eris t ic v ar ia t io n so m ewh a t in d et ail in wh ich he will find that ,

wh ile each S pec ies is b u ilt u po n it s o wn so m ewhat pec u liar an d

d efin it e pa tt ern yet th is pa tt ern is s u bj ect t o m a ny a n d pro fo u nd
,

alt erat io n s and t he o rgan is m is fr eq u ent ly ab le t o exi s t u po n an

,

o th er and m u c h d is t o r ted p la n a ll o f wh ich goes far t o ward en light ,

en in g t he s t u d ent as to t he v ariat io n s that m ay b e expect ed in t he

o rga n ic wo r ld S t u d ies in m eris t ic v aria t io n ar e u s efu l to t he s t u

.

d ent of thremm at ology n o t so m u ch fo r th eir o wn s ak e as fo r t he

,

light th ey S h ed u po n t he n a t u re and m a nn er o f v a riat io n .

Ex a mples of meri s ti c v ari ati on E x am ples o f m er is t ic v aria .

t io n are to b e fo u nd at ev ery hand I n t he d ou bling o f fl o wers .

and the s t oo ling o f grain in increas ed o r red u c ed n u m bers o f ,

fi nger s and t oes in t he fo u r lea v ed c lo v er and t he branch ing

,
-

hab it o f m any p la nt s — ev erywh er e a r e S een a lt era t io n s in t he

,
'

c u s t o m a ry pla n o n wh ich nat u re d oes it s wo rk .

F o rt u nat ely an ext end ed and v a lu a ble c ollect io n of m eri s t ic

v ar ia t io n s m os t ly am o ng an im als has been m a d e by Ba t es o n 1
,
.
,

He lis t s his data u nd er 8 8 6 h ead ings each r ec o rd ing fro m o n e ,

t o s ev eral a u th ent ic cas es .

E q u ally c o m plet e dat a c o v er ing p lant s hav e n o t been c o llect ed ,

t ho u g h it is a mo ng plant s th at m er is t ic v ar iat io n is m o s t c omm o n .

I nd eed it is so c o mm o n and so ev id ent that fo rm a l c o llect io n is

,

hard ly n ec es sa ry The st u d ent is th erefo re referred t o pla nt life

.

o u t o f d oo r s a n d t o Ba t eso n s c o llect io n fo r a f u ller s t u dy o f th is

’

im po rtant s u bj ect a bare o u t lin e o f wh ich as a g u id e b eing a ll

, , ,

that is att em pt ed h ere .

1 B t M t i l f th S t dy f V i t i
a es o n , a er a s or [M m ill
e 8: C u o ar a on ac an o .
,
 M ER I ST I C V A R I AT I O N 39

SEC TI O N II — M ER I ST I C V A R I AT I ON I N LI N EA R SE R I ES

V ert eb re . A m o ng
fi s h es and s nak es v ar iat ion in t he n um ber
of v ert ebr a : m a b e v ery great I n mam ma ls it is s m aller b u t
y .

et d is t inct a s in t he f o llo wi ng exa m p les in s tanc es o f wh ich

y , , ,

acco rd ing t o Bat eson c ou ld b e m ult ip lied ind efin it ely

,
.

E HED
1
ERINAC EOUS EU R OP/EU S ( T H GE HOG )

Cerv ica l Do rsa l Lu m ba r Sa c ra l C oc cygea l TO T AL

I t will b e n o t iced that I and 5 d iff er o n ly in t he d o r sa l regio n ,

wh ich fact h o wev er a ffect s t he t o tal b u t that 4 and 8 d iffer bo th

, , ,

in t he sacral and t he c o ccygea l with o u t a ffect ing t he t o tal .

C o m m ent ing o n the ph en o mena o f an add it io nal lu m bar o r

s acra l v ert ebra Ba teso n s ay s ,

The e i a t gr
gg e t
si t h t ( i
s ro n e f t h
sui k i d ) th ms on a n c as s o s n e nu

b er o f v erteb rae h as b een i n c rea s ed b y s i m ple ad d i t io n o f a n ew seg men t

b eh in d , a fter h
t he fas io n o f a g ro win g wo r m ; the v a ria t io n o f v ert eb r a:
th u s seems a S im p le t in
g h B h
u t t ere is ev id en c e o f o t er
. i nd s , w ic h k h h
p la i n ly s h o ws h
t is V ie w o f the m a t ter to b e q u ite i n a d eq u a te .

What th is ev id enc e is he pro ceed s t o S h ow by s u cc eed ing

exa m p les a few o f wh ich are reprod u ced h ere
,

I n a S k elet o n o f Py t/zon t igr is ( N o 60 2 M u s eu m of Me Co lleg e

2 '

.
,

of S u r
g eo ns ) t he v ert ebr ae a r e n o r m a l u p t o t he 1 47 th i nc l u s iv e .

T he 1 4 8 th and 1 4 9th are h o wev er abn o rm ally s h o rt fr om fro nt

, ,

t o back s u gges t ing arr es t ed d ev elo p ment with im perf ect s epara
,

t io n alth ou gh each v ert ebra bears a n o rmal r ib on eith er s id e

,
.

1
B t M t i l
a eso n , t p O3a er a s , e c I b id pp
.
, 3 05 . I .
9 . . 10 -
1 .
4 0 VA R I AT IO N

Pa s s ing back ward in the sa m e S pec imen t he I 6 6 t h v ert ebra is ,

s een to b e n o r m a l o n t h e left b u t d o u b le and b ear ing t wo r i b s o n

t he right th u s g reat ly cro wd ing t h e r ib s o n that S id e

,
T he I 8 st h .

v ert ebra is r epo rt ed in t he s a m e c o nd it io n bo th being d o u bled ,

o n t he right S id e and S ingle o n t he left ( s ee F ig .

F o llo wing Ba t eso n giv es t wo ex a m p les o f t he rev er se c o nd i

1
,

t io n na m ely with d u plic ity o n t he left s id e and an o th er with

, ,

d u p lic ity o n t he right S h o wing ,

c lear ly that m er is t ic v ar iat io n in

o n e s id e o f a b ilat era l s y m m etry

may o r m ay n o t in v o lv e the o th er
S id e .

R ib s V a r ia t io n in t he d o r sa l .

reg io n n ec es sar ily in v o lv es t he

r ib s A s id e fro m th is all ev i .
,

d enc e go es to S h o w that part ial

d iv is io n o f the rib s is m u ch m o re
c o mm o n than is va riat io n in the
n u m ber o f v ert ebr a I n m an .
,

fo r exa m p le t he cart ilage is fre ,

q u ent ly d iv id ed fo r a co n s id erable
d is t anc e ( I 5 in ) back from t he . .

F M i i i i i t b
s t ern u m o ft en in v o lv ing a r ea l
IG
3. . er s t c v a r a
t on n v er e ra

d bl
ou igh t id
e on r A ft B ts e er a es o n
,

b ifu rcat io n o f t he rib it self

.

“
Home ot ic v aria ti on in v ert eb r a a n d rib s T h es e m a y b e o u t
1
.

lin ed a s fo llo ws ( a ll ln m a n exc ept as n o t ed ) ,

I . Cer v ic a l r es e m bli ng d or s a l : ba c k wa rd lt o m a os is ’
. T he d is
t in gu is hin g ch a ract er o f d o r s a l v ert ebra is t he bearing o f ribs ,

b u t th is ch a ract er is o ft en a s s u m ed by n eighbo ring c er v ical ,

being c o mm o n o n t he s ev enth and n o t u nkn o wn o n t he s ixth .

O f fi fty s ev en c a s es exa min ed by S tr u th er s fo rty t wo s h o wed

-
,
-

ribs o n bo th s id es and fift een o n on e s id e o n ly s h o wing a t end ,

en cy t o pr es er v a t io n o f sy mm etry T he c om plet en es s o f d ev elo p

.

m ent ranges all t he wa y fro m t he m eres t r u d im ent s ( rare) to a

perfect r ib c o nn ect ed by cart ilage to t he s t ern u m (als o rare) t he ,

co mm o n es t fo rm end ing free o r being j o in ed by cart ilage to t he

fi r s t tr u e rib .

B t 1 M t i l t
a es o n , p 5 I b i
a er a s , e cd p.
p 6, .8 10 .
3 . . 10 4 2 .
 M E R I S TI C V A R I AT I O N 4 1

2 . D orsa l r es em b l i ngf w a r d c er v ica l :

Izo m a o s i s N o t so or ’
.

com mo n as abo v e b u t S tru t hers d es cr ib es a s pec imen in wh ic h

,
l

t he fi r s t pa ir o f rib s is d efect iv e On t he left sid e t he r ib .

ex t end s b u t t wo fifth s o f t he wa ar o nd wh e r e it rt ic l t
y u a u a es ,

with a pr oc es s on t he sec o nd rib On t he r ight s id e it j oin s the .

c nd rib abo u t on e inch beyo nd the t u berc le A s s ev en n or

se o .

ma l c e r v ical v ert ebra are pr esent in th is s pec im en it is to b e ,

regard ed as a mod ifi ed d o r sal ra th er than an ext ra cerv ical ass u m

in g t he c haract ers o f t he d o rsa l as in t he prec e ding cases
, .

3 D
. o rs a l r e s e m b l i n
g l u m b a r F r e q.u e nt ly t h e t w e lfth r ib
in m an is ru d imentary in wh ich ca se t he las t d o rsal v ert ebra
,

ass u m es the fo rm an d gen eral appearanc e o f a lu m bar .

4 L
. u m b a r r e s em b li ng d o r s a l C a s e s. o f a th irt ee nt h r ib ar e

no t u nkn o wn b u t a re mo r e rare than t he red u ct io n o f t he

t welft h .

5 H
. o m a o ’
s i s b e t w e en l u m b a r s a c r a l
,
a n d c occyg ea l ,
T h e la s t .

lu m ba r m ay u n it e o n o n e o r bo th s id es with t he sacra l in wh ich ,

case the lu m bar d ev elops proc es ses to as s is t in t he su ppo rt o f

the il iu m On t he c o ntra ry t he firs t sacral m ay rema in d etach ed
.
, ,

th u s becoming pract ically a lu m bar S imila r relat io n s obtain b e .

t ween t he sacral a n d the coccygeal .

A c arefu l s t u dy o f th is wh o le s u bj ect d ev elo p s t he follo wing

fact s
I T hat an increa s e in t he n u m b er o f pa rt s in o n e regio n m ay
.

or m a n ot a ffect t he t o ta l n u m ber in t h e s er ies

y .

2 That c o n seq u ent ly a change in n u m ber in o n e regio n may

.

or m a n ot b e acc om pa n ied by changes in o th er reg io n s o f the

y
s a m e series ; that is changes in t he d o r sa l d o n o t im ply changes
,

in eith er the c erv ica l o r t he lu m bar .

3 T
. hat h o m o e o s is i n v e rt e br a and r ib s is c o nfi n e d t o m em ber s

co nt igu ou s ; that is if a cerv ical resem b le a d o rsal it will b e that

, ,

cerv ic al ly ing n ext t o t he d o r s al s er ies .

4 T
. hat t h e t en d e ncy i s f o r an e xtra m e m b er t o r es em b le

so m ewhat t he m em ber s o f t he n ext r egio n ; that is an extra ,

d o r sal is lik ely t o resem ble a lu m bar o r a c er v ical if n ot to ,

en t irely replac e it s u gges t ing that it ar o s e at t he en d and n o t

, ,

in t he m id d le of t he d o r sa l s eries
,
.

1 B at eso n , Ma t erials , etc .

, p . 1 09.
 V A R I AT I O N
4 2

5. T hat
f o rw a rd ho m a o s is i n o n e r egio n is n o t n ec es sa ri ly

att end ed by fo rward ho ma o s is in o th er reg io n s o f t he s a m e

s e ries .

6 T hat in gen era l ( es pec ial ly i n m a n wh ere th is has b een

.
,

m o s t s t u d ied ) fo r ward h o m oeo s is is at t end ed by a t o t a l increas e

o f t h e s eries and back ward h o m a o s is by a d ecreas e
,
.

7. T hat an abn o r m a lity o n o n e s id e m a

y or m ay no t b e

attend ed by a like a bn o r m ality o n the o th er th o u gh t he t end ,

ency is s tro ng ly t o t he pres e r v a t io n o f b ila t e ra l s y m m et ry .

8 T hat wh en o n e p a rt res em b les a n o th er it is t he m em be r

.

ly ing c o nt ig u o u s ; th a t is a d o rs a l v e rt ebra will res em b le a c er

,

v ica l o r a lu m bar n o t a s ac ra l and ly in g b et ween t he s t a m en

, ,

and t he leaf a re the peta l a n d s epa ] and all int er m ed iat e g ra da

t io n s eith er pres ent o r o b lit era t ed
,
.

M eris t ic v aria ti on i n s pin al n erv es B ra nch es fro m t he s p in al .

c o rd em erge bet ween t he v ert ebra s o th a t in gen era l the sys ,

t em o f s p ina l n erv es is de t er m in ed by t he v e rt ebra A s id e f ro m .

th is h o wev er t he em ergenc e o f t he bra nch es v a ries great ly

, ,

bo th in n u m ber and in c o nfo rm at io n ev en wh en t he v erteb ra ,

are n o r m a l .

F ii rb ri n ger s s t u d ies i n b ird s s h o w that the m in im u m n u m

1 ’

b er o f s p in a l n er v es f o r m ing t he brach ia l p lex u s ( s u pp ly ing t he

wings ) is three b u t in s o m e s pec ies it ris es as h igh as s ix
, .

Mo reo v er in so me in s tanc es the n u m ber v aried fro m fo u r to

,

fi v e wi t/t i n a s i ng le sp ec ies a n d in o n e ( the p igeo n ) t he v ari a

,
.

t io n was fro m fo u r to s ix A s m ight b e s u s pect ed t he t wo s id es

.
,

a re o ft en d iff er ent ly s u pp lied F o r ex a m p le in o n e s pec i m en

.
,

( g oo s e) t he p lex u s was f o r m ed o n t he ri ght s id e by n e rv es X V I

“
,

xvn xv m
,
and x xx wh ile o n t he left s id e it rec eiv ed a s t rand
, ,

fro m t he x x t h n erv e in add it io n to th ese .

F ii rb ri n ger s tab les s h o w th a t in s o m e s pec imen s o f t he goo s e

’

t he wing s were s u pp lied by t he n erv es x v t o x xx wh ile in ,

o th ers th ey were s u pp lied by t he x v n t h to t he x x t h I n the .

d o v e t he bra ch ial p lex u s wa s fo rm ed by the x th branch es of

t he s pina l n erv e in s o me s pec im en s by t he x u t h t o the x v t h ,

in o th ers and in o n e c a s e by t he n t h to t he x xv th as an
,

i nt er med iary .

B t M i l1 t —
9 35
a es o n ,
pp a t er a s , e c .
, . 1 2 1 .
 M ER I STI C V A R I AT I O N 43

Herrin gh a m d issect ed
their o rigi n the n erv es fo rm ing
1
to
the bra ch ia l p le x u s in fift y fiv e h u m an s u bj ect s ( th i r ty t wo f eta l - -

and t wenty three ad u lt ) Q uo t ing from his wo rk B at eso n s ay s

-
.
,

T he o ig i f t he l r e w t d i th i ty t w
n o e fo rt
u nar n erv as r ac e n r -
o c as s , u een

b ein g a d u lts It i n fo u r d iff eren t wa s

th e u ln ar n erv e was fo un d to a r ise
( ) .
y .

Mo s t c o m m o n ly it a rises from the v m t h a n d I x th ; t is o c c u rred in t wen ty h

h
t ree c a s es W ith the v m t h a n d I x th is so metim es c o m b in ed a s tran d from
.

the fi l t h, as s h o wn in fi v e c as es
( fo u r feta l , o ne a d u lt
) . In t h ree fetal
c as es it a rose fro m the v m th o n ly, and in o ne fetal a n d o n e a d ul t c a s e fro m
the v u t h a n d v m th . In sev eral c a s es the b ra n c h fro m t he m th was
v

m u c h l a rger tha n t h a t f om r the xx th , b u t the rev erse was n ev er m et with .

S im ilar c o nd it io n s were fo u nd elsewh ere with m a n the go rilla , ,

babo o n ,
and ch i m pan zee a n d the f o llo wing p rinc ip le was set ,

fo rth : A ny g i v en fi b er m ay a lter i ts p os i ti on r ela ti v e to tire

v er teb r a l c olu m n , b u t wi l l m a i n ta in i ts
”

p os i t i o n r ela t i v e to otlzc r fi b ers .

Hom e os is in in sect s an d ot h er s mall

an im a l s T he rep la c em ent. pa rt of one

by a n o ther wh ile c o mm o n a m o ng plant s,

( m o d i fi ed lea v es and s t em s ) is c o m par ,

at iv el
y rar e in an im al life I t is h o wev er .
, ,

by n o m ean s u nkno wn and s o m e s trik ing ,

exa m p les a re q u o t ed fro m B at eso n t o

s h o w t he rem arkab le m ann er i n wh ich a

perfect part m ay ar is e in a m o s t u n u s u al F I G 4 Ho m oc m ic v a ria t io n

w“ ? sa "8 m a n ten n a
plac e a m o ng which a re the f o llo wing
,
2
n o rm a ; l l ef t a n te n n a

b ea rin g a fo o t A and If,

S pec imen s o f s awfly CI m be r i l la r is in
'

<
.

I aI

h h the
w ic
.

left an ten n a en d ed in “
.

a well fo rm ed -
l
en a rge d — A ft. er B a t es o n

f ot ha i
o , v n
g a p a ir of n o rm al c la ws a nd the p la n /u la b etween t h em
F
( gi . R ig t h a n ten n a n o rm a l .
8

2 . A m ale b u m b leb ee ( B om b u s v a r ia bil zs

) ta en ink M u n ic h s h o wed the
left an ten n a “
p a rtiall
y d ev eloped as a foo t,
”
b ea r in g “a
p a ir of re ul a rl
g y
fo rm ed c la ws li ke the c laws o f th e foo t .

A m ale s pec im en of Z yg a n a lzpen d u la “ i u p ern u m era r

'

3 .
fi p o s s ess n
g a s y
win g a ris in g in suc h a p o s i tio n a s to s u
gges t t h at it rep la c ed a leg F
( gi .

T he ex t ra wing was o n t he left s id e and p ro ec tedj fro m the u n d ers id e o f the

b o d y a fter the ex a c t fa s io n o f t he leg, h wh ic h was a b s en t . T he s pec im en

B a tes o n , Ma teria l 1 3 5— d
1 s , etc 1 38 9 Ib i 1 47
.
, pp . . .
p . .

3 Ib i d .
pp . 1 4 6- 1 55 . Pro fes s o r B a t es o n h
v o uc es fo r t h e gen u in en es s o f t is h
s
p ec im en , wh ic h he h l
ims e f c a ref u ll y ex a m in e d l th
, a ou
g it b h
e on e
g t o D l
r K raa t z d . .
44 V A R I AT I O N
b elo n gs to M r . R ic ha rd s o n , a nd wa s ex a m in ed b y Pro fes s o r B a teso n as

c losel y as wa s po s s ib le wit o u t h rem o v i n g the h a irs , to wh ic h the o wn er

j
o b ec ted . I t is well k n o wn t a t h su
pe rn u m era r y
wings ma y a rise wit the h
n o rm al n u m b er o f leg s . I n t is h c as e the c lo s es t ex a m in a tio n fa iled to rev ea l

ev en a r u d im en ta ry leg, a nd h
t ere wa s c er

ta in ly no em p ty s o c k et or o th e r su
gges t io n

t h at th e res t O f the leg h a d b een lo s t .

”

4 . S pec im en of P a li uu m s p en ic i ll a t us wit h
an
“
a n ten n a-l i k e fl agellu m g ro wi n
g up fro m
the s u r fa c e o f the l f
( e t ) eye .

5 . T he fem ale c ra yfi s h has n o rm all

y a p a ir

o f o v id u c a l o p en in gs o n the b as es of the a n te

p en u lt im a te wa lk in g legs
p a i r o T h is s pec i
f .

m en po s s es sed in a dd itio n a p a ir al s o o n the

F I G 5 S u p e rn u m era r wi n g o n
. . y en ul tim a te, s h o win i rreg u la r se m en ta t io n
eft s i e — A ft e r B a t es o n
l d p g g .

6 A n o t h er s pec im en , a ls o o f the c ra yfis h ,

.

po s s ess es a n ex t ra
p a i r o f o v id u c al Open in gs , a s in the la s t, ex c ep t t h at h y
t e
were pla c ed o n the las t p a ir o f legs , s k ipp in g the pen u ltim a te I t is n o te .

wo rt h y t h a t th is is the n o rm a l po s itio n fo r t he sex u al o rga n s Of the m ale,

ex c ep t t h a t the Op en i n gs were pla c ed i n t h eir o wn p ro per p o s it io n o n the leg

a n d no t a t the po s ter io r s u r fa c e o f the j o in t a s the m a le Open in gs a re

”
.

7 . B a teso n h im s elf ex am in ed 5 8 6 fem a le c ra yfi h s fo r a b n o rm alit ies of

o v id u c al Open in gs . Of this n u m b er he fo u n d 5 6 3 were n o rm al an d 23

a b n o rm al, as fo llo ws
I . E x t ra ov i d l uc a o
p en in g on l eft pe l
n u t im a t e leg 7
2 . E x t ra ov id l uca O
p en in g h t p l t im t l g
o n ri
g en u a e e IO

3 . E x t ra ov id l uc a O
p i en n
g b t h p on l t i mo t l g en u a e e s I

4 . E x t ra ov id l
uc a op i en n
g b t h p
on l t i m o t d l en u a e an as t l egs I

5 . S in g l e ov id l
uc a o p i en n
g l f t idon l y
e s e on
3
6 . S in g l e ov id l
uca o p i en n
g ig h t id ly
on r s e on I

To t l b
a a m l p im
nor a s ec en s 23

B a teso n repo r ts b ut o n e a b n o rm al s pe c im en o u t o f 7 1 4 m a les ex a m in ed

b y him , a nd h
t is a b n o rm a li t y c o n s is ted in the a b s en c e O f a g en era t iv e o pen

in g o n the r ig h t s id e .

8 . A mong ea r t h wo rm s will b e fo u n d m a n y c a s es o f im per fec t seg m en ta

t io n , s h o wi ng m o re r in gs o n o n e s id e t anh u po n h
t he o t er, o ften s u gg es ti n g

a s p ira l r a t er t h ha n a series O f r in gs . G rea t irreg u lar ity is als o fo u n d in the

p o s i t io n O f g en era t i ve Open i n gs , a s to w h eth er p a i red or s i n gle, 1 alt h o gh
u

the male pa r ts a re alwa ys po s ter io r to the fem ale, w h a tev er the nu m b er of

the r in g o n w ic h h
ei t er is b o rn e h .

Cerv i cal fi st u l a and au ri cu l ar appen da ges in ma mmal s .

2

C erv ic alfi s t u la a re o pen ings in the n eck occ u rring s ing ly o r in ,

pair s and lo cated anywh ere fro m t he med ian lin e ba ck ward as
B t M t i l
1 t pp 5 6 66
a es o n , I b id pp
—
74 8
a er a s , e c .
, . 1 -
1 .
3 . . 1 1 0 .
4 6 V A R I AT I O N

On e of t he mos t
rema rkable ca ses ev er d es cr ibed is th a t Of
an infant bro u ght to G u y s h o s pital in ’
A n o th er wa s
Of a ch ild ha v ing a well d ev elo ped s u pern u m era ry a u r ic le o n
-

each s id e o f t he n eck ( s ee F ig T h es e appendages were .

ea s ily re m o v ed and pr o v ed t o b e ent irely c u tan eo u s th o u gh ,

each wa s s er v ed by a s m all art ery .

Wh eth er c er v ic a l fi s t u la a re t o b e regard ed a s rema in s o f

u nc lo s ed gill s lit s o r wh eth er th ey a re to b e rega rd ed a s rep et i
,

t io n s Of the ext erna l c a r in a n y ev ent th eir pres enc e s h o ws a

,

pro n o u nc ed t end ency to repeat c ert a in charact eris t ic s tr u ct u res

in th is part ic u la r reg io n o f t he b o dy .

Gro wth s Of th is character a re by n o m ean s c o nfined to m a n .

C er v ic a l a u ric les ( t he s o called wa tt les

-
a re c o m m o n in s h eep ,

esp ec ia lly m eri n o s T h ey a re well kn o wn in go at s and a re ex

.

c eed in gly c o mm o n in m a ny s trai ns Of u n i m pro v ed s win e S trange .

a s i t m a y s eem th es e repet i tio n s o f t he ea r app end ages a re u n

,

kno wn in eit her the h o rse o r t he o x .

Merist ic repeti t ion in ma mma 2

On e o f t he c hief d is t ing u is h
.

in g f eat u res Of m a m m als is m ilk s ecret io n S peak ing gen erally .

,

th is occ u r s at s o me po int o r po int s o n eith er s id e of the v entra l

s u rf a c e o f the bo dy o n lin es r u nn ing fro m the ar m p it t o t he

gro in I n s wine and in d o gs it is d is trib u ted thro u gh o u t the

.

ent i re ext ent o f th es e m a mm a ry lin es I n catt le h o rs es go at s .

, , ,

s h eep et c
,
it is c o nfin ed t o t he rear extrem ity o f t he lin e and
.
, ,

in t he elep ha nt it is a s d ec id ed ly f o rward t he u dd er being lo cat ed ,

at t he ar m p it I n t he hu m an b eing t he po int o f n o r m al act iv ity

.

is relat iv ely f u rth er back ( d o wn ) than in t he elephant b u t yet ,

ab o v e t he m idd le .

T h is latt er p o int is es tab lis h ed by t he fact th a t s u pern u m erary

n ipples are fo u nd bo th a bo v e and belo w t he n o r mal T he fact .

that n o les s than three s u pern u m erar ies hav e been fo u nd abo v e
ind icat es that t he n o rm a l m am m a a re perhap s fo u rth in a f u ll
s er ies. I t is t o b e n o t ed in th is c o n n ect io n h o wev er that in , ,

m o s t ca s es s u pern u m era r ies a r e s it u at ed belo w rat her th a n abov e

t he n o rm a l T h ese s tr u ct u res v a ry a ll t he wa y fr o m m ere n ipp les
.

resem b ling wart s and ent irely u nacc om pan ied by m am m ary t is s ue

1 Ba tes o n , M a t eri a l s , et c .
, p . 1 7 8 .
9 d
Ibi .
pp . 18 1 —1 95 .
 M ER IST I C V A R I ATI O N 47

up to f o r m ed o rga n s fu lly f u nct io nal C u r io u s ly en ou gh su per

well -
.

n u m erary m am m a are mo re c o m m o n in m en than in wo men .

Bat es o n q u o t es B ru c e as hav ing fo u nd in 2 3 1 1 fem a les f ou rt een

1

cases p e r c ent
) and in 1 6 45 m a le s f o rty s e v en,
ca s es -

p e r c ent ) I n an.o th er ser ies 3 1 5 s u bj ect s wer e exam in ed s h o w ,

in g t wenty fo u r cas es -
pe r c ent ) n in e t ee n b eing m a le and ,

fi v e f em ale Bard eleb en is a lso q u o t ed as hav ing exam in ed 2 7 3 6

.

recru it s (all m ales o f c o u r se) I n th is s er ies 6 3 7 cases

,
“
.

p e r c e nt ) w e r e s ee n 2 1 9 b e ing o n t he r ight s i d e,
2 4 8 o n t h e left , ,

an d 1 7 0 o n bo th s id es .

T he larges t n u m ber o f s u pern u m erary m a m m a ev er reco rd ed

was in a s u bj ect d es cr ib ed by N eu geba u er 2
T h is pa t ient had .

fiv e pa irs Of n ipples o f which t he fo u rth n u m bered fro m abov e , , ,

wa s t h e n o rm a l Wh en the ch ild was being s u ck led m ilk oo zed

.

from eac h Of the u pperm o st pa ir b u t all oth er s u pern u m eraries ,

y ield ed m il k o n ly with pres su re .

E xt ra t ea t s in c ows a re t o o c o m m o n t o n eed m ent io n exc ept t o

call att ent io n to th eir exc es siv e n u m ber T he c o w R o se fa mo u s .
,

fo r her r ec o rd a t t he Illin o is S tat io n had in a ll eight m am m a

3
, ,

s ix Of wh ich wer e fair ly well d ev elo p ed th ou gh o nly fo u r were ,

fu nct io nal I t is n o t ic eable that su pern u m eraries are n ea rly

4
.

always po s t er io r to t he n o rm a l o r else co n s t it u t e a d o u b ling o f o n e

o f t he n o rm als Ev ery m ilk er kn o ws by s ad experienc e that th ese
.

s u pern u m eraries ar e n o t o n ly c o mm o n b u t fr eq u ent ly f u nct io nal .

A c lo s e s t u dy Of th is su bj ect s ho ws that r epet it io n o f th ese

pa rt s m ay b e by pai rs or s ingly that t he repeat ed part s m ay b e
o n t he sa m e o r o n d iffer ent lev els that th ey m ay b e o u t Of lin e ,

being in so m e ca s es v ery n ear t he m ed ia n and t hat t he n o r m al ,

n ipple m ay b e d o u b led Fro m t he la tt er fact we f u rth er establis h .

t he p o int that m eris t ic v ar iat io n m ay o cc u r in t wo way s eith er ,

by add it io n to t he s er ies or by d iv is io n Of a n o rm al n u m ber We .

s h all fi n d t he s a m e in t eeth .

1 Ba t eso n , Ma teria l s , etc ., pp . 1 82 —1 83 .

2 Ib i d .
p . 1 83 .

3 S ee B u lleti n A ’
o . 66 .

4 T he s e s u p ern u m era ries were no t s y mm et ric a ll y p l ac e d . h

O n th e rig t s i d e the
two ex t ra t ea t s were p l d b h i d th t w f
ac e e n e o u n c t io n a l , as is c o m m o nly
th e cas e

b ut on l
t h e eft s i e d on ly p m y w
on e su ern u era r as so p l ac e d wh il
, e t he o t h er was
b et ween t he t wo fu n c ti lt t
ona ea s .
 8 V A R IA I ION
’ ‘

Merist ic v ar iat ion in t eet h . As Bat es o n rem ark s ,

“
Teeth
ar ise by s pec ial d ifferent ia t io n a t po int s alo ng the jaw a s m a m m a ,

ar is e by S pec ial d ifferent iat io n a t po int s alo ng t he m am m ary line ,

and we s hall s ee that with t eeth as with m am m a these point s

o f s pec ial d iffer en t iat io n m a fr eq u ent ly lie o u t s id e t he n o rm a l
y
r egio n th at t hey a re s u bj ect t o increa s e o r d ecrease in n u m ber
, ,

and that t he increas e m ay b e d u e eith er to t he add it io n o f a m em

b er t o t he series t o t he int erpo la t io n o f a m em ber o r to t he
, ,

d iv is io n Of a n o r m al m em ber .

Befo re c o n s id er in g s pec ia l ca s es it is well t o n o t e that t he

s im ila r ity bet ween t he r ight and left ja ws is that Of o rd ina ry

bilat eral s y m m etry b u t that th ere is a lso a k ind Of sy m met ry

, ,

n o t v ery c los e b u t s t ill m ark ed bet ween t he d ent it io n of t he ,

u pper and that Of t he lo we r w I s h o u ld b e f u rth er n o t ed

ja t .

that in m any a n im als as in t he s h a rk a lliga t o r etc t he d ent i

, , ,
.
,

t io n c o n st it u t es a s er ies in wh ich t he separa t e t eeth d iffer from

o n e an o th er m a in ly in s ize B u t m a m m a ls fo r t he m o s t part a re
.

h et erod o nt ; t hat is t he s eries is bro ken u p int o gro u ps wh ich

,

d iffer am o ng th em selv es th o u gh t he m em ber s o f t he separat e ,

gro u ps resem ble o n e an oth er c lo s ely T h u s t he inc iso rs are .

q u it e d ifferent fr o m t he can in es wh ich in t u m d iffer from the ,

prem o lar s a n d t he m o lar s T he d ifferent inc is o r s h o wev er .

, ,

a re v ery m u ch a lik e and t he s a m e is tr u e o f t he can in es and t he

,

v ar io u s m o lar s a n d
p r em o lar s M er is t ic v a r ia t io n in a het ero e
g .

n ou s s er ies lik e th is is m an ifest ly m u ch m o re c o m plic a t ed than

in a s im p le s eries lik e t he m a mm a o r t he r ib s W ith th is intro .

d u ct io n att ent io n will b e ca lled to a few s pec ial exam ples q u o t ed

fro m t he 2 3 7 ca s es that hav e been c o llect ed by Ba t es o n 1
.

1 . On e h u n d red a nd fi fty two -

a d ul t s k u ll s of an t h ro p o id a pes s h o wed
t welv e c a s es o f ex t ra tee t h O ew a i . n as n n c is o r , o n e was a n o m al o u s , a n d the
h
Ot er s were m o la rs Thi i e ly 8 p
s s n ar er c en t a b n o rm a l, a s aga i n s t 4 2 5
—
.

O ld W o rld m o n k eys t h t h o wed b t t wo

a s u c a s es O f ex tra teeth les s th an
,

one h alf O f 1 p er c en t .

2 . A d u lt o ra n g , wit h an a d d i tio n al p o s ter i o r m o la r on bot h s id es ab o v e

a nd o n the le ft s id e b elo w . N o t ra c e of ex t ra m o la r o n r ig ht si d e wa s d is
c o v ered , “ h
t o ug h th e e i r s a lm o s t a s muc h r oo m fo r it as o n the left s id e
.
”

E x tra m o la rs per fec t b u t s lig tl h y s m a ller th an th e nor mal .

1 B a t es o n , M a t e ria l s , e tc . ,
pp . 1 95 —2 7 3 .
 M ER I S TI C V A R I AT I O N 49

3 . S k u ll ( o ra n g ) No . 2 04 3 a , Ox fo rd M useu m , is n o rm al ex c ep t as to the
sec o n d
p rem o lar i n the u pper a w
j (p Bo th th es e tee t h a re m iss in
g fro m
their p o pe pla c e T h ere i ple t y o f s pac e o the left s id e b t so mewh at
r r . s n n u

le s t h an th e o rm al o n the righ t id e T he m iss ing too t h o f t he ig h t s id e

s n s . r

i p rese t in the s k ull b t in s tea d Of b ein g in i ts p ro pe plac e it s ta n d

s n , u
p r s u

f om the oo f Of the m o th wit h in the a rc a d e im med ia tely i f on t o f the

r r u n r

righ t c a i n e d alm o t e a c tl y o t he le el o f the sec o n d i c iso b ei g i

n an s x n v n r, n n

the p r em a x i lla a t s o m e d is ta n c e in fro n t of the m ax illa ry s u t u re .

D is c u s s ing
th is case Bat eso n Ob serv es ,

T h t th i too th i
a t ally the s d p em l wh ic h has b y som e m e
s ac u s ec o n r o ar an s

b ee h ifted i t th i po i ti
n s th b
n o d b t wh t s It hs th on ere c an e no ou a ev er . as e

e a t f
x c m f the se ord p m l od i Of f ll i It t d
c on ly ti
re o ar an s u s ze . s an s n ea r v er

c ally, b ut is a li ttle in c lin ed to ward s the o u ts id e . T he c a n in e is , b y t he

gr o wt h o f this too t h ,h tly sepa rated f om the sec o d i c iso
s lig r n n r, a n d the fi rs t

prem o l ar is c o nseq u en tl y p us h ed als o so m ewh t f rt h er b ac k a u . Hen c e i t

h a ppen s tha t the d ias tem a spac e fo r the s ec o n d p rem o la r o n the ri
g h t s id e

is not of fu ll s ize . T is h s ho ul d be u n d ers tood , as it m ig t o t erwis e b e h h

im agi n ed t h at the c o n t a c tio wa d e to a r n s u c o m plem en ta ry i n c rea s e i n the

s i ze o f the o th er teet h o f wh i c h t h e e is no e , r v id en c e.

Th e m is s ing pr emo lar ft s id e was n o t v is ible b u t o non t he le ,

“

t he left s id e Of t he pa lat e th ere wa s a v ery s lig ht elev at io n at a '

po int h omo logo u s and sy m m etrical wit h that at wh ich t he sec o nd

prem olar o n t he right s id e was plac ed A s m a ll p iec e o f .

bo n e was h ere c u t a way with t he res u lt that a t oo th o f abo u t t he ,

sam e s ize and f o rm at io n as p was fo u nd im bedd ed in t he b o n e

2 "
.

I n th is cas e th er efo re t he u p per pr em o lars o n b o th s id es had

, ,

“
trav eled away fro m th eir pr o per po s it io n s and t ak en u p n ew
and sy mm etr ical po s it io n s in t he palat e ant er io r t o t he c an in es ,
.
"

A s Bat eso n pert in ent ly rem a rk s ( ita lic s and pa renth es is m in e) ,

Th e fact s o f th is ca s e go t o s h o w th a t t he g er m of a toot/z c on
ta i n s wi t/t i n i ts elf a ll t/ze elem en ts n ec ess a ry to i ts d ev elop m en t i n
i ts own tr u e f or m [ev en in a n abn o r m a l po s it io n ] pro v id ed o f ,

c o u rse that n u tr it io n is u nres tr ict ed T h is is a s ign ific ant po int .

”

Of pec u liar int er es t t o s t u d ent s o f thr em m at o lo gy n o t b ec a u s e Of ,

it s bear in g u po n d ent it io n b u t b eca u se o f t h e light it a ffo rd s u po n

t he ba s is o f v a riab ility a n d t he u l t i m a te u n i ts o f v a r iat io n .

4 . G o rilla fro m the C o n go , wit h a fi ft h i n c is o r s ta n d in

g alm o s t in the
m idd le o f the lo wer j a w. I t has the c ha ra c teri s t ic c hi s el s h ape o f the in c iso r ,
b u t i t is t u rn ed h al f ro u n d s o h
t a t the pla n e O f i ts c h i el
s s t a n d s o b liq u el y .
”
 V A R I AT l O N
5 0

5 . D og wit h lo wer j aw and tee t h n o rm al , b u t wit h u pper c a n in es imper

fec tly d iv id ed . T he d iv is io n was m o re c o m p lete o n the ri
g h t s id e, fo rm in g
p ra c t i c a ll y two c a n in es s ta n d in g in lin e wit h the reg ula r teet h .

6 . D og wit h a w d o u b led , b o t
fi rs t p rem ola r in rig t s id e o f u pper j h h teeth
h
b ein g n o r mal in s ape, the a n terio r s om ew a t the la rger h .

FI G 7
. . M eris m in t eet h : c a n in es p a r t ia ll y d i id d v e . A fter B a t es o n

7 . D og wit h a n ex tra p rem o la r o n bot h id s es a b o v e a n d b elo w, the d en ti

tio n fo rm u la b ein g
S 5
p
5 S
8 S led ge d o g :
. A ll teet h n o m i al , ex c ep t left u
pper p h ”
. T is too t h n or

y
m a ll has two ro o ts . Here it is re
p res en ted b y t wo teet h ea c h h a
,
v in
g one

ro o t .

9 . A b s en c e fi rs t p rem o la r freq u en tly q u i te

Of c o mm o n k
in E s im o d ogs ,
su
gges t in g a b reed pec ul ia r i t y .

10 A m o ng d o m es tic d o gs s u pern u m era r m o la rs were fo u n d in twen ty

. y
h k
eig t c as es o u t o f 3 4 5 s u lls ex a m in ed , a s fo llo ws , the n o rm al d en t i tio n o f
1

the d o g a s to m o la rs b ein g t wo a b o v e a n d t ree b elo w ( m h

14
3 on bot h id s es an d o n one si d e
a
m on bo t h ids es

m on o ne s
l1 id e

m } a nd o n o n e si d e on ly
m 1 on bot h id s es

m 4 on o ne s id e on ly
T h is s tro ngly fo r m u la m 2 wh ich is that Of
su gges t s t he ,

Ot o cyo n ( La la nd c s d o g ) a n d o f the fo s s il A m ph icy o n t he s u p ,

po s ed d oglike pro gen it o r o f t he bears I t calls t o m ind t he .

f u rth e r rema rkab le fact s t hat Ot o cy o n it s elf v aries fro m m 2

t o m } and that i t is t he o n ly m a m m a l o u t s id e t he m a rs u p ial s

1 Ba t es o n , M a t eria l s , e tc .
, p . 2 20 .
 M E R I ST I C V A R I AT I O N 5 1

that ev er has f o u r m o lar s o n b o th ja ws wh ich go es far t o ind i ,

1

cat e a m ars u p ial anc es t ry .

R em em b er i ng that th e t eeth a re c o n s id ered a s o n e o f t he few

mo s t reliab le ba ses fo r c las s i ficat io n t he rem arkab le v ariatio n in ,

th ei r n u m be r charact er and po s it io n throws n o litt le light o n

, ,

t he m ann er in wh ich v ariat io n b eha v es which is t he ch ief r eas o n ,

fo r th ei r e xt end ed no tic e h ere .

S u pern u merary eyes T he d ev elo p m ent o f ext ra eyes s eem s

.

to b e c o nfi n ed to i n sect s wh ich a ffo rd a n u m be r o f exc ellen t

,

exa m p les o f t he d ev elo p m ent o f n o r m a l t is s u e in abn o rm al

s it u at io n s .

B at eso n s N o s 4 1 9 t o 4 2 1 a re all cas es of t he d ev elo p ment

’
.

o f a th i rd e e in C o leo pt era I ev ery ca s e th es e ext ra ey es a re

y n .

q u it e d is t inct fro m t he n o r m a l I n N o 4 1 9 the s u pern u mera ry . .

wa s s mall a n d la y a b u tt ing a ga in s t b u t d is t inct fro m t he r ight

e e I t s c o lo r wa s bro wn is h y ello w while t he n o r m a l e e wa s
y .
y ,

b la c k I n N O 4 2 0 the ext ra eye was o n t he left s id e b u t q u it e

. .

ind e pend ent o f t he n o r m a l ey es wh ich were exact ly a lik e I n ,

.

N O 4 2 1 t he extra eye wa s o n t he left s id e o f t he h ead wh ich

.
,

wa s rat h er les s d ev elo p ed tha n t he r ig ht T his eye is bo rn e .

u po n an i rreg u lar ch it i n o u s lo o p ha v ing a d ia m et er o f abo u t ,

m m T h is loo p is attach ed t o t he s u b s tanc e o f t he h ead

.

befo re and beh ind and th es e t wo attach m ent s a re d is tant fro m

,

each o th er abo u t 1 mm T he d ia met er Of the eye is ab o u t 2 5 m m

. . .
,

th u s o cc u py ing the f u ll s u rfac e of t he lo o p and its fac eting is ,

s a id t o b e n o t q u it e reg u lar and fin er a n d s light er than that Of

“
,

t he n o r m a l eye I t is th u s a v ery g o o d a ttem pt at a f u nct io na l

.

th i rd eye .

S u per n u merary w in gs in i n sect s B at es o n 2

repo rt s and de .

s cr ib es fift een ca s es o f extra wing s a m o ng i n s ect s s o m et i m es ,

on o n e s id e s o m et imes o n t he o th er b u t gen era lly if n o t a lway s

, ,

s m a ller than t he n o rm a l ; s o m et im es p la in ly id ent i fi ed with t he

fo re wing m o re freq u ent ly wit h t he p o s terio r ; o cca s io nally n o r

,

m a l in c o lo ring and s ca ling b u t a s a r u le abn o r m a l I n o n e in

,
.

s tanc e i t t o o k t he f o r m o f a la rg e u pr ight s cale and in an o th e r o f

a winglik e appendage t o t he left anter io r wing .

Lyd k k 1 L ib y f N t l Hi t y p 5 8
e er , ra r o a u ra s or , . 0.

B t 3 M t i l t
a es o n ,
pp 8 — 8 5
a er a s , e c .
, . 2 1 2 .
 VA R I AT ION
5 2

Meristic v ariati on appendages a ffo rd goo d

in horns .
1
T hes e
m at er ia l fo r s t u d ies in v ar iat io n T h ey so met imes c o n s is t o f .

h o rny matter ( catt le s heep rhino c ero s etc ) and s o m et imes o f

, , ,
.

tr ue bo n e as in t he d eer Th ey so met im es pers is t thro u gh life

,
.

( catt le s h eep goat s etc ) and s o m et im es are p erio d ically s h ed

, , ,
.
,

a s with the ant ler s o f t he s t a g T h ey s o m et i mes as i n catt le .

, ,

hav e a b o ny cas e wh ich is a t ru e o u tgro wth o f t he s k u ll b u t

, ,

o ft en a s in t he rh i n o c e r o s th ey ha v e n o c o nn ect io n what ev er
, ,

with t he b o n e b en eath A ga in t he ant ler s wh ich a re bo ny s ep

.
, , ,

arate wi th a c lean s car fro m t he bo ne o f the s k u ll as a leaf ,

s t em part s fro m it s t wig .

T he m eri s t ic v a riat io n s o f h o rns a re n o les s rem ark a b le than

th eir s u b s tant iv e v a riat io n s j u s t ment io ned Th ey a re fo r the .

m os t part s y m metrically p lac ed in pa irs o n eith er s id e o f t he

s k u ll j u s t ab o v e t he ey es th o u gh t he h o rn o f t he rh ino c ero s is
,

bo rn e u po n t he n os e and th erefo re u po n t he m ed ian lin e .

V ariat io n in n u m ber o cc u rs eith e r s y m m etrica lly o r as ymm et

ric a ll I f t h e rh in o c ero s has an extra h o r n i t wi ll b e j u s t
y .

abo v e and o n the med ian lin e wit h t he n o rm al S heep may hav e .

an ext ra pair j u s t ext ern al t o ( beh ind ) t he no r ma l o r there may 2

,

b e three o n o n e s id e and t wo o n t he o t her I n t he latter cas e the .

th ird h o rn will b e a litt le o n e ly ing bet ween t he n o r m al a n d t he

mo re o rd inary extra b o rn I n s t ill o th er ca s es acc o rd ing to
.
,

B at es o n a d o u b le c o re will b e f o u nd inca s ed i n a k ind Of

,

d o u b le barreled s ingle h o rn
-
.

A m o ng catt le n o increas e in t he n o r m al n u m ber o f h o rn s is

kn o wn t o t he writ er b u t th eir ent ire ab senc e is c o m mo n,
.

I nd eed the rea d in es s with wh ich t h e p o lled cha ract er appears is

,

as t o n is h ing pa rt ic u larly as it is a s s oc iat ed with a pec u liar pro m

3
,

in en c e ( t he po ll) ly ing b et ween and o ft en s light ly below the

no r mal bas e o f t he h o rns I n catt le meris t ic v a riat io n in ho rn s
.
,

s eem s t o b e a s s o c iat ed n ei th er wi th d iv id ed h o r n s o r ext ra

pro ngs .

1B ta es o n ,M t i l t pp 8
a er a s , e c
5 8 7 .
, . 2 -
2 .

9F h
o u r- db d
o rn e t k w B t
ree s a re n o M t i l t p 8
un
5 no n . a es o n , a er a s , e c .
, . 2 .

1It i s w ll k w f t t h t if i th
a e -
no n acp t b p ll d a t h h l met er a ren e o e e o rn s a re a os

t i
c er a n t b o b t i
e a th ff p i g
s en n d St e O in hi W il d W h it C t t l
s r n f , an o re r, s e a e o

G rea t B it i r y th i id
a n , sa s th t t h
ere s
p k t t
evl h ben c e l a es e ar ca e av e een s ev era

t im es a lt t l y p ll d
ern a e d h o d i th i i l
e an i th p k
o rn e s n ce e r n c o s u re n e ar s .
54 V A R I AT I O N

has t he al n u m ber Of d igit s b u t t he th u m b has three in s tead

u su ,

o f t wo pha lang es th o u gh it s general s hape is n o r m al I n the

,
.

left hand th ere is m u c h c o nf u s io n in t he r eg io n Of t he th u m b .

Th ere is a n extra d ig it b u t i t s t ru e cha ract er is n o t so ev id ent

,
.

I t is s h a rp lik e a finger b u t f u nct io n s a s a t hu m b

,
I nt erna l t o
,
.

th is is a th u m b lik e s u pern u m era ry with a tr u e nail b u t fro m ,

i t s po s it io n it is f u nct io n les s ( s ee Fig .

F IG .
9 . M eris tic v a ria t io n in t h e h dan ht d l
: rig an eft h d
an s of the s a m e in id
v i d l h
ua , s o win g on th e l e ft h dan a d pli ti
u ca on Of t he fo re fin ge r at the
e x p en s e O f t h e t h u mb , an d on t he r igh t h d an an ex t ra j o in t in t h e t h u mb .

A f t er B a t es o n

A fifth rea l finger ma k ing s ix d igit s in a ll is n o t u n c o m

, ,

mo n I t s tr u e p s it io n h o wev er is by n o m ean s a lway s eas y

. o , ,

to d et er m in e .

S p ea k ing gen era lly extra d ig it s m ay ar is e in three way s

, ,

eit her by add it io n t o t h e s e r ies o f an o u t s id e m em b er n ext t he

th u m b o r t he litt le finger by t he in s ert io n o f a m em ber at s o me ,

p o int with in t he s eries o r by the d o u b ling Of a m em ber J u s t, .

wh ich ha s tak en p lac e i n any giv en c a s e is n o t a lway s ea s y t o

d et erm ine .

R ed u ct io n in t he n u m b er o f d ig it s is c o m m o n and t a k es plac e
in three way s — b y t he lo s s o f a n o u t s id e m em b er ( gen era lly
,
 M E R I S TI C V A R I AT I O N 55

t he th u m b wh en the rad iu s is ab sent ) by the s u ppres s io n o f

, ,

a m em b er wi th in t he se ries ( ectro dact ylis m ) o r by t he u n io n of ,

two o r m o re m em b er s ( s yndacty lis m ) .

S yn d acty lis m m ay o cc u r in a ll d egrees fro m m ere webb ing ,

to a rea l b o ny u n io n as in t he ca s e o f s o lid ho o f ed h o gs F ig 1 0
,
-
. .

F IG . 10
. D egrees of s y d
n ac t ylism in d igit s : th e e n era
g l h s ape o f t he m em b er is
p d
res erv e ev en w h en o n e d igit i s s u p p ress e d . A fter B a t es o n

e xh ib it s a cas e B in wh ich t he n o rm al s hape is preser v ed in

, ,

t he ab s enc e o f a m em b er (ectr o dacty lis m ) with n o th ing t o s u g ,

ges t a u nio n ; that is to say d igit s I I I and I v s eem to b e f u lly

,

repres ent ed by a s i ngle d igi t n o rm al in charact er b u t rep lac ing

,

t wo m em b ers o f t he u s u a l s eries .
 6 V A R I ATI ON
5

To f u lly apprec iat e the s ignificanc e o f th is s u bj ect we n eed

to rem ind o u r s elv es o f ev o lu t io na ry h is t o ry wi th res p ect t o d igi t s .

Ma n for exa m p le has n o rm ally fi v e d igit s in a ll extrem it ies

, , .

The s a m e is tru e o f t he bat T he o x has o n ly t wo t oes and t he

.

h o rs e b u t o n e yet th ere are r u d iment s Of o thers in bo th cas es

, ,

s tr o n gly s u gges t ing that at s o m e rem o t e p e rio d t he n u m ber m ight

hav e been great er .

A ll th ing s c o n s id er ed it loo k s a s if fo r s om e u n exp lain ed and

, ,

at pres ent u n expla inable reaso n an im al life in m o s t o f its h igh er

,

fo rm s had been o r iginally co n s tr u ct ed u po n a pla n o f fi v e a s

r ega rd s the extrem it ies T ru e m any if n o t m o s t s pec ies hav e
.
, , ,

lo ng s inc e d epa rt ed m o re o r les s wid ely fro m t h e o rigin a l plan ,

and yet t he n u m eral fi v e is as d is t inct ly charact er is t ic o f t he d igit s

in an im als a s it is o f peta ls in t he ro s e fam ily a m o ng p la nt s .

Ho w th is n u m ber ha s been g ra d u a lly red u c ed to a final form ,

s o m et im es o f t wo a s in ca tt le s o m et im es o f o n e as in h o r s es
, , , ,

is a chapt er in d ev elo p m ent that belo ngs t o t he anc ient h is t o ry

o f ev o lu t io n
. Mo r eov er it is a chapt er that for o b v io u s rea so n s
, ,

m u s t b e read back ward s and rec on s tr u ct ed fro m it s f ragm ent s .

I t will a s s is t in t his r ec o n s tr u ct io n and what is o f m o re

, ,

c o n sequ enc e to t he br eed er it will thro w m u ch light u po n the

,

m ann er o f d ev elo p m ent and t he u n it Of v ar iab ility if t he s tu ,

d ent will c o n s id er t he pres ent co nd it io n and ev id ent anc es try o f

a few charact eris t ic s pec ies with res pect t o d igit s .

Bat s ha v e fi v e d igit s in b oth wing and leg tho u g h t he th u m b ,

is m od ifi ed in t o a s tro n g c la w .

B ird s ha v e three d ig it s in t he wing n am ely 1 t he th u m b , , , ,

wh ich ma k es t he s o c a lled bas tard wing ; 1 1 and 1 1 1 wh ich mak e

-
,

t he tru e win g ; I V and V m is s ing ,

R ad iu s and u lna are both
.

present I n the leg t he fib u la is a m ere s plint ly ing by the

.
,

t ib ia
.

T h ere is b u t o n e m etatarsu s b u t it is large and h ea v y end ing

, ,

in thr ee p u lley lik e su rfac es ov er wh ich play t he t end o n s that a re

-
,

attach ed to t he three t oes d irect ed fo r ward ( 1 1 I I I and I V ) T h is , , .

plan s u gges t s that t he three m idd le m etatar sals o f t he n o rm al

foo t hav e h ere bec om e u n it ed alo ng t he s ha nk int o o n e b u t ,

with three su rfac es pres erv ed fo r attach m ent Of d igit s Mo s t .

b ird s hav e also a toe beh ind T h is is rega rd ed as d igi t 1 b u t

.
,
 M E R I S TI C V AR I AT IO N 57

no bird has s ho wn a trace o f V A dd ed t ogeth er th is all m ea n s

.
,

t hat bird s h a v e los t d igit v from t he leg if th ey ev er posses sed it , ,

and I v and V fro m the wing with I in a fa ir way to u lt im at ely

,

d isappea r fro m bo th wing and leg exc ept wh en f u nct io nal in t he ,

la t t er .

The ca t has n o rm ally fo u r d igit s ( 1 1 t o v ) o n each foo t a ll ,

with t hree p ha lan ges an d a ll f u rn is h ed with c laws Bes id es th is

,
.
,

I is r epresent ed on t he fo re fo o t by a po llex ( th u m b) o f t wo

phalanges an d a n on retra ct ile c law while o n t he h ind foo t the

,
-
,

hallu x ( great toe) is r ud im entary co n s is t ing o f a s m all bo n e ,

a rt ic u lat in g with t he c u n eifo rm b u t bear ing n o c la w Of all .

an im a ls as id e fro m m an t he ca t is t he m o s t s u bj ect t o s u per n u

, ,

m erary d igit s es pec ially o n t he fo re fo o t I n t he gr ea t m aj o r ity of

,
.

ca ses the d o u bling is in t he regio n o f d igit 1 Oft en t he extra mem .

b er is s haped n o t lik e it s n eighbo r s b u t rath er as if belo ngi ng

, ,

to t he o ppos it e foo t th o u gh som et im es it is ind ifferent

,
Fo r .

ex ha u s t iv e m at erial o n th is s u bj ect s ee Ba t eso n Mat er ial s for t he

—
, ,

S t u dy o f V ariat o n pp 3 3 3 2 4
i 1, . .

S peak in g gen erally t he d og tr ibe has fi v e t oes in fro nt (d ig it

,
1

1 n o t t o u ch in g t he g rou nd ) and fou r beh ind ( I ab s ent ) .

The sea l ha s fi v e d igit s o n all extrem it ies th o u gh t he ha n d is ,

m od ifi ed int o t he fl ipper and t he foo t is b u t s light ly f u nct io nal

,

and ev id ent ly well on t he road to ext inct io n .

T he whale gen erally has fiv e d igit s in fro nt in c as ed in s k in

to fo rm a fl ipper th o u gh t his n u m ber is o ft en r ed u c ed t o fo u r
, ,

and in all cas es 1 1 and 1 1 1 hav e m o re than the u su al n u m ber of

j o int s The o n ly trac es o f a h ind lim b are a few s mall bo nes
.

benea th the sacral regio n and occasio nally a part o f a lim b 2

.

I n t he m anat ee and t he d u go ng t he pr oc es s has go n e fa rt h er .

T h o u gh th es e aq u at ic m am m a ls ha v e exc eed ingly serv ic eable

fl ippers with fiv e d igit s yet t he h ind leg has been ent irely los t
,
.

T he v ert ebra in t he s acral regio n are n ot u n it ed and ev en t he ,

pelv is is represent ed on ly by a pa ir of s plint bon es th ou gh som e ,

fo s sil fo r ms s how a ru d im ent ary femu r or th igh bo n e 3

.

1 E xc ep tin g t he A fric an h u n tin g d o g, w ic h h has fo u r ( y

L d ek k er, L ib ra ry of

N a tura l His to ry, p .

Th is is sim i la r to t he l wi n gs in t he cas e o f t h e N ew Z ealan

o ss o f d a p t eryx.

L yd ekk er, l
L ib ra ry o f N a t u ra H is t o ry p 1 1 5 6 , . .
 8 V A R I AT I O N
5

T he bea r ha s fiv e t oes all r o u nd with a n add it io nal c la w in

, ,

d ig it I I beh ind wh ich he u ses fo r c o m b ing

,
.

I n m ic e and rat s d ig it 1 in fro nt is r u d im entary T h is ca se is .

u n iq u e b ec a u s e in m o s t in s t a nc es wh er e a d ifferenc e is n o t ic e ,

able t he red u ct io n in d igit s ha s proc eed ed farth er beh ind than

,

befo re .

S nak es es pec ia lly t he large o n es o ccas io na lly s h o w ext erna l

, ,

v es t iges o f h ind legs and int ernally ar e freq u ent ly fo u nd trac es

,

n o t o n ly Of t he pelv is b u t lik ewis e o f t he th igh bo n e o r f em u r

1
.

Th is s h o ws c lea rly tha t t he s nak e is a so m ewh a t rec ent fo rm

d ev elo ped fr om lizard lik e an c es t o rs with lim bs t he h ind pa ir of
-
,

wh ich m u s t h a v e b een p lac ed n o t far fro m the m idd le po int o f

t he m u ch elo ngat ed b ody
- T h is V iew is s trength en ed by t he fact
.

that as a ru le b u t o n e lu ng is d ev elo ped s h o wing that t he body is ,

m o re s lend er th a n fo rm erly .

Of a ll s t ud ies in d ig it s t he m o s t int eres t ing is that o f t he

u ngu lat es o r h o o fed an im a ls I t is a ls o t he m o s t pro fitab le.
,

beca u s e t h e m aj o rity o f o u r v a lua ble d omes t icat ed an im als are

inc lu d ed in th is c la s s ifica t io n .

T he int eres t ar is es fro m t he fa ct th a t o u t o f th is s t o ck ha v e

d ev elo ped t wo v ery d ifferent fo rm s o f feet V iz t he t wo t o ed (as ,
.
-

catt le) and t he o n e t oed ( as h o rs es ) b o th ev id ent ly hav ing

-
,

d es c end ed from fi v e t oed a nc es t o r s each by a proc es s o f it s o wn

-
, .

F o r exam p le cat t le s h eep d eer p ig s et c ha v e t wo t o es ( I I I

, , , , ,
.
,

and I v ) well d ev elo ped int o a s er v ic eable foo t with t wo o th er s ,

( 1 1 and v
) s t a nd in g beh ind n o t t o u ch in g t he g ro u nd
, ( p ig s ) O ft e n ,

r ud im entary (d eer ) a n d freq uent ly repres ent ed m erely by s plint s

,

( catt le) . O cc a s io nally a ll trac e o f th es e d ig it s is lo s t

( gi ra ffe
) .

On t he o th er hand t he h o r s e and his k ind h a v e b u t a s ing le

,

to e b u t o n eith er s id e is a well d ev elo ped s plint t he r e -

,

m a in s o f t he s ec o nd a n d fo u rth m etacarpals in fro nt and o f t he

c o rres po nd ing m etat a r s als beh ind Th ey are h owev er with o u t .
, ,

fu nct io n al s ig n ificanc e b e ing a ttach ed o n ly abo v e and ext end

,

in g d o wn w rd with s lend er s h a ft s a n d fr ee end s n o t s u pplied

a

with d ig it s .

I n th is co nn ect io n it is t o b e n o t ed th a t t he ext inct pr o t o h ipp u s

Of t he U n it ed S t a t es and t he h ippar io n o f E u r o p e b o th d ec id ed ly ,

1
L yd kk L ib
e y f N t l H i t y p 5 35
er , ra r o a u ra s or , . 2 .
 M ER I STI C V A R IAT I O N
59

hor selike an im als an d rega rd ed as ances t or s of the m od ern ho r se ,

had each thr ee t oes that pro bably rea ch ed v ery n ear t he gr ou nd .

Pas s ing s t ill f u rth er ba ck (d o wn ) in geologic t im e and loo k ing

for a s t ill mor e remo t e anc es t o r we get beyo nd wha t ca n b e ,

called a tru e h o r se as can the prot oh ippu s and t he h ippar io n, .

B u t yet th ere is am o ng th es e lo ng e xt inct fo rm s s u ffic ient h or se -

lik e c haract er t o s u gges t anc es try as with t he fo res t h o r s e and ,

d es ert h o r se o f t he Wh itn ey find in Wy om ing fo rt y inch es h igh ,

and t hr ee t oes d own A s we progres s in th is d irect io n h o wev er

1
.
, ,

t he t oes increas e in n u m ber t o fo u r and ev en fi v e c lear ly ind i ,

ca t in g that the m od ern h o rse has d ev elo ped fro m a fiv e t oed an ec s -

to r l ik e t he t ippu s t welv e t o s ixt een inch es h igh and a ll t o es

,

d own a lso d iscov ered in Wy om in g wh er e it fl o u r is h ed acc ord ing

, ,

to O s bo rn so m e thr ee m illio n y ea r s ago o r th er eabo u t s

,
11
.

I f we begin with t he m od ern t wo t oed s pec ies and att em pt to -

read th eir s t o ry back ward s we soo n land am o n g t he sam e fou r ,

o r fi v é t oed prim it iv e f o r m s j u s t m ent io n ed fo rc ing t he c o nc lu

-
,

s io n th a t t he On e t oed and t he t wo t oed s pec ies of recent t im es

- -

hav e each d es cend ed from fi v e t oed progen it o rs ind eed we may -

, ,

ev en b eliev e fr o m t he s a m e fi v e t oed pro gen it o r s -

.

T he mann er of th is d esc ent is n o t d iffi c u lt t o trac e by t he

com pa r iso n of m od ern s pec ies with s im ilar ext inct fo rm s in su c
c ess iv e d o wn wa rd back w ard ) g l ic t im I a lm t h
( eo o g es n o s t e .

lowes t t ert ia ry rock s o f bo th N o rth A m er ica and E u ro pe o cc u r

fos s il rem ain s of large u ng u lat es T h ese Co ryph od o n s were
“
.
”

s u pp lied with fiv e t oed f eet m u ch lik e t he elephant o f t o d ay

-
,

that has su rv iv ed by v irt u e o f his t eeth and in s pit e o f his feet .

A s c end ing to t he M io c en e T ert iary we find large u ng u la t es ,

s t ill r em a in ing b u t d ig it 1 is go n e wh ile t he m etacarpa l ( o r m eta

, ,

tar sal) has bec om e m u ch length en ed and the th ird and fo u rth
m em ber s great ly s trengt h en ed n o t o n ly in th eir o wn d ev elo pm ent ,

1 H en ry F Os b o rn O r ig in
.
, an d H is t o r y of the H o rs e .

3 T he l d th h
ev e o p m en t o f
f m t e o rs e ro a n a n c es o r o n l y tw l e v e t o s ix t een in c h es

h igh d wit h fi t
an ll d w i th bve t i o es , a o n, s e es n s tanc e o f p g
r o r es s iv e ev o u t io n l
o f wh i h w h
c y ke wl dg D b tl
a v e an th no e e
. ou es s l
e ev o u t io n o f o t h er s p ec ies ha s
b l
een n o t d d d f i ti g b t f
es s ex en e an a sc n a n , u o no o th er c a s e d o we p o s s es s s o
co m pl t h i t y t h k f wh i h
e e a s or i l g, an s or c a re n ar e m ea s u re d u e to t h e g en ero s i ty
o f t h l t W ill i m C W h i t y
e a e d t th l b
a . ne an o e a o rs o f Pro fes s o r H en ry F Os b o rn . .

S l h p
ee a s o c t ii
a . x , s ec . .
60 V A R I ATI O N

b ut regard s th eir art ic u lat io n wi th t he small bo n es abov e

a ls o a s .

T h is foo t is n o w o n the r oad to beco m ing ind iff erent ly eith er a

t wo t oed o r a on e t o ed fo rm d ep end ing u po n wh eth er I I and v
- -
,

r ed u ce t ogeth er or wh eth er 1 1 1 tak es t he lead .

I n th is c o nn ect io n c erta in int er m ed iat e or s trand ed fo rm s are

o f n o litt le int er es t For exa m p le the elephant has fi v e t oes in
.
,

fro nt with fo u r and somet im es three beh ind T he rh inoc eros has
,
.

three bo th befo re and beh ind b u t the ext inct fo rm oft en had ,

fou r The tap ir wh ich is also rega rd ed as a rem nant o f anc ien t
.
,

life pr eser v ed u nt il t he pres ent has gen erally three t oes th o u gh , ,

s o m et im es f o u r and occas io n a lly t wo I n any case h o wev er d igi t .

, ,

I I I is larg es t and s y m m etr ica l in it s elf s h o wing a ffi n ity with t he ,

lin e o f d es c ent t hat has d ev elo ped t he s ingle t oed fo rm s -

.

T he ca m el ha s t wo t oes wh ile t he n early relat ed ch ev ro ta in ,

has fo u r t wo being red u c ed

,
T he h ippo po ta mu s ha s f o u r s h o rt
.

to es a ll d o wn all h oo fed and part ly webbed s h o wing a ffi n ity

, , ,

with t wo t o ed fo rm s in that the s y m met ry is ab ou t a lin e d ra wn

'
-

bet ween d igit s I I I and I V T h is is the s a me plan as that of the . .

pg i e
,
xc e pt that i n t h e latt e r t h e f o o t i s m o re c o ntr a ct e d t h e ,

t oes being flatt ened o n the in s id e and t he s ec o nd pair n o t to u ch

in g t he gro u nd .

T he kangaro o is an o m a lo u s hav ing fi v e t o es in fro n t and in ,

general f o u r beh ind of wh ich I v is m u ch the larges t ; v is s mall , ,

and I I and I I I a re m u ch red u c ed and inca sed in a c o mmo n

i nt eg u m ent .

With th is brief s u rv ey o f s pec ific d ifferenc es in res pect to

d igits c erta in ind iv id u a l d ev iat io n s will hav e an add ed m ean in g
, .

B at es o n g iv es us t he f o llo wing 1

1 .H h i g pe m y t
o r se av n su i id e f igh t f
rn u er a r f t p e m
o e on ns o r o re oo , r su

a b ly d ig i t I I It ti . l t d wit h ar cu a et b o e i th l w an ex ra w f the n n e o er ro o

c ar u s a n
p d w p id d
as w i th hro v f e e b oath idoo , e m b l i
c on v g th e x s es , res n

h f f
oo o
( e
a n as sF ig se .

2 .F l h i g tw toe
oa av n e ch f o f t o t h e wi e
s on a m al
o re T he c
oo , r s nor . ar

p us w i t h
as i c as e
n m s
a l b t th t n or t e , w ue e g i b
e ex ra e t h o
e s r a a n o rn on

i id e
ns d w e p an id ed with
er ro vm ll h f a s a oo .

3 H . h a io rs e
g d i m
v n t y d i
a ru
g i t i i d e
en a r f l f t h i d f t
on T h
nsi o e n oo . s

a g i
a n res u l t f m li
sg h t d
ro l p m
a s t f d i g i t I I
ev e ow h i c h i t
en he m o t om , s os c

m o n c aus e f p l yd tyli m m g h
o o ac e s a on o rs s .

1 F r i ti
o i th f t f t h h
v ar a on n e B t
ee o eM t i l t
o rs e, s ee
pp 36 3 7a eso n , a er a s , e c .
, . 0- 2.
62 V A R I AT I O N

6 . h righ t fo e f t b ea i g tw c o mplete d igit s y mm et ic a lly

F o a l wi t r oo r n o s r

d e elo p ed ea c h b e ri g w ll f m ed h
v , f th t a e fl tte ed O the i
a n e e -
or oo s a r a n n nn r

sid es a n d c e t w d ea c h th e li k e t h
u rv e o f th e
o t io d c tyle ( c t t le
ar o r os ar a s a ,

T h i i ll t a te c o d it i s
( 3) us r s n on

j ust m en t io n ed ( s ee F ig .

C att le h eep and p igs a f ,

s ,

fo rd d ev iat io n s n o les s int er

l
eS t1 n g
F IG . 1
3 . Fo o t of h o rs e : d igi t III sup

I . C a lf ha v i ng t h ree d i g i ts on
p r es s e d d igit
, s I I an d I v d l
e v e o pe d .

A ft er Ba t es o n
rig ht fo r e foo t, b o r n e on a s in
g le

c o m mo n b on e a fte r th e fa s h io n of the b i rd s and fu l l y

s
ym m e tri c a l
( s ee F i
g .

2 Hei fer a v i n g t
. h h ree fu ll y d ev elo ped to es on

ea c h hi nd li m b . I n t ish c as e th e s u p er n u m era r y was

c lea rly d igi t II .

3 . C a l f wi t h “ s
u per n u m era r y to e p la c ed b e t ween
the d ig i ts O f th e r ig ht m an u s ( fo re fo o t
) . h
T is to e
ha d a h o of an d s eem ed ex

tern a lly to b e per fec t, b u t o n

d is s ec t io n i t was fo u n d to
c o n t a in no o s s i fi c a t io n , b ut
was en ti rel y c o mp o s ed of

fi b ro u s tiss u e a nd fa t
”
.
11

4 . C o w, fu ll-
g ro wn , r ig t h
fo re foo t wi t h
fo u r d ig i ts
a r ra n ged in two g ro u p s o f
two ea c h ( s ee F ig .

h T is is c lea r l y a c ase in
wh i h c the i n c rea s e is d u e,
no t to the r ea p pea r a n c e o f

a n a n c ien t los t toe li ke I t or

v, b ut ra t he r to the d ou b li ng
(
f t/i e n or m a l d igvt r I l l a nd

1 V 111 r o ug lz or d i n a ry m er i stic
1
1
v a r ia tion .

5 . C a l f, left hi nd foo t
wi th fiv e to es , “
an i n n er
FIG . I 4 . h
R ig t f o re fO O t Of ca lf : t h ree digit s
g ro u p of two to es c u r v i n g
p res en t, ea c h su
pp l i d e wit h bot h fl ex o r an d
to ward ea c h ot he r, and an
ex t en s o r t en d ons . A ft er B a t es o n
o u ter r Of h
t ree, of
g o u p
1 Ba t es o n , M a t eria l s , etc .
, pp 3 7 3
.
-
390 .
2 Ib i d .
p 377. .

3 No c as e is b et ter t h an h
t is t o su
gg est c a u t io n to th e s t u d en t o f ev o u t io n . l
W h en a n ex t ra t o e a p p ea rs a m o n g t h o se fo rm s w h o s e a n c es t o rs were k no wn to
 M E R I ST I C V A R I AT I O N 63

wh ic h the m id d le o n e was a lm o s t b il a tera lly s

ym m et r i c a l, w ile th e h h o o fs
o f th e o t h er two t u r n ed to wa r d i t
1 ”
.

B at eso n ay s of the p ig that he kn o ws o f n o cas e o f po lyd ac

s

tylis m in th e h ind feet A ll cas es d es crib ed a re o f t he f o re feet .

,

and t he extra t oes a re o n t he int ern al s id e

o f t h e d igi tal s eries .

S y n dact ylis m in ca tt l e, s h eep , an d i

p gs .

By t h is ter m is meant a real u n io n o f

d igi t s I I and 1 1 1 int o a s in gle b o ne inca sed
i n a s i ngle h oo f as i n t he s o lid h o o f ed h o g s
,
-
.

A cc o rd i ng t o R os enberg a s q u o t ed by ,

B at es o n i n t he n o r m a l s h eep t he m eta
2
,

carpa ls I I and V are d is t inct in the

em bry o n ic s tat e aft erward s c o m plet ely ,

u n i t i ng
( f u s ing ) with 1 1 1 and I V
3 T h is .
"

thro ws s o m e light u po n the wh o le q u es

t io n a s t end ing to expla in n o t o n ly c erta in
,

cas es o f p o lyd a cty lis m b u t all c as es o f

s n d a c t lls m
y Y
1
A ga in q u o t i n g B
‘

a t es o n
dt h
.

F IG . 1 5 . Sai b e t h e rig t
o

I A yo u n g o x a v in g th e two d ig i ts o f the h 10 1 3 10 0 1 0 1 C° W = ( 118 11 8 111

h
.

r igh t fo re fo o t c o m pletel u n i ted y .

“NO 8 1 0 11 1 5 O f N o ea c
1 .

1 B a t es o n ( f m
B 1p?
“ ro
2 C a l f : ea c
. fo o t a v in g o n l
y h
o ne o o f,
h h
h a ll the b o n es were n o rm a l
8 anq u e )
th o u g .

3 . S a me as a b o v e, ex c ep t t h at in t he fo re fo o t th e n o rma l d igits ( I I I a n d
Iv ) were c o m p letely u n i ted , b ea ri n g a s i n gle h oo f . T he s a m e c o n d i tio n

wa s fo u n d b eh in d , ex c ep t t hat the h oo f was m o re po in ted .

4 . A fo r e foo t and a hi nd foo t of th e s a me i n d iv id u a l i

( p g) , i n w i c h h
the t wo c h ief d igi ts we re c o m p letel y u n ited , V iz . rep res en ted b y a s i n g le

s er ies Of b o n es .

ss ess a
g rea t er n u m b er o f d igit s, h b it l with m y t g d it
i t is a ua an re a r

—t h
o o
p as a c as e

Of a t a v is m , e rea p p ea ra n c e o f al g l t h t B t h w i it i th
on -
os c a rac e r . u o s n e c as e

of m an w h en a s ix t h o r e v en s ev en t h d ig it p p Th i m t b m i tia iea rs ? s us e er s c v ar a

t io n a n d n o t a t a v is m , b ec a u s e n o i t d p i s x f -

yoe t h b
s ec es o an so r as ev e r een

d d
es c r ib e o r i ts ex is t en c e s u s p ec e t d M i ti
. i ti th f
er s i t l im it d
c v ar a on, er e o r e, s no e

tol t h
os c a ra c t ers o r t o n u m b e rs o n c e n o r m l b t m yg f i f it h
a , u a o ar n ex c es s o e er .

H th
e re, en , is n ee d f di or s c ri m in a t io n , fo r ev en t he ap p ea ra n c e o f a c h a ra c t er t h a t

h as b een o nc e l o st is n o t a b so u t e ev i l d en c e o f a t a v is m .

1 B a t es o n , Ma te ria l s , et c .
p 3
. 8 1 .
1 Ib i d p 3 3
. 8 . .

3 Th e fo rm er fro m fa i in g ev er t o l u n ite , th e l a t t er f ro m a c o n t in u a t io n o f the

f u s in g p ro c ess to i n c u e 1 1 1 a n d I V l d .

4 T h at is , I t u n it es wit h 1 1 1 , a n d v u n it es wi t h Iv d u ri n g d l
e v e o p m en t .
64 V A R IATI O N

Oth er ar ca s es a re g iv en tho u gh the latt er is t he

and s im il ,

onl o n e d es c ri bed in wh ich t he s yndacty lis m is c o m p let e in a ll

y
fo u r feet I t is h o wev er gen era lly s im u ltaneo u s in fo re and
.
, ,

h ind f eet 1
.

A b sen ce of par t s in a lin ear ser ies M en with hand s bu t n o .

ar m s with feet b u t n o legs a re n o t u nkn o wn Wh eth er t he

, ,
.

m is s ing par t s a re rea lly dr o pp ed o u t o f t he s e ries o r wh eth er ,

th ey were o rig inally pres ent b u t

s u ffered a b o rt io n d u ri ng e m bry

o n ie d ev elo p m ent being rep re ,

s ent ed at m at u ri ty by r u d i m entary

part s is u nc erta in tho u gh zo Olo , ,

gis t s wo u ld inc lin e s tro ng ly to t he

latt er V iew T he exact fact wo u ld .

hav e an im po rtant bearing u po n

t he u n i t o f v ar iab ility t he nat u re ,

o f h er ed ity a n d t he m ann er o f ,

d i ff erent iat io n .

Wh a t ev er t he fact in th is re
gard v a ria t io n s o f th is o rd er are ,

m a n ifes tly ra re a s c o m p a red wi th

t he increa s e o r d ecrea s e in s trict ly

m u lt ip le part s I n o th e r wo rd s .
,

wh ile c n s id era b le d ev iat io n in t he o

F 6 M i ti i i i l
IG . 1 .
p t t
er s c reg n
e
u m b er Of s i m ilar m em be rs
on n e :
( as
h
ri
g t l g f b t
e l op t dee e re ea e In

— fi 5 c o m mo n d
t ip l i
r t A ftca e B t
. er
n g
a es o n
e rs
) 1 1 1 1 S ex c ee ,
“

i n gly u nc o m m o n fo r an ent i re
gro u p (as t he hand ) to b e o m itted rarely fro m t he en d o f ,

t he s e ries ( as t he f o o t o r hand ) and s t ill m o r e rarely if ev er , ,

fro m t he m idd le o f the series as wo u ld b e t he cas e in a tr u ly ,

m is s ing a rm ( h u m er u s rad iu s and u lna ) b u t with t he hand , , ,

present c o m ing d irect ly fro m t he bod y

,
.

Ex t ra l egs T he repet it io n o f a m em b er a s c o m p licat ed a s a

.

leg is extrem ely u n u s u al b u t by n o m ean s u nkn o wn T h e wr it er .

B t1
g i
a es o nm y im i l
v es an s h w i t h m
a r c a s es ,p l i i t y
ea c f i t w so e ec u ar o s o n .

S l id h
o f d p ig
-
oo e s f q tl y
a re s een d t p i t
so re wid l y
u en m
an d b th
a o n s so e re ove o

i t im
n d p e an ( m t is d b
ac e
y A i t
en t l o ne d p t d f m
r s o m y e ang i f
re t h
or e ro an re ons o e

ea rth) ( B t s ee M t i l a es o n , t p 3 )8 7 t h t
a er a s , e c t h i w . l d
, . m t b a i t i s ou s ee o e a var a on

th t h
a ft i
as o f h
en a r s e n a res .
 M ER ISTIC VA R I ATION 65

saw o n e s ep c im en o f a leglik e appendage gro wing fro m the left

sid e o f t he n eck o f a ca lf n ear t he po i nt o f t he s h o u ld er The .

leg was n o t m o re than t wo th i rd s t he u s u a l length and wa s ,

t wis ted and f u nct io nless tho u gh it t erm inat ed with a h oo flik e ,

gro wt h 1
.

E xt ra leg s a re c o m mo n in in s ect s s o met im es thro u gh o u t th eir ,

ent ire length s o met im es d o u b ling at t he f em u r s ee F ig

, ( .

SECT I ON III — M E R I STI C V A R I AT I O N A N D B I LAT E RA L

SY MM ET R Y

Mer is t ic ariat io n a m o ng pa ired o rga n s and th o s e s tand ing

v

s ingly o n t he m ed ian lin e thro ws n o litt le light u p o n t he nat u r e

of b ilat era l s y m m etry and a ls o inc id enta lly u p o n t he m anner o f

v a riat io n .

S pe ak ing gen era lly pa ired o rgan s m ay d o u b le o n eith er s id e

,

s eparat ely o r o n bo th s id es d igi t s leg s wing s th y m

( o r e a
y , , ,

u n i t e i nt o a s ingle o rgan wi th it s ax is o n t he m ed ian lin e (h o rs e

s h oe kidn ey ) .

M o s t Of the exam p les o f mer is t ic v ariat io n already g iv en a re

o f r e pe t i t io n in pa ired o rgan s i n b ilat era l s y m m etry I t rem a in s .

t o ca ll att ent io n t o t he o ppo s it e c o nd it io n t he f u s io n o f a p a ir ,

into a s i ngle o rga n s tand i ng o n t he m ed ian lin e

I .A good e a mpl f th i i t h a t f
x eb k h i g the h
e o s m s o a ro uc av n o rn s co

po un d de f f ll y
orh lf t h ei l ug t h i t a i g le b
a e m t dri g t h
en n o s n
“
a s an n on e

m id d le lin e 3
( s ee F ig .

2 . A y ho
b ee a v in
ne
g th e t wo c o m1’
p ohu n d ey es u n i ted in to o n e at the to p
o f th e head wi t no
g h
r oo v e o r lin e o f d iv is io n b etween t em
‘
h .

P i r en d s o f k id n e u n i ted i n m a n , fo rm i n g a y h h oe k id ey
3 . os ter o
( ) o rs es n

wi th h ree
t ren al art er ies on ea c h s id e . T is h c as e is in s h a rp c o tra t to
n s

Ba tes o n N o 4 0 7 , wit a s in gle la rge

’
s . h k id n ey o n the left a n d two s m aller,
on e b elo w the o t er, o n the rig t h h .

1 T ish s p ec im en is d esc rib e d fro m m em o r y , as it was s een b efo re t h es e p h e

h o m en a were ma t t ers o f p ers o n a in t eres t l .

2 Ba tes o n Ma t eria ls etc , p 4 60

, , . . .

3 Ib i d
p 4 6
. 1 . .

4 Th e c o m o n in
p u g o f e e d
s has a rea b ee n m ey
n t ione I t a ppa reln t dyo c c u rs d . ly
on ly d
in in s ec ts , b u t is a go o exa m p le o f t he ev e o p m en t o f ig iff eren t ia te d l h hly d d
l
t iss u e in ab n o rm a s it u a t io n s , i u s tra tin g n o t o n ll
m eris t ic v a ri a t io n b u t f u n c t io n al ly
v a ria t io n as we ll .
66 V A R I AT I O N

C o n v ers ely npa ired o rgan s s tand ing o n the med ian lin e m ay
,
u

b e d iv id ed s o a s t o fo r m a p a i r o f o rg a n s s y m m et ric a lly p lac ed .

I t s h o u ld b e n o t ed th a t in genera l a s ing le o rgan s tand ing o n

t he m ed ia n lin e a s t he n o s e is s y mm et rica l b o th with refer enc e
, ,

to i t s elf and to t he m ed ian line b u t ,

that fo r t he m o s t pa rt pa ired o rga n s ,

th o u gh s y m m etr ic a l with ref erenc e t o

t he m ed ian lin e a r e n ot tlzem s el v es ,

n ec es s a r i ly sy m m et r ica l b od i es
( ear s

ar m s b a n d s ) I n o th er cas es o f pa ired ,
.

o rga n s h o wev er (eyes k idn ey s ) t he , , ,

m em b ers d o ha v e a k ind o f s y m m etry

o f th eir o wn .

A ga in n o th ing is m o re c o m m o n , ,

es p ec ia lly a m o ng p la nt s than t o find ,

a s ingle o rga n o n t he m ed ian lin e

appea ring a s a pa ired o rga n in c er
ta in in d iv id u a ls o r in n ear ly relat ed
s p ec ies o r v ar iet ies .

Ba t eso n g iv es a s e xa m p les o f t he
la s t t he p o s t er io r pet al in Ver on ica ,

wh ich in m o s t rela t ed s pec ies appears

F
a s a p a i r o f p eta ls ly ing o n eith er s id e
1G 7
1 °

org t
an s :h t w h fet h i o o rn s o s
O f t he m 1 d d 11 11 n c
ro eb kuc i di t i gl
a re u n t e n o a s n e
' °

b m f
ea or id bl
a di c o nsA ft er g iv ing n u m ero u s in s tanc es o f
e ra e s

b " f1 m d 1h y p
t a n c e’ ‘ a e
d iv is io n o f m ed ian o rga n s in fi s h es
a' 3 e se '

amt e A ft B t er a es o n
a n d i n in s ect s h e c i t es a u th o r ity fo r
'

s ay ing th a t T he o rgan s m o s t o ft en d iv id ed in m a n a re t he
“

s t ern u m n eu ra l arch es u t eru s p en is et c

,
and o f th ese s pec i
, , ,
.
, ,

m en s m ay b e s een in any p a th o lo g ica l m u s eu m 1

O rgan s m o re .

rarely d iv id ed a re the t o n g ue ep iglo tt is u v u la a n d c entral n eu ra l , , ,

canal 2
T h es e la tt er a re in rea lity ca s es o f a x ial d u p lic ity 3
"
.

1 B a t es o n , M a ter ia l s , et c .
, pp 4 5 0 4 5.8 -
.

2 l gy i t h t b h f b i l gy wh i h t t f b m liti
T era t o o s a ran c d it
o o o c rea s o a no r a es , an

a ff d m
or s y an f m i ti
c as e s Thio t dy h
e x t re b id
e v ar a d o ns . s s u as een cons ere as

i
c ur o us th th
ra p erfi t b l d y
an h b
ro m lait i m
e, a n i g t b et , a s suc a no r a es a re c o n o e

re gar d d e f qas t l y d ret d f i

u en g m i t m yy pt
ue t
o ah t t i e ec t v e
t er , a e ro v e a a t t en on o

c as es o f th i d m y f i h t h k y t t h l t i f q ti i l i g th
s or er a u rn s e e o e so u on O u es ons n vo v n e

un it fo i b il i y
v ar a t . B M t i l t p p 5 59 566
11 a t es o n , a er a s , e c ., . .
F IG . 18 . D o ub l h dd
e- ea e l
t u rt e c o m p a re d wit h t he u s u a l S p ec im e n s t wo t o t h r ee

d y
a s o ld . N o te effec t o n s h ll p l
e a t es . h
I n t is s p ec i m en the m o v e m en t s of

t h e legs o n o p po s i t e s i es d were no t we ll coordi n a ted . A fter B a t es o n

68 V A R I ATI O N

F ig . 1 h o ws a case of d o u ble h ead in the t u rt le Many

8 s .

s im ila r in s t a nc es ha v e b een d es cr ib ed b u t th is is es p ec ia lly i nt er ,

es t ing b ec a u s e the t wo h ead s s ee med to act ind epen d ent ly and ,

it is sa id th ere was n o c o nc erted a ct io n bet ween t he f eet o f the

two s id es T he s a m e ph en o m ena o f d o u b le m o n s ters a re s ai d
.

to b e freq u ent ly n o t ed in fi s h hatch ing es tab lis h m ent s A m o ng

-
.

s n a k es s o m e t wenty ca s es a re rec o rd ed o f c o m p let e o r part ia l

d u plic ity n ea rly alway s Of the h ead S ev era l o f th ese were a n i

,
.

m a ls o f go o d s ize and m u s t h a v e led an ind ep end ent ex is tenc e

,

fo r s o m e c o n s id era b le t im e 1
.

S im ilar ca s es o f d o u b ling a re kn o wn in b ird s and ev en in m a m

m a ls b u t a m o ng th es e h igh er a n im a ls t he pract ica l d i ffi c u lt ies
,

in s u s tai n ing ex is t enc e with extrem e abn o r m al ity a re v ery great ,

and th ey c o m m o n ly d o n o t lo ng s u rv i v e .

B et ween th is d iv is io n o f a s ing le o rgan ly ing o n t he m ed ian

lin e and t he d o u b ling o f s o im p o rtant a p a rt as t he h ead th ere ,

s eem s t o b e n o c lear li n e o f d em arc a t io n Th is d o u b lin g may .

ev en g o f u rth er a s in t he ca s e o f the S ia m es e t wi n s u nt il t he
, ,

s p ec im en is r egard ed a s es s ent ia lly t wo i nd iv id u al s u n i t ed by

s o m e s o rt o f attach m ent .

S ECT I ON IV — SYMM ET R Y I N V A R I A B LE PA R T S

With o u t a d o u bt m eris t ic v a riat io n in o ne o rga n o f the body

is lik ely b u t n o t c erta in to b e acc o m p a n ied by abn o r m a lity in
, ,

ano th er F o r exa m ple a v a ria t io n a mo ng t he d ig it s o f the fo re

.
,

fo o t is lik ely t o b e a s s o c iat ed with a s im ilar v aria t io n beh ind s t ill ,

m o re lik ely o n t he o ppo s it e s id e b u t n o t p o s it iv ely with eith er ,

.

A gai n th ere is so m e s u gges t io n o f s y m m etry with in the part

,

it s elf in wh ich t he v a riat io n o cc u rs A goo d exa m p le Of th is is .

B a t es o n s N o 4 9 5 ( Fig
’
. .

T h is is a left hand an d the f o u r extra fi ngers s eem t o rep re

,

s ent no t t he th u m b o f th a t h a nd b u t the fing ers O f t he o pp o s it e

( r ight ) hand th u s s ee m
,
i ng t o a i m a t a k i nd o f s ec o ndary s
ym
m etry with in the m em b er 2
.

I n t he d es cript io n o f th is ca s e we a re t o ld that th is d o u ble

hand and ar m were v ery m u s c u lar s o that it wa s n o t po ss ible ,

B t1 M t i l
a es o n ,t p 5 6
a er a s , e c I b id p 33 5
.
, . 1 .
9
. . .
 V A R I AT I O N
7 0

in m a n . A gai n , wh ile
d igita l v ariat io n is exc eed ingly c o m mo n
in ch ick en s i t is ra re in b ird s genera lly a n d is a lm o s t u nkno wn
, ,

in d u ck s and gees e wh ich hav e lo n g b een d o m es t icat ed .

T he fa llacy ab o v e a llu d ed t o s eem s to ha v e aris en fro m t he

fact that d o m es t ic a ted s pec ies a re bett er kn o wn than wild o n es ,

and that c ertain v ariat io n s at leas t a re m o re lik ely to b e pres erv ed .

I n any ev ent th ey a re m o re s tro ngly im pres s ed u p o n o u r att en

t io n T he t ru th s eem s t o b e that v ariab ility d epend s u po n the
.

nat u re o f t he part and t he relat iv e s tability o f t he s pec ies in q u es

t io n n o t u p o n its d o m es t icat io n o r its plac e in t he s ca le o f life
,
.

W e can th erefo re a v a il o u rs elv es o f a n y m at eria l bearing u po n

t he general q u es t io n wh erev er it m ay b e f o u nd h o p ing h o wev er , , ,

fo r t he ear ly c o m ing o f t he t im e wh en t he v a riat io n s with in t he

part ic u lar fi eld o f t hrem m at o lo gy s hall b e bett er k n o wn and mo re
acc u rat ely d es crib ed .

A symmet rical dev el opm en t in symmet ri cal pa rt s The ca s e o f .

t he nar wha l illu s trates a fact in v ariat io n wh ich th o u gh s eld o m ,

s o apparen t is d o u bt les s o ft en p o t ent ia lly pres ent and if s o is

, , ,

c ertain ly to b e reck o n ed with by t he breed er .

I n th e na rwha l t he ca n in e t o o th ( in t he m a le o n ly ) d ev elo p s
.

a s a t u s k o ft en atta in ing a len gth o f s ev en o r eight f eet o r ha lf

, ,

t he length o f t he b o dy T he pec u lia rity is that n o r m a lly o n ly

.

t he lef t t u s k d ev elo p s and in t he few ca s es s een i n wh ich bo th

,

a re d ev elo p ed t he r ight t u s k is s p ira lly t wis t ed fr o m l o r i let

f t t g e ,

exact ly lik e th e left t u s k and n o t i n t he o pp o s it e d i rect io n as

, ,

we s h o u ld exp ect What is s t ill m o re as t o n is h ing is th a t n o ca s e

.

lzas b een d es c r ib ed i n rv lzic lz t lze I t t a s k wa s d ev elo ed

ev er
g
ri/ p
a lon e in s t ea d of t he left . E it her b o th a re d ev elo ped o r t he
left o ne o n ly and
,
in t he fo rmer ca s e th ey a re es s ent ially bo th
left t u sk s .

S ECT IO N V — M E R ISTI C V A R IAT I O N IN R A D I A L SE R I ES

E xc ept in t he lo wer fo rm s rad ial ser ies a re characteris t ic of

,

plant ra th er than o f an im a l life I n t he b ra n ch ing o f s t em s and

.

t he part s o f fl o wer s m em b e rs o f rad ial s e ries a re ev ery wh ere

,

abo u t u s T h eir v a riat io n s a re alway s int eres t ing ( d o u b lin g of

'

fl o wers ) and o ft en exc eed in gly v alu ab le (s t o o ling o f gra in ) .

 M E R I ST I C V A R I AT I O N 7 1

B o tan is t s wo u ld y that w hat sa s eem t o u s a s rad ial s eries ,

with t h e m em b er s s tand ing o n t he sa m e h o r izo ntal lev el a re in ,

mo s t cas es rea lly s h o rt en ed s t em s br ing ing th es e part s int o a ,

rela t io n wh ich is appar ent rath er than act u a l as wo u ld happen if ,

we c o u ld t eles c o pe any lo ng s t em u nt il t he lea v es r eg ular ly dis ,

po sed alo ng it s length s h o u ld co m e t o occ u py t he sam e plan e ,

1
.

I n th is V iew o f t he ca s e t he petals o f fl o wer s a n d t he bra nch

in g O f s t ems a s in t he s t o o ling o f gra in wo u ld b e exam p les o f
, ,

lin ear s er ies v ery m u ch s h o rt en ed ra th er than o f rad ia l s er ies ,

acc o rd ing to t he s tr ict es t d efi n it io n o f t he t er m For o u r p u rpo s es .

,

h o we v er th is s tru ct u ral po int m ay b e wa iv ed a n d all apparent

, ,

c as es o f rad ial s y m metry tr eat ed as act u a l .

O b s erv at io n s ind ic a t e and exper im ent s S h o w that m em ber s o f

s u c h s eries m a b e increa s ed in n u m b er a lm o s t ind efin it ely A l l
y .

t h e m em be r s m ay b e d o u b led s im u ltan eo u s ly ( a s fi v e peta ls

incr eas ed t o ten ) o r any o n e m em b er (o r ig inal s egm ent ) m ay
,

d ou b le or ev en tr ip le o r it may b e en t irely s u ppres sed with o u t ,

reference to o th er m em ber s Of t he s er ies .

Th e n a t u ra l m et hod o f d o u b ling s eem s t o b e fo r c ell d iv is ion

t o pr o c eed o n e s t ep bey o nd t he n o r m a l g iv ing r is e t o t wo in s t ead ,

of one I f th is occ u r s in a ll t he m em b er s ( p eta ls ) th en t he

.
,

m em ber s will a ll b e d o u b led a s ten in s t ead o f fi v e ; if o n ly in ,

part th en o n ly that po rt io n will b e a ffect ed m ak ing s ix s ev en

, , , ,

o r ev en eight in s t ead o f fi v e T h u s we ha v e c lo v er ru nn ing a ll

.

t he wa y fro m t h e n o r m a l thr ee u p t o a s h igh as s ev en leafl et s .

Ma n ifes t ly if c ell d iv is io n pr oc eed s t wo s tages bey o nd t he

n o rm al each o f t he t win p a ir aga in d iv id ing
,
it will res u lt in ,

1L ea v es a r e a r ra n
g d i g l d
e n re
p t h t
u ar or m f p l
er u t on d i g
e s e t s o a n s a cc o r n o

a y
s t ms e t i t t i
co ns g t h m u t h mn t i l e i a
fi 3 3
e a t
ca i w h
s er es , i h t h , , . e c .
, n c e

nu m t
era o r i d i at n th c mb es f i e nu it d th t m t
er o c rc u s h l f
a ro u n e s e o r ea c a ea

d i t ly
rec th o v er wi t h wh i h t h
e one t w t
c t d d th d
e co un mi t
as s ar e th , an e en o na or e

nu mb f l
er o th t wea v es ld b p d i a ou h e i it
as s e I t th n f s uc p a c rc u . er e o r e re re

t
s en s th mb e nu f m mb er o i wh l f e er sh t d tn a or f th i
o h t
a s o r en e s ern o s c a ra c er.

C o rn f , mpl o r ex a b l g w i
e, t h l l t h
e on m s, m b f t
a h o g f
er mi l y
e t er s o e ra s s a , o

th f e ti
r ac
4 on b i l t p
-
t h p
u l f
u t won T h ie m b
an o t h o. g h t t h
s nu er r u n s ro u ou e

p l an t d w
, h i
an l t h em b f w
e nu f er o t h ro b
s m O y y f
c o rn o n l y f m e co a v ar ree ro

e igh t t tw oty f en -
f
o ur , d d m b
n o ca se o
q / an w I o nu b erp t d ro s za s ev er een r e or e .

Thi f t ts d t t m l im i t t
ac en s o se so e w yw d s th i g
o ev en s o m i ti a i
ar a n as er s c v ar a

ti on, wh i h m c s ee t h s n ev erp d d o av e f ro w i t
uc eh d d
an ea r o m b f
c o rn an o nu er o

ro w s Thi
. m m s s ee l wh w s a rv e o us id th h
en i t w k wi t h d igi t
e cons er e av o c or s s

an d wit h e m pl i t d
v en s o c o t t ca e h d a s ru c u re a s a ea .
 V A R I AT I O N
7 2

q uadr u pling that m em ber o f the s eries I f h o wev er o nly on e o f .

, ,

th is pair s h o u ld d iv id e again we S h o u ld th en hav e on e plu s t wo , ,

o r thr ee n ew part s in p lac e o f t he o n e that wa s n o rm a l

,
A ga in .
,

if all fo u r s h o u ld s tart and o n e abo rt it wou ld lik ewis e res u lt ,

in three d ev elo ped m em bers in s t ead of fo u r that s h o u ld hav e

appeared .

A ll th es e v ario u s pr oc es s es may tak e plac e b u t what ev er t he ,

final res u lt a n d what ev er n u m ber u lt im at ely d ev elo ps t he m eth od

, ,

is that o f d ou b l i ng tli roug /z c ell d iv is ion giv ing r is e na t u ra lly t o ,

ev en n u m ber s Od d n u m ber s a re explainable h o wev er by s u p

.
, ,

po s ing that on e o f a pair c o nt in u es th e proc es s o n e s t ep farth er

than it s t win or else that o n e of t he m em ber s fa ils t o d ev elo p
, .

I n th ese wa y s an o rigin al m em ber of a rad ial series m ay at any

t im e d ev elop int o t wo three fo u r o r m o re an d if all t he m em
, , ,

ber s tak e part a tr u e d ou bling res u lt s

,
.

M eristic v ar ia t ion an d cell di v i sion I n t he las t analy s is th ere .

,

fore v ar iat io n in t he n u m ber of m em bers in a rad ial ser ies is

,

red u c ible to q u es t io n s o f c ell d iv is io n I nd eed we m ay go fu rth er .

,

and n ot e that all cas es of m eris t ic d ev iat io n ar is e in th is m ann er

that t he preserv at io n of t he n o r mal n u m ber of m u lt iple part s
d epend s u po n s u cc ess fu l c ell d iv is io n u p t o a c ertain ( n o rm al)
point and it s abr u pt c essat io n at that po int and that all so rt s of
abn or malit ies m ay arise thro ugh exc ess iv e m u lt iplicat ion thro u gh ,

abo rt ion or throu gh so me o th er d is t u rba nce of t he proc es s o f

,

c ell d iv is io n .

T h is V iew of t he ca s e h elp s to exp lain why it is that m eris t ic

v ar iat io n s in ra d ial s er ies are a m o ng t he ea s ies t t o exp lai n o f all

v ar iat io n s wh ich m a pres ent th em s elv es t o t he br eed er

y .

Con sid erat io n s o f th is chara ct er m ak e c lear the f u t ility and

s ho r t s igh t ed n c s s o f app ea ling to r ev er s io n o r atav is m t o expla in

what m ay b e a m ere inc id ent in c ell d iv is io n an inc id ent m o re , ,

ov er that m ay n ev er hav e o cc u rred in phylogeny m ay n ot b e

, ,

ev en c om m o n in o nt ogeny an d is th er efo r e n ot to u nd u ly im ,

pres s t he observ er 1
.

1 T he
s e t erm s wi ll b e fr eq u en t ly use d in t h e t ex t . P hy l o g en y r efers to the
d e l pm
ve o en t o f t h e S p ec ies , o n t o gen y to t he d l
ev e o p m en t of t he in d iv id u a l .

T he l t t a er is s u pp o s e d in a g en er a l w yt
a o r ep ea t t h e s t ep s o f t h e fo r m er , t h o ug h
wi t h t h i s v ie w o f t he m a t t er im p o r t a n t g a p s are o f freq u en t oc c u rren c e .
 M E R I STI C V A R I AT I ON 73

SEC TI ON V I — I M PORTA N C E OF M ER I ST I C V A R I ATI ON

N oth ing is of m o re
d irect ben efit to man than the st oo ling of
gr ain and t he d ou bling o f fl ower s is o f pr ime im po rtance to
,

s t ud ent s of t he bea u t if u l D igital v ariat io n and ind eed m os t o f

.
,

t he exam ples a m o ng an im als are n o t o n ly o f n o pract ica l u se b u t

,

th ey c o n s t it u t e d efo rm it ies that wo u ld at o nce b e elim inat ed from

t he fi eld s o f any in t elligent s t ock m an .

Th eir s t u dy is h owev er u sef u l t o the s t u d ent in t wo way s

, ,

fi rs t as s h owing him that frea k s a re by n o m ean s u nc omm o n

,

an d th erefo re n o t to b e s pec ia lly pr ized ; and sec o nd to S h ow ,

t he mann er in wh ich v a riat io n o perat es and the S ize o f t he u n it

in v olv ed t ogeth er with som eth ing of it s relat io n s to o th er and
,

s im ilar u n it s in t he sam e body The carefu l st u d ent will n o t

.
,

th erefor e was t e h is t im e in trying to establish a race of solid

,

h oo fed h ogs t he fir s t t im e a S pec imen of t he k ind t u rn s u p in

his yard b u t he will u t ilize t he info r m at io n affo rd ed in m eris t ic
,

v ariat io n g en erally t o ad v anc e his u n d ers fan d in g o f the m ann er

in wh ich v aria t io n beha v es and o f t he r elat io n s that o btain bet ween

t he s ev era l part s o f a h igh ly d ifferent iat ed bo dy .

The p u rpo se at th is t im e is to s ec u r e a m a s s of charact er is t ic

fact s o n wh ich fu t u re s t u d ies may b e based Mos t o f the danger s .

o f er ro n eo u s proc ed u re in th is field ar ise fr o m a pa u c ity o f well

a u th ent icat ed in s tanc es and fr om r es trict ed v iews of th eir real

S ig n ifi canc e .

S u mmary M er ist ic v ar iat io n refer s to d ev iat io n s in the p lan

.

or pa tt ern o n wh ich the o rgan is m is b u ilt I t s c entra l th o u ght is

.

sy m m etry S ym m etry m ay b e rad ia l with t he mem ber s id ent ical

.
,

o r it may b e b ilat era l with o ppo s it e m em ber s a s o pt ica l im ages ,

t he o n e o f the o th er D is t inct io n s o f right and left ar ise fr om

.

th ose of d or sal and v entra l and hav e reference t o t he relat io n of

,

t he ind iv id u a l t o the ou t sid e wo rld Organ s s y mm etrically plac ed

.

m ay or m ay n ot hav e a symm etry o f th eir own b u t t he ten ,

d en ey is for a pa rt to es tablis h so m e kind o f s ym m etry with in

it s own m em bers .

Wh en part s are mu lt iplied th ey m ay b e lik e the o th er m em

b ers o f the ser ies in wh ich th ey arise o r th ey m ay im itat e th o se ,

Of n eighbo rin r i h o m oeo sis R peat ed part s are espec ially

g se es
( ) e .
 V A R I AT I O N
74

su bj ect to m eris t ic v ariat ion The gen eral plan is pr eserv ed b u t

.
,

wid e v ar iat io n in t he d eta ils is c o mm o n a s in t he n er v e branch es ,

fro m t he s p inal c olu m n .

T he pa rt r epea t ed m ay b e S im ple lik e a d igit ; o r it may b e ,

a n ent ir e gr o u p a s a wh o le hand o r an ent ir e leg

,
.

M er is t ic v ariat io n ha s it s s ea t in c ell d iv isio n I t is of litt le .

u t ility in an im a ls th o u g h h igh ly u s ef u l in p la n t s b u t it s phe ,

n o m en a ar e v a lu a ble for t he in s ight th ey aff o rd int o t he m a n

n er o f v ar iat io n t he gen era l p er s is t enc e o f p lan and t he u n it o f

, ,

v ariab ility .

Ex erci ses L et t he s t u d ent giv e t en s epa rat e exam ples o f

.

m er is t ic v ariat ion n o t m ent io n ed in t he t ext an d d es cr ib e each

fu lly s t at ing a ll tha t is in v o lv ed of s ym m etry h o m oeosis etc
, , ,
.

A DDI TI ON AL REFEREN CES

V A R I AT I ON . A co c k wi t h n o s p u rs o n the leg , b u t wit h well d ev elo pe

-
d o n es

o n ei t h er id e o f the
s c omb . By E . S . D ex ter . S c ien c e, V I I , 1 36 .

V E G ET A B L E T E R A T O L OG Y . B y M a x well T . M as ters .
 CH A PTE R V
FU NCTI ONAL V AR IATI ON

By fu nct io nal v ariat io n is m eant a d ev iat io n n ot in fo rm o r in

,

t h e n u m ber o f part s b u t in t he f u nctio n s that th ey perfo rm T he .

liv ing a n im a l (or plant ) n o t o n ly i s s om eth ing b u t it d oes so m e

,

th ing and pla nt s and an im als d iffer amo ng th em selv es n o t o n ly

,

in what th ey a r e b u t in what th ey d o .

Ea ch po rt io n of a h igh ly d ifferent iat ed o rgan ism has its o wn

pec u lia r act iv ity wh ich is es sent ial ly d iff er ent from that o f any
,

o th er part of t he sa m e o rga n is m . Th es e act iv it ies are n ot c on

s tant b u t v ar iab le ; a n d ina s m u ch as m any an im als and n o t a

few p lant s a re k ept n o t fo r th eir appeara nc e b u t fo r what th ey

c an d o any d ev ia t io n s in th eir perfo r m a nc e ab ility a re o f prim e
,

im po rt anc e t o t he br eed er who is bent u po n th eir increa sed

,

effi c iency and th eir perm an ent im prov em ent for the serv ic e
of m an .

N ow plant s and an imals a re c o n s id ered as h igh or low in th eir

d ev elo pm ent acc ord ing to the d egr ee o f d ifferent iat io n or d iv is io n
o f labo r bet ween th eir d iff er ent part s . I n t he pro t ozo on t he f u nc
t io n s o f life are few and it s relat io n s to the en v iro n m ent a re
,

s im p le . A cco rd ingly it s act iv it ies are ex ert ed and its o b ligat io n s

t o life d is charged by t he c o m m o n m as s o f u nd iff er ent iat ed p ro
t o plas m perhaps with o u t so m u ch as a s t o m ach r epro d u ct io n
, ,

being eff ect ed by a d irect d iv is io n o f t he wh ole ma ss .

I n h igh er o rga n is m s ( m etazoan s ) h o wev er lif e is m o re c o m p lex

, ,

and t he res po n s ib ilit ies o f ex is ten c e are h eav ier T he se are m et

.

by s pec ia lized s tru ct u res s u ch as the m ou th to take food the

, ,

s t o m ach and int es ti n es t o d isso lv e and prepare it fo r u s e t he ,

liv er to c o n v ert c erta in po rt io n s into s pec ia lly u s ab le f o rm ,

t h e lu ngs t o ab s o rb air the b loo d v es sels to ca r ry it and the

,

d iges t ed foo d to all part s of the body wh ere each ext ract s what
,

it n eed s and can u se .

 6 V A R I ATI ON
7

T h en th ere a re o rga n s a s the k idn ey s wh os e f u nct io n is to , ,

rem o v e was t e prod u ct s that wo u ld o th erwise acc u m u lat e and

d es troy t he bo dy Th ere a re o th ers as t he u dd er an d v ario u s .
,

gland s wh o s e f u nct io n is to m an u fact u re s o m e part ic u lar pro d u ct

,

t o b e u sed eith er with in o r with o u t t he bo dy T h ere is a sy s t em .

he m u s c u lar ) fo r m o v ing t he b o dy as a wh o le o r fo r t he ex erc is e

( t
o f an o f i t s p a rt s and a n et wo rk o f n er v es f o r m ing a r eady and
y ,

rap id m ean s o f c o mm u n icat io n T h ere is a h eart t o driv e t he .

b lood and perhap s a bo ny s k elet o n to h o ld the c o m plicat ed m as s

,

t ogeth er .

N o w t he act iv it ies o r f u nct io n s o f th es e v ario u s part s a re by

n o m ean s c o n s tant and in v ar iab le fro m day to day I n o th er .

wo rd s th ere is pro bab ly as m u ch d ev iat io n in f u nct io n as in

,

fo rm and fo r t he p u rpo se o f the far m er it is ev en m o re

,

im p o rtant .

Ev ol u tion n ot a st u dy in morphol ogy on l y T he m is tak e is .

o ft en m ad e o f d efin ing ev o lu t io n a s exc lu s iv ely a s t u dy in m o r

p h o lo gy
1
I t m e an s m o.r e than that L iv ing b eing s a re s o m e .

th ing bes id es fo r m and th ei r ev o lu t io n so m eth ing m o re than t he ,

d ev elo p ment o f th eir fo rm ; ind eed in their s erv ic e to man bo th ,

an im als and plant s a re v alu ed les s fo r th eir s tr u ct u re than fo r

th eir f u nct io n wh ich is what th ey can d o A nd so it is that
,
.

1 l
Th e p ro b em o f ev e o p m en t is a n d l ac k no w e l dg d e m o rp h lo o gic a lp ro b em l ”

—C
.

. B D a v en p o rt , E x p erim en ta M o rp
. h l gy P t I P f l o o , ar , re a c e .

Th p ti
e conc e h l l d d ton i t d
ere a i fi
f l tu t e t of s Tnoh i d f cu o ac c o un or . e ea o

l ti
ev o u d l pm t
on or ev e o
pp d t t h
en l d as o m p t i o se f p i ol e o er as s u on o s ec a c rea

ti onw fi t as rsd t th anno unc e y l f th a igh t th

e v er t y b tw
c ose o t e e een c en u r , u as no

g en erall y b f t h p b l i
e o re t i l t h p e
p u f
c un t h O i gi f S
e a p i f t t
ea ra n c e o h e r n o ec es , a er e

m idd l f the o i t th t
e n n e een y A t t h t t im b i l gi t w
c en u r h i fly a e o o s s ere c e con

d i l ifi t i
c e rn e n c as s b d p ca t
o n as l t t as e f m It i
u o n ex e rn a t t g s ru c u re o r or . s no s ra n e,

th f
ere o re, th t t h d i ia h ld h fi t i
e s c u ss o n s d th b t t l
ou i id t t
av e rs a r s en , an e a es nc en o a

ne w t tli g
, s ar d i th p p l n , an mi d, n i l gi
e o th y h b fi t f gh t
u ar n , s ac r e o us eo r av e een rs ou

ou t i th fi l d
, n f m p h l gy
e e o or o o .

G d ll y h w
ra ua b i l gi t b g
, o t
ev er , o o th m
s s l m e an d m o c o n c e rn e s e v es o re an o re

with i t l t t ( h i t l gy ) d q it t t h i
n ern a s ru c u re s o o p i ,t hany f, d t
u h m e o e r su r r s e, e oun e

l
s e v es t ill wi th i
s th fi ld f nl ti Th e e m o t di ev o u i f ti
on .
(p hy i en ca e s u es n u nc on s

l gy ) h wi g
o o , s l i l y th t t h i t i m t t f d l p m t d b j t
o n c o n c us v e a s, oo, s a a er o ev e o en an su ec

t i ti
o v ar a d h di ty I t i th f
o n an e re t .ly s b t f th b
ere o re n o d on e rro n eo u s u or e ree er

d g an l y m i l di g t
ero u s id
s ea th t dy
n fo cons l ti er fi d t th fi l d
e s u o ev o u o n as c o n ne o e e

o f m p h l gy wh i h i
or o o t th , l i c s no t him th p im y m
e exc u s v e n o r if t o ev en e r ar an es a

ti on o f th g t p i eip l rea f l ti T h ir nc e o l i
ev o u f f ti
on . w ll ere s ev o u o n o u nc o n as e as

ev o lu tio n o f fo rm .
 8 V A R I AT I ON
7

a re n o t t he s a m e fo r any two s pec ies ( fu nct io nal v ariat io n bet ween

s pec ies ) .

A gain n o t wo ind iv id u a ls o f t he s a m e s pec ies can b e d epend ed

,

u po n to gi v e exact ly t he s a m e q u a lity o f m ilk fo r h erd r ec o rd s ,

s h o w that t he m ilk o f d iff er ent c o ws v a r ies nat u rally fr o m les s

than 3 per c ent to m o re th a n 6 per c ent fat (f u nct io nal v ar ia

l

t io n bet ween ind iv id u a ls ) N o r is th is d epend ent u po n t he f oo d.

s u pp ly fo r all a u th o r it ies agr ee tha t t he pro po rt io n o f fat t o

,

o th er so lid s is d epend ent u p o n t he ind iv id u a l a n d n o t u p o n her

feed Mo reo v er d ifferenc es n ear ly as wid e as th ese q u o t ed m ay

.
,

b e fo u nd with in the lim it s o f a s ing le h erd and th erefo re u nd er

id ent ica l c o nd it io ns a s t o feed .

S t ill aga in t wo ind iv id u a ls o f t he s a m e breed will pro d u c e

,

rad ica lly d ifferent a m o u nt s o f m ilk o r fa t wh ich ev er is meas u red , ,

fro m id ent ical a m o u nt s o f t he s a me k ind of f eed Th is ha s been .

rep eat ed ly and c o nc lu s iv ely s h o wn by Pro fes so r F ra ser o f t he

U n iv ers ity o f I llin o is 2
Pro bab ly n o fact in an im a l phy s io lo gy is
.

o f m o re far reach i ng i m p o rtan c e than is th is m ark ed i n s tanc e o f

-

f u nct io nal d ifferenc e bet ween ind iv id u a ls .

T hree exp erim ent s were c o nd u ct ed in t he att em pt t o d et er

m in e t he limit s o f th is d ifferenc e bet ween c o ws c o n s id ered go o d
en o u gh fo r a p lac e in a c o m m erc ia l h erd I n t he fi r s t 3
Ev a .

pro d u c ed 4 8 per c ent m o re m ilk and 1 1 pe r c ent m o re b u tt er

in n in ety o n e d a y s than d id J an et and in d o ing s o c o n s u m ed n o
-
,

m o re gra in and b u t p e r c ent m o re r o u ghn ess T h e s e c o ws .

were b o th mat u re were fres h o n t he s a m e day and n eith er s u f

, ,

fered acc id ent d u ring the exp erim ent yet Ev a pro d u c ed 1 0 5 7 ,

p o u nd s o f m i lk and 1 2 p o u n d s o f fat o u t o f an extra feed o f

1 1 2 po u nd s o f hay and c o rn s t o v er a d ifferenc e great er than ,

-

any margin o f pro fit t he dairy m an m ay ho pe to rea lize .

T he sec o nd experim ent was a c o m par is o n b et ween R o se a ,

nat iv e c o w n ine years o ld a n d N o ra a nat iv e c o w s ix years old ,

‘
, .

1 T he ac t u a l ran
ge in m i lk is fa r grea t er t h an t h es e fi g u res . S in g l e mi l ki
n gs

h av e b een k no wn t o ru n as lo w as p e r c en t fa t , an d Jers ey c o ws n ea r th l
e c ose

of l ac t a t io n o f t en g i v e m i l k with p e r c en t fa t .

9
S ee B u lleti n A o 5 1 a n d B u l leti n A
’
.
'

o . 66 , l
A g ric u t u ra l E xp erimen t S t a tio n,
y ll
U n iv ers it o f I in o is Ma y, 1 8 98 , .

3 I i
b d1 )
( 5 p 03
. 1 . .

B u lleti n N o 66 , U n iv ers it
. y of ll
I in o is , N o v em b e r, 1 90 1 .
 FU N C TI O N A L V A R I AT I O N
79

R os e c om m enc ed A pril 1 3 and N o ra May 2 2 1 8 99 and bo th , , ,

were milk ed fo r a f u ll t welv e mo nth s Bo th were in good h ea lth .

,

and bo th c o nt in u ed in good flo w u nt il the la s t R os e av eraging ,

o v er 1 8 po u nd s o f m ilk per day and N o ra n ear ly 1 4 po u nd s fo r

t he las t sev en day s o f t he t es t Ea ch c o n s u m ed all t he f eed s he

.

cared to tak e t he o nly res trict io n being that it s c om po sit io n

,

was t he s a m e fo r bo th N eith er was in any sen s e beefy b u t

.
,

R o s e ga in ed 1 8 1 po u nd s and N o ra 1 6 5 po u nd s fro m A u g u s t 1
t o A pril 1 s h o wing that th ey were ev id ent ly wo rk ing at o r near
,

th eir li mit o f m ilk prod u ct io n T he g rain fed was c o rn meal .

an d o il m ea l and t he r o u ghage c o n s is t ed o f c lo v er hay

,
rape , ,

green c o rn and c o rn s ilage alway s in c o m binat io n with o n e o r

, ,

m o re o t f h e f o llo w i ng — glu t en meal wh eat bran and gro u nd

, , ,

o at s .They wer e n ev er o n pa s t u re d u ring t he exper iment and , ,

a s has been s tat ed t he f eed was id ent ica l in q u a lity fo r bo th

, .

R o s e c o n s u med s light ly t he h ea v ier rat io n and y ield ed d ec id

ed l
y t he larg e r pro d u ct bo th in m ilk and fat T h e fo llo wi ng .

table exh ib it s t he t o tal feed c o n s u med and the prod uct yield ed
fo r t he ent ire perio d of t welv e m o nth s z l

COM PA R AT IV E M I LK PROD U CTI ON ON BAS IS OF FOOD CONSU M ED

FEE D MI L K B UTT R E '

C a s t in rba l fo r m th is m ean s that R o s e was ab le to pro d u c e

ve

47 p er c ent m o r e m ilk and 8 9 p er c ent m o r e b u tt e r than N o ra ,

with t he c o n s u m pt io n o f b u t p e r c ent m o r e f eed R ed ue .

i n g bo th to t he s a m e ba s is o f f o o d c o n s u m ed it ap pear s that ,

with a giv en a mo u nt o f feed f or ev ery 1 00 p ou n d s of m i lk g i v en

by N ora , R os e g a v e p ou n d s a nd f or wa y 1 00 p o u n ds o f
1 Fee di n p o u n d s of d ig es t ib le n u t rien t s . B u t t er rec k d
one at p er c ent

wa t er , a ddi
n g o n e s ix t h to t he b u tter fa t . Per c en t o f d iff l l d
e ren c e c a c u a t e on

N o ra as a b as e .
80 V A R I AT I O N

bu tter f at ( or b u tter ) p r od u ced by N ora , R os e p r od uc ed 1 80 7 .

p ou n d s . Fo r p u rpo s es o f m ilk prod u ct io n , th erefo re f eed wa s ,

wo rt h c ent mo re wh en fed to R o s e than when fed to

per

N o ra and fo r b u tt er pro d u c tio n i t was wo rth 8 0 per c ent m o re

,
.

T h is th en is t he tr u e m ea s u re o f the f u nct io nal d ifferenc e

, ,

bet ween th ese t wo c o ws and it is good and s u ffi c ient gro u nd ,

o n wh ic h t o ba s e breed in g o pe rat io n s F u rth er it is to b e n o t ed .

,

that th is is n o t t he d ifferenc e bet ween a goo d c o w and a poo r

o n e b u t be t wee n t wo go o d c o ws ; fo r N o ra prod u c ed po u nd s
o f b u tt er wh ich a s Pr o f es s o r F ras er rem ark s
, ,
is n ear ly three ,

t im es t he a v erage y ield ( 1 30 po u nd s ) o f c o ws in t he U n it ed
S tat es and a lmo s t o n e ha lf mo re than t he a v erage y ield ( 2 5 0
,

po u n d s ) o f what a re c o n s id ered pro fitable c o ws in I llino is .

I t may b e add ed at th is writ ing ( 1 906 ) that R o s e th o u gh u s ed ,

in m any experi ment s and exh ib it ed at v ar io u s s tate fa ir s and

at t he S t Lo u is Expo s it io n is s t ill liv ing ha le and h ea rt y at
.
, ,

s i xt een y ears o f age and is s t ill an ec o n o m ic a l pro d u c er o f m ilk

,
.

S he ha s an a v era ge y early r ec o rd o f 3 84 p o u n d s of b u tter f a t

fo r ten y ears and th o u gh s he has been in m any t es t s s inc e t he
1
,

o n e j u s t repo rt ed s he ha s n ev er b een b eat en b u t o nc e T hat .

was in t he f o llo wing ca se wh ich bear s f u rth er o n t he pres ent ,

po int T hree c o ws were in th is t es t with R o se — T ina C lay s

.
,
’

Q u een kno wn to b e a poo r c o w and t wo nat iv es kn o wn as

, , ,

N o 1 and N o 3 s u p po s ed t o b e t wo o f the fo u r bes t c o ws bo u ght

. .
,

fo r experim ent a l p u rpo s es o u t o f a h erd o f o n e h u n dred R ed u c ed .

t o t he s a me f eed ba s is a n d tak ing t he y ield o f Q u een a s 1 00 , ,

th a t o f N o 3 wo u ld b e repres ent ed by 1 2 1 o f R o se by 3 04 and

.
, ,

of No 1 by 3 1 2. T h is is a ra t e o f m o re than three to o n e aga in s t

.

t he poo r c o w o r o v er t wo a n d o n e ha lf to on e b etween g ood c o ws

,
-

on the s a m e f eed ba s is .

T h is d ifferenc e in t he efli c ien c y o f ind iv id u al c o ws is d epend

en t n o t s o m u ch u p o n d a ily d i ff erenc es as u po n t he ab ility fo r

lo ng t i me perfo r m anc e
-
S o me c o ws will g iv e a h ea v y y ield fo r
.

three o r fo u r m o nth s and go dry in s ix o r s ev en m o nth s o th ers

,

will g iv e a pro fit a b le y ield a lmo s t c o nt in u o u s ly B o th extrem es .

a re d ec ept iv e T he h erd s m an will a lmo s t c erta in ly o v errat e the

.

1
Si th b
nce w w itt R h
ea mpl t d tw l
ove asy r d f 7 58en ose as c o e e a e v e- ea r rec o r o 2

poun d f m i l k
s o d 3 6 p d f b tt f t
an
g 0 o un s o u er a as a n a v era e.
 F U N CT I O NA L V A R I AT I O N 81

f o rm er and u nd erra t e t he latt er so pro n e a re we to remem ber ,

s t r i k ing and m a x i m u m d a t a .

T h es e a re n o t is o lat ed and p ec u lia r c a s es Pro fes s o r F ra s er .

o f t h e U n iv er s i ty o f I llin o is t es t ed 5 5 4 c o ws in 3 6 c o m m erc ia l

dai ry h erd s o f the s tat e fo r a f u ll perio d o f t welv e mo n th s each .

He f o u nd that t he bes t 2 5 per c ent of t he wh o le n u m ber t es t ed

we r e a b le t o pro d u c e an a v erage o f 3 0 1 po u nd s o f b u tt er fat per
y ear wh ile the 2 5 per c ent o f lo wes t efli c ien c y were a ble t o pro
—
,

d u c e an a v erage o f b u t po u nd s a rang e o f c o n s id ,

e ra b ly m o r e than t wo t o o n e T he pract ic al s ign i ficanc e o f th is

.

d i ff e r enc e is po int ed o u t by Pro fess o r F ra ser as fo llo ws : I f it

c o s t s th irt y d olla rs a year to feed t he poo rer c o ws and th irty
ei g h t d o llar s a y ea r t o f eed t he b ett er o n es th en at pres ent pr ic es ,

a h e rd o f t wenty fiv e o f t he latter will prod u c e as m u ch n et p rofi t

-

as w o u ld a th o u s and o f t he f o r m er A litt le ca lc u lat io n will s h o w

.

t h e im m en s e sa v ing in lab o r i n k eep ing t he s m a ller h erd and what , ,

is e q u ally s igni ficant t he relat iv ely s m a ller in v es t m ent in an im a ls

, ,

feed and barn s and t he s m aller v o lu m e o f b u sines s gen erally

, , .

T he fac u lty o f pro d u c ing a h igh y ield o f m ilk m a n ifes t ly

d epend s n o t o n ly u po n t he act iv ity o f t he m a m m a ry g land s b u t
a ls o u po n t he c a pac ity o f t he s t o m ach t o hand le a large a m o u nt

Of f eed and t he ab ility o f ev e ry o rgan o f t he b o dy t o d is charge

,

i t s n o rm a l f u nct io n s reg u lar ly and t o end u re t he wea r and t ear

o f s u s ta in ed ex ert io n u nd er h ea v y pr es s u re T h is part ic u lar .

f u nct io n o f m ilk pro d u ct io n is th erefo re a k ind o f res u ltant o r

, ,

algebra ic s u m o f m any bo dy f u nct io n s a n d we s h o u ld n o t ex pect ,

t o find it s m ax im u m exc ept ra rely and in few ind iv id u a ls A .

s i m p ler f u nct io n p ra ct ica lly ind epend ent o f o th er s wo u ld th ere

fo re b e u nha m pered by th eir weakn esses and it wo u ld reach its ,

m ax im u m in a h igh er pro p o rt io n o f ind iv id u a ls .

V ar i at ion in meat produ c t ion T hat t he s a m e p rinc ip le is .

o perat iv e in m eat pro d u ct io n is ab u ndant ly s h o wn by ex peri

m ent s S t eers were fed sepa rat ely fro m ca lfh o o d to f u ll m at u rity
.

at t he M ich igan E xperim ent S tat io n The exper im ent was c o m

l
.

m en c ed as a br eed t es t by Pr o f es s o r S a m u el J o hn so n and c o m ,

p le t ed by t he w ri t er a s a t es t o f i nd i v id u a l d i ff e r enc es in ab il i ty
t o p u t o n ga in i n pro p o rt io n t o f eed c o n s u m ed .

1
B ll ti N
u e n69 A g i l t
o . l E p im t S t ti Mi h ig
, r cu u ra x er en a on , c an .
82 V A R I AT I O N

G A IN IN PROPO RT ION TO FE ED C ONS U M ED

S T R W E IG H T AT BEG I N N I N G
PO U N D S OF GRA IN To

GA
EE
ON E OF IN

D ifferenc es th is cha ra ct er a re f u rth er S h o wn by Pro fes so r

of

M u m fo rd s experim ent s with t he v ario u s m ark et grad es o f s t eers l

’
.

Feed ing catt le a re d iv id ed in t he m ark et s int o s ix grad es fro m ,

fancy select ed d o wn to inferio r A car lo ad ( S ixt een ) o f each o f .

th es e s ix gr ad es (n inety s ix a n im als in all) were fed o n t he s a me

-

rat io n fo r a pe rio d o f 1 7 9 day s The an im als were a ll nat iv es .

,

th o u gh t he bett er grad es S h o wed a m u ch h igher perc entage o f

go o d b lo o d than d id t he lo wer The fo llo wing tab le s h o ws t he .

relat iv e ab ility o f thes e s ix g rad es o f s t eers to hand le feed and

c o n v ert it int o ga in
R E LAT IV E E F FIC I ENC Y OF D I FFER ENT G RA D ES OF ST EE RS

RA G A IN PE R TO TA L G A IN TO TA L D xv MA T Dav M A TT R E PE R
G D ES
S T R
EE 16 STEERs T ER CO N S I '
III E D PO U D GA
N OF IN

Here is a v ariat io n o f o v er 3 1 per —

c ent (4 60 3 5 0) in t he t o tal
ga in of S i xt een s t ee rs u nd er eq u a l o pp o rt u n it ies ,
and , what is

1 f
’
. l l
B u lleti n A o 90 , A gri c u t u ra E xp erim en t S t a t io n , U n iv ers it y of ll
I in o is .
 FU N CT I ON A L V A R I ATION 83

mo re gn ificant a d ifferenc e of ov er 30 per c ent in t he feed

Si ,

req u ired fo r a po u nd of ga in This s ho ws t he inferio r feed ing .

q uality o f t he lo wer grad es d u e part ly to age and part ly to lack

,

o f breed ing .

Composi ti on of corn as influ en ced b y f u n ct i onal dev i a t ion On e .

h u ndred and s ixty three good s eed ears were selected fro m a
-

s train o f c o rn kn o wn as B u rr s Wh it e Th ey pr es ent ed t o the

l ’
.

e e n o m o re d ifferenc es than a re u s u al wi th s eed c o rn T hr ee

y .

ro ws of k ern els from each were analyzed fo r pro t ein and also
fo r fat A S a res u lt t he pro t ein in t he v ario u s ears was f o u nd
.

t o ra nge fro m p er c ent t o p er c e nt and t h e fat fr o m

p er c ent t o p er c ent .

C OM POS IT ION OF COR N FR OM 1 63 D IFFER ENT EARS

CO R N
P R O TEI N
u y D R A T Es
P R O TE I N

1
h d
T is was th e fo u n a t io n o f D r H o p in s wo r in c o rn b ree in g
. k ’
k d at t he Uni
v ers it y l
o f I lin o is , as re o r t e
p in B u lleti n A 7»d55 a n d B u ll eti n A o 1 00
.
’
. . I t wi ll be
f u rt h di
er sc uss e d la ter on .
84 V A R I A TI O N

C O M POS ITI ON OF C OR N F R OM 1 63 D I FFE R E N T EA RS -

Con tin u ed

CO RN
A S I1 P RT
O E IN
C AR BO CO R N
A SII P R T
O EIN
I I v D R A T Es l\ o
Y
.
86 V A R I AT I O N

I t is s u ffi c ientpres ent p u rpo s e t o n o t e that th ere was

fo r t he
a d ifferenc e o f p er c e nt ( 1 5) i n t he pr o t ein c o nt ent
o f t he h igh es t and t he lo wes t ear s o f t he 1 6 3 ana ly zed ; o f

p er c e nt —9 3)
0 i n t he s e ed p lan t ed ; and o f p e r c e n t
in t he cro p har v es t ed t he firs t y ear .

I t is n o t t he int ent t o p u rs u e th is s u bj ect f u rth er at th is

point I t will b e f u lly d is c u s s ed u nd er h ered ity and u nd er p lant
.

breed ing b u t en o u gh ha s been q u o ted to S h o w that th es e f u nc

,

t io n a l d ifferenc es a re b o th d is t i nct iv e and h ered i tary and that in ,

it all the ear is t he u n it to s u ch an ext ent th a t it is ent irely p rac

t ic a b le t o p er m an ent ly infl u enc e f u nct io nal d i fferenc es by s elec
t io n I nd eed th is has b een d o n e a lready to s u ch an ext ent that
.

c o rn has b een prod u c ed with a h igh er pro t ein c o n t ent than wh eat .

V ar iat i on in su gar produ c t ion S u ga rs o f v ario u s k ind s a re .

pro d u c ed by m any p la nt and an im al a ct iv it ies C ertain plant s .

exc el i n th is part ic u lar f u nct io n and a m o ng th es e wid e d i ffer ,

en c es ha v e b een f ou nd lead ing t o m a rk ed a n d p er m an ent i n

,

creas e in the a m o u nt o f s u gar pro d u c ed The b eet fo r exa m ple .

, ,

th o u gh o rig in ally pr o d u c ing b u t fro m 4 to 6 per c ent o f s u gar ,

ha s b een s o i m pro v ed a n d it s s u ga r prod u c in g act iv it ies ha v e -

b een s o increa sed as to y ield s pec imen s c o nta in ing as h igh as

2 5 p er c ent o f su gar and wh o le cr o p s a v eraging 1 4 per c ent .

C a n e is a ls o v ariab le and ev ery o n e fa m iliar with t he m a p le

,

kn o ws th a t c ertai n trees will y ield a large a m o u nt o f exc eed

in gly s weet s ap wh ile o th ers y ield b u t litt le wh ich li tt le may
, ,

b e eith er s weet o r ta s t eles s — ind eed ev en b itt er , ,

.

V ar i ation i n s peed , sc en t , an d organ i c act i v it i es gen er all y .

On e h o r s e is fa s t er o r m o re end u r ing tha n an o th er , not so

m u ch fro m c o n fo r m at io n a s fr o m inh eren t act iv ity and po wer

o f end u ra nc e S o m e d o g s a re es p ec ia lly k een in s c ent o th er s
.
,

a re d efect iv e and t he h ear ing i n s t inct is m u ch bett er d ev elo p ed

,

in s o m e ind iv id u a ls (d o g s c a t s h o rs es catt le b ird s et c ) than

, , , , ,
.

in o th ers 1
.

Mental q u alit ies p ers o n a l t as t es a n d int ellect u al a b ility in

, ,

gen era l are c o nd it io n ed n o t u po n c o nf o r m at io n b u t u po n t he

1 I t is m o re t h l ik l y t h
an e at s o m e o f t h es e d ifferen c es a re c o n n ec t e d with the
d egree of d l pm t f
ev e o en o c ert a in p o rt io n s of t he n erv o u s s y ste m . Th y e a re

n o n e the l ess fu n c
ti l h w
ona , o ev er .
 FU N C TI ON A L V A R I AT I O N 87

pec u liar a ct ion of cert ain part s pos s es sed by the rac e in c o m
mo n b u t wh o se s pec ia l f u nct io n ing in each part ic u la r ca s e
,

det er m in es t he plac e of t he ind iv id u al in t he sc a le o f life .

V ariati on i n v i tal f u n cti on s F o r pres ent p u rpos es t he ani m al .

body m a y b e rega rd ed as a c o lo ny of organ s ea ch end owed ,

with it s own pec u liar f u nct io n t he lif e o f t he wh o le and o f ,

ev er
y m e m b er being d epend ent u po n t he d eg ree o f s u cc ess

with wh ic h ea ch po rt ion d oes its wo rk The wh o le is th erefo r e .

, ,

as s tro n g as i ts weak es t m em ber and wh en t he wh o le is p u t t o ,

wo rk in serv ic e fo r m an that se rv ic e will d epend n o t o n ly u po n

,

the f u nct io na l act iv ity o f t he s pec ia l o rgan in v olv ed as t he ,

u dd er o r t he m us c u lar s y s t em b u t also u po n t he s u cc ess f u l ,

d is charge of all v ital f u n c t io n s wh en s u bj ect ed to the u nnat u ra l

s t ra in i n v o lv ed in wo rk ing u nd er pres s u re T he po int at wh ich .

the m a ch i n e will break d o wn o r fa il to d o s u cc ess f u l wo rk is ,

th erefo re a m att er of relat iv e s trength o f part s and in t he las t

, ,

an aly s is t he lim it ing element in perfo r manc e is n o t infreq u ent ly

on e o r m o re v ita l f u nct io n s wh ich experienc e S h o ws are as ,

v ariab le as a re o th er and fro m t he b iolog ical s tandpo int less

, ,

impo rtant charact ers .

T h e beat o f the h ea rt in m a n fo r exa m p le th o u gh s t ead ily

, ,

d ecrea s ing in rap id ity fro m infancy to old age yet v a ries b e ,

t ween d ifferent in d iv id u als at m a t u r ity all t he way fro m les s

th an fi fty t o mo re than eighty beat s per m in u t e A th let es tell .

u s that the s lo w beat is ch a ra ct eri s t ic o f lo ng d is tanc e ru nn ing -

and s u s tained eff o rt gen era lly b u t that ind iv id u a ls o f th is o rd er ,

are ill adap ted t o s h o rt d is tanc e ru nn ing o r o th er wo rk req u i r ing

-

q u ick res po ns e to s t i mu lu s .

T h ere is a m ark ed d i fferenc e in t he d ig es t iv e p o wer s o f

d ifferent an im als and s o m e ind iv id u a ls s tar v e b eca u s e t he
,

s t o m ach and int es t in es a re u nab le to d is s o lv e s u ffic ient foo d to

m eet the d em and s o f t he b o dy — and th ere a re all d egrees o f ,

s tar v at io n Oth ers with exc ellent d iges t io n b u t with lim ited
1
.

po wers of as sim ilat io n fa il to m ak e u se o f t he f u ll s u pp ly that

1 W id e s t u dy of m en , l
an im a s , a n d p l a n ts wi ll rev ea lm y
an c as es in w ic h h t he
d d lh
in iv i ua as a c c u s t o m e d it l f t
se o a n ab n o rm a ll y sma ll f d
oo s u pp ly . T he effec t

is n o t n ec essa ri ly fa ta l , b u t i t is s h o wn in re d uc e d o u tp u t eit h er o f l ab o r, bo dy
p ro d u c t, or ot h er fo rm o f o rga n ic ac t iv it y .
88 V A R I AT I ON

is p u t int o t he c i rc u lat io n I n t he ca s e o f R o s e and N o ra , p re

.

v io u s l
y c i t ed , what d id N o ra d o with t he exc ess o f f oo d c o n

s u me d D iges t io n exper iment s with ind iv id u als ind icat e n o s u ch

d ifferenc es in d iges t io n a m o ng h ea lthy a n im als as the d ifferenc es
in m ilk y ield that a re kn o wn to ex is t bet ween c o ws T he o n ly .

c o nc lu s io n is th a t in a cas e lik e th is the s u rp lu s foo d pas sed o n

o u t o f t he b o dy lay i ng exc es s i v e lab o r u po n t he excret i ng o rgan s
,

a s well a s i nc u rring los s u p o n t he o n e who pro v id ed t he f eed

1
.

F ro m th is it will b e s een that t he excret ing o rgan s th em

s elv es act a s a k ind o f s af ety v a lv e and that m u ch d epend s ,

u p o n th ei r relat i v e ab ili ty t o d is charg e th eir f u nct io n s well .

T h is th ey a re bett er adapt ed t o d o i n s o m e ind iv id u a ls tha n in

o th er s b u t that ev ery eff o rt is m ad e to k eep u p with d em and s
,

is ev id enc ed by the fact that if o n e k idn ey is lo s t t he o th er

act s fo r b o th u s u a lly increas ing in s ize S peak ing gen era lly a
,
.
,

c o w will g iv e as m u ch m ilk fro m three t eat s as fro m f o u r .

Wh eth er th is is fro m c o m pen s at io n as with t he lo s t k idney o r , ,

wh eth er i t is tr u e o n ly beca u s e c o ws a re s eld o m wo rk ed u p t o

th eir lim it s the data at b a n d d o n o t d et er m ine
, .

T he trem end o u s increa se in the act iv ity o f t he s a liv ary gland s

o n t he part o f t o bacc o ch ewer s t he increa s e i n S ize o f m u s c les ,

thro u gh u s e and the m ar v elo u s d ev elo p m ent o f s k ill in eye and

,

hand by lo ng c o nt in u ed pract ic e as in t he p lay in g o f m u s ic a l

-
,

in s tru m ent s read ing o n the part o f t he b lind etc

,
— th es e a re ,
.
,

a ll fa m iliar exa m p les o f f u nct io na l d ev elo p m ent thro u gh p ra e

t ic e T hat th is d ev elo p ment is o r may b e great er in s o m e ind i

.

v id u a ls than in o th er s is t oo well kn o wn to n eed m o re than

pas s ing m en t io n in th is c o nnect io n .

R es is tanc e t o d is ea s e and in v a s io n s o f t he an im a l ec o n o m y
gen era lly d iffe r grea t ly in d i fferent ind iv id u a ls S o m e a re a b s o .

lu tely imm u n e t o c ert ai n d is eas es o th ers p ec u liarly s u s c ept ible , .

I t is a m att er o f c o m m o n o b s erv at io n that in fi eld s o f c o rn

k illed by fro s t an occas io nal s talk rema ins green and u na ffect ed ,

s h o wing u n u s u al p o wer s o f res is tanc e d u e to s o m e c o n s t i t u ,

t io n al d i ff erenc e W ith o u t a d o u bt th es e a re t he d iff erenc es

.

1
h
T is is c o n c lu s iv e p roo f o f h
t he fa c t t a t a pp et ite is n o t a s afe g u i e t o t he d
amo u n t o f fo o d th a t c an b e p ro fi tab ly
c o n s u me d
T he m o s t t a t ca n b e s ai is
. h d
h
t at it is a goo d i di n c a t io n o f bo dy use a mo n g a n im a s l h
w o s e effi c ien c yi k
s no wn .
 o V A R I AT I O N

V ar iat ion in f ertili t y . C erta in

b ird s regu lar ly pro d u c e t wo
egg s o th er s three a n d s t ill o th er s f o u r b efo re i nc u bat io n
, ,
T he ,
.

av erage hen f ollo wing her nat u ra l hab it lay s a sett ing
, ,
“ ”

en t o f o u rt een ) and th en s u s pen d s fo r inc u bat io n T h cr op

( t e .

o f o v a has b een la id and t im e is req u i red fo r an o th er t o c o m e

,

f o rward The Ma ine s tat io n h o wev er has s u cc eed ed in great ly

.
, ,

increa s i ng t he pro d u ct io n o f eggs and has pro d u c ed o n e hen that

,

ha s la id t wo h u ndred and fi fty o n e eggs d u ring a s ing le y ear -

.

M o s t c o ws pro d u c e o n ly fi v e o r S ix ca lv es m any o n ly o n e o r ,

t wo and s o m e n o t any yet Old Gra n ny ( N o 1 in t he Gallo way

, ,
.

Herd Boo k ) prod u c ed t wenty fi v e t he la s t o n e in her t wenty -

,

n inth year T he d ifferenc e bet ween reg u lar and S hy breed ers
.

is t he d iff erenc e in t he f u nct io na l act iv ity o f t he repro d u ct iv e

o rga n s and n ext to perfo r m a n c e ability it is t he m o s t im po r
,

tant charact er in t he eyes o f t he breed er Ev en longev ity it self .

is n o t i t s eq u a l fro m t he s tandpo int o f the im prov er b eca u se ,

q u a lity cann o t b e s a id t o ex is t in t he rac e u n les s the ind iv id u als

that po ss es s it a re s u ffic ient ly fert ile t o in s u re its eas y and
c erta in perpet u at io n .

A ccu m u l ation of f u n cti on al v ariat ion s Ha v ing s h o wn t he .

m ar v elo u s d iff erenc es i n f u nct io n al act iv ity b et ween d iff erent in

d iv id u als a n d hav ing s h o wn that th es e d ifferen c es a re h ered itary
, ,

a s in c o r n it f o llo ws n ec es s arily that f u nct io na l v ariat io n s m ay b e

,

acc u m u lat ed int o tru e breed d is t inctio n s and that s trains o f ,

an im als and p lant s m ay b e permanen t ly es tab lis hed with exc eed
in gly h igh effic iency in d es ired lin es ; ind eed th is has b een already ,

acc o m p lis h ed th o u gh we a re s till far s h o rt of what is po s s ib le

, .

F o r exa m p le t he b eef breed s a re m o re ec o n o m ica l pro d u c er s

,

o f m eat than a re t he da iry breed s and t he c o n v er s e is t ru e a s ,

to m ilk pr o d u ct io n T h es e t wo f u nct io n s hav e th erefo re been

.
, ,

largely s eparat ed alo ng breed li nes B u t it canno t b e s a id t hat

.

o n e b eef breed is m o re effi c ient than an o th er in m eat pro d u c

tio n o r that any o n e d ai ry breed s tands o u t preem inent ly as

,

t he m o s t ec o n o m ical pro d u c er o f m ilk T h is i s pa rt ly b eca u s e.

breed d ifferenc es hav e b een largely b u ilt u p alo ng lin es o th er

than tho s e o f effi c iency a n d part ly b eca u s e all breed s c o nta in
,

m any i nd iv id u a ls o f lo w effic iency in th eir o wn breed characters

an d h igh effic iency in th o s e o f o th er breed s So me J erseys .
,
 FU N CT I O N A L V A R I AT I O N 9 1

therefo re ,
a re b ett er
feed ers than c ertain S ho rth o rn s and th ere ,

are t o b e f o u nd a m o ng t he latt er breed ind iv id u als that a re

m o re ec o n o m ical pro d u c ers o f m ilk th an a re m any o f the Jer

se
y br eed .

I t has been s aid that n o rac e was ev er tak en by a part bred -

h o rse o v er o n e that was rac ing bred Wh eth er literally tru e o r .

not ,
it is s u b s tant ial ly c o rrect s o i nt en s ely ha v e the rac ing
,

ab ility and in s t inct b een d ev elo ped in c ertai n breed s .

T h es e fact s to geth er with what is kn o wn as t o c o rn and beet s

, ,

S h o w c learly that m u ch e t rem a i n s to b e d o n e i n the way o f

y
d ev elo p ing f u n ct io nal act iv ity th ereby increas ing effic iency ,
.

SECT I ON I I — V A R I AT I O N I N T H E D EG R E E OF A CT I V IT Y
OF N OR M A L FU N CT I ON S WI T H I N T H E SA M E I N D I V I D U AL

Of no c ient ific o r ec o n o mic in t eres t than the d ata giv en

les s s

in t he las t s ect io n is t he fact that f u nc t io na l v ariat io n is by n o

m ea n s c o nfined to d ifferenc es b et ween ind iv id u a ls T he prin .

c ip le app lies th o u gh to a less ext ent to t he ind iv id u a l i t s elf

, , ,

wh o se act iv it ies a re n o t c o n s tant b u t v ariab le fro m day to day

an d thro u gh o u t it s life .

Daily flu ctu ati on in n or mal fu n cti on s I t will b e f o u nd u po n .

in v es t igat io n that t he o rd ina ry f u nct io n s o f the bo dy a re u n ex

p e c te d ly i rr egu lar E v en t he h eart a
. ct io n is n o t ab so lu t ely

c o n s tant it is S lo wer wh en the b ody is at res t than wh en it is

in act io n and is s u bj ect t o gr eat acc elerat io n in c erta in d isea s es .

A ll o rgan s o f t he bod y wo rk bett er s o m e day s than o th ers ;

i nd eed th ere is a d is t inct p er iod ic ity with each
,
— a p er io d o f ,

increa se f ollo wed by o n e o f m ax im u m act iv ity and that aga in by

, ,

o n e o f s u b s id enc e Th is is t he way o rgan s res t

. T h es e p erio d s .

a re ev id ent ly o f d iff e rent length s and it is th eref o r e o n ly occa ,

s io n ally that t he b ody a s a wlzole is a t i ts ma x i m u m A goo d .

exa m p le o f th is per io d ic ity with f u nct io na l d ev iat io n fro m day t o

day is aga in that of m ilk S ecret io n wh ich as has been rem ark ed
, , , , ,

is a k ind o f res u ltant o f a ll the act iv it ies o f t he b o dy T he fo l .

lowi n g tab le giv es t he v ari at io n s in t he q u ant ity and charact er

o f m il k fro m a S in g le c o w fo r a p erio d o f o n e m o nth
1
.

1 B ll a N 5 A g i lt l E p i m t S t ti U i it y f I ll i i
'

u e u o. 1 , r cu u ra x er en a on, n v ers o no s .
 V A R I AT I O N

o m o o
‘
T o M M M '
T C
I N N

xo xo w xo o xo rx xo cm o

N . N u
: N ~ 00 Q M M ~ m -r u st ; m m
v i
O
N

d md d d d d d o d d mo o
0 O 0

o o o o o o o

v fl m o
m '
f ffi ffi f
 V A R I AT I ON
94

C o m m ent is h a rd ly n ec es s ary t o s h o w t he i rregu lar wo rk ing

o f t he an im a l m ach in e wh ich in t his ca s e wa s a m at u re, s tro ng , ,

and h ea lthy c o w T he per c ent o f fat v aried a ll t he way fro m

.

to with in the s pac e o f t wenty -

fo u r h o u rs .

I nflu en ce of age u pon f u n cti on al act iv it y . At b irth the

vi tal f u nct ion s and th os e o f gro wth a re at th eir ma xim u m A t .

th is t im e gro wth s eem s to b e proc eed ing with t he en ergy “

o f em b ry o nic d ev elo p m ent I t c o nt in u es at a m ax im u m fo r a

”
.

t ime grad u a lly d ec lin ing in rat e u nt il m at u ri ty wh en gro wth

1
, ,

o f t he gen era l b o dy c ea s es alth o u gh fo r c e rtain part s (h air , ,

teeth h o rn h oo f reprod u ct iv e c ells etc ) it c o nt in u es n early o r

, , , ,
.

q u it e thro u gh lif e I n case o f acc id ent m any part s n o t c o m

.

m on ly ind u lging in gro wth aft er m at u rity a re ab le t o regen erat e

wi th m o re o r les s s u cc es s (bo n e S k in b loo d v es s els n erv es ) , , , ,

b u t a mo ng t he h igh er o rga n is m s gen era lly gro wth pra ct ically

c eas es at m at u rity I t is fo r th is reas o n th a t f eed ing en t erpr ises
2
.

a re m os t pro fi tab le with y o u ng a n im a ls that a r e s t ill gro wi n g .

A t th is age f u nct io na l act iv ity is great er and gen eral bo d ily

effi c iency h igh er .

On e reas o n why the lo wer grad es o f f eed ers a re les s ec o n o m

ica l prod u c ers than t he bett er grad es is that the an im a ls a re
o ld er t he d iff er enc e in age b et ween fancy s elect ed and inf erio r
,

s t eer s being o n e t o t wo y ears .

S trength and end u ra nc e a re ev id ent ly at th eir m ax im u m

s o m ewhat aft er m at u ri ty alth o u gh wit h the pas s ing of that ,

age is a ls o los t s o m eth ing o f t he po wer o f rap id rec u pe rat io n

and repair .

T he reprod u ct iv e f u nct io n s u nd ev elo ped u nt il a c o n s id era b le ,

period aft er birth atta in a h igh d egree o f en ergy if n o t th eir

, ,

m ax im u m so m ewhat b efo re f u ll m at u rity is reach ed T h ey th en

,
.

d ec lin e and fa il entirely in o ld age T h ei r d u ra t io n is th erefo re

,
.

c o n s id era b ly less than t he lif e o f the ind iv id u al o ft en dropping ,

belo w 5 0 per c ent o f t he f u ll life period .

1Th h wi g t f g wth t d iff t g h

e c u rv e s o n ra e o t b ro ffi i t ly a e re n a es as n o een s u c en

w k d
or et b t ou gh ,h b d u t
en o u i d i t t h t t
ash m i
eenm m t f
one o n ca e a e ax u ra e o

gro w t h i tt i d f
s aw d y f
a net b i t h ad t h tet h i m
a im m
s a i er g i r an a s ax u s n ev er a a n

reac h d e .

2T ti t i
rees c o n i i t nue m o t t t l t d i g lif ill t t
n c rea s e n s ze, o so e ex en a ea s ,
ur n e, u s ra

i g m k d d iff
n a ar b tw
e t h h igh p l
eren c e t d e im leen e er a n s an an a s .
 FU N C TI ONA L V A R I AT I ON 95

A t what period the reprod u ct iv e fu nct io n s a re mos t s u cc ess

fu l in pr o d u c i ng y o u ng o f a h igh o rd er o f exc ellenc e wh eth er in ,

yo u th midd le age or old age is no t d eterm ined th ou gh it is a

, , , ,

po i nt o n wh ic h light is bad ly n eed ed I t is c ertain th a t the .

pract ic e o f u s ing yo u ng and im m at u re s ires is alm os t u n iv ers al ,

es pec ia lly a m o ng catt le T hat th ere is dang er in c o nt in u ed

.

reprod u ct io n fro m im m at u re an imals ev en thou gh th ey a re sex

,

u a ll v igo ro u s th ere is gra v e rea so n t o fear and e t in genera l

y , y , , ,

repro d u ct io n ant edat es m at u rity .

T h e d u rat io n o f pro fitab le s erv ic e d ep end s o f c ou rs e u p o n , ,

t he n a t u re o f t he f u n ct io n in v o lv ed The life s erv ic e o f a rac ing

.

h o rse is m an ifes tly les s tha n that of a wo rk h o rse and t he life o f ,

a m eat a n imal is les s than that o f o n e k ept fo r m ilk .

I n flu en ce of ex ercise : u se an d dis u se T he b en efic ia l eflec t of

.

u s e i n d ev elo p ing and p erf ect ing the f u nct io n s o f t he b o dy has

been rec o gn ized fro m the m os t anc ient t imes A th let es tra in .

for th is p u rpos e M u s ic ian s pract ic e fo r m any h o u rs ev ery day ;

.

i nd eed th ei r ch ief lab o rs aris e fro m the n eed o f c o n s tant and

s ev ere e x erc is e o f t he m u s ica l fac u lt ies in o rd er to ach iev e any

c o n s idera b le d egree o f perfect io n and to h o ld it aft er it has

been acq u ired .

Ho rses d es ign ed fo r rac ing a re wo rk ed alm o s t fro m the firs t

i n o rd er to m ak e t he m o s t o f any nat u ra l ab ility t o tro t o r ru n .

C o ws a re b eliev ed to b e m o re effic ient pro d u c ers o f milk if th ey

begi n at t wo years of age and are k ept so to s peak in c o n s tant , ,

p ract ic e and barrennes s is believ ed to b e les s lik ely if h eif ers

,

a re bred early than if left t o attai n m at u r ity with ou t pro d u c

in g y o u ng .

D ar win d is c o v ered that t he wing bo n es o f wild d u ck s as c o m

pared with th eir leg bo nes were relat iv ely h ea v ier than th ose of
ta me d u ck s c o rres po nd ing to th ei r res pect iv e hab it s of life
,
.

The ar m o f the b lack s m ith and t he wing o f t he o s trich ; t he

remarkable leg of t he kangaroo and t he remai n s of that of the
whale ; t he bra in p o wer o f the b u s y m an and that o f t he S lu g
gard ,
— th ese and s c o res o f exam ples that m ight b e c ited S h ow
n o t o n ly that ex erc is e d ev elo p s q u ick en s and p er f ect s t he b o dy
, ,

f u nct io n s b u t th ey S h o w too that th eir v ery ret ent io n o r lo s s

, , ,

d epend s in t he lo ng ru n u po n th ei r c o n s tant and rat io na l u se .

 6 V A R I AT I O N
9

B lind peo ple acq u ire a q u ick en ed and an ed u cat ed s en se o f

h ea ring and a t o u ch that a m o u nt s a lm o s t t o a s eparat e s en se .

I n t he S a m e way t he d ev elo p i ng o f t he b o d ily f u nct io n s and

act iv it ies gen era lly d epend s u po n j u d ic io u s u s e (exerc is e) ; and
t he s k ill o f t he t ra in er and t he r es u lt s he is a b le t o ach iev e .

d epend n o t o n ly U po n his kn o wledge o f meth o d s t hat will m o s t

c ertain ly in s u re the ex erc is e o f t he d es ired part s b u t a ls o u po n
his j u dgm ent as t o ho w s ev ere a n d pro tract ed that ex erc is e
s h o u ld b e in o rd e r t o s ec u re t he m ax im u m effect s o f u s e and

n o t i nc u r t he d es tr u ct iv e c o n s eq u enc es o f o v eru s e T h is is as .

tru e in t he feed in g yard as u po n t he rac e t rack and applies as ,

well t o t he ra is ing o f goo d a n d pro fitab le feed er s a s to t he d ev el

o p ing o f rac ing h o rs es t he ed u c a t io n o f dr iv er s and s add ler s t he
, ,

t ra in ing o f h u nt ing d o gs t he try ing o u t of h o ming pigeo n s

,
“
,

o r t he t each ing o f canar ies t o S ing by n ev er al lo wing the y o u ng

b i rd s to h ear a fa ls e n o t e .

I n flu en ce of feed u pon f u n ct i onal acti v it y T he relat io n o f

.

t he a m o u nt o f f eed to it s ec o n o m ic a l c o n s u m pt io n is a s u bj ect
n eed ing caref u l in v es t igat io n En o u gh is kn o wn to warrant the
.

a s sert io n that an im als c an and d o learn to tak e am o u nt s fa r

larger than can b e rea lly u s ed Wh en a s t eer c o n s u m es o v er
.

a b u s h el o f c o m a day he has s imp ly fo rm ed t he eat ing hab it

a s t he res u lt o f a m o rb id app et i t e n o r is th is appet it e a n ind i
,

cat io n o f bo dy n eed s o r a gu ara nty o f it s p o wers to ec o n o m ica lly

c o n v ert t he feed int o m eat m ilk o r la bo r
, ,
.

I t is s ign i ficant that s t eers v ery grad u a lly bro u ght int o f u ll
feed will nev er tak e th es e en o rm o u s a m o u nt s Pro fes s o r H W . . .

M u m fo rd o f t he U n iv ers ity o f I llin o is find s that u nd er s u ch c ir

c u m s ta n c es t wenty p o u nd s o f gra in per day is a ll th e an im a l

will tak e .

C o n s u m pt io n o f extrem e a m o u nt s is th erefo re ev id enc e o n ly

, ,

o f t he q u ant it ies o f f eed t he d iges t iv e tract c a n carry and d is

charge with o u t c ala m ity o i i ts po wer to secret e gas tri c and

,

o th er d iges t iv e j u ic es and o f t he ab ili ty o f t he excret ing o r gan s

,

to elim i n a t e u n u s ed and u n u s ab le s u rp lu s fro m t he b o dy

I n t he c a s e o f R o s e and N o ra t he latt er c o n s u m ed t he s a m e
feed as t he fo r mer b u t ret u rn ed b u t litt le m o re tha n half as
m u ch . S he was u nd o u bt ed ly fro m t he s t a ndp o int o f ec o n o m y
, ,
98 V A R I AT ION

SECT I O N I II — M O D I FI C A T I O N O F N O R M A L FU N C TI O N S
BY EXT ER N A L O R OT H ER I N FLU E N C ES

To what xtent a re no r ma l f u nct io n s d epend ent u po n fa v o rab le

e

en v iro n ment and ho w d o th ey r es p o nd t o changed c o nd it io n s ?

A few n o tab le fact s wi ll thr o w s o m e light u p o n th is all im p o rt a nt -

q u es t io n .

Gal l s A n in s ect s t in g s a p lant

. U nd er t he in flu enc e o f t he
.

po is o n a ga ll is fo r med I f th is ga ll b e S h o wn t o a b io lo gis t he
.

will at o nc e s tate with c erta inty t he p lant wh ich prod u c ed t he

gall and the in s ect that m ad e the inj u ry so d efin it e in f o rm and ,

appearanc e is t he res u lt ing gro wth and s o d is t in ct is it fro m

any n o r mal gro wth o f the u n inj u red plant .

I n o th er wo rd s s pec ia lized p lant t is s u es s u bj ect ed t o a c erta i n

,

inj u ry prod u c e a n ew k ind o f gro wth a lm o s t a s s pec i fic a s wh en

o perat ing u nd er t he la ws o f h ered ity Here t he f u nct io na l ac t iv i
.

t ies o f t he p lant a t Me affec ted poin t ha v e been n o t d es troy ed b u t

per m an ent ly a lter ed in s u ch a m anner as to g iv e rise to n ew s tru e
t u res o f d efin it e fo rm and o ft en with s pec ific ch emical pro pert ies ,

as i n t he ca s e o f n u tga lls
1
c o nt ain ing 30 to 8 0 per c ent o f tann in .

I n ca s es o f th is k ind a n ew f u nct io n has been d ev elo ped s u dd en ly

o u t o f o ld m at eria ls a n d it at o nc e giv es ris e t o n ew and d is t i nct
,

pro d u ct s bo th s u b s tant iv e and m o rph o lo gical

,
.

Ab n ormal ov ergr owt h of disordered an imal t i ssu es W e ha v e .

j u s t n o t ed that v egetab le t is s u es s u bj ected t o s pec ific inj u ries

o ft en s u ff er s u ch a d era ngem en t o f th ei r f u nct io n s a s to ca u s e

t he pro d u ct io n o f an abn o r m a l b u t characteri s t ic o v ergro wth o f

t he inj u red part Q u it e s im ilar is the res u lt o f t he in v a s io n
.

o f t he an im a l ec o n o m y by s pec ific g er m s fro m with ou t a s in ,

the ca s e o f B a c i llu s tu b er c u los is Here t he gro wth s ( t u b erc les )

.

res u lt ing fro m the apparent att em pt to ency s t t he fo reign m ateria l

a re s u ffi c ient ly charact er is t ic to s erv e as a n a m e fo r t he d is ea s e

( t u b erc u los is ) .

T u mo rs g en era lly wheth er m a lignant o r b en ign a re per v er s e

, ,

o v ergro wth s o f n o r m a l t is s u es o f t he b o dy wh o s e o rd inary ,

1 A ll fo rm u la s fo r go o b d l k
ac writ in g in k i nc lu d e ga ll
n u ts
( o r n u t ga ll s ) as t he
c h d
a ra c t eris t ic in g re ien t T . h o s e c o mm o n l y u se d a re p ro d uc e d on t he o ak b y t he
s t in g o f the a llfl
g y( y p g
C ni s a llw t i n c tor i re
-
) .
 FU N CT I O N A L V A R I ATI ON 99

f u nct io n s hav e b een d eranged by s o m e cau s e ext ernal o r int er ,

n a l and wh o s e act iv it ies ha v e b een d iv ert ed t o t he pr o d u ct io n o f

,

abno r m al b u t cha ract eris t ic t iss u e with n o typica l t erm inatio n

t o it s gro wth .

N ea r ly a ll t is s u es o f t he bo dy a re s u bj ect ed t o th is d erange
1

ment o f th eir o rd ina ry f u n ct io n s res u lt ing in t he s u s pen s io n of ,

a ll ac t iv it ies exc ept that o f gro wth wh ich proc eed s with t he
en e r gy o f em b ry o n ic d ev elo p m ent and c o nt in u es ind efini t ely .

T h o u gh o ft en s u pp lied wi th s pec ial b loo d v es s els and a s y s t em

of n e rv es t h es e gro wth s a re ent i rely f u nct io n les s and th erefo re
,

u s eles s to t he bo dy B eing para s i t ic th ey a re a lway s a drain

.

u po n it s reso u rc es and o ft en fro m th eir nat u r e o r po s it io n th ey

,

co n s ti tu t e a real m enac e to its ex is t enc e .

A t u m o r repres ent s a b it o f d ifferent iat ed b o dy t is s u e th at

fo r s o m e reaso n o r o th er ha s aband o n ed it s cha ract eris t ic f u nc
tio n s c u t loo se fro m a ll res trai nt s o f h ered ity s et u p an ind e
, ,

pend ent ex is tenc e o f its own at t he expen se o f t he c olo ny of

wh ich it has been a res p ectab le and d ependab le m em ber and ,

has n o w d ev o t ed a ll its reso u rc es t o gro wth wh ich as ha s b een , ,

s a id pr o c eed s with t he en ergy o f em bry o n ic d ev elo p m ent res u lt

, ,

in g in n o th ing b u t f u nct io n les s m as s es o f li v in g m att er s tro ngly ,

s u gges t ing a rev er s io n t o pr i m it iv e u nd iff er ent iat ed t is s u e .

U nd er c o nd it io n s n o t well u nd ers t oo d a ll s o rt s o f abn o rm al

gro wth s m ay a ppear I n th is way an ant enna m ay appear wh ere
.

an eye ou ght to b e or it m ay en d in a fo o t in s t ead o f a feeler

”
,
3
.

T he writ er kn ew a y o u ng lady o f c u lt u re and o f n o litt le nat u ra l

bea u ty exc ept fo r the fact that gro wing fro m o n e ch eek was a , ,

t u ft of c oars e b lack hair three o r fo u r inch es lo ng Her no rm a l .

ha ir was bro wn and her c o m plex io n c lear What f u nctio nal d is .

t u rb a n c e c o u ld ha v e g iv en ris e to s u ch a gro wth is as m y s t erio u s

a s it was u nf o rt u nat e .

Oss ific at io n is a nat u ra l pro c es s b u t u nd er t he infl u enc e o f ,

exc es s iv e s trai n it m a pro c eed to an abn o rm a l ext ent as in

y ,

S pa v in wh ere t he ent i re h o ck j o int b ec o m es s o lid thro u gh t he

,

o ss ificat io n o f t he fl u id thro wn o u t a s t he res u lt o f i nj u ry .

1 M u sc l es , fa t t y ti
s s u e, c o n n ec t iv e t is s u e, b one c a rt ilag e, n erv es , g la n d s, b l d
oo

v ess els , t he c o v eri n g of t he b ra in , et c

—4 7
.

‘
1 Ba tes o n , M a t eria l s , et c .
, p . 1 5 1 .
3 Ibi d .
pp . 1 46 1 .
 00 V A R I AT I ON

D erangement s a mo re
f u nda m enta l nat u re o ft en aris e d u r
of

in g em b ry o n ic d ev elo p m ent res u lt ing in m o n s t ers o f a ll d egrees

,

o f a bn o r m a li ty Terat o lo gy has litt le int eres t t o t he b io lo g is t

.

genera lly b ecau se th es e abn o r mal car icat u res o f life c o ns t it u t e

,

n o th ing b u t s po ra d ic o ffs h oo t s o f the s pec ies D ev elo p in g .

fro m d ef ect iv e germs and ha v ing n o c o nnectio n with the lin e o f

d es c ent th ey a re o f litt le int eres t to t he ev olu t io n is t T h ei r
,
.

int eres t to t he t h rem ma t o lo g is t lies in th ei r bea r i ng up on f unc

t ion a l a c t iv i ty a nd tire d eg ree f
o c er ta i n ty w i t/r w/zic /z spec i al i z ed
t i s s u es m ay b e d epen d ed p
u on to d i s c /ea rg e t/zei r leered ita ry a nd

u nc t i ons
p p
r o er f .

V ariat ion du e t o t h e su ppression or f ail u re of t he r eprod u cti v e

fu n cti on s . abd o men o f the c rab Ca rc in u s m d na s n o rm ally
T he
has s ev en s egm ent s I n t he f em ale th es e a re d is t in ct
. I n the .

m ale the abd o men is m u ch narro we r and the d iv is io n s bet ween ,

t he th ird fo u rth and fifth s egm ent s a re o blit erat ed

, ,
M ales .
,

h o wev er inhab ited by t he pa ra s it e S a cc u l i n a d o n o t d ev elo p

,

s ex u a l character s a n d in th em t he s eg mentat io n is c o m p let e as

, ,

in t he fem a le 1
.

A y o u ng male is cas t rat ed T he part s remo v ed a re in no .

s en s e v ital and th ey s eem ing ly h av e n o c o n n ec t io n wit h o th er

,

o rgan s o f t he bo dy A ll t he b o d ily f u n c t io n s exc ept th o s e o f

.

reprod u ctio n pro c eed b u t n ot a s bef o re I n gen eral t he d ev elo p

,
.

ment o f t he s h ou ld er s and n eck will b e arres ted and th ey will ,

rem ai n lighter and finer T he v o ic e t he n erv o u s t em pe ra ment

.
, ,

t he d is po s it io n and t he gen era l act iv ity o f t he bo dy a re all

,

a ffect ed .T he m ane o f h o rs es will b e th inn e r fi n e r and s h o rt er , ,

.

T he ha ir o f fac e and neck in catt le will b e fi n er a n d les s c u rly .

I n ho g s t he t u s k s an d s h o u ld er p lat es d o n o t d ev elo p T he gro wth .

o f t he h o rn s is s t o pp ed in s h ee p b u t in catt le t he o n ly effect is ,

t o m ak e th em s ligh t ly lo n ger and a litt le m o re S lend er ap ,

p r o a c h in g t he f em a le typ e T h e h ind e r part s o f t h

.e b o dy as a
wh o le d ev elo p rath er m o re in ca s trat ed than in ent ire an i m als ,

and th ere is a gen eral appro ach t o t he fo r m o f the fema le I t .

is n o t ewo rthy in th is c o nn ect io n that t he s a m e genera l effect

fo llo ws t he failu re o f t he s ex u al po wers with ad v anc ing age ,

exc ept that t he b o dy d ev elo p m ent has a lready tak en p lac e .

B t M t i l 1 t p 95a es o n , a er a s . e c .
, . .
102 V A R I AT I O N

Pla nt s ,
and an im als t oo fo r that m att er gro wing in c old ,

c lim a t es o r u nd er hard c o nd it io n s s u ffer pro fo u nd changes t o ,

wh ich th ey b ec o m e acc u s t o m ed (acc lim at ed ) and wh ich a re ev er

af ter ward c o n s t it u t io n al W e bec om e acc u s t om ed t o c o ld o r to h eat
.

and a re t hereaft er les s a ffect ed by extrem es R ec ent calo rim et er .

t es t s s ho w that t he t em perat u re o f t he h u m an body is lo wes t fro m

three t o fi v e o c lock in t he mo rn ing and h igh es t from o n e t o thr ee
’

in t he aft ern o o n th u s fo llo wing fa irly c lo se t he m in im u m and

,

m ax im u m o f o u t s id e t em perat u res T h es e c o nd it io n s c o n t in u e .

ev en if t he s u bj ect wo rk s at n ight and s leep s in t he dayt im e .

T wo c o n d it io n s t en d t o prod u c e hard an d s pi ny gr o wt h in
v egetat io n T h es e a re int en s e light a n d extrem e d ryn es s Bo th
. .

a r e fo u nd in tr o p ic a l r eg io n s and wh en th ey o cc u r t ogeth er th eir

,

m ax im u m r es u lt s fo llo w as to har s hn ess an d s p in es T h ese c o nd i .

t io n s ca n b e v erifi ed in t he labo rat o ry s h o wing conc lu s iv ely ,

that the charact er of grt is in a m ea s u re at lea s t d epend ent

, ,

u po n s u rr o u nd ing s .

S peak ing gen era lly p lant lic e reprod u c e parth en ogen et ically
,

d u ring t he gro wing sea so n o f t he s u m m er and d u r ing th is t ime ,

o n ly W ing les s fem a les a re pr o d u c ed W ith t he approach o f cold .

weath er h o wev er a winged b is ex u al bro od is pr od u c ed t hat liv es

, ,

ov er wint er .

T h ese c o nd it io n s c a n b e pr od u c ed art ific ially in t he green ho u s e

at any t ime by lower ing t he t em pera t u r e and allowing t he plant s
o n wh ich t he lic e f eed t o dry u p T h u s we m ay sa y that wings
.

and sex may b e d ev elo ped at will by t he m a n ipu lat io n o f t he c o nd i

t io n s o f life .

T he s o ca lled c o n v ers io n o f o n e s pec ies int o an o th er by in flu

-

en c in g it s en v iro n m ent ha s b een largely o v er s tat ed and et t he

y ,

fact s are that wh en S c hm an kewit s c h grew A r tem ia s a lin a in

1

wat er wh o s e s alin e c o nt ent was gr ad u ally increa sed t he ca u da l ,

fi n s and th eir br is t les progres s iv ely d egen era t ed u nt il in m any ,

ca s es th es e append ages had d isappea red t he an im al th u s a s su m

, ,

in g t he chara ct er o f A m i l/za u s en i i wh ich n o r m ally liv es in wat er s

.
,

o f extr em e d en s ity Thes e experim ent s wer e u nd ertak en beca u s e

.

h e s eem ed t o ha v e o b s erv ed th is tran s fo rm a t io n t a k ing plac e

nat u rally in a la k e cr o s sed by a d am and which wa s inhab it ed ,

B t M t i l
1 t p 96
a es o n , a er a s , e c .
, . .
 FU NC T I ON A L V A R I AT I O N 1 03

by bo th s pec ies the on e abov e the o ther Th is dam bro ke

, .
,

m ix ing t he t wo S pec ies b u t in three y ear s o riginal c o nd it io n s

,

were pract ica lly res t o red T he exper im ent s wer e u n d ert ak en .

to lea r n wheth er rea l tran s fo rm at io n had t a k en p lac e o r wh eth er

t he r es u lt had been bro u ght abo u t by s elect io n .

The experim ent s eem s c o n v inc ing and t he po int is f u rt h er ,

s tr ength en ed by t he fact s tha t S c hm an kewit sc h res t ored the

ca u d al fi n s by red uc ing the solu t io n and that wh en the r ed u ct io n ,

was carried below the n ormal of A sa l i na with in thr ee gen era .

,

t io n s t he las t segm ent o f t he body d iv id ed aft er t he fas h io n of

A b ra nc l uj
. ous an o th er relat ed S pec ies
, T he fact s are t he m o re .

rem a rkab le as A s a l i na
. reprod u c es par th eno gen et ica lly wh ile ,

A b ra ucl uj
. ou s is n o t kn o wn t o d o s o .

Bio logis t s are extr em ely car ef u l n ot to as su m e t he ab so lu t e c on

v er s io n o f o n e s pec ies int o an oth er by any s u ch d irect m eth od s

as hav e h er e been n o t ed t he q u es t io n being rath er wh eth er th ey

, , ,

are all good s pec ies ; b u t that S ingle charact er s ar e pr o f ou nd ly

infl uenc ed by changed c o nd it io n s and c om e to res em b le t he sam e ,

chara ct er in an oth er and relat ed S pecies is too well estab lish ed ,

t o b e lo nger q u est io n ed What light th is m ay finally throw u po n

.

t he o rigi n o f s pec ies is pro b lemat ical b u t it s erv es t he present ,

purpose in s h o wing the power of env iro n m ent to pr o fo u nd ly

m od ify t he f u nct io na l act iv it ies o f liv in g beings .

If a c u tt ing o f willo w c u rra nt o r o th er s u itab le gro wth b e

, ,

plan t ed in the ea rt h roo t s will s tart fro m t he part belo w t he

,

grou nd and leav es an d branch es from the part abo v e I f n ow

,
.
, ,

it b e c u t o ff at t he s u rfac e o f t he gr o u nd and the to p po rt io n b e

plant ed again it will aga in tak e roo t at the n ew point o f s ev er
,

ance wh ich had befor e bo rn e lea v es The proc es s m ay b e c o n .

t in u ed ind efin it ely o r u nt il t he p iec e is u sed u p s h o win g that

, ,

roo t s or lea v es m ay b e d ev elo ped at will at any po int alo n g t he

c u tt ing acco rd ing as it is plac ed belo w o r abo v e grou nd
,
.

I f a c u tt ing b e plant ed roo t s will d ev elo p o n ly at t he lo wer en d

,
.

I f h o wev er before p lant ing it b e c u t int o t wo p iec es each will

, , ,

d ev elop roo t s on the part belo w gro u nd and in m any s pec ies th is ,

will occ u r ev en if t he p iec es b e in v ert ed and plant ed to p d o wn 1

.

1 Th i i m
s s t l ik l y t t k p l
os e i y
o g w ad l
e l ik l y i ld w d S
ac e n o un oo , es s e n o oo . ee

M o rga n R egen era t io n pp 7 1 9 I

, , .
-
.
1 04 V A R I AT I O N

A m aple tree gro wing in U rba na had forked int o t wo n ea rly

eq u a l part s abo u t s ix f eet fr om t he gr o u nd On e part wa s s p lit .

d o wn and t o rn o ff in a h ea v y s t o rm wh en it was s een that roo t s ,

had d ev elo ped in t he cr o tch and were ev id ent ly at wo rk u po n t he

s o il that had b lo wn fro m t he s treet and t he m o is t u r e that had

acc u m u lated from ra in s .

The writ er was excav at ing fo r a bas em ent A b lack ch erry .

tree st ood so m e ten feet fr om t he lin e o f t he wa ll I n tak ing .

th is o u t many of t he roo ts were sev ered th eir c u t end s being left ,

in the bank o f u nd is t u rb ed earth I n a few day s th es e c u t end s

.

were c loth ed with a gro wth o f gr een lea v es Here was t is s ue .

that u nd er n or mal co nd it io n s f u nct ion s o n ly as roo t s yet u po n

, , ,

o cca s io n r ea d ily g iv es r is e t o bo th leaf a n d s t em .

A ll plant s and m o s t a n im als m a inta in d efin it e relat io n s t o light ,

and if free to m o v e or ient th em selv es with r eferenc e to the

1
,

d irect io n o f t he light ray s s tr ik ing the s u rface o f th eir bod ies .

A plant bend s t oward t he wind o w b eca u s e o f the c o ntract ing

effect o f t he light u po n t he pr o t o p la s m o f o n e S id e o f t he s t em .

Many larv ae a re n egat iv ely h elio tro p ic ; that is t he light ed S id e ,

o f t he body is m o r e irr itab le and th ey m ov e a way fr o m t he light ,

co m ing to res t o n ly in dark plac es wh ere th ey feed a n d m at u re , .

Oth er s are po s it iv ely h elio tr o p ic wh en h u ngry and n egat iv ely

h eliotrop ic wh en fed S u ch lar v ae will c lim b tr ees and feed u po n
.

t he lea v es o r b u d s u nt il fi lled wh en b ec o m ing s en s it iv e to light

, , ,

th ey d es c end and h id e in t he grou nd u nd er r u bbis h o r in any , ,

o th er p lac e s h ield ed fro m t he s u n T h is act io n has b een erro n e

.

o u sl
y attr ib u t ed t o a s em i int elligent in s t inct-
I t is n o th ing b u t .

f u nct io nal d epend enc e u po n ext ern al s t im u li Th is is n o t t he .

place to pu r su e the su bj ect in d etail — wh ich will b e d o n e wh en ,

d c
is u ss ing n t nct d fl ex A ct o n
“
I s i an R e i — b u t it is t he plac e t o ,
”

no t e the wo nderfu l d epend enc e o f c ertain n o rm al f u nct ion s u po n

ext e n a l infl u enc es
r .

A n an im al is in v a d ed by a fo reign germ and s u ff er s from

d isea se I t is in m os t cases ev er aft er imm u n e t o that d isease
. .

What change has been wo rk ed in t he an imal ec o n om y ? W e

1 By o ri en t a t io n is m ea n t t he di r ec t io n h h
in w ic t he l o n g a x is o f t he bo dy is

h
b ro u g t to res t wit h r efer en c e t o s u rr o u n di g b di
n o es o r in fl u en c es , s u c h as gr av
it y o r l igh t .
 1 06 V A R IAT IO N

o th erwise wo u ld b e lef t with o u t f oo d T h ere is th erefo re a .

, ,

part ic u lar s eq u enc e in th is c o m plicat ed p erfo rm anc e that m u s t

b e o b s er v ed o r fa ilu re res u lt s ; and fa ilu re is fatal t o t he ex is t
enc e o f b o th S p ec ies .

M o reo v er th i s a c t is p erfo r m ed b u t o nc e in t he lif et im e of

,

t he m o th who has n o kn o wledg e o f t he ac t s o f her pred ec es s o rs

, ,

and is th erefo re n o t pro c eed ing fro m s im u lat ion ; n o r has S he

o pp o rt u n ity t o learn t he f a t e o f her o ffs pring and pro fit by

t he exp erienc e .

N o w t h e tr u th is th a t u n erri ng a s is th is p erf o r m anc e a , ,

goo d m any o v u les es c a pe fro m failu re o f t he egg t o hatch o r ,

fro m o th er c a u s es and th u s t he y u cc a is ab le t o m at u re s o me
,

s eed T hat c o m p licat ed pro c es s es o f th is k ind a re n o t alway s

.

carried o u t in pro per s eq u enc e and f u ll d eta il is S h own by

caref u l s t u dy o f d ifferent ind iv id u a ls as po int ed o u t es pec ially ,

by Pro fesso r C S C randall in his s t u d ies o f t he app le and plu m

. .

l
c u rc u lio .

C aref u lt dy o f th es e c o m p licat ed act s and o f the bo dy

s u

f u nc t io n s in gen eral m u s t c o n v inc e t he s t u d ent n o t o n ly o f th eir

n ic e adj u s t m ent b u t ( what is o f eq u al c o n seq u enc e) o f th eir
exc eed ing v ar iab ility and i rr eg u lar ity with i n lim it s that c ert a in ly

are by n o m ean s narro w .

C as es o f d o u b le perso na lity in wh ich t he ind iv id u al

“
,

b ehav es fo r a t ime a s an o th er and d is t inct ly d ifferent perso n ,

a re t o o well kn o wn t o req u ire mo r e than a pas s ing n o t ic e .

Th es e a re in s tanc es in which an ent irely n ew s et o f f u nct io n s

is bro u ght int o p lay — d is t inct fro m t he n o r mal yet wo rk ing
, ,

t o geth er to t he acc o m p lis h m ent o f d efin it e end s .

B u t a fe w o f t he m any m od i fic a t io n s o f n o rm al f u nct io n s
’

hav e been m ent io n ed T h is is n o t t he p lac e t o exh au s t t he .

s u bj ect On ly en o u gh has been g iv en t o S h o w t he s t u d ent that

.

ev en t he h igh ly s p ec ia lized f u nct io n s a re s u bj ect t o t he la ws o f

v ar iat io n T he m att er will b e mo re c o m p let ely c o v ered u nd er

.

C a u s es o f V ariat io n
”
.

1 B u lleti n A
'
o 8
9 , A gric u t u ra l l E xp erim en t S ta t io n , U n iv ers it y of ll
I in o is
—
.
,

pp .
5 00 5 0 2 . T he ac c o u n t o f the d ifferen t wa y s h h th
in w ic ree d iff
eren t fem a l es

f m d th e ir wo r k
is g iv en in f ll d “
T ra n s m is s io n of Mo d ific a t io n s ”
p er o r e u un er ,

s ec t io n o n Ha b i t an d I n s t in c t .
 FU N CT I ON A L V A R I ATI ON 1 07

SEC TI O N IV — N O R MA L FU N CT I ON S EXER C I S ED
U ND ER A B NOR M A L CON D I TI ON S

T he ma m m ary gland n o rmally c o nfined to females is c om , ,

m o u ly fu nct io n les s u nt il aft er p regnancy ; b u t by m an ip u lat io n

o f the u dd er h eifers and o th er fem ales may b e mad e to y ield
,

m ilk with o u t beari ng yo u ng A ga in ru d imentary m a m mae .

, ,

pres ent genera lly in m ales are occ as io nally acc ompan ied by ,

c o ns id erab le d ev elo pm ent o f m a mm a ry t issu e n early alway s ,

b u t no t n ec ess ar ily f u nct io n less 1

.

Mo s t rem arkable o f all ma mm ary gland t is su e has been ,

-

kno wn to d ev elo p in extremely u n u s u al p lac es u po n t he bod y .

Ma mm ary t u mo rs in t he ax illa ( ar m p it s) a re d es cr ibed as o f

c o mmo n occ u rrenc e in ly ing in wo m en 2
T h ese t u mo rs hav e -
.
”

no d u ct b u t in s q u eezing th ey y ield bo th c o lo s tru m and

,
“

m ilk fo llo wing in t he s a me o rd er as fro m n o r ma l m a m mae

”
, ,

and oo zing thro u gh the s ki n at the s it u atio n s of t he sebac eo u s

follicles .
”

Besid es th ese th ere is ind is p u tab le ev id enc e o f t he pr esenc e

“

o f a ma mm ary g la nd o n t he th igh o n t he ch eek on

t he acro mio n ( S h o u ld er po int ) and in t he lab iu m m aj u s .

I n the t wo las t ca s es t he m a m m ary nat u re o f t he gland

was prov ed by m icro s c o p ic exam inat io n 3 ”
.

S im ilar c o nd it io n s m ay b e pro d u c ed a rt i fic ia lly by graft ing ,

and all so rt s o f abno rm a lit ies m ay t es t ify to the pers is t enc e

with wh ich h ighly s p ec ialized t issu e c o nt in u es to d is charge it s
f u nct io n s o ft en u nd er t he mos t d is c o u raging c irc u ms tanc es
,
1
.
‘

F o r exa m p le Hu nt er and D u ha m el graft ed the s p u r o f a

,

yo u ng c o ck int o h is c o m b wh ere it c o nt in ued to grow to its ,

n o r m al s ize 5 “
Bert t ra n sp lant ed t he ta il o f a wh it e rat t o
.
”

t he bo d y Of M u s d ecu m a n us ( t he c o m m o n bro wn ra t ) wh ere it ,

c o nt in u ed a liv e 5
T he s a m e expe ri men t er bent ov er t he tail
”
.

1 D r Ho t t es ,
. a p ers o n a l frien d of t he wri ter, k ne w a y o ung m an in G erman y
who was su c k li n
g an i n fa n t .

2 B a teso n , M a t eria l s , etc .

, p . 1 8 5.
3 Ib i d .
p . 1 87 .

1 As w h en a p iec e o f m a m mar y gl d w an as gra f t e di t n o t h e ear o f a gu in ea p ig ;

wh en t he p ig b ec a m e p regn a n t t h gl de an c o m m en c ed t o s ec r et e
—7 9
.

5
Mo rga n, R egen era tio n p p , . 1 78 1 .
I 08 V A R I AT I O N

of a ra t and g raft ed it back int o its o wn bod y A ft er it had .

u n i t ed he s ev ered it at t he n o r m a l ba s e and th u s pro v id ed t he

an im al with a rev ersed ta il

“
He fo u nd h o wev er t hat t he .
, ,

tail o f the m ou se d id n o t gro w as well in the bo dy o f the rat and

wo u ld no t u n it e at all with t he b o dy o f eit her t he d o g o r t he cat 1
.

Bo rn s u cc eed ed in u n it ing t he a nt er io r and po s t er io r part s o f

the tadpo les o f t wo d i fferent gen era o f fro g s ( R a na esc u len ta
fo r ant erio r and B om b i na tor ign eu s fo r p o s t er io r ) T he c o m b i .

nat io n liv ed fo r ten day s wh en it was k illed beca u s e o f path o ,

lo gica l C hanges 2
.

I n t he s a m e way Har ris o n m ad e u p an ind iv id u al o f t wo

s pec ies ( R a n a v i r es c en s and R a na pa l u s tr i s ) T h is he k ept aliv e .

u nt il aft er it s tran s fo r m at io n i nt o a fro g each ha lf reta in ing

“
,

t he c haract er is t ic f ea t u res o f t he s p ec ies t o wh ich it b elo ngs 2 ”

.

T h is being tru e it is n o t s u rpr is ing that m any v ar iet ies o f

,

app le can b e gra ft ed into the s a m e tree to p E xa m p les o f t his .

so rt mig h t b e m u lt ip lied i nd efin i t ely a s in t he m ak ing u p o f ,

wo rm s by graft ing t o geth er p iec es o f t wo d ifferent S pec ies in ,

wh ich each p iec e pres erv es it s s pec ific charact ers ; b u t en o u gh

has been giv en t o S h o w t he p ers is t enc e with wh ich s p ec ia lized
t is su e c o nt in u es to d is c harge its nat u ral f u nct io n ev en u nd er t he
hard es t o f c o nd it io n s 3
.

T he C irc u m s tanc es u nd er wh ich liv in g m att er c a n d is charge

it s n o r m a l f u nct io n s u na lt ered eith er in ch a ract er o r i n d egree , ,

and t he lim it s bey o nd wh ich th ese f u nc t io n s m u s t c ease o r

u nd erg o a lt erat io n — all th is i s n o t o n ly a q u es t io n o f d eep
,

b io logical int eres t b u t it is o n e o f s pec ial s ign ificanc e in breed

in g b eca u s e it thro ws n o li tt le light u p o n t he rea l nat u re and
,

cau ses o f v ariab ility a s u bj ect u p o n wh ich we so rely n eed in

,

fo rm at io n if v a riat io n s a re ev er to b e c o ntro lled eith er in t heir

d ev elop ment o r in th eir tra n s m is s io n .

Su mmary We a re t o rega rd v ariat io n s in f u nct io n as well

.

a s in f o r m o f act iv it ies as well as o f s tr u ct u re o f what an

, ,

an im al o r plant d oes as well as what it is , .

T he b o dy f u nct io n s a re n o t c o n s tant b u t v ari ab le T h ey a re ,

.

v ari ab le a s b et ween d ifferent ind iv id u a ls and a lso with t he s a m e

M g — d
I b id h p i pp 59—
1 R g
o r an , ti pp 7 8
e en era 79 on , I
. b i 1
pp 8 318 5 .
9 . . 1 -
1 .

1 c a . 89 G ft i g
x, d R g
1 ti1 ra n an e en era on.
. .
,
 CH A PTE R V I
MU TA TI ONS

SECT I ON I D I ST I N CT I ON B ET WEE N M UTA T I O N

AN D OR D I N A R Y V A R I AT I O N

T he d ev iatio n s fro m type h eret o fo re c o n s id ered a re t ho s e o f

i nd iv id u als rath er than o f gro u p s Wh et her q u ant itat iv e s u b .
,

s t a n t iv e m eris t ic o r f u nct io n a l t h ey repres ent the fl u ct u at io n s

, , ,

o f ind iv id u a l m em ber s o f a s pec ies o r a v a riety abo u t t he n o r

m al type o f t he rac e n o t n ec es s a rily ex hib it ing any t end ency

,

t o d epart per m an ent ly fro m that type .

T he s t u dy o f th es e d ev iat io n s s h o ws t hat wh ile n o two ,

i nd iv id u al s a re alik e yet t he d epart u res o f c erta in ind iv id u als

,

in o n e d irect io n a re c o m pen s at ed by d epart u res o f o th er ind i

v id u als in t he o pp o s i t e d irect io n I n o th er wo rd s the m em b ers .
,

o f a rac e c lu s t er c los ely abo u t what m a b e ca lle d a c ent r f

y e o

fl u ct u at io n wh ich is in mo s t cas es c o mpa ra t iv ely s tat io nary

, , , .

Beca u s e o f th is f a c t we m ay ha v e a rela t iv ely fi x ed typ e ,

ind icat ing a prac tica lly s tat io nary ra c e ev e n in t he m id s t o f ,

c o n s id erab le ind i v id u al d ev ia t io n l
.

M u tat io n s o n t he o th er hand m ark s u dd en a n d d is t inc t

, ,

d epart u res fro m typ e T he pend u lu m s wings b u t d oes n o t re

.
,

t u rn A n ew c en ter o f fl u ct u a t io n is es tablis h ed fro m wh ich

.
,

ind iv id u a ls d ev iat e in a ll d irect io n s a s bef o re I t is n o t that .

t he o ld c ente r is aba n d o n ed — fo r t he m as s Of ind iv id u a ls s t ill

,

c lu s t er abo u t it as b efo re b u t i t is tha t a n ew c ent er is es t ab

,

lis hed abo u t wh ich a n ew gro u p c lu s t e r s and all c lo s e o b serv er s

, ,

rec ogn ize a t o nc e th a t a n ew type ha s b een b o rn int o t he wo rld

1 h
T is fa c t is l y f i g p i ll y t im l b d
ex t rem e c o n us n I t h m id t
, es ec a o an a ree ers . n e s

of wi d e v ar a i ti d wi t h b t f w i d i id
ons an l li i g t y
u e t im th b
n d
v ua s v n a an one e, e ree er

is o ft en una b l t t ll wh t h
e o e hi g el g ( t
ery p ) i ism p i g t
en era a v era e or e s ro v n , re ro

g ra di n g, o r s an t d i g till T h i m t will b ll d d t g i d Typ d

n s . s a t er e a u e o a a n un er e an

V aria b i it l y .
"
 M UTATI ONS I I I

and a n ew rac e is es tablis h ed o n t he ea rth This n ew gro u p .

and its c ent er c o ns t it u t e a m u tat io n and t he ind iv id u als are ,

s po k en o f a s m u tant s V ariat io n has bec o m e d is c o nt in u o us as

.

well as c o nt in u o u s .

T h es e s u dd en o ffs h oo t s fro m es tab lis h ed S pec ies were n o t ed

by D ar win who ca lled th em S po rt s He c o n sid ered h owev er
, .
, ,

that n ew s pec ies are fo rmed o nly by the S lo w b u t c o nt in u o u s

act io n o f select io n wo rki ng wi th o rd inary fl u ct u at io n s (c o nt in uou s
v ar iat io n s b ild ing w typ a li t l at a t im thr gh h
) u , u p n e e s t e e o u t e

g rad u al acc u mu lat io n of S light b u t fav o ra b le d ev iat io n s

—
.

His so ca lled s po rt s were th eref o re m y s t erio u s and fro m ,

t he fact that u nd er nat u ral c o nd it io n s th ey genera lly d isa pp ear

rap id ly by c ro s s i ng he was led to attach litt le im po rtanc e to
,

th ese s u dd en d epart u res fro m t he es tab lis h ed type La t er 1

.

resea rch es h owev er hav e g iv en th em u nexpect ed sign ificanc e

, ,
.

The d is t ingu is h ing f ea t u re o f a m u tat io n is that th ere a re n o

i n t er med iat es bet ween t he o ld type and the n ew wh ic h was ,

th erefo re attained no t by S lo w d egr ees b u t by a s u dd en lea p ;

that th ere is b u t a S light t end ency to rev ert to the o ld f o rm ,

b u t that if rev ers io n tak es p lac e at all it is c o m p let e at o nc e an d

the ret u rn is t o t he o ld type and n o t t o an int erm ed iat e f o r m .

T he m u tant is d is t inct ly a ca se o f d is c o nt in u ity .

SECT I ON II — EXA M PLES OF M U TAT I ON

T he c lass ic exa mp les o f m u tat io n are the weep ing willo w and
t he n ectari n e T h ey a re t o b e regard ed h o wev er a s fa m ilia r
.
, ,

illu s t rat io n s o f genera l princ ip les wid ely o pe ra t iv e and g iv ing

rise no t to few b u t to ma ny d is t inct types .

Wh en a seed germ inat es it p u t s ou t two s pro u t s On e is .

posit iv ely geo tro p ic that is it res po nd s to t he f o rc e o f gra v ity

,

and gro ws d o wn wa rd int o t he so il d ev elop ing the roo t sy s t em , .

T he o th er is n egat iv ely geo tro p ic ; that is it gro ws u p ward ,

1 T he s t u den t o f ev o u tio n s l
h ld g i th ou p t i
a n t h t t h t
e c on c e
yp on a e e o f a rac e

is no t afi xe p d i t f
o n m w h i h
ro d i t i dc i t ev a
; i t i
ons rat h t h
a e s ra er e c en t er o f

gra vi t y fo llat h i d ie id nl v f t h ua s O it t l
e ra c e, t i s d
exacp di g oc a on e en n u po n t he
e t
x en t d di
an ti
rec f i d i id
on o l d i ti
n v h ifti g ligh tl y f m t im
ua ev a on s , s n s ro e t o t im e
wit h t h th t i fl
e c a u s es a n i b il ity
u en c e v a r a .
1 12 V A R I A T I ON

again s t grav ity and with an energy s u ffi c ient t o maint ain it in

,

a fa irly u pright po s it io n d ev elo p ing s t em s b ranch es and leav es , , ,

.

Occa s io na lly t his latt er geo tro p is m fa ils and t he branch es ,

hang d o wn ward fo r m ing a weep ing v ariety T his is es pe

,
.

c ia lly c o mmo n in t he willo w and t he b irch t h o u gh by n o m ea n s ,

u nkn o wn in o t her trees n o tab ly the elm m ap le and beech , , ,

.

C u t lea v ed v ariet ies and fan to p s o ccas io nally ari s e

- 1 “ ”

s u dd en ly a ll o f wh ich m ay b e pres e rv ed by g raft i ng o r by b u d

,

d ing S O that with pro pe r att en t io n we m ay h a v e weep ing c u t

, ,

lea v ed and fan to p v ariet ies n o t of a few b u t o f m any Spec ies

,
-
,

o f tr ees and s hr u b s a lth o u gh t he read i n es s with wh ic h a part ie

,

u lar m u tat io n may appear in o n e s p ec ies is n o g u aranty o f it s

appea ranc e in ano th er .

A tree wh ich has alway s b ef o re b o rn e peach es m ay s u dd en ly

b ear n ectari nes o r mo re lik ely a s ingle b ranch may m ake the
,

d epart u re t he rem aind er Of t he tree c o nt in u ing to bear peach es

,

as befo re I n any ev ent t he m u tat io n m ay b e pro pagat ed by b u d

.

o r po s s ib ly by s eed — in wh ich latt er ca s e a n ectari n e b earing

,
-

tree res u lts T h is tree may bear nectarin es all its life o r it may
.
,

o cc a s io na lly b ear peach es o n t he wh o le o r a p o rt io n o f its t o p .

T he S ign ificant f a ct is that th ere is n o int ermed iat e b et ween t he

p each and t he n ectarine a n d yet t he o n e may arise at any t ime ,

fro m t he o th er T he m u tat io n fro m peach t o n ectarin e is c lear

?

c u t and d is t i nct and the rev ers io n fro m n ectarine to peach

,

wh en it occ u rs is eq u a lly c o m p let e .

T he apr ic o t a ppears to b e relat ed t o t he p lu m m u ch a s

the n ecta ri n e is to t he p each I n bo th ca s es the m a in s t o ck s .

( p e ach e s and p l u m s ) e x is t i n m any v ar ie t ies and t he m u tat io n s ,

( n e ctar i n es and apr i c o t s ) i n b u t fe w I n t he ca s e o f t h e f o r m e r .

( t he peach ) t he m ain s t o ck is d o wny wh ile the m u tant is gla ,

bro us o r d es t it u te o f d o wny c o v ering I n t he latt er h o wev er

, .
, ,

t he c o nd it io n s a re rev er s ed fo r t he m a i n s t o ck ( t he p lu m ) is ,

glabro u s while it is the m u tant that is d o wn y

, .

1 A f t p t
an -
i o i wh i h th b
ree s one h n b pp
c i t id e f t ra n c es a re o rn e o n o os es es , a er

th f h i
e as f on o I m ll f
c o rn . t pl t ti
n a s b l gi g t t h w it
a o res i f an a on e on n o e r er s a an

t p li d
o n wg w t
en , n o id ro b l p p ti
n o co ns era e ro or o ns .

1 I m h
n as th p
uc hi as id
e d th m i
ea c s thc o ns t i i
ere id as e a n ra c e, e n ec a r n e s sa

to i
ar se f m th p ro h b y m t tie Th
eac f wh p hu a on.b p ere o re en ea c es a re o rn e u on

t i
n ec a r n e t th rees i id dt b
e c ase f
s c ons i ere o e on e o rev ers o n .
 V A R I AT I O N
1 1 4

I n t he a me genera l mann er c o lo r is lik ely to fail and no th in g

s , ,

is m o re c o m m o n in nat u re than a lb in o v a r iet ies T h u s we h a v e .

o u r wh it e c u rrant s tra wb erry ras pberry and ev en t he black

, , ,

b er ry — a lmo s t ev ery t hick et a ffo rd ing its exa m ples a n d s peak

,

in g elo q u ent ly o f the freed o m with wh ich nat u re creat es n ew

fo r m s and if we will o n ly o pen o u r eyes to see what is going o n
,

abo u t u s we S ha ll learn m u c h o f ho w it is d o n e
,
.

A lb in is m a mo ng an im a ls is ev en m o re c o m m o n than a mo ng
p la n t S M en nd o gs cat s h o,
r s es c a tt le S h eep
,
b ear s rabb
, i t s , , , , ,

rat s m ic e and m any o th er s pec ies a re d is t ing u is hed by a lb i n o

, ,

v ariet ies .

T h es e d is t inct io n s m ark ed th o u gh th ey a re aris e d o u bt les s

, ,

fro m t he s im p les t cau s es F o r exa m p le if an an imal fo r any .

,

reas o n fails to s ecret e p ig m ent in t he n o r mal m ann er it is fro m

n ec es s ity a n a lbin o and if t he failu re is h ered itary a n a lbin o
,

ra c e is lik ely t o b e es tab lis h ed alth o u gh u nres tricted breed ing ,

great ly red u c es it s pro bab ility thro u gh cro s s ing with o th er fo r m s .

S ECT I O N I I I EX PER I M E NTS O F D E V R I ES 1

H u go D e V ries pro fes s o r o f bo tany in t he U n iv e rs ity of

,

A m s t erda m lo ng ago b eca me c o n v inc ed th a t D ar win s th eo ry

’

o f t he o r ig in o f s pec ies thro u g h t he g rad u al acc u m u lat io n o f

f o r t u it o u s v ariat io n s is n o t t he o n ly m ean s o f creat ing n ew types .

D a r win ta u ght n o t o n ly that ex is t ing types had been pres e rv ed

by s elect io n beca u s e t h ey in s o me way fitt ed the c o nd it io n s Of
life b u t that t he int er v en ing s pac es b et ween S pec ies and v ariet ies
,

represent ext inct io n s thro u gh t he agency o f nat u ra l s elect io n .

D e V ries ca m e t o b eliev e tha t in m any ca s es at lea s t t he n ew , ,

type S prings s u dd en ly fro m t he o ld with o u t gradat io n and with o u t ,

int erv en ing f o r m s and that wh ile s elect io n m ay s hap e u p t he

,

n ew typ e and p erhap s t he b ett er fit i t fo r ex is t enc e et t he

y ,

s elect iv e pro c es s is in no way res p o n s ib le fo r it s o r ig in I nd eed .

,

o n e o f t he ear lies t ev id enc es t o his m ind t hat n ew types o ft en , ,

arise with o u t t he agen cy o f s elect io n was t he n o tab le fact that ,

n ew f o r m s ar is ing s p o ntan eo u s ly in nat u re a re fo r t he m o s t

1 H g D V i Sp i d V i ti th i O igi b y M t t i O C
u o e r es , ec es a n
[ p t
ar e es , e r r n u a on en o ur

P b l i h i g C mp y C h i g ]
u s n o an , ca o .
 M UT AT I ONS 1 5

part pro mpt ly ext er m inat ed by t he rigo rs o f nat u ral s elect io n ,

wh ich t herefo re c o u ld n o t ha v e been t he C h ief agency in th ei r

c rea t io n .

A cc o rd ing ly he c o nc eiv ed t he id ea o f c u lt iv ati ng a few u n s table

f o rm s u nd er c o nd itio n s s u ch as wo u ld pro tect and pres erv e any
m u tat io n s that m ight arise h o p ing in th is way t o thro w s ome ,

light o n t he o rigin o f n ew types and t o d et erm in e wheth er in the

o ri gi n o f s pec ies nat u ra l s elect io n wo rk s pr i nc ip ally u po n ind i

v id u a ls o r u po n types .

Ex peri men t s wit h t oadflax ( Linaria v u l garis ) T h ese experi

1
.

m ent s were d es ign ed t o t es t t he o ri gin o f t he pelo ric fo rm 2

The .

t o ad fla x was ch o s en firs t b eca u s e t he pelo ric fo r m is kn o wn to

,

hav e arisen repeated ly and s ec o nd beca u se the change inv o lv ed ,

is S light s tru ct u rally s peak i ng T h ese t wo c o n s id erat io n s ga v e

, .

reas o n fo r the h o pe t hat if t he s pec ies were p u t u nd er carefu l

Ob s erv at io n and c o ntro l he ( D e V ries ) m ight b e present at the ,

t ime wh en nat u re prod u c es an o th er o f th ese rare changes .

”

T he exper iment s c o mm enc ed in 1 88 6 with n o r ma l p lant s bear

in g on e o r t wo pelo r ic fl owers
“
a s is c o mm o n wi th m o s t i nd i
”

v id u a ls o f th is g en u s T he roo t s were p lant ed in t he ga rd en

.
,

and fl o wered and seed ed in 1 8 8 7 T h is sec o nd gen erat io n was .

grown fo r three years pro d u c ing in 1 8 8 9 0ne and in 1 8 90 t wo

3
, , ,

pelo r ic s tr uct u res T he s eed s o f th es e were s a v ed and prod u c ed

.

t he th ird gen erat io n in 1 8 90 — 1 89 1 A mo ng s o m e th o u s and s of .

blo ssom s in th is generat io n th ere wa s o n e fiv e s p u rred fl o wer -

.

T h is was po llinat ed by hand and lu ck ily pro d u c ed “

abu ndant
fru it with eno u gh seed s fo r t he ent ire c u lt u re o f 1 8 92 and they
, ,

o n ly were s o wn
4 ”
.

1 D e V ries , S p ec ies a n d V a rieties , et c , pp 4 64 48 7 . .

-
.

9 T he n o rm a lfl
o wers o f t h e t o a d fla x a re exc ee in g d ly uns ym m et rica l . A si de
f ro m b ea ri n g a s h t or s p u r, t h y e are d es c rib e d as c o n s is t in g o f a l
t wo - ipp e d c o ro lla ,

t h e lo wer l ip s d i g d t h l b d wi t h b
s p rea n an l g
ree- o t
e , a as e s o en a r e d as o n ea r ly
l
c o s e t he th ro a t Pl
.
”
t b i g
an s h ym m t i l fl w
ear n s uc d t dfluns e r ca o ers a s o oa a x, s n ap

d ra go n , etc .
, k w
a re no i nlly t p d
oc c as o n ap l i t h t i y mom ro uce e or c, a s, s et ric a l ,

flo wers N o . t ly t h t b t p l i
on a , i ti
u t k w
e o r c v ar ed th es a re n o un no n , an es e exp eri

m en ts were d ig d t l t h m
es ne o so v e f th i igi e an n er O e r or n .

3 T h e to a dfl iax bi s a i l en n a .

4 Pelo ric fl w o f th i
ers O
p i smm l ys t i l
ec es b t i y
a re c o on s er e, u n an c as e a re

d ep en d en t u p t
on ar ifi i l f t il i t
c ai T h ery b y t e
za i ll
on d p.t d t p e a re na ur a a e o res erv e

t h ems elv es .
 I 1 6 V A R I AT I ON

U p to th is po int
xper im ent each gen erat io n req u ired
in t he e

t wo y ear s a s t he to a dfla x is a b ienn ia l n o t b lo o m ing u nt il t he

, ,

s ec o nd ear A ft e r th i h o we v er t h e s eed ling s wer e s tart ed

y s .
, ,

u nd er g la s s and t ran s p lant ed t o t he gard en in J u n e By th is .

mean s t he n ew p lant s were m ad e to pro d u c e fl o wers and s eed s

t he fi rs t y ear .

A b o u t t wenty p lant s o f th is ( fo u rth ) gen erat io n were s ec u red ,

and u n d er th is treat m ent m os t o f th em pro d u c ed s eed t he fi rs t

y ear On ly o n e pelo ric flo wer wa s o b s erv ed h o wev er in t he
.
, ,

ent ire lo t T he p lan t b earing th is flo wer and o n e o th er were

.

pres er v ed and all o th ers were d es troyed T h ese t wo fert ilized

,
.

each o th er freely and pr o d u c ed 1 0 cc o f s eed b u t n o m o re .

,

pelo ric fl o wers appeared I t is fro m th is pair o f p lant s ho w .

,

ev er that a p elo r ic rac e fina lly s pr u ng

l
,
.

I n 1 8 94 ab o u t fifty p lant s wer e in flo wer T h ere wa s n o rea .

s o n fo r c o n s id er ing th es e p la n t s any mo re pro m is ing t han p re

v io n s s o wing s exc ept that s tray p elo r ic fl o wer s were o b s erv ed

“
,

in s o m ewhat larger n u m b er s than in prev io u s gen erat io n s ,

elev en p lant s b ear ing o n e o r t wo o r e v en th ree s u ch a b n o r , ,

m a lit ies D e V ries wis ely rem a rk s that th is c o u ld n o t b e “

”
.

c o n s id ered a s a rea l ad v anc e S inc e s u ch p lant s m ay o cc u r in ,

v a ry ing th o u gh o rd i nar ily s m a ll n u m b er s i n ev ery g en erat io n

”
.

Ho wev er bes id es th es e elev en ind iv id u a ls each bear ing o n e

, ,

o r t wo abn o r m a l fl o wer s t lzer e wa s a s i ng le p la n t bea r i ng on ly ,

p e lo r ic
fl o wer s T h e m u tat io n h a d a.r i s e n and D e V ries
“
w a s

pres ent at t he t im e .

T his p lant wa s caref u lly k ept a ll o th er s being d es tro y ed and , ,

t he n ext y ear it b lo o m ed a ga in b ea r ing o n ly p elo r ic fl o wer s It , .

wa s tr u e t o i ts typ e I n th is c o nn ect io n D e V ries s a y s .

He e we h e the fi t
r p i m e avt l m t t i f m l i t
rs ex p leri c n a u a on o a nor a n o a e or

rac e . T wo fa c ts were c lea r an d si m ple : [fi rs t] the a n c es try was k n own

1 I t has b een sa i d th at t he fl o wers o f o n e p l a n t a re s t e ri e l to p o ll en f ro m t h e
sa m e p l an t . D e V ries a s c e rta in e d by c a ref u l ex p e rim en t t h at t hi s is t r u e in ab o u t

50 p er c en t o f t h e c a s es , s o t h a t , th o u g h m h h igh
a g uc f f t i l it y i t er d e ree o er ex s s

b et ween in iv i d d l th ua s an with i n t he i d i id
n l b l t b
v ua , b tw
a so u e ll a rren n es s e een a

fl o wers o f th m pl
e sa e a n t c an n o t be t d
a s s er e Th p i t i. t ig ifi e t i th
o n s no s n c an n e

p res en t c o n n ec t io n , b u t it is im p o r tan t d m t ti g t h t f t il ity d t il it y

as e o n s ra n a er an s er

a re n o t a lw y a s in di rec t p p
ro o r t io n to c o n sa ng i i t y d t h t t h
u n g h l , b
an d a , ou c o se ree

i ng m a y b e c o m m o n in fert i e in ly l t i t i
c er a n s ra n s , it b y m f l l w t h t it i no ea n s o o s a s

l y
a wa s in fer t i e ev e n in t he sa m el s t ra in .
l I 8 V A R I AT I ON

New t ype persi s t en t N ext he u n d ert o o k to

. d et er mine to what
ext ent th es e m u t ant p elo ri es wo u ld breed tru e in o rd er t o ”

c o m pare t he pro po rt io n with t he prev io u s ly as c ertained 1 per

c ent I n th is he enco u nt ered d iffi c u lty beca u s e o f t he h igh
.

d egree o f s terility o f pelo ric fl o wers He had in a ll s o me t wenty .

plant s and po llin at ed art ific ia lly m o re than a th o u s and fl o wer s

,
.

Of th es e he s ay s

N ot a S in gle o n e a ve a n o rm al fru it, b u t s o m e s m all a n d ru d im en ta r

g y
c a ps u les were p rod u c ed b ea ri n g a few seed s F ro m t h es e I had 1 1 9 flo wer .

in g pla n ts , o u t o f wh ic h 1 06 were pelo ri c a n d the rem a in d er ( I 3 ) o n e s p u rred -

.

T he grea t m a j o rit y ( s o m e 90 per c en t ) were t h us S h o wn to b e tr u e to th eir

n ew ty e W h et h e t he 1 0 per c en t rev er t ing o n es were tru l a ta v is ts c a u s ed
p . r y
b y s tra y po llen grain s fro m a n o t h er c ul tur e c an n o t o f c o ur s e b e d eterm in ed
wit h s u ffic ien t c er ti tu d e .

T h is xperi ment d eter min es n o t only the d is t inctnes s o f t he

e

n ew typ e and t he s u dd enn es s Of it s f o r m at io n b u t it s es s ent ia l ,

p u rity as well fo r it bred tr u e in 90 per c ent o f t he cas es wh ile ,

t he pro bab ili ty o f o r ig ina l m u tat io n wa s S light c erta in ly n o t O v e r ,

1 pe r c ent .

T he t o t al lack o f int er m ed iat e s t ep s in t he c o ntro l experi

ment s is s ign i ficant T h ei r a b s enc e in nat u re is n o t les s s o ( fo r
.

if th ey were pres ent a s tran s it io n s t ep s t o ward t he f o r m at io n o f

pelo ric rac es th ey wo u ld c erta in ly b e d is c o v ered part ic u lar ly
, ,

wh en we rem em b er th a t t he s pec ies is a p erenn ial ) and t he c o n ,

c lu s io n s eem s in ev itab le that t he tran s it io n is abru pt and t he ,

n ew typ e rep eat ed ly re f o r m ed is with o u t d o u bt t o b e regard ed

,
-
,

as a tr u e m u tat io n .

The c o m m o n s n a pd rago n wh os e fl o wers a re exc eed ing ly u n

,

s y m m etr ica l a lso ha s a p elo ric rac e B u t t he s napd rago n is “

.
,

s elf f ert ile and so is it s p elo ric v a r iety

-
,
o b s e rv es D e V ries , .

Th es e m u tat io n s a re th erefo re m u ch m o re ea s ily pres erv ed and ,

a re as we s h o u ld exp ect m o re c o m m o n than i n t he t o ad fla x

, , ,

s o c o m m o n and s o d is t inct a s with o u t a d o u bt to giv e ris e t o

, ,

real hybr id s with t he o ld fo rm .

What is tr u e of toad fla x and s napdrago n is h eld t o b e tru e o f

u n s y mmetrical fl o wer s g en era lly ; na m ely a s tro ng t end ency t o ,

g iv e r is e fro m t ime t o t im e t o p elo ric v ariet ies n o t by grad u al ,

change o f t he parent s t o ck b u t by s u dd en o ffs et o r m u tation , .

 MUT AT I O N S 1 19

Ex perimen t s t ion of dou b le flowers A ft er

in t he p r od u c .
1

rem ark ing that mu tat io n s o cc u r a s o ft en amo ng c u lt iv at ed as

a m o ng wild p lant s D e V ries dro p s t he c au t io n th at in all experi
,

m e n tat io n o f th is o rd er hybri d is m m u s t b e caref u lly g u ard ed

aga in s t He o b serv es too that wh it e v a riet ies seem c o m pa ra
2
.
, ,

t i v e ly o ld a s th ey a re c o m mo n in t he wild s tat e wh ile d o u b le

, ,

fl o wers a re ra re in t he wild s tat e and c o rres po nd ingly rec ent ,

in d icat ing th eir o rigi n u nd er c u lt iv at io n and th u s mak ing t he ,

m a tt er o f d o u b ling a fa v o rab le charact er with wh ich to c o nd u ct

i n v es t iga t io n s u po n m u tat io n .

I n t he exp eri m ent s u p o n pelo ric t o ad fla x n o th ing n ew wa s

att em pt ed The o bj ect was to repeat what nat u re was kno wn
.

o ft en t o ha v e d o n e b u t s o to c o ntro l c o nd it io n s as t o
,
b e th ere
wh en it happened n ext t ime .

I n th is exp er iment h o wev er D e V ries d et er m in ed t o att em pt

, ,

a n ew m u tat io n that is to try to s ec u re d o u b le fl o wers wh ere

, ,

th ey had n ev er been o b s erv ed in nat u re He acc o rd ingly ch o s e .

t he c o rn m arigo ld ( Clu y s a n tlzem u m s eg etu m ) c o mm o n in t he ,

gra in fi eld s o f c entra l Eu ro pe and it s c u lt iv at ed v a riety g r a n d i , ,

fl o rum T h e .n u m ber o f ray fl o ret s is v ar iab le in b o th b u t is , ,

o n an a v erage th i rt een in t he wild and t wenty o n e in t he c u lt i

,
-

v at ed T h is ind icat ed t he latt er a s t he m o re fav o ra b le fo r t he

.

expe ri m ent and it was th erefo re ch o s en ; b u t i t is far fro m p u re

, ,

fo r m any o f it s h ead s ha v e a s few a s th irt een ray s On ly s ix .

o u t o f t he fi rs t lo t o f three h u ndred p lant s reach ed an a v erage

o f t wenty o n e a n d th es e were s elect ed a s t he f o u ndat io n

-
,
.

T he s eed s o f each o f th es e were s o wn s epa rately F iv e ga v e .

proo f o f being s t ill m ixt u res with t he wild fo rm and were re

je c ted T h.e o ffs pr ing o f the s ixth p lant av e raged t wenty o n e -

ray fl o ret s and aft er c o u nt ing s o me fifteen h u ndred h ead s the

,

t wo p lant s were s elect ed wh o s e s econdary h ead s m ad e t he b es t

s h o wi ng T he pro geny o f th es e p lant s a lso a v eraged t wenty o n e
.
-
,

1 D e V ries , S pec ies a n d V ariet ies , etc 8

pp 4 9 5 5
1 -
.
, . .

3 I f a n ew fo rm is a m u t a n t it wi l l b ree dt ru e to l
it s e f in t he g rea t ma j o rit y
o f c as es a n d hyb id i p erfo rc e r ze wi th t he o rigi n a l s to c k . I f, on t he o t h h d
er an , it
is an di y hyb id it wil l
or n ar r , not d t
b ree ru e, b ut will o b s erv e t he p i ip
r nc l e of

M d l l w ( t b di
en e s
’
dl t
a o e sc u ss e a er , b
) y wh i h c a c ert a n i d efi n it e p e rc e n t a e is o f t h e
g
origi l typ
na Th it i mp es . us s co a ra ti ly y t
ve eas o as c ert a in w h the er a n e y
w t p e is t he
p od u c t o f a s in gle rac e
r b y m u t at io n o r of two rac es by hyb ridiza t io n .
I 20 V AR IAT I O N

and D e V ries
c o n s id ered that t he s train was n o w p u re and that
n o f u rth er s elect io n c o u ld b e o f a n y a v a il .

On e o f th es e t wo p lant s wa s d is t ing u is h ed by pro d u c ing t wo

s ec o ndary h ead s with t wen ty t wo ray s wh erea s g en erally o n ly -
,

t he t er m ina l h ead r ea ch ed so m any as t wen ty on e t he o th er -

,

retro grad ing o ft en as lo w as to th irt een T h is exc ept io nal p lant .

was d is t ing u is h ed o n ly by th es e t wo s ec o nd a ry h ead s I t s ter .

m ina l h ead had b u t t wenty o n e ray s and t he a v erag e o f all it s -

,

h ead s was n o t exc ept io na lly h igh ; b u t n o o th er p lant o u t o f

h u ndred s had ev er pro d u c ed s ec o nd a ry h ead s with m o re th a n
t wenty o n e ray s a n d it was fro m th is p lant that the d o u ble
-
,

fl o weri ng lin e d ev elo p ed three y ears la ter .

T h is p la nt a ppeared in 1 8 96 I t s s eed was so wn in 1 8 97 . .

T he la rg es t n u m b e r o f ray s in t he t er m ina l h ead s u dd en ly

i ncreas ed fro m t wenty o n e to th i rty fo u r ; n ext y ear -
to -

fo rt ye ight ; n ext to S ixty s i x ; a n d d u ri ng th is t i m e t he -

general a v erage fo r all t he h ead s increased remarkab ly N o .

i nd icat io n o f d o u b ling ha d h o wev er yet appeared T he im

, ,
.

pro v ement wa s s u ch a s fo llo ws select iv e breed ing with flu c tu at

i g v i ii
n ar ab l ty — im pro v emen t by grad u a l change and with o u t
,

m u tat io n s .

L at e in the s ea so n ( S ept em ber ) o f th is y ear ho w

ev er three s ec o ndary h ead s appeared o n o n e p lant with a few
,

ray fl o ret s s cattered o v er the d is k T he m u tat io n anx io u s ly .

a wai t ed fo r s ev en y ears had s u dd en ly appeared in th is s m all ,

belat ed way t o wa rd the c lo se o f the gro wing seaso n and in ,

a m ann er that wo u ld ha v e es c a ped t he a ttent io n o f any b u t

t he m o s t pa in s tak ing i n v es t iga t o r and that wo u ld ha v e in v i ted
1
,

ext er m i nat io n in nat u re .

T h is wa s in 1 8 99 T he h ead s were o f c o u rs e p o llinat ed with

.

o th er and i nf er io r flo wer s b u t i n 1 900 t he h igh es t n u m b er o f

,

ray s ro se to o n e h u nd red and in 1 90 1 reach ed two h u ndred , .

He rem ark s S u ch h ead s a re a s c o m p let ely d o u ble as a re

,
“

1
Th t d t will
e s u t th t
en y fl w f th no e d f pl t w
a ev er f lly o er o o u s an s O an s as ca re u

exa mi d ne d th t i
, an y th fa d ti f t h m t ti w i
n e v er c as e e i oun a on o e u a on as n an ncon

s p i c uo u s
p l t t i tan b , l k
c er a n d b y o l b T
e o v er o oh b i l e c as u a o s erv ers . e o v o us ess o n

i th t
s a ly th m t
on f l d y t m ti
e os mi ti
c a re u wil l d t t t h f
an s d
s e a c ex a na on e ec e o un a

ti t
on s oc k il y d
,
it
s o ea s p t i i t h
o es
g l m
es c a d
e no di ly i i t
ce n e en era ass an s o rea s

l t
os l i lt d dp t t d
u n es s so a e an ro ec e .
I 22 V A R I AT I O N

happen ed rath er than to ent er j u s t aft erward and o n ly in tim e

,

t o n o t e res u lts wi th n o ev id enc e a s t o meth od s .

Of all t he h u ndred s o f p la nt s c u lt iv at ed he f o u nd t he ev en ing

prim ro se m o s t fert ile in d is t inct s train s bo th in the wild and,

in t he c u lt iv at ed s tat e . Oth er s pec ies ga v e ris e t o v ariet ies

freely b u t n o o th er appeared t o b e s u ffic ient ly m u tab le t o giv e
,

rise freely to what he rega rd ed a s elementa ry s pec ies .

Of all t he prim ro s es Gi no tlrera L a m a rck ia na c o m mo n ly ca lled

, ,

L a m arck s ev en ing pr im ro s e was t he m o s t pro lific in d is t inct

’
,

fo r m s and acc o rd ingly th is was ch o s en by D e V ries fo r s pec ia l

,

att ent io n in his experim ent s I t is d es cribed as a s tat ely

.
“
.

plant with a s t o u t s t em att a in ing o ften a h eight o f

,
m or .

mo re Wh en n o t cro wd ed the m a in s t em is s u rro u nd ed by a

.
,

large c irc le o f s m a ller branch es gro win g u p ward fro m i t s b a s e

s o as t o f o r m a d en s e b u s h T h es e branch es in th eir t u rn hav e
.

n u mero u s lat era l branch es .C o ntrary t o th ei r c o ngeners ,

th ey a re d ep end ent o n v is it ing in s ect s fo r po llinat io n .

Ord inar ily th is p ri m ro se is a b ienn ia l pro d u c ing ro s ett es in

,

the fi rs t year and s t em s in t he s ec o nd y ea r Bo th t he ro s ett es

.

and t he s t em s a re h igh ly v ariab le in na t u re pro d u c ing a n u m ,

b er of d is t inct rac es s o me o f wh ich S h o w a m ark ed ab ility to

,

h old th ei r o wn u nd er nat u ral s u rro u nd ings wh ile o thers a re too ,

wea k t o end u re .

M any o f th es e D e V ries regard ed as n ew s pec ies E xperi .

m ent s t o d et er m in e th is p o int were c o m m en c ed with s t o ck d is

c o v ered n ear Hilv ers u m a n d three p la n s were fo llo wed fi rs t to
, ,

tran s plant t he apparent n ew s pec ies int o t he gard en wh enev er

t he n ew rac e wa s s u ffi c ient ly s tro ng ; s ec o nd t o r eprod u c e weak ,

rac es by s o wing s eed s fro m ind ifferent p lant s gro wing wild ;
“ ”

th ird t o s o w the s eed s fro m t he introd u c ed plant s

,
T h es e .
“

v ario u s m eth o d s he add s ha v e led to the d is c o v ery o f o v er

“
, ,

a d ozen n ew t ypes n ev er prev io u s ly ob s er v ed o r d es c ribed .

T h es e n ew p lant s a re d iv is ible a cc o rd ing to D e V ries int o

, ,

fiv e d iff erent h ea d s o r gro u p s ( 1 ) th o s e that a re ev id ent ly

”

t o b e c o n s id ered a s v a r iet ies i n t he na r r o wer s en s e o f t he

wo rd ,
represent ing retro grad e d ev elo p m en t ; ( 2 ) progres s iv e
elem entary s p ec ies wh ich a re s tro ng the par ent
“
as as

s p ec ies pr gr iv e elem entary s p ec ies w ak r t

( 3) b h an
”
o es s u t e e ,
 MUTAT I ON S 1 2 3

the parent an d apparent ly n o t d es t ined to b e s u cc es s f u l

,
“

(4 ) c e rta in f o r m s that a re
“
o rg a n ica lly i nc o m p le t e ; ( 5) in
” “

c o n s tant f o r m s
”
.

Grou p ( I ) retro grad e v ariet ies Of th is c lass t he fo llo wing

,
.

three fo rms were d is c o v ered a ll prod u c ed in nat u re as well as ,

i n t he gard en
0 le v ifolia t he s moo th lea v ed v ariety c o n s tant fro m s eed and
.
,
-
,

n ev er r ev ert ing exc ept fro m cro ss ing A S s t ro ng and fert ile as .

the paren t .

0 b rev is ty lis t he s h o rt s ty led fo rm

.
, I n th is t he o v ary is so
-
.

p lac ed that it is reach ed by v ery few po llen t u bes Th u s wh ile .

t he p lant is v igo rou s i t is b u t ind ifferent ly pro d u ct iv e o f s eed s ,

and as D e V ries s ay s ma ny [caps u les ]c o nta in ed no seed s at

all f ro m o th ers I ha v e s u cc eed ed in s a v ing o n ly a h u ndred seed s
from thou sand s o f caps u les T h es e s eed s h o wev er reprod u c e
.
, ,

t he v a riety with o u t rev ers io n s t o L a m a reb i a n a .

0 n a n el la the d warf
.
,
“
a m o s t attra ct iv e litt le p lant
, ,

v ery s h o rt o f s tat u re r each i ng o ft en a h eight o f o n ly 2 0 3 0 c m

,
— .
,

o r les s th a n o n e f o u rth o f that o f t he parent The fl o wers .

a re as large as th os e o f t he parent ; t he lea v es a re m u ch

s m aller and with n o rev ers io n in s eed li ng s ev en in rep eat ed ,

and s u cc ess iv e generat io ns .

G rou p pro gress iv e elementa ry s pec ies and v igo ro u s ; two ,

fo r ms d is c o v ered
0 g iga s t he giant d es erv ing it s na m e n o t fro m being h igh er
.
, ,

than it s parent b u t becau se it is s o m u ch s t o u t er in all

,
“

res p ect s . The s t em s a re o f t en t wic e as th ick as in t he parent

L a ma r ck ia n a ) and t he int rn d h rt r and h l a
( , e o e s a re s o e t e e v es

mo re n u m ero u s c o v ering t he s tems with a d en s er f o liage

,
T he .

fl o wer s a re larger and t he seed c a ps u les a re s maller and fi lled

,

wi th fewer b u t larger s eed s t han in t he parent plant I t has a .

s tro ng t end ency t o rem a in a b ienn ia l .

0 r u b r i n er v is the red v eined fo r m I n th is t he v ein s o f t he

.
,
-
.

lea v es a re d is t inct ly t inged with r ed and the fr u it s a re s treak ed

wi th red T he p lant s a re in m a n y way s a c o u nterpa rt o f the
.

giant exc ept fo r t he red t in ge and d is t inct ly ligh ter fo liage

, .

T h is latt er pro bab ly acc o u nt s fo r t he m ark ed t end ency on t he

part o f thi s fo r m to bec o me an ann u al L ik e t he giant th is f o r m .
,
.
I 24 V A R IAT I O N

is tr u e t o type wh en gro wn fro m t he s eed and it s rec u rrenc e is ,

far m o re c o m m o n than is t hat o f g ig a s wh ich is extrem ely rare , .

Gro u p progress iv e elementary s pec ies b u t weak ly Two ,

.

fo rm s :
0 a lb i d a t he a lb in o with wh it is h narro w leav es
.
, ,
appar , ,
“

ent ly i ncapab le o f pro d u c in g s u ffic ient q u ant it ies o f o rga n ic

foo d .T he s eed ling s a re exc eed ingly d elicat e and if left t o ,

th em s elv es will b e s peed ily o v ergro wn by th eir mo re v igo ro us

n eighbo r s ; b u t if tran s p lanted and giv en the bes t of care th ey ,

m ak e fa irly v igo ro u s p la nt s t he s ec o nd y ear c o m pari ng fa irly ,

well with t he parent s t ock b u t bear in g fewer s eed s T h ey .

c o me tru e ev en to t he th ird generat io n and the type remai n s

d is t inct .

0 ob long a t he narro w lea v ed fo r m I t

.
,
m ay b e gro wn eith er-
.

a s an ann u a l o r a b ienn ia l I n t he fi rs t ca s e it is v ery S lend er .

and weak bearing o n ly s m all fru it s and few s eed s I n the a lt er

,
.

na t iv e ca se h o wev er ( b ienn ia l) it bec o m es d en s ely branch ed

, , ,

bea ring flo wers o n q u it e a n u m ber o f rac em es an d y ield ing a

f u ll harv es t o f s eed s .
”

T he in v es t igat o r s ay s
W h e w gi
e av the d
no i p ti f
v en w f m wh i h d i es c r
ge on o sev en ne or s c v er

i d iff e t w y f m th p
n er n a t ty p s A ll w
ro b l t ly
e t
a ren t f m e e re a so u e c o ns an ro
'

s ee d . H d ed un th
r d f s d li g m y h
or o us a n i b t th ys o s ee n s a av e a r s en , u e

a lw y a me t
s co d e e t t th
ru e an ig i l 0 L m
n ev bi
r r v er o e or na . a a rc ana .

He add s t he rem ark that th ey ha v e inh erit ed t he c o nd it io n o f

m u tab ility t o s o m e ext ent and a re ev id ent ly th em s elv es ab le t o
pro d u c e n ew fo r m s b u t that th ey d o so b u t rarely
, .

T wo o th er f o r m s b elo ng t o t h is gro u p 0 s em i la ta and ,

.

0 lep ca p a
. to r — b u t th eir charact ers d o n o t m eri t S pec ial
,

d es c ript io n .

Gro u p f o rm s o rgan ically inc o m plet e °

0 la ta is a p is t illat e v ariety w h o lly d ep end ent fo r fert iliza

.
,

t io n u p o n o th er f o r m s and it had therefo re n o o ppo rt u n ity to

,

es tab lis h it s type wh ich h o wev er freely a ppeared I t is a lo w

“
,
.
, ,

p lant b u t with d en s e f o liage and lu x u r iant gro wth

,
“
Its .

pres enc e can b e d et ect ed in t he s eed ling by t he broad S in u at e “

,

leav es with ro u nd ed t ip s B eing p is t illat e it pro d u c es s eed .

,

o n ly wh en cros s p o llinat ed in wh ich ca se it s charact ers a re ,

1 26 V AR I AT I O N

p la nt s The types that aro s e at d ifferent t imes ha v e already

.

been d es cribed b u t c o n s id erab le int eres t and n o litt le pro fi t

,

attach es to the d etails o f t he experiment es pec ia lly in re ,

g a rd t o t he o rd er and m ann er o f t he appearanc e of t he n ew

types T he fo llo wing is abs tract ed fro m the experim ent er s

.
’

acc o u nt o f o n e o f th es e fo u r exper im ent s ru n n i ng thro u gh ,

eight gen era t io n s

1
.

B eg inn ing in t he f a ll o f 1 8 8 6 he t o o k n ine large ro s ett es o f

0 L a ma r c k ia n a fro m t he fi eld and p la nt ed th em in t he g a rd en
. .

T he s ec o nd generat io n wa s s o wn in 1 8 8 8 a n d fl o wered in 1 8 8 9 .

The s eed pro d u c ed fift een th o u s and s eed lings o f wh ich ten ,

were d iv ergent at o nc e fi v e la ta and fi v e u a n el la N o int er

,
.

m ed iat es appea red T h ey ca m e int o ex is t enc e at o nc e

“
s ay s
”
.
,

D e V r ies f u lly eq u ipped with o u t prepara t io n o r int erm ed iat e

,
“
,

s t ep s N o s eries o f gen erat io n s n o s elect io n n o s tr u gg le fo r

.
, ,

s elect io n wa s n eed ed I t was a s u dd en leap in t o an o th er typ e

.
,

— a s po rt in t he bes t acc eptat io n o f t he wo rd 2 .

”

T he th ird gen erat io n o f ten th o u s and p lant s s h o wed three la ta

and three no n e/la bes id es o n e r u b r in er v is ,
.

Gro wing expert in d et ect ing m u tant s at an early s tage he ,

d is c o v ered 3 3 4 y o u ng p lant s o u t o f o f t he f o u rth g en er

at io n T h is is ab o u t 2 5 per c ent Of th es e 1 7 6 were . .

o b lon 73 la ta 6 0 n a n ella 1 5 a lb i da 8 m b r i n er v is , 1 s c i n t il
g a , , , ,

la u s ,and 1 g zga s .

The larger n u m b er a n d wid er range o f m u tant s d is c o v ered

th is y ea r a re to b e as cribed to gro wing s k ill in d et ect ing th em
at an ea rly age M an if es t ly s u ch im m en s e n u m bers m u s t b e
.

great ly red u c ed at t he earliest p os s ib le dat e and with o u t d o u bt ,

s o m e g o od f o r m s wer e o v er lo o k ed in t he ear lier gen erat io n s .

A ft er th is ( fo u rth ) gen era t io n t he n u m b er o f s eed ling s wa s

great ly red u c ed with t he effect o f r ed u c ing the n u m ber o f
,

m u tant s and a lso t he chanc es o f t he rarer f o r m s app earing at

all ind eed g iga s n ev er appeared aga in and s c i n ti lla ns n o t aft er
,
-
,

1 D e V ries S p ec ies a n d V a ri et ies etc , p p 5 4 9 5 5 6 -

, , . . .

2
d
I t ma y o c c u r t o t he s t u en t t o o b ec t t o t h e c o n c j l u s io n o n t he g ro u n d th at

t he p a ren t s to c ktk
a en fro m t he fi e ld m a
y not l h
it s e f av e b een p u re I f, o wev er, . h
th e s to c k ha d b een in an
y s en s e hyb id r , t he d epa rt u res s h o ul dh
a v e b een , ac c o r d
in g t o M en d l
e s
’
law, m o re t h an t en ; but not h in t is or in la t er gen era t io n s d id
eit he r p a ren t s t o c k o r m u ta n t b e h av e li k e a hy b id i t h i
r n s res p ec t .
 M UTAT I ON S 12 7

t he s ixth gen eratio n . The ent i re t t he eight gen erat io n s

res u l s of

of breed ing a re giv en in t he fo llo wi ng tab le .

EIG HT G EN E R AT IONS OF A MU TAT IN G STR AIN OF EV EN IN G PR IM ROS E

( 0 L.a m a r ckia n a )

G EN E R A B ub r
r
n er m :
b La
m
tu na
a r ck
N e ru d a L a ta
S c i n ti l

I n t he pin io n o f the experim ent er h ere a re n u m bers en ou gh

O

and types s u ffic ient ly d is t inct to warrant the en u m erat io n of

c erta in la ws o r princ ip les that appear to go v ern t he appeara nc e
o f m u tant s es pec ia lly in t he s pec ies u nd er o b s erv at io n
,
Th is .

D e V ries att em pt s t o d o b u t with o u t pres u ming to say ho w

,

c los ely th ey m a y app ly t o o th er s tra in s o f p lant s o r an im a ls .

Laws of mu tab il i ty f or ev en in g pri mroses . De V ri es ’

ex peri

ments . bas is o f his experim ent s with t he ev en ing prim

On t he
ro s e the in v es t igat o r annou nc es t he fo llowing la ws o f m u tab ility
as app ly ing t o that s p ec ies :
1

1
“
.T hat n ew elementary s pec ies appear s u dd en ly with o u t ,

i nt ermed iat e s t ep s A S proo f he p o int s o u t that n o in terme

.

d iat e fo rm s appeared to fill the gap s and that n o s elect io n was ,

n ec es s ary to es tab lis h the typ e .

2
“
.N ew fo rm s s pring lat era lly fro m t he ma in s t em The .

c u rrent c o nc ept io n c o nc ern ing t he o rigin o f s p ec ies (o r n ew

fo r m s gen era lly ) as s u mes that s pec ies a re s lo wly c o n v ert ed int o
o th er s The c o n v ers io n is as s u m ed t o a ffect all t he ind iv id u a ls
.

in t he s a me d irect io n and in t he s am e d egree T he b irth .

1 D e V ries , S pec ies an d V a ri eties , etc .,

pp . 8
55 5 5
-
7 T
. h es e la ws , w i hl e

an n o u n c e d fo r t he ev en in
g p rim ro s e, a r e wit h out d o u b t o f wi d e if not gen era l
app li
ca tio n .
 I 28 VA R I ATI ON

of a pec ies nec es sarily seemed to in v o lv e t he d eath of the

ne w s

o ld o n e at leas t the o ld m erged int o t he n ew

,
”
.

T he exp er im ent er po int s o u t h o wev er th a t thro u gh all t he , ,

proc es s o f o r igin at ing a d o zen o r mo re d is t inct fo r m s t he parent ,

s t o ck c o nt in u ed u nchanged and s t ill c o n s t it u t ed t he p rinc ipal ,

s trai n o f a ll t he p ri m ros es and fro m th is he d ed u c es the law

l
,

that m u tant s a re lat erals .

3
“
N ew. e le m entary s p e c ie s atta i n th e i r f u ll c o n s tancy at
o nc e C o n s t a ncy is n o t t he resu lt o f s elect io n o r o f im pro v e
ment I t is a q u ality o f it s o wn
. I t can n eith e r b e c o n s tra ined .

by select io n if it is ab s ent fro m t he beg inn ing n o r d oes it need ,

an y nat u ra l o r art ific ia l a id if it is pres ent .

D e V ries rem ark s that sc i n t i lla ns repeat s it s chara ct ers in

b u t part o f it s o ff s pring and t hat he ha s tried t o d eliv er it ,

fro m th is inc o mp let en es s o f h ered ity b u t in v ain T he in s ta .

b ilit y seem s t o b e h ere a s per m an ent a q u a lity as t he s tability

in o th er in s tanc es E v en h ere n o s elect io n ha s b een ad eq u at e
.

t o change the o r ig ina l f o r m He regard s it a s it s elf in a s tat e .

o f in s tab ility .

4 S o
. m e o f t he n ew s t ra i n s a r e e v i d en t ly e le m entary s pec ies ,

wh ile o th ers a re to b e c o n s id ered as v a riet ies

”
.

Elementary s pec ies a re regard ed a s po s s es s ed o f pro gress iv e

cha ract ers b u t v ariet ies a s d iffering fro m th eir parent s t o ck in
,

b u t a S ingle cha ract er and that i n t he way eith er o f an a s su m p ,

t io n o r o f a lo s s The elementary s pec ies is th erefo re a n ew

.
, ,

aggregat io n o f characters wh ile t he v ar iety is s im ply the o ld ,

fo r m m in u s a s ing le charact er Wh eth er th is d is t inct io n h o ld s .

,

rem ain s to b e d et erm ined M u ch o f t he arg u ment t u rn s u po n .

what is t o b e c o n s id ered as a charact er and wh en it is lo s t Fo r .

1 A n a t u ra l c o ro ll y t th i
ar o s o b s e rv a t io n is t o rem ar k u p o n t h e err o n eo u s p o
p u

la r as s u m p t io n th t a f im i l
o s a r a n d c o n t em p o ran eo u s fo rm s t h e m o re p rim i t iv e a re
n ec es s a ri l y t h p g it f th m
e ro en ly o rs o l e h y o re n ea r p erfec t F o r e.x a m p e , i t is as t i l
as s um d t h t if e l t i i t th m m t b th d i t d
a ev o u on s ru e d t f th en an us e e rec es c en an o e

ap e .B t t h u p t h h
e a
g y ld
e, i t i ll
ou
p d h
v er i tod d,i g is t s an a e, a n e s no esc en n n o

an y th i g b t p Th gh id tl y d l p d f m th m igi l t k t
n u a es . ou ev en ev e o e ro e sa e or na s oc a

so m t im e d i e an m w y wh t h n so by e b y m y m t ti
a , e b dy ker w one or an u a o ns n o o no s ,

y tet h g p e b t wa i e id t l
eeny fi d us d t s ev
g wi g len b i g b i d g d
xe an no ro n es s or e n r e

at an
yp i t G
o n d .l t i oo
g d l t
ev o ud f m o n re a rt d b y t i f s re a e or s as c o n n ec e es o con

s an
g i i t
u ny b t wh , t h dui t w het i m er l ik l y
reci d i ,t i
o r,
g t m a s o re e , n rec , ru n n n o so e

ex ti t nc mm co t ly i will t t m p t t
o n a n c es o r, o n
y a no v c e a e o sa .
1 0 V A R I AT I O N
3

u nd er c u lt iv at io n that n o th ing is kn o wn of th eir o r igin Of oth ers .

,

o n t he c o ntrary t h e life h is t o ry is well kn o wn

, .

Wh en E u ro pean s peo p led A mer ica th ey nat u rally bro u ght

wit h th em th eir fr u it s th eir v eget a b les th eir grain s th eir gr as s es
, , , ,

and th eir d o m es t ic an ima ls T he n ew c o u ntry wa s r ich in nat iv e .

s p ec ies bo th plant and anim al b u t the Eu r opea n s pec ies had

, ,

t he ad v a ntage o f being bett er kn o wn a n d b ett er adapt ed t o t he

s p ec ial n eed s o f m an A cc o rd ingly wh erev er t he in trod u c ed
.
,

v a r iet ies s u cc eed ed the c o rres p o nd ing nat iv e types wer e n eg

,

lec t ed ; b u t wh en t he E u ro pean v a r iet ies fa iled th en t he nat iv es ,

were d ev elop ed I t is fro m th is la tt er Cla s s that som e im po rtant

.

o b s er v at io n s m a y b e m ad e
1
.

The gooseb erry ?

T he large E nglis h go o s eberry was t oo t end er
fo r t he A m er ican c lim at e and with a l was exc eed ingly liab le t o ,

m ild ew N at iv e va r iet ies fl o u r is h ed wid ely in t he fo res t s

. U n fo r .

t u n a t ely t he v a r iet ies b ea r ing t he la rges t b err ies wer e exc eed ing ly
th o rny bo th on b u s h a n d fru it S id e by S id e h o wev er with th es e
,
.
, ,

prick ly so rt s were s m o oth v a riet ies fr ee fr o m prick er s bo th o n ,

”

fr u it and b u s h T h ese were freely tran s p la nt ed to t he ga rd en s

.

o f t he p io n eer s and f u rn is h ed an acc eptab le fr u it I n go od t im e

3
.

th ey d ev eloped im pro v ed so rt s — fi rs t t he Ho ug ht o n a s eed , ,

ling o r iginat ed by A b el Ho u ght o n o f L ynn Ma s s ach u s ett s s o m e , ,

t im e in t he ear ly fo rt ies T h en c a m e t he D own ing a s eed ling of .

,

t he Ho u ght o n fir s t d es cr ib ed in 1 8 5 3 t he fr u it o f wh ich is s aid

, ,

to b e “
t he larges t yet kn o wn b ein g ab o u t t wic e t he s ize o f t he ,

Ho u ght o n s seed lin g it s par en t it is pale or light gr een with o u t

’

, ,

an
y b lu sh and s m oo th
,
T h e S k in is v ery th in an d t he fr u it as
.

d elicat e and t end er as any E u r o pean go oseberry on it s nat iv e

s o il . T he flav o r and aro m a a re perfect .

Ba iley o b s erv es in th is c o nn ect io n “

T h is berry n o w kn o wn , ,

as t he D o wn in g is t he s tand a rd o f exc ellenc e in A m erican

,

goo seb err ies and is pro bably gr own m o re ext en s iv ely than all
,

o th er v a r iet ies c o m b in ed ; and et it is o n ly t wo r em o v es fr o m

y
t he wild s p ec ies
”
.

1 L H B il y E
. . l ti
a fe , N ti F it [
vo uT h M m i ll C
on m p y o ou r a ve ru s e ac an o an ,

N w Y k]
e or .

9 I b id p p 3 8 9 3 99 -
. . .

3
T h e wri t er r em em b er s v er y w ll e as a b oy s ea rc hi ng t he wo o d s, an d es p e

c i a lly t h e s wa m p s , of h
M ic iga n fo r t h es e s m o o t h v a r iet ies fo r t ra n s p l a n ti n g
.
 M UTAT I ON S 1 3 1

IS n ot th is a na t u ral m u tat io n in t he trues t sen se o f t he t erm

I f n o t th en it is m erely a q u es t io n o f t erm in o logy and d efin it io n
, .

T he fact rem ai n s tha t it aro s e s u dd en ly a d is t inct type and , ,

rem ain s tru e with n o charact er ist ic s of a hybr id We n eed a t er m .

for th is so rt of th ing wh ich is o ccas io nally o cc u rring ev ery wh ere

in nat u re in o u r gard en s and in o u r h erd s and I kn ow o f n o n e
, ,

bett er th en t he o n e already in u se m u tat io n .

Th e st rawb er ry 1
The wild s tra wberry gr ew ev ery wh ere in
.

north er n N o rth A mer ica T h ere were not o n ly m an y d is t inct.

types of t he red b u t like t he nat iv e ras pberry and t he blackberry

, , ,

it had ev erywh ere it s alb in o rac e G ood progr es s had b een m ad e .

in t he c u lt iv at io n o f t he n at iv e s t ra wberries and with o u t d o u bt ,

good v ariet ies wou ld in t ime ha v e d ev elo ped b u t the introd u ct io n

o f t he C h ilean berry ( the par ent o f m os t present v ar iet ies ) s eem s

t o hav e pu t a s t o p to th is T he m os t promis ing o f a ll n at iv e

.

s tra in s was t he Frag a r ia C/ uloen s zs a nat iv e t o Or ego n an d t he

'

Pac ific co as t ; b u t a s Ba iley o b ser v es ,

“
t he ga rd en pro geny o f ,

it s S o u th A m er ica n branch is a lready so goo d that th ere is litt le

reas o n fo r r et u rn ing t o the wild fo r a n ew s tart Here is a c u r i .

o u s in s tanc e o f t he su cc es s iv e s u pp lant i ng o f v ariet ies E u ro p ea n .

so rt s wer e v anq u is h ed by d ev elo p m ent s o f N ew England nat iv es .

T h en t he wild type of Orego n ca m e int o t he s tr u ggle and

threa t ened t o s u pplant th em bo th fo r it was fu ll o f pro m is e ,
.

B u t it s pros perity was it s o wn d efea t fo r it s o wn C h ilean ,

bro th er has n ow s u pplant ed ev eryth ing in that it is t he s t o ck

wh ich is f u rn is h ing o u r im prov ed v ar iet ies A n y s t u d ent o f th is .

s u bj ect will r ec og n ize t he c om parat iv e r ead in es s with wh ich th es e

n ew ty pes s pr ing u p .

Th e b lack b erry ?
The blackberry gr o ws wild bo th in A m er ica
and in Eu rope b u t is sa id to b e c u lt iv at ed o n ly in N o rth A m er ica
,
.

I t is n o t m o re than fi fty y ears s inc e im pr ov ed v ar iet ies wer e

introd u c ed and it s real c ult iv at io n d a t es o n ly fro m abo u t 1 8 7 5
,
.

T h ere are t wo pr inc ipal types o f the wild b lackberry gro wing
in the n o rth ern U n it ed S tat es : ( I ) t he “
h igh b u sh lo ng an d ”

lu s c io u s lov ing t he s had e r epr es ent ed in it s c u lt iv at ed types

, , ,

acco rd ing to Ba iley by the Taylor and t he A nc ient Br it o n ;

,

L H B il y E
1 . l ti f N ti F it p p 4 4 4 3
. a e , vo u on o o ur a ve ru s, . 2 -
2.

I b id p p
3
98 33. . 2 -
0.
 V A R I AT I O N
1 3 2

( )
2 a a ller v a r iety gr owing in s u nny Open plac es and bearing
sm ,

s m a ll ro u nd lo os e gra in ed fru it s r ipen ing lat e a n d exceed ingly

, ,
-
,

so u r .T h is type is r epr es ent ed in c u lt iv a t io n by L a wt o n ,

K ittat inny S nyd er A ga wa m Er ie and o th ers N eith er o f th ese

, , , ,
.

y ield ed read ily to c u lt iva t io n and res traint and t his fact s er v ed ,

in an early day t o earn an ev il repu tat io n fo r what Pr o fes s o r

C ard calls th is gy p s y o f t he fr u i t s N ev erth ele s s th ey y ie ld ed .
,

t o p er s is t ent e ffo rt s and ha v e giv en r is e a s Ba iley p u t s it to a

“
, , ,

h o s t of v ar iet ies v ery m any o f th em wild ing s o r ch anc e ,

b u s h es fo u nd in fenc e ro ws .
”

T he fi rs t na m ed v a r iety was t he D o rch es t er introd u c ed abo u t

-
,

1 84 1 I t s exact o r igin is u nkn o wn th o u gh it s o r iginat o r ( pr o b

.
,

ab ly Captain L o v ett ) is kn o wn to ha v e tran s plant ed wild plant s

fo r m any s u cc es s iv e y ear s Wh eth er th is fi r st c iv ilized gyps y
.

was a s po rt o r s im p ly a s tra in im pro v ed by s elect io n is n o t n o w

capa ble o f pr oo f and yet its c o n stancy is go od pr esu m pt iv e
,

ev id enc e .

VVils on s Ea rly was kn o wn in 1 8 5 4 t he Holc o m b in 1 8 5 5 and

’

, ,

in 1 8 5 7 t he L a wt o n ( firs t c alled N ew R o ch elle) was introd u c ed ,

being at o nc e d ec lared s u per io r to t he D o rch es t er Of th es e t he .

Wilso n was d isc ov ered in t he wild abo u t 1 8 5 4 by J ohn W ils o n

“

o f B u r lingt o n N ew J ers ey ,
and t he L a wt o n fo r m erly N ew ,

R o ch elle wa s fo u nd in t he t o wn o f N ew R o ch elle N ew Y o rk
, , ,

by L ewis A S ec o r T h ese t wo s tra in s hav e giv en r ise to

. .

n u m er o u s d is t inct m od ern v a r iet ies T he loo s e c lu s t er s tra in s

“
.
-

a re r ega rd ed by h o rt ic u lt u r is t s as t he d es c en d a n t s o f t h e W ils o n .

T he o r ig in o f c erta in o th er v a r iet ies s eem s t o b e a s fo llo ws

I n 1 8 7 0 M r W illia m Parry o f N ew J er s ey
.
“
s elect ed a , ,

h ealthy yo u ng D o rch es t er a n d plan t ed it in t he Sa m e h ill with a

s tr o n g h ealthy W ils o n s E a r ly o r breed er s o c a t ed fa r a way
’

, ( f ) l ,

fro m an y oth er blackberries 1

I n 1 8 7 5 t he s eed fro m s o m e o f
.

t he larges t b err ies gr o wi ng o n t he W ils o n were pla nt ed On e .

plant o n ly was regard ed a s va lu able an d all o th er s were d estroyed ,

.

T h is n ew s tra in wa s na m ed W ils o n J u n io r T h e fr u it wa s large “

.
,

ear ly and v ery fi n e a n d S O pro lific th at in 1 8 84 o n e acre ”

“
, ,

y ield ed 1 b u s h els o f fr u it by the s id e o f fiv e acres o f Wilso n s ,

’

Ear ly in t he s am e fi eld with t he s a me c u lt u re wh ich a v era ged , ,

B il y E 1l ti f a eC lt i t d F i t p 3 6
,
vo u on o o ur u va e ru s, . 1 .
 V A R I AT I ON
1 34

it is , m u s t b e cred it ed with ha v ing prod u ced many n ew for m s

s po ntan eo u s ly .

Grapes .
l
N o rth A m er ica is
a nat u ra l v in eyard says Bailey , ,

and yet with t he m os t s k illf u l and per s is t ent att em pt s to c u lt i

v at e t he E u ro pea n v ar iet ies fo r win e m ak ing th ey ha v e n o t s u c ,

c eed ed U nd er th es e c irc u m stanc es n o th ing is m o re nat u ra l

.

than that v a luable n at iv e v ariet ies s h o u ld arise pro v id ing t he ,

capac ity was inh erent in t he s pec ies .

J o hn A d lu m wr ot e abo u t 1 8 2 3 “
The way is to dro p m o s t
, ,

ki n d s of fo reign v in es at o nc e and seek fo r t he bes t k ind s o f o u r

larges t nat iv e grapes He is t o b e rem em bered fo r t he intro
”
.

d u ct io n Of t he fam ou s Cata wba wh ich was fou nd wi ld in t he

“
,

wood s of B u nc o m be C o u nty in extrem e west ern N o rth C aro lina

in 1 80 2 .

The Cata wba is , th erefo re almo s t c ertain ly a s port of t he ,

wild grape gro wing in pro f u s io n in that regio n I n 1 84 3 cam e .

t h e D iana a seed ling o f the Cata wba

,
.

I n 1 8 4 0 M r E W B u ll b o u ght a h o u se in Co nc o rd
. . . S om e .

s eed ling s o f wild grapes s prang u p abo u t it o n e of wh ich fr u it ed ,

in 1 8 4 3 I t was S O exc ellent in q u ality that all o th er s were

.

d es troyed and the n ew v ar iety was named t he C o nc o rd T h is .

seed ling has s inc e g iv en u s t he W o rd en Moo re Ear ly Po c k lin g , ,

t on ,
Eat o n and R ock land o f wh ich t he fir s t ha s lo ng b een
, ,

famou s The C on c o rd it s elf a m u tant seem s to hav e been

.
, ,

pec u liarly r ic h in po s s ib ilit ies for s t ill o th er races .

I n the y ear 1 8 2 1 Ho n o rab le Hu gh W h it e th en in the j u n io r ,

c las s in Hamilt o n C ollege N ew Y o rk plan t ed a s eed ling v ine , ,

in the gro u nd s o f Pro f es s or N o y es o n C ollege Hill wh ic h s t ill , ,

rem ain s and is the o r iginal C lint o n

,
.
”

T h es e a re o n ly a few o f the m a n y v a riet ies o f grape of A meri

can o r igin trac ing d irect ly to wild nat iv e s t o ck
, .

Lost possib ili t i es Had o th er d o m es t icat ed p lant s and an im als

.

bro u ght fro m E u ro pe s u cc eed ed les s ad m irably what en rich ment ,

m ight hav e c o m e throu gh t he nat iv e fl o ra and fa u na o f A m er ica !

T he prai rie ch ick en wo u ld ha v e b een im pro v ed if the d o m es t ic
hen had n o t s u cc eed ed T he t u rk ey wa s a n ew th ing and was
.

th erefo re seized u po n T he b u ff a lo wo u ld n o t n o w b e ext in c t

.

B il y E l ti f
1 a N ti
e , F it vo u
p p
— 6 on o our a ve ru s, . 1 12 .
 M U TAT I ON S 1 35

if catt le had acc lim ated les s s u cc ess f u lly N at iv e gra in s o th er .

than m aize wo u ld hav e been d ev elo ped had it n o t been fo r th is

c o m pet i ti o n and nat iv e gra sses ha v e n o t liv ed u p to th ei r po s s i
,

b ilit ies T h is is thro u gh n o fa u lt o f th ei rs th o u gh we s t ill lack

.
,

t he b es t A mer ican gra s s

”
.

S ECT I ON V — ECON OMI C SI G N I FI C A N C E OF M UTATI ON S

B eca u se of t he way wardn es s

po rt s t he im po s s ib ility
of s ,

o f pred ict ing th ei r app earanc e t he read i n ess wi th wh ich th ey

,

d is appear wh en int erbred wi th t he parent s t o ck and th eir v ery ,

freq u ent inab ility to reprod u c e at all — b ecau s e o f all these c on ,

s id erat io n s i t has bec o m e fa s h io nab le t o d ec lare s po rt s in gen eral

t o b e of S light ec o n o m ic i m po rtanc e and u n wo rthy t he br eed er s

’

s erio u s att ent io n T he o n ly c o u r se left o p en fo r im pro v em ent is

.
,

therefo re t he s lo w o n e o f grad u al acc u m u lat io n thro u gh s elec

,

t io n of m in u te b u t fav orable v ariat io n s acc o rd ing to the th eo ry ,

o f D arwi n .

T he b es t o f ev id enc e ex is t s h owev er fo r b eliev ing that th is

, ,

is a has ty an d u n warrant ed c o nc lu s io n and that m a n y if n o t , ,

i nd eed m os t o f o u r really v a lu ab le n ew typ es hav e aris en s u d

,

d en ly a s m u tatio n s and n o t grad u a lly thro u gh infin it es im a l d iffer

en c es as is c o m m o n ly s u pp o s ed
,
T he exp er iment s o f D e V r ies
.

and the A merican v ariet ies o f fr u it s bo th c o me n ear en o u gh to

t he o ri gi n o f typ es t o mo re than warra nt th is V iew o f t he s it u a
t io n and to a ffo rd gro u nd fo r t he greatest h o pe that u n s u s pect ed
po ss ib ilit ies s t ill ex is t in m an y if n o t m o s t d o m es t icat ed s pec ies ,

po s sib ilit ies o f s po ntan eo u s ly g iv ing o ff v ariet ies repres ent in g

es s ent ially n ew c o m b inat ion s o f t he charact er s o f t he s pec ies and

c o n s eq u ent ly pos s es s ed o f d iffe rent and p erhap s enhanc ed ec o

no m ic v a lu e T he wo rk of L u th er B u rbank and o f o u r c o m m er
1
.

c ial s eed s m en add c o nfi r m at io n t o th is h o pe wh ich if well , ,

fo u nd ed pro mis es n ew m eth o d s in breed ing and v a s t ly increas ed

,

po s s ibilit ies fo r im prov ement .

The s m all n u m b er s i n v o lv ed in an im a l breed ing red u c e en o r

m o u sly the C han c es of m u tat io n s appear ing a n d yet n early ev ery
1 W . S Harwo o
.
, in
dTlre Cen t u ry Al ag a zi n e, M a rc h and A p ril, 1 90
5 l
; a s o N ew
C reation s in ll
Plan t L ife [T he Ma c m i an C o m pa n , y
 6 V A R I A TI ON
1 3

s p ec ies h a s thr o wn o ff it s a lb in o v a r iety wh ich in m o s t cas es is

,

eas ily pro paga t ed Ho rn les s catt le o ccas io nally appear in n early
.

all br eed s and t he type is c o m pa ra t iv ely eas y o f pres erv at io n

, .

I t is m o re than lik ely that the d ifferent types in t he larger

breed s wh ich breed ers find s o d iffi c u lt to break u p a re in
, ,

reality q u it e d is t inct .

I n f u t u re chapt ers d ealing with t he m eas u rement s o f v aria t io n

and the s tat is t ic s o f hered ity in genera l it will ap pear that ev en
,

fix ed types a ffo rd s u ffic ient d ev iat io n to k eep a breed er b u s y

wi th select io n in o th e r wo rd s that t he an im a l breed er d ea ling
, ,

a s he is wi th s m a ll n u m b ers will a lway s fi nd s u ffi c ient v ar iat io n

,

t o lead him to s u p po s e that he is gett ing res u lt s o f h is s elect io n

ev en wh en he ha s n o t S h ift ed t he c ent er o f v ar iat io n t he s light

es t . M u ch that pa s s es fo r breed ing is n o th ing m o re than th is

i neffect u a l m u lt ip licat io n and i t is n o t t oo m u ch t o s ay that
,

h u ndred s o f breed ers and t ho u s and s o f an im als hav e liv ed and

d ied wi th o u t a ffect in g t he breed in t he s lightes t .

T he wri t er is s tro n g ly o f t he o p in io n that wh ile s elect io n is a

p o werf u l agent fo r S hap in g u p and fin is h ing o ff a fairly
“ “

acc eptable type and wh ile it is the o n ly m ean s o f d ec id ing

,

what s ha ll liv e and what S h all d is appear yet m u ch o f t he real ,

ad v anc e in bo th an imal and p lant breed ing is lik ely to c o me

thro u gh d is t inct o ffs et s wh ich a re n o w called m u tat io n s and ,

wh ich in D ar win s t im e and u nt il rec ent ly were erro n eo u s ly if

’

n o t repro achf u lly d e n o m in a t ed

,
s po rt s .

S ECT I O N V I — B IO LOG I C A I SI G N I F I C A N C E OF
,

MU TAT I O NS

T oo m u ch m y s t ery has rro u nd ed th is matt er o f s po rt s

su ,

a n d th ere has b een a too r eady t end ency to ev o k e t he a id o f

la t ent chara ct ers t o exp la i n th is and almos t ev ery o th er u n u s u al

phas e o f ev o lu t io n .

I n tr u th th ere is no m o re my s t ery abo u t m u tat io n s than

,

abo u t h ered ity in gen eral wh ich is a c o m p licat io n o f m y s t eries

, .

I t is n o t a q u es t io n o f lat ency b u t o f relat iv e pro m in enc e o f

charact ers of the po s s ib le los s o f a rac ial pec u liarity or what
, , ,
 3 V A R I AT I ON
1 3

If a rac e is pro d u c ed by hybrid izat io n th en a n ew c o m

n ew ,

b inat io n o f charact ers has been eff ect ed and it is fa ir t o a s s u me ,

that t he c o m binat io n is richer in poss ib ilit ies a n d po s s es s es a

larger n u m b er o f charact ers tha n d id eith er parent M u t a t io n .

t each es that n ew as s o rt ment s o f charact ers m ay take p lac e in ,

s o m e ca s es at lea s t with o u t hyb rid izing

,
.

I f a ra c ia l charact er as c o lo r o r ha ir i n es s is lo s t we rec o gn ize

, , ,

t he n ew typ e and na m e it a s a n ew creat io n I t m ay b e m o re .

v a lu able t o u s th a n it s parent b u t i t m u s t b e rec o gn ized b io lo g

,

ic a lly a s hav ing lo s t s o m eth ing t o wh ich i t was rac ia lly en t it led .

A gain if all n o r m al charact er s acq u i re a n u n u s u a l d ev elo p

,

ment relat iv ely o r ab s o lu t ely as in giant s o r if th eir d ev elo p

, , ,

m ent is abn o r ma lly arr es t ed as in d wa rf s we aga in rec o gn ize

, ,

t he n ew d epart u r e and it is a goo d m u ta t io n

,
.

S t ill aga in if c erta in ch a ract ers o n ly u nd ergo C hange in d ev el

,

o p m en t wh ile o th er s rem a in n o r m a l th en rela t iv e v al u es a re

, ,

changed the effect is altered and we rec o gn ize a d ifferent type

, , .

T h is t o o is a g o o d m u tat io n pro v id ed t he n ew relat io n p ers is t s

, , , .

A ll th es e changes c a n b e wo rk ed with t he n o r m a l charact ers

o f t he rac e with o u t t he intro d u ct io n o f n ew charact er s o r ev en
,

t he s u ppo s it it io u s a id o f la t ent chara ct er s S o b erly c o n s id ered.

,

th es e changes a re n o ne o th er than t he s t u d ent o f b io logy wo u ld

exp ect u n les s ind eed ra c ia l ch a ract e rs a re b o u nd t o geth er m u ch
,

m o re r igid ly than pres ent ev id en c e wo u ld lead u s t o s u s pect .

S u mmar y N o t all v aria t io n s a re c o nt in u o u s and c o nn ect ed

.

wi th t he typ e by in s en s ib le d iff erenc es S o me d ev iat io n s a re d is

.

c o nt in u o u s with a t end ency fo r f u t u re v a riat io n s n o t rev ert ing t o

,

t he m a in ty pe b u t c lu s t ering ab o u t a n ew c e nt er o f v ar iab ility th u s ,

S ett ing u p a n ew typ e S u ch a d ev iat io n is ca lled a m u t a nt and

.
,

n ew s tra in s m a ar i s e in th is m ann er a s well a s by t he s lo wer

y ,

D ar win ian m eth o d o f h et ero gen eo u s v a ria t io n o u t o f wh ich n ew ,

types are es tab lis h ed by t he S lo w pro c es s of s elect io n .

Bo th t he exp erienc e o f breed ers — es pec ia lly with n ew v arie

t ies in A m e ca
r i — and n u mero u s in s tanc es o f experim ent al ev i
d enc e s h o w c o nc lu s iv ely that n ew s train s n o t o n ly m ay b u t in ,

act u al pract ic e do o riginat e in th is m ann er s u dd enly and c o m

, ,

p le t ely w i th
, o u t any appar e nt pr epa r at io n and w ith li tt le t end e ncy
t o rev ert t o t he o r ig ina l o r m a in typ e wh ich c o nt in u es a s befo re
,
 M U TA T I O N S 1 39

Mu tant s lik e th eir parent types are s u bj ect to fl uct u at in g

, ,

v ariab ili ty wh ich is a n ec ess a ry la w o f repro d u ct io n and th ey

— —
, ,

m ay b e im pro v ed that is s haped u p b y j u d ic io u s selec ,

t io n b u t th ei r p rinc ipal c ha racters and mai n t rend were fi xed

,
'

wh en the type arose .

AD DITI ONAL REF ERENCES

A LB IN I S M A c ri tic a l s t u d y o f i ts B y E Pa n ta n elli E xperi men t

.
( c a u s es .
) . .

S ta ti o n R ec ord , X V , 5 5 .

A T A V IC M U T A T ION T O M A TO By C A W h i te S c ien c e, X V I I ,
—
O F T HE .

—
. . .

6
7 7 8 2 2
34 3 5 1 °

A T A V IS M IN T HE POTA T O . By S . R h o d in . E xperim en t S ta tion R ec o rd ,

X I, 7 1 0 .

D E TE R M I N A T E M U T A T ION S .
( D e V ries an d o t ers h q u o ted .
) By M . M .

M etc alf . S c ien c e, X X I , 3 5 5 3 5 6 -

.

E V OL U TION A ND A D A PT A T I O N By T . . H . M orga n . S c ien c e, X I X,

2 2 1- 2 2 5 .

E V OL U T I O N W IT HO U T M U T A TIO N . B y C B D av en po rt. . . S c ien c e, X I X , 2 1 5 .

I N HE R I T A N C E OF Mo N S TR O S I T I E s .
( E x
peri men ts of twelv e yea rs .
) By
Hu go D e V ries .E x perimen t S ta tio n R ec o rd , X 1 , 5 4 6 .

M U TA T I O N AND S E L E C T I O N ( W a t c a us es m u ta tio n s ? A re t
. h h ey all in
one d irec tion ? ) B y M M M etc alf
. S c ien c e, X I X , 7 5 7 6
. .
-
.

M U T A T I ON I N T HE T O M A TO B y C A W ite S c ien c e, X I V , 8 4 1 8 4 4
. . . h .
-
.

M U TA T I O N T HEO R Y ( A r ev iew o f S pec ies a n d V a ri et ies

) B y C B D a v en

—
. . . .

p o rt .S c ien c e, X X I I , 3 69 3 7 2 .

MU T A T I O N T HEO R Y O F D E V R I ES .
( T went y eigh t
- lec tu res b y the a u t o r h
at the U n iv ers ity of C alifo rn ia , E xp erim en t S ta tio n R ec o rd ,
XV 1. 7 4 5 .

M U T A T I O N T HEO R Y OF D E V R I ES . By D . T . M c D o u ga l . E xperimen t
S ta tion R ec o rd , X I I I , 3 2 4- 6 1 9 ; X IV , 2 2 6, 5 2 6 .

M TA T I O N T HE O R Y O F O R G A N I C E V O L T I O N ( A b r ief b u t po in ted s u r
U U .

v ey o f the s u b j ec t
) B y W E C as tle o f Harv a rd U n iv ers i ty
. S c ien c e,
. . .

X X I , 5 2 1 - 5 4 3 ; from s ta n d poin t o f an im al b reed in g, 5 2 1- 2

5 4 ; from
s ta n d po in t o f c ytolo 8
gy, 5 5 5
2 2 .

M U TA T I O N S IN PL A N T S . By D . T . M c D o ugal . A m eric an N a tu ralis t,

X XX V I I , 7 3 7- 7 7 0 ; also in E xp erim en t S ta tio n R ec o rd , X V I , 2 3 .

O R I G I N O F S P E E
CI S B y H u go D e V r ies S c
. ien c e, X V , 7 2 1 -
7 2
9 . .

O R I G I N OF S PE C I ES T HR O G H S E L E c T I O N C O N T R A S T ED W I T H U T HEI R

OR I G I N T HR O UG H A P PE A R A N C E OF D EPI N I T E V A R I ET I E S . By T H . .

M orga n . Popula r S c ien c e M o n t ly, L X V I I , 5 4—66 h .

PR EPOT E N C Y U
O F I N D I V I D A L S w1 T H A B N O R M A L V A R I A T I ON OR M UTA
T IO N .
(A s tu d
y of c a ts with ex tra toes .
) By H . B . T o rrey . S c ien c e,
X V II
 0 V A R I A TI O N
1 4

S OM E C A U SES OF S A L T A T OR Y V A R I A T ION . By C . H E igen m a n n

. . P ro
c eed i n gs o f the A m eric a n A s s o c ia tio n fo r the A d v a n c em en t o f S c ien c e,
1 00 ,
9 X L IX, 207 .

S PO R T S .
( A ut ho r c o n c lu d es h
t ere a re o t er la ws t a n h h M en d el s ’
a nd C alto n ’
s .
)
B y C B:D a v en po rt
. . S c ien c e, X I X , 1 5 1 als o in E x perim en t S ta tio n
R ec o rd , X V , 7 5 3 .

S PO R T S O N G R A PE V I N E S . By J . C . T a lb a c k . E x peri m en t S ta tio n R ec o rd
XV , 478 .

S PO R T S ; T I I E PE A C H N EC T A R I N E Jo u rn al o f the R o al Ho r tic u ltu r al S o

- . y
c iety, X X V I , 5 96 — 598 ; a ls o i n E x erim en t S ta tio n R ec o rd , X I V , 4
p 5 .

U
T HE M T A T I O N O F LY C O PE R S I C M B y C A W ite Po p ula r S c ien c e U . . . h .

M o n th ly , L X V I I 1 5 1 ,
-
1 62 .

T HE M U T A T I O N T H E O R Y .
( A d efen se .
) By T h o m as L C asey. . S c ien c e,
X X I I , 3 0 7— 0
3 9 .

T H E OR I G I N OF A W HI T E BLAC KB ER R Y . B y Lu t he r B u rb a n k . E x per
im en t S ta tio n R ec o rd , X I V , 1 07 1 .

T HEO R Y OF M UT A T IO N S . By A . A . W . Ho b rec h t . Po p u lar S c ien c e

M ont h ly , LX V , 2 0 5- 2 2 3 .
 CHA PTER VI I
THE MECHANI SM OF DEVE LOPMENT A ND DIFFER ENTIATI ON

B efo re pec ific inq u iries can b e pro fitably m ad e int o the ca u s es
s

o f v ar iat io n i t is n ec es s ary t o b ec o m e f a ir ly fa m ilia r with what

is kn o wn o f t he es sent ia l c o n s t it u t io n o f liv ing m a tt er and o f its

m anner o f gro wth and d ifferent iat io n .

A ft er att en tio n ha s b een bes t o wed fo r a t i me u po n th es e c o n

s id erat io n s it will b e ev id ent t o t he s t u d ent that h ere in the
, ,

inn er wo rk ings o f liv ing matter a re f u nda m enta l c au s es o f p ro

,

f o u nd v ariat io n s ev en in pro t o p las m s eem ingly the m o s t s table

,
.

S ECT I ON I — PR OT OPL A S M Y
T H E PH SI C A L BA SI S
OF L I FE

Pro t o p la s m is a gen eral na m e fo r a ll m att er that is end o wed

with life b u t t he s t u d ent m u s t n ev er f o rget that t he b io lo gis t
, ,

lik e t he C h em is t is d ea lin g with m a tt er c o m p o s ed o f well kn o wn

,
-

ch em ic a l elem ent s u n it ed in d efin it e pro po rt io n s T he kn o wn .

d ifferenc es bet ween liv in g and n o n liv ing m atter a re fo r o u r

-
,

p u rpo s es t he fo llo wing

,

1 .L iv ing m att er is end o wed with a m y s terio u s f o rc e ca lled

lif e
.

2 .M a tt er s o end o wed ha s a m u ch m o re c o m p lic a t ed ch em ica l

c o m p os it io n th a n has n o n liv ing matter o r th a n can b e m ain
-
,

t a in ed aft er t he life pr inc ip le has d ep a rt ed L iv ing m att er at

.

d eath th erefo re break s u p (o r d o wn ) int o o rd inary ch em ical

, ,

c o m po u nd s T he tru e c o n s t it u t io n o f liv ing m a tt er c a n no t

.
,

th eref o re b e d et er m ined by any kn o wn m eth o d s o f analy s is

, ,

wh ich rev ea l o n ly t he elem ent s in v o lv ed b u t n o t th eir exact

r elat io n s d u ring life .

3 .M att e r end o wed w i th li f e i s ab le t o appr o pr iat e t o it s elf

o th er o u t ly ing m att er and t o incr ea s e it s b u lk thro u gh gro wth .

1 4:
 THE M ECHA N ISM OF D EV ELOPMENT 1 43

4 . T h is
gr o w th i s n o t o f b u lk m e r ely b u t i t is att end ed by ,

different iat io n so that o n e pa rt is d is t inct ly d ifferent from

,

ano th er .

5 A
. s gr o w th proc ee d s b i t s o f th i s b u lk a r e thr o wn O ff each ,

Of wh ich c o n s t it u t es a n ew ind iv id u al capab le of i nd epend ent

ex is t enc e reprod u ct io n
,
.

6 T he n ew ind iv id u al is s u bs ta n tia lly b u t n ev er exact ly

.
, ,

lik e t he o n e fro m wh ich it arose and h ere lie t he ch ief m ys ,

t eries o f breed i ng I n repro d u ct io n th ere a re no d u plicat es

. .

N o th ing appro ach es th is in the i n o rga n ic wo r ld s a v e c ry s tal

liza t io n .C ry s tals add m att er t o th eir b u lk and th u s may b e
s a id t o gr ow Mo reov er the matt er is add ed in an o rd erly
.
,

m a n n er r es u lt ing in a k ind of d efin it e s tru ct u re with exact

,

angles always t he same b u t n o th i ng lik e d iff erent iat ion ex is t s

, .

On e part o f t he c ry s t al is lik e an o th er ; it has n o p o wer o f

reprod u ct io n and is po s sess ed of n o fo rc e c o m parable with life .

T he s t u d ent s h o u ld early learn th a t t he fi eld o f b iolo gy is

d is t inct b u t he s ho u ld also f u lly realize that it lies with in an d
,

n o t o u t s id e t he rang e o f ch em is try and that liv ing m att er is no t ,

freed fro m its o rd ina ry afli n ities by reas o n of its assoc iat io n with
life b u t o n t he c o ntrary it c o nt in u es as befo re to b e s u bj ect t o
,

t he o rd inary phy s ical and ch em ica l relat io n s o f m att er gen era lly .

I f he can d o th is he will s im p lify many o f his d ifli c u lt ies

, .

SECT I O N I I —T HE C ELL T H E U N I T OF ST R U CTU R E

If a b it o f liv er bo n e woo d o r any o th er fo r m o f plant or

'

, , ,

an im al t is s u e b e exa mined u nd er the m icro sc o pe it will b e fou nd ,

to po s s es s a d efin it e s tr u ct u re and to c o n s is t o f a large n u m b er

,

o f s epar a t e d iv is io n s each fi lled with a gelat in o u s m a s s ca lled

,

pro t o plas m T hese separat e d i v is ion s o r c ells a re apparent ly

.

alik e throu gh o u t the s u bs tanc e o f any part ic ular t is su e — as

—
,

the liv er b u t th ey d i ffer great ly in d iff eren t t is s u es o f t he

,

s a m e b o dy ( bo n e m u s c le bra in ),
B io lo gis t s ha v e b een u n willing
,
.

t o c o n s id er the ind iv id u a l a s t he u n i t b eca u s e he is to o large ,

and his s tr u ct u re and act iv it ies a re to o c o m p licat ed T h ey hav e .

,

th erefo re ch osen to rega rd t he in d iv id u al as a c olo ny o f m any

,

and v ario u sly d ifferent iat ed c ells .

1 44 C A U SES O F V A R I AT I O N

In a s su m ing t he c ell as a u n it m a ny s tr u ct u ral d iffi c u lt ies

were s o lv ed t o the ent ire s at is fact io n o f t he anat o m is t b u t the ,

phy s io lo g ic a l and ev o lu t io na ry pro b lem s were c o m p licat ed ra th er

than s imp lifi ed beca u s e tl us en ti re colony of ma ny d ifleren t c ells
' '

a nd a c t i v i ti es lzea r t, l u ng s , liv er , m u sc les , n erv es , etc .

, w i th
tlzc i r m a ny a nd d i v er s e f u nc t i ons p
s ra ng or i gi n a lly f r om a

s in
g l e c el l ; m o reo v er ,
th is c o lo ny will thro w Off a s u cc es s io n o f
s ing le c ells each o f wh ich wi ll u nd e rg o s p ec ific an d o rd er ly
,

d ev elo p m ent a n d fi nally p r od u c e a c olony like l li o p a r en t T h is .

being tru e tlze cell ca n n ot b e reg a rd ed a s tlze u lti m a te u n it of

,

liv ing m att er u n les s we a s s u m e s o m e k ind o f u n ity b et ween all

,

t he c ells ; s o m e k ind o f int erc ellu lar fo rc e t o in su re that d iffe r

en t ia t io n S h a ll t a k e p lac e at t he pro p er p o int s and s tages ,

o th erwis e t he o r ig in a l c ell wo u ld d ev elo p int o a lu m p o f pro t o

p la s m o r int o a c o lo ny o f c ells a ll a like .

The s ing le c ell fro m wh ich a n ew ind iv id u a l is t o d ev elo p

( t h e
“
g er m c ell o r m o th er c ell
) is gi ft ed with p o t ent ia lit ies

fo r t he ent ire b eing with a ll its c o m p licat io n s o f s t ru ct u re and

,

with all i ts v ariety o f f u nct io n B io lo gis t s at o n e t i me were

.

inc lin ed to regard th is ger m c ell as t o t ip o t ent that is able ”

, ,

t o d ev elo p int o a lm o s t any k ind o f s tr u ct u re d ep end ing u p o n

t he s u rro u nd in g s T h is v iew c o u ld n o t h o ld beca u s e d ifferent
.

germ c ells u nd er id e n t ica l c o nd it io n s o f life d ev elo p each into

it s o wn s p ec ies .

T he ca u s e o f d i fferent iat io n th erefo re lies p ri m a r ily with in

, , ,

a n d t he ger m c ell is t o b e reg a rd ed a s g ift ed with u n li m it ed

po wer s o f d ev elo p m ent on ly wi t/t i n tlze clea ra etc rs tlza t b elong to

flee s
p ec i es .

S pec i fic pro t o pla s m is t heref o re po s s es s ed o f s pec ific pro per

, ,

t ies a s tr u ly a s is any ch em ica l s u b s t a nc e and a ll t he ch a racters

'

o f s tr u ct u r e o r f u n c t io n belo nging to t he m at u re ind iv id u al

a re t o b e regard ed a s i n s o m e wa nh erent i n t he germ
“ "

y i .

T he c ell is th eref o re lik e the ind iv id u a l t oo la rge and t o o

, , ,

c o m p licat ed t o b e c o n s id ered as t he u lt im a t e u n it o f liv ing

m att er .

T h is v iew is u ph eld n o t o n ly u po n th eo ret ic a l gro u nd s b u t

a lso by t he kn o wn f a ct s o f i t s c m p lic a t ed s tr u c t u re a n d it s re
o

m arkab le beha v io r d u ring c ell d iv is io n and gro wth — a s u bj ect ,

 1 46 C AU S ES O F V A R I AT ION

“
chro m o meres o f o th ers the mic ro so mes o f s t ill o th ers , ,

and t he id s o f Weis mann “ ”

.

L y ing g en e ra lly in the cyt o p la s m j u s t o u t s id e the n u c leu s will

c ommo n ly b u t n o t alway s b e fo u nd an extremely m in u t e h igh ly
, ,

s ta in ing b o dy t he c entro s o m e ab o u t wh ich wh en d iv is io n is

, , ,

abo u t to o cc u r t he n ear b y m att er is thrown int o rad iat ing lines

,
-

lik e ir o n fi ling s ab o u t the p o les o f a magn et giv ing t he wh o le a ,

k ind of s tarlik e appea ranc e .

T h es e a re t he po rt io n s o f the c ell m o s t c o nc ern ed in c ell

d iv is io n s and th eir s pec ial charact ers a re m os t pro n ou nc ed and
,

t he d ifferenc es m os t d is t inct j u s t prev io u s t o the a c t o f d iv is io n ,

and leas t well mark ed in t he c ell d u ring its res t ing s tage “ ”

b et ween d iv isio n s .

T he act u al pro c es s o f c ell d iv is io n wh ereby o n e c ell giv es

rise to t wo and by wh ich gro wth is attained is es sent ially as
follo ws
Wh en d iv is io n is abo u t to tak e p lac e t he chro m at in m atter
( gr a n u les ) a ss u mes t he app earanc e o f a fi n e n et wo rk ru nn ing

throu gh t he m as s of the n u c leu s t he g ran u les look ing like b ead s ,

s tr u ng u p o n a thread T h is n et wo rk c o m m o n ly th o u gh n o t
.
,

alway s c o nd en s es int o a r ibbo n o r thread ( the s p ireme) wh ich

, , ,

h o wev er s peed ily break s u p tran sv ersely into a d efin it e n u m

,

b er o f s eg m ent s gen erally in th e fo r m o f s h o rt ro d s s tra ight o r

, ,

c u rv ed ( the chro mo s o mes ) Wh eth er o r n o t the ret ic u lar o r n et .

wo rk f o r m pa s s es thro u gh t he s p i rem e s tage t he res u lt is alway s ,

t he s a m e ; n a mely t h e chr o m at in m att er b ec o m es d iv id ed int o

,

a d efinit e n u m ber o f chro mo s o m es Here a re t wo remarkable .

a n d s ign i ficant fact s ; fi rs t tlze n u m ber of c lzr om os om es is c on ,

s ta n t in a ll i n d i v id u a ls o
f tlze sa m e s ec i es ;
p and s ec o nd “
i n a l l ,

1
ec ies a r i s i ng by
’
s ex ua l r e rod u c ti on tlze n u m b er is
s
p p ev en .

Wh ile t he c hro mat in ma tter has been engaged in break ing u p

o r d o wn f o rm t he chr o m o s o m es an o th er S ign ifican t pro c es s
( ) t o ,

1 W il s o n
T h e C ell, p 6 7
, T he a u t o r ere giv es t h e n u m b er o f c ro m o s o m es
. . h h h
h
C ara c t eris t ic o f c erta in s p ec ies as f o o ws : s o m e o f t he s a rk s ,
3 6 ; m ll
o u s e, h
d g ly
sa la man er , t ro u t , a n d li , 2 4 ; o x, gu in ea p i , a n d o n io n 1 6 ; g ras s o
pp e r, 1 2 , h
A s ca r is , 4 or 2 ; t he c r u s t ac ea n A r tem ia , m a n , 1 6 o r p o s s ib
1 68 ;
3 2 I ly
n t is . h
c o n n ec tio n it is wo rt hy o f n o te t h at v a r iet ies o f t h e s a m e S ec ies o f t en
p i ffer in d
t h e n u m b er of h
t eir c h ro m o so m es , t h e s ign ific a n c e o f w ic h h
v a ria t io n h as n o t

y et b een d eterm in e d .
 T HE M EC HA N I SM OF D EVE LOPM E NT 1 47

has been go ing on The c entro s ome has d iv id ed and the t wo

.

n e w bo d ies d eriv ed fro m it ha v e separa t ed and m igrat ed to

Oppo s i t e S id es o f the n u c leu s each s u rro u nd ed by i t s rad iat ing

,

lin es in which c o nd it io n th ey a re kn o wn a s
,
“
a s t ers ( s tars ) .

D u ri ng thi s m igrat io n t he a s t ers a re gen era lly ( n o t alway s )

v is ib ly c o nn ect ed by lin es b u t in eith er case by t he t im e th ey
,

hav e rea ched o ppo s it e S id es Of the n u c leu s th ey will b e s een to

lie at o ppo s it e end s o f a s pind le s haped b ody ( the a m ph ias t er ) -

c o n s is t ing of lines am o n g wh ich lie t he chro moso mes

,
.

Matt ers a re n o w ready fo r t he fin al and S ign i fican t act s o f

c ell d iv is io n The chro mo so mes arrange th em selv es en d t o en d
.

a lo ng t he eq u at o r of the s pind le and at right angles to its ax is ; ,

wh ereu po n each C hro m os o m e sp l i ts leng tlzwis e o n e gro u p o f ,

halv es m igrat ing to o n e as t er (c entro so me) the o th er to t he ,

o th er wh ere each c lu s t ers abo u t it s o wn c ent er f o r m in g a n ew

, ,

n u c leu s with its c en tro s o me T he c ell wall n o w bec o m es c o n

.

s t ri c t ed d iv id ing the cyt o p las m appro x im ately eq u ally (s o m e

,

t imes v ery u neq u ally ) bet ween the t wo n ew c ells and the d iv is io n ,

is c o m p let e The res t ing s tage en s u es d u ring wh ich preparat io n

.
,

is m ad e fo r an o th er d iv is io n ind eed t he c entro so m e o cca s io n ally ,

d iv id es in ant ic ipat io n of t he next d iv is io n ev en b ef o re all the

, ,

d etails o f the firs t d iv is io n a re c o m p lete Fo r a graph ic o u t lin e .

o f t he c o m p let e pr o c es s o f m it os is s ee Fig s 2 0 and 2 1 . .

Th is in gen era l is the pro c ess o f c ell d iv is io n wh ich with

, , ,

m o re o r les s v a riat io n att end s a ll gro wth ,

T he s ign ificant fact s .

bro u ght to light in th is c o m p licat ed pro c es s a re : ( 1 ) that the

n u m ber Of chro mo so mes is c o n s tant fo r all ind iv id u als with in
t he s pec ies ; ( 2 ) that fo r a ll fo r m s ar is ing by s ex u al rep ro d u c
t io n t he n u m ber is ev en ; ( 3 ) that h o wev er its d etails m ay v ary ,

c ell d iv is io n c o n s is t s es s ent ia lly in a s plitt ing o f t he chro m o

s o m es by wh ich each dau ght er c ell sec u res (apparent ly ) an
,

exact eq u iv a lent o f what is rec eiv ed by t he o th er da u ght er c ell

o f t he s a m e d iv is io n C ell d iv is io n is th erefo re n o t a lu m p
.

d iv isio n of the c ell m as s b u t it is m eris t ic in s u ring a s trict ly

, ,

q u alitat iv e d iv is io n in wh ic h o n e ha lf o f ea ch C hro mo s o me
d es c end s to eith er da u ght er c ell l
.

1 Th m f t h
es e s a e dd d ig ifi
ac s wh
av e a id
e sd i n tic anc e en c on s ere n c o n n ec on

wit h th g m
e ll p d ti
er ce s , re d th p b l m
ro uc f h d ity
on , an e ro e s o ere .
1 48 C A U S ES O F V A R I AT I O N

Fig . 20 . D ia gra m s s h o win g t h e p ro p h a s es

of m it o s is

A, ll with
res t in g c e l
ret ic u a r n u c leu s a n d tru e l l
n u c eo u s : a t c t he att ra c t io n S phere c o n ta i n in
g
t w o m
cen t ro o s es . B, ea r ly p ro
p ha se : t he c h ro ma t i n fo r m in g a con t i n u o u s p
s i r em e,

l l ill
n u c eo u s s t presen t ; a b o v e,
( ) t he
C,
D ,
t w
a m ph ia s te r
o d i fleren t t ypes o f a ter
a . l
h es : C , d is a p ea r an c e o f t he ri ma r y s i n d le, d iv er gen c e o f t he c en t ro so mes to
p p
ro a s p p p
l ll
Op po s it e po les o f t he n u c eu s ( exa m p les , s o m e p la n t c e s , c lea v a ge s t a ges o f ma n y eggs )

D , pers is t en ce o f t he p ri ma ry s pi n d le ( to fo rm in so me ca ses the c en t ra “

l
l
fa d i n g Of t he n uc ea r memb ra n e, in gr o wth o f t he as t ra l ra ys , seg men ta t io n o f t he s p i reme
l
t h read t o fo rm t he c h ro m o s o mes ( exa m p es , epid er ma c e s o f sa a ma n d er, fo r ma t io n o f l ll l
t he po l ar b od ies ) . E, l a t er p ro p ha se o f t ype C : fa d in g of t he n u c l ea r me m b ra n e at t he

po es, l fo r m a t io n o f a ne w s p in d le in s id e t he n u c leu s ; p rec oc io u s sp l i ti

t n g Of t he c hr o mo
s o m es ( t he l a tter n o t c ha ra c t er i s t ic o f t his t ype a o n e) l .
“
I t he m it o t ic figu re est a b is hed ;
. l
e
p , t he eq u a to r i a l pl a te o f c h ro m o s o m es A fter
. I ls o n
1 5 0 C A U SES O F V A R I A T I O N

a ls o b e s im ilar to that o f the m o th er c ell fro m wh ich th ey a re

d es c end ed .

F or all c ell d iv is io n wi th in t he s a me t iss u e th is latt er is tru e ;

that is to say in t he gro wth o f liv er t iss u e bo n e t is s u e o r any
, , ,

o th er s p ec i fic s tr u ct u re t he c ells app ear t o b e id en t ic a l and

their res u lt ing gro wth s a lik e B u t it m u s t n o t b e f o rgo tt en that.

th es e d ifferent t is s u es all aro s e o riginally fro m a s ing le c ell ; in

o th er wo rd s that s o m e c ell d iv is io n s a re att end ed by d z
, f er en t i

a t i on Wh en ? Ho w ? Here lies t he ch ief my s t ery o f v ariat io n

. .

T he m echan is m o f c ell d iv is io n wo u ld s eem t o b e s pec ia lly

d es ign ed to p rev en t d ev iat io n and to in s u re ab so lu t e tran s miss io n

F ig . 22 h l gi
. Pa t o o cal m it o s is in ep i d erm al c e ll s of sa a l ma n d er c a u s e d b y p o is o n s

A, as ymm t i l m it
e r ca o s is a fter t ra t men t wit h l
a n t i pyr in s o u t io n ; B , t ri po l ar m it o s i s
after t rea t men t with p o t a ss ic i o did l
s o u t io n . A fter Wilso n ,
fro m G a l eo tt i

fro m mo th er c ell to da u ght er c ell I t d o es n o t acc o u nt fo r o r .

ind eed appear to ad m i t o f d ifferent iat io n o f t is s u es o r v a riat io n

in gro wth B u t d iff erent iat io n d o es tak e p lac e and v ar iat io n is
.

a fact t o b e i n s om e way exp la ined .

I rr egu l ar iti es in c ell di v ision N o t a ll c ell d iv is io n it is tru e.

, ,

p ro c eed s with the r egu lar ity and perfect io n o f pla n ind icated in
the d es cript io n wh ich is the c o m m o n m eth o d in h igh er an im als
,

and p lant s and may th erefo re b e r ega rd ed as fa irly typical T he .

kn own abn o r m al c a s es a re o f s ev eral d is t inct k ind s

1 A sy m m et r ica l m i tos is in wh ich t he chro m o s o m es a re n o t
.
,

eq u a lly d is t rib u t ed t o t he d au gh t er c ells m o s t o f th em m a s s ing ,

 T H E M EC HA N I SM O F D EV ELO PM E NT 1 5 1

a t o ne po le ,
so me o f th em perhap s being los t a lto geth er in t he
m as s o f t he cyt o plas m .

2 . M u l tip ola r m i tos i s in wh ich t he

n u m ber of c entro so mes is
,

m o re than t wo and the res u lt ing da u ght er c ells three or m o re .

B o th th es e abno r m a l proc es ses h o wev er a re charact eris t ic o f , ,

a bn o r m a l gr o wth s s u ch as C anc er s and t u m o r s and a re th erefo r e

, ,

c o n s id ered as path o lo g ica l I t is a s u gges t iv e fact t hat s u ch .

i rregu lar it ies m ay b e art i fic ia lly pro d u c ed by p o iso n s and o th er

F ig . 2
3 . Pa t h l
o o gic a l m it o s es in h u m a n c a n c er c e ll s

A, as ymmet ri ca l l
m it os is wit h u n eq ua cen t ros o mes ; B , a ter s ta ge, s ho win g u n eq ua d ist ri l l
l
b u tio n of the c h ro mo so m es ; C, q u ad ri po a r m it o s is ; D , t ripo la r m itos is ; E , ater stage ; l
l ll
F , t ri n uc ea te c e res u ti n g l
A ft er W i so n , fro m G a eo t t i
. l l
C h em ical s u b s ta n c es s u ch as ch lo ral q u in in n ic o t in ant ipyrin , , , ,

c o c ai n etc ( S ee F igs 2 2 and

,
1
. .

3 A
. m i t o t ic d i v i s i on ? — that is d iv is io n w ith o u t t he f o r m a ,

t io n Of t he a m ph ias t er o r the s plitt i n g o f the chro m o so m es .

T his fo rm o f c ell d iv is io n is eff ect ed by c o n s tr ict io n res u lt in g ,

s i m p ly in a lu m p d iv is io n o f t he m as s o f t he n u c leu s with o u t ,

referenc e to q u a litat iv e c o n s id erat io n s I n th is cas e t he da u ght er .

c ells wo u ld n o t pres u mab ly b e alik e T h is f o r m o f c ell d iv is io n

, ,
.

W il
1 T h C ll pp 97 98
so n, I b id p
e e 4 , .
, . . . 11 .
 C A U SES O F V A R I AT I O N
1 5 2

is of rare occ u rrenc e is n ev er kno wn in em bryo n ic t is s u es and , ,

is charact eris t ic o f t is s u es o n t he way t o ward s d egen erat io n 1 “

.

4 Q u
. i t e g en era lly u n i c el l u la r o rg a n is m s d is p lay e xtr em e

i rr egu lari t ies in m it o s is s o m e s pec ies o m itt ing o n e and o th er s ,

ano th er o f the pro c es s es typ ica l in h igh er s pec ies Pro m inent .

a m o n g th es e d ev iat io n s is t he failu re of chro m at in g ran u les to

u n i t e t o f o r m d e fin it e chro m o so m es I n plac e o f th is t he ind i .

v id u a l g ran u les th e m s elv es d iv id e s u gges t ing that

f i s s io n o
f ,

is tlze el em en ta ry a nd es s en t ia l n u c lea r
tlt e g r a n u les f ea t u re o
f
2
d i v is ion .

Oth er m in o r d ev iat io n s a re kn o wn th o u gh m u ch o f t he ,

fi eld is yet u n wo rk ed Th ese may acc o u nt to s o me ext ent fo r .

d ifferent iat io n d u ring c ell m u lt ip licat io n and yet s o far as is at ,

pres ent kn own all proc es s es that d o n o t acc o m p lis h an eq u itable

d iv is io n o f the chro m at in gra n u les thro u gh t he s plitt in g pro c es s
a re loo k ed u p o n a s d is t inct ly path o lo g ica l N or ma l cell d i v is ion .
,

s eem s be i n tlt e i n teres t f d ifier eu tia

'

tlzer e
f ,
o r e to o c on s ta n cy , n ot

t ion , a n d wlza t p ower i t is tlza t p r od u ces on e s o r t o

f tiss u e f rom
a n otlzer , as m u s t lzapp eu i n tlt e d ev elop i ng em b ry o, is s ti l l a

my s tery .

S ECT I O N V — PHY S I O LOG I C A L U N ITS

T h is d iffic u lty has led to t he as s u mpt io n of s o m e s o rt o f phy s

io lo gic a l u n it s s o m e o f wh ich a re act iv e at c erta in s tages o f
,

d ev elo p m ent o thers at o th er s tages and the chro m at in g ran u les

,

wh o s e q u a litat iv e d iv is io n is in mo s t ca s es caref u lly in s u red

a re q u it e gen era lly r egard ed as t h e rep os it o r y o f th es e u n it s

and the c o m mo n v eh ic le o f h ered ita ry trans m is s io n S u ch were .

the gemm u les o f D ar win t he s t irp o f Ga lt o n t he id io p la s m o f

3
, ,

N i geli and the d et er m inant s o f W eis m ann

,
4
.

1 W il s o n , T he C e ll , pp . 1 1 6- 1 2 1 .
2 Ib i d .
p .
90 .

1 S ee D a rwin , A n im a l s an d P l a n ts , c h pa . xxv n .

4 W eis ma n n ’
s l
e a b o ra t e t h eo r y of h d ity
ere re a r
g d d
e t he germ p as m as t he l
o rigi n a l s u b s ta n c e, of w ich h t he bo dy i s t h e n a t u ra l ex
p a n s io n . h
T is “
a n c es t ra l

g erm p l h gi g
as m h g bl d is l g th p i
unc an d n , unc an ea e, a n , so on as e s ec es en u r es , is
im m t l or aHe g d t h i g m p l m
. m p i d l t im t l y f b i p h
re a r s s er as as c o r se u a e o
“
o o rs
”

l( if b
e ) wh i h m
ea rers y b p k f l i
, i g m l c l Th b i p h a e s o en o as v n o ec u es . es e o o rs , o r

u lt im t it
a e un mb i d i d ly m s, i t
a re c od t m i t wh ne n an o r er an n er n o
“
e er n an s,
”
o se

ac t i ity
v t d l p m t d t m i wh t th p t i l p t h ll b T h
a ev e o en e er n es a e ar c u ar ar s a e. es e
1 54 C A U SES O F V A R I ATI O N

u n it s req u ired to expla in the f u nct io n and ach iev ement s Of t he

ger m plas m — the b it of v italized m att er that h o ld s with in its
,

s u b s tanc e all t h e p o t ent ialit ies o f its part ic u lar k ind o f lif e .

S u mmary T he cau s es o f v ariat io n a re c lo s ely c o nn ect ed

.

with t he m echan is m o f gro wth and d i fferent iat io n The c ell is .

the u n it o f s tr u ct u re and all gro wth is by c ell d iv is io n ; b u t it

is no t t he u n it o f d ifferent iat io n o f d iff erent part s o f t he bo dy ,

becau s e all part s ar is e fro m o n e o r iginal c ell the germ c ell , .

C ell d iv is io n s eem s ad m irab ly adapt ed to in s u re ab s o lu t e t ran s

m is s io n wi th o u t v ariab ili ty Of any k ind B u t b o th d ifferent iat io n .

and v ariability a re f a c ts We s eek th eref o re a phys io lo gi c al

.
, ,
“

u n it mo r e m in u t e than t he c ell wh os e ac t iv i t ies and po ss ib ly ,

wh o s e c o m b i nat io n s with o th er phy s io lo g ica l u n i t s o f d i fferent

pro pert ies a re able to bri ng fo rth fi rs t d ifferent iat io n withi n t he
bo dy and later d ifferenc es b et ween d ifferent ind iv id u als .

ADDITI ONAL REFEREN CES

C HR O M O SO M E V E S I C L E S IN M ATU R ATION . By W M . . S m a llwood . S c ien c e,

X X I , 3 86 .

C Y TO LO G I C A L FE AT U R ES OF F E R TI LI Z AT I O N . By W . H . B lac m an k .

Pro c eed i n gs of the R o a l S o c iet y , y

Lo n d o n , L X I I I , 4 00 4 0 1 -
.

F E R T I L IT Y O F EG G S A FT E R R E M O V A L O F C OC By L G K . . . J a rv i s .

E x peri m en t S ta t io n R ec o rd , X I , 6 7 1 .

L A ws O F E M B R Y O N I C D E V E L O PM E N T : T HE L A w O F V O N B A ER . By
Ot to G lase r . S c ien c e, X V , 97 6 9 8 2 — .

M EC H A N I S M OF D E V E L O P M E N T B y W illia m T u rn er,
. F R S
. . . Po p u la r
S c ien c e M o n t h ly L V I I , 5 6 1 5 7 5
,
-
.

O N T O G E N ET I C AN D P I I Y L OG E N E T I C V A R I ATIO N By H F Os b o rn
—
. . . .

S c ien c e, I V , 7 86 7 89 .

PR O B L E M O F D E V E L O P M E N T . By E . B W ils o n
. . S c ien c e, X X I , 28 1 —2 93 .

PR OT O P L A S M I C S T R U CT U R E . By E . B W ils o n
. . S c ien c e, 1 1 , 8 93 - 8 99 ;
x1 33 45-

S OM E OB S E R V A T I O N S AND C O N S I D E R A T ION S U PO N M A T U R A T I O N PH E
NO M EN A OF G ER M C ELL S . By T . H . Mo n tgo m ery . B io lo gi c a l
B u lletin , V I , 1 904 .

STR U CTU R E AND FO R M A T I O N OF PU S C E L LS . E x perim en t S ta ti o n

R ec o rd , X I V , 1016 .

V IT A L IT Y O F PO L L E N .
( R o s es , twe n t y -
two da ys ; c liv ias , h
t ree m o n t s h .
)
E x peri men t S ta tio n R ec o rd , X I I I , 6 2 0 ; ( B ea r , t irty d a y )
s X V h
, 8 7 2 .
 CHA PTER VI I I
IN TER NAL CA U SES or V ARI ATI ON

Whi le t he cau ses of v ariat io n a re bo th internal a nd ext erna l

t o t he o rgan is m t he fact s o f t he las t chapt er m u s t s at is fy
,

t he s t u d ent o f breed ing pro b lem s that m any o f t he pr oc es ses

att endant u po n gro wth and reprod u ct io n a re fru itf u l so u rc es o f
v a riab ili ty
. I t is t he p u rpo s e o f t he pres ent chapt er to d is c u ss
these int ernal in fl uen c es s om ewhat at length T h ey a re o f t wo .

k ind s — ( I ) tho se affect ing the ind iv id u a l o n ly and ( 2 ) tho se

, ,

a ff ect ing t he rac e as a wh o le I t is exped ient t o d is t ing u is h

.

bet ween th es e t wo c las s es and t he chapt er will be d iv id ed int o

,

t wo part s c o rres po nd ing to th ese d is t inct io n s as fo llo ws

, ( )
1 ,

i nt er n al infl u enc es a ffect ing primarily t he ind iv id u a l ( 2 ) int ernal

in flu enc es a ff ect ing t he rac e as a wh o le .

I —I N TE R N A L I N FL UE N CE S A FFE C TI N G P R I M A R I L Y
THE I N D I VI D UA L

SECT I O N I C E L L D I V I S I ON

Gro wth is t he resu lt o f c ell d iv is io n Man ifes t ly th erefo re

.
, ,

a ll d ifferenc es in S ize o r i n patt ern a re int im at ely d epend ent u po n

t h e ext ent and reg u lar ity o f th is proc ess .

Morph ological v ariation du e to cell di v ision What ev er in flu .

en c es u nd er lie t he ph en o m ena o f m it o s is a ll q u es t io n s Of f o rm
,

o r S ize a re ab s olu t ely d epend ent u po n t he ext ent t o wh ich c ell

d iv isio n an d its att endant gro wth proc eed T he ind iv id u al c ells .

in gi ant s a re no t larger than th os e Of n o r m a l s pec imen s b u t th ey ,

a r e mo re n u m ero u s ; and i n d warf s th ey a re n o t s m a ller b ut ,

f ewer in n u m ber What energies d ec id e ho w far c ell d iv is io n

?

s hall proc eed and wh en i t s ha ll s t o p in t he ca s e o f each s eparat e

W il
1 T h C ll pp 3 8 8 3 8 9
son , e e , .
-
.

1
55
 1 5 6 C A U SES O F V A R I AT I ON

o rgan we d o n o t kn o w Fo od and c lim at e u nd o u bt ed ly ex ert a .

gen era l infl u enc e a s we s ha ll see b u t a lt ogeth er as id e fro m th is

, ,

th ere m u s t b e pro f o u nd int erna l fo rc es o r int errelat io n s h ips ,

u p o n t he n o r m a l ex erc is e o f wh ich a ll typ ica l res u lt s d epe nd .

C o n s id er t he d ev elo p ment Of a n o rm a l ind iv id u a l fro m t he

fert ilized o v u m to mat u rity The c i rc u ms tanc es req u ire no t o n ly .

that a rm leg and bo ne h eart liv er and brain ar is e at the pro pe r

, , , , , ,

t ime an d plac e b u t a ls o that t he att endant c ell d i v is ion s in ea clt

,

r oc eed to tlt e r e u is i te n u m b er a n d Men s to I f h n u m be r b e

p q p t e .

t o o few a d warf is t he res u lt ; if t oo large a g iant ; and if t o o

, ,

few in s o m e part s ( a rres t ed d ev elo p m ent ) o r too large in o th ers

( hyp e rtr o phy
) t h e i nd iv id u a l i s thro wn o u t o f pro po rt io n and is
,

rec o gn ized as m o re o r les s o f a m o n s tro s it y acc o rd ing to the

d egree o f d is pro po rt io n T o b e s u re all th ese th ings occasio n .
,

ally happen an d yet in t he m aj o rity o f cas es t he proc es s o f

, , ,

c ell d iv is io n is adj u s t ed with a n ic ety that is n o th ing s h o rt o f

m arv elo u s ; in any ev ent t he res u lt s s ec u red th o u gh v a ry ing , ,

s o m ewhat in t o ta l d ev elo p m ent a re yet a lm os t ab so lu t ely ,

pro p o rt io n al
Wh a t ev er m ay b e t he c o n tro lling f o rc e to d ec id e at what po int
c ell d iv is io n in each c as e s ha ll s t o p and wh en t he ind iv id u a l as
a wh o le S ha ll c ea s e t o gro w the p lain phys io lo g ical fact is that ,

a ll c o n s id erat io n s o f S ize ( q u ant itat iv e v ar iat io n ) a re f u n d a m en

tally th o s e o i c ell d iv is io n .

T he c es s at io n o f gro wth at m at u rity d o es n o t im p ly th e lo s s

o f p o we r o f c ell d iv is io n bec a u s e m o s t f o r m s o f lif e plant o r a n i
, ,

m al h a v e m o re o r les s p o we rs o f regen erat io n if a part is lo s t

,

o r i n j u r ed I f a leg o f a s a la m and er b e c u t a way it will s peed ily

.
,

b e res t o red b o n es and a ll as goo d a s n ew A ta il o f a lizard is

, , .

read ily b ro k en o ff s ep a rat in g n o t bet ween two v ert ebrae b u t at

,

t h e m idd le o f a v ert ebra ( i n s o m e s p ec ies gen era lly t he s ev enth

ca u dal) Wh en the tail regen erat es h o wev er t he v ert ebra : d o

?
, ,

1 h
A t t is p o in t t h e aut h or q u es t io n s h is o wn s ta t em en t . As a m a tt er of fac t ,
the d a ta in v o l dh
ve av e no t b een s ub m it t e dt o l
a b s o u te mat h em a t ic a ld eterm in a

t io n . We do not k n ow w h the er t h e n o r m l d
a ev ia t io n in s i ze d ue to v a ria tio n in
ce ll d i v is io n is t h e s ame fo r a ll s p ec ies ; nor d o we k no ww h the erin gia n ts an d
d w f ll p
ar s a a rt s b ear the sa me l
re a t iv e p ro p o r t io ns as in n o rm a l
S p ec im en s ;

i d d th
n ee , ere is gro u n d
f o r b e i ev i n g l t h at t h y e do no t . I n t h e m o s t gen era l
h w
s en s e, o ev er, t he s ta tem en t is t ru e .
2 M o rgan , R egen era t io n , p 1 98 . .
1 5 8 C AU SES O F V A R I ATI O N

Meri sti c v ariat ion in gen er al du e t o c el l di v i sion A ll d ifferen .

t ia tio n in v olv ing n u m bers o f d u p licat e part s ma nifes t ly has it s

s eat in c ell d i v isio n A n ad d it io nal d iv is io n at t he po int o f o rigin
.

o f t he s er ies d o u b les t he n u m ber b u t an ext ra d iv is io n at t he ,

po int o f o rigin o f a m em ber add s a pa ir if bo t h dau ght er c ells ,

d ev elo p o r a single mem ber if b u t o n e d ev elo ps

,
.

When the n u m ber in a m eris t ic series is ev en t he series is

eas ily c o nc eiv ab le as ha v ing ari sen fro m a c o rres po nd ing n u m b er

o f c ell d iv is io n s Fo r exa m p le .

2 in the series , 1 d iv is io n
4 i n t he s eries , 2 d iv is io n s
6 in t he s eri es , 2 d iv is io n s , wit h one p a ir d iv id ing aga in

8 in the seri es , 3 d iv is io n s

10 in the s eri es , 3 d i v is ion s , wit h o ne p a i r d iv id in g a

ga i n

I f t hen u m ber o f mem bers is Od d it is o n ly n ec es s ary to a s s u me ,

that on e o f the ev en n u m bers failed to d ev elo p o r what is m o re

—
, ,

lik ely that o n e o f a pa ir ind u lges i n add it io n a l d iv is io n

,
it s ,

s is t er m em be r rem a in ing s i ng le ; th u s

3 m em b e rs ,
I d iv is io n , one mem b er d iv id in g ag a in

5 m em b er s ,
2 d iv is ion s , o ne m em b er d iv id ing ag a in

7 m em b ers , 2 d iv is io n s , two mem b ers d iv id ing aga in

9 m em b ers , 3 d iv is io ns , o n e m em b er d iv id i n a
g g a in

T he freq u ent rec u rrenc e o f fiv e as a d igi tal n u mber is o n e o f

t he m y s ter ies in creat io n and it s s ing u lar p er s is t enc e is an o th er
, .

I t is h o wev er s u bj ect t o m any d ev iat io n s as wa s s een i n t he

, , ,

C hapt er o n Meris t ic V a riat io n

“
ev en i n t h e ro s e fa m ily th ere

is an o cca s io na l lo s s o f o n e o f the m em ber s .

T he freq u ent pres enc e o f S ix d ig it s is n o t t o b e exp lai n ed by

rev ers io n as n o bo dy s u ppo ses that n u m ber ev er to ha v e been
,

C ha ract e ri s t ic i n any S p ec ies — a fact that s h ou ld b e n o t ed by

,

s o m e o f o u r friend s who a re a lway s r eady t o i n v o k e t he aid o f

atav is m to explain ev ery abn o r ma lity .

Meris t ic v ariat io n lik e o th er d ev iat io n s a ris ing fro m ext ern al

,

cau ses is to so m e ext ent h ered itary and capab le o f being in

, ,

flu en c ed if n o t ab s o lu t ely c o ntro lled thro u gh s elect io n

,
NO ,
.

o th er meth o d is kn o wn a s id e fr o m t he fact that ext ernal i nj u ry

,
 I NT ER N A L C A U SES OF V A R I ATI ON 1 59

to m a ny p la nt s and c ertai n a nimals res u lt s in b u dd ing and m u l

t iplic a t io n o f part s We c u t t he m a in s tem o f a s m a ll tree o r
.

s hr u b in o rd er t o increa s e t he n u m b er o f s id e branch es So me .

what S im ilar ly inj u red part s a re o ft en d o u b led in regenerat io n

,
?

I n th is way lizard s m ay b e m ad e t o prod u c e an increas ed n u m ber

o f t oes and ev en d o u b le f eet leg s and ta il I t is s u pposed that
, ,
.

d ou ble feet som et imes seen ev en in m am ma ls may b e pro d u c ed

, ,

by a f old o f t he a m nio n c o ns tr ict ing the m idd le o f the beg in

“

ni ng o f the y ou ng leg 2
in t he em bryo T h is h o wev er is c u ri .
, ,

ou s rath er than v a lu ab le t o u s as it t end s t o exp la i n abn o r m alit ies ,

ra ther than to po int a way to pract ical im pro v ement .

I rregu lariti es in cell di v i sion 11 cau se of v ariat ion ?

T he char
ac ter is t ic act in c ell d iv is io n s eem s t o b e t he s p li tt ing o f t he

chro mo so mes (o r chro m at in gran u les ) and t he m igrat io n of exact

eq u iv a lent s to eac h n ew da u ght er c ell s tro ng ly s u gges t i n g that ,

the as so rt m ent o f phy s io lo g ical u n it s ( what ev er they may b e)

rec eiv ed by o n e dau ght er c ell is an exact d u p licate o f that rec eiv ed
by t he o th er th u s in s u ring an o rd erly and s y s t em at ic d ev elo p
,

ment thro u gh a s trict ly q u alitat iv e d iv is io n o f h ered itary s u b

s tanc e at ea ch a n d ev ery s tage o f gr o wth .

The wh o le m ech an is m o f m it o s is s eem s adj u s t ed t o th is en d ,

and if t he as s u m pt io n is t r ue it s s igni ficanc e can hard ly b e o v er

es ti m at ed .If th is caref u l adj u s t ment o f t he mecha nis m o f c ell
d iv is io n is nec essa ry t o o rd erly d ev elo p ment it is m an ifes t that ,

any s u b s tant ial d ev iat io n is lik ely if n o t c ertain to res u lt in , ,

v ariat io n m o re o r les s pro f o u nd S u ch d ev iat io n is charact eris t ic .

o f a m it o t ic d iv i s io n g en era lly and it is mo re than c o nc eiv able ,

that the o rd inary pro c es s is s u bj ect to o ccas io na l s lips S o me .

chro mat in gran u le m ay fa il to d iv id e at t he pro per m o m ent and

m ay pas s o v er to o n e da u ght er c ell en t ire o r c o n v ers ely it
‘
, , ,

m ay ind u lge in an extra d iv is io n S u b s tant ia l d ev iat io n s in the .

proc ess are kn own t o occ u r n o t ra rely b u t freq u en t ly F o r ex .

a m p le the s p litt ing so met imes tak es p lac e in t he s p ireme s tage

, ,

s o met imes aft er t he f o r m at io n o f t he chro m oso m es ; s o m et im es

1 M o rgan , R egen era t io n pp , . 1 3 7 - 1 39 .

1 Ib i d .
p 39
. 1 .

3 S ee p rev io us c h a p ter .

4 h
T is is k no wn to o c c u r in c erta in in s t an c es in m a t u ra t io n .
1 60 C A U SES O F V A R I AT I O N

the c entro s o me d iv id es befo re the res t ing s tage m o re c o mm o n ly ,

a ft er ward T a k in g it a ll in a ll h ere is a n exc eed ing ly c o m p li

.
,

c a ted pro c ed u re o n ly s em i m ech a n ica l and t h eref o re s u bj ect t o

,
-

d ev iat io n s A bs o lu t e c o n s ta n cy d em an d s n o fa ilu re in t he fi na l
.

o bj ec t o f exact q u a li tat iv e d iv is io n b u t t he s t u d ent s ees m any ,

p o s s ib ilit ies fo r u n eq u a l d iv is io n a n d th erefo re fo r d ev iat io n in

gro wth I S th is t he f u nd am ent a l ca u s e o f m u tat io n s ? On e th ing
.

is c erta in — liv i n g f o r m s a re m ad e u p o f elem ent s and t h es e

, ,

elem ent s a re s u bj ect t o s trang e c o m b in a t io n s thro u gh o u t t he

e n t i re range o f p lant a n d an im a l lif e and t he fact s s eem t o t each ,

t hat fro m t im e t o t im e c o m b inat io n s m a y a r is e that a re ent i rely

n ew . M o reo v er wh o le u n it s s eem o cca s io n a lly to b e lo s t o u t

,
“
,

a s wh en h o rn ed c a tt le s u dd en ly g i v e ris e t o p o lled s t ra in s ha iry ,

s p ec ies t o s m o o th v ar iet ies c o lo r ed t o a lb in o etc , ,

.

C o n v er s ely d o v it a l elem ent s lik e ch em ica l ra d ic les as s u m e

,

n ew c o m b inat io n s fro m t im e t o t im e g i v in g r is e t o n ew char ,

a c t ers a n d n ew typ es wh ich we c a ll m u ta n t s ? W e d o no t “ ”

kn o w a n d yet we feel t he c o n v ict io n that at th is po int we a re

,

v ery c lo s e to t he o r ig in o f charact ers the ca u s e o f m u tat io n s ,

and o f v a ria t io n in gen eral .

M an ifes t ly in so far a s ir reg u la ri ties in c ell d iv is io n m ay b e a

,

ca u s e o f v aria t io n t he m att er lies abs o lu tely beyo nd o u r c o ntro l

,

exc ept that lin es i n wh ich it is b eliev ed t o o cc u r ma y b e a v o id ed

i n s elec t io n Here is a fi eld h o wev er t o o far bey o n d o u r pres

.
, ,

en t k n o wled g e to ad m i t o f a n yth in g m o re than the m eres t

m ent io n W e c o nfid ent ly b eliev e that t he f u t u re will s h ed mo re

.

light o n th is o b s c u re s u bj ect .

S EC TI O N II —B I S EXU A L R E PR O D U C TI ON A F U N D A M ENTA L

C A U S E O F V A R I AT I O N

A m o ng h igh er a n ima ls and pla nt s t he n ew ind iv id u al is the

d irect p rod u ct o f t wo o th er s t he m a le a n d t he fem a le parent
, ,

a n d is o f n ec es s ity d iff erent f ro m eith er b ein a pr d ct f

g o u o ,

b oth I n bisex u al reprod u ct io n t h erefo re b io lo g ist s rec ogn ize a

.
, ,

f u ndam en t a l ca u s e o f v a riat io n S light if t he pa ren t s a re Of like

,

blo o d lin es extrem e if o f rad ic a lly d ifferent a s in h ybrid is m

, ,
.
1 62 CA U SES O F V A R I ATI ON

A ll that is in v olv ed in fert ilizat io n is n o t well u nd ers t ood b u t ,

it s es s ent ial f eat u r e is t he u n i on o r f u s i on of tlze n u c lea r m a t ter

( c lt rom os om es ) f o tb c g er m c ells f r om t wo p a r en ts to f or m tlze

c lea v ag e or s eg m en ta t i o n n u c leu s wlt os e s u bs e u en t
q g r o w l /c a n d

f rt l d n u c le i o tlze b ody T h is
”
d i v is ion s i r is e to f a ll i i tlze
“
g v e e z e .

o v u m b ec o m es th eref o re t h e fi r s t c ell Of t he n ew being wh ich

, , ,

inh er it s d irect ly and eq u ally a po rt io n o f the n u c lea r m att er fro m

bo th parent s s o that ev ery n u c leu s of t he ch ild m ay c o ntain
“
,

n u c lear s u bs tanc e d er iv ed f ro m both parent s 1

Here th en is
”
.
, ,

t he av en u e o f a ll inh er it a nc e and as t he n ew ind iv id u a l is a k ind , ,

of b lend o f b o th parent s we s ee in fert iliza t io n an in it ia l and ,

prim ary cau se o f v ar ia t io n .

T h is is t he o n ly f orm o f v ar iat io n rec o gn ized by W eis m ann in

his ea rlier wr it ing s as in an y s en s e h er ed itary A ll d ev iat io n s .

in d ev elo pm ent d u e to ext erna l cau ses were c o nc eiv ed to a ffect

t he b o dy ( so m a plas m ) o n ly ex ert ing n o infl u enc e u p o n t he
2
,

a nc es tral germ p la s m T r u e he lat er ann o u nc ed t he th eo ry o f

3
.
,

g erm inal s elect io n in wh ich a k in d o f s tru ggle fo r ex is t ence is

,

c o nc eiv ed as tak ing plac e bet ween t he bio ph o rs ( phy s io lo gica l

u n it s ) by wh ich s om e pro s per and m u lt ip ly exceed ingly wh ile
,

o th er s a re cr o wd ed o u t ent irely T h is wo u ld giv e an o th er ca u s e

1 ‘
.

o f v a r iat io n with in t he germ plas m o f each ind iv id u al .

B io lo g is t s gen erally rec o gn ize int ern al ca u ses o f v ar iat io n

o th er than th es e and et t h is u n io n Of t he chr o mo s om es from
y ,

d ifferent ind iv id u als tak ing plac e at each n ew gen erat io n m u st b e

regard ed a s a v ery effect iv e m ean s o f introd u c ing v ariab ility .

E v en if the o ffs prin g of a s ingle parent as in parth en o gen es is , ,

s h o u ld b e a n exact d u p licat e o f t he parent wh ich it is n o t , ,

ev ery o n e wo u ld rec o g n ize t he fact that t h e blend ing o f h ered i

tary s u b s tanc e fro m t wo par ent s m u s t of n eces s ity prod u c e an

ind iv id u al with a n ew c o m b inat io n o f fac u lt ies .

I t is a v a riat io n h o wev er c o nfin ed n o t o n ly to t he charact er s

, ,

o f t he rac e b u t als o t o t he fam ily po s s es s io n s o f t he pa rt ic u lar

parent s Bisexu al reprod u ct io n cann o t b e loo k ed u po n as a m ean s

.

W il
1 T h C ll p
son , 8 e e , . 1 2 .

3 S m pl m o a i t mas dt p t th p
s a t pl
er m f t h b dy i
use o re r es en e ro o as s o e o n

g en eral d i t
as i t f m t h
s nct p t f t h ro l ll ( g m p
e ou l m ) u o e s ex u a ce s er as .

W i m
3 e s T h G m Pl m
an n , h p ie er as , c a . x .

W i m
4 e s G mi l S l ti
an n , (p mph l t)
er na e ec on a e .
 I NT ER N A L CA U SES O F V A RI AT I ON 1 63

of intr od u c ing charact ers int o t he rac e and wh ile it is m an i

n ew ,

fes t ly a fr u itf ul so u rc e of n ev er end ing c o m b inat io n s of rac ia l

-

charact ers in n ew ind iv id u als yet v a riat io n s so introd u ced are

,

c o m parat iv ely slight exc ept wh en t he t wo par ent s belo ng to sepa

rat e lin es .

Fert iliza t io n of t he ov u m is so m eth in g m o r e than a s t im u lu s

t o gr owth I t is a rea l u n io n of m at er ia l bo d ies phy s iolo gical
.
,

u n it s or what ev er th ey m a b e c alled r epres ent ing t he h er ed ita ry

, y ,

s u bs t an c e o f bo th par ent s B isex u al r eprod u ct io n is th er efor e

.

n o t o n ly a g uaranty of tran s m is s io n o f rac ia l charact er s b u t also

an as s u rance o f inh eritanc e with som e v ar iat io n .

Con t rol Here is a f u ndam ental cau se o f v ar iat io n pract ica lly
.

u nd er t he c o n tro l o f t he br eed er thro u gh s elect io n T r u e his .

,

kn o wledge an d his j udgm ent are in s u ffi c ient t o in su re him aga in s t

m is tak es in m at ing and it is also tr u e that th er e are m any o th er
,

infl u enc es at wo rk to prod u c e v ariat io n s b u t th is is t he field in ,

wh ich t he breed er can ex ert t he larges t infl u enc e and it is by ,

s elect io n that t he gr eat es t res u lt s in im pr ov em ent hav e b een

at ta in ed u p to dat e .

matin g, an d assortat iv e matin g

1 2 ?
S ex u al sel ecti on , pref erenti al
Po werfu l th ese infl u ences in d irect ing t he tr en d o f v aria
as are

b ility th ey yet belo ng t o gen eral ev o lu t io n beca u se th ey a re ele

,

m ent s in nat u ral s elect io n and th ey ha v e n o plac e in t he pres ent

,

d is c u s s io n .

S EC TI ON III — MATU R ATI ON A N D T H E R ED U CT I ON OF

T H E C HR O MOSO M ES A C A U SE OF V A R I ATI ON

F ert ilizat io n is a proc es s wh o s e in ev itable c o n s eq u enc e wou ld

seem t o b e t he n u c lea r m a tter i n d efi n i tely for
“
p il i ng p
u o f
if, wi th each gen erat ion t he Chrom osom es ( o r phy s io logical
n ew ,

u n it s ) o f t he o n e par ent a re a dd ed t o th o s e o f t he o th er it wo u ld ,

s eem that in t im e t he r es u lt ing n u c lear m att er wo u ld s peed ily

beco me u n m anageably large and inco nc eiv a bly c o p x m le — an ,

ev ent c ertain to f ollo w exc ept for a s er ies o f v ery rem arkab le

1 D a rwin , O r ig i n o f S p ec ies , s ee I n ex d .

3 Pea rs o n , G ra mm a r o f S c ien c e , p p 4 2 5 4 2 8.
-
.

3 Ibi d pp 4 2 9 4 3 7
. .
-
.
1 64 C AU S ES OF V A R I A TI O N

fact s occ u rring j u s t prev io u s to fert iliza t io n and by wh ic h tlze

n u m b er o f c /zr om os o m es i n b ot/z tlze m a le a n d f em a le

g er m cells i s
r ed uc ed to on e lza lf tlt e u sua l or s o m a t ic n u m b er , so th a t th eir
u n io n at fert iliza t io n res t o res the tru e n um ber o f chro m o so m es
typical of the rac e T h u s if t he s o m a t ic n u m ber Of chrom o so m es
.
,

is S ixt een t he n u m ber in t he ger m c ells at fert iliza t io n will b e

,

eight each o r S ixt een aft er f u s io n o f t he n u c lei

,
T h is proc es s by .

wh ich t he n u m ber o f c hro m o s o m es is ha lv ed in the germ c ell is

kn o wn as red u ct io n and is s u ppo s ed to b e the s ign ific a nt feat u re
,

o f t he m at u rat io n pr o c es s by wh ich t he m a le and fem ale ger m

cells a re prepared fo r u n io n .

Paral leli s m in t h e sex es Ma t u rat io n a n d it s att end a nt p he .

n o m en a o f r ed u ct io n in t he n u m b er o f chro m o s o m es is a s u bj ect

that m u s t b e c on s id ered s eparat ely in t he m a le and t he fem ale ,

and yet th ere ex is t s a s trange para llelis m wo rt hy o f n o t ic e To .

q u ot e W ils o n 1

Re t h h h w th t m t ti
c en res ea r c f m t th as m t y pe
s o n a a u ra on c o n or s o e sa e

i b th
n o e S t t d i th m t g
s ex s . l t m th i p ll l i
a e f l n e os en e ra er s s a ra e s as o

l w : I b th
o s n th fi ol d ti i th e mb
s ex es f h m m e i na re uc on n nu er o c ro oso es s

eff ec ted i n t he c o u rs e o f t he la s t t wo c ell d iv is io n s , or m a t u r a tion d i v i szb u s

[as t e h y a re c a lled
] , b y w ic h h t he d efi i t i n ve
g er m c ells a r is e, ea c h of th e
fo u r c ells t h us fo rm ed h a v in g b ut h alf th e u su al n u m b er of c h ro m o s o m es .

I n the fem a le b u t o n e Of the fo u r c ells

[ res u lti n
g fro m t he t wo m a tu ra tio n
d iv is io n s ] fo r ms th e ov um p ro p er ,
w ile h th e o t he r t h ree, k n o wn as t he

p ola r bod ies , 2

a re m in u te , ru d i m en t a r y , an d i n c a p ab le of d ev elo p m en t . In
the m a le, on t he ot h er h a n d , a ll fo u r of t h e c ells b ec o m e fu n c t io n a l s
pe r

m a to zo a . h
T is d i fferen c e b e t ween th e t wo s e xesis p ro b a b ly d ue t o t he
p hy s io lo g ic a l d i v is io n of lab o r b et ween t he g er m c e lls , t h e s per m a to zo a

b ei n g m o t ile a nd v er y s m a l l, h w i le th e e
gg c o n t a in s a la rge am o u n t of

p ro to p la s m a n d yo l k , out o f wh ic h t he m a in m ass o f t h e em b ryo n i c b od y is

fo rm ed . I n t he m a le, t h ere fo re, a l l o f th e fo u r c ells m ay 3
b ec o m e fu n c
tio n a l in th e fem a le t he fu n c t io n s of d ev elo p m en t h av e b ec o m e res t ri c ted

to b ut o n e o f th e fo u r, w ile th e h ot he s h
r av e b ec o m e r u d i m en t ar
y .

1 W il s o n , T he C e ll , p . 2 34 .

2 T he aut h or is h er e s p ea ki ng s p ec i fic a l ly o f rep r o d u c t io n in a n im a s , l as

p l an ts do fo r m p o a r b o ies
no t l d . T he d i fferen c e i n p la n t an d a n im a l r ep ro d u c

t io n is , h o wev er m o r e in fo r m t
, h a n in s ig n i fic a n c e .

3 T he a u t h or sa y s m ay b ec o m e f u n c t io n a l. I I e m ea n s b y t is t a t eac h h h of

t he fo u r ce ll s ( s p er m a t o zo a ) a ri s in g f ro m t h e l a s t t wo d iv is io n s is c a p a b e l of

l
fer t i i zin g an o v u m , w i hl e o f t he fo u r ce ll s a r is i n
g f ro m t h e l as t t wo di v is i o n s in
t he fem a l e on ly o ne is c a pa b e o f l b e in g fer t i i ze l d .
1 66 C A U SES O F V A R I ATI O N

fo r t he la s t t wo , m at u ra t io n ,
d iv is io n s I n th is c o nd it io n the
or .

egg c ell rem a in s u n t il n ear t he t im e o f f ert ilizat io n wh en t he ,

pro c es s o f m at u rat io n pro per t ak es p lac e .

T he S ign ificant d eta ils o f th is int eres t ing s eries o f changes a re

c o nc ern ed with the n u c leu s and a re s u bs tant ially as IOIIOW S ’

D u ring t he lo ng res t ing s tage preparat o ry to th es e fina l d iv is io n s

t he n u c leu s incr ea s es i n b u lk and t he chro m at in m att er a ss u mes
t he ret ic u lar f o r m charact er is t ic of t he res t i ng s tage o f d iv id in g
c ells in general I n th is c o nd it io n t he n u c leu s is kn o wn as t he
.

“
germ inal v es ic le U p t o t his p o int t he n u m ber o f chro m o
.

s o m es is t he s a me a s that o f t he bo dy c ells in gen era l T h ei r .

id ent ity is o f c o u r s e n o w lo s t b u t as t he t ime fo r t he fi rs t

, , ,

m at u rat io n d iv is io n arr iv es in s t ead o f t he sp i r em e o f o rd ina ry

,

m it o s is break ing u p i nt o t he u s u al n u m ber o f chro mo s o m es ,

th ere appear m o re o r les s s po ntan eo u sly a n u m b er o f prim ary

c hrom at in m as s es in t he fo rm o f rod s rings o r V s haped bo d ies , ,
-
,

each o f wh ich u lt i m at ely break s u p int o f ou r s m aller b o d ies .

T h es e gro u p s o f fo u r a re a lway s on e lza lf t/ze u s ua l or soma tic

n u m ber o c /zr om os om es
f .

Wh eth er the chro m at in m as ses appear in the f o rm of ro d s ,

ring s or o th erwis e t he final res u lt seem s to b e alway s the sa me

, ,

n a m ely t he break ing u p o f each int o f o u r s m aller b o d ies eith er

, ,

by two lo ng i t u d i na l d i v is ion s or by on e ( t he fi rs t ) long i t u d i na l

a n d on e tra ns v ers e T he d et ails d iff er in d ifferent s pec ies and
.

hav e been wo rk ed ou t in b u t few cases I t is n o t im po rtant .

h ere to t rac e t he bewild ering d ifferenc es b u t rath er t o d es crib e ,

typ ical beha v io r ?

Hav ing a ss u med th is c o nd it io n t he n u c leu s n o w m ig rat es to

t he m ar gi n o f t he c ell each o f the gr o u p s o f fo u r ( t et rad s in rod
,

s hap ed ca s es ) s p li tt in g int o t wo s m aller gro u p s o f t wo each

( dyad s
)
?
T h e m a s s n o w d i v id es o n e pa i r fr o m e ach gr o u p ,

1 Fo r a fu ll di s c u s s io n o f th e di ff eren t fo rm s of d
re u c tio n , s ee W il s o n , The
Ce ll , V, 2 3 3- 2 8 7 .

9 T he t erm s “
t et ra d an d
“
dy d
a o f c o u rs e a
pp ly on ly in t h e c as e o f ro d

s h ap e dm as s es . In t h e c as e o f ri n gs ( c o mmo n l )
in d V h
a n im a s
p d m an -
s a e as s es

( c o mm o n l
in p a n ts ) t he p a rt in g in t o f ourt k a
p l ges d ll y t
ac e h w k pra ua as e or ro

c eed s , hl
w i e in th e c as e o f ro s t he d di i i i t f
v s on n o k pl o u r tal y d th es a c e ea r an e

p ar t s a re di s t in c t fro m t h e fi rs t . In th i f m t i
s or th a tw d i i i
on t k e p l o v s o ns a e ace

m uc h m o re ra p i d l y th an in t h e c as e o f i g wh i h p l it
r n s, d d i id
c s l wl y T h an v e s o . e
 I NTER NA L C AU S ES OF V A R I AT ION 1 67

remain ing in t he c ell t he o th er pas s ing o u t s id e fo rm ing t he , ,

fi rs t po lar bo dy wh ich may o r m ay n o t u nd ergo f u rth er d iv is io n

, .

T he po rt io n n o w rem a in ing wi th in t he c ell c o n s is t s o f gro u p s

of two each i n s t ead o f f o u r and th ei r n u m ber is o f c o u r s e t he
, ,

s a me as b ef o r e na mely o n e ha lf t he s om at ic n u m b er o f C hro m o
, ,

s o m es I mm ed ia t ely n o w with o u t as s u m i ng t he res t ing s tage

.
, ,

t he dyad s o r gr o u p s of t wos t u rn on e f o u rth aro u nd tak ing a

, , ,

po s it io n at right angles to t he m argi n of the c ell and a t onc e ,

d i v i d e ag a i n , on e m em ber f
o w eb p a i r rem a i n i ng b elzi n d i n tlze
egg cel l , tlze otlzer p a s s i n ou t , f or m i n tl s ec on d ola r b ody
g g ze p .

T he fi r s t polar body carried a way on e h alf t he n u c lear m att er ,

and the rem ai n in g half has no w been d iv id ed eq u ally bet ween

t he s ec o nd p o lar b o dy and t he m ain c ell wh ich is n o w ready fo r ,

fert ilizat io n and is fro m th is t ime on s po ken of as the ov u m .

N eith er p olar bod y carries any apprec iab le q u ant ity o f cyt o
p las m an d bo th a re d es t in ed to d egenerat e and d is appear T he
,
.

fi rs t o n e h o wev er c o nta in ing ha lf Of t he t o t al n u c lear matt er

, , ,

c o m m o n ly d iv id es o nc e s o that the fi rs t polar body represent s ,

n o t o n e b u t two c ells
, t he fi rs t and the th ird po la r b o d ies ,
o
.

T he t o tal res u lt th en of th is c om p licat ed proc es s s eem s t o

, ,

b e t he eq u al d iv is io n o f t he chro m at in m att er bet ween t he

o v u m capab le o f f ert ilizat io n and three p o lar b o d ies d es t in ed
, , ,

to ext inct io n .

A grou p o f f o u r c ells th u s aris es — na mely t he mat u re egg , ,

( o v u m) w h ich
,
aft er f ert il izat io n g iv es r is e t o t he em bry o and ,

three s m all c ells or po lar bo d ies ( incapab le of f ert ilizat io n ) ?

wh ich tak e n o part in the f u rth er d ev elo p ment are d iscard ed , ,

and soo n d ie with o u t f u rth er change T he egg n u c leu s (of the .

o v u m pr o p er ) is n o w r eady fo r u n io n with t he s p er m n u c leu s ?

wh ich proc es s is kn own as f ert iliza t io n

—
.

“
I n s o m e ca ses fo r exa m p le in the s ea u rch in — t he p o lar
b od ies are fo rm ed b ef o re fert ilizat ion wh ile t he egg is s t ill in ,

t et ra d fo rm is a lwa s c y h o s en fo r d esc rip tio n b ec a u s e t he d et a i s l a r e c a pa b e o f l

m o re d efin it e s ta t em en t . W ha t ev er t he f o rm o f t he m as s es , h wev er t h e fi n a
o , l
res u lt s eem s a l wa y s t he s a m e name ly d , a re u c t io n to o n e h l
a f t he u s u a n u m b er l
of c h ro m o so m es , a n d t hi s b y the m e th d f di
o o v is io n a n d ex t r u s io n .
1 In ra rein s t an c es t he p o a r b o l di h es a v e c o m m en c e dt o se
gm en t, b u t t e h y
n e v er p ro c ee f a r i n e d
v elo m en t
p d .

3 Wils o n , T he C e ll
, pp . 2 3 6- 2
37 .
F IG . 24 . D ia gra m s s h win g th e
o es s en t ia l fac ts in t he m a t u ra t io n of t h e eg .

T he s o m a t ic n u m b er o f C h ro m o s o m es is d
s u pp o s e to b e fo u r

A , i n it ia l p ha s e : t wo tet ra d s ha v e been l
fo r med i n t he ge r mi na v es c ic le B , the t wo .

tetra d s ha v e b een d ra wn u p a bo u t t he p l
s in d e t o fo rm t he e ua t o ri a
q p a te o f the fi rst l l
po la r m ito t ic figu re C t he m ito t ic figu re has rota ted in to po s it ion ea v in g the rema in s
.
, , l
o f t he
g e r m i n a l v es ic e a t
g l
v D ,
fo r m
. a t
. io n o f t he fi rs t p o a r b od
y : m c h tet ra dl
l
d iv i des in to t wo d yad s E , fi rs t po ar b o d y fo rmed : two d ya d s in it a n d a so in the
. l
egg . l
F , p repa ra t io n fo r t he sec o n d d iv i s ion G , sec o n d po a r b od y fo rm in g a n d the
l
fi rst d iv id i n g : ea c h d ya d d iv i des in t o t wo s in g e c hro moso mes H , fi n a res u t : three . l l
p ola r b o d ie s a n d t he ma t u re o v u m , e a c h c on t a in i n g t wo s i n g e c h ro m o so mes , l
o r ha lf

t he so mati c nu mb er ; c, t he egg c en t ros o me, whic h no w degen era tes an d is lost . A fter
Wil so n
1 7 0 C A U S ES OF V A R I ATI ON

A t t he proper s tage each s per mat o cyte u nd ergo es two d iv i

s io n s ( m at u rat io n d i v is io n s ) int o f o u r c ells ca lled s p er m at id s
, ,

each of which d ev elo p s a ta il and b ec o m es f u nct io na l in wh ich ,

fin is hed c o nd itio n it is kn o wn as a s per m at ozo on wh en it is ,

ready t o ent er and fert ilize t he ripened o v u m .

T he h is t o ry and d is trib u t io n o f t he chro m at in ma tter in t h e

.

male is id ent ical with that in the fem a le s o that each s p er m a ,

t o zo on i nh eri t s on e f ou r t/z t/ze c/zro ma t i n m a tter a nd on e ha lf t/ze

c /zr om os o m es o f t he o rig ina l c ell I n p lant s t he pro c es s d iff ers
.

b u t t he gen era l res u lt s a re t he s a me .

S ign ifi can c e of r edu ct ion On t he female s id e three f o u rth s o f

.

t he chro m at in m atter has b een extr u d ed in t he p o la r b o d ies and ,

t heref o re lo s t to t he lin e of d es c ent Wh eth er red u ct io n tak es

.

plac e by extru s io n o r by rearrangement o n e th ing is c ertain ,

‘

wh en t he s ec o nd d iv is io n is tran s v ers e and p o s s ib ly wh en it is

,

lo ngit u d ina l it res u lt s in an u n eq u a l d i v is ion of p/zy s i o log ic a l

,

u n i ts if the id ent ity o f t he chro m o s o m es and t he chro m at i n

,

g ran u les has any mean ing I f th is d iv is io n b e anyth ing else

.

than s tric tly q u alitat iv e then the ext ru s io n o f t he po lar bo d ies

,

mean s a los s o f s o m eth ing q u a litat iv e o n the f em a le s id e .

O n t he m ale s id e t he lo s s is n o t ab s o lu t e b eca u s e a ll fo u r ,

c ells are f u nct io nal b u t if red u ct io n has t he m ean in g we attach

,

t o it th es e f o u r s p e r mat o zo a a re n o t id ent ica l b u t d iff erent in

,

t he h ered i tary s u b s tanc e wit h wh ich th ey a re pr o v id ed .

S ign ifi cance of f ertili za tion Here th en a re t wo s ex u al c ells

.
, ,

ready for u n io n Each has lo s t large po rt io n s of its chro m a t in

.

m att er t he ev id ent v eh ic le o f tran s m is s io n and each bring s t o

, ,

t he u n io n b u t o n e ha lf t he n u m b er o f chro m o s o m es c ha rac t eris

t ic of i ts s pec ies s tro ng ly s u gg es t ing a lo s s o f c erta in chro m at i n
,

gran u les and t he h ered itary q u alit ies th ey repres ent ed .

Wh en f u s io n o f t he n u c lei o f th es e t wo ger m c ells tak es plac e

at fert ilizatio n h o wev er t he act o f u n io n aga in r es tor es thef u ll
, ,

a nd p r op er n u m ber of c /zr om os onzes wh ich will rem a in cha rac

,

t eri s tic o f t he n ew ind iv id u al thr o u gh o u t i t s lif e and wh ich it ,

will hand d o wn t o po s teri ty alway s thro u gh t he s a m e c o m p li

,

c a t ed m eth o d we ha v e att em pt ed t o d es cr i b e T he n u m ber of .

chro m oso mes is ev id ent ly k ept c o n s tant by the c o m plicat ed

proc ess o f red u ct io n d u r ing m at u rat io n and by fert ilizat io n
 I NTER N A L CAU SES O F V A R I AT ION 1 7 1

a ft er ward ; b u t what abo u t th eir charact er ? I n what c o nd it io n

ha v e th ey emerged fro m th is seem ing ly inc o mpreh en s ib le tang le
I s n a t u re as caref u l to preserv e th ei r q u a lity a s it is th eir n u m ber ?
What o ppo rt u n it ies fo r pro fo u nd v ariat io n C ertai n ly if
chro m at in m atter has any f u nda mental mean ing and if chro mo ,

s o mes a r e in any way representat iv e of phy s io logica l u n it s ,

an d if th ey i n th eir t u rn a re in any wa r epres entat iv e o f ra c ial

y
cha racters th ese pro c es ses m u s t ha v e so m e m ean ing in v a ria
,

tio n C erta in ly we hav e been v ery n ear in all th is to the m at erial

.

bas is of tran s mis s io n o f rac ial charact ers and to f u ndamenta l and
,

in it i al cau s es o f v ari at io n.

S ometh ing has been lo s t in t he t wo p ec u liar d iv is io n s attend

in g mat u rat io n S o m e d efi n i t e gro u p ings o f h ered itary s u b s tanc e
.

hav e d is appeared from the lin e o f d es c ent T h ey c ou ld n o t hav e

.

represent ed o rd inarily d efin it e po rt io n s of a b o dy b u t they m u s t

, , ,

represent so meth ing Wh at chanc es fo r acc id ent A n d in the

.
,

light o f th es e marv elo u s p hen o m ena d o we wo nd er that ind iv id

,

u als are so met i mes b o rn min u s a leg an a rm o r s o me o th er part

, ,

D o we wo nd er that v ita l part s a re s o o ft en a ffect ed and that o n e ,

t hird of ou r ch ildren die in infancy How m any d ie b efo re b irth ,

and how m any mo re d ie at s o me s tage in em bryo

Ev id ently all that is req u ired t o m ak e a liv ing being is a
fairly perfect d ev elo p ment of t he v i ta / pa r ts q u ite regard les s o f ,

t he pres enc e o r a b s enc e o f t he m any o th er rac ia l charact ers

that s h ou ld b e pres ent in t he p erfect ind iv id u al I s it su rpris ing .

that perfect in d iv i d u als are so few an d that d efect iv es a re

,

freq uent ly so far fro m the type ? Here is m at erial fo r s t u dy

o n t he part of c rim in o lo g is t s and c o u rt s o f j u s t ic e a s well as ,

s t u d ent s o f meth o d s o f ec o n o m ic i m pro v em ent .

R edu cti on an d f ert il izat ion in pl ant s I t m ay b e s a id in

.

genera l that in an im als the ev id enc e t end s to t he as s u m pt io n

that red u ct io n tak es p lac e at the extr u s ion o f the sec o nd po lar
body ; that each gro u p ( rod ring o r V s hap ed b o dy ) is in reali ty
, ,
-

a d o u bled (b iv alent ) chro moso m e and that t he fi rs t p olar body

,

remo v es o ne half of each (s plit ) chro moso m e wh ile t he n ext ,

rem ov es ev ery alt ernat e chro m osom e .

Wh ile t he fact s of red u ct io n in p lant s are yet in a h o p eles s

tangle it is s afe to s ay that t he ev id enc e t end s to s h o w that no
,
1 7 2 C A U S ES OF V A R I AT I O N

t ru e po lar b o d ies a re f o r m ed b u t th a t t he ch ro m o s o m es s u d ,
1

d en ly a ppear in red u c ed n u m b er a t t he fir s t d iv is io n a s if i t ,

were effect ed d irect ly by t he s eg m ent a t io n o f t he s p irem e

thread (o f the m at u r ing germ c ell) int o half t he s o ma t ic n u m ber
o f chro m o s o m es .

Wh ile d et a ils v a ry grea t ly b o tan is t s rec o g n ize t wo s tages in ,

t he d ev elo p m ent o f t he f em a le g e r m c ell o f t he p la nt n eith e r ,

o f wh ich is i d ent ical wit h m at u ra t io n i n an im a ls t ho u gh t he fi r s t ,

is fa i rly c o m parab le th eret o The fi r s t s t a ge o r S po ro gen es is

.
, ,

fo llo ws aft er that act iv e m as s ing o f f o o d m a t erial wh ich m ark s in

bo th p la nt and an im al t he preparat io n fo r red u ct io n A t th is .

t ime t he n u c leu s d iv id es q u ick ly int o f o u r daught er n u c lei each ,

o f wh ich is s u pp lied with ha lf t he n u m b er o f chr o m o s o m es that

charact er izes t he s pec ies T he prec is e m eth o d s f o llo wed o u t a re

.

m att er s o f m u ch d is p u t e a mo ng b o t a ni s t s b u t t he s ign ificant ,

fact is that red u ct io n is acc o m plis h ed at th is s tage .

O f th es e f o u r da u ght er n u c lei n o n e a re extr u d ed b u t three ,

o f th em d egen erat e in t he cyt o p la s m wh ile t he f o u rth incr ea s es ,

in s ize t o fo r m t he em bryo s a c wh ich with o u t wa it ing fo r f ert i , ,

liza t ion as a m o ng an im a l s c o nt i n u es t o d iv id,

e — c o m m o n ly ,

t wic e g iv ing rise t o eight s u b n u c lei wh ich arran ge th em

,
-
,

selv es in d efin it e p o s it io n s T wo o f t hes e s u b n u c lei rem a in

.
-

n ear t he c ent er o f t he em bryo s a c a n d g iv e ris e t o t he end o

s p er m ; three m igrat e t o t he e xt rem ity n ea r es t t he p o int o f

a ttach m ent with t he p is t il and o n e o f th es e ( and o n e o n ly ),

t he s o c a lled egg n c le u s
- u — u n it es with t he n u c leu s o f t he

po llen gra in to fo r m t he fert ilized ger m ; t he three rem ain ing

m igrat e t o t he o th er ext rem ity o f t he em bry o s ac an d c o nc ern
th em s elv es wit h es tab lis h ing a foo d s u pp ly with t he pa rent p la n t .

On t he m a le S id e t he pro c es s is s im p ler T he p o llen n u c leu s .

d iv id es o n e h a lf fo r ming t he po llen t u be alo ng wh ich t he o th er

, ,

ha lf tra v els d iv id ing aga in at so m e po int befo re u n it ing with t he

,

egg n u c leu s o f t he e m bry o s a c T h ese d iv is io n s a re ev id ent ly .

red u c ing d iv is ion s a s t he po llen gra in n u c leu s bri n gs t o the

,
-

u n io n a red u c ed n u m be r o f chro m o so m es .

D i p t d b y C h mb l i
1 s u e wh b l i a th t th
er a n , gg w it
o h it t h ep l e v es a
“
e e s ree o ar

b dio t it t
es c o n s g t i d i t l y
u es a m b
en era
p l w i
on t h t h
rec g m t p hy ti g co a ra e e a e o c en era

ti i pl l d X i i
” "
on n S B t i l G
a n ts . tt ee X X X I X 39 ;
o a n ca a ze e, , 1 s ee a s o u n er en a, n

th i t st ex .
 C AU S ES OF V A R I AT I ON
1 74

T he lat er d is c o v ery o f t he m echan is m o f m at u rat io n and o f

t he extr u s io n o f t he p o lar b od ies was a s t art ling c o nfi r m at io n o f
Weis m an n s pred ict io n and went far to fi x his th eo ries o f h ered
’

it y in t he m ind s o f m any b io lo gis t s I n th is c o nn ect io n W ilso n .

v ery pert i n ent ly rem ark s

T he fu lfillm en t of W ei s m a n n ’
s p red i c ti o n is o n e o f t he m os t in teres ti n g
res u l ts o f rec en t c yto lo gi c a l res ea rc h . I t h as b een d em o ns tra ted i n a m a n n er
w ich h seem s to b e in c o n tro v ertib le t h at the red u c in g d iv is io n s po s tu la ted
by W eis m a n n a c t u ally o c c u r , h
t ou g h no t p r ec is el yi n th e m a n n er c o n c ei v ed

b y hi m , b u t it rem a i n s q u i te a n o pen q u es tio n wh eth er t h ey h a v e the

s ig n i fi c a n c e a t trib u ted to t em h by W eis m a n n .

J ust wh en t he
red u ct io n occ u rs is n o t kno wn I t was at fi rs t .

as s u m ed that it o cc u r s in c o nn ect io n wi th t he extru s io n o f t he

s ec o nd p o lar bo dy an a ss u m pt io n b ased u po n t he d ev elo p ment

,

o f p a rth en og en et ic egg s B u t p lant s d o n o t f o r m p o lar bod ies

-
.
,

and aga in t here is great u nc erta inty as to wh eth er t he rod s

( t et rad s ) ri,
ng s o r V S hap ed b o d ies ( wh o se n u m b er is ha lf t he
,
-

u s u a l n u m b er o f chro m o s o m es ) a re t o b e r egard ed a s repres ent

in g t he u s u al n u m b er o f chro m o so m es s p lit and arranged in

pairs —
,
in wh ich cas e t he s ec o nd po lar b o dy wo u ld acc o m plis h
the red u ct io n ; o r wh eth er the chro m o s o mes as f o rm erly kn o wn
n ev er em erge fro m t he n u c leu s o f t he oocyt e so that the id en ,

t ity o f t he chro m o so mes is in so me way lo s t a n d t he red u ct io n

is effect ed at th is ear ly s tag e by s o m e s o rt o f int erna l f u s ing o r ,

perhap s by an ent ire rearrangement o f chro mat in gran u les On .

th is p o int the ev id enc e is c o nf u s ing b u t o n t wo s ign ificant ,

po int s th ere is n o d o u bt t he lo s s o f chro m at in m att er o u t o f

,

the lin e o f d es c ent and a red u ct io n o f t he chro m os o mes in the

,

ger m c ells t o o n e ha lf t he so mat ic n u m ber 2

.

Composit ion of t h e chr omosomes 3

I n v iew o f t he im p o rtant .

o ffi c e o f t he ch ro m o s o m es and t he m any th eo r ies o f h ered i ty

bas ed u p o n th eir nat u re and c o n s t it u t io n in v iew als o o f th eir ,

ev id ent i m p o rtanc e in all s t u d ies o n inh eri tanc e and v ariat io n ,

1 W ils o n , The C e ll , p . 2 46 .

“
3 By s o m a t ic n u m b er is m ea n t t he n u m b er c ha ra c t eris t ic o f t he s o m a ,

the bo dy
in gen era as l
is t in c t f ro m d the g er m in a l ma t t er w h ose f u n c t io n is n ot

g ro w t bhu t r ep ro u c t i on d .

l
3 W i s o n T he C e
, , pp
2 94 ll 0
3 4 .
-
.
 I NT E R N A L CA U S ES OF V A R I AT I O N 1 75

i t m ay b e well to no t e the s u bs tanc e o f what is rea lly kn o wn

abo u t their act u al c o n s t it u t io n .

T hat t hey a re n o t m as s es o f h o m o gen eo u s matt er is c erta in ,

an d that th ey c o n s is t o f n u m be r s of s m a ll gran u les capab le o f

m u lt ip licat io n a n d d iv is io n seem s eq u ally c erta in To q u o t e .

W ilso n
T h e fa c ts a re n o w well es ta b lis h ed ( ) 1 h
t a t in a la rge n u m b er o f c ases

the h ro m a tin th read c o ns is ts o f a se ie o f g a u les ( c h omoso m e )

c r s r n r s

i m b ed d ed i a nd h eld to gether b y the li n in s b s ta n c e ; ( 2 ) th a t the s plit

n u

ti n g o f the c h ro m oso me is c a u s ed b y the d i i io n o f th ese m o re elem e ta y

s v s n r

b od ies ; ( 3 ) tha t the c h om a t i g ra ins m a y d i id e a t the t im e wh en the

r n v

s pi reme is o n ly j ust b egi n n ing to emerge fro m the re ti c u lu m or res ti n g

s ta ge .

B ecau se th ese fact s th ere arises the s tro nges t t end en cy to

of

attach ind iv id u ality to t he chro m at in gran u les an d to c o nc eiv e

th em as b u ilt u p of d efi ni t e th ou gh o ft en d iv er s e phy s iolo gical
, ,

u ni t s t hu s c o n s t i t u t ing a s em i m echan ical b a s is fo r h ered ity and

,
-
,

i nc id ent ally fo r v ariatio n as well T h is a s s u m pt io n W eis m ann .

and o th ers hav e m ad e Wheth er the fact s s h o uld b e pu s h ed to

.

th is extreme int erpretat io n is in t he o p in io n of t he a u th o r as

, ,

e t u nc ertain T h e fact a re extrem ely s u gges t iv e t o s a t h

y . s
y e ,

leas t and it is c erta in ly n o t too m u ch to believ e that at th is

,

po int we hav e t o u ch ed t he phy s ical bas is o f life and in som e

fas h io n the v ery roo t of inh eri tanc e and v ariat io n ; i nd eed we
may proc eed u p o n t he c o n v ict io n that tra n s m is s io n is a f u nct io n
o f t he chro m at i n gran u les .

R edu ct i on as a cau se of v ar iati on The m o s t rem arkab le and .

s u gges t iv e fact ab o u t liv ing b eings is t he n u m eric a l c o n s ta ncy

o f ch ro m at in u n it s ( chro m o s o m es ) fo r each s pec ies and the ,

mos t remarkable and s u gges t iv e o f a ll t he v ita l proc es s es is

th eir red u ct io n befo re fert iliza t io n I f a s we s u pp ose t he .
, ,

chro moso mes a re the phys ical bas is o f inh er itanc e th en in tb e ,

loss o f clzr om a t i n m a tter at m a t u ra tion l i es a f u n da m en ta l c a u s e

f
o v a r i a t i on , a n d o n e q u it e in d epend ent of the ef e f ct s of f ert ili
za t ionaft erward .

R ed u ct io n wo u ld s eem t o b e a proc es s ca lc u lat ed t o in s u re

that n o t wo germ c ells ev en fro m t he s a m e ind iv id u a l s h o u ld
, ,

ev er b e a lik e and th is is t he m o s t ev id ent rea s o n fo r t he

,
 1 7 6 C A U S ES O F V A R I AT I ON

es s e nt ial d ifferenc es in ch ildren o f t he s am e parent s ev en in ,

t he cas e o f t win s
1
.

I t is tru e o f c o u r s e that n o t wo ind iv id u a ls ev en t wi n s can

, , , ,

b e d ev elo ped u nd er c o nd it io n s o f life exact ly id ent ical and yet

t he d ifferenc es o f c o n d it io n ca n n o t acc o u nt fo r t he fact that ,

wh ile o n e bro th er res em b les h is fath e r an o th er m ay res em b le ,

T wi id
n s a re c o n s d i i g f m
ere p
as a r s n t i t h ro f m l t i
s e a ra e o v a, as
p l n e c as e o u e

b i t h ( p ig d g
r s s, o d f
s, an h ib iti g t h d i ti t b p t d
, o c o u rs e, a s ex n e ev a o ns o e ex ec e

f m d iff
ro t g m
ere n d d i t i t f t il i t i
er s an s i lit t
nc g
er ll y
za o n , as n e rs en era .

S m t wio e h w ns, o ev e r, l y l ik ( id t i l twi ) t gg t t h

a re s o n ea r a e en ca ns as o su es e

p o ssib il it y f t h i
o h i g e r i f
av n m i l
a r s en
g m i ro m wa s n
y p t d i
e ov ut n so e a se a ra e n o

h l a v es t it a fi t l s rsg h
c ea v ah l f d e, l p i ea c
g i d i iad l T h
ev e o i i w i n an n v ua . s v e s

ev id tl y f
en d b y G dd
a v o re d Th m e (
es a n Th E l t i o f
so n S p s ee e vo u on o ex , .

I f t wi h l d b d l p d i th i m
ns s ou e ev e o eth y w n l d id t l y b f t h
s a n n e r, e ou ev en e o e

n ea res t p ib l oss im il it y f e s th y ar p , t b or t e m re d
res enb t i gl u o ne ovu an u a s n e

f t il i t i
er za on .

Th i p s ib i l it y i
oss pp t d b y th s su p i
or e m t f R E e exd er d W l ten s o o u x, n res , an a er,

i wh i h
n c h bl t m
ea c f t h tw
as o ll t g f th f g m im ( t l w y )
ere o e o -c e s a e o e ro so et es no a a s

is ca
p b la f d e o l p g i i
ev e ot p n f t i d i i
n o a d l D i
e r ec h w k
n i g w i t h v h i ua . r es c , or n ec no

d m er s, t b l i h d th
es a s m e f t e wh i h sa e l w ll k w i th
ac s , c a re a s o f e no n n e c ase o

A mp / i r ox u s ( W i l s ee T h C ll p so n, e e , .

C o n v ers e ly wh tw
, f t il i den o f er hi ze A ova i o dh s ea i u rc n, or sc a r s , a ere ac c

d t ll y th y m y d l p i t
en a , e a ev e o mb y f n o l d im i ( L b
an e r o o u n us u a en s o n s s ee oe ,

S t diu es i G n l Phy i l gy P t I I p
e n era s o o , ar , .

Wh h w
en , o th b l ev er, t m e t
as o p t d b t
eres a re n o f t h m i k
se i ll d
a ra e , u one o e s e

by a h t d d l ( R ) th
ea e n ee e i joux d h lf l , e d l p b t it p d
un n u re a a o ne ev e o s, u ro u c es

a t th b t
e es ki d f h lf l a n o ( igh t a l f h l f)a rv a r t i i g or m d ll ye t a ,
“
con a n n one e u ar

f ld
o , one au d it y p it ( W il or T h C ll p so n , Ch D i e e h M g , . d un , r es c , or an , an

F i h l w k i g wi t h t
sc e , or n
p h gg c en oh w f
o re e d t h t i s, l t o d b l t m
e v er , o un a so a e as o eres

o f t h tw e f o o urigh t ll t g d l p d d f t i l
or e -
ce s a es ev e o h i g ly e
“
e ec ve e,
a rv z av n on

fo u r, tw o , o r o n e ro w f wim m i g p l t o s Al C mp t
n f
a es d th t i th
.
"
so ra on oun a n e

c as e o f th m i e g t
ar ne p d I l a s e ro i l t d bl t m
o r a n a s sa f tw ll f
so a e as o e res o o -c e or our

ce ll t g s a m
es s e
g t d en el y i f f mi g p t f
e xa c t as ti or mb y n dg ar i o a n en re e r o , an av e r se

t f
o rag m t f
en s o l t t
a mp l t dw f
a rv a , n o o co i th h i d m ( W il
e e ar s as n e ec no er so n,

T h C ll pe e , .Th i t t mp t t f ms a ti e i d i id l f m p ti
o or en ly
re n v ua s ro a or on o n

o f af t il i d gg
er ze l t i g i t ft d i d w f
e , res u n as o m t th w i t
en o es n ar s, s ee s o e r er a

p l
ro c es s c o s e l y k i t ag n t i o re
( w h i h en e ra i h pton R l ti
c S t b il it y
s ee n c a er o n e a ve a

an d I t b i l it y f L i i g
ns a o v n d w ld m t i an d bt t
ou it s ee o ra s e ou s as o s

s u c c es s u f l i t h h igh
o c c u rren c e n im l e er a n a s .

Th ou
g h t h no i g p i
ea r n ll y p es t h p i
ec a t i q u t i ont h m t t
e f t wi
o n n u es o n, e a er o ns

in ttl
ca i e iq s un d w t hy f m
ue an ti or Th k i d f twi
o en on .k w i ree n s o ns a re no n n

ca t tl e: ( ) t h r t w i me
y b b ns t h f m l
a d e b t h o m l e
; ( )
a e an t h o no r a or 2 e s ex es

m y b d iff
a e t d
e ren m l; an ( 3) b th m y h m l
nor a or i wh i h o alw y e a es , n c c as e o ne a a s

ex h ib it th p l i b m l ity k w
s e ec u ar a no r f a m ti no th n i t
as a l g ree- ar n,
’
e n e rn a o r ans

a re m l b t tha e, u t l e ex ern a y g f m l
a c c ess o r d th or an s l a re di e a e, a n e re a re a s o ru

m t y f m l d t
en a r e a e ( G dd
uc s d T h m e Thes E an l t i f So p so n , e vo u on o ex , .

Thi i s s ak i d f h m p h d i i m d t i m m l y pp d
n o er a ro t s , an no , h ifas s co on su ose ,
“
a e er

t wi wit hn b ll a u .
 1 7 8 C AU S ES O F V A RI AT I ON

female and her ger m c ell a re in v o lv ed B u t ind iv id u als th u s .

aris ing thro u g h u n is ex u a l repro d u ct io n v a ry w id ely a fa ct ea s ily ,

cred it ed wh en the ph eno mena o f red u ct io n a re rem em bered .

Pa rth en ogen es is bei ng lim it ed to lo wer a n i m als t he rang e and ,

chara ct er of v ariat io n s a re fo r the m os t part d iffi c ult of d et ect io n

and meas u rement I t is kn o wn h owev er that great d ifferenc es .
, ,

i n s ize o cc u r a m o ng ind iv id u a ls parth en o gen et ica lly pro d u c ed ,

a n d charact er s g en erally a re s o v ar iab le in s u ch i nd iv id u a ls

as t o lead t o t he s tat em ent that t he v ariab ili ty o f o ff s pri ng

n ev er ma tes b u t o n c e) s he is , o f c o u rs e, c a p a b le o f la yi ng o n ly u n fert ilize d or d ro n e

eggs
"
. h
I n t is wa y, in c ro ss in g t h e , d ron es an d t he wo r k ers may ac t u a l lyb e of

differen t b ree s d .

P l an t lic e rep ro d uc e p a rt hen o gen et ic a l ly d u rin g t he s u m m er s ea s o n , p o

r d u c in
g
on ly f m l ; b t e a es u as t h e t em p era t u re lo wers wi t h a p p ro a c hi n g au t u m n a m ix e d
b d f b th m l
ro o o o a es an d fema les h h p m ti g p d a p p ea rs , w ic th l , u g on a n , ro u c es e on

li v d wi t
e , d i e n er- n g gg I t
ur n i e t w st hy
. t h t st h no e
p t h org t i gg a f e ar en o en e c e s o

b eesd l pm l ev e o ly wh i l t h
a es on , f p l t li e d l p f m l d i g t he
ose o an ce ev e o e a es ur n

su mm b th er, o pp i
s e x es a
g t m
ea r n p p h
as a u u n a ro a c es .

It i l t w t hy t h t
s a so n o e ord t h tifi i l h t f g h a un er e ar c a pp im t
ea o reen o u s es , a ro x a

i gp p t l
n er e u a mm su d iti p t h g e i
er c o n ons, tiar i d fi it ly
en o en s s c o n d m l n u es n e n e , an a es

a re n o tp d d l th p l t b m b d ly d i d p
ro uc e u n es s e an s ec o e a r e u .

P the g
ar no i diffe en es s g t l y i d
rs g reaI t i pp
n d t
e ree b . m p l t i s su ose o e co e e n

t i mi
c er a n t t
n u e c r u s a c eans a ndi m y tif m n g w anh i h ro m l ers a h on c no a es a v e ev er

b een f d It i
o un .
”
l s
“
i th
s eas o n a
p h i (p
”
l t l
n i ) d
e a p ts i l i t
anh ce , an
“
ar a
”
n e

h ybone d i
ee a n m “
f the l w
n so e o im l wh i h o er a n t t h em l
a s m lly c are n o s e v es n o r a

p art h en o
g t i b
en e t h e
c, l tui avw h i h re a Ov es i l pc t h a re g .
”
i h c c as o n a ar en o en es s as

b een f eq r t ly u en b d A o s erv e m p l i th e ilk m t h i wh i h N

. n e xa e s b s m o , n c u ss au

fo un d th t t f a ou of til i d gg
1 1 0 2 u n er d l p d
ze e s p t 22 t i ev e o e u o a c er a n

p i
o n t I t
. i pp s su d t h t i osel l a f p n at h c as es o
g i m y e gg
ar f i l t
en o d len es s
p an s a o ev e o .

I th i n ti
s c o n n ec it i t w t hy
on d
s no e t m ly or gg t i
an t h t
ex re m g gh i eh su es ve a a on er

an im l —f ga s h ro d e
s, en s , m mm l — t he
an v en f t il i d
a a s m i llyu n er ze ov u o c cas o n a

b gie n s seg m t t i en a p d
o n , n ev eri g f h w ro c eee i t
n p t h e
a r, g o t i ev r, o n s ar no en e c c o u rs e.

Th t d
e s u t h l d de t d th t i ll p b b il it y l g mb f egg
en s ou un rs a n a n a ro a a ar e n u er o s

f il t d
a o l p i tev e o m p l t i d i id l
n o co e e i th m t
n v ua s f l p th
ev en n e os s u c c es s u ar en

o g en es s ,i j t d m y f t ili d
us as o an f il l g th w y ( e W i m
er ze ov a a a on E y e a s e e s an n , ss a s

on He ed ity I r , Th , ll d g ere a re a f p th g i
e rees o f m t h p far t l y en o en es s , ro e er ec

s u c c es s u f ld w t o n o zero .

B i g p
ea r n th u
g on l b
e j t
en era t h e i su t t iec g a re
p im t f L
n eres h i n ex er en s o oc n

ar t ifi i l p t h
c a ar g i
en o p i ll y
en es s , es f t h ec a h i o m ll y b i
e s ea u rc l b t w h i h
n, nor a s ex u a , u c ,

a ft er im me i i rs o n li l ti
n a sa f h igh d
n e so u it y on o d b q t t en s t an su se u en re u rn o nor

m l e w t
a s a mm
a er, d gm t ti
co en c e d ft w d d
se en a l p d li i g l
o n an a er ar ev e o e v n e.
a rv a

M g a i m
n es u d p t an i m lt p
o as s u d m t ff t i th gh i g
sa s ro v e os le y
ec v e, ou n en era an

t me t il th t i t/m w f m f i l i d
'
t rea n av a s t a s erv es p t i
o ro
q f fi t
r a ro a t / or ont re a er ro ze u n er ze

eg g ( e L eb i A m i
se o n j l
er ca n ff P l y i l
ou r n a
g I I I 4 3 4
( ; l L
a s o o b S t d i v,s a so oe u e
—
, ,

i G
n en era l Phy i l gy P t I I p p 5 7 6 6 6 63 8—
s o o , ar , 69 ; M
. t h d pp 7 7 7 ;
-
2 , 66 2 e o s, . 2

G dd e
e d Th m
s an E l ti
o so n , f S vo u
pp 1 8 3on o ex , .
 I N TE R N A L C AU SES O F V A R I A T I O N 1 79

as ex u ally reprod u c ed is no t immen s ely red u c ed belo w t he v ari “

ab ility o f t he rac e 1
.
”

I n t he h o neybee o n ly t he male s ex is prod u c ed pa rt heno ge

n et ic a lly I n p lant lic e it is c o m mo n ly t he f em ale alo n e u nd er
.

h igh t em pera t u re and bo th sex es u nd er lo wer Weis man n bred

, .

sepa rat ely t wo v arieties o f Cyp r is r e p ta n s f o r s o m e s ev en y e ar s ,

c o v ering mo re than fo rt y gen erat io n s and m any th o u s and ind i

v id u als .On e v ariety A was light in c o lo r ; t he o th er B was
, , , ,

dark N o ma les were ev er d is c o v ered in eith er and it is s u p

.
,

po sed that th is s pec ies prod u c es o nly parth en ogen et ically Whi le .
,

fo r t he m os t part the d es c endant s of each were extremely a lik e

, ,

y e t m i n u t e d iff erenc es in v ariab ly ex is t ed .

N o t o n ly was th is tru e b u t in 1 8 8 7 three y ears after t he , ,

expe rim ent c o m m enc ed s o me i nd iv id u a ls o f t he dark green

“
,

v ariety B appeared in t he aq u ariu m with t he light v ariety

, ,
.

T he s a m e v ariat io n app eared a s ec o nd and a th ird t ime and in ,

t he las t in s tanc e int er m ed iat e f o rm s c o u ld b e m ad e o u t In .

1 8 9 1 an o th er c as e oc c u rr ed and in t he s a me y ear it s c o n v ers e

—
,

appeared a few typ ica l light i nd iv id u als a m o ng t he dark

,

c o lo ny tha t had bred tru e many y ears “ 3

D oes th is experi .
”

m ent a ls o thro w light o n t he o rigi n of v a riet ies and were th ese ,

m u tat io n s ? Ho wev er th is m ay b e it c learly s h o ws t hat v ari ,

ability is n o t ent irely d epend ent u po n sex u al u n io n and that ,

ev en d is t i nct v ar iet ies m ay aris e wi th o u t t he i nt erv ent io n o f s ex .

T he fi rs t s ign ificant fact in m at u rat io n o f parth en ogen et ic

egg s is that th ey pr o d u c e b u t o n e po lar b o dy
3
.

F ro m th is po int o n t wo a lt ernat iv es s eem po ss ib le I n the .

fi rs t p lac e a s ec o nd p o lar b o dy app ear s to b e f o r ming in t he

,

u s u a l m ann er and t he s eparat io n o f t he n u c lear m att er tak es

p lac e b u t ins t ead o f pas s ing o u t o f t he egg it rema i ns be/zi nd

, ,

f u s i ng g
a a i n w i t]: th e n u c leu s f
o tb e egg pr o pe r w h ich s traight ,

way u nd ergoes d ev elo p m ent with it s chro mo s o mes i ncrea sed to ,

1 Pea rs o n , G ra m m a r of S c ien c e, p p 4 7 2 4 7 3
.
-
.

3 W eis ma n n , Ess a y s on d y
Here it , I , 1 6 1 - 1 64 .

'
3 h
Oft en t is p o a r b o l dy d d
iv i es giv in g t he a p pea ra n c e o f t wo , b u t a s ec o n o n e
, d
is n o t fo rm e d
I t is q u ite rem a r a b e t o u g en t ire
. k l h
c o n s is t e n t in t h e c as e o f
, h ly ,

h h
a p is, y
o n e b ees , a n d c ert a in o t er fo rm s t a t
p ro u c e b h
o t s e xu a a h
n d as ex u d h ll y
y h d l
a ll , t a t t he f er ti lized egg s p ro u c e t wo p o a r b o ies b u t t he p a r t h en og en eti c eggs d
on ly on e.
1 80 C A US ES O F VA R I AT ION

t he pro per n u m ber I n th is cas e t he s ec o nd po lar body appears

.

in t he r ole o f a m a le elem ent so we m ay s peak o f t h is as a k i nd ,

of
“
f er t i l iz a t ion by tlze s econ d pola r b ody .

I n t he th er f o rm o f parth enogen es is h o wev er th ere is litt le

o , ,

su gg es t io n o f a s ec o nd po lar bo dy ; c erta in ly n o fo r m a l s epara

t io n and lat er f u s ing o f n u c lea r m a tter tak es p lac e On t he c o n .

t ra ry d ev elo p m ent tak es p la c e d i rect ly u po n t he extr u s io n o f

,

t he fi r s t po lar b od y and i t is S ign ificant that i n d i v id u a ls a r is i ng

,

i n tlzis way p oss ess b u t lza lf t/ze n u m ber of c lzrom os om es a s c om

r ed wi t/z tlt os e a r is i ng by s ex u a l r ep r od u c tion o r by tlze m et/rod
p a

j u st d esc ri bed c ies (as in A rt em ia )

. So m e r epr o d u c e pa r
s pe
1

t hen o gen et ic a lly by b ot/z m eth o d s g iv ing ris e t o t wo d is t inct ,

v ar iet ies o ne wi th ha lf t he n u m be r o f chro m o so mes C haract er

,

is tic of t he o th er “ .

M uta ti on as r ela t ed t o red u cti on an d f ertil i zat ion M u tant s .

s eem t o b e d epart u res charac t e r ized by a s u dd en lo s s o f s o me

rac ia l cha ract er o r by it s po s s es s io n in s o m e u n u s u a l d egree .

T h ey d o n o t appear to b e end o wed with ch a ract ers n ew t o t he

rac e exc ept wh en art ific ia lly prod u c ed by hybr id izat io n
,
.

I f t he pro c es s o f red u ct io n m ean s t he lo s s o f h ered it a ry

m at er ia l and if fert ilizat io n m ean s its res t o rat io n and if eith er
, ,

m ean s in any s en s e n ew c o m b inat io n s th en we can s ee in t he ,

t wo ph en o mena tak en t o geth er o r ev en s ing ly ab u ndant o p po r , ,

t u n it y fo r t he m o s t pro f o u nd v aria t io n s ; ind eed a d m itt ing th eir ,

po s s ib ility thro u gh th es e c a u s es t he wo nd er is t hat th ey a re n ot ,

et m o r e c o m m o n and infin it ely m o re rem arkab le

y .

I f th ere is in a ny s en s e h o wev er S light a q u a litat iv e lo s s , ,

thro u gh red u ct io n th en by t he la w o f chanc e t he t im e is c ertain

,

t o c o m e wh en s o m eth ing u n u s u a l will appear I s it n o t m o r e .

than lik ely that h ere lies a fr u itf u l so u rc e o f s weep ing cha n ges ,

a s well a s o f th e m o re o b s c u re d iff erenc es th a t a re ev ery wh ere

abo u t u s A n d is it n o t lik ely that s t ill great er and m o re fre

?

q u ent changes wo u ld pres ent th em s elv es were it n o t that fert ili

za t ion is fo r t he m o s t part r es t r ict ed t o c o m parat iv ely narro w

li nes ?

1 W il s o n , T he C e ll , pp . 282 —
2 83 .

2 A rt emis t h us v a r ies f ro m 8 4 t o 1 68 , di
ac c o r ng to t he p a rt ic u ar l m et h d
o

o b s erv e d .
1 82 CA U SES OF V A R IAT ION

liab ility to intern alchange and n o t at all as is erro neo u sly s u p ,

p os ed fo r t he inh eritanc e of acq u ired cha racters beca u s e t he

, ,

charact ers in q u es t io n were no t acq u ired in t he o rd ina ry “

acc eptanc e o f t he term — th ey were the resu lt o f internal n o t

, ,

ext ern al im p u lses

,
.

SECT I O N V — I N FLU EN C E OF T HE C ON D I T I ON O F THE

G E R M U PO N D EV E LOPM ENT

S tal en ess . Bo th t he ma le
and the fem a le germ c ells a re
capab le o f liv ing fo r a c o n s id era b le t ime after m at u rat io n s o ,

that f ert ilizat io n may b e so m ewhat d elayed ; ho w lo ng is no t

k n o wn and what t he effect of d elay m ay b e is n o t f u lly
,

u nd er s t ood .

E xperi ment s by V ern o n u po n t he o v a and s per m at ozo a o f t he

s ea u rch in o f d iff erent ages fro m n in e t o fo rty fi v e h o u r s i n d ic a t e
,
-
,

that t he S ize o f the larv a is in so m e d egree d epen d ent u po n the

fres hn ess o f t he germ at f ert ilizat io n T he res u lt s o f a n u m ber
‘
.

o f tr ia ls were as f o llo ws

1 With s ta le o v a and s tale s per m t he res u lt ing larv ae d iff ered

.

b u t s light ly fro m t he n o r ma l ( in wh ich bo th were fres h ) .

2 With fres h o v a and s t ale s per m t he la rv ae were d is t inct ly

.

larger pe r c ent ) .

3 W
. i th s t a le o v a and fr es h s pe r m t he la rv ae w e r e d is t i nct ly
s m a ller than wh en b o th were fres h c nt
p er e ) .

I t is c ertai n that t he ab o v e c o m b inat io n s as t o s ta lenes s a re

poss ib le in t he fert ilizat io n o f m am m als by m at ing and of plant s
by p ollinat io n Wh eth er the res u lt s a re the S a m e and wh eth er
.

t he d ifferenc es pers is t throu gh lif e a re o f c o u rs e u nkn o wn T he , ,

.

fact s rec o rd ed a re s u gges t iv e b u t wh eth er th ey will ev er b e ,

u s ef u l r ema i n s t o b e d et er m in ed .

I n di v idu alit y of t h e ger m T hat s u cc es s iv e germ c ells fro m

.

t he s a m e ind iv id u a l m ay b e s u b s t a nt ia lly d ifferent e v en as id e ,

fro m c o n s id erat io n s o f m at u rat io n is a f act beyo nd q u es t io n ,

.

T he c a r o f c o rn lik e it s ta s s el m at u res fro m t he ba s e u p ward

, ,
.

T he tip k ern els a re no t o n ly y o u nger b u t d ec id ed ly s ma ller than

th eir half s is t ers at the bas e The d ifferent pea s in a po d a re
-
.

V 1 V i ti i A im l
ern o n , d Pl t pp
ar a on
5
n 8 n a s an an s , . 10 -
10 .
 I N T ER N A L C AU SES OF V A R IAT I ON 183

q y d ev eloped On e of t he t win pai r o f oat s is m o re o r

no t e u a ll .

les s u nd ev elo ped I s th is d ifferenc e in s ize d u e to s eas o n foo d

.
,

su pp ly r oo m o r t o s o me pec u liari ty in the ger m

, ,
I t may b e lack
o f r oo m i n po d b ea ring p lant s b u t it cann o t b e that in t he ca s e
-
,

of c o rn The s tro ng pres u m pt io n is in the o p in io n of t he writ er

.
, ,

that thes e d iff erenc es in s ize a re part ly d u e t o d iff erenc es in food

s u pp ly b u t mo re largely t o inh erent d i fferen c es in t he ger m s .

SECT I ON V I —X EN I A , OR FERTI LI ZAT I ON OF T HE

E N DOSPE R M, DOU BLE F ERTI LI ZAT I ON

If o ne k ind of c o rn
fert ilized by ano th er the mixt u re will
be ,

s h o w t he fi r s t y ea r Fo r exa m p le if wh it e and y ello w c o rn b e

.
,

plant ed s id e by S id e the wh it e c ars will ha v e many yello w g ra in s

, ,

s h o wi ng at o nc e t he eff ect s o f cros s fert ilizat io n T h es e o ff .

kern els a re t he m ix ed s eed s b u t reas o n in g fro m a na logy we , , ,

s h o u ld n o t expect t he m i xt u re t o appear u nt il t he gra in s a re

plant ed and the genera t io n of m ix ed breed ing is at hand The .

v is ib le part o f t he k ern el is n o t t he germ ; end o

“
it is t he
s pe rm ,
o r s u rro u nd ing po rt io n wh ich s erv es as food fo r t he
,

s pro u t u nt il t he y o u n g p lant ha s es tab lis h ed it s elf I t is rela t ed .

t o the germ m u ch as t he wh it e o f egg and i ts S h ell a re relat ed

to t he y o lk Fert ilizat io n is o f t he g erm
. Ho w th en d o th ese .
, ,

o u t s id e part s bec o m e a ffect ed

I t will b e rem em bered that in the an im al the fem ale ger m

giv es rise to o n e m at u re f u nct io nal c ell t he o v u m and th ree , ,

n o n f u n c t io na l
-
t he p o lar bo d ies ; that t he m a le c ell giv es ris e
,

also to f o u r mat u re c ells t he s perm at ozoa all fu nct io nal and

, , ,

that the n u c leu s o f the o n e u n it es d irect ly with that of the

o th er wi t/t o u t i n ter v en i ng n u c lea r d i v is ion s .

I n plant s h o wev er it is f o u nd t o b e s u b s tant ia lly d ifferent

, , .

The mat u re fem a le c ell c o rres po nd ing to the o v u m in s t ead o f

, ,

a wait ing f ert ilizat io n c o nt in ues its act iv ity u nd ergo ing gener
, ,

ally two (so met im es m o re) d iv is ion s o f the n u c leu s g iv ing rise ,

t o eight o r so m e o th er c o rres p o nd ing n u m b er o f

,
s u b n u c lei -
,

which rem a i n flo at ing with in the cyt o p la s m I t will b e rem em .

bered that of th ese eight s u b n u c lei o n ly o n e is capable of f u nc-

t io n in g as an egg n u c leu s ; a ls o that t wo o th ers rem ai n n ear

1 84 C A U SES O F V A R IA TI O N

t he c ent er o f t he em bryo sac to fo r m t he end o s per m I t will .

b e remem b ered a ls o that t he po llen n u c leu s u nd ergo es a s ec o nd

, ,

d iv is io n d u ri ng it s progres s d o wn t he po llen t u be and befo re

u n it ing with t he egg n u c leu s .

Of th is d i v id ed n u c leu s o n e po rt io n u n it es with t he S ingle

f u nct io nal mem be r o f t he fema le gro u p m ak ing t he ger m and , ,

i n cas es s u ch as a re n o w u nd er c o n s id era t io n t he o th er j o in s
with t he min o r mem b ers c o nc erned with t he d ev elo p m ent o f
t he end o s pe r m I n th is way by m ean s o f t h is k ind o f d o u b le
.
,

fert ilizat io n t he end o s perm is it s elf a ffect ed and t he c ro s s ing is

,

ev id ent the fi r s t y ear .

Of c o u r s e th is V is ib le e ffect u po n the end o s per m is o f it s elf

p u rely t ra ns ito ry ha v ing n o infl u enc e u po n t he line o f d esc ent
, .

T he rea l eff ect o f po llinat io n is m a n if es t ly a s i n all o th er fer t i ,

liza t io n c o nfin ed t o t he germ

,
.

Wh ether t he t wo fert ilizat io n s a re s im ilar as to c o m pa rat iv e

infl u enc e o f t he t wo parent s n o o n e kn o ws n o r d o es it great ly ,

m a t t er T he eff ect u po n t he end o s per m enab les u s t o d et ect

.

t he cro s s if it is capab le o f d et ect io n and to rem o v e t he c o n

, ,

t a m in a ted s eed s if we d es i re t o reta in p u r ity I f t he o bj ec t b e .

t o s ec u re cro s s ing we s ha ll o f c o u rs e s u b s eq u ent ly d ea l wit h

,

t he pro d u ct s o f t he cro s s bred germ wh ic h o n ly a re S ign i ficant

-
,

fro m t he breed er s s tandpo int ’

.

J u s t what s pec ies ind u lge in th is d o u ble f ert iliza t io n is n o t

well kn o wn I t is ho wev er well es tab lis h ed in a large n u m be r
.
, , ,

and the proc es s is s u ppo s ed to b e c o mmo n rath er tha n u n u s u a l .

Effec t of crossin g u pon f ru it in gen er al What t he lay m an ca lls .

fr u it is c o m mo n ly n o t t he end o s per m that has been u nd er d is

c u s s io n b u t t he th ick en ed and m u ch d ev elo ped fl es hy rec eptac le

o n wh ich t he s eed s a re ho m e I t has been c la imed that th ese .

part s a re d irect ly infl u enc ed the fi rs t y ear by cro s s ing so that the ,

charact er o f s tra wb erries app les pear s m elon s s q u a s h es etc

, , , , ,
.
,

d epend s m u ch U po n t he s o u rc e o f t he po llen u s ed in f ert ilizat io n .

T h is c la im ha s n ev er b een well s u b s tant iat ed by d irect experi

m ent .D r B u r rill o f t he U n iv ers ity o f I llin o is t ells m e that he
.
, ,

cro s sed C res c ent s t ra wberries bo th with t he S ha r p les s and wit h

a wild berry es pec ia lly select ed fo r its in s ign ificant wo rt h less ,

fr u it N o bo dy was able t o d et ect the d ifferenc e in the res u lt ing

.
I 86 C A U SES OF V A R I AT I O N

i n s tan c e Lo rd Mo rt o n m are m ent io n ed by D ar win T h is

’
l
of s .

m are bo re a c o lt by a qu agga wh ich was o f c o u rse s triped a ft er ,

the m ann er o f his S i re S he aft erward s b o re t wo c o lt s by a

.

s ta llio n bo th o f wh ich were sa id to hav e been m ark ed with

,

bars o n s h o u ld ers and legs s u ppo sed ly s h o wing t he effect s o f

t he q u agga u po n t he o ffs pring o f t he s ta llio n .

Pro f es s o r Ewart of Ed i nb u rgh has rec ent ly rep eat ed th is

, ,

exp er im ent o n an ext end ed s ca le with res u lt s S h o wing n o trac e ,

o f t he q u a gga b ey o nd his o wn o ff s pring R ec ent in v es t igat io n s

2
.

in c o nt em p o rary lit erat u re thro w g ra v e d ou bt u po n t he ess ent ia l

acc u racy of t he data at D arwin s hand T hey seem to S h o w ’
.

that t he s u ppo sed resem b lanc e o f t he s tallio n c o lt s to t he q u agga

wa s exc eed ing ly fanc if u l pro bab ly being n o th ing b ey o nd what
,

appears freq u ent ly in yo u ng h o rs es o f t he p u res t parentage ,

d u n c o lo red h o rs es a s a r u le s h o wing m o re o r les s t end ency t o

-

s tri pes and bar s .

I t is o n e o f t he b es t ev id enc es o f t he p o wer o f trad it io n that

th is s ingle ins tanc e happen ing m o re than a h u ndred years ago
, ,

has d o n e d u ty ev er s inc e to pro v e an exc eed ingly d o u btf u l

th eo ry and an alm o s t u nacc o u nt a b le b elief I t is rema rkab le .

that s o u nc ertain a c irc u m s t a nc e and o n e s o eas y o f repet it io n , ,

wi th u n iv ers a l expe rienc e t end ing c o n s tant ly to throw light u po n

t he s u bj ect s h o u ld ha v e b een s o exc es s iv ely o v erwo rk ed
,
It .

s h o ws as n o o th er i n s tanc e has ev er s h o wn
,
t he per s is t enc e o f ,

trad it io n t he ext ent o f cred u lity in the presenc e o f the ph en o m

,

ena l and t he willingn es s o f m en to repeat an a s s ert io n o r ev en

, ,

an o p in io n u nt il by m ere repet it io n it c o mes t o ha v e all the

,

fo rc e o f a u th o r ity T he th a nk s o f t he wo rld a re d u e to Pro fes s o r

.

E wart fo r his exc ellent wo rk in d is po s ing by d i rect experiment , ,

o f a c itat io n that has d o n e da m age lo ng en o u gh I t is to b e .

h o ped that t he q u es t io n may at leas t b e h eld o pen u n t il so m e

s o rt of p o s it i v e ev id enc e is bro u ght f o r ward that is wo rthy t he

cred enc e of caref u l s t u d ent s .

Tel egony in dogs D og fanc iers a re pretty gen era lly cred it ed
.

wi th believ i ng in t elego ny es pec ia lly in ca s e o f fi rs t m at ings , .

1 S ee D a rwi n , A n im a l s an d P a n tsl un d er D o m es t ica t io n , c h a p . x iii , p . 1 7 , of

s ec o n d diti
e on by A pp l et o n .
(Q uo te d fro m P b i/os op / nca l Tr a n s a c tion s , 1 82 1 ,
’
2 B r eed er s G a zette, X LI ,
p . 1 009 .
 I NT ER N A L C AU SES OF V A R I ATI ON 1 87

The bes t s t u d ent s h owev er in s is t that v ery litt le real ev id enc e

, ,

has been pro d u c ed o n the s u bj ect and mi n e at a ll t end ing t o ,

prov e the ex is t enc e o f th is in flu en c e 1

.

With a v iew to t es ting s o mewhat t he real ext ent o f th is b e

lief t he a u th o r addres sed lett ers t o the bes t kn o wn d og fan
,
-

c iers o f t he U n it ed S tat es Of th irty sev en an s wers rec eiv ed

.
-
,

o n e wri t er is a believ er in t elego ny ; six so mewhat m ild ly

express u nc erta inty ; t wo a re n o n c o mmitta l ; and t wenty eight - -

a r e o u t s po k en aga in s t t he th eo ry T he mos t o u t s po k en o f th em
.

a ll is a well kn o wn fanc ier o f lo ng exper ienc e

-
J u dging fro m .

th is s m a ll n u m ber it wo u ld s eem that th is belief a mo ng dog

,

fanc iers has b een ov errated .

Pr oof b y t h e met hod of in stance W ith o u t a reas o n ab le d o u bt .

belief in t elego n ic infl uenc e res t s u po n s t ray in s tanc es d iffic ult ,

o f u nd ers t and i ng by th os e who happ en ed t o b e t he o b s erv er s ,

and has t ily acc epted as ev id enc e N o w n obody s h o u ld b e mo re .

caref u l than the breed er t o j u dge acc u rat ely the nat u re and
v alu e o f ev id enc e A s ingle i n s tanc e may b e goo d n egativ e
.

t es timo ny b u t it is seld o m wo rth m u ch as po sit iv e ev id enc e

, .

The prod u ct s o f breed ing a re s o m any and s o v ariou s and the ,

ca u ses o f v ariat io n are so n u m ero u s and so c o m plicated that a ,

part ic u lar res u lt can seld om b e as s igned to the o perat io n o f any

s ingle cau se I t is mo re lik ely t he m ix ed o r c o m po s it e res u lt
.

o f m any infl u enc es b o th i nt ern a l and externa l ; and in o rd e r to

,

kno w t he effect o f a s ingle ca u s e it is n ec essary to iso late t he

cas e if po ss ible o r if n o t to reso rt t o t he exam inat io n o f la rge
, , ,

n u m bers o f cases s u bj ect to v ary ing d egrees of infl u enc e and

, ,

th u s ind i rectly to es tim at e t he effect o f any s pec ia l ca u s e o f v a r i

at io n Fo r exa mp le s tr ipes and bars were o nc e c o m m o n c o lo r
.
,

m ark ing s o f h o rses as th ey a re n o w o f a s ses es pec ia lly zebra s

, ,

and q u aggas Co n seq u ent ly a c ert ain pro po rt io n o f c o lt s what

.
,

ev er t he parentage will b e b o rn wi th trac es o f s h o u ld er and leg

,

m ark ings N ow u nd er t he la ws o f chanc e a c erta in p o rt io n o f

.
, ,

th ese will b e the d irect o ff s pring o f s triped o r barred s ires a n d ,

will attract n o att ent io n t he m ark ings b eing c o n s id ered h ered i

,

tary By the s a m e law o f chanc e a c ert ai n (sm aller ) po rt io n will

.

b e the o ff s pring of parent s n o t b a rred and a s t ill s m a ller n u m b er ,

1 P r oc eed i ng s tire R oy a l S ociety , L X , 2 7 3

o
f .
 88 C AU S ES O F V A R I AT I O N

will b e t he pro geny of u nbarred r

and o u t o f da m s on ce m a ted
S i es

T h is s m a lles t p o rt io n get
'

w i t/c ba r red s i r es f or o tlzer of sp r i ug .

,

t ing its bars n o t by d irect d es c ent b u t by r ev er s ion will m o s t ,

lik ely b e erro n eo u s ly c o n s id ered t o ha v e d er iv ed th em fro m the

barred m ale n o t th eir s ire T he s am e is tr u e o f o th er m ark ings
.
,

and o n s u ch ev id enc e as th is t he t h eo ry o f t elego ny ha s been

b u ilt u p and s o far a s proo f go es it res t s o n n o b ett er fo u n d a
, , ,

t io n a s yet I n o rd er t o s ec u re ev id enc e a m u nt in g to pro o f i t

. o ,

is n ec es s a ry t o exa m in e la rg e n u m b ers in v o lv in g b o th p o s i t iv e a n d
n ega t iv e ev id enc e in o rd er to sec u re tru s t wo rthy a v erages VVhen
,
.

ev er th is ha s b een d o n e t he th eo ry o f t elego ny f a i ls o f s u pp o rt .

Tel egon y in man T h e s tat is t ica l m eth od ha s b een applied by

.

Pea r s o n in t he c a s e o f m a n
1
He t geth er with G a lt o n po s
.
,
o ,

s ess es data c o v er in g h u ndr ed s o f ind iv id u a ls in E ng lis h fa m ilies .

He rea s o n ed that if t he s ire ex ert s a p er m a n ent infl u enc e u po n

t he d a m t end ing t o a ss ert it s elf in a ll f u t u r e o ff s pr in g th en th is
, ,

infl u enc e m u s t b e in a s en s e c u m u la t iv e s o that t he y o u nger s o n s

,

in t he fa m ily will t end t o res em ble t he fa th er s lig ht ly m o re th a n

will t he o ld er s o n s c o nc eiv ed b efo r e s u ch infl u enc es h a v e bec o m e
,

es tab lis h ed .

His s t u dy c o v ered 3 8 5 bro th er br o th er s a n d 4 5 0 s is t er S is t ers ,

tak en t wo and t wo I n s o m e in st a nces th ere wa s c o n s id erab le

.

d ifferenc e in ages a n d in o th er s t hey were s u cc es s iv e ch ildren

, .

His d a ta c o v er ed b o th h eight and a rm len gth and aft er m ak in g ,

t he u s u a l allo wanc es fo r s ex and a ge Pea r s o n c o nc lu d es th a t so ,

fa r as th es e ch a ract er s a re c o nc ern ed n o s t ea dy
,
t elegon ic
infl u enc e ex is t s .

A ga in t he m any s u cc es s iv e m a rriages o f bo th c o lo r ed an d
,

wh it e wo m en t o m en o f o p po s it e c o lo r S h o u ld a ffo rd n u m ero u s
exa m p les o f t elego ny were it a c o n s eq u ent ia l f o rc e in h ered ity .

Sci en t ifi c ob j ecti ons t o t h e t h eory of t el egon y If t elego ny .

ex is t s it s infl u enc e o v er h er ed it a r y charact er s m u s t b e exp la in ed

, ,

s o far as pres ent kn o wledge go es in o n e o f three way s : 1 m

, ( ) s o e

eff ect u po n t he t is s u es o f t he f em a le s u ch as will infl u enc e

f u t u re o v a in t heir mat u ra t io n o r t he emb yr o in it s d ev elop m ent ;

( )
2 s o m e th ing li k e a part ia l f ert il iza t io n o f im m at u r e an d u nd e
v elo ed ov a
p i
,
n s u ch a w a
y a s t o infl u enc e th e ir charact e r at
1 P r oc eed i n g s qf t/ze R oy a l S oc iety , L X , 27
3 .
1 90 C A U S ES O F V A R I AT I O N

m are t ime o f S erv ic e and o f c ou r s e ar e extrem ely car e

at t he ,

fu l t o pr o t ect her fro m u nplea sa nt c o lo rs o f any so rt 1

T he .

h old o f th is th eo ry u po n t he po pu lar m ind is t he bes t exa m p le

a fford ed by breed ing o f t he strength o f t rad it io n T he s u ppos ed .

rea s on o n wh ich it res t s has slight bas is in fact T he c o ntact .

bet ween t he m o th er and the fet u s is n ot s o int im at e as is pOpu

la rly s u ppo s ed T he fet u s is ab so lu t ely d epend ent u po n t he
.

m o th er fo r n o u ris h m ent it is tr u e and it lies fl oa t ing in it s , ,

fl es hy incas em ent wh ich is in int im a t e c o ntact with t he t is s u es

,

o f the u t er u s ; b u t th er e is n o o rga n ic c o nn ect io n n o n erv o u s ,

int errelat io n what ev er .

A nyth ing wh ich wo u ld c u rtail o r s h u t o ff n o u r is h m ent wo u ld

o f c o u r s e inj u r e o r d es tr o y t he f et u s I t is a lso s u bj ect t o o th er .

acc id ent s as bec o m ing entangled in it s o wn c o rd wh ich m ay

, ,

th u s d iv id e a lim b o r ca u s e s tra ngu lat io n a ll o f wh ich ho w , ,

ev er is q u it e a s id e fro m t he m att er in po int

,
.

T he r ea l q u es t io n is wh eth er an d t o what ext ent t he fet u s is , ,

infl u enc ed by pec u liar it ies o f n ou r is h m ent d u ring it s d ev elo p

m ent I t wo u ld o f c o u r se b e inj u red by po iso n s and t he d a n ger
.
,

from ad m in is t ering an aes th et ic s is gr eat b u t th is d is c u s s io n is ,

lim it ed t o t he d irect effect o f m enta l im pres s io n s .

T he ind ifferenc e o f t he f et u s to it s so u rc e o f n o u r is h m ent is

s h o wn by an exper im ent o f Hea pe perfo rm ed fo r an o th er p u r
2
,

po se b ut throwing light u po n th es e q u es t io n s I n th is exper im ent

,
.

t wo s egm ent ing o v a wer e o bta in ed fr o m an A n go ra d oe rabb it

wh ich had b een fert ilized by an A ngo ra b u ck th irty t wo h o u r s -

prev io u sly and were im m ed iat ely tran s ferr ed t o the u pper en d
,

o f t he F a llo p ian t u b e o f a B elg ian har e rabb it wh ich had b een

fert ilized three h o u rs befo re by a b u ck o f t he sam e bree d as

h erself I n d u e c o u r s e th is Belgia n hare d oe gav e birth t o s ix
.

y ou ng Fo u r o f th es e res em b led h ers elf and her m at e b u t the

.
,

o th er t wo wer e u nd o u bt ed A ngo ra s Bo th o f t he A ngo ra s

3
.

were b o rn b igger and s tr o n ger th a n any o f t he o th er y ou ng and ,

F 1
g d
or ll t
a i f ll good i t co ec M il on S t k B
o d i g a e e n s a n c es , s ee es , oc ree n ,

pp 8 — 2
9 5 1 l2t y ig h b h d
o r co ns u l an ne or oo o rac e

—
.
, .

V 2 V i ti
er n o n , i A im l d Pl
ar a t pp on 9 n ; l Pn w d i g qf a s an an s, . 1 1 1 20 a so r ee u s

t/ze R o y a l S o c i ety , X L V I I I , 4 5 7 .

8 T h e A n go ra s wer e c h a ra c t eri ze d an d ea s i ly d is t in g u is h d e by t h eir l o n g,

si l ky h a ir a n d t h ei r h ab it o f s wa in g t hey h d
ea fr o m s id e to s id e .
 I NT ER NA L CA U SES OF V A R I AT I ON 1 9 1

th ey all alo ng m a intain ed th eir s u prem acy in th is d irect io n

'

W ha t ev er th is exp er im ent pro v es o r d oes n o t pro v e it S h ows ,

c o nc lu s iv ely t hat a fert ilized A ngo ra ger m preserv es and d ev elo p s

it s inh erent charact ers perfect ly in an exc eed ingly fo reign en v i
ro n m en t o n wh ich it ev id ent ly d ep end s o n ly fo r n o u r is h m ent
,
.

Ment al im pres s io n s and n erv ou s c o nd it io n s a re c o m m o n ly ln

v o ked t o exp la in birth m ark s and o th er na t u ra l abn o rmalit ies s u ch ,

a s t he los s o f a fi nger . I n th is c o nn ect io n t wo fact s ar e t o b e

c aref u lly c o n s id ered fir s t th ere is c ertain t o occ u r a large n u m
,

b er of m ar k s s tra wb er ry c u c u m ber and oth ers ) and m any

”

, ,

m alfo rm at io n s o f o n e k ind o r an o th er S carc ely an ind iv id u al is

.

abso lu t ely free fro m so meth ing of t he k ind A gain m oth er s are .
,

s u bj ect ed t o a ll so rt s o f s ight s s o u nd s and exper ienc es d u r ing t he

, ,

m any week s of em bry o n ic d ev elo p m ent a n d it wo u ld b e s t range ,

ind eed if o u t o f t he th o u s and s o f cas es s o m e c o r res po nd enc e

bet ween mark s and experienc e c o u ld no t b e figu red o u t es pe ,

c iall
y by o n e wh o s e belief is fi xed and who ha v ing t he ca s e at ,

hand n eed s o n ly to fi nd the pro per co rres po nd ing experienc e

,
.
”

T he law o f c hanc e a lo n e will in s u re an o cca s io nal c o rr es po nd enc e

bet ween the t wo ,
— ent irely en o u gh t o s tart t he trad it io n and

t o m a in ta in it aft er ward A s in th eo r ies c o nc ern in g t he co ntro l

.

o f sex any th eory s tat ed will b e v er ifi ed ha lf t he t im e b eca u s e

,

th ere is b u t o n e alt ernat iv e s o h ere wh ile t he a lt ernat iv es are

, ,

m o re t he c o rr es po nd enc e is c ertai n s o m et im es t o a ppear u nd er

,

t he la w o f chanc e a lo n e .

A n o th er fact t o b e reck o n ed with is that if t he fet u s were s o

sen s it iv e t o m enta l im pres s io n s a s t o req u ir e t he d is p lay o f

pro perly co lored b lank et s — if fem ales were so s u s c ept ible a s

,

th is t o s u rr o u nd ing S ight s what a j u m b le o f c o lo r s o u r d o m es

,

t ic an im a ls wo u ld s peed ily d is play I n t he o p in io n o f t he wr it er

.

th is trad it io n has n eith er a s c ient ific bas is n or well es tablis h ed -

in s tanc es a n d it is t im e it n o lo nger o cc u p ied t he mind s o f

,

breed ers to t he exc lu sio n o f far m o re im po rtant m att ers I n th is .

c onnect ion it is worthy of rem ark that if t he a v erage breed er

were half as fam iliar with im po rtant fact s and ha lf a s att ent iv e ,

t o th eir bear ing u po n h is o perat io n s as he is fa m iliar with an d

,

att ent iv e to fl oat ing trad it ion s we s h o u ld hav e a far sm aller

,

pro po rt io n o f wo rth les s an imals .

1 92 C AU S ES OF V A R I AT I O N

S ECTI ON I X — R EV ER SI O N A N D A TA V I S M

T h es e t wo t er m s a re u s ed to d es ignat e charact ers appearing

i n the o ffs pring b u t no t V is ib le in t he parent s R ev ers io n is .

u s ed t o i nd icat e res em b la nc e to a c o m parat iv ely n ear b y anc es t o r -

,

n o t t he parent wh ile ata v is m refers to exc eed ingly remo t e

“
,

a nc es t o rs s o m et im es o f o th er and fo u ndat io n s pec ies

,
.

T h u s if a da s h o f im p u r ity o f b loo d enters a h erd it will

, ,

appear o ccas io na lly fo r m any gen erat io n s T h is wo u ld b e .

s p o k en o f a s a rev ers io n t o the s t range b loo d If a s i re o r

“
.

d a m ha s s o m e pec u liar charact er lik e wh it e s t o ck ing s in h o rs es , ,

a pec u liar h o rn in catt le o r a hab it in m an i t is likely to appear

, ,

fro m t im e to t ime in f u t u re genera t io n s ev en after its real ,

o r ig in is fo rgo tt en T h is is a rev ers io n E ng lis h breed s o f cat t le

. .

a re d ev elo ped fro m t he a nc ient wild wh i t e catt le o f B ri ta in and ,

t he o ccas io n a l appeara nc e i n a ll th es e breed s o f a wh it e ca lf

with red o r bro wn ears lo wer leg s and t a il bru s h is t o b e
, ,

expect ed I t is a rev er s io n n o t a pro o f o f m ix ed b loo d

.
, Of .

c o u rs e t he a n imal s o m ark ed is u seles s fo r breed ing p u rpo s es ,

b u t n o repro ach t o t he h erd and n o n e n ec es s ar ily t o t he d a m that

,

prod u c ed it fo r rev ers io n s fo r t he m o s t part seem to c o m e s in gly

,
.

A ta v is m o n t he o th er hand go es farth er back

,
Fo r exa m p le
, .
,

ma m m a ls d u ring th eir early em bry o n ic d ev elo p m ent s t ill S h o w

tra c es o f t he gill s lit s th u s betray ing th eir u nd o u bt ed o ne t ime
,
-

c o nn ect io n with t he s a me s t o ck wh ich ga v e rise to t he aq u at ic

an im a ls T h es e gill s lit s o cc a s io n a lly pers is t fa iling to c lo se
.
, ,

and g iv e ris e to t he abn o r m a lity kn o wn as c erv ica l fi s t u la I t

“
.

is an u nd o u b t ed a ta v is t ic abn o rm a lity — an extrem e cas e of ,

c o u rs e bec a u se o f it s ant iq u ity

,
.

C as es o f th is k i nd a re t o b e caref u lly d is t ingu is h ed fro m m ere

meris t ic v ariat io n s Fo r exa m ple t he s u dd en app earanc e o f a
.
,

three t oed ho rs e wo u ld b e regard ed a s ata v is t ic for a ll h o rses

-
,

o nc e had three t oes ; b u t a s ixth d ig it in m a n is c ert a i n ly n o t

ata v is t ic fo r we hav e n o ev id enc e that m a n ev er pos s essed

,

n o rmally m o re than fi v e d igit s The crim inal in s t inct in c erta in .

m en is u nd o u bt ed ly ata v is t ic s h o wing n o t s o m u ch a d eligh t

,

in ev il d o ing as a n ent i re ab s enc e o f t he h igh er s en s e o f right

d o ing.
1 94 C AU SES O F V A R I ATI O N

Galt o n has end ea v o red to a s s es s m ath em at ica lly t he fract io n

o f d i rect i nh er itanc e o r m o r e acc u rat ely t he S im ilari ty b et ween
, , ,

t he ch ild and it s v ar io u s anc es t o r s F ro m his s t u d ies he c o n .

c lu d es that t he V is ib le o r d o m inant cha ract er s o f t he ch ild a re ,

on the a v er ag e i nh eri t ed that i s c o rres po nd with th o s e o f t he

, ( ,

v ar io u s anc es t o r s ) ro u gh ly a s f o llo ws
1
, ,

F ro m the immed ia te pa re n ts
F ro m the g r a n d pa ren ts
Fro m the g rea t g ra n d p a ren ts
-

F ro m the g rea t , g r ea t-gr a n d p a ren ts

E arlier a n c es t o rs i n p r o po rt io n

Pear son , wo rk ing with larg er n u m bers and d iv er se cha ra ct ers ,

c o nc lu d es that C alto n s fract io n o f d irect inh eritanc e

’
is
t o o h igh and i s inc lined t o b eliev e i t n o t ab o v e
,
fo r b lend ed
charact ers T he s u bj ect will b e p u rs u ed farth er u nd er The
.
“

La w o f A nc es tral Hered ity b u t th is glim p s e into t he nat u re o f

,

inh eritanc e is t he bes t m eth o d kn o wn to the a u th o r t o d is s o lv e

t he a l mos t s u pe rnat u ra l m y s t ery that has b een thro wn aro u nd
rev er sio n and it s nat u ral c o ro llary lat ent chara ct ers T he s t u dy , .

can b e p u rs u ed n o farth er at th is po int b u t what has been s aid ,

will s er v e t o s h o w that rev er s io n ( regres s io n ) is a fert ile ca u s e

o f v ar iat io n a s ca lc u lat ed fro m t he t h I will
yp e o
f t e p a r en t t .

s erv e a ls o a s an i ntr o d u ct io n prepar ing t h e s t u d ent fo r t he m o re

,

s er io u s s t u dy o f inh er itanc e lat er o n wh en we s ha ll learn that ,

t he real type fro m wh ich a ll d epart u res s h o u ld b e reck o n ed is

, ,

t he typ e of t/ze ra c e and n o t t he s p ec ia l typ e o f t he parent o r

, ,

ev en o f t he m id par ent -
.

S EC TI O N X — I N DI V I D U A L C HA R A CT ER S D E PE N D ENT

U PO N S EX

T hat bo th
x es p o s ses s a n d tra n s m it all the cha ract ers o f the
se

rac e is a well es t ab lis h ed fact in ev o lu t io n I t is a ls o tru e that

-
.

t h e part ic u lar charact er s t o u nd erg o d ev elo p m ent and t he ext ent ,

1 V ern o n , V a ria t io n in A n i ma l s an d l
P a n ts , p . 1 23 ; P r oc eed i n g s f
o tb e R oy a l
S oc i ety , l
L X I , 4 0 1 , 1 8 97 ; G a t o n , N a t u ra I n er it a n c e, p 1 9 1 l h . . T is h d o es no t

m ea n h
t a t ev er y d d ll h
in i v i u a l wi in eri t in t is p ro p o rt io n , b u t t h h at t he f ra c t io n s
e xp res s a v era ges .
 I NT ER NA L C A U SES OF V A R I ATI ON 1 95

to wh ichthey will d ev elo p d epend s v ery m u ch u po n the sex o f

t he i nd iv id u a l Ho w m u ch allo wanc e to m ak e o n acc ou nt o f s ex
.

in c o m pari ng o n e ind iv id u al with an o th er o f a d iff erent sex we

are in m o s t cas es u nab le t o s ay n o t fr o m t he im po s s ib ili ty o f
,

kno wing b u t fro m t he fact that in res pect to mos t charact ers
,

t he m atter has n o t yet been wo rk ed ou t I t is eas ily po s s ib le .

,

h o wev er .

F or exa m p le in r es pect t o s tat u re m en a re 8 per c ent ta ller

, ,

tha n wo men so that wh en the h eights o f the latt er a re m u lt i

,

p lied by 1
t he t wo a re s tr ict ly c o m parab le and n o t bef o r e ,
.

Wh en th is is d o ne the d zj r enc e d u e to s ex has been eli m inat ed

'

fl e ,

and the s tat u res o f men and wo men m ay b e d irect ly c o m pa red .

I n gen eral m ales and fem ales exh ib it the s a me charact ers
, ,

b u t in v a rying d egrees For exa mp le t he wo m an as well as t he

.
,

ma n has hai r o n t he fac e b u t in les s a mo u n t ,

t he m a le as well
as the f em ale has n ipp les b u t th ey a re r u d im entary ,
A mo ng .

m a m m a ls and t he d o m es t ic an im a ls gen erally t he ma le is h eav ier

in fro nt gen erally o f a m o re ro b u s t b u ild and c o ns id erab ly
, ,

la rger than t he f em a le a d is t inct io n that by no mean s ho ld s

,

in ani mal life gen era lly .

I n the present s tate o f kn owledge we s im p ly kno w that t he

general appearanc e o f t he ind iv id u al and its character d ev el
Op men t are largely d epend ent u po n i t s s ex ; b u t t o what exact

ext ent rem ai n s in m o s t ca s es t o b e d et er m in ed and t he d et er ,

m ina t io n m u s t b e m ad e bef o re we can c o m par e ind iv id u a ls o f

d ifferent s ex es with any d egree o f acc u racy With o u t a d o u bt .

d is t inct io n s in s ex hav e been greatly o v erwo rk ed t he d iff er ,

en c es b ei ng mo s t ly o f d egree rath er tha n o f k ind and far les s

3
,

c o n seq u ent ia l than has been s u ppo s ed .

I nd iv id u a ls d epr iv ed o f th eir sex u a l o rgan s by cas t rat io n o r

by s paying do no t d ev elo p th eir pri m ary s ex u al charact er s T he .

cas trat ed male is no t a f em ale as is so m et imes erro neo u s ly ,

believ ed b u t a m ale arres t ed in his d ev elop ment ; and t he s payed

,

f emale is an u nd ev elo ped f em ale A s wo u ld b e expect ed bo th .

,

1 Th es e d
a ta a re t he res u lt o f G alt o n
’
s s tu dy o f t he s ta t u re o f E n g is l hp eo p le.

l
S ee G a to n , N a t u ra l I n erita n c e h .

3 I t is i l
d
e t o a tt em p t t o p ro v e h
t a t c e r ta i n c h a ra c t ers , as i d e f ro m t h os e o f

rep rod u c t io n , a re es ec ia
p ll y id en t ifi e d with o n e s ex .
1 96 C AU S ES OF V A R I ATI O N

tak e o n t he s ec o nda ry s ex u al cha ract ers ( th os e d o m inant in the

o th er s ex ) prem at u rely y o u n g
1
.

M any ent ire ind iv id u als n ev er d ev elo p s tro ng ly the prim ary
cha racters o f th eir o wn s ex T h ere a re effem inate males and .

m a s c u line f em ales th ose in wh ic h the charact ers of t he o pp o

,

s it e s ex a re u n u s u a lly d ev elo ped I t is n eed less to s ay that s u ch .

i nd iv id u a ls a re n o t the bes t parent s .

II I N TE R N A L I N FL U E N CE S A FFE C TI N G THE
R A CE A S A WH OL E
Ov er again s t th o se ca u ses that m ay o perate in t he cas e of
each i nd iv id u a l t o warp it s d ev elo p m ent a re t o b e c o n s id er ed

th o se that infl u enc e the rac e as a wh o le t u rn ing t he line o f ,

d esc ent mo re o r les s perm anently as id e fro m fo rmer chann els .

S o m e o f t hese infl uenc es a re c learly d efin ed and ea s ily rec o g

n ized ; o th ers a re pro b lem at ica l t he d is c u s s io n n o t ha v ing e t
y ,

pas s ed beyo nd t he s tage o f a pla u sible th eo ry .

T he s t u d ent o f threm m at o lo gy and t he breed er s h o u ld b e

alway s mindf u l tlra t tlre pu rpos e of a ll g ood b reed i ng is n ot s i mp ly
to lrold r/ra t
te we a lr ea dy lea v e b u t to p rod uce n ew typ es b etter
a da p ted tlra n tlre o ld to tlre p u rpos es qf m an A cc o rd ingly any .

and a ll lin es
that pro mise any h o pe of su cc ess s h ou ld be as s i du
o u s ly in v es t igat ed .

S ECT I O N X I — R EL A T I V E F ER T I L I T Y , O R G EN ET I C
S E L EC T I ON 1
'

T heassu m pt io n that all m em bers of a ra c e a re eq u ally fert ile

i n s e and i n ter s e (o f th em s elv es and bet ween each o th er in all
d irect io n s ) is n o t o n ly has ty b u t dangero u s ly inc o rrect To .

q u o t e Pearso n Fert ility is n o t eq u a lly d is tribu t ed a mo ng all

,

ind iv id u a ls .

1 Th e en t ire a n im a l wit h in c reas in g a ge, it s o wn s ex u a l c h a ra c t ers a b a t in

g,
b egin s t o t a k e on t h ose of t he o t h er s ex . T h us t he h en gws s p u rs , t h e c o w
ro

be ll o ws an d p a ws t he di t r , wo men g ro w s c a n ty b ea rd s , a n d o ld m en s v o ic es
’
gr o w
l ight .

3 Pea rs o n , G ra mm a r of S c ien c e, pp 3 7 6, 4 1 4 , 4 3 7 - 4 49. 4 62

. .
 1 98 C A U SES O F V A R I AT I O N

u n les s the tend ency s ha ll b e held i n by the g r ea ter a tt ra c t

c hec k

i v eness f g loi ht er e es ,
y r e er en t i a l m a t i n
p f g T h is w o u ld b e a .

lo ng and S lo w proc es s , b u t it wo u ld a v a il m u ch to red u c e th o u gh ,

it c o u ld n ev er ov erc o me the eff ect s o f t he ,

h igher fert ility o f the
darker eyed ind iv id u als
-
.

Pearso n c o llect ed 4 44 3 cap s u les o f wild po ppy 1

T h ey s h o wed .

t he fo llo wing d is tr ib u t io n arranged acc o rd ing to t he n u m b er o f

s t ig m at ic band s

T he larges t n u m ber o f caps u les ( 9 5 4 ) had 1 3 band s and the

next larges t n u m ber had 1 2 V ery few had s o m any as 1 8 o r 1 9 .
,

o r s o few as 5 6 o r 7 The type n u m b er o f band s is th en 1 3

, ,
. .

He prov id ed rec eptac les and k ept t he s eed s o f each gro u p

s eparat e He s ay s :
.

h
.

T my g a t
o p i e re w e s ur m y er s p t le, f 1o2 d
ev1 3 w e
r, e r ce ac s or an r

filled p wi t h th
u t t f f w
e c o n en s o
y p l e t h e f 1 1 d 1
v er 4 m e e ca su s, os or an or

t d ily th
ar e f
, 10 os d 5 o ly with em p t y i g
or an 1g t m b en f p le n a rea nu r o ca su s,

wh ile I o ld h d ly g t
c u y e d
ar t ll f m teh an p l w i th
s e y m a y a ro os e c a su es v er an

or v e y f w b
r d ; th y we e p
e an s ti lly te i l e T h typ r p l were
ra c ca s r e . e e ca s u es

m
en o r ly fe tile [wh ile] t h
o us r with , m d e te d e i tio f m i t
os e ev en a o ra v a n ro

[w e er
] e l ti l
r
y e
a e b
ve l t l y
or i f e t
v i le n a so u e n r .
3

T h isbeing t ru e the po ppy has bec o m e ab o u t as s table as is

,

po s s ib le fo r it s h igh es t fert ilit y is with its m o s t n u m ero u s po pu

,

lat io n Th is p lant was gro wing wild in nat u re Ob v io u s ly t he

. .

great b u lk o f s eed s d is t rib u t ed wo u ld b e of the type n u m ber 1 3 ,

o r n ear it and t he m a s s o f d es c endant s wo u ld aris e fro m s eed s

,

c lo s e to t he typ e What chanc e n o w wo u ld th ere b e in nat u re

.

fo r a large c o lo ny o f S ix o r s ev en ba nd ed s tra in s t o aris e ? V ery - -

litt le u n less th ey happen ed t o po s s es s s o m e d ec id ed ad v antage

,

i n t he s t ru gg le fo r ex is t enc e in wh ich ca s e t he type wo u ld ,

s peed ily S h ift in that d irect io n ; b u t a s lo ng as t he h igh es t

fert ility rem ained with t he h igh er n u m ber o f band s the rac e ,

wou ld b e u n s tab le .

1 Pea rs o n , G ra mm a r of S c ien c e, p p 4 4 3 4 44.

-
.

3 Ib i d .
p 4 44
. .
 I NT ER N A L C A U S ES O F V A R I AT I ON 1 99

Su ppos e it were t he p u rpo s e o f ma n t o d ev elo p a po ppy with

fewer o r with m o re than the nat u ra l n u m ber o f
, ,
band s sa
y ,

s ev en o r s ev ent een U nd er what d is ad v antage he wo u ld wo rk

.

as lo ng a s the f ert ility rem a in ed relat iv ely lo w ! and he wo u ld

n ev er s u cc eed u n les s he s epa rated t he plant ings fro m t he m o re

pro lific type T h is is genet ic select io n
. .

B reed ers are c o n s tant ly o perat ing aga in s t t he d rag o f infert ility
with o u t kn o wing it and a re as o ft en wo nd ering why bett er
,

res u lt s d o n o t fo llo w es pec ially wh en o n ly approv ed m at ing

,

is pract ic ed Co n s id er t he m ath em at ic s in v o lv ed in s ay three

.
, ,

lin es o f d es c ent o f d iffer ent d egrees o f p ro lific ac y F o r t he .

s ak e o f s im p lic ity in illu s t rat io n let u s s u ppo s e three c o ws were

liv ing i n a h erd t o geth er On e o f th es e c ows raises t wo calv es

.

and bec o m es barren ; ano th er raises fo u r bef o re she c eases to

breed and ano th er s ix Fo r the s ak e o f fu rth er s im plic ity let u s
,
.

s u p po s e that o n e ha lf t he calv es a re f em a les and that each ,

dau ght er d es c endant ex act ly repeat s t he perfo rm anc e o f her

d a m and th en b ec o m es barren Ho w will t he acc o u nt s tand in
.

a few generat io n s 1

C U M U LAT IV E EFF ECTS OF FERT I L l I Y

’
As S HOW N sv T HE R ELAT IV E
‘

N U M BE R OF FEM A LE D ESC EN DANTS OF Co ws OF V AR IOU S

D EG R EES OF FERT I LITY

G EN E RA T I ON S

C o ws C Aw as

T h ist a bu lar pres entat io n S h o ws that aft er fiv e gen erat io n s

o f th is k i nd o f breed ing th ere wo u ld b e b u t on e f er ti le c o w o f

t he fi rs t o rd er in t he h erd 3
wh ile if a ll had b een k ept th ere
,

wo u ld b e 3 2 o f t he s ec o nd o rd er and 2 4 3 o f t he th ird What an .

1 l
Th ere is n o o n ger a n y o u b t t a t fert i it isd h l y an in h eri tab le c h a ra c t er .

h
3 T ere m i
g h
t b e a n
y n u m b er o f liv in
g b ar ren on es if th e s t ra in h appen s t o be
a fav o rite a n d is lo n g liv e -
d .
2 00 CA U SES O F V A R I AT I ON

o ppo rt u n ity fo r s elect io n in t he latt er in s tanc e with no ne what ,

ev er in t he f o r m er ! T he fi r s t u nt i mely d eath wo u ld rend er t he

line ext inct wh ich is pe rhap s t he b es t fat e that c o u ld o v ertak e

,

a rac e wh ich at bes t is able o n ly to h o ld its in it ia l n u m ber go o d .

Of c o u r s e art ific ia l c o nd it io n s ha v e been a ss u med in o rd er t o

bring o u t t he princ ip le I t d oes n o t wo rk o u t in th is reg u lar
.

and ev id ent mann er in o u r h erd s b u t t he p rinc iple o f genet ic

,

s elect io n is at wo rk nev erth eles s I t wo u ld b e fo rt u n a t e if it

,
.

were m o re ev id ent fo r t he h erd s a re few that d o n o t c o nta in a

,

larg e pro po rt io n o f f emales that c o ntrib u t e n o th ing to t he r ea l

lin e o f d es c ent th o u gh th ey n o w and th en g iv e b irth to exc ep
,

t io n al ind i v id u a ls T he q u a lity is goo d b u t t he ra t e o f repro

.
,

d u ct io n is to o lo w .

Ho w m any a breed er ha s s pent fru it les s years in in effect u al

attem pt s to b u ild u p a s train exc ellent in it self b u t es s ent ially
inf ert ile ! W itn es s t he fat e o f that rem arkab le fa m ily o f s h o rt
h o rn s the D u ch ess T h is fa mo u s fa m ily in i ts glo ry was n ev er
,
.
, ,

s u rpa s s ed ,yet i t wa s gen et ic s elect io n that ext er m inat ed t he

lin e F o rt u nat e ind eed is the breed er who kn o ws th is pri nc ip le

.

and realizes it s f u ll po wer wh en ev er he fin d S h im self o ppo sed

by it .

T he s t u d ent m u s t n o t get t he im pres s io n that g en et ic s elec

t io n is an enemy o n ly A pro lific line t end s a s s tro ngly to
.

es tab lis h and m a inta in i t s elf a s d oes a barren o n e t o r u s h h ead

lo ng t o ext inct io n G enet ic select io n is th erefo re a friend po w

.

erfu l fo r g o o d a s well as an en e my p o werf u l fo r ev il ; b u t i t is

,

a s q u iet and u n o btr u s iv e in t he on e r elat io n a s it is i n s id iou s

in t he o th er T he breed er ha s o n ly to b e et erna lly c o n s c io us o f

.

the fact that if he is t o s u cc eed he m u s t ha v e n u m b er s n o t ,

oc cas io nal b i rth s b u t reg u lar and gen ero u s

, T h en he m ay b e .

s u re that he is n o t try ing t o d o a th ing o n wh ich nat u re ha s

s et t he s ea l o f her d is appro v al thro u gh n o n pro d u ct io n Ho wev er-

.

wo rthy and h o wev er v alu ab le intri n s ica lly t he s trai n m ay b e it is ,

wo rth les s u n les s he can prod u c e i t with c erta inty and in any
d es ired n u m bers “
Beware o f t he S hy breed er and trea s u re t he
.
,

o ld f em a le that breed s reg u larly and t ru e T h is d o ct rin e es tab

.

lis h es a c oOpera t io n with nat u re that in s u res res u lt s and with ,

o u t it g en et ic s elect io n will wo rk aga in s t u s n o t fo r u s ,

.
2 02 C A U SES O F V A R IA T I ON
.

Weis m a nn b eliev es that in nat u re the d eath po int has been

1

fi x ed at an age m o s t pro fitab le to the ra c e a s a wh o le That is .

to s ay i t is b es t fo r t he ra c e ( I ) that o n ly t he s tro nges t s h o u ld

,

s u r v iv e t o t he br eed ing age ; ( 2 ) that th es e s h o u ld li v e a s lo ng

as th ey a re ab le to r epr o d u c e ; and that ( 3 ) th ey s h o u ld th en d ie

a n d c ea s e t o oc c u py roo m and c o n s u m e f oo d wh ich wo u ld o th er

wis e b e a v a ilab le fo r t he s u s t enanc e o f m o re ro b u s t i nd iv id u als

engag ed i n repr o d u ct io n T h is fi xes the d eath po i nt th eo ret ic
.

ally a t the c es s at io n o f repro d u ct io n exc ept in s u ch s pec ies a s ,

th o se in wh ich the y o u ng need t he care o r t he ed u cat iv e as s is t

anc e o f t he mo th er I n th ese t he th eo ret ical d eath po int wo u ld b e
.

at the m at u ri ty o f the las t y o u ng T h is o f c o u rse is in referenc e .

t o wild s p ec ies and Weis m ann believ es that nat u re has es tab
,

lis hed t he d eath p o int in c lo s e c o rres po nd enc e to th is princ ip le .

Ho wev er that m ay b e th ere is a pro b lem h ere fo r the breed er , .

I t is fo r him t o fix t he d eath lim i t well bey o nd t he period o f the

p a r t ic u la r s er v ic e r e u i r ed
q I n nat u r e th er e is b u t o n e o bj
. ect in

life —
,
s elf preserv at io n and r eprod u ct io n
-
On o u r far m s th ere .

a re o th er o bj ect s T he h o rs e is fo r labo r and his s er v ic eab le

.
,

age a s well a s his d egree o f intelligenc e n eed s t o b e length en ed

a s m u ch a s p o s s ib le I n nat u re early a n d ra p id reprod u ct io n is

,

a f u ll eq u iv alent fo r lo ngev ity I t is n o t s o on ou r farm s wh ere .

,

t he ind iv id u a l c o u nt s fo r mo re and ev en ra p id repr od u ct io n ,

cann o t tak e t he p lac e of lo ng life and fa ithf u l serv ic e .

S EC TI O N X I V — BATH M I C I N F LU EN C ES

Do pec ies po s s ess inh erent t end enc ies to v ary ? I f a rac e
s

c o u ld b e s u rro u nd ed by po s it iv ely u nchanging c o nd it io n s if it ,

c o u ld pro d u c e as ex u a lly and if all types were eq u a lly v igo ro u s

,

and eq u ally fert ile wou ld i t r ema in c ons ta n t ? S o m e v ariat io n

,

wo u ld aris e thro u gh red u ct io n b u t th is wo u ld b e h et ero gen eo u s , ,

that is n o w in o n e d irect io n no w in an o ther The real qu es

, , .

t io n s the bath mic ev olu t io n is t as ks a re th ese : I s th ere a t end

ency fo r t he type t o drift ind ep en d ent o f s elect io n o r s u rro u nd ing
,

infl u enc es ? A re it s d ev iat io n s cha ract erized by a c o nt in u a l b ias

1 W eis m an n , Es sa y s on H ered it y I , , 1 1 1 —
1 6 : s ee al
3 so Pears o n , C ha n c es of

D ea t h , pp . 1- 42 .
 I NT ER N A L C A U SES OF V A R I A T I O N 2 03

in fav o rit e d i rect io n s ? D oes it v a ry progres s iv ely becau se

im pelled in th es e d irect io n s by gro wth fo rc e o r o th er inh erent
“

en ergy ? A re s pec ies h eld t o th ei r pres ent s tandard s by o u t s id e

infl u enc es ? o r if n o t , h eld a re th ey d rift ing in s p it e o f u s ?
,

I s t he life pri nc ip le c o n s tant o r per iod ic in it s act iv it ies ; and

a re th os e i nterna l en erg ies that v i ta lize m att er and that d et er m in e

v i f e i —
d e elo p ment and d fer nt iat o n a re th ey ind ifferent as t o the ,

trend o f t he type o r d o th ey ru n m o re eas ily in s ome chann els

,

than in o th ers ? I s v ariat io n in so me s en s e s u bj ect to and d irected

by a nat u ra l b ia s ? Th is is t he fi eld o f bath m ic ev o lu t io n and 1
,

th es e a re t he q u es t io n s in v o lv ed N o o ne is m o re int eres t ed in .

th eir d is c u ss io n than is t he breed er o f d o mes t icat ed f o r ms .

Two princ ipa l th eo ries c o v er ing t he field o f bath m ic ev o lu t io n

hav e been pro po sed bo th incapab le of ab so lu t e pro o f as all s u ch
, ,

th eo ries m u s t b e b u t bo th o f int eres t to t he breed er

, .

Accel erat ion or reta r dati on of growt h force T h is pri nc ip le is .

ann ou nc ed by Co pe as a n int erna l and ev er pres ent cau se o f

2 -

progress iv e ev o lu t io n r u nn ing thro u gh a ll f o r m s of life and

,

beneath all o rd inary infl u enc es im pelling u nn o t ic eably b u t irre ,

s is t ib l in c erta i n d ir ect io n s I t is aft er a ll acc o rd ing to th is

y .
, ,

au th o r t hem os t s u bt le and m o s t po t ent ca u se o f d epart u re from

,

type The h o rse has u nd ergo n e s t eady pro gres s iv e d ev elop m ent
.

fro m an an im al o f t he s ize o f a jack rabb it u p t o his pres ent

pro po rt io n s and perfect io n T h is is d u e acc o rd ing to Co p e .
, ,

n o t so m u ch t o s elect io n a s t o a c o nt in u o u s perhaps a lm os t ,

u nprec ed ent ed a c c elera t i on qf g r owth f orce

,
.

T h is th eo ry att empt s t o exp la in m u ch o f ev o lu t io n thro u gh

t he en ergy o f gro wth th u s thro wing in t o t he d is c u ss io n a
,

dyna m ic elem ent c o m m o n ly n eg lect ed by ev o lu t io n is t s I n th is .

c o nnect io n Pears o n pert in ent ly rem ark s

T he e i r th i g m e (
s no n l ) orie t ifior i iess
g i h t
u n sc
g o w t h n c n us n an n eren r

fo rc e to e p l i x the e l
a n h ge i li i g f m th i
s c u ar c an i g the f
s n e v n or s an n us n o rc

of g r avita ti i h e e t i m tte t
on n r n p lni t he ad l pmr t f pl
o ex eta y a n ev e o en o an r

1 P G mm
ea rs o n , ra f S i pp 3 7 5 3 7 7
ar o Th t m
c en c e, b th m i . h -
. e er
“
a c as ere

use dd o es t i
no l d g t i l ti
nc u e en e y tch l
s e ec t i g t i t
o n o r an l o e r s e ec ve a en , n ern a or

t
ex ern a l b , th
ec a u s e ff t f ll
e e h i flec s o t dt
a m t suc d n u en c es en o co e o a res t a n

no t t o con s t it t u e a ti l bi c on nu a as .

O rigin o f t he F itt es t , p p 6 ; Pri ma ry Fa c t o rs

3 C o pe, 1 8- 8
3 0, 1 90- 1 92 ,
3 9 39
-
.

o f O r
gan ic E v o lu t io n , pp 4 7 3 4 94
- . .
2 04 C A U SES OF V A R I AT I O N

s y s tem s from n eb ulm . T he ul tim a te ac tio n o f v i tal u n its in ea c h o th e r s

’
p res

en c e wo uld b e no m o re, no r less , of a m y s ter y t h a the ltim a te

n u ac tio n of

m a terial u n its . T he real o b ec tio nj to ba th m ic e o l tio lie

v u n s n ot in a ny

a
p r i or i reas o n ag a in s t a n in he ren t
g r o wt h fo rc e, b u t to t he o b v io u s his to r
ic al fa c t t h at s uc h a fo r c e ha s b een u sed to c o v er a ll s o rts o f o b s c u re reas o n

in g a n d ev en s h eer f oo lis h n es s . S c ien c e wo u ld welc o m e ab o v e al l th in gs a

d es c rip tio n of the a c tio n b etween v i ta l u n i ts as s im ple as the la w of g ra v i
ta tio n , p ro v id ed i t g ve a c a u s al a c c o u n t o f v a ria t io n an d the welc o m e
'

a ;
wo u ld b e n o n e the les s S in c ere if the a c tio n s h o wed th a t v a ri a tio n was b iased
a n d th a t ev o lu t io n wo u ld b e irrev ers ib le ev en wit h a rev ersed seq u en c e o f

p hy s ic al en v iro n m en ts .
1

C o pe th eo ry o f acc elerat io n o r retardat io n o f gro wth fo rc e

’
s

is o f c o u rs e m erely q u a n ti ta ti v e and wo u ld exp la in any d ifferenc es ,

that m ight arise thro u gh eit her s ize o r pro po rt io n s o f part s o r ,

fac u lt ies d epend ent u po n s u ch pro po rt io n s I t d oes n o t att em pt .

t o exp la in t he int ro d u ct io n o f cha ract er s and if it can b e in any ,

way c o ntro lled n o m eth o d ha s yet been po i nted o u t .

W e all a llu d e to t he s a me genera l th o u ght wh en we u se t he

wo rd s v it ality and c o n s t i t u t io n to d en o t e n o t s o m u ch ten
a c it o f lif e a s V igo r o f gro wth and we a ll rec o gn ize that s o m e
y ,

ind iv id u als and s o m e lin es po s s es s th is fac u lty in m u ch h igh er

d egree than o th er s S o m e ind iv id u als n ev er s u rv iv e t he emb ry
.

o n ie s tag e o th e rs d ie in i nfancy s t ill o th er s r ea ch f u ll m at u ri ty ,

and a few pers is t t o an ad v anc ed age A s d eath c o m es o nly with .

t he fa ilu re o f s o me v ita l f u nct io n t he ind iv id u a l m ay p ers is t lo ng ,

aft er he is s t ripped o f ev eryth ing that m a k es lif e enj o yab le .

I t is so with rac es S o m e s eem end o wed with ph en o m enal

.

v igo r wh ile o th er s a re pres erv ed fr o m ext inct io n o n ly wi th the

,

grea t es t d iffi c u lty and by the na rro wes t m argin n o t o n ly becau se ,

o f lo w f ert ility b u t a ls o by r ea s o n o f inh er ent lack f v igo r Th

q e .

ex is t enc e o f th es e int erna l fo rc es is n o t a m att er o f d o u bt and th eir ,

ofl i c e in d irect ing v ariat io n is an int eres t ing and v alu ab le pro blem
wh ich t he pres ent s tat e o f kn o wledge is in s u ffi c ient to s o lv e .

Ort h ogen esis C los ely ak in to C op e s c o nc ept io n is Elm er s

’ '
.

th eory o f o rth o gen es is ( straight cr eat ion ) or as he calls it

3
, , ,

“
d efin it ely d irect ed ev olu t io n .
”

1 Pea rs o n , G ra m m a r S c ien c e p p 3 7 5 —
of
3 7
, 6 . .

3 E im er, O n h
O r t o gen es is a n d t h e I m p o rt an c e o f N a t u ra l S e l ec t io n in S p eci es
Fo rm a t io n ( p a mph l t e s, 6 a es )
5 p g O
[ p en C o u rt P u b lis in
g C o m p a n h y] .
2 06 C AU SES O F V A R I AT ION

th is V iew o f the case charact er s bad and good d ev elo p

In
t ogeth er The wors t o n es a re elim inat ed b u t m any u nd esirable
.
,

o r ind iffer ent o n es ar e left beh ind as n o t being s u ffi c ient t o t u rn

t he sca le aga in s t t he ind iv id u al o r t he rac e T h u s m any u nd es ir .

able charact ers linger ih a ll rac es and what is m o re to th e po int , , ,

u tility is n ot the ca u se qf ei ther the o r ig i n or the p er s is tenc e

4
c ha r a c ter s , but on ly of thei r ob l i ter a t i on when su
fi ic i en t ly d etr i

m en ta l to d es tr oy the i nd i v id u a l a nd theref or e cu t o f d esc en t in

tha t pa r t ic u la r l i n e .

T he wr it er hares t he Opin io n that th is is t he tr u e lim it o f the

s

s elect io n pr o c es s u nd er n a t u r e a n d that t he pr es enc e o f u nfa v o r ,

able charact er s argu es fo r th eir hav ing ar isen fro m ca u s es q u it e

ind epend ent o f s elect io n .

I n o u r yard s and fi eld s we c o ntro l select io n acc o rd ing t o what

ev er s tandard s we m a p lea s e t o es tab lis h b u t if u nfav o rab le char
y ,

a c t ers d ev elo p in nat u r e wh er e select io n a im s d ir ect ly at life will

, ,

th ey n o t b e lik ely to d ev elo p also u nno t ic ed u nd er o ur o wn s elec , ,

t io n es p ec ially wh en we d o all with in o u r po wer to preserv e life ?

,
1

T he pres enc e o r ab s enc e o f a pr inc iple a s id e fro m select io n ,

y et r es po n s ib le fo r t he pr es enc e o f cha ract er s t u rn s v ery largely ,

u po n t he q u es t io n a s t o wh eth er aft er a ll th er e a re well e s tab lis h ed , ,

-

in s tanc es o f chara ct er s ind epend ent o f u t ility and th er efo re o f ,

s elect io n T he pres enc e o f s u ch chara ct er s is ea s ily s h o wn

. Fo r .

exam ple what is t he u t ility o f t he s cro t u m a m o ng m a m m als ?

,

Wo u ld it n o t ha v e b een bett er with s h eep fo r in s tanc e if t he , ,

t es t ic les had rem a in ed with in t he abd om inal c av ity wh ere th ey ,

d ev elo p and wh ere th ey wo u ld b e safe in st ead o f d es cend ing

, ,

int o an ext ern al sa ck expo sed t o freq u ent inj u ry ? U nd o u bt ed ly

it wo u ld hav e been bett er fo r i n d i v id u a l s fo r m any hav e n ot o n ly ,

lo s t th es e o r ga n s b u t th eir liv es a s well fr om th is u nfo rt u nat e

, ,

po s it io n b u t the n u m ber los t is n ot s ufiic ien t to s er iou s ly afi ec t

the ra c e 2
I n o th er wo rd s s elect io n ha s a im ed at th is v u ln erable
.
,

1 I t is n o t ic eab e t l h
d ti t a t n a t u re a l o ws rep ro l uc on o g o on u n res t ric t e d , an d

th l y b y th
en s a s M t ff
e m i ll i
d th i wh lons . a n ca n n o a or s o es a e l d es t ru c t io n o f n u m

be rs .H k t p
e s ee t d i b l b i th
s o ki d
rev en un es ra e r s, a n of a d l
v a n c e s e ec t io n h
t at
has b t h it
o d tg d it d i d
s a v an at g es a n s sa v an a es .

3 Th i h w th t wh t i b d f th i d i id l i
s s o s a a s a or e n v ua s no t y b
n ec es s a r i l a d fo r t h e

ra c e l y wh t i b t f th
c o n v ers e ,
i ft h d
a s es or e ra c e s o en ar u
po n or ev en fa ta l t o th e

ind i id l T h i i t he
v ua .
y e f
s l csti v er ess en c o se e on .
 I N T ER N A L C A U SES OF V A R I AT I ON 207

poin t m an y t imes an d in n u m ero u s cas es with eff ect b u t mam

, ,

m als as a rac e hav e been able to end u re t he hand icap else th ey ,

wou ld lo n g s inc e ha v e bec o me ext inct .

T his S h o ws that s om e ca u ses o th er than u t ility are res po n sible

for t he appeara nc e an d c o nt in uance o f rac ial charact er s ; tha t
t eleo logy is not a u n iv ersal princ iple and that the f u nct io n o f
1
,

select io n is res tr ict iv e no t crea t iv e ,

.

Oth er charact er s n o t t eleo lo gical may eas ily b e m ent io n ed

1 The pec u liar m in u t e m ar kings o n d iat o m s and o n o th er
.

inc o n s pic u o u s o rga n is m s .

2 The green c olo r o f leav es d u e sim ply t o t he fact that

.
,

ch lo rophy ll is gr een T h is is n o m o re o f an inh erent n ec ess ity

.

than that coal s h ou ld b e black o r gold yello w .

3 T h e
. d ig ital n u m b er fi v e w h ich ru n s g en era lly thr o u gh
v ert eb ra t es wh ich has o ft en been m od ified and o ft en left intact
,
.

Certa in ly t he o rigin al n u m ber fiv e c o u ld n o t ha v e been teleolo g

ical I t is n ot en o u gh t o as s u m e that changed c o nd it io n s m ight
.

hav e rend ered an organ d etrimental wh ich was o nce u sef u l .

T h ere are too m any o b v io u s ly u seless cha ract ers .

4 T. h e ph o s ph o r e s c enc e o f p e lag i c an im a ls
3
and t h e pe a r l o f ,

the oy s t er wh ich is d u e to inj u ry

,
I s th is bea u ty u s ef u l o r is it .

acc id ental ? 8

5 T. h e br i ght c o lo r o f d ee p s ea fi s h es I s it any m o r e s ig
-
n ifi .

cant than the (acc id enta l) c o lor o f ch lo ro phy ll bear ing leav es 1 -

6 T he h o rn s o f s tags
. u s ef u l in batt le b u t weapo n s as
, ,

dangero u s to the po s sesso r as to his en em y .

T he lis t m ight b e ext en d ed in d efi n it ely Eim er s argu ment is

’
.

that ch a ract er s s u ch as these hav e been prod u ced n ot throu gh

select io n b u t in s p it e o f it and thro u gh t he agency o f o r gan ic ,

gro wth in d efin it e d irect io n s wh ich is or thog en es is I t wo u ld b e ,

.

d iffic u lt to b e alway s c ertain that n o basis o f u t ility ex is t s or

ev er has ex is t ed s im ply b eca u s e it is n ot n o w ev id ent y et n o ,

1 T he d oc t rin e t h at d ev elo p m en t is in li
ne wit h u ti l ity an d t h a t ev er t y hi ng is
u s efu l is kno wn as t eleo l gy
o .
”

3 Eim er O rga n ic Ev o u t io n , p x iii

, l . .

3 h
Eim er, O r t o gen es is p 1 0 , . .

1 I n c erta in lea v es o f b r ig t c o lo r h the c hl o ro p hy l l is u n a b le to d o m in a t e the

s t ro n ger s h d a es o f o t h er c h em ic a l s u b s t an c es , a n d t h e l eaf is not g reen b ut s o me

ot he r c o lo r.
2 08 C A U S E S O F V A R I AT I O N

caref u l s t u d ent o f ev olu t io n d o u bt s a n y lo nger th a t th ere are

m any m is fit s in nat u r e Wh eth er t hey ha v e aris en a s Eim er
.
,

as ser ts by r easo n o f org a n ic g rowth and wh eth er th ey a re ev i

, ,

d en c es o f d efi n i tely d i r ec ted d ev ia ti on is q u it e an o th er m att er

, .

T h ere is n o d o u bt o f t he per s is t enc e o f a cha ract er o nce s tart ed ,

ev en in t he fac e o f s elect io n b u t wh eth er it b e n ec es sary to

,

in v o k e t he a id o f an int erna l d irect iv e fo rc e to exp la in it is a

q u es t ion u po n wh ich mo re ev id ence is so rely n eed ed I t is .

exc eed in gly im po rtant fo r t he br eed er t o kn o w and rec o gn ize all

t he inh erent t end enc ies with wh ich he m u s t fi na lly reck o n an d ,

it m ay b e n ec es sa ry to go bey o nd phy s io logic a l u n it s c o rres po nd ,

in g t o ch em ical at o ms o r mo lec u les and in v o k e so m e fo rm of

,

“
gr o wth fo rc e co rres po nd ing to ch em ical en ergy to exp la in the
m y s t er ies o f d ev elo p m ent A n y th eo ry h o wev er that will ev en
.
, ,

reaso nably acc ou nt for th ese m y s t eries m u s t b e in t he present ,

s tat e o f kn owledge largely an a ss u m pt io n and let t he a s s u m

, p ,

t io n b e a s s im ple as po s s ible u n t il we can d efend it s c om plex ity

by a mas s o f well es tablis h ed fact s I n t he op in io n o f the wr it er
-
.

t he ex is t enc e o f s u ch a pr inc iple a s o rth ogen es is is m o r e than

pro blem at ical exc ept as it is an expres sio n o f t he relat ion s
,

that nat u rally o btain bet ween phy s io lo gical u n it s what ev er th ey ,

m ay b e .

SECT I ON X V Y
PH SI O LOG I C A L U N ITS

T he “
gem m u les of t irp o f G a lt o n, the
D ar win , t he “
s

id io p la s m igeli t he b io ph o r s “
“
d et erm inant s and ”
of N a
“ ” ”
'

, , ,

o f W eis m ann and t he phy s io lo gical u n it s o f o th er

“ “
id s
” ”

wr it er s a re a ll att em pt s t o exp lai n inh er itanc e o f d efi n it e q u a li

t ies by as s u m ing that the ger m c ell wh ich pass es o v er fro m
parent t o o ff s p rin g at t he t im e o f proc reat io n is c o m po sed of
d efin ite u nit s o f liv ing m att er each with its s pec ific pro pert ies
, ,

a mo ng wh ich a re n u t rit io n and m u lt ip licat io n wh ich t o geth er ,

w t —
c o n s t it u t e gro h and c o n s id ering t he s eparat e pro pert ies o f
,

t he d ifferent u n it s o f wh ich a giv en i nd iv id u al is c o m po s ed

gro wth in d efi n ite d i rec tion s .

I n s u ppo rt o f th is gen era l th eo ry it m ay b e u rged that the ind i

v id u a l is wha t he is v ery larg ely b ec a u s e o f i n ter n a l q u a li ties .

C o rn and wh ea t gro w S id e by S id e dra wing th eir n o u r is h m ent

,
2 10 CA U SES OF V A R I AT I O N

S o meth ing o v er eighty d is t inct k ind s o f m att er kn o wn ; a re

th eref o re s o m e eighty k ind s of at o m s a re a s s u m ed and th is ex ,

h a u s t s t he p os s ib ilit ies s o far as u n lik e l i he at o m s a re c o nc ern ed

u n les s o th er at o m s a re s u b s eq u ent ly d is c o v ered o r cr eat ed )
( .

B u t th is d oes n o t exha u s t t he po s s ib ilit ies o f m att er fo r th es e ,

ato ms c o m b ine t o geth er fo rm ing n ew u n it s (c alled molec u les )

,

with d is ti nct pro pert ies T h u s N a c l ( s o d iu m ch lo rid ) is d iffer

.

en t in ev ery way fro m ei th er t he s o d iu m o r t he ch lo r i n at o m s

that ha v e u n it ed to prod u c e it I n th is way thes e (eighty ) .

v ar io u s at o m s effect a ll s o rt s o f c o m b i n at io n s many o f th em ,

exc eed ing ly c o m p lex each c o n s t i t u t ing a n ew m at eri a l u n it a

1
, ,

s u ffi c ient ly large n u m ber o f wh ich c o n s t i t u t es a m ea s u rab le

q u ant ity of a s u b s tanc e wh o s e rea l c o m po s it io n c o u ld rarely b e
pred ict ed by any o f its v is ib le pro pert ies The fo llo wing tab le .

o f ch em ica l f o r m u lae is pr es ent ed fo r t wo p u rp o s es ( )

I t o S h o w
t he exc eed ing c o m p lex i ty o f o rd inary m ater ia ls ( )
2 to s h o w h o w
c ertain g r o ups qf a tom s (as C H2 o r C OzH) beha v e as u n it s
2
,

eff ect ing pr o f o u nd changes i n t he pro p ert ies o f th ei r c o m po u nd s .

I t is ev id ent that the pos s i ble c om b i na ti on s ev en with t he few ,

mos t c o m m o n at o m s a s C H O ,
N F e N a K P S —, are , , , , , , , ,

pract ically infi n ite wh en th ey a re ab le t o o rgan ize th em s elv es

int o larger u n it s g iv ing ris e to c o m p lex s er ies lik e t he tab le on
,

the fo llo wing p a ge

3
.

I n th is tab le the rad ica l C OQH r u n s thro u gh t he ent ire s eries ,

giv ing a k ind o f gen et ic q u ality to t he c o m po u nd s wh ile s pec ific ,

d ifferenc es acc o m pany t he v a ry ing nu m bers o f C a n d H at o m s

pres ent with t he rad ical I t is to b e n o t ed h o wev er that th es e
.
, ,

C and H at o m s a re in d efin it e pro po rt io n to each o th er na mely , ,

C l l z 1 that is fo r ev ery a t o m o f C th ere will b e on e m o re tha n

n n + ,

t w ic e a s m a ny a to m s of H + C O2 H — a ll o f wh ich is extrem ely , ,

s u gges t iv e a s ear ly s t ep s in t he wo rld o f o rgan ized m att er .

1 T he c o m p o s it io n of s t r c lm in e,
y C 21 H 22 N 2 0 2 , an d h t at of gra p e s u ga r,

C u f f s-
20 1 1 , a re bo t h ex c ee d i gl yn s im
p l e as c o m pa re d with m an y k no wn s ub

s ta n c es .
3 S uc h gro u p s o f a t o m s t h at mo v e h
t o ge t er a r e k no wn as
“
ra di l ca s . T h y e

a re in ev er y s en s e u n its an d a re ca p a b e o f rep a c in g o r o f l l di pl
s a c in g o t h er a t o ms

in t h eir c o n s t ru c t io n s .

11 We a re t o l d b y th h e c em is t s t h at m o re t h an o n e h d d th
un re o u san d s ep a ra t e

c o mp o u n d s a re n o w kn o wn .
 I NT ER N A L CA U SES OF V A R I AT I ON 2 1 1

M ONOBAS IC AC IDS OF T HE A C ET IC SER IES , Cn Hm fl COgH

S o u R ea FO R MUL A

red an ts , n et t es l
v in e
ga r

d
o x i a t io n o f o ils

ra n c i db u t t er

l
v a eri a n ro o t

ra n c i db u tt er

o xi d a tio n o f cas to r o il

ran c i db u t t er

geran i u m l ea v es

ra n c i db u t t er

o il o f ru e

y
b a b erri es
c oc o a n u t o il

nut m eg b u t t er
A g a r i c u s i n teg er (a fu n gu s )

p a lm o i l

ta ll o w

b u t t er

b eec hwv o o d ta r

b ees wa x

Ob ser v e t he exc eed ing c o m p lex ity o f these c o m po u nd s and ,

n o t ic e that they s tand in d efin ite series with u n if o rm d ifferenc es

there a re n o break s in the series and n o m is sing m em bers u nt il
we reach C 2 1 H43 N o t e to o that th es e c o m p o u nd s may th em
.

selv es c o m b in e wi th o th er s ; th u s s t eari n t he cha ract eris t ic fat,

o f ta llo w is C H
, 3 5 (
C 18
H35 02 )3
i n wh i ch ca
,
s e 3 H fr o m t he ac id
rad ica l and 3 HO fro m t he g lyc eri n rad ica l ha v e u n it ed t o f o r m
( )
H
3 ( z ) O ,
o r wat er .

A bo v e ev eryth ing els e in th is s eries n o t e t he wid e d ifierenc e

'

i n p hy s ica l p r oper t ies l ig ht d lfier enc e i n c hem ic a l

'

a r is i ng f r om a s
2 I 2 C A U S ES O F V A R I AT I ON

con s t i tu t i on . xam p le it is s ign ific a nt that in th is s er ies

Fo r e ,

bo t h the s o lu b ility in wa t er and t he ac id s treng th d im inis h as

t he pro po rt io n o f carb o n increa s es .

I n add it io n t o the pro pert ies o f n on li v i ng u n i ts t he th eo ry o f -

h s i olog ica l u n i ts as t he ba s is fo r s pec i fic ch a ract er s in liv ing

p y
m att er req u ires o n e d is t inct iv e and add i t io na l q u a lity na m ely ,

lif e with it s att n ant ph n

,
e d e o m ena — t he po wer o f n u tr i t io n ,

and gro wth B u t what are n u trit io n and gro wth ? Co n s id ered
.

in gen era l t er m s n u tr it io n is S im p ly t he po wer o f o n e ch em ica l

,

c o m po u nd ( t he liv ing) to ent er int o t he c o m po s it io n o f ano th er

( t he f oo d ) and break it u p and r eadj u s t it s elem ent s o n a ba s is

lik e i t s o wn lea v ing t he res id u es t o tak e care o f th em s elv es

,
.

T h is readj u s t m ent o f t he n o n liv in g f o o d t o t he c o m po s it io n

-

o f t he liv ing p lant o r an im al we ca ll n u tr it io n and it m ean s ,

es s ent ia lly an inc rea s e o f t he liv ing at t he expen s e o f t he no n

liv ing ; in o th er wo rd s g r o wth thro u gh t he n u m erica l increa s e

,

o f liv ing u n it s .

T h ere is o ft en rea dj u s t ment in t he n o n liv ing wo rld wh en t wo -

c o m po u nd re b r gh t t o geth er
s a o u — t he weak er g iv ing way t o
,

t he s tro nger a ffin ity — b u t th ere is n o s u ch wh oles a le

,
carry “

in g o v er o f m att er fro m o n e t o an o th er a s in t he ph en o m ena

”

we ca ll n u t rit io n and gro wth T h is is a tru e in v a s io n o f t he

.

n o n liv ing by t he liv ing wo r ld tran s f err ing m a tt er a lm o s t ind ef

-
,

in itely u n t o it s elf n o t o n ly pr es erv ing i t s o wn id ent ity in the

, ,

m eant ime b u t im pres s ing it u p o n the appro priat ed m at eria ls

as well .

T h es ephy s io lo gica l o r V ital u n it s a re th erefo re c o nc eiv ed to

b e t he s m a lles t liv ing u n i t s lik e m o lec u les in n o n li v ing m att er
,
-
,

exc ept that th ey a re far m o re c o m p lex i n c o n s t it u t io n and a re

en d o wed with t he p o wer o f s elf m u lt ip licat io n throu gh n u t ri t io n

-
.

T h is req u i res gro wth and d iv is io n aft er t he m an n er wh ich has

been n o t ed in chro m at in gran u les exc ept o n a scale infi n it ely ,

m o r e m in u t e .

T h is c o nc ept io n o f t he act io n o f liv ing u n it s has it s s im ilit u d e

in t he n o n liv ing -
C ry s ta llizat io n is a gro wth in t he s en s e o f
.
,

increas e o f S ize b u t it is n o t att end ed by tran s fo r m at io n s eq u al

,

to th o s e in liv ing m att er F u rth er mo re cry s ta llizat io n is gro wth

.
,

with o u t d iffe rent iat io n exc ept as t o geo m etric fo r m

, E ith er .
2 14 C A U S ES O F V A R I AT I ON

of e xtreme impro b ab ility On t he o th er ha nd if infl u enc es are .

1
,

d is c o v ered wh ich a re really effic ient in sett ing bo u nd s t o v aria

b ili ty th en th ey m ak e t he tran s f er o f charact ers fro m parent t o
,

o ff s pri ng s o ab so lu t ely c ertai n reg u lar a n d fix ed a s to s eem to , , ,

leav e litt le o r n o po ss ib ility o f v ariat io n .

T h is latt er is t he ca s e with t he hypo th es is o f phy s io lo gi c al

u n it s . Weis mann rec ogn ized its lim itat io n s and pro po sed t he
th eo ry o f ger m inal s elect io n t o acc o u nt fo r v ariat io n as well as
3

inh erit anc e .

T h is th eo ry a ss u mes that t he b io ph o r s o r th e phy s io lo gi ca l ,

u n it s by what ev er na m e th ey m ay b e ca lled a re engaged in a ,

k ind o f s tru gg le amo ng th em selv es with in the ger m m u ch as ,

are p lant s and an im als in t he larger wo rld o u t s id e .

A n y th eo ry o f phy s io lo gica l u n it s m u s t inc lu d e th eir ab s o rp

t io n o f food and th eir po wer o f self m u lt iplicat io n I f th es e -
.

act iv it ies proc eed at a u nif o r m ra t e fo r each u n it in v o lv ed th en ,

n o v ariat io n wo u ld res u lt fr o m th is m u lt ip licat io n b u t if pr o p o r

t io n s change o r if t he v itality v a r ies th en v a riat io n wo u ld n ec os
, ,

s a rll
y r es u lt fro m th es e ca u s es a lo n e N o w th es e act iv i t ies m u s t.

b e eith er c o n s tant o r v ariab le Weis m ann a s s u m es that th ey .

a re v ar iab le ; that th es e u n it s o f v ario u s relat iv e n u m b ers and

s trength s a re c o m pet it o r s a m o ng th em s elv es o n e with an o th er , ,

fo r food ; and that th o s e mo s t en erget ic in f oo d ab s o rpt io n and

capab le o f t he mo s t rap id m u lt ip licat io n will n o t o n ly b e the
mo s t v igo ro u s b u t th ey will a ls o ex is t in relat iv ely the larges t
n u m bers Th ey will th erefo re t end the m o re to im pres s th eir
.

charact ers u po n t he lat er d ev elo p m ent o f t he ind iv id u al .

U n d er th is v iew o f the cas e t h e ba lanc e o f po wer is c o n

”

s t an t l s h ift ing a lw ay s in fa v o r o f t he m o s t v igo rou s and

y ,

ra p id ly m u lt ip ly ing u n it s Believ ers in phy sio lo g ical u n it s m u s t

.

eith er f o llo w W eis m ann in th is c o nc ept io n o r els e a s s u m e o n

t he part o f t he u n it s ab s o l u t ely eq u a l p o wers o f n u trit io n and

m u lt ip licat io n fo r m u lt ip licat io n th ere m u s t b e if s u ch u n it s
,

av a il anyth ing in the rOle o f inh eritanc e .

1 A ll t h i g n id
s d i h it
c o ns e re d , t
n i ti
er i t h m y t y Th
anc e an no v ar a on s e s er . e

w d
on er i t t h t i d i id
s, no a l y nb t t
v h t t h
ua sy f ll w
v ar , l l
u y t h
a y d e o o as c o se as e o

t h t yp
e f h
e o t t wh i h t h y b l g
e ra c e o c e e on .

3 W i m e s O G mi l S l ti
an n , n er S
na f D fi it V i t i
e ec on as a (p m o u rc e o e n e ar a on a

p h l
e t,
d
s ec o n d it i e O
) [ p C on t P b l i h ien
g C m po ur y ] u s n o an .
 I NTER N A L CAU SES O F V A R I ATION 2 1 5

Weis m ann remind s u s that by fa r the larges t part of “

q ualitat iv e mod ificat io n s res t o n q uan t itat iv e changes .

A d eter m inant say s he

“
m u s t b e c o mp o sed o f h et ero
"

, ,

ge n eou s b io ph o r s and o n th
,
ei r n u m e r ica l pro p o rt io n r ep os es ,

acc o rd ing to o u r hypo thes is th eir s pec ific nat u re I f th is p ro

,
.

po rtio n is a ltered so also is t he cha ra ct er of the d et er minant ;

,

and f u rth er : fo r fl u ct u at io n s o f n u triment and t he s tr u ggle

“

fo r n u triment with its s eq u ent preferenc e o f t he s tro n ges t

, ,

mu s t tak e p lac e b et ween t he v ario u s s pec ies o f b io ph o rs as

well a s bet ween t he s pec ies o f d et erm inant s B u t cha n ges in .

the q u ant itat iv e ra t io s o f t he b io ph o rs app ear t o u s q u ali tat iv e

changes in the c o rres po nd ing d et er minant s 1
A n d again “
By .
"

a s elec tion of the h i n d r e d h m i s s i v ely m od i

f e r r e t o t e g er s p g
r o r e

fi ed i n a ma n n er p w
corr es on d i ng
i th the p rod u c t i on qf a d e n i tely
fi
d i rec ted p rog ress iv e v a r ia tion of the pa r t 3
I n th is way Weis
”
.

man n wo u ld “
exp lai n Eimer s o rt h o gen es is ; b u t it is n o te
” '

wo rthy that n on e of the theor i es y et p ropos ed wi ll a cc ou n t f or the

or ig i n a l i n t rod u c tion f
o a n ew c ha r a c ter in the ra ce, wh eth er
represent ed and tran s mitt ed by a phy s iolo gical u n it o r n o t .

G er minal select io n wou ld pro v id e fo r changes in relat iv e p ro

po rt io n s of cha ract ers an d ev en fo r th eir u tt er ext inct io n b u t
, ,

n o t fo r th ei r intro d u ct io n u n less ind eed charact er s may o rigi

, , ,

nat e by n ew c o m binat io n s o f old element s .

On e is a lm o s t fo rc ed to the c o nc lu s io n that in nat u re loss

o r m o d i ficat io n o f charac t ers is far m o re c o m m o n than th ei r

o r ig in and intro d u ct io n I t loo k s a s th o u gh mo s t o f t he cha n ges

.

arise in th is way yet it is c o nc eiv ab le that an ent irely n ew

,

q u ality m ight aris e thro u gh a relat iv ely S light m od ificat io n o f

the ch em ica l o r phy s io lo g ical m ak e u p o f t he V i ta l u n i t s I t is
-
.

s een and rec o gn ized that in the n o n liv ing wo r ld a s light change -

in the ra d ical is f o llo wed by a pro fo u nd a lt eratio n in phys ical

and chemical pro pert ies and that th is s weep ing change m ay b e
,

ind u c ed by c o m parat iv ely s light and ev en ext ern a l cau ses .

May n o t t he s am e b e tru e o f v ital rad icals o r u n it s and may ,

n o t n ew charact er s arise m o r e r ead ily than we s u ppo s e a ll per ,

hap s ou t of elem ent s fewer and t ra n s fo r mat io n s s im pler than

, ,

we hav e h ith ert o im agined

1 W i m
e ss O G mi l S l t i
an , n er ( na d d it i e ec 6
) pp 4 4 7 on I b id
s ec o np 35 e on , .
- .
3 . . .
2 1 6 CA U SES O F V A R I AT I ON

Con t rol of in t er nal c au ses affec t in g the r ace as a whol e .

What ev er ca u s es o f th is nat u re m ay b e at wo rk in o ur fi eld s

and yard s — and th ey a re t o b e reck o n ed with
,
o u r c o ntro l ,

o v er th em is s ec o ndary and ind i rect T h eir effect s if pres ent

.
, ,

a re at o nc e i n s id io u s and s weep ing .

W e can b e m indf u l o f t he effect s o f gen etic s elect io n and the

s elect iv e d eath rat e and pro v id e aga in s t th em at lea s t to a
, ,

large d egree I f g ro wth f o rc e and o rth o gen es is a re a ls o fo rc es

.
,

th ey can b e as s is t ed o r h eld in ch eck by select io n b u t th ey can ,

n ev er b e ab so lu t ely c o ntro lled and if germ inal s elect io n is a fact ,

i t is go ing o n ent irely in d epend ent o f any c o ntro l wh ich pres ent
kn o wledge enab les u s to ex erc is e exc ept thro u gh s elect io n .

S u mm ary A ll that is in v o lv ed in h ered ity is c o nta in ed in a

.

m in u t e b it o f liv ing m att er pa s s ed fro m parent to o ff s pring and ,

wh o s e d ev elo p m ent will c o n s t it u t e the n ew in d iv id u a l T he im .

p u lse to d ev elo p m ent th erefo re and it s f u nda m enta l po ss ib ilit ies

, ,

a re f o rc es int erna l t o t he ger m and t o t he liv ing o rga n is m .

I t is n o t d iffi c u lt to s ee m any ca u s es o f v ariat io n in t he

int er na l proc es s es kn o wn t o b e in v o lv ed in the act iv it ies o f
liv ing pro t o pla s m Gro wth is t he res u lt o f c ell d iv is io n wh ich
.
,

s eem s t o pro c eed u p o n p lan s calc u lat ed to in s u re q u a lit a t iv e

as well as q u ant i tat iv e eq u ality as bet ween t h e da u ght e r c ells .

A n y d ev iat io n fro m t he p lan h o wev er — and d ev iat io n s a re

, ,

kno wn to o cc u r — m u s t resu lt in v aria t io n T h is is es pec ia lly

,
.

tr u e in t he red u ct io n pro c es s wh ich is cha racteris t ic o f m at u ra

t io n in bo th s ex es and wh ich pro bab ly lies at the bas is o f b u d
,

v ar iat io n a n d o f m any m u tat io n s .

F ert ilizat io n and s ex u al u n io n a re pro c ess es calc u lat ed to

eff ect n ew c o m b inat io n s o u t o f t he elem ent s in v o l v ed th o u gh ,

t he p o s s ib ilit ies in th is d i rect io n wo u ld b e ra p id ly red u c ed by

c lo s e breed ing o r by any o th er c irc u m s tanc e which s im p lifi es
t he anc es t ry .

D o u bt les s t he c o nd it io n o f t he germ ha s s o me infl u enc e b u t ,

it is n o t well u nd er s t o o d T he ph en o m en o n o f x en ia o r d o u b le
.
,

f ert ilizat io n in c erta in p lan t s ca u s es t he s eed c o at s to v ary t he

,

fi rs t y ear in the s am e d irect io n as the ger m Telego ny is a .

myth and int ra u t eri n e infl u enc es a re d o u bt less lim it ed t o th ose

,
-

o f n u tr it io n exc ept in ca s es o f d is a s t er
, .
 2 18 CA U SES OF V A R I ATI ON

E V OL U TI ON ON PR E D E T E R M I N E D LI N E S . B y T D A C oc . . . k erell . S c ien c e,
X II I, 3 1 312 1- .

E x PE R I M E N T A L S T U DY OF V A R I A T I O N . By J . C . E war t . R epo r t o f the

B r i tis h A ss o c ia tio n fo r th e A d v a n c emen t of S c ien c e, LX X I , 666

6 80 .

E x PE R I M E N T s IN C R OS S I N G C OR N AND W A TE R M E L ON S . B y F C C a rd . .

an d G . E A d a ms
. E xper im en t S ta tio n R ec o rd , X I I I , 7 4 0
. .

E x PE R I M E N T S U PO N T HE I N F L U E N C E OF T H E S E x U A L C E L L S U PON T HE

S OM A T I C . . W . Fi eld
B io lo g i c a l B u lletin , I I , 3 46 - 3 4 7
By G . .

H ET E R OT V P I C A L D I V I S I O N I N M A T R A T I O N B y T H B r c e R epo rt U . . . y .

o f th e B ri tis h A ss o c ia tio n fo r the A d v a n c emen t o f S c ien c e, 1 90 1 ,

pp 6
. 8 5 6 8 7 — .

HY B R I D I Z A T I O N OF C O R N A N D W A T E R M E L ON S B y F C C a rd E xperi . . . .

fn en t S ta tio n R ec o rd , X 1 92 8 , .

HV B R I D I Z I N G M E L ON S : E F F E C T I N S U G A R C O N T E N T Experim en t S ta ti o n .

R ec o rd , X V I , 2 29 .

HY B R I D I Z I N G ZE BR A AND HO R S E .
( E x per i men ts of B a ro n d e Pa ra n a i n
B ra zi l E x per i m en t S ta tio n R ec o rd , X I ,
) .
9 7 2 .

I N D IV I D U A L I T Y O F C HR O M O S O M E S B y H Metc alf . . . Proc eed in gs of the

N eb ra s k a A c ad emy o f S c ie n c e, 1 90 1 .

I N II E R IT A N C E IN PA R T HEN OG E N E S I S . B y E rn es t W arren . Pro c eed ings

o f the R o yal S o c iety, LX V , 1 54 -
58 1 .

OR G A N I C S E L E C T I ON . By J . M . B a ld win ( S c ien c e, I V , 7 2 4 7 2 5 ; V , -

634 E D C ope ( S c ien c e, 1 1 , a n d H F Os b o rn S c ien c e,

. . .
( .

V 1. 5 83
”

O R T II OG E N ET I C V A R I A T I O N . By H . F . G ad o w, C a mb rid ge . S c ien c e,
X X I I , 6 3 7 - 6 40 .

O R T H OG E N E T I C V A R I A T I O N IN C E R T A I N M E x I C A N S PE C I E S O F L I ZAR DS .

B y H F G ad o w Pro c eed i n gs of the R oyal S o c iety, L X X I I , 1 09- 1 2 6

—
. . . .

PH Y S I O L OG I C A L S E L E C T I O N . By G .
J . R o m a n es . S c ien c e, V I I , 606 60 8 .

PR E M A T U RE FE R TI LI Z ATI ON I NJ U R I OU S TO FR U IT .E x per im en t S ta tio n

R ec o rd , X I V , 6 3 4 .

PR O B L E M O F D E V E L OPM E N T . By E B . . W ils o n . S c ien c e, X X I , 2 8 1 - 2 93 .

S E C O N D A R Y S E x U A L C HA R A C T E R S ( A s tu d
y o f the effec ts . o f c as t r a tio n .
)
B y S G S a ttu c k an d C G S eligm a n n
. . h Pro c eed in gs . . . of the R o yal
S o c iet , L y
XX I I I , 49 5 8
-
.

S EX UAL U
R E PR O D C T I O N ; D I S T I N C T I O N F R OM A s E xU A L . B y R ic h ard
Hertwig ( tra ns la ted b y W C C u r tis ) . . . S c ien c e, X 1 1 , 94 0 94 6 -
.

S ON G OF BI R D S K E PT F R OM T HE I R OW N S PE C I E S B y W illia m E . . D .

S c o tt . S c ien c e, X I X , 1 54 .

STE R I L E FR U IT B LO S SO M S . B y S A B ea c . . h . N ew Y o r k S ta tio n B u lleti n ,

C LX I X , 3 3 1 3 7 1 -
.

S T E R I L I T V O F C A TT L E , C A U S E S OF . E x perim en t S tatio n R ec o rd , X 1 , 2 89.

T E L E G ON Y . By J . C . E wa rt . Pro c eed in gs of the R o yal S o c iety, L X V ,

2 43 - 251 .
 I NT ER N A L CA U SES O F V A R I ATI ON 2 19

T E L EG ON Y ( D I S P R OV E D ) . By J . C . E war t . Po p u lar S c ien c e M o n th l y ,

LV I I , 1 2 6- 1 3 3 .

T E L EG ON Y ( D I S PR O V E D
) : A CCO U NT OF E WA R T
’
S E xPE R I M E N T W I T H
Z E B R OI D S . By J . C . E wa r t . B reed ers
’
G a zet te, X L I , 1 00 9 ; also in
T ra n s a c tio ns of th e Hig h la n d A g ri c u ltu ral S o c ie ty , 1 90 1 , pp 8 1 1 34 ;
.
-

and in E x per imen t S ta tio n R ec o rd , X I , 1 07 7 ; X I I I, 2 7 5 ; an d X I V ,

76 .

T E L E G ON Y ( D I S P R OV E D ) : E x PE R I M E N T W I T H G U I N EA PI G S . By C . S .

M in o t . R epo rt of the B r itis h A ss o c ia tio n fo r the A d v a n c emen t of

S c ien c e, 1 906 , p 606

. .

T HE C H R O M O S O M E S IN HE R E D I T Y . By W . S S u tton
. B io log ic al
B u lleti n , I V .

T HE G E R M PL A S M . By A . W eis man n . 1 v ol .

T H E PH Y S I C A L B A S I S O F HE R E D I T Y . By K . H E igen m an n
. . Po pu lar
S c ien c e M o n t ly, LX I , 3 2 —
44 h .

T H EO R Y O F HE R E D I T Y A N D T E L E G O N Y . By J . C . E wa r t . N a tu re, LX ,
33 0 —3 3 3
X E N I A , E xA M PL E IN A PP L E . By L . H B a iley
. S c ien c e, I V , 4 98 4 99
.
-
.

X EN I A , OR D OU B L E FE R TI LI Z AT I ON R v iew o f wo r k o f D e V r ies ,
.
( e

C o rren s , and W eb b er ) . E x p er im en t S ta tio n R ec o rd , X I I , 4 2 1 , 7 1 7 ;

als o X I I I, 6 20 .

XENIA IN MAI ZE . By Hu go D e V ries . E x peri m en t S tat io n R ec o rd , X I ,

1016 .
 CH A PTER I X
EXTER NAL INFLU ENCES AS CAU SES OF V ARI ATI ON

I t was lo ng n o t ed and the m os t cas u al o b s er v er cann o t

ago ,

fail to d isc o v er that ind iv id u als o f the sa m e s pec ies v ary grea t ly
,

acc o rd ing to th eir en v i ro n m ent m ean ing by that t erm all t he

,

ext ern a l c o nd it io n s o f lif e s u ch a s c lim at e f oo d fr iend s en em ies

, , , , ,

and all th o se o u t s id e infl u enc es fav o rab le o r u n fa v o rab le a mo ng

, ,

wh ich t he in d iv id u a l find s it s elf bo rn and with wh ich it m u s t ,

liv e u po n t he b es t t er m s po ss ib le if it wo u ld liv e at all T hat .

th es e ext ernal agenc ies ex ert a d irect effect u po n liv ing matt er
is b ey o nd q u es t io n a n d it rem a in s to g iv e att ent io n t o t he natu re
,

and ext ent o f th is infl u enc e as a part ial an s wer to the q u es t io n

we wo u ld s o lv e the d epend enc e O f o rgan ized liv ing m att er
,

u po n t he ext erna l wo r ld fo r t he n a t u re and range o f i t s act iv it ies .

A nyth ing we m ay learn u p o n th is po int will b e a c o ntrib u t io n to

t he s t ock o f kn o wledge o u t o f wh ich we s ha ll o n e day d et er m in e
a ll t h e ca u s es o f v ar ia t io n .

W ith o u t a d o u bt t he great b u lk o f v ariab ility is d u e to

ca u ses int ern a l to the o rgan is m m a in ly in the fo rm o f inh erited
,

t end enc ies Pears o n af ter exhau s t iv e s tat is t ical in v es t igat io n s

.
, ,

rem ark s T he ind iv id u al c o ntain s with in it s elf o wing t o a bath

,
“
,

mic la w o f gro wth a v ariab ility wh ich is it s elf q u it e s en s ib le

, ,

being 80 o r 90 per c ent o f the v ariab ility o f t he rac e 1

.

E v en th en h o wev er th es e int erna l in flu en c es a re d epend ent

, ,

u p o n o u t s id e c o nd it io n s fo r th ei r o ppo rt u n ity A b o rn g iant .

m u s t ha v e f o o d in ab u ndanc e b u t n o a mo u nt o f fo o d wo u ld
,

m ak e a g iant o u t o f a d wa rf N o r will it a v a il to a wak en lat e

.
,

in lif e f o rc es that o nc e m ight ha v e been act iv e S o m e d warf s

,
.

a re th eref o r e b o rn and o th er s a re pro d u c ed by in su ffi c ient f o o d .

T he ext erna l c o nd it io n s o f lif e a ffect v aria b ility in fo u r d is

t in c tly d iff erent way s ( )
I thr o u gh nat u ra l s elect io n in fl u enc ing ,

1 Pea rs o n , G ramm a r of S c ien c e, p 4 7 3

. .

2 20
222 C A U SES O F V A R I ATI O N

A m erican d growers generally s eem to ha v e s ett led u po n

s ee

K an sas as the s po t m o s t fav o rable t o the d ev elo p ment Of the

h igh es t q u ality in the wat er m elo n and it is acc o rd ingly the ,

fav o rite s eed prod u c ing locality D ar win t ells u s that the s eed
-
.

o f the Per s ian melo n y ield s n ear Paris a fru i t inferio r t o the

poo res t mark et k ind s b u t at Bo rd eau x yield s d elic io u s fru it

,
1
.
"

E u ro p ean v ariet ies o f g rapes fa iled s o u tt erly in ea s t ern N o rth

A m erica as t o nec es s itat e the d ev elo p ing o f v a riet ies fro m t he
nat iv e V in e .

I nd ian c o rn d ev elo p s loca l v ariet ies wi th extrem e rea d in es s ,

b u t th ey s eld o m s u cc eed wh en t ran s ferred ev en s h o rt d is tanc es ,

at leas t u nt il t ime en ou gh fo r acc lim at izat io n has elap s ed The .

writ er s ent a s tandard wh i t e Illin o is c o rn ripen ing in ab o u t a hu n ,

dred and t wenty day s and capab le o f m ax imu m y ield s (sev enty fiv e -

b u s h els per acre) to b e gro wn in M ich igan W is c o n s in M aine and

, , , ,

M is s is s ipp i 3
I n M a in e it fa iled t o ripen b u t at all o th e r p o int s
.
,

it ripen ed in abo u t a h u ndred day s pro d u c ing s m all inf er io r , ,

ears a lto geth er wo rth les s a s a c o m m erc ia l cro p

, T hat it s h o u ld .

h u rry thro u gh its period of gro wth at the no rth was n o t s u rpris
in g b u t that it s h o u ld d o t he s a me at t he s o u th wh ere i t had
, ,

ev en m o re t ime at its d is p o s a l than at h o m e is u nacc o u ntab le , .

Wh eat o n the o th er hand is a c o s mo po litan cro p and wh ile

, , ,

v a riet ies s u cc eed bett er in s o m e lo ca li t ies than i n o th er s et a

y ,

n ew v ariety s eld o m fails and so met imes s u cc eed s ev en bett er

,

than in the locality whenc e it cam e Ho wev er it is alto geth er .

,

lik ely t hat n o kn o wn wh eat gro wing regio n eq u a ls E ngland in -

nat u ra l ad v antages fo r max im u m y ield s Th is is s u ppo s ed t o b e .

d u e to t he h u m id at mos ph ere and c lo u dy s k ies d u ring t he lat er

s tages o f gro wth in s harp c o ntras t to t he b right s k ies and ho t
,

d ry ai r o f A m erica at th is s ea s o n I f th is b e the tr u e partial .

explanat io n o f the o cca s io na l ph en o m ena l and a lway s h igh y ield s

i n Great B rita in we may h o pe fo r eq u al res u lt s some day in t he

3
,

s im ilar c lim at e o f O rego n .

1 D ar win , A n im a ls an d Plan t s , I I , 2 64 .

3 W is c o n s in , 2 00 miles n o rt h; M ic iga nh , 2 00 m iles n o rt h an d 1 00 m iles eas t ;

Main e 300 m il es
, n o rt h and 900 mi l es eas t ; Mis s iss ippi 4 5 0 miles s o u th
, .

3 The a v era ge w h ea t y i ld
e of th e U n it ed S ta t es is b etween 1 2 a n d 1 3 b u s h els
pe r ac re, h l th
w i e at o f G rea t B ri tain is a lmo s t exa c t l y 30 , an d a ma xim u m of 90

b us h els has b een repo rte d .

 EXT E R N A L I N FLU EN C ES A S CA U SES OF V A R I ATI ON 223

T he xperim ent s of B on nier u po n th is gen eral s u bj ect are

e

c las sic He d iv id ed dand elio n h elianth u s an d m any o th er

1
.
, ,

p lant s growin g in t he v a lley an d p lant ed on e h alf o f eac h i nd i ,

v id al p lan t o n t he m o u ntain at an elev at io n o f 2 3 00 t o 2 4 00

u
m et er s leav ing t he o th er h alf in t he v alley ( how m u ch belo w
,

he d oes n o t say) A s t he d iv i sio n was m ad e b y a v ert ica l c u t

'

thro u gh t he fl es hy roo t t he t wo ha lv es m u s t hav e been p rac t i

,

cally alike .

He fou nd that the p o rt io n s o n t he m o u ntain d ev elo ped p lant s

m u ch sm a ller than th o se in t he v alley s T he d ifferenc e was .

m ain ly in the lengt h o f the int ern od es n o t in th eir n u m ber ,

.

T he lea v es o f t he d a nd elio n were les s than o n e fifth a s lo ng

wh en dra wn t o s c ale and t he fl o wer s ta lk s were n o t on e t enth as
,

lo n g T he m o u ntain p lant s in g en era l d ev elo ped h igh er c olo r s

. .

D arwi n t ells u s that t he med ic inal q u a lit ies o f d igi talis a re

eas ily a ffect ed by c u lt u re and that the wo od of the A mer ”

ican loc u s t tree ( R ob i n ia ) wh en gro wn in E ng land is n early

wo rth les s a s is that o f the o ak tree wh en gro wn at the C ap e o f
,

Good Ho pe 3
T he s a me a u th o r q u o t es S ir J E Tenn ent as . .

s ay ing that in t he Bo tan ic Gard en s o f C ey lo n t he app le tree

“

s end s o u t n u m ero u s r u nn ers u nd ergro u nd wh ich c o nt i n u a lly

‘
,

rise int o s ma ll s t em s and form a gro wth arou nd the parent

,

tree 3
If th is b e tr u e its nat u rally s light t end ency t o s pr o u t
’
.
,

has in th is loca lity d ev elo p ed int o a pro no u nc ed hab it .

S h eep es pec ia lly the m erin os a re c o s m o p o litan and yet th ey

, , ,

s u cc eed n o wh ere els e a s in N ew Z ea land On t he o th er hand .

,

acc o rd ing to D ar win th ey s eem n o t t o s u cc eed i n the Wes t

I nd ies o r o n t he wes t c o as t o f A frica wh ere t he wo o l disa p ,

pears from t he wh ole body exc ept o v er t he T he wri t er

has seen t he s a m e th ing in B razil exc ept that t he bes t c lo th ed ,
-

po rt io n o f the bod y was t he back o f the neck t he s a me s po t o n ,

wh ic h t he v ic u na bears it s fu r .

T he s tat em ent is freq u ent ly m ad e that fat ta iled S h eep rap id ly -

lo se th is charact er wh en rem o v ed fro m th eir nat iv e s a lin e pas

tu res b u t t he ass ert io n n eed s c o nfi r m at io n fo r the writ er ha s
, ,

1 C B o n n ier, R ec
. h erc he s ex p l
l a d a p t a t io n d es p la n t es n u
é rim en ta es s u r
’
c l im at

alp in . A nna l °
ll
es d es s c ien c es n a t u re es , 7 S é rie, T o m e X X , 1 8 95 .

3 D a rwin , l l
A n im a s a n d P an ts , I I , 2 64 3 Ibi .
p 2 66 d 1
.I b i .I , . d . 1 02 .
2 24 C AU SES OF V A R I AT I O N

s ee n a res pectab le d ev elo p ment o f fat wh en the s h eep were k ept

u nd er o rd inary c o nd it io n s .

On e o f t he m o s t rem arkab le and s eem ingly bes t a u th ent ic a t ed

in s tanc es o f t he ev il i nfl u enc e o f loca lity u po n charact er d ev el
o m en t is t he a lm o s t u n i fo r m f a ilu re to m a int a i n t he q u a lity o f
p
c ertain Englis h breed s o f d ogs wh en bred in I nd ia We a re .

ind ebt ed agai n t o D ar wi n fo r t he remarkab le s tat em ent that in

1

that c o u ntry the b u lld o g rap id ly lo ses his feroc ity and o f all ,

d o gs the h o u nd s d ec line m os t ra p id ly .

I n s tanc es m ight b e m u lt ip lied ind efin it ely b u t t wo th ings ,

m u s t b e bo rn e in m ind by t he s t u d ent wh en d ea ling with th is

c la s s o f fact s fi rs t th ere is t he greates t o ppo rt u n ity fo r erro r
,

o r exagg erat io n fr o m i n exact o b s erv at io n and rep o rt ; s ec o nd ,

t he p lant o r an i ma l is exp o s ed to a m u lt it u d e o f n ew c o nd it io n s
wh en t ra n s plant ed t o a n ew lo ca lity o n ly a p o rt io n o f wh ich a re ,

i nh erent i n t he c o nd it io n s o f life C o mmo n ly t he breed e rs o r .

att endant s are n o t fam iliar with t he n ew fo rm and d o n o t a ffo rd

pro per c o nd it io n s as in n o t gi v ing s u itab le foo d t o an ima ls o r
,

in fa iling t o affo rd s u fli c ien t roo m to large gro wing v a riet ies -

o f p lant s
2
.

A ft er m ak ing d u e allowanc e h o wev er fo r a ll th es e c o n s id era

, ,

t io ns the fact rem ains that t he c o nd it io ns o f life ev id ent ly d o

,

ex ert a s tro ngly m o d ify ing i nfl u enc e u p o n d ev elo p m ent .

Loc alit y a comprehensiv e t er m T h ere is li tt le u s e in att em pt .

i n g t o d et ermi ne t he exact infl u enc e o f each s eparat e lo ca lity .

T he t erm is an exc eed ing ly c o m preh en s iv e o n e inc lu d ing m any ,

th ing s — c limat e by wh ich we m ean n o t o n ly t em perat u re

, , ,

mo is t u re and light b u t th eir c o m parat i v e pro p o rt io n s in th a t

, ,

part ic u lar s po t ; s easo n by wh ich we m ean t he s u cc es s io n of

,

c li mat ic c o nd it io n s ; foo d ( bo th as t o q u ant ity and q u a lity ) o n ,

wh ich t he creat u re ab so lu t ely d ep end s n o t o n ly fo r life b u t a lso ,

fo r gro wth hab it s o f life rad ic a l changes in wh ich m ay b e f o rc ed

,

u po n t he an ima l by i t s hab itat .

T h is is all t oo c o m p lex fo r pro fitab le s t u dy and d is c u ss io n .

W e m u s t separ a t e lo ca lity int o it s elem ent s and d et erm ine if we ,

1 D a rwin , A n ima l d Pl
s an a n ts , I , 39.

9 I t wi ll b d i th i
e n o te n s c o n n ec t io n t h at mo s t of t he in s ta n c es c i t e d a re t h o se

of d et eri o ra t io n .
226 C A U S ES O F V A R I ATI ON

and that u nd er d ev elo p m ent d u e to in s u flflc ien t food is q u it e

-

d is t inct fro m d wa r fi ng es pec ially a m o ng an im als in that t he

, ,

bo dy d o es n o t d ev elo p pro po rt io nat ely I n u nd erfed ca lv es fo r .

,

exa m p le t he h ead o u tgro ws t he res t o f t he bod y t he leg s a re

, ,

lo ng and t he j o int s a re la rge

,
.

I n gen eral f u ll f eed m ean s n o t o n ly increa s ed s ize b u t early

,

m at u rity a s well wh ich is o f ev en great er c o n s eq u enc e B eca u s e

, .

o f t he larg e pro po rt io n o f f o o d n ev er r ec o v er ed in ga i n it is ,

m an ifes t that any s h o rt en ing in t he p erio d o f d ev elo p m ent t e

s u lt s n o t o n ly in i m pro v ed q u a lity b u t a ls o in t he s a v i ng o f f eed ;

in o th er wo rd s early gain s a re ec o n o m ic a l gain s and th ey t end

,

t o h igh er q u a lity .

Effec t u pon fert ili t y T he a m o u nt and ch a ract er o f f oo d o ft en

.

ex ert pr o f o u nd phy s io lo gical i nfl u enc es F o r exa m p le t he fer

.
,

t ilit y o f the f em a le h o n eyb ee is m ain ly d u e to f oo d t he s t erile ,

wo rk ers and t he f ert ile q u een s d ev elo p ing fr o m t he s a m e eggs .

I f ev en aft er t he wo rk er egg s a re hatch ed and t he larv ae well

,

d ev elo ped th ey b e tak en fro m t he wo rk er c ells pu t i nt o q u een

, ,

c ells and fed q u een s f oo d th ey will d ev elo p int o q u een s

,
’

, ,

a fact o ft en tak en ad v antage o f by the bees th em s elv es wh en by

acc id ent all t he pro s pect iv e q u een s ha v e been lo s t Here fert ility .

is larg ely a m att er o f f oo d a lth o u gh an o cca s io n al wo rk er is

,

known t o prod u c e eggs T h is gen era l d ifferenc e bet ween wo rk er

.

and q u een m u s t th erefo re b e regard ed as o n e o f d ev elo p m ent

d epend ent m a in ly o n t he f oo d s u pply .

S peak ing gen erally exc es s iv e foo d s u pply lead s t o infert ility
,

a m o ng bo th p lant s and an i mals The fo r m er v egetat e lu xu ri .

o u sl b u t th ey d o n o t b lo s s o m and fru it s o ab u ndant ly a s u nd er

y ,

a fu ll b u t mod erat e su pply o f plant fo o d .

Whether t he eff ect in q u es t io n is d u e to o v erfeed ing o r t o o

s o m e o n e o r t wo elem ent s in part ic u lar is n o t well es tab lis h ed .

E n o u gh is kn o wn h o wev er t o j u s t ify t he a s s ert io n that extrem e

, ,

pro po rt io n s o f n itro gen prod u c e lu x u rianc e in s t em and leaf at

t he expen s e o f fl o wer and fru it b u t th ere is exc eed ing d o u bt
,

wh eth er th is effect wo u ld f o llo w a well ba lanc ed f oo d s u pply -

wit h plenty o f ph os ph o r u s .

E xc es s iv e feed ing o f an i mals es p ec ially fem ales t end s t o fatty

, ,

d egenera t io n o f t he es sent ial s ex u al o rgan s and c o n s eq u ent ly to ,

 EXT ER NA L I N FLU ENC ES AS C AU S ES OF VA R I ATI ON 2 2 7

t rility and th is res u lt is has t ened if t he foo d c o ntain s u n u s u al

s e ,

pro po rt io n s o f carbohydrat es es pec ia lly s u gar ,

.

Effec t of feed u pon v ar iab il ity I t is a c o m m o n belief t hat .

plant s and an im als a re m o re v ar iable when well fed than o th er

wi s e D o u bt les s th is is tr u e s o far at lea s t a s appearanc es go
.
, ,

fo r o n ly u nd er s u c h fav o ra b le c o nd it io n s a re all the fac u lt ies

that a re bo rn int o t he exc ept io n al i nd iv id u a l ab le to d ev elo p and
bec o me v is ible Wh en a rac e is liv ing u nd er med io cre c o nd it io n s
.

th ere is a d ead lev el in d ev elo p ment T he med iocre ind iv id u als .

hav e relat iv ely t he bes t chanc e and few will r is e abo v e t he c o n

,

d it io n o f med iocrity .

Wh eth er f u ll feed is a d i rect s t im u lant to v ariab ility o r o n ly

brings po t ent ia l d ifferenc es to t he s u rfac e is th erefo re an o pen
q ues t io n The proc ed u re ind icat ed is h owev er in eith er c as e the
.
, ,

s am e ; n a mely to pr o v id e m ax im u m c o nd it io n s if t he breed er
,

expect s t o rea lize the ut m o s t fro m his b es t i nd iv id u a ls o r h o p es

ev er t o find v ariati o n s wo rth pres erv ing .

Herein lies t he fact that well bred an im als o ft en req u ire m o re

-

feed than th eir s cru b relat iv es I t was u po n that po int that t hey
.

—
d epart ed from th eir k ind n o t that th ey c o ntra ct ed to ex is t on
les s feed b u t that th ey were ab le t o hand le m o re feed and pu t
,

i t to goo d u se If t he p u rpo s e o f t he breed er were to d ev elo p

.

ra c es with a m in im u m ma int enanc e rat io n it c o u ld b e d o ne ; b u t ,

we k eep d o m es t ic an ima ls n o t fo r th eir s o c iety b u t fo r what th ey

can d o —
,
fo r what th ey can m an u fact u re o u t o f c o rn o at s and , ,

hay We im pro v e cro ps n o t to see u po n ho w poo r land th ey m ay

.
,

liv e b u t rath er t o i ncrea s e th ei r a b ility to c o n s tru ct v a lu ab le f o o d

,

m at er ia ls fro m t he mi n era l elem ent s o f t h e s o il and t he in o rgan ic

c o n s t it u ent s o f the at mo s ph ere N o t m in im u m o f c o n s u m pt io n
.

b u t ec o no mic c o n s u m pt io n is th erefo re t he v irt u e s o u ght .

Ev il effec t s of ov erf eedi n g T he p lant will s eld o m s u ffer fro m

.

ab u ndanc e o f foo d alth o u gh it is n o t im po ssib le E xc es s of

,
.

n itro gen ca u ses rank growth o f s t em b u t ph o s ph o r u s is need ed ,

for s eed So meth in g ak in t o a ba lanc ed rat io n is d o u bt les s bes t

.

fo r p lant as well a s an im a l yet t he f o r m er has m o re o f s elect iv e

,

ab ility than the latt er .

T he an im a l is ea s ily o v erfed and if s o t he inj u ry is lik ely to

,

b e per m an ent I t res u lt s ( I ) in d is o rd er s o f t he d iges t iv e tra ct ;

.
2 28 C A U SES O F V A R I A T I O N

( )
2 in d is o rd e r s o f t h e e xcr e t o ry o rg a n s ;
( 3) in e xc ess i v e fat ;

(4 ) in s t e ri li ty e s p e
,
c ia ll y i n f em a le s thr o u gh fatty d e
,
g e n e rat io n
o f t he es s ent ia l s ex u a l o rgan s .

Q u ali t a t i v e e ff ec t s f r o m t h e n a t u r e o f t h e f ood T h e c o lo r .

o f p lant s and fl o wer s and ev en o f a n i m a ls is s a id o ft en t o b e

, ,

d irect ly infl u enc ed by th eir f oo d b u t it is exc eed ingly d ifli c u lt ,

a lway s to separat e fact fro m trad it io n in th is part ic u lar fi eld ,

beca u s e t he av erage pers o n has an exaggera t ed c o nc ept io n o f

t he infl u enc e o f fo o d u po n t he c o n s t it u t io n o f t he o rga n is m ,

o ft en believ i ng that a m eat d iet fo r exa m p le inc lin es t o f er o c i ty

, ,

and u nc o ntro llab le t em per generally D ar win ment io n s t he l

.

pra ct ic e o f feed i ng h em p seed to b u llfin c hes to darken th eir

c o lo r ; the fat o f a c ertain fis h to parro t s ca u s ing th em to ,

b ec o m e bea u t ifu lly v a riega t ed with red and yello w feathers .

”

He a ls o s tat es that t he s h ells o f m o llu s k s a re largely infl u enc ed

by t he a m o u nt o f lim e i n t h e wat er in wh ic h th ey gro w a n d
'

B ea ] s u cc eed ed in gro wi ng b lu e hydra ngeas fro m p ink s t o ck by

o cc as io na l app licat io n o f a lu m wat er t o the roo t s .

T hat t he fla v o r o f m ilk a n d egg s is largely d epend ent u po n

foo d has lo ng been kno wn a s ha s t he s pec ific effect o f c ertain
,

foo d s u po n the t ext u re a n d fla v o r o f m eat M u m fo rd fo u nd at .

Illin o is that t he fat o f p o rk fed u po n large a m o u nt s o f c o tt o n

s eed pr o v ed o n ch em ica l exa m inat io n
,
t o c o nta i n a pro po rt io n
,

o f tr u e c o tt o n s eed o il s h o wing th a t a p o rt io n at lea s t o f t he

-
,

fo o d had been carr ied o v er and s t o red u nchanged in t he t is s u es .

T hat th is is t h e exc ept io n o r p erhaps m o re c o rrect ly t he ,

les s er res u lt is pro v ed by t he fact that in gen era l t he id ent ity o f

,

t he fo o d is lo s t in t he nat u re o f t he o rgan is m s h o wing that t he ,

o rgan is m and no t i t s f oo d d o m inat es res u l t s T he s a me m eat .

bec o mes d og o r man ind ifferent ly ind icat ing that fo o d c o m ,

po u nd s a re n o t as a r u le carr ied o v er b u t rath er s u ffer pro fo u nd

d is ru pt io n being red u c ed if n o t t o th eir elemen ts a t lea s t to c o m
,
”

, ,

parat iv ely s im p le c o m po u nd s t he en ergy s et free in the d is ru p

,

t io n being u t ilized in t he s u b s eq u ent wo rk o f c o n s tru ct io n 2

.

T he h igh er o rgan is m s a re c o m para tiv ely i nd epend ent o f th eir

foo d s o far as q u alitat iv e changes a re c o nc ern ed T h ey s eem .

1D wi
ar A im l
n, n d Pl
a s ant II (
an s , d d iti ) 69 7 s ec o n e on , 2 -
2 0 .

2
E y l p d i B it
nc c o e
z a i r XIX
an n c a , ti l Phy i l gy ar c e s o o , 2 1.
2 3 0 C A U SES OF V A R I AT I O N

freed th em selv es fro m t he bo ndage d u e to t he acc id ent o f b irth

plac e being ab le to m ov e a bo u t and th erefo re to es tab lis h in t he
,

wid es t s en se an ind epend ent ex is t enc e .

N o t with s tand ing all th is and aft er m ak ing allo wanc e fo r t he ,

gro s s er infl u enc es o v er lo wer o rga n is m s t he fact yet rem ain s ,

that t o a s light ext ent and t o a s light ext ent o n ly t he an imal

, , ,

is infl u enc ed by t h e charact er o f his food T hat th is infl u enc e is .

larger u po n t he prod u ct s o f t he bo dy th a n u po n the bod y it self

is c ert a in and u po n t he s u bt ler q ua lit ies lik e flav o r rath er th a n , ,

t he m o re es s ent ia lly b io lo gic a l charact er s s u ch a s s tr u ct u re .

SECT I O N I II — T HE E FFECT O F M O I ST U R E U PO N
D EV E LO PM E NT

A n im a ls a r u le ar e q u it e ind epend ent of mo is t u re pr ov id ing

as ,

th eir d irect n eed s fo r wat er a re s at is fi ed in t he way o f bo dy

c o n s u m pt io n in add it io n to that n eed ed t o red u c e the effect s o f
int erna l h eat by ev apo rat io n ‘
.

Plant s on t he o th er h a nd d o n ot ha v e t he c irc u lat ory sy s t em of

an im als and th ey d epend u po n wat er tak en in by t he roo t s a nd
, ,

ev apo rat ed by t he lea v es t o act ua lly carry f oo d t o a ll pa rt s o f t he ,

s tr u ct u r e T h eir n eed fo r wat er is th er efo r e far abo v e t he am o u nt s

.

n ec es sary fo r act u al c om po s it io n Fo r exam ple it r eq u ir es the .

,

ev a po rat io n o f s o m eth ing lik e t he eq u iv a lent o f eight inch es o f

s tand ing wat er o v er t he ent ir e fi eld to m at u r e an a v era ge c o rn cr o p .

1
It i ft s l y t gh t th t im l
o en erro n eo u s m b
au f d ta an a s co n su e car o n ac eo u s oo s o

t i t h b dy t m p
s us a n e t wh il th t t h i t h t ll f d
o e era u re, t ll y til i d i
e e ru s a a oo ac u a u ze s

b k ro d w d it
en o
g y t f
n an T h i g y
s en er i d i p sed f iree t h . w y s en er s s o se o n ree a s

fi t t
rs , l igh t t t i ff t i g t h
o a s ex en mb i t i n e f th ec l m t f th f d
n e rec o na on o e e e en s o e oo

i t
n o th d i gl y mp l p t p l m f t h b dy it p d t ;
e exc ee n co ex ro o asd by o e o or s ro uc s s ec o n ,

t h b dy
e m
o f i t p t i th f m f i t
o r so e o s l f ar s t n l w k ; thi d
e or o n ern a or o ex ern a or o r, r ,

it i s ra di t d i t h f m f h t f l w i t it y I th i w y t h b dy i f t y
a e n e or o ea o o n en s . n s a e o s a ac o r

th t i
a t tly p d i g h t wh i h m t b d i p d f th t t will b
s c o n s an ro uc n ea c us e s o se o or e s ru c u re e

f d i g t h i —b y
,

d t y d Th
es ro e b t tw w y
. ere a re u di tio a d by
s o p o n s, ra a o n an ev a o ra

ti on T h b dy i t h
. e t
o tly p d ig di
s us c o n s an t tl y l i g h t It
ro uc n an s as c o n s a n os n ea . s

t
ac u a l t m p t i t i t b m t h i g b th t f t h
e e ra u re s c er a n o e so di g m di me n a ove a o e s u rro u n n
’

e u
—h w m h will d p d p t h l ti b t w
,

o uc e en th u p
on id it y fe re a
p d t i on e een e ra o ro uc on

an d h f il i t it f
e di ti
ac d esp t
ori raT h b ady t
o n an m p t ev ai t h o ra on. e o e era u re s us a

k i d f l g b i m d it d p d p
n o a e ra c su g , tan i t y f e d i
ent i sS m u llon a rea v ar e o c on o ns . a

an im l d i t p id l y l g
a s ra a e l ra p id l y f , d i
ar t i i
e o n es f ess ra t i , or ra a on s a s u r ac e ac on,

an d th i f i l i p p ti
e r s u r ac e t th b l k
s es s H gn d i t l wl y b
ro or on o e u . o s ra a e s o , eca u s e

o f t h i b l k t f f t Th y d
e r an e t w t
o a d th
. e il y o no h td s ea , an u s a re eas o v er ea e .
 EXT ER NA L I N FLU E N C ES A S C AU SES OF V A R IATI ON 23 1

Wat er or t he lack of it is th erefo re in m any cou ntries an d in

, ,

m an y s ea s o n s t he li m i t i ng elem en t ; that is t o say t he y ield is ,

limit ed n o t by t he a v a ilab le f ert ility o r t he ab ility o f t he cro p b u t

by t he mois t u re present .

I n exc es s iv ely wet s ea so n s cro ps a re n o t o r io u s ly s o ft that is , ,

lacki ng in s u b s tanc e J u s t what t he d iff erenc e is has n o t been

.

well es tablis h ed b u t s ize has been attain ed at t he expen s e of

,

q u ality T h ere is good reaso n t o a ssu me that it is the res u lt of

.

abu ndance of wat er lead ing t o fu ll c ellu lar d ev elo pment b u t

, ,

d efic iency of ev apo ra t io n and tran s fusio n o f food d u e to C lou dy

s k ies res u lt ing in a lack o f act u a l dry m att er
,
.

Efi ect of m oi st u re i n t he a t mospher e I t is sa id that m o is t .

at mos ph eres pr od u c e fin en ess of ha ir or fu r in an imals and d eli

cat e foliage in plant s and that a dry at mos ph ere inc lin es to a
,

har s h dry coat and to s piny gr owth in plant s U nd er nat u ral

, .

c ond it io n s ho wev er mois t u r e is o ft en assoc iat ed with c oolnes s and

, ,

s h ade and dryn es s with gr eat h eat and int en s e light

, Certain it .

is that fur bear ing an im a ls are f o u nd in c oo l c lim at es and that

-

v egetat io n is d elicat e in t he t em pera t e r egio n b u t har s h dry , ,

and spiny in t he arid sect io n s T hese fact s are well kn own and .

u n iv er sally rec o gn ized b u t ho w m u ch is d u e t o m o is t u re a lo n e

,

cann o t well b e d et erm in ed in nat u r e .

R eso rt ing to d ir ect exper im ent h owev er we find that the sa m e , ,

pla nt m ay b e grown with or with ou t s pin es acco rd ing t o the

d egree o f mois t u re in the s u rr ou nd ing at m o sph ere S pin es are .

u nd ev elo ped lea v es as th o rn s ar e a b o rt iv e s t em s

,
and anyth ing ,

that ch eck s gr owth t end s to th eir prod u ct io n That th is is m ain ly .

t he r es u lt of a dry at m o s ph ere h o wev er is eas ily s h own in t he

, ,

labo rat o ry .

Lot helier has m ad e n u merou s o b ser v at io n s in wh ich ind iv id

u als of t he s am e s p ec ies wer e plac ed s id e by s id e s o m e expo sed ,

freely to t he air and oth er s k ept m o is t u nd er a glass s had e .

U nd er c o nd it io n s s u ch as th is “
B er ber zs v u lg a n s bo re n o n
' '

s pin es c ent leav es in a m o is t at m o s ph er e b u t s p in es a lo n e in a ,

perfect ly d ry o n e A gain t he s h oo t s wh ich in Lyr zu m ba r ba m m

'

.
, ,

Ulex E u r opwu s etc ,

wo u ld n o r m ally h a v e for m ed th o rn s by
.
, ,

arr es t ed d ev elopm ent and s clero s is ( hard ening) in a v ery dam p ,

at m o s ph ere c o nt in u ed to gr ow and elo n ga t ed int o leafy b ranch es

,
.
2 3 2 C A U S ES OF V A R I ATI O N

M icro s c o p ica l ex a m inat io n s h o wed that in t he dry a ir s pec im en s -

“
t he pa lisa d e c ells wer e well d ev elo ped and th er e wa s a s pec ia l
co n so lidat io n o f fibro u s t is s u es 1
T he sa m e a u th o r c o nt in u es
”
.

Ag i tha n, mm w t e co d P / g m it mm i wh e g wi g i
on a er ree zr a es co un s, n ro n n

th i ig t d
e u n rr f th N i l
a e ll y f m t
a rea s o t d g wth wi t h y
e e va e , or s a s un e ro v er

s h t d h pp i t d l
or an s ar Cl t th N il
-
h w
o n e it g w ea v es . o se o e e, o ev e r , ro s

i
n ne o r t f t h ig h with l g l
en ee lm t tly li k e the p l
, t i on ea v es a os ex a c an s n

E g li h i
n s r v e rs .

A ll o b s er v at io n s go t o s h o w that t he n u m ber o f v es s els in the

fi b ro v a sc u lar s y s t em is grea t er in t he aer ia l than in t he aq u at ic
-

fo r m s o f t he sam e s pec ies and the ev id en c e in gen era l s eem s

2
,

c o nc lu s iv e that t he n o t o r iou s ab u ndance o f s p in es in tro pical

v egetat io n is d u e pr im ar ily t o a dry at m o s ph er e a s s is t ed t o ,

s o m e ext ent n o d o u bt by t he r etard ing effect o f int en se light

, ,

U po n g r o wth .

A s ig n ificant fact in th is c o n n ect io n po s s ibly attrib u table t o ,

t he s c a rc ity o f wat er po ss ib ly t o t he lack o f h ea t is t he well kn o wn

, ,
-

ph en om en o n that plant lic e pro du c ing fem a les o n ly d u r ing the ,

s u m m er b egin with appro ach ing a u t u m n t o pro d u c e m a les and

, ,

that u nd er t he perpet u al h eat of t he gr eenh o u s e t he in sect s o h

s er v e s u m m er hab it s ind efin it ely u n les s t/ze la n ts on ru /t ie/t t/t ey
p
d
a r e f ee i ngr
a r e a llo wed to b ecom e d ry .

A s is well kn o wn k ept fo r lo n g per iod s if ,

s eed s m a
y b e

th o ro u gh ly dr ied I n th is c a s e t he v ital act iv it ies are red u ced to

.

a m in im u m b u t pro ba bly n o t ent irely s u s pend ed b eca u s e s eed s

, ,

will n o t la s t ind efin it ely C erta in lo wer fo rm s o f plant a n d a n i .

m al life ha v e a m ark ed po wer o f a ppa rent ly s u s pend in g life

thr o u gh d es icc at io n a n d r es u m ing it s act iv it ies aga in with s u fli
c ient m o is t u re 3
.

T he act u al infl u enc e o f wa t er u po n d ev elopm ent is n o t yet

well u nd er s t ood exc ept that it is o n e o f the a b s olu t e c o nd it io n s
,

o f life and b eing a fl u ct u at ing elem ent o ft en lim it s d ev elo pm ent

, , ,
.

T he s t u d ent s h o u ld fu lly apprec iat e t he bea ring o f all th is u po n

t he m att er in hand Tb e d eg r ee 4 d ev elop m en t qf a n in d i v id u a l
at m a tu ri ty i s n ot a eom
p lete i n d ex to lzis i n /t er ited d i a m eters
‘
.

1 V em o n , V a ria t io n in A n im a p H en s lo w , O rigi n l s an d P l a n ts , . 2 64 ; s ee a s o l
l
o f P a n t S t ru c t u res p 4 0 1 V e rn o n V a ria t io n in A n im a ls an d P a n ts , p 2 6
5 l
C B D a v en po rt E xperi m en ta M o rph o o gy Pa rt I pp 59—
. .
, , . .

3 . .
, 65 l l , , . .
 C A U S ES O F V A R I AT I ON
2
34

S pec ific rat e o f v ibra t io n so that wh en th is is reprod u c ed by a ,

v ibrat ing c o rd o r p lat e exp lo s io n o f t h e s u b s tanc e m ay o cc u r

,
.

I od id o f n itro gen is o n e o f th ese s u bs tanc es wh ich is explod ed

by a h igh n o t e 1
L iv i ng pro t o p la s m is n o exc ept io n t o t he
.
”

gen eral r u le that s pec ially u n stable c o m po u nd s a re sen s it iv e to

co nt a ct .

Effec t of cont ac t u pon t h e met ab oli sm of prot opl asm 2

I t is a .

well kn o wn fact that ph o s ph o resc enc e is increa s ed by m ech a n ic al

-

irr itat io n S o tr u e is th is that t he wat er thro wn fr o m the pro

.

peller wh eels of a s t eamer in tro pical regio n s lo o k s lik e liq u id

fire and a br is k breeze m o v ing o v er t he s u rfac e su gges t s at a
,

d is tanc e t he wh it e foa m o f the s u rf .

C o nta ct also ex ert s a s tr o ng ly st im u lat ing infl u enc e u po n s ecre

t io n s n ot o nly with lo wer o rga n is m s wh ich s eek attach m ent s and
, ,

t he gla n d s o f in sect iv o ro u s plant s b u t with h igh er a n im a ls as well 3

, .

Efi ec t of cont act u pon mov emen t T he fir s t eff ect o f m echan ical .

d is t u rbanc e in prot o plas m is t o ch eck all m ov em ent M in u t e or .

n is m s tra d es c ant ia ha ir s etc c ea s e th eir pr o t o plas m ic m o t io n

g a , ,
.
,

by the irritat io n o f m o u nt ing u nd er t he c o v er glass I n h igh er

. .

plant s a s u dd en jar ca u ses c es sat io n o f m ov em ent and o ft en a

retreat o f t he pro t o plas m to o n e S id e o f t he c ell so charact eris t ic
as to b e s po k en o f as fr ight
“
.

I f an a m m b a with p seu d o po dia o u t is t o u ch ed o r irr itat ed it ,

im m ed iat ely a s s u m es t he s ph erical fo rm and in gen era l t h e effect ,

o f C o ntact is t o ca u s e pr o t o p la s m t o c ea s e m o t io n and a s s u m e

appro x im at ely the s ph erical fo r m o r in o th er wo rd s to oc c u py , , ,

t he lea s t S pac e po s s ib le B u t t/t is is to a ll in ten ts a n d p u rpos es

.

con tr ac t ion and t he gen eral pr inc ip le m ay b e laid d o wn th a t ex

,

t ernal c o ntact ca u s es c o ntract io n es pec ially n o t iceab le in m u s cle , ,

wh ich is pa r ex cellen c e the c on tr a c t i le ti ss u e .

T h is c o ntra ct io n m os t c o m m o n ly a n d o f n ec es s ity Opera t es , ,

at fi rs t to d ra w the o rga n is m a way fro m t he irritat ing bo dy

( n e gat iv e t hig m o tax is ) b u t if the b o dy b e lo ng o r la rge s o that
, ,

lo c o m o t io n c o nt in u es th en the s id e n ext t he f o reign b o dy will

,

1 Th is s u gges t s , as t h e a u t h o r o b s erv es ,
p h
t he en o m en a o f h ea ri n g .

3 C B D a v en p o r t , E xperi m en t a M o r
. . l p h l gyo oP , a rtI pp 98 99
, .
-
.

3 l k
A s is we l n o wn , th e eif er t a t has h h n ev er
p ro d d
uc e a c a f ma
y b l
e ma d e to

giv e mi lk m ere ly b y p ers is t en t m an ip u l a t io n o f t he u dd er .

 EXT E R N A L I N FLU E N C ES A S C A U SES OF V A R I AT I O N 2 35

be Sh o rten ed and the o rgan is m will m o v e in a c u rv e that will

s p eed ily br ing i t i nt o act u a l c o nt a ct I t is n o t ic eab le too that .
, ,

real c o ntact being o nc e es tab lish ed is b ro ken with d iffic u lty

'

, , .

M any lo wer an im als a s in aq u aria c o m ing in c o ntact with a p lain

, ,

s u r fac e m o v e alo ng that s u rfac e u nt il th ey reach a p o int wh ere

,

s id e and b o tt o m o r wh ere t wo s id es j o in and wh ere th ey can ,

plac e th eir bod ies in c o ntact wi th t wo s u rfac es T h ey a re lik ely .

n o w t o m o v e a lo ng the groo v e f o r m ed by t he t wo s u rfac es u nt il

a c o rner is reach ed wh ere c o ntact o n three s id es is po s s ib le .

Here if anywh ere t he o rgan is m will c o m e to res t I t is o n ly

, , .

that it is mo re c om f o rt ab le
“
that it m ov ed u nd er the molar
im p u lse u nt il it reach ed a p o int wh ere f u rth er m o v em ent and
mo re c o m p let e c o ntact were a lik e im po ss ib le Ev en h igh er .

animals c o me to the h igh es t s tat e o f res t wh en in c o ntact with

fo reign bo d ies o n as m a n y S id es as po s s ible .

Efi ect of c on t act u pon directi on of mov emen t t higmotax is , ,

or s t er eot ropis m
1
I t is a well kn o wn fact that roo t s gro wing in
.
-

r u nn ing wat er grow ups trea m n o t d o wn s tream and that m any

, ,

fis h at t he breed ing s ea s o n a re po s s es s ed o f an i rres is t ib le im

p u ls e to m o v e aga in s t the c u rrent ( rh eo tax is ) Th ey th erefo re
2
.
'

as c end t he s tro nges t c u rrent s lea p waterfa lls and s u rm o u nt

, ,

ev ery p o s s ib le o b s t a c le in u p s trea m m o v em ent s — a pas s io n ,

wh ich u lt im at ely carries th em t o th eir breed ing gro u nd s i n s ha l

lo w wat er I t is a t th ese t im es that s alm o n p ile th em selv es u p
.

ev en a b o v e t h e wat er lev el and that th ey will f o llo w any d ec o y

that lead s aga in s t t he c u rrent ev en int o h o peles s trap s,

.

T h u s m ay ext er nal agent s ex ert a s tro ngly m o d ify ing in flu

enc e u p o n s u ch es s ent ia l act iv it ies o f liv ing m att er as t he
c o ntra ct ility o f pro t o p las m res u lt ing in d efi n it ely d irect ed mo v e
,

ment s thro u gh th ei r c o ntro l o f m u s c u lar c o ntra ct io n A s we .

s ha ll s ee c o nta ct is n o t t he o n ly infl u enc e capab le o f s t im u la t

,

in g c o ntract ility o f pro t o p las m and c o ntro lling t he d irect io n o f

mo v em ent ; o n t he o th er hand m u s c u lar t iss u e is expo s ed to ,

t he exc i t ing infl u enc e o f a great v ariety o f c irc u m s tanc es b o th

interna l and ext erna l t o t he o rgan is m any o n e o f wh ich will ,

i nd u c e t he charact eris t ic react io n o f th is so rt o f t is su e wh ich is ,

c o ntract io n .

1 C B D. .
p t E
a v en
p i m t
orl M,p hx l gy Per t I
en a
p 5 or I b id p 09
o o , ar , . 10 .
1 . . 1 .
2 3 6 C A U S ES O F V A R I AT IO N

SECT IO N V — E FFECT O F G R AV I T Y U PO N L I V I NG
G EOTR O PI S M 2

A ger m inat in g s eed s en d s o u t t wo s pro u t s F ro m what ev er .

po s it io n th ey em erge o n e gro ws d o wn ward in res po n s e to grav

,

ity the o th er u p ward in o ppo s it io n

, I n o th er wo rd s t he ro o t is .
,

po s it iv ely and the s t em is n egat iv ely geo tro p ic ; that is t o s ay ,

each c o nt a in s with in it s elf s o m e q u a lit y th a t

p u t s it int o d efi n it e rela t io n with t he c ent er o f

t he earth b u t in o pp o s it e d i rect io n s ,
.

T hat th is t end ency is so m eth ing c o n s id er

a b le is s h o wn by t he fact that it is c a pab le o f

ex ert io n ag a in s t pr o n o u nc ed r es i s t a nc e a s in ,

b u rro wing thro u gh the s o il o r pers is t ing aga in s t

m ech a n ical o b s tru ct io n s .

T h is d efin it e relat io n t o gra v ity s eem s to ex

er t it s elf i n t he m ann er o f in wa rd f o rc es res p o nd

in g to o u t s id e c o nd it io n s ; fo r if a p iec e o f
s t em b e a lt ered i n it s p o s it io n f u t u re gr o wth ,

rea dj u s t s it s elf as pro m pt ly as po s s ible in

1 C BD a v en po rt, E x p erim en ta M o rp o o g
. l h l y Pa rt I ,
—
.
,

1 1 2— l
1 2 4 ; a s o Pa rt I I , p p 3 9 1 4 0 2 , f ro m w ic h h mo s t of
pp . .

t he fac ts h ere c it e d a re ta k en .

1 T wo s e ri es of t erm s a re in u s e, o f s u b s t a n t ia ll y t he
“
sa m e mea n in g : one ( s ee
“
geo t ro p is m and
g eo t ro ic
p
wit h en di n gs d eri v e df ro m t h e G ree k m ea n in g to tu r n th e

o t h e r s ee (
“
geo t a x is an d
“
geo t a c t ic G ree k en di n
g s

s ign if yi ng to a r r a n g e. W e th u s h a v e a so l “
c h em o t ro p is m ,
”

wit hdj t i h m t
it s a ec ve
“
c e o ro p ic ,
”
o v er a ga in s t
“
c h e mo
t i
ax s , h m t ti
"
h li
“
c e o ac c,
”
e o t ro
p ism,
"
and
“
h li
e o t a c t ic ,
”

t
e c T h l tt
. di g w e a er en n s ere s u pp li de wh en t he effec t of

c h m i l g ity d l igh t
e c a s, ra v , an u po n f r ee-m o wingr o rg a n is m s ,

F ig . 25 . In fl u en c e o f as b a c teria , s wa s fi rs t n o t e w ic eff ec ts
wa rm s p o res , et c .
, d h h ,

g e o t ro
p i s m : be led t o efi n it e d
a rra n gem en t s b u t wh en s im i a r effec ts l
h a v io r o f a
g ro w d l
were n o te u po n a rger o rga n is m s n o t free to m o v e li e , k
in g p u m u le in l d l
t h e fi xe p an ts , b u t w ic h h
m a n ifes t e t he effec t b y tu r n i ng , d
h
ri g t in g its elf af t er t h e term “
t ro p is m
”
c ame in t o u s e, s ign if yi ng a t u rn in g .

b ein g p a c e in a l d h
T is term an d it sd e ri v a t iv es s eem t o exp res s m o re ac c u

h o r i zo n t a
p o si l ra t el
y w h at rea ll y h a pp en s in m o s t c a s es , an d t h y e a re

t io n as
, at 1 .
c o m in g to b e p refer re d in a ll gen era li d ze c on s i d era t io n s .

A ft er C B D a v en . . Hen c e in t h e tex t t h e t erm g eo t ro p is m is p referr e to d

p o rt , fro m S t ras g eo t a x is a n d s im i ar l ly i n mo s t p l ac es fo r all c o rr es po n d
b u rger in g t erms .
 8 CA U SES OF V A R I ATI ON
2 3

A t
s em of willo w s ev ered
fro m it s pa rent plant and s uspend ed ,

apex u p ward in a m o is t at mo s ph ere will o f c o u rs e s en d o u t

, ,

s h oo t s fro m i t s a p ex and ro o t s fro m it s ba s e I f a r ing o f bark .

b e rem o v ed fro m t he midd le o f t he s t em th en s pro u t s will is s u e ,

fro m t he ap ic al extrem it ies of t he s ect ion s and roo t s fro m the

ba s al en d N eith er o f th ese experiment s d et er m in es wh eth er
.

grav ity o r po lar ity is ch iefly in s tru m ental in the pro d u ct io n o f

s t em and ro o t b u t if t he p iec e b e in v ert ed and s u s p end ed ape x
,

d o wn ward (in a m o is t at mo s ph ere) we s ha ll get s o m e light o n ,

the t wo fo rc es ext erna l and i nt erna l

,
.

U nd er th es e c o nd it io n s t he ap ical en d n o w d o wn ward will , ,

y e t pro d u c e s tem s b u t th ey will change th ei r d i rect io n wi th ref

,

erenc e t o t he ax is and p o int up wa r d wh ile t he ba s a l en d will ,

pro d u c e roo t s b u t th ey will ext end d o wnward I n th is ca se

,
.

ea ch en d ha s pro d u c ed i t s charact eri s t ic gro wth and ea ch has ,

res po nd ed to gra v ity in the u s u a l way ( s ee Fig exc ept that .

,

if t he p iec e b e o f t he o ld er woo d ro o t s will appear thro u gh o u t

,

t he ent ire length T h e f o rc e that fix es t he cha ract er o f gro wth

.

appears to b e int erna l ; that wh ich fix es it s d irect io n appears to

b e m ain ly ext ern a l .

I f a bego n ia leaf b e p lant ed in t he gro u nd o r s u s pend ed in

m o is t air what ev er it s po s it io n ro o t s will s tart fro m t he ba s a l
,

en d o f t he s t em a t i ts p o i n t of s ev er a nc e and aft er ward s h oo t s ,

will aris e j u s t ab o v e t he po int o f o rigi n o f t he ro o t s the bo dy o f ,

t he leaf with e ring a way ab ant b t n

l “
B
( y o ve is m
. e e wee t he

o r igi n o f t h e ro o t a n d t he ap ex o f t he leaf what ev er i t s po s it io n ) ,

.

B y th is we s ee th a t t he s t em has a d is t inct p o lari ty prod u c ing ,

S pro u t and ro o t each at t he pr o p er en d ; th a t t he leaf ha s n o

,

tr u e po lar ity pro d u c ing pri ma rily o n ly roo t s b u t that wh erev er

,

and h o wev er pro d u c ed a d is t inct geo tro p is m charact erizes bo th

,

s t em and r o o t .

T h ere is litt le geo tro p is m in leav es o r in t he h o r izo ntal s t em s,

o f m any p lant s r u nn ing a lo ng o r j u s t b elo w t he s u rfac e o f t he

gro u nd Ho wev er the s t em s and roo t s pro d u c ed at t he n o d es

.
,

o f th es e u nd ergr o u nd s t em s a re b o th geo tro p ic .

Geot ropis m i n an i m al s G eo tro p is m is m u ch les s m ark ed with

.

an im als than with p lant s W e m ay s ay that o n ly fi x ed o rgan is ms

.

1 Mo rgan R egen era tio n pp 7 4 7 6

, , .
-
.
 EXT ER N A L I N FLU EN C ES A S CA U SES OF V A R I ATI ON 2 39

wo u ld b e lik ely to d ev elo p d ec id ed geo tro p is m ; or , c o n v ersely ,

we may s ay that o rgan is ms wi th m ark ed geo tro p is m wo u ld b e
lik ely to bec o me fi xed I n eith er ev ent les s geo tro pis m wo u ld b e
.

expect ed a m o ng ani m als and eith er as s u m pt io n wo u ld s q u are

,

with t he fact s M any lo wer o rgan is m s t ru ly an im al a re d is t i nct ly

.

geo tro pic h o wev er and m os t an imals s how a d ec id ed preferenc e

, ,

as to p o s it io n with r ef erenc e to gra v ity Bo th with lan d ani m als .

,

h igh o r low and wi th fis h as well the v entra l s id e is carried

, ,

d own ward and the ant erio r po rt io n in gen era l u p ward

,
.

Effect of geotr opism u pon prot oplasm T he pro t o p las m o f c ells .

,

plant o r an im al is n o t h o m ogeneo u s T he n u c leu s is h eav ier

,
.

than t he cyt o p las m and t ogeth er with ch lo ro phyll g ran u les an d

,

s tarch gra in s tend s t o s ett le t o t he lo wer s id e o f t he c ell gi v ing ,

it a k ind of po lari ty d u e t o gra v ity “

I n m any o v a t he yolk .

s ink s t o t he lo wer p o le and t he cyt o p la s m fl o at s o n t o p in ,

what ev er pos it io n t he egg m ay b e h eld — a fact wh ich

”

u nd o u bt ed ly has an im p o rtant eff ect u p o n d ev elo p m ent

1 ”
.

Gen eral effect of gr av i t y u pon d ev el opmen t T h ere is n o roo m .

fo r d o u bt as t o the pro fo u nd eff ect o f gra v ity u p o n d ev elo p m ent .

Ho wev er th is infl u enc e o f gra v ity has been c o nt in u al and c o n

,

s tant o n all exi s t i ng s p ec ies fo r u nt o ld g en era t io n s and it m ay ,

b e loo k ed u po n a s ha v i ng already ex ert ed the m ax i m u m o f its

infl u enc e u p o n all f o r ms o f life .

The efi ec t o f gra v ity u po n d ev elo p ment ha s th eref o re lo ng , ,

ago reach ed t h e po s it io n o f a c o n s tant f o rc e t o b e r eck o n ed with ,

an d is n o w t o b e regard ed a s a fi x ed fact o r i n d ev elo p m ent ra th er

than as a pres ent cau s e o f ind iv id u al d ev iat io n t o b e s t u d ied ,

m o re fo r t he s ak e o f learn ing t he d egree o f d epend enc e o f liv

in g m att er u po n o u t s id e f o rc es rath er than as a d i rec t m ean s o f
fu rth er change .

SEC TI ON V I — EFFECT O F L I G HT U PO N LI V I N G M ATTER

In ch lo ro phyllac eo u s p lant s t he a mo u nt o f carb o n fi x ed

a ll ,

and therefo re the t o ta l o f gro wth in the sen s e o f increa se in dry

m att er is in a lmos t d irect pro p o rt io n to t he expan s e o f leaf s u r
,

fac e and t he a mou nt of light that f alls u p o n it .

1
C B D p t
. E p .im t l M
a v enp h l gy
or P t
,
I p
x 4
er en a or o o , ar , . 1 1 .
2 40 C A U SES O F V A R I AT I ON

L ight has th er in flu enc es h o wev er than th o se exert ed

o , ,

thro u gh t he fixat io n o f carbo n Fo r exa m p le s tro ng s u n light .

,

t end s t o ch ec k gro wth in t he s en s e o f increas e in b u lk a nd wh en ,

th es e t wo effect s o f light a re c o m b in ed a s th ey a re in the ,

tro p ic s they giv e u s nat u ra lly t he s lo w gro wing genera lly s mall
,
-
, ,

and extremely d en se woo d o f t he lo wer lat it u d es .

B r iefly light lik e gra v ity ex ert s s p ec ific effect u po n m att er

, , , .

M any o f t he effect s o f gra v ity ( p o s it iv e geo tro p is m ) m ay b e

regard ed as aris ing fro m t he elementary pro pert ies o f m att er ,

fo r nat u ra lly a ll m att er is attract ed by and appro ach es a s n ear ly ,

a s po s s ib le to t he s u rfac e o f the earth ; th a t is m a t t er in g en era l

, , ,

may b e s aid t o b e p os it iv ely geo tro p ic S en s it iv en es s t o light .

,

h o wev er s h o u ld b e regard ed as d u e to t he s pec ial c omp ou nd s

,

that c o n s t it u t e liv i ng m att er rath er than as a pro perty of ma t ,

t er in gen era l fo r m att er in gen e ra l is ind i fferent t o light

, .

L ight ex ert s i nfl u enc e u po n liv ing m att er es pec ia lly p lant s in , ,

three d is t inct ly d ifferent ways : ( I ) thro u gh it s h eat ray s a ffect ,

in g t em perat u res ; ( 2 ) thro u gh t he s o ca lled ch em ica l ( act in ic ) -

ray s ca u s ing d efin it e ch em ical react ion s in t he pro t o p la s m ;

,

3) thr o u gh t h e l u m in o u s ra y s i nfl u enc ing es pec ia lly t he d i r ec tion

,

o f gr o wth in th o s e part s that a re s o fix ed a s t o b e incapab le o f free

mo v ement C ertai n o f th es e i nfl u enc es a re wo rthy o f s o m ewhat

.

ext end ed c o n s id er a t io n .

Ch emical effect s of l ight 1

Wh ile m att er in general in its .

s im p ler c o m p o u nd s is q u it e i nd i ff erent t o light et c e rta i n c o m

y ,

po u nd s a re n o t o rio u s ly d epend ent u po n i ts infl u enc e ; that is

to say m any c o m b inat io n s a re effect ed m o re r ead ily and o th er s
,

on ly in t he pres enc e o f light ( ph o t o s ynth es is

) .

Ox idat io n o f v egetab le o ils is m u ch m o re ra p id in day light

than in darkn es s Hydro gen and ch lo rin u n it e exp lo s iv ely i n
.

Me p res enc e of lig /t t C h lo rin pa s sed thro u gh alc o h o l in s tro ng

.

s u n light u n i t es with it fo rm ing ch lo ral hydra t e and ch lo rin

, ,

c o m po u nd s generally are s en s it iv e t o light .

T he wh o le fi eld o f ph o t o g ra phy is d epend ent u po n t he act io n

o f light u p o n t he ha lo gen s a lt s o f s ilv er go ld p lat in u m and , , ,

o th er m eta ls d u e to the s o c alled ch em ica l ra ys ext end ing

“ -
”

1 C B D. .
p t E
a v en
p im t l M
or
p
, h l gxy P t I
er pp 6 — 6
en a
5 f m w h i h
or o o , ar , . 1 1 1 , ro c

m t f th i
os o e d t h i h di g t k
n s ta n c es u n er s ea n a re a en .
242 CA U S ES O F V A R I AT I O N

alc o ho l d ec o lo rize and th en with io d in wh ich fo r m s it s char

to , ,

ac ter is t ic b lu e with s t a rch T he s pectr u m bet ween E and D 1

.

i nc lu d es t h e u pp er p a rt o f t he red t he o range and the lo wer , ,

part s o f t he y ello w t he th e rm ic rath er than t he act in ic

,

po rt io n o f t he s pectr u m .

A m o ng b o th p lant s and an imals light ha s an im po rtant in flu

enc e u p o n c o lo r T he ch lo ro phy ll o f p lant s is fo rm ed o n ly in
.

it s pres enc e a n d it is int i m a t ely c o nc ern ed i n t he pro d u ct io n of

,

p ig m ent s in t he s k in N o t o n ly that b u t t he arrangem ent and

.
,

po s it io n o f p ig m entary m a tt er wh eth er lying next the su rfac e ,

i ff —
and ell d u s ed th u s giv ng c o o r to t he an i al o r ly in g
w i l m —
c o llect ed in m a s s es d eeper in t he s k in a n d ha v ing litt le effect
u po n t he c o lo r a re d u e la rgely t o t he d irect effect o f light
,

falling u po n t he s k in o f t he an im a l I n th is way c erta in an imals .

,

a s t he cha m eleo n a re capab le o f exhib it in g a c o n s id erab le range

,

o f c o lo r s g iv i n g r is e t o t he fict io n that th ey a re ab le t o i m itat e

,

any c o lo r n ea r wh ich th ey may b e s it u at ed 2

.

I t has b een c u s t o ma ry t o C it e t he fact that c a v e an im a ls a re

freq u ent ly les s h igh ly c o lo red than th eir c o ngeners o f t he land ,

as ev id enc e that c o lo r is f u nda m enta lly d epe nd ent u p o n light .

Th is ca n no t b e t ru e exc ept i n a v ery gen eral s en s e A ll m at e rial .

s u b s tanc es ha v e s o m e relat io n t o light and th er ef o re ha v e s o m e

c o lo r What t he c o lo r o f a bod y m ay b e is th eref o re d epend ent

.

pr imarily u po n it s c o m po s it io n and in th is sen s e its c o lo r m ay ,

b e s a id t o b e acc id ental — a rem ark that is a s tr u e o f C h lo ro phy ll

,

a s i t is o f go ld o r s ilv er o r o f red wh it e o r y ello w brick , , ,

.

B u t wh en t he part ic u lar c o m p o u nd happ en s to b e o n e lik e

ch lo ro phy ll o r a pig ment that can b e f or med on ly i n the p res en ce
,

of l tg/t t th en and th en o n ly c a n c o lo r b e s a id t o d ep end u p o n

,

t he pres enc e o f light D eep s ea fi s h es a re o ft en h igh ly c olo red

.
-

r ock s h idd en in t he ea rth ha v e th eir charact eris t ic t int ; the

b lo o d o f v ert ebrat es is red n o t fro m the presenc e o f light b u t ,

fro m t he presenc e o f c o m po u nd s of iro n I n all th es e cases .

t he c o lo r ar is es fr o m a s u b s tanc e in n o s en s e d ep end ent u po n

light fo r its fo rm at io n and ex is t enc e and t he cas e is d is t inct ,

1 The v ib ra t io n s at t hi
s p o in t a re a p p ro x im a t e ly 5 5 2 x 0
1 1 2
p e r s ec o n d ( 52 5 ,

1 C B D a v en p o rt , E x p er im en ta
. . l Mo rp h o lo gy Pa rt I pp , ,
. 1 9 2 —
1 94 .
 EXTE R NA L I N FLU E NC ES AS CAU SES O F V A R I AT I ON 2 43

fro m o n e in wh ich t he c o lo r is d u e to a s u b s tanc e formed o n ly

by the aid o f light I t is o n ly in th is latt er c as e that light m ay
.

tru ly b e S a id to b e a d i rec t ex terna l c a u s e of v a r ia t ion .

E xa mp les o f th is a re fo u nd in t he c o lo r ing o f fr u it eith er ,

u nd er n o rm a l c o nd it io n s o r in fru it ph o t o g raphy a proc es s

“
,

by wh ich p ict u res may b e mad e t o appear o n h igh ly c o lo red

fru it by s had ing with a s creen d er iv ed lik e a nega t iv e fro m t he
p ict u re to b e tran s ferred 1
.

The s u n i s s u ppo s ed t o ex ert a d irect effect u p o n t he s k i n ,

ra nging fro m the tan o f the wh it e ma n to t he d a rk c o lo r o f the

tro pical rac es T h is s eem s an ill adaptat io n and so it is as
.
,

rega rd s the h eat fo r b lack o bj ect s are war mer than wh it e o nes ;
,

b u t t he adaptat io n is n o t t o t he h eat ray s b u t t o t he ch em ical ,

fo r black p ig m ent is a lmo s t t o ta lly n o n act in ic Henc e we may

-
.

sa
y that dark s
-
k i nn ed p eo p le h av e lo s t so m eth ing in h eat res is t

anc e b u t th ey ha v e gained what is f m o re c o n s eq u enc e

,
o — a ,

s c reen aga in s t t he act in ic ray s

2
.

L ight exh ib it s it s m o s t cha ra ct er is t ic effect u p o n the eye o f

h igh er an im a ls I t h ere giv es rise to t wo remarkab le act io ns
.
,

m u sc u lar c o ntra ct io n o f t he i ris by wh ich the a mo u nt o f light

,

ad m itt ed is reg u lat ed and a n erv e s t im u lu s wh ich f o r m s a d efi

, ,

nit e i mage o n t he ret ina as u po n a m irro r and wh ich is perfect ly

, ,

c o m preh end ed by the m ind Wh eth er t he c olo rs and images

.

s een by a ll ey es a re ab s o lu t ely id ent ica l is o b v io u s ly a matt er

that can nev er b e d et erm ined I t is o f c o u rse s afe to ass u me

.

that t he images in s o far as f or m is c o nc ern ed a re id ent ical

, , ,

beca u se t he o u t lines are d u e t o t he m echan ical la ws of refra c

t io n b u t the colors a s compre/zen ded m ay b e d u e i n part if n o t
, ,

ent irely t o phy s io lo gica l pec u liari t ies

,
T hat is t he c o lo r wh ich
.
,

t o o ne is red may loo k t o him as y ello w d oes to an o th er — a ,

su ppo s it io n ent i rely p lau s ib le wh en we remem ber that with

s o m e ind iv id u a ls s o u n d a lway s s u gges t s c o lo r as well so that ,

t he n a me J o n es im m ed iat ely s u gges t s b lack or r ed o r so me , ,

o th er c o lo r d ifferi ng with d iff er ent ind iv id u a ls

,
What relat io n .

o r c oOrd i na t io n b et ween the a u d it o ry and t h e o pt ic n erv es can b e

res po n s ible fo r th is so rt o f m ix ed im press io n we d o n o t kno w .

p 38 1
1 L i ter a ry D i gest, S ep tem b er 1 6, 1 90 5 , . .

1 Ibi d
Oc to b er 7 , 1 90 5 , p 48 5
. . .
2 44 C A U S ES O F V A R I AT I O N

V ital li mi t s . C h lo ro phy llac eo u s

p lant s are ab s o lu t ely d epend
en t u p o n light for th ei r v ery ex is t enc e b u t para s it ic p lant s , ,

and an imals in gen era l a re n o t d epend ent u po n light in any ,

v ital s en s e beca u se th ey lik e an im als s u b s is t u po n h igh ly

, , ,

o rgan ized m at eria ls i n wh ich t he fi xat io n o f carbo n has b een

a lready acc o m p lis h ed by o th er o rgan is m s 1

.

A ll kn o wn fact s ind icat e that an ima l life i n gen era l is es s en

t ially su cc es s f u l i n t o t a l darkn es s M u les ha v e b een k ept in .

m in es fo r t wenty years and b ey o nd t em po ra ry s en s it iv en es s o f

,

t he ey es n o effect wa s perc ept ib le Pr is o n ers ha v e s pent th ei r .

liv es in d u ngeo n s A ll em bry o n ic d ev elo p m ent in ma m m als tak es

.

p lac e in t he t o tal darknes s o f the m o th er s bo dy I t is d o u bt les s ’

.

n o t too m u ch t o s ay that light ha s n o effect wh a t ev er U po n t he

v i ta l f u nct io n s o f t he h igh er an im a ls ; that i t is a s u n es s ent ia l

in th is res pect to an im a ls a s it is ind is pen s ab le t o p la n t s .

B acteria as a ru le n o t o n ly d o n o t n eed t he light and fl o u ri s h

, , ,

bes t in d a rk n es s b u t s tro ng s u n light is a lm o s t u n ifo rm ly and

,

q u ick ly fata l ; ind eed d irect s u n light is rec o gn ized a s o ne o f

,

t he m os t s u cc es s f u l ger m ic id es ( wh ich is o f it s elf t he pr inc ipa l

rea so n why p lenty o f light s h o u ld b e pro v id ed wh erev er d o mes t ic
an im als a re k ept ) T h is fact is illu s tra ted by in o c u lat ing a gela
.

t in p lat e u n ifo r m ly with bact e ria a s B ac i llu s a n t/t r a c i s c o v er ing , ,

t he p lat e with a p iec e o f b lack pap er o u t o f wh ich s o m e patt ern

is c u t as t he lett er E and expo s ing it a ll t o s tro n g s u n light fo r
, ,

a few h o u rs I f the p lat e is th en p u t i n t o a n inc u bat o r t he bac

.

t eria will gro w exc ept o v er t he area expo s ed to t he light in

, ,

wh ich area th ey ha v e been k illed 2

F ro m t he well kn o wn fact .
-

that bact eria i n a v a cu u m a re n o t a ffected by light we c o nc lu d e

that d eath is d u e t o o x idat io n in t he pres enc e o f light a phe ,

n o m en o n c o m m o n i n o rgan ized c o m p o u nd s .

Ligh t ri gor T he mo v em ent s o f pro t o plas m in gen era l a re

.

retard ed o r ev en s t o pped i n t he pres enc e o f int en s e light

, , ,

so that rigo r pr ec ed es d eath T h ere is th erefo re ( fo r p lant s ) .

an o pt im u m m in im u m and m ax i m u m i nt ens ity a n d bet ween

, , ,

h
T is is writt en in g en er a l ter m s , a n d rega r es s
1 dl of t he fac t t h at c ertain

h
lo wer o rga n is m s , w o s e n ea res t re a t iv es a re is t in c t l d ly a n im a l th, em s e v es l b ea r
c hl o ro p hy ll .

1 C B D av en p o rt , E xp erim en t a Mo rp
. . l h l gy
o o , Pa rt I , pp . 1 7 1 —1 7 2 .
2 46 C AU S ES O F V A R I AT I ON

than in light and a few pro m inent a m o ng whi ch a re the po as

, , ,

ger m inat e mo re rea d ily in the light as d o a ls o t he s po res o f fern s ,

a n d t he s eed s o f t he m is t let o e
l
.

E v id enc e as t o wh eth er s u n light infl u enc es t he gro wth o f a n i

m als is i n c o nc lu s iv e E xper im ent s ha v e b een c o nd u ct ed with
.

tadpoles s nails t he eggs o f c ertain fis h and with the y o u n g o f

, , ,

h igh er an ima ls b u t wh ile S light ly bett er gro wth

,
repo rt ed

“h ite
'

G reen

F IG . 27 . Effec t of l igh t u po n ra t e o f g r o wt h : s en s it iv e p la n t s g ro wn fo r t r ee h
and a h lf m a on th i s n red ,
g reen , w i t e,h an d b l ue l h
ig t . A fter C B D a v en
. .

p o r t, fro m F la m m a rio n

d u ring day light it is n o t c ertain that o th er c o nd it io n s d id n o t

,

c o ntrib u t e to t he r esu lt s E xper im ent s in f eed ing fatt en ing a n i

?

m a ls in darknes s and in light fail to es t a b lis h s p ec ia l d ifferenc es .

A ll c o n s id erat io n s po int t o the c o nc lu s io n th a t light ex ert s a

s tro ng ly m o d ify ing in flu enc e u p o n liv ing m att e r in gen era l b u t ,

that the h igh er an im als a re s u b s ta n t ially free fro m its d irect

eff ect exc ept t o s o m e ext ent in t h e m a tt er o f c o lo r .

1
C B D .
p t E p. i m t l M
a v enp h l g y P t
orI I,pp 4 9 xd er
4 4 en a or o o , ar , . 1 an 2 .

I b id p p 4 5 4 6
1 . . 2 - 2 .
 EXT E R N A L I N F LU EN C ES A S C AU S ES OF V A R I ATI ON 247

I nflu en ce of li ght u pon t he di recti on of l oc omotion or of gr owt h ;

heliot ropis m I r ri tab ility t o light is o n e o f t he
. pro pert ies o f
pro t o p las m T h is react io n is genera lly in t he f o r m o f c o ntrae
.

t io n as with m u sc le fiber res u lt ing in a s h o rt en ing of t he S id e

,
1

next t he so u rc e of light W ith free mo v ing p lant s and an ima ls .

-

th is giv es d irect io n to loc o m o tio n and th ey grad u ally s wing ,

abo u t u nt il bo th sid es are eq u ally light ed wh en if m o t io n , ,

c o nt in u es t he creat u res will o f nec es s ity pro gres s to ward the

,

light T h is is p o s it iv e h elio tro p is m

. Q u ick mo v ing fo rm s are .
-

o ft en ca r ried int o t he s o u rc e o f light by th ei r v e ry im pet u s ,

befo re the repellent effect of great h eat has t ime to act I n .

th is way mo th s fly int o t he cand le and a re k illed wh ile s lo wer ,

mo v ing fo rm s a re ch eck ed by t he h eat in t im e t o s a v e th em

s elv es I n th is latt er ca s e t he f u t u re m o v ement s will b e a
.

res u ltant o f t he po s it iv e h elio tro p is m a n d n egat iv e th er mo tro p

is m by wh ich t he in s ect s a re h eld at a c erta in rad iu s c irc ling
,

abo u t the s o u rc e o f bo th light and h eat as if n o t ab le t o leav e ,

it as i nd eed th ey a re n o t
,
.

N egat iv ely h elio tro p ic f o r m s lik e earth wo r m s a re th o s e wh o se , ,

pro to plas m d oes n o t c o ntract in the pres enc e o f light b u t o n t he , ,

c o ntra ry expand s T h es e a re carr ied a way fro m the light and

,
.
,

if in th eir wand eri ng s a light ed area is appro ach ed th ey a re

, , ,

u nab le t o ent er i t .

Wh en t he o rgan is m is n o t free to m o v e a s in the cas e of ,

s t em s o f h igh er p lant s t he eff ect will b e m an if es t ed in the d i r ce

,

ti on of g r owt/t wh ich is a ll t he r es p o n s e t o h elio tro p is m po s

,

s ib le u nd er t he c irc u m s tanc es I n cas es o f th is k ind t he s t ern .

will b end t o ward o r a way fro m t he s o u rc e o f light acc o rd ing

, , ,

a s t he p lant is n egat i v ely o r p o s it iv ely h elio tro p ic u nt il a ll ,

s id es a r e eq u a lly light ed in wh ich p o s it io n it will rem ai n d u ri ng ,

gro wth a s do o ther fo r m s d u ring loc o m o t io n

,
.

T h is p lac i ng o f t he b o dy (o r s t em) wi th referenc e to t he

v a rio u s tro pis m s is t echn ically kn o wn a s o rientat io n and ”

1 A s mo s t of t h e exa m p es t o l fo ll o w a re c o n fi n e dt o the l o wer l

a n im a s , a n d t o

p l a n ts in w ic h h p r o t o
p l as m is c o m pa ra t iv e ly exp o s e d , i t is we ll t o rem in d the

d t th
st u en at t he h igh er a n im a ls a re n o t d es t itu t e o f t h e sa m e r
p po ert ies , an d t h at

th y he a v e o n e ex p o s e d regio n p l i l
ec uy ar s en s it i v e to lig ht , n ame ly th e iris of t he

e e,
y w h os e m u sc l es c o n t rac t p ro mp t l y d un er its in fl u en c e.
2 48 C AU SES O F V A R I ATI O N

th is part ic u lar o r ientat io n with referenc e to t he rays o f light ,

wh eth er para llel t o th eir d irect io n o r t ran s v er s e ha s rec eiv ed ,

t he s p ec ia l na m e o f ph o t o tax is 1
.

Effecti v e rays A ll exp er im ent s ind icat e that t he b lu e ray s .

a re t he effect iv e o n es i n pr o d u c ing h elio tr o p ic effect s

?
Some
o rgan is m s s eem s en s it iv e t o t he y ello w b u t bo th red and u ltra ,

v io let a re a lik e i n o p erat iv e Helio tro p is m is th erefo re an effect .

d ec id ed ly d u e t o t he l u m i n ou s ray s .

Con t rib u tary con dit ion s ? Helio tro p is m is d epend ent u p o n a
v ar iety o f c o nd it io n s b o th ext erna l a n d i nt erna l t o t he o rgan is m .

1 T h ere is grea t u n c ert a in t y as t o t erm s . T he on e j ust

q u o te d p h o to ta xis

is in it s ro o t a p ro t es t a ga i n s t t he o ld t erm “
h e lio t ro p is m in th at it rec o
g n izes

[l g/ct as t h e a c t iv e ag en t ra t h er t h an t h e s u n ( b eli es
) , w ic h h is a s o u rc e n o t o n l y of

l igh t b ut of h ea t a n d c h em ica l en erg y as we ll ; an d it rec o gn i zes , to o , th t th a es e

i fl
n u en c es a re ex e rt e d as lzglzt , q u it e in d ep en d en t o f t he s u n o r an
y ot h er pa rt ic u
la r s o u rc e .

O f la te t h ere ha s b een a s t ro n g hi di p
t f
s o s it io n
th ld t o s u b s t it u t e t s ro o or e o er

t m gi i g
er , p /
v nw t t p i m
us i p l f
o ro h l i t
s
p i m n T h d
ac e o i p d t t h ie o ro s .
”
ose s o se o s

i w w ld g f th
v e ou d di o im i t b t w
ar er a n th sc r m m t t h t pp
na e e t een ose ove en s a a ea r o

wit h f t th di f th y wh i h i p h t t i d th
”
occ u r re e ren c e ti o e r ec on o e ra s, c s o o ax s , an ose

wh i h c m d wit h
a re f a e t th i t i ty f ill m i
re eren c e ti
o wh i h i
e h
n en s
p t p o u na on , c s
“
o o

a t hy . Th”
t d t l
es e s u g i t h f
en s a s o rec o t t h t i i t b i l it y n zed t h e q t
ac a rr a an e c o ns e u en

m ove m t wh t h en s , p i t i e g er t i d p
os d y
v e o r ne m h p a t h i t
v e, i t ye en v er uc u on e n en s ,

so th t ag i m or t h t an s g t
s i l y h a li t i
a re n e a
p t h i g h i t ve i t y p ei t i o rol y c a n en s a re os ve

h li t p i t l w i t it y gg t i g m idd l p i t t wh i h t h g i m will
e o ro c a o n en s , su es n a e o n a c e o r an s

b h ld
e e with d p
, as im l t h t ee -
ms ea an th a s f t igh t b t i k t
a co e n ea r e s u r ac e a n u s n o

c o ns id b l d p t h i t h b igh t l igh t f d y t h w t
era e e s n e r ti g o Th i a , e a er ac n as a sc reen . s

t
n e u ra lp i t ti fi d
o n d it i i d ib d
o r sa s e con
p h t t on s es c r e as o o o n us .
”

It m y b a y t e n ec es s a r
g i l l t h o di t i
rec o t i w h
n ze a t h b j t i es e s nc o ns en e su ec s

b tte d t d b t it i m th lik l y th t t h d iff t b h i

er u n e rs o o , u s o re an e a es e eren e a v o rs a re

on l y d iff t m if t ti e ren f th
an m
es a t l i i t b il it y t l igh t
o ns o e sa th pe n a u ra t f rr a o on e ar o

p t
ro o
p l m i as
g l d n t h t t h
en e ra ll, d an
p h t t a i e so c a
p h-
t t e o o ax s o r e v en
“
o o onus

is on ly th d it i f th g i m ft it h b gh t it l f i t
e con on o e o r an s a h p it i
er as ro u se n o s uc os on

th t a i i t b il ity i b l
one rr a d b y th ( wh i h i l w y
s a anc e y wit h b il t l
an o er c s a a s ea s a era

s y m m t y ) th t it i k i d f lim ti ti
e r , or a s a n
q i d o b y t h
ac c
p t p l a m t za on ac u re e ro o as o

th p e ti ar l i t ity t h d
c u ar n en s a an .

Th t h e ti l b j ti
eo re t dica to f ec t th
on o i t i ly
rec d Th e
re eren c e o e su n s c er a n s o un .

a tt ti
ra c ( on p l i ) i
o r re w it h fu s on t [ g/ t
s li g h t d t t t h
re eren c e o l r as , an no o e s u n as

it s s o u rc e. B t th i p é miu t ly t h
e su n s re f l igh t d f
n en th i e so u rc e o , an or s reas o n ,

an d wit h p p ad t ro d i ger u nf t h f t th ers a n th n

p f o f e p p ac s , e au or re ers or our ur o s es

t o adh t th i gl ld t m t l t d th p
ere o e s n e o er t
, t t
a f k
eas wl d g un er e res en s a e o no e e,

f or if p p l y ro d t d it m b t ti ll y
er un ers o o s ee t d s su ll p s an ti l a c o rr ec an s erv es a ra c ca

p urp f
o s es i l y w ll a r e .

1
C B D p t E a v en
p i m t l
or M p h lx
g y er P t Ien a
pp 0 0 3 ; L
or b o o ar 2 1 -
2 oe

l Phy i l gy pp 9—
. .
, , , .
,

S t di u i G es n en era
3 s o o 2 1
—
, . .

I b d pp
8 i 96 0 . . 1 2 1 .
 CA U SES O F V A R I AT I ON
2
5 0

I n t he y h igh es t an im a ls litt le b u t the eye is s en s it iv e t o

v er

light m o s t part s b eing pro t ect ed by a h ea v y ep id erm is o r o th er

,

c o v ering s o that h elio tro p is m in it s s tr ict es t sen se is in th em

,

limit ed t o the v is u a l part s I n lo wer an im a ls h o wev er with .

, ,

bo d ies les s pro tect ed light ex ert s a c o ntro lling infl u enc e u po n
,

m ov em en t s .

I nflu en ce of li ght u pon t h e dir ec ti on of l ocomot ion .

1
I t ha s
r ady been exp la in ed that so me an imals exh ib it h elio t ro p is m
al e

o n ly at c ert a in p er io d s o f th eir liv es o r o n ly in c erta in c o nd i ,

t io n s as wh en h u ngry Oth ers h owev er a re c o n s tant ly and

,
.
, ,

u n if o r m ly s en s it i v e T he c o m m o n h o u s e fly is p o s it iv ely h elio
.

tro pic wh ile t he la rv ae of the s a m e hatch ed in the dark soo n

, , ,

bec o me s tro ngly negat iv e and s o c o ntin u e wh ile in the larv a l ,

c o nd it io n ?

T h is d ifferenc e b et ween t he larv a l and ad u lt s t age is c o mmo n ,

and led L oeb at fi rs t to s u ppo s e it to b e a gen eral princ ip le ,

a c o nc lu s io n inv alidat ed by the fact that cat erp illars and th eir
im agoes b eha v e a lik e ?

B o th m o th s and b u tt erfl ies a re p o s it iv ely h elio tro p ic b u t ,

m o th s are “
att u n ed to a lo wer int en s ity T h is with th eir .
,

m o re rap id fl ight is res po n s ib le fo r th eir wh o lesa le d es tru ct io n

,

by t he nak ed fla me wh ich t he S lo wer flyin g b u tt erfl ies av o id as

,
-

a so u rc e o f h ea t .

The t en d ency o f m any s m all an im als to creep int o crev ic es as

if to h id e m u s t no t b e u nd ers t oo d a s ev id enc e o f n egat iv e h elio
tro p is m m u ch less a s ev id enc e o f t imid ity I t is o ft en d ue S im p ly
, .

t o c o ntact i rr itab ility ( s t ereo tro p is m ) fo r it is a well es tab lis h ed ,

-

fact that liv ing m atter is s en s it iv e to con ta c t with o th er so lid

s u b s tanc es T h is is t he pri n c iple d is c u s s ed in S ect io n I V and
.

t he o n e that gen era lly lies at t he ba s is o f t he h u dd ling t o geth er

o f i nd iv id u a ls o r o f th eir cro wd ing i nt o c o rn er s o r cr ev ic es
, .

L o eb bro u ght o u t th is pri nc ip le v ery n ic ely with s o m e n egativ ely

helio tro p ic bu tt erfl ies wh ich wedged th emselv es c lo sely bet ween
t wo p lat es o f g la s s i n tlze p r es en c e of lzglt t s h o wi ng ho w o n e ,

tro p ic in fl u enc e in th is case c o ntact irritab ility is c o mpet ent

1 C . B . l h y
D a v en p o rt , E xp erim en t a M o rp o lo g , Pa rt I , pp 1 80 2 1 0 ; Lo eb , —
—
.

d
S t u ies in hy
G en era l P s io lo g y
p p 1 1 1 4 , f ro m w ic m o s t o f t h e ex a m ples
. h h
a re c it e d .
1 Lo b
e , S t u ies in G en era l P d s io lo gy, p 2 0 hy . .
EXT ER N A L I N FLU EN C ES AS CAU SES OF V A R I ATI ON 2 5 1

to o v e rc omean ppo s it o e b ut one weak er ,

— in th is cas e nega
t iv e h elio tro p is m .

I t is a n o t ewo rthy fact that irritab ility t o light wh ile a pro p ,

ert y of pro t o p las m i n gen era l is m o re pro n o u nc ed in s o m e ca s es ,

than in o th ers ev en with in t he s a m e o rgan is m s Th is is t ru e

,
.

n o t o n ly in t he ey es o f an im als b u t in gen era l t he o ra l ( ant erio r ) , , ,

en d o f ey eles s an im a ls is m u ch m o re s en s it iv e t o light than is

the abo ra l 1
a s a lso is t he d o r s al s u rf ac e m o re s en s it iv e than

the v ent ra l . L ight th eref o re o perat es s tro ngly t o infl u enc e n o t

o n ly t he p os i t ion b u t t he loc o mo t io n o f an im a ls a s well .

The fo llo wi ng c o nc lu s io n s fro m L oeb u po n t he i nfl u enc e o f

light are v alu ab le ?
I n s u b s tanc e th ey a re
I. The d epend enc e of an im a l mo v ement s on light is in ev ery
po int t he s a me as t he d epend enc e o f p lant mo v ement s o n the
s a me s o u rc e o f s t im u lat io n
”
.

1 .
“
The d irect io n o f t he m ed ian p lan e o r t he d irect io n o f
t he pro gr es s iv e m o v em ent s o f an an im al c o inc id es with t he
d i rect io n of the ray s o f light .
”

2 .
“
T he mo re refra ng ib le ray s o f t he v is ib le s p ectr u m a re
m o re effect iv e than t he less refrangib le ray s .

3 . L ight o f a c o n s tant i nt e n s i ty act s a s a c o n s tant s t i m u lant .

4 . H eli o tr o p is m is i n a larg e m e a s u r e d e p e nd e nt u p o n t h e

i nten s i ty o f light d iffer ing fo r d ifferent an im a ls

,
.

5 . H e lio tr o p ic m o v e m e n t s o cc u r o n ly b et w ee n c e rta i n l im i t s

o f t em p era t u re
”
.

II “
T he o rientat io n o f an an im al t o ward a s o u rc e o f light
.

d epend s o n t he fo r m o f t he bo dy j u s t a s t he o rientat io n o f a ,

p lant to light d epend s o n t he fo r m o f the p lant .

”

1 . S y m metrica l p o int s on the su rfac e o f d o rs iv entral an im a ls

pos sess eq u al irritab ilities .

2 . Th e ir ri tab ility o f the o ra l p o le o f an an im a l is d ifferent

“

fro m t he i rritab ility o f the d o rs al po le and is gen erally greater ,

.

3 . T h e i rr i tab ili ty o f t h e v e ntra l su rf ac e is d i ff er ent fr o m t h e

irritab ility o f t he d o rs a l s u r fac e

”
.

“
T h ese three c o nd it io n s tak en t o geth er ca u s e d o r s iv ent ra l
a nimals to p lac e th ei r m ed ian planes in the d irect io n o f the
L b S tu
1 di i G l Phy i l gy pp 7 6— 78
I b id pp 8 —
oe , es n en era s o o , . .

1 . 84 . 1 .
 C AU S ES O F V A R I AT I O N
2 5 2

ray s and to mo v e t o ward o r a way fro m t he s o u rc e o f light in

th is d irect ion .
”

4 E
. y eles s an i m a ls be ha v e i n th is res pect lik e an ima ls ha v

in g ey es
”
.

III Helio tro p ic irr itab ility o f an an imal m an ifes t s it s elf

.

freq u ent ly o n ly a t c erta in epo ch s o f its ex is t enc e .

”

—
1 4 I n winged ant s th is epo ch is at t he n u pt ia l fl ight ; in
.

plant lic e it is wh en the wing s are pres ent ; in the larv ae o f

llf us ca v om i tor ia n egat iv e h elio tro p is m is m o s t pro m in ent wh e n
t he larv a is f u lly gro wn ; and i n a large n u m b er o f an im a ls t he
irr itab ility is o ppo s it e in t he larv a l and i n the ad u lt s tages .

5 B
. o th n ight and day L ep id o pt era a re p o s i t iv ely h elio tro p ic ,

and th eir h elio tro p is m is S im ilar to that o f ev ery o th er po s it iv ely

h elio tro p ic a n im a l The perio d o f S leep o f t he n ight L epid o p
.

t era h o wev er fa lls in the dayt im e and o n ly fo r th is reas o n is

, , ,

th eir h elio tro pis m m an if es t ed exc lu s iv ely at n ight 1

.
”

IV “
I n m any an im als h elio tro p ic i rri tab ility is c o nn ect ed
.

with s ex u a lity A nt s a re s en s it iv e o n ly at t he t im e o f t he
.

n u pt ial fl ight a n d in bo th ant s and Lep id o ptera t he males a re

,

m o re s en s it iv e than t he fem ales .

V T he b e h av io r o f an an im a l d epend s o n t he s u m t o t al o f
.

it s d ifferent f o rm s o f irri tab ility

”
.

V I M a ny an im a ls a re
.
“
c o m pelled t o o rient th eir bo d ies
again s t the s u rfac es o f o th er s o lid bo d ies o r to bri ng th eir ,

bo d ies in c o nt act with o th er s o lid b od ies o n as m a ny s id es

“

a s p os s ib le ( s t ereo tr o p is m ) .

V I I A n im als . m ay b e f o rc ed by light to m o v e fro m d iff u s ed

light int o s u n light and to r em a in exp o s ed t o t he h igh t em p era
t u re o f the s u n light ev en th o u gh it m ay ca u s e th eir d ea th
,
.
”

C o n s id er ing a ll t he ir ri tab ilit ies and tro p is m s to wh ich

an im als and p lant s a re expo s ed and to wh ich th ey react it is ,

n o t n ec ess ary t o app ea l to i n s ti n c t o r ev en t o t he n em o u s

,
r

sy s tem t o exp la i n a ll m o v em ent s o f an im a ls n o r is i t well to

, ,

a s c ribe to th em s u ch anthro po m o rph ic q u a lit ies a s lo v e o f light ,

d is tas t e fo r darkness preferenc e av o idanc e c u rios ity rea so n o r

, , , , ,

o th er s u ch ba s es o f h igh er i nt elligent act io n .

1Th e q u es t io n n a tu ra l ly y
a ris es , W hy is t h e d a t im e t h e p erio d fo r l
s eep in a

p o s i t iv e e lio ly h
t ro
pic a n im a ? T h e a l
n swer h a s n o t
yet b een gi v en .
 C A US ES OF V A R I AT I O N
2 54

q u ality is o ften fatal as in t he case o f m o th s I n gen era l ho w

,
.
,

ev er m att er s ha v e lo ng s i nc e b ec o m e adj u s t ed t o th es e react io n s

,

as t o o th er nec es s it ies g o v er ni ng t he b eha v io r o f liv i ng m att er .

2 . A ll exp eri m ent s i nd icat e a h igh d egree o f v a riab ility a m o ng

ind iv id u als n o t o n ly as regard s t he d egree o f res p o n s e t o helio t
,

ro p is m b u t als o a s regard s t he eff ect o f all o th er o u t s id e in flu

,

en c es ev en that o f p o iso n s
,
.

A m o ng exa m p les f u rn is h ed by t he ext end ed i n v es t igat io n s

int o th is s u bj ect we h av e s pac e t o n o t e b u t t wo
,
.

T he a m m b a wh ich repres ent s ab o u t t he s im p les t f o r m o f

,

an im al life is an exc ellent m ed iu m fo r illu s trat ing s en sit iv eness

,

t o light Fig 2 8 exh ib it s t he m o v em ent s o f o n e o f th es e b it s o f

. .

liv ing m att er u nd er t he infl u enc e o f light wh o s e d irect io n as , ,

S h o wn by t he arr o ws wa s o cca s io na lly changed ,

t he figu r es i n ,

d ic at in g t he h o u r — al l o f wh ich is s tro ng ly s u gges t iv e o f t he

,

pro c ess o f driv ing S h eep .

T he o th er exa m ple t o b e n o t ed 1 8 the effect o f lig ht u po n t he

po sit io n o f the ch lo ro phyll gran u les in the leaf c ells u nd er d if ,

feren t d egrees o f illu m i nat io n wh eth er o n t he ext erio r walls t he, ,

parti t io n o r the int erio r walls

Con di ti on s t h at d et er mi n e h el iot r opism S o m e o rgan is m s a re
.
,

po s it iv ely h elio tro p ic in o n e int en s ity and n egat iv ely so in

ano th er ; so m e a re po s it iv e at o n e t em perat u re and nega t iv e at
an o th er ; s o me a re po s it iv e in a c ertain c o nc entrat io n (of sea
wat er) and n egat iv e in ano th er and the gen e ra l pr inc ip le m ay ,

b e s tat ed that d ecrea s ing t he c o nc ent rat io n ha s the s a me effect

as i ncr ea s ing t he t em p erat u re I t th u s app ear s that the s a me
1
.

o rga ni s m can o ft en b e m ad e p o s it iv ely o r n egat iv ely h elio tro p ic

at will by alt ering t he s u rro u nd ing c o nd it io n s o f life .

S ECT I O N VII — I N FLU EN C E O F T E M PER AT U R E U PON

LI V I N G M AT TER

T he relat io n o f h eat t o liv ing m att er is m ain ly b u t n o t exc lu ,

s iv el
y q u ant i tat iv e ; that is t o s a
, y t he effect o f h eat is pri nc i ,

p ally u po n t he ra te o f gro wth a n d act iv ity I n gen eral each .

,

s p ec ies ha s it s m ax im u m ab o v e wh ich pro to plas m bec o mes

,

L b S t di 1
i G oel Phy i l gy p p
, u 65 —94
es n en era s o o , . 2 2 .
 EXT E R N A L I N FLU E N C ES AS CA U SES OF V A R I AT I ON 2 55

inact iv e ( h eat r ig o r ); i t s m i n i m u m b elo w w h ich all

,
act i v i ty
c eas es ; and it s o pt im u m that po i n t at wh ich gro wth is m os t
,

rap id Certa in fact s in th is c o nnect io n a re n o t ewo rthy

.

1 . The ma x im a m in im a and o pt i ma a re n o t t he s a m e fo r
, ,

d ifferent s pec ies .

2 . Pro t o p las m is k illed if carried m u ch abo v e t he m ax im u m ,

the o rgan is m d ec o m po s es a n d is d es tro y ed .

3 . T e m p e rat u r e s b elo w t he m i n i m u m a re n o t fata l exc e pt i n

t he pres enc e o f m o is t u re wh ich o n c o n v ers io n int o ic e d es troy s

, , ,

t he s tru ct u re o f pro t o p la s m by t he act o f expan s io n .

4 . T h e o pt im u m at w h i ch gr o w th i s m os t ra p i d is n e ar e r t h e

u ppe r than t he l wer li m it

o .

5 . B o th t he o pt i m u m and t he m ax im u m m a
y b e ra is ed by
caref u l m eth od s in v o lv ing grad u a l acc lim at izat io n .

S pecifi c effect of h eat u pon pr otoplas m 1

Begi nn ing at t he .

o pt im u m and d ecreas ing b o th wa y s to the lim it s it m ay b e s a id ,

in gen era l that pro to p las m ic act iv ity is in pro po rt io n t o t he t em

p e ra t u re T.h is i s t ru e o f t he a m o u nt o f o xygen ab s o rb ed o f ,

carbo n d io x id ev o lv ed o f ch lo ro phy ll f o rm ed and o f carbo n

, ,

fixed —
,
in o th er wo rd s o f m etab o lis m ,
T he s am e is tru e a s .

to m o v em ent s o f pro t o p la s m and i t s i rritab ility t o light c o ntact , ,

or o th er s t im u li T he f o llo wing t a b le exh ib it s t he n u m ber o f

.

electri c S h ock s per s ec o nd req u ired at d i ff er ent t em p erat u r es t o

prod u c e t etan u s in t he n eck m u s c les o f a to rt o is e ?

EFFECT OF T EM PER ATU R ES U PON A N I MA L A CT IV IT IES

S u oc x s PER S ECO N D RE

T EM P E R A TU R ES TE M P E RA TU R E S Q UR D I E To P RO DU CE

TE TA U N S TI T A N US

Effect of h eat u pon t h e r at e of gr owt h in pl a n t s .

3
The relat io n
of t emperat u re to plant gro wth is well s h o wn in t he tables o n
t he fo llo wing page 4
.

1 C B D a v en po rt , E xp erim en ta l M o rp o lo g Pa rt I , pp 2 2 2 2 3 1 h y -

—
. . . .
,

1 Ib i pd2
3.0 .
3 I i
. b P a t
r I I , pp 4 5d0 4 60 .
1 I b i p 4 5
. 1 .
‘
d . . .
2 56 CA U SES OF V A R I AT I ON

G R OWT H IN M I L LIM ET E RS OF T HE PLU M U LES AN D T HE R AD IC LFS OF S EE D

L I NG S G R OWN U N D ER D IFFE R E NT T E M PE R AT U R ES FOR 4 8 H O U RS ( Sa c hs)
P LUMUL ES R AD L
IC ES

T em pera tu re C . M a ize Bea n T e m p era t u re C . Ma ize Bea n

39
-
4 -5 47
34
3 0
28
55 o

2 . 2 22

59
-
7

M A x I M A M IN I M A , ,
AND OPT I M A FOR V A R IO US SPEC I ES A R R A NG E D

A C C O R D IN G To T HE O PT IIMA 2

S P EC I E S O PT I MUM MINI MUM M A MUM

XI

ll
B a c i u s p hO S p h o res c en S
l
Pen ic i li u m
l
Phas eo u s m u lt iflo ru s ( ea n )
1
1 b
l
Phas eo u s m u lt iflo u r s f
( b ea n )
Pis u m s a t i v u m ( pea )
4

l
S in a p is a b a ( w it e m u s ta r )
1 ‘
h d
Lep idi u m s a t iv u m 1

L in u m u s ita t is s im u m ( a x )
‘
fl
L u p in u s a b u s 1 l
l
l I o rd e u m v u ga re ( b a r e )
1 ly
l
T ri t ic u m u ga re ( w ea t )
v 1 h
Y eas t
ll
B a c i u s s u b t i is l
B ac ter iu m t erm o
Z ea m a is 1 ( I n ia n c o rn ) d
Z ea m a is ( I n ia n c o rn )
1 d
‘
C u c u rb ita p ep o ( go u r ) d
l
Ba c i lu s ra m o s u s
ll
Bac i u s an t ra c is h
ll
Bac i u s t u b erc u o s is l
l h
B ac i lu s t e rm o p i u s hl
1 G ro wt 9 6 o u rs hd u ri n g h .

1 C B D a v en po rt , E x p erim en ta M o rp
. . l h o lo g y , Pa r t I I , p 4 5 4 . .

3 R a ic d l e .
4 Pl umu el .
2 5 8 C A U S ES OF V A R I AT I ON

Effect of h eat u pon gr owt h an imals

in a n i mal s .
1
A ll larger lan d
hav e acq u ired fac ilit ies fo r m a intain ing pract ically a c o n s tant tem
p er a t u re T h is is n o t tr u e fo r a ll a n im a ls m any o f wh ich lik e
.
, ,

m ari n e s p ec ies a re n o tab ly d epend ent fo r t h eir t em pe rat u re u p o n

,

t he a cc id ent o f en v i ro n m ent in wh ich res p ect th ey d iff er b u t litt le

,

fro m plant s I t rem a in s t o n o te th erefo re what i n fl u enc e h eat

.
, ,

m ay ex ert u p o n t he gro wth o f an im a l lif e s o c o nd it io n ed a s t o b e

d epend ent u po n t he s u rr o u nd ings fo r it s t em perat u res .

I N C R EAS E IN L EN G T H ( M I L L I M ETER S
) OF Y OU NG T AD POLES OF FRO G AN D

OF TOA D , U N D ER D I FFER ENT T EM PE R AT U R ES , F R OM T HE 2 4 T H

T o T HE 8
4 TH HOU R T ER
AF H AT C HING 1

TE M P RA T R
E U ES
AVE RA GE
PB M PE R A T U R B S
AV E RA GE G R WT
O H

C .

T oa d

T wenty n in e -
th irty
o n e d egrees s eem s t o b e ab o u t t he
to -

o pt i m u m t em p erat u re fo r b o th fr o m wh ich t he tab le s h o ws that

,

t he land liv ing t o ad pro s p ered rath er b ett er u nd er t he h igh er

-

t em p era t u res than d id t he fro g a s he c erta in ly s u ffered m o re ,

u nd er t he lo we r b u t that in b o t/t th e rat e o f gro wth wa s s u b

,

s ta n t ia ll
y i n p rop or t i on to flee temp era t u r e .

The a u t ho r o f t he tab le r ep o rt s that t he i nt erv a l

bet ween
fert ilizatio n a n d h a tch ing in c o d v aries fro m th i rty days at a
temp erat u re o f to th i rt een day s at a t em p erat u re o f 6 —
° °
8
that h erring v ary fro m fo rt y d ays at 2 to elev en day s at
°

a n d that t he t im e req u i red fo r t he fr o g t o atta in a

°
10

d ev elop ment at wh ich t he h ead and ta il a re s harp ly d efined is at ,

S ix day s ; at o n e day .

1
C B D .
p t
. E p i m
a v ent l M p h l
or
gy,P t I
x I er
pp 4 57
en a—
4 60 or o o , ar , . .

I b id p 4 5 7
1 . . .
 EXT ER NA L I N F LU ENC ES A S CAU SES OF V A R I AT I ON 2 59

Bi rd s d ev elo p n ly at h igh t em perat u res T he no rmal tem

o .

p e ra t u re for the ch ick is F é ré

1
i nc u bat ed at t em p erat u r es

v ary i ng fro m 3 4 T he ind iv id u als were all exami n ed at

°
to
t he s a me a b s o lu t e t ime and t he fo llo wing fig u res expres s t he
,

perc entages o f d ev elo pm ent atta in ed tak ing that of 3 8 as th e ,

°

s tandard :
I

T he au th o r remark s that so m e d o u bt attach es to the fig u res

u nd er and and c alls o u r att entio n to the fact that
so m ewh ere n o t far ab o v e 4 1 t he s er ies wo u ld bec o m e zero
°
.

B u t fo r t he range g iv en t he d ev elo p ment is in pro po rtio n to t he

,

t em pera t u re altho u gh the h ighes t giv en

,
is c o n s id erab ly
abo v e that atta ined u nd er nat u ra l c o nd itio n s Tlze g r owi ng clzicb .

theref ore does n ot, in na t u re, a clziev e i ts op ti mu m

Effect of h eat u pon t h e di r ecti on of gr owt h , — t h erm otropis m .
1

With o u t go ing into the meth od s o f in v es t igat io n it appears that , ,

i nd ep end ent ly o f t he infl u enc e o f light o r o th er tro p is m s plant s ,

a re o ft en p o s it iv ely o r n egat iv ely th erm o tro p ic largely acc o rd ing

t o t em p erat u res T he p lu mu le o f s eed ling m a ize fo r exa m p le

.
, ,

is kn o wn t o b e p o s it iv ely th ermo tro p ic at o rd inary t em perat u res ,

wh ile t he rad ic le is p o s it iv e b et ween 1 5 and in d iff erent at

°

and n egativ e abo v e T he ind ifferent point with the bean

.

( ra d ic le) is gi v en at 2 2 .

T he s u bj ect is li tt le u nd ers t oo d and th o u gh t he im p u ls e o f ,

thermo tro p is m is wea k as c o mpa red with that o f h elio tro pis m o r
geo tro p is m it is s u ppo sed to b e o n e wh ich inc lines the o rgan is m
,

t o a lign it self in acc o rd wi th t he d i rect io n o f h eat ray s a lth ou gh ,

it is tr u e that th er mo tro p ic p lant s a re sen s it iv e t o c o nd u ct ed a s

well as to rad iant h eat .

Temperat u r e li mi ts of l ife ?
A ll experi ment s i nd icat e that as
t he t em perat u re ris es abo v e t he m ax im u m t he fi rs t effect is h eat

1 C B D av en p o rt , E xp erim en ta M o rp
. . l h l gy
o o ,
Pa rt I I , p 4 5 9
. .

1 I
d
b i pp 463 4 6 7
. .
-
.

1 F o r ext en e d dd
isc u ss io n , a n d fo r t ab es l o f t em p era t u re li m it s , s ee C B D a v en
—
. .

E i m l M rp ho lo P I
'

p o rt , xp er en t a o
gy , a rt , pp . 231 2 49 .
2 60 C A U SES O F V A R I AT I O N

r igo r ,
wh ich n pas s es int o d eath T he m o re rapid the rise
so o .

t he lo wer t he d eath p o int and t he mo re g rad u a l t he ris e t he ,

great er the res is tanc e A ga in if t he t em perat u re d oes n o t rise .

,

t o o h igh t he h eat r ig o r m ay g rad u a lly pa s s o ff and a ctiv ity m ay

, ,

b e res u m ed ev en at t em p erat u res wh ich a t fi rs t were f o llo wed

,

by ent ire s u s p en s io n o f act iv ity and ev en by r igo r T h is is t he , .

fi rs t s tage in the pro c es s of acc lim at izat io n .

S ach s fo u nd that a s en s it iv e p lant k ept at 4 0 C fo r o n e h o u r

°
.

s u ff ered lo s s o f s en s ib ility d u r ing t wenty m in u t es aft er r em o v a l .

R a is ed s lo wly t o s en s ib ili ty was o n ly t em po rar ily lo s t b u t ,

5 2
°
pr o v ed fat a l I m m e r s e d i n w at e r
. h e at ri g o r oc c u rr ed at a ,

t em perat u re 5 to 1 0 lo wer 1
Ho f m eis t er fo u nd that ha irs
° °
.

fro m the s t em a n d leaf o f E cba lli u m ag res te s h o wing liv ely ,

m o v em ent wh en grad u ally ra is ed fro m 1 6

° ° °
,
I7 C to 4 0 C -
. .

“
beca m e m o t io n les s b u t that aft er o n e o r t wo h ou rs m o v e ,
“

m ent ret u rn ed and wa s v ery v io lent C o o led and ra is ed aga in t o .

45
°
C t he .pr ,
o t o p l a s m w a s m o t io n le s s at fi r s t b u t a ft e r s ev e n ,

t een m in u t es mo v ement s rec u rred b u t were n o t rap id 1

.
”

T he v ita l lim it v ar ies great ly with t he s p ec ies T h u s ro u gh ly .

,

s p eak ing fo r bact eria i t is 4 5 C ; for c rypt o ga m s gen era lly

°
.
, ,

45
°
w i th an o cc a s io na l o n e at fl o w e ri ng p lant s ,

45
°
pr o t o z o a 4 0 w i th a fe w a s,
h igh °
as m o l
lu s k s 3 0 ,
°
—
wo rm s 4 5 5 0 ( tard ig rad es d r ied ,
°
c ru s °
, ,

ta c ea n s 2 6 — 4 ,
i n s ect s 2 7
°
— 4 3 fi s h 2 7 4 0 s,
a la m and°
e r .
,
°
-
°
,

fro g 4 0 d o g rabb it and m a n 4 4 4 5

,
°
— , v er tebrat e , ,
°

m u s c le 4 0 T h is s eries exh ib i t s a wid e range o f res is tanc e

°
,

t o exc es s iv e h eat yet few o rgan is m s can end u r e m u ch abo v e

,

A ll exper im ent s and o b s erv at io n s ind icat e that d eath fro m

h igh t em perat u res is ca u sed by c o ag u lat io n o f the a lb u m en of
t he pro t o p la s m a c i rc u m s tanc e s h o wing that al b u men carries
,

int o i ts V ita l relat io n s it s o rd ina ry pro perty o f c o ag u lat io n by

h ea t T h a t liv ing m att er c o ntain s m a ny ea s ily c o agu lable pro
.

teid s n o lo nger ad m it s o f d o u bt and th a t th eir c o ag u lati o n ca u s es ,

d eath is ev id enc ed by t he fact that o nc e in th is c ond it io n th ey

d o n o t ret u rn t o th eir n o r m a l s tat e .

1 C B D a v en p o rt E xp e ri m en ta M o rp o o g
. .
, l h l y , Pa rt I , p . 2 32 .

1
Fo r fu t a b es f ro m w ic ll l
t hes e a b s t rac t s h h a re ma d e, s ee C . B D a v en po rt ,
.

E x perim en t a l M o r p ho l gy
o , Pa rt I , pp . 2 3 4- 2 3 7 .
262 CA U SES O F V A R I AT I O N

lo nger and u nd er m u ch lo wer t em perat u res than we s h ou ld at

firs t s u ppo se .

Su b s tan ti v e v ariati on du e t o t emperatu re ; c ol or mar k i n gs .

Ear ly in t he cti o n it was rem ark ed tha t t he effect s o f t em

Se

e t re a re q u a li tat iv e as well a s q u ant i tat iv e W ith o u t d o u bt

p ra u .

t em perat u re exert S a c o ntro lling infl u enc e u po n t he c o lo r o f

b u tterflies as has been d et er mined by a n u m ber o f d irect ex
,

p er im en t s .

Fo r e xa m ple Va n essa lev a na and V p r orsa were lo ng regard ed

,
.

as d is ti nct s pec ies charact erized by a y ello w a n d

“
L ev a n a is .
-

black patt ern o n t he u pper s id e o f t he wings wh ile p r or s a ,

has b lack wings with a bro ad wh it e t ran s v ers e band and d eli
cat e y ello w lin es ru nn ing para llel to t he m argin s 1
L at er th is
”
.

wa s rec o gnized a s a cas e o f s ea s ona l d i m orp/zis m t he y ello w ,

and b lack lev a n a b eing t he s pri ng broo d and t he dark er p r o rsa

-

being t he s u m m er broo d ; that is to say lev a na emerging in , ,

t he s pri ng breed s im med iat ely pro d u c i ng a s u mm er broo d

, ,

(p r o rsa
) and ,
th is br oo d in t he s a m e wa
y g iv es r is e t o a g en era

tio n wh ich pas ses t he wint er in t he C h ry sa lis fo r m emerg ing in ,

t he s pri ng as lev a n a T h u s th es e t wo s p ec ies a re pro d u c ed

“
.

fro m the s a me s t ock t he d iff erenc e being that o n e pas s es the

,

C h ry s alis s tage in t h e s u m m er t he o th er in t he wint er ,

.

T hat th is d ifferenc e is o n e o f t em perat u re was pro v ed by

d irect experim ent D o rf meis ter s u cc eed ed in pro d u c ing pr ors a
2
.

d irect ly fro m p r or sa by t he app licat io n of war m th t o t he p u pae ,

an d
“
by the app licatio n o f c old he o bta in ed fro m lev a na n o t
t he p u re lev a n a fo r m b u t o n e int er m ed ia t e b et ween i t and
,

p ro rs a,
”
an i nt er m ed iat e o cc a s io na lly o b s er v ed i n n a t u r e and

kno wn as V par i ma ?
.

Weis ma nn repeat ing t he experiment f o u nd that by u s ing

, ,

lo wer t em perat u res lev a na c o u ld b e pro d u c ed d irect ly fro m

lev a na and he add s
,
T he c o n v ers e exp erim ent wa s als o o cca
,

s io n ally su cc es s f u l the p u p ae o f t he wint er gen erat io n b eing

,

fo rc ed to assu me t he s u m m er f o r m by t he infl u ence o f a h igh er

t em perat u re d u ring o r S ho rt ly aft er p u pat io n ,
3
,
.
”

1 W eis m an n , G erm P l as m , p 3 79
. .

1 V ern o n , V a ria t io n in A n ima l s an d Plan t s , p . 2 33 .

3 Ibi d . ; a so l W eis m an n , G erm P l as m , p 3 79

. .
 EXT E R N A L I N FLU E N C ES AS C A U SES OF V A R IAT I ON 263

Here t em perat u re x rt a c o ntrolling infl u enc e u po n

s eem s to e e
p igment f o r m at io n alth o u gh Weis mann is caref u l to info rm u s
,
1

t hat the t wo pattern s d o n o t c o rres po nd that if we were to

s u per po s e o n e u p o n t he o th er it wou ld s eem that the
“ ” “
,

b lack parts in p rorsa d o n ot c o rres po nd to the y ello w o n es in

lev a na and that the wh ite band in the f o r m e r d oes n o t c o rre
,

s po n d to
[ei th e r ] a y ello w o r a
[] b lack part in the latt er T h i s .

ba nd is o n the c o ntra ry en ti rely wa n t i ng in lev a na and is

, , ,

repres ent ed by bot/t b la c k a n d y ellow r eg ions 1

.
”

A ga in Weis m ann experiment ed with P oly om m a t n s p ic/wa s ?

,

a s pec ies d is trib u t ed o v er the wh o le o f the t em pe rat e an d

“

c o ld er part s o f E u ro pe a n d A s ia T o ward t he n o rth ( in .

”

G ermany ) the u pper s u rfac e o f t he wings is o f a beau t if u l “

redd is h gold c o lo r h enc e it s po p u lar name fi re b u tt erfly

-
,
” “
,
.
”

Bu t he s ay s F art h er s o u th t he redd is h go ld c o lo r is m o re o r
,
-

les s th ick ly d u s t ed with b lack and S pec imen s fro m S ic ily , ,

Greec e o r J apan o ft en d is play o n ly a few redd is h go ld s cales

,
-
,

t he gen era l app earanc e b eing a lm o s t b lack

”
.

“
I n G erm a n y th is b u tt erfly is d o u b le brood ed and the t wo -
,

gen erat io n s a re s im ilar b u t in c erta in d is trict s o f sou th ern

,

Eu ro pe t he fi rs t gen erat io n is redd is h go ld t he s ec o nd -

, ,

wh ich fl ies in m id s u mm er and is kn o wn as the v ariety eleu s ,

hav ing t he wings well d u s t ed with b lack A s in G er m any .

d u ring exc ept io nal ly ho t s u m mers ind iv id u als with a black is h

tint ha v e repeat ed ly been cau ght t ogeth er with the o rd inary
fo rm and Weis m ann o b serv es that it wo u ld seem the “

bu tt erfly bec o m es red wh en exposed to a m o d erat e t em perat u re

and b lack wh en t he h eat is great er .

A tt em pt s t o pro d u c e th es e f o r m s at will h o wev er by regu la , ,

t io n of tem perat u re o nly part ially s u cc eed ed B u t the c o nd it io n s .

were s ev ere T h ere wa s n o c o m m o n anc es t o r Weis m ann u nd er

. .

took t o pro d u c e t he s ou tltern f or m fro m t he nor tlt ern s tock and

v ic e v er s a I n sect s reared fro m G er m an b u tt erfl ies b u t k ept in
.

h igh tem perat u res were in many in s tanc es d us t ed with black ,

bu t r es em b led tlze d a rk es t or m s f tue s ou t/zer n

n on e o f ” mm f o

eleu s . Co n v ers ely , b u tt erfl ies ra ised in c oo l t em pera t u res fro m

1 W i m
e s G m Pl m p 3 7 9
an n , er as , . .

1
I b id p p 3 99 4
. .
-
00 .
2 64 C A U S ES O F V A R I AT I ON

N eapo litan t ck were light er in c o lo r than in th eir nat iv e hab i

s o

tat b u t no n e were s o light c o lo red as t he o rd ina ry Ger m an

,
“ -

fo r m 1
T h is d iff erenc e he a sc ri b es to t he c u m u lat iv e infl u enc e
”
.

o f t he nat u ra l s ea s o na l t em pe rat u res and is q u ick to pro t es t ,

again s t its i nt erpretat io n as ind icat ing an inh eritanc e o f acq u ired
charact er s He calls it a ca se o f int ernal s elect io n as bet ween
.

wi nt er and s u m m er d et er m in a nt s .

Ho wev er that is o f n o c o n seq u enc e in t he pres ent c o nn ect io n

,
.

T he fact s h ere giv en S h o w bey o nd a d o u bt that o u t s id e t em p era

t u res ex ert a d irect effect u po n s o im po rtant a ch a ract er as
c olo r Wh eth er th is occ u rs by C h em ic a l d is t u rbanc e in p ig ment
.

fo r m at io n by int ernal s elect io n o r by o th er m ea n s d oes n o t

, ,

great ly m atter h ere T h e re is s o me ev id enc e t end ing t o Sh o w

.

that t he lig ht c olo r o f po lar an im a ls is d u e to t he d irect ac t io n

o f c o ld T h is if tru e argu es for C h em ic a l act io n u po n p ig m ent
?
, ,

a s t h e ca u s e o f c o lo r changes d u e t o t em p erat u re .

Temper atu re an all per v adi n g influ en ce T em p erat u re d iffers

-
.

fro m mo s t o th er ext erna l f o rc es in being fo r m any s pec ies at , ,

lea s t a n a ll p erv a d i ng in fl u enc e

,
-
H igh er an im als and p la nt s .

a re th em s elv es c ent er s o f h eat pro d u ct io n and in g en era l th eir ,

t em perat u res a re t he a lgebra ic s u m o f th eir o wn h eat prod u ct io n

and t he h eat o f th eir s u rro u nd ings L o wer o rga n is m s h o wev er .
, ,

a re v ery largely d ep e nd ent u po n th ei r en v i ro n m ent fo r t heir

t em perat u res and in cas es o f t his k ind t he ent ire pro t o p las m o f
,

t he b o dy is a ff ect ed .

S EC T I O N V I I I — E FF EC T O F C H E M I C A L A G EN T S U PON
P R O T O PL A S M I C A C T I V I T Y
A ll d ev elo p ment all d ifferent iat io n a n d a ll f u nct io nal a ct iv ity
, ,

o f liv ing o rgan is m s a re t he r es u lt o f pro t o p la s m ic a ct iv ity ; b u t

pro t o plas m ic act iv ity is in the la s t analy s is ch em ic al act iv ity

, , ,

and it is c ert a in ly s u bj ect t o m a ny o f t he laws c o ntro lling o rd i

nary ch em ical react io ns I t is n o t ic ea b le in the s t u dy o f v it al
.

1 W eis ma n n ,
G erm P l as m , pp .
9
3 9
—4 00 .

1 T he wri t er h as so me h
w ere rea d th at an im a s l on S h ipb o ar d b ec o m e ra p i d ly
l igh t er in l
c o o r as t h e c o a t b ec o m es ex
p ose dt o in t e n s e co ld , b u t h e is u n ab e l to
v erif y th e rep o rt .
2 66 C A U SES O F V A R I AT I O N

bro min in the presenc e o f wat er act fa tally u po n all o rgan is m s “

by S plitt ing [t he]wat er fo r m ing hydro ha logen c o m po u nd s and ,

-
,

lea v ing t he o xygen t o u n it e with the liv ing pro t o p la s m 1

.

Hy dr ogen A m oeb ae s u bj ect ed t o an at mo s ph ere o f hyd ro gen

.

fo r t wenty fo u r m in u t es b ec a m e m o t io nles s s o m e ha v ing a s s u m ed

-
,

t he S ph er ica l f o r m T he sa me gen eral res u lt f o llowed in trad es

.

cant ia hairs b u t fro m t he fact that n orm a l act iv ity was res t o red
,

by ad m itt ing air it was a s s u m ed that t he res u lt s aro s e n o t fro m

any inj u rio u s effect o f hydro gen b u t fro m t he exc lu sio n o f o xygen ?

Ox i ds of ca rb on — CO2 an d CO T h es e t wo o x id s o f carb o n
, .

ha v e v ery d iffer ent effect s u po n pro to pla s m T he fo rm er lik e .

,

hydrogen seem s t o act o n ly by exc lu d ing o xygen d ea th when

, , ,

it res u lt s b eing d u e m a in ly t o as phyx ia wh ile t he la tt er k ills by

, ,

attack ing the pro t o plas m d irect ly ?

Cat alyt ic poi son s 4

A large n u m ber o f u n s tab le carb o n c o m
.

po u nd s n eith er ac id n o r b a s ic and th erefo re n o t characterized

,

by int en se ch em ica l act io n are yet v io lent p o is o n s Here b elo ng , .

t he an aes th et ic s a s ch lo ro fo r m ch lo ra l ethy l eth er a lc o h o ls etc

, , , , , ,
.

T h es e u n s table c o m po u nd s a re charact erized by a liv ely “

c o nd it io n o f mo lec u lar m o v ement ( N ageli) wh ich is c o n s id ered ,

to d is t u rb t he n o r m a l mo v em ent s o f the pro t o p la s m

“
or to ,

lead to ch em ica l tran s f o rm at io n s in t he u n s tab le alb u m en o f t he

pro t o pla s m ( Lo ew) .

Cata lyt ic s u b s tanc es a re s u ppo s ed to ex ert th eir act io n n o t by

ent eri ng int o and eff ect ing n ew c o m b i n at io n s b u t by d is t u rb in g ,

thro u gh th eir m ere presenc e t he us u al behav io r o f bod ies I t ,

.

is in th is way that pro t o p la s m s u ffers i n th eir pres enc e T h u s .

hydro ch lo ric and pru s s ic ac id s u n it e o n ly at h igh t em pe ra t u res ,

exc ept in t he pres en c e o f v ar io u s eth er s wh en th ey will u n it e ,

ev en at T he v igo r o f th is cata lyt ic act io n is in pro po rt io n

t o t he m o lec u lar c o m p o s it io n 5
T h u s in t he m ethan s eries with .

C H3 as t he bas e we ha v e C H, C s C 3 H8 etc i n wh ich , ,

.
,

1 C . B D a v en p o rt , E xp eri m en ta
. l M o rp h o l o
gy Pa rt I ,
, p 4 . .

1 Ib i d .
p 5
. .

1 I b id .
p 6
. .

4 A f ree ex t ra c t f ro m C . B . D a v en po rt , E xp erim en t a l M o rp h o l gyo , Pa rt I ,

pp 7
.
. 8 .

T he fa c t s h ere s ta te d a re ta k en a lm os t l it era ll y fro m C . B . D a v e n p o rt,

Ex peri m en ta M o rp ho l l gy
o , Pa rt I , pp 7 8 .
-
.
 EXT E R N A L I N FLU E N C ES AS CA U SES O F V A R I AT I O N 2 67

t/ze p ois onou s a c t ion i n cr eas es u p to a c er ta i n li m i t i n p r op or t ion

Me f n u m b er ab C a tom s , wh ile “
th is lim it t he c o m
”
to o o v e

po u nd s a re m o re s tab le and a re m o re ind i fferent ,

as fo r e xa m ple
1111 1 3 11111 C H
( ZI M 1 0

A ga in , in ch a series if the H at o m s b ec o m e rep lac ed by

su ,

o n e of t he ha lo gen s t he po is o no u s pr o p ert ies c o rres po nd i ng ly

,

increa s e ; th u s : 1

C H4 m ar s h ga s inn o c u o u s
, ,
.

C HaCl s light ly a naes th et ic

,
.

C H Cls ch lo ro fo r m po werf u l an ae s th et ic
, , .

CC 14 v ery dangero u s s t u pefy ing i n v o lu ntary m u s c les

, ,
.

C h lo ro fo rm and eth er a ffect all pro t o p las m b o th p lant and ,

animal h igh er as well as lo wer T h ey s eem to prod u c e at fi rs t

, .

( t w o t o fi v e m in u t es i n a 2 5 p er c e nt w at e r s o lu t io n ) a v e ry “

int en s e exc it em ent in t he m o v em ent o f t he pro t o p la s m fo l ,

lo wed by “
s tro ng v ac u o lizat io n and th en t he cyto plas m grad ,

u ally bec o m es imm o b ile and d ies if the infl u enc e is c o nt in u ed ,

.

I n a s im ilar way t he v ar io u s a lc o h o ls ex ert s t u p efy ing eff ect s in

pro po rt io n to t he n u m b er o f C H, rad ica ls pres ent and c a rbo n ,

d is u lph id ( C5 2) is o ne o f t he mo s t po werfu l cata lyt ic po iso n s .

Pro t o p la s m is th erefo re s u bj ect to catalyt ic d is t u rbanc es i n ,

wh ich i t is no t d ifferent fro m o th er and m o re o rd inary ch em ica l

m at erials — a fact in it s elf exc eed i ngly S ign ificant t o t he s tu
,

d ent lo o k ing fo r f u nda mental ca u s es o f v aria t io n .

Poi son s which f orm sal t s ?

T h es e a re ac id s and bas es wh ich
L oew believ es as s tat ed by D a v enp o rt
,
u n it e
[d i rect ly
] w i th ,

t he pro t ei n s u b s tanc es o f t he pro t o p la s m pro d u c ing s a lt s

”

, ,

d is t u rbanc es that o f c ou rse s o o n lead to d eath T hu s fo r m ic “

.

ac id ev en in s m a ll per c ent s
, p er c ent t o pe r c e nt ,

— prev ent s the d ev elo p ment o f bact eria On t he o th er hand

,

.
,

so m e pro t o p las m ha s acq u i red a r es is tanc e to o rgan ic ac id s t he ,

v in egar eel liv ing in 4 e c e nt ac et ic ac i d and t h e g land c e lls

p r ,

o f s o m e m ari n e Gas tro p o da s ec ret e fr o m 2 per c ent t o 3 p er

c ent o f s O a s trengt h wh ich is fa t a l t o mo s t pro t o plas m .

1 The h a lo gens —fl u o rin , c hl o rin , b ro m in , an d io in d —f o rm a g ro up of sub

s t an c es o f v er y s im i a r l c h em ic a l p ro
p er t ies , b u t fo rm , in t h e o r er d n a me d , a

d ec r eas in g s eri es as t o c h em ic a l en erg y

—4
.

1
C B D av en po rt , E xp erim en ta M o rp
. . l h l gy
o o ,
Pa rt I , pp . 12 1 .
2 68 C A U S ES O F V A R I ATI O N

N a geli xp erim ent s ind icat e that wat er d is t illed in c o p

’
s e
1

e v ess els o r s tand ing f o u r day s in o th er v es s els wi th t welv e

p r ,

C lean c o ppe r c o in s p er each li t er o f wat er wa s fata l t o S p i ro ,

gyra tho ugh t he pro po rt io n of c o pper to wat er was b u t 1 to

,

T h es e rea ct io n s res u lt ing in d eath rath er than in m o d i fi ed

,

act io n a re im po rtant n o t as S h o wing p ri m a ry ca u s es of v aria

, ,

b ility b u t a s pro v ing aga in that li v ing pro t o p la s m is s t ill s u bj ect

t o m a n y o f t he ch em ic a l a ffi n i t ies th a t c o ntro lled it s elem ent s
bef o re th ey beca m e o rgan ized int o liv ing m att er I t m u s t b e .

rem em bered in th is c o nnectio n that m any o f th ese s u b s tanc es

attack o n ly liv ing pro to plas m hav ing n o act io n u po n d ead p ro ,

to p la s m s h o wing that at d eath t he h igh ly c o m p lex m at e rials

,

h a v e t o s o m e ext ent at lea s t bro ken d o wn

, ,
.

T he act io n o f s o m e p o is o n s lik e n ic o t in is pro po rt io n al t o

’

, ,

t he d ifferent ia t io n o f n erv o u s s u bs tanc e o th er s lik e c o ca in ,

and atro p in fi rs t exc it e and th en pa ra lyze t he c entral n er v o u s

,

s y s t em o f v ert ebrat es b u t act a s v io lent p o is o n s u po n u n d iff er

,

en t ia t ed pro t o plas m ( Pro t o zo a )

?

Tox ic poi son s I t is n o t the g er m that k ills b u t rath er it s

.
,

s p ec i fic t o x in that d eranges s o m e o f t he v ita l f u nct io n s b ey o nd

end u ranc e T he d i re effect s o f ger m d is ea ses a re th erefo r e d u e

.

n o t so m u ch t o t he o rgan is m s th em s elv es a s t o th ei r c o n s tant

m a n u fact u re with in t he b o dy of a ch em ical p o is o n wh ich t he

pro t o plas m cann o t end u re and preserv e i ts n o rm a l f u nct io n s I t .

m ay d ie in t he att em pt o r it m a y s u cc eed and b ec o m e ac c li ,

m a t ed b u t wh ile t he s tr u ggle is o n t he b o dy f u nct io n s will b e

, ,

c o n s id era bly d is t u rbed I t is S ign ificant that c o m po u nd s s im ilar

.

t o th o s e o f d is ea s e pro d u c in g bact eria ha v e been

-
e xtract ed fro m

t he s eed s o f s o m e phan ero ga m s fo r exa m p le ric in fro m t he s eed s ,

o f t he ca s t o r o il b ean et c -
I n t his c la s s m ay c o m e t he p o is o n s
,
.

s ecr et ed by c erta in an im a ls a s t he ra tt les nak e and c o bra fata l , ,

t o v ert ebrat es b u t i nn o c u o u s t o I nf u s o r ia and F lagellata .

I t is also n o t ewo rthy th a t t he b lo o d s eru m o f o n e s p ec ies

rap id ly d is s o lv es the c o rp u s c les o f an o th er ( red and wh it e) and ,

is th eref o re inj u r io u s o r fa ta l acc o rd ing to t he a m o u nt s pres ent ?

1
C B D p
. t E p i
. m t l
a v enM p h lor
g y P t I p ,4 x er en a or o o , ar , . 1 .

I b id p p
1
3 d 4 . . 2 I b
anid p 2 .
3
. . 22 .
2 7 0 C AU SES O F V A R I AT IO N

I nsect T he po is o n
pois on s .
—
bees fo r mic ac id is fatal— of

t o in s ect s and s m a ll an im a ls and i n s u ffic ient q u ant ity to t he ,

larger v ert ebrat es inc lu d ing m a n th o u gh freq u ent s t ing s o f a

, ,

m o d erat e n u m b er lead rap id ly to acc lim at izat io n I t is n o t e .

wo rthy t oo in th is c o nn ect io n that t he s t ing o f t he i n s ect

, ,

( m u d w a sp fo r exa m p le) d o es n t a lway s k ill b u t o ft en m e rely

,
o

para lyzes s o th a t the creat u re s to red with the egg will rem ain
,

a liv e to fu rn is h fo o d t o t he larv ae so me week s lat er .

Gall s Ga lls a re the d i rect res u lt o f t he s t ing o f an in s ect

.
,

lea v ing a s pec i fic p o iso n to act u p o n a part ic u lar f o rm o f pro t o

p las m The res u lt is n o t d eath b u t a d iv erting o f the act iv it ies
.

int o ent irely n ew chann els D ar win s tates that n o less than .

fifty eight k ind s o f gall a re pro d u c ed o n t he s ev eral s p ec ies o f

-

o ak s by C yn ip s with it s su b gen era and M r B D Wa ls h s tat es -

,
. . .

tha t he can add m any mo re 1

.
”

D ar win f u rth er rem ark s that m any gall in s ect s are exc eed ing ly
s m a ll and th a t c o n s eq u ent ly t he dro p o f p o is o n th ey inj ect m u s t
,

b e exc eed ingly m in u t e ; m o reo v er it is n ev er inj ect ed b u t o nc e , .

T he gro wth that fo llo ws h o wev er is s p ec ific and c o nt in u o u s , ,

.

He q u o t es Wa ls h as s ay ing Galls affo rd go o d c o n s ta nt and , , ,

d efin ite cha ract ers each k ind k eep ing as tr u e to f o r m a s d oes
,

any ind epend ent o rgan ic being 1

and he ca lls o u r attent io n to ,
”

t he fact that s ev en o f t he t en d is t inct ly d ifferent ga lls prod u c ed

o n t he willo w a re by in s ect s wh ich th ou gh es sent ia lly d is t inct ,

s pec ies et res em b le o n e an o th er s o c lo s ely that i n a lm o s t all

y ,

cas es it is d iffic u lt and in mo s t ca ses im po s s ible to d is t ingu is h

t he f u ll gro wn in s ect s o n e fro m an o th er
-
T he d ifferenc e in the .

q u a lity o f t he po is o n s ecret ed by in sect s s o n early alik e cann o t b e

great yet it is s u ffic ient to giv e ris e to galls wid ely d ifferent
,
.

L a s t and n o t lea s t he ment io n s that

, Cy n ips f ec u n da t r ix has
,

b een kno wn to pro d u c e in t he T u rk is h oa k to wh ich it is n o t ,

pro perly attach ed exact ly the s a me k ind o f gall as o n t he

,

Eu ro p ean oak T h es e latt er fact s apparent ly pro v e that t he

.

n a t u re o f tb e p o ison is a m ore p o werfu l ag en t i n d eter m i n i ng

the f or m f
o the g a ll tlza n i
[ ]s M e spec ific c /za ra c ter o
f t b e t ree
3
w/ uclt is ac ted on
'
”
.

1 D a rwin A n im a ls , an d l
P a n ts , I I , 272 .
1 Ib id .
p . 27
3
.

1 Ib i d .
( s ec o n d diti e on
) , 2 73 . T he i ta lic s a re m in e .
 EXTER N A L I N FL U E NC ES AS C AU SES OF V A R I AT I ON 2 7 1

Tu mor s . T h ose
rgro wth s o f v ario u s part s o f t he bo dy
ov e ,

called t u m o rs a ris ing fro m ca u s es n o t well u nd ers t oo d hav e

, ,

their s pec ific charact ers as tr u ly as if d eriv ed fro m inh erit

anc e Wh eth er the charact er o f t he t u mo r is d eriv ed primar ily
.

fro m t he t is s u es a ffect ed o r fro m so me o u t s id e S pec ific ca u s e is

n o t kn o wn b u t i t i s a s ign ific a n t f a et t/za t tlze p r o top la s m w/t ic /z
, ,

d er i v ed i ts c lta ra c ter s or igi na lly by i n /zer i ta nc e lza s u n d erg o ne ,

p er ma n en t a nd d efi n i te a lter a ti on t/zr oug lt t/ze op era t i on f

o ca u s es

a bs o lu te ly d is tinc t f r om i nlzeri ta n c e, wlzetb er i n tern a l or ex ter na l

to t/ze org a n is m , t/t u s wi ng t/te p os s i b ili ty of d i v er ti ng tlze en er

s no

g i es o f i n lt er i ted m a teri a l i n to a bs ol u tely n ew cha n n els t/t r oug /z

app a r en t ly s lig /zt c a us es .

At th is po int it is well to call ou r a tt ent io n to t he pro fo u nd

changes ( perm anent v ariat io n s ) wro u ght u po n t he c o n s t it u t io n
o f t he ind iv id u a l by s u ch i nt ernal ext er na l c irc u m s tanc es a s v ac c i -

nat io n o r t he inj ect io n o f ant it o x in as well as to t he im mu n ity ,

ac q u ired throu gh a S ingle attack o f an infect io u s d is ea se .

Germinat ion of seeds I t is a well kn o wn fact that c erta i n

.
-

ch emicals acc elerate and o thers retard t he pro c es s o f germ ina

t io n J u s t why th is is tr u e is no t c lear o n any o th er gro u nd
.

than that o f t he o rd inary s u s c ept ib ility o f gro wing pro t o p las m

t o s t i mu lant s and s edat i v es T he proc es s o f germ d ev elo p ment .

req u ires in add it io n to what is c o ntain ed with in t he s eed o n ly

, ,

o xygen wat er and a fa v o rab le t em p erat u re ; t he infl u enc e o f

, ,

o th er ch emica ls m u s t b e ind irect .

Ch emota xi s an d ch em otr opis m

1
T he infl u enc e o f ch em ical .

s u b s tanc es u po n t he lo c o m o t io n o f free m o v i ng o rga n is ms is t ech -

n ic all
y kn o w n a s ch e m o tax is and th eir i nfl u enc e u p o n t he direc ,

t io n of gro wth ( in p lant s ) is kn o wn as ch emo tro p is m ?

T he
d is t inctio n is hard ly wo rth o b s erv ing fo r o u r p u rpo s es b eca u s e ,

bo th ph eno m ena arise fro m a d irect infl u enc e o f ch em ica l s u b

s tanc es u po n liv ing pro t o p la s m eith er att ra ct iv e o r rep ellent ,
If .

att ract iv e i t ( t he pro t o p las m ) will m o v e t o ward the m at eria l in

,

1C B D .
p . t E i
a v en
p m t l M
or , p h l x er
gy P t I 3
en a 4 5 ; P t
or I I pp o o , ar , 2- ar , .

33 5 3
-
4 ; L b 2 P hy i l
oe
gy f t h
, B i pp 5
s o o 8o8 9 0 8 e 8 6 88
ra n , . 0, -
, 1 1 , 1 -
1 .

1A h sl dy b as a rea t d th m d i ti t i
een no e i ,
ft e sab d b tw
e s nc on s o en o s erv e e een

g t i
eo a x s , t h i fl e f g n it y u en c e o
p l m t i ra v d g t u p i mon i t i
o cofl o o n , an eo ro s , s n u en c e

u p on t h d i t i e f g recwt h ; l on ob t w h
ro l i t i d h
a so l i t p
e i m een e o ax s an e o ro s as c o v er

i g
n c o rres
p d i g i fl on f t
n h n l
( g )
i h t
u en c es o e sun .
2 7 2 C A U S ES O F V A R I AT I O N

q u es t io n pro v id ing it is free to d o s o (as in t he C ase o f the

,

lo wer p la nt s and al l an im a ls ) o r it s gro wth will b e d irect ed ,

t o ward it if ( as in t he C as e o f h igh er p lant s ) it is fi x ed and u n

ab le to m o v e T he fo r m er is s t rict ly s peak ing chem o t a x is ; the
.
, ,

latt er is ch em o tro p is m F o r o u r p u rpo s es it is a d is t inct io n

.

with ou t a d ifferenc e b eca u s e in t he latt er ca se t he p lant is

,

u nab le t o ind u lg e in lo c o m o t io n and p e rf o r m s t he n eares t po s ,

s ib le act in changi ng t he d i rect io n o f gro wth Bo th a re lo c o .

m o t io n u nd er t he c i rc u m s tanc es ; b o th a re d u e t o t he s a m e
ca u s e — a
,
ch e ca a n ity
m i l ffi o r a tt act io n
r — and they d iffer ,

beca u se o f d ifferenc es in t he o rga n is m s a ffect ed in res pect to

t he p o wer o f lo c o m o t io n no t beca u s e o f d iffer enc es in t he nat u re
,

o f t he f o rc es in act io n T he t wo t er m s are th erefo re fo r o u r

.
, ,

p u rpo ses s yno ny m o u s as d en o t ing a po wer o f att ract io n bet ween

,

pro t o p la s m and c ertain o rd inary ch em ica l c o m po u nd s s u ch a s t o

ca u s e t he pro t o p las m to appro ach if it b e free to m o v e o r if n o t , , ,

t o gro w i n that d i rect io n wh ich is all that c an b e d o n e u nd er

,

the c irc u m s tanc es t o s at is fy t he a ffin ity .

C h em o tax is o r ch em o tro p is m a s t he writ e r pref er s to ca ll

,

it 1
,
ha s b u t a S lend er h o ld u po n h ig h er an im a ls I t appear s t o .

b e lo c a lized in t he n o s t ri ls and to m an ifes t it s elf o n ly in t he

s en s e o f s m ell agreeab le o r o th erwis e ; b u t in lo wer o rgan is m s
, ,

ev en i n m any in s ect s i t is apparent ly n o t c o nfin ed t o a m in u t e

,

f ract io n o f t he s u rf ac e b u t perv ad es t he wh o le o rgan is m with

,

an infl u enc e that is all b u t o v erp o wering I t m ay o f c ou rs e b e .

a id ed o r o pp o s ed by o th er tro p is m s a s gra v ity o r light in wh ich , ,

c as e t he t o tal res u lt is the algebraic s u m o f a ll the en ergies

o p erat iv e b u t that ch em o tro p is m is a fo rc e t o b e r eck o n ed with
,

in d ev elo p m ent is a fact n o t to b e d o u bt ed .

Ex ampl es of ch emotr opis m 1

En glem a nn n o t ic ed that bac .

t eria u nd er a c o v er g la s s will g ath er a lo ng t he m argin o r if ,

green algae b e intro d u c ed th ey will c lu s t er abo u t t h em s o lo ng

a s th ey a re pro d u c in g o xyg en b u t in the d arkn es s th ey will n o t ,

1 Th e en di ng
“
ta x is is fro m t h e G r ee k , m ea n in g a s s o rt m en t o r a rra n
g em ent ;
t he en di ng t ro p is m is f ro m t ro p ic ,
”
t o t u rn . ( Ih em o t ro p is m is p ron o u n c e d
k t m d t rOpi zm
’
- -
.

l h
1 U n t i o t e rwis e n o te d wh at fo ll o ws is a f ree t h ou
g h t no ex a c t t ra n s c rip t fro m
d a ta gi v en in C . B D a v en po rt
.
’
s Ex p eri m en ta M o r l p h l
o o
gy P , a rt I , p p 3 2 - 39
. .
2 74 C AU SES O F V A R I AT I O N

an
y k i nd c ertain ly incapable o f ex erc is ing anyth ing lik e in
,
is
t elligen t C h o ic e T he charact e ris t ic m o v em ent m u s t h a v e been
.

d u e t o t he s p ec i fic ch em ic a l act io n o f t he j u ic es o f t he m eat
u p o n t he pro t o p las m o f t he m u s c le fi b e r s o f t he t en t a c le wh ile ,

t he p a p er had n o s u ch act io n and h enc e n o m o v em ent f o llo wed .

T h is m o v em ent and n o n m o v em ent lo o k lik e int elligenc e o r -

in s t inct a n d ha v e o ft en
p a s s ed fo r o n e o r t he
o t h er t o t h e i n fi n i t e c o n

f u s io n o f t he s u bj ect ?

L u m b r ic i f et i d i
e liv e
i n t he d ecay ing c o m po s t
o f o ld s tab les and pr o b ,

a b ly t he ch em ical nat u re

o f c ert a in s u b

s t a n c es c o n

t a i n ed in t h e
c o m p s t h o ld s o

th em th e r e ,

F IG 9
. S
2 t i
.m l t i g ff t
u a f t
n i e h m i
ec l o
p m
c er a n f o rc w h
e en o n
ca s e u on us
“

l ti
c u ar ac
p i on : af m t l id ec e o
p t h t t eal f h a lf O f t h
a u on
e
e en a c es o

th ti i
e ac im l t
n an s t th i ti u a esd it i d w i t t h
e r ac o n an s ra n n o e
I ) O t t o m Of
1
t 16
im il l y p l d i i p ti ‘
m thou A pi. f p p ec e o a er s ar ac e s no e ra ve

b o x is c o v ered
with m o is t b lo tt ing paper and t he o th er ha lf with a th in lay er
o f c o m po s t a ll t he wo r m s will gath er o n t he c m p o s t S id e
,
o .

D ec ap it a t ed wo r m s b eha v e in t he s a m e wa y 1
so th a t t he ,

e ff ect is d u e t o a gen era l infl u enc e n o t t o n erv e c ent ers

“
.
,

L o eb s ay s 3
I h a v e o ft en p lac ed p iec es o f lea n m ea t and
,
“

p iec es o f fa t fro m t he S a m e an im al S id e by s id e o n t he W ind o w

S ill b u t t h e fly n ev e r fa iled t o la i t s egg s o n t he m eat and n o t
, y
H tr d f c r w t t cc ra ar
”
o n t h e fa t ( e
. ie o o u s e i h o u ,
s u e s s t o is e l v e
a ,

in t h e fat ) I t can ea s ily b e s h o wn th a t lar v ae o f t he fly a re po s i

.

t iv ely ch em o t ro p ic t o ward s c ert a in ch em ic a l s u b s tanc es wh ich

a re f o r m ed fo r i n s t a nc e in d ec ay ing m eat o r ch ees e b u t wh ich
, , ,

a re n o t f o r m ed in fa t T h e s u b s t a nc es in q u es t io n a re pro bab ly
.

v o la t ile n it r o g en o u s c o m p o u nd s and t he ch em ica l effect s o f

“
,

1Th i b j t will b p
s su ec d f th e d I
u rs u e ti t d R fl
ur er u nA ti er ns nc an e ex c on .
1L b Phy i l gy f h B i
oe , s o o
p 9 o t e I b id
ra n ,
pp .
—
86 8 7 0. 1 . . 1 1 .
 EXT ER NA L I N FLU E NC ES AS CAU SES O F V A R I AT I ON 2 75

t he d iff u s ing mo lec u les o n c erta in element s o f t he S k in infl u enc e

t he t en s io n o f t he m u sc les ca u s ing mo t io n ,
.

T he fema le fly is attra ct ed by m eat t he s a m e as a re lar v a and , ,

a s s oo n as t he fly is s eat ed o n t he m eat ch em ica l s t im u li s eem

t o thro w int o act i v ity t he m u s c les o f t he s ex u a l o rgan s and ,

eggs a re d ep o s i t ed o n t h e m eat T h is ch em ica l s t im u lu s is .

abou t all th ere is o f the wo nd erf u l in s t inct by wh ich in sect s ”

a re led alway s to d epo s it th eir eggs in exact ly the r ight p lac es .

Th es e and s im ilar exa m p les S h o w t he eff ect o f c erta in ch em

ic als u po n free m o v ing o rgan is m s -
I t rem a in s t o illu s t ra t e th ei r .

eff ect u p o n t he d i r ec t i o n ofg r ow th a m o ng p lant s wh ich a re n o t ,

free to m o v e .

Firs t o f all we a re t o no t e t he effect o f c ertain C h em ica ls u po n

,

t he t entac les o f in s ect iv o ro u s p lant s D arwin n o t ic ed that wh en .

“

dro ps o f wat er o r s o lu t io n s o f n o n n itro gen o u s c o m po u nd s a re -

plac ed u po n t he lea v es o f t he s u nd ew D ro sera t he t entac les , ,

rema in u n in flec ted b u t wh en a dro p o f a n itro gen o u s fl u id s u ch ,

a s m ilk win e a lb u m en inf u s io n o f ra w m eat s a liv a o r is ing la s s

, , , , ,

is p lac ed o n t he leaf t he t entac les q u ick ly bend in ward s o v er

,

t he dr o p 1
D arwin f o u nd that o f n i n e s a lt s o f a m m o n ia tried
”
“
.
,

a ll ca u s ed infl ect io n and o f th es e t he ph o s phat e wa s t h e m o s t

,

po werf u l and that s o d iu m s alt s in gen eral ca u sed infl ect io n

,
“

with extrem e q u ickn es s T h is act io n o f n itro gen o u s ph o s

.
,

phat ic and o th er ch em ica ls c o m mo n t o an ima l life was t he

, ,

sa m e u p o n t he t entac les o f in s ec t iv o ro u s p lant s as u p o n t he

t entac les o f lo wer an ima ls su b s is t ing u po n the s a m e k ind o f

foo d T h u s plant and an im a l tiss u es appear to b e s u bj ect t o
?

t he s a m e gen era l la ws in th is regard .

F ro m th is p o int o f d ep a rt u re c o m m o n t o b o th p lant and a n i ,

m al we no t e that t he an i ma l free t o m o v e d o es s o in res p o n s e

, , ,

t o th is c la ss o f s t im u li Wh a t d o es t he pla n t d o that cann o t

.

mo v e ev en a s t entac les m o v e I n o th er wo rd s ho w d oes c hem o t

, ,

ro p is m a ff ect t he d i r ec ti on o w l a m ng gh r p lant
fg r o t z o hi e s

R o o t s in gen eral a re s u ppo s ed t o gro w t o ward o xyg en and ,

po llen t u bes will c ertainly t u rn to ward t he s t ig m a o f t he fl o wer ,

1 C B D a v en po rt E xp erim en t a M o rp o o g , pp 3 3 5- 3 3 6
. . , l h l y . .

1 S ee t he ex e ri
p m en t o n a c t in ia n s
p re v io u s
q u o t e f ro m L o eb P ly d hy i l gy
s o o of

—
,

t he B ra in , p p 49 5 0 . .
2 7 6 CA U SES O F V A R I AT I O N

t he s u pp o s mon
being that s u gar is the s pec ial attract ing s u b
s tanc e in t he latt er cas e I t is n o t ewo rthy that t he p ollen t u be
.

is att ract ed n o t s i m p ly to it s o wn s t ig m a b u t also t o t he pis t il

and t he o v u le o f o th er s pec ies ev en of a d ifferen t gen u s with ,

wh ich it is u nable t o u ni t e ?
D a v enpo rt s ay s 1 '

T he res ul ts of ex peri men ta tio n u po n c h emo tr op ism S h ow tha t v ario us

s u b s ta n c es ma y d irec t the gro wt h o f s u c h elo nga ted o rga n s as ten d rils , roo ts ,
‘

a n d h yp h a o f plan ts I n m an y in s tan c es i t c an b e s h o wn t h a t the

d irec tio n o f growt h is o n the wh ole a d v a n ta geo u s to the o rgan ism In .

o t h er c as es , h o wev er , the respo n s e s eem s to ha v e n o rela t ion to a d ap ta tio n .

The im med iat e ca u se

change of d irect io n is exc ess iv e
of
“

gro wth o n o n e s id e d u e to exc es s iv e im b ibit io n o r to exc ess iv e

,

o r r es tr ict ed a ss im ilat iv e act iv i ty .

T he ev id enc e s eem s c o nc lu s iv e that t he ch emica l element s

c o n s tit u t ing liv ing m att er hav e n o t ent irely los t their o rd inary
a ffin it ies and pro pert ies ; that pro t o plas m is in many res pect s
s u bj ect to t he la ws o f o th er ch em ica l c o m po u nd s ; that it s ac t iv

it ies m ay b e acc elerat ed o r retard ed ; that th ey m ay b e t em p o

ra ril o r ev en p er m a n ent ly m o d i fi ed in charact er by ch em ica l
y
a lt erat io n o r ev en ent irely d es tro yed by ent ering int o n ew and
,

s trange c o m b inat io n s with s u rro u nd ing s u b s tanc es Here is a .

real ca u se at leas t o f occa o na

,
s i l V a riat n
io — po ss ibly o f th o se ,

s u dd en chang es we ca ll m u ta t io n .

R hyt h mical con t r acti on of mu scl e in stit u t ed b y c ert ain ch em

ical su b s t an ces . T he i rritab ility

and c o n s eq u ent c o ntract io n of
m u sc les d u e to infl u enc es of a ch em ica l n a t u re hav e already been
no t ic ed as in the c a s e of the m o v ement of the t ent ac les of the
,

act in ian the attract iv e o r rep u ls iv e effect o f c erta in od o rs and

, , ,

t o s o m e ext ent in t he p lac ing o f in s ect egg s in

,
“
exact ly t he

right s po t .

I t is n o w well kn o wn that c erta in s alt s ex ert a s pec ific act io n

u p o n m u s c le exc i t ing ev en rhyth m ic c o ntract io n
, A s lo ng ago .

a s 1 8 8 1 B ied er m ann d is c o v ered that if t he m u s c le o f a fro g b e

caref u lly exc is ed at a lo w tempera t u re (0 1 0 th en weight ed °

-
°

and d ipped int o a pe r c e nt s o lu t io n o f s o d i u m C h lo r id c o n

tain ing als o s mall a m o u nt s o f s od iu m ph os phat e an d sod iu m

1 C B D a v en p o rt , E x p erim en ta
. . l M o rp h o lo
g y , p 339
. .
1 Ib i d .
p 34 3
. .
2 7 8 C AU SES O F V A R I ATI O N

t he and there fo r ming with it s s u b s tanc e d efin it e c o m

mu sc le
po u n d s and he believ es t he acc elera t ing effect o f H o r O H is
,

d u e t o th eir cata lyt ic act io n in fac ilitat ing t he fo rm at io n o f th es e

c o m po u nd s I n th is c o nn ect io n i t is t o b e rem em bered that the
.

s er u m o f the b od y wh ich bath es t he m u s c les is a lway s in h ea lth , ,

s tr o ngly s alin e and S light ly a lka lin e .

F u rth er expe rim ent s c lea rly s h o wed that t he s alt s o f po tas
s iu m and th ose o f calc iu m m a gn es iu m s tro nt iu m m anga n es e
, , , ,

and c o balt t end s tro ngly to prev en t c on tra c t ion th is b eing es pe ,

c ia lly tru e i n t he ca s e o f p o ta s s iu m and ca lc iu m f o rc ing the ,

c o nc lu s io n that cer ta in d efi n ite s u bs ta nc es a re nec es sa ry to c on

t rac t ion tha t c er ta i n ot/t ers t en d to a cc eler a te a n d s t i ll ot/zer s to
r eta r d t/t is c /za ra c ter is t ic a c t i v i ty o f m u scu la r lis s u e .

A rtifi cial part h en ogen esi s t h rou gh ch an ges in t h e su rr ou n din g

solu t ion ? T he ind ef a t igable lab o rs J acq u es L oe b u po n th is of

s u bj ect ha v e n o t o n ly thro wn m u ch light u p o n t he es s ent ia l

feat u res o f fec u ndat io n b u t inc id entally t hey ha v e a ffo rd ed

,

resu lt s o f h igh v alu e ln d et erm in ing t he nat u re and range o f

'

ext erna l infl u enc es u p o n t he C haract eri s t ic act iv it ies o f liv ing

m att er ?

I t had lo ng b een kn o wn that m any o f t he eggs o f sea

u rch i n s arthr o p o d s a n d m ari n e wo r m s ev en w/zen u nf er t il iz ed
, , , ,

wou ld if left fo r a c o m pa ra t iv e ly lo ng t im e in sea wat er b egi n

, ,

t o s eg m ent reach ing t he t wo and s o m et im es t he fo u r c elled

,
-

s tage L oeb and lat er M o rga n f o u nd that if t he c o nc entra

.
,
“
,

t io n of t he s ea water b e ra ised s u ffic ient ly by t he add it io n o f

c ertai n s alt s a seg m entatio n o f t he n u c leu s takes p lac e with
,

ou t any s eg m entat io n o f pr o t o p la s m cyt o p la s m S u ch egg s

[ ] .
,

h o wev er wh en bro u ght back int o no rm al s ea water d iv id e into

,

as m any c ells a s th er e were pref o r m ed n u c lei I n n on e o f

3 ”
.

th es e experim ent s d id t he c ell d iv is io n lead t o the f o r m at io n

“

o f a b las t u la A h eap o f c ells at t he b es t ab o u t s ixty were

.
, ,

fo r med and then ev eryth ing s t o pp ed A s in t he ca se o f t u mo rs

, .

1 d l hy l y
Lo eb , S tu ies in G en era P s io o g , Pa rt I I , p p 53 9 69 1 .
-
.

h
1 T es e in v es t i
ga t io n s a v h
e b een p u b il d
s h e fro m t im e t o t im e, es p ec ia in th e ll y
l
A m er i c a n jou r n a l of P b y s iology , a n d a t er ( 1 90 5 ) in b o o
r
fo rm u n er t h e t it e, k d l
d
S t u ies in G en era l Phy i l gy s o o .

1 Lo eb , S t u di i G es l Phy i l gy P t
n en era s o o , ar .
—
I I , pp 5 40 5 4 1 .
 EXT E R NA L I N FLU E N C ES AS C AU SES OF V AR I AT I ON 2 79

and gal ls h ere was growth with ou t sys t emat ic d iff erent iat io n ;
,

c ell d iv is io n with o u t t he fo rm at io n of an em bryo .

E nc o u raged by t his d egree o f seg mentat io n and by his experi

ment s u po n irri tab ility o f mu s c le Loeb t ried a great v ariety o f ,

s o lu t io n s in v ariou s d egrees of c o nc entrat io n in the h o pe of

, ,

carrying t he segmentat io n far enou gh to prod u c e real emb ryos

and li v e free mo vi ng larv a,
-
.

He was grea t ly ha m pered by t he fact that u nfert ilized eggs

d o no t fo r m m em bran es as d o f ert ilized s o that gro wth t end ed ,

t o b e fo r m less and ev en wh en as s u ming d efin it e f o r m in t he

,

blas t u la s tage th ere were o ft en fo rm les s masses o f d iv id ing

1

matt er ly ing t o o n e S id e .

B riefly s tat ed t he fo llo wing facts d ev elo ped d u ri ng t he pro g

,

ress of th is sys temat ic experiment

1 I n a so lu t io n o f so d iu m C h lo rid t he eggs were u nab le t o
.

reach ev en the b las t u la s tage .

2 With t he add itio n of MgC l, h owev er blas t u la were fo rmed

.
, , ,

b u t th ey d id n o t mov e Wh en aft erward p lac ed in n o rm al sea .

wat er mov em ent soo n app eared .

3 W
. i th thr ee ch l o ri d s ( N a K and C a )
“
t h e e gg s n o t ,
o n l y ,

reach ed t he b las t u la s tage and s wam arou nd in the mo s t liv ely

way b u t th ey reach ed t he ga s tr u la an d ev en the p lu t eu s s tage
, ,

with t he exc ept io n h o wev er that pract ica lly n o S k elet o n was , ,

fo rmed 1
S u ch larv a li v ed abo u t ten day s
”
. .

4 T
. h e add i t io n o f a trac e o f r e s u l t e d in t h e f o rm at io n
o f a S k elet o n b u t it wa s n o t q u i t e n o r ma l
,
I t was m ad e n o r m al .

by add ing a t rac e o f Mg Cl, .

Th
1 ly t g
ree ea r h t i t i f th
s a es a re c ly d l p m t f ll mb y
a ra c er s c o e ea r ev e o en o a e r os

( )
I t h m e/ m lb
or n a , o r y t g i w h i
u h ll d
erri i i g i s a i t e, g l nb l c ce v s on v es r s e o a o u ar

m f
as s o d d llro u n h m e ce l di ti t li k t h g p
s , ea c o re o r b h
es s s nc , e e ra es o n a unc or

th d f m lb y ; ( ) th bl t l t g i wh i h th
e s ee s o a u err 2 e t ll b
as u a m s a e, n c e o u er c e s ec o e

con d d h wi g d i ti t t l y
ens e , s o n a th b ls t d m ;
nc d ( 3) th g t l
o u er a er , e as o er an e as r u a

t g
s a i wh i h
e, n id b c m one s
p h d i (
e i gi t
ec o d ) es w l d
usp h
e i n nv a na e , as o n e ou us n a

h ll w b b e b ll with hi th mb f m i g k i d f m th d t m h A f w
o o ru r a s u , or n a n o ou an s o ac . e

f m
or s n ev erg t b y d t h i t
e g b t
e onm t p q i k
s s al y t h g he,i t d iffu t i os ass u c ro u , eren a

tion p d i
ro c ee
g p i d ly I nh ig h ra im l t.h tn l y ( t d
er an m )g i i a s e o u er a er ec o er v es r s e

to th ki e s d it n pp dan g t h s i a ( d end m )a t t hes , i t e l g n n er en o er o e n ern a o r a ns .

Am gon h i
s ea u rc w h i h w h ns, d p
c i m t eret h t t ere u n
g i k w er ex er en e n ex s a e s no n
—t h t g f f m i g l
,

as t h p1 t e u en s , e s a e o It w thi t g
ree- th ov n p i a rv a . as s s a e e ex er

m t d i d t p d
en er es re o ro u c e.
1L b S t di oe i G
, l Phy i l gy P t I I pp 58 5 586
u es n en era s o o , ar , . .
2 80 C A U SES OF V A R I AT I O N

5 . xp eri m
A ll
ent s i nd icat ed that it is i m p o s s ib le to s ec u re
e

m o re than t he b eginn ing o f s eg mentat io n fro m an u nfer t ilized

egg wi t/t ou t ra is i ng tlze c o nc en t ra t i on o f tlt e s ea wa ter .

6 . t his p u rp o s e Mg Cl2 was p ec u liarly effectiv e and

B u t fo r
n o rmal p lu t ei ( free s wimm ing larv a ) d ev elo p ed fro m u nfert ilized
-

egg s ly ing in n o rm a l s ea wat er aft er ha v ing la i n fo r t wo h o u rs

i n a s o lu t io n o f M g C 12 o f pro per s trength ?

7 T.h e s e eff ect s s eem ed to b e d u e to t he increa s ed c o n een

t ra tio n in t he s ea wat er bri ngi ng abo u t i ncrea s ed os m o t ic pres

,

s u re and res u lt ing in a lo s s o f wat er o n t he part of t he eg g .

T h is loss o f wat er s eem s t o b e t he act iv e ca u s e o f rap id s eg

m entat io n and a v ariety o f s u b s tanc es were d is c o v ered whi ch
,

were ab le t o br ing it ab o u t .

8 The princ ipal d i fferenc e n o t ic eab le in t he p lu t ei was that

.

tho se d ev elo ped fro m f ert ilized eggs s wa m freely a t the top o f
t he wat er wh ile th os e d ev elo p ed fr o m u nfert ilized egg s
,
were
a ll at t he bo tt o m o f t h e d is h and u nab le t o r is e
”
.

9 E.xp e ri m e nt s u p o n t he m ari n e ann el id C ha t o p t eru s

1
i n d i
c a t ed that art i fic ia l d ev elo p m ent is ea s ier than wi th t he s ea

u rch in b u t that it is ach iev ed by a d iff erent s o lu t io n I n t he

'
.
,

wo rd s of t he expe rim ent er W e m ay s ay that Cha to p ter u s ,

po ss es ses a h igh er d egree of parthen o genet ic t end ency than t he

A rbac ia [s ea u rch in ] eggs 3
and if t he s ea wat er c o nta in ed
“
,
”

o n ly a s light ly great er pro po rt io n o f K we s h o u ld find that ,

C ha t o p t eru s wa s n o r m ally pa rt hen o gen et lc 4 ”

.

10 I f c erta in f o r m s are prev ent ed fro m b ec o m ing pa rt hen o

.

genet ic by t he c o ns t it u t io n o f the s ea wat er we m ay infer that ,

tho se wh ich a re nat u ra lly parth en o genet ic a re so by t he c o n s t i

tu t io n of t he b loo d o r t h e s ea wat er enab ling t he egg t o d ev elo p 5
.

1 1
“
T he bridge b et ween t he ph en o m ena o f nat u ra l and
.

art ific ial parth enogenes is is fo rm ed by th o se an imals in wh ich

1 Lo eb , S t u dies in G en era l Phy i l gy P t I I p 6 4
s o o , ar , . 2 .

1 d
M ea had l dy f d th t if p
a rea o un at R C ]i dd d t er c en s a e o s ea wa t er th e
u n fer t ili d ze eggs o f C h t pt
a oth w t th i p l
eru s b di
ro wh il
ou e r o ar o es , e the a dd it ion

of
p er c en t N Cl p d
a dro uceh ff t no L b S t di
suc e ec .
”
oe , u es i n G en era l
Phy i l gy
s o o , Pa rt I I , p p 6 56- 6 5 7. .

1 Lo eb , d
S t u ies in G en era P l hy i l gy
s o o , Pa rt I I , p p 6 5 4 —
655 . .

4 Ib i d .
p 66 5
. .

5 I b id .
p 683
. .
282 C AU SES O F V A R I ATI ON

S EC T I O N I X — EF F EC T O F SA LI N E SO L U T I O N U PO N
D EV EL O PM E N T I N A Q U A T I C A N I M A L S

S ea wat er d iff ers fro m fres h wat er in two part ic u lars salini ty ,

and d en s ity bo th of wh ich exert m ark ed infl u enc e u po n an im al

,

life and b et ween wh ich it is o ft en d iffi c u lt to d is crim inat e A .

goldfi s h p lu nged int o s ea wat er at fi rs t S h o ws V io lent in c oOr

din a ted m ov ement s b u t S h o rt ly bec o m es im mo b ile and rises
”

t o t he s u rfac e by v irt u e o f it s lo we r s p ec ific g ra v i ty 1

.

T he effect o f fres h wat er u po n m arin e o rgan is m s is eq u ally

s t rik ing. T h ey go i m m ed iat ely to t he bo tt o m and m o v e with
d ifli c u lty S wim ming an ima ls s wim bad ly if at all and s m all
.
,

fi s h es h av e to mak e m u ch ex ert io n to ris e t o t he s u rfac e 1

.

On m any m ar in e an im als as m o llu s k s and fi s h fres h wat er , ,

a ct s a s an an a s t het ic t he m o llu s k s s oo n y ield ing t o pa ra ly s is

, ,

t he fi s h app earing t o s u ff er fro m lack o f ai r T he res p i rat o ry .

mo v em ent s b ec o m e d eep and rap id T he t iss u es bec o me .

s wo llen s o that s o ft b o d ied an im a ls a re v is ib ly d ef o rm ed — in

-
,

fi s h es t he eyes a re fo rc ed o u t t he foo t o f ga s tro po d s s wells the , ,

bloo d c o rp u s c les s well u p and b u rs t a n d m u s c u lar t is s u e may ,

i ncrea s e as m u ch as S ix t im es in v o lu m e 1
.

Many o f th es e effect s a re c learly d u e to d ifferenc es in pres

s u re wh ich m ay a m o u nt t o ma ny at m o s ph eres b u t it rem ain s ,

t o s epara t e so fa r a s m ay b e t he eff ect s o f s alin ity fr o m th o se

, ,

o f s p ec ific gra v ity .

R ath er s tart ling C la im s ha v e b een m ad e fro m t im e t o t ime

as to t he c on v ers i on of on e sp ec i es i n to a n ot/zer by a lt ering t he

d egree of salin ity F u rth er in v es t igat io n seem s t o S h o w that in

.

a ll s u ch ca s es int er m ed iat e f o r m s a re kn o wn t o o cc u r wh ich ,

argu es that t he t wo fo r ms wh ich had been rec o gn ized a s d is

t inct were n o t b o th goo d S pec ies ; that is t o s ay it is a cas e o f ,

o n e s p ec ies wi th wid e v ar iab ility a s t o c erta i n charact er s n o t that ,

o f t wo d is t inct and well d efi n ed s p ec ies I f h o wev er d ifferenc es

-
.
, ,

in s a lin ity a re eff ect iv e i n bringing abo u t a lt erat io n s in ev en a

s ingle charact er t he fact is o f int eres t h ere n o m att er what
, ,

s p ec ific lin es S h o u ld b e d ra wn by t he b io lo g is t .

1
C B D .
p t
. Ea v en
p i m t l M p h l
or g y P,t Ix er
p 79 en a or o o , ar , . .

I b id p p 7 9 8
1
. .
, 0 .
 EXTER N A L I N FLU EN C ES AS C AU S ES O F V A R I AT I ON 283

The a ll c ru s tac ean s A r tem ia sa lina and A m i lluzu s en zz

sm .

hav e been rec o gnized as d is t inct t he fo rmer liv ing in b rack is h ,

and t he latt er in s t ill mo re c o nc ent rat ed wat ers the t wo d iffer ,

in g m ai nly in t he n u m ber and lengt h o f bris t les bo rn e at t he

extrem ity of t he ca u dal fi n s .

Ea rly in t he sev ent ies S c h man k ewit s c h p u b lis h ed an acc ou nt

o f t he m u t u al c o n v ers io n o f each f o rm i nt o t he o th er b u t t he ,

fact s as giv en by him ha v e been great ly o v ers tat ed as freq uent ly ,

happens in repet it io n T h ey a re su ffi c ient ly S ign ificant as fi r s t

.

repo rted and it seem s well t o giv e t he o rigi na l s tat ement as

,

rec o rd ed by Bat eso n wh ich is as fo llo ws

1
,

Th e lt l goo sa K y l ik w d i i d d b y d m i t
a n, u a n , ppe as d v e a a n o an u r an a

lo we p t ; the wa t
r ar i the l t te b ei g a t ated wi th al t wh ile the
ers n a r n s ur s ,

wa ters of th e u p per p art were less sal t . By a s p ri n g fl o od i n the yea r 1 8 7 1

th e wa ters o f the u pper p a r t o f the
‘
la k e s wep t o v er th e d a m a n d. red u c ed
th e d ens i ty of the lo wer wa ters to 8 B au m é ab o u t s p an d
°
g . .

h
in t is wa ter g rea t n u m b ers o f A sa li n a t en . h a ppea red , p res u m ab l
y h av i n g
h
b een was ed in fro m the u p per pa r t o f the la ke or fro m th e n eig h b o ri n g

s al t po o ls . h
A fter t is th e d am wa s m ad e go o d a n d th e wa ters o f th e lo wer
k
la e, b y ev a po r a t io n , b ec am e m o re an d more c o n c en tra ted , b ein g , in t he
s u mm er o f 1 8 7 2 , 1 4 B ( a b o u t s p g
° °
in 1 8 7 3 , I B B ( a b o u t s p g
. . . .

in A u g u s t, 1 8 7 4 , B ( ab o u t s p g a nd la ter in t . . h a t yea r

the sa lt b eg a n to c rys ta llize o u t I n 1 8 7 1 th e A rtem ia [as firs t. c a rri ed

o v er ]h ad c au d al fins of
go od s ize, b earin g eig ht t o twelv e, ra rely fifteen
b ris tles , b u t wit h the progr ess i v e c o n c en tr a tio n o f th e wa ter th e gen era
tio n s o f A rt em i a p ro gres s iv ely d egen era ted , u n t il a t th e en d o f th e s u m m er
o f 1 8 7 4 a la rge pa rt o f t h em h ad n o c a u d al fi ns , th u s p r es en tin g the

c ha r a c ter o f A . m i l/za u s en i i . F I S C HE R AND M I L N E E DWA R DS

-
.

B at es o n add s
A was pro d u c ed ex perim en ta ll b y g ra d u a l c o n c en tra tio n
s i m i la r s eries y
o f wa ter, lead i n g to th e ex t rem e fo r m res em b l i n g A m i l/za u s en i i I t wa s . .

fo u nd also tha t i f the a n im als wit o u t c a u d al fi n s were k ept i n wa ter w i c

h h h
was gra d u ally d ilu ted , a fter so m e weeks a p a i r of c o n i c al p ro m i n en c es ,

ea c h b ea ri n g a s i n g le b ris tle, a ppeared a t the en d o f the a bd o men .

The xperiment er als o relat es that by breed ing s a l i na in s t ill

e

mo re d ilu t ed wat er he atta in ed a fo r m res em b ling S cha ffer s

’

gen u s B ranch ip u s B u t t he princ ipa l d ifferenc e bet ween t he

.

genera is that in A rt emi a the las t s eg m ent is abo u t t wic e a s

1 Ba t es o n , M a t eria ls , etc .
, p 96. .
2 84 C A U SES OF V A R I ATI O N

lo ng ac h of t he o thers wh ile in B ranch ip u s it is d iv id ed I t

as e , .

is extrem ely S ign ificant that th is d iv is io n s h o u ld b e pro d u c ed in

A rt em ia by c u lt u re in c o m parat iv ely fres h wat er b u t t he fact is ,

n o warra nt fo r t he a s sert io n that o n e gen u s can b e c o n v ert ed

int o an o th er by a lter ing the en v i ro n ment I t ra th er ca s t s d ou bt.

u po n t he W is d o m o f a C la s s i ficat io n wh ich es tab lis h es g en eric d is

t in c t io n s u po n d ifferenc es s o s light and s o eas ily bro u ght abo u t .

T he s a m e experim ent er s t u d ied s pec ies o f t he gen u s D aphn ia ,

and fo u nd in th eir cas e also c o n s id erab le s t ru ct u ral and phy s i

“

o lo gi ca l chang es t he fres h and s a lt wat er f o rm s d iff eri ng in his

,
-
,

o p in io n by charact ers u s u a lly h eld to b e s pec ific

1
.
,

B at es o n s t u d ied t he c o m m o n c ock le a m o llu s k ev erywh ere , ,

present in t he A ra l\ Sea and it s o u t ly in g wat ers o f d ifferent

d egrees of s alin ity On e of t he lak es ( S hu m ish K u l) o n it s
.

wes t er n s h o re exh ib it ed n o les s than s ev en d is t inct t erra c es ,

h eld to repres ent s u cc es s iv e s tages o f t he wat er lev els d u ri ng

it s lo ng pe ri o d of dry ing u p with c o rres po nd ing increa s e in
,

s alin ity T he m o s t n o t ic eab le d ifferenc es in t he s h ells tak en

.

fro m th ese s u cc es s iv e t er rac es and pres u m ably d u e to in c reas

,

i n g s a lin ity a re o u t lin ed as f o llo ws

,
1

1 . A d im i n u t io n i n t he t/ziek nes s o f t he s h ells fi rs t apparent ,

in t he th ird t errac e I n th e s ev enth t errac e th is change was s o

.

m ark ed that t he s h ells were a lm o s t h o rn y and th ei r weight was ,

n o t a th i rd o f that o f t he S h ells fro m t he fi r s t t wo t erra c es .

2 . D im in u t io n o f t he s ize o f t he b ea k [wit h t he lo wering o f

t he lev el] .

3 . High c o lo rat io n [Th e.a u th o r d o e s n o t s t a t e w h ich w a

y
t he changes ra n wheth er u p o r d o wn th e t errac es b u t he re
, ,

m ark s th a t all t he s h ells o f a g iv en t e rra c e were v ery n ear ly

“

a lik e in t ext u re t hicknes s and d egree o f c o lo rat io n

, , .

4 .Gr o o v e s b e t w e e n t he rib s appearing o n t he in s id e o f t he

s h ell a s r idges with rectang u lar fac es .

5 .A gr e a t d i m i n u t io n i n ab s o lu t e S iz e o f t h e S h ells o n t he

lo wes t t errac e .

6 A lt e rat io n in pro po rt io n o f lengt h t o breadth rangi ng

.
,

fro m 1 to i n t he S hells o f t he fi rs t t errac e t o 1 t o in

th o s e o f t he s ev enth and 1 t o o n t h e s ho res o f n eighb o r ing

1 V V i ti
ern o n , i A im l
ar a on n nd Pl t p
a s an 75an s , I b id pp
. 2 .
75
1
7 6 . . 2 , 2 .
2 86 C AU S ES OF V A R I AT I O N

U pper l imi t s of dev el opment n is d ev elo p ment

. I n what se se

c o nd it io ned u po n u se D o es u s e S im p ly enab le t he part to

?

atta in it s no r ma l and p roper d ev elop m en t to wh ic h it is ent it led ,

u nd er t he la ws o f h ered i ty o r d oes it s ti m u la te dev elop men t

,

bey on d tlze n o rm a l 1
.

S o me b io lo gis t s at o nc e as s u m e t he latt er to b e imp oss ible

and t hat any u n u su a l app earanc e is a cas e o f ata v is m I t is tru e .

t hat in times lo ng pas t there m ay hav e ex is t ed d u ck s that walk ed

and o t hers t hat flew m o re and better t han t ho s e wh ic h D arwin
exa m i n ed b u t wh en d id nat u re prod u c e a r u nn ing o r a tro tt ing _

ho rs e as goo d a s t he o ne o f to d ay ? T o what rem o t e anc es t o r

d o o u r v io lin is t s and o u r p ian is t s o we t heir S k ill and what wa s ,

t he i n s tr u m ent o n wh ich t hey acq u i red it ?

T he acc o m pany ing c u t is a fac s im ile o f a pro per ly att es t ed let
t er wr itt en with t he f eet by a y o u ng wo m an t wenty three y ea rs -

o ld who los t b o th arm s at t he a ge o f t en A m o ng her o th er ac .

i m S he n u m b er s c u tt ing s ewing thr ad i ng w

p l s h e nts
( e he
, r o n

need le) d rawing s weep ing and a great v ariety o f ho u sewo rk

, , ,
?

Here is a ca s e o f p u tt ing part s to an ent irely n ew us e ,

d em and ing a n ic ety o f adj u s t ment that was n ev er acq u ired ev en

in o ne o u t o f a m illio n o f t he anc es t o rs C o u ld th ere b e bett er .

ev id enc e o f t he fact that few ind iv id u a ls ev er u s e and th ere ,

fo re few ev er d ev elo p m o re t han a f ract io n o f the capac it ies bo rn

,

i n th em ; that t he po s s ib ilit ies o f life a re s eld o m rea lized and ,

that n ev er are all fac u lt ies d ev elo ped to th ei r u t mos t in any

S ing le ind iv id u al ?

A ll th is is c lear b u t it is n o t s o ea s y to d et erm in e wh ere t o

dra w a line and say A ll d ev elo p m ent belo w th is is d ue to
,
“

inh erit a nc e an d a ll a b o v e t o u s e T he tr u th wo u ld s eem t o b e

.

th a t d ev elo p ment d epend s u po n bo th inh erent t end enc ies and

ext ern a l c o nd i t io n s a ff o rd ing o ppo rt u n it ies fo r th eir ex erc is e ,

and that t he m ax i m u m of d ev elop men t is rea c/zed on ly w/zen

b ot/z a r e a t tlzei r op t i m u m T h ere is m u ch f o rc e in t he wo rd
.

o pt i m u m T o o m u ch ex erc is e t o o m u c h f o o d t o o mu ch
“
.
, ,

t em perat u re o r t oo m u ch o f any o f t he c o nd it io n s o f life is

,

as u nf o rt u nat e a s t oo li tt le .

1 Th
e au th o r saw m wh w t wi th hi f t b t th y w
on e an o ro e tt h ed
s ee , u e ere a ac

di
rec tly t
o t h b dy wi t h
e o , l g Wh t no e s a ev er .
 EXTER N A L I N FLU E N C ES A S C A U SES O F V A R I AT I O N 287

. l y h hyd
F IG 30 illu s tra t es t he a b i i t o f a ig l l
ev e o p e d
p a rt to p erf o rm a n ex t rem ely
u n u s u a l s erv ic e
. Th e a b o v e l ett er was writ t en wit h t he feet in s t ea d f with
o

h d
th e an s

I t is as f u t ile to attem pt to d ec id e wh et her int ernal o r ext er

n al c i rc u m s tanc es a re m o r e h elpf u l in d ev elo p m ent a s it is t o

att empt to say wh ether foo d o r h eat is m o re es s ent ial t o life .

B o th a re ab solu tely n ec es sa ry and it s u fli c es pres ent p u rpo s es

,

i f t he s t u d ent u nd ers tand s that ext erna l c o nd it io n s ev en to t he ,

matt er o f ex erc is e a re f u nda men t a lly es sen tia l t o f u ll d ev elo p

,

ment an d that t he limit ing elem ent

,
“
m ay b e fo u nd ext erna l
t o t he o rga n is m a s well as int erna l We s ha ll n o t b e able t o
.

as sert ho w m u ch is d u e to each sepa ra t ely n o r can we d et erm in e

,

t he c oefl i c ien t t o b e a ss ign ed t o u s e and t o d is u se b u t we a re ,

s afe in res t ing a s s u red that fo r m any part s d ev elop m en t is f a i r ly

p pr o o r t i o n a l t o ex erc i s e,
at leas t wi th i n t he li m i t s o f i nh er i tanc e .
2 88 C A U S ES OF V A R I AT I O N

Disap pearin g d isappearanc e of leg s fro m s nak es

or gan s . T he
and fro m whales t he lessen ing o f the fo re ar m s of the kanga roo
, ,

and o f t he wing o f c ertain bird s the lo s s of t oes fro m m any ,

m a mm als and ru d i m entary c o nd it io n s gen era lly arg u e fo r t he

, ,

grad u al d isappea ranc e o f a part that is n o lo nger u s ef u l and no

lo nger u sed .

T he fi rs t s tages o f th is d is appeara nc e can b e u nd ers t ood as

aris ing t hrou g h t he ces sa tion of s elec tion and t he res u lt ing pan
m ix ia by wh ic h inh eri tanc e is fro m t he g en era l a v er ag e o f the
1
,

wh o le ra c e in s t ead of fro m a s elec ted lo t as h eret o fo re r es u lt

, , ,

in g n ec es s arily in d egen erat io n a s c o m pared wit h a s ta ndard

s u s ta i n ed by rig id s elect io n L at er s tages m ay b e exp la in ed by .

rev ers al o f select io n wh en t he h ith erto u sefu l o rgan has n o t ,

o n ly bec o m e u s eles s b u t in so m e way d et ri m enta l T h is wo u ld .

acco u nt fo r s t ill f u rth er d egeneracy and is as far as the princ i ,

p le o f u s e and d is u s e app lies

?

S pac e cann o t b e tak en h ere fo r t he en u m erat io n o f i n s tanc es

s h o wing t he eff ect s o f u se and d is u s e T hey m ay b e f o u nd o n .

ev ery hand and th ey ab ou nd in b o o k s o n g en era l ev o lu t io n

?
,

Hypertr ophy U n u s u al en largem ent o f a part is t echni ca lly

.

kno wn as hypertrophy Two kind s a re rec o gn ized f u nct io na l

.
,

h ypertrophy wh en a part is enlarged thro u gh u s e ; and c o m

,

p e n s a t i n g hyp ertr o phy w h ich tak es p lac e when o n e o rgan , ,

bei ng rem ov ed o r bec o m ing f u nct io n les s an o t her en larges ?

,

T he v o lu ntary m u sc les o f t he hand and a r m gro w large

thro u gh h eav y u se b u t t he m u s c les o f the fingers o f a m u s ic ian
,

d o n o t u nd ergo hypertro phy th o u gh t he t o ta l a mo u nt o f wo rk ,

m ay b e v ery large I t is o n ly wh en m u s c u lar wo rk is d o n e aga in s t

.

great res is tanc e that en largement o f t he m u sc les takes p lac e 5

.

1 A term c o in e d by W eis m an n an d d en o t in g
“
u n iv ers a l c ro s s in g,
”
l it l ly
era

a ll m ixe d .
”
S ee W eis m a n n , Es s a y on H ere it d y I, 91, 1 4 1 a sol R o m a n es , D ar
—
,

win an d A f ter D a rwi n , Pa rt I I , p p 2 9 1 3 06 . .

1 ll d
Fo r f u er isc u s s io n o f is a p pea rin g o rga n s , d s ee n ex t c h a p t er .

D a rwin , O rigin o f S pec ies ( s ix t h

3
e d it i on ) pp
, . 1 08 —1 1 2 [D . A pp l et o n 81
y l l
C o m pa n ] a s o A n i m a s a n d P a n t s l un d er D o m es t ic a t io n ( s ec o n d dit i
e on
)
[D A p p l t 8 : C
e on m p y ] o an : in gen e ra l II, 2 8 5 - 2 93 , 3 5 3
4 4 6 ; in ra b b it s , I, 1 29
—
.
, ,

134 ; i d n k 1
uc 99 3 s, , 2 01 .

4 M o rga n R egen era t io n pp

, , . 1 1 5
-
1 18 .

5 Of c o u rs e th e e ff ec t s o f u s e a re n o t l im it d t e o in c reas e o f s ize ; t e h y a re y
f u ll
as n o tic ea b e l in n ic et y of a dj u s t m en t .
2 90 C A U S ES OF V A R I AT I O N

SECT I ON X I EXTER N A L I N FLU E N C ES AS C A U S ES O F

V A R I AT I ON I N T Y PE
T he t d ent m u s t d is t ing u is h c learly bet ween the infl u enc e
s u

o f ext erna l c o nd it io n s u p o n an o ccas io na l ind iv id u al and th ei r

eff ect u p o n t he typ e o f t h e rac e T h ere a re three p o s s ib le way s

.

in wh ic h t he en v iro n m ent m ay res u lt in a m o d ificat i o n of type :

1 ) by a ff ect ing a ll i nd iv id u a ls in t he s a m e way ; ( 2 ) by s elec
(
t io n ; ( 3 ) by t he inh eritanc e o f t he m o d ificat io n s d ue to c o nd i
t io n s o f life I t rema in s to ex a m in e each s o mewhat caref u lly
. .

A ll in div idu al s affect ed in t h e sam e m ann er , t h u s in flu encin g

t h e type di r ec l y t The m o d ify ing eff ect s o f t he c o nd it io n s o f life
.

ha v e been q u it e f u lly n o ted I f b u t few ind iv id u als a re a ffect ed

.
,

it is man if es t that t he type will n o t b e s er io u s ly changed ; b u t

if o n t he o th er hand ev ery i nd iv id u a l is a ffect ed and in t he
, , ,

s a m e way th en t he typ e is t o that ext ent d u e t o t he c o nd it io n s

,

o f lif e .

Fo r exa m p le S ize is d irect ly in flu enc ed by t he f o o d s u pp ly

, ,

and increas e o f S ize in a rac e c o nt em p o ran eo u s with a bett er

,

fo o d s u pply may fa irly b e attrib u ted to the fav o rab le infl uenc e
,

o f f u ll f eed act ing u p o n a ll i nd iv id u a ls a lik e .

S ize is a ls o in her it ed s o that t he lim it s o f d ev elo p m ent a re

,

d u e t o t wo i nfl u enc es act ing t o geth er — inh eri tanc e and foo d

,

s u pp ly .I t is o ft en exc eed ing ly d iffic u lt t o d et er m in e ho w m u ch

to attr ib u t e to t he o n e infl u enc e and ho w m u ch t o t he o th er .

What is tr u e o f S ize i n th is res pect is tr u e o f ev ery o th er char

act er t hat is in t he s lightes t d egree d epend ent u po n en v iro n ment
fo r it s d ev elo p m ent A cc o rd ing ly m u ch u nc ert a i nty prev ai ls as
.

t o t he c o m parat iv e i nfl u enc e o f inh eri tanc e and en v iro n m ent .

T he rac ia l type can b e d et er m i n ed o n ly by t he s t u dy o f t he ind i

v id u a ls c o n s t it u t ing t he m a tu r e p o p u la t io n ; b u t th ei r d ev elo p

ment is t h e r ut
e s l o f t o s et s o f ca u s es
w — t he o n e o f h ered i ty
, ,

t he o th er o f en v i ro n m ent bo th c o ntr ib u t ing to t he s a me effect

,

and bo th c o nt in u ou s t h ro u gh life .

U nd er t he o ld V iew ev ery ind iv id u al wa s regard ed a s t he t e

s u lt no t o n ly o f what wa s b o rn int o i t b u t a ls o O f t he d irect

infl u enc e o f its en v i ro n m ent I nd iv id u a ls c o n s t it u t e the type

.
,

and so i t is that when t he c o nd it io n s of life a ffect a ll ind iv id u als

 EXTER N A L I N FLU EN C ES AS CAU SES OF V A R I ATI ON 2 91

a lik e th ey n ec es sarily infl u enc e t he type of t he rac e b u t t o ,

what ext ent cann o t b e t o ld with o u t s o me m eth o d o f s u btra ct ing

t he no rm al res u lt s o f s im p le i nh eri tanc e N o meth od of d o ing
.

th is has ev er been dis c ov ered and m u ch u nc ertainty has alway s

,

prev ailed as to what pro po rt io n of ex is t ing types s h o u ld b e

cred it ed to t he c o nd it io n s of life .

The p uzzli ng pro b lem is great ly S im p li fi ed if we enlarge t he

meani ng o f t he wo rd “
inh eri tanc e t o c o v er n o t o n ly t he lin es
”

o f po ss ib le d ev elo p ment b u t t he f u ll capac i ty a s well I n th is

.

v iew o f t he cas e the i nd iv id u al and t he rac e a re u nd er s t oo d

as po ss es s ing h ered i t a ry capac i t ies fo r d ev elo p m ent far bey o nd

that fo r which o ppo rt u n it ies a re lik ely ev er t o b e a ffo rd ed by

en v i r o n ment . T h is largely remov es t he c o nd it io n s o f life fro m
am o ng t he f u n d a m en ta l ca u s es o f v ar iab ility and relegat es th em

t o t he r eal m o f pa s siv e and pe rm is siv e th o u gh n ec ess ary req u i

, ,

s it es fo r t he f u ll d is p lay o f h ered i tary po wer I t t end s s tro ng ly

.
,

too to regard m o s t ind iv id u a ls as so m eth i ng less than wo u ld

,

b e ind icat ed by th eir h ered i tary po s s ib ilit ies and to c o n s id er

,

en v i ro n m ent in gen era l as t he lim it ing n o t t he s t im u lat ing

‘

, ,

elem ent in d ev elo p m ent With th is v iew the writer is inc lined
.

t o agr ee th ou gh i t n ec ess a rily red u c es t he ext ent o f d i rec t

,

influ enc e o f t he en v i ro n m ent . T he s t u d ent is n ev er to fo rget

that whatev er may b e t he infl u enc e o f su rro u nd ing c o nd it io n s
,

u p o n o n e f o rm o f lif e t he s a m e infl u enc es a ffect o th er s pec ies

,

d ifferent ly th u s S h o wing that th eir cha racteris t ic eff ect is c on

,

d it io n ed u po n t he p ower o f t he o rgan is m t o react a c o nd it io n

,

t hat is em inent ly internal and inheritable .

Sel ecti on as a cau se of v ariat ion in ty pe Ma n y i nd iv id u a ls

.

a re u nab le t o m eet the c o nd it io n s o f lif e with wh ich th ey find

them selv es s u rro u nd ed and in the att empt bec o me ext inct
,
.

Th is is s electio n and it is m an ifes t that the prev a iling typ e o f

,

the rac e as it ex is t s at any m o m ent bei ng m ad e u p o f select ed

,

ind iv id u al s is so meth ing d zf eren t fro m that wh ich was bo rn

,

int o t he rac e I t is a ls o m an if es t that o n ly a li m i t ed p o rt io n o f

.

ad u lt s wi ll reprod u c e s o that select io n throu gh t he ext erna l c o n

,

dit io n s o f life ex ert s a s tro ng infl u enc e u p o n type .

Select io n is th eref o re o n e o f t he m o s t po werf u l — if n o t the

—
mos t powerf u l in flu en c es kn o wn in the mo d ificat io n o f s pec ies ,
2 92 C A USES OF VA R I AT I ON

and it may b e f airly s pok en o f as a pri mary ca u s e o f v ariat io n in

type T h is d oes n o t m ak e it h o wev er lik e t em pera t u re o r foo d
.
, ,

s u pp ly a f u nda m en tal ca u s e o f v ar iat io n in liv ing m att er

,
T h ere .

is n o bas is fo r s elect io n u nt il d i ff erenc es ha v e appeared fro m

o th er ca u s es Wh en t he s elect io n is m ad e based o n th es e d iffer
.
,

e n c es i t is c erta in t o a ff ect the type and it is th eref o re a ca u s e

, ,

o f type v ariat io n ; b u t it wa s n o ca u s e o f t he o ri gi n al d i ff erenc es

o n wh ich i t s act io n wa s pr ed icat ed and it is in n o sen se an

,

o rig ina l ca u s e o f v ar iabi lity T he c o nf u s io n o f m ind wh ich ca u ses

.

s elect io n to b e regard ed a s a ca u s e o f v ari at io n ha s ar is en fro m

a failu re to d is t ingu is h c learly bet ween t he ind iv id u al and t he

type 1
— b et ween t he m at eria l o n whic h we can wo rk thro u gh
s elect io n o n t he o n e hand and t he fin is h ed pro d u ct o n t he o th er
,
.

V ariat ion in ty pe t h rou gh t h e in h eri tance of m odifi cat ion s d u e

to en v i ron m en t Wh eth er ind iv id ual mod ifica t io n s d u e to en v iro n
.

m ent ( acq u ired charact ers ) are inh erit ed will b e d is c u s s ed in a

following chapt er I t is the p u rpo s e h ere o n ly t o S h ow t he nat u re
.

o f t he eff ect o f s u ch inh er itan c e in c a s e it d oes occ u r .

I f t he m od ificat io n s d u e t o ext ernal infl u enc es S h o u ld per

c hanc e b e inh erit ed ev en in the s light es t d egree th en t he effect s
, ,

o f m od ifi cat io n wou ld lik e th o s e o f s elect io n b ec o m e c u m u lat iv e

, , ,

and t he type wo u ld t he m o re rap id ly c o nfo rm to t he en v iro n m ent

and the m o r e ra pid ly es tablis h a fi t with t he c ond it io n s o f life .

I t is ev id ent that th is is a d iffic u lt fi eld M er ely thro u gh t he

.

ex igenc ies o f m ai nt enanc e th o s e C h a ract er s will d ev elo p bes t that

ar e m o s t fa v o r ed by t he c o nd it io n s o f lif e th u s br inging abo u t ,

a k ind o f m a s s res po n se to t he exact io n s o f t he en v ir o n m ent .

A ga in by t he pr inc iple o f s elect io n o n ly th o s e in fa ir acc o rd with

, ,

t he en v ir o n m ent will liv e and th is brings abo u t a s t ill c lo s er fit

,
.

I f n o w m od ificat io n s d u e to en v iro n m ent were f u lly and com

, ,

l etely in /zer i ted t he adj u s t m ent to c o n s tant c o nd it io n s wou ld

p ,

s peed ily b ec o m e so exact a n d c o m p let e a s t o lea v e n o ro o m for

select io n .

F ew b iolo gis t s a re bold en o u gh t o c la im th is extrem e d egr ee

o f inh eritanc e o f m o d ific a t io n s d u e t o en v ir o n m ent Many d eny .

it in toto T ha t n eith er extrem e is r ight is c o m parat iv ely ea sy

.

o f a s g oo d proo f a s we a r e g en era lly ab le t o br ing in a ffa ir s b io

logical b u t wh er e b et ween lies t he tr u th it is m o s t d iffic u lt t o

,
2 94 C AU SES OF V A R I ATI ON

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—1 5 8
. .

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U
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s ec o n d , s ho r ta
g e o f po t a s s i u m ) . E x per im en t S ta t io n R ec o rd , X I V ,

E FF EC T OF WATER CONT ENT U PO N D E V E LO PM E N T OF W HE A T , OATS,

C LOV E R , ETC . E x peri men t S ta t io n R ec o rd , X I I I , 1 2 5 —1 2 6 , 44 1 -
631 .

E X PE R I M EN TA L Z O OL O G Y . By J . H . M o rga n . C ha p ters 11 a nd 1 1 1,
pp . 12

I NJ U R IOU S E FF E C T OF FR E E ZING ON D E V E LO P M E N T O F T HE EMB RYO

I N T HE EG G O F T HE HE N . E x peri m en t S ta t i o n R ec o rd , X 1 , 5 7 7 .

V A R I A T I ON OF A N T HR A x B A C I L L L S W HE N E R E D
'
N DER D I FFER ENT U
CO N D IT I ON S . E x per i m en t S ta tio n R ec o rd , X I V , 2 93 , 9 1 6 .

V A R I AT IO N IN N I T R OG E N C O N T E N T OF W HE A T . E xperi m en t S ta t io n
R ec o rd , X I I I , 4 5 1 .

V A R I A TI ON I N T U B ER C I . E B A C I L L I W HE N FO UND IN D I FF E R E N T E N Y I
R ON M E NT . E x perim en t S ta tio n R ec o rd , X I V , 1 12 1 XV, 1 88 .

V A R I AT ION S CA U SED BY FE R T I L I Z AT ION . E x peri m en t S ta tio n R ec o rd ,

X I V . 34 7 .

1 h
T is e xc e ll en t v o u m e l was not
yet o ff t h e p re s s w h en t hi
s co
p yw as re
p p a re d .

I t is es p ec ia lly rec o m m en d d e .
 CH A PTE R X
R ELATIV E STABILI TY AND IN STABILITY OF LIVING MATTER

In o rd er gu id e the st ud ent of breed ing in fo rm ing his c o n

to
c ept io n s as t o what ma and w hat m a n o t b e acc o m p lis h ed in
y y
t he wa y o f m od ifying t he fo rm o r f u nct io n o f d o m es t icat ed an i
m a ls and plant s ev eryth ing is v a lu ab le wh ich thro ws light u po n
,

t he d eg r ee of fi x ed n ess in liv ing m att er ; that is to s ay in t he ,

relat io n s that happen to hav e beco m e es tablish ed bet ween the

es s ent ial charact er s o f ex is t ing S pec ies .

Wh en t he s t u d ent fo r a t im e best o ws caref u l st udy u po n v aria

t ion an d c omes t o r ea lize ho w rad ical are som e o f t he d epart u res
from type and how s weeping a re s ome o f the d ev iat io n s from the
no r mal he is led to feel in s t inct iv ely that liv ing matt er ex is t s in
,

a s tat e of extrem e in s tability as rega rd s bo th fo rm and f u nct io n ,

an d t ha t alm o s t anyth ing is lik ely to h a ppen .

Wh en h owev er he co n s id er s that thro u gh it all d is t inct types

, , , ,

are pr es erv ed and wh en he n o t es t he s ing u lar per s is t ence of c er

ta in cha ract er s thro u gh all the v ic iss it ud es o f t im e and ev olu t io n ,

rea ppea ring gen erat io n aft er gen erat io n wh en th ey were s u ppo sed
to ha v e been lo ng los t and in m any ca s es linger ing aft er th eir
,

u sef u ln es s is pa s t and a ss o c iat ed charact er s ha v e been b lo tt ed

ou t,
— wh en he c o n s id er s all th is t he carefu l s t ud ent will realize
,

t ha t stability and in s t ability are relat iv e t erm s and he will begin

,

serio u s ly to i nq u ir e int o t he d egr ee o f s tab ility o f t he v ar io u s plan s

u po n wh ich m att er ha s been o rgan ized and v italized . I t is th er e

l

fore profitable to in q u ire som ewhat f u lly int o the relat iv e s tab il
ity or in s tab ility o f th o s e c o m po u nd s that ar e end o wed with life ,

an d in t o th eir r elat iv e a b ility t o m ainta in th eir int egr ity and d is

charge th eir fu nct ion s u nd er c o nd it io n s bo th n o r m al and abn o r m al .

Th e u t ility o f th is inq u iry res t s in t he light it m ay throw u po n t he

ext ent t o wh ich char act er s that ha v e b ec o m e typica l a re fix ed and

u nc han geable and t he ext ent t o wh ich th ey m a b e m od ifi ed

, y .

1 95
2 96 C A U SES O F V A R I A T I O N

S EC T I O N I — EV I D EN C E F R O M ST A B I L I T Y O F TY PE

A lt ho u g h no t wo ind iv id u a ls a r e a li ke . and a lt ho u
g h a gi v en
C ha ra cter d i ff er s g rea t ly in d iffe rent m em b e rs o f t he ra c e yet ,

th ere is a s pec ific t v pe th a t is a lways and ev e ryw h ere prese nt ;

th o u g h t he elem ent s a re e xc eed in gly v a ria b le yet t he res u lt an t ,

c o m p s it e is rem a rk a b ly c o n s tant as c o m pa red wit h o t h er t ypes

o .

I n o t h e r wo rd s when we c o m pa re h o rs es wi t h h o rs e s we a re
,

i m pres s ed with t he fa ct o f v a r ia b ili tv b u t wh en we c o m pa re ,

h o rs e s with c a t t le o r ev en wit h a s ses th en we a re led t o m a rv el

, ,

at t he fi xit y a n d pe rs is t enc e o f type I n a ll it s v a riat io ns the .

,

h o rse is s t ill c lea rly a h o rs e W id e and pro fo u nd as is v a riability

.
,

it is yet well with in lim it s and c er ta in types c o nt i n u e with

,

s ingu la r pe r s is t enc e .

T he Hu bb a rd s q u as h and t he M o rga n h o rs e a re goo d ex am

p les o f per s is t enc e o f type With a ll t he v a riab ility o f t he C u
.

c u r b ita c ea t he H u bbard s q u as h pe rs is t s d is t inct i n type and

, ,

q u a lity I t m ix es freely wi th o th er types b u t it s ch a ract ers a re

.
,

e v id ent ev en th en and i t po s s es s es a s i ngu lar ab ility t o free i t self

,

fro m s u ch ad m ixt u r es and ret u rn a gain to t he o riginal .

T he Mo rgan h o r s e is a breed es tab lis h ed by a S ing le ani m al ,

and yet a h u nd red yea rs aft er t he d ea th o f J u s t in Mo rga n wh en

, ,

t he pe r c ent o f his b loo d is o f n ec es s ity s light t he M o rga n ,

C ha ract er s s t ill s t and o u t c learly c o n s t it u t ing a ty pe alm o s t as

,

d is t inct as that o f any ex is t ing breed Th is is the so litary kno wn .

in s t a nc e o f t he f o u nd i ng o f a breed by a s in g le anc es t o r It .

illu s trat es in a pec u liar wa y t he o cca sio na l extreme per s is t enc e

o f a type o nc e f o r m ed a n d is in m ark ed c o ntras t to t he read in es s
,

with wh ich o th er types break u p and d is appea r .

The o ld t ime pe rs is t enc e o f t he s lo p ing ru m p i n t he Berk s hi re

-
,

o f t he narro w C h es t in t he Po land C h in a o f lack o f d epth b eh in d

-
,

in t he po ny b u ilt Heref o rd o f d efic ient cro p s in t he S ho rt ho m

-
, ,

th es e and S i m ila r d efect s that m ight b e m ent io n ed illu s trat e

the s trength wi th wh ich c e rt a in ch a ra ct er s c o nt in u e ev en in t he ,

fac e o f t he mo s t po werf u l o ppo s itio n and arg u e s tro ngly for ,

s tab ili ty o f typ e .

I t is a ls o a gen era l f a ct th a t s pec ies h o ld th ei r t ypes with

es s ent ia l s u cc es s u nd er a g reat v a r iety o f c o nd it io n s bo t h ,
2 98 C A U S ES O F V A R I AT I O N

SEC TI O N I I — EV I D EN C E FR O M M U T A B I L I TY
OF S PEC I ES

S pec ies d o h o wev er alwa y s rem ain u nchanged On the

no t , ,
.

c o ntrary th ey freq u ent ly exhi b it a pro gres s iv e d ev elo p m ent

,

tru ly m arv elo u s Ho rs es fo r exa m p le a re t rac eab le back ward

.
, ,

by eas y s tages and well d efi n ed c o nnect ing link s t o a t ime far -

beyo nd t he appea ra nc e o f m a n u po n t he earth t he lin e end ing ,

FIG 3 1 . . C o m pa ra t i v e d ra win gs o f s k llu s, feet , an d t eet h of p hi

re s t o ric h o rs e
s ,

s h o win g l
ev o u t io n a r y d l
e v e o p m en t . R ep ro d uc e d b y p erm iss io n fro m
O rigin an d H is to r y of t he H o rs e b y H F Os b o rn . .

in o n e of t he earlies t m a m m als ,
a t o ed creat u re n o t litt le fi v e -

m u ch larger than t he d o m es t ic cat .

N o les s than t welv e s tages in th is ev o lu t io n a re well kno wn ,

and repres ented by s pec im en s m o re o r les s c o m plete 1

1W il li m D M tth w f t h A m i
a . a M e m f N t
, o l Hi t y t i l
e er c a n u s eu o a u ra s or , ar c e

H th E
o rs e, l ti f i E y l p d i A m i
e vo u on o ,Th i i
”
n f th b t
nc c o a a er c a n a . s s one o e es

and f th
on e o w t t f th d
e ne es l pm t f t h h
ac c o u n s o di h b e ev e o en o e o rs e, an s C o s en e

c ause o f it ib ilit y d li b il it y T h
s ac c es s tl i an
gi w h il t m
re k d b
a
y . e ou ne v en , e no ar e

q t
uo a t i i
ons,
p t i l ly bst t
rac f t h
ca f an a s rac o e r e eren c e .
 R E LA T I V E ST A BI LI T Y OF LIV I NG M ATT ER 2 99

1 . Hy m cotlzer i u m . S k u lln ly F o u nd in Lo nd o n c lay o f

o .

t he Lo we r Eo c ene ( earlies t m a m ma ls ) S pec im en i n B ri t i

.s h
Mu s eu m .

2 . E o/zipp u s . M u ch
better kn o wn c o m ing fro m the Lo wer ,

Eo c en e o f Wyo m ing and N ew Mex ic o Teeth lik e t he f o rm er ; .

fo u r t oes o n the fro nt foo t with a s plint of the fifth three t oes ,

beh ind with a s plint o f ano th er s h o wing that c o n s id erable

, ,

d epart u re had already tak en p lac e fro m its ev id ent fiv e t oed -

anc es t ry Height of an im al 1 2 to 1 6 inches

. .

3 . P ro t o r ol pp
u u s ( Wy o m i ng F o u r c o m ,
p le t e t o e s i n

fro nt and three beh ind ; no s p lint s ; s kelet o n o f the s ize o f a

s m all d o g .D es c ribed by C o pe as the fo u r t o ed h o rs e -
.

4 . O r o l n
pp u s O n ly .part s o f ja w s and t ee th b u t th e s e S h o w ,

s o me ad v anc e S pec im en at Y ale U n iv ers ity

. .

5 .E p pp
i lz z us
( U pp e r E o c e n e N e w W o r ld ) O n l
,
y i nc o m p l et e .

s pec im en s f ou nd b u t m u ch t i me has elap sed and c o p s id era b le

,

d ev elo p ment is no t ed es pec ially in t he t eeth The t oes a re

,
.

s t ill f o u r and three b u t the c ent ra l t oe is bec o ming m u ch larger

than t he S id e t oes .

Co llat era l branch es o f the S a m e perio d in t he Old Wo rld

( P a leo t h e r i u m and P a lo p lo t heri u m ) had three t oes o f n early

eq u a l S ize o n ea ch f oo t T h ey were v ery ab u ndant at th is t im e

.

b u t s eem to ha v e b ec o m e ext inct — a fat e that o v ert o o k m o s t

,

o f t he branch es o f th is f ert ile and pro gres s iv e s t em .

6 M es a/ upp us ( W h it e R iv er F o r m at io n ) T hree t oes o n each

. .

fo o t the mi dd le m u ch t he larges t t he S id e t o es bea ring litt le

, ,

weight . By th is t im e t he an ima l ranges in s ize fro m t he c o y o t e

t o t he s h eep and t he m o lar t eeth a re well d ev elo p ed
,
A ll part s .

o f t he s k elet o n kn o wn Fifth t o e represent ed by a s p lint

. .

7 .A nc /z i tlz er i u m ( L o w e r M io c en e) F ou nd b o th i n E u ro.p e

and in A merica M u ch lik e its pred ec ess o r b u t larger with

—
.
, ,

bett er d ev elo ped t eeth May b e o n e o f t he s id e branch es .

“

rath er than in t he d irect lin e of t he m od ern h o rse .

8 Hypo/ upp u s and P a ra / upp u s A c o m p let e S k elet o n o f Hypo

. .

h ippu s was fo u nd 1 90 1 by t he Wh itn ey Exped it io n n ear

, ,

Pawn ee B u tt e Co lo rad o ,
I n t he f o ref oo t s m a ll ru d iment s s t ill
.

repres ent t he fi rs t and fifth t oes b u t t he s p lint s a re go n e ; ,

t he sec o nd and fo u rth d igit s s t ill t o u ch t he gro u nd th o u gh ,

FI G .
3 2 . Pro gr ess i v e ev o lu t io n in th e h o rs e th e lo wer fi g u re is a f u ll s ize
-
dm d lo e

of the t ip p u s in c o m p a ris o n wi t t he sh k ll f th m d h
u o e o ern o rs e, s h wi g
o n

h
t a t t he s k ll
u l
o f t he a t t er h l
o rs e is a rge r t h t h t i b dy f it
an e en re o o s an c es t o r.

F ro m s p ec i m en in A m eri c a n M u s eu m f N t
o l Hi t y N
a u ra s or , e w Y o rk .

C o u rt es y of D irec t o r H F Os b o rn
. .
 02 C AU SES O F V A R I AT I O N
3

ab ru pt en d The mod ern h o rs e h o wev er aft er his lo ng and

.
, ,

t o rt uous ev o lu t io n seems d es t ined fo r a pro lo nged and n o table ,

ex is t enc e D u ri ng the p ro gres s iv e chang es in t he f eet t he

.

leg has been great ly length en ed the j o int s m o d ifi ed fro m t he ,

loo s e ball and so ck et t o t he fi rm er h inge j o int and t he t eeth

“

hav e bec o me exc eed ingly serv ic eab le I t is a n o t ic eable fact .

that t he po wer of a s pec ies to with s tand t he v ic iss it u d es o f ex

treme lap s es o f t ime d epend s v ery largely u po n t he ab ility o f
it s feet it s leg s and it s t eeth t o u nd ergo m o d ificat io n
, ,
I t has .

been s a id that the elephant ha s S I ICCeed ed in m aintain ing him

self to t he pres ent in s p it e o f h is f eet and by v irt u e o f his exc el ,

lent t eeth and his remarkab le pro b os c is .

I t is n o t ic eab le that the larger part o f th is ev o lu t io na ry h is

t o ry of t he h o rse has been wo rked o u t fro m s pec im en s fo u nd in
wes t ern A meri ca b u t n o o n e believ es that t he m o d ern h o rs e
1
,

is an A merican a ni ma l T h is ev o lu t io n seem s t o hav e pro c eed ed .

U po n s u b s tant ially parallel lin es in the eas t ern and t he wes t ern

c o nt inent s wh ich were d u ring its pro gress u n it ed by a bro ad

, , ,
‘

s trip o f land in t h e reg io n o f A la s ka ; b u t so meth ing s eem s t o

ha v e happened to the A merican branch an d it is believ ed tha t ,

we a re ind ebted to the Eu ro pean and A s iat ic branch fo r t he h o rs e

o f the pres ent .

I nd eed S o u th A m er ica is repres ent ed by a fos s il fo r m ( Hip

,

p id iu m ) w h o s e f ee t r e s em b led E q u u s e xc ept that th ey w ere ,

s h o rt and s t o u t I t s t eeth res em b led th o s e of Plio h ipp u s ( Mu s eo

.

N a c io na l B u en o s A yres ) T h is f o r m had ev id ent ly ad v anc ed

,
.

n early to that of t he present b u t peris h ed in the gen era l d is as t er , ,

what ev er it was that o v ert o o k the A merican h o rs e fo r it left

, ,

n o d es c endant s that p er s is t ed u nt il h is t o ric t imes .

Ca u ses of t h e ev olu ti on of t h e h orse A s is rem ark ed by .

M atth ew the ev o lu t io n of t he h o rs e adapt ing it to liv e o n the

, ,

d ry plains pro bably went hand in han d with t he ev olu t io n of

,

the p lai n s th ems elv es A t the c o m menc ement o f m a m malia n

”
.

O1 k wl dg f th
ur no l ti f t h h
e e o i l g ly d th
e ev o u
t i d f t on o e o rs e s ar e ue o e n e a

ig b l l b
a e f P f
a H
o rs o
y F O b D i
ro ess o rt f t h Am i M
en r m . s o rn , rec o r o e er c an u s eu

o f N t l Il i t y
a u ra d t t h m g ifi
s or t g, it y f th l t W illi m C
an o e a n c en en ero s o e a e a .

W h it y t hne gh wh i h
, t i
ro u
p l t i d igc ifi t d i i
ex en s v e exw o ra o n s an s n ca n s c o v er es ere

m d i W y mi g
a e n d th o
gi n f w t an Am i Th
o t d
er re t f thi ons o es ern er c a . e s u en o s

su b j t wil l
ec g l y w it Pea f O b m
er f th
a m i g f ll p t
a ro es s o r s o
’
s or co n u re or .
 0
3 4 C AU SES O F V A R I ATI O N

lif e t he M iss is s ipp i alley was j u s t em erging fro m t he G u lf o f

v

Mex ic o and t he pla in s o f wes t ern Eu ro pe a n d A s ia were lo w

,

and wet T he c lim at e was m o is t a n d tro p ica l s t im u lat ing d en se

.
,

and lu x u riant gr o wth of giant v egetat io n ev en as far no rth as

Green la nd With t he Tert iary ca m e a gen era l elev at io n u s h er
.
,

in g in a c o m parat iv ely c o ld d ry c lim ate fa v o rab le t o gras s es , ,

and t he ha rd er v egetat io n generally W ith th is cam e gras s y .

pla in s and the ev o lu t io n of rac es with goo d t eeth and exc ellent
f eet and legs fitt ing th em t o a lif e in t he o pen
,
.

Wit h th es e pro fo u nd changes in nat u re o th er fo r m s u nd er

went a d ev elo p m ent s imilar t o that o f t he anc es t o rs and o th er
relat iv es of t he h o rse Many o f th es e as ou r catt le S heep s wine
.
, , , ,

et c . d ev elo ped a t wo t oed foo t and s o m e as t he rh inoc eros

,
-
, , ,

s t o pped at t he three t o ed s tage b u t n o n e o f th em beca m e s o

-
,

h igh ly s pec ialized as t he h o rs e .

Here was a great lin e o f d es c ent c o nt in u ing alm o s t fo r ages , ,

and t erm inat ing in m any h igh ly s pec ia lized s pec ies that a re s t ill
fl ex ible B u t it gav e rise o n t he way d o wn (o r u p ) to m any kno wn
.
,

and d o u bt less t o many u nkno wn branch es that becam e ext inct ,

thro u gh so me gen era l d isas t er o r m o re lik ely b ecau s e o f th eir , , ,

inh erent inab ili ty t o d ev elo p a ll t he charact eris t ic s n ec es s a ry to

meet changi ng c o nd it io n s Fo r exa m p le as t he t eeth d ev elo ped .
,

i nt o m o lars fi tt ed fo r grind ing t he ev er hard en ing f o rage s o m e -

,

s pec ies s ecret ed c em ent in t he v a lley s th u s s u ppo rt ing the hard

and gri nd ing ridges ; o th ers d id n o t and it is S ign ificant that in ,

t he latt er ca s e n o s pec ies end u red 1

T he elep han t a lo n e o f his .
,

k ind has persis t ed to t he pres ent and if th is is beca u s e o f his

, ,

t eeth and in s p it e o f body and f eet wh ich a re ill adapt ed to

, ,

m o d ern c o nd it io n s it s er v es t o S h o w o n ho w S lend er a thread

,

t he life o f a s p ec ies o ft en hangs .

Pres ent ex is t i ng land s pec ies a re t o b e rega rd ed as repres ent ing

lin es o f d es c ent nat u ra lly end o wed with an u n u s u ally h igh d egree
o f fl ex ib ility a ll t he m o r e s tab le and les s adaptab le f o rm s ha v ing

peris h ed o ff t he earth in the lo ng s t ru ggle to k eep u p with t he

1 I t i w thy f s m k t h t th
or o t
re l pl i
ar f S th A m i
a m t
e c en ra a ns o ou er c a s ee o

h d lpd h
av e ev e o l ik im l ( L i t pt ) l i g it l t l t
e a o rs e- e an a dd l p
o er n a , os n s a era o es a n ev e o

i g th h i g j i t
n e d l g th
n e d li m b ; b t it
o n an en d l p d m t i it
en e u n ev er ev e o e ce en n s

g i
r n d wh
ers ,i h m i d i f i
c re d w a ne t i d
n er o r,
p t h t i t li
an b m e a re n o sur r se a s ne ec a e

ex t in c t .
 06 C A U SES O F V A R I AT IO N

D u ringth is t ime th ere has been (s u p posed ly ) n o ad mixt u re

with d o m es t icated h erd s and yet th ere appears o ccas io nally , ,

ev en in a D ev o n h erd a whit e calf wh o se ea r s lo wer legs and

, , ,

bru s h o f ta il a re m ark ed with t he tawny red o r bro wn of the

wild anc es t o r and wh o se m att ed c u rly ha i r u ps ta nd ing h o rn s
, , , ,

and pec u liar fac ial expres s io n bes peak his rev ers io n t o t he
ear ly type .

T h is s ingu lar pers is t enc e o f cha ract ers o nc e typ ica l arg ues
s tr o ngly fo r s tab ility if c o n s id ered fro m t he s tand po in t o f t he

anc ient cha ract er b u t it s peak s n o t les s plai n ly fo r ins tab ili ty if
,

c o ns id ered fro m the s ta nd po int of t he n ew ( present ) type .

T he v er m if o r m append ix t he pers is t enc e of t he ta il in mos t ,

m a m ma ls — th ese and sc o res o f S imila r in s ta nc es att es t t he

,

s t u b b o rn res is tanc e o f a s tr u ct u ral part to t he ext inct io n that is

in ev itab le ; and t he cas e o f t he “

hea rd of t he t u rk ey c o ck
”

illu s trat es t he fact that a c o m b inat io n of what ev er o rd er o nc e ,

s ta rt ed t end s s tro ngly t o c o nt in u e ev en th ou gh u seless an d

, ,

u n m ea n ing .

S EC TI O N IV — EV I D E N C E FR O M D I SA PPEA R A N C E
O F PA R TS

T he rga nic wo rld is f u ll o f in s tanc es of s tru ct u ral pa rts

o

lingerin g lo ng aft er th ei r u s ef u ln es s ha s largely o r q u it e d isa p

p e a r ed and aft
,
er ent i rely n ew relat io n s ha v e been es tab lis h ed

a mo ng as soc iat ed cha ract ers ?

T he h ind legs o f t he pyth o n and t he wha le already ru d im en ,

ta ry and represent ed o nly by bo nes in t ernal t o t he s u rfac e and ,

th ose o f t he sea lio n and t he seal ev id ent ly d isa ppearin g in t he ,

s a me ma nner ; t he wi ng o f t he apt eryx r ed u c ed to t he m eres t

'

t rac e h idd en in t he plu m age and that o f t he o s trich p lai n ly fol , ,

lo wing a lo ng to t he sa m e fat e ; t he feta l ha ir o f t he wha le and ,

1 h
T is is en t irel yi d
n ep en de n t of t he q u es t io n o f t he in fl u en c e o f u t il it yi n t he

o rigin an d d ev elo p m en t of a n e h
w c a rac t er I t has b een t he fas io n t o ass u m e . h
h h
t a t n o n e b u t u s efu l c arac ters wi l o rigin a te l
T h e wri ter o n t h e c o n t ra ry, in c lin es
.
,

l h h h
t o t h e b e ief t a t a n y c arac ter wil l a ris e w o s e e em en ts a re p res en t in t h e o rgan l
is m , q u it e i rres pec tiv e o f it s us ef u ln ess an d t a t it wi c o n t in u e u n less p rev en ted
, h ll
l h h ly l
b y se ec tio n , a lt o u g ma n if es t it wil n ev e r a t ta in ma xi m u m p ro min en c e exc ept
h h
t ro u g t he c u m u la ti v e effec t o f t h e s elec t iv e p roc ess .
 R E LAT I V E STAB I LIT Y OF L I V I N G MA T TER
3 7
0

its f u nct io n less t eeth red u ced to t he primi t iv e c o n ical fo rm yet ,

oft en acc o m pan ied by r u d imentary s u c c ess io n als p int

1
; t he s l
bo nes of h o rses s ilent wi tnesses o f what has befallen thei r n eigh
,

b o r d igi t s and s t em proph et s o f th ei r o wn ext i nctio n t he ru d i

,

m enta ry m u s c les o f t he sc alp and t he ea rs in m an th ese and ,

nu m berless s im ilar v es t iges of the pas t remind t he s t u d ent of the

tenac ity with wh ich v ital act iv it ies pers is t when o nc e es tablis h ed ,

prod u c ing a ru d im enta ry leg o r wing o r lesser part fo r in d efi , ,

n ite generatio ns aft er its u sef u lness has c eas ed and all select ion
has d isa ppeared .

SEC TI ON V — EV I D EN C E FR OM T HE D I R EC T A CTI ON
OF T H E E NV I R ON M ENT

A ll t he d nce goes to S ho w that t he c o nd it io n s o f life ex ert

ev i e

a d irect and po werf u l act io n u po n the ext ent o f d ev elop ment

and the n at u re o f t he f u nct io n s o f liv i ng m att er We hav e o nly .

t o reca ll the cha ract eris t ic react io n s o f c erta in ch em ic al s u b

s tanc es u po n pro t o p la s m t he infl u enc e of light gra v ity and
, , ,

t em perat u re and the effect o f ev en so slight a c irc u ms tanc e as

,

c o ntact to realize that liv ing m atter is exc eed ingly d epend ent
,

u po n it s env i ro n ment I f we s h o u ld f o rget fo r t he mo ment that

.

rac ial li n es are preser v ed d is t inct in s p it e of the en v iro n men t ,

we might read ily c o me to believ e with t he L a m arck ian s that the

c o nd it io n s o f life are the s u preme fact o rs in d i rect ing ev olu t io n
and in fi x ing t he type .

To ent e r f u lly int o t he d isc u s s io n of th is phas e of the q u es t io n

at th is po int wo u ld b e to repeat what has been s a id as to ext er
n al ca u s es o f v ariat io n o r t o ext end t he ex a m ples c it ed and
, ,

s pac e can b e a ff o rd ed for n ei th er The s t u d ent s attent io n is

’
.

in v i t ed t o th is pha se of t he q u es t io n h owev er fo r he s h o u ld , ,

acqu ire a j u s t and reas o nable c o nc ept io n of the relat iv e s tability

o f liv ing m att er as s h o wn by t he ab u ndanc e o f t he c lass of ev i

denc e here hinted at rath er than fu lly explo it ed .

1 h h d d
The fac t t a t s u c c ess io n al teet a re fo rm e in a ru im en ta r s t at e a rgu es fo ry
t he h h l h h h
fac t t at t he t eet p res en t a re th e fi rs t o r m i k t eet , w ic , b ein g fu nc tio n less ,
ar e no t d h d d y
isc a rge an d rep lac e as in mam ma ls gen era ll , b u t , o n the c o n t ra r , y
p ers is t th gh
ro u lif e
.
3 0 8 C A U SES OF V A R I AT I O N

SECT I ON V I —EV I DE N C E FR OM A C C LI M AT IZ AT I ON

Co n s id erab le light is thro wn u po n the relat iv e s tab ility of

liv ing m att er and t he ext ent t o wh ich it res p o nd s t o external
infl u enc es by t he ph eno mena o f acc limat izat io n and t he k indred
ph en o mena of im m u n ity .

A ccli ma t iza t ion t o ch emic al s , especial l y poison s .

1
I t is c lear
that t he pro t o p las m o f d iff erent o rga n is ms is d is s imilar W e .

s ee th is in t he d iff erent r eact io n s to t he s a m e C h em ical agent .

N o t o n ly is t he react io n o f t he v ario u s s p ec ies u n lik e b u t in ,

d iv id u als o f the s a me s pec ies fro m d ifferent lo calit ies d iff er

wid ely 1 ”
.

The c o m m o n po iso n iv y pro d u c es extreme i rritat io n with m os t

perso n s — t he effect b eing rec u rrent fo r m any years
,
wh ile ,

with o th ers i t is ent i rely inn o c u o u s T o ev en t he m o s t v i r u .

lent inf ect io u s d is eas es m any ind iv id u a ls a re ent i rely im m u ne ,

and with nearly a ll d iseases of th is c las s a s ingle attack res u lt s

in immu n ity fo r life thro u gh s o m e k ind o f acc limat izat io n ,
In .

m u ch the s a m e m ann er in d iv id u als bec o m e acc u s t o m ed to t he

s t ing o f b ees a n d exp erienc e litt le inc o n v en ienc e fr o m what
,

wo u ld hav e ca u s ed int en se s u ffering t o mo s t o th ers o r t o th em ,

selv es b efo re acc lim at izat io n Phy s ic ia n s find i t n ec es s ary t o

.

change th ei r remed ies freq u ent ly b eca u se th ey s o o n lo se th eir

cha racter is t ic effect u po n the pat ient .

I n lo wer o rgan is m s t he s a m e ph en o m en o n is n o t ic eab le T h u s .

,

“
few bact eria can res is t 1 per c ent o f N aaC Og a n d ev en t he ,

extrem ely res is tant A s car is liv es o n ly [fro m ]fi v e t o S ix h o u rs i n a

p e r c ent so lu t io n o f th i s s a lt b u t L o e w h a s f o u nd i n
“

Owen s L ak e C a lifo rn ia ( an a lka lin e wat er c o nta in ing a m o ng

'

, ,

o th er th ing s p e r c ent
,
N a
2
C O a)
n u m e r o u s li v ing I nf
,
us o ria ,

C o pepo da lar v a o f Ephyd ra and m o ld s 3 ”

.
, ,

A solu t io n o f ac etic ac id p e r c e nt s tr o ng k ill s t he t enta

c les of D ro sera b u t t he v in egar eel liv es in a 4 per c ent s olu t io n

,
.

M os t pro t o p las m is extrem ely s en s it iv e to ac id s gen erally es pe ,

c ia ll
y t o H C l and H 2
8 0 4 ;
b u t t h e ga s tr i c j u ic e i s larg e ly H C l ,

and the gland c ells of s o me m arin e Gas t ero p o d a s ecret e HQS O4

1
C B D .
p t E
.
p i ma v ent l M orp h l, g y x P t
erI en a
pp 7 3or o o , ar , . 2 -
2 .

I b id p
1
7 . . 2 .I b id p 8 1 . . 2 .
 IO CA U SES OF V A R I AT I O N

imm u n ity t hat m ight b e xpres s ed a s 4 00 wh en c o mpared with

e

t he no r m al as I T he i ncrea s e wa s s lo w at fi rs t b u t had reach ed

.
,

t wenty t im es t he n o r m a l with in eight day s aft er wh ich it ro s e ,

rap id ly
?

S t ill m o re s ign ificant a re t he fact s c o nn ect ed with t he prepa

rat io n o f d iphth eria ant it o x in fro m t he h o rs e I n th is proc es s .

R o u x firs t m ix es t he fi ltrat e o f t he bac illu s with io d in t o red u c e

i ts v iru lency On ly o n e q u art er o f a c u b ic c ent i met er o f th is
.

“
io d ized t o x i n is u s ed fo r t he fir s t inj ect io n b u t o n t he th ir ,

t een t h day a f u ll c u b ic c ent im et er is u sed On t he s ev ent eenth

“
.

day o n e f o u rt h c u b ic c ent im et er o f t he p u re t o x i n is inj ected ,

and th is is grad u ally inc reas ed in a m o u nt t ill o n the fo rty fi rs t -

day 1 0 cc is inj ect ed and o n the eight ieth day n o les s than
.

2 5 0 cc T he v i ru lency o f the la s t d o s e m u s t ha v e b een s o me

.

fi v e th o u s and to t en th o u s and t im es great er than that o f t he firs t

d o s e s h o wing t he ext ent t o wh ich t he c o n s t it u t io n o f the h o rse
,

had bec o me acc lim ated t o t he po iso n ?

D a v enpo rt and N ea l reared t wo lo t s o f s t ent o rs — o n e in p u re ,

water t he o th er in a s o lu t io n o f
, p e r c e nt Hg zC l A ft e r .

t wo day s bo th lo t s were p u t int o a k illing so lu t io n p er c e nt

Hg C lz) The res is tanc e perio d o f the lo t reared in p la in wat er was
.

o n t he a v e rage b u t 8 3 s ec o nd s wh ile that o f t he lo t gro wn in t he ,

weak s o lu t io n was 3 04 s ec o nd s Oth er expe r im ent s ga v e lik e.

res u lt s and fro m th es e in v es t igat io n s t he princ ip le was d ed u c ed

,

t/za t the r es is ta nc e per i od v a r ies d i r ec t ly wi t/t tlze s t r eng th of til e

s ol u t ion i n w/ziclt tlt e p r otop la s m ha s b een c u l ti v a ted
?
A S wo u ld
b e expect ed if t he c u lt u re so lu t io n s h o uld chanc e t o b e o n ly j u s t
,

o u t s id e t he letha l p o int it m ight per m an ent ly wea k en with o u t

,

d es tro ying life in wh ich c as e s u cc es s f u l acc limat izat io n wo uld

,

n o t f o llo w ; t he an im a l wo u ld s u cc u m b thro u gh weakn es s and ,

t he pri nc ip le s tat ed wo u ld n o t h o ld go o d .

I n th es e experi m ent s n o d eath s o cc u rred h enc e t he res u lt s

1 ‘

m u s t b e attr ib u t ed ent irely t o a lt era t io n o f the pro t o p la s m and n o t

at all to s elec t io n A ll th es e c o n s id erat io n s po int c o nc lu s iv ely
.

1C B D
.
p
. t Ea v en
p i m t
or l M ,p h l x
g y P
er t I en a
p 9 or o o , ar , . 2 .

1
Q t d
uo e f m V ro( V i t i i A
ern o n im l d P l
ar a t p on nw h q n t a s an an s , . o u o es

f m C
ro k h k T t B k f B t i l gy 896 p 5 8
ro o s an
’
s ex -
oo o ac e r o o , 1 , . .

1C B D
.
p
. t Ea v en
p i m t
or l M ,p h l x
g y P
er t I en a
p 3 or o o , ar , . 0
.

1I b id p 3 . . 1 .
 R ELAT I V E ST AB I LIT Y OF L I V I N G M ATT ER
3 1 1

to t he fact that ev en s o s ta b le a n d p ec u lia r a c o m ou n d a s

p l i v i ng
p r oto
p la s m ma
y u n d er o p r o
g fou n d a l ter a ti o n
f r o m c a u s es en t i r e ly
ex ter n a l t o i ts elf .

Mo reo v er , th is
t at io n is n o t m erely t em po ra ry b u t is
al er ,

o ft en la s t i ng ev en p er m an ent I n t he ca s e of Ehrlich s m ic e
’
.
, ,

already referred to ind iv id u als wh ic h had acq u ired a n im m u n ity

,

repres ent ed by 2 00 and were th en k ept on no r m a l f ood f or

,

m on tlzs wer e s t i l l f ou n d res is ta n t c er ta i n ly to tire ex ten t of 5 0

, ,

ho w m u ch m o r e wa s n o t d et er m in ed T h i i i acc o rd with
( s s n

t he expe r ienc e o f ars en ic and o p iu m eat er s and o f alc o ho l and

t o bacc o u s ers t he c o n dit io n is mo re o r les s perm an ent and the ,

body c rav es t he s pec i fic d ru g wh ich is n o lo nger po iso n o u s , .

I t agrees t oo with t he exper ienc e in i m m u n ity fro m d is eas e

, , ,

wh ich is freq u ent ly pe rm an ent thro u gh life aft er o n e attack .

A ll exper im ent s and exp erienc e agree that i m m u n ity fro m o n e

po iso n wh eth er nat u ral o r acqu ired is n o gu a ranty o f im m u n ity
, ,

i n any d egree fro m an o th er .

A cclim a tization t o hi gh t emper at u res A ll expe rim ent s ind i .

cat e that d eath fro m extrem e t em perat u res is d u e to c o agu la

tio n o f the pro teid s o f the pro t o p las m and that o rd inar ily t he ,

c oagu lat io n po int is n o t far abo v e t he h igh es t nat u ral t erres tr ial
t em perat u res S O that m o s t pro t o p la s m s a re u nab le t o res is t
,

t emperat u res abo v e abo u t 4 5 C hav in g apparent ly bec o m e °

.
,

n ic ely adj u s t ed to nat u ra l c o nd it io n s as gen erally enc o u ntered .

Y et o rgan is m s a re f o u nd in t he S O c a lled b o ili ng S p ri ngs at -

t em perat u res o f and ev en wh ich is n ear that

o f bo iling wat er
?
Mo rgan m ent io n s s p ec ifically L ept o thr ix in ,

t he K arls bad s pring s at 4 4 n o s t o c s and Pro t oc o cc u s

°
to ,

fo rm s in t he geys ers o f C alifo rn ia at Os c illaria in t he

Y ello ws t o n e Park s pri ngs at 5 4 t o in t he Ph ilipp in es at
°

in I s ch ia at and in I c eland at He rem ark s that

th ese t em perat u res may b e so m ewhat t oo h igh b ec a u se ho t ,

s pri ngs a re c old er at t he edges t ha n a t t he c ent er ; et t he h eat

y
is extrem e and far ab o v e t he nat u ral res is tanc e o f a n y kn o wn
,

fo rm o f pro t o plas m ?

1C B D a v en po rt , Exp eri m en ta M o rp o o g , Pa r t I p 3
. . l h l y , .

1 Mo r a n E v o lu t i n an d B D v en p o rt , E x p erim en ta l
g , o A d a p t a t i o n , p 3 2 0 ; C a . . .

h l y
Mo rp o o g , Pa rt I , p 2 5 2 ( tab es ) . l .
3 I 2 C AU SES O F V A R I AT I O N

T he a m e a u th o r m ent io n s s nails liv ing in F ranc e at 3 5 to

s
°

3 6
°
C and i n Pad u a at
,
r o t if ers at K ar ls bad at 4 5 t o
°

fro gs at Pise at and t he c ru s tac ean Cyp r is ba ln ea ria at

Ha mm a m Mec ko u t in at 8 1
°
-

A ll th es e a re liv ing at t em perat u res m u ch ab o v e t he d ea t h

po int of th eir neares t relat iv es and we are fo rc ed t o t he c o nc lu ,

s io n that th ey ha v e d es c end ed fro m anc es t o rs liv ing in t em pe ra

t u res n o t abo v e 4 0 and n o t at all ab le to end u re the extrem e °

t em perat u res in wh ich t he pres ent gen erat io n s liv e and th riv e .

A cc lim at izat io n t here m u s t ha v e been i n so m e way and at s o m e ,

t im e and that of extreme d egree in v olv ing pro fou nd c ha nges i n

, ,

t he pro t o p la s m .

E xperim ent s thro w int eres t ing light u po n t he m ann er o f ac c li

ma t iza t io n D u t ro c het f o u nd that if t he p lant N it ella wa s
.

p u t i nt o w at er at t h e c u rr ent s in t h e pro t o p la s m were

s t o pped b u t s oo n bega n aga in

,
I f p u t n o w int o wat er at .

th ey again s t o pped m o v ing b u t in a q u art er of an h o u r began ,

o nc e mo re If th en p u t int o wat er at
. t he c u rrent s s lo wed
d o wn b u t bega n aga in lat er
,
1
.
”

D a v en po rt and C a s t le reared t wo lo t s o f tad po les fro m t he

s a me lo t o f rec ent ly la id egg s On e lo t was k ept at 2 4 to
°
.

t he o th er at B o th lo t s d ev elo ped n o r m ally t he

° “
2 5 C .
, ,

fo rmer m u ch mo re rap id ly A t t he en d of f o u r week s bo th .

lo t s were t es t ed fo r h eat v igo r by g rad u a lly h eat i ng t he wat er

in wh ich th ey were c o nt ai ned T h o s e reared at 1 5 beca me
°
.

m o t io n les s at 4 1 o r b elo w ; wh ile o f th o s e reared at 2 4 t o 2 5

° ° °

( 1 0
°
h igh e r ) n o tadp o le d ie d u nd
,
e r t h e a v erage increa s ed

res is tanc e being The experi m ent ers rem ark that thi s ln
creas ed resis tanc e was d u e in n o s en s e to s elect io n fo r n o ,

d eath s occ u rred d u ring the perio d o f acc lim at iza t io n I t m u s t .

ha v e been d u e to changes wro u ght in t he pro t o plas m T h is c o n .

d it io n was mo re o r les s las t in g fo r t he d ifferenc e in res is ti ng ,

po wer th ou gh les sen ed was s t ill no t ed aft er sev enteen days

, ,
’

so j o u rn in c oo ler wat er
3
.

1 Mo rga n , Ev o l u t io n a n d A d pta a tio n , p .

3 2 0 ; C . B D a v en p o rt , E xperim en ta l
.

M o rp h o y P l
a r t I , p 2 2 ( ta
og
5 e,s . bl ) .

1 C B D a v en p o rt , E xp eri m en t al Mo rp
. . h o lo gy . Pa rt I , p . 2 53 .

1 Ib i p 2
54 d . . .
 C AU SES O F V A R I A T I ON
3 14

Acc lim at izat i on t o elect ricity . I t is m a tt er

c o m m o n kno wlof

edge that ind iv id u a ls wo rk ing abo u t electric ity and a cc u s t o m ed

t o freq u ent s h o ck s acq u ire a h igh res is tanc e po wer E xact data .

a re n o t at hand b u t o f t he gen era l fact th e r e c a n b e n o d o u bt

,
.

Accl imati zati on in gen eral I t is m a t t er o f c o m m o n exper ienc e

.

that a mo ng the h igh er an im als and plant s extrem e changes in

lo ca lity m ay b e f o llo wed by a n y o n e o f three separat e c o n s e
u en c es : fir s t u n pr ec ed ent ed pr o s p e ri ty fr o m extrem ely s u i t
q ,

able c o nd it io n s c o nd it io n s ev en mo re fav o rable than in the

,
-

nat iv e habitat ; sec o nd ab so lu t e failu re fro m S h eer inab ility to

,

end u re t he C hang ed c o nd it io n s ; and th ird a p e rio d o f s t ru ggle , ,

f ollo wed by a m o re o r les s c o m plet e adaptat io n to n ew c o nd i

t io n s ,
a proc ess kn o wn ev ery wh ere a s acc lim at izat io n .

T he kno wn f act s c o nc e rn ing ad a ptat io n i n lo wer f o rm s s eem

t o app ly i n t he S a m e d i rect io n if n o t t o t he s a m e ext ent t o
, ,

ev en t he h igh es t kn o wn S p ec ies F o r exa m p le t he b ienn ial o f

.
,

t he t em perat e r egio n s b ec o m es p erenn ia l in t he tro p ic s wh ere ,

a ll d is t inct io n s o f th is o rd er b lend and d is app ear and wh ere all

p lant s liv e u nt il th ey d ie fro m o th er than c lim at ic cau ses T h is .

is a s u ffic ient exp la nat io n fo r t he m o re ready fo rm at io n o f fl es hy

roo t s and s t em s in no rth ern regio n s and t he les ser s t o ra ge o f foo d
,

res erv es in the trop ic s wh ere gro wth t end s to b e c o nt in u o u s

, .

I t has a lway s b een h eld that p lant s hav e h igh er p o wers o f

acc lim at izat io n than ha v e an im a ls D o u bt less th is is tr u e at .
,

lea s t so far a s p u rely c lim a tic c o nd it io n s a re c o nc ern ed b eca u s e ,

an imals b eing free t o m o v e and gift ed wit h mo re o r les s in telli

,

genc e a re ab le in a c o n s id erable d egree to a v o id t he f u ll effect s

,

o f c lim at e by s eek ing S h elt er o r pro t ect io n o f s o m e s o rt T h ei r .

o wn b o d ies a re fi tt ed t o m a int a in ev en a ga i n s t f earf u l o dd s a

, ,

nearly c o n s tant t em p era t u re S o fa r as t em pera t u re is c o nc ern ed

.
,

th eref o re an im als d o n o t need t he sa m e d egree o f adaptat io n as

,

d o p lant s wh ich m u s t end u re t he bes t th ey can t he acc id ent o f

pos it io n .Wh eth er t he a n i m al po s s es s es t he s a m e capa c i ty fo r
acc lim at izat io n wh eth er it is c ap a ble o f u nd ergo ing t he s a me d e
,

gree o f alt era t io n and d ev elo p ing t he s am e res is t a nc e is an o ther ,

q u es t io n b u t experienc e with p o is o n s t end s to s h o w that t he an im al

,

o rg a n is m is n o t i nfer io r t o t he v eget a b le in p o int o f ad a ptab ili ty

wh en p u t t o t he act u a l t es t and c o m p elled t o fac e t he in ev i tab le .

 R E LAT I V E S TABI LITY O F LI V I N G M ATT ER
3 5
1

Spa c e d oes n o t perm it t he en u m erat io n of in s ta nces of acc lima

tizat io n o f d o mes t ica t ed an im als and p lant s R eferenc es a re f u ll .

of da t a o f th is so rt , an d c o m m o n exper ienc e s u pp lies a m a s s

of a dd it io nal info rm at io n T hat bo th an im al and plant S pec ies

.

po sses s an d exerc ise large po wers o f adaptat io n to ext ernal co nd i

t io n s is n ow n o lo nger q u est io n ed T he o n ly d ou bt is as to the .

m eth od Wheth er t he wh o le effect is prod u c ed throu gh ind iv id

.

ual mod ificat io n wh eth er it is largely t he res u lt o f select io n o f

,

ha rd ier strain s o r wh eth er th ere is som e inh eritanc e of mod ific a

,

t io n o f th is s o rt is yet a n Open q u est io n .

With ou t a d o u bt ind iv id u al m od ifica t io n has been n u d eres t i

m at ed I t has been sa id tha t t he po wer s o f ind iv id u al adaptat io n
.

are S light an d that r ea l acc lim at izat io n is s ec u red thro u gh t he

select io n of t he few ind iv id u a ls nat u ra lly end o wed with po wer s

of h igh res is t a nc e Fo r exa m p le it is a well kn o wn fact that

.
,
-

wh en an ent ire field o f c orn ( m aize) has s u ffered from a k illing

fros t a few s ta lk s h ere an d th ere are u naffect ed The same fact .

has been no t ed am o ng ga rd en v egetab les and fru it s of v ariou s

so rt s m any o f wh ich a re m ent io n ed by D a rwin I n C hanging
?
,

S pr ing t o wi nt er wh eat all b u t a few s ta lk s a re k illed t he fi rs t

wint er b u t th ose rem a in ing gi v e r is e t o wint er s train s

, .

T h ese and S im ilar fact s ha v e giv en rise to t he id ea that a c c li

mat izat io n is really thro u gh t he select iv e proc es s and ba sed on ,

inh erent d ifferenc es in c o n st it u t io n o f t he o rgan ism rath er th an

u po n ad aptab ility R ec ent res earch es h o wev er hav e s h o wn
.
, , ,

with in lim it s an u n expect ed d egr ee o f adaptab ility in t he ind iv id

,

ua l and t he exa m p les already c it ed es tab lis h t he fact that acc lim a
,

t iza t io n is largely t he res u lt o f mod ifi cat io n o f pro t o p las m Ho w .

far th is m od ifi ca t io n is po ssible and h ow far it is perm an ent and ,

wh eth er it is in any d egr ee inh erit ed a re n o w t he ab so rb ing ,

q u es t ion s That th ere is a h igh d egree of perm an enc e at leas t

'

.
,

in so me cas es is well es tab lish ed Wheth er t he m od ifica t io n s a re

,
.

t o an y ext ent inh erit ed is a s u bj ect that will b e d is c u ssed in t he

n ext ch a pt er .

T wo fact s are wo rthy o f n o t e in all s t u d ies in acc limat izat io n

fir st the v ar iat io n s are f u nct io nal n o t m o rph olo gical ; sec ond
, , ,

t he ext erna l fo rces in v o lv ed a re s u ch a s ex erc ise an a ll per v a d i ng -

D wi 1 A im lar d Pl t
n, d D m ti ti I I 99 3 0
n a s an an s u n er o es ca on, , 2 -
0 .
 16 C AU SES OF V A R IAT I O N
3

infl u ence . Ha
chem ica l act io n light elec t ric ity
t, a ll a r e ih , , ,

flu enc es t hat pen et ra t e t o t he v ery c o n s t it u t io n o f t he pro to

plas m ev en of t he germ perh a ps T he m o s t S ign ifican t b iologi ca l
,
.

fa c t in th ese s t u d ies is t he wid e ra nge o f ad apta t io n and t he

exc eed ing ly d iv er se c o ndit io n s u nd er wh ich liv ing m att er m ay

con t in u e to d is c harge it s n o rmal f u nct ion s eith er u nalt er ed or

—
,

b u t S light ly mod fi ed
i and t he next mo s t s ign ifican t fact is t he
,

m o re o r les s per m a n en t alt era t io n in f u nct io nal act iv ity thro u gh

changed c o nd it io n s of t he o rd er m ent io n ed ?

SEC TIO N V II —EV I D EN C E FR OM R EG ENERATION

A ll n imals and plant s h a v e m o re o r les s power t o res t o re los t

a

o r inj u red pa rt s I t is n o t ic eable a n d perha p s S ign ifi ca nt tha t

.
, ,

t he po wer o f regen erat io n is ro u gh ly s pea k ing in in v er se ra t io , ,

t o t he d egr ee o f d ifferent ia t io n ; that is t o sa y lo wer o rgan is m s ,

hav e in m any cas es at lea s t alm o s t u n lim it ed powers of regen era

, ,

t io n wh ile a mo ng t he h igh es t s pec ies the a b ility to res t o re los t

,

pa rt s is v ery S light .

T he bearing o f th is m att er u po n t he q u es t io n in hand — tire

r ela ti v e s ta b il i ty or i ns ta b i li ty qf li v i ng ma tter is o f c ou r se not
in t hef ac t o f regen era t io n b u t in t he charact er o f t he regenera t ed
pa rt as com pared with t he o rigin al and in the gen eral beha v io r ,

o f t he o r ga n is m wh en occas io n fo r regen erat io n aris es Mos t o f .

t he in s tanc es gi v en in th is c o nn ect io n a r e d ra wn fro m M o rga n s

’

exc ellent wo rk R egen era t io n ,

.

R egen er ati on in an imal s ?

T rem b ley Re a u mu r

Bo nn et a n d S pa llan za n i
( 1 768
) w e r e ea r ly i n v es t i at o r s
g
in th is s u bj ect T h eir exper im ent s hav e been o ft en repea t ed an d
.

th eir o bserv a t io n s ext end ed u nt il the field has been well wo rked
a n d t h e lim it s o f regen era t io n fa ir ly well es tab lis h ed .

I f t he foo t Of a s a la m and er b e c u t o ff a n ew o n e regen era t es If .

t he ent ire leg b e r em ov ed it regen erat es and a t wlza tc v er po in t t he ,

1 Fo r a dd iti ona l re feren c es on ac c l im a t iza t io n s ee D a rwin ’

s Origin of S pe
c ies , an d h is A n im a ls an d Plan t s un d er D o m es t ic a t io n I I, 2 9 5-
3 5;
0 V ern o n ,
—
.

V a ria t io n in A n im a l s an d Plan t s , p p .
3 79 8
3 7; Ba i ly e , S u rv iv a l of the U n li ke
( d d it i ) W eis ma n n d
S t u ies in t h e T h y of D esc en t ,
—
s ec o n e on pp 0
3 7, 3 1 0, 3 2 0 ; eo r
—
, .
,

pp .
55 5 6 22 M o rga n E v o lu t io n, and A d a
p t a t io n , pp .
3 9
1
3 5 : 2
1 M g or an, R egen era t io n , p p . 1 - 1 0.
3 18 C A U SES O F V A R I A TI O N

2 . I f t wo or if t hree s egm e n t s b e c u t O ff ,
a n eq u a l n u m b er w ill
be res t o red ( see F ig 3 6 C D ) .
, ,
.

3 I.f f o u r o r fi v e b e cu t o ff g en era llv b ut , ,

n o t a l wa v s , a c o rr e
s po n d in g n u m b e r c o m e ba ck ( see F I g 3 6 E .
, , F) .

F IG .
35 . T h e m et h d
o of
g r o wt hi n reg en era t io n

l l
A , ta i o f F u n d u l u s b eter oc li t u s : c u t sq ua re y, it re gen era t es as a t B ; c u t o b iq u e y, as a t C l l .

l
I n D a n d E a re s ho wn regen era t io n fro m t wo o b iq u e c u t s ; 6 , ta i o f s ten o p u s , regen l
l
c ra t in g fro m sq ua re c u t a t H a n d fro m o b iq u e c u t a t I A fter M o rga n
.

4 . If si x or m or e b e n ly f o r
c ut
u o r fi v e c
o ff,
o m e back
o .

T h ese are t he s egm ent s c o n s t it u t in g t he h ead pro per a ll o f wh ich ,

a re res t o r ed b u t if t he c u t b e m ad e farth er back t he int erm ed iat e ,

 R E LAT I V E S TAB I LITY OF LI V I N G MAT TER
3 19

segment s are n o t res t o red and t he wo r m rem a in s s h ort en ed

,
“

through ou t the res t o f it s life ( s ee Fig 3 6 G H ) .

, , .

0
FIG .
3 6 . R e en era t io n o f ea rt
g hw
or m , A ll olobop b or a f a ti da
’

A, n or l -
l
ma worm ; B F , regenera t ion a fter remo v a o f o ne, t wo , th ree, fou r, an d fiv e hea d
segmen t s res pec t iv e y ; l l
G , an teri o r t hird c u t o ff, on y fi v e segmen t s rest o red ; H , cu t
l
in t wo in t he mid d le, o n y fi v e rest o red ; I , c u t posterior t o t he midd e, t a il m a tter l
rest o red A fter M o rga n
.

5 . I f t he far
cut e n o be
u gh back t o r em o v e t he r epro d u ct iv e

o rgan s th ey a re n ev er ref o r m ed and t he wo r m remain s inca

“
, ,

p ab le o f r epr o d u c i ng i t s elf .
”
 0 C A U SES O F V A R IAT I O N
3 2

6 . I f the cut b e m ad e
beyo nd that is po s terio r to t he m idd le , , ,

b o th pa r t s will reg en era t e T he po s t erio r pa rt h o wev er will re

.
, ,

genera t e S lo wly and with difli c u lty n o t h ea d b u t t ail s eg ment s , , ,

a n d t he an o m aly a ri s es o f an ea rt h wo rm wi th t wo ta ils and

n o h ead T h ere is th u s a k i nd o f o la ri t by w h ch ant r r ”

.
p y i e i o

s eg m ent s m ay aris e o n ly fro m c o m p a rat iv ely ant erio r regi o n s ,

a n d by wh ich po s t eri o r regi o n s m u s t o f n ec es s i ty g iv e ris e t o

po s t erio r segm ent s T he S a me S ign ificant fact s a re n o ta ble in

.

o th er s pec ies b o th an i m a l a n d p lant ( s ee F ig

,
.

7 . T h is s a m e r ela t io n bet ween t he n u m be r o f s eg ment s c u t

o ff fro m t he ant erio r en d and t he n u m ber that is reg en erat ed

s eem s to h o ld go o d thro u gh o u t t he wh o le gr o u p o f ann elid s ,

alth o u gh t he m ax im u m n u m be r th a t c o m es back m ay b e d iffer

en t in d iffer ent s pec ies T h u s in L u m b ric u lu s S ix s ev en o r
.
, ,

e v en eight n ew s eg m ent s c o m e back if m o re than that n u m be r ,

a re rem o v ed T h ere is th eref o re a d efin it e t erm in at io n t o

1 "
.

t he po wer o f r egen erat io n ev en in s pec ies with h igh regen er ,

at ing po wers .

8 “
. I f we ex a m in e t he m eth o d o f regenera t io n fro m th e po s
teri o r end o f a p iec e o f an ea rth wo r m we fi nd that wh en s ev era l ,

o r m any po s t eri o r s eg m ent s h a v e been r em o v ed a n ew part c o m es

back c o m po sed at fi rs t o f v ery few s egm ent s

,
1
t he la s t o f wh ich ,
”

c o ntains t he n ew Open ing fo r t he d iges t iv e t ract La t er add i .

t io na l s eg m ent s a re f o r m ed ju s t a lrea d qf tire la s t s eg m en t q u it e ,

aft er t he n o r m a l manner o f inc reas ing t he length in m any s pec ies

o f ann elid s .

9 T.h e ant e r io r p o rt io n d oes n o t po s s es s t he po wer o f r egen

c rat io n u n le s s it c o n s i s t s o f a c o n s id erab le n u m b er o f s egm ent s

( S ee F ig .
37 A —
,
E ) .

10 . a s h o rt piec e ( three to s ev en s eg ment s ) o f the a n ter i or

If
po rt io n of o n e wo r m b e g raft ed in a r ev ers ed pos it ion u po n t he
a n ter i or en d o f a n o th er wo r m t he g raft ed pa rt will regen erat e a,

h ead o f abo u t t wo s egm ent s fro m it s free en d wh ich was o rigi ,

nally t he po s t erio r extremity b u t o f an ant erio r p iec e ( see F ig , .

37 F
, ) T h is. s h o ws t he ent i r e po we r o f r ev e r s a l a s t o relati v e

po s it io n of seg ment s a h igh d egree o f adapta b ility

, .

1 M o rga n R egen era t io n p 9

, , . .

1 Ib i d .
p 9 . .
3 2 2 C A U SES O F V A R I ATI O N

Lateral r egen er at i onxa m p les th u s far gi v en are o f regen

. A ll e

c rat io n length wis e and o f t he res t o rat io n o f part s s i m ilar to th os e

,

rem ov ed b u t aris ing fro m m att er d ifferent ly d ifferent iat ed

,
.

I f t he p lanarian b e s p lit length wise eith er thro u gh the m idd le

,

o r well to o n e s id e each p iec e will r egenerat e

,
I f a hydra b e .

FI G 38 . . R egen era tio n in th e p l an a ria n

A, n or l
ma worm ; B , B 1
, regen era t io n o f an ter io r l
ha f ; C, C 1 , regen era tio n of pos teri o r ha fl
l
0 , c ross p iec e ou t o f m id d e ; D i , D 1 , D 3 , D 4, regen era tio n o f sa me ; E , o ld heed ; E 1 ,
E 1 , E 1 , regen era t ion o f sa me ; F , o ld hea d c u t o ff jus t b ehin d the eyes ; P I , regenera t io n
o f n ew hea d on po s teri o r en d o f s a me A fter Mo rga n
.

split t he edges of the t u be will c o me t o gether and j o in mak in g

, ,

a t u be o f s m aller d ia m et er en larging lat er I f the leg o f a sala

,
.

m and er b e s p lit and a part rem o v ed it will b e replac ed s h o wing

, ,

th a t regenerat io n is lat era l as well as lo ngit u d inal .

I f h o wev er t he bo dy o f t he s a la m and er s h o u ld b e s p lit bo th

, , ,

p iec es wo u ld d ie n o t pr im arily fro m lack o f regenerat i ng po wers

,
 R E LA T I VE ST A B I LI TY O F L IV I N G M ATT E R 3 2 3

b ut fro m s h eer inab ility to s u ppo rt life Man ifes t ly a ll regen er .

at io n is a s tru gg le and a prereq u is it e t o it s acc o m p lis h ment is

,

an as s u red ba s e o f s u pp lies and u n inj u r ed v ita l o rgan s .

R egen erat i on b y t ran s f ormat ion ‘

R egen erat io n in s o m e o f .

t he lo wer an ima ls is acc o m p lis h ed throu gh rearrangem ent o f o ld

s u b s tanc e as well as thr o u gh t he add it io n o f n ew m at er ia l .

F IG 39 . . R egen era t io n of S t en t o r cut in t o t h ree p iec es , as a t A

B :t his ro w sho ws re gen era t io n o f a n terio r p iece . C :th is ro w s ho ws re gen era ti o n o f m idd l e

p iec e D :th is ro w s ho ws regen era t io n o f po s ter io r p iec e T h is regen era t io n is effec ted
l—
. .

fi rs t of a ll by rea r ra n ge men t o f ma teria , ea c h p iece ha v in g b een su pp l i d with

e a po r
tio n of the l
n u c eu s . A fter M o rga n , fro m G r u b er

If a s h o rt p iec e b e c u t fro m the s t em o f a hydra the fi rs t s t ep

in t he f o r m at io n o f a n ew ind iv id u a l fro m t he p iec e is t he c lo s ing
o f t he end s and a s hrink in g o f d ia m et er th u s m ak ing a c lo s ed c lin
y ,

der b u t m u ch s ma ller than t h e s t em fro m wh ich it wa s c u t

,
In .

a day o r t wo fo u r t entac les appear at o n e en d and s h o rt ly t he ,

piec e has a s s u med t he cha ract eris t ic f orm and p ropor tion s o f a
c o mplet e hydra aft er wh ich it m ay increa se i n s ize T he sa m e
,
.

is tru e of a p iec e o f a p lanarian o r of a S t ent o r (s ee F ig .

1 M o rgan , R egen era t io n pp , . 1 3

- 1 5 .
3 4
2 c a s es o r V A R I AT IO N

he fi r s t p roc es s s e m s t o b e t o as s u m e t he c ha ra c t e ris t ic
e

o r m af t e rw a rd t o i n c rea s e in s iz e ; n o t o n i v t h a t b u t re ene ra
f ,
g ,

ti o n is po s s ib le in t he e n t ire abs n e f f
e
c o o o d , as w ill b e s ee n

la t e r all o f wh ic h ind ic a t es m o re o r less e x t e ns iv e t r a n s

. a
f or

m a t io n of ma t e ria l .

R egen er at ion in emb r yos a nd eggs .

l
T h ere is m u c h reaso n to

b e liev e tha t regenera t io n es pec ia iiv b v rm g e m e nt is m,

o r e ,

pro no u nc ed in t he em b ryo n ic t ha n in t he a d u lt s t a te Th e fro g .

d o es n o t re genera t e t he le g b u t t he ta d p le d s I f t he b las t u la ,
o o
e
.

o f t he s ea u rch in b c u t in t o t wo p iec es ea c h w ill d e v elo p int o a

e
,

p e rf ect b u t a bn o r m a lly s m a ll e m bry o

,
I f t he p a rt s b e se pa ra t ed ,
.

a t t he t wo o r t he f o u r c elled s t a g e ea ch is c a p a b le o f d ev e lo p i ng
-
,

in t o a p rf ect em b ryo b u t a t t he eig h t c elled s ta g e t h ey a re n o t

e

ca pa b le o f s u ch d ev elo p m ent .

I f ea c h c ell o f t he t wo c elled s t a ge is c a p a b le O f d ev elo p in g -

int o a c o m p let e i nd iv id u a l th en t he m a t erial at th is s t a ge m us t ,

b e ind i fferent th a t is u n d iff eren t ia t ed,

I n o t h er wo rd s if t he .
,

fir s t c lea v a ge b e c o n s id ered a s d iv id in g in t o rig ht and left h alv es ,

and ea ch ha lf is c a pa b le o f d ev elo p ing in t o a wh o le t he cas e is ,

exac t ly lik e th a t O f r egene rat io n i n t he s p li t p la n a ria n ; i f ho w ,

ev er t he fi rs t s egm e n t a t io n b e c n s id e red a s d i v id in g i nto a nte

,
o

rio r a n d p s t e rio r r egio n s a n d if ea ch m ay d ev elo p a n i nd iv i d u al

o , ,

it is exa ct ly s im ila r to t he c a s e o f t he regen era t io n Of a wo rm

that has been c u t in t wo cro s s wis e int o ant erio r a n d po s t erio r
h a lv es I n all c a s es s o m e rea dj u s t m ent o f m a t eria l is in v o lv ed
. .

T h is s epa ra t io n is ea s ily eff ect ed in t he s ea u rch in by s hak in g ,

in wh ich c a s e ea c h pa rt b elo w t he e ig ht c e lled s t age d ev elo p s an

,
-
,

in d iv id ua l I f in t he fro g t he p a rt s ev en a t t he t wo c elled s ta ge
.
,
-
,

b e s epa ra t ed ea c h c o lla p s es ; if in s t ea d o n e h a lf b f i l led by a

,
e

n eed le t he u n inj u red p a rt d ev elo p s at fi rs t a ha lf em b ry o aft er

, ,

ward s m ak in g m o re o r les s s u cc es s fu l po s t gen erat io n o f a

wh o le em b ry o 2
I t is th eref o re a pe rfect ly well e s tab lis h ed fact
.
-

that in c ert ain s pec ies at leas t a part o f an egg o r em bry o is

, ,

capa b le Of d ev elo p ing int o a perfect ind iv id u a l .

To wh a t ex t ent t h is s epa ra t io n o f s eg m ent ing eggs at t he

t wo c elled s t a ge m a y t a k e p la c e in n a t u re is O f c o u r s e u n kno wn
-
,

M g R g 1 i or
p 8 ; l
an, h p ei pp 6
e n e ra t o n ,
4 1 a so c a x 2 1 -
2 1

6—
. . .
, .

I b id pp 2 . . 2 1 22 1 .
 6 C A U SES O F V A R I A TI O N
3 2

pa rt s t han do th o se wh ich a re mo re h igh ly d ifferent iat ed S ti ll .

t he latte r a re n o t d es t i t u t e o f thi s fac u lty O f rep lac em ent T he .

t eeth o f m any v ert ebrat es a re s h ed o nc e a n d rep la c ed ra rely a

s ec o nd rep lac em ent o cc u r s I f t he o x los es t he h o rn t he lo s s is
.
,

perm an ent b u t t he s tag s h ed s his ann u ally each s u cc es siv e pai r ,

a ris ing fro m t he s a m e s car o r b u d b u t ea ch pro v id ed with an

,

a d d i t iona l p r ong . By what i nh e rent q u ality Of th is part ic u lar

s po t a re we t o exp lai n thi s ann u a l change i n t he chara ct er Of

t he part res t o red ?

Bi rd s s h ed th ei r p lu m ag e and m any ani m a ls th ei r h ai r a n n u
, ,

ally as trees s h ed their lea v es and Oft en t he n ew growth d if

, ,

f ers m at erially fro m the Old The milk t eeth a re si m pler than
.

t he per m a nent s et ; t he c o lo r o f t he fo al and t he fa wn chan ges

wi th m a t u ri ty ; and t he s hape o f t he c o ty led o n gi v es litt le i nd i
cat io n Of what t he real leaf will b e N o bo dy s eeing t he u m brell a
.

li k e fir s t leaf o f t he bas s woo d wo u ld s u s pect what t he lat er

lea v es will b e n o r wou ld o n e s u s pect t he c lo v er u nti l t he
,
thi rd
leaf appea rs .

R epai r o f i nj u ry a m o ng h igh er a n i m als s eem s t o b e exc eed

ingly li m it ed . Ho wev er wo u nd ed m u s c les c an fill u p to s o me
,

ext ent ; c u t n erv es m end s lo wly ; s ev er ed b loo d v es s els r epa i r

them selv es and res to re c irc u lat io n t o t he pa rt ; liv er ki dney , ,

gland s and t iss u es generally hav e s u flic ien t po wer of regenera

,
‘

t io n to c lo se wo u nd s and to replac e los t po rt io n s m o re o r les s

perfect ly b u t nearly alway s with a s c a r bro ken bo nes will kni t
,

o r a s m a ll piec e remo v ed wi ll b e r es t o red b u t an ent i re bo n e c u t

,

o ff will n o t b e r ep lac ed Of a ll part s t he s kin p o s s es s es the h igh es t

.

po wer Of res t o rat io n pro bab ly bec a u s e it is no rm ally in act iv e

,

g r o wt h fr o m b e n ea th t o re p lac e t he pa r t s w o rn o ff fr o m ab o v e .

Th e ch ar acter of t he regen erat ed part T he regen era t ed pa rt .

may c o m pare with t he o ri gina l in any o n e Of fo u r d i fferent way s

I
. I t m ay b e exact ly lik e t he o riginal as in t he leg o f a s ala,

m and er (h olo m o rph o s is ) .

2. I t m ay b e lik e t he o rigi nal exc ept s m aller in s ize ( mero

m o rph o s is ) .

3. I t m a
y b e d i ff e r e nt fr o m t h e o ri g i na l b u t l ik e s o m e o th er
part Of t he body as wh en an ant enn a replac es an eye ( h et ero
,

mo rph o s is ) .
 R ELAT I V E STA B I LIT Y O F L I V I N G M AT TER 3 7
2

4 .I t m ay b e u n li k e any n o r m a l s tr u ct u r e o f t h e bo dy ,
as w h e n
n ew leg is u n lik e any o th er leg o n t he bo d y
1
a ( n eo m o rp ho s i s ) .

With res pect t o t he t is s ues fro m wh ich regen erat ed part s a rise
t wo d i s t i nct cas es a re t o b e n o t ed
I .Wh ere t he part regen erat ed s p rings fro m t iss u e o f t he
s a me k ind req u i ri ng o n ly an ext en s io n o f gro wth as wh en an
, ,

inj u red m u s c le is repaired .

2 .Wh ere t he regenerat ed part s prings fro m t is s u e o f a t o tally

d ifferent o rd er as wh ere a sev ered leg is res t o red fro m t he c u t
,

o u t ward o r wh ere t he len s o f an e e ari s es fro m t he i ris re

, y ,

qu iring d ifferent iat io n a s well as gro wth 2

T h is s u bj ect Will b e .

pu rs u ed f u rther u nd er t he sect io n o n Origi n Of N ew C ells a nd

“

Tis s u es .

Effect of t emper atu re u pon r egen er ation .

8
Plan arian s were c ut

in two tran sv ers ely at pha rynx N o regen erat io n t oo k plac e

t he .

b elow 3 C Of six s pec imen s k ept at th is t emperat u re o nly

°
.

o n e r eg en erat ed and that inc o m p let ely t he eyes and brain

, ,

being inc o mp let e aft er s ix m o nth s The t em perat u re at wh ich .

regenera t io n t oo k p lac e mo s t rapid ly was a t wh ich a n ew

h ead fo rmed in fo u r and s ix t enth s day s A t it req u ired

-
.

eight and a h a lf day s t o c o m p let e t he h ead at 3 2 regenera t io n °

c o m m enc ed b u t d eath o cc u rred in abo u t s ix days ; at 3 3 re

,
°

genera t io n was s light and at 34 n o n e t oo k p lac e d eath occ u r

,
°
,

ring with in three d ay s Oth er s pec ies s h o wed a s im ilar ra nge

.

for o pt im u m m in im u m and m ax im u m
,
‘
,
.

I nflu en ce of food u pon r egen erati on 5

Wh ile regenerat io n tak es .

plac e mo re rap id ly with a f u ll food s u pp ly i t n ev erth eles s p ro ,

c eed s wi t ho u t it I n th is c a s e t he n ew gr o wth app ear s t o b e d e

.

ri v ed n o t fro m s u rp lu s fo od m at er ial b u t fro m the pro t oplas m

it self res u lt ing in red u ct io n in s ize
,
.

I f a p lanarian b e k ept fo r s ev era l mo nth s with o u t foo d i n ,

th is s ta rv ed c o nd it io n it g ra d u a lly s hri nk s in s ize ev en t o on e ,

th irt eenth o f the n o r m a l (s ee F ig .

I f a s tarv ed wo r m b e c u t in t wo p iec es ea ch will reg en erat e , ,

th o u gh m o re s lo wly than if fed the n ew part increa s ing in s ize ,

at t he expen s e Of t he O ld .

1 M go r an , R g ti e en era
p 4 I b id
on,pp 6— 7 2 I b id pp 7
— 9
3 2 2 5 2 2
—
. . . . . . . .

3 I b id p. .
5 20 . I b id pp 6 7 . . 2 2 .
3 2 8 C A U S ES O F V A R I AT I ON

A s Mo rgan remark s T he gro wth of t he new part at the ,

1 “

exp en s e o f t he o ld t iss u es is a ph en o men o n o f t he great es t i m

po rtanc e an exp lanat io n o f wh ich will i n v o lv e I th ink the mos t

, , ,

f u nda mental q u es tio n s pertain ing to gro wth To th is rem ark it .

”

may b e add ed that t he ph en o men o n is a ls o o f t he great es t i m po r

tanc e in its b earing u po n the rela t iv e s tab ility Of liv ing m att er .

I n s o far as pro t o p las m is wor ked ov er to s erv e n ew p u rp o ses it ,

s h o ws wo nd erf u l elas t ic ity b u t t he fact that

t he o rgan is m pres er v es o r c o m p let es i t s p lan
a t a lm o s t any exp en s e ev en to i t s elf bet ray s , ,

a wo nd erf u l pers is t enc e o n t he part of the

o r ig i na l p lan .

I t is kn o wn that t he s t a rv ing cat o r d og

will rep lac e a large s hare o f the d ry m att er
o f t he b o dy with wat er and s ac ri fic e all o th er ,

act iv it ies to the v ita l pro c es ses Plant s gro w .

in g with n o n itro gen s u pp ly s a v e what is

c o nta in ed in t he s eed s will soo n reach t he
max im u m Of p o s s ib le d ev elo p ment b u t will ,

c o nt in u e to pro d u c e n ew lea v es at t he ex
pen s e Of the o ld o n es as t he ra pid ly grow
2
,

in g s t alk o f t he c en t u ry p lant f eed s on it s

Efi t é f t ec s ”

“ ti p t
on h p l
u
th i ck
on
en ed e
lea v es o r t he b eet and carr o t f eed
a ,
,

inar an o n th eir th ick en ed r oo t s .

A w ll f d w m ; B th
3
, e -
e Effect s of li gh t u pon regen erat ion
or ,
e I n t he .

i
ssggfitt iif;
E ca s e Of p lant s o n ly d o es light s eem tO a ffect
4

regen era t i o n and i n th em t he i nfl u enc e

O

f f
or m th
ou r d on s an ,

M t“
i
d y
hlls; app a
e ar s to b e c o n fi n ed to t he b lu e
s- “
ray s .

Herb s t Ob s er v ed that wh en the eye o f c er

ta in C ru s tac ea is c u t Off s o m et imes an eye and s o m et i mes an
ant enna is regen erat ed E xp er im ent s were c o nd u ct ed t o see
.

wh eth er light m ight b e t he fact o r wh ich d et er min ed wh eth er t he

1 Mo rgan R egen era t io n , p p

, .
—
27 29 .

3 T h e writ er s a w t h e o ri
gi n a l c l o v er
p lan t s in t h e fi rs t R o tha m s t ed s eri es fo r tes t
in g t h e n it ro gen -ga t h eri n g p o we rs o f ro o t t u b erc les . On e of t hes e p l a n ts ha d no

s o u rc e O f n it ro en
g b u t t he s ee d . I t wa s t wo y ea rs O ld , s t i ll p ro d u c in
g ne w l ea v es

as t h e O ld o n es di d d e wn b u t it h ad
o , n ev er b lo s s o m e d , no r was it ab e t o l p ro

d uc e mo re h
t a n fo u r lea v es a t an y t im e
—
.

3 Mo rga n R egen era t io n pp

, , . 2 9 30 .
4 Ib i d .
p 7
. 8 .
33 0 C A U SES O F V A R I AT I O N

in U rbana , I llin o is , ght een inch es o r m o re in d ia meter I t

ei .

fo rk ed abo u t s ix feet fro m the gro u nd A h eav y s t o rm s p lit .

o n e Of t he lim b s a wa y wh en it wa s fo u nd that a d en se n et
,

wo rk Of ro o t s ha d f o r med i n the m o is t s o il th a t had c o llect ed

in t he fo rk .

F IG 4 1
. . R eg en era t io n in res p o n s e to gra v ityi n a n im a l o rga n is m s : t hi
s c rea t u re

( A n ten n a /a r i a a n ten m na )
'

r es emb es l p l t
an s in it s h a b it Of g ro wt h , b ein g
a tt a c h d;
e it a ls o l
res e m b es h
t em in it s res p o n s e to g ra v i t yi n re en era ti o n
g .

A f t er M o rga n fro m L o eb
,

4 . p
I f t he
iec e b e s u s p end ed ap ex d o wn t he s h o o t s will s t ill ,

s tart fro m t he ap ic a l en d b end ing u p ward a n d ro o t s will s tart

‘

, ,

n o t o n ly fro m the b a s a l en d n o w u pp er m o s t b u t th ey will s tart

, ,

fro m the wh o le len gth Of t he s t em a n d b end d o wn ward s h o wing ,

that the fo rc e d et er m in ing wh eth er s h o o t o r roo t s hall b e pu t o u t

 R ELAT I V E ST A BI LIT Y OF LI V I N G M ATT ER 33 1

is largely i n terna l , b u t that t he infl u enc e d et erm in ing t he d i rec

t io n of gro wth is ex terna l
t he grav ity ( s ee F ig 2 6 B ) , .
,
.

5 T
. h is p o lar d iff e r e nc e is m os t e n e rg e t ic in y ou n
g s t e m s ,

g rad u a lly les sen ing in Old er gro wth th o ugh the t end ency rem ain s ,

in q u it e Old p iec es .

6 I f i nt ern o d es o n ly a re u s ed s o me p lant s will regen era t e

.
,

and o th ers will n o t ; b u t if th ey d o regenerate the t end ency is ,

fo r roo t s t o appear fro m t he bas a l en d and lea v es fro m t he ap ical ,

whatev er the p o s it io n 1
'

7 I.f a p ie c e o f t h e r o o t o f a p o p lar b e s u s p e nd ed v e rt ica l ly in

a mois t cha m ber apex d own wa rd leav es and s h oo t s will appear

, ,

fro m the ba s a l n o w t he upper en d ; if s u spend ed ba sal en d

2
, ,

d own ward the s h oo t s will s t ill arise fro m th is ( bas al) en d

,
3
.

8 C erta in p lant s a s t he bego n ia a re ab le to pro d u c e n ew

.
, ,

p lant s fro m ev en a s ing le leaf if set in mo is t sand I n all cases .

,

s o far as kn o wn roo t s fi r s t aris e at t h e ba s e o f t he leaf s t em o r

, ,

m id rib s ect io n and later s h o o t s ar is e o n t he ap ica l s id e of t he

,

roo t s what ev er t he p o si t io n
,
.

9 C. u r io u s ly if t h e le af b
, e t a k e n fr o m a b e g o n ia j us t r e ady
to fl o wer t he n ew p lant will fl o we r v ery q u ick ly aft er b ec o m i ng
,

es tab lis h ed b u t wi th few lea v es and litt le gro wth ; b u t if it b e

,

tak en fro m a p la nt j u s t o u t o f fl o wer the gro wth will b e great er ,

and t he perio d lo nger befo re fl o wering fo llo ws .

10 When piec es Of s tem a re s u s pend ed v ert ica lly apex u p

.
,

ward p o lar ity and gra v i ty act t o geth er ; wh en s u s p end ed ap ex

,

d o wn ward th ey act o ppo s it ely T he two fo rc es m ay to s o m e .

,

ext ent b e s eparat ed by em p lo y ing d iff eren t po s it io n s

,
Fo r ex .

a m ple if t he piec e b e h eld o b liq u ely ap ical en d t he h igh er t he

, , ,

b u d s alo ng the upper s id e will d ev elo p m o re than th o s e o n the

lo wer s id e ; and if i t b e p lac ed h o rizo nta lly all t he b u d s o n what ,

is no w t he u pp er s id e o f t he s t em will s tart b u t th o s e at the ,

apical en d will gro w m o re rapid ly .

I f h eld in an Ob liq u e po s it io n with ba s al en d h igh er d iffer ,

en t res u lt s f o llo w ; b u t in g en era l it is s h o wn that wh ere p o lar ity

1 Mo rga n , R egen era t io n p 7 4 , . .

2 It s h oul d be rem em b e re d th at t he b as a l en d o f t he ro o t is t h e en d n ea res t

th e s t em .

3 Mo rga n , R egen era t io n p 7 5 , . ,

 C A U S ES O F V A R I ATI O N
33 2

and grav ity are o ppo s ed to each o th er the fo rm er is by far t he

s tro nge r f o rc e
1
.

I I .I f a lo ng p iec e Of s t em b e s u s pend ed by bo th en d s roo t s ,

will s t art freely a lo ng t he c u r v ed lo w er s u rf a c e Of t he ben t s t em .

I f n o w t he be nt s t em b e rev er s ed ro o t s will n o t o n ly f o r m les s

,

freely b u t th ey will mo s t ly b e a lo ng t he lo wer o r u n d er s id e Of

,

t he arch t he c o nc a v e in s t ea d Of t he c o n v ex s id e o f t he c u rv e
, ,

an d c lu s t ered c lo s e r t o t he bas al en d
2
.

SEC TI O N VIII — I N T ER N A L FA C T O R S I N R EG EN ER AT I O N

Ev en a mo ng p lant s it is n o t ic eab le th a t int er n a l influ enc es

a re s tro ng ly in v o lv ed in t he regen era t iv e proc es s es I nd eed .
,

o n e po int o f d iff erenc e b et ween p la nt s and a n i m a ls is that t he

latter regen era t e d i rect ly fro m t h e c u t s u rfac e wh ile t he fo rm e r ,

regen era t e by mean s Of b u d s thro wn o u t at t he s id e and j u s t

beh ind t he po int Of s ev eranc e .

A gain i t is to b e Ob s erv ed that th o s e o rga n is m s wh o s e

,

regenerat io n is in flu enc ed by gra v ity a re t he a ttach ed s pec ies ,

th a t m ai ntai n hab it u al and c o n s tant relat io n s t o g ra v ity b u t that ,

th o se s pec ies wh ich lik e m o s t an im als m o v e a b o u t freely

, , ,

regenerat e acc o rd ing to in ter na l i nflu ences exc ept in so far ,

as v i ta l pro c es s es a re in v o lv ed th ro u gh f o o d t em pe ra t u re o r , ,

ch em ical c o n d it io n s I n o th er wo rd s t he s t u dy o f regenerat io n
.
,

is es sent ia lly a s t u dy o f int ernal f o rc es and ev en a m o ng p la nt s ,

s u bj ec t t o th e infl u enc e o f gra v ity t he i nt ern al fact o r s a re yet

t he d o m inant o n es Th ey a re well wo rth s t u dy th erefo re as

.
, ,

bearing u po n the s u bj ect o f th is ch a pt er and u po n t he inh erent

fo rc es Of o rgan ized and liv ing m att er .

Pol ari t y and het eromorp hosis I f a s h o rt p iec e b e c u t fro m

.

t he ant erio r en d o f an eart h wo r m t he m a in p iec e regen erat es

pro m pt ly the s m a ll p iec e with d iffic u lty o r n o t a t all ; and t he
,

s a me is tru e as t o t he po s t erio r en d A ga in if t he s h o rt p iec e

.
,

regenerate at a ll it d oes n o t c o m plet e it s elf by gro wi ng a n ew

,

wo rm b u t by gro wi ng a r ev er s ed lzea d ( o r ta il) lik e it s elf s o

, ,

th a t we may hav e an ind iv id u al with t wo h ead s and n o tai l o r ,

v ic e v er sa I n o th er wo rd s t he p o larity is rev e rs ed ; fo r if t he
.
,

M g 1 R g or tian ,
p 7 8 e I b
en era id pp
on, 7.
9— 8. . . 0 .
 C A U S ES O F V A R I A T I O N
3 34

If an inc is io n b e m a d e in t he
an act inia n t en tac les de
si of ,

wi ll ar is e a t th a t p o int and th ey will s eize p iec es o f m ea t and

,

pres s th em aga in s t the s tern wh eth er a m o u th wa s fo rm ed o r ,

1
not
( s ee Fig .

a p lan a rian b e s p lit length wis e int o r ight a n d left h a lv es

If ,

regen era t io n tak es p lac e wh eth er t he h a lv es b e eq u al o r u neq u al .

T he c o m p let ed wo rm will b e at fir s t b o t/z na r r ower a n d s lzo r ter

than t he n o r m al b u t with t im e and ,

feed it will eq u al o r ev en ex c eed

t h e o r igina l Ho w the n erv e c o rd s .

a n d gen ita l d u ct s a r e f o rm ed a n ew ,

es pec ia lly fro m t he p iec e so fa r t o

o n e s id e a s t o inc lu d e n o n e Of th ei r

s u b s t a nc e is a m y s t ery c lo s ely a k i n ,

t o th a t O f o rigi nal d i ff erent ia t io n

fro m t he fert ilized o v u m I f the .

s lic e b e t a k en well t o o n e s id e n o ,

h ea d m a tt er is in c lu d ed I n s u ch .

a c a s e the h ea d appea r s fi r s t at

FIG L t l t i t 1 11 3 f i l " 0 1 t he p iec e 13 11 1 13 1 3 1 11 l

43. .
g a e ra re e n era on : a c u ,

i th
n id f es ti ieo p d d a s s u m es i t s pro p er p o s it io n
an a c n an ro uce
.

l mp f t t l wh i h w l d
a c u O en a c es c ou
R e en eratl on f rom an Ob h u e
s eize a ie of m ea t and e it g q
p c e
p r ss

a ga in s t t h e s i d e o f th e bo dy , e v en su rf ace I f t he 1 3 11 01 a ta dpo le
t h o ug h no mo u t hw ere rem o v ed b y a n 0 b 1iq u e c u t
p res en t .
be ,
A ft L b
er oe
t h e regen era t ed t a il will at fi rs t
s tand a t ri g/i t a ng les to tlze c u t a n d o b liq u ely t o t he ax is Of t he

t a il A S gro wth pro c eed s h o wev er the t a il gra d u a lly s wings

.
, ,

int o lin e wi th the ax is o f t he bo dy .

I f t he h ead o f a p lan a r ian b e rem o v ed by an Ob liq u e c u t it will ,

begi n to regenerat e at the fo rward part m ak ing the h ead a t firs t ,

s tand at r ig/i t a n les to tire c u t and n o t i n lin e wi th the bo dy

g ,
.

I f a p iec e b e tak en fr o m t he m idd le o f a p la n a r ian by t wo

para llel b u t Obliq u e c u t s ( fo r ex a m ple ru n n ing back ward from ,

left to right ) the h ead will b egi n to regen era t e fro m t he a nt erio r
,

en d at i t s f o r w ard ( lef t ) s id e wh ile t he t ail will begin a t t he ,

po s t erio r extrem ity b u t o n the oppos ite s id e t he fi n al res u lt being ,

c o m p let e res t o ra t io n o f the n o r m a l s h ape .

L b Phy i l gy f t h B i
1 oe p 5
. s o o o e ra n , . 2 .
 R E LAT IV E S TA B I LI TY OF L I V I N G M ATT E R 335

T hat t he n ew m at erial
pro d u c ed in regen erat io n is at fi rs t
—
to t ipo t ent that is capab le o f m o re t han o n e d ifferent iat io n
,

is eas ily s h o wn I f a s h o rt piec e b e c u t fro m t he m idd le o f a

.

p lanarian and if aft er the n ew m at eria l has b eg u n to fo rm the

, , ,

wh o le m a ss b e s p lit length wis e bo th halv es will d ev elo p h ead s ,

d i rect ly ; b u t if the s plit is n o t mad e u nt il j u s t befo re t he

“

fo rm a t io n of a h ea d th en each h a lf p iec e pro d u c es at fi rs t a

,

half h ead that c o m p letes it s elf lat er at t he c u t s id e

,
1
.

A gai n if t he h ead b e c u t fro m a p lanarian and t he b o dy b e

,

s p lit fo r a d is tanc e t he s p lit wi ll h ea l and a s ing le h ea d will

,

regenerat e ; b u t if a s lic e b e tak en o u t o f the m idd le lin e o f t he

bo dy o r if o th er wise the t wo p iec es b e prev ent ed fro m f u s ing
, ,

th en fwo lzea a s will regenerat e o n e o n each p iec e T h ese t wo

' 2
,
.

h ead s may lat er p u ll apart with s u ffic ient fo rc e to t ear t he bo dy 8

.

A ll th es e ph en o m ena rev eal great capab ility o f readj u s t ment

as t o m o re o r less d iff erent iat ed t is s u e T he m o re t he m att er is .

s t u d ied t he mo r e we d is c o v er that t he li n e b et ween regen erat io n

and d ev elo p ment is o n e of d egree rath er than o f k ind and that ,

d iff erent iat io n in bo th cas es wh eth er n o r m al o r abno rm al res ts

, ,

u p o n ca u s es v ery i m pe rf ect ly u nd ers t oo d b u t c lo s ely ak in to

those o f d ifferent iat io n in genera l .

S ECT I O N IX — EV I D E N CE F R O M G R A FT I N G
Wh en t is s u e o f o n e k ind pla nt o r an imal is r em o v ed from
, ,

it s c o nn ect io n s and s et int o t is s u e o f t he sam e o r o f a d ifferent

k ind and a u n io n tak es place s o that gr owth en su es t he u n io n is
, ,

called a graft F ro m o u r s tandpo in t n o litt le int eres t attach es to

.

t he v ar iety o f c o nd it io n s u nd er wh ich graft s m ay b e effect ed and ,

to the fact that t he gr owth u po n t he graft is lik e t he part s et i n

and n ot lik e the t is s u e that s u ppo rt s it — t he h o s t act s o n ly a s ,

a ffo rd ing s tand ing roo m and foo d s u pply to t he graft A gain if .
,

t wo d iss im ilar p iec es a re j o in ed each pr es er v es it s id ent ity ,

.

Gra ft ing is co m pa rat iv ely eas y a m o ng plant s th o u gh t he s pecies ,

that may b e j o in ed are lim it ed A m ong an im als it is m o re d iffi .

c ult b u t by n o mean s im po ss ible

, .

M g 1
R g or ti
an , p 49 e I b
en erai d p 5
on, . .
9 . . 0 .

L b Phy i l gy f th B i
3 oe p 8
, s o o o e ra n , . 2 .
33 6 C A U SES OF V A R I AT I O N

Bo rn su cceed ed
n it ing tadpo les o f t wo d ifferent s pec ies o f
in u

fr ogs R a na esc u lm ta fo r t he a nt er io r part and R a r m /is fo r t he

,
.

po s t er io r T h is mad e u p an im al liv ed s ev ent een day s Harr iso n

.
“ -
”
.

s u cc eed ed in k eeping an ind iv id u a l m ad e u p o f R w r cs c m s and

'

R pa l u s t r zs u nt il it s tran s fo r m at io n fro m a tadpo le int o a fro g 1

'

I n t he s a m e way eart h wo rm s may b e b u ilt u p o f t wo or m o re

ind iv id u als o f t he sa m e o r o f d ifferent s pec ies and t he piec es u s ed ,

in th is b u ild ing u p may ev en b e r ev er s ed s o that t he m idd le part ,

o f t he m ad e u p wo r m m ay hav e it s po s t er io r pa r t plac ed ant er i

-

o r ly o r t he r ev er s e
,
Wo rm s m ay b e m ad e with t wo h ead s with
.
,

o r with o u t a ta il o r with t wo ta ils with o r with o u t a h ead

, , ,

th o u gh t he latt er fo r o bv io u s r easo n s can liv e b u t a s h o rt t im e

,
2
,
.

R ibbert graft ed a po rt io n o f m am m ary gland int o t he ea r o f

t he gu in ea pig wh er e it gr ew and wh en t he pig beca m e pregn a nt
, ,

t he gr aft ed t is s u e s ec r eted m i lk
3
.

The wh ole s u bj ect o f graft ing ev en m o re than that o f r egen ,

era t io n s h o ws t he wo nd erf u lly per s is t ent nat u re o f d i ffer ent ia t ed

,

t iss u e th o u gh it s h o ws als o t he v ariety o f c o nd it io n s u nd er wh ich

,

it s act iv it ies may b e d is charged .

S ECT I O N X — EVI D E N CE F R O M T H E O R I G I N OF N EW
CE LLS A N D T I SS U ES
N ew t is s u e m a y ar is e in r egen era t io n in three d is t inct way s
1 .I t m ay ar is e from t is s u e o f it s o wn k ind that is , t he t iss u e
prod u c ed in regen erat io n m ay arise fro m t he same k ind o f t iss u e
in t he o rga n is m .

2. I t m ay ar i e n o t fro m lik e t is s u e
s —,
o f wh ich n o n e m a
y b e ,

pres ent — b u t fro m t he s a m e po int o f o r igin and in t he sam e

,

m ann er a s d u ring em bry o n ic d ev elo p m ent .

3 I
. t m ay ar is e fr o m a s o u rc e ent ir ely d iff er ent fr o m that o f
em bry o n ic d ev elo p m ent and in th is ca s e m ay b e eith er n o rm al
,

o r h et er o m o rph ic .

E xa m ples o f t he fi rs t c la ss are ev ery wh ere at hand ; ind eed ,

th is is t he mo s t o rd in a ry fo r m o f regen erat io n and m any ha v e ,

erro n eo u s ly s u ppo s ed it t o b e t he o n ly f o r m O rd in a rily m u s c le .

,

1 M o r an,
g R egen era t io n , pp . 1 84 -
185 .
2 Ib i d .
pp . 1 7 1 -
1 73 .

3 L o eb , P hy l gy
s io o of t h e B ra in , p . 2 06 .
33 8 C A U S ES or V A R I AT I O N
c o u r se o f em bryo n ic d ev elop m ent is in v o lv ed in the m y s t ery o f
o rd inary d iff erent iat io n fro m t o t ipo t ent o r ind ifferent t is s u e but
regen erat ed m att er wh ich ar is es from t iss ue in n o wis e in v olv ed
i n em b ry on ic d ev elop m en t wo u ld s eem t o b e o u t s id e o f t he ia flu
enc e o f h er ed ity I n t he la tt er cas e it m ay res u lt in n o r ma l t is
.

s u e a s in t he len s o f t he e e o r in h et er o m orph ic gro wth as in

, y , ,

the prod u ct io n o f an ant enna in s t ead o f an eye o r o f a foo t in s t ead ,

o f an ant enna .

S EC T I ON XI EVI D E N C E F R O M D EV E LO PM E NT AND

D I FF E R E NT I AT I O N
F rom t he fert ilized o v u m t o t he f u lly d ifferent iat ed ad ult in
d iv id ua l o f t he h igh es t s pec ies is a lo ng s t ep N oth ing is m o re .

ev id ent than that a ll t/ ns d z i i i / f f

ze r es u lt o
' '

fi er en t a t on s t o rc es

r es i d en t wi t/t i n t/ze s in le c ell o

g f M e ov u m What ev er o ffic e is .

d ischarged by o u t s id e agenc ies and what ev er d ev iat ion s or mod i

,

fi c at ion s are prod u c ed th ereby t he i mp u lse t o d ifferent iat e and

,

t he d i r ec t i on of d ifferent iat io n aris e fro m fo rc es int erna l t o t he

o rgan is m . M o reo v er th is d iffer ent iat io n g reat as it is in t he
, ,

fi n is h ed prod u ct wh en trac ed back ward m erges by im perc ept ib le

,

s had es ea ch int o t he n ext pr ec ed ing u nt il t he u nd iffer ent iat ed,

o vu m is r each ed at t he en d o r b eginn ing ) o f t he s er ies

( .

What is the nat u re o f th es e int ernal fo rc es and what are the ,

agenc ies that s et th em in mo t io n a re t he ch ief my s t er ies in an i,

mal and p lant d ev elo p m ent T hat a s ingle ger m c ell s im ilar in
.
,

it s es s ent ia l nat u r e to any o n e o f t he t is s u e c ells o f wh ich t he

body is co m po s ed t hat s u ch a c ell can carry with it t he s u m
,

t o tal o f t he h eritage of the s pec ies that it can in t he cou r se of ,

a few day s or week s giv e r ise t o a m o llu s k o r a m an is t he ,

gr eat es t m ar v el o f b iological s c ienc e 2

.
”

“
I n att em pt ing t o analy ze t he pr o b lem s that it in vo lv es c o n ,

t in u es W ils o n ,
“
we m u s t fr o m t he o u t set h old fa s t to t he fact
o n wh ich Hu x ley in s is t ed that t he wo nd erf u l fo rm at iv e en ergy of
,

t he ger m is n o t im pre s s ed u po n it fr o m with o u t b u t is inh erent ,

in t he egg a s a h er it age fro m t he parenta l life o f wh ich it was

o r ig inally a part .T he d ev elo p m ent o f t he em bry o is n o th ing n ew .

1 W il T h C ll p 4 3
so n , e e , . 0 I b id p 3 96
.
3 . . .
 R ELATI VE ST A B I LITY OF L I V I N G MATT ER 339

I t in v olv es breach of c o nt in u ity an d is b u t a c o nt in u at io n o f

no ,

t he v ital pr oc es s es go ing on in t he par enta l body What giv es .

d ev elopment it s m arv elo u s charact er is t he rapid ity with wh ich

it proc eed s and t he d iv ers ity o f t he res u lt s att ain ed in a s pan
so brief .

We can d efin e the ch ief m y st ery of d ev elopm ent a s ly in g in

t he fact s o f d zfi er e ntzat ion an d d e i m i gr wth b
'

fi n te ter na t ion t o o u t ,

if we as k lzow t he im p u ls es go v ern ing s u ch c om p licat ed res u lt s lie

la t ent in a s ingle c ell and lzo w th ey o perat e in o rd erly s eq u en c e
,

beginn ing and end ing at the pro per moment we hav e as k ed t he ,

fi n al qu es t ion an d in approa ch ing it say s Wilso n we m ay

, , ,

as well m ak e a frank c o nfes s io n o f ig n o ran c e

”
.

A bo u t all we can h o pe to d o in t he pr esent s tat e o f kn owledge

is to throw light u po n t he q u es t io n o f c om parat iv e s tab ility o f
o rga n ized matt er and n o t e t he c o nd it io n s u nd er wh ich o n e ki nd

of c ell giv es r is e to an o th er o f a d iffer ent o rd er On th is point .

certa in fact s o f d ev elo pm ent hav e an im po rt ant bearing .

Cell div ision A ll d ifferent iat io n d u r ing d ev elop m ent is by

.

c leav age o f the germ c ell The ph enom ena o f d iv is ion and m it o
.

s is are in gen eral t he sa m e in the c lea v ag e o f t he g er m c ell and

t he d ev elop m ent o f t he em bry o as in t he d iv is io n o f o rd inary

t iss ue c ells exc ept that the body c ells for t he mos t part giv e
,

rise o n ly t o o th er s like t/zemselv es wh ile tho se of t he germ and

,

t he em b ry o gi v e r ise n ot o n ly to o th er s lik e th em s elv es b u t t o

ma ny d ifier en t k in ds as well .

T h is d is t inct io n is relat iv e ra th er than ab s o lu t e for we remem ,

b er that in c ertai n s pec ies a s m a ll part ev en a leaf is ab le by , ,

regenera t io n to gi v e rise to a n ew ind iv id u al in v o lv ing d ifferen ,

tiat io n s i mi lar if n o t eq u a l t o that o f em bry o n ic d ev elo p ment .

Ho wev er in many s pec ies and wi th m o s t t is s u es t he po wer o f

,

typical d ifferent iat io n appears to b e los t in t he firs t d ev elo p ment .

I n th is c o nn ect io n it is well t o rem ind o u rs elv es that in many

s p ec ies t he a m o u nt o f ch ro mat in m att er in t he s o m at ic c ells is

d ifferent fro m that in t he germ c ells 1

.

Geometr ical character of cleav age T he earlies t c leav age o f .

t he germ c ell is go v ern ed by t wo d efi n i t e g eo m etric princ ip les

annou nc ed by S ach s : ( r) t he c ell typically t end s t9 d iv id e int o
1 Wil T h C ll p 4 6 (
so n , e e , . i g B2 i i t ig t i
c o n c ern n A i )
o v er
’
s n v es a o ns o n sc ar s .
34 0 C A U SE S o r VA R I A TI O N

q al part s ; ( 2 ) each n ew p lan e o f d iv is io n t end s to i n t ersect

e u

t he prec ed ing o n e at a right ang le

1
.

A typ ic al c lea v age o f a s ph erica l egg wo u ld b e fi rs t v ert ical , , ,

d iv id ing int o r ight and left halv es ; s ec o nd also v ert ical b u t at , ,

right angles to t he fi rs t d iv i d ing into d o rs al and v entral po rt io n s ;

,

th ird h o rizo ntal d iv id ing int o anterio r and po s t erio r part s ;

, ,

aft er wh ich all s o rts o f irreg u larit ies m ight b e expect ed inc lu d ing ,

c leav ages para llel with t he s u rfac e c u tt ing in two the lo ng c ells ,

that f o rm erly ext end ed t o t he c enter The egg o f t he ho lo thu .

rian lik e those o f S ynapta pro c eed s regu larly u n t il as many as

, ,

5 1 2 c e ll s ar e re ac h e d 2
w h i le o th e r,
s b e c o m e i r r e g u lar a lm o s t at
o nc e s o m e q u adrant s d iv id i ng m u ch m o re ra p id ly than o th ers
, .

V ar iat ion in t he rat e of div i sion I f d iv is io n pr o c eed s wi th .

p erfect reg u larity t he n u m ber o f c ells will o f c ou rse fo rm an

,

increas ng g i e o m et ca
r i l s er i e s w h e r t io is t wo
o s a — 2 4 8 16 , , , , ,

3 2, 6 4 1 2 8
,
2 5 6 5 1,
2 1 0 2,4 e t c S ,
u ch a s e r i e s h a, s b ee n . m e n

tio n ed ext end i ng t o 5 1 2

,
.

T h is u n if o r m s eries is rarely rea lized h o wev e r o wing t o i rreg , ,

u la r it ies i n t he rat e o f d i v is io n ; fo r exa m p le N ereis regu larly ,

giv es ris e t o the s eries 2 4 8 1 6 2 0 2 3 2 9 3 2 3 7 3 8 4 1 4 2 , , , , , , , , , , , ,

“
after wh ich t he o rd er is m o re o r less v ariable 3
.

I n s o m e po rt io n s o f t he d iv id ing c ell t he c lea v age pro c eed s

th erefo re with m u ch great er rap id ity than in o th er s n o r is t he p lan ,

u n if o r m i n all c as es th o u gh t he res u lt s ach iev ed m ay b e s u b

,

s t a n t ially id ent ical I n gen e ral t he rat e o f d iv is io n is m o s t rap id

.

in t he u pper h emis ph ere o f t he o v u m and in s o m e in s tanc es it p ro ,

c oed s v ery s lo wly with lo ng pau s es in t he lo wer giv ing great

, , ,

irreg u lar ity in t he S ize o f c ells .

Temporar y effec t of ou t side influ en ces u pon cleav age D ries ch .

p lac ed eggs o f sea u rc hin s u nd er pres s u re s u ffic ient t o flatt en the

s ph eres to d is k s I n th is po s it io n the a m ph ia s t er s a ll a s su m e
“
.

t he po s it io n o f lea s t res is tanc e i e para llel t o t he flatt en ed s id es ,

. .
,

and the egg s eg ment s as a flat p la t e o f eight sixt een o r th irty , ,

t wo c ells T h is is t o ta lly d ifferent fro m t he n o r m a l fo rm o f

.

c lea v age yet s u ch egg s w/zen relea s ed f r om p ress u r e a r e capa ble

o f d e v e lo
p m e n t a n d g i v e r i s e t o n o r m a l e m b f
y os
4
W i th ou t .
”

“ il
1 T h C ll p 3 6
'
so n , e e , I b id p 3 66
. 2 .
3 . . .

I b id p 3 64
2 . . . I b id p 3 7 5 I t li mi 4 . . . a c s a re ne .
 C A U S ES o r V A R I AT I O N
34 2

part s of the bo dy ) the sec o nd s ett ing o ff d o rs a l and v ent ra l

,

areas ; wh ile t he t hird whic h is h o rizo ntal d iv id es int o ant erio r

, ,

and po s t erio r p o rt io n s .

T he proc ess is t he s a me in m any s pec ies and wh erev er t he ,

“

egg ax is can b e d et er m in ed by t he acc u m u lat io n o f t he d eu t o

plas m in t he lower Item isp /zere t he egg n uc leu s s oo n er o r lat er lies

ecc entrica lly in t he upper lcem isp /cer e and the p o lar b o d ies a re ,

fo rm ed at the upp er p ole .

”

I n s u ch cas es o f d is t inct p o larity the c lea v age p lan es are prae .

t ically pred et erm in ed and t he prod u ct s o f d iv is io n ha v e each

,

t hei r d efi ni t e role in d ev elo p m ent Thu s the u pper h em is ph ere .

so c a lled an im a l po le) is t he s eat o f m o s t act iv e d iv is io n wi th

( -
,

s m aller c ells wh ich d ev elo p int o c erebra l d o r s a l and ant e rio r

, , ,

po rt io ns o f t he bo dy ; wh ile the lo wer h emis ph ere ( v egetab le o r

n u t rit iv e pole) d iv id es m o re s lo wly it s c ells a re larger and they , ,

d ev elo p int o the alim entary o rgan s and t he p os t erio r and v entral
pa rt s gen erally 1
.

W h ile th is ru le is n o t ab solu t e in a ll s pec ies it yet ind icat es a ,

broad gen era l princ ip le that lies at t he v ery thres h o ld o f d ev el

o p m en t n a mely t hat t he or igi n a l i mp u ls e to d i r ec t ion of g r owtlz
,

lies i n t/ze p ola r i ty f

o Me

Cau se of polarity in t he ov u m Grav ity wo u ld s eem to b e t he .

co ntrolling cau s e in es tab lis h ing a k ind o f po larity in t he o v u m .

T he n u c leu s being o f lo w s pec ific gra v ity t end s to lie ecc entrica lly ,

nearer the u pper s id e o f t he o v u m and t he h ea v ier d eu t o p las m ,

s ett les t o t he lo wer s id e t he part s arrang ing t h em s elv es acc o rd

,

in g to relat iv e weight lik e s tarch gran u les o r o th er c ell c o nt ent s

, .

M o reo v er Bo rn ha s s h o wn that if fro gs egg s b e fa st en ed in “ ’

an ab n o rm a l po sit io n — e g u p s id e d o wn o r o n t he s id e
,
. .
,

rea r rangement o f t he egg m at erial tak es p lac e a n d a n ew ax is “

is es ta b lis /zed i n t/ze egg 2

S ch u ltze d is c o v ered that if t he egg
.
“

b e t u rned u p s id e d o wn wh en in t he t wo c elled s tage a wh o le -

em bryo ( o r ha lf o f a d o u b le em bry o ) m ay ari s e fro m each b las t o

m ere in s t ead of a ha lf em bry o a s i n t he n o r m a l d ev elo p ment

, ,

and that t he ax es o f th es e em bry o s s h o w n o c o n s tant relat io n

to o n e an o th er
2
M o rgan lea rn ed t hat eith er a half emb ryo
“
”
.

W il1 T h C ll pp 3 7 8 3 7 9
son , e e , .
, .

I b id p 4
9 . . 22 .
 R E LAT I VE STA B I LI T Y o r L IV I N G M A T TE R
343

or a wh o le h f s ized d warf
al -
m ight b e f o rm ed , a cc or d in
g to tlze
l
os i tion o tb e b la s tom er e
f
”

p .

Cau ses of differen ti ati on . T h es e fact s s h o w

1 . T hat a primary ca u se o f d ifferent iat io n lies in t he po la rity
of t he ov u m .

2 . T hat
th is polarity is d u e p ri marily to gra v ity sepa ra t ing
it s h eav ier fro m i t s light er part s .

3 T hat
. t h e e gg is at fir s t i n m any ca s e s at le a s t i nd iff e re nt , , ,

and that its po larity may ev en b e changed aft er h av ing o nc e

been well es tab lis hed .

4 T hat
. if t h e s e g m e nt i ng o v u m m a
y b e s e pa rat e d i nt o i t s

bla s t o m eres and each may pro d u c e a c o mp let e ind iv id u al (A m

,

p h io x u s th
,
en t he e gg as a w h o le is n o t o n l y t o t i p o t e nt
b u t it s earlier s eg ment s a s well a re each capab le of prod u c ing
a ll t he part s o f t he b o dy .

5 T h e
. p o lar i ty and t h e d ev e lo p m ent o f a part a r e i nfl u e nc e d
largely by it s p os i t ion with r ef er enc e to otlzer a n d esp ec ia lly
la rg er p a r ts .

R o u x f o u nd that when in t he t wo c elled s tage of the frog s egg ’

-
, ,

o n e o f t he b la s t o m eres wa s k illed by a ho t n eed le t he r ema in ing ,

o n e d ev elo ped a ha lf em b ry o h erea s in many cas es a s in gle

8 w
,

blas t o mere is kno wn to b e ent irely capab le o f pro d u c ing a wh ole

em bry o z ff r eed f r om i ts n ez /zb ors
g .

A ga in a sect io n fro m an earth wo r m t end s t o regenerat e

,

head matt er at it s ant erio r en d and ta il matt er at its pos t erio r

en d ; b u t if a s m a ll p iec e b e gra ft ed ev en i n a r ev er s ed p os i tion , ,

o n t he ant er io r en d o f a larger p iec e it will regen erat e h ead ,

m att er f r om i ts pos ter ior ex t r em ity s h o wing ho w t he p o lari ty o f ,

t he s m a ller p iec e is s o to s p eak o v erc o m e by that o f t he larger

‘
.
, ,

A c o n s id erat io n o f th es e fact s lead s W ils o n t o q u o t e Hert wig

a s f ollo ws : T/ze r ela t i v e p os i tion of a b la s tom er e i n the whole

a s s em b lag e d eterm i n es i n ener a l wiza t d ev elop s f r om i t ; if i ts

[ ] g
b e clea ng ed , i t g i v es r is e to s omething d ifler en t in other
'

p os i t i on
5
word s , its p r ospec ti v e v a lu e is a f u nc ti on of i ts p os i t i on .

1 The d iff eren t c e ls o f l t he ea rlier c lea v ages ( t he 2 4 1 6-c elle d s tages , et c .

)
a re k n o wn as b l as t o m eres .

2 W ils o n , The C e ll , p .
42 3 .
4 M o rga n R egen erati o n , p 8
, . .

3 Ibi d .
p 3 8
.0 .
5 W ils o n , T he C e ll , p .
41 5 .
3 44 C A U SE S OF V A R I AT I O N

6 Wh eth er the ab ov e s ta t em ent is ab s olu t ely o r o n ly rela

.

t iv ely tru e t he fact is c lea r th a t liv ing m a tt er will o ft en go

,

throu gh its changes a nd c o m p let e it s cyc le o f d ev elo p ment

u nd er extrem e hard s h ip F o r exa m p le in s o m e s pec ies a b la s
.
,

t o m ere fro m t he t wo o r fro m t he f o u r c elled s t a ge m ay d ev elo p-

i nt o a wh o le ( d warf ) em b ry o o r i nt o a half em b ry o wh ile in ,

o th ers ( ct en o ph o r e) each o n e o f t he fi r s t eight m a y d ev elo p i nt o

a sy m m etrical and th eref o re c om p let e ind iv id u al b u t la ck ing tlze ,

1
u ll nor m a l n u m b er o s w i mm ing l
f f p a tes .

7 . T he c o nc lu s io n is na uv o id a b le that liv ing m att er is end o wed

with wo nd erf u l ela s t ic ity and p ers is t enc e o f p la n o r an a pp ro xi ,

m at io n t o it ev en u nd er m o s t ad v ers e c o nd it io n s pro v id ing o n ly

, ,

th ese c o nd it io n s d o n o t d es troy life .

A n d s o we ret u rn t o t he o r iginal q u es t io n s : Ho w m u ch o f
what t he ind iv id ual is t o b e is d u e t o inh eritanc e ? Ho w flex
ib le is t he p lan ? To what ext ent a re m od ificat io n s p o s s ib le and ,

t o what ext ent a re v ari at io n s i nh er it ed

Su mmary I n gen era l t he fact s o f d iff erent iat io n and o f regen
.

c ra t io n t o g eth er with t he p er s is t enc e o f an es tab lis h ed typ e in

,

carrying f o r ward inh erited q u alit ies ev en u nd er m o s t ad v erse

c o nd it io n s ind icat e extreme s tab ility o f liv ing m att er
,
.

O n t he o th er hand t he fact s o f acc lim at izat io n argu e fo r t he

,

alt erab ility of pro t o pla s m bo th as to c o n s t it u t io n and a s to fu nc

,

t io n inc lin ing o n e s tro ngly t o t he b elief that s u ch a lt erat io n may

,

bec o m e m o re o r les s per m an ent with t he o rgan is m .

A lt o geth er t he c o n v ict io n is f o rc ed u po n u s that t he act iv i t ies

o f liv ing m att er pr o c eed u p o n a p lan in s p i red fro m with i n and

aris ing fro m the nat u re of the o rgan izat io n b u t th a t th is plan ,

has acc o m mo dat ed it s elf t o su rro u nd ing c o nd it io n s in the pas t

and is ent irely c om pet ent t o d o so a ga in wh en ev er su ffic ient
occas io n a ris es pro v id ed o n ly that t he n ew d em and s b e n o t t o o
,

s weep in g o r t o o s u dd en ly im p o s ed .

T h ese fact s n eed t o b e c o n s tant ly I n t he m ind o f t he farm er ,

who m u s t b e prepared fo r c o m pa rat iv ely s weep ing ch a nges fro m

apparent ly s light c a u s es .

We pas s n o w t o a d is c u s s io n o f t he q u es t io n a s t o wh eth er o r
n o t ind iv id u a l m o d i fica t io n s m ay b e tra n s m itt ed .

“ ill '
Th C ll p 4 8
so n , e e , . 1 .
 PA R T I I I T R A N S M I S S I ON

U p to th is p oint the s t u dy has been c o nfined to the na tu r e

and kin d s of v ariat io n s that may arise t ogeth er with t he ca u s es
,

that lead t o th eir appearanc e We c o me n o w to inq u ire wltetlter

.
,

and to wlza t ex ten t th ese v a riat io ns are tra ns m is s ib le

,
.

Th is is t he all im po rtant q u es t io n b eca u s e v ar iat io n s t hat

-
,

a re n o t t ran s m i tt ed a re m an ifes t ly o f n o c o n seq u enc e exc ept

t o t he i nd iv id u a l Wh ether d es i ra b le or u nd es irab le th ey hav e

.
,

n o o ppo rt u n ity to a ffect t he rac e a s a wh o le eith er fa v o ra b ly

or u nfa v o rab ly . We may t heref o re d is rega rd in all q u es t io n s of ,

rac e i m pro v ement any and a ll d ev iat io n s that a re n o t tra ns

,

miss i b le
.

Th o s e that are tra n s m itt ed h owev er a re by that fact d es t ined

, ,

t o ex ert a mo re o r les s per man ent infl u enc e for goo d o r ev il .

A cc o rd i ngly we cann o t kno w too mu ch a b o u t th is c la s s o f v ari

at io n s and t he c i rc u m s tanc es u nd er which th ey d es c end fro m

genera t io n to generat io n and u lt im ately c ome to characterize if ,

n o t t o d o mi nat e t he t yp e o f t he rac e
, .

T he s t u d ent a n d t he breed er a s well is lik ely t o bec o me c o n

, ,

f u sed by t he c easeles s pan o ra ma o f characters and v ariat io ns

present ed to his v iew as generat io n s c o me and go in v o lv ing ,

s o met imes li t era lly m u lt i t u d es o f i nd iv id u als o f all s had es o f

differenc e He m u s t learn t o pas s th is pro c es s io n befo re his

.

menta l v is io n with t he f u ll realizati o n that a large p o rt io n o f all

that he sees will ha v e n o perm anent infl uenc e u po n the rac e .

He m u s t bec o me ac u t e in d et ect ing the s igni ficant fact o r s wh ich ,

a re th o se o n ly that a re c o nn ect ed wi th t ran s m is s io n .

T he s t u dy o f rac e im pr o v ement is th erefo re es s ent ia lly a

, ,

s t u dy o f t he la s th a t go ern tran s m is s io n
w v — a ll that ha s go n e
,

befo re being preparat o ry to t hat s tu dy U po n t wo q u es t io n s .

t he breed er m u s t fi x his att ent io n Wita t is tra ns m i tted a nd

, ,

b ow is tb c tr a ns miss ion a cc o m
p l is hed
347
 C HA PT E R X I

TR ANSMI SSI ON OF MODIFI CA TI ONS DU E TO EX TER N A L

INFLU ENCES

S ECT I O N I I NT R OD U CTO R Y
d is c u s s io n o f the ca u s es o f v a riat io n a c lear d is t inct io n
I n t he
was m ad e bet ween ca u ses i nt ern a l and cau s es ext ernal t o t he
o rgan is m T h is d is t i nct io n s h o u ld s t ill b e b o rn e in m ind th o u gh
.
,

t he pr inc ipa l d is c u s s io n and all o f t h e c o ntro v er s y ar is e in c on

n ec t io n with e xt ernal ca u s es .

V ar i ati on s du e t o ca u ses in tern al t o t h e ger m ar e t ran s mitt ed .

So far as t he writ er is aware n o o n e has ev er d is p u t ed o r ev en , ,

q u es t io n ed t he c o rrectnes s-o f th is pro po s it io n I t is f u nda ment al

,
.
,

if n o t ax io m at ic that any change tak ing p lac e in t he s t ru ct u re

,

o f t he g er m inal m att er wh ich is pa s s ed o n fro m gen erat io n

,

to gen erat io n wh ich is t he b earer o f a ll chara ct er s and t he

—
,

exc lu si e b as is o f all tran s

v m is s i n
o that any su ch alteratio n ,

will o f n ec es s ity m ak e it s elf m an ifes t at o nc e and ind efi n it ely ,

af t er ward ; ind eed as lo ng a s t he change in t he c o n s t it u t io n of

,

t he ger m c o nt i n u es T h is is t he p u rpo s e o f a ll s elect iv e m at ing

.
,

— to s ec u re a germ end o wed with as m any as p o s s ib le o f what

a re c o n s id ered to b e d es i rab le v ar ia t io n s t end ing to m ax im u m ,

d ev elo p m ent and as few a s po s s ible that are u nd es irab le and

, ,

th es e with a m in im u m d ev elo p m ent S u ch a germ s h o u ld .

d ev elo p a n embryo and fina lly an ad u lt ind iv id u a l with t he

, ,

h igh es t o btai nable d egree of exc ellenc e .

T o c o ntro l t he ger m is t he p u rp o se o f a ll s elect io n I t i s t he .

o bj ect o f a ll breed ing W e p ract ic e lin e breed ing and ev en

.
,

in breed ing t o g iv e i n ten s i ty a lo n g c erta in lin es ; we reso rt t o

-
,

mix ed breed ing ev en t o cro s s breed ing t o intro d u c e n ew v ari

, ,

a tio n s and t o end o w t he rac e with great er flex ib ility .

I t is in t he ger m th a t m u tat io n s aris e T he ca u s es a r e .

o b s c u re,
b u t it is h ere in t he c o n s t it u t io n o f t he ger min al
,

348
35 0 TR A N SM I SSI ON
S t rict ly peak ing th erefo re it is t he charact er and n o t it s
s , , ,

m od ificat io n wh ich is tra n s m itt ed and what we d es ire t o kn o w

, , ,

is wh eth er t he mo d ificat io n o f a chara ct er in a part ic u lar ind i

,

v id u al t end s to b ec o m e pe r m an ent ; that is t o s ay wh eth er a ,

charact er that has u nd ergo ne m od ificat io n will b e tran s mitt ed in

it s mod ified o r in it s u n m od ified f o r m .

I n th is c o nn ect io n t he s t u d ent m u s t b e rem ind ed that we

ha v e n o way o f j u dg ing what charact ers o r what m od ificat io n s
a re b orn int o an i nd iv id u a l exc ept by t he d ev elo p m ent th ey

atta in in his o wn perso na lity aft er reach ing t he a d u lt s tage 1

F o r exa m p le wh en n o t ing an a v erage ind iv id u a l w

.

e cann o t s a
, y
wh eth er he is o n e that wa s exc ept io n a lly well b o rn with fa ir ,

o pp o rt u n it ies fo r d ev elo p m ent ; wh eth er he was o n ly fa irly well

b o rn b u t with exc ept io nal o ppo rt u n it ies ; o r wh eth er he is an

,

av erage b o th as to b irth and o ppo rt u nity A ll three c o m b ina .

t io n s wo u ld pro d u c e abo u t an av erage ind iv id u al .

On t he c o ntra ry if he b e an exc ept io nal ind iv id u a l we a re

, ,

fairly s afe in as su m ing that he was bo th well bo rn and well c on

d it io n ed b u t if he b e o n e o f the lo wes t ind iv id u a ls that he was ,

bad ly bo rn and has liv ed u nd er hard c o nd it io n s b o th o f wh ich , ,

o p erat ing t og eth er m ak e ev en an av era ge d ev elo p ment im po ss ib le

,
.

Of a m at u re ind iv id u a l we may s ay fi rs t that a ll t he ev id ent , ,

character s o f wh ich he s tand s po s s es sed were bo rn int o him ,

b u t that t he d ev elop men t th ey ha v e atta in ed is partly a m att er

o f b i rth and part ly a m att er o f t he ext erna l c o nd it io n s o f life .

His d i ff erenc e in d ev elo p ment as c o m pared with the mean o f t he

rac e tak en at t he ad u lt s tag e is his d ev iat io n o r v a riat io n ( fo r

we u s e t he t er m s int erchangeab ly ) and t he q u es t io n we a re n o w

,

a sk ing is A re th ese ind iv id u a l d ev iat io n s tran s m itt ed o r are all

, ,

chara ct ers tran s m itt ed acc o rd ing to a d ead lev el o f inh erita nc e ,

lea v i ng all d ev iat io n s t o b e acc ou nt ed fo r a s m att ers of i nd iv id u al

atta in m ent throu gh m o re o r les s s u cc ess f u l d ev elo p ment ?
Adaptati on s T he m od ify ing eff ect o f t he c o nd it io n s o f life
.

is s o pro f o u nd that in nat u re ev ery wh ere bo th a ni m als and

p lant s har mo n ize with th eir en v iro n ment alm os t perfect ly t hu s ,

1 W h en t he d d l
i n iv i ua d h d l
b ec o m es a b ree er his in erit e q u a it ies wi l b e fa irly l
we ll i d i
n c a te d . h
b y h is p o wers o f t ran s m is s io n I n t is wa y an an im al tru may ly
b db
ree et t er h
t a n h e is im s e f h”
l .
 T R A N S M I SS I O N O F M OD I F IC A T I O NS 1
35

giv i n g the appea ranc e o f ha v ing been s pec ially creat ed fo r th ei r

pa rt ic u lar s u rr o u nd ings W e know en o u gh o f t he ca u ses of
.

v a ria t io n h owev er t o realize t hat th is fi t b et ween the animal

”

, ,

o r t he p la nt and i t s en v ir o n ment has b een br o u ght abo u t by the

d i rect act io n o f o u t s id e c o nd it io ns u po n o rganis ms s o mewhat

plas t ic and capable o f bec o ming adapt ed to a wid e range of
c irc u s tances
m ,
— all i nd iv id u als no t po s ses s ing th is adaptab ility

hav ing lo n g s inc e d is appeared by nat u ral s elect io n .

N o th ing is mo re s im p le and nat u ra l than t o a s su me t hat

wh en t he i nd iv id u a l has acq u ired s o me mod i ficat io n thro u gh
t he i nfl u enc e o f t he en v iro n m ent it will t ran s m it th is m o d ific a
t io n t o i ts d es c endant s and that what was at firs t im pressed
,

fro m with ou t grad u ally bec o m es hered itary exh ib it ing all t he ,
'

c u m u lat iv e effect o f t ran s m is s ib le q u alit ies A ll appea ranc es .

a re in fav o r o f s u ch an a ss u m p t io n and it is t he S im p les t and

,

mo s t d irect exp lanat io n o f t he ph en o mena o f adaptat io n an d o f

t he well kn o wn ha rm o ny that alway s ex is t s b et ween a s p ec ies
-

and its en v iro n m ent .

B u t th ere are at lea s t tw o ot/zer meth od s by wh ich the en v i

ro n m en t im pr es ses it self s tro n g ly u p o n t he s pec ies — bo th of ,

wh ich are alway s at wo rk b o th o f wh ich ach iev e large res u lt s

,

o f an exact ly s im ilar charact er and appearanc e and wh ich m u s t ,

th erefo re b e eith er s u btract ed o r f u lly acc o u nt ed fo r b efo re we

a re warra nt ed in as s u m in g r es u lt s d u e to d irect t ran s mis s io n .

The fi rs t o f t hes e is t he d i rect effect o f t he c o nd it io n s o f lif e

act ing separately u po n all t he ind iv id u als o f each gen erat io n
and all in the sa me d irect io n T h is has a ll the appea ra nce o f
.

tra n s m issio n b u t it is n o t c u m u lat iv e and can brin g ab o u t n o

, ,

better adaptat io n in the s pec ies and no great er t o tal change

, ,

tha n can b e wrou ght for each ind iv id u al d u r ing its lifet ime .

The o th er is d u e to t he select iv e effect o f t he en v i ro n ment ,

wh ich is c u m u lat iv e and s o far as we can s ee ab u ndant ly ab le

, , ,

to acc o u nt fo r al l t he ph en o mena o f adaptat io n and fo r t he f u ll

effect s o f en v i ro n ment .

Th e en v i ron men t al ways sel ecti v e So m e in d iv id u a ls of ev ery

.

gen erat io n fail u tt erly t o end u re t he c o nd it io n s o f lif e and th u s ,

failing th ey leav e n o d esc endant s The n ext generat io n th ere

,
.
,

fo re bein g d esc end ed fro m the mo re adaptab le ind iv id u als is

, ,
35 2 TR A N SM SS I ION

m o re n early in harm o ny with t he en v iro n ment and th is c u m u ,

lat iv e infl u enc e c o n s tant ly ex e rt ed gen era t io n a ft er gen era

, ,

t io n s lo wly b u t s u rely m od ifies the ra c e i n tlze d i rec t io n of tlee

,

en v i r on men t gi v ing aga i n t he a pp ea ranc e o f inh er it ed m od ifi c a

,

t io n s N o w in n a t u re th is selec t iv e in flu enc e is a lway s at wo rk

.
,

so met im es by act u a lly d es tr o y ing t he les s adapt iv e ind iv id u als ,

m o re o ft en by a ff ect ing n o t life b u t fert ility o r lo ngev i ty o r , ,

b o th thro u gh litt le n o t ic ed b u t in s id io u s agenc ies

,
-
.

T h is is m o reo v er t he m o s t p o werf u l o f all t he mean s by

, ,

wh ich en v iro n m ent in flu enc es s p ec ies I t ent irely o u t s t ri p s t he .

res u lt s o f d irect in flu enc es u po n ind iv id u a ls and is f u lly ab le ,

t o acc o u nt fo r all o rd in a r y c a s es o f en v i ro n m enta l in flu en c e

1
.

Th e sou rce of t h e d iffi cu l ty Here is an ev er pres ent all .

-
,

po werf u l in flu enc e b end ing s pec ies int o h a rm o ny with th eir en

,

v iro n m en t and i t s effect s m u s t b e f u lly el im i n a t ed o r acc o u nt ed

,

fo r befo re we can d et er m in e wh eth er a res id u e rem a in s t o b e

attrib u t ed to d irect t ran s m is s io n T h is is the s o u rc e o f t he .

greates t d iffic u lty in att em pt ing t o lea rn wh eth er m o d i ficat io n s

a re d ir ect ly t ran s m i tt ed .

T h is s elect iv e pro c es s o f t he en v iro n ment res u lt s in t he i nd i

rect trans m iss io n o f s u ch m od ificat io n s as increa s e t he chanc es
o f the i nd i v id u a l in t he s tr u gg le fo r ex is t enc e ; b u t what fa rm er s

want t o kn o w is wh eth er m o d ificat io n s pro d u c ed by t he c o n

d it io n s o f lif e a re d i r ec t ly t ra n s m i tted a s s u e/t ind epe nd ent ly ,

o f t he q u es t io n wh eth er th ey in cr ea s e o r d ecrea s e t he C hanc es

o f life in t he i nd iv id u a l and ind epend ent ly o f all q u es t io n s o f ,

s elect io n .

1 l e i n d iv i d u a l and tlze typ e . Th e s tu dm u s t b e c ear as

en t l to w h a t c o n s ti

t u t es t y i d i id l m t b
yp e
. E v er d d h
n v ua t t l
us t tw e c o n s i ere wit referen c e o a eas o

ge n era ti th t wh i h it b l g
o ns , d th
e one o t T h t yp c f t
e on
y s an e n ex . e e o a ra c e a an

p a r ti l m m t i fi d b y th p
cu ar o enl q l it i f l l i t d l t m m b
s xe e d
e rs o n a ua es o a s a u e e rs , a n

fo r thi p p lsl i d i
ur id l f
o se a
q l w ig nh t vd h
ua s t d iff
a re o e ua e , an one c a ra c er o r er

en c e i g d
s as t h E y
oot h i g as tanoi fi i g t h
er .t yp f v eri t i g n c o un s n x n e e o a n ex s n

en era tio n
g .

But w h th en e n ex t g en er a t ion is c o ns i d ere d it i

, s no t so . On ly th o s e in d i id
v ua ls

co un t wh i h c s u c c ee d in r e r od u c i n g ,
p and on l y th o se d iff e ren c es c oun t th t a are

t r a ns m i s s i b l e . T h e i n i tia l o r n a tu r a l y
t p e, th f
e r e o re, as s ec u r e d by t ra n s m is s io n ,

is so me h d d
w a t iff e ren t in s u c c ee i n g gen era t io n s , n o r is it ev er fu exp ress e in ll y d
a d lt
u . l y
m emb ers T h e ex is t in g a n d v is ib e t p e o f a n y ra c e is t erefo re, a t b es t ,
h
b ut an in c o m p l e t e ex
p res s io n o f it s p o s s ib il it i es .
354 T R A N S M I SS I ON
as proo f o f t he inheri ta nc e o f acq u i red chara ct ers t he s t u d ent ,

m u s t b e o n h i s c o n s t a n t g u a rd a n d he m u s t b e ex trem ely c ar e ,

fu l in t he gen e ra liza t io n s he al lo ws h i m se lf t o m a k e in t hi s pa rt i e
u la r fi e ld O f s t u d
y .

T hi s s u bj ec t ha s b ee n gr ea t ly c o m p lica t ed b v a mas s o f tra d i

tio n t ha t has gr o wn u p aro u nd it T he a v era ge ma n a s s u m es .

th a t t he effect s o f en i ro n m en t a re t ra n s mi t t ed He d oes n o t
v .

s t o p t o i nq ui re se ri u s ly i n t o t he s u bj ect He c o n s id ers a ll
o .

d o u b t o n t h is p i n t a s f oo lis hnes s ; b u t t ha t wh ich he c o ns id ers

o

as g d pro o f is in m a n v c a s es a t leas t anyt h i ng b u t proo f

oo , , .

A s if t h is were n t en o u g h t he n a t u ra lis t o n h is pa rt has s t il l

o ,

f u rt he r m u dd led t he m a t t er b v a m o s t u nf o rtu na t e an d c o n
f u s ing n o t t o sa y ina cc u ra t e u s e o f t e rm s T ra d it io ns m ay b e
, ,
.

ign o red fo r p u r po s e s o f s t u d v b u t n o t so wit h t erm i no lo gy , ,

wh ich s h o u ld b ex a ct a n d a cc u ra t e T he t wo m o s t u n f o rt u na te
e
.

t e rm s t h a t e e r c rept i nt o ev o lu t io na ry s t u d ies ar e
v c o ngen i tal
an d a c q ui red .

Char act ers con gen i tal an d acq u i red I t is a c u s t o m wi th .

ev o lu t io n is t s t o as s u m e t h a t ev e ry a d u lt i nd iv i d u al is i n po s s es

s io n o f t w o k i nd s o f c h a ra c t e rs : fi rs t t h o se whi ch were b o rn ,

in t o i t ( t he c o n gen i t a l sec o nd t h o s e w h ic h were f o rc ed u po n ,

it b v t he c o nd it io n s o f life o r p ic k ed u p as t he res u lt o f ex peri

enc e ( t he a cq u i red ) T hev t h en proc eed t o t he v e ry n a t u ral
.

as s u m p t io n t ha t t he c o n gen i t al c h a ra c t ers h a in g been d eriv ed ,

v

b v inh eri t a nc e wi ll in t u rn b e t ra n s m it t ed ; a f t e r whi ch t hey

,

ra ise wh a t wo u ld see m to b e t he fin al q u es t io n A re t he ,

a cq u i red ch a ra c t e rs t ra n s m it t ed T h u s is t he fi eld Of d isc u s

?

s io n s t ak ed o u t a n d n o s u c h b a t t le ro v a l ( o f wo rd s ) was ev e r

fo u gh t o u t in m od e rn t im es — exc ept o v er q u es t io n s t heo lo gic al

,

— as has b en waged o er t he q u es t io n o f t he in h eri tan c e (o r

e
v

t ra n s m iss io n ) o f a cq u ired c h a ra ct e rs
‘
.

1Th i i s h q s ti h h d i id d h
e u es t o n L m t kiat asd th v e t e n ee - a a rc an s an e n eo

D wi i
ar d lm
n a n s. an h i a h f l i
os t t i f h l
e e n t re w ty y ost o ev o u t o n s ts or t e ast t en ea rs

h av e
g
ra nd h m e l t e id s h
e v es h f h
oni dione
p sd h e or t e ot er o t s s u te an av e

a ll w d h m l
o e t e b id fi d wi h
s e v es t o h e h f h h
en t i il
e mp t one or t e ot er o t e os t e ca s.

As t h e s pec ia l o p po n en t o f t h e t h eo ry o f t h e t ra n s m ixs i o n o f a c q u i red c h ar

ac t ers . t h e s t u d en t s ho u l d rea d W ei s m a n n pa rt ic u la rlv h is E ssa v s

. on Heredi ty ,

h is G erm Pla s m . an d h is G e rm i n a l S e ec t io nl .

Th e m os t v l
igo ro u s a n d a c c es s ib e if n o t t h e m os t a b e c a m p io n o f th e
. l . h
t ra n s m is s io n is t s is p e r h a ps R o ma n es in h is Ex a m in a t io n o f “ e is m an n is m a nd
'
 T R A N SM I SS I O N O F MO D I F I CA T I O N S
35 5
The w ri t er d oes
pro po s e to ent er t he d is pu t ed t errit o ry
not

at th is po int fo r the reas o n that he d oes no t believ e in acc ept

,

in g fo r th es e p u rpo s es t he f u nda m ental d is t inct io n bet ween

“
c o ngen ital and acq u ired chara ct ers The u s e o f the term “
.

“
c o ngen ital as d is t inct fro m acq u ired is mos t u nfo rt u nat e
“
.

A n y charact er pres ent at b irth is by d efin it io n c o ngen ital T he , ,

.

lis t wo u ld inc lu d e n o t o n ly t he cha ract er s typ ical o f t he rac e b u t

als o ma lfo rmat io n s and d efo rm it ies d u e to c o nd it io n s i n u tero o r
t o u nkn o wn ca u s es T h u s m en ha v e b een b o rn with o u t feet yet
.
,

t he chanc es o f tran s m i tt ing t he d ef o r m ity a re extrem ely remo t e .

A n a rm o r a leg m ay b e m is p lac ed d u ring em bry o n ic d ev elo p

ment and m alfo rm at io n s res u lt wi th o u t es s ent ia lly bea ring u po n
the c o nd it io n s that a re s u ppo s ed t o c o ntro l i nh eri tanc e .

M an ifes t ly t he m o s t pro fitab le d is t inct io n to Ob ser v e regard

in g t he i nh eri tanc e o f v ariat io n s is a s t rict d is c ri m inat io n b e
t ween th ose that ha v e arisen t/zr o ug /z ca u s es afi ec tzng t/ze g er m
’

p la s m d i re ct ly o n t h e o n e h a nd ( b l
,
a s t o g en ic ) and th os e that ,

affec t t he bo dy d u ring it s d ev elo p m ent u po n t he o th er ( so mat o ,

g e n i e ) N o w. an acc id ent t o t h e e m b r y o i n u te r o is a s m u ch
ext erna l t o t he g er m p la s m — t he inh er it ed s u b s tanc e o n wh ich

v l —
d e e o p ment d epend s as is an acc id ent after b irth and th ere ,

fo re d is t inct io n s b et ween c o ngenital an d acq u ired charact ers a re

extremely m is lead ing es pec ia lly a m o ng m a mm a ls ; ,
les s s o o f ,

c o u rse a mo n g b ird s in wh ich n o s u ch th ing as b irth ex is t s

, ,
.

h is D a rwin an d A ft er D a rwi n . C o p e, in his Prim a r yF a c t o rs o f O rga n ic E v o l u t io n ,

may b e d wit h p fi t
rea ro , as m a y Eim er in O rga n ic E v o l u t io n .

Bo th f ti l im t
ac o ns c a o be di sci
p l es a n d ex p o n en t s o f D a rwin , b u t t he o pp o n en ts
of t ra n smi i f m th s s on , ro eir e x t rem e a p ea
p lt l
o s e ec t io n , h a v e c o me to be k no wn as

n eo - D a r win ia n s , and t he a d v o c a t es of t ra n s m is s io n h a v e a c c ep t e d t he n a m e o f

n eo - La m a rc h i b l i f i t h f m ti
ki an s , f ro m t e r f th
e e n ie m t
or a ve i fl n u en c e o e en v ro n en ,

wh i h w t h d i t i g i h i g f t
c as e f L m
s n k i w f l ti
u s n ea u re o a a rc
’
s v e o ev o u on.

I ge n l it m y b id th t wh il th
n era a m y
e sa t bl a , p tei t h
e re a re an no a e exc e ons, e

zo ol gi t t
o d t id wit h t h
s s en D wi i
o s e
g i t t h id e n eo -f t ar m i i
n ans a a ns e ea o ra n s ss o n ,

wh il t h b t i t d l i g wit h fi d
e e o an s s , d f ea it y m h m
n
pxe l tani f o m n ec ess uc o re as c or s,

ten d t g with th o o L m li e n eo - a a rc t a n s .

I thn p i i f
e ot h w i t n on L l yd M
o
g i be
y f trh er, m to t i f ort yan s ar e os sa s ac o r

i t ig t
n v es d w it
a or th i
an d q ti r er p on i ll y is h iv ex e H b i t u es d on, es ec a n s a an

I t i t wh i h h
ns nc , ld b c d by s y
ou f l t d e t reaf th i b j t Th
ev er c a re u s u en o s su ec . e

i q i y i
n u r d t d t fi d t t h t th
s con uc e t t
o
p n ou t d ie p ru t ,
h no o ro v e or o s ro v e e

i h it
n er f i
an c e o
q d h t eac d h iu re l i c a ra ct t drs , anp g 3 7 3 2
s c o n c u s o n s as s a e on a es 0 -
2

o f th t l m m y w ll
a vo u g g e t h tt
a t i f
e t h t hen g h at f e l d
e a w
en h t
on o e ou u rea er , a ev er

hi p s l i w th p i t i di p t
ers o n a v e s on e o n s n s u e.
 6 I R A N S MI SS I O N
' ‘

35

In ev er y cas e b i rd o r m a m m a l p la nt o r anim al t he n ew i nd i
, , ,

v id u al begi n s wi th t he f er t i l i z ed g er m n o t a t s o m e la t e r period
,

cal led b irt h and fo r o u r p u rpo ses t he d is t inct io n s h o u ld n o t b e

wh eth er t he v a riat io n wa s im p lant ed b ef o re o r af ter b irth b u t ,

wh eth er i t s ca u s e was i nt ern a l o r ext ernal to t he germ .

A tt ent io n s h o u ld b e fix ed u po n t he ger m p las m t he phy s ical ,

bas is Of lif e and t he o nly kn o wn av en u e o f t ra n s m is s io n fro m

o n e gen era t io n to t he n ext and t he d is t inct io n s h o u ld b e c lea r ly
,

m ad e bet ween v a riat io n s d u e to changes i n it s s t ru ct u re fro m

int ernal o r o th er ca u s es a n d th o s e chan ges o f t he o rgani s m
,

d u e t o infl u enc es ex ert ed d irect ly u po n t he o rgan is m d u ri ng it s

d ev elo p ment A s th is d is c u ss io n proc eed s it s h o u ld b e c lea r ly
.

b o rne i n m ind that n o charact er c a n b e tra ns m i tted n o m att er ,

ho w s tro ng ly pres ent u n less t/ze g er m i n a l m a tter is i n s om e waj

,
'

i l eeted N o th ing els e p a s s es o v er fro m p a re nt t o

p r ev o u s y fia .

o fi s p ri n g and n o o th e r m ed iu m o f t ran s m is s io n is p o ss ib le
,
T he .

s t u dy is ,
th erefo re c learly d efined D o m od ificatio n s as s ue/t
,
.
, ,

a ffect t he ger m d irect ly and s o bec o m e t ran s m itt ed o r if n o t

, , ,

d o t he s a m e i nfl u enc es that a ffect t he d ev elo p ing ind iv i d u al

a ls o a ffect t he g er m i na l m a t ter i n t he s a m e d irect io n gi v i ng all ,

d es c endan t s an in it ial trend o r m od ificat io n similar to the o ne

im pres sed u po n t he parent ?

SEC T I O N I I EV I D E N C E F R O M T H E N AT U R E OF
V A R I AT I O N

I n the pin io n o f t he writ er it is f u nda m ent ally wro ng bo th

o ,

lo gic a lly and b io lo gic a lly t o c o nc eiv e o f t he ind iv id u a l as m ad e

,

u p o f t wo s et s o f f a c u lt ies o n e inh er it ed
,
-
and t he o ther
acq u ired T he d is t inct io n n o t o n ly d o es n o t res t u po n goo d
.

gro u nd b u t in it s app licat io n t o t he fact s o f lif e it lea d s to m os t

,

u nfo rt u n at e c o nc ept io n s and t o m o s t erro n eo u s c o nc lu s io n s It .

is th is f u nda m ent a l m is c o nc ept io n o f t he f u nct io n o f t he en v iro n

m ent that is r es po n s ib le fo r m o s t O f t he f o ggy th ink ing whi ch
mark s t he c o nt en t io n o v er t he q u es t io n o f inh erit a nc e o f acq u ired
charact er s and wh ich no wh ere els e bea rs s u ch u nfo rt u nat e fru it
,

as in t he fi eld o f p ract ic a l a ffa ir s I n gen e ra l ev o lu t io n i t d oes

.

n o t m att er great ly wh eth er acq u ired charact ers a re inh erit ed

 l R A N S M l SS I O N
' ‘

35 8

l s a m e ra c e tlt e me c ha r a c ter s lzer ei n d o ra c ia l

of t ze p oss es s sa

v a l u es ex is t a n d lzer eo
y a r e r a c ia l d is t i nc ti on s es ta His/zed .

V aria ti on pra c t icall y confi n ed t o racial char acter s . In a ll t he

exa m p les o f v a ria t io n th a t h a v e b een c it ed ,

— and th ey ha v e
p u rpo s ely been m a ny and in all t he cas es ,
— that oc c u r in ou r
fi eld s and yard s v a r ia t i on is c onfi n ed to ra c ia l c/za r a c ters This
,
.

is t he e xp e rienc e o f breed ers ev ery wh ere When d ealing with .

catt le breed ing a ll v a ria t io n is o f ca ttle c lza r a c ters W e d o no t .

exp ect and we d o n o t find a m o n g catt le t he appea ranc e o f char

ac t ers b elo ng ing t o h o rs es s h eep p igs d o gs o r ch icken s , , , , ,

exc ept as t hey a re p o s s es s ed i n c o m m o n T he s a m e is tr u e o f .

o th er s p ec ies and wh en cha ra ct er s are po s s es sed in c o mmo n

,

t he v ar iat io n i n each ca s e is well with in t he range o f t he s pec ies

in q u es t io n .

T hat is t o S ay v a riat io n s i n c a tt le a ll appea r a m o ng well

,

kn o wn and lo ng es tab lis h ed chara ct ers that d is t inct ly belo ng t o

-

c a tt le s u ch as the h ead h o rn s legs c o lo r u dd er q u ant ity o r

, , , , , ,

q u ality o f m ilk etc A m o ng ch ick en s t he v a ria t io ns a re a mo ng

, .

ch ick en ch a ract ers s u c h as t he s hape and c o lo r Of feath ers ; s ize

, ,

c o lo r and q u ality Of t he egg ; q u ality o f m eat etc

, , .

W e find n eith er ch ic ken charact ers appearing a mo ng cattle

n o r c a tt le ch a ra ct e rs app ea ri ng a m o ng ch ick en s T he hen can .

n o t gi v e m ilk n o r c a n t he c o w b ear f eath er s

,
T h ere is n o i nt er .

change o f charact ers bet ween s pec ies eith e r by b irth o r by ,

acq u is it io n aft e r ward N o t o n ly th a t b u t ev en wh en t he s a me

.
,

ch a ract er is po s s es s ed by t wo d is t inc t s pec ies it s v a r ia tions i n

ea c /z a r e wel l w i t/t i n tlt e ra n
g e o f tlze p a r tic u la r s p ec ies . Fo r
e xa m p le t he legs o f c a tt le and ch ick en s a re b u ilt u p o n t he s ame
,

ge n e ra l p la n b u t th ey ha v e d rifted far apart and d o n o t o v erlap

, ,

ev e n i n th ei r v ar ia t io n s T he leg o f a h o rs e and t he leg o f a c o w

.

a re o n n ear ly t he s a m e p lan and et n o o n e wo u ld m is tak e t he

y ,

o n e fo r t h e o th e r n o m att er what t he ra n g e o f v ar ia t io n
, Ev en .

c o lo r d ev iat io n s a re alway s with in c erta in d efin it e lim it s .

No su ch t hi n g a s an a cq uir ed chara c t er V a riat io n is th ere .

,

fo re a con d i tion n o t a t/zi ug I t is t he s ta te of a ra c ia l clza ra c ter

, ,
.
,

n o t t he res u lt o f t he intro d u ct io n o f a n ew o n e ind eed v ariat io n ,

by the intro d u ct io n Of a po s it iv ely n ew charact er is if n o t ,

u nkn o wn a mo ng u s a m att er th a t belo ng s t o gen era l ev o lu t io n

,
 TR A N SM I SS I O N O F M O D I FI CA TI ON S
3 59

and to the o rigin o f ra c es I t is n o t a m att er that pra ct ically

.

c o nc erns t he th rem ma to lo gis t who is wo rk ing with es tablis hed

,

ra c es and charact ers wh o se v ar iat io n s are fa ir ly well c irc u m

s cri bed . T he app ea ranc e o f a po s it iv ely n ew chara ct er a m o ng
any o f thes e rac es wo u ld b e cau s e fo r pro fo u nd as to n is h ment .

U nd er the pres ent s tat e Of kn o wledge and fo r o u r p u rpo ses , ,

we may say that th e re a re n o s u ch th ings as acq u ired charac

t ers in any pro per s en s e o f t he t erm I t is a figu re of s peech
,
.

at bes t and a mos t u nf o rt u na t e o n e at that

, , .

By acq u ired charact er a s t he t er m is c o mmo n ly em p loyed

”

, ,

is a lw ay s m eant o n e o f t wo th i ng s — ( )
I d i ff er enc es, in t h e

d egree o f d ev elo p ment o f o rd inary ra c ial chara ct ers o r ( 2 ) the ,

pec u liar u se to wh ich the ind iv id u al has pu t his nat u ra l end o w

ment s u nd er his s pec ia l c o nd it io n s o f life .

T h es e a re d ifferenc es in d eg ree n o t in k in d ,
To s peak o f .

th em as charact ers is to dign ity th em with a t er m wh ose mean

in g is em in ent ly q ua li ta tiv e n o t q u an t itat iv e and th is it is that
, ,

has b u ilt u p t he fa ls e c o nc ept io n that ind iv id u a ls o f t he s a m e

breed o r rac e d iffer fro m each o th er in so m eth ing that is real ;
—
that ind iv id u al d ifferenc es are all qu a litat iv e wh ereas with in , ,

t he rac e th ey are q u ant itat iv e merely

, .

T o s peak o f th es e d ifferenc es wh ich a re o n ly d ifferenc es in

,

d egree o f d ev elo p ment o f o rd inary rac ial cha ract ers o r at m os t ,

o nly d ifferenc es in beha v io r o f o rgan s and part s kn o wn t o b e

able to res po nd to v ariou s s t im u li an d to f u nct io n so m ewhat

d ifferent ly u nd er d ifferent c o nd it io n s —t o s peak o f d ifferenc es,

su ch a s th es e p er s o na l acq u is it io n s as acq u i red charact ers is ,

t o u s e t he t erm “
charact er in an u nfo rt u nat e and s ingu larly
”
-

m is lead ing s en se .

N o w a d ifferenc e wh ich is n o th ing mo re than a d egree o f

d ev elo p m ent of a well kn o wn charact er is n o t in it self a n ew
-

c lza ra c ter I t is n o t in th a t s en se an acq u is it io n

. I t is m o re .

in t he nat u re o f a rea lizat io n o f what was befo re a po t ent ia l

po s sibility .

N eith er is a lza oi t ent it led to t he t er m n ew or acq u ired , ,

c haract er Hab it refers o n ly t o t he c u s t o m ary u s e o f nat u ra l

.

fac u lt ies S o me cha ract ers lik e th o se a s s oc iat ed with the p ro

.
,

d u ct io n o f t he ga s tric j u ic e fo r exa m ple ha v e b u t a narrow

, ,
3 6 0 T R A NSM I SSI ON

f u nct io nal range and a re q u it e o u t o f c o ntro l o f t he ind iv id u al ;

o th er s
,
lik e t he bra in and t he h a nd a re c a pa b le Of f u nct io n ing,

in m any d irect io n s — s o m any that n o lif et im e is lo ng en ou gh

, ,

o r i t s n eed s and e x p erienc es v a r ied en o u gh to exha u s t th ei r ,

p os s ib ilit ies .

I f t he s t u d ent h o pes t o f o llo w t he m a zes o f ra c ial chara ct ers as

th ey t ra v er s e t he gen erat io n s lik e thread s thro u gh t he pa tt ern s
,

o f a f a br ic he m u s t n o t c o nf u s e eith er h is t er m in o lo gy o r his
,

i d eas by attach ing s o i m p o rtant a c o nc ept io n a s charact er

to what is n o th ing m o re tha n an ind iv id u a l m a n ifes tat io n O f
chara ct er d ev elop m ent o r the pa rt ic u lar pers o nal u se t o wh ich
,

m any ra c ia l charact er s m ay b e p u t .

T he fa c u lt ies o f t he ind iv id u a l a re lim it ed i n k i n d t o t he

fac u lt ies o f t he rac e and in d eg ree to t he int en s ity o f th ei r
,

inh eritanc e and t h e c o n d it io n s o f lif e T h ere is n o cas e in .

wh ich an ind iv id u al o f o n e ra c e ha s p ick ed u p o r o th erwis e

acq u ired a charact er that b elo ng s to a n o th er rac e and no t t o
his o wn . A ll his ach iev em ent s a ll his c a pac it ies a re wit hi n
, ,

t he lim it s o f h is rac ia l charact er s and t he c o nd it io n s c o ntro lling

th eir d ev elo p m ent .

Th e in di v i d u al i s in a c t u al posses s ion of al l t h e c h ar act ers of

t h e race T hat t h is is tr u e is s h o wn by breed ing e xp erienc es
.

ev ery wh er e fo r in all c a s es t he ind iv id u al tra n s m it s t o so m e

,

d egree all t he ch a ract ers of his rac e M ilk s ecret io n is a char .

act er lim it ed to m a m m als a n d f u nct io n al o n ly in t he fem a le sex ,

et ev ery da i ry m an kn o ws that t h e b u ll will tran s m it m ilk i ng

y
q u alit ies as s u cc es s fu lly as will the c o w .

Ou r exp erienc e with rev ers io n s — th o se lo ng lo s t chara c

“ -

—
t ers t h at ret u rn to p lag u e u s is c on v inc ing proo f of the fact
that ev ery chara ct er o f t he rac e is p oten tia lly present in ev ery
ind iv id u al wh et h er t he d egree o f d ev elo p m ent b e m u ch o r li tt le
, .

I n n o o th er m ann er c o u ld th ey b e s o lo ng and s o p ers is t ent ly

pres erv ed i n t he rac e T h eir repeat ed appea ra nc e lo ng aft er
.
,

th ey hav e c ea s ed to b e typ ical o n ly S h o ws that th ey were nev er

,

tr u ly lo s t .

T he ind iv id u a l is th erefo re b o rn wi t h a ll th e po s s ib ilit ies o f

, ,

t he rac e t o wh ich he b elo ngs T h o s e wh ich s ha ll d ev elo p and

.

fix t he type will d ep end u p o n t wo c o n s id era t io n s — fi rs t t he , ,

 6 2 T R A N S M I SSI ON
3

T hi s , an d ntrod u ct io n o f n ew charact ers is the bas is

n ot t he i ,

o f v a riat io n b et ween ind iv id u a ls o f t he s a m e ra c e D ifferenc es .

bet ween rac es m ay b e eith er q u alitat iv e o r q u ant itat iv e o r ,

bo th ; b u t d ifferenc es between ind iv id u als of the s a m e rac e are

es sent ia lly q u ant i tat iv e .

Q u antitat iv ely that wh ich is t ran s m itt ed fro m parent to o ff

,

s pri ng is a c ert a i n capac ity fo r d ev elo p ment B u t p o s s es s io n

.

o f t he capac ity fo r d ev elo p ment is n o gu aranty that d ev elo p m ent

will fo llo w Wh eth er o r n o t it will f o llo w d ep end s u po n t he

.

nat u re of the c o nd it io n s of lif e and wh eth er th ey will affo rd t he

,

o ppo rt u n ity fo r d ev elo p m ent .

T he lim it s of d ev elo p m ent o f any ra c ial charact er are fi x ed ,

th erefo re by two facto rs fi rs t t he in it ia l im p u ls e bo rn int o t he

, ,

ind iv id u al t h e int en s ity o f wh ich is a m att er o f breed ing ; and

,

s ec o nd the att it u d e o f t he en v ir o n m ent wh eth er fa v o rab le o r

, ,

u nfa v o rab le .

M an if es t ly with any ind iv id u a l t he h igh es t d ev elo p m ent will

b e in th os e charact er s wh ose i nh e ri t ed int en s ity is s t ro nges t and
fo r wh o se d ev elo p ment t he en v iro n ment is m o s t fa v o rab le N ext .

in o rd er will c o m e th o s e wi th h igh int en s ity b u t wh ich a re fo rc ed

t o s tru ggle aga in s t an u nfa v o rab le en v i ro n m ent a s well as th o s e ,

wh o se inh erit ed int en s ity is les s Weak es t o f a ll will b e t he d e

.

v elo p men t o f th o s e charact er s wh o s e inh eri t ed int en s i ty is lo w an d

fo r wh ich the en v iro n m ent is es pec ially u nfa v o rab le A s we ha v e .

s een n o m att er what may b e t he en v i ro n ment n o d ev elo p m ent

, ,

will tak e p lac e exc ept a lo ng lin es that a re c learly rec o gni zed
a s with i n rac ia l po s s ib ilit ies and th eref o re d u e t o tran s m i tt ed

im p u lses .T h ese c o nt ingenc ies c o v er a ll ca s es o f v ari at io n

bet ween ind iv id u als and the real q u es t io n befo re t he s t u d ent
,

is n o t wh eth er acq u ired charact ers a re inh erit ed b u t it is th is ,

Will t he extreme d ev elo p ment o f a rac ial chara ct er u nd er u n u s u

ally fav o ra b le c o nd it io n s o f lif e au g ment ev en t o t he slight es t
d egree t he tra n s mi tted ten d ency f or d ev elop men t in til e n ex t g en
c ra ti on o r a re int en s it ies t he pr o d u ct o n ly o f changes int er na l

to t he germ p la s m

S tated m o re bro ad ly t he q u es t io n is th is : D o es the d ev elop

men t attain ed by t he ind iv id u a l infl u enc e h is p o wers o f tran s m is
s io n ? T h is q u es t io n s h o u ld s tand o u t c lear c u t in t he s t u d ent s
'
-
 TR A N SM I SS I ON OF MOD I FI C AT I ON S 6
3 3

mind . I t is not wh eth er

an ind iv id u al with s trong t end enc ies is
a bett er breed er than o n e with weaker t end enc ies that is
,

c o nc ed ed ; it is n o t wh ether a rac e liv ing u nd er a fav o rab le

en v iro n m ent fl ou ris h es bett er than o n e liv ing u nd er hard c o n

d itio n s — ,
that is c o nc ed ed too fo r nat u ra l select io n is inev i
, ,

tab le ; b u t t he q u es t io n is wh eth er t he i nd i v id u a l will b e the

,

better o r the wo rse as a breed er beca u se o f the sp ec ia l d ev elop

ment he has acq u ired .

T he an s wer to th is will d ec id e t he q u es t io n wh et her we s hall

keep o u r m eat breed ing an im als in h igh o r in mod erat e fl es h ;
-

wh eth er we m u s t d ev elo p t he s peed o f ou r rac ing s ta llio n s an d

mares ; wh eth er a giv en s ire o r d a m is a bett er breed er aft er
s peed is d ev elo p ed than was t he s a m e ind iv id u a l wh en green .

I t will d et erm in e t he wh o le m att er o f t he im po rtanc e o f d ev elop

i ng breed ing s t o ck n o t o n ly as a mean s o f inc reas ing nat u ra l
,

capac ity b u t as a mean s of int en s ify ing the po wers o f tran s

,

mis s io n .

F o rt u nately we a re n ot with o u t fact s beari ng u po n th is

v ex ed q u es t io n ; b u t t he wh o le fi eld is exc eed ing ly d iffi c u lt and ,

reliab le ev id enc e is eagerly so u ght I t is the mo re d iffi c u lt to

.

s ec u re b eca u s e o f t he ev er pres ent and a lway s p o we rf u l i nfl u enc e

-

o f s elect io n .

T he part ic u lar m od ificat io n s ( acq u ired charact ers ) that ha v e

been m o s t d is c u s s ed and who se t ran s m iss ibility has been ad v o
c a ted o n the o n e hand o r d en ied u p o n t he o th er a re o f f o u r

d is t inct k ind s
1 .M u t ilat io n s d u e to inj u ry o r d es tru ct io n o f ra c ial cha ract er s
aft er th ey hav e reach ed fu ll d ev elo p ment .

2 .Habit s o f lif e ar is ing o u t o f t he ex igen c ies o f ex is t enc e .

3 .S tr u ct u ra l p ec u liar it ies d u e t o u s e and d is u se .

4 .A daptat io n s t o c li m at ic c o nd i t io n s .

T h es e a re c o m mo n ly all c o n s id ered a s acq u is it io n s in t he

s en s e o f add it io n s t o ra ci a l chara ct eris t ic s I n t he v iew a d v o
.

c a t ed by t he wr it er th ey a re a ll red u c ib le eith er t o d i fferent

d egrees o f d ev elopmen t o f rac ia l cha ract ers o r to the u ses to ,

wh ich th es e a re p u t u nd er t he c o nd it io n s and ex igenc ies o f

life T h ey will b e c o n s id e red in t he o rd er na m ed a lway s with
.
,

t he q u es t io n u pp er m o s t in m ind A re th ey t ran s m is s ib le
,
 6
3 4 T R A NSM I SS I O N

S ECT I ON III — EV I D E N C E F R OM M UTI LAT I O N S

M u t ilat io n is the f o rc ib le remo v a l o f a part aft er it has d e

v elo ped o r at lea s t t he d es tr u ct io n o f th o s e part s wh ich are f u lly
,

en d o wed with t he p o wer o f c o m p let e d ev elo p m ent U n fo r t u .

na t ely in th is fi eld t he m o s t ab s u rd s t o ries ha v e ga i n ed cred enc e

, ,

and th ei r po p u lar acc eptanc e has d o ne m u ch t o o b s c u re t he

wh o le s u bj ect S o m e o n e o wn ed a cat wh o se ta il was p inch ed
.

o ff in a d oo r and s tra ight way all her k itt en s were ta illes s

, A .

few semi trad it io nal s t o ries lik e th is a re m ad e t he f o u ndat io n fo r

-

believ ing in the t ran s m iss io n o f m u t ilat io n s t he fact s being fo r ,

go tt en that fo r gen erat io n s it has been the c u s t o m to rem o v e

t he ta ils fr o m la m b s with n o S ign yet o f tailles s s h eep as a ,

res u lt and that c irc u mc is io n has been pract ic ed by m any t ribes

,

from the rem o t es t t imes and by t he J ews c ertainly fo r fo u r ,

th o u s and years apparent ly with o u t effect u p o n t he nat u ral d e

,

v elo p m en t of part s Certa in ly if any eff ect has been prod u c ed

.
, ,

it is n o t ev id ent and is s o s m a ll a s to b e c lass ed a m o ng n egli

,

g ib le q u ant it ies fa lli ng ent i rely o u t s id e t he fi eld o f p ractica l

,

res u lt s .

T he ta il being a po rt io n o f t he v ert ebra l c o lu m n m ight b e

, ,

exp ect ed t o lo ng res is t a ll infl u enc es t o ward it s s u ppres s io n ;

b u t t he prep u c e is an u n im po rtant and rec ent s t r uct u ral add i

t io n yet s t ill it lingers d es p it e pers is t ent and s y s t em at ic
, ,

remo v al by fo rc e .

T he q u es t io n lies d eeper than t he s u rfac e A m u t ilat io n lik e .

,

any o th er d ifferenc e in o rd er to b e tran s m itt ed m u s t fi rs t , ,

effect t he ger m p la s m wh ich is the o n ly m at eria l carried o v er , .

I f D ar win s th eo ry o f g em m u les were tru e th en it m ight b e c o n

’

c eiv a b le that a d efect iv e part wo u ld n o lo nger pro d u c e it s s hare

o f t he germ p la s m and that it wo u ld c erta in ly d is appear fro m

,

t he rac e and that a t onc e

,
The fact that p ers is t ent ly m u t ilat ed .

part s d o n o t d isappear is goo d pro o f n o t o n ly that m u t ilat io n s

a re n o t tran s m itt ed b u t that the th eo ry o f gem m u les is i nc o rrect
, .

Fo r t he m o s t pa rt b elief in t he inh eri tanc e o f m u t ilat io n has

,

res t ed n ot u po n exp erim enta l ev id enc e b u t u po n in s tanc es in

, ,

wh ich nat u ra l d efo r m it ies in t he o ffs pring c o rres p o nd to m u t i

la t io n s in t he parent s S u ch c o rres po nd enc e is a s s u m ed to b e
.
3 6 6 TR A NS M I SSI ON

T he su cc ess io n s of red and black representing ga in a n d lo ss

a re s o irreg u lar and s o c on f u s ing that t he p lay er fa ils t o d et ect

th e rat io o f m o re than 5 p er c en t that is aga in s t him and c o n ,

s eq u ent ly d o es n o t rea lize that t he lo nger he p lay s t he m o r e

c ertain are his lo sses N o r d oes he rea liz e th a t in the lo ng ru n

.

n o th ing is m o re ab s o lu tely c ert a in than t he la w of ch a nc e T he .

d ev iat io n is n o t great at any po int aft er t he fi rs t few thro ws ,

a n d herein lies the rs t d ec ept iv e q u a lity o f a ll ga mes o f chanc e

fi .

I f t he lett ers Of t he wo rd inc o m preh en s ib ility

“
b e t o s sed
”

int o t he air i n s u ch a m ann er that th ey m u s t fa ll i nt o a lin e t he ,

chanc es o f th ei r fa lling in t he pro per o rd er t o s pell t he wo rd

c o rrect ly are exc eed ing ly rem o t e y et i t is b ou n d to [cappen if tlze
,

tri a ls a r e long en o ug /t con ti n u ed .

T h es e s im p le fact s t each u s n o t to attach too mu ch im po r

t a nc e to occas io n al occ u rrenc es h owev er s trange o r apparent ly
,

im pr o bab le . T h ey t each u s t oo t h at th ere m ay b e n o s pec ia l

, ,

ca u s e at t he bo tt o m o f t he o cc u rrenc e b eyo nd t he m ath em at ical

la w o f pro bab ility The t o s s ing of c o in s s h o ws why it is fo r

.
,

exa m p le t hat ev ery th eo ry fo r t he c o ntro l o f sex that ev er has

,

been o r ev er can he in v ent ed has been repeat ed ly v eri fi ed .

T h ere is b u t o n e alt ern at iv e and ev ery a s su m pt io n of ca u s e

, ,

h o wev er abs u rd is c ertain to c o me tru e

,
Ita lf tlce ti me wh ich ,

i s s u ffic ient proo f fo r m os t p eo p le who d ep end u p o n m em o ry

i m pres s io n s rath er than u p o n ab s olu t e rec o rd s .

Pro o f by t he m eth o d o f in s tanc e is th eref o re extrem ely

h azard o u s S o m eth ing m ore than the m ere fact of c o inc id enc e
.

is n ec es s a ry i n o rd er to es tab lis h a ca u sat iv e relat io n with any

v ery h igh d egree o f c erta inty .

Wh en th erefo re a d efo rm ity in a ch ild c o rres po nd s to a m u ti

, ,

lat io n in a parent we a re n o t warrant ed in at o nc e as s u m ing a

ca u s at iv e relat io n W e are to remember that d efo rm it ies o f all
.

k ind s are c o m parat iv ely c o m m o n ; that mu tilat io n s a re exc eed

in gly s o ; that freq u ent ly a m u t ilat io n will r es em b le a nat u ral
d ef o rm ity o r an inj u ry ; and that occa s iona lly u nd er t he laws of ,

pro bab ility t he m u t ilat io n o f the parent will resemb le t he

,

d efo r mity in t he o ff s pring th u s s ugges t ing d irect tran s m is sio n

, .

We a re to remember t oo that the law o f chanc e mu s t fi rs t b e

, ,

s at is fi ed b ef o re we can a s s u m e ca u s at io n .
 T RA NSM I SSI ON O F M OD I FI C AT I ON S 6
3 7

The m os t d irect way o f proc ed u re is h o wev er no t to end eav o r

, ,

t o elim inat e t he la w o f chanc e b u t by d irect experiment

, , ,

to learn wh eth er a s u ffic ient n u m b er o f o cc u rrenc es can b e

es tab lis h ed t o c learly exc eed i n n u m ber any p o s s ib le c o inc id enc e .

Ex perim en tal ev i d en ce on in h eri tan ce of mu t ilat ion s . T he

fact s j u s t giv en s h o w c o nc lu s iv ely the hazard o f fra m ing th eo ries
o n chanc e o cc u rrenc es and d emo n s t rat e t he p ract ica l wo rth les s
,

nes s o f all b u t experimental ev id enc e in t he s t u dy o f inh eri tanc e .

U nfo rt u nat ely b u t litt le ev id enc e o f th is k ind is at hand and , ,

s o far a s i s kn o wn t o t he wr it er that wh ich is at hand is c o nfin ed

,

t o art i fic ial inj u ries t o t he n erv e with exc ept io n o f that a lready
,

c it ed in s u ch pra ct ic es as d ock ing and c irc u mc is io n .

R o m an es o u t lin es sev en c las s es o f abn o rma lit ies that appeared

in t he Off s pri ng o f g u in ea p igs c o rres po nd ing t o th o s e art ific ia lly
prod u c ed in t he parent s by B ro wn S eq u ard and his ass is tant s-
.

T h ey a re in brief as fo llo ws 1

I. A ppeara nc e o f ep ilep s y wh en parent s ha v e been rend ered

,

ep ilept ic by an inj u ry t o t he s p ina l c o rd .

2. S a m e wh en t he inj u ry had b een t o t he s c iat ic n erv e

,
.

3. C hang e i n t h e s hap e o f c a r i n an im a ls b o rn o f par e nt s i n

wh ich s u ch a change was t he effect o f a d iv is io n o f the c erv ica l

s y m path et ic n e rv e .

4 . P art ia l c los u r e o f th e e y e lid s in y o u ng b o r n o f pare nt s i n

wh ich that s tat e o f t he eyelid s had been ind u c ed by s ect io n o f

t he c er v ica l sy m path et ic n erv e o r t he rem o v a l o f t he s u p er io r
c erv ical gang lio n .

5. E x o phtha lm ia i n y o u ng b o r n o f parent s in wh ich a s im ilar

pro tru s io n of t he eyeba ll had b een prod u c ed by inj u ry to the

res t ifo rm bod y .

6 Gangren e o f t he ear s in a n im a ls wh o se parent s ears had

’
.

been a ffect ed by inj u ry t o the res t ifo rm body .

7 . A b s e nc e o f t o es in y o u ng wh os e par ent s had eat en O ff

th eir t oes wh ich had bec o me an aes th et ic by reaso n of the
,
“

sect io n o f t he s c iat ic n er v e alo n e o r o f that n erv e and t he c ru ra l .

8 V a rio u s m o rb id s tat es o f the s k in and ha ir c o rres p o nd ing

.

to a s im ilar c o nd it io n o f t he parent s wh ich had been brou ght o n

b y an inj u ry t o t he s c iat ic n erv e .

1 R o ma n es , D a rwin an d A ft er D a rwin , I I , 1 03 - 1 2 2 .
 6 8 T R AN S M I RS IO N
3

I t is no ta b le that a ll th ese ex pe ri m ent s a re bas ed u po n i nju ry

to the n er v e and a re o f a d egr ee o f s e v er i t v lik ely t o a ffec t t he

en t i re o rgan is m se rio u s ly I f h we e r it i s t ru e th a t inj u ry o f

.
,
o v ,

an
y k ind i n t he paren t lea d s as B r w n S e , q u a rd a n d R o m an es
o -

e v id e nt ly s u p po se t o c o rr es po nd i n g d ef o r m ities i n t he o ff s pri ng
, ,

t he f ac t is exc eed i ng ly s ign i fican t T h is is a fi e ld h o wev er

.
, ,

q u it e d ifferent fro t a t m h O f o rd i n a r
y i nj u r ies — s u c h as t h e

rem o v al o f a b o rn o r a ta il pro d u c ing n o c o n s t it u t io nal d is tu rb

,

anc e and lead ing to n o o rga n ic c hanges F u rt h er ex periment s .

a re grea t ly n eed ed t o c o nfir m d e ny o r m od ify t he res u lt s o f

, ,

B ro wn S eq u a rd
-
. I n t he m ea nt im e it s eem s a lm o s t i nc red ib le
t hat so m u ch erro neo u s trad it io n s h o u ld ha v e gr o wn u p s u r
ro u nd ing th is m a tt er o f in herit ed m u t ila t io n s es pec ially wh en ,

t he wo rld fo r u nkno wn gene ra t io n s has a lm o s t i n v a ria b ly s ee n

p e rf ect ch ildr en b o rn fro m o n e ar m ed o n e leg ged and o th er wise

-
,
-
,

m u t ilat ed pa rent s I nd eed if O ffs pri ng i nh eri t ed t he o rd in a ry

.
,

m u t ila t io n s o f t h eir paren t s t he wo rld wo u ld ha v e b ec o m e lo ng

,

s i n c e a c o llect io n o f m o n s tro s i t ies wh ich wo u ld p u t t o s ha m e t he

rare s pec im en s n o w c o llect ed in d i m e m u s e u m s .

I nh erita n ce of di seas e T he o ld trad it io n o f inh eritanc e o f d is

.

ea se is lo ng s i nc e d is pro v ed and th o s e d is eas es o nc e th o u ght

,

t o b e i nh e ri t ed a re n o w kn o wn t o a ri s e n o t fro m i nh eri tanc e

b u t fro m inf ect io n aft er bi rth wh ich fo r Ob v io u s rea s o ns is ex

,

t rem ely ea s y bet ween parent and O ff s pr ing .

T he wea k en in g effect o f wa s t ing d is ea s es u po n t he parent s ,

and t he in flu enc e o f th is weak en ing u po n t he c o n s t it u t io n o f t he

o ff s pri ng,
ind u c ing pred is po s it io n t o d is ea se is h o wev er q u i t e , , ,

a no th er m att e r . T hat m a ny o f the effect s o f s u ch a d iseas e o f

t he pa rent s will wo rk in j u ry and weakn es s to t he O ff s pri n g will
b e rea d ily a d m itt ed ; t ha t s u ch o ff s pri n g will b e t he m o re s u s
c ep t ib le to attack fro m d is ea s es o f a ll k ind s will h a rd ly b e

d en ied ; b u t wh ether it will b e p ec u l ia r ly s u scep t i b le to tlte spe

c iu l d is ea s e tlt a t wr o ng /ct lt a v oc w i t/z tlt e pa r en t is a q u es tio n

o n wh ich we n eed m u ch m o r e ev id enc e .

U p t o dat e th is po int ha s n o t i n t he o p in io n o f t he wr ite r

, ,

been es tab lis h ed A lth o ug h i t is tru e th a t c erta in fa m ily li nes

.

a re s p ec ia lly s u s c ept ib le t o t u berc u lo s is it is n o t yet s ho wn ,

wh eth er th is s u s c ept ib ility is t he res u lt o f inro ad s o f th is s pec ial

 T RA N SM I SS I O N
37 0

that is inc o nc eiv ab le u nd er any law o f phy s iology that is known

o r that can b e im ag in ed .

T he n o n d ev elo p ment Of part s is a n o th er and q u it e a d ifferent

-

m att er I f t he n o n d ev elo p ment b e d u e to a d efect iv e ger m i t

.
-
,

o f c o u r s e d oes no t c o m e u nd er t he pres ent inq u i ry I f h o wev er .

, ,

it b e d u e t o an inj u ry at an ear ly s tage res u lt ing in arres t ed

,

d ev elo p ment it may o r m ay n o t b e eq u iv alent to a m u t ilat io n

,
.

I f t he no n d ev elo p m ent b e d u e to t he d es tru ct io n o f c ells as

-
,

in ch em ica l d eh o rn i n g it is to all int ent s and p u rpo s es a m u t ila

,

t io n as c o nd it io n s were pres ent fo r f u ll d ev elo p m ent I f t he

,
.

n o n d ev elo p ment b e d u e t o m al n u tr it io n t he cas e is d iff erent

-
, ,

and b elo ngs a m o n g cas es to b e c o n sid ered lat er .

T he es s ent ia l weakn es s o f t he wh o le th eo ry that m u t ilat io n s

m ay b e tran s m itt ed lies in t he fact that t he charact ers in q u es t io n
a re pres ent f u lly d ev elo ped and f u nct io n a l u nt il remo v ed by v io
, ,

lenc e all o f wh ich is c o nc lu s iv e ev id enc e o f a nat u ral capac i ty

,

for c o m p let e d ev elo p ment I n v iew o f o u r inab ility t o c o nc eiv e

.

ho w t he remo v al o f a part can po s s ib ly a ffect t he c o rres po nd ing

po rt io n Of an u nd ev eloped and ev en u nfert ilized germ ; in the
absenc e o f reliab le experimental data and with t he c ertainty tha t ,

were t he inj u ries d u e to the m u lt it u d e o f acc id ent s o cc u rri ng t o

a ll f o r m s o f life tran s m i tt ed a ll s pec ies wo u ld s o o n b e d is fig u red

by an o v er h elming a s s of inh erit ed m u t ilat io n s

w m — in v iew o f ,

a ll th es e fact s we a re c erta in ly war rant ed i n f eeling a s s u red that

inj u r ies t o t he f u lly d ev elo ped bod y a re n o t tran s m itt ed .

S EC T I O N I V — E V I D EN C E FR OM FOO D SU PPL Y
T h is is a ry d ifferent m att er fro m mu t ilat io n Of a ll t he
ve .

c o nd it io n s o f life th is is pa r ex c ellenc e t he lim it in g element n o t

, , ,

o n ly in b o dy bu ild ing and in f u nct io na l act iv ity b u t in c o n s t it u ,

t ion a l v igo r a s well .

T hat t he a m o u nt o f f oo d av a ilab le is a c o ntro lling fact o r in

t he d ev elo p m ent o f s ize is a m att er t o o well kn o wn to req u ire
d is c u s s io n I n t he pres enc e o f ab u ndant foo d an im a ls and plant s
.
,

o f a ll s p ec ies at t a in th ei r m ax im u m s ize and th ei r m ax i m u m

d ev elo p m ent in a ll res pect s I f t he s u pply b e lim it ed t he effect

.
,

is in v aria b ly s een in u nd er d ev elo p ment ev en th o u gh t he t o tal

-
,
 T R A N SM I SSI O N OF MOD I FI C AT I ON S 37 1

amou nt c o n s u med is many t imes gr eat er than that act u ally u sed
in bo d y b u ild ing .

I n acc lim at ing t o a s h o rt en ed f oo d s u pp ly t he an im a l o r p lant

is f o rc ed n o t so m u ch t o m ak e m o re ec o n o m ical u s e o f what it
,

can o btain as to red u c e the scale o f liv i n g and act u a lly t o aec o m
p lis h le ss i n t he wa y o f g r o wt h and f u nct io n al act i v ity g en er a lly .

A s tar v ing an im a l o r p lant will m ak e t he mo s t o f a ll a v ai lable

food b u t in add it io n the an im al will replac e a large s hare o f the
,

dry matt er of the bod y with wat er and red u c e it s act iv ity to a
m in im u m b efo re it s u cc u m b s and a s ta rv ing p lant will s t ill p u t
,

fo rth n ew lea v es u sing the s u bs tanc e o f the o ld to no u ris h

,

the n ew .

I f t he h o rtage in foo d c o m es bef o re d ev elop ment is c o m p let e

s

it s eff ect is s een in u n d er d ev elo p m ent o r po s s ib ly in arres t ed

-
,

d ev elo p ment — rec o gn ized by the farm er u nd er the t erm

,

s t u nt ed S o m et im es t he c o nd it io n is o n ly t em po ra ry b u t
“ ”
.
,

mo re Oft en it is p erm an ent wh en n o a m o u nt o f f oo d lat er in life

,

will a v a il t o repa i r t he da m age d o n e by s h o rtage d u r ing d ev el

o p men t F arm ers acc o rd ing ly rec o gn ize t he p erio d o f gro wth
.

as a c rit ical period a n d u n ifo rm ly say that if any liv e s t o ck

,

is to b e s h o rt o f feed let it b e t he o ld er o n es Sad expe rienc e .

has ta u ght t he i rr epara b le ev il o f s h o rtage in foo d d u ring

d ev elo p ment .

U nd er n o u ri s h ment s t rik es at t he v ery ro o t o f l ife as well as

-

at d ev elo p ment What is tru e o f ind iv id u a ls seems tru e o f

.

rac es U nd er n o u ris h ment is fo llo wed by a lo wer ing o f t o n e

.
-

and a lessened rat e o f liv ing wh ile f u ll feed and m ax im u m c o n ,

d it io n s o f lif e gen erally ind u c e great pro t o p la s m ic act iv i ty and

rapid c ell d iv i sio n res u lt ing in m ax im u m s ize and m ax im u m
,

fu nct io na l act iv ity in all part s of t he s tr u ct u re .

A ll experienc e go es t o S h o w that weak en ed parent s p lant o r ,

an im a l giv e rise to yo u ng that a re lo w in v igo r and s lo w o f

,

growth S eed c o rn that is belo w the n o rm al in v itality tho u gh

.
,

it may germ inat e will g iv e ri s e t o weak and s lo w gro wing p lant s

,
-
.

T h ere a re all d egrees o f v igo r and i nt en s ity o f t he v ita l

proc es s es fro m zero u p and n o th ing seem s m o re po t ent than
, ,

the f o o d s u pp ly i n in flu en c ing th is m att er that lies at t he ba s is

o f a ll d ev elo p m ent and a ll f u nct io na l act iv it
y .
 T R A N S M I SS I O N
37 2

As t em pera t u re is an per v ad ing infl u enc e wi th s maller

a ll-

o rga ni s m s s o is f oo d a n a ll p erv a d ing in flu enc e wi th a ll o rgan

-
,

is ms large o r s m a ll W i t h o u t d o u bt it ex ert s a c o ntro lling effec t

'

.
,

u po n t he q u a lit y O f ger m in a l m att er pro d u c ed a s it d o e s u po n ,

it s q u ant ity and u po n t he m ax im u m o r m in im u m d ev elo p m ent

,

Of t he bo d y .

C o ns t it u t io na l v igo r wh ich is t he m o s t v alu ab le as s et o f any

,

p la nt o r an ima l is a h eritage wh o s e s ea t is in t he germ fro m

,

wh ich it was d ev elo ped S u ch a ge r m c o u ld b e pro d u c ed o n ly

.

by a v igo rou s h ea lthy well no u ris h ed parent A nyth ing wh ich

, ,
-
.

wea k ens th is parent c o n s t it u t io na lly wh ich lo wers its t o ne , ,

red u c es it s v ita l po wers and lo wers it s ra t e o f li v ing m u s t

, ,

o f n ec es s ity a ffect t he q u a li ty o f any g er m ina l m att er i t m a

y
prod u c e and t he c o n s t it u t io na l v igo r o f its d es c endant s .

We d o n o t pe rm it th is c o nd it io n t o any large ext ent in o u r

d o mes t icat ed s pec ies p lant o r an im a l fo r we rea lize t oo well
, ,

t he c o n s eq u enc es b u t we ha v e o n ly to lo o k a m o ng t he u nd e rfed
,

c las s es and rac es Of h u m an s t o s ee t he ev il effect s Of mal nu t ri

t io n in weak en ed c o n s t it u t io n s lo w v ita lity pred is po s it io n to the
, ,

ra v ages o f d is ea s e and g en e r a l in efl
,
i c ien c y wh erev er any great
f u nct io n al act iv ity phy s ic al o r m en ta l is req u ired T hat th is
, , .

c o nd it io n is tra n s m itt ed d oes n o t ad m it o f a rea s o nab le d o u bt .

On t he o th er hand ra c es that a re well n o u ris h ed fo r m any

,

generat io n s u nd ergo m a x im u m d ev elo p ment Th is has been t he .

expe rienc e with a ll d o m es t icat ed a n im a ls a n d p lant s F o r t he .

mo s t part th ey ha v e b ee n pro v id ed with a ll t he fo o d th ey need ed ,

and th ey hav e res po nd ed with a d ev elo p ment s u ch as nev er

c a me to th em u nd er nat u ra l c o nd i t io n s .

N at u re n ev er pro d u c ed s u ch s pec im en s as o u r m od ern b eef

o r m ilk breed s o r o u r d raft h o r s es Ou r a c hiev em ent a s breed .

ers is d u e th eref o re t o s o m eth ing b es id es s elect io n and we a re

, , ,

f o rc ed to o n e o f three c o nc lu s io n s
1 T hat nat u re n ev er pro d u c ed a p erf ect s pec im en ; that is
.
,

that nat u ral c o nd it io n s were nev er s u ffic ient ly fa v o rable to allo w

t he ind i v id u a l to rea lize t he f u ll d ev elo p m ent t o wh ich his nat u ral
end o wment s ent it led him .

2 T hat with each inc rem ent o f ga in thro u gh s elect io n es tab

.
,

lis hin g a h igh er gen era l a v erag e a n ew c en t er o f v ariat io n

,
“
 T R A N SM I SS I O N
3 74

th is ma tt er o f c o n s t it u t i n a l igo r wh ich s o la rgely d epend s o v ,

u po n n u t ri t io n t ha t t h e breed e r c a n a ff o rd t o m a k e n o mi s tak e

a t th is po i n t .

I n fl u e nc es that s t ri k e at t he ro t o f t he v i ta l p rinc ip le what o ,

e v e r th a t m a b e a r e fa r rea c h in g in t h e i r c o n s eq u e nc es T o
y ,
.

m a int a in t he v it a l po wer s a t a m a x im u m is o n e Of t he p rim e

o bj ect s i n a ll breed i n g and t ha t th is is t o a la rge extent a ,

m a tt er o f n u t rit io n is a fa ct th a t s h u ld b e f u lly apprec iat ed o

by him who h o pes to m a int a in u n im pa ired t he v a lu a b le rac ia l

ch a ract ers fo r whic h he breed s his a nim a ls a n d his p la nt s .

T h ere is n o bett er m a x im fo r t he breed er than t his : t he

res u lt s Of goo d feed a re t ra n s m itt ed t o t he o ffs p ring in t he fo rm
o f a v igo ro u s c o n s t it u t io n and large po wer s o f a s s im ilat io n and

o f s er v ic e .

SEC TI O N V EV I D EN C E F R O M A C C LI M ATIZ ATI O N

I t is kn o wn fa ct t ha t t he ind iv id u a l a cq u ires by experi

a we ll -

enc e a h igh d egree o f res is t anc e to t e m pera t u re po is o n s o r , ,

o th e r a d v er s e c o nd it io n s o f life ; that th is m od i fic a t io n is m o re

o r les s
p e rm a n e nt wi th t he i nd i v id u a l and t hat i n g o d t im e t he o

ra c e a s a w /t o le b ec o m es acc li m a t ized to changed b u t pers is t ent

c o n d it io n s I s t h is ra c e ac c lim a t izat io n in any way t he resu lt

‘
.

o f tra n s m is s i n o f t he acc lim at iza t io n o f t he ind iv id u a l ?

o

1 It s h ld b
ou e c ea r l ly un d e rs to o d th at
“
ac c l im a t iza t io n is not c o n fin e d to
a d v ers e c o n di i t o ns ; it m a y l
re a t e as we ll d
t o a a pt a t io n t o i m pr o v ed c o n d it i o n s,

s uc h as in c rea s e d fo o d su
pp l y . I n d e ed . t h e t erm is in t en dd e t o c o v er a c c om m o

d a ti o n t o an
y (
“

I ma g e i n ex ter n a l r a n d / t i o n s «
f on, w h th e er fa v o ra b l e o r u nf a v o ra b e, l
g ra d l ua o r su d d en .

A s is we ll k n o wn , ac c l im a t iza t io n is m o re s u c c ess f u l if t he j s u b ec t io n be
g ra d u a l; b u t in , an
y e v e n t , o n e o f t wo res u t s l wi ll fo l l w if
o th i d i id
e ln v ua is no t

ki ll de in t he p ro c es s ( I ) it wi ll b ec o m e p er m a n en t ly l t a e re d d will t h
, an erefo re

di sc ha rge it s fu n c t io n s in a mo d ifi d e m a n n er ; o r ( 2 ) i t wi ll ac q u i re by exp eri en c e

so h igh a d eg ree o f res is t a n c e a s to be ab e to l res u m e its usua l a c t iv it ies a f t er

t he fi rs t d is t u rb a n c e and a f t erwa r d to di s c ha r
g e i ts n or m a l f u n c t io n s in s p it e o f

a d v e rs e c o n d it io n s .

T h ere a re t h us t wo ki d n s of ac c l im t i t i
a za h h th f
on : one in w ic e u n c t io n s or

a c t iv it ies a re m o d ifi e d , t he o t h er in wh i h t h i
c d i id l e d i n v ua s u c c ee s n res is t in
g the
c h a n ge d con di i t o ns , an d t h e re fo re i p
n i g it m l f
res e r v n ti s no r a un c o ns . Of t h e
t wo . t h e l a t te r is p e rha ps t h e m o re c o mm Th f m
on b t y th
. e or er e ra s e m o re con

s t it u t io n a l c h a n ge, t h e a t t er t h e g rea l t l t i ity i g i t i

er e as c n o r a n za on .
 T R A N SM I SSI O N OF M OD I F I CA T I ON S 3 75
I

To d et er m in e wh eth er in t he acc lim a t izat io n o f a rac e agen '

c ies a re in v o lv ed o t her than select io n o perat ing u po n ind iv id u als ,

it is n ec es s a ry eith er t o elim inat e t he res u lt s d u e to s elect io n o r

els e t o d is c o v er ca s es in wh ic h i t d o es n o t o cc u r Wh ile t he .

firs t is all b u t im po s s ib le of acc o m plis h m ent with an y feeling o f

as su ra nc e t he s ec o nd is in t he op in io n of t he wri t er ent irely
, , ,

feas ib le es pec ially in c ertain lin es

,
.

The im p o rtanc e o f t he wh o le q u es t io n and the d iffic u lty o f

sec u ri ng reliab le data a re s u ffi c ient exc u s e fo r i ntro d u c ing a

s o m ewhat f u ll d is c u s s io n o f c erta in t o p ic s wh ich a ff o rd ev id enc e

u po n t he q u es t io n at hand .

Ex t en t of acc lima tizat i on T he p o wer o f t he ind iv id u al to

.

adapt it s elf to changed c o nd it io n s is s o m eth ing marv elo u s a s ,

has b een s een u nd er t he s u bj ect o f ca u s es o f v ar iat io n and in t he

d is c u s s io n o f relat iv e s tability T h is s a m e elas t ic ity o f o rgan i
.

za t io n is c haract er is t ic o f rac es a s a wh o le By m ean s o f t his .

adaptab ility m any s pec ies o f bo t h an im als and p lant s hav e

t o ta lly c hanged th eir hab itat and with th is change h av e u nd er
,

go n e t he m o s t s weep ing alt erat ion s The whale is d ev elo ped .

fro m a land m am mal and is s u ffering d egen eracy as a c o n s o

q u en c e . S w i n e ha v e b een adapt ed fro m a d iet o f r o o t s and

fl es h to o n e m ain ly Of grain Ho rs es and c a tt le in th ei r wild

.

s tat e s u b s is t ed ent i rely o n pa s t u re b u t wit h u s th eir d iet is

,

fro m 2 5 t o 7 5 per c ent gra in S h eep are m o u nta in an im als b u t

.
,

t hey ha v e been adapt ed t o t he rich es t pa s t u res and the lo wes t

plain s T he t u rk ey n at iv e to N o rth A m er ica is m ak ing his
.
, ,

way o v er all the earth as ch ick en s ha v e s catt e red bro adcas t

,

fro m t heir nat iv e hab itat in s o u th ea s t ern A s ia .

T he po tat o nat iv e t o the m o u nta in s o f Per u is n o w gr o wn

, ,

ev erywh ere in t em p erat e r egio n s th o u gh it n ev er s u cc eed ed in

,

acc lim at ing in the t ro p ic s exc ept in high a lt it u d es T h ere a re .

ev id ent lim it s o r els e t he p oss ib ilit ies a re no t et e xha u s t ed

, y .

C o rn ( m a ize) will end u re b u t s light change i n lo ca lity with o u t

s u ff e ring s erio u s ly yet aft er a few y ear s i t app ear s t o rec o v er
,

t o ne and s u cc eed I n th is way the c u lt u re o f th is cr o p has b een

.

grad u ally m o v ing n o rth ward in t he U n it ed S t a t es u nt il n ow it ,

is f u lly acc lim at ed in regio n s in wh ich a q u art er o f a c ent u ry ago

it s c u lt u re was im p os s ib le S O s en s it iv e is the c o rn plant to
.
 6 T RA N SM I SSI ON
37

c lim at ic change b u t so read ily d oes it adj u s t it self t hat n ew

, ,

v ariet ies can b e intr o d u c ed s u cc es s f u lly pro v id ed th ey a re gi v en ,

c o n s id erab le t ime to acc lim at e o n a s mall s cale befo re th eir pro

d u ct io n u nd er fi eld c o nd it io n s is att em pt ed .

Wh eat has ext end ed alm os t o v er t he eart h exc ept in extreme ,

lat i t u d es .T he v ariet ies ha v e b ec o m e s o well fix ed in t he v ario u s

regio ns t hat when bro u ght fro m long d is tanc es th ey seld o m thriv e
at firs t S o m e s train s n ev er s u cc eed b u t o th er s acc lim at e per
.
,

fec tly in n ew loca lit ies V ariet ies m ay b e ch a nged read ily fro m
.

s p ri ng to wi nt er s o rt s and t he h ib ernat ing hab it b ec o m e fix ed

, ,

a s in o th er b ienn ia ls .

I m p o rt ed an im als a re s eld o m f ert ile u nt il acc lim at ed I t is .

s ai d ho wev er that c erta in breed s o f t he d o g n ev e r acc lim at e i n

, ,

I nd ia s u ffic ient ly well t o preserv e th eir d is t inct iv e rac ia l c ha rac

t er s On t he o th er hand s pec ies o cc a s io nally pro s per bett er in
.
,

n ew lo ca lit ies t han i n o ld o n es G enera lly s peak ing d is tanc e .

,

m ak es less d ifferenc e th a n a lt it u d e t em perat u re s u n lig ht and , , ,

f o o d s u pp ly T he ev id ent princ ip le in v olv ed is t he infl u enc e

.
,

fa v o rab le o r o t herwise o f c ert a in elem ent s of c lim at e u po n t he

,

d ev elo p m ent o f rac ial ch a ract ers .

A ccli m at iza ti on t o t em per atu res I t i s a well kn o wn fact that .

-

if we br ing t o get h er and p u t u nd er t he s a m e c o nd it io n s p lan t s

o r c u tt ing s o f t he s a m e s pec ies b u t g r o wn i n d iff erent lat it u d es

o r a lt it u d es a n d th eref o re hab i t u a lly exp o s ed t o wid ely d iff er ent

,

t em pera t u res th o s e fro m t he m o re n o rth ern lo calit ies and t he

,

h igh er alt it u d es will b e t he fir s t to pu t o u t b u d and leaf .

D e C and o lle o f S wi t zerland a n d Ba iley o f N ew Y o rk ha v e , ,

b o th c o nd u ct ed ext en s iv e ex perim ent s in th is d irect io n and with ,

t he s a m e r es u lt s
1
.

T he f o r me r t oo k a m o ng o th er s c u tt ing s o f t he p o plar t he
, , ,

t u lip tree and t he cata lp a b o th fro m M o ntpellier and from

, ,

G enev a and plant ed t hem at t he la t t er plac e i n glass es o f wat er

,

wi th s a nd a t t he b o tt o m I n ev ery c as e th o s e tak en fro m

.

G en ev a t he c o ld er loc ality lea fed o u t fi rs t T he d iff erenc e in

, ,
.

the ca s e o f t he p o p la r wa s ab o u t t wenty t hree d a y s i n t he c as e -

,

o f t he t u lip tr ee a b o u t eigh t een day s and in t he cas e Of t he ,

catalpa t wenty d ay s .

B il y S i l f t h U lik
1 a e pp
, 6—
9 3 u rv v a o e n e, . 2 01 .
 8 T R A N S M I SSI O N
37

T he c o rn fro m N ew Y o rk was ev id ent ly t he bett er seed ,

becau s e it s fi nal perc ent age o f ger m inat io n wa s h igh er .

T h is fact m ight acc o u nt fo r s o m e po rt io n o f t he d ifferenc e in

pro m ptness o f germ inat io n b u t we a re inf o rmed that d u ring t he
,

ent ire m o nth Of t he exp eri m ent t he p la nt s fr o m t he n o rt h ern

grown s eed were the larges t and m o s t v igo ro u s of any

“
T h ey .
”

wer e ev id ent ly ah ead in th eir d ev elo p m ent B ai ley rem ark s that .

no t o n ly c o rn gav e t he m os t m ark ed res u lt s in fa v o r o f the

“

n o rth ern sa m ples b u t th ere was genera lly a s imilar d iff erenc e
,

in t he wat er melo n s and b ean s wit h n o t o n e c o nt ra ry res u lt

,
.

B o nn ier m a d e o b s erv at io n s with Teu cri u m S co r od on ia ( wood

“
1

s age) fo r eight year s Wh en gro wn o n t he h igh alt it u d es o f t he

.

Py ren ees it pro d u c ed s h o rt er s t em s dark er g reen and m o re ha iry

,
-

leav es and m o re c o mpact infl o res c enc e than wh en gro wn o n lo wer

,

land S eed s ga t hered fro m th ese p lant s and s o wn in Paris aft er

.
,

tlzr ee y ea r s in t he n ew hab itat pro d u c ed elo ngat ed s tem s wit h ,

les s hai ry and bright er green lea v es o r p lant s v ery s im ilar t o

-
,

t ho s e fro m seed s Obtain ed in the n eighbo rhoo d of Paris .

T he s am e experim ent er c o llect ed s pec im en s o f o th er s pec ies ,

bo th in alp ine and in arct ic regio n s and f o u nd that th ose o f t he ,

latt er reg io n had “

m o re ro u nd ed c ells and larger interc ellu lar
s pac es
2
.

T hat rac es as well a s ind iv id u als acc lim at e t o t em perat u re is

ea s ily s h o wn and well kn o wn b u t a s t he proc es s is g en era lly
,

acc om pan ied by s elect io n th ere is d iffi c u lty in fi nd ing in s tanc es

free fro m its infl u enc e and it is pract ic a lly im po ss ib le to a s ses s
,

and d ed u ct its res u lt s I t is n o t im po s s i ble h owev er to fi nd

.
, ,

cas es fairly s at is fact o ry .

The o r igin and h is t o ry o f t he S h et land p ony is n o t kno wn ,

and yet it is pract ically c ertain that it s s ma ll s ize is part ly t he

res u lt of a c old c lim at e and s canty f eed U nd er th es e ad v erse .

c o nd it io n s th ere mu s t hav e been v igo ro u s n a t u ral s elect io n We .

s ha ll learn i n t he chapt er o n Hered ity that this s m all size

“

c o u ld d o u bt les s b e acc o u nt ed fo r by progressiv e s elect io n b u t ,

t he q u es t io n h ere is wh eth er pro gress iv e acc lim at izat io n is n o t

also in v olv ed .

V 1 V i ti
ern o n , i A im l
ar a on n d Pl t
n
p 3 f
a sm w
anh i h t he a
an s , t . I 2, ro c c c o un

i t k
s a en . I b id p 3 3 2 . . 1 .
 T R A N SM I SS I ON OF MOD I FI C AT I O N S 3 79

th ings c o n s id ered t he c o n v ict ion is fo rc ed u po n u s that

A ll ,

t he S het land s s u ffered a pro gress iv e d im in u t ion b ec a u s e o f lo w

t em perat u res o r sh o rt feed o r bo th o r else t hat th ese no rth ern
, ,

fo rm s liv ing u nd er hard c o nd it io n s lagged behind th eir mo re

, ,

fo rt u nat e n eighbo rs in t he general i ncrease in size that has

att end ed t he ev o lu t io n Of t he h o rse k ind genera lly .

T he t em pera t u re of ho t s p ri ng s v aries all t he way fro m 5 0 t o °

9 8 C 1
°
S o far. as kn o w n th ey a r e a ll inhab i t ed by li v i ng o rg a n
is m s The pro t o p la s m o f o rd inary p lant s and an im als cann o t en
.

d u re a t em pera t u re abov e 4 D eat h q u ick ly follo ws t he att em pt

t o ra ise it m u ch abo v e th is po int and the n ea res t relat iv es o f ,

these ho t s prings s pec ies are n o exc ept io n t o t he genera l ru le

-
.

Y et t he fact rem a in s that t he ho t s prings a re inhabit ed and ,

by crea tu res so s m all t hat t heir t em peratu re m u s t b e t he sam e

as that o f t he wat er s in which th ey liv e Ho w ha v e t hes e wat ers .

bec o me peopled wit h o rganis m s liv ing in t em pera t u res ten to

fifty d egrees higher than c ou ld b e end u red by t he s t ock fro m
wh ich th ey m u s t ha v e d es c end ed ? 2

N o a mo u nt of s elect io n c o u ld acc ou nt for t he fact fo r th ere ,

a re n o o th er kn o wn s pec ies liv ing i n an en v i r o nm ent appro ac h

in g th es e t em pera t u r es T here m u s t hav e been p r og r es s i v e d e

.

v elop m en t o f s o m e fa s h io n and fro m s o m e ca u s e .

U p o n th is po int t he experim ent o f D a llin ger is bo t h s ign ifi

cant a n d v alu ab le He rea red Flagellata in an ov en where
3
.

c o ntrol o f t em perat u re was ab solu t e Beginn in g at C. .

,

he t oo k f ou r m o nth s in whic h t o ra ise the t em perat u re throu g h

5 . a pr eca u t io n n o w kn o w n t o b e u n n ec e s s ary a s F l ag e llata ,

will end u re a q u ick rise to

W hen t he t em perat u re reach ed 2 3 t he o rgan is ms began “ °

dying b u t soo n c eased and aft er t wo m o nth s t he t empera t u re

, ,

was ra is ed half a d egree m o re A ft er a t im e it reach ed .

when th ey a ga in began to d ie and fo r eight m o nth s the tem per ,

at u re c ou ld no t b e raised ev en a q u art er of a d egree abo v e th is

1
C B D .
p t. E p i m
a v en t l Mor p,h xl er
g y P t
en aI pp 5
— S
or o o , ar , . 2 0 2 I .

3
Th t mp e t ei ligh t ly i d iff
era u re v a r es st p t f h t p i g b i g l w
n eren ar s o o s r n s, en o er

n ea rth dg e e e
—
.

3
C B D p t E p i m
a v en t l M
or p h x l er
g y P t
en a I pp 5 or 54 ;
o o V ar 2 2 2 ern o n ,

—
.
. .
, , ,

V i ti
ar a i A im l
on d Pl t pp 3 7 9 3 8 ; j
n n a s an an s , l f til R y l tlI i
. 0 p i l
ou r n a o e o a c r osc o ca

Soci ety , V I I , 1 91 .
 8 0 T R A N SM I SSI ON
3

po int I t seem ed at th is t ime t hat a s tat io nary po int had

.
“

been reac hed b u t u lt im at ely D allin ger was ab le t o m ak e s light

,

add it io n s t o t he t em perat u re and proc eed ing by s lo w s tages , ,

“

and fo r sev eral yea rs he s u cc eed ed at las t in reaching

,

wh en t he exper im ent was t er m inated by an acc id ent .

T he exact length o f t im e em p loyed i n th is exper im ent is n o t

s tat ed b u t it is s u ppo s ed by V ern o n t o b e appro x im at ely s ix
,

ear s T h u s th es e o rgan is m s were b r ed fo r m any gen erat io n s

y .

d u ring t he experiment and it is really a c as e of rac e a cc lim at i

,

za t io n .T he o rga n is m s were m o nad s it is tr u e wh ich m u lt iply , ,

by fi ss io n s o that as D av enpo rt s tat es the h igh t em perat u res

, ,
“
,

acted u po n t he s a m e pro t o plas m at t he en d o f t he exper im ent

a s at t he b egi nn i ng I s th ere any rea so nab le d o u bt t hat t his
”
.

is t he proc es s by wh ich o rgan is m s o f th is charact er hav e ga i ned

acc es s to o u r ho t s p rings ev en u nd er nat u ra l c o nd it ion s ?
I n th is expe ri m ent three p o int s a re n o t ewo rthy
T h ere were c erta in “
s t ick i ng p o int s s o t o s p eak t hat ”
I .
, ,

were d iffi c u lt to get o v er b u t aft er th ese were pa ss ed add i t io nal

, ,

i ncrea s e of heat wa s ea s ily end u red T he t em perat u re o f 2 5 5

°
.

was o ne of th es e s t ick ing po int s .

2 . I t was f o u nd th a t t he pro c es s o f acc lim at izat io n d id n ot

bec o m e grad u ally s lo wer and m o re d iffi c u lt wit h t he h igh er tem
e r a t r e s for w a s t he m o s t d i ffi c u lt t em p erat u re en c o u n
p u ,

t ered req u i ring eight m o nth s t o s u r mo u nt wh ile t he ri s e fro m

, ,

4 7I.
°
t o 8
5 3 w
°
a s m ad e i n s e v e n m o nth s and that fr o m 6 1 I ,
.
°

t o 7 0 in a few m o nth s ( n u m b er n o t s tat ed ) s h o wing that t he

° 1
,

lim it s o f v ariab ility were n o t reac hed and s u gges t ing that t he ,

exp e ri m ent m ight ha v e b een c o nt in u ed m u ch lo nger and t he ln

creas e p u s h ed m u ch fart her .

3 . O rgan is m s a cc lim at ed t o 7 0 d ied o ff w h e n r

°
et u r n ed t o t h e

o ri gina l t em pera t u re o f I s h o wi ng that t he m o d ificat io n o f

t he pro t o p la s m wa s n o t o n ly pro f o u nd b u t a ls o p er m anent In .

o th er wo rd s , h ere is a s pec ies wh ose tem perat u re has been

rais ed thro u gh s o m any d egrees and its pro t o p la s m S O alt ered , ,

that it can n o lo nger end u re its o rigi na l no r ma l t em perat u re .

I t has been tak en ent i rely o u t o f it s fi eld and p lac ed in an o t her

so far rem o v ed a s t o h a v e n o c o n n ect io n with it s f o rm er s tat e .

1 V V i ti
ern o n , i A im l
ar a on d Pl nt np 3 80 a s an an s, . .
 8 TR A N SM I SSI ON
3 2

in gen eral to p oiso n s o f all so rt s g rad u a lly ad m in is t ered is ,

well kn o wn 1
.

S o far as we a re ab le to j u dge im mu nity to i nf ect io u s d is eas es

'

is pro d u c ed in t he s a m e way On e attack o f c ert a in d is ea s es .

s er v es t o r end er t he i nd iv id ua l im m u n e th ro u gh life T he .

q u es t io n that int eres t s u s at t his po int is t his : I s th is imm u nity

in any s en se o r t o any d egree tr a ns m i tted to tlze d esc en da n ts ?
, ,

I t is c la imed by s o m e that if a so w rec o v ers fro m a ca s e o f

ho g c ho lera s u ff ered wh ile carry ing y o u ng t he p ig s will b e b o rn ,

wi th a high d egree o f res is tanc e if n o t ab s o lu t e i m m u n ity ; t he ,

id ea being that th ey acq u i red i n u ter o fro m t he b lo o d s er u m o f

t he m o th er t he s a me k i nd o f im m u n ity t ha t c o u ld b e pro d u c ed
by i n oc u lat io n .

T h ere is to o litt le experi m ental ev id enc e a s yet and we kn o w ,

t oo litt le o f t he rea l nat u re o f a cc lim at izat io n to wa rrant p o s it iv e ,

c o nc lu s io n s Wha t is kn o wn ho wev er is s u ffic ient to ra is e so m e

.
, ,

i nt eres t ing and exc eed ingly s u gges t iv e q u es t io n s .

I f im m u n ity can b e prod u c ed in t he m o th er by the repeat ed

inj ect io n o f t he ch em ic a l prod u ct s o f d is ea s e and if s u ch im ,

m u n i ty b e p erm an ent th en why s h o u ld n o t t he y o u ng wh o s e

, ,

b loo d s eru m is d er iv ed fro m the mo th er b e als o o f t he s a me ,

c haracter ? Wh enc e c o m e o u r nat u ra l im m u n es

“
a re t hey

mu tant s o r a re th ey t he pro d u ct s o f im m u n izing infl u enc es

,

fro m t he pa rent age R ec ent in v es t igat io n s s eem to ind icat e

?

s pec ific q u a lit ies in b lo o d s er u m and it may v ery well b e

2
,

that b loo d relat io n s h ip mean s mo re than we hav e hit hert o

“ ”

s u ppo s ed .

T o what ext ent i m m u n ity is p u rely a ch em ica l q u es t io n and ,

t o what ext ent it is c o nn ect ed wi th t he p o wer o f t he wh it e

c o rp u s c les to attack and d iges t in v ad ing o rgan is m s we d o n o t ,

kn o w I n s o far as it d epend s u po n o r a ffect s t he s er u m o f t he

.

b lood it may well b e a tran s m iss ib le q u ality fro m the f ema le

m a m ma l if n o t fr o m o th er parent s .

1C B D. .
p t E p i m
a v en t l M or p,h l x
g y er P t
en aI pp 8 or; V o o , ar , . 2 2 ern o n ,

V i ti
ar a i A im l
on n d Pl t n p p 3 86 3 8 7
a s an an s , .
- .

3Wh th b l d en m f e oo
p i i
s erui j t od i t
one s t h i
ec es f t h s n ec e n o e v e ns o an o er,

th e m t i j i
os ff t
n ur o us id ft e t f ll w
ec s a re s ad th o ll d p ip it i
en o o o , an e s o -
ca e rec n

tes t m t
s ee t b l i h th f
s t t h t d iff
o es a s e i bl ac d a m f d iff t p
eren c es i n oo s eru o eren s ec es

a re p f d
ro o u n S B l d I.m m i t y eed B l oo d R l t i unh ip yb
an N tt ll oo i w e d e a ons , u a , rev e

in S c i en c e, Oc t o b er 2 8 , 1 904 .
 T R A N SM I SS I O N O F M O D I FI CAT I ONS 8
3 3

Ch emical p in io n o f the
act ion of n orm al secr et i on s . I n t he o

writer th o s e who d is c u ss t he s u bj ect o f t he tra n s m is s io n of

,

mo d ificat io n s (acq u ired charact er s ) a s if it were a s ing le is s u e ,

and d is m is s it in toto as impo s s ible u po n the th eo ret ica l gro u nd

that n o s u ch m o d ificat io n s c o u ld by any m anner o f mean s a ffect
t he ger m p las m — th o s e who d is c u s s and d is m is s t he matt er in
th is m anner c o m m it a f u nda mental err o r in o v er lo o k ing the
fact that as a wh o le th is is a bro ad q u es t io n o r rath er a ser ies
, , ,

o f q u es t io n s and that t he liv ing o rgan is m s exp o s ed as th ey

, ,

a re fo r gen erat io n s t o o u t s id e c o nd it io n s ab s o lu t ely es s ent ia l t o

th eir ex is t enc e present many po int s o f c o ntact in wh ich th ey

,

a re exc eed ing ly s u s c ept ib le to infl u enc e T h ey m ak e t he fata l .

erro r t oo o f fa iling t o d is t ing ui s h b et ween th o s e c irc u m s t a nc es

, ,

that a ffect o n ly t he ext erna ls o f t he b o dy and th os e all per v ad ing -

infl u enc es that a ff ect t he v ery c o n s t it u t io n o f t he o rga n is m .

F o r exa m p le it is n o w well kn o wn that t he perf ect wo rk ing

,

o f t he b o dy a s a wh o le d ep end s u po n t h e pr es enc e o f s p ec i fic

s ec ret io n s o f c erta in o rgan s m any o f wh ich were o nc e th o u ght ,

to b e f u nct io n les s D es tr u ct io n o f t he thyro id g la n d at o nc e

.

arres t s n o t o n ly the phy s ica l b u t t he m enta l d ev elo p m ent I n .

c hild ren it s d egenerat io n res u lt s in retard ed m ent al d ev elo p m ent

and ev en in id io cy a cala m ity that can b e a m elio ra ted a n d ev en
, ,

av ert ed by t he inj ect io n o f thyro id s u b s ta n c e o f an im a ls Ba u

,
.

mann f o u nd that t he s ecret io n s o f th is g land a re chara ct erized by

t he pres enc e o f io d in wh ich is f o u nd n o where els e in t he b o dy
,
1
.

On th is s a m e gen era l q u es t io n V erno n aft er s p eak ing o f t he ,

freq u ent ly fat a l eff ect o f rem o v ing t he thyro id gland u n les s t he ,

anim al b e fed thy ro id s u b s tanc e rem ark s as fo llo ws 2

,

E t i p tio
x r f th a p l
n o
g l d l t
e su i m h m o e p
ra ren a e d y d t h
an s res u s n uc r s e ea ,

an d he g i th
re a i j t i a nf e t t
e m y d
n ec l y th f t l
on o i e E ti
x ra c s a e a e a a ss u . x r

p a t ion o f the pa n c reas c a u ses the p rod u c t io n o f s ev ere d ia b etes , and u lt i

m a tel y d ea t h , but suc h an effec t m ay b e a v o id ed b y the g ra ft in g o f a

po rt io n o f ex c is ed g la n d i n the peri to n ea l c av it yo r the tis s u es A g a in ,
.

ex t i rpa t io n o f t he to ta l k
id n ey s u b s ta n c e of the d o g lead s , n ot to a d i m in
is hed s ec retio n o f u r in e, b u t to a la rgel y in c rea se d sec retio n , a c c o m p a n ied

by a ra p id was t in g a wa y wh ic h s o o n en d s fa ta lly Hen c e the k id neys m a y

, .

po ssess a n in fl u en c e o n the m eta b o lis m o f the w h o le b o d y , a s well as th eir

1 I o eb , P hy i l gy f th
s o o o e B ra in . pp . 2 07 - 2 08 . E xt ra c t s of t hi s an d ot he r

la
g n d s a re n o w g l ly p p
re u ar re a re d at o u r l l
a rg er s a u g ht h er o u s es .
2 V ern o n , V aria t io n in A n im a l s an d Pl an ts , &
pp 3 5 3.60 -
.
 8 T R AN S M I SS I O N
3 4
o b v io u s sec reto r y fu n c tio n . Th e s p lee n a p pea rs to ha ve an in tern al sec re

t io n h h
w ic is of in fl u en c e in s e t t in
g fr ee th e p a n c rea t i c ferm en t Fi n a.ll y ,

ex t ra c ts o f v a rio u s n e r v o us t iss u es d
b ra in , s p i n a l c o r , a n d s c ia t ic n erv e
h av e b een fo u n d , w he n y i j ec ted to p od
i n tra v en o u s l n , r uce a d is ti n c t fa ll of

b lood p ress u re, w ils t1

h th ose o f t he p i t i t y b od y p odu ar r u ce a m ar k ed ri se .

Here is a bas is fo r po s s ib le tra n s mis s io n o f s u ch d is eas es as

m ight b e c o nn ect ed with n o r m a l s ecret io n s a s it c ertai n ly is fo r ,

any ext ernal in fl u enc e th a t c o u ld per manent ly a ff ect eith er th eir

cha ract er o r th eir q u ant ity T h is o pen s a wid e fi eld fo r t he po s
.

s ib le and p er m an ent infl u enc e o f ca u s es o f v a riat io n ly ing o rigi

na lly o u t s id e t he ger m b u t wh o s e effect s a re o f s u ch a nat u re

,

as t o m ak e t hem s elv es felt thr o u gh o u t t he ent i re o rgan is m and ,

t o infl u enc e no t o n ly it s d ev elo p m ent and act iv ity b u t it s p o wer

o f tran s m iss io n a s well .

A k in t o th is is t he p o s s ib le effect o f s u c h ch em ica ls a s alc o

ho l wh ich has s pec ific relat io n s t o pro t o p la s m and is o n e o f
,

t ho s e infl u enc es t h a t a pparent ly a re c a pa b le o f penet ra t ing to

t he u tt er m o s t lim it s o f t he o rga n is m \V itho u t d o u bt o th er m at e
.

ria l elem ent s o f f oo d and d ri nk ex ert f u nd a m enta l i nfl u enc es

o f a ch em ica l n a t u re wh o s e eff ect s m a rea c h t he g e rm ina l m at

, y
t er and t hu s o f n ec es s i ty d es c end fr o m gen era t io n t o gen erat io n ,

to t he d is t inct m o d ificat io n o f t he rac e .

How races accli ma t e Ho w d o rac es b ec o m e acc limat ed ? .

T h ere a re at lea s t fi v e m eth o d s c o m pet ent t o exp la in t he

proc es s
I T he a cc li ma t iz a ti on of a l l tlze i n d i v id u a ls o f a rac e eac h
.
,

o n e in t he s u cc es s iv e g en e ra t io n s s ep a rat ely ;

2 S elec t ion o b lit erat ing s u ch ind iv id u a ls a s a re u nab le t o

.
,

acc limat e s u cc es s f u lly th u s res trict ing d es c ent t o t he fi t t es t ;

,

3 T. h e d i r ec t t r a n s m is s i o n o
f i n d i v i d u a l m od ifica t i on s
( a c

q u ired character s ) at leas t in s o m e s lig ht d egree t he acc u m u la

, ,

t io n o f wh ich u lt im at ely prod u c es c o m plet e acc lim at izat io n in

t he rac e ;

4 I.t is p o s s ib le th a t t h e s a m e ca u s es wh ich i nd u c e m od ific a

t io n s in t he ind iv id u a ls m ay a ls o ex er t i nflu en c es s o d eep s ea ted -

a s to af ec t tlze g er m p la s m d i r ec t ly a n d in th is wa pr d c e a ll
y o u

the appea ra nc es o f inh er itanc e o f m o d ific a t io n

D 1
p b b
uel y t p ifi
ro t ia t h h
o s t ec c ac on on e ea r .
3 8 6 TR A N SM I SS I ON

t he ger m and t he pro t o plas m o f t he d ev elo ped bo dy are es s ent ially

d iff erent has lo n g s inc e been d is pro v ed Bo th are s u s c eptib le .

t o any infl u enc e that can reac h th em and in c li m at ic c o nd it io n s ,

genera lly we ha v e ab u nd a nt infl u enc es o f th is k ind .

M ay we n o t c o n s id er a s es t a b lis h ed t he p os s ibili ty t hat t he

ger m it s elf and th erefo re d es c ent m ay b e d irect ly mod ifi ed by
, ,

ext erna l infl u enc es ? Ho w far t h is m ay go and what infl u enc es ,

a re i nc lu d ed is an o th er s u bj ect
,
and o n e ca lling fo r the m o s t ,

caref u l s t u dy and req u iri ng the m o s t reliab le data as a basis fo r

a n int elligent o p in io n .

T he pres ent s tat e o f kn o wledge is in s u ffi c ient t o ent i rely

s o lv e t he pr o b lem b u t th ere is add it io n al ev i d enc e wo rth
,

c o n s id erat io n .

SEC TI O N VI — EV I D E N C E FR OM HA BI T A N D I N ST I N C T .

I S I N ST I N CT I N H E R I T ED HA BI T ?

Hab it and i n s t inct

bo th r efer to t he u s e wh ich ind iv id u als
m ak e o f th o s e r a c ia l charact ers that a re capable o f act io n It .

is a m att er o f c o mm o n kn o wledge that an o ft repeat ed ac t -

s p eed ily b ec o m es a hab i t wit h t he ind iv id u a l and as s u c h , , ,

repeat s it self a lmo s t m echan ically ; so t hat what was at fi rs t

a n ic e ad a pt a t io n o f m eans to en d s h o rt ly b ec o mes litt le mo re
than refl ex act io n .

B u ild ing u p o n th is fact i t is a p la u s ib le a s s u m pt io n t hat

,

what is hab it wi th o n e gen erat io n b ec o m es in s t i nct i n t he n ext .

I t is a s weep i ng b u t ea s y gen era lizat io n that n o d is t i nct li n e

can b e dra wn b et ween in s t inct and reas o n ; that in s t inct is

1

inh er it ed hab it and rea s o n inh eri t ed in s t inct m o d i fi ed by ind i

,

v id u a l exp erienc e .

T h is is t he po s it io n tak en by m any o f t he o ld er nat u ra lis t s ,

es p ec ia lly R o m an es who d efi n ed i n s t i nct as ,

refl ex act io n int o
wh ich th ere is im po rt ed t he elem en t o f c o n s c io u s n es s 3
an d
”

reas o n as t he fac u lty wh ich is c o nc erned with t he int en tio nal

“

adaptat io n o f m ean s to en d 2
.
”

1 l
R o m a n es , A n im a I n t e igen c e, p I 5 ll . .

2 Ib i dp .1 7. T is v o
. l u m e ishp er a p s t h e m o s t h e xt rem e ex o n en t o f
p t he idea
of in h eri t e dh a b it , a n d o f ll
in t e igen c e as ly i n
g at t he b as is o f a ll an i ma l a c t iv it y.
 T R A N SM I SSI ON OF MODI FI C AT I O NS 8
3 7

T h u s t he who le qu es t io n is u p again in th is c o nnect io n Will .

th e hab it o f t he ind iv id u al b e t ran s m itt ed to it s o ff s pri ng ? I s

t he hab it o f o n e gen erat io n t he in s t inct o f t he n ext ? I f s o t hen ,

we ha v e a c as e o f t ran s m is s io n o f m o d i ficat io n s ( inh er itanc e o f

acq u ired charact er s ) of t he mos t d irect and c ertai n k in d .

T he an s wer to t he q u es tio n is i m po rtant fo r it s own s ak e b u t ,

m o re es p ec ially for t he light it may thr ow u po n t he ma in q u es

t io n n ow in hand To arr iv e at a s afe ans wer it is nec essa ry to
.

g iv e m o r e than a p as s i ng n o t ic e t o t he n a t u r e of i ns ti n c ti v e ac ts ,

and t o c rit ically d et erm ine whet her in s t inct is b u ilt u po n habit
o r hab it u p o n in s t inct .

Natu re of in st i n ct iv e acts Mo s t act s o f int elligent beings

.

a re perf ormed fo r a part ic u lar p u rpo s e and fo r d efin it e end s .

Mo s t s u ch act s a re c o ntro lled by a great er o r less d egree o f

p u rpo s efu l adaptat io n of mean s t o en d that in v o lv es knowledge
o f res u lt s ba s ed o n exp eri enc e and mergi ng nat u ra lly int o hab it
,
.

Hab it is t hen t he c u s to m a ry u s e t o wh ic h the ind iv id u al p u t s t he

part s with wh ich it is end o wed by nat u re aft er th ey hav e reach ed
,

f u ll d ev elo p ment I ts ch ief int eres t to u s at th is po int arises

.

fro m t he fact that ano t her ind iv id u al m ight p u t t he s a m e part s

t o a d ifferent u s e wh ile in i n s t in ct iv e act s u s e is a f u nc t ion of
,

s t ru c t u r e and b u t o n e s et o f act io n s is po s s ib le exc ept u nd er

,

great ly changed c o nd it io n s .

T he ter m i n s t inct iv e is app lied to th o se act s wh ich a re per

“

fo r med wi tlcou t p r ev ious exper ience and perhap s u nd er c irc u m

s tanc es that prec lu d e a ll kn o wledge o f what the res u lt will b e ;

so t hat a s far as t he i n d i v id u a l is c o nc ern ed t here is and can b e

, , , ,

n o c o n s c io u s n es s o f p u r p o s ef u l act io n o r o f adaptat io n o f m ean s

t o en d and yet t he act io n o ft en c o m p licat ed in the extrem e

, , ,

may b e em inent ly adapt iv e and exh ib it ev ery app ea ra n ce o f bein g

t he act o f a m o s t int elligent b ei ng Y o u n g m am ma ls s u ck with
.

the lip s yo u ng wat erfo wls swim and d iv e b u t land b ird s do n o t

,

yo u ng s q u irrels h id e o bj ect s and ev en in a roo m go t hro u gh t he

,

mo t io n s o f d igging and b u rying ; yo u ng c hick s ha v e th eir o wn

pec u liar c ry and peek at s h in ing o bj ect s ; b ird s b u ild th eir nes t s
,

wi t hou t in s t ru ct io n o r as s is tanc e fr o m old er o r m o re experienc ed

ind iv id u als .T h es e a re in s t inct iv e act s of t he s im p les t o rd er ;
m any o t hers ho wev er a re extrem ely c o m p licat ed Fo r exa m ple
, ,
.
,
 8 8 TR A N SM I SSI ON
3

an in sect b u rro ws a cav ity and lay s it s egg I t th en attack s .

an o th er in s ect s t ings it s o as to pa ra ly ze b u t n o t to k ill it d rags

, ,

it to t he b u rro w t u ck s it in n ext t o t he egg wh ere i t will s erv e

,

as f oo d t o t he la rv a s ea ls a ll u p and g o es a way, , .

T he y u cca m o th em erges fro m t he c o c oo n j u s t as t he y u c c a

o pen s it s fl o wer s each fo r a s i ng le n ight ,
The f em ale c o llect s .

po llen fro m o n e fl o wer and knead s it int o a b u nd le S he th en .

fl ies to an o th er m a k es a p u nct u re and lays her egg a m o ng t he

, ,

o v u les aft er wh ich s he da rt s to t he s t igm a and s t u ff s t he

,
“

po llen pellet int o t he f u nnel s haped o pen ing 1

I t is s u ppo s ed
-
.
”

that th is is t he o n ly way in wh ich t he in s ect r epro d u c es and the ,

ch ief way in wh ich t he y u cca is fert ilized .

Here d efin it e and im po rtant end s a re d epend ent n o t o n ly

u po n c o m p licat ed act s b u t a lso u p o n t he s er ia l o r d er of tltei r p er

r ma nc e a ll wi th o u t prev io u s kn o wledge o r expe rienc e o n t h e

f o ,

part o f the agent fo r t he fem ale in m o s t cas es is perf o rm ing

,

th is act fo r the fi rs t t im e and in m any cases will n o t liv e t ill the

,

egg s hatch Giv en any am o u nt o f int elligenc e th erefo re s he

.
, ,

c o u ld n o t kn o w t he fina l res u lt o f her o wn ind u s try al th oug h ,

t he ent i re pro c es s ha s ev ery appea ra nc e o f i nt elligent ev en d elib ,

c rat e act io n
,
I t is n o t ic ea b le a t o nc e that in s t inct s o f th is s o rt
.

a re c o nc ern ed wi th th os e act s wh ich lik e r epro d u ctio n a re f un da , ,

m ental t o rac e pres erv a t io n .

I s in stinct f ou n ded on h ab it ? T he o u tc o m e o f m o s t in s t in c
t iv e act s is s o c learly t he pres erv at io n o f life and t he goo d o f
t he s pec ies the act s th em s elv es a re o ft en s o extrem ely c o m p li
,

c a t ed th eir s eparat e s t eps a re s o n ic ely adj u s t ed to t he final

,

en d ,
th eir proper s erial o rd er is s o acc u rat ely o b s erv ed the ,

appearanc e o f d eliberat e p u rpo sef u l act io n is s o gen u ine t he

, ,

need o f int elligent d irect io n at so m e per io d o f t he o rgan izat io n o f

t he s e ries is s o apparent t he i m p o rtanc e o f t he end s ach iev ed is
,

so o b v io u s and t he s im ilari ty bet ween the in s t i nct o f a rac e and

,

t he hab it o f an i nd iv id u a l is s o c lo s e th a t m any nat u ra lis t s ha v e

leaped t o t he c o nc lu s io n that i n s ti n c t is in h eri t ed li a b i t in o th er
wo rd s that wh a t ha s b een f o u nd b en efic ial i n t he expe rienc e o f
,

ind iv id u a ls ha s b ec o m e hab it u a l with th em and thro u gh th eir ,

d es c endant s it has bec o m e t he hab it o f the ra c e Th is po sit io n .

1 A f ree t ra n sc rip t fro m M o rgan , H ab it and I n s tin c t , p . I 4 .

 0 T R A N SM ISS IO N
39
with it s n eighbo r s c a r t he baby with it s o wn fis t o r the you ng
’

, ,

la m b with a lo ck of woo l wh ich if t he m o th er b e y o u ng and , ,

in exper ienc ed it may s u ck u nt il i t s tarv es

,
Y o u ng ch ick s will .

p ick u p and s wallo w at firs t what ev er att ract s th eir att ent io n
—b it s of co lo red yarn o r nas ty caterp illars as read ily as
,

, ,

goo d wo rm s ; b u t th ey rapid ly learn by experienc e 1

.

T he i n s t inct o f t he yo u ng ch ick is t o s trik e at any s m all

o bj ect that attract s its att ent io n eith er by it s c o lo r o r i t s m o v e

m ent s ; yet t he fi rs t att em pt s a re extremely a wk ward and s el

d o m res u lt in a catch t he b ill go ing s o me d is tanc e to o n e s id e
,

o f t he o bj ect Grad u ally h o wev er it learn s by experienc e to

.
, ,

s trik e acc u rat ely

2
.

Walk ing fly ing s wimming a n d ta lk ing a re in s ti nct iv e act s

, , ,

wi th s pec ies po ss es s ing the req u isi t e mechan is m ; yet t he fir s t

att em pt s a re exc eed ingly cru d e and m u ch experienc e and prae ,

t ic e a re req u ired befo re effect iv e pro fic iency is d ev elo ped .

A fa i r s t u dy o f the s u bj ect can b u t c o n v inc e the s t u d ent that

i n s t inct is at fi rs t b u t litt le m o re than an im p u ls e t o act io n in
gen era l wh ich h owev er rapid ly s hapes u p int o well o rd ered
, , ,
-

pe rfo rm anc e u nd er t he c o rrect iv e infl u enc e o f expe rienc e .

I n st i n c t i v e acts ar e n ot perf orm ed in t h e same way eit h er b y

al l indiv id u als or b y t h e same i n di v i du al at su ccessiv e perf orm
an ces . To watch t he c o m p licat ed act s o f t he egg-lay ing i n s t inct
in m any in sect s is fi b ec o m e c o n v inc ed that th is m arv el
at rs t to

o u s s eq u enc e o f ev ent s is a ss u red o n ly by t he h igh es t in telli

genc e o r by an in s t inct that is u nerring in it s d irect iv e po wer ;

y e t a li tt le f u rth er s t u dy will c o n v inc e t he s t u d ent that th es e

c o m p licat ed in s t inct iv e acts a re no t a lway s carried fo rward o n

t he typ ica l p lan n o r a re th ey al way s s u cc es s f u lly ex ec u t ed
, On .

t he c o nt ra ry i m po rta n t ev en s ig n ificant s t ep s a re o ft en o m itt ed

, , ,

fro m t he s eries and d iff erent ind iv id u als d iffer great ly in t he

,

d egree o f th o ro u ghnes s and t he rap id ity with wh ich t he wo rk is

carried fo r ward .

F o r exa m p le C randall end ea v o red to n o t e acc u rat ely all t he

,

s t ep s in t he proc ess o f m ak ing t he p u nct u re and lay ing the e

gg
o f t he p lu m c u rc u lio wo rk ing u po n t he app le 8
.

1 Mo rga n Hab it an dI n s t in c t , p p 40- 4 4 , 5 0 3 Ibi

pp 3 7 , 4 7 d
—
, . . . . .

B u ll eti n ’ . , ll
A o 98 I in o is E xp eri m en t S t a tio n , p p 500 504 . .
 T R A N SM ISS I O N OF MOD I FI CAT I ON S 39 1

Of many att em pt s to wa tch and rec o rd the proc es s from

fi rs t to las t o n ly three were s u cc es s fu l in c o v eri ng t he ent ire
,

period The res t were fragm entary c o v ering o n ly p o rt io n s of

.
,

t he proc es s T h is was o wing t o t he d iffi c u lty of k eep ing t he

.

i n sect u nd er foc u s fo r t he fift een to t wenty fi v e m in u t es re -

q u ired fo r the c om p let e o pera t io n with o u t d is t u rb ing it s wo rk .

T he rec o rd o f the o b serv er is as f o llo ws

I n th e fi rs t o b serv a tio n the fem a le m ov ed ab o u t th e apple fo r s ev era l

sec o n ds , keeping the en d o f her b ea k i n c o n ta c t wi th the s u r fa c e, as if

see k in g a fa v o ra b le s po t . W h en was d ec ided u po n , th e
the ex a c t s po t

m in u te j a ws a t the en d o f the sn ou t b eg an a ra pid m o v emen t wh ic h q u ic k ly

m a d e a n open in g t h ro ug h the s k in T h is o pen in g was n o la rger th an n ec es
.

sa ry fo r a d m iss io n o f th e t ip o f the b ea k N 0 s k in was remov ed ; i t was

.

s i m p ly to rn a n d t h rus t as id e to gi v e a c c ess to the p u lp b elo w Later, as .

the ex c av a ti o n p roc eed ed , t he b ro k en s k i n was seen as a s o rt o f fri n ge

a ro u n d the b ea k a t th e s u r fa c e o f the fr u i t A s so o n as ex c a v atio n in the
.

p p
u l was c o m m en c ed , the b e a k was d efl e c ted b a c k wa rd so t h a t the w o r k
was c a rri ed o n u n d er the ins ec t, j us t b en ea th the s ki n a n d n ea rly parallel
wi th the s u rfa c e A s the work ad v a n c ed , th e o pen in g t h ro u gh the s k in
.

b ec a me s lig h t ly en la rged b y la ter al m o t io n s o f the b ea k T he p u lp was .

a ll ea ten as ex c a v a ted D u ri n g th e p roc ess th e b ea k was n o t o n c e wi t h

.

d rawn , n o r was t h ere an y c es s a tio n o f m o tio n W h en the ex c a v a tion o f

.

t he c a v ity was c o m p leted the b ea k was wi t h d r awn b y a q u i c k m o tio n , the

i ns ec t t u rn ed j
ab o u t, ad u s ted the t ip o f the a b d o m en to the o pen in g and
d epo s i ted an egg , wh ic h was fo rc ed to the ex t rem i ty o f t he ex c av a ti o n by
th e ov ipositor . T he ins ec t n o w res ted witho u t m o tio n for
m in u tes ; two
t h en , t u rn i n g aga in , p roc eed ed to c u t the c res c en t in fro n t o f the egg .

T h is c res c en t p u n c tu re was n o t wh o lly a s epa ra te pu n c tu re, b u t, s ta rtin g in

th e o rigin al o pen ing t ro u g h h k
the s in , was cut la terally in either d irec tio n ,
p a r t ly b y the j a ws an d p ar tly by c ro wd in g th e b ea k fi rs t
way a n d th en on e

t h e o t h er . T he d irec tion o f the b ea k was b u t li ttle d efl ec ted fro m the per

p e n d i c u la r , a n d the c u t wa s m ad e as d eep a s th e len g t h o f th e b ea k wo u ld
allo w . T he pu lp to rn a wa y i n m a k i n g th e c res c en t was ea ten j us t as was ,

d o ne i n ex c av a ti n g the egg c av i t y . T he c res c en t c o m p leted , the in sec t

walked away, d rew the legs c lo s ely u n d er the b o d y, an d s ettled d o wn ,
a p pa ren tly to s leep T he time o c c u pied i n the p ro c es s d es c rib ed was d is
.

t rib u ted a s follo ws

E xc a v a tin g egg ca v it y
D ep o s it io n o f egg

R es t
C u t t in g t he c resc en t

To ta l
 TR A N SM I SSI ON
39 2

T he egg c a v i ty was c y lin d ric a l with a ro u n d ed b o tto m an d b y mea s u re

, ,

m en t was fo u n d to b e in c h i d ep th T h e egg wh en d epo s i ted v ery

n .

n ea r ly filled t he c a v it y .

T he s ec o n d o b s erv a tio n of the c o m p lete p ro c ess wa s nea rly id en t i c a l

with the o ne d es c rib ed . T he i n sec t s p en t no ti m e i n c h o o s i n g t he ex ac t

s po t, b u t wen t to wo rk at o nc e . I t wo r ked in a m o re leis u rely wa y a n d d id

not ex c a v a te as d eep a n egg c a v i ty . E lev en m i n u tes wer e s pen t o n the
c a v i ty, two m in u tes i n d epo s it in g t he egg , two in res t, a n d fo u r in c u ttin g
t he c res c en t, a to ta l o f n i n eteen m i n u tes T he egg c a v it y m eas u red 5 .

in c h h
i n d ep t an d wa s c o m p letely filled b y the egg O n c o m plet io n o f t he .

p ro c e ss th e i n h
sec t mo v ed a s o r t d is ta n c e a n d im med i a t el
y b eg a n a s ec o nd

c a v it y.

E ss en t ia l d iff eren c es from p ro c ed u re in the two p rec ed in g c as es were

h
n o ted in the t i rd c o m p lete o b s erv a tio n E x c a v a tio n o f the egg . c a v i ty was

the s a m e, ex c ep t t a t i t h
was d eeper in th e p ulp a n d o f g rea ter ex ten t A fter .

d epo s i ti n g the egg the b eet le tu r n ed , a n d wi th h er bea k wo rk ed th e egg

b a c k to the b o t to m o f the c a v i ty Tli en slt e beg a n tea r i ng of bits of s k i n
.

a n d pu lp , wl uclt wer e ca r e l
f ly p k d i b l l l w
'

u a c e n , a ov e t t e e
gg, u n t i t ze ca v i ty as

o llo win g th is , the c res c en t was c u t i n m u c h the sa m e m an n er as i n

1
u ll F
f .

th e p rec ed in g c as es T h en s he a ppea red to m a k e a fin a l i n s pec t io n , a nd

.

add ed s o m e fur t h er p a c k i n g ab o v e the egg F in a lly the wo r k a p peared to .

b e s a t is fa c to ry a n d s he wa lk ed awa y a n d b ega n a s ec o n d p u n c tu re T he .

t im e c o n s u med i n th is p ro c ess was lo n ger t h a n in th e o t h ers , a n d wa s d iv id ed

as follo ws :

E xc a v a tin g egg c a v it y
D epo s it in g egg

Pla c in g t he eg wit h the b ea k

k
Pac in g t h e c a v it y
C u t t in g t he c resc en t
h
Fin is in g t o u c hes

T o t al

A m on g the m a n y c ases wh ere o n ly pa r t o f t he p r o c es s was ob serv ed

so me a n o m a lies were n o ted . I n two c a ses the i n sec t walk ed awa y im

med ia tely a fter d epo s i tin g the egg a nd mad e no c res c en t c u t . In t h ree
c a s es b eetles were s een to c u t c resc en ts an d , m o v in g a s hor t d is ta n c e,
b egi n ot h er p u n c tu res . T h ese c res c en ts h ad n o egg c av i ties a n d n o eggs

were d epo s ited in h

t em . I n two c as es eggs were fo u n d d epo s i ted d irec tly
in c res c en t c u ts , n ei t er o f h wh ic h h ad the u s u al e
gg c a v it
y M a r k ed v a ria.

tio n in d ep th o f the egg c a v i ty was freq u en tly ob s erv ed . N o t i n freq u en tly

the c av i ty is so s h allow th a t the tip o f th e egg p ro t ru d es , a n d so m etimes
i ts d ep t is h n ea rly eq u a l to twi c e the len gt o f the egg P a ck i ng tlze eg g h .

ca v ity wi t/t i l i m m b u n i v er sa l , r ac tic e; o ften

p ec es
of p p
u s a c o o n , u t n ot p
h
t is is n e lec ted , ev en w ere t he c a v i t
g i s d ee
p h y .

1 l
I ta ic s a re min e .
 T R A N SM ISSI O N
3 94

t he t wenty prev io u sly rec o rd ed o b ser v a tio n s n o two agreed and ,

n o t o n e ga v e a c o m p let e acc o u nt o f what m ay b e ca lled t h e

typic a lly in s t inct iv e proc es s B ea ring a ll th es e fact s in m ind we

.
,

a re n o t t o s u ppo s e th a t s t ill m o r e c o m p licat ed pr o c es s es a re

s u cc ess f u lly carr ied f o r ward in ev ery in s tanc e a s we ha v e been ,

led t o infer . I n s t inct is th erefo re n o t u n erring n o r d oes it

, , ,

in s u re u n ifo r m i ty o f proc ed u re I t loo k s as if i m po rtant link s

.

in t he cha in o f im p u ls e m a y b e o m itt ed o r if the se r ies is int er

, ,

ru p t ed for any ca u s e ,
it m ay b e res u m ed at a lm o s t any po int .

C erta in it is that d ifferent s er ies a re far fro m u n ifo r m and that

v ery m any o f th em fa il o f th eir ev id ent and fina l o bj ect .

I n sti n cti v e ac t s ar e n ot al wa ys ada pt iv e t h eir ori gi n is t h er e

f or e n ot i n pu rposef u l act s . I t is o f n ec es s ity tr u e that i n mos t
c a s es in s t inct iv e act s a re fo r t he g oo d o f t he s pec ies a n d th ere ,

fo re by d efi n it io n adapt iv e If th ey were n o t fo r the good o f

, ,
.

t he s pec ies th ey wo u ld s peed ily lead t o ext inct io n wh ich is b u t ,

an o th er way o f s ay ing th a t in s t inct s n o t adapt iv e ha v e lo ng

s inc e b een b lo tt ed o u t by s elect io n a lo ng wi th t he i nd iv id u als
,

in wh ich th ey aro s e .

T hat s u ch n o n a dapt iv e in s t inct s e xis t h o wev er is eas ily

-
, ,

s h o wn .Fo r exa m p le wh en t he m o t h fl ies int o t he fla m e a n d is

,

k illed by n o s tretch o f the im aginat io n c a n th is b e called an

,

ad a pt iv e act ; n o r can it b e c o nc eiv ed as h a v ing aris en in t he

p u rpo s ef u l act s o f its anc es t o r s I t n ev er c o u ld ha v e s erv ed
.

any b u t an ev il p u rpo se to the rac e and t he o n ly element that

,

n o w s t a n d s in t he wa o f t h e u tt er ext inct io n o f rac es po ss es s ing

y
th is in s t inc t is t he rela t iv e in freq u ency o f t he nak ed flam e so ,

that c o m pa rat iv ely few ind iv id u als s u ffer to o few to affec t ,

eith er t he r a c e o r it s in s t inct s W e m u s t th erefo re lo o k far th er

.
, ,

than habit fo r the o r ig in o f in s t inct s .

I n st in c t s origin a t e in reflex ac ti on n ot i n h ab i t I f a p iec e o f

.

m ea t b e la id u po n t he t ent a c les o f an act in ian th ey will imm e

d ia tely c o ntra ct info ld i ng t he m ea t and carry ing it i n t o t he
,

m o u th .I f a p iec e o f pap er b e la id u po n t he t ent a c les n o act io n

fo llo ws ; b u t if t he paper b e c o v ered with t he ju ice o f m eat t he
act io n is t he s a m e a s if the p iec e were v a lu a b le fo r food .

T enta c les th a t ha v e ari s en fro m a wo u nd in t he s id e o f t he bo dy

will rea ct to m ea t and m ea t j u ic e a s d o t he n o r m a l parts ,
 T RA NSM I SSI ON OF M OD I FI CAT I ON S 3 95

in fo ld ing and pres s ing t he piec e again s t the bo dy as if end eav o r .

in g to t u ck it int o a mo u th th o u gh n o n e is pres ent

,
1
Here it is .

n o t int elligenc e that a ffo rd s t he bas is o f m u s c u lar c o ntract io n

b u t clzem ic a l a c tion o f t he meat j u ic es u p o n t he m u s c le fib er s of

t he t entac les T he s a m e pri nc ip le gov ern s m o t io n and s ecret io n
.

in in s ect iv o ro u s p lant s to wh ich n o o n e wo u ld as cr ibe ev en t he

,

elem ent s o f int elligenc e Od o rs exc ite t he s a liv ary g land s and
.

m ak e the m o u th wat er b u t it is c o ntact that s tart s t he sec re

,

t io n of gas tric j u ic e in the s t o mach .

L ight s t im u lat es s pec i fic react io n s in m any fo rm s o f pro t o

plas m and many t iss u es c o ntract u nd er it s infl u enc e I t is th is
,
.

c o ntract io n that cau ses the b end ing o f s t em s t o ward the light .

T he iris o f t he eye c o ntra ct s n o t by n er v o u s im p u ls e b u t by

,

t h e act io n o f t he light ca u s ing d irect ly m u s c u lar c o ntract io n

, , , .

L oeb repo rt s that he ha s o ften o b ser v ed the c o ntract io n of t he

2

i ris o f d ea d s hark s u nd er t he i nfl u enc e o f light s ev era l h o u rs

after d eath wh en s ign s of d ec o m p o sit io n had already begu n

,

t o appear .

L o ng bod ied in sect s if ly ing with t he s id e t o t he light will

-
, , ,

becau se o f th is hav e th eir bod ies bent with the c o ncav e s id e

, ,

n ext t o the so u rc e o f light if po s it iv ely h eliotro p ic Wh en ev er .

th ey m ov e in th is co nd it io n th ey m u s t o f n ec ess ity m ov e in a
c u rv ed in s t ea d of a s tra ight lin e u nt il s u ch t im e as th ey are
,

h ead ed d irect ly t o ward the light F ro m that t ime t he body is .

eq u ally illu m inat ed on bo th s id es I t th erefo re bec om es and.

rem a in s s tra ight so that f u t u re m o t io n m u s t b e in a s traight

,

lin e any d ev ia t io n being q u ick ly c o rrect ed by t he u n eq u a l illu m i

,

nat io n o f t he body I t is th is s eries of fact s arising from the

.
,

nat u ral r elat ion bet ween light and prot o pla sm and n ot c u rio sity , ,

that acc o u nt s fo r t he fly ing o f t he m o th int o the cand le N ega .

t iv ely h elio tro pic a n im als wo u ld o f c o u r se b eha v e in exact ly t he

o ppo s it e m ann er b u t for t he sa m e g en eral r ea so n s
,
.

I n s ect s and s m a ll wo rm s a re sa id t o b u rro w int o dark plac es

for t he p u rpo s e o f h id ing T h is cann o t b e tr u e fo r u nd er d irect
.
,

exper im ent s m all an im als o ft en em erge fr om d a rkn es s t o light ,

fro m h id ing to expo s u re u nd er t he im pelling fo rce o f an in

,

s t in c t to br ing th eir bo d ies int o con ta c t with a s m any s u rf ac es as

1 L b Ph y i l gy f t he B i pp 4 8—
oe , s o o o
54 ra n ,I b id p 4 . .
2 . . 0 .
396 T R A N SM I SSI ON

po s s ible I n a b o x they will crawl a lo ng the bo tt o m t ill th ey

.

com e t o a s id e Here th ey can t o u ch t wo s u rfac es and mo t io n

.
‘

will th en b e alo ng t he g roo v e wh ere t he s id e m eet s t he bo tt o m

u nt il th ey r ea ch t he c o rn er wh er e a th ird s u rfac e j oin s wh en
, ,

th ey are lik ely n o t to t u rn the c o rn er ( u nles s im p elled by som e

s tr o nger in s t inct ) b u t t o c o m e to r es t in c o n ta ct with thr ee

s u rfac es th is affo rd ing appar ent ly

,
t he h igh es t atta inab le sat is
, ,

fact ion I f a t u b u lar o pen ing b e fo u nd in sect s o f th is in s t inct

.
,

will cro wd int o it if po s s ib le o r at lea s t m ak e t he att em pt

, ,
1
.

T h is in s t in ct to s eek great er c o m f o rt by gett ing s n u gly plac ed

in c o n tact with fo reign bod ies is pr es ent in t he h igh er an im als
and in m an a n d it acc o u nt s for m any o f t he m ov em ent s an d
,

r es t ing po s it ion s so c o m m o n ly s een .

T hat th is act io n is n o t t he res u lt o f a p u rpo s e t o h id e is

ev id enc ed by t he fact that n eith er light n or darkn es s has an
y
effect u po n it and that t he in s t inct is n o t changed ev en by t he
,

rem ov al o f t he brain I n nat ur e o f c o u r s e plac es t hat will sat is fy

.
, ,

th is in s t inct a re generally s h u t a way from light and in sect s so ,

bes t owed are a lso h idd en — a fact that ha s g iv en r is e t o t he

,

trad it io n th at t he pu r po se w a s t o s ecr et e th em selv es fro m pred

a t or en em ies O f c o u r s e t he trad it io n it s elf is n o t c o n s is t ent

y .
,

for int elligent an im als s eek ing foo d soo n learn t he fa v o r it e ha u nt s

o f th eir pr ey T h ey th erefo re kn o w prec is ely wh ere to loo k fo r
.

th em an d s peed ily t u rn th em o u t .

W e ha v e a lready s een that pro t o p la s m m ay b e exc ited to

act io n by a great v ariety o f ext ernal agent s (light h eat elec , ,

t ric it y ch em ical s u b s tanc es ) ; that the charact er o f pro to p las m ic

,

act iv ity may b e m o d ifi ed by c ert ain o f t hese external fo rc es ,

n o tab ly light and ch em ica l att ract io n ; that t he d irect io n o f

m o v em ent o r o f gro wth m ay a ls o b e infl u enc ed by t he s a m e
c las s o f agent s (light ch em ical s u b s tanc es h ea t gra v ity ) and
, , ,

that in all th es e way s the act iv it ies o f liv ing beings a re largely
d epend ent u po n t he nat u re o f t he o u t s id e fo rc es with wh ich th ey
c o m e int o c o ntact Here lies t o a v ery large ext ent the in it ial
.
, ,

cau se o f th o s e ext ernal act s we o rd inarily s peak o f as in s t inct s ,

and t he rem ain ing elem ent s o f ca u s at io n a re to b e s o u ght in the

in t ern al m echan is m o f t he creat u re I n all lik elih oo d it is n ot .

L b Phy i l gy f th B i p 93
1 oe , s o o o e ra n , . .
 9 8 T R A N SM I SSI ON

an d c o m p licat ed than th o s e s em i m echan ica l act s c o nn ected with

-

t he d epo si t io n o f egg s and t he care o f y o u ng wh ich a re th em ,

s elv es i n th eir es s enc e n o t far rem o v ed fr o m v it a l pro c es s es .

A ll m o t io n is red u c ib le t o i r r i ta b ili ty and c on tr a c t ili ty o f

r oto la s m as i t s u lt im at e ca u s e and any i m p u ls e that will
p p , p r o

d u c e th is effect will lead to act io n Fu rth erm o re th is act io n .

,

m u s t fro m t he nat u re o f t he ca s e b e charact eris t ic o f t he

, ,

o rgan is m and i ts p ec u liar m echan is m .

I t has b een h eld that all m u s c u lar act iv ity m u s t ha v e it s o rig in

in a n erv e s t im u lu s o f s o m e s o rt T hat th is is erro n eo u s is s elf
.

ev id ent M u s c le t is s u e u n s u pp lied with n er v e is s t ill c o nt ract

.
, ,

ile and t he im p u ls e can s t ill b e s u pp lied fro m a v a riety o f

,

s o u rc es ( h ea t light elect ric ity c o ntact )

, , ,
.

N er v o u s im p u ls e is b u t on e o u t o f m a n y s t im u li t o m u s c u la r
c o ntra ct io n o r o th er act iv ity o f liv in g pro t o p la s m th o u gh i t is
, ,

t he m o s t ra p id and di rect and t he o n e m o s t rea d ily u n d er c o n

,

tro l of t he m ind a n d the will R es tat ing the pro po s it io n n er v o u s

.
,

m echan is m is n o t n ec ess ary to m o t io n n o t ev en t o c o o rd inat ed ,

m o t io n o f a h igh d egree b u t i t is n ec essa ry t o the h igh es t c o or

,

d in a t io n o f t he m o s t c o m p licat ed o rga ni s m s ; it is n ec es s a ry a s
t he m ea n s by wh ic h t he will m a y q u ic k ly reach a ll part s o f t he
m ach in e and d i rect o r s et a s id e mere i n s t i nct iv e m o t io n ; it is
nec essary fo r t he realizat io n o f all the po s s ib ilit ies o f wh ich a
h igh ly o rgan ized s tru ct u re is capab le ; it is n o t n ec es s ary to
act io n o r ev en t o a h igh d egree o f c o m p licat io n in act io n
,
.

An y effec t i v e im pu l se will serv e t o s t i m u l a t e t o acti v it y

an d ,in gen eral wi t h al l com pl icat ed act ion s each act b ecomes ,

t h e impu l s e f or t h e n ex t I f t he h eart o f a fro g b e c u t i nt o s ev

.

era l p iec es th es e will a ll b ea t rhyth m ica lly b u t t he n u m ber o f ,

c o ntract io n s will v ary in t he d ifferent p iec es The s in u s v eno s u s .

beat s m o s t ra pid ly and t he n u m ber o f it s c o ntract io n s in a u ni t

,

o f t im e eq u a ls that o f t he h eart b ef o r e it wa s d iv id ed Tla cs we .

s ee tlza t tlze wlzole lt ea r t bea ts in tb c r lcy tlzm o f tlze pa r t tlt a t lza s

tlze m ax i mu m n u m b er o f c o n tr a c t io n s p er m i n u te . Fr om i lt is we
’
m us t a ss u m e tlta t tlt e coor d i na t i on f
o tlt e lzc a r t s a c t i v i ty is d u e to
tlt e f a c t tlza t tlce p a rt w/ziclz c o n tr a c ts m os t f r eq u en t ly
f orc es t/te
1
otlzer i n tlte s a m e r lcy tlcm
”

p a r ts to con tr a c t .

1 Lo eb , P hy s io lo g y of t he B ra in , p . 25 .
 T R A N SM I SS IO N O F M OD I FI CAT I O N S
3 99

T h is h ows n o t o nly that a c ent er of c o ord ina tio n ex is t s in

s ,

t he o rgan it s elf ind ep end ent of n e rv e c ent ers b u t that t he

, ,

t i on
of p on e a rt becom es tlze i mp u ls e f or ex c i t i n
g a c ti o n i n i ts
n ei lzb o r i n t
g gp a r .

Hydro med u sae d ifferen t rat es o f p u ls at io n were u n ited in

of

pair s in L oeb s lab o rat o ry by the pro c es s of graft ing Wh en the

’
.

u n io n wa s c o m p let e a lo ng n ear ly a ll the c u t edge t he wh o le

beat s ynchro n o u sly b u t when the u n io n c ov ered b u t a s mall

,

area the two beat s eparately with a d iff erent rhyth m 1

.

T he h eart o f t he a s c id ian is an elo ngat ed t u b e beat ing s o a s ,

t o s end the b loo d a lt e rn at ely fro m left t o r ight a n d fro m right

t o left ; that is the im p u ls e s eem s t o o rig inat e at o ne en d fo r a
,

t im e and th en after s ev eral h u ndred b eat s t o s h ift to t he o th er

, , ,

en d I t was f o u nd that the area o f im p u lse was c o nfin ed t o a

.
“

S h o rt sect io n at ei th er en d ; that each o f th es e s ect io n s if c u t ,

a way c ont in u ed to beat rhyth m ically b u t that the lo nger m idd le

, ,

part seemed incapab le of c o ntract io n with ou t ext ernal s timu lu s .

C o m ment ing o n the fact L oeb s ays : 2

,

The e pe imse t i t
ex m s t
ro m e le e o
n s, m fo d o
s ee b t t h t th e , av n roo r u a

c h ge i the d i ec tio i the c t ti

an n r f the id i
n s h
n t i d eteo n rac on o as c an
’
ea r s r

m i ed b y e h f th two e d getti g the ppe h a d l tern tely

n ac o e n s d fo c i g n u r n a a an r n

the o th e c e t t ac t i
r it h yth m f
n er time
o Th i “
gent t i g s th e r pp or a . s n u er

h a d migh t p ib ly me
n ot h i g m
o ss e th a th t e e d g i
an n the timen or n a on n a ns

i whi h to e d ff
n c w e f c o tr ti
s n o b efo e the th e e d b gi
a av o ton ac on r o r n e ns

c on tr c t F
a th i it i
. l y ec essa y th t a i gle h e t b e t o f the le d
or s s on n r a s n ar a a

i g e d b e d el yed o f il e ti ely p h e ome o t h t l o pp

n n a r a n r io
, a n n n a a s a ea r s o c c as n

ally i the h m a h e rt
n u n a .

L oc o mo t io n in t he earth wo rm is by a series of elonga t io n s and

c o nt ract io ns o f the s u cc es s iv e s egm ent s of the bo dy in reg ular ,

o rd er fro m t he fro nt back ward N o b o dy ev er s u ppo s ed th is s eri al .

o rd er t o b e c o ntr o lled by c o n s c io u s i nt elligen c e b u t i t h as b een ,

as s u med t o b e d u e to t he c o ntrol o f ner v e fibers from t he gang lia

alo ng t he d o rs al s u rfac e o f t he bo dy I f h o wev er the wo r m b e .
, ,

c u t in t wo and the part s u n it ed by thread s lo c o m o t io n is ent i rely ,

s u cc es s f u l ev en th o u gh a c o n s id erab le s pac e int er v en es b et ween

,

t he p iec es I n th is ca se c o ntract io n proc eed s back ward seg

.
,

m ent by segm ent as in u n inj u red wo r ms s u ffering n o s pec ial

, ,

1 Lo eb , P hy s io lo
gy of th e B ra in , p p . 2 6—7 2 .
9 Ibi d .
pp . 2 7- 2 9 .
 00 T R A N SM I SS I O N
4

i nt erru pt io n at t he po int wh ere t he n erv e c o nn ect io n is sev ered .

The thread s erv es pe rf ect ly t o carry t he i m p u ls e o v er fro m t he

las t s eg m ent o f t he ant er io r p iec e to t he fi rs t o f t he p o s t erio r .

T he u n av o idab le c o nc lu s io n is that at lea s t fro m th is p o int back

ward each s egm ent d eriv es i ts im p u ls e no t f r o m n er v e fi bers b u t
m /ze s eg m en t ju s t a lzea d i n o th er wo rd s that t he m o t io n o f
f r o t ,

o n e part bec o m es a s t im u lu s t o appr o pr iat e act io n in a n eighbo r

in g part .

I t is rep o rt ed that R ibb ert tran s p lant ed a m ilk gland t o t he

ea r o f a g u in ea p ig and that wh en t he ind iv id u a l b eca me preg
,

n a nt t he gland c o mmenc ed to s ecret e milk What ev er t he .

nat u re o f t he s t im u lu s it was c learly i nd epend ent o f n erv e

,

im p u ls e 1
.

I t is a well kn o wn fact that if an ant b e remo v ed fro m it s

-

n es t fo r a t im e and th en p u t back it will b e crit ic ally exa m ined , ,

b u t will b e rec eiv ed aga in ; if h o wev er a s tranger ant b e intro , ,

d u c ed it will at o nc e he attack ed and k illed

,
Ho w is t he d iffer .

enc e d et ect ed ? T h is q u es t io n is largely an s wered by t he fact

that if the ant b elo nging t o the n es t b e s m ea red wi t/t tlzeju ices
,

of a c r u s lzed s t ra ng er i t will at o nc e he attack ed and k illed as

,

wo u ld t he rea l s tranger 2
C lear ly it is by t he o d o r that t he ant s
.

d et ect t he d ifferenc e m u ch as the d o g rec o gn izes his mas ter in

,

day light o r in the darkn es s o r fo llo ws a t ra il alo ng a cro wd ed ,

s tr eet A ll t his s h o ws that ev en a s light ca u s e lik e o d o r m ay

.
, ,

s er v e t o s tart t he o p erat io n o f a tra in o f m o s t rem arkab le re

fl ex es wh ich o nc e s t a rt ed proc eed s a u t o m at ically each act

, , , ,

o perat i ng a s a s t im u lu s to t he n ext .

T h is fact is f u rth er illu s t rat ed in the ca s e o f d ogs d epr iv ed o f

large po rt io n s o f t he n erv o u s sy s t em I nd iv id u a ls that ha v e .

lo s t t he s p inal c o rd “
alm o s t u p to t he m ed u lla may liv e for ”

years and perfo rm all n o r mal f u nct ion s 3

.

G olt z ent irely r emo v ed b o th h em is ph eres o f t he brain from a

f u ll gro wn d og T he fir s t effect s o f so v iolent a n exper iment are
-
.

appar ent ly d isa s tro u s b u t if s k illf u lly d o n e t he s h o ck soo n su b

,

s id es and a ll n or ma l n c ti ons o i b d d b
f u f t re o y p r oc ee as f
e or e .

T hat is t he a n imal perfo r m s t he ext ernal act s o f eat ing u rin a

, ,

t io n d efecat io n et c t he sa m e as wh en in po s sessio n o f the brain

, ,
.
, ,

1 L b Phy i l gy f t h B i
oe , s o o
p o 6 I b id pp
e ra n , .Ib id p 4 3
20 .
2 . . 2 2 0- 2 2 1 .
3 . . .
 0 2 T R A N S M I SS I O N
4

wo nd erf u l se ries of wh ich it is it self b u t a part ,

an d of wh o se
beginn ing an d end it is alik e ign o rant .

T he in s t inct t o s u c k cann o t b e c o nc eiv ed as fou nd ed o n hab it ,

beca u se s u ck ing is n ot a life hab it with any ind iv id u al I t is .

pract iced b u t a br ief seaso n aft er birth and th en aband o ned , ,

lea v ing n o trac e o r ev en im p u ls e beh ind .

I t is d iffic u lt for u s to d isso c iat e th es e c om p licat ed a ct s from

t he id ea of int elligent c o ntro l Y et m any o f th em a re perfo rm ed
.

by plant s in wh ich th ere is n o t s o m u ch as the beginn ings of a

,

n erv ou s sy s t em t he im pu ls e tra v eling fro m c ell t o c ell as it is

, ,

ent ir ely capab le o f d o ing in an im a l t is s u e b u t at a rat e ea s ily ,

o v ert a k en by n erv e tra n s m is s io n wh en ev er t he latt er is s u perim

,

posed by the will or o th erwise .

A ga in we m u s t c o n s id er t he exc eed ingly co m plicat ed nat u re

,

and s erial o rd er of the v ital pro c es s es gen erally Mo s t of th ese .

proc es ses it is tr ue as id e fro m c o p u lat io n t he lay ing o f the egg

, , , ,

and the care of the y o u ng a re carried o n i ns i de t/ze body a n d

, ,

tlzere f or e o u t of gs z lt t f
o tlze o b s er v er I f we c o u ld by s o m e .

m enta l m icr os c op e s ee n o t o n ly t he p u ls at io n o f t he h eart the ,

m o v em ent s of the s t om ach a n d int es t in es and t he d is charge o f ,

acc u m u lat ed s ecret io n s b u t a ls o t he int ernal act s of secret io n

,

go ing on with in the v ario u s gland s of t he body with t he a s s oc iat ed ,

pro t oplas m ic mot io n and cell d iv is io n act iv ely acc om pa n ied by ,

t he caref u l d iv is io n o f t he chro m o s om es int o exact ly eq u al po r

t io n s q u a litat iv ely as well a s q u ant itat iv ely — if we c ou ld s ee
all th is as we see ext ern a l in s t inct iv e act s we s h o u ld b e led to ,

m arv el at t he m y s t ery and t he c o m plic at io n o f v it a l act iv i ty in

gen eral \Ve sh o u ld in v o lu ntar ily s eek a ba s is o f int elligent c on
.

t rol wi t/t i n t he o rga n ism wh er eas we kn o w that t he pro per

,

plac e to s eek ca u sat io n is ou t s id e t he creat u re in t he fo rc es

that s h aped it s d ev elo p m ent and in t he h igh er po wer that en
d o wed it all with life wh os e charact er is t ic act is m otion an d
,

appro pria t io n of n ew m at erial W e are th erefo re n o t t o b e .

, ,

m isled by t he c o mplex ity o f a ct s ha v ing t he appeara nc e of

int ellig enc e .

We hav e b e en led to proj ect intelligenc e too far d o wn t he

s ca le . I t m ay if pres ent tak e h o ld o f and ov erru le m o s t ( n o t all)
, ,

in s t inct iv e act s b u t t he v a s t m aj o rity of o rgan ic act iv i t ies go on

,
 TR A NSM ISSI ON OF MOD I FI C AT I ON S 0
4 3

with o u t it Y o u ng th ings hav e litt le or n o s en s e of fear at

.
1

b irth and at the firs t c o n sc io u s n ess of fear m ak e n o th ing lik e

,

int elligent d iscr im inat io n T he y o u ng ch ick so L loyd Mo rga n

.
,

t ells u s is as m u ch afra id o f a fly ing n ews paper as o f a hawk

, ,

and has n o preconc eiv ed n ot ion of the danger from bees wh ich ,

can s t ing as c o m pared with that from fl ies wh ich are harm les s
, , .

I t s fir s t fear s are o f “
largi s h th ings b u t exper ienc e rap id ly ,

info r m s it o f s pec ific th ings I t has at fi r s t n o apprec iat io n o f

.

wa t er as wat er b u t wh en it acc id entally peeks at a s h in ing dr o p

,

and wet s t he bill t he presenc e o f t he wat er s tart s t he s eries of

,

refl ex es and t he ch ick h o ld s u p it s h ead to s wallo w or if a d u ck , , ,

it h
s ov e s l — each o rgan is m react s in it s o wn c ha rac t eris t ic

—
, .

m ann er and both learn rap id ly by exper ience

,
2
.

Hab it s foll ow an d are fou n ded u pon instin ct s T he tru e o rd er .

seem s t o b e that t he ear lies t att em pt s a r e in s t inct iv e b u t that ,

th ey are rapid ly s haped u p and perfect ed by exper ienc e and in ,

th is cond it ion th ey bec om e habit u al I t als o appea r s that n o .

exact lin e can b e dra wn b et ween what we ca ll in s t inct and what

is n o th ing b u t res po n s e to s t im u li Man if es t ly we hav e applied .

t he wo rd “
in s t inct t o th os e refl ex act s that ha v e t he appearanc e
of being p u rpo s ef u l I t wo u ld apply eq u a lly as well to m any act s
.

c lear ly refl ex and wh en it is s h own that th es e so c alled in st in c

,
-

t iv e act s are th em selv es o n ly refl ex es and can b e perfo rm ed per

fec tly in t he ab s enc e o f a ll po s s ib le applicat io n o f int elligen c e ,

eith er o n t he part o f t he ind iv id u al o r o f it s anc es t o r s it appea r s ,

tha t o u r pres ent u se o f t he word is a c o n v en ienc e rath er than a

fa ir m ark of a sc ient ific d is t inct io n .

T he th eo ry th er efo r e wh ich plac es hab it at t he po int o f o r igin

, ,

o f in s t inct iv e act s c lea r ly tak es it o u t o f it s pr o per o rd er in t he

s eries I t is a pr operty o f t he ind iv id u a l rath er than o f t he rac e

.
,

exc ept a s we s peak of t he hab it o f a rac e m ea n ing th er e ,

by t he hab it d ev elo ped by all or mos t o f the m em ber s o f the

ra ce T he c ommon d ev elo pment o f s u ch a hab it is n o t u nnat u ral
.

s inc e all of t he ind iv id u a ls o f t he rac e po ss es s t he s am e o r gan s

1 Th e a c t io n o f t h e h ea rt a n d o f t h e s ec r et in g o rg a n s is b e y d
on t h e c o n t ro l of

t he wi ll , an d h
t a t o f t h e in t es t in es is l argely so ; b u t m an yp a rt s o f t h e o rga n is m

are u n d er c o n t ro l eit h er o f t h e wi ll , of im p l u s es in tern a l t o t h e p a rt , o r o f c a u s es

ext ern a l t o t h e o rgan is m . 9 Mo rgan , H ab it an d I n s t in c t, pp 40- 90
. .
 0
4 4 TR A N SM I SSI ON

and are for the m os t part s u bj ect ed t o t he sa me s u rr ou nd ing

c ond it io n s .

C aref u lly exam in ed th is fi eld int eres t ing a s it is o f it self is

, , ,

ba rr en o f ev id enc e u po n t he t ran s m is s io n of habit s th o u gh it ,

is t he on e m os t o ft en appea led t o fo r pr oo f of t he in h eritanc e

o f acq u ir ed c haract er s The error lies in a ss u m ing a cau sat iv e
.

relat io n bet ween t wo act s wh ich are sim ilar I t is tru e that t he .

hab it s of t he in d iv id u al and the in s t inct s o f the rac e are s imilar .

T h ey c o u ld n ot b e o th er wise s eeing that bo th d epend u po n t he

,

pres enc e of su itable o rgan s with ou t wh ich the pa rt ic ular act

,

wo u ld n ot b e po s s ible and with wh ich it is c erta in t o appea r

,

wh en ev er s u itable s t im ul u s is enc ou nt ered b u t hab it is fou nd ed

u po n in s t inct and n o t in s t inct u po n hab it .

The c o nc lu s io n wh ich seem s in ev itable is th is : tlza t b reed ers

n eed ha v e no f ea r tlza t tlt e li a b i t of a n i n di v id ua l, as s u e/t , wi ll be
i n ker i ted by i ts oflspr i ng b u t tlzef a ct tlza t tlze li a b i t d ev elop ed a t
'

a ll is s u ic i en t r ea s on
fi f or k n o w i n
g tlza t i ts d ev elop m en t is a lway s

p oss i b le in tlt ef a m ily li ne, w/t en ev er s u i ta b le c on d i tions a r i s e

.

T he att ent io n o f the st ud ent is called t o the fact that wh ile

we ha v e s h o wn t hat in s t inct is n o t t he res u lt o f hab it a n d th ere ,

fo re that it s ex is tence is n o proo f of the tra n sm is s io n of habit u al

act s yet th is d oes n ot s h ow wh eth er or n o t the habit u al u se o r
,

n o n u s e of a part will a ffect t he int en sity o f t ran s mis s io n o f that

-

part or the t end ency t o m ak e u se of it Hav ing pu t in st inct

, .

beh ind u s wh ere it properly belo ngs we hav e n o w to inq u ire into
,

t he effect s o f u se and d is u se .

SECT I ON V I I EV I D E N CE FR OM U SE A N D D I SU SE

The q u es t io n is n o t wh eth er u se d ev elo ps and d isu se lead s to

n o n d ev elo p ment o r d egen eracy
-
T he fact s on that po int are
.

already well kno wn The inq u iry is wh eth er the effect s of u se

.

and d is u se a re tran s m itt ed ; wh eth er th eir infl u enc e is d irect ,

n o t m erely ind i rect by r end ering t he ind iv id u als les s o r m o re

ab le to meet the d emand s of select io n ; wh eth er s pec ializa t io n

o f a part is i n any wa d e t o u s e a s id e fro m it s effect in
y u ,

d ev elop ing ind iv id u als separat ely and as id e fro m it s c o nnect io n

with i ncreas ed rigo r of s elect io n ; wh eth er gen eralizat io n and
 06 T R A N SM I SSI O N
4

Weis m ann s xact wo rd s c o nc ern ing t he two v ariet ies of

’
e

b u tterflies — t he dark er I ta lian and t he l ght er G erm an

i — a re

as f o llo ws
1
T he t wo v ariet ies may ha v e o riginat ed o wing t o
a grad u al c u m u lat iv e infl u enc e o f the c lim at e t he s light effect s ,

o f o n e s u m mer or wint er ha v ing b een tran s m itt ed and add ed t o

from gen era t io n to gen erat io n ; and aga in 2

I n m any o t her
”

,
“

an imals and plant s infl u enc es o f t em pe rat u re and en v iro n ment

m ay v ery p o s s ibly pro d u c e p er m an ent h ered itary v ariat io n s in a
s i m ilar m ann er and t ill aga i n 3
M any ar i t i f p lant
”

; s v e es o s ,

m ay a ls o b e d u e wh olly o r in part to t he s i m u l ta neou s v ar ia t io n

o f c o rres p o nd ing d et er m inant s in so m e part o f the s o m a and in

t he ger m p la s m o f t he repro d u ct iv e c ells and th es e v ariat io n s ,

m u s t b e h ered itary Temper a tu re a n d n u tr i t ion i n i ts w id es t

.
,

tb c wk ole b ody f tb c l t/ze s om a t i c c el ls a s

'

s ens e, a fl ec t o
p ant
1
well t/t e g er m c ells
” ‘

as .

In all th is it m u s t b e
n o t ed that Weis man n limit s act io n o f
th is k ind to s u ch ext ernal infl u enc es as a re ab le to pen etrat e the
o rgan is m and a ffect t h e ger m p la s m d ir ect ly ; wh eth er an in flu

enc e d oes th is is t o b e d et er m in ed i n ev ery ca s e by d i rect ex

e i m en t and is n o t t o b e a s s u m ed fr o m t h e eff ect o f the s a me

p r ,

infl u enc e u po n mere b ody d ev elo p m ent .

T h es e c i tatio n s fro m Weis m ann wh ile n o t es pec ia lly beari ng ,

u po n t he t o p ic o f u s e and d is u s e a re i ntr o d u c ed b eca u se his ,

po s itio n as the lead er in o ppo s it io n to t he th eo ry of t he tran s

m is s io n of mod ific a t io n s d u e to ext erna l infl u enc es is o ft en m is
u n d er s t oo d T h ey are intro d u c ed at th is p o int b eca u s e it is
.

c o nc ern ing u se and d is use that the mo s t v igo rou s d is c us sion s

ha v e arisen .

T h o s e believ ing in the tran s m itt ed effect s o f u se and d is u se

base th eir belief m ain ly u po n the meth o d o f pro o f by in s tanc e ,

and m o s t o f th em c it e in s tanc es that were far b ett er o mitt ed .

A s lo ng a s o ne s id e d epend s u po n s im p le en u m era t io n and ,

t he o th er m a in ly u po n ab s t ra ct r ea s o n ing we are n o t lik ely t o ,

g e t ah e ad A s a
. ll f o r m s o f a cq u i re d charact e r s a re d i s c u s s ed
togeth er it is pract ically im po s s ible to c it e referenc es d ea ling
,

exc lu s iv ely with u s e and d isu se ; b u t in o rd er that the s t u d en t

1 W im e sG m Pl m p 4 —
an n , er
4 6 I b id p 4 6
as , . 00 0 .
3 . . 0 .

3 I b id p 4 5 . . 0 . I t li mi 4 a cs a re n e.
 TR A N SM I SSI O N OF M OD I FI CATI O N S
4 7
0

may d o his th ink ing so m e o f the princ ipal referenc es relat

o wn ,

in g in part t o u s e and d is u se a re giv en in t he f oo tn o t e 1

.

I t is im po rtant that we kn o w t he tru th in th is m att er if pos

s ib le I f t he p erf ect io n o f d ev elo p m ent that c o m es o n ly with
.

u s e is to any ext ent tra n s m itt ed th en we m u s t t p ee d

p u o u r s ,

h o rses thro u gh a lo ng c ou rse o f t ra in ing and d ev elop th em fu lly

befo re we m ay h o pe that th ey will tran s m it m ax im u m s peed
-
.

I f th is th eo ry b e c o rr ect th en t he h eifers fro m a m at u re c ow

,

that has been lo ng in milk a n d m ad e rec o rd will b e capab le ,

o f d ev elo p ing i nt o b ett er m ilk er s than wo u ld b e p oss ib le if

t he d a m had n ev er m ad e extrem e rec o rd s o r than wo u ld b e ,

po ss ible with the earlier calv es fro m the sa me c o w (befo re the

extrem e r ec o rd s were m ad e) Man ifes t ly t he a ge o f t he b u ll .

will n o t c o u n t as he is i ncapab le o f d ev elo p ing th is part ic u lar

,

character A ll he can d o is to tra n s m it u nau gment ed and

.
,

u nchang ed t he h ered itary fac u lt ies o f m ilk pro d u ct io n exact ly a s

,

th ey d esc end ed to him I n meat pro d u ct io n o f c o u rse as in s peed

.
, ,

t he c as e wo u ld b e d ifferent as b o th s ex es m ay b e c o nc eiv ed as ,

capab le o f add ing to o r d etract ing fro m t he rac ial int en s ity
by reaso n o f th eir o wn d ev elo p ment o r lack o f d ev elo p ment .

T h is s u bj ect is n ow u nd er in v es t igat io n and wh ile t he p o int ,

wo u ld s eem to b e ea s y o f d et erm i nat io n yet it in v o lv es t he care ,

fu l s t u dy o f all t he pro geny o f m any ind iv id u a ls b o th b efo re and

aft er d ev elo p ment On th is point h owev er ev id enc e may b e
.
, ,

expect ed at n o v ery d is tant dat e .

Effect of dev el opment u pon t ran s mission of speed in horses .

In a rec ent series o f art ic les Ca s per L R ed field tak es t he ,

2
.

po si t io n that t he effect s o f s peed d ev elo pm ent a re tran sm i tt ed ,

and he c it es n u merou s in s tanc es calc u lat ed to s h ow that m ax i

m u m s peed is tran s m itt ed o n ly fro m s i res wit h lo ng an d
1
Ag i t t mi i
a ns W i m
ra n s G m Pl m p p 3 9 4
ss o n : I f
es f
an n , er as , . 2- 10 . n av o r o

t ran s m is s io n : R o m a n es , D a rwin an d A ft er D a r win , I I , 60 -

2 87C o p e, Ori gin o f
the Fittes t , p p 1 94 - 2 03 , 40 5- 4 2 1 , an d Pri ma r F ac t o rs y o f O r a n ic E v o u t io n ,
g l
—
.

pp 2 4 6 3 84 ; E im er, O rga n ic Ev o u t io n , p p
- 1 53 1 7 3,
- 2l05 2 2 1 N o n -
Pa rt is an
—
. . .

l yd
L o M o rga n , Hab it an d I n s tin c t, p p 2 80 3 2 2 ; V ern o n , V a ria t io n in A n im als
.

an d Pla n ts , pp 3 5 2 3 7 0
.
-
.

3 M r R ed fi eld s t eo ries
’
h a re b es t s et fo rt h in a s eri es o f a rt ic es en t it e l l d Ev o
2 5—
.

lu t io n o f t h e S et ter, in A m er i ca n Fi eld , L X I I , N o s
”
, a n d L X I I I , N 0 5 1 9 . 27 .
-
.

h y h
T e a re fu rt er s et fo rt in a s eries en tit led, h B reedin g t he T ro t ter, p u b lis e h d
in Tli e Hor sema n , X X V , N o s . 1 9-
34 .
4 08 T R A N SM I SS I ON
h o no ra ble ra c ing reco rd s The s t u d ies wo u ld b e mo re c o nc lu
.

s iv e if th ey inc lu d ed larger n u m b ers o f exa m p les and if th ese

—
,

were thro wn int o t o c la s s es

w o n e go tt en a ft er t he r ec o rd s
,

were m ad e t he o th er go tt en by the s a m e s i r es be
, fo r e th ei r
d ev elo p ment .

M r R ed field c o nc eiv es that t he s ire o r d am that is c o n s tan t ly

.

wo rk ed u p t o a s af e limit d ev elo p s th ereby a larger s t ock o f

what he c alls dyna m ic fo rc e and that tra n s m is s io n is in p ro
”

po rt io n to the ext ent o f th is fo rc e present at the t ime o f pro c re

at io n T h ere is n o need o f in v o lv ing t he s u bj ect with n ew t er m s
. .

What is in M r R ed field s mind is d o u bt less t he s a m e tho u ght

.
’

that lies at the bas is o f C o pe s th eo ry o f growth f o rc e wh ich is

’

o n e o f the s tro nges t o f what m ay b e ca lled t he dyna m ic th eo ries

o f ev olu t io n Ev erybo dy rec o gn izes a dyna m ic ba s is i n tra n s

.

mis sio n t hat wh ich is c o nn ect ed with t he int en s ity of t he

,
-

v ital pro c ess es M any fo rc es ca u se that int en s ity to v a ry and

.
,

t he im po rtant q u es t io n is wh eth er ex erc is e u s e extrem e d ev el , ,

o p m en t in the ind iv id u al is o n e o f th em M r R ed field s art ic les

'
. .
,

may b e read wit h pro fit wh eth er o r n o t a ll his c o nc lu sio n s will

s tand is an o th er matt er T he art ic les a re ch iefly u s ef u l fo r t he
.

large m a s s of fact s pres ent ed wh ich g ood as th ey m ay b e a re, , ,

no t e t su ffi c ient t o m ai nta in h is th eo ries o r t o an s wer the q u es

y
t io n that h o rs em en wo u ld lik e to ha v e sett led .

Cert ai n o u t s id e c o n s id erat io n s m u s t b e b o r n e in m ind in

s t u dy ing th is s u bj ect

I . T he b ett er t he s i re as t o s peed t he b ett er will b e his ,

o ppo rt u n ity t o get s peed fo r t he m o re n u m ero u s will b e his

,

o ff s p ring and t he bett er will b e t he c las s o f m ares o ff ered T he .

s a me p rinc ip le h o ld s tr u e a s to t he d am fo r o n ly a goo d o n e is ,

wo rth t he fee fo r a h igh c las s s ta llio n ; in o th er wo rd s t he s i res

-
,

and da m s with rec o rd s hav e bett er o ppo rt u n it ies to prod u c e

than d o th ose eq u ally goo d b u t with o u t rec o rd s Beca u se of .

th is fact the get of on e an imal m u s t b e c o m pared n ot with tlza t ,

b u t wi l l: lzzs wn of
’

f
o a no t/zer , a a la ter o r ea r l i er da te .

2 . S peak i ng gen erally

bes t h o r s es lat er in life
, t he get of t he ,

aft er th ey a re kno wn t o b e v alu able will b e bett er train ed and ,

better d ev elo ped than t he get o f t he sa m e an imals earlier in

lif e and in the h and s o f mo re o rd inary h o r s emen .
 lo TR A N SM I SS I O N
4

lim b s ; t he wi ngs hav e go n e fro m t he apt eryx and appear t o b e

go in g fro m the o s trich ; ey es o f ca v e in sect s and fi shes a re in
m any ca s es im perf ect o r r u d im entary ; h o rs es catt le s h eep , , ,

and h og s hav e los t t oes that belo nged to their anc es t o rs and ,

part s gen erally wh ich are f u nct io n les s are ev id ent ly d isappeari ng .

How n o w a re part s los t wh en o nc e th ey hav e b ec o me u seles s ?

, , ,

Econ omy of Nat u re n ot t he reason for l os s of part s . I t is

s o me t im es s a id that a part n o lo nger u s ed is rem o v ed by N at u re
in t he int eres t o f ec o n o my T h is is bad s c ienc e N at u re is n o t
. .

ec o n o m ic a l S he no t o n ly s u ppo rt s m any expen s iv e and u s eles s

.

part s s u ch as tremend o u s h o rn s and t u s k s b u t she o ften p ro

, ,

d u c es n ec es s a ry prod u ct s in wa nt o n pro f u s io n s u ch as po llen

and fat Oth er and d eeper cau ses are at wo rk — ca u ses mo re
,

.
,

rat io na lly c n ect ed with t e fact o f l fe

o n h s i — than any su ch ,

anthro po m o rph ic rea so n as ec o no my Wh a t ev er may b e tru e .

as t o ec o n o m y o f gro wth i t is a fact n o t a pri nc i p le ; a res u l t

, , ,

n ot a ca u se .

How part s di sa ppear W e a re r easo nably int elligent u po n a

.

part of t he proc es s of d egen eracy and d is appearanc e at leas t in ,

t he mo re act iv e p o rt io n s o f t he b o dy A s lo ng a s a part is in .

u se it s c o n s tant m o v em ent s incr ea se t h e flo w o f b loo d to that

,

part and it enj oy s the extrem e d ev elo p ment that c omes o n ly with
m ax im u m n o u r is h m ent and u n if o r m h ealthy ex erc ise When .
,

h o wev er the part is n o lo nger u s ed the flo w o f b lo od is lessen ed

, ,

a n d it s u ff er s fr o m les s en ed n o u r is h m ent I n t hi s way t he fi rs t .

s te s o f d eg en erat io n a re ea s ily acc ou nt ed fo r

p .

I f t he p a rt ha s b een u s ef u l h eret o fo re it has o f c o u rs e been ,

s u s ta in ed by s elect io n B ei ng n o lo nger u s ef u l th is infl u enc e is

.
,

withdra wn and breed ing is h enc ef o rth t o tally with o u t ref erenc e
,

t o th is part ic u lar part ; that is t o s ay th ere is ab s o lu t e c ess a ,

t io n of select io n o r pa n mix ia a s to th is part U nd er th is c o n

,
l
,
.

d it io n the av erage parentage wo u ld b e lower t han h eret ofo re th u s ,

acc ou nt ing fo r a s t ill f a r t/t er s t ep in the d o wn ward pro c es s .

To all th is is o ft en add ed t he ad v erse effect o f s elect io n ,

wh en fo r any rea s o n that infl u enc e is t u rn ed ag a i n s t the part .

T h is “
rev ers al o f s elect ion o f c o u rse c o mes o nly wh en a pa rt
o nc e u s ef u l ha s b ec ome no t m er ely u s eles s b u t d et ri menta l In .

1 R o m a n es , D a rwi n an d A fter D a rwin , I I , 97 —

1 00 , 2 91-
3 06 .
 TR A N SM I SSI ON O F MOD I FI C AT I ON S I !

su ch nt th ere en s u es a thi rd s tage of d egen eracy Obv io u s ly

ev e .

t he f u ll effect s o f s elect io n wi ll d ep end v ery m u ch u po n t he

chara ct er of t he part and it s c o nnect io n with the v ital interes t s
o f t h e s pec ies .

T he i nfl u enc es j u s t m ent io n ed will s u flic ien t ly acc o u nt fo r ex ‘

treme d egeneracy o f a o nc e act iv e part b u t th ey will n o t acc o u nt ,

fo r ab s o lu t e d isappea ra nc e A n y ch a ract er u nd er s u ch c o nd i t io n s
.

wo u ld d ecreas e to a lo w m in im u m wh ere it s presenc e bec o m es

in s ign i ficant and th ere i t wo u ld rem a in b u t i t wo u ld no t ab s o
, ,

lu t ely d is app ear exc ept throu gh s o m e o th er agency T hat .

characters ev en th o se wh ich were o nc e im po rtant d o ent irely

, ,

d isa ppear lea v i ng n o t ev en r u d imenta ry pa rt s is ev id enc ed by

, ,

t he d is appearanc e o f the leg s o f s nak es b u t that t he lat er s tages ,

a re extremely S lo w is s h o wn by t he ru d imentary leg b o n es o f

t he pyth o n and t he wha le .

Degen er at i on of eyes in cav e -

dwelli n g an d deep -
sea s pecies .

I t is a well kn own fact that t he eyes o f ca v e fi s h es and in sect s

-

o ft en e xh ib i t a ll grad es o f d egen e ra t io n fro m n ear t he n o r m a l ,

d o wn to the m eres t r u d iment s Being enti rely u seless u nd er .

t he c o nd it io n s o f life s elect io n is s u s pend ed and N at u r e is

, ,

ha v i ng her way wit h t he rem nant s o f what was onc e a h igh ly

d ev eloped o rgan .

D eep s ea fi s h es a re eith er in the s ame c o nd it io n o r els e a re

-

s u pp lied wi th en o rm o us ey es o f a k ind ev id ent ly fitt ed t o per

ce iv e light rath er than t o m ak e d is t inct and c lear c u t im ages -

.

T he grad u a l failu re o f part s lik e t he t o es that ha v e go n e fro m

t he h o r s e o r t he apparent ly d is app ea ring v erm if o rm appe nd ix
, ,

m ight b e attrib u t ed t o s o me fa ilu re o r im p erfect io n in the germ

b u t the in s tanc e o f d is appearing lim b s is t o o c learly c o nn ect ed
wi th d is u s e and that o f d is app eari ng ey es wi th lack o f what
‘

m ay b e ca lled es s ent ial c o nd it io n s o f d ev elo p m ent t o b e exp la ined ,

wh olly o n t he gro u nd o f i m perfect io n fro m with in as t he f u n da

mental ca u s e o f d egen eracy ‘
.

T he fina l d is appearanc e of a u s eless part is c erta in ly d u e t o

s o m e fact o th er than t h e withdra wa l o r ev en t he rev er s a l o f

s elect io n T h ere m u s t b e m o rph o lo gical u n it s of s o m e k ind

.

1L igh t i id
s ev t ly ti l t th
en igi f th
es s en it i
a p t wo l l t h y
e or n o e s en s ve s o e ca e e e,

es p i
ec al ly i t h
n f m ti f p igm t
e or a on o en .
 12 T R A N SM ISS I O N
4

res id ent with in the ger m fro m wh ich th ey slo wly d isa ppear
,

wh en n o lo nger fa v o red by t he c o nd it io n s o f life and n o lo nger

s u s ta in ed by s elect io n T h es e v ita l u n it s if ev er d is c ov ered will
.
, ,

b e f ou nd t o b e c lo s ely c o nn ected with t he or igin o f charact ers

as well as wi th th ei r pres erv atio n .

W e s hall s ee lat er that w/zen a cita ra cter is u n d erg oi ng rigor

,

our s elec t i on n e w and l g

u/t er v a l u es tha n ev er be
f o r e a r e c on s ta n t l
y
a pp ea r i ng . May r
n otev er s e b e a ls o tr u e
t he ,
— na m ely that a
,

chara cter o n the d ec line may present it s s u cc ess iv e d ecreasing

v a lu es b eca u s e o f i nfl u enc es ent irely int ernal

That d is app earanc e of part s i s n o t d u e ent irely t o d is u s e is

s h o wn by t he fact that the proc ess c o nt in u es lo ng aft er t he fact

o f d is u s e c o u ld ha v e t he s light es t infl u enc e Where a ru d i .

m entary t ibia f u rth er d egen erat es t o a ru d im entary pelv ic b o ne ,

t he q u es t io n o f d is u s e is c erta in ly no t in v o lv ed ; n eith er is u s e
o r d is u se i n v olv ed in breed ing fo r h igh o r lo w o il in c o rn fo r ,

exa m p le . M an if es t ly so m e b iolo gic al princ ip le is in v o lv ed that

has n o t yet b een d is c o v ered and id ent ifi ed and as it is ev id ent ly , ,

a p rinc iple fu nda m ental to tran s m iss io n and v ariat io n it s isola ,

t io n is exc eed ingly impo rtant .

Charact ers n ot depen den t u pon adaptati on G en erally s peak .

in g th ere is a c lo s e c o rrelat io n bet ween t he d ev elo p m ent of a

,

cha ract er and it s u s ef u lness to the ind iv id u al and t he s pec ies 1

.

T h is fact has gi v en ris e to t he i m pres s io n that all charact ers

a re d ep end ent u po n t eleo lo g ica l pr inc ip les fo r th eir ex is t enc e .

N o great er erro r c o u ld b e m ad e I t is tr u e that in nat u r e s elec

.

t io n o perat es m os t ly alo ng u t ilitarian lin es and in th is way aft er ,

a t ime it bri ngs mo s t cha ract ers int o line with t he great es t
s erv ic e a n d t he c lo s es t adaptat io n ; b u t s elect io n m a o pera t e in
y
any d irect io n ev en to the d isad v antage o f a s pec ies I n th is
, .

case h owev er t he res po n se is to s elect io n n o t t o u t ility

, , ,
.

N ei th er is d ev elo p m ent a lo ng u t ilitarian lin es n ec es s arily

tr ue o f th o se charact ers that lie o u t s id e the fi eld of select io n
o r o f th o s e u p o n wh ich it o p e rat es t oo rarely t o i mpr ess i t s elf .

F o r exa mp le the in s t inct t o fly t o ward a s o u rc e o f light

,

wo u ld ext erm inat e c erta in s p ec ies if nak ed fi re were m o re gen er

ally enc o u nt ered T hat the t es t ic le in m am mals s h ould hav e
.

K w 1 m g b i l gi t
no n a t l
on l gyo o s s as e eo o .
4 14 T R A N SM I SSI ON

m eet s i t by as s u m ing that the charact er s that d is t ingu is h t he h igh er

rac es were in s o m e way im pr es s ed u po n t he o ri gi na l pro t o p las m

fro m with o u t ,
w/zile t/ze r em ote a nc es tor s were y et i n tb e s in
g le

c elled s tag e . T h u s d o t he m o s t rad ica l

ct io n is t s bec o m e s ele

L a m arck ian s o f t he p u res t k ind when d riv en far en o u gh back 1

.

B u t is th is v io lent a s s u m pt io n n ec es s a ry ? Life in t he s ingle

c elled s tage is n o t f u n da men ta lly d ifferen t fro m life in t he c o lo ny
f o rm A c ell is a c ell in eith er cas e a n d it s act iv ity what it —
can d o — is d epend ent part ly u po n it s anc es try and part ly u po n
.
,

t he c o nd it io n s o f lif e wh ich a re i t s o ppo rt u n it ies

,
T o b e s u re .
,

t he s ingle c ell is m o re d epend ent u p o n t he extern al wo r ld and ,

react s mo re c o m plet ely to t em pera t u re and o th er ext ernal fo rc es ,

than d oes t he la rger c o lo ny o f h igh ly s pec ialized u n it s ( c ells) ,

b u t th is is a d ifferenc e i n d egree rath er than in k ind .

I n the la s t analy s is we a re dri v en to t he c o nc lu s io n eith er

that all characters were creat ed and im plant ed in the o rigi nal
pro to plas m ,
so t hat liv ing m att er ev en in it s s i m p les t f o r m
, ,

l
h as a l t e ph o t en t a l t es o f t e h gh es t f o r m o f life
i i i h i — o r els e ,

that the pec u liar ch em ical c o m p o u nd s that c o n s t it u t e liv ing mat

ter a re ab le n o t o n ly t o ent er int o d efin it e relat io n s with t he
wo rld at large b u t als o p erh a p s t o effect fro m t ime t o t ime
, , , , ,

n ew c o m b inat io n s a m o ng th em s elv es th u s a c q u i r ing n ew o r

,

great ly m od ifi ed charact ers s o met im es c o nfo rm ing nat u rally t o

,

s u rro u nd ing c o nd it io n s so m et im es n o t
,
T o th is latt er v iew t he .

wr it er s tro ng ly inc lin es .

A ch em ic a l elem ent a s iro n a cq u i res n o n ew charact ers

, , ,

th o u gh it behav es d ifferent ly u nd er d ifferent c irc u m s tanc es .

S u lph u r is ext rem ely s en s i t iv e t o s u rro u nd ing c o nd it io n s and ,

m any o f t he o rgan ic c o m p o u nd s d epend alm o s t ent irely for th eir

pro pert ies u po n the c o nd it io n s u nd er wh ich th eir p ec u liar c o m
b in at io n s were eff ect ed .

B u t wh en lif e ent ers t he fi eld all t he c o m p licat io n s a re in fi

nit ely m u lt ip lied and wh en we a re d riv en t o the las t d itch all
,

m u s t agree t h at the charact er s po s s es sed by liv ing m att er a re

t o a large ext ent an d in s o m e way an expres s io n o f t he c o nd i

t io n s o f life Ho w th es e c o nd it io n s impres s th em s elv es n ot o n ly

.

u po n t he in d iv id u al b u t a lso u p o n t he rac e is in so me c ases at ,

1 W eis ma n n , G erm Plas m , pp 4 1 5- 4 1 6 . .

 TR A NSM I SSI ON O F M OD I FI C A T I ONS
4 5 1

leas t ( t em pera t u r e f oo d po iso n s a nd m any

,
c h em i ca ls, n o t ,

po iso n s) bo t h ev id ent and eas y to u nd ers tand ; in o th ers it is

,

o b s c u r e a n d u nc erta i n t o t he las t d egree .

Degen eracy an d ori gi n con t rast ed Of o n e th ing we a re c er .

tain : cha ract er s a re d isappearing bef o re o u r v ery ey es and ,

wh ole ra c es a re bec om ing ext inct What d oes t h is m ean I s the .

wo rld growing po o rer in po s s ib ilit ies I s s pec ia lizat io n rea lized

o n ly at t he expen s e o f less en ed adaptab ili ty t o n ew c o nd i t io n s

lat er o n ? I f a part o r a charact er n o w u s eles s d egen era t es and

d is appears will it ev er c o me back o r will a n ew o n e ar is e to tak e ,

i ts p lac e if n ec es s ity fo r i t s pres enc e S h o u ld ret u rn ?

T h es e a re large q u es t io n s q u es t io n s that we cann o t an swer ,

b u t that we m u s t th ink ab o u t and tak e int o c o n s id erat io n in the

s t u dyi ng and an s wer ing o f ea s ier and s m a ller q u es t io n s .

I n t he m ean t im e we will rem em b er that so les flo u n d ers and , ,

t he fla t fi s hes gen era lly are d ev elo p ing a n ew s ty le o f liv ing and 1
,

that their ey es are tak ing a n ew po s it io n wit h ref erenc e to t he

o th er b o dy part s We will n o t f o rget that all an im als that liv e
.

in ( u nd er) the wat er if o f m u ch s ize a re o f o n e gen eral s hape

, , ,

t he s hape o f lea s t res is tanc e t o wat er T hat th is is in d epen d .

en t o f s elect io n is s h o wn i n t he h is t o ry o f t he wha le and o f the

few land m am mals that t o o k t o t he wat er and wh o s e t ran s fo rm a

t io n m u s t hav e been c o m parat iv ely rec ent .

We will remem ber that wh ile m os t n ew charact ers a re“

,

b u t n ew c o m b inat io n s and d ifferent adj u s t m ent s o f o ld o n es ,

th ere is aft er all progres s iv e d ev elo p m ent s h owing the inf u s io n

, ,

o f s o m eth ing p ract ica lly n ew and d z f e r en t H igh e r li f e d i ff e r s .

fro m lo wer in k ind as well as in d egree That it s prings from .

t he lo wer is c erta in and that s o meth ing has been add ed in t he,

proc es s is n o less c e rta in .

T he wo rld is f u ll o f lo wly f o r m s o f lif e T he s p ec ies o f s ingle .

c elled o rgan is m s that a re k nown pro bably far o u tn u mber ex is t ing

1 T h es e fis h es , of w ich h th ere a re a n u mb er o f s p ec ies , a re s y mm et ric a l , or

n ea rl y so , in t h e em b r o a n d y l it t l tim ft w d th t t fi t th y wim
fo r a e e a er ar , so a a rs e s

li k e an y o t h er fis h Th wi m m i g b l dd
es i d f ti
n h w a d h tly th y
er s e ec v e, o ev e r, a n s or e

ll y l ft id d w — t h gh m
.

t i ru to o ne Si d e an d li t h b tt m g
e on e o o , en era e s e o n, ou so e

d d l
in iv i u a s a re rev ers e d I t h i p i t i t h l ft ( w l w ) y t l p w d
. n s os on e e no o er e e ra v e s u ar

t w d th
o ar e ot h er s id t il t h t w
e, u n y li id b y id
e th igh t
o e es w th e s e s e on e r , no e

upp i d er, s e .
 16 TR A N SM I SSI ON
4

s pec ies h igh ly d ev elo ped o rgani s m s I s th is the ra w m ate

of .

ria l o u t o f wh ich n ew s pec ies s hall b e ev o lv ed lat er o n t o t a k e

t he p lac e o f what is n o w s o rap id ly bec o m ing ext inct ? I t is a
m o v ing pan o ra m a and we kn o w n eith er t he b egin ni ng n o r t he
,

en d . I t is all a part o f a v as t p lan on wh ich t he u n iv erse is

fo u nd ed by t he C reat o r We cann o t d ou bt that t he mo s t c o m
.

p lex element o f it all is lif e n o r can we d o u bt that t he mo s t

,

charact eris t ic pro perty of liv ing matt er is its p rog ress i v e d ev el
o p m en t b ringing to light n ew act iv it ies a n d es tablis hing n ew
,

relat io n s with the wo rld o u t sid e Wheth er th is pro perty is wholly

.

i nt erna l and dyn a m ic o r d u e at leas t t o s o m e ext ent t o ou t s id e

, , ,

i nfl u enc es th is is t he ch ief m y s t ery

,
.

SECT I O N I X V A R I ATI O NS D U E TO C A U S ES N OT A F FECT

I N G T H E G ER M A R E N OT T R A N S M ITT EI)

T h is m u ch is a . lo gical
c o nc lu s io n T h is m u ch is f u nda mental
.

and ax io mat ic The ger m inal matt er is the o nly material that
.

pa s s es o v er fro m generat io n to generat io n T he fert ilized germ .

is the o nly h eri tag e and i t is t he o n ly a v en u e o f tran s m iss io n

, .

What ev er fac u lt ies o r end owm ent s the ind iv id u al may p o s sess ,

he can tran s mi t o n ly th os e wh ich can fin d lo dg m ent and repre

s en t a t io n in t he ger m and wh eth er t he cau ses of germ inal
,

changes are internal or ext ern al o r b o th n o v ariat io n can b e , ,

tran s m itt ed u nl es s it a ffect s the germ o r u nl es s the ca u se that

ind u c ed t he v aria t io n a ls o a ffect ed t he germ .

V ariat ion s du e t o ca u ses in tern al t o t he b ody b u t ex ternal t o

t he ger m are n ot t r an smit ted ntirely po s s ible that c ertai n

. I t is e

v ariat io n s eith er o f s tru ct u re o r o f f u nct io n m a ar i s e f ro m

, y ,

irreg u larit ies in c ell d iv is io n d u ri ng d ev elo p ment d u e t o lo cal

rath er tha n t o ger m ina l ca u s es Fo r exa m p le it has already
.
,

been no t ed that c ells may at any t ime d iv id e int o u n equ al po r

t io n s o r int o three in s t ead o f t wo part s g iv ing rise to abno rm al
, ,

if n o t to path o lo gica l c o nd it io n s .

A lmo s t all po rt io n s o f the bo dy a re s u bj ect t o t hose o v er

g ro w th s ca lled t u m o r s and wh ile it is n o t
, y e t d efin it ely kn o w n
wh eth er the ca u ses o f abno r ma l gro wt h a re ext ern al o r int erna l
t o t he o rgan is m t hey c e rtai n ly are n o t locat ed in t he germ
, .
4 1 8 T R A N SM I SSI ON

q ues t io n wh eth er extrem e d ev elo p ment t end s to a ffect the germ

and t o bec o me i nh erit ed is a q u es t io n beset with m any d ifli c u l
t ies T he great es t o b s tac le to th is s t u dy is t he ev er pres ent
.
-

fact of s elect io n wh ich rap id ly b rings abo u t a c lo s e c o rres p o nd

,

enc e b et ween o rgan is m s and th eir s u rr o u nd ing s N o reliab le .

c o nc lu s io n can b e d ra wn u nt il th is infl uenc e is acc o u nt ed fo r

o r elim i n at ed .

I n f u rth er p u rs u anc e o f th is s t u dy we n o w pas s t o m o re d ef

i n it e d is cri minat io n s a s to type and t o m o re exact and crit ica l
d is t inct io n s c o nc ern ing v ariab ility as expres sed n o t in ind iv id ua ls
b u t in n u m bers s u ffi c ient ly large t o b e f a irly i nd icat iv e o f t he ra c e .

A D DITI ON A L REF EREN CES

A N E X A M I N A T I ON O F W EI S MA NN IS M . By G .
J . R om a n es . 1 v ol .

D A RWI N AND A F T ER D A R W I N . By G .
J . R o m a n es . 2 v o ls .

D E V E L OP M E N T A ND E V O L U T I ON . By J . M . B ald win . S c ien c e, XV I,

8 1 9- 8 2 1 .

E N V I R ON M E N T AND I TS E F F ECT ON T HE T R A N S M I T T I N G PO W E R O F
SE EDS . By W W . . T ra c y
S c ien c e, X I X , 7 3 8- 7 4 0
. .

E X PE R I M E N T A L Z OO L O G Y B y T H M o rga n C ap ters I V h and V , pp

—
. . . . .

43 6 1 .

FO U N D A T I ON S OF Z OO L OG Y . By W . K . B roo k s . 1 v ol .

HE R E D ITY AND I NS T I N C T . By J . M . B ald win . S c ien c e, I I I , 4 3 9- 4 4 1 ,

55 8—5 59
I N HE R I T A N C E O F A C Q I R E D C HA R A C T E R S U
B y E D C o pe S c ien c e, V , . . . .

J h
6 3 3 6 3 4 ; b y o n M c F a rla n d , I b id , 93 5 94 5
-
.
-
.

I N H E R I T A N C E O F A C Q I R E D C HA R A C T E R S ( E U
xa m les
p ) B y F R H er . . . .

ri c k . S c ien c e, V I I , 2 80 .

I N HE R IT A N C E O F A C Q U IR ED C HA R A C T ER S . By D . E . H u tc h en s
N a t u re, L X X I , 8 3 .

N AT URE OF CAN C ER S AN D A B N O R M A L G R O WT Hs , AND T R A N S M I SS I B I L I T Y

OF S A ME . By E . B . B as fo rd h . Pro c eed in gs of the R o al y S o c iety,
Lo n d o n , L X X I I I , 66- 6 7 .

R I G HT A N D L E FT E Y E D N E S S -
. By G . M G o u ld
. .S c ien c e, X I X , 5 9 1 - 5 94 .

T H E H E R E D I T Y O F A C Q I R E D C HA R A C T E R S U . B o s to n M ed ic al a n d S u rgi
c al o u J
rn a l ,
C X X X V I I , 4 2 —
7 42 8 .

U S E I N HE R I T A N C E
- . D irec tio n of Ha i r i n M a n an d A n im als and its A ppli
c a tio n to D a rwin is m . By W . K id d . S c ien c e, X V , 1 42- 1 4 3 ; als o in

S c ienc e, X X , 4 0 1 —4 7
0 .
 CH A PTER X II

TYPE AND V AR I A B ILITY 1

E n o u gh h as been sho wn in earlier c hapt ers t o c o n v inc e the

s t u d ent t h at v a riab ility is an in ev i tab le acc o m pan im ent o f bo th

reprod u ct io n and d ev elo p m ent and th erefo re that v ariat io n is to ,

b e expect ed a m o ng liv ing being s ev ery wh ere .

B ef o re anyth ing lik e a c o m prehen s iv e id ea o f tran s m is s io n

can b e d ev elo ped th erefo re it is n ec es s ary to s t u dy n o t ind i , ,

v id u a ls b u t gro u p s and t o es tab lis h d efi n it e c o nc ept io n s a s to

, ,

type and v ariab ility I n th is as in a ny o th er crit ical s t u dy o f a

.
,

rac e we proc eed charact er by character and a re caref u l to

, ,

inc lu d e en o u gh i nd iv id u als to b e fai rly repres entat iv e o f t he rac e

as a wh o le
2
.

A fa r m er p lant s an ea r o f c o rn say t en i nch es in length .

W hat he get s is n o t a c ro p o f ears all t en inch es lo ng b u t a ,

gro u p o f c a rs ranging in length all t he way fro m perhap s three

o r f o u r i nch es u p t o elev en o r t welv e o r ev en a litt le m o r e T he ,
.

s a me pr inc ip le will h o ld if t he ea r that is p lant ed is n in e inc hes

lo n g in s t ead o f t en exc ept that the d is tr ib u t io n will b e d ifferent

, ,

len gth s r u nn ing in gen eral S light ly lo wer ; that is to s ay t he

, , ,

length o f ea r o f the o ff s pr ing is n o t t he s a m e as t hat o f t he

paren t b u t it c o n s t it u t es a d is trib u t io n ext end in g b o th abo v e
,
“

and belo w that length S o far as is kn o wn th is pri nc ip le o f tra n s

.
,

m is s io n h o ld s tr u e in all rac es a n d fo r a ll charact ers S tat ed in .

m o re gen era l t er ms we m ay s ay tlza t t/ze ofi sp r i ng a s a w/t ole is

'

n ot t/ze sa m e a s t/te i m m ed ia te p a r en ts , b u t tha t i t co ns t i tu tes a

d is tr i b u t i o n ex ten d i ng f r o m n ea r t/ze lower to app r ox i m a tely t/ze

upp er l i m i ts o f t/ze ra ce . T/zi s gs ug es ts a t on c e tlze id ea o

f typ e

a nd tha t d ev ia t ion f r om type w/ziclz we ca ll v a r ia b i li ty .

1 S ee B u ll eti n N o . 1 1 9 A g
,ri c u t u ra l E x l
p eri m en t S t a t io n , U n iv ers i t y of I ll i n o is ,

b y t he au t h or of h
t is t ex t .

3 Ha v in g di s c o v ere d t h typ t
e e as o s ev era l im p o rt a n t c h a ra c t ers , it wo u ld th en

b e p o s s ib l l
e t o s e ec t a t yp i l i d i id
ca n v u a l.

4 19
 0 T R A NSM I SS I O N
4 2

SECTI ON I T PE Y
What ou r c n ow is typ ears will
on c ept io n of e ? I f ten inch -

n o t pro d u c e ten i nch ear s b u t s o m eth ing els e-

and n o t o n ly , ,

s o m eth ing els e b u t a c o n s id erab le v ar iety o f length s ; and if

what we get ext end s bo th abo v e and b elo w t he parent th en we ,

arriv e at o nc e at a d o u ble c o nc ept io n as to type ; that is to say ,

t he ty pe o f t he o ff s pr i n g is n o t t he s a m e a s that o f the parent .

T he ty pe o f the parent is v ery d efin ite repres ent ing an id ea l ; ,

b u t if t he o ff s pri ng is d is t rib u t ed b o th ab o v e and belo w t hat

id ea l so me being bett er and s o m e n o t so goo d th en a c lo s e
, ,

analy s is of t he real charact er o f that o ffs pring bec o mes n ec es s ary

in o rd er t o m ak e any j u s t c o m par iso n b et ween the t wo o r t o ,

arriv e at any ad eq u at e c o nc ept io n o f type in a m ix ed po p ulat io n ,

ev en in o n e ar is i ng fr o m a s elect ed anc es try .

A c o ncret e ca se will ser v e b es t t o illu s trat e t he p rinc ip le

i n v o lv ed I n t he y ear 1 906 s o m e Lea rn ing c o rn was ra is ed o n
.

goo d gro u nd fro m seed ears exact ly ten i nch es in length A .

rand o m s a m p le o f th is cro p c o n s is t ing o f 3 2 7 ears ga v e t he

1
, ,

fo llo wing d is t rib u t io n as t o length

On e ea r was inch es lo ng o n e was i n ch es t wo were , ,

i nch es three were ,

inch es n i ne were inch es eight , ,

were inch es t welv e were inch es n in et een were

, ,

inch es th irty t wo were

,
inch es f o rty were
-
inch es , ,

S ixty s ev en -
were inch es s ixty three were inch es ,
-
,

th irty eight were 1 0 inch es t wenty o n e were

- inch es eigh t ,
-
,

were inch es t wo were inch es and o n e was

, ,

inch es lo ng 2
.

1 By a ra n d o m s am p e l is m ea n t a s u ffic ien t p o rt io n o f t h e w h o le, ta ke n so

mu c h at ra n d o m as to fai r ly r ep res en t t h e en t ire c ro p , o r

“
p o p u la t io n ,
”
as t he
t ec h n ica l ph
go es ras e .

3 M eas u rem en ts m ig t b e t a h k en at q u a rt er in c h es wi t h a s eem in g l y h igh er

d egree o f ac c u rac y , b ut rep ea te d t ria s l h s o w h

t at the s a me fin al r es u t s l f ll w
o o

wh eth er m eas u remen t s a re t a k en a t t h e q a rt er in c

u h o r at h lf i h T h m i
the a nc . e a n

p o in t is t h a t t h e n u mb ers s h ll a be s u ffi c ien t a n d th at th mp l h ll b p
e sa e s a e re re

s en ta t iv e .
Ju dgm en t m u s t di t c a t e a s t o t he a c c u ra c y f th mpl b t th m
o e sa e, u e nu

d
b er ep en s d l i b il it y d i
u po n the d egree of re a es re d T h i m t t wi l l b f ll y
. s a er e u

d d d th b j t f p b b l
isc u ss e u n er b t
e su ec o ro a e erro r, u ex p i h w
e r en c e st h t i t doi s a n s u es
wi t h ll t
c o rn lt b h d wit h f m
exc e en res u s c an e a ro 2 00 t 3 00 o d e y f i
ea rs , an
a v r r

res u lt m y gs l ly b h d with h lf th t mb
a en era e a a a nu er.
 T R A N SM I SS I ON
4 2 2

therefo re ,
we kno w at o nc e w/ta t is Me n a tu r a l type 1
o f til e rac e

or v a r i ety s o far as t he chara cter in q u es t io n is c o n c erned and ,

wlzen this is d eter m i n ed f o r a n u m ber of i mp or ta n t c/za ra c ter s we

s /za ll lza v e a g ood k n o wledg e of fire r a c i a l typ e a s a wlzole T h u s .

we m ight o bt ai n t he m o d e fo r c i rc u m ferenc e n u m ber o f ro ws , ,

weight c o lo r o f gra in per c ent o f c o b o r any o th er d esired

o f ea r , , ,

character and ha v ing d o n e s o a typ ica l ea r of t/zis v a r iety c ou ld

,

be d e fi n i te l
y d e s c r i b e d W e th u s . arr iv e at an acc u ra t e id ea o f

ty pe in a gen eral po p u lat io n and o f it s d efi nit e m ea s u rement ,

.

The empirical an d t h e t h eoreti cal mode I t is ev id ent by ih .

s pec t io n o f t he freq u ency tab le that if m ea s u rem ent s had b een

t ak en at t he q u art er inch o r so me les s f ract io n t he h igh es t fre , ,

q u en ey wo u ld hav e fallen n o t at t he n in e inch po int b u t S light ly -

abo v e it fo r the n ext freq u ency abo v e ( 6 3 ) is greater than t he

,

n ext o n e belo w that is t o say t he mo d e is to so m e S light ,

ext ent d epend ent u po n t he s ch em e o f m eas u rement s ad o pt ed .

A n y s ch eme expres sed i n n u m b er s eith er wh o le o r fract io nal , ,

is o f c o u rs e by nat u re d is c o nt in u o u s and t he m od e aris ing fro m ,

s u ch a s ch em e is at b es t o n ly an appr o x i mat io n I t is th erefo re .

called t he em p ir ical mo d e I f all po s s ib le v a lu es were repre .

s ent ed h o wev er as is d o ne wh enev er t he th eo ret ical c u rv e is

, ,

p lo tt ed c o rres po nd ing to the freq u ency d is trib u t io n s u ch a c o n ,

t in u o u s c u rv e will find t he t ru e o r a s it is c a lled t he th eo ret ical

, ,

m o d e I t is n ec es s a ry t o rec o gn ize th is d is t inct io n a ltho u gh

.
,

in pract ica l breed ing o perat io n s t he em p ir ica l m o de a ris ing

fro m c o n v en ien t mea s u rem ent s is s u ffic ient ly acc u ra t e .

Th e coemcien t of mode or modal coemcien t ,

2
I t is n o t en o u gh .

s im p ly t o d et er m in e wh ich v a lu e ha s t he h igh es t freq u ency ,

ev en th o u gh th is g iv es u s the type ; we d es i re t o kn o w a ls o

what p r opor ti on o f a ll t he ind iv id u a ls t end s t o d ro p int o t he type

1
It i id t t h t t h f q
s ev en a y d i t ib t i
e re d th
u en c f ths r ty p uf on an , ere o re, e e o an

a d lt p p l t i
u o u a i on m t hi g d iff
s so e t f n m h t wh i h w b
e ren i t th
ro t a c as o rn n o e ra c e.

W h t th t m y h
a a a b av ew een k w M y i di id l d id t
e ca n n e v e r no i . an n v ua s no s u rv v e,

an d h d t e l pm t f ll w i fl
ev e o en o a d b y i m t Th fi l
as n u en c e lt en v ro n en . e na res u as

re p t
res en ed i d lt i
n a d i uid l i ll w
n v ua s id s a e c an c o n s er .

T h w it
9 e rh er th i
a s n e v e r s een
p i d I t i f l it tl
s ex res s o n u s e
q i . s o e c o n se u en c e n

g l
en e ra l t i
ev o u b t i
on , f m u h si g iofi i thucmm sl gy wh th
n c an c e n re a to o , ere e
b d d i t k w wh t p p t i
ree er es res o no f hi
a im l
ro
p l
or ton f om t s an a s or an s con or o

typ e I h
. th
a v e, f t k
ere o re,th l ib t y f i g it h
a en e er th o fi i tus n , as er e, e coe c en

o f m d o m d l
e, o r fli i t
o a co e c en .
”
 TY PE A N D V A R I AB I LI TY 4 3
2

of t he rac e Th is is ea s ily d et er m in ed in the fo rm o f a ra t e

.

p e r c e nt by d i v id ing t he h igh es t freq u ency by t he t o tal n u m b er

o f v ar ia t es T he h igh es t freq u ency in th is cas e is 6 7 wh ich is

1
.
,

ov er 2 0 per c ent o f t he t o ta l n u m b er o f ear s mea s u red .

T h is we ca ll it s m o da l c o effic ient beca u se it i nd icat es t he p er

c entage o f t he t o tal po p u lat io n that c o nfo rm s to type in res pect
t o th is cha ract er The m od al c o effi c ient o f s o me o th er v ariety
.

m ight b e q u it e d ifferent s ho wing that a h igh er prop o rt io n o f

,

o n e v a riety m a c o nf o r m t o type than o f an o th er ; o r what is

y ,

t he s a m e th i ng that o n e v ariety m ay b e m o re c o n s tant and

,

tru er to type Tb e m od a l c oefi c ien t tlzer ef or e is a n in d ex of

.
, ,

r ela t i v e c on orm i ty to t
f ype a v a lu ab le b i
,
t o f kn o w ledg e fo r

p u rpo ses of s elect io n .

M oda l coefli cien t par tl y depen den t u pon t h e sc h eme of measu re

men t s adopt ed I f th es e mea s u rem ent s had been tak en to the
.

q u art er inch there wo u ld ha v e been t wic e as m any freq u enc ies

and ea ch wo u ld hav e been repres ent ed by c o rres po nd ing ly fewer
ears . T he h igh es t freq u ency th erefo re wo u ld ha v e been n o t 6 7
, ,

b u t appro x im at ely ha lf that n u m b er 33 o r th er eab o u t s

, ,

and t he mo dal c oeffi c ient wo u ld ha v e been n o t 2 0 per c ent b u t

n ear 1 0 per c ent T h is being t he case mod al c oeffi c ient s are n o t
.
,

d irect ly c o m para b le exc ept wh en aris in g fro m t he s a me sy s tem

m r em en ts r aft er t he c o effi c ient has been d i v id ed by
of e a s u o ,

the width o f t he c las s ; th u s 2 0 5 eq u als 1 0,

J .

Fo r t he p u rpo s e o f c o m par ing t he v ariab ility o f rac es we u s e

t he c oeffi c ient o f v ariab ility to b e d es cribed lat er The mo da l
,
.

c oeffi c ient is ch iefly v alu able fo r c o m pari ng o n e type with

ano th er wit/t i n t/ze ra ce wh ich is a ll that is req u ired in o rd inary
,

breed in g .

Practi cal v al u e of t he f req u en cy di st ri b u t ion , t h e mode, an d

th e modal coeffi cien t . pract ical im po rtanc e of t he info r ma

T he
t io n a ffo rd ed by th ese v alu es m u s t b e apparent By mean s of .

the freq u ency d is trib u t io n t he breed er is enab led at any t im e ,

wh en he can s ec u re s u ffic ient n u m be rs t o s pread o u t befo re his ,

ey es a go o d and fa ir r epres entat io n o f t he who le p o p u lat io n o f

t he v ariety o r rac e he is breed i ng with res pect t o any charact er ,

wh ich he can m ea s u re o r acc u ra t ely es t im at e .

1
By i t i m
v a r a es t t h i d i id
s l m ean d ( i th i 3 7
e n )
v ua s ea s u re n s c as e 2 ca rs .
 T RA NSM I SSI O N
4 4
2

Wh en he has as c ertain ed its mod e he kno ws what is the

nat u ral ty pe fo r m o d e ind icat es type ; a nd he th en kno ws by
,

ho w m u ch i f any it d iff ers fro m t he ty pe wh ich he has ch os en

, ,
1

as t he s tandard fo r s elect io n By th is he may j u dge wh ether .

and to what ext ent he is o perat ing at v arianc e with nat u re .

Th e mean T h ere is s t ill an o th er c o nc ept io n o f typ e as to

.

th is d is tr ib u t io n and that is t he av erage o r m ean as it is

, ,
“

t echn ically called I t will b e n o t ed that t he d is .

V l V '

t rib u t io n d o es n o t d ec lin e u ni f o r mly b o t h abo v e x 1

and belo w the mo d e ; that is to s ay th ere a re x

,
o on

s ix v alu es b elo w and o n ly three abo v e — fro m 4 x 4

,
1

wh ich we c o nc lu d e that t he a v erage length o f ear

i 2
is so mewhat d i fferent fro m t he m o s t u s u a l length
By m u lt ip ly ing each s epara t e length by t he nu m
i s x

x 9
3
.

b er o f e sar o f that length and add i ng t he pr od u ct s 6 5 3 5 - X 2 -0

hat s t he s a m e th ing add i ng t o geth er t he

’

Z
c z
( w i
z f
'

o r , ,

s
x 32
len gth s o f all t he ears ) and th en d iv id ing by t he
t o tal n u m ber o f ears we find t he a v erage o r x 4 , , 0 :

mean len gth to b e 3 i nch,

e s x 6 7 .

A cc o rd ingly we ha v e t he fo llo wing fo r t he d e 9 5 x 63 5 98 5 - -

x 38
terminat io n o f t he mean : Mu lt iply each v alue
’
x 21
by I ts freq u ency add t he res u lt s and l l d e t he , x 8 ,

s u m by t he n u m b er o f in d iv id u a ls o r v ariat es
3
x .
2

A pp lyi ng th is pri nc ip le to t he ca se in hand we X 1 2 -0 t

ha v e the resu lt seen in the acc o m pany ing table : 4

3 7 88 7 2 2 -0

w
-
t 32 7 :

(be
l
Of
I t is c u s t o m ary t o d ro p o ff ex t rem e ly o ut l yi g n l
v a u es in gs: a
t he di s t rib u t io n , y i th i
b ut ev e if id en t l g m
n s c as v er yl ar e n u

b ers h db a t k th
een b l k w ld h e b e fi ll d ; t h t i
a en es e an s ou f i he
av e n e a s , ea rs o nc s

and i h w ld h
nc es b ouf d; h av e ll l een i l d ed h
o un en c e a v a u es a re nc u ere.

H 1 i ere th s p t
an o i f ert yp e T
co nc e h i d l f
on t h o b d wh i h . he e ea o e ree er, c

ac c e
p t hi
s t
as d d is k
s an i d f ar m i
, s a b i n t y pe
o
q i t d i t i
ec o n o t c or u s n es s , u e s nc

f m t h b i l gi l t ype i d i ted b y t h m d
ro e o o ca nTh p p ca f ll b edi g i e o e
. e u r os e o a re n s

t b i g th tw
o r n el t get h
o as c os e
p ib l e o er as o ss .

By m
3 i h ea n m t th s it h m t i l
ere e ge wh i h i t h
ean e ar
g e em t
ca av ra , c s e a v era os

comm ly on pt d d th
ac c e ymb l f wh i h i M F
e an e s di o io f diff e t c s . or a sc u s s o n o er n

e A pp di
m m
ave ge ra s, s e en x .

( V )
?
f L

1 T h l g b i f m l w ld b M i wh i h
" 1 ’ '
e a e ra c or u a ou e n c

f j
l, ; t h f a re
q i V e V re V t h
u en c es ,
p t i e 1.l d
g th e
’
e r es ec v v a u es , an n

numb f i d i id
er o l mn v d ua s eas u re .

I 1 th i n t bl V t
s d f
a e l m g it d
s an s — i th i
o r v a u es , l gt hor a n u es , n s c as e en ,

and f t d f s an f q s y th
or re mb u en c f i ti
, or
( e )
e nu f h p ra t
er oe v ar e es a rs o eac se a

m em e t
ea s u r Th wh l
n . l m d i te h i lly k n w as f eq e cy
e o e co u n un er s c n ca o n a r u n
4 2 6 T RA N S M I SS I ON

t dy o f v ariab ility it is wo rse t han u seles s t o s t u dy a

I n t he s u

few s catt ered ind iv id u a ls h ere and th ere What we s eek is a .

meas u re o f what m ay b e ca lled t he a v erage t end ency t o d ev iat e

fro m type S o me ind iv id u als d ev iat e b u t litt le o th ers mo re
.
, ,

and s t ill o th ers v ery m u ch and we seek a m eas u re o f th is n o n ,

c o nfo rm ity to type To find th is we m u s t s t u dy g r oups o f in .

d iv id u a ls s u ffi c ient ly large t o b e representat iv e o f th eir ra c e .

T h is bri ng s u s back t o t he freq u ency d is trib u t io n and what it

can t each as to v a riab ility 1
.

im p ress io n t h at if t he la w of h
inf a llib le a ll o ff s p rin g wo u l b e o f a
ere d ity we re d
y
c o m m o n t p e, an d t a t a n
y ep a r t u re f ro m t h eh t p e o f t h e ra c e, d
v a riet , o r b ree y y d
d d h
is t o b e rega r e as b y s o m u c a fa i u re o f ered it a n d a c o n c es s io n t o v a ria t io n l h y .

T he t ru t h is t h a t a ll t ran s m is s io n is h et ero gen eo u s in t he s en s e t h a t t he in di

v id u a ls o f a n
y ra c e, h h
w et er pa ren ts o r o ffs pri n g, be l on
g n o t to a fi xe d typ e b ut
t o a freq u en c yd is t rib u t io n s im i a r to l t he o n e n o w u n d er disc u ss io n . T he i d ea o
f
typ e t/ ms a r is es o u t o
f tire d i s tr ibu ti on , an d it c o ns t it u tes a c o n v en ien t b as e fro m
w ich ht o rec k on d ev ia t io n .

T h e c ief h c o n c ep t io n t o res t in t h e m in d of t he s t u d en t a t t h e p res en t s t a ge o f

ma t t ers is h wh t t at, a ev er t h e p a ren ta ge, t h e o ffs p rin
g wil l c o n s t it u t e a di s t rib u t io n

ex te d i g t h
n n gh ro u a co nsi d l
e ra b e ra n ge, a n d t h at the p a ren t it s e f l a ls o l
b e o n ge d
an d w d w f m
as ra n ro s o me p o rt io n of a f req u en c y di s t rib u t io n n o t v er yd ifferen t
fro m th t f th a o e ra c e in gen era l .

V i b il it y i
ar a th s, erefo re, n o t t h e o pp o n en t of h ere d it y b u t its in ev itab le
ac c o m pa n im en t in t ra n s m is s io n , and o u r p ro b lem is to d ev is e h d f
met o s o ac c u

ra tel
y m eas u rin g a n d exp ress in g it s ran ge an d ex t en t in an y p ar ti l i t
c u ar n s an c e .
1 No a po lo g y is m a d e fo r in t ro d u c in
g t h e s o -c a ll de s t a t is t ic a l m t h d f t dy
e o o s u

at t his p o in t ; l y l i b l m th d f
firs t , b ec a u s e it is t he o n re a e e o o a tt ki g p b l m
ac n ro e s

i t
n mi i
ra n s d d b
s s o n , an th i m th d i
s ec o n ywh , mi g i t
ec a u s e s e o s ev er e re c o n n o u se

a m g on f l t d t
ca re u Th s u d i g d d th t d t h ld b q i d
en s . e rea er s u r e , an e s u en s ou e re u re ,

no t t o d th i p t i
ev a e f th s bj t b
or on th m th d f t
o tm
e su t m y ec ec a u s e e e o o rea en a

h pp t b
a en f m il i
o e un aO th th h d h i g d t f m ili i h im lf
ar . n e o er an , e s ur e o a a r ze se

no t l y with t h m th d f w k b t wit h th p i t f i w i l d I f h wil l

on e e o o or u e o n o v e nv o v e . e

d th i b t h
o s, i b il it y
o d l t
v ar a l ti an d h d ity i g l wil l
a er o n c o rre a m o n an ere n en era co e

to h wm
av e a ne i g d f m
ea n n ti
, lan d mp h one i th ar th o re ra ona an co re en s v e an e

h y ti
az no l d f m th
o ns ev o v e y t m ti t dy f i l t d i d i id l T h
ro e uns s e a c s u o so a e n v ua s . e

p ir ncip l i les d f
n v o ve t h m t p t
a re im p l or d i t h i l
e m t os
y t t i ar s e, a n n s e e en a r rea se

ev er y ff t will b m d t t t th b j t f m th t d p i t f th
e or e a e o rea e su ec ro e s an o n o e non

m t h m ti l
a e a d ca rea er .

F th
or i f th
e c o n v en en c e o wh m y t p lit t l f t h
ose m o f a c a re o u rs u e a e ur er s o e o

th m e t i t l y m t h m ti
o re s r c l p tai ei l ad A p ca
p d i h
conce b p ons nv o v e , an en x as een re

p d
a re b y D H L R i
r. t f t h. m t h. m t i l
e z, o d p t m t f t h U
e i i t
a y f e a ca e ar en o e n v ers o

I llin o is .

Th e l m th d f t dy f p b l m i h d ity d i t i t f m t h
s ta t is t ic a e o o s u o ro e s n ere , as s nc ro e
t i t l y b i l gi
s r c l w i t d d b y D F i G l t f E gl d ( N t l
o o ca , as n ro uc e r
. ra n c s a on o n an s ee a ura

I h it
n er a n c e, d ft w d m h t d d by K l P
an d th
a er ar uc ex en e ar ea rs o n an o ers

(s ee es p i ll y G
ec am m f S i d Ph il p h
ra i l T t i
ar of t h R y l c en c e an o so ca ra n s ac ons o e o a

S i ty )
oc e It i . w mi g i t h mm
s no th t co
q t ly j l n n o s uc co on us e a a u a r er o u rn a
 Y
T PE A N D V AR I A BI LIT Y 4 2 7

A ga in t h e c o ncret e ser v es well as a med iu m fo r t each in g a

pr inc ip le I n th is c o nn ect io n we refer o nc e mo re t o o u r d is t ri
.

b u t io n o f 3 2 7 ear s and n o t e that ev ery ea r i n t/ze lot d ev ia tes

,

s o m ewhat fro m t he m ean o f i nch es T he range and ext ent .

o f th is d ev iat io n a re S h o wn in t he f o llo wing tab le c o lu m n D ,

.

T he pract ica l q u es t io n n o w is t o red u c e

D EV IATI O N o r 3 2 7 EA RS
th is c o lu m n o f d ev iat io n s to a s in gle ex
o r C OR N FR OM TH E I R
press io n d eno t ing the v ariab ility o f the M E A N L E N G T H o r
po p u lat io n o f wh ich th is d is trib u t io n is I N C HES
repres entat i v e M an i fes t ly when th i s I S
.

V
,

d o ne the v ari ab i li ty o f th i s d l s trl b u tl o n

,

can b e c o m pared d irect ly and exact ly with

t hat of any o th er and at the pr esent o r , 4 -0

any f u t u re t ime T wo meth o d s of p ro .

4 5

c ed u re a re po s s ib le in th u s sec u ri ng a k ind

o f gen era l expres s io n fo r t he a v erage

a mo u nt o f d ev iat io n g iv ing ris e to t wo ,

s i m i l a r b u t s l i g h t l y d iffer ent v a lu es ,
12

9 '
na mely t he av erage d ev iat io n and t he
,

s tandard d ev iat io n .

Th e av erage dev iati on If each d ev i . 67

at io n (c o lu m n D ) repres en t ed an eq u al 95 6 3
n u m ber o f ears th is s in gle expres s io n ,

c o u ld b e read ily d eriv ed by add in g t he 13 ;

2

d ev i at io n s and d i v id i ng by t he t o t a l 2

n u m ber B u t th es e d ev iat io n s d o n o t
.

repres ent eq u al n u m b ers o f c a rs T he 3 7 .

2

d ev iat io n fo r exa m p le repres en ts b u t o n e c a r wh ile n o , ,

less than t welv e ears d ev iat ed inch es belo w t he m ean a n d

t wo d ev iat ed i n ch es abo v e with o th ers u n ev en ly d is tr ib u t ed , .

Man ifes t ly eac h d ev iat io n S h o u ld fir s t b e m u lt ip lied by the

n u m ber o f c a rs in v o lv ed as in t he s u cc eed ing table : ,
1

( B i m
o t ib
e r ) i d a t d
s t t h
ev o e p t f t t i t i
o l t d i i
e re o r s o l t i t h i
s a ll y
s ca s u es n ev o u on , ec n ca

k w
no n asbi m t y o e r .
”

1Wh th d
en i ti e i t b
ev a b t i d i t h i w y th m i
on s o eo ig i d i
a ne
g n d d s a e n us s n s s re a r e .

D i di t
‘1 n th d i ti f th
c a es e ev a l g p f m th m m m
on o e s ev e ra f ro u s ro e co on ea n o

th e ra c e, i h Th
nc esf . mpl th fi t us, d i t th d iff
or ex a e, e rs c ar ev a es e eren c e

b tw
e een
3 i h nc d i
es a n h w h i h b i nc
g b l w t
es , o r h m i w i t t c , e n e o e ea n , s r en

wit h t h g t i
e ne a ig ; l
ve s t h
n i
a so h l g d i t
e 2 1 ea rs -
nc es on ev a e or

i h
nc f m th m
es ro e d b i g b
ea n , an th m w w it it p it i
e n a ove e ea n , e r e os v e.
 8 T RA N SM I SS I ON
4 2

f D D] T he res u lt o f th is calc u lat io n is that the

1 x t o tal d ev iat io n o f 3 2 7 ears fro m th eir a v erage
0 X length is i nch es s o m e abo v e a n d s o me ,

1 x
belo w t he m ean I f n o w we d iv id e . by
0 x

2 x 3 72 t
,
he n u m b e r o f ear s i n v o l v ed we ha v e ,

x inch es wh ich is a g o o d expres s io n o f

3 3 33 ,

9 x t he a v erag e d ev iat io n o f th is p a rt ic u lar po p u

8 x la t io n I f an o th er v ariety S h o u ld g iv e a la rger
.

12 x

19 x
q u o t ient we s h o u ld c o n c lu d e it t o b e mo re
,

v ar iab le I n th is m ann er we m ay red u c e t he

.

3 2 x

0 x v ar iab ility o f a wh o le p o p u lat io n t o a s i n g le

4
67 x expres s io n .

63 x
S tan da rd dev ia t ion M ath em at ic ian s hav e .

3 8 X

21 x
ano th er meth o d o f ca lc u lat ing v ariab ility I t .

8 x
d iffers fro m the o n e j u s t d is c u s sed in o n ly o n e
2 x d etail ; n a mely t he d ev iat io n s a re sq u a r ed
,

I x bef o re mu lt iplicat io n by th eir res pect iv e fre

3 2 7 e n c ies as in t he tab le wh ich f o llo ws
q u ,

Th e l
c o u mn ma r k d
d b y q i g th
e I ) 8 is i s ec u re s ua r n e v ar o us d ev ia t io n s , t h us

el im i ti g th m i
na n eig F mpl
n us S n . o r ex a e, x

1 T h l m
e co u m k d D f i b t i d b y m l tip ly i g th
n’ ar e s o a ne u n e s q u a re dd ev ia t io n s

eac h b y it p t i
s resf qec y F
ve m
re
p l u en c x 9 . o r exa e,

1 Th G e k pit l igm ( 23) i t h

ree ca a s l ig f mm ti
a s e u s ua s n o su a on in ma t h em a tics .
 0 T R A N SM ISS I ON
43

by th is a m ou nt and a rr iv e at t he tru e
m ean 3
C o n s id eri n g n ext t he s tand a rd d ev iat io n I n c o lu m n f V
"
1
G)
we ha v e a s t he s u m o f t he pro d u ct s o f t he S ev era l fre

u en c ies i n t o th eir res p ect iv e d ev iat io n s fro m t he a s s u m ed m ean

q ,

d eriv ed o n t he sa m e p la n as wh en wo rk in g fro m t he t ru e m ean

—
V G) ( V -
G) ’ V -
m :

I
D D iv id ing
by t he t o ta l n u m b er ( 3 2 we ha v e 3 7
2
—
.

c o rres po n d ing to t he q u o t ient 1

3 7
2

o f t he prev io u s ca lc u lat io n wh en wo rk ing fr o m t he tr ue m ean .

T he c o rrect io n m a d e i n t he m ean wa s b u t as we a re
n o w d eal i n g with S ec o nd p o wer s it S eem s b u t nat u ra l that t his

a m o u nt s h o u ld b e s q u ared befo re it can b e tak en fro m t he q u o

t ient T h is will b e j u s t ifi ed by a m ath emat ic al proo f in
t he A ppend i x T he s q u are o f
. 0 1 7 1 is -
or .

On t he ot h h d
er an , s ho ul d th me su o f t he p o s i t iv e d ev ia t io n s exc ee d th e sum

of the n ega t iv e d i ti
ev a on s it w l d i d i
ou n c a te t h a t o u r as s u m e d l
v a ue is t o o s m al l
an d we s h ld
ou a dd t h e c o rrec ti on i d
n or er t o a rri v e a t t h e t ru e m ea n .
 Y
T PE A N D V A R I A B I L I T Y 43 1

We hav e therefo re as a c o rrect io n on acc ou n t of t he tr u e m ean

T h is grees v ery n early with t he v a lu e

a prev io u s ly
fo u nd b u t th is s ho rt er meth od is t he mo re acc u rat e beca u se
, ,

fewer d ec im als hav e b een lo st The s q u are roo t o f

. is
t he s tandard d ev iat io n s o u ght agreeing p erf ect ly with
,

t he f o rm er v al u e and d eriv ed by a v ery m u ch s h o rt er m eth o d .

T he s t u d ent will no t e that t he d ifferenc e in t he t wo m eth o d s

is ess ent ia lly th is : in t he latt er we d ea l o n ly with d ev ia tion s ,

wh ile in t he f o rmer ent ire v a lu es a re in v o lv ed I t is t ru e that .

d ev iat io ns a re tak en fro m an ass u m ed m ean b u t t he c o rrect io n,

is acc u rat ely m ad e and t he wh o le o p erat ion can b e carried

,

fo rward n o t o n ly with s maller n u m bers b u t a lso with o u t the

lo s s o f d ec im a ls n ec ess ar ily i n v olv ed in t he m o r e d irect b u t far
m o re lab o rio u s and o n t he wh o le les s exact met hod fi rs t gi v en .

T he fi rs t meth o d is u s ef u l fo r exp o u nd ing t he pri nc ip les in

v o lv ed b u t t he lat er is far pr ef erab le fo r act u a l u se n o t o n ly o n
, ,

acc o u nt o f it s brev ity b u t o n acc o u nt o f it s increas ed acc u ra cy

,

as well .

A v erage dev i ati on an d stan da rd dev iat ion t

con r ast ed . Th es e
t wo e xpres s io ns fo r v ariability res t u po n t he sa me ar ith m et ical
p rinc ip le b u t t he latt er has d ec id ed m ath em at ical ad v antages
,

o v er t he f o r m er fo r m any p u rp o s es and is t he o n e u n iv er s a lly

u s ed by m ath em at ic ian s The o n ly ad v antage in t he a v erage

.

d ev iat io n lies in the sim p ler ca lc u lat io n as n eith er s q u ares n o r

,

roo t s a re in v o lv ed W ith t he s h o rt ened m eth od o f fi nd ing t he

.

s tandard d ev iat io n h o wev er th is ad v antage is s light

, ,
.

I t mak es litt le d iff erenc e which is u sed in pract ic e pro v id ed ,

t he S a m e m eth o d is a lway s em p lo y ed The res u lt s o bta ined d i ffer

.

c o n s id erably as c o m par ed with T he s tandard

d ev iat io n is a lways larger than t he a v erage d ev iat io n beca u se o f
t he s q u ar ing o f t he s ev era l d ev iat io n s I t th u s exaggera t es the
.

wid er d epart u res fro m type as c o m pared with t he meth od s

em p loy ed in find ing t he a v erage d ev iat io n Th is b ei ng t ru e
.
,

res u lt s o btain ed by the two pro c es s es can n ev er b e c o m pared ;

t hat is wh en d ealing with v alu es d es ign ed to expres s v ariability
,

we m u s t a lway s kn o w wh eth er a v erage d ev iat io n o r s tandard

d ev iat io n is m eant .
43 2 T R A NS M I SS I ON

T he breed er ch oo se eit her meth od b u t hav ing o nc e

m ay ,

ch o sen a ll his rec o rd s m u s t b e m ad e in t he s a me way I n

,
.

as m u c h as t he s tandard d ev iat io n has d is t inct math em at ica l

ad v antages o v er t he s imp le a v erage d ev iat io n and inas m u ch as ,

it is the on e c o m m o n ly em p lo y ed in math em at ical lit erat u re it ,

is t he o n e that will a lway s b e emp loyed in th is t ext Wh en .

,

therefo re a meas u re o f v ariability is ment io n ed it will b e the

,

s tandard d ev iat io n and no t t he a v erage d ev iat io n that is m eant

1
.

M eanin g of stan dard dev iation T he s tandard d ev iat io n is a .

goo d meas u re o f v ariab ility fo r t he cha ract er in q u es t io n and

t he rac e i n v o lv ed I t th erefo re a ff o rd s a r elia b le ba s is f or com
.

p a r i ng t/t e v a mabil i ty of on e rac e wi t/c tha t of a nother a s to tire

clza rac ter u n d er c on s id era tion , or of on e c lt a ra c ter wi t/t t/za t of

a no tlzer c i t/t er i n tlze sa me or i n d zf eren t ra ces .

In a sc ertain ing the s tandard d ev iat io n t he mea n o r av erage , ,

o f t he rac e was tak en as t he bas is and a ll d ev iat io n was reck o n ed

,

fro m that I t is man ifes t h o wev er that if t he m od e b e taken

.
, ,

as a bas is and d ev iat io n reck o n ed fro m th is f o llo wing t he s a m e

, ,

m eth od s as fo r d et erm in ing t he s tandard d ev iat io n a so m ewhat ,

d ifferent v alu e will res u lt Wh en ev er as in m o s t cases t he

.
, ,

m od e d oes n o t c o inc id e with the m ean th is will repres ent the ,

d ev iat io n fro m the prev ailing typ e wh ich is o ft en o f mo re prae ,

t ical im po rtanc e to t he breed er t han the d ev iat io n j u s t d es cribed ,

es pec ially t o t he breed er who pro po ses to d ea l with i nd iv id u a ls

s elect ed with refer enc e n o t t o t he m ean b u t t o t he prev a iling

type .

I n t he sa me m anner t he breed er may calc u lat e the d ev iat io n

fro m his o wn id eal type o r s tandard and in th is way a sses s the
d ev iat io n no t fro m t he pres ent mea n o r type b u t fro m t he o n e
,

he h o pes to es tab lis h I n th is way he is ab le to k eep acc u rat ely

.

info rm ed fr o m y ear to y ear as t o t he progres s he is m ak i ng and

the d egree o f s u cc es s that i s fo llo wing his s elect io n s .

The wri t er has n ev er seen eith er o f th es e d et er minat io ns

u s ed b u t th ey a re es pec ia lly v alu ab le to the br eed er who d es ires
,

t o k n o w ho w a c erta in v ariety d ev iates o n t he a v erage fro m it s

1 I n t he s ec t io n o n Pro b ab ilit yC u rv e ( pp
A en ix id
) t will b e s hwn t hat the
o

s ta n d d d
ar ev ia tio n is on t he wh lo e p refera b le fro m t he p u rel y mathema t ic al
s t an dp i t
o n .
4 34 T R A NS M I SSI ON

with that o f any o th er rac e plant o r an imal and fo r any charac , ,

t er o f wh ich acc u rate m ea s u rement s can b e tak en and a fre

q u eney d is trib u t io n b e c o n s tru ct ed l

.

Pr acti cal appl ica t ion of th e coefli cien t of v ariab il i ty . T he

va lu e o f th is ter m lies in the fact that it a ff o rd s an acc u rat e
m ean s o f c o m pari ng d irect ly the v ariab ility o f o n e freq u ency
d is trib u t io n with that o f ano th er no ma tter wlza t tlze clea rac ter , ,

w/zetlzer i n tb e sa m e d ifleren t i n d i v id u a l s, b et ween s i m i la r

'

or or

o r u n li k e s p byec ies th i . Th u s ,
s m ea n s we ma
y c o m par e t h e v ar i

ab ility o f the leng th o f an ear o f c o rn with that o f its weight ;

t he v ariab ili ty o f it s c irc u m f erenc e with that o f it s n u m ber o f
ro ws o r with that of any o th er m ea s u ra b le charact er
,
.

W e can also in th is way c o m pare t he v a riab ility o f ea rs o f

, ,

c o rn with that o f th eir s ta lk s ; with th a t o f t he length o f h o rse s ’

leg s o f what t hey can p u ll o r o f t he rat e at wh ich th ey can

, ,

tra v el ; with that o f the h eight o f men th eir weight length o f , ,

arm s meas u rement s o f the h ead

,
— ind eed wit h that o f any , ,

o bj ect liv ing o r no n liv ing that po s s es s es v ariab le charact ers

,
-
,

that can b e acc u rat ely meas u red wh eth er by feet inch es , , ,

po u nd s o r by any o th er u n it that can b e d ev ised

,
?

By mean s o f th is c o effi c ient t he breed er m ay n o t o n ly a s c erta in

wh eth er o n e charact er is m o re v ariab le than an o th er b u t by ,

tak ing th is c oeffic ient freq u ently as ann u ally fo r the s a me v ar iety ,

o r u nd er d i ff er ent c o nd it io n s he can kn o w t he v ar iab ility o f t he ,

s a m e charact er a s in flu enc ed by t i me o r c irc u m s tanc es .

I n gen era l th es e d et e rm inat io n s enab le t he breed er to kn o w

wh ich way h is v a riet ies a re dr ift ing o r wh eth er th ey a re s tand ing ,

1Cl l y if t h m d
ea r th
, te d d f l ti h b
o e or d
e s an b i ar o s e ec on as een u s e as a as e n

l l t i g t h d i ti t h th m l h l d b
ca c u a n e ev a o n, den b e sai l e va ue s ou e u se as a ase n ca c u

l ti g h
a n ffi i
t ; th
e co eit i p ib l t
c en t usffi i t f s i b il it y f m
oss e o s ec u re a c o e c en o v ar a ro

an y d i d t y
es rep w ll f m t h
e as m e as ro e ean .

2Th ffi i t
e coef i b il it y h
c en b w k d
o t f
v ar a l g mb f as e en or e ou or a ar e nu er o

c h t i m
a ra c ers i h w i th f ll wi g t b l ( V
n an , as s s o n V i ti
n i e o o n a e S ee ern o n , ar a on n

A im l
n d Pl
a s an t p an s , .

N l gt h
o se en 9 49 H d b m dth - ea

N b
o se d th rea 7 57 U pp m l gt h - er a r en

N h ight
o se e F m l gt h o rear en

F h d h ight
o re ea e U pp l g l gt h er e en

U d n j w l gt h
er a en L w l g l gt h o er e en

M ou th b rea dt h Foo t len gt h

Hea d en gt h l
 TY PE A N D V A R I A BI LIT Y 435

s t ill in s pit e o f v igo ro u s selec t io n ; wh eth er in his select io n he

is o pera t ing with o r agai n s t n at u re ; wh eth er the type is b ec o m
in g a litt le m o re fix ed o r wh eth er t he t en dency is mo re and
m o re t o s catt er ; wh eth er the m ean o r gen era l av era ge of exc el
lenc e is appro ach ing o r rec ed ing fro m t he d es ired type wh eth er ,

as t im e pa ss es v ariab ility is les s en ed o r i ncreas ed ; wh eth er as

, ,

t he res u lt o f s elect io n n ew v alu es are c o mi ng in at t he u pp er

,

en d . I n s ho rt by th es e ca lc u lat io n s he may kn ow wh eth er he is

,

mak ing real pro gres s o r is o n ly dabb ling in the wh i rl o f v ari

,

ability that is inev itable wi th all liv ing th ings wi th o u t in flu ,

c u ei ng at a ll t he trend o f t he ra c e I t is n eed less to s ay that

.

m u ch o f th is latt er so rt o f ineff ect iv e breed ing is going o n all

abo u t u s ev erywh ere The s tat is t ica l meth od is t he o nly kno wn
.

m eth od o f sec u ring acc u r a te kno wledge o f ty pe and v ariab ility ;

fo r type is n o t s i m p ly W hat we d es ire b u t it is what we act u a lly ,

g e ,
t a n d any br eed e r i
'
s wo rk i ng in t he dark w ho d oes no t k n o w
t he real nat u re o f th e wk ole p op u la ti on of t he rac e he breed s .

SECT I ON III — PR A CT I C A L HI NTS ON T HE TA K I NG A N D

G R OU PI N G OF M EA SU R EM ENTS

I t is h igh ly im pro bab le that in m eas u r ing any part o f an

,

o rga n is m a p erf ect ly acc u rat e mea s u re is o bta in ed

,
I t m ay b e .

v ery eas y t o mea s u re t he length o f a n o bj ect acc u rat ely t o 1 inch ,

o r to inch o r ev en t o
,
inch if the r u ler is s u flic ien t ly
,
‘

acc u rat e ; b u t fi na lly in t ry ing to m eas u re to a s h igh a d egree

,

o f acc u racy as p o ss ib le we c o m e to a p o int wh ere o u r ru ler

,

fails u s A ga in the o bj ect m ea s u red m ay b e of s u ch a nat u re

.
,

that it is fu t ile to try to tak e m eas u rement s beyo nd a c erta in

d egree o f acc u racy F o r in s tanc e it wo u ld b e u seless to t ry to
.
,

m eas u re the length o f an ea r o f c o rn t o i nch S im ilar ly in .

,

weigh ing s u b s ta n c es it is p os s ib le with a goo d balanc e t o tak e

,

m eas u rement s to ou nc es o r t o t enth s o f m illigra m s if extreme

acc u racy were d emand ed b u t finally in s t riv ing aft er acc u ra cy , ,

t he balanc e fa ils u s and we m u s t grant that it is h igh ly im p ro b

,

able that the weight wh ich we rec o rd is perf ec tly acc u ra t e .

Wh ile we th u s S ee that we canno t attain absolu t e acc u racy in

any m ea s u rement s yet threm m at ology is no exc ept io n al fi eld in
,
43 6 TRA NSM I SS ION

th is res pect and experienc e has s h o wn that meas u rem ent s can b e
,

eas ily tak en o f s u ffi c ient acc u racy t o i n s u re v ery reliab le res u lt s .

Fo r o bta in in g t he freq u ency d is trib u t io n o f a po p u lat io n with

res pect to so me m eas u rab le charact er it is impract icab le to lay ,

d own s pec i fic r u les in regard t o t he acc u racy of meas u rement s .

Th is m u s t b e sett led largely by experienc e and c o m mo n sen se .

I t may h o wev er b e sa id h ere that i n m eas u ri ng a large po pu

, ,

lat io n u nd er t he free act io n o f t he la ws o f pro bab ility s u b s ta n

, ,

t ially as m any m ea s u rem en t s will b e tak en too large as a re

taken too s m all Henc e S light erro rs in m ea s u rem ent s d o no t
.

apprec iab ly d is t u rb t he m ean and v ariab ility o f the po p u lat io n ,

b eca u s e t/zey ten d to ojs et ea cli otlzer

’

Sch eme of mea su remen t R ev ert ing to . freq u ency d is t he

t rib u t io n o b ta i n ed fro m meas u rement s o f c o rn it will b e n o t ed ,

that th is po p u lat io n is d is trib u t ed in c lass es o r gro u ps which

d iffer by a half inch in lengt h F o r the pu rpo se o f fo r ming th is
.

freq u ency tab le th ere wo u ld th en b e n o o bj ect in t ak ing the

mea s u rem ent s c loser than t he n eares t half inch This ra is es t he .

q u es t io n o f the in c lu s iv enes s o f a c la ss in gro u p in g a po p u lat io n ;

that is S h o u ld th ese m eas u rem ent s ha v e been tak en at the
,

q u art er inch o r perhap s at t he ev en inch ? I l ere aga in no hard , ,

and fas t ru le can b e laid d o wn ; b u t it m ay b e sa id that in

genera l and fo r the bes t res u lt s the c las s ra nge s h o u ld b e mad e
, ,

j u s t large en o u gh fo r s o me v a riat es to appear in each c las s ,

exc ept ,
po s s ibly in a few n ear the extrem es o f the range of
,

ria b ilit
y w h en t he t o tal p o p u lat io n is n o t v ery large .

S o met im es t he b es t u n it fo r m ea s u rement and gro u p ing

will b e ev id ent b u t freq u ent ly s o me prelim ina ry wo rk is n ec es
,

sa ry in o rd er t o d ec id e it Fo r exa m p le in beg inn ing t he

.
,

s tat is tical wo rk with c o rn we at fi r s t t o o k m eas u r ement s t o t he

q u art er inch with gro u p ing s acc o rd ing ly b u t fo u nd n o res u lt s

, ,

d iff erent either as t o mean o r v ariab ility fro m th o se o btain ed

, ,

wh en meas u rem ent s were tak en t o t he ha lf inch and b u t s light ly ,

d ifferent fro m th o s e tak en at t he ev en inch A cc o rd in gly we .

a re u s ing t he ha lf inch m ea s u rem ent s fo r extreme acc u racy and

-

t he ev en inch fo r ro u gh er wo rk .

M u ch j u dg ment m u s t b e u s ed i n d ec id ing u po n t he s ch eme

o f m ea s u rem ents to b e ad o pt ed and t he gr o u p ing s t o b e m ad e .
43 8 TR A NSM I SSI ON

T he real na t u re o f pro bab le erro r and the meth o d s fo r

d ed u c ing t he fo rmu las fo r its calc u lat io n will b e c ov ered in
t he A pp en d i x wh ich is d ev o ted es pec ially to m ath em at ical
,

m eth o d s and c o nc ept io n s b u t en o u gh s h o u ld h ere b e s aid to

,

gi v e the s t u d ent an intelligent id ea o f what is m ea nt by pro bab le

erro r and t o acq u a int him with t he bare fo r m u las fo r it s c alc u

lat io n as t o t he v alu es h ere u nd er d is c u ss io n .

The pro b a b le erro r ( d en o t ed by :t E ) is a pai r o f d iv ergenc ies

ly in g o n e abo v e and the o th er b elo w t he v a lu e d et erm in ed an d ,

o f wh ich we can s ay with c o nfid enc e that t he re is an ev en chan c e

that the tr u e v alu e lies bet ween th ese limit s T hese n u m bers
1
.

a re n u merically eq u al b u t o n e is rega rd ed as p lu s t he o t her as

, ,

m in u s ( i E ) and t he t wo d efi n e a ra nge with in wh ich o u t of a

, ,

v er large n u m ber o f d et er m inat io n s at lea s t ha lf t he t ru e v a lu es

y ,

wo u ld b e fo u nd T h is b ein g t he c as e we may say o f any s ingle

.
,

d et erm inat io n that the chances are ev en th a t any erro r in v o lv ed

will no t fall o u t s id e t he lim it s s et by :t E I t is o b v io u s th ere .
,

fo re that the s m aller t he pro bab le erro r t he narro wer th is ra nge

, ,

t he great er c o n fid en c ewe s h o u ld p lac e in o u r d et erm ina t io n and ,

th e s m a ller a re the chanc es o f a large erro r ha v ing been m ad e .

T he expres s io n “
pro bable erro r may b e m is lead ing I t is .

not as m ight b e s u pp o s ed fro m t he wo rd s

,
t he m os t p r ob a ble ,

err or The mo s t pro bab le v a lu e is o u r d et er m inat io n and t he

.

m os t pr ob a b le err or is zer o N eith er d oes the pro bab le erro r

.

fi x the limit s of erro r b u t it is an extrem ely goo d m eas u re of

,

acc u racy in that it fix es a range abov e and belo w the d et erm ined
v a lu e s u ch that t he c /za nc es a re ev en that t he tru e v a lu e lies

wi t/zi n t/zis ra n
g e .

T h u s , if
a series o f calc u lat io ns res u lt s in a fi nal n u m ber
with a pro bab le e rro r o f i 1 2 it m ean s that o u t o f a great .
,

n u m b er o f ca s es t he tr u e v a lu e o f o n e ha lf will lie bet ween

1 2 ) and .
( 74
2

I f an o th er ca lc u lat io n in v o lv ing larger n u m bers o r mo re ac

c u rate meth o d s s h o u ld res u lt in t he s a me v a lu e b ut a ,

pro bable erro r o f o n ly :i: 04 th en the tr u e v alu e has an ev en

.
,

chanc e o f ly ing bet ween and wh ich is a v ery

1Th i f l
ere s , o h th t th t
c o u rs e, a so a n ev en c l li t id
an c e th a e ru e v a u e es o u s e e

sa m l im it
e s.
 TY PE A N D V A R I AB I LITY
4 39

narrow m argin gi v ing u s m u ch c o nfid enc e in the d eter m inat io n

, ,

with o n ly a s m a ll cha n c e o f being wid e o f t he tru th .

Of c ou rse the erro r in a d eter m in at io n has a ls o a n ev en

chanc e o f ly ing o u t sid e the lim its set by t he pro bab le erro r ( E ) ,

b u t t he follo wing tab le will s h o w that it is v ery u n lik ely that

t he err o r is m any t im es a s great as E Th u s the chanc es that .

t he tr u e v alu e lies wi t/t i n t he range s et by l: 2 E etc a re

: ,
.
,

as fo llows
1

i E the c h a e
: e e e nc s ar v n

i 2 E the h
s ce e t 1 c an s ar o

l 3 E the c h a e a e 2 1 t 1
:: nc s r o

i 4 E the h a
zz e e 142 t 1 c nc s ar o

i 5 E the c h a c e
: e 13 10 t 1 n s ar o

t 6 E the c h a e
z re t 1 nc s a o

j 7 E the c h
; ce a e t 1an s r o

i 8 E the h
: e e c t 1
an c s ar o

3 9 E th e h a c e
; e b t c n s ar a ou

I t is e xtrem ely impro bable th erefo re that an erro r will b e , ,

m any t i m es a s larg e a s the pr o bab le erro r Fo r in s tanc e it is .

,

pract ically c ert ain that t he erro r is no t as large as 9 E sinc e ,

t he tab le s h o ws that the chanc es are abo u t a b illio n t o o n e in

fav o r o f its being s maller than 9 E .

T h u s by giv ing a lo ng with any res u lt t he c a lc u lat ed p r oba b le

, ,

err or the read er ma kn o w what d egree o f c o n fid enc e is to b e

, y
p lac ed in t he res u lt s .

Fo r a graph ic illu s tra t io n o f the m ean ing o f :l: E s u ppo s e in ,

the fo llo wing figu re t he li n e A B represent s o u r d et er m inat io n

—
,

’ ’
and t he lines a b and a b the locat io n o f E and E

N ow th is m ean s that if A B is n o t the t ru e locat io n o f t he v a lu e

in q u es t io n t he chanc es a re ev en that th is locat io n is n o t o u t s id e
,

t he lim i t s s et by t he lin es a b and a b repres ent ing :t E

’ ’
.

1 C B D a v en po rt , S ta tis tica l M et
. . h o ds , p . 14 .
44 0 TR A N SM ISS IO N

If no w we set o th er lines lik e t he fo llo wing at :i: 2 E,

' ’
1
a, a A a a

' ’
1
b, b B b b

th en we kno w fro m t he table that t he chanc es a re to 1

that t he tr u e po s it io n o f A B is n ot o u t s id e the lin es a l b l an d a b
‘ ’
,

eac h r em o v ed t wic e t he pr o bab le err o r fro m the d et erm inat io n .

Pr ob ab l e er ror of mean T he pro bab le erro r o f t he mean is

.

based u po n t he s tan da rd d ev iat io n as we no t ic e by t he fo llowing ,

fo rm u la 1
s ta n d a rd d e ia t io n v
ziz o . 674 5
V n u m b er o f v a ria tes
or EH t
: 74 5
V;
S u b s t it u t ing fo r t he cas e in ha nd ,
we hav e
i: 0
: . 674 5 i: o 0 4
: .
7

T he t d ent canno t fa il to n o t ic e the o v erwh elming infl uenc e

'

s u

o f n u m ber s in c o ntro lling t he v a lu e o f E o r to rea lize that if ,

t he n u m ber o f d et e rm in at io n s s h o u ld bec o m e infini t e E wo u ld ,

bec o me zero .

Prob ab l e error of stan dar d dev iat ion A cc o rd ing t o meth od s .

o f d ed u ct io n to b e d is c u s s ed lat er t he pro bab le e rr o r o f an y

, ,

d et erm in at io n fo r s tandard d ev iat io n is fo u nd by t he follow

in g pro c es s : d iv id e the s tandard d ev iat io n by t he s q u are roo t
o f t wic e t he n u m ber o f v a riat es and m u lt ip ly the res u lt by

I n the cas e in po int we hav e as t he s tandard d ev ia tio n

wit h 3 2 7 v ariat es S u b s t it u t ing th ese n u m b ers in t he fo rm u la

.
,

we hav e 1 . 28
_ .
z: l
X 327

1 S ee C B D av en p o rt , S ta t is t ic a Met o s , p 1 5 T h e m et
. . l h d . . h d
o h
b y w ich t he
c o n s ta n t d ll
is o b t a in e wi b e ex p la in e in t he A ppen ix d d .

1 The fo rm u la fo r
p r o b a b e er ro r l
o f s t a n a r ev ia t io n is d dd
i : 0 67 4 5 1 zn
S ee C B D a v en po rt , S ta t is t ic a
. . l Met h o ds , p . 16 .
 2 T R A N S M I SS I O N
44

If a c irc le b e d ra wn abo u t t he c ent er with a rad iu s eq u al to

t he a v erage d ev iat io n o f A th is c irc le will fa ir ly well repres ent ,

his m ark s m an s h ip ; that is to s ay t he a v erage d is tanc e o f a ll his ,

S h o t s fro m t he c ent er is t he s a m e a s if th ey all lay o n th is c i rc le .

I f t he m ark s m an s h ip o f B is n o t so go o d as that o f A his av erage ,

is great er and t he c irc le c o rres po nd ing ly larger 1

.

N o w n eith er o f th es e c irc les is an a bs olu te ind ex o f t he m ark s

m an s h ip o f ei th er A o r B u n less an infin ite n u m ber o f s h o t s ,

ha s b een m ad e ; that is to s ay if o n ly a few s h o t s h a v e been ,

fi red t he pro bability o f erro r is grea t if we as su m e thes e c irc les

,

t o b e f u lly repres entat iv e o f t he m ark s m an s h ip b eca u s e th ere ,

is practical c ertai nty th a t s u cc eed ing s h o t s will b e eith er bett er

o r wo r s e ; ind eed th ere is a lway s a chanc e that t he n ext m a b e
, y
a lu cky s h o t and lo wer the d ev iat io n o r a wild o ne and ra ise it ,
.

W e s h o u ld th erefo re c o nc eiv e of t wo o th er c irc les ly ing

n eighbo r to each o f th o s e repres ent ing the ca lc u lat ed d ev iatio n s .

Th es e a re repres ent ed in the c u t by t he light lin e c irc les and -

giv e a graph ic m ean ing to the pro bab le erro r Th ey a re d ra wn .

s o that i f t he h ea v y c irc les d o n o t acc u rat ely represent A s and

’

B s m ark s man s h ip th en the chanc es a re ev en that t he tru e

'

po s it io n o f the c irc le repres ent ing A s m ark s m a n sh ip fo r exam '

p le lies s
, o m ew h e r e b et w ee n t he light lin es r e pr e s ent ing t h e

pro bable erro r of t he c o m p u tat io n The ch anc es are o f c o u rs e .

a ls o ev en that it lies be y o n d th es e l im it s ei th er w it h i n o r o u t s id e ,
.

Ob v io u s ly t he s m a ller t he pro bab le e rro r the great er t he c o n fi

,

d enc e to b e plac ed in the calc u la t ed d ev ia t io n T he s tu d ent is .

ca u t io n ed h ere that in th is illu s trat io n the pro bable erro r

“ ”

refers n o t t o A s o r B s fa ilu re to hit t he target b u t to o u r

' ’

1 T he q u es t io n m a y b e ra is e d as to w h th e er t h ere is not a b ett er m eas u re of

mar k s ma n s i
p th a n t h e h
a v era ge d ep a rt u re o f t he s h o ts f ro m t h e b u l l ‘
s -e e
y . Fo r
in s t a n c e, wit h t he bu ll ‘
s -e e as a c e n t er,
y we m a y d es c rib e c irc es t l h ro u
gh eac h of

t he s ho ts of A, an d c o n s t r u c t a c i rc e l h thwit e a v era ge a rea o f th es e c irc es l fo r

it s
'

a rea . h
T is c i rc e l m ay t hen be l td
s e ec e as a mea s u re of A '
s mar k s m an s h ip
i n s t ea d of l
t h e c irc e ab o v e di
s c u ss e d . The ra di us o f h
t is c irc e c a n l be o b ta n e i d
b y ta ki n
g the s
q u are root of t he m ea n s q u a re o f t he d i ev a t io n s f ro m t he b u ll ’
s

eye . It is not i mpo rta n t fo r us to di sc u s s h ere t h e l ti

re a ve m er it s of th es e t wo

ma ho d s of m ea s u rin g m a rk s m a n s h ip
we rec o gn ize t a t , b u t it is im po rt a n t t h at h
t he met o h d
exp ain e l d
in t h e t e x t is b a s e u p o n w a t we m a y we c a “
a v era ge d h ll ll
dev ia t io n fro m a n i ea
”
d l hl h
w i e t a t s u gges t e in t is fo o t n o te m ay we l b e ca l e
, d h l l d
s t and ar dd
ev ia tio n f ro m a n i ea l
”
d .
 Y
T PE A N D V A R I A BI L IT Y 443

ca lcu la t i on a s s u m ing t he c irc les to represent the d ev iat io n o f

in
eac h wh en o n ly a lim it ed n u m b e r o f s h o t s had been fi red
,
.

I t is th is d ev iat io n and n o t o u r pro bab le e rro r that is to b e

t ak en as expres s ing erro r in mark sm an s h ip fo r with an infin it e ,

n u m ber o f sh o t s o u r E wo u ld d is appea r b u t the erro r in mark s ,

m an s h ip o r the d ev iat io n o f A o r B wo u ld n ev er d isappea r W ith .

infin ity it wo u ld ha v e a n exact and fi x ed v a lu e with E 0 ,

.

I t is o b v io u s that t he d ev iat io n s ab o v e al lu d ed to a re d ev ia
t io n s fro m an id ea l ( t he c enter o f the target ) wh ich is c o m par
ab le to t he select io n type in p rac t i
c a l br eed ing . I t is o b v io u s too , ,

tha t th ese h ea v y c irc les represe nt

t he mean s o f a ll t he s h o t s fi red by
the t wo m ark s men b u t th ey d o n o t
,

represent th eir d is tr i b u tion T hat .

is t o say A fo r exa m p le m ight

, , ,

ha v e p u t m o s t o f his s h o t s o r all ,

o f th em fo r that m att er at a u n i ,

fo r m d is tanc e fro m t he b u ll s eye '

-
,

no ne h itt ing t he c ent er and no ne F 44 L t t h h y l i A d IG . . e e ea v n es an

go ing wild ; in o th er wo rd s his B p t t h l l t d m k

re res en e c a cu a e ar s

m h ip f A d B p ti l y ;
,
an s o an res ec ve
S hOOt m g m 1gh t hav e b een v ery th t h ligh t l i A d B
p pen e n es e an e

u n if o r m ,
b u t n eith er v ery goo d wil l p t th p b b l re res en e ro a e erro rs

n o r v ery bad o wing eith er to b ad

i th m p t i t h t li n A d e as s u on a n es an
,
B p t t l y t h m k mre res en ru e ar s an
m ark sm an s h i p o r to a bad ly S i ght ed h ip f A d B s o an

g u n. N o w i t is c o nc ei v ab le that

an o th er m ark s m an C s h o u ld s u cc eed in m ak ing an a v erag e

, ,

id ent ica l ith that o f A b u t in a v ery d ifferent m anner

w ,
— so me ,

o f t he S h o t s h itt ing t he b u ll s e e and s o m e g o ing wild T h

'
es e
y
-
.

t wo m en th en d o s h o o t ing o f an ent irely d ifferent c las s t he on e

, ,

fro m t he o ther that is m ak e a v ery d ifferent d is trib u t io n ev en

, ,

th ou gh th ey win the s am e m ea n I f n o w fro m th is m ean (rep re .

,

s ent ed by t he h ea v y c irc les ) we c alc u lat e s ta n dard d ev iat io n

,

with res pect to a ll t h e s h o t s fi red we th en ha v e a c o nc ept io n o f ,

d ev iat io n c o rres po nd ing exact ly t o that o f t he o rd inary s tandard

d ev iat io n ; na m ely d ev iat io n fro m the m ea n o f all t he v ariat es
, .

T h u s we illu s t ra t e bo th d ev iat io n fro m m ean a n d d ev iat io n fro m

an id eal t o geth er with t he pro bab le erro r in v olv ed
, .
 T R A NSM I SS I ON
44 4

SECT I ON V — COM PAR AT I V E T PE Y AN D V A R I A BI L IT Y

FOR D I FFE R E NT C HA R A CT ER S I N T H E SA M E
POPU LAT I ON

If a v ariety of charact ers in the sa m e po p u lat io n b e c rit ica lly

s t u d ied it will b e f o u nd that each ha s it s o wn type a n d v ar iab ility

Fo r exa m p le i n t he p o p u lat io n a ris ing fro m t he ten inch ear s

,
-

already d isc u s sed it was fo u nd that o th er charact ers v a ried as

fo llo ws
T Y PE A ND V AR IA B ILIT Y FOR FOU R C HARACT E RS OF C OR N G R OWN FR OM

SEED EARS T EN I NC HES LON G

M E A N I N C H ES
STA N DAR D D av u Co n n lc u mr
‘
or

I N C H ES VA RlA mu
U
, ' '
1 10N , n
' ‘
OR .
O PE R C EN T
OR N C ES
Ou N c n s

Le n g t h o f ea r f 1
C i rc u m f eren c e o f ear j; 1
W eigh t o f ea r i i
N u m b er o f ro ws i }
z;

F ro m th is we see that each s eparat e charact er tak es its o wn

ty pe and v ar iab ility F o r exa m p le c o rn is m u ch mo re v a ria b le
.
,

as t o weigh t p e c ent ) than as t o a th er cha ract e r

r n
y o

mea s u red Th is is t o b e exp ect ed b eca u s e weight is t o s o me

.
,

ext ent t he res u ltant o f b o th len g th and c irc u m f erenc e and wo u ld

th u s partak e o f the v a r iab ility o f b o th B u t we n o te a ls o th a t .

th is c o rn at lea s t was m u ch m o re v ariable a s to length than a s to

c irc u m ferenc e p e r c e nt a s c o m p a r ed w i th p er c ent ) .

A ga in we n o t e that th es e ear s a re m u ch m o re v ariab le a s to

n u m ber o f ro ws than as to c i rc u m ferenc e by wh ich we inf er ,

that the width o f t he k ernels is far fro m u n ifo r m else t he t wo ,

wo u ld mo v e t o geth er .

T h is ra is es t he wh o le q u es t io n o f t he relat io n o r b o nd b et ween
d ifferent chara ct ers a s u bj ect to b e d is c u s sed in t he su cc eed in g
,

ch a pt er o n Co rrelat io n I t is s u ffi c ient h ere to remark that

.

t he typ ica l ea r o f th is cro p o f c o rn is inch es lo ng ,

inch es in c irc u m fe renc e ha s ,

ro ws o f k ern els and weigh s ,

o u nc es I f we sh o u ld p ick su ch an c a r it c o u ld s tand a s
.
 TYPE A N D V A R I A BI LI I Y
' ‘

44 7

SECT I ON V I I — I N D I R ECT EFFEC TS O F S ELECT I O N U PON

TY PE A N D V A R I A BI LIT Y

I n t he tab le j u s t giv en we n o t ed o n ly the d irect effect s u po n

t he chara ct er u nd ergo ing s elect io n I t n o w remain s t o c o n s id er .

ho w rig id select io n o f o n e charact er may a ffect o th er and p re

s u m ab l
y c o rrelat ed charact ers T o th i s en d we c o n s t ru ct a
.

ta b le s h o wing t he phy s ical charact er s o f t he fo u r s t ra in s o f

c o rn n o w u nd er d isc u s s io n remem bering that th ese s tra in s a ll
,

d ev elo ped fro m t he s am e o riginal s t ock and that select io n was

c o n fined t o t he ch em ical charact ers pro t ein and o il lea v ing the , ,

phys ica l and phy s io lo g ica l charact ers free to take care o f
them selv es .

I N D 1R ECT EFFECTS R ES U LTS

’
OF SELECT ION : OF SEV EN Y EA Rs SELEC
T 1O N FOR C HE M I CA L C OM POS IT IO N U PON PHv s1CA 1 . C HA RA CT ERS .

Au . FO U R ST RA 1 Ns D E V ELOPE D FR O M T HE SAM E OR IG I NA L STOC K l

LE NG T H 0 1? E A 11 C 1 RC U M R E R E NC E

V A R 1 s Tv

M ea n S ta n d D ev . . M ea n S tan d D e v
. . Co ef V ar
. .

at : a: = 1= t

a; a: a: a:

= 1= a: a: a:
1 .
39 t

a: 1 . 3 0 a: a: a: a:

N UM B R E OR R o ws W 11 10 HT OP EA R S

V A R w Tv
I ! 12

Mean Stan d D ev . . Co ef V a r
. . Stan d D ev
. . Coef V ar. .

a 1 . 85 a: a: 04 a: a:

= 1= a: -
1= 10 a. a:

2 .08 z 0 04 . 07

a: x a: 08 z t

Discu ssion of cr it ic a l s t u dy o f th is tab le rev eals

data . A
so me s ign ificant fact s c o nc ern ing t he ind irect effect s o f select io n

S ee B u l leti n N o . 1 1 9, U n i v ers it y of ll
I in o is A gri c u ltu ra l Experim en t S ta tio n .
44 8 TR A NSM I SSI ON

u po n cha ract ers o th er tha n th o s e u nd er s pec ial c o n s id eratio n .

S u ch a c ri t ica l s t u dy d ev elo p s the fact that th ese fo u r s t rai n s

d iffering in pro tein and in o il c o ntent ha v e d ev elo ped also int o
fo u r d is t inct s train s regard ing t he p u rely phys ic al cha ract ers ,

length c i rc u m feren c e etc

, ,
.

1. Higlz a n d lo w p rotein T he ea r o f t he fo r mer is t he

‘

s h o rt er ( c o lu m n I ) and t he s m a ller s tandard d ev iat io n and c o

,

effic ient o f v ar iab ility S h o w i t t o b e les s v ariab le as t o length

( c o lu m n s 2 and I t is a ls o s m a ller in c i rc u m f erenc e

( c o l
u mn with a les s n u m b er o f ro ws ( c o lu m n and light er in
weight (c o lu m n I t is a lso les s v ariable in ev ery res pect ,

bo th relat iv ely (c olu m n s 2 5 8 and I I ) and absolu t ely (c ol , , ,

u m u s 3 6 9 and , , ,

2 .s g/z a nd low oi l I n t h e s a m e m ann er we learn that t he .

h igh o il ea r is a s h o rt er and a s m aller ea r than the lo w o il b u t

- -
,

that i t is more v a r ia ble as t o length a n d les s v ariable as to

c irc u m ferenc e (c olu m n s 1 to T he lo w o il ea r is rap id ly b ec o m -

in g a t welv e ro wed v ariety wi th a lo wer d ev iat io n than any

-
,

o th er ( c o lu m n I t is t he h ea v ies t th o u gh n o t t he lo nges t o f , ,

t h e f ou r select ed s tra in s .

3 .T li e f ou r s el ec t ed s t ra i ns O f th e s e f o u r s t ra in s d ev elo p e d .

fro m t he sa me o rig in a l s t o ck t he lo w pro t ein ea r is the lo nges t ,

-

and the h igh o il t he sh o rtes t The h igh o il is also the mo s t

-
.
-

v ar iab le as to length c o lu m n
(
T he lo w o il is t he larges t and t he h igh pro t ein is t he s malles t
- -

in c i rc u m ferenc e T he latt er is als o t he lea s t v ariab le and t he

.

lo w pro tein is t he m o s t v a riab le a s to c irc u m ferenc e

-
.

T he h igh o il c o rn has t he larges t n u m be r o f ro ws with t he

-
,

lo wes t v ariab ility and t he lo w o il t he fewes t ro ws with t he

,
-
,

lea s t s tandard d ev iat io n b u t t he great es t c oeffi c ient o f v ariab ility .

T he lo w o il is t he h ea v ies t and t he h igh pro t ein t he light es t

- -

ea r b u t t he lo w pro t ein is t he m o s t v a riab le a s t o weight

,
-
.

I t wi ll be n oted tha t t/zc s e

'
tlze n a tu ra l r es u lts
'

a ifl c r c nc es a r e
of
s elec t in
g ,
n ot f or Ma c p a r tic u la r clza r ac ter s , bu t f or c/cem ica l

con ten t . T h ey
th erefo re t he ind irect effect s o f select io n
a re ,

and b ring u p aga in t he wh ole s u bj ect of c o rrelat io n which will ,

b e treat ed f u rth er i n a lat er chapt er .

TY PE A N D V A R I A BI LI TY 45 1

m
-
0.
0
H H
m c
H H H H H 0 0
w
u
3 S m
m
a
n
«
m m 3 n m
o 0 0 0 0 0 0

n m
« u
0
.
o
0 d
H H
v
H H H H H m m
e 0 m 3
m m
n

d 0
.

0
.
3
0

0 0 0
n
7 n
d 0 d
H H H H H H
0 « « « u m
9 Q m
.
G c. w
m n d
9 0
. ? a u

w « 0
n m m
o o 0.
d d 0
H
0
H H H 0
H H H H H
m 0
9 i
h
u 0 o m
3 m ? 6
.
a
.
n
.
?

m v w « 0A .

m 9
o 0.
o d 0
H
u
H
-
H
o
H H H H H H H
m m « m
t v

m
.

m n . m w w w
« ? o
.
0
.
.
.
0. o o
d
.

& w w fi
.

u “ “ “ u

°P° N
45 2 T R A N SM I SSI ON

S PE C I A L E x E R C I S E s

T h e s tu d en t s h ou ld hav e ex ten s iv e in m a in g freq u en c d is tri

p r a c t i c e k y
k
b u tio n s o f d iff eren t s pec ies a n d i n wo r in g o u t s ta n d a rd d ev ia tio ns an d
c oefl i c ien ts o f v a ri a b ili t y . y
He will th u s n o t o n l b ec o me fam ilia r wi th the
m ethod s of wo r k b u t he will
, a c q u ire n ew c o n c ep tio n s o f th e wh o le j
s u b ec t

o f v ar iab ili t y a n d type The . h

tea c er s ho u ld req u ire t he s tu d en t at t ish
po i n t t o m a k e o igi n al s tu d ie
r s on his o wn a c c o u n t , an d to pro sec u te the
wo r k u n ti l he b e c o m es en ti rely fa m ilia r b o th wi th the m ethods an d wi th the
c o n c ep tio n s i n v olv ed .

AD DITIONAL REF ERENCES

T HE G R A M M A R O P S C I E N C E . By K a rl Pea rso n . C h a p ter X .

V A R I A T ION IN A N IMA LS A ND PL A NT S . By H M V ern o n

. . . C h ap ter I .
 54 T R A N S M I SS I O N

pres enc e o f s u n light and the fix ing o f carbo n ; and by su ch

o th er r elat io n s a s a re in v o lv ed in d i rect ca u s a t io n .

On t he o th er hand s u ch a relat io n as d eafn es s a mo ng t eleg

,

ra pher s o r b lindn es s a m o ng c iv il engi n eer s o r lo c o mo t iv e d ri v er s

is u nkn o wn b eca u s e t he c o nd it io n s a re s u ch that t he chara ct ers

,

in q u es t io n a re m u t u ally exc lu s iv e .

I n gen era l h o wev er c o rrelat io n fa lls s o mewh ere b et ween

, , I

and u n ity and o n o n e s id e o r the o th er o f t he zero po int that is

, ,

a d egree of relat io n sh ip ex is t s wh ich is n eith er ab so lu t e d en o t ,

in g d i rect ca us at io n n o r n egat iv e s ign ify ing m u t u a l exc lu s io n

, ,
.

Fo r exa m ple a h igh d egree o f c o rrelat io n ex is t s bet ween length

,

o f c o b and weight o f c a r I t d oes n o t a m o u nt to u nity h o wev er

.
, ,

fo r t he c irc u m ferenc e a lso c o ntri b u t es t o weight .

Mo s t res u lt s in liv ing o rgan is m s a re the effect o f m ix ed

ca u ses and fo r th is reas o n c o rrelatio n s a re mo re c o m p licat ed
,

tha n may at fi rs t appear Fo r exa m ple many if n o t mos t goo d

.
, , ,

c o ws hav e a capac io u s barrel and a roo m y u dd er and men

“
,

hav e been led t o a ss u m e a perf ect c o rrelat io n bet ween th ese

s p ec ia l charact er s and m ilk pro d u ct io n ; wh erea s t he tru th is

that t he c o rrelat io n th o u gh h igh is s o meth ing les s than u n ity

, , ,

b eca u s e go od c o ws a re k no wn with s ma ll barrels a n d with in c o n

s p ic u o u s u dd ers Here is a real n eed fo r acc u rat e meth o d s of
.

d et erm in ing what d egree o f c o rrelat io n act u a lly ex is t s T he .

av erage man a s k s wh eth er o r n o t two charact ers a re c o rr elat ed ,

and expect s a pos it iv e an s wer Y es o r N 0 ; wh erea s t he q u es t io n

s h o u ld b e T o what ext ent d o t he t wo cha ract er s app ear to geth er ?
,

exp ect ing fo r an an s wer a fract io n ly ing so m ewh ere b et ween

zero and u n i t s ay p erhap s 4 0 to 60 p er c ent as in c o rrelat ion

y , ,

o f length t o weight o f ear .

The s t u d ent m u s t d is t ingu is h clear ly bet ween c o rrelatio n and

mere a s s oc iat io n Fo r exa m p le we m ig ht a s k t he q u es t io n wh eth er
.
,

black p ig s a re m o re s u bj ect to ch o lera than a re p igs o f o th er

c o lo rs T he fi rs t s t ep wo u ld b e t o es tablis h a rat io bet ween the
.

n u m ber o f d is ea sed p ig s and p igs in gen era l T h is rat io wou ld .

n o w expres s t he chanc es that a par t ic u lar pig irres p ect iv e o f ,

c olo r will b e a flIic t ed with th is d iseas e

,
‘
that is by that o pera
, ,

tio n o f ind epend ent pro bab ility wh ich we ca ll chanc e If n ow .

we find u po n inq u i ry that u nd er t he s a me c o nd it io n s t he rat io

 C O R R ELA T I ON 45 5

of C holera s u bj ect s to b lack pigs is h igh er than the ra t io to pigs

in gen eral th en we s h o u ld c o nc lu d e that an act u a l pos it iv e c o rre
,

lat io n ex is t s bet ween t he black c o lo r and th is part ic u lar d isea se .

On the o th er hand if th is rat io s h o u ld b e b elo w t he rat io o f

,

p igs in general th en we s h ou ld c o nc lu d e that b lack p igs a re

,

les s s u s c ept ib le t o th is d is ea s e than a re p ig s o f o th er c o lo r s and ,

that a n egat iv e c o rrelat io n ex is t s as s u ming a lways eq u al oppo r ,

t u n it ies for i nf ect io n T h is is t h e o n ly c o r rect m eth o d of s t u dy

.
,

and it wou ld n o t b e s afe to c o nc lu d e that black p igs a re pec u l

ia rly s u s c ept ible s im p ly beca u se m o s t of t he p igs that d ied u n d er
o u r o b s erv at io n happen ed t o b e b lack fo r b lack p ig s a re m o r e ,

n u mero u s in t he cho lera b elt than a ll o th ers c o mb in ed and ,

u nd er pr o bab ility a lo n e th ei r ab s o lu t e m o rta lity m u s t b e h igh er .

Wh en expres s ed in the fo rm o f a rat io h o wev er the tru th c o mes , ,

t o t he s u rfac e 1
.

S im ilar ly we m ay as k the q u es t io n wh eth er d iff erent s pec ies

o f p lant s o r an im als t end t o att ra ct o r r epel each o th er wh en

thro wn to geth er in the s ame terr it o ry Here aga in the fi rs t .

s t ep is t o fi n d t he ra t io o f a s so c iat io n u nd er free Op era t io n o f

i nd epend ent pro bab ility a rat io based o n the relat iv e n u m bers
,

o f in d iv i d u als o f t he s p ec ies in q u es t io n and t he ext ent o f t erri

t o ry fi rs t wh ere n o o ppo rt u ni ty fo r as so c iat io n is po s s ib le and

, ,

S ec o nd wh ere s u ch a ssoc iat io n is po s s ib le I f the t wo rat ios

,
.

d iffer th en we infer that so m e d egree o f c o rrelat io n ex is t s

,
.

S ECT I O N I I — C A LCU LAT I O N O F CO EFFI C I ENTS O F

C OR R E LAT I O N

Wh en t he presenc e o r absenc e o f the chara ct ers in q u es t io n

is ab so lu te as red o r b lack ha ir pres en c e o r ab s enc e o f h o rn s
, , ,

th en the c o rrela t io n is expressed by a s ingle rat io as we ha v e ,

s een B u t m os t ca s es a re n o t o f th is extrem e s im p lic ity ; fo r

.

exa m p le it is s a id that wh it e cat s a re d eaf

,
I f n o w a ll wh it e .

cat s are d eaf th en t he c o rrelat io n bet ween alb in is m and los s o f

,

h eari ng power is ab so lu t e and is expressed by t he c oeffic ient I ,

.

1 Per h a ps it oug ht to he rem a r k d th t th i ill

e a s u s t ra t io n is ta ke n p u rel y at

ra n d o m , as n o s t u di hes av e b een m d
a t
e as th l
o e re a tio n b etween c o lo r an d t h is
p a rtic u la r di s eas e.
 6 TR A N SM ISS I O N
45

S u ppo s e, h o wev er , that o u t cats tak en at ra nd o m

of 1 000

2 0 a re wh i t e , 1 0 a re d ea f and 6 are bo th white and d ea f

,
.

I s th ere c o rrela t io n ? N o w acc o rd ing t o th is a s s u m pt io n the

, ,

pro bability o f a cat being d ea f with o u t res pect to c o lo r is

10 4 -
1 000 or I
,
t o 1 00 ; b u t t he pro bab ility o f a whi t e c a t
bei ng al so d eaf a m o u nt s to 5 2 0 o r 1 S ho wing a h igh c o rr e , ,

lat io n bet ween alb in is m and d eaf nes s .

B u t to d eri v e an e xact expres s io n fo r th is c o r relat io n is no t

so s im p le as it m ight seem A cc o rd ing to t he c o nd it io n s wh ich
.

we ha v e a lrea dy la id d o wn and i n c o n s is ten c ywit h o th er ph as es

,

o f t he pro b lem o f c o rr elat io n in gen eral a n y express io n wh ich ,

we m ay ad o pt as an efli c ien t m eas u re o f t h is c o rrelat io n s h o u ld

b e s u ch a fo r m u la as will b ec o m e zero wh en t he t wo cha ract ers
are ind i ff eren t to each o th er ; will bec o m e I wh en t he t wo m o v e

t o gether perfect ly ; and will bec o me I when th ey are m u t u ally

exc lu s i v e .

Y u le has giv en an elegant meas u re o f th is a sso c iat io n o r

l
,

c o rrelatio n wh ich sat is fi es th ese

,

c o nd it io n s To d ev elo p th is fo r
.
Cm N W . or 11 1 "

m u la he arra nges t he po p u lat io n

as in t he acc o m pany in g d iag ra m D f ea

with res pect to t he cha ract ers

N t d f
in q u es t io n ( d eafnes s a n d c o lo r )
o ea
.

Th en the m ea s u re o f a ss o c i
at io n bet ween d ea fn es s and t he wh it e c o lo r is expres s ed by

In general if we hav e a po pu
,

M Pm “ M Am “
lat io n arranged with ref erenc e to ?

t he pres enc e o r ab s enc e o f t wo

N P R ES E N T
cha ract ers M and N in n u m
, ,

b ers a b c d t he arra ngement

, , , ,
N A B S EN T
wo u ld s tand t he s a m e a s ab o v e ,

(Id be
and t he fo r m u la wo u ld b e . If care b e tak en to a rra n ge
ad bc
the tab le so that oc s ha ll be n u merica lly les s than ad, then the

1 P il ilo sop nica l Tr a n s a c tio R oy a l S oc i ety . C X I V ,

ns o
f t/1 c 2
5 3 9
7 1-
.
45 8 T R A NSM I SSI ON

SECTI ON I I I —THE C OR R E LA T I O N TA B LE

To d et erm ine t he d egree o f c o rrelat io n bet ween any t wo

charact ers in any rac e a so c a lled c o rrelat io n table is c o n
,
-
"

s t ru ct ed o u t o f t he mea s u rem ent s o f t he t wo cha ract ers a s f o u nd

in a large n u m b er o f i nd iv id u a ls o n e charact er being rec o rd ed

,

i n c o lu m n s and t he o th er in ro ws T wo rec o rd s are th u s m ad e

.

o f t he s a m e i nd iv id u al o n e fo r each charact er
,
S u ch a table .
,

wh en fi n is h ed c o n s is t s o f a d o u b le sy s t em o f arra y s each
, ,

d epend ent on t he o th er and f ro m wh ose m eans and s tandard

,

d ev iat io n s the m u t u al relat io ns h ip s can read ily b e wo rk ed o u t .

K n o wing th is relat io n s h ip and t he v alu e o f o n e o f t he char

ac ters we a re enab led t o c alc u lat e t he c or res po nd ing m ea n v alu e

o f t he o th e r . T he ad v antages o f th is fo r p u rpo s es o f select io n are

o b v io u s
. T he m eth od is bes t illu s t rat ed by an act u a l exa m ple .

Fo r in s tanc e i t is ev id ent that t he weight of c a rs in c o rn d e

,

pend s part ly u po n th eir length and part ly u po n th eir c irc u m fer

enc e . To what ext ent fo r exa m p le d oes it d epend u po n lengt h
, ,

I n o rd er to an s wer th is q u es t io n d efi n it ely a large n u m ber of

ear s tak en at rand o m a re bo th weigh ed and m ea s u red and t he ,

data a re arranged in tab u lar f o rm as d es c ribed abo v e appearing ,

as f o llo ws

COR RELAT IO N BET W EEN WE IG HT AND LE NGT H or EAR

E A M I N G COR N )
( L

WE I G HT OP E AR S IN Ot '
N CE s

7 8 9 10
 C OR R E LA TI ON 4 59

Ar rays of th is ta ble each ea r is

a correlati on ta b le . In
rec o rd ed in t he pro p er s q u are t o represen t bo th it s weight and
i ts length T his being the ca se all t he ear s o f t he sa m e weight
.
,

that a re a ls o of the sa m e leng tlz are rec o rded t o gethe r in the

s a m e s q u are T h is mean s that t he v ario u s r ows a re freq u ency
.

d is trib u t io n s o f weight wit/z respec t to leng tlt (as 1 6 I I 2 6 , , , ,

I I, 8 6 I t he freq u ency d is t ri b u t io n c o rres po nd ing to t he

, , ,

length inch es ) and all t he colu m ns are freq u ency d is trib u t io n s

,

o f length wit h res pect to weight S u ch freq u ency d is tri b u tio n s .

wi th res pect to a c orrela ted cha ract er a re techn ica lly kn o wn as

“
ar ra y s The enti re ta b le therefo re may b e loo k ed u po n as
.
, ,

m ad e u p of t wo s y s tem s o f pa ra llel ar ra y s with res pect t o the

t wo characters in q u es t io n Tli cy a r e i n n o r esp ec t d ifieren t f r om
.

a ny o tb er f r eq u ency d is tr i b u tion s a nd tlzei r m ea n s , s ta nd a r d

dev ia t ions , v a r ia b i l i ty , a nd o tlzer d eter m i n a tions a r e ca lcu la ted

by tlze sa m e m et/rod s as g i v en i n tb c la s t c lzap ter .

SECT I ON I V —T H E C O R R E LAT I O N COE FFI C I ENT

A m ere in s pect io n c o rrelat io n table j u s t giv en s u gges t s

of t he
that in gen eral S h o rt ears a re light ears and that lo ng ears a re
, , ,

h eav y ears ; b u t what we s eek is a s tat is t ical c o n s tant wh ich

will b e a meas u re o f th is c o rrelat io n and wh ich ind icat es to ,

what extent t he weight o f c a rs can b e pred ic t ed fro m th eir

length s T he c oefi c ien t of corr ela t ion is s u ch a c o n s tant and
.
,

wh en d et er m in ed i t will b e d eno t ed by r .

A d is c u ss io n o f the m ath em at ica l th eo ry o f c o rrelat io n will

b e g iv en in the A p pend ix b u t it S h o u ld b e s a id h ere as bef o re
, , ,

that the c oeffi c ient alway s tak es s o m e v a lu e bet ween I and

—I I f r + 1 th ere is s a id to b e perfect po s it iv e c o rrela
.
=

t io n ; that is the t wo charact ers a re ca u s ally c o nnect ed I f

,
.

r — 1 there is pe rfect n egat iv e c o rrelat io n ; that is th ey ,

a re m u t u a lly exc lu s i v e I f n o c o rrelat io n ex is t s r . o in d ic a t ,

=
,

in g t he t wo charact ers as b eing ind i fferent to each o th er and

mo v i ng ind epend ent ly I n nearly all ca s es so m e act u a l c o rr ela
.

t io n ex is t s and in a general way we m ay s ay that t he c o rrela

, , ,

t io n s h o u ld b e j u dged by the v alue wh ich r tak es bet ween zero

and u ni ty .
 6 0 T R A NSM I SS I ON
4

Meth od of fin din g correlati on c oefii cien t . The m eth od o f c a]

c u lat in g t he c o rrelat io n c o efli c ien t ( r ) is exh ibit ed in c o nn ect io n

wi th t he tab le o n page 4 6 1 s h o wi ng a repres entat iv e ca s e fo r , .
,

c o n v en ienc e c o nt in u ing t he s a me c o r relat io n table a lready c o n

s tr u ct ed T h o u gh t he m eth od is so m ewhat c o m p licat ed it is
.

giv en in f u ll .

I t is h igh ly im po rtant t o get at o nc e a gen era l c o nc ept io n o f

th is table and o f the m eth od o f pro c ed u re A ll t he c om p u ta .

t io n s s h o wn exc ept th o se in v o lv ed in t he c o lu m n h ead ed SP

ha v e to do on ly with fi nd ing tlte m ea n s a n d s ta nd a rd d ev ia tions
o f t he po p u lat io n wi th res pect t o t he t wo charact er s i n q u es t io n ,

acc o rd ing to the meth od f u lly treat ed in t he la s t chapt er ; that

is t o s ay t he c o lu m n s o f fig u res h ead ed f “ f V
,
l
D D} , L, L, ,

f D
L
z
L ,
D w D }
a re al l s elf,e xp lanat ou
ry t o,
-

any o n e fa m iliar with the m ean ing o f o rd inary algebraic sy m bo ls ,

a n d who kn o ws ho w t o find t he v ar iab ility o f a po p u lat io n by t he

meth o d s already gi v en .

T h ere r em a in s the c o lu m n o f figu res h ead ed SP wh ich it ,

s eem s wo rth wh ile t o exp la in in d eta il and wh ich is t he o n ly

s p ec ia l f eat u r e in t he d et er m inat io n o f t he c o rrelati o n c o effi c ient .

Ea ch n u m ber in th is c o lu m n repres ent s t he s u m o f t he prod u ct s

o f t he c o rres po nd ing length and weight d ev iat io n s fo r ev ery

i nd iv id u al in the h o rizo nt a l arra y to wh ich t he n u m b er belo ng s .

T o s h o w ho w th es e n u m b ers a re c o m p u ted s elect fo r exa m ple , , ,

t he h o rizo nta l ar ray m ark ed 1 0 and we s h all s h o w ho w t o fi nd ,

t he n u m b er
I n th is ca s e
1 0 t he m ea n , DL o f ro w 1 0 .

T h en
I 3 8 ( 2 3) 1 8
6 ( 5 -3 ) 4 (M ) 2
(7 43 1 -0

A llth er n u m bers o f the c olu m n h ead ed S P a re fo u nd in the

o

s a m e way and t he t o ta l is wri tt en s y m b o lica lly a s

,

R d f
1 ea b m i g th f q
su Ir , y f w igh t ;
”
ea n n b V b W m e re u en c o e s su 11
’
su , ean

i g th f q
n e y f w igh t m l tip li d b y t h l i w igh t ; f b
re u en c o e s u m e e v a ue n e s
“
su ea n

i g th f q
n e rey f l gt h t
u en c o en , e c .

Th9 l ig ifi
e rea f EP i b t h w b y t h
s n ca n c e o p i Z D D W t hst es s o n e ex ress o n L , a

i th
s, m f th p d t
e su o f b t h d i ti
e ro f ll t h i d i id
uc s o l i th t b l
o ev a o ns o a e n v ua s n e a e.

I t i w it t
s r i i
en w y b t lw y wit h th
n v ar o u s b m
a s, i g u a a s e a ov e ea n n .
4 6 2 T R A N SM I SSI O N

T he v a lu e o f r is f ou nd by m ean s of t he fo r m u la
”
2D
7
flo u ”
r ;

S yst emati c ar ran gemen t of proc es s wh ich work . T he wh o le ,

s eem s s o m ewhat c o m p licat ed is aft er all q u ite s im p le To , .

recapit u lat e it a mo u nt s to m u ltiply ing the fig u res in each s q u are

,

by bo th th eir o wn d ev iat io ns ( that is by their d ev iat io n a s to ,

length and th ei r d ev iat io n a s to weight ) and th en add ing a ll t he ,

res u lt s and d iv id ing by t he wh o le n u mber ( o f ears ) m u lt iplied

by t he prod u ct o f t he t wo s tandard d ev iat io n s ( See fo r m u la .

S D ‘D "f

j In perf o r ming t he act u al wo rk , h o wev er it is

-

,
n a Lo
'

h igh ly im po rt ant to hav e a s y s t emat ic s ch eme fo r carrying o u t

t he c o m p u ta t io n s i n o rd er to a v o id c o nf u s io n in the so m ewhat
c o mplicat ed d eta ils I t has s eem ed d es irable th erefo re to
.
, ,

present t he m att er in t he fo r m o f a d etailed d es cr iptio n o f t he

v a rio u s s t ep s i n v o lv ed .

Fi rs t s tep Ha v ing giv en t he c o rr elat io n tab le o f t he po p u la

.

t io n we fi rs t add t he freq u enc ies in the arra y s with res pect to

,

b o th charact ers ; that is add the n u m bers in c o lu m n s and ro ws ,

o f t he tab le T h is gi v es t wo freq u ency d is tri b u t io n s o f the t o ta l

.

po pu lat io n t he o n e wi th res pect t o length o f ears ( f )

,
and the ,

o th er with res p ect t o weight o f ear s

f
T he o n e with res pect t o lengt h has the freq u enc ies 4 5 I 4 , , ,

T he one with res pect to weight ha s the freq u enc ies 4 ,

22 ,
2 7,
50 , 47, 7 1 , 7 5, 7 1 » 75 8 8 ,, 6 5, 37, 8, I 3, 4.
S ec on d s te
p . Fo r ach
e of these freq u ency d is tri b u t io n s
( c olu m n f 1 ,
and ro w f t he m ea ns a nd tb c s ta n d a r d d ev ia t ions
m u s t be c a lc u la ted . T he m eth o d
th ese ca lc u lat ion s of m ak ing
is t he s a m e a s t he o n e u s ed fo r m ean and s tandard d ev iat io n s
in gen eral I t ha s al ready been f u lly exp la in ed and th erefo re
.
,

n eed n o t b e repeat ed h ere A s y s t em a t ic arra ngem ent o f the .

wo rk is s h o wn in c o n n ect io n with t he t a b le T he r es u lt s a re .

m ean len gt h M L

st an d a rd d e ia t io n in len gt h v 01 .

m ea n weight bi ”
.

stan d ard d e iat ion i n weight 3 v a " . .

 COR R ELATI ON 6
4 3

l ird ps te . T h is is t he
n ly par t o f tohe w o rk that is r ea lly
n ew . I n d o ing t he sec o nd s t ep we fou nd t he d ev iat io n s o f
t he c las ses fro m t he m ean length and m ean weight T h ese a re .

rec o rd ed u nd er D and D ,
F o r exa m p le in ro w I we fi nd
”
.
,

that 4 ears were each 3 inches lo ng ; that is to say th ey ,

d ev iat ed inch es fro m t he m ean length t ak en at in

s t ead o f 5 in o rd e r t o s a v e lab o r W e n ext tak e t h e n u
. m
b er in each s q u are o f the c o rrelat io n tab le and m u lt ip ly it by
the c o rres po nd i ng d ev iat io n s b o th in weight and in length ,

thereby sec u ring a prod u ct wh ich is t he res u lt of the f u ll n u m

b er o f v ariat es in v olv ed and of th eir d ev iat io n in res pect t o
bo th chara ct ers .

Fo r exa m p le wh ere the c o lu m n h ea d ed 9 ou nc es and t he

,

ro w labeled 6 5 inch es cro ss each o th er o cc u r s t he n u m b er

.

8 wh ich ind icat es that 8 ears o f t he po p u lat io n weigh ed 9

,

o u nc es and were inches lo ng I n o th er wo rd s these 8

.
,

ear s each d ev iat e I 7 o u nc es ( ro w D W c o lu m n 9) fro m mean

.
,

weight o f ear s and inch es fro m t he m ean length (c o lu m n

,

D ro w Henc e fo r th is n u m ber 8 we fo rm t he p rod u ct

8( W ith o u t regard to labo r we S h o u ld ,

find s u ch a prod u ct fo r ea clz n u mber in tlze c or r ela ti on ta b le .

I f no w a ll th es e prod u ct s b e add ed and the res u lt d iv id ed by

t he prod u ct o f t he t wo s tandard d ev iat io n s i nt o t he n u m ber o f
v ariat es in t he t o ta l po p u lat io n we s ha ll o bta in the c o rrelat io n
,

c oeffi c ient o r the i nd ex o f c o rrelat io n wh ich we seek

,
.

T he s y s t em at ic way o f car ry ing o u t th is wo rk is t o rec o rd t he

res u lt s o f th is o perat io n fo r each h o rizo nt al array u nd er t he
h ead ing S P and then add th ese res u lt s fo r the array s to o btain
,

t he fi na l res u lt wh ich is s y m bolica lly ind icat ed by

ED D W o r s u m m at io n D D W
L , L .

T o illu s t rat e the meth od o f c a lc u la t io n a few o f t he pro d u ct s

rec o rd ed in c olu m n EP will b e s h o wn .

F o r exa m p le with a s t he m ean length and

, as t h e

m ean weight we ha v e fo r t he array c o rres po nd ing t o length

,

4 i nch e s ,
x 3
7 7 x 5
—3 8 h
.

. x —6 7
. x 5
T is g iv es

s7
. x 1
 6
4 4 TR A N SM I SSI ON

F o r t he array c o rres p o nd ing to 5 i nch es ,

x 2

x 4
x x 7 h
T is giv es
x 2

x 4

T reat array s in a s imilar m ann er and fi na lly d iv id e the

all , , ,

su m o f all t he pro d u ct s th u s o bt ai n ed ( that i s by t he

prod u ct o f t he t wo s tandard d ev iat io n s a n d the n u m ber of
v ari at es b ei n r e u l a l wa s to l l l d i i i
, g c a f y p r es er v e t ze f u s t nc t on a s

to p lu s a n d m i n u s s ig n s T h is g iv es
.

49 7
4 -2
t he r
c o r ela tio n c oeffi c ien t .

993 ( 1 - 5 7)
T he math em atica l d eriv at io n of th is c oeffi c ient as a meas u re
o f c o rrelat io n in v o lv es t oo m u ch m ath em at ic s t o b e g iv en h er e .

I t m ay b e n o t ic ed fro m t he c o m mo n s en s e s tandpo int h o wev er

-
, ,

that it seem s to b e a goo d meas u re o f c o rrelat io n To app re .

c ia te t he mean ing o f th is c oeffi c ient it s h o u ld b e recalled that

,

we tak e the pro d u ct s o f b o th d ev iat io n s fo r ev ery in d iv id u al in

t he tab le add th es e pro d u cts
, and d iv id e the res u lt by the
,

n u m ber o f ind iv id u als T h is giv es the a v erage o f all the

.

prod u ct s o f bo th d ev iat io n s We th en d iv id e th is a v erag e

.

p r od u c t o f t h e i n d iv id u a l d ev iat io n s by t he pr o d u ct o f t h e t wo

s ta n d a r d d ev ia ti ons th u s S ec u r ing an expres s io n wh o s e v alu e is

,

infl u enc ed by t he d ev iat io n o f b o th charact ers with ref erenc e

each t o t he o th er .

I t req u i res b u t litt le m ath em at ical in s igh t to s ee that if t he

c o rrelat io n is po s it iv e and c o ns id era b le po s it iv e v alu es o f the
,

t wo charact er s c o rres po nd and n egat iv e v a lu es c o rres p o nd and

f u rth er that all t he prod u ct s o f d ev iat io n s a re po s it iv e T h is ’

.
,

m ak es fo r a large c o rrelat io n c oeffi c ient On t he o th er hand if

.
,

no c o rr elat io n ex is t s fo r any v a lu e o f o n e charact er we may

,

expect in t he lo ng ru n eq u a l and o pp o s i t e d ev iat io n s o f t he

o th er charact er wh ich m ak es t he s u m o f pro d u ct s o f d ev iat io n s

,

v ery s m a ll T h is c o m m o n sen se ex a m inat io n ind icat es t he real

.
-

nat u re o f the c o rrelat io n c oeffic ient .

Fou r tlz s tep F ind the pro ba b le erro rs in t he d et erm in ed v alu es

. .

T h o s e in t he mean s and s tandard d ev iat io n s a re c o m p u t ed by

4 6 6 TR A NSM I SSI ON

b efo re by t he prod u ct of the n u m ber of v a riat es and the t wo

s tandard d ev iat io n s Fina lly we su bt ract fro m th is res u lt t he
.

pro d u ct s of t he t wo c o rrect io n s to o u r g u es ses in find ing m ean s ,

aft er d iv id ing that pro d u ct by t he pro d u ct o f t he s tandard d ev i

a t io n s o f t he t wo sy s t em s o f v ar iat es .

We s ha ll pres ent o n p age 4 6 7 an illu s t rat io n o f th is S h o rt er

m eth o d u s ing fo r the p u rp o s e t he c o rrelat io n bet ween length
,

and c irc u m ferenc e o f c a rs o f L ea ming c o rn I n th is G and C .

,
.

a re t he g u es s es at t he c la ss m ark n eares t t o t h e m ea n o f t he

po p u lat io n as to length a n d c irc u m ferenc e res pect iv ely .

L et M and M b e t he m ea n length and c irc u m f erenc e res pec

L C

t iv ely and C and C t he c o rrect io n s to C and G wh ich gi v e

, L C L

M and M s o that M
L C, G l C and M , GL
-
C L C
: .

A ls o let D and D repres ent d ev iat io n s of c las s m ark s

L
' '

fro m the g uess es G and G res pect iv ely

, C
.

U n les s o n e carr ies th ro u gh a larg e n u m ber of d ec ima l p lac es

t he m eth o d prev io u s ly d is c u s s ed is n o t o n ly v ery la b o rio u s b u t
it is m u ch les s a cc u rat e than t he s h o rt er m eth o d h ere d es cribed .

SECT I ON V — T HE R EG R ESS I ON C OE FFI C I ENT

F ro m the c o rrelat io n c o effic ient and the s tand a rd d ev iat io n s

wi th res p ect to t wo cha ract ers i t is eas y to o bta in what is kno wn
as t he regress io n c o effi c ient T o o bt a in the regress io n c oeffi c ient
.

o f t he weight o f ear s r elat i v e t o th ei r leng th s m u lt ip ly t he ,

c oeffic ient o f c o rrelat io n by t he s tandard d ev iat io n o f weight ,

and d iv id e t he pro d u ct by t he s tandard d ev iat io n o f length .

T h is giv es fo r t he regres s io n o f weight relat iv e to length

, ,

S im ilarly , t he regressio n of length relat iv e to weig ht is

U se of t h e regression regress io n c oeffic ient is

coefi cien t . T he
u s ef u l fo r pred ict io n ; that is to s ay if we kn o w t he d ev iat io n ,

o f o n e cha ract er fr o m it s m ean th is c oeffi c ient will enab le u s t o

,
 C OR R ELAT I ON 6
4 7

COR RELA T ION STU D Y ( LE A M I NG COR N C R OP, 1 9 5

0 )
C I R C U H P E R EN C E .
L

(D L ) f
’ 2
wd ) “

1 00 00

v
s 3

s
0 u T « a «
s
« 0. Y
u .

6 é
.

d m
.

m
:
0
q v o o
. . w n m » . n m
u
\ I I I l
I

v 0 0
« n n

d d
.

«
b
u ” m w v «
t « h 0 T

v
.

e
.

0 o
o
« n 3
.

.
S
m
S o m m 0 m
w u m n e o
e d
. .

U d - 0
m
Q h 0 n
.

” C
4 6 8 TR A NSM I SSI ON

p red ic t what will

pro bab ly b e t he d ev iat io n o f the c o rrelat ed char -

act er fro m its mean T h u s s u ppose we select c ars wh ich d ev iat e

.
, ,

sa t wo i nch es fro m the mean length o f c a rs ; that is wh ich a re

y , ,

t wo i nch es ab ov e t he a v erage : t he r egres s io n c o effi c ient

of weig/zt r ela t v e i to leng th i n d ica tes tlta t we s lcou ld ex p ec t s u e/c

ea r s to be a b ou t nc es fro m the mean that is 2 x

ou ,

To b e mo re general if we s elect ears wh ich ha v e any d ev iat io n

,

x fro m t he m ea n length we S h o u ld exp ect th eir d ev iat io n s in

,

weight to c ent er abo u t a v a lu e x fro m t he m ean weight of

ear s fo r t he wh o le p o p u latio n .

T he regres sio n c oeffi c ient is th u s a fix ed rat io b et ween d ev ia

t io n s of c o rrelat ed cha ra cters so that kno wing ho w m u ch o n e o f
, ,

t he charact ers d i ffers fro m it s m ean in any u n it o f m ea s u rem ent ,

sa
y i nch es we a re enab led t o pr ed ict ho w m u ch t he a s s o c iat ed
,

character d epart s fro m i ts m ean in i ts u n it of m eas u rement say ,

in p o u nd s . T h u s if a regres s io n c oeffi c ient o f weight u po n s tat

u re is ,
sa
y w e kn o w that any d epart u re fro m t he m ea n

s tat u re will b e fo llo wed by a d epart u re t im es as great in

res pect to weight u sing in b o th ca ses t he sa me u n it s as were
,

u s ed in ca lc u lat i ng t he c o effi c ient fo r exa m p le f eet and p o u nd s , ,

inch es and o u nc es o r ev en inch es and po u nd s if th es e were t he

, , ,

u n i t s act u a lly u s ed in c o m p u t i ng t he r egres s io n c oeffic ient .

SECT I ON V I —ST U I ) I ES I N SPE ED R ECOR D S OF

T R OTTE R S

S t u d ies were m ad e o f tro tt ers po s sess ing rec o rd s of

2 3 0 o r b ett er ,
i n o rd er
to learn th ei r d is trib u t io n a s t o s p eed ,

and the po ssib le c o rrelat io n o f s peed with c o lo r and m o re par ,

t ic u la rly with sex ( s ee tab les pages 4 6 9 and ,

T h e data were tak en fo r each q u arter s ec o nd and t he rec o rd ,

m ad e a s cro ll o v er fo rt y f eet lo ng T he m att er is h ere c o n.

d en s ed to d ifferenc es o f o n e s ec o nd in o rd er to bri ng it into

s u i tab le s pac e .

T he o rigi nal rec o rd s ho wed t wo s trange p ec u lia rit ies A lmo s t .

in v ariab ly t he larges t n u m b er o f rec o rd s was fo u nd o n t he firs t

q u art er sec o nd after t he ev en m in u te as et c that is
, .

to s ay t he r ec o rd s were n o t ev en ly s pread o r as m ath em at ic ian s

, , ,
47 0 TR AN SM I SS I O N

sa
y ,
th ey were n o t s m oo th and if t he c u rv e were p latt ed th ere
,

wo u ld b e an u nacc o u n ta b le h u m p at ea ch q u art er sec o nd .

The sec o nd pec u liari ty ha d referenc e to t he las t rec o rd 2 3 0 , .

By t he p rinc ip le j u s t s tat ed t his n u m ber s h o u ld ha v e been a bo u t

0
7 p er c e nt of t h e n u m ber r ec o rd ed at 2 O n t he c o ntrary ,

it was v ery m u ch grea t er O n t he pri nc ip le ru n n ing thro u gh

.

t he res t o f t he rec o rd s we s h o u ld pred ict t he n u m be r at 2 3 0 to

b e abo u t 7 2 7 wh ereas t he n u m ber act u a lly rec o rd ed is 1 09 7
, ,

s h o wing c o nc lu s iv ely that s o me 3 7 0 h o rs es had been a d m itt ed

to t he 2 3 0 lis t that rea lly belo ng ed at 2 3 o i o r s lo we r ; a ll o f

wh ich s h o ws ho w s tat is t ical s t u d ies b ring to t he s u rfac e as n oth
in g els e will t he n at u ra l i rreg u la ri t ies o f o b s e rv a t io n s o r what ev er
abno rm a l fact s c o nnect th em s elv es wi th o u r in v es t iga tio n s The .

table o n page 4 69 is s u gges t iv e in many way s .

R ELA TION or S Ex A ND C O LO R To SPEE D As E x p RESSE D IN

R AT E PE R C ENT

A ND a w l AND BE L O w 2 :1 0 AND BE Lo w
D ES C R PTI ION I

Per Ce n t N u m b er I Per C e n t Per Cen t

4 4 93

Co lo r
'

T o ta l

F ro m t he
table ab o v e it appears that th ere is litt le relat ion
b et ween s peed and eith er s ex o r c o lo r I t is t ru e that as we .
,
 COR R E LAT I ON 47 1

read across t he ta ble we see that the percentage o f females fa lls

,

S light ly as we get i nt o h igh s peed s b u t t h e fa ll is v ery S light , .

T he perc entages as to c o lo r v a ry b u t s light ly exc ept in b lack , ,

wh ich d ec id ed ly increa s es with h igh s peed and in gray wh ich as , ,

d ec id ed ly fa lls o ff .

A pp ly i ng Y u le s for m u la to the s t u dy o f th es e fig u res let u s

’

, ,

fo r exa m p le tak e the bay c o lo r and i nq u i re as to t he m ea s u re

,

o f c o rrelat io n bet ween th is c o lo r and h igh s p eed

T o tal n u m b er o f b ays
N um b er of b ays at o r b elo w 2 :1 5
—2 16

T o tal n um b er o f er fo rm ers
p
T o ta l n u m b er o f per fo rm ers a t o r b elo w 2 :1 5 -
2 :1 6

N u mb er not b a ys at or b elo w is 9 1 5 5 0 2 413

N u m b er of b ays ab o v e 2 1 5
-
2 :1 6 is 7 3 76 5 0 2 68 7 4
N um b er not b a ys ab o v e is 4 1 3) 6090

A rrang ing th ese v alu es acc o rd ing Y u le s fo r m u la we hav e

'
to ,

N OT B A v

An o v s i
z zl s

Th is g iv es as a m eas u re of t he as s o c iat io n of t he bay c o lo r

and s peed ,

99 90 x 5 0 2 68 7 4 x 4 1 3
6 0 90 x 5 0 2 68 7 4 x 4 1 3

While th is res u lt h o ws that the bay s ha v e f u rn is h ed

,
S

s light ly m o re than th ei r s har e o f t he h igh s p eed tro tt er s it is -

,

d o u btf u l wh eth er th is c o effi c ient is large en o u gh t o enab le u s to

assert any d ec id ed c o rrelat io n .

I n m u ch t he s a m e way exha u s t iv e s t u d ies s h o u ld b e m ad e in a ll

lines o f breed ing and at any expen s e o f t im e a n d lab o r in o rd er
, ,

that we may po s s es s o u rselv es o f relia b le info rm at io n in regard

to as m any d eta ils as p o s s ib le c o nc ern ing t h e relat io n s o f n o tab le
characters in o u r m o s t v alu ab le rac es .
 T R A N SM I SSI O N
4 7 2

S u mmary . C o rrelat io n is
gen era lly a relat iv e m att er and ,

im pres s io n s are exc eed ing ly d ec ept iv e N o th ing b u t ac t u al c a l .

c u lat io n will S h o w t he extent to wh ich cha ract ers rea lly mo v e

t o geth er and t he im po rtanc e o f th is kn o wledge is a m ple rec om
,

pen se fo r a ll t he labo r in v o lv ed A s will be seen lat er t he sa m e .

,

m eth o d s gi v e u s th e o n ly reliab le m eas u re o f h ered ity .

S PE C I A L E N E R C I S E S

A ga in let th e s t u d en t a c t u a ll y d o th e wo r k of fi n d in g c o rrela tio n c o effi

c ien ts u n ti l he a c q u i r es fa c ili ty in o p er a t io n a n d a d is ti n c t c o n c ep tio n o f

wh a t is in v o lv ed .

ADDITIONA L REF ER ENCES

C O R R E L A TI ON IN R YE . E x peri m en t S ta ti o n R ec o rd , X I I I , 241, 64 1 .

C O R R E L A T I ON IN T HE PA R T S O F C O R N . By A . A . B r ig h am , G ottin gen .

E x p eri me n t S ta tio n R ec o rd , V I I I , 4 86 .

C O R R E L A T I O N I N W HE A T E x peri men t S ta tio n R ec o rd , I X , 5 5 3

. .

C O R R E L A T I O N M A T HE M A T I C S S c ien c e, X X I I , 3 09- 3 1 2
. .

C O R R E L A T I O N O F C HA R A C T E R S C OR N G erm a n E x perim en t S ta ti o n
IN .
( .
)
R ec o rd , X V I , 46 1 .

C O R R E L A T IO N OF S E E D S AND COLOR or FR U IT . E x peri m en t S ta tio n

R ec o rd , X I , 93 2 93 6 — .

C OR R E L A T I ON O R T HE M ENT A L AN D P HY S I C A L C HA R A C T E R S IN MAN .

B y A lic e Lee a nd K a rl Pea rso n . Pro c eed in gs R o ya l S o c iety, LX X I ,

1 06 - 1 1 4 .

C O R R E L A T I ON T H EOR Y . S c ien c e, X X I , 3 2 —
35 .
4 74 T RA NSM I SS I ON

of it s and each es tab lis h es and m aintain s apparent ly fa irly

o wn , , ,

d efin it e relat io n s to c erta in o f it s as s o c iat es (c o rrelat io n ) wh ile ,

with referenc e to o th er s it s eem s ind ifferent if n o t ind ep end ent .

I n di v idu al s in h erit di fferen tl y A ll ind iv id u a ls o f the s a m e .

rac e po s sess t he sa me c ha ract ers b u t in d ifferent pro po rt ion s , ,

and n o t wo ind iv id u a ls ev en fro m t he sa m e parent s a re alik e

, , .

S o m e po rt io n o f th is d i ff erenc e is o f c o u rse d u e to d ev elo p m ent

acc o rd i n g to t he c o nd it io n s o f life yet all ev id enc e goes t o s h o w ,

that aft er f u ll a llo wanc e is m ad e fo r th is fact o r nat u ral d iffer

, ,

en c es ex is t that c a n b e d u e o n ly t o inh er itanc e .

T hat each ind iv id u al is in po s s es s io n o f all t he chara ct er s o f

t he rac e is ev id enc ed by t he fact that h is d es c endant s p o s s es s
th em and that he tran s m it s fa r m o re charact ers t han a re d e
v elo ped s u ffi c ient ly t o b e n o t ic eab le in his o wn p er so nalit y .

Lat en t char act er s T h u s cha ract er s m ay b e pres ent b u t

.
,

u nd ev elo ped lat ent Ga lt on as sert s that lat en t chara c

“ ”
or , .

t ers are n o t v ery n u m ero u s ; b u t it is c ert a in that m any

“ l "

charact er s m ay rem ain u nd ev elop ed thro u gh life and yet b e

t ran s m itt ed perfect ly F a m iliar exa m p les a re t he o ccas io nal
.

s ecr et io n o f m ilk by t he m ale s ex already allu d ed t o and , ,

t he t ran s m iss io n o f t he m ilk ing q u a lity by b u lls a s well a s

by c o ws A ll th ing s c o n s id ered it is s afe to say th a t t he V is i
.
,

b le and f u lly d ev elo ped charact ers o f an ind iv id u a l c o n s t it u t e

b u t a s m a ll pro po rt io n o f his real po s sess io n s E s pec ially m ay .

th is b e s aid o f a h igh ly d ifferent iat ed rac e .

I n heri ta n ce n ot l imit ed t o sex I t ha s been a fa v o ri t e s ay ing .

that c erta in charact ers a re tran s m itt ed to o n e s ex b u t n o t to the

o th er T h ere is n o ev id enc e o f any s u ch lim it a t io n s t o inh erit
.

anc e The limitat io n s o f sex m ay a n d d o prev ent th e d ev elop

.

m en t o f m any charact er s that we kn o w t o b e p o t ent ia lly present ,

s o far as inh eri tanc e is c o nc ern ed beca u se th ey can b e t ran s ,

m itt ed I n th is res pect t he relat io n o f t he m a le m a m mal to m ilk

.

se cret io n is n ot d ifferent fro m that o f t he f em ale that has n ev er

y et b o rn e y o u ng T h e fac u l ty is lat ent
.
2
o r u nd ev elo p ed in bo th , ,

1 l
G a t o n , N a t u ra l I n h eri t a n c e, p . 1 87 .

1 T he t erm “
l a t en t is u n fo rt u n a te . It c on v e s y t o o s t ro n g l y th e s en s e of

k
lu r in g U n ev elo p e
.
“
d d is t h e s en s e t a t o u g h ht t o a t tac h to th i
s u nf o rt u n at e

h
t erm t a t has n o w b een use d to o lo n g to be d i l dg d
s o e .
 H ER ED ITY 475

c ases T h ey d iffer o n ly in the fact that with the fem ale t he

.

d ev elo p m ent is eas ily bro u ght a bo u t wh ile in t he m ale it is ,

d iffi c u lt and in mo s t cases im po ssib le 1

.

Bela t ed in h eri ta n ce I t is well kn o wn that all charact ers d o

.

n o t d ev elo p c o nt em po ran eo u s ly T h u s t he s ex u al charact ers .

bec o me d ev elo ped j u s t befo re f u ll s tat u re is att ained and with ,

t he fai lu re o f t he pri m ary sex u al charact er s with ad v anc ing age

c o mes t he d ev elo p ment o f many o f t he pec u liarit ies o f th e o th er
s ex . T hen it is th at t he hen c ro ws t he h u m a n f em a le gro ws ,

m o re hair and t he v o ic e o f t he m a le b ec o m es effemin at e T he

,
.

t erm belat ed in h eritanc e th o u gh fix ed in o u r lit erat u re is

“
, ,

u n f o rtu nat e I t is belat ed d ev elop men t that is meant I nh eri tanc e

. .

c o mes o n ly at o r rather b efo r e b irth ; b u t d ev elo pm en t is c on

d it io n ed u po n m any fact o rs — a m o ng wh ich age and s ex a re
, ,

im po rtant b u t n o t t he p ri nc ipal c o n s id erat io n s

, , .

Bl en ded an d ex cl u siv e in heri ta n ce Perhap s t he fi r s t and .

mo s t n o t ic eab le fac t is that so me character s b lend wh en brou ght

t ogeth er by tran s m is s io n wh ile o ther s rem ain d is t in ct being
, ,

apparent ly m u t u ally exc lu s iv e T h u s s k in c olo r in m a n blend s .

read ily t he cro s s bet ween wh ite and n egro being n ea r ly alway s
,

o f so m e s had e int er m ed iat e b et ween th o s e o f t h e parent s ,

“
almos t n ev er s po tt ed I n Sho r tho m catt le and in J ers ey s the
l
.

c o lo r s freq u ent ly if n o t gen erally b lend wh ile in t he Ho ls t ein

, , ,

F ries ian th ey a lway s rem a in d is t inct I n h o rs es t he b lend is .

c o m mo n b u t in h og s it is pract ically u nkn o wn so that in a

, ,

litt er o f p ig s fro m a b lack and a wh it e parent t he c o lo r s will

rema in d is t in ct ; so me m ay b e black s o m e wh it e and o thers , ,

s po tt ed b u t n o n e will b e r o an s o r gray s
, .

The s a m e d is t inct io n h o ld s a s to cha ract ers gen era lly S o me .

t imes the o ffs pring will b e int erm ed iat e bet ween the parent s ,

s h o wing a b lend ; and aga i n it will res em b le o n e o r the o th er ,

o r els e exh ib it t rac es o f b o th each d is t inct and separat e ,

.

F o r exa m p le s o far as t he m att er has b een s t u d ied t he b lend

, ,

is mo s t p erf ect a s t o s tat u re and pro bab ly a s to S ize in gen eral

3
, ,

b u t eye c o lo r d o es n o t read ily b lend 4

n or do
“
t em pers 5 ”
or ,

1 T he s t u d en t is rem in d d th
e at m il k s ec ret io n a m o n g m a les is n o t u n n o wn k .

3 T h e s p o t t e s in is d k n o t a b s o lu t e ly un k n o wn a m o n g u m an s , h
o wev er h .

3 G al t o n , N a t u ra I n l h erit an c e, 8
p 9 . .
1 Ib i
p 1 4d5 .
5 Ib i
.
p
. 2 33 d . . .
47 6 T R A N S M I SS I O N

“
tas t es . T h is
being t ru e we a re o ft en d isappo int ed in t rying ,

to m od ify o r t o n e d o wn a v ic io u s d is po s it io n by m at ing with

o n e o f m ild er t em per t he pro geny t end ing to f o llo w t he a v erage
,

o f t he rac e o r els e t o b e a s v ic io us a s t he o bj ect io nab le paren t

, .

Part icu lat e in heri ta nce, in herita n ce b y t ype, or hit b y b it .

C ha ract ers ft en s o a re o c lo s ely as so c iat ed ( c o rrelated ) as to

m o v e in c o m p a ny so that wh o le gro u p s of cha ract ers appea r
,

a n d d is appear t o geth er ev en wh en th ere is litt le o r n o kn o wn ,

cau s at iv e relat io n bet ween th em Wh eth er th is is merely acc i .

d ental assoc iat io n o f character s n o t m u t u ally exc lu s iv e and c ertain ,

to happen oc ca s io na lly u nd er t h e law o f chanc e o r wh eth er it ,

is d u e rath er to so m e d eeper ly ing p ri nc ip le is pe rhap s u n c er

, ,
-
,

tain ; b u t it is su rely tru e that m an fo r exa m p le ru n s in types , , ,

and whoev er has tra v eled mu ch o r has enj oyed a fairly ext en ,

s iv e acq u a i ntanc e has m et m any p eo p le o f n o b loo d relat io n s h ip

, ,

in p lac es wid ely s epara t ed who yet were c learly o f t he s a m e ,

type and wh ose s im ilar ity beca m e mo re ev id ent u po n c loser

,

acq u aintanc e ‘
.

C learly charact ers a re n o t a lt o geth er ind epend ent o n e o f

,

an o th er and o ften t he great es t d ifli c u lty is enc ou nt ered in

,

break ing u p a gro u p so m e m em bers o f wh ich a re d es irable and

,

o th ers o bj ect io nab le S o inh eri tanc e is o ft en b it by b it as if

”
.
,

t he u n i t o f tra n s m is s io n were larger a n d m o re c o m p lex than t he

s ingle charact er ; a s if a k i nd o f perm an ent partn er s h ip were in

f o rc e T he b io lo gi cal bas is o f all th is if it rea lly ex is t s will

.
, ,

pro bab ly rem ai n fo r a lo ng t im e h idd en b u t c oefli c ien ts o f ,

c o rrelat io n a ffo rd at lea s t a meth od fo r d et erm ini ng t he d egree

and t he pers is t enc e o f th is c o partners h ip .

Pol ymor ph is m an d sex u al di morphi sm M any ra c es i n s t ea d .

,

o f s h o wi ng a ll i nt erm ed iat e g radat io n s fro m o n e extrem e t o the

o th er in res pect t o s ize fo r exa m p le o r c o lo r o r any o th er

, , ,

charact er o r as so c iat io n o f cha ract ers will exh ib it t wo three , , ,

o r m o re f o r m s o r typ es s o d iff erent and d is t inct a s o ft en to b e

,

1I n p ra c t ic al b reedin g o p era tio n s t he grea tes t n ee ex is ts fo r d exa c t k no d

wle ge
o f c o rr ela t e d h
c a ra c t ers T he m et o s giv en in t he p rec e in g c
. h d d h a p ter en ab le t he
stu d en t t o d t mi
e er ne q u a n t it a t iv e l y th e rea l ex t en t o f c o rre a t io n , a n d l b ree de rs

sh ld p
ou ro s ec u t e m o s t in d u s trio u s l y th e st u dy of h
t is s u b ec t , j u n ti l th y
e well
a re

i f m d
n or e as t o th e rea l rela t io n s of a ll v alu ab le c h a rac t ers o f d m
o es t ica t ed

an ima ls an d p lan ts .
47 8 TRA N SM I SSI ON
charact ers ( s ee Mend el s la w) Wh eth er i n goo d t ime th ey ’
.
, ,

wi ll b lend o r will r em a in d is t in c t giv ing ris e to po ly m o rph ic

, ,

fo r m s with in the breed is an im po rtant q u es t io n i n wh ich the

,

breed er is alway s d eep ly int eres t ed If the po ly m o rph is m c an .

b e rem o v ed he o f c o u rs e d es ires t o d o i t ; if n o t he m u s t m ak e
, ,

t he b es t o f it and c ea s e wa s t ing t im e o v er t he u nattai nab le .

I n a ll s u ch ca s es t he breed er is t o s at is fy h im s elf as q u ick ly

a s p o s s ib le wh eth e r t he po ly m o rph is m is t em po rary o r p er m a

nent ; and if it b e pe r m anent he will d o well to ch oo se the ,

type he is to breed and aband o n the effo rt t o b lend it with

,

ano t her — ,
ih o th er wo rd s he m u s t b e c o nt ent to sec u re his ,

res u lt s g rad u ally by s elec tio n ,

.

SECTI O N I I —STAT I STI C A L M ET HOD S O F STU D Y OF

H ER E D I T Y

U nt il rec ent ly n o phas e o f ev o lu t io n has been s o bad ly s t u d ied

a s h ered i ty T he c o m m o n m is tak e ha s b een t o n o t e a few
.

rem arkab le ind iv id u a ls and exc ept io nal in s tanc es and fro m ,

t hes e att em pt to d ed u c e t he laws o f d es c ent

“
I n th is way .

po p u lar c o nc ept io n s o f hered ity hav e g ro wn u p m any o f wh ich ,

a re exc eed ingly erro n eo u s n o t t o s a y f a n t as t ic ,

.

We ha v e o nly rec ent ly learn ed t hat s t u d ies and c o nc lu s io n s

based u p o n ind iv id u al in s tanc es a re wo rs e tha n u seless becau se of
t he extrem e ran g e o f v ar iab ility and t hat t o d eter m in e t he fact s ,

o f h er ed ity with any d egree o f reliab ility we m u s t s t u dy w e ra c e as ,

a w hole a n d n o t s im p ly t he s eparat e i nd iv id u a ls that c o m po s e it

, .

A ll t h is m ean s that t he la ws o f d es c ent a re to b e d isc o v ered

by a c ri tica l s t u dy n o t o f ind iv id u als b u t o f en tir e pop u la tions
, , ,

o r at lea s t o f pro po rt io n s o f po p u lat io n s s u ffic ient ly large to b e

s af ely repr esent a t iv e .

U nfo rt u nat ely t he app licat io n of t he s tat is t ica l m eth od to t he

,

s t u dy o f th is s u bj ect is c o m pa rat iv ely n ew and as it is extremely ,

labo rio u s t he acc u m u lat io n o f a large ma ss o f m at erial will of

,

nec essity b e a so mewhat s lo w proc es s l

.

1G lt a won th fi t t as pp l y e
p t d y m t h d t th
rs o a t dy f h res en dity -
a e o s o e s u o ere ,

b t P
u d th
ea rso n an f ll w do d e rs id o b l l it
o te ,i an m l ti g
a c o ns era e era u re s ac c u u a n ,

to wh i h im p t t dd it i
c o r an b i g
a
p i d ly m
o nsd T h a re
q t ly j
e n l ra a e . e u a r er o u rn a

Bi m l l wi d
o e n t d t
s th t dy f t h i
ev o e b j t b y th
o e s u t ti t i l m et h d
o s su ec e s a s ca o .
 H ER E D IT Y 4 79

U nfo rt u nat ely again the fi rs t and mo s t exha u s t iv e s t u d ies

,

were m ad e ou t s id e o f o u r fi eld and m o s t ly in that o f h u m an ,

chara cters so that the bes t m at erial fo r t he s t u dy o f h ered ity

,

lies in th is fi eld B u t lat er s t u d ies in wid er fi eld s lead u s

.
, ,

c o nfid ent ly t o believ e that t he sa m e genera l princ ip les c o ntro l

t ran s mis sio n s ev ery wh ere in all ra c es and with a ll cha ract ers
, .

A cc o rd ingly t he writ er will em p lo y any s t u d ies that hav e been

m ad e in what ev er fi eld s that o ffer v alu ab le m at erial eith er fo r
, ,

elu c idat i ng pr inc ip les o r fo r i llu s trat ing meth o d s o f s t u dy T he .

fu lles t data o f a ll a re tho s e c o llect ed by Galt o n d eali ng princ i ,

pally with s tat u re and th ey will b e freely em ployed fo r bo th

,

pu rpo ses .

SECT I O N II I — T HE R EG R ESS I ON TA BLE

As a lready seen (chap x ii) wh en a repres entat iv e po pu la

.
,

tio n o f any rac e is arranged in t he fo r m o f a freq u ency d is t r ib u

t io n we a re ab le to d ed u c e exc eed ingly acc u rat e expres s io n s fo r
v ariab ility .

I f n ow th is d is trib u t io n b e s eparat ed a n d as s or ted a cco rd ing

to pa r en tag e we s ha ll hav e a s eries o f d is tr ib u t io n s each o f
, ,

s im ilar parentage t h e wh o le pres ent i ng t he bes t fac ilit ies p o s s i

,

b le fo r the s t u dy o f h ered ity S u ch a tab u lar arrangem ent c o n

.

s tit u t es what is ca lled a regress io n tab le and inas mu ch as

“ ”

a ll regres s io n tab les pres ent t he s a m e g en era l feat u res we loo k ,

u po n th em with c o nfid enc e a s a ff o rd ing r eliab le data fo r the

s t u dy o f th is m o s t im po rtant b u t o th er wis e m o s t d iffi c u lt and

apparent ly self c o nt rad ict o ry s u bj ect

-
.

I n all regres s io n tab les a s ca le o f v alu es ( m ea su rem ent s ,

weight s n u m erals o r o th er v a lu at io n ) is pro v id ed at o n e s id e

, ,

fo r the parent s and a c o rres po nd ing s cale a lo ng t he t o p fo r t he

,

o ffs p ri ng o r v ic e v er s a
1
.
,

T he o ff s pri ng c o n s id ered as ad u lt s is th en d is tri b u t ed each

, , ,

ind iv id u al being rec o rd ed o ppo s it e t he v a lu e repres ent ing his

1 Ob v io u s l y
t he pa ren ta l m ea s u rem en ts m ay b e a rra n ge d l a o ng t h e t o p an d
h
t o s e o f t h e o ff s p ri n g a t th e s i e E v er d . y o b s erv er fo l l ws his
o fa n c in t is y h
res pec t , a n d as a m a t ter of fa c t t he t a b l es a re ma d e in b o t h wa y s . T h e writer
has b ec o m e m o re ac c u s t o m e t o t he d one d es c rib e d in t he t ex t, an d fo r no o t h er

reas o n p ref ers t o a r ra n ge t he pa ren ta l l

v a u es a t t he s i e d .
 8 2 T R A N S M I SS IO N
4

I t is not lik ely

chara ct ers beh a v e prec is ely a s d oes that a ll

s tat u re ind eed i t is kn o wn th a t th ey d o n o t ; b u t a ll s t u d ies

, ,

o t o S h o w that charact er s o f ev e ry k ind o b ey t he s a m e g en e ral

g
la ws in d es c ent and a ll regres s io n ta b le s th a t h a v e ev e r been
,

prepa red exh ibit t he s a me genera l feat u res T h is tab le th ere

‘
.
,

fo re wh ile p rim arily fo r t he s t u dy o f s t a t u re m ay b e c o n s id ered

, ,

as typ ica l o f regr es s io n ta b les and d ed u ctio n s m ad e fro m it , ,

agr eeing as th ey d o with th o s e m ad e fro m all o t her s im ilar

ta bles whatev er the rac e o r t he cha ract er may b e s a fely
, ,

acc ept ed as exh ib iting f u ndam enta l la ws in h ered ity Bec a u s e .

a ll regres s io n tab les a ff o rd t he sa m e d ed u ct io ns they m ay b e ,

s ta t ed in t he fo r m o f gen e ra l pri nc ip les as o u t lin ed in t he

fo llo wing sec t io n s .

SECT I O N I V L I K E PA R ENTS BEG ET U N LI K E OFFSPR I N G

A N D , C O N V E R S EI X , L I K E O FFS PR I N G M AY B E
B EG OTT E N BY U N LI K E PA R E NTS

th is tab le the h eight s o f the pa rents a re in t he ro ws 5 to m

In ,

and tho se o f t he ch ildren (as ad u lt s ) in t he c o lu m n s 2 t o 1 5 I t .

will b e s een at o nc e that t he o ffs pr ing o f parent s o f any g iv en

s l igh t d
m o ifi ca t io n s a re n ec es sa ry o n ac c o u n t o f t he p a ren ta l a n c es t r y , b ut s uc h
c o r rec t io n s a re n o t n ec es s a ry fo r p res en t p u rp o s es .

G a to n l ca ll s a t t en t io n t o a n er ro r in t h e fi rs t ro w of c h il d ren an d m id pa ren ts , -

sa yi ng t h a t a n err o r was in t ro d u c e d so me w h ere in t h e o r igi l t bl

na a es , w ic h h
c an n o t

no w be c o rrec t e d . I t is o b v io u s th at fo u r c i h ld ren c a n n o t h fi av e v e m id
pa ren t s~
,
”

he sa y s, b u t t he n u m b e rs a re s o S m a ll as t o b e gen era ll y d i d d sc a r e , an d h en c e t he

l
ta b e is rep ro d uc e d , e rro r a n d a ll . He a dd s T he b o tt m li
o
( f ne o u rt e en c h il d ren

wit h one m id p a ren t ) . w ic

-
h h l k oo s s u s p ic io u s , is c o r rec t ( N a t u ra l I n h eri t a n c e,

p .

In l l
c a c u a t io n s g e n era ll y t h e e x t rem es ( a bo v e or or b elo w or

a re di sc a r d d e b ec a u s e t h e n u m b e rs a re s m a ll and b ec a u s e exa c t m ea s u re
m en t s a re n o t giv en . In l l
c a c u a t in g t h e g en era l m ea n , h o wev e r, t wo v a l u es h av e

b een d et erm in e d , o ne wit h out t h e ex t rem es , t he o t h er b y in c l di

u ng t h e ex t rem es ,
as s u m in g t h at m ea s u rem en t s ab o v e a v era g e d ab o v e a v era ge d
be l ow a v e ra ge d and be l o w a v era ge d The ass u m p t io n is
en t ire ly gr a t u it o u s , b u t it affo r d s a b as is fo r u s in
g t he ex t rem es , a t lh o u gh , as is
n o t ic e d , it m a k es b ut s l igh t d iff e ren c e in t h e res u lt s .

1 R egr es s io n t ab l d
m a y b e p repa re fo r a ny c a ra c ter t a t c a n b e m eas u re ,
es h h d
weig h d
e , c o u n te d , o r in a n y wa y a c c u ra t e et e rm in e It on ly d a pp en s t a t d . ly h h
s tu di es in h u ma n s t a t u re h
a v e b e en t h e m o s t c o m
p et e o f a n
y, a n d a re t erefo re, l , h
u se dh e re .
 H ER ED IT Y 8
4 3

h eight a re n o t a me b u t o n t he o th er hand t hat th ey c o n

t he s , , ,

s t it u t e a d is tr ib u t io n b eg inn ing b elo w t h e parentage and ext end ,

in g to a c o n s id erab le d is tanc e a b ov e it with t he la rges t n u m ber ,

o f ind iv id u a ls n ear t he m idd le o f t he d is tr ib u t io n c lo s e t o b u t ,

n ot id ent ica l with parenta l h eight .

T h u s t he 6 8 ch ildren o f 2 2 parent s inch es h igh ( row e)

a re d is tr ib u t ed fro m b elo w t o ab o v e i nch es a rang e o f ,

I I i nch es wi th t he grea t es t n u m ber ( 1 8 ) s light ly belo w the

,

parental h eight A n y o ther ro w tak en at rand o m will S h o w

t he s a m e d is tri b u t io n in th e s tat u re o f the o ff s pri ng Hered ity .
,

th erefo re i nv o lv es s o m ething b es id es t he infl u enc e o f the im me

,

d iate parent wh ich acc o rd ing t o all s t u d ies seld o m exc eed s 5 0
, , ,

p e r c e nt o f t h e t o ta l i nfl u enc e o f t h e anc e s t ry lea v i ng t he o th er ,

half to b e acc o u nted fo r by anc es t o rs farth er back l

.

N o t o n ly is i t tr u e that lik e parent s pro d u c e u n lik e o ffs pri ng ,

b u t t he c o n v erse is a ls o tru e — that lik e o ff s p ring m ay res u lt ,

fro m u nlike parent s T ak e any c o lu m n o f t he tab le at rand om

.
,

as c o lu m n 1 0 c o nta in ing t he d is trib u t io n o f t he 1 6 7 ch ildren o f

,

t he u n ifo r m h eight inch es ,

T h es e m en o f ev en h eight .

were pro d u c ed by parent s ranging i n s tat u re a ll t he way fro m

inch es d o wn to less than inch es — a ra n ge o f 8 ,

inch es .To b e s u re the parenta l h eight that pro d u c ed t he

,

grea tes t n u m ber (4 8 ) was inch es n o t far fro m t he c o mm o n ,

h eight o f t he o ff s pring ( ro w 0 c o lu m n 1 6) and alm o s t exact ly ,

t he a v erage h eight o f a ll t he parent s ( ro w o c o lu m n b ut ,

t he crit ica l s t u dy o f th is and a ll t he o th er c o lu m n s will c lear ly

S h o w that t he sa m e k ind o f o ff s p ri ng may b e pro d u c ed by

great ly d ifferent parents .

T h es e fact s s h o w c lear ly that t wo s i res o r da m s o f eq u ally

fav o ra b le appeara nc e m ay ha v e s pru ng fro m v ery d ifferent
anc es try Th ey bo th belo ng to a d is trib u t io n c o v er ing a c o n
.

s id era b le ra n ge and if o u r s elect io n is t o b e effect iv e we n eed

,

t o kn o w ev eryth ing p o s s ib le o f t he ent i re gro u p to wh ich t he

pros pect iv e parent b elo ngs o r at leas t b e int elligent as to t he ,

po rt io n o f the d is trib u t ion fro m wh ich he is drawn .

1 E v ery in iv i d d ua l o f l
b is ex u a p a ren ta ge ha s a to tal o f 2 04 6 a n c es t o rs h
wit in
t en gen era t io n s h h
W et er t es e a n c es to rs rep res en
. h a t m an ifferen t in iv i t th yd d d
l h h
u a s, o r w et er s o m e are o f t-rep ea t e , d d
ep en s u p o n t h e c o s en es s o f b ree in
g d l d .
 8
4 4 TR A N SM I SS I ON

SECT I ON V R EG R ESS I O N . IN G E N ER A L , TH E O FF
S PR I N G I S M OR E M ED I OC R E T HA N T H E PA R ENTS ;

T HA T I S, WHA T EV E R T H E PA R E NTA G E, T H E O FF
S PR I N G EX H I BITS A ST R ON G TEN D E N C Y TO

R EG R ESS T OWA R D T H E M EA N OF T H E R A C E

A glanc e at any regres sio n tab le S h o ws an u nev en d is trib u

tio n o f the po p u lat io n with t he larges t n u m ber s near t he m idd le
,

o f t he tab le exh ib it ing a s tro ng t end ency t o c lu s t e r ab o u t t he

,

cent er N o t o n ly is th is so b u t if a ny parental ro w (e t o m ) b e
.
,

caref u lly s t u d ied t he f o llo wing po int s will b e no t ed

,

I
. T he m ean o r a v erag e h eight s o f t he ch ild ren ( c o lu m n 1 8 )
a re in n o ca s e t he s a m e a s t he h eight s o f t he parent s C o m pare .

c o lu m n 1 8 with c o lu m n I .

2. Wh en t he parenta l h eight is a bov e t he mean o f the rac e ,

that is t o say inch es and u p ward (e t o g ) th en t he m ea n ,

h eight of t he ch ildr en is so meth ing les s than the h eight o f the

pa rent ( s ee any row fro m c to g ) .

3
. B u t w h en t he par enta l h eight i s be l o w t he m e an o f t h e

rac e —,
6 7 5 in ch es and les s (it t o m )
. th en t he m ean o f the ,

ch ildren is great er than the h eight o f the parent ( see any ro w

fro m It t o m) .

To illu s trat e : in ro w e a re r ec o rd ed t he h eight s o f t he 6 8

ch ildren o f 2 2 m id parent s inch es h igh
-
I n c o lu m n 1 8 we .

s ee that t he m ea n h eight o f th es e 6 8 ch ildr en wa s inch es ,

or a h eight o n e inch b elow t he par entage and by that m u ch ,

n ea rer t he gen era l m ean o f t he rac e A ga in in ro w k are .

,

rec o rd ed the v ario u s h eight s o f the 66 ch ildren of 1 2 m id paren t s -

i n c h es tall I n c o lu m n 1 8 we s ee that t he m ean o r a v erag e

.

h eigh t o f th ese 66 ch ildren was n o t 6 5 5 inch es a s in the c ase .

,

o f t he parent s b u t inch es o r
,
inch es g rea ter and by that
, ,

m u ch nearer t he gen eral m ean of t he rac e than were t he h eight s

o f th eir parent s .

T h is p ri nc ip le o f regres s io n o r as it is so m et im es ca lled the

, , ,

“
d rag of t he rac e repres ent s t he p u ll o f t he anc es to rs
,
“ ”

beyo nd t he im m ed iat e parent s By th is we see that on tlze .

,

m lzole o ff s pr ing a re les s exc ept io na l th an their parent s ; o r

, ,
 3 6 TR A NS M I SS I ON
4

T h is m ed iocrity is , th eref o re a th ing always to b e rec ko n ed

,

with by t he breed er who h o pes to atta in u n if o r m s u cc ess wi th

im pro v ed s t rai n s He cann o t free h im s elf fro m it s i nfl u enc e
. .

We s hall s ee that its p u ll is n o t less than 5 0 per c ent The fail .

u re t o kn o w th is fact and t he willin g n es s t o res t t he c as e with

,

the imm ed iate par en ts and t o a s s u m e that lik e fath er lik e s o n

“

is the way o f h ered ity o r t o acc ept pu r i ty of b lood ( ped igree) a s

,

s yn o ny mo u s with u n iform i ty o f yp
t e — th is is t he o n e f e rt ile,

ca u se o f the great es t failu res in s t o ck breed ing The o nly s u re .

bas is o f u n ifo rm s u cc es s lies in a u n ifor m ly exc ellent anc es try

fo r at leas t fi v e o r s ix gen erat io n s back 1
T h en t he drag o f
“
.

t he rac e will bec o m e a f riend and n o t an enem y o f im pro v e

ment ; b u t n o m a t t er ho w exc ellent t he anc es try it can nev er
‘

, ,

eq u a l t he ex c ep ti ona l p a ren t I n o rd er t o m ak e t he m o s t o f him

.
,

th erefo re th is drag s h o u ld b e red u c ed t o a m in im u m

,
.

SECT I O N VI —T HE M EA S U R E OF H E R ED I TY

Now th is regres s io n is the p u ll of the anc es t ry back o f the

parent and it is the bes t argu m ent fo r the fact that i nheri tanc e
,

is part ly fro m t he rac e and n o t exc lu s iv ely fro m t he i m med iat e

parent C learly we need a m eas u re o f t he d egree o f resem blanc e
.
,

bet ween the o ff s prin g and the imm ed iat e parent s o that we m ay ,

kn ow ho w m u ch to cred i t to t he parent and ho w m u ch to cred it

t o t he back anc es try thro u gh regress io n S u ch a m ea s u re o f .

resem b lanc e bet ween m id parent and o ffs p ring will b e a good
-

mea s u re of h ered i ty and it is ca lled t he c oefi c ien t of lzeredi ty

, .

Th e coefi cien t of heredi ty Fo rt u n at ely th is i n v o lv es n o n ew

.

c o nc ept io n s and no n ew m eth od s The regressio n table is no th .

in g mo re n o r less than a spec ia l f or m of c orr eb ztion ta b le in wh ich ,

in s t ead o f in v o lv ing t wo charact er s in t he s a me s et o f ind iv id u als ,

we seek t he c o rrelat io n bet ween t wo s et s o f relat ed i nd iv id u als

wi t/z r esp ec t to the s a m e c lza ro oter .

Th u s , in t he tab le o f s tat u res we hav e in fact a c o rrelat io n,

tab le bet ween m id parent s and so n s with respect to s tat u re an d

-
,

i t s c o rrelat io n c oeffic ient ( r ) is f o r t/zem a c oeflzc ien t of b eredity

2
'

1 S L w f A
ee a o t l H dity t i f th i h pt
n c es ra ere , s ec . x v o s c a er .

1 Th ffi i t f
e coe c en l tio i ywh d t d b y th l t t
c o rre a on s ev er ere en o e e e er r .
 H E R E D I TY 8
4 7

The c oeffic ient of h ered ity is th erefo re no th ing m o re no r les s than

t he c o rrelat io n c oeffic ient ( r ) o btai ned fro m a regres s io n tab le
in wh ich t wo set s of ind iv id u als relat ed by d es c ent a re tab u lat ed
with res pect to t he s a m e charact er The m eth o d s o f fi nd ing
.

t he c o effic ient o f h ered ity are prec isely t he sa me a s th o s e al ready

d escribed fo r find ing t he c oeffi c ient of c o rrelat io n ; ind eed the ,

c o rrelat io n c oeffi c ient o f a regres sio n tab le is tlze c oefi c ien t of

I t is m an ifes t that th is c o rrelat io n tab le m ay b e c o n s tru ct ed

n o t o n ly bet ween m id parent s and o ff s pr i ng b u t bet ween fath er s
-
,

and so n s bet ween g randfath ers and g rand so ns bet ween m o th ers
, ,

and so n s (o r da u ght er s ) bet ween u nc les and neph ews b et ween

, ,

bro th ers and s is t ers bet ween bro th ers and bro th er s and ind eed
, , , ,

bet ween pers o n s c o n nect ed by a ny t ies of c o n sangu in ity what

ev er d i rect o r ind irect
,
I n each ca s e the c o rrelat io n c oeffic ient
.

bec om es a goo d mea s u re o f h ered itary resem b lanc e .

If a regres s io n ta b le b e c o n s tr u ct ed bet ween f a t/t ers and

m ot/t er s a c o rrelat io n wo u ld s t ill b e f ou nd th o u gh t he t wo a re
, ,

u ni t ed by n o b loo d li n es exc ept th o s e c o mm o n t o t he rac e i n

gen eral S u ch c o rrelat io n c o m es ent irely thro u gh s elect io n and

.
,

it s c oeffi c ient ( r ) is c o m mo n ly ca lled the c oeffi c ient o f cro s s

h ered ity o r asso rtat iv e m at ing
“
I t is a go od m eas u re Of t he
.

d egree o f selectio n in v olv ed in m at ing .

The tab le o n t he n ext page gi v es s o m e o f t he c oeffic ient s o f

h ered i ty that hav e been d et er m ined fo r d ifferen t relativ es .

Pear so n rem ark s “

We see th at o n the a v era ge t he i nt en sity
o f parenta l c o rrelat io n is abo u t to o f grand parental ,

abo u t to and o f frat ernal abo u t ,

to — t he
latt er c o rrelat io n being so m ewhat red u c ed wh en t he frat ern ity
c o n s is t s of mem bers o f o pp os ite sex es .
”

R egr ession coemcien t The regress io n c oeffi c ient h ere is

.

c o m p u t ed exact ly the s a me as t he regres sio n c oeffi c ient fro m

any o th er c o rrelat io n table and it has t he s ame u ses na m ely fo r
, ,

p r ed ic ti on that i s t o sa
y fo r exa m p,
le kn o wi n g t h e ,
d ev iat io n o f

a grou p o f m id parent s fro m th eir m ean what d ev iat ion s hall

-
,

we expect o n the part o f th ei r o ff s pr ing ?

The u se of t he wo rd “
regres s io n in t he t erm regress io n
”

c oeffi c ient is lik ely to lead t o c o n f u sio n We m u s t n ot as s um e .

4 8 8 T R A NSM I SS I ON

COEFF1C 1ENTS or H E R ED IT Y FOR D I FFER ENT R E LATION S HI PS ‘

R s LA r ION sm p
'
M AT R A L
E I CH A RA T R
C E CO R R LA T
E I ON

h
F at er a n d s o n l h
E n g is
F th
a er a n d d a ug ht er E gl i h
n s

M th
o er a n d s o n E gl i h
n s

M th
o er a n d d a ug ht er E gl i h
n s

M th
o er a n d s o n N th A m
or eric an d
I n ia n s
M th
o er a n d d aug ht er N th A m
or eric a n In di ans

S ire a n d fo a l Th h
o ro u
g b re dh o rs es

D am an d fo a l Th h
o ro u
g b re dh o rs es

G ran d s i re a n d o ffs p rin g Th gh b

o ro u re dh o rs es

G ra n d s i re a n d o ff s p rin g B t h
as s e o un d
B ro th er a n d b ro t h er E gl i h
n s

B ro th er a n d b ro th er N th A m
or e ric a n d
I n ia n s
Co lt a nd c o lt Th h
o ro u
g b re dh o rs es

S is t e r an d s is t e r E gl i h
n s

S is t er a n d s is t er N th A m
or e ric a n d
I n ia n s
Fill y and fi ll y Th h
o ro u
g b re dh o rs es

B th
ro er an d S is t er E gl i h
n s

B th
ro er an d s is t er N th A m
or e ric a n d
I n ia n s
C lt
o an d ll y
fi Th h
o ro u
g b re dh o rs es

Wh l o e b re th ren B t h
as s e o un d s

that c o effi c ient o f regres s io n is a d i r ec t m eas u re o f the p u ll

t he
o f t he back anc es try o r t hat it d i rect ly m ea s u r es t he d is s i m ilari ty
,

bet ween parent and o ffs pring a s S h o wn in t he regres s io n tab le .

I n fact we n o t e that wh en regres s io n is p erf ect th en t he c oeffi

, ,

c ient o f regres s io n is zero ; a n d wh en th ere is no regres sio n

( pe rf ect c o rr elat io n ) th e n t h e r egr es s io,n c oe ffi c ie nt i s l arg e .

T h is is bro u ght o u t i n t he d iagra m o n t he f o llo wing page .

Th is d iagra m is a g eo m etr ica l exh ib it o f t he r egres s io n tab le

o f s tat u res ( pag e 4 8 0 I i im p ly a r pr d ct i n f that tab l
) t s s e o u o o e ,

o m itt ing the freq u enc ies and p u tt ing in cro ss es r p

( )
x t o e t e

s ent t he m ea ns o f t he h o r izo nta l array s 2

c o lu m n
(
1 Pears o n , G ra m m a r S c i enc e, p p 4 5 8 460
of .
— .

1 T h e m ea n h
o f o ffs p ri n g i n t is t a b e is 68 in c l h es , an d we h a v e ass u m e d t h is
v a lu e t ob e, fo r t h e p res en t p u rp o s e, n ea r en o u gh t o t he m ean o f mid -p a ren ts an d
o f t he ra c e t o t a e t he k h
o rizo n ta l an d v ert ic a l i n es ma r e 68 as pas s in g t ro u gh l k d h
t h e m ean o f t he ta b le .
490 T R A N SM I SS I O N
s lope is o u r c oeffic ient o f regress io n and its v a lu e is expres sed ,

by the rat io o f P N to P 11] .

Ga lt o n fi nd s th is rat io wh en d ealing with mid parent s and

1
,
-

m id filial s ta tu r es to b e appro x imat ely as 2 to 3 ;

-
,
that is to
t he fi lia l d ev ia t io n fro m P ( the c o mm o n mean ) is o n t he
sa
y ,

a v erage o n ly t wo th ird s as wid e as the m id parenta l d ev iat io n -

.

I call th is rat io o f 2 t o 3 [he s ay s ]t he rat io o f fi lial regress io n

’
.

I t is t he pro po rt io n in wh ich t he s o n is o n the av erage les s , ,

exc ept io na l than h is m id parent T hat is the d ev iat io n o f t he

-
.
,

s tat u re o f ch ildren fro m t he m ean o f t he ra c e is o n ly a b o u t t wo

th ird s as wid e a s that o f th eir m id parent s -

.

S ECT I O N V I I — T H E M EA N O F T H E O FFSPR I N G N OT
N EC ESSA R I LY T H E SA M E A S T HE M EA N OF
T H E PA R E NTA G E

At fi rs t th o u ght it wo u ld seem ax io m at ic that o n the av erage , ,

t he o ff s pr ing a s a wh o le wou ld b e the s a m e a s t he parent s ,

u n les s t he rac e is u nd erg o ing chang e I n t he tab le o f s tat u res .

,

h o wev er by c o m paring c o lu m n s 1 6 and I 7 ro w 0 we see that

, , ,

t he m ean o f a ll t he parent s wa s inch es ( n o t c o u nt ing ex

tremes o r ,
inch es inc lu d ing extremes ) wh ile t he mean of ,

the ad u lt ch ildren wa s b u t inch es inch es inc lu d ing

extr em es ) T h is ind icat es a lo s s i n s tat u r e o f o v er a half inch
.

in a s ing le gen erat io n u n les s s o m e o th er i nfl u enc e is at wo rk to

,

c o u nt eract th is d is crepa ncy T h a t c o u nt erac t ing infl u enc es a re

.

at wo rk we s ha ll s h o rt ly d is c o v er b u t the fact rema in s that the ,

tlze ri n
i w
'

mea n f
o fip g
o s s s el d o m i d e n tic a l i t/c t/ze m ea n of til e

p a r en tag e . T h is fact is to b e c o n s tr u ed a s m ean ing o n e (o r bo th )

of th ings
t wo eith er that t he rac e is u nd erg o ing tran s f o r m a
,

t io n o r els e that all g rad es are n o t eq u ally pro d u ct iv e

, .

I n th is c o nnect io n it is t o b e n o t ed fir s t that is n o t to , ,

b e tak en a s t he m ean o f t he g en er a ti on t o which th es e m id pa rent s -

belo ng ; that mean m ight ha v e been eith er les s o r m o re becau se ,

n o t a ll m em b er s o f a g en erat io n bec o m e parent s n o r is t h e ,

parent po p u lat io n a rand om d raft fro m the generat ion s of the

m id pa rent s
-
.

G lt N t l I h it
1 a p 97
on, a u ra n er an c e, . .
 H E R ED I TY 49 1

No fact is be tt er kno wn a m o n g s tat is t ic ian s than that wiv es

d iffer fro m da u ght ers and mo th ers d iffer fro m wiv es — wh ich ,

mean s that all wo m en ( da u ght ers ) d o n o t m ar ry and that n o t ,

all wiv es a re m o th er s ( select io n ) ; that is to say parent s a re ,

a select ed d raft fro m the ent ire po p u lat io n and we s h o u ld n o t ,

expect t o find th ei r m ean t he s a m e a s that o f th ei r o ff s pr ing ,

wh ich c lo sely appro ach es that of t he gen eral po p u lat io n .

SECT I ON V I I I — EXT R EM ES O F A R AC E R ELATI V ELY LESS

PR O D U CT I V E T HA N T H E M EA N S

I n t he tab le o f s tat u res a s tro ng t end ency is ev id ent t o ward

increas ed h eight and a s t ill s tro nger o n e t o ward d ecreas ed
s tat u re ( s ee s ect ix Pro gres s io n
.
,
Th is being tr u e t he rac ia l ,

d is t rib u t io n wo u ld rap id ly s pread if n o t ent irely d iv id e int o t wo

, ,

rac es g iant s and d wa rf s u n less prev ent ed by som e princ ip le s u ch

, ,

as nat u ra l s elect io n T h is princ ip le in th is part ic u lar in s tanc e is

.

ev id ent ly relat i v e inf ert ility a princ ip le eas ily d ed u c ed in at

,

lea s t two way s

1
. On e h u nd red and three ch ild ren a re rec o rd ed at o r belo w
and o n ly o n e parent belo w ( s ee tab le o f s tat u re s ) .

C lear ly mo s t s h o rt ch ildren d o n o t bec o m e par ent s els e the rac e ,

wo u ld ra p id ly d egen erat e as t o s ize T h is agrees with c o m m o n

.

o b serv at io n wh ich is that d warf s d o n o t m arry

,
Wh en h owev er
.
, ,

t he princ ip le is app lied t o d egen e racy and cr im e t he ca s e is d if ,

feren t fo r crim inals o ft en prod u c e m o re than thei r n o rm a l rat io

, ,

and ma ny o f th eir o ffs pring by the princ ip le o f pro gres s io n

,

( s ee s ect i x
). a r e fr,
ightf u l d e g e n erat es .

To what ext ent g iant s m arry is n o t s o c learly i nd icat ed by

th is table b u t that they are les s fert ile than the av erage wh en
,

th ey d o m arry is c lea rly s h o wn by c o m paring c o lu m n s 1 6 and 1 7 .

2
. T he av era ge o f fert ility in t his tab le a s a wh o le is a lm o s t
exact ly 4 } ( 9 2 8 + 2 0 5 ) per m id parent Th is a v erage is well
-
.

s u s ta i n ed in t h e lo wer and m idd le s tat u res ris ing in m any ,

cases to 5 per m id parent ; b u t it rapid ly les sen s in the h igh er

-

s tat u r es fo r fro m 7 0 5 u p it is appro x im at ely 3

,
. T h is to o .
, ,

agrees with c o m m o n experienc e na mely that ex t rem es of a ra c e

, ,

a r e ener a ll
g y le ss
f er ti le tit a n tb c m ea ns I nd eed .it is c o mm o n,
ly
49 2 T R A N SM I SS I O N

f that
er t i l i ty fi x es typ e u n le s s it s eff ect s happ, e n t o b e o v erc o m e

by so me o th er fo rm o f s elect io n ; that is to say t he type o f ,

h igh es t fert ility will b ec o me mo s t n u mero u s and will therefo re ,

nat u rally d et erm ine t he type o f the rac e .

I t is ev id ent ly t he les sen ed fert ility o n the part o f extremes ,

and t he h igh er breed ing po wers o f the m ed ioc re ind iv id u als ,

tha t in nat u re as s is t select io n in h o ld ing t he pr inc ip le o f pro

g re s s io n i n ch e ck th u s g e n e ra ll y ,
b u t n o t a lw ay s pr ev ent i ng, ,

t he s p litt i ng u p o f rac es i nt o s m a ller v ariet ies ; ind eed type is ,

m ai n ly t he res u lta nt o f relat iv e f ert ility and s elect io n — v ery ,

largely o f t he f o rm er T h is is why breed ers o f h igh ly i m prov ed

.

rac es mu s t need s loo k well to fert ility fo r at that po int th eir ,

fi rs t tro u b les will aris e all regress io n tab les in d icat in g that they
,

will n ev er s u ffer fro m lack o f v a riab ility as seen in t he fo llo wing ,

s ect io n .

S ECT I ON IX — PR OG R ESS I O N . PA R ENT S I N G EN ER A L PR O

D U C E A FEW I N D I V I D U A LS MO R E EXT R EM E
T HA N T HE R A C E

What tru e o f a v erages is n o t nec es sari ly tr u e o f ind iv id u a ls

is
o r o f s elect ed gro u p s R egres s io n app lies t o t he ma s s n o t to
.
,

t he s epa rat e ind iv id u a ls that c o m po s e it 1

.

Parent s in general pro d u c e i nd iv id u a ls bo th inferio r and

s u per io r t o th em selv es f o r m ing a freq u ency d is tri b u t io n wh ose
,

m ean lies so mewh ere bet ween that o f t he parent and t he gen eral
m ean of the rac e B u t t he i n d i v id ua ls ext end bo th way s fro m
.

th is mea n and so me of th em lie well b eyo nd t he ra nge repre

s ent ed by t he parentage On th is po int s ee any ro w in any
.

regress io n ta ble as ro w e in the table o f s tat u res ( page

,

F o r exa m p le in the cas e at h and b u t 5 m id parent s and po s

, ,
-
,

s ib l o n ly 4 ( 1 0 o r 8 par ent s ) a re ab o v e t he h eight i nch e

y , s

( S ee ro w b ) b u t 3 1
,
ch i ldren a r e r ec o rd ed at o r ab o v e ( s ee

c o lu m n s 1 4 and T he n u m be r o f extreme ch ildren is th u s

m o re than three t im es a s great a s t he n u m be r o f extreme

1 Fo r exa m p l e, we h av e s h o wnh t a t , o n t he a v era ge, o ffs p ri n g a re d

m o re m e ioc re
t h an h
t eir p a ren ts ; b u t fo r a h igh ly l s e ec t e d o ffs p rin
g ( 7 2 -
in c h h
s ta t u re) we s o u ld

fi n d t he pa ren ts to be n ea rer medio c rity t ha n the o ffs

p rin
g .
 HER EDITY 4 95

pec ent with a m od e of 1 1 an d an u pper lim it (o n e ear)

r , ,

o f 1 4 pe c ent
r — mo re acc u rat ely
,

T he res u lt o f on e year s select io n m ad e li tt le ev id ent im pres

’

s io n Th e s ec o nd y ea r s cro p wa s n o t m u ch b ett er T he m od e
’
. .

was t he sa m e t he a v erage s light ly lo wer b u t t he d is tri b u t io n

, ,

became so mewhat ext end ed with the appea ranc e of two h igh er ,

v al u es repres ented by o n e ear each .

I n the t hi rd year wi th s t ill bett er seed ,

t he d is tr ib u
t io n ext end s s t ill farther b u t t he n ew v al u e is d own n o t u p
, , .

W e n o te howev er a genera l increas e o f t he h ig her v alu es and

, , ,

the m od e has m o v ed u p a no tch 1

.

I n t he f o u rth y ear wi th s t ill bett er s eed t he

lo wer v alu es les s en and so me d ro p o ff ent irely On e n ew v alu e .

appea rs A ll the u pper freq u enc ies are increased ; t he m o d e

.

has go n e u p t wo n o tch es and we n o w ha v e n o les s than 2 8 c a rs

,

a s goo d o r be tt er than t he extrem e ea r o f 1 8 96 .

T he sa me princ ip le c o nt in u es in 1 90 1 wh ich was an exc ep ,

t io n a lly g oo d y ea r fo r pro t ein and t he th eo ret ica l m o d e go es u p

,

nearly t/zr ee po int s reach ing a par with t he s ingle exc ept io nal
,

ea r o f t he f o u ndat io n s t oc k I t so happen s that th is y ear t he

.

n u m b er o f ca rs exam ined was t he s a me as that o f t he fo u ndat io n

s t ock and o f th ese ears n o less tlza n 47 o r 2 8 per c en t wer e , ,

tlze eq ua l of tlze ex c ep ti on a l fi rs t ea r The effect s o f s elect io n .

sett le back s light ly t he n ext y ear b u t in t he s u cc eed ing s ea s o n ,

los t gro u nd is rec ov ered and th ere is prod u c ed t he exc eed in gly
ex c ept io na l ea r wh ich ha s s inc e pro v ed a remarkable
,

parent .

T he pri nc ip le o f pro gr ess io n is s t ill f u rth er illu s t rat ed by the

next table s h owing t he effect of select io n bot/z ways T h is is
,
.

t ak en fro m D r Ho pk in s s o rigin al dat a in breed ing fo r h igh o il

.
’

and for lo w o il fro m the s am e f ou ndat io n s t o ck The s t u d ent .

s h ou ld n o t e the d ec is iv e m ann er in wh ich th es e gen e rat io n s

s epa rat e th em s elv es fro m t he f o u ndat io n s t o ck and fro m each

o th er as s elect io n go es o n N o th ing c o u ld s h o w m o re c o nc lu
.

s iv ely that u nd er i nt en s e and p er s is t ent s elect io n n ew v al u es

appear freely as t he rac e b ec o m es lib erat ed fro m t he h ea v y drag of

1 It c an be s een b y in s p ec t io n t h at the t h eo ret ic a lm d o e is n o t so h igh as t he

em piric a l mo d e, 1 2 .
498 TR AN M
S I SSI O N

is no t no w g roh a p x ii; lea d s t o t he c o nc lus io n

wi ng les s ( s ee c .

tha t t he u p pe r li m it s i n t his b reed in g e x pe ri m en t w ill b e se t b y

s m e f a c t o r o t he r th a n t he fa il re f v a ri a b ilit y ; i n d ee d i t is '
t o
o ,

t he o p in io n o f t he writ e r t ha t t he p ri n c ip ie o f p r gr es s io n is a b le o

a lway s t o a ffo rd a ll t he m a t e rial t he b reed e r w ill n eed a n d t ha t ,

t he li m it s o f i m p ro v em ent will b e se t wh en t h ey a re s et by , ,

so m e b io lo gi c a l m ech a n ic al o r o t h er c o n s id e ra t io n s ent irely

, ,

as id e fro m t he fa ilu re o f v a ria b ility t o p rese n t n ew u ppe r v alu es

o n wh ic h to b as e se lect io n T he re m a rk a b le fa c t a b u t p r gr es
1
. o o

s io n is th a t t he d is t ri b u t io n s a re n t d is t o rt ed b u t a re a ll typ ic a l o

o f t he ra c e ( see t a b le p a ge ,

Th is fa ct o f pro g res s io n b et ray s a u n iq u e p ri n c iple i n hered ity ,

o r ra th e r in v a ria b i lity b eca u s e pro g re s io n is o v e r a ga i n s t a n d

,
s

in s p it e o f t he d rag o f t he rac e T ho s e ind i id u a ls th a t ha v e

”
. v

o v e rlea ped t he li m it s o f t he rac e ha v e n o t o n lv e xc eed ed th ei r

o wn pa rent s b u t by m u cli m o re h a v e th ey exc eed ed t he c o m pa ra

,

t iv e m ed ioc rity o f their o t h er a nc es t o rs Pro g res s io n c a n no t

?

th erefo re b e exp la ined by any p rinc ip le o f m ea n pa rent age

, .

I t res t s u p on a p rinc ip le f u nd a m entally d is t inct and is t o b e ,

rega rd ed as t he res u lt o f th o se fo rt u it o u s c o m b ina t io n s o f phy s io

lo gica l u n it s wh ich we m ay ex pec t t o o c c u r fro m t im e to t ime in
t he c o m p lic a t ed proc es ses att e nd in g repro d u ct io n and d i fferent i
at io n a n d o n wh ich mo re wi ll b e sa id in Sect io n X I I o f t his
,

chapt er .

T h is s u gges t s what will b e la t er f o u nd t o b e a fa ct n a mely , ,

that in a la rge sen s e h ered ity fo llo ws t he la ws o f pro b a b ili ty and ,

in proc es s o f t i me we m ay expect a ll p os s i b le com b i na t i o n s o f t he

elem ent s that m ak e u p charact er s and o f t he ch a ra ct ers that ,

m ak e u p t he rac e t he larges t pro po rt io n o f wh ich will c lus t er

,

abo u t a c o m mo n po int wh ich we ca ll t he type b u t a few of , ,

wh ic h will in ev it a b ly appear at t he extrem e lim it s o f t he range

o f po s s ib ilit ies .

1 The g rea t es t m en a c e to e x t re m e im p ro v em en t is , as ha s a rea l dy b een sa i d ,

l e s s en e d
fe r t i it l y
A c c o r in g . d to Pea rs o n a ll e v i d en c e
p i n t s t o t h e fa c t t
o h a t v a ria

b ilit y wi l l n e v er be re d uce d b y m o re t h an ab o u t 1 1 p e r c en t . S e e G ra m m a r of

S c ie n c e, p 4 8 3 . .

1 T h e s t u e n t wi d ll no t fa i t ol n o te h t th
t a e a n c es t o rs o f t he exc ep t io n a lp a r en t

a re o f n ec es s it ym o re d
m e io c re t h an th t p a a ren t . T h y will th
e en e x e rt t h ei r in flu

e n c e a g a in s t , n o t in fa v o r o f, p ro gres s io n .
 H E R E D IT Y 4 99

SECT I ON X —THE EXC EPT I O N A L I N D I V I D U A L A R I SES

EIT H ER FR OM M ED I OC R I TY OR F R OM T H E
EXC EPT I ON A L PA R EN T

F u rth er i n sp ect io n table o f s tat u res s h o ws that of the

of t he

7 2 ch ildre n r e c o rd e d ab o v e s i x f eet i n h eig h t ( s ee c o l u m n s 1 3 ,

14 , 4 2 o r o v,
e r o n e ha l f w e re pro d u c e,
d by m id par ent s -

inc hes o r u nd er ; that appro x imately 2 2 were pro d u c ed by

mid parent s les s than o n e inch ab o v e t he m ean o f t he ra c e
- 1

i nches s ee ro w 0 c o lu m n 1 7 ) that n o les s than 1 2 o r o n e

, , ,

si xth o f all wer e pro d u c ed by m id parent s rec o rd ed at o r b elo w

,
-

t he mean o f t he rac e ; an d that 1 was pro d u c ed by a inch -

mid parent
-
.

F ro m th is we s ee that exc ept io na l ind iv i d u als may aris e eith er

fro m exc ept io nal o r fro m med iocre parent s T he pro bab ility .
,

h o wev er is great ly in fa v o r o f the fo r mer S ix 7 2 5 inch m id

, . .
-

parent s prod u c ed 1 3 exc ept io nal ch ildren o u t o f a t o ta l of 1 9 ,

c o n sid eri ng ev eryth ing abo v e s ix feet as exc ept io n al Of th is .

n u m ber 6 o r o v er 30 per c ent exc eed th eir own parent s in

, ,

h eight Th ou gh the 69 5 inch m id parent s prod u c ed a h igh er

. .
- -

n u m ber o f except io n ally ta ll ch ildren 4 1 par e nt s w e re

in volv ed i ns t ead o f 6 Th is is les s than 3 per c ent o f t he t o ta l
.

c hildren in s t ead o f 6 8 per c ent as in t he cas e o f t he ,

pro geny of t he taller parent s .

I t is at th is p oint that t he po li t ica l s c ient is t and t he threm

mato logis t rec ogn ize d iff erent p rinc ip les Bo th a re int eres t ed in .

exc ept io n al i nd iv id u a ls A s we ha v e seen th ey m ay b e had

.
,

eith er fro m t he gen era l p o p u lat io n o r fro m a h igh ly s elect ed

pa rentage The breed er ch oo ses the latt er beca use he cann o t

.

a ff o rd to su ppo rt so large a po p u lat io n fo r s o few exc ept io nal

i nd iv id u a ls He tak es t he h ig hly s elect ed parentage becau s e
.

t he p r op or t ion o f extrem e exc ellen c e is h igh er and b eca u se it s ,

dra g is less He is d es iro u s o f u s ing m in im u m n u m bers fo r

.

ec o n o mic rea so n s .

The po lit ica l s c ient is t is lim ited by n o su ch c o n s id erat io n s .

I f he reso rt t o select io n (elect io n ) each t im e a r u ler is to b e

1 I l di g
nc u ly
n on h l f f th ff p i g p d d b y p t
one a o e o s r n dd t
ro uc e aren s r ec o r e a
 00 T RA N S M I SS I O N
5

chos en he will alway s ha v e good mat eria l at hand and as he is

, , ,

n o t s pec ially int e res t ed in pro geny he is no t c o nc ern ed wi th t he ,

dra g W hat he want s is i nd i v id u a l ser v ice

.
”
.

If o n t he o th er hand he ad o pt s t he p lan o f h ered itary so v et

, ,

eign ty he will d eal wit h few fa m ilies at a t i me ; and whi le if

, ,

they a re extrem ely well bred to b egin with a large p ropor tion -
,

will b e exc ept io nal yet a glanc e at t he u pper lines o f th is tab le

,

will b e en o u gh t o ind icat e that he will b e c o nf ro nt ed by a good

many h ered ita ry ru lers who a re far fro m exc ept io na l ( see es pe
c ially ro w e) He ha s tak en a u seless haza rd and th is is t he
.
,

inev itab le hand icap o f an h ered itary m o narchy F ro m t he s t and .

po int o f ev o lu t io n t he pri nc iple is wro ng .

The abo v e o u ght to mak e it c lear why t he breed er an d t he

po lit ic ian s h o u ld ad opt oppo s it e meth od s I f s erv ic e al o n e is .

want ed it is bett er t o find it than t o breed i t ; and that is why

,

it is o ft en bett er to b u y a part ic u lar type o f an im al than t o

attem pt to pro d u c e it es pec ially if t he type is at all u n u su al as
, ,

in t he c as e o f t he fi re ho rs e .

SECT I ON X I FR AT ER N A L V A R IA BI L I TY , OFFSPR I NG
OF SA M E PAR E NTS NOT I D ENT I CA L

The f pring o f lik e parent s a re n o t o n ly u n lik e b u t the s uc

of s ,

c es s iv e o ff s p ring o f t he so m e parent s v ary wid ely T he o nly .

data c o m piled o n th is im po rtant fact a re c o ntained in the table

o n the f o llo wing page fr o m G alto n s s t u d ies in s tat u re
’
, .

This tab le pres ent s all the charact eris t ic s o f t he o rd inary

regressio n table b u t in a d egree slight ly less pro no u nc ed T h is .

s h o ws that t he s a me la ws o f r egres s io n and pro gr ess io n apply

wi t/t i n t he fa m ily a s app ly b et ween fa m ilies .

A llu d ing to th is s ign ificant fact G alt o n rem ark s : 1

,

I t ppe rs th t th e i
a a di c t h a d it y l t i
er b e twee the pe so al
s no re ere ar re a on n r n

p a re t
ns d t he p an l c h ild e c ep t p h p th gh li ttle k ow c h
ers o n a , x er a s ro u -
n n an

ne ls o f ec d y im po ta c e b t t h a t the m a i li e f h
s on ar r dit y co e
n , u n n o ere ar nn c

t io n un ite the et o f eleme t o t o f wh ic h the pe

s s s l p re t h d b ee
n s u rs o n a a n s a n

evo v l ed wi th th e et o t o f wh i h th e pe so l h i ld w
s u e o l ed c r na c as v v .

T h i i why it i
s s im p t t i h e d it y i q i y t d e l wit h f
s so o r an tern iti n re ar n u r o a ra es

1 l
G a t o n , N a t u ral I n h eri t a n c e, pp . 1 9- 2 0 . I ta i c sl are m in e .
5 0 2 TR A N SM I SS I ON
r a t/ter Ma n wi th i n d i v i d u a l s , and wi th la r ge fr a tern i t ies ra t h er t h an wi t h
s mal l o n es . W e ou gh t , fo r ex a m p le, t o c o m pa re t he
g p r ou c o n t a in in
g b o h
t

p a ren ts a n d a ll t he u n c l es a n d a u n ts wi t h h t a t c o n ta i n i n g a l l t he c h ild ren .

Here is y gis t o f the wh o le m atter s ho wing t he f o lly

t he v er ,

o f d ea ling with ind iv id u a ls in q u es t io n s o f breed ing A ll t he bes t .

ev i d enc e s h o ws th a t s elect io n ba s ed u p o n t he i n d i v i d u a l with o u t ,

regard to the gro u p to wh ich he belo ngs will n ev er res u lt in ,

c o nc entrat io n o f exc ellenc e I t is o n ly by pers is t ent s elect io n .

,

based o n t he gro u ps as a wh o le ( p u ri ty o f ped igree in t he s en s e

o f u ni fo r m ity o f typ e ) th a t we s h a ll ev er f rc e ev en t he fa m ily
,

fro m t he d rag o f t he rac e a nd m ak e rea l pro g res s in im pro v ement, .

Haro ld and M is s R u s s el t he s ire a n d d a m o f M a u d S were , .

,

o wn ed at W o o d b u rn fo r m a n y y ea rs b u t o f all th ei r ge t o n ly o n e ,

M au d S d ev elo ped h ig h s peed Vi hy T he q u es t io n canno t b e

.
,
.

an s wered any f u rth er than i t c a n b e i nferred fro m t he p rinc ip le

j u s t s tat ed and fro m t he well kno wn meth od s o f c lea v age o f t he -

n u c leu s in c ell d iv is io n and in m a t u rat io n ; b u t with th es e fact s

in m ind we s lzou ld i mrd ly exp ec t tlza t t wo i d en t ica l i n d i v id u a ls
,

wou ld ev er a r i s e, ev en f r om t/ze sa me
p a r en ts .

Th e ear l ier offs prin g li v e lon ger t han do t he you n ger c hil dr en
of t he same par en ts . Ha v ing es ta b lis h ed t he fact that s u cc es s iv e
f pring o f t he s a me parent s a re d ifferent th ere rem ai n s the
of s ,

tas k of d eterm in ing to what ext ent th is frat ernal v ariability is

heterogeneo u s a n d to what ext ent it m ay b e c o rrelat ed with age
,

o r wi th s o m e s im ilar c i rc u m s tanc e t hat t end s t o thro w t he o ff

s p ring int o a r egu la rly grad ed s eries o f s o m e s o rt .

On th is po int we kn o w b u t litt le To t he eye th is v ariab ility .

appear s q u it e h et ero gen eo u s b u t s t u d ies in lo ngev ity fo r ex , ,

a mp le ha v e f u lly es tab lis h ed t he fact that in m a n t he old er ch il

,

,
h v v —
dren o n t e a era ge li e lo nger that is ha v e lo nger liv es , ,

than d o t he y o u nger o n es o f t he sa m e fa m ily Th is d i fferenc e .

,

as b et ween t he o ld es t a n d t he y o u n g es t a m o u n t s t o n o les s t ha n ,

fou r y ears Wh eth er t he s a m e o r a s im ila r d ec line tak es p lac e

1
.

in o th er charact ers and fa c u lt ies o n ly exha u s t iv e s t u d ies c o u ld

d et erm in e T h ere is n o reas o n t o d o u bt that general p ri nc iples
.

app ly eq u ally to m an and t o o th er an im a ls exc ept ing in s o far ,

1 S ti l
ee a r I h it f th D
c e on ti f Lif
n er by B t d P
anc e o e u ra on o e, ee o n a n ea rs o n ,

B i m t il
o e r V l I P t I pp 5 —
'
a ,
76
o .
, ar , . 0 .
 H ER EDITY 5 3
0

as s o c ial and o th er s omewhat art ific ial c o nd itio ns of life m ay

interv en e and we s hall an x io u s ly a wa it t he resu lt s o f f u rth er
,

in v es t igat io n int o the charact er and range o f d ifferenc e bet ween

o ff s pring o f t h e s a m e parent s .

I n di v idu ali ty What ev er t he resu lt s o f in v es t igat io n in th is

.

d irect io n and what ev er grad u al ad v anc e o r d ec lin e m ay b e

,

es tab lis h ed a m o ng t he m em b e rs o f t he s a m e fa m ily on Me a v er

a e t he fact rem a in s that v ariab ility is largely h et er o gen eo u s as

g ,

bet ween i nd i v id u a ls and that a m ark ed ind iv id u ality perv ad es

,

all o ff s p ring ei th er o f t he s a m e o r o f d ifferent parent s

,
.

T his les s ened d ev iat io n bet ween m em bers o f t he s am e fam ily

as c o m pared with d es c endant s in gen era l is d u e t o t he fact that ,

a m o ng bro th ers no t o n ly bo th im med iat e parent s b u t a ll a n ces

,

tor s a re id ent ica l The d ifferenc es that d o ex is t with i n t he s a m e

'

fa m ily s erv e to S h o w the wid e d i v erg enc ies pos s ib le wi t/z t/ze sa m e
lzer ed i ta ry elem en ts alth o u gh in s t u dy ing ad u lt s s o m e a llo w
, , ,

an ec s m u s t alway s b e m ad e fo r d ifferenc es in d ev elo p m ent d u e

t o ext e rn a l ca u s es Wh ile m em b ers o f t he s a me fam ily a re in gen
.

era l reared m o re n early a lik e than a re m em b er s o f d iff eren t fam

ilies yet in a larg e s en se ev ery ind iv id u a l ha s a life h is t o ry o f his

,

o wn q u it e d is t inct and in m any s en s es d iff er ent fro m that o f any

o th er i nd iv id u a l o f h is o wn o r o f any o th er gen erat io n N o w t his .

life h is t o ry a ffect s d ev elo p m ent and acc o u nt s fo r s o m e o f t he

d ifferenc es b et ween ad u lt ind iv id u als N o t all of the v ariability .

with in a fa m ily can th eref o re b e a s s ign ed t o h ered itary in flu

, ,

en c es b u t that a large s hare o f it is d u e to s u ch infl u enc es is

,

rend ered extrem ely lik ely by the well kn o wn fact that s u cc es s iv e -

ov a,
s per m at o zo a o r p o llen gra in s fro m t he s a m e i nd iv id u a ls
,

a re n o t a lik e eith er i n th ei r g en es is o r i n th ei r b eha v io r aft er

ward .T he v ery m echan is m o f m at u rat io n s tro ng ly s u gges t s

pro fo u nd q u alitat iv e d ifferenc es and t end s to m o d ify o u r as
,

s u m pt io n that a ll ch ildren o f t he s a m e parent s po ss ess id ent ica l

h ered itary elem ent s T he experienc e o f breed ers ev ery wh ere is

.

that Off s pring o f t he s a m e ind iv id u a ls are n o t S light ly d ifferent

b u t in gen era l th ey a re wid ely d iff erent Wh eth er and to what .
,

ext ent th es e d i ff erenc es can b e less en ed by s elect io n and by

,

relat iv e p u rity o f anc es tral ga met es a re q u es t ion s o n wh ich light

is s o rely n eed ed .
5 4
0 T R A N SM I SS I O N

SECT I ON XI I —C HA RA CT ER S T EN D T O C OM BI N E I N
D E FI N ITE M AT H E MAT I CA L PR OPOR T I O N S

T he t d ent wo rk ing with large po p u la t io n s m u s t b e s tr u ck

s u

by t he rem arkab le S im ilarity o f the general feat u res o f a ll i re

q u eney d is tribu t io n s and arrays B ro ad ly int erpret ed t his s u g .
,

ges t s a s tro ng m ath em at ic al bas is in repro d u ct io n .

I n a ll f o r m s o f life hal v ing and d o u b ling a re bas ic proc esses .

T he n u m ber t wo th erefo re as a m ath em at ical c o nc ept io n lies

, , ,

at the bo tto m o f a large s hare o f o u r b iolo g ic al pro b lem s es pe ,

c ially th os e o f v ar iab ility and a litt le c o n s id erat io n will s h o w

,

t hat t he u s u a l fo r m of the freq u ency d is trib u t io n is the nat u ral

res u lt o f t he repro d u ct iv e proc es s es —ind eed that t he fact s of , ,

v ar iab ility la r el th ou gh n o t exc l u s i v ely f o llo w t he o rd inary

g y , ,

m at hem at ica l la ws o f c o m b inat io n s and pro bab ilit ies .

Th e mix in g of pu re for ms To illu s t rat e th is f u nda menta l fact

.

let u s u nd ertak e to f o llo w t he h is t o ry o f two charact er s bro u ght

t o get her for t he fi rs t tim e an d the m ann er in wh ich th ey will
,

nat u ra lly appear in the o ff s p ring .

To p u t the m att er in it s S im p les t fo r m let u s s u ppose a h erd ,

o f p u re b lack s t o m eet and m ing le wi th a h erd of eq u a l n u m bers

( )
o f p u re red s and that th ey breed t o geth er with o u t r es tra int —t hat
, ,

is wi t/tou t s elec t i o n T h ey will th en m ate ind is cr im inat ely ; t hat

, .

i s to s ay a b lack f em ale will m at e i nd ifferent ly with a b lack o r

,

a red m a le s o m et im es wit h o n e and s o m et im es with t he o th er

,
.

T h is being t ru e o n e ha lf the o ff s prin g o f t he b lack females

,

will b e p u re b lack (d es ignat ed by B ) and o n e ha lf will b e m ixed

1
,

b lack and red (d es ignat ed by B R ) .

T he s a m e pri nc i p le app lies t o the red f em ales wh o s e progeny ,

will in lik e m ann er b e eq u ally d iv id ed b et ween t he m ix ed o ff

, ,

s pri ng and t he p u r e r ed s .

E xpres s ed in tab u lar fo r m we s h o u ld th en ha v e

Fo r ev ery 2 00 o ffs p ri n g o f b lac k fem a les 1 00 B 1

1 00 BR
1
Fo r ev ery 2 00 o ffs p r i n g o f red fem a les 1 00 BR 1 00 R
o ffs p ri n
1
T o t al d is t rib u t io n , 4 o o g 1 00 B 1
2 00 BR 1 00 R
1
I n p ro por t io n o f B 1
2 BR R

1 h l d
T is p rin c ip e was fi rs t a n n o u n c e b y Q u et elet , 1 84 6, in Let t res s u r la t é o rie h
d es p ro b a b ili t es
'

S ee V ern o n , V a ria t io n in A n im a s a n d P an t s
.
p 1 2 l l , . .
 0 6 TR A N SM I SSI ON
5

bino m ial B R exp and ed t o t he fo u rth po wer acc o rd ing t o t he

b ino m ial th eo rem .

S u cceedin g popu lati on s f oll ow t h e law of t he b in omia l t h eor em

ex cept as in t erru pted b y sel ect ion or differ en ce s in f er ti li ty In .

t he breed ing o f th is th ird generat io n in ter s e we fi nd the n u m be rs

bec o m ing rap id ly c o m p licat ed b u t fro m the fact that it alway s ,

follo ws t he bino mia l t heo rem we can wri t e t he n o rm al d is t rib u

t io n fo r t he f ou rt h gen era t io n o f d esc end a nt s o f any pai r o f
characters a s fo llows 1

A nalyzing th is
fo u rt h gen erat io n po p u lat io n we find
-
,

1 T hat n o less th a n n in e c o lo r c o m b inat io n s a re repres ent ed

.
,

ran gi ng all the way fro m p u re b lack t o p u re red .

2 T hat t he freq u ency n u m be rs repres ent ing t he v ario us c om

.

b in at io n s 1 8 2 8 5 6 7 0 5 6 2 8 8 1 f o r m a s ym metrical fre
, , , , , , , , , ,

qu enoy d is t rib u t io n who s e t o tal is 2 5 6 o n ly t wo ind iv i d u als o f ,

wh ich a re p u re .

3 .T hat f u t u re br e ed i ng s w o u ld b e c o m e rap i d ly c o m p li cat ed ,

b u t that we s h o u ld a l way s ha v e o n e p u re b lack an d o n e p u re

red with a l l p oss i b le c om b i na t i ons between t he t wo
,
.

4 T hat
. t h e act u a l c o lo r c o m b i nat io n o f t he ind iv i d u al cann o t
in mo s t ca ses b e inferred fro m appea ranc es F o r exa m p le th ere .
,

is b u t on e rea l black and b u t o n e real red in t he wh o le po p u la

B “
t io n Ho wev er t he 2 8 E A will look lik e black s becau s e t here
.
, ,

hav e been S ix inf u s io n s o f b lack t o b u t two of red wh ile the ,

rev ers e is tr u e o f ano th er eq u al n u m b er 2 8 B R The 7 0 B R2 6 ‘ ‘

,
.
,

wh ich is t he larges t n u m ber o f all c o n s t it u t ing n early o n e th i rd ,

o f t he ent i re po p u lat io n is eq u ally ba lanc ed as t o c o lo r t end en

,

c ies b u t will app ea r to b e of the c o lo r that is m o s t n o t ic eable

, ,

in th is cas e pro bab ly a dark red .

Comb in ati on s of t hree cha r act er s T ho u g h t he n u m ber s bec o me .

mo re rap id ly c o m p licat ed t he s a m e pr inc ip les app ly wh en d ea l

,

in g wit h three o r m o re cha ract ers F o r exa m p le s u ppo s e we .

,

i ntrod u c e a th ird c o lo r wh it e W e s ha ll th en hav e as t h e res u l t

,
.

of t he fir s t m at ing t he f o llo wing :

1 The s t u d en t ca n eas i ly v erif y th es e figu res b y t he p l l dy

an a rea out li d
ne .
 HER ED ITY 0
5 7

L
M A ES

Offs p rin g, fema le B

Ofls p rin g, fem a l e R
Offs p rin g fema , l e W .

T o ta l

Here we hav e aft er on e ind is cr iminat e m at ing all aro u nd a

, ,

t o tal of n ine an ima ls o f whi ch o n e is p u re black o n e p u re red

, , ,

and o ne p u re wh it e wi th t he o t her s ix d iv id ed int o three

,

gr ou ps each o f two c o lo rs c o m b in ed
,
— that is all the po s , ,

s ib le c o m b ina t io n s .

I f n o w th is gen erat io n b e bred int o i t s elf n ew and s trange ,

c o m b inat io n s are inev i table giv ing rise to a c om p licat ed pop u la ,

t io n as fo llo ws
,

T A B LE or OF FS PR ING SHOWI N G T H E FOU RT H G EN ER AT ION I N T HE

A TT EM PT To CO M BI N E T HR EE C HA R A CT E RS

8 1 11 115 or S AM E B R EE D ING As D AM S

DA u s
2 BR 2 B W 2 IV R

B 1 B 4 1
2 B R BM ” 2 RU V 2 1
B R W B M! ”

2 BR 2 B” ? 4 B A” 2 B 11"1
4 B 1 W R 4 BR 1 lV 2 BR u
’
1

R 1 B1
R 1 R 1 2 BR W
1 2 R W 3
R 1W 1

W W W W 4 BR W B W
1 B 1 B R1 1 1V 1 1 1
2 B 2 B 4 R 2 4 B 2

R W B1 R W W 4 R W R W
1 1 1 1 1 1
'

2 2 4B R 1V 2 R 4 B R 1V 2

W1 B 1 W1 2 BR W 1 R 1 W 1 2 B lV 1
1 f
2 R W 3
’
W 1

[2 R W
B 1
+ 6B 1 W 1 + 4 B R1 + 12 BR 1 W
12 BR IV 1
+R 1
4R 1V 1
+6R 1 W +4BW +4RH
1 3 ’3
I
-
Wfl— to ta l , 81

Here ghty o n e ind iv id u als of n o les s than fifteen d iffer

a re ei -

en t c o lo r c o m b inat io n s all effect ed with in t wo gen erat io n s fro m

,

p u ri ty Ou t o f t he eighty o n e ind iv id u a ls three and t hree o n ly

.
-
, , ,

a re as p u r e as if n o ad m ixt u re had been att em pt ed su gges t i ng ,

that a c erta in s ma ll p ropor tion will a lways rema in u n m ix ed i n

lzeterog en eou s b r eed i ng , no m a tter lzow long c on ti n u ed .
 8 T R A N SM I SSI ON
5 0

A ll t he res t are m ix ed in c o lo r n o m att er what th ei r appea r ,

anc e Of th es e t he 4 B R and t he 4 B W will m o s t lik ely

.
3 3

appea r b lack as will a ls o i n a ll pro bab ility t he 1 2 B R IV 2

, , , ,

beca u se the B element s a re c learly in t he m aj o rity S im ilarly .

,

eq u a l n u mb er s will app ea r red and o th er eq u al n u m bers will

,

s eem t o b e w h it e — u n les s b o th c o lo r s app ear as i n r o an s an d

, ,

p iebald s .

T h ere a re three s et s o f s i x ea ch ( 6 B R z “ 6 B W’ “ and , ,

6 R W ) in wh ich b u t t wo c o lo r elem ent s a re present b u t in

1 2
,

wh ich t he appearanc e will pro bab ly b e fix ed by the c o lo r that is

mo s t pro n o u nc ed and wh ich th erefo re t end s t o d o m inat e the, ,

o th er ; th u s t he 6 B R will appear as b lack o r v ery d a rk red

2 1
.

T h u s it is that a ppearanc es a re o ft en d ec eiv ing and that ,

wh ich loo k s lik e a h et ero gen eo u s j u m b le is aft er a ll an o rd erly , ,

c ollect io n o f m ath em at ic a lly exact c o m bi nat io n s A s ch eme lik e .

t he abo v e s er v es t o S h o w t he exc eed ing ly c o m p lic a t ed ye t ,

o rd er ly s y s t em s that n ec es s ari ly ari s e in b is ex u a l reprod u ct io n

, ,

wha tev er the cha r a c ters i n v olv ed c o m plex ity that increas es ,

rapid ly ind eed almo s t inc o nc eiv a b ly a s gen era tio n s m u lt iply
, , .

B eca u s e o f th es e fact s reprod u ct io n wo u ld b e red u c ed to a

pro b lem in pro bab ilit ies and we S h o u ld hav e all poss ib le c o m b i
,

nat io n s present ed were it n o t fo r the fact that s elect io n is

,

alway s at wo rk to elim inat e c ert a in u nfav o red fo rm s and that ,

d ifferenc es in fert ility s er v e to giv e c ertain c o m binat io n s s t ill

f u rth er ad v antage o v er o th er s Ho wev er we a re no t to ov er .
,

loo k t he fact that ev en th o u gh c erta in v a lu es b e wi thdra wn

,

fro m s u ch a d is t ri b u t io n t he laws o f pro bab ility c o nt in u e t o ap

,

pyl t o t h e r e m a in ing v a lu es wh o s e c o m b inat io n s will t ak e p lac e

,

a s b ef o re and i n t he en d g iv e r is e t o a d is tri b u t io n n o t v ery

,

d ifferent in f o r m fro m that wh ich wo u ld hav e ari sen if n o

v al u es had b een r em o v ed .

A n u lt imat e c o nfi r mat io n o f th is s tat em ent is f o u nd in t he

fact that mos t f req u en cy d is tri b u t ion s a r e f a i r ly sy m m etr ica l ,

a nd tha t on e la rg e en o ug lz to b e f a i r ly s m oot /t , w/ta tev er t/ze

n u m b er o f i ts ter m s or t/ze s iz e f
o i ts f re
q u en c i es ,
ca n b e c los ely
re r od uc ed
p b y exp a n d i n
g a b i n o m i al I f t he d is tri b u t io n b e .

s y m m etrica l , t he t er m s o f t he b in o m ia l s h o u ld b e n u m erica ll
y
eq u al (B R, or 12 + l) b u t if it s m o d e is no t near t he m idd le
 T R A NSM I SS I O N
5 10

Dis t in ct ion s b et ween in heritan ce an d dev el opmen t . T h ere is

s t ill ano th er reas o n why t he tr u e nat u re o f an ind iv id u al cann o t
alway s b e d et ect ed by appea ranc es and th is is fo u nd in the ,

relat iv e d ev elop m en t o f inh eri t ed cha ract ers .

Fo r exa m p le s u ppo s e that in t he illu s t ra t io n gi v en o n page

,

5 0 6 B and R r epr es ent s iz e in s t ead o f c o lo r I f B r epr es ent s .

grea t s ize and R extreme s m allnes s th en by all t he p rinc ip les , ,

o f inh eri tanc e t he 2 8 B R o f t he s ch eme prev io u s ly d is c u s s ed

6 2
,

a re bo rn fo r so m eth ing abo v e t he m ean s ize wh ich wo u ld b e ,

represent ed by t he 7 0 B R ‘ ‘
.

S u ppo s e h o wev er that thro u gh in s u ffic ient feed m an y o f

, ,

th es e ind iv id u als fail to dev elop t he f u ll s ize wh ich is th eir b i rth

right S u ch ind iv id u a ls th en app ea r s m a ll lik e t he B R o r
.
’ G
, ,

perhaps ev en t he R 8
.

So it c o m es abo u t that th es e t wenty eight ind ivi d u a ls th o u gh -

,

bo rn alik e as to t end enc ies with res pect to s ize and each repre ,

s ent ed by f o rm u la B R et v e ry d i ff erent wh en exa m in ed

’ ‘
a re
y ,

aft er d e elopm en t wh ich o f nec es s ity d epend s u po n t he c ond i

v
,

t io ns o f life A n d so it is th a t o wing t o d ifferenc es in d ev elo p

.
,

ment relat iv e q u a lity as we i nf er it fro m t he appeara nc e o f t he

,

a d u lt is b u t a r o u gh and o ft en m is lead ing i nd icat io n o f t he char

a c t ers act u a lly pr es ent thro u gh inh eri tanc e R elat iv e s trength .

o f ch a ract er s a s inh eri t ed m ay b e kn o wn with c erta inty o n ly

thro u gh an int imat e kno wledge o f ped igrees Th u s a bu yer o f .

an ad u lt an imal wo u ld ha v e grea t d iffi c u lty in s electing fro m ,

appearanc es o n ly the ind iv id u al bo rn with great es t t end ency to

,

d ev elo p u n u s u a l S ize .

Th e mat hem ati cal n ature of descen t n ot d u e en ti rel y t o b isex ual

reprod u c ti on . tr u th o f th is s t a t em ent we d ed u c e fro m t he
The
fact that o ffs pring as ex u a lly pro d u c ed v ary o n t he s am e plan as
d o ind iv id u a ls th a t a re b is ex u a lly pro d u c ed I t is t o b e i nferred .

als o fro m the f u rth er fact already allu d ed t o that s u cc ess iv e

, ,

ff p
o s r i ng o f t/ze s a m e pa r en ts a r e n ot a l i ke, b u t f orm a d is tr ib u ti on
tlze s a me g en er a l ou t li n es a s t/za t tb e tota l p op u la t i on
f
o f
o .

T h is h o ws th a t the m ath emat ical element in reprod u ct io n is

S

to b e s o u ght n o t o n ly i n b is ex u a l u n io n b u t farth er back a ls o ,

in t he fact s o f c ell d iv is io n a n d t he s p litt ing of t he chro mo so m es ,

i f n o t ind eed in th eir v ery c o n s t it u t io n .

 H ER ED IT Y 5 1 1

Wha t is it that is tr ansmitt ed ? Ev olu t ion ary lit era t u re abo u nd s

in c h t erm s a s t end enc ies
su rev ers io n anc es tral b ias ,
”

,
" “
,
”

and m any o th ers wh ich im p ly an intang ib le so meth ing back of

t he im med iate parent The c o m mo n impres s io n o f tran s miss io n
.

is of s o meth ing hand ed d own o r pas sed o n fro m o ne genera ”

t io n to t he next and that t he v is ib le cha racters represent the

,

inh eri tanc e I t is ev id ent h o wev er tha t that wh ich is tran s

.
, ,

m itted is n o t t he charact er b u t ra th er the elem ent s o u t o f wh ich ,

t he character is b u ilt u p and that th es e element s are capab le o f ,

many and v aried c o m bin at io n s .

What th ese element s m ay b e lik e and what the u lt imat e u ni t s ,

o f v ar iab ility may b e wh eth er chro mos o mes o r s o me i nfin it ely

,

s m aller c o m po n ent we do n o t kno w Phy s io logica l u n it s ha v e

, .

n o t b een d is c ov ered b u t t he la ws u nd er wh ich th ey c o m b in e t o

,

fo r m chara ct ers and u nd er wh ich t he characters c o mbine to fo r m

,

i nd iv id u als with i n rac ial limit s — th es e hav e b een s u ffi c ient ly ,

s t u d ied to warrant t he a s s u mpt io n that th ey f o llo w es s ent ia lly

t he o rd i nary m ath emat ica l princ ip les o f per m u tat io n s and c o m

b in atio n s wo rk ing u nd er t he la ws o f pro bability 1
I n o th er wo rd s .
,

1 By c o m b in a tio n s is m ean t t he n u mb e r o f diffe e r nt gr o u i

p n gs t h a t c an be
d
ma e f ro m a gi v en n u mb er o f o b ec ts wit o u t r ar j
o t he a rra n gemen t o f t h e h eg dt
h h
m emb ers T u s, wit a , b, c, d , ta en t ree a t a t im e, we m ay a v e fo u r c o m
. k h h
y
b in a tio ns , n a mel , a bc, a bd, o ed , bed ; o r, ta en two a t a t im e, we a v e s ix c o m k h
y
b in a t io n s , n a mel , a b, a c , a d be, M , ed Ea c o f t es e c o m b in a tio ns m ay a v e
, . h h h
t wo o r m o re p erm u ta t io n s d
ep en in g u p o n t he or d er in w ic
, d
t he n u mb ers h h
s ta n d h
; t u s , th e c o m b i n a t io n a bc is c a pa b e o f t he p erm u t a t io ns a be, u eb, ba c , bca , l
T he n u m b er o f c o m b in a t io n s p os s ib e l wit h a gi v en n u mb er o f u n its d ep den s

u po n the n u mb er t a k en in ea c hg ro u in
p g .

The gen eral fo rm u la is

n (u I ) ( n

I 3 r

h h l
in w ic n is t he t o ta n u m b er an d r is t h e n u m b er in eac h g ro u p .

d
T he n u mb er o f p erm u t a t io n s , o r iff eren t a r ra n gem en ts , p o ss ib e a lso ep en s l d d
u p o n t he n u mb er i n eac g r o u p T he n u mhb e r of
p erm u t a t io n.s o f o b ec t s ta en j k
two a t a t im e is n ( n ta en t ree a t a t ime i t is n ( u k I ) (n h a n d so

on ; k ta en r a t a t im e it is t h erefo re n u ( 1 ) (n 2 ) ( n r

Wh en a ll t he n u mb ers en t er in to eac hp erm u ta t io n , t h en t h e fo rm u la a m o u n ts

t o t he l
m u t ip lic at io n t o g et h er o f a ll t he mb ers fro m u n it u p t o
n a t u ra l n u , y t he

n u mb er i ts elf ; t h at is , t he n u mb er o f p erm u ta t io ns o f fi v e lett ers , a , b, c , d , e, is

eq u a l tO I X 2 X 3 X 4 X 5 = 1 20 .

To giv e an illu s t ra t io n of t he p rob ab i l it y o f a n ev en t, if a en

p n yb e t o ss e d th e

o dd s ar e ev en h h d
t at ea s l
wi l b e u p, b ec a u s e t h ere is b u t o n e a lt ern a tiv e. T h is
 l R A N SM IS SIO N
'

5 I 2

it is t he elem en s t
c ia l c h a rac t e r s t h a t a re t ra ns m itt ed and
o f ra ,

o u t o f t h es e e le m e n t s a ll p os s ib le c o m b in a t io n s ari se So me .

c o m b in a t io n s a r e u n s u it ed t o t he c o nd i t io n s o f life a n d o th ers

a re r elat iv ely o r ab s o lu t ely infe rt ile m a k i ng b la nk s po t s in t he ,

s y s t em wh ich o th e rw is e wo u ld b e m a t h e m a t ic a lly c o m p let e and

s u b s t a nt ia lly reg u la r .

Ev en with th ese o m is s io n s h o wev er t he d is tri b u t io n s int o , ,

wh ich c ha ract er s f a ll lend th em selv es t o o rd in a ry math em at ical

meth od s o f s t u dy giv ing t h e s tro nges t gro u nd fo r t he c o nfid ent
,

be lief th a t the la ws o f h ered ity will n o t lo ng c o n t in u e t o b e

regard ed a s u nexpla ined my s t eries s u bj ect to all s o rt s o f ,

exc ept io n s and rev er s io n s b u t that cha ra ct ers in d es c ent will ,

b e f o u nd t o o b s erv e as well d e fi n ed a n d well kn o wn m ath em at ica l - -

p rinc ip les a s d o chem ical elem ent s in th eir s im ila r b u t v as t ly

p ro b a b l y
i it we exp res s a s 1 On t he o t h h d
er an , if a di ce be t h ro wn , t he c h an c e

of an
y p a rt ic u a r s i l d e c o m in g u
p is b ut
a f or t , h e re a re S i x
p o ss i l
b i it ies . If a

wa ge r b e l id t h
a at t he n u m b er 3 c o m es u p , t h e o dd s wi ll b fi e ve t o o n e ag a in s t it ,
fo r it s C h an c es a re o n e o u t o f s ix . If t wo di c e a re t h w th
ro h n, e c an ce o f a
3
c o m in g up on ea c h at t h e s a m e t im e is J
; x l , or
3 ,
1
; b t th h u e c an c e of two
( lg/T r en t n u m b e rs , as 3 an d 4, c o m in g u p t o ge t h er is d b l d Thi
ou e . s is b ec a u s e
t he c h anc e o f o ne di c e t u rn in g u p el f/[ C f a
3 or a 4 is not
g , b ut 3 a f t er w ic h h th e

s ec o nd di ce m ust su
pp l y t h h e
p p
r i b t A
o er m a t e,
d l w h
g 1 1
ose c a nc e s u , an , x z .
5 .

Th i m s th t m t h l
ea n s g a t h i t w ei ll h pp on r u n f s ev en y igh t a en o n c e o r e v er e een

th w ro th s,g h it ou t b fi d t l y
c an n o p d i t d t h
e t it
con w i l l enh pp reth c e a a en on e

eig h t th een th t h i t y i th
, e y
r t h
-
s x
p t i ,l t h
o r on an w o er ar c u ar ro .

Th w d e h m i t y h i d iff t l t t
or
“
c e s r By t h p i ip l
as n n e eren f p m e ers . e r nc e o er u

t tia j t l id d w th
o ns us a i ol tt n, p b
es e n n e l f x e
3 ers
x 4 a re c a
x 5 x 6 a e o 1 2 x

x 7 8 x 9 x , d i
orff t g m t l y e ren f wh i h wi ll p l l t h
a rra n e en s , o n one o c s e e

w d or h m i t y I f th f t h l tt
c e s r .
, f th i w d h
e re o re, ld b t d i t t h
e e ers o s or s ou e o sse n o e

i
a r d l ft t
an f ll i t
e o
g a d n o g a t h m l
ro o v e i l
ani b
a rra n
y h e e s e v es n ne c a n c e,

th dd w l d b
e o s ou t g ei t t h l t t t k
o i g1 at h a ns
p p e
g me t ers a n e ro e r a rra n e en

t o p ll
s t eh w d ; be t if th
or t i g h u l d b t i
e doss ni t i S t oui t h t i e con n ue , s c er a n a n

th l g
e on th y w ru n l d f ll i t t h p p
e ou d t p ll t h i p t i l w d
a n o e ro er o r er o s e s ar cu ar or .

S o o n er o r l t th fa e r, if t h h d iff f m
e re o re, th
e c an c e t i t
ers ro zero , e ev en s s u re o

h pp ; d f t h i
a en an or th i g i s m reas o n t i th
no h n s if ly o re c er a n an c an c e, on a

su ffi i t c en mb f p
nu ib il i i b
er o ff d d oss t es e a or e .

T h m i d i l t i th p
e n s os f l g n mb
e d th j d gm t
re s e n c e o ar e nu e rs , a n e u en 13 c o n

f d wh t h im p b b l h pp
use en e y ro t t h t at h e l t t
a w l
en s , d l
e t im t l y
a
p l l es e e e rs ou u a e s e

th i w s d by po r h i u re c t t i
anc e t t k s an
p l e v en
; t l y
c er a ni t h i o a e ac e no on s s so ,

b t u !b l g
i n e it will h p p
in ru n f y a t h w
en o n c e m d o r ev er ro s a e .

A l it t l f l t dy f t h p
e c a re u ib l
s u mb i tio e f
o ss f w l me co t na on s, e v en o a e e e en s ,

an d f th o t i f p
e c e r a n o c c u rr en c e oib l th g h i m p b ossb l e, ev ent w il l ou ro a e, ev en s ,

l d t h t d t t w k wit h i b l i l ge mb wit h g t l y i e ed
ea e s u en o or v ar a es n ar nu e rs rea n c r as

con fid en c e.
 T R A NS M I SS I ON
5 I4

and it aim s to pred ict the charact er o f the o ffs pri ng when th ese
hyb rid s a re bred t o geth er .

Ho w will th is hybri d breed ? W ill it c o nt in u e p u re o r ,

will it break u p int o it s c o m po n ent part s ? T h is q u es t io n

M end el s la w attempt s t o an s wer and t he ess enc e o f the la w
’

can b e s tat ed in t wo pro po s it io n s

1
. Wh en crossed fo rms o r hybri d s (B R ) a re bred t ogether
, , ,

th eir o ffs pri ng will n ot all res em ble the cro s sed parent s b u t o n e ,

fou rth o r 2 5 per c ent will b e lik e t he o riginal p u re paren t B

, , ,

ano ther fo u rth will b e lik e t he o th er o rigina l p u re parent R and ,

o n e ha lf o r 5 0 per c ent
,
will res em b le the cross ed fo rm s so
, ,

that the o ff s pring o f the cros s will t end t o as s u me t he o riginal

“
general f o r m o f B + 2 B R + R Of th es e the p u re in d i
2
.
“ ”

v id u als will breed as tr u e a s to tice clza r a c ter i n q u es t i on as if

their anc es t o rs had n ev er b een s u bj ect ed t o cross ing ; and t he

5 p
0 e r c e nt o f cr os s e
,
s w h e n br e d a m o ng th e m s el v es will aga in ,

s p li t u p i nt o p u re and cr oss ed f o rm s in t he pro po rt io n o f I ,

s o that B R bred with B R will gi v e o ff s pri ng r epres ent ed by

B + 2 B R + R ind efin it ely

° I n o th er wo rd s t he o ff s pri ng o f
1 2
.
,

hybrid s will n o t a ll b e hybrid s b u t th ey will as s u me t he s a me

,

genera l pro po rt io ns that a re a s s u med wh en pu re fo rms a re

allo wed to breed t ogeth er ind is criminat ely .

I f th is th eo ry b e t ru e it s h o ws t he im po s s ib ility o f breed ing

,

a cro s s tru e to it s o wn type o n acc o u nt o f it s innat e t end ency to

,

s p li t u p int o it s o rigi na l p u re o r u ncr oss ed f o rm s a t en d ency ,

wh ich has lo ng s inc e been enc o u nt ered by breed ers am b it io u s

to fi x a f o rt u nat e cro s s and wh ich ha s go n e far t o c o n v i nc e t he
,

po p u lar mind o f t he d ifli c u lty in effect ing a real cro s s .

T he s ec o nd f u nda menta l in Mend el s la w is t he d is ti nct io n

’
2
.

bet ween d om i na n t and r ec es s i v e charact ers I f t he chara ct ers .

in q u es t io n were ev en ly “
ba lanc ed and eq u ally d is c ernible ”

, ,

2 “
th en in a po pu lat io n lik e B + 2 B R + R t he B wo u ld b e 2

c learly d efin ed say b lack T he R wou ld a lso b e c learly d efi ned

, .
1
,

sa red and t h e 2 B R wo u ld b e so me k ind o f b lend o r m i xt u re

y ,

o f t he t wo ; i n o th er wo rd s s u ch a p o p u lat io n wo u ld b e eas il
, y
as s o rt ed int o three gro u ps in t he pro po rt io n o f 1 2 :I .

On the o th er ha nd su ppo s e o n e charact er t o b e s t ro ng a n d

,

eas ily n o t ed as a red c o lo r o r a s tro ng h ea v y s t em while t he

, , , ,
 H ER E D IT Y 5 51

o ther is xtremely delicat e as a light s had e o f blu e easily los t

e , ,

in t he red o r a lightn es s o f f o liage ea s ily o b s c u red by a

, ,

h eav y s t em N o w u nd er c irc u m s tances s u ch as th es e t he les s

1
.
, ,

n o t ic eab le o r rec essiv e cha ract ers will be v is i b le on ly in tlze

, ,
”

ind i v id ua ls tlta t a re p u r e r ec ess i v es , a ll otlzers bei ng dom ina ted by

l li o m or e p rono u nc ed c/za r a c ter . T h u s , if D
tan d s fo r d ominant s

( r e d p e ta l o r s tr o ng s t em ) and r i o r rec es s iv e light b lu e petal

( -

o r d elicat e f o liage) th en t he act u al d is tri b u t io n wou ld b e 0 +

2
,

a s b ef o r e ; b u t in th is d is t rib u t io n thr ee o u t o f f o u r
2
2 Dr + r

i nd iv id u als wo u ld b e d is t ingu is h ed by the d om i n a n t cha ract er ,

wh ile the les s a ss ert iv e t he rec ess iv e cha ract er wou ld b e o h

, ,

s c u red exc ept i n t he 2 5 p er c en t in wh ich it ex is t s u n mix ed

with the d o m inant Henc e an ind iv id ua l s h o wing a rec ess iv e

.

character may b e k n o wn at o nc e to b e p u re b u t we canno t ,

t ell by looking at ind iv id u als s h o wing the d o minant cha ra ct er ,

whi ch a re p u re and wh ich a re m ix ed W e kn o w in fact th a t .

, ,

we ha v e b o th fo r m s in t he pro po rtio n o f I t o 2 b u t in a cas e ,

o f th is k ind 7 5 per c ent wo u ld app ea r t o b e d o m in ant wh ile o nly ,

2 5 per c ent wo u ld a ppear t o b e rec es s iv es I n rea lity 2 5 per .

,

c ent a re p u re d o m inant s the o th er 5 0 per c ent b eing apparent ly

,

d o minant s b u t act u ally mix ed — a fact that im med iat ely b e

, ,

c omes ev id ent wh en th ey a re bred a mo ng th em selv es as th ey ,

at o nc e giv e rise to t he charact eris t ic d is tri b u t io n with the ,

pro per 2 5 per c ent o f p u re rec es s iv es .

I nas m u ch as few p ai rs o f C ha ract ers a re eq u ally ba lanc ed and

eq u a lly ab le t o m ak e th em s elv es ev id ent it is gen era lly tru e ,

that any generat io n fro m cros sed parentage will S h ow 7 5 per

c ent d o minant ( really 2 5 p u re and 5 0 m ix ed ) and 2 5 per c ent
rec essiv e in s t ead o f the typ ical 2 5 5 0 and 2 5 that wo u ld
, , ,

appear if the charact ers were eq u ally ev id ent and eq u ally

a ssert iv e .

Dist inc ti on b et ween c har act ers an d in div id u al s T he read er .

will b e mis led if he tak es ind iv id u als int o c o n s id erat io n h ere

i n s t ead of cha ract ers T he ent ire d is c u ss io n app lies to charac
.

t ers tak en s ing ly and wh en we S ay o f an ind iv id ua l arising fro m

,

hybrid parents that he will breed p u re we m ean o n ly as to ,

”

1 It i l
s c ea r th t aly h
on t c th t m t ll y
a ra c ers l i a bar e d i u ua exc u s v e ca n e use n

ex per i m
en t i n g ill
on or t t i t
u s ra
g h i bnj t s su ec .
 6 T R A N SM I SS I O N
5 I

t he g l e d i a m eter i n q u es t i on ;
s in J ersey s fo r e xa m p le hybrid s , of

and Sho rth o rn s wo ul d n o t breed p u re ind i v id u a l s o f eith er breed ,

th ou gh pu re J ersey and p u re Sho rt ho m c/za r ac ters wo u ld ap pea r

freely .

I f we a re t o c o n s id er many ch a ra ct e rs at o nc e we ma y eas ily ,

sa t is fy o u rs elv es a s to t he chanc es o f an ab so lu t ely p u re i n d i

v id ua l ari s ing o u t o f a hyb rid anc es try .

I f o n e f o u rth o f t he po p u lat io n aris i ng fro m a hybrid an c es try

can b e s aid to b e p u re as to a s i ng le charact er t he q u es t io n ,

arises as to what pro po rt io n o f th is n u m ber can b e c o ns id ered

as p u re with res pect t o t wo charact ers .

F ro m t he fact t hat th is s ec o nd charact er enj oy s t he sa me

c hanc es as to p u rity as did the fi rs t we c o nc lu d e that o n e fo u rth ,

o f t he n u m ber o r o n e S ix t eenth o f t he wh o le ( w ll b
, 1 i)
x i e ,

p u re as t o t wo charact ers By t he s a m e reas o n ing we k now that .

l l i
x x carr ied to any n u m be r o f t er m s will expres s t he chanc e

o f an i nd iv id u al being p u re wi th res pect t o t he c o r res po n d i ng

nu m b er of cha racters I f m any ch a ract ers a re in v o lv ed th ere.

,

fo re t he chanc es o f a p u re i n d i v id ua l aris ing o u t o f mi x ed

,

breed in g a re exceed i ngly s light and o u r chanc e o f b eing ab le t o ,

pick h im o u t by his appea ra nc e is s t ill m o re remo t e

”
.

Ex perim en tal ev iden ce ‘

I t rema in s n o w t o i nq u ire s o m ewhat
.

caref u lly int o the ev id enc e u po n wh ich th ese pro pos it io n s are
fo u nd ed .

Mend el s fi rs t expe rim ent s were c o nd u ct ed with ga rd en peas

’

and c o v ered t he fo llo wing chara ct ers : 2

1 .D ifferenc es in f or m of r ip e s eed s eith er r o u nd and ,

s m oo th o r wi th s ha llo w wri nk les or els e angu lar a n d d eep ly ,

wrink led .

2 .D ifferenc es in the co lo r of s eed a lb u men (end os perm )

eith er pa le y ello w br ight y ello w o ra nge o r green , , , .

3 .D i ff e renc e s i n t he c olor o f tire s eed c oa t — wh it e g r ay , , ,

g ra y bro wn leath er bro wn with o r with o u t v io let s po tt ing

, .

4 .D if f er enc es i n f o r m of p p
r i e od — w h e th e r inflat ed o r ,

c o n s trict ed b et ween seed s .

1 Fo r a t ra n s l a t io n of M en d el s ’
o ri gi n a l rep o rt , s ee Ba t es o n , M en d el s
’
Prin
c ip les o f H ere it d y pp 40
—1 03 fro m w ic h h are t a k en th e d a ta h erein gi v en

—
, .
, .

1 Ib i d .
pp .
45 4 6 .
 18 T R A N SM ISSI O N
5

Th e secon d gen er at i on , Wh en h o wev er

b r ed f rom h yb ri ds .
, ,

t hes e hybrid s were bred a mo ng th em s elv es , t he

rec es s iv e cha rac
t ers ca me int o ev id enc e c o ns t it u t ing in all cases appro x im at ely
,

on e f ou r th of tb e of sp r i ng lea v ing t he o th er 7 5 p er c ent t o b e

,

apparent ly d o m inant act u ally 2 5 per c ent p ur e d o minant and

, ,

5 p
0 er c e nt appar e nt ly d o m i nant b u t r ea l ly m i x ed .

T h u s i n E xperi ment 1 ( as to fo r m o f s eed ) fro m 2 5 3 hybrid s

, ,

73 2 4 s ee d s w e re o bta i n e d i n t h e s ec o nd tr i a l y e ar A m o ng th e m .

54 74 w e r e r o u nd o r r o u nd i s h and 1 8 5 0 w e r e ang u lar — a rat io ,

of to I .

I n E xp eri m ent 2 ( as to c o lo r of e nd os perm ) 2 5 8 cro s sed ,

plant s y ield ed 60 2 2 y ello w and 2 00 1 green — a rat io o f

,

to 1 .

Distrib u ti on th es e experim ent s both

of char a ct ers . In e ach of

k i n d s of s eed wer e u s u a lly f ou nd i n tb e s a m e p od s h o wing that ,

t he ov u le and n o t t he p o d is t he u n it
,
N o t o n ly wa s that t he
,
.

case b u t the pro po rt io n o f three d o m in a nt s to o n e rec es s iv e

,

h eld o n ly in t he lo ng ru n and d id n o t h o ld fo r ind iv id u al plant s

, ,

a s is s een in t he fo llo wing tab le giv ing t he c las s i ficat io n o f t he

Off s pri ng o f t he fir s t t en p lant s in ea ch exp erim ent

‘
.

E x PE R I M E N T N O . 2, CO LR
O o r1

P LA N TS
l-
1x PE R I M E N T X O I , PO R M 0r S EE D
EN D P OS E R M

R o und A n g u lar Y e llo w G reen

F ro m th is it appears that t he d o m inant a lway s exc eed s t he

rec ess iv e in n u m ber b u t that t he pro po rt io n o f 3 t o 1 is
,

1 B a t es o n , M en d l e
’
s Prin c ip l es o f d y
Here it , p 53
. .
 HER E D ITY 5 19

no t ma inta in ed in each ind iv id u a l p lant A s to wheth er th is .

c o mes fro m d iffi c ult ies in id ent ify ing and c las sify ing d o u btf u l
s pec im en s , o r fro m so me b io lo g ica l rea s o n , Mend el d oes n o t

e xpres s an o pin io n th o u gh t he po int is im po rtant

, .

I n o th er expe ri ment s t he ra t io bet ween t he d o minant s and

the rec ess iv es wa s in all cas es a ppro x i mat ely 3 t o I I n Ex peri .

ment 3 (a s to c olo r o f seed c oat s ) it was to 1 ; in E xperi ,

ment 4 (a s to fo r m o f po d s) i t was to 1 in E xperim ent 5

,

o c o lo r o f u nri pe po d s ) it was i n E xpe rim ent

( as t t o I ,

6 (as t o p o s it io n o f fl o wers ) it was to I and in Exper iment

,

7 ( as t o length o f s t em ) it wa s t o 1 b u t t h e
, n u m b e r s were ,

relat iv ely s mall ( 7 8 7 and 2 7 7 ) as c o m pared with t he n u m bers

i n v olv ed i n E xper iment s 1 and 2 .

Th e t hi rd gen er ati on secon d f rom hyb rids , A cc o rd ing t o .

Mend el ‘
tho se fo rm s wh ich in t he fi rs t gen erat io n maintain
,
“

the rec ess iv e cha ract er d o n o t f u rth er v a ry in t he sec o nd gen era

t io n as regard s th is cha ract er ; th ey rem ai n c o ns tant in th eir
o ff s pri ng .

“
I t is o th erwise with th o s e wh ich p ossess the d o m inant
charact er in t he fi rs t gen era t io n ( bred from hybrid s ) Of 2
.

these t wo th ird s y ield o ffs p ring wh ich d is play t he d o m inant and

,

rec es s iv e charact ers in t he pro po rt io n o f 3 t o I a n d th ereby ,

S h o w ex a ct ly t he s a me ra t io a s t he hybrid f o r m s wh ile o n ly one ,

th ird rem ain s with the d om inant charact er c on s tant T hat is .

to say o f t he 7 5 per c ent appa r en t d o m inant s o n e th i rd o r 2 5 per

, , ,

c ent of t he wh o le breed s p u re d o minant s s h o wing that th is

, ,

pro po rt io n is act u ally what it appears to b e na mely p u re d o m i , ,

nant s wh ile two th ird s o r 5 0 per c ent of t he wh o le y ield bo th

, , ,

d o m inant s and rec es siv es in pro po rt io n of 3 to 1 s h o wing th eir ,

es s ent ial ly hybrid o r cros sed nat u re and that th ei r d o m inanc e ,

is apparent rath er than act u al The s eparat e experiment s y ield ed .

rat io s as fo llo ws 3

I n E xperim ent 1 a m o ng 5 6 5 p lant s ra ised fro m ro u nd s eed

,

1 9 3 y ield ed rou nd s eed s o n ly and rem a i n ed th erefo re c o n s tant ,

in th is charact er wh ile 3 7 2 ga v e bo th ro u nd an d angu lar s eed s in

,

1 B a tes o n , Men d el s Pri n c ip

’
l es o f Hered ity p 5 5 , . .

1 T h at is , wit h t he 75 p e r c en t a p p a ren t l y d mi t o n an .

1 B a tes o n , M en d el s Pr in c ip
’
l es o f Here d ity p p 5 5 58
, .
-
.
5 2 0 TR A N M I S SS I O N
t he propo rt io n o f 3 to 1 The n u m ber o f the hybrid s as c om pared.

with the c o n s tant s was th erefo re to 1 I n E xperi ment 2 it .

was to 1 —
in E xp eri m ent s 3 7 t he n u m bers were s m all b u t ,

appro x im at ed the s a me rat io .

F ro m th is Mend el s tat es t he f ollo wing c onc lu s io n regard ing

t he o ff s pri ng of hybrid s
1
( i ta lic s h is)

It is no w c lea r tha t the hy br i d s f or m s eed s ha v i ng on e or other o f the two

dfi r en th ti ng c h a r a c ter s , a n d o f thes e on e ha lf dev elop a
g a in the hy b r id

f or m , whi le the oth er ha lf y i eld pla n ts whi ch r em a i n con s ta n t a n d r ecei v e

the dom i na n t or r ecess i v e ch a r a c ter s ec ti v ely in eq u a l n u m ber s

( r es
p ) .

T h is
c o nc lu s io n wh ich s eems t o b e in a cc o rd with his res u lt s
, ,

is c learly aga ins t t h e p o s s ib ility o f eff ect ing a real cro s s bet ween
chara cters that behav e as d o th os e in q u es t io n .

S u b seq u en t gen erati on s On t his po int M end el say s .

T he p o p t io i wh i h the d e e d
r or n n t o f the h yb id d e elo p
c sc d plit n an s r s v an s

u p i tnh e fi t d ec
rs dang e e t i
s o p e m
on b l y h ld g n od f ra a ll b ns r su a o s o or su se

q eu t g
n e e tni E
ra p e i
o nsm e t
. 1 d 2 h axe l er d y b ee n s ied th gh
an v a r a n c a rr ro u

i ge e t io
s x n ra 3 d 7 t
ns ,h o g h fi
an e d 4 5 d 6 r t hu g h f o v , an , ,
an ro u ur

and no d epa r tu re fro m the has b een perc ep tib le

r u le . T he o ffs p r in g o f the

h yb rid s sep a ra ted in ea c h gen era t io n , i n th e r a t io o f 2 1 1, i n to h yb rids

a n d c o ns ta n t fo rms .

T hat is t o saf p ring o f hybrid s o n e f ou rth r es em bled

of t he o f s
y ,

o n e p u re par ent and ev er aft erward br ed tr u e with r es p ect t o t he

charact er in q u es t io n ; o n e fo u rth resem b led the o th er and also

bred tru e ; and o n e ha lf s t ill rema in ed hybrid b u t i ts of sp r i ng , ,

in it s t u rn f ell apart aft er t he s a m e rat io 1

, 2 1 .

When more than t wo char act ers are in v ol v ed Mend el c o n .

d u c ted in v es t iga t io n s with p lant s d iffering in a n u m ber o f char

ac ters s im u ltan eou s ly and c o nc lu d es a s f o llo ws
3

Tha t the of sf r i ng of the hy br i d s i n wh ich s ev era l ess en tia l ly dif er en t

ch a r a c ter s a r e com bi n ed r e
p h m s er i es com bi na tions in
r es en t t e ter s
f
o a o
f
wh i ch the dev elopm en ta l f s er i es
p d
o if
f er en t ior ea ch
a t i n
g cha r ac ters a r e a ir

as s oc ia ted I t is dem on s t r a ted a t the sa m e ti m e tha t the r ela tion g ea ch p a i r

”

d
o ifi r en t ch a r a c ter s i n h br id
'

f e
y u n ion is i n d ep en d en t of the other d ife r en c es
i n th e two or i
gi n a l p a r en ta l s toc ks .

If n rep resen ts the n u m b er o f the d ifferen tia ti n g c h arac te rs i n the tru e

o rigi n al s to c ks , 3
"
g iv es the n u mb er o f ter m s o f the c o m b in a tio n series , 4
"

1 B a t es o n , Men d el s
’
Princ ip les of Hered ity , p 5 7 .

—
.

3 Ib i d .
p 57. .
3 Ib i d .
pp 64 65. .
5 2 2 T R A N M I SS I S ON
t he la w o f chanc e . T h is m ean s that ea c h
p a ren t
p r od u c es s u cc es

s i v ely
g er m
f c ells o bo th that
cha r a c ters hybr i d ( B and R ), so

fo rm s prod u c e p o llen s per mat ozo a o v a etc o f bo th o r igina l , , ,

.
,

k ind s wh ic h th erea ft er c o m b in e by the law o f chanc e ; that is

, ,

B o f o n e parent u n it es with eith er B o r R o f t he o th er pro d u c ,

i n g ei th er p u re B s o r B R S ; and a ls o in a large n u m b er o f
’ ’

in s tanc es s o m e R s u n it e wit h B s pro d u c ing hybrid s and o th er s

’ ’

, , ,

with R s p ro d u c ing p u re R s fro m hybrid pa rent s T h is is t he

’ ’
.
,

th eo ry o f ga m etic p u rity a n as s u m pt io n n ec es sa rily in v o lv ed as

1
,

a fu nda mental c o nc ept io n in Mend el s law wh ich in t he o p in io n ’

, ,

o f M end el h i m s elf app lies o n ly t o ch a ract er s that d o n o t b lend

, .

Pr ov in g or di sp rov in g M en del s l aw A go o d m any in v es t i ’

.

gat o r s a re t ry ing o ft en wit h n u m b ers inad eq u at ely s m a ll to

, ,

pro v e o r d is pro v e Mend el s la w as a gen eral princ iple o f h ered ’

ity I t m ay n o t b e ou t o f p lac e t o ca ll t he att ent io n o f t he

.

s t u d ent t o t he u s eles s n es s o f th is att em pt and to d irect his ,

att ent io n t o t he pro b lem s c o nnect ed with th is q u es t io n which

really c all fo r f u rth er and m u ch ex t end ed s t u dy -
.

Fir s t o f a ll M end el s law a s a gen era l pro po s it io n n eed s n o

’

f u rth er proo f T he experiment s o n wh ic h it was f o u nd ed hav e

.

been caref u lly repeat ed by D e V ries by C o rren s and by Ts ch er , ,

m a k , who agree as t o the c o rrectnes s o f his res u lt s T h ey hav e .

b een repeat ed u po n many o th er s pec ies with u n ifo rm res u lt s in ,

m o s t c as es and n o n ew ev id enc e is wo rt h n o t ing that is n o t

,

fo u nd ed o n n ew s pec ies and that d oes n o t inv o lv e relat iv ely

large n u m b er s .

B es id es all th is t he f u nda m enta l c o nc ept io n o f Mend el s la w

’
,

r es t s lik e G alto n s la w o f anc es t ral h ered ity u po n the inev itab le

,
’

m ath em at ic a l relat io n s in repro d u ct io n a s o u t lin ed in the prev io u s

s ect io n T h is la w aris es th eref o re as a s pec ia l cas e o u t o f
“
.
, ,

Th t m g m t
1 e er i mi g t b
a e e d y y m wi th g m ll
s co n o e u se , as s non ou s
“
er ce ,

to m th i
ea n f t ili d g m with t
e u n er gze d t er W h it i f t ili d it i
, ou re ar o s ex . en s er ze s

s p k
o f
en o y g tas a z o e.

Th i th l g g
us , n e f th ant m w h ld
ua e o
y t h t
es e h hy beri d p s, t e s ou sa a ea c r a ren

p d
ro t w
u c es k i d f go m t
n B d
s oR d m ia t
e es , d i an w h t , or o n an an r ec es s v e, o r a ev e r

c h t h b
a ra c ers mb i d N w m g m t
ave een c o nef th B k i d wil l
. o it wit h
so e a e es o e n un e

g m
a t Be esp d i ,g y gro t uc n t h
z wi ll
o es i t w i t h R g m t p o ers un e a e es , ro

d i g yg t
uc n BR (
z o es d ) ; d t ill th R g m t wil l it wit h R g m t
c ro s s e an s o er a e es un e a e es ,

p d
ro i guc ny g t R R
z
( l o es p ) O f m t h m t i
a so l i
u ret y f . t i p p a e a ca n ec es s , o r an en re c u

l ti th p p ti will b
a on es e ro or BB onsBR RR e as 1 2 1 .
 H E R E D IT Y 23

the general p rinc ip le that the bino m ial c oeffi c ient s represent po p
u la t io n s in genera l I f th ere is n o b lend th en t he c oeffic ient s
.
,

represent t he pro po rt io n s of d is t inct charact er element s I f a .

blend has oc c u rred then t he c oeffic ient s s t ill represent the pro
,

po rt io n s with in t he blend T here is and can b e n o u ncertainty

.

u po n th is f eat u re o f t he cas e T he d o m inanc e o f so m e cha ract ers

.

o v er o th er s is a m att e r o f c o mm o n o b s erv atio n and ma b e

y
acc epted as a matt er of c o m m o n s en se .

T he feat u re o f M end elis m o n wh ich f u rth er light is n eed ed is

the m at t er o f ga m et ic p u ri ty a b io lo gica l elem ent o f t he pro b
,

lem wh o se u n iv er sal tru th is n o t yet es tab lis h ed b u t wh ich seem s ,

n ec es s ary to a rat io nal explanat io n of t he law as to separat io n

int o d is t inct fo r m s ind eed Mend elis m in it s pres ent f o r m s eem s
,

to m ean s u b s tant ia lly that hybri d ind iv id u als d o n o t pro d u c e

hyb rid germ s b u t rath er that th ey prod u c e s u cc es s iv ely t he
,

pu re germ s o f bo th lin es o f parentage .

I t m ay b e c o n s id ered that M end el s law is well es tab lis h ed fo r

’

c ertain charact ers ; that is fo r characters that b lend Th is may

,
.

b e h o wev er o n ly an o th er way o f ad m itt ing it s tr u t h fo r th os e

, ,

s pec ies wh ich m a inta in ga m et ic p u r ity — th ose in wh ose germ s

t he d ifferent charact ers d o n o t m ix ( b lend ) o r in wh ich if th ey , ,

d o m ix t he pro c es s is v ery s lo w T h is in t u rn is an o th er way

,
.

o f s ay ing that M end el s law is a d em o n s trat ed fact t he o n ly

’

u nan s wered q u es t io n being To what s pec ies and cha ract ers
,

d oes it app ly ? and the an s wer t o th is d epend s u p o n t he ext ent

o f ga m et ic p u ri ty . I n t he Op i n io n o f t he writ er th is is the
g r eat u nan s wer ed q u es t io n and h e
,
r e is t he o bj ect o f i nq u i ry

t oward wh ich all i nv es t iga t io n o f M end el s la w s h o u ld b e ’

d irect ed namely to d is c ov er to what s pec ies and to what char

, ,

ac ters it s a pp licat io n s a re li m it ed .

U po n th is p o int it is wo rth wh ile t o n o t e that cro s sed f o rm s

fall int o o ne of three c las ses so far as appea ra n c es a re c o nc erned
,

( )
1 t he hybr i d m a
y r es em b le o n e o f i t s p u r e p a r ent s s o
“ ”

c losely a s t o b e ind is t ingu is hab le fro m it ; ( 2 ) it may b e a k ind

o f int erm ed iat e bet ween t he t wo d iff e r ent par nt ; ( 3) i m
( ) e s t a
y
b e q u it e d is t inct fro m eith er parent .

Of th ese th ree c las s es t he fi rs t is c learly M end elian wh ile the

—
,

sec o nd and th i rd a re d o u btf u l t he sec o nd exc eed ingly s o

,
.
5 24 TR AN M I S SS I O N
Bo th t he c nd and th ird es pec ially t he f o rmer s u gges t a
se o , ,

b lend and there is m u ch in t he experienc e of breed ers t o ind i

,

cat e that c ertain charact er s d o blend m ak ing a s u cc ess f u l u nio n , ,

ent i rely aga in s t M end el s la w wh o s e o p erat io n s ind icat e t he

’

n o n f o r m at io n o f s tab le hybr id s
-
.

T he wr it er v ent u res t o u rge th eref o re no t t he att em pt to , ,

pro v e o r d is pro v e th is great princ iple b u t t he end eav o r to learn ,

and d efin e t he lim it s of it s act io n T hat it app lies t o cross es .

generally th ere is the great es t rea so n to believ e and t o it s gen ,

eral tr u th m a ny a breed er can t es t ify wh en he has s een so m e o f ,

his fa v o rit e prod u ct io n s rev ert t o th eir o riginal f o r m s b efo re his

v ery ey es .

Ex peri men t s i n c ross in g J apan ese wal tzin g mice with al b in os .

l

D a rb is hire c o nd u ct ed ext en s iv e experim ent s with th is cros s ,

pro d u c ing th o u s and s o f ind iv id u a ls wh ich are f u lly c las s ifi ed in ,

t he o rigi na l t o wh i ch referenc e is h ere m ad e

,
I t is m o re wo rk .

o f th is k i nd that is n eed ed S pac e f o rb id s g iv ing m o re than a

.

brief ou t lin e o f so me of the m o re charact eris t ic c o nc lu s io n s o f

the exp er im ent er

1 W he th
. n f w lt i g m i c i
e ra c e o a d wi t h alb i
z n m i c e wh i h d o
e s c ro ss e no c

no t w lt a th w l t i g h b i t d i p pe
z, e a z n i th a lti g ysa g so t h t
ars
a n e res u n oun ,

w lt i g i
a z n m pl t l y e e i
s co i M d ls
e e r c e ; th
ss v e
y e n l o f enth e e
’
sen s e e c o or

h yb id i lw y d k th
r s s a a s t ar l ,i i bl
e c oa g ll y m
c o or i t e o f
s v ar a e, en er a a x ur

wild g y ra d wh it an th h t
e, f th e e c t b i g d i ti
a ra c er o tl y l ted c oa e n s nc c o rre a

wi th c h t t
a ra c ers m it t d b th b y the lb i
ra n s e d b y th e
o l ed p e t a no an co or ar n .

i th d m M d f
’
Th ere s p p e
us no i ro e i r l o e n an c e ye nc l o en e s s n s e, s o a r as o r

an d t
c oa l ce ed t he h yb i d d iff e i g
c o o r a re c o n rn , lw y i g e c l r d
s r n a a s n a , o or, a n

g en erall y i t o lo
n c oa f m b cth p r, t ro o a ren s .

2 Wh
. th en hy b id p d c d f om the
e r s ro d e c ib ed
u e e p i edr c ro ss s r ar a r

t get h e
o th e e
r, lt t yr su g h ib
an i t e e
g g
ou n t i i tex t h e g p a s f r a on n o re ro u s so ar

a s eye c o lo r a n d c o a t c o lo r a re c o n c er n ed , i n to two so fa r as rega r d s the

walt zi n g h ab i t h
The p en o m en o n o f s eg rega ti o n is c lo s el s im ila r to t a t
. y h
d es c r ib ed b y M en d el a n d i n c o lo r, w et er o f e es o r o f fu r , the p ro po r h h y
y
t io n s a re c lo s el i d en t i c a l wi t t o s e o b serv ed b y h im , h h
a q u a r ter o f the

yo u n g res em b l in g t h ei r a lb i n o g ra n d p a ren ts , ha lf rep res en t in g h

t ei r hyb r id
pa ren t s , a n d a q u a rter res emb li n g h
t ei r wa lt zin g g ra n d p aren ts in s o h
fa r t a t
h y h
t e av e p i n k ey e s a nd th e s a m e- c o lo r ed fu r , b u t d ifferin g fro m an y o f

th eir im m ed ia te a n c es t o rs i n t he r a n g e o f c o a t c o lo r ex ib i ted h .T he p r o
p o r t io n o f i n d i v i d u a ls w i c h h ex h i b i t t he w a l t zi n g h a b it i s les s t ha n o n e fi fth
o f the wh o le nu m b er of y o u ng, a nd is not a M en delia n propor ti on .

1 B i om etr ika , V o l I I I , Pa rt I , pp
. . 1-
5 1 .
5 2 6 TR A N S M I S SIO N
b et ween f p ring and g ra ndparent and s o o n ind efin itely ; b u t
Of s ,

it wo u ld need t o b e d o n e fo r ea ch charact e r s epa rat ely T h is .

in v o lv es i m m en s e la bo r and m o reo v er we d o n o t o rd ina r ily

, , ,

o s s es s s u ffi c ient ly a cc u rat e i nf o r m a t io n a b o u t a nc es t o r s b a ck

p
o f t he parent t o en a b le u s to m ak e s u ch c a lc u lat io n s We .

s eek ,
th erefo re s o me expres s io n fo r th is rela t io n an d s u ch
, ,

an expres s io n wh en gen era lized wo u ld c o n s t itu t e a law o f

anc es t ra l h ered ity .

How mu ch influ en ce b el on gs t o each separ ate an ces t or If

i nh eri tanc e is fro m t he ra c e o r in a m o re part ic u lar s en s e fro m
,

t he fa m ily gr o u p th en in pract ica l breed ing a ffai rs we need a

, , ,

mea s u re o f t he infl u enc e o f each anc es t o r in o rd er t o k n o w ho w

m u ch im p o rtanc e t o attach to t he parent and ho w m u ch to
attach t o t he s ev era l anc es t o rs back o f the parent .

On e gen erat io n back t he t o tal h eritage res t ed in t he t wo

im med iat e pa rent s and ro u gh ly s peak in g is to b e r egard ed a s
, , ,

d iv is ib le eq u ally bet ween th em Two genera t io n s ba ck it res t ed .

in f o u r grandparent s and aga i n wa iv i ng c o n s id erat io n s o f p re

, ,

po t ency o n e fo u rth o f t he h eritage cam e fro m each T he th ird

,
.

generat io n ba ck t he h eritage was d iv id ed a m o ng eight great

gra ndpare t pr s u ab ly o n t he a v erage eq u ally
n s ,
e m ,
— th a t is , , ,

o n e eighth c o m ing fro m each S t ill an o th er rem o v e and n o les s

.
,

than s ixt een great grea t grandparent s c o ntrib u t ed to t he s tream

- -

that m ad e t he fina l h eritage and it is fair to a s s u m e that each ,

ind iv id u al c o ntrib u t ed it s s hare na m ely o n e s i xt eenth ,

.

N o w t h es e s a m e s ixt een ind i v id u a ls ha v e c o nt rib u t ed a ls o t o

the pro d u ct io n o f m any o th er lin es o f d es c ent I f we ha d all .

t he d es c end a nt s o f th es e s ixt een great gr ea t grandparent s o f the - -

part ic u lar ind iv id u al we h a v e in m ind th ey wo u ld c o n s t it u t e a ,

large po p u lat io n and we ha v e n o kn o wledge a s t o wh ere in th ei r

, ,

freq u ency d is t rib u t io n with res pect to t he c haract er in qu es t io n ,

o u r s p ec ia l ind i v id u a l m ig ht b e f o u nd B u t o f all t he d es c endant s .

o f th es e S ixt een gr eat great g ra ndp a r ent s o n ly eight o f t he n ext

- -

generat io n c o ntrib u t ed to the pro d u ct io n o f the ind iv i d u al we

ha v e s pec ially in m ind ; aga in in the next g en era t io n o n ly fo u r , ,

hav e so c o nt rib u t ed and la s t o f all o u t o f t he large po p u lat io n

, , ,

d es c end ed fro m th es e s ixt een anc es t o r s t wo o n ly h a v e prod u c ed ,

o u r ind iv id u a l N o w t he law of anc es t ra l h ered ity is d es ign ed

.
 H ER ED ITY 5 7
2

to e nable u s to pred ict what on the a v erag e s h o u ld b e t he , ,

prod u ct of th is selected anc es t ry .

I n h eri tan ce compl ex T he h eri tage fro m the parent is n o t

.

theref o re a s im ple th ing b u t rath er a c o m plex s trea m o r m o re

, , ,

pro perly t wo s treams that meet fro m d iffer ent d ir ect io n s each
, ,

m ad e u p o f c u rrent s c o ntri b u t ed fro m m any trib u tar ies Ho w .

m u ch n o w has each t rib u tary (anc es t o r) c o nt rib u t ed t o the

gen era l and c o mpo sit e m ixt u re that we call the her itage ?
G a lt o n m ad e t he fi rs t att empt t o an s wer th is q u es t io n and
l
,

annou nc ed as t he law of anc es tra l h ered ity that the t wo immedi

a te parent s c o ntri b u t ed b et ween th em o n e h a lf f h
( 5)
0 o t e .

ef ec t i v e h er itage t he g randparent s o n e f o u rth

, the great

g randpar e nt s o n e eighth and s o o n so that t he effe ct iv e ,

c o ntrib u t ion s o f t he s u cc es s iv e gen era t io n s wou ld b e represent ed

by the fract io n s i l 16 1 et c and t
, h e tota l h eritage wo u ld
, , .
,

b e repres ent ed by t he s u m o f th es e fract io n s wh ich ext end ed , ,

to infin i ty wo u ld eq u a l 1 th u s acc o u nt ing fo r t he t o ta l h eri tage

, ,
.

T h is general la w app lies t o g enera t i on s n o t t o i n d i v i d u a l ,

a n ces tors and th es e fract io n s s h o u ld b e s t ill aga i n d iv id ed by t he

,

n u m ber o f anc es t o rs in each generat io n in o rd er t o d et er m ine

the fra ct io na l s hare c o ntrib u ted by each i nd iv id u al anc es t o r .

T he f o llo wing tab le exh ib it s t he fract io na l c o ntr ibu t io n o f each

genera t io n and o f each ind iv id u al anc es t o r acc o rd ing to t he law
of anc es tra l h er ed ity a s s tat ed by Gal t o n .

EFFE CT I V E HERIT A G E CO N TRI U TE B D BY EAC H G ENE R AT I ON AN D BY

EAC H SE PA RA TE A N C ES T O R A C C OR D I N G To T HE LA w OF

A NC EST RA I . HER E D ITY AS ST AT ED BY G A LT O N

GEN E R A TI ON E P PB C T I V B C T R I TI N ER ANCBS E B B E CT I V E C O N T R I TI O
A N E TO R
ON BU ON U MB OF BU N

BA WA RD
CK OF E A CH G E N E R A TIO N ToRs INVO L VE D on E ac h C S

or

or 6 . 2
5 7:
J

1
Z
‘
or
3 ; ,

a i r o r 0- 9
3
1
73
(
or
I 02 1

1 l
G a t o n , N a t u ra l In h eri t an c e, pp .
—
1 34 1 3 7 ; P r oc eed i n g s o
f t h e R oy a l S ociety ,
L X I , 40 2 .
 28 T RA N S M I SS I O N
5

Th is ries ( 5 i 5
se ,
et c ) carr ied t o i nfin ity wo u ld acc o u nt
» ,
.

fo r t he t o ta l h eritage and if th ese fr a ct io n s a re c o rrect ly tak en

,

t he infl u enc e o f each s eparat e anc es t o r wo u ld on the a v erag e , ,

b e repres ent ed by t he fract io n s in t he la s t c o lu m n .

T h is la w
“
at fi rs t so mewhat arb itrarily d eriv ed and an
, ,

n o u n c ed with c o n s id erab le h es itat io n ha s r ec eiv ed gen era l s u p ,

po rt fro m lat er in v es t igat io n s and all res earch es m ath em at ical , ,

a s well a s o th erwis e t end t o es tab lis h it s s u b s tant ia l acc u racy

,
.

T he fi rs t ann o u nc em ent was ba s ed u p o n s t u d ies in s tat u re .

S o m ewh a t lat er o ppo rt u n ity was a ffo rd ed to m ak e exhau s t iv e

s t u d ies fro m a large n u m ber o f B as s et h o u nd s o f d is t inct c o lo r s

and of sev eral gen erat io n s o f kno wn anc es try Th is s t u dy is .

repo rt ed in f u ll by G alto n and the res u lt s c o nf o r m s u b s tant ially

l
,

“
la w wh ich as Gal t o n o b s er v es i s s tr ict ly c o n so n ant
”
t o t he , , ,

with t he o b serv ed b ina ry s u bd iv is io n s o f t he ger m c ells a n d t he ,

c o nc o mitant ext ru s io n an d lo ss o f o n e half o f the sev eral c o ntri

b u t io n s fro m each o f t he t wo pa rent s t o t he ger m c ell of t he
o ff s pr ing
2
.

T h ese B a s s et h o u nd s were es p ec ia lly fa v o rable fo r a s t u dy o f

th is k ind Th ey a re of t wo c o lo rs o n ly lem o n and whit e o r
.
,
“
,

th ey m ay b e m ark ed in add it ion with a th ird c olo r ( black ) in ,

wh ich ca s e th ey a re kn o wn as tr ic o lo r .

I t is s a id that ind iv id u a ls a re d is t inct ly o f o n e o r t he o th er

C la s s and that tran s it io na l s p ec im en s a re v ery rare
,
The ped i .

grees and c o lo r d es cript io n s ha d b een caref u lly k ept by S ir

Ev erett M illa is who had o rig inat ed t he par tic u lar s t o ck
, .

So me 8 1 6 3
h o u nd s o f kn o wn c o lo r were d esc en d ed fro m
parent s o f kno wn c o lo r ; in 5 6 7 c as es the c o lo rs o f t he grand
3

parent s also were kn o wn and in 1 8 8 cas es the c o lo r s were kn own

,

fo r th ree g en erat io n s back A s s ign ing fract io nal v alu es t o

.

parent s gra ndpa rents etc acc o rd ing t o C a lt o n s law of a n

, , .
,
’

c es t ra l h ered ity and ca lc u la t ing wh a t t he d es c endant s s ho u ld b e

,

u nd er t h e la w i t app ea red that a cc o rd i ng t o th eo ry th ere s h o u ld

, , ,

ha v e been 1 80 tric o lo r h o u nd s d es c end ed fro m th o se wh os e

anc es to rs were kn own fo r three generat io ns I n fact 1 8 1 s u ch .
,

1 P r oc eed i ng s of th e R oy a l S oc i ety , L X I , 4 0 1 4 1 2 -
.
9 Ib i d .
p 403
. .

3 N o t 8 1 7 an d 5 7 7 , as p rin t e d
T ere s eem s to b e
. h a n err o r o f I in t h e fi rs t ro w.

P r oc eed i n g s of th e R oy a l S oc i ety , L X , 409 .

 TR A NSM I SS I O N

A s Pearso n s ays
,
what now a re h8 and
that is wha t
E; ,

d ev iat io n fro m t he mea n o f o ffs pring in gen eral (11 3 ) is to

b e expect ed in t he o ff s pri ng o f th es e part ic u lar parent s and ,

what is th eir v ariab ility o r s tandard d ev iat io n with res p ect t o

th eir o wn m ean Ca s t in s t ill m o re general t erm s t he ,

q u es t io n s a re th ese Ho w will t he o ffs pring f ro m s elected parent s

d iffer from o ffs pring in general and ho w will they d iffer a mo ng
,

th emselv es
N o w th ese a re f u nda m ental q u es t io n s in breed ing and th ei r ,

an s wer in v olv es the f ollo wing add it io nal c o nc ept io n s all wit h ,

referenc e to the charact er in q u es t io n :

1. T he s tandard d ev iat io n fo r fath ers in gen era l
2 . The s tandard d ev iat io n fo r m o th ers in gen era l
3 . T h e s tandard d ev i at io n fo r o ff s pr ing i n g en eral

4 . T he c o e ffi c i ent o f h e r ed i ty b e t wee n fath e rs and o ff s p r i n g

reck o ned as s o n s
5. T h e c o e ffi c ie nt o f h e r e d i ty b e t w een m o th er s and o ff s p ri ng ,

also recko ned as s o n s ( r 2) .

6 T he c oeffi c ient o f c o rrelat io n ( cro s s h ered i ty ) b et ween

.

fath ers and mo th ers d u e to asso rtat iv e mat ing ( rs)

,
.

Galt o n c o n sid ered that inh eritanc e fro m t wo parent s is s u b

s ta n t ia lly eq u i v a lent t o i nh eri tanc e fro m a m id parent wh ich
”
-
,

s h o u ld b e t he m ean o f t he t wo aft er tran s m u t ing t he f ema le

v a lu es ( mea s u rem ent s ,

fo r exa m p le) int o m a le eq u i v a len ts by
m u lt ip ly ing th ose v a lu es by t he rat io o f t he m a le to the f em a le
m ea n fo r t he charact er in q u es t io n
,
.

Pear s o n o n t he o th er hand d eals fir s t with d ev iat io n s a n d

, , ,

by the d ev iat io n s o f t he parent s fro m parent s in gen eral he

att em pt s to predict t he d ev iat io n o f th ei r part ic u lar o ffs p ring
fro m t he mean o f o ffs pring in general wh ich is t he s a me as s ay ,

in g “
fro m the m ean o f t he rac e .

Wh ile Galt o n th u s art ific ially b u ilt u p a m id parent t o t ak e -

the p lac e o f t he t wo parent s Pears o n d ev elo p ed fi rs t t he th eo ry

of
“
biparental inheritanc e tak ing int o acc o u nt t he m ean s
,

and v ariabilit ies o f t he parent s t he c oeffic ient of as so rtat iv e

,

mat ing a n d t he c o effic ient o f c o rrelat io n b et ween o ff s p ring and

,

parent s th u s lead ing t o t he f o llo wing fo r m u la fo r b iparental

,

inh eritanc e
 H ER ED I TY 53 1

Where hl and h, a re t he d ev iat io n s o f parent s fro m t he m ean

o f parent s fo r t he C haract er i n q u es t io n is t he d ev iat io n o f ,

t he o ffs pr ing o f th ese parent s fro m o ff s pr ing in gen era l a is

'

, ,

t he s tandard d ev iat io n o f fath er s fro m t he m ean o f th eir gen er

at io n a , is t he s tandard d ev iat io n of mo th ers fro m the mean
,

o f th ei r g enerat io n a is t h e s tandard d ev iat io n o f o ff s pri ng in

s ,

general as to the chara cter in qu es t io n r l is the c oeffi c ient of ,

hered ity bet ween o ff s p ring and parent s ( parent s being tak en
as eq u i po t ent that is mak ing r, r is t he c o effi c ient o f
, 8 ,

as so rta tiv e mat ing .

N o w fo rm u la ( I ) m ay b e wri tt en as fo llows

0I
I
“
i
“ "

1's

or, again ,
it m ay b e writt en

Inth is fo rm t he fo r mu la is m ad e u p o f fo u r fac t o rs Of these .

t he fir s t r epres ent s t he m id paren ta l d ev iat io n ; t he s ec o nd t he

-
,

v ar iab ility o f m id parent s ; t he th ird t he c o rr elat io n c o effi c ient

-
,

bet ween m id parent and o ffs pri ng ; and the fo u rth t he s tandard
-
,

d ev iat io n fo r t his part ic u lar chara ct er I f n o w each o f th ese .

, ,

fact o rs (exc ept 3 ) b e represent ed by a single lett er we ha v e

0 ,

t he f o llo wing :

h, the m id pa ren tal d ev ia tio n ,

-

the v a ria b ility o f m id p a ren ts, -

c o rrela tio n c o effic ien t between m id pa ren t -

V r8
an d o ffsprin g .

Fo r d eriv a tio n o f t hi s fo rm u la, s ee A ppen ix d .

ll
7 T he o ffs prin g wi , o f c o u rs e, fo rm a n a rra y , an d th e h , is t h e d ev ia tio n o f it s
m ea n fro m t he m ean o f o ffs p rin g in gen era l .

h
1 T is las t exp ress io n d
is in v ert e b ec a u s e we d es ire to u s e S in t he d en o m in a to r.
5 3 2 TR A N SM I SS I O N
W e ar e able to pu t fo r m u la I ) in t he fo r m of a regress ion
n ow

eq u at io n giv ing t he v alu e 11 3 a s fo llo ws

,

A, R
if
wh ich is t he d ev iat io n of th is s pec ial pop u lat io n from the m ea n
of t he race .
1

T h is f o r m o f express io n o f f o r m u la ( I ) has t he ad v antage of

s im p lic ity I n s t ead o f t he d ev iat io n s (h1 and h, ) o f t wo parent s
.

with v ariab ilit ies a l a n d o z we n o w ha v e t he d ev iat io n ( H ) o f a ,

s ingle art i fic ial m id parent m ad e by fi r s t tran s m u t ing f em ale -

d ev iat io n s int o m a le v a lu es by m u lt ip ly ing by t he rat io o f ma le

t o f em ale v ariab ilit ies fo r t h e C haract er in q u es t io n and th en
tak ing t he m ean o f t he m a le and t he tran s m u t ed fema le v alues 2
.

T h is is Pears o n s m id parenta l d ev iat io n ( H )

’
-
.

S is t he p o rt io n o f th is f o r m u la wh ich in v o lv es t he v a r ia b i l i ty
of pa ren ts fo r it d ep end s u po n a , and u p o n t he c o efli c ien t o f
,

as s o rtat iv e mat ing ( ra) and when a ssoc iat ed with H as it is in ,

t he f o r m u la it may b e loo k ed u po n a s express ing the v ariab ili ty

o f t he m id parent -
.

L ik ewise R is t he p o rt io n o f t he f o r m u la wh ich in v o lv es the

c o rr ela t i o n b et ween parent and o ff s pr ing and fro m t he f o r m o f ,

eq u at io n ( 4 ) it m ay b e lo o k ed u p o n as t he c o effi c ient o f c o rrela

t io n b et ween O ff s p ring a n d m id pa rent -

.

I f we n eglect t he c o effi c ient o f a s s o rtat iv e mat ing mak ing ,

r 0 t h e f o llo wing c o nc lu s io n s m ay b e d ra wn fr o m fo rm u la s
3 ,

(3) and and t h e v a lu es o f R and S

1 T he v ar ia b ility o f t he m id parent ( S ) is eq u al t o that o f

.
-

fath ers d iv id ed by V 2 .
‘

1 E xp erimen ta ld et erm in a t io n s s h o w t h at fo r m o s t c h a ra c t ers t h us fa r in v es ti

g a t e d t he re
g r es s io n c o effi c ien t of o ffs p ri n g as c o m p a re d wit h m id p a ren ts is-

ab o u t so t h at we m a y wri t e, in gen era l , ha II ; o r, in h

o t er wo r s, if a d
m id p a ren t
-
d ev ia t es a c ert a in a mo u n t th e o ffs
p ri n g m ay b e ex p ec t e d in gen era l
to d ev ia te of t h a t a m o un t f ro m t he m ea n of t he ra c e .

h 1
( 2 h
) H l
9 T is is the hl of the fo rm u a .

2 09

0l v i + r8
T h at is , in S if as s o rt a t i v e ma t in g b e dis rega r d ed , r . b ec o m es
2
01 V1 ” l
ze ro an d the fo rm u l a b ec o m es b ut V I 1 , an d we h av e
2
 I RA N S M I SS I O N
' ‘

5 34

in l g , ,
by 1
w
1 6
h ich h e acc o u nt s fo r t he t o t al h e ri tag e T h is .

f ra ct io na l infl u enc e o f the d ifferent generat io n s th erefo re m ay , ,

b e acc ept ed a s the bes t g en era l s tat ement p o s s ib le o f t he la w

o f anc es tra l h ered ity T he infl u enc e o f ind iv id u a l anc es t o r s
.
,

waiv i ng a ll c o n s id era t io ns o f s pec ial prepo t ency wo u ld b e fo u nd ,

by d iv id ing th ese fract io ns by the n u m b er o f anc es t o rs o f that

generatio n ( gby 8 3 ; fo r great grandparent s ) ( S ee table
1 = -
.
,

page 5 2 7 fo r an ext end ed s tat em ent o f t he f ract io nal in fl uenc e

,

o f gen erat io ns and s epara t e anc es t o rs ) .

T he v ar iab ility o f t he o ff s pri ng o f an anc es t ry s elect ed fo r an

ind efin it ely large n u m be r o f g enerat io n s b ack is gi v en by a
fo rm u la which is m erely an exten s io n o f t he fo r m u la fo r t he
v a riab ility o f the o ff s p ring o f t wo s elect ed parent s If we .

as s u m e C alto n s c oeffic ient s in t he la w o f anc es tral h ered ity

’

t he f o r m u la fo r t he gen era l ca s e m ay b e wri tt en a s f o llo ws :

2 f l f a 7 "

z \ /é ( 2 (2V 5 )
”
( 2 x /Ey
i n wh ich l 2 3
r ,
r ,
c o effic ient s o f c o rrelat io n
7 . r” ,
a re t he

b et ween o ffs pring and t he fi rs t s ec o nd th ird n t h m id , , ,

parent s U s e will b e m ad e o f th is f o rm u la in treat ing o f the

.

red u ct io n o f v ariab ility by s electio n .

SEC T I O N X V LI M I T TO T H E R ED U CT ION OF

Y V A R I A B I LIT

We o ften s peak of fix ing t he type by s elect io n m ea n i ng ,

by that t he r ed u ct io n o f v a riability A ll rec ent s t u d ies ho wev er .

, ,

t S h o w that we d o n o t great ly red u c e v ariab ility by s elect io n

g o o ,

h o wev er m u ch we a lt er t he type .

I n t he rec o rd s o f c o r n breed ing it will b e remem bered that ,

wh ile t he pro t ein and o il c o nt ent s rap id ly res po nd ed to s elec tio n ,

r i a b i l i ty changed b u t litt le nd d
2
et t he c oe fi c i en ts o
f i i t
y v a ee ,

is t he experienc e ev ery wh ere that v ariab ility is b u t s light ly

red u c ed by s elect io n .

T h is experienc e acc o rd s with m ath em at ica l th eo ry I t will .

b e s h o wn in t he A pp end ix that in genera l t he v ariab ility o f an , ,

a rray is o bta i n ed fr o m t he s tandard d ev iat io n o f o ff s pri n i

g n

t bl
1
S P eeG mm f S i
ea rs o n ,
p 4 8
ra S ar o
p 4 46 c en c e, . 2 .
3 ee a e, . .
 H E R ED ITY 5 35

general by mu lt ip ly ing th is s tandard d ev iat io n by V I r l ; tha t z

is in s y m bo lic langu age

2 g v es t he v a riab ility
2
,
a
s
r,,
i

( s tandard d ev iat io n ) o f an arr ay o f o ff s pri ng wh o s e c o rrelat io n

with t he select ed parent is r s and i n which the v a riab ility of ,

o ffs pri ng in gen eral is a s .

The n u m eri ca l v a lu e o f th is v ariab ility in a giv en in s tanc e

d epend s u po n t he v alu e o f r N ow experiment al ev id enc e , .

s h o ws that t he c o rrelat io n b et ween parent and o ff s pr ing r an ges

a ll t he wa
y fr om w i th li tt le o r n o a s s o rtat iv e m at ing u p to ,

abo u t with t he h igh es t s elect io n o f bo th parent s that has yet

been ach iev ed (see ta b le of c oeffi c ient s of h ered ity page ,

N o w in o u r fo rm u la
2 s u s s u b s t it u t e th es e
"
1
a V I r le t
,
v alu es :

When r 1
2 a V I
s
a
s ; that is i n th is ,

cas e wh en o ne parent is select ed we get an o ffs pri ng o nly abo u t

,

5 p er c e nt le s s v ar i ab le than t h e o ff s pr i ng i n g e n era l .

W e hav e al ready seen (page 5 3 3 ) t hat when t wo parent s are se

lec ted a s s u m ing th em to b e eq u ipo t ent t he f o r m u la fo r t he v ari
, ,

2
2 r
a b ility of the o ffs pri ng of select ed parents is 2 a
s
1
1

1 r
s
Let u s now m ak e the
as su m pt io n as befo re ; na mely tak e
sa m e ,

r
l
fi rs t a s fo r pan ga m ic m at i ng and aga i n a s for the c as e ,

of perfect as so rtat iv e mat ing .

3
2 r
—
and th en 2
If 0 0, 1 bec o mes
1 r
1
.
3 r
s
a
s 1

I 0
s elect io n of bo th parent s o u t of a rac e dev elo ped by panga mic
m at ing will res u lt in the red u ct io n o f v aria b ility by o n ly abo u t
1 0 per c ent .

2 I f r
.
1
0 5 and r
s
I — that i
.s w i th p e rf ect a s
, s o rtat iv e ,

mat ing and with the h igh es t c o rrelat io n fo u nd i n h igh ly bred

rac es ,

bec omes
53 6 T R A NS M I SS I ON
a ll of wh ich m eans that t he c los es t selectio n o f bo th parent s
( pe rfect a s so rtat iv e m at ing ) cann o t res u lt in t he red u ct io n o f

v ar iab ility by m o r e than abo u t 1 3 per c en t .

M o reo v er if t he ent i re back anc es try b e select ed t he v ari

, ,

ab ility will n o t b e m u ch red uc ed belo w th is po int I n c o nnec .

t io n with the law of anc es tra l h ered ity ( page 5 34 ) we gav e a

fo r m u la fo r t he v a riability o f t he o ffs pring o f an anc es tra l li ne
s elect ed back fo r an ind efin it ely large n u m b er of gen erat io n s .

T h is fo r m u la is

I
Vi V 5) E) 6)
“ ; “
2
( 2
( 2 N
( 2 N

in wh ich E is t he ar iability o f t he o ffs pring o f th is s elect ed

v

anc es t ry a is t he v ariab ility o f o ffs pring in gen eral fo r t he

,

po p u lat io n fro m wh ich s elect io n is m ad e and r , r s r s r a re , , , ,

m
, ”

t he c o rrelat io n c o effi c ient s o f o ffs pri ng a n d fi rs t s ec o n d th i rd , , ,

n t h m id parent s -
.

F o r pan ga m ic m at ing r l rs r s r m ay b e tak en as

, , , , "

’
v; ( V 5 )
:
( V 5
)
:
( v z
)
S u b s t it u t i ng th ese v alu es in we g et

o
u r w h en
2
(7

z a a ,

wh ich m ean s that in t he c as e o f pa n ga m ic ma t ing t he v ariability

is red u c ed o n ly abo u t 1 1 pe r c ent by s elect ing t he ent i re
anc es try .

B a s in g his rem a rk s o n th es e fact s Pears o n s ay s t hat t he ,

1 0 t o 1 3 p er c ent r ed u ct io n o bta in ed by t h e s elect io n o f t wo

l 1 I
‘f T he s e ri es t o In fi n i t y Is a g eo m et ri c a l r
p g
o res si o n w h ose
3 I?
I
”

sum is fo u n d in t h e usua l m an n er b y div i in g t h e fi rs t d t erm by 1 m in u s t h e ra t io .

 8 T R A N SM I SS I O N
53

to b e med ioc re on t he one hand or s o me thi ng m o re tha n

med ioc re on t he o th er .

Assu ming medi ocri ty b ey on d some defi ni te poin t in th e b ack

G alto n s fo rm o f t he law o f anc es tral h ered ity m ay
’

an cestr y .

b e writ ten

to i nfin ity , in wh ich h has t he mea ni ng

d efin ed o n page 5 3 3 .

I f t he v ar iab ilit ies o f s u cc es s iv e gen erat io n s b e ta k en a s

eq u a l t he f u ller s tat em ent o f th is la w as gi v en o n page 5 3 3
,

red u c es at o nc e to th is s im p le fo rm .

A s t he s im p ler f o r m o f s tat em ent giv es a goo d appro x im at e

v alu e we s ha ll fo r t he s ak e o f s im p lic ity and eleganc e o f res u lt s
, , ,

b e c o nt ent h er e to i n v es t igat e what gr o ws o u t o f th is la w in t he

way o f es tab lis h ing a chara cter fo r wh ich s elect io n and breed i ng
a re b eing carried o n .

1 . I f we a ss u m e med ioc ri ty back of t he im m ed iat e parent s ,

we m us t m ak e

T h en I! 0 -5 HI ;

that is ,h al f the d es ired cha ract er is present in the o ffs p ri ng

o ne .

2 . I f we as s u m e m ed io cri ty back o f t he grandparent s we ,

m u s t mak e

T h en h H, 11 2 .

I f we hav e a fix ed s tandard o f s ele ct io n ,

HI Hs , an d

I! HI .

3 . a
I f we
s su m e m ed io cr ity back of the great grandparent s
-
,

we m u s t m ak e

T h en

and with a fix ed s tand a rd o f s ele ct io n

fro m wh ich it H1 .
 H E R E D I TY 5 39

Ifth is s a me line o f argu m ent b e carried on , so that the fou rth

genera t io n of back anc es try is select ed ,

1: 0 -93 75 H I .

L ik ewis e, if t he fifth generat io n be sele ct ed ,

h 111 .

A n d if a s ixth generat io n be s ele ct ed ,

h H1 .

T he si gn ificant po int in all th is is that six generat io n s of

s elect io n
,
ev en on a m ed io cre s t oc k es tab lis h t he s elect ed ,

charact er to with in abo u t p er c ent T h e f u ll s ig n i fi canc e o f .

th is point will appear lat er .

F in ally if s elect io n o f t he charact er o f d ev iat io n H , b e m ad e

,

fo r n gen erat io n s and if we may a s s u m e m ed io crity in the

,

anc es try bey o nd t he n th generat io n the a m ou nt of the charac ,

ter es tab lis hed is giv en by

1
+ +
I
H. x
4 8 2

Mak i ng n o ass u m pti on s as to mediocri ty in b ack an cestr y .

I t has been s h o wn bo th by experimental and by theo ret ica l

m eth od s that if m id parent s with charact er H , a re s elected t he
-
,

o ff s pring will o n t he a v erage

,
exh ib it ab o u t H 1 ? o f t he
,

C ha ract er in q u es t io n T he inq u i ring read er will as k h ere why

.

th is d iffers fro m t he o b ta in ed fro m G alt on s la w It

’
.

s h o u ld b e rem em b ered that 0 5 H 1 is what we o bta in ed by .

ass u m ing med ioc rity bac k o f the fi rs t m id parent s I n genera l -

.
,

if we select parent s o f charact er H 1 th eir s pec ia l a n c es try will ,

exh ib it th is C h a ract er t o a gr eat er d egr ee than anc es t ry in

general fro m wh ich the select io n is m ad e I t is th erefo re o n ly .

c ommo n s en se to expect a h igh er v alu e than H , u nd er t he

present as su m pt io n .

Gra nt ing then if we can m ak e n o as sert io n abo u t back

, ,

anc es try that an o ffs pri ng will exh ib it

,
o f t he d ev ia t io n o f

is the su m o f the geo m et ric a l p g

ro r ess io n
(
3
2
1
4 g
t S ee p 5 33
.
; a ls o P r oc eed i ng s 4 th e R oy a l S oc iety , L X I I , 396 .
54 0 TR A N M I S S SION
s elect ed mid parents Pears o n has es ta b lis h ed by t he th eo ry
-
, ,
1
of

m u lt ip le c o rrela tio n the fo llo wing res u lt s

,

I .If s elect io n b e m ad e o f fi rs t and s ec o nd gen eratio n s of

anc es t ry with n o kn o wledge as t o b ack anc es try

, ,

h H, H2 .

I f we ha v e a fix ed s tandard o f s ele ct io n , H,

t hen E 1.

2 . If ct io n b e m ad e o f three genera t io n s
s ele o f a nc es try
u nd er s im ilar c o nd it io n s a s t o b a ck anc es try ,

h 11, H2 H, ;

with a fi x ed s tandard
h H, .

3 . If s ele ct io n b e mad e of fo u r gen era t io n s of anc es try ,

h H, H, H, IL ;

with a fi x ed s t a nd a rd
h 1 4 H, .

4 . S im ila rly , fo r a s ele ct io n of fiv e genera tio n s ,

h H, H,, H, H, H, ;

and with a fi x ed s tandard

h 1 7 H , .

5 . Fin a lly , fo r a s ele ct io n o f S ix gen era t io n s ,

h H, H, 2 5 0 H, H,
H, H0 ;
and with ou r fi x ed s tandard H, ,

[1 H1 .

It h o u ld b e n o t ed that the c o effic ient s wh ich we o b tain are

s

appro ach ing m o re and m o re G alto n s c o efli c ien t s in t he law o f ’

anc es t ra l h ered ity wh ich m ea n s that if select io n b e carried

,

o n fo r a v ery la rg e n u m b er o f gen erat io n s it d oes n o t m att er

wh eth er the back anc es t ry o f o u r s elect io n b e m ed iocre o r

abo v e med ioc rity .

1 th e R oya l S oc iety , L X I I , p p
P r oc eed i ngs f
o .
397 3 98
-
.
 54 2 TR A N M I SSI S ON
to m at e wi th o u t s ele ct io n I n th is trea t m ent we s hall as su me
.

m ed io crity in th is back anc es try o f t he n s elected gen erat io n s

of anc es try .

A s was s een o n p a ge 5 39 , t he C h a ract er is es tab lis h ed in t he

a m ou nt giv en by H, ,
and we s hall n ow S h o w that if

th is of s f pring be allo wed to breed with o u t fu rth er s ele ct io n it

I
W i ll breed tr ue to I of t he s ele cted cha ract er .

I n t he fi rs t gen erat io n of of s f p ring a ft er no s ele ct io n we

s h o u ld h av e
1 I

of t he chara ct er q u es t io n
H , in .

T h is s eries may b e writt en as

I
+ l = I
;
‘

a m o u nt o f t he charact er present is th erefo re u nchanged

T he .

I n t he s ec o nd gen erat io n o f o ff s pring aft er n o s elect io n we

s h o u ld ha v e

I 1

— x

= I

so that a mo u nt o f t he ch a ract er pres ent is aga in u nchanged

t he .

T he m eth o d h ere u s ed can b e ext end ed to any n u m ber o f

generat io n s W e may s h o w th a t if it b e t r ue fo r t he r t h gen era
.

t io n of o ff s pring bred with o u t select io n it will b e tru e fo r t he ,

h generat io n If l of t he charact er has appeared in

( r + I ) t . I
2
;

r generat io n s th en i n the next gen erat io n

, t he amou n t of t he
charact er s ho u ld b e gi v en by
 H E R ED ITY 543

2 2 2 2 2

1 I 1 1
1 sm ee
2 2 2 2

1
wh ich s h ows that I
”
of t he d es ired cha ract er will b e present
2

in any generat io n Henc e the s t ock will alway s breed tru e t o

.

t he per c ent o f t he charact er es tab lis h ed .

I t m ay b e rem em bered i n t he ab ov e th a t m ed io cri ty ha s been

as s u med in the anc es try ba ck of t he n s elect ed generat io n s of
o ff s pri ng ; b u t if th is wer e n o t a s s u m ed we ha v e s een that aft er ,

a few gen erat io n s we o bta in appro x im at ely G alto n s c oefli c ien ts ’

.

Henc e with o u t a s s u ming m ed io cri ty back o f n genera t io n s we

, ,

m ay s afely say th a t the o ff s pri ng will breed tr u e t o t he a mou nt

o f t he cha ract er es tab lis h ed by s elect io n .

T he fo llo wing tab le present s t he a mo u nt o f a cha ract er es tab

lis hed by s elect io n o f 1 2 3 4 5 and 6 gen erat io n s o f anc es t ry
, , , , ,
.

To illu s tra t e ho w th ese breed tru e we m ay t a k e t he si m p les t

case wh ere o f the charact er is es tab lis h ed by s elect ing o n e

generat io n S u ppo se th en th a t a gen erat io n is pro d u c ed with

.
, ,

o u t s elect io n T he a m o u nt o f t he charact er pres ent will b e

.

giv en by

which illu s t rat es t hat s a t bility is es tab lis h ed .

5 44 T R A NSM I SI S ON

E FFEC T OF C ON TI N U E D SEL ECT ION U PON V A R IA B I LI TY A ND TY PE ‘

F R A C T IO N A I ON I BU C T R TION A o rr N C ES TR
O S R A TI O or T
R A IO OF

R G EN E R A TI N
. ,

V A IO U S O S Fm “ , TO Fm “ , TO

SE L EC TE D I N I TI A L

T Y PB V A R I A BI L I TT

.
5 1 22 . 2 92 7

.
50 1 5 . 2 55 3 . 1 4 59

2 5 07 1 2 76
.
5 00 2 . . . 07 2 9

.
5 000 . 2 5 0 1 . 12
53 . 06 3 8 03 6 5

2 5 00 0 062 7 0182
.
5 000 . . 12
5 . . 03 1 9 .

S ECT I ON XV I I I — D U R ATI ON OF V A R I ETI ES, BR EEDS,

A N D FA M I LY ST R A I N S

How l on g can a desir ed b reed or famil y b e ret ai n ed ? T h ere is

a po p u lar b elief th a t v a rieties wear o u t and that breed s m u s t ,

o f n ec es s ity b e c o n s tant ly reenf o rc ed by n ew m at erial o r by n ew

c om b inat io n s to t ak e t he plac e of wo rn o u t s t ock -

.

The fact s j u s t pres ent ed h o wev er c lea rly ind icat e that if a , ,

typ e d oes n o t rem ain tr u e ind efin it ely eith er it is the fa u lt o f

ad v ers e s elect io n acc id ental o r o th er wis e o r els e it is d u e to
, ,

s o m e phy s ic a l o r b io lo gi ca l ca u s e fo r t he typ e o nc e o bta in ed , , ,

nat u rally breed s tr u e .

A gain fro m t he fact that v ariab ility is n o t great ly red u c ib le

, ,

we a re s afe in a s s u m ing that types o nc e es tab lis h ed by s electio n

will n o t o n ly rema in tr u e b u t a re capab le o f s t ill f u r ther d ev elo p
m ent if we bes t o w add it io na l att ent io n and s elect io n a n d t ha t ,

t he u pp er lim it s o f im pr o v em ent a re fi x ed if fix ed at a ll by , ,

s o m e c irc u m s tanc e o th er th a n v ariab ility I t m ay b e b io lo gic al .

—lik e los s o f fert ility o r rev ersal of s electio n — o r it m ay b e

,

m echan ical b u t t he ca u s e what ev er it m ay b e that s et s a limi t

, , ,

t o im pro v em ent is n o t c o nn ect ed with v ari ab ility .

I n t he la s t analy s is h o wev er we are b o u nd t o ra is e t he q u es

, ,

t io n wh eth er all types c a n b e ind efinit ely ma inta ined ev en by t he ,

1 Pea rs o n , G ra mm a r of S c ien c e, p 4 8 5 . .
 D ifferent ind iv id u als of a m e anc es try inh erit d iff erent ly
t he s ,

a n d in g en era l t he b eha v io r o f C ha ra ct er s i n tran s m is s io n s u g

ges t s that th ey a re in so m e way m ad e u p of c o m b inat io n s s o ,

that a h igh d egree o f v ariab ility is in ev itab le ev en with t he ,

s a m e element s ; a s fo r in s ta n c e a gr eat v ar iety o f c o lo r effec t s

, ,

c an b e pro d u c ed wi th t h e s am e three pri mar ies red b lu e and

, , ,

yello w .

S o m e C haract er s b lend and o th ers a re m u t u ally exc lu s iv e ,

each t end i ng t o preser v e i t s i d ent ity On th is acc o u nt as well

.
,

as fro m o th er ca u s es s u ch as
,
relat iv e fert ility rac es o ft en ,

exh ib it d is t inct p o ly m o rph is m I nh er itanc e is n o t s o m u ch c o n

.

n ec t ed with s ex a s i s p o p u larly s u pp o s ed C ha ract ers o ft en

.

d o n o t d ev elo p u nt il lat e in lif e T h is is n o t to b e regard ed as

.

belat ed in h er itanc e b u t a s belat ed d ev elo p m ent .

T he o nl y pro p er way t o s t u dy t he pr inc ip les o f h ered ity is by

s tat is t ical m eth o d s u s ing gro u p s i n s t ead o f s ingle ind iv id u al s
, ,

fro m which n o gen eral c o nc lu s io n s can b e s afely d rawn .

T he regres s io n tab le bri ngs o u t c lear ly t he fact th a t lik e

paren t s beget u n lik e o ff s pring ; that in genera l t he o ff s pring , ,

is mo re m ed iocre than t he parent b u t th a t fo r s elec ted o ff s pring

,

t he anc es try is c o m parat iv ely m ed io cre ; that t he c oefli c ien t of

h ered ity b et ween t he n eares t relat iv es is seld o m abo v e
that t he m ea n of t he o ffs pri ng is n o t n ec es s a rily t he s a me as t he
m ean o f t he parent ; that t he m ean s o f a rac e a re it s m os t fert ile
po rt io n s ; that i n genera l a few o ffs pring exc eed t he prev io u s
, ,

limit s o t e ra c e
f h ,
— that is pro gres s a way fro m t he type if
,

fav o red by s elect io n ; that exc ept io nal ind iv id u als m ay a ris e
eith er fro m exc ept io na l o r fr o m m ed io cre parentage ; and that

s u cc es s i v e o ff s pr ing fro m t he s a m e parent s a re n o t id ent ic al .

N o th ing is Clearer than that t he in ev itab le c o n s eq u enc es o f

b is ex u al repro d u ct io n and of the m anner o f gro wth by the halv
in g o f t he c ell c o nt ent s is t o in s u re th a t cha ract er c o m b inat io n s
eff ect ed in th is m ann er a re br o u ght t o geth er in d efin it e m ath e

m a t ic al pro p o rt io n s n o t far fro m th os e expres s ed i n t he expan

s io n o f a b in o m ia l T h is is t he rea l fo u ndat io n o f M end el s law
'

,
.

fo r C hara ct ers that d o n o t b lend and it a lso express es t he rela

,

t iv e pro po rt io n s o f charact ers that d o b lend .

T he s t a t is t ica l m eth o d s of s t u dy enab le u s t o d ev elo p t he law

o f a nc es tra l h ered ity wh ich agrees c lo s ely with exp erim ental
,
 H ER ED ITY 54 7

d nc e and wh ich s h ows the d egree to wh ich the v a rio u s gen

ev i e ,

o rat io n s ha v e c o ntri b u t ed t o the res u lt s .

C o nt in u ed s elect io n will s hift the typ e i n a ny d es i r ed d i rec tion ,

and aft er a few generat io n s i t will b r eed tr u e i n i ts n ew f orm .

A s v ariab ility is n o t great ly red u c ed by s elect io n th ere is a lway s ,

opp or t u n i ty i r ovem en t s o far as v ariab ili ty is c o nc ern ed

f or mp .

S PE C I A L E x E R C I S E S

1 . E x er c ises in grea t v a r iety i n fo rm in g regress io n ta b les a n d in d ed u c in g

the c o n c lus io n s .

2 . I n v es tiga t io n s i n to the Opera t io n s o f M en d el s law, b y the

’
ex a m i n a

t io n a n d id en ti fi c a t io n o f la b o ra to ry m a ter ia l an d if pos s ib le b y the a c t u al

r a is i n
g fo rm
o f c ros s ed
s fo r th e p p
ur os e .

3 S pec ia l a n d d efin it e a p pli c a tio n s o f the la w o f an c es t ra l

. h ered i ty
to the p ro b lem s o f the b reed er, es pec ia lly i n c ross i n g , gra d in g, an d lin e
b reed in g .

ADDITIONAL REF ERENCES

A LT E R N AT I V E I N HE R IT A N C E . By K a rl Pears o n . Pro c eed i n gs of the

Ro y al S oc iety, L X X I I , 5 0 5 5 1 0 -
.

AM E R ICAN T R OTT I N O R E C OR D S A S D AT A F OR HE R E D I T Y S T U D I ES .

B y F ra n c is G a lto n . Proc eed in gs of the Ro y al S o c iety, LX I I ,

3 1 0- 3 5 1 .

B A T E SO N O N PE A R S O N
’
s CO N C EPT I O N OP HE R E D IT Y . Proc eed in gs of

the R o yal S o c iet y , LX I X , ; Pe a r so n s a n s w

1 93 - 2 0 5
er , LX I X , 45 0
’
.

C HA N C E S OF D E AT H .B y a rl Pea rs o n K S c ien c e, V I , 3 2 8 3 3 0 .
— .

C ON T R I B U T I ON OF S E V ER A L A N C E S T O R S T o OPP S PR I N G B y F r a n c is .

G a lto n . Pro c eed i n gs

R o ya l S o c iety, L X I , 4 0 1 4 1 3
o f t he -
.

C RRO EL O
A T I N B E T W E E N LO N G EV IT Y A N D F E R T I L I T Y By a rl Pea r . K
so n Pro c eed in gs o f the R o ya l S oc iety, L X V I I , 1 5 9 1 7 9, 3 3 3 - 3 3 7
.
— .

C R I T E R I O N To T E S T T HE O R I E S O P H E R E D I T Y By arl Pea rso n . K

Pro c eed i n gs o f t he Ro y al S o c iety, L x x l l l , 2 62 - 2 80 .

D O V A R I ET I E S R U N OU T ? B y J C ra ig. . G ard en in g , 1 8 99 , pp . 2 7 8 —2 79
,
i n E x peri m en t S t a tio n R ec o rd , X 1 , 1 5 2
als o .

E x P E R I M E N T A L E V I D E N C E U PO N M E N D E L S L A w H Loc k
’
By L
—
. . . .

L X X , 60 1 6 0 2 ; b y
N a tu re, a r l Pea rs o n , 6 2 6- 6 2 7 K .

E x P E R I M E N T A L S T D I E S I N HE R E D I T Y U
C o rn R ep o r t . of the R o yal
S o c iety, 1 90 2 , p . 1 60 ; als oin E x per i m en t S ta ti o n R ec o rd , X V I I , 6 3 4 .

E x PE R I M E N T A L Z OOL OG Y B y T H M o rga n C ap ters V I a n d V I I , h

—1 66
. . . .

pp . 66 .

E X PE R I M E N T S IN C R OS SI N G W H IT E AND B L A C K OA T S . By ] H . . W ilso n .

N a tu re, 1 90 4 , p 4 3 ;
. E x 1
p e r i m en t S ta ti o n R ec o rd , X V I , 4 6 2 .
54 8 T R A N SM I SSION

E Y E C OL O R
. IN MA N . h h
P ilo s o p ic a l T ra n s a c tio n s of the R oyal S oc iety,
C X CV , A , 79 1 50 -
.

U
FO R M L A F O R R E G R E S S I O N . B y Pea rso n and Y u le . Pro c eed i n gs of the
Ro y al S o c iety , LX , 4 7 7 4 89 -
.

HE R ED I T Y OP C O A T C HA R A C T E R S IN PI G S AN D R A B B IT S . By W . E .

C as tl e S c ien c e, X X I , 7 3 7 —
.
7 3 8 , 98 6 .

HI STO R Y O F T H E D E V E L O P M E N T O P T HE Q U A N T I T A T I V E ST U DY OP
V A R I A T I ON . By C B . . D av en po r t S c ien c e, V I I I , 86 4 ; Pro c eed ings
.

o f the A m er ic a n A ss o c ia tio n fo r th e A d v a n c em en t o f S c ien c e, 1 900 ,

X L IX, 1 97 —
2 00 .

H Y B R I D OR A N G E S . By Web b er and S wi n gle . S c ien c e, X V I I , 262

2 63 .

HY B R I D W HE A T S . By W .
J . S p illm a n . B u lletin N o . 1 1 5 , Offi c e of Ex
pe S ta tion s ; als o i n S c ien c e , X X , 6 8
ri m en t .

I N HE R IT A N C E I N C OA T C O L O R , T HO R O G HB R E D U HO R S E S . B y B la n
cha rd . B io m etri ka , I,
3 6 1 —3 6 4 ; b y a r l P ea rs o n , K h
P i lo s o p ic al h
T ra n s a c t io n s of th e R o ya l S oc i et , C X C V , A , 1 —49 y .

I N II E R I T A N C E OF FE R T I L I T Y .
( R a c e ho rs es an d the h u man r a c e.
) By
Ka rl Pea rs o n . S c ien c e, I X , 2 8 3 - 2 86 .

I N HE R I T A N C E OF M E NT A L C HA R A CT E R S IN MAN . By K a rl Pea rs o n .

Pro c eed i n gs of th e R o y al S o c ie t y , LX I X, 1 53 - 1 55 .

LA T E N T C HA R A C T E R S A ND R EV E R SION . By W . E . C as t le . S c ien c e,

LA w O P A N C E S T R A L HE R E D I T Y K ka
'

. By a rl Pea rso n . B io metri , l l,

2 1 1 - 2 2
9, 2 3 I 2 3 6 -
.

LA w OF HE R E D I T Y . By C . B . D a v en po r t . S c ien c e, V I I , 1 58 -
16 1 .

LA W O F R E V E R S I ON . By K a rl Pea rso n . Pro c eed in gs of the R o yal

S o c iet y, LX V I , 1 4 0- 1 6 4 , 2 4 1 - 2 4 4 , 3 1 6- 3 3,
2 3 4
2 -
3 7
2 .

LA W S OP A N C E ST R A L HE R E D I T Y . By K a rl Pears o n . S c ien c e, V I I,
3 3 7 3 3 9. 5 5
- 1- 5 54
L A ws or HE R E D I T Y OP G A LT ON AND M EN D EL , AND S OM E L A ws
G O V E R N I N G I M P R OV E M E N T B Y S E L EC T I O N B y W E C as tle Pro . . . .

c eed in gs o f th e A mer ic a n A c a d e m y o f A rts a n d S c ien c es , X X X I X ,

2 2 1- 242 .

LI M I T S or V A R I A T ION IN PL A N T S .
( A u t ho r sa ys v a ria tio n is in ma the
m a ti c a l ra ti o .
) By J . W Ha rs h b erger
. . S c ien c e, X I I I , 251 .

L ON G E V I T Y AND T HE S E L EC T I V E D E A T I I R A T E . Pea rso n a nd B eeto n .

Pro c eed in gs o f the R o ya l S o c iet y , LX V, 2 90 - 30 5 .

M A T HE M A T I C A L C ON T R I B U T I O N TO T HE T HE O R Y O P H E R E D I T Y . By
K arl Pea rs o n . Pro c eed in g s o f t he R o ya l S o c iet y, LX X I , 2 8 8- 3 4
1 .

M AT HE M A T I C A L E V O L U T I ON . By K a rl Pea rso n . Pro c eed in gs of the

R oyal S oc ie t y , LI V , 3 2 9 .

M AT HE M A T I C A L E V O L U T I O N . By K a rl Pea rs o n . Pro c eed i n gs of t he

R oya l S oc i e ty , L X I V : G e n e t i c S ele c t i o n , 1 6 3 - 1 6
5 ; I n h erita n ce of

Fertili ty, 1 6 5 66 ;
- 1 I n eri t a n c e o f F ec u n d i ty ,h1 66 - 1
6 7 .
55 0 T R A N SM I SS I O N

R E G R E S S I O N , H E R E D I T Y , PA N M I x I A By K a rl Pea rso n Proc eed in gs of

—
. .

the R o ya l S oc iety, LI X , 6 9 7 0 .

R EPR OD U CTI V E S E L EC T I ON By . K a rl Pea rs o n . Pro c eed i n gs of th e

R o y al S o c iet y , L I X , 3 0 1 - 3 04 .

S EC O N D -
G EN E R ATI ON HY B R I D S . B y Ha ls tea d an d Kel se
y. N ew Je yrs e

E x per im en t S ta tio n R epo r t, 1 90 2 , pp 3 7 7 3 95 ;

. E x per i m en t S t-
a t io n

R ec o rd , X V , 1 5 2 .

S K E W V A R I A TI ON B y . K a rl Pea rs o n . Pro c eed in gs of the R o yal S o c iety,

L V I I , 2 5 7 2 60 -
.

S T A T E M E N T OF M E N D E L b ib l io g ra p y ) B y W E C as tle
LA w W h h
'

’
S .
( i t . . . .

S c ien c e, X V I I I , 3 96- 4 0 5 ; a lso b y L H B a ile , X V I I , 4 4 1 - 4 5 4 . . y

T E L EG O N Y I N M A N By a rl Pea rs o n
. K
P roc eed in gs o f the R o yal .

y
S o c iet , L X , 2 7 3 2 8 3 -
.

T HE S T A T I ST I C A L U D Y O F E V OL U T I ON B y C B D a en po rt
ST . . . v .

Po p la S c ie n c e M o th ly L I X 44 7 460
u r n , ,
-
.

V A R I A BI L IT Y OF I N DI V I D U A L A N D R AC E B y K a rl Pea o n Pro c eed rs .

i gs o f the R o y a l S o c iety LX V I I I 1 —
.

n 5 3 7 2 3 73 , , ,
-

V A R I A T I O N A N D C O R R E L A T I ON I N M A N — C I V I L I Z E D A s C O M P A R E D
.

W I T H PR I M I T I V E R A C E S . B y Ka r l Pea rs o n S c ien c e, V I , 4 9 5 0
.
-
.

W ON D E R HO R S E S AND M E N D EL I S M .
( S ev era l gen era t io n s o f o rs es h
with v er y lo n
g m a n es a nd ta i ls .
) B y C B D av en po r t . . . S c ien c e, X I X ,
1 5 1- 1 53 .
 CH A P TE R XV
PR E POTENCY
“

T hat a ll pa rent s a re n o t eq u ally po werf u l in im press in g

rac ial C haract ers is a fact well known to t he m eres t n o v ic e in
breed ing I t is d is t inct ly s h own in all regres s io n tables and
.
,

t he rea s o n fo r it is C lear ly seen in t he m ath em at ica l nat u re o f

reprod u c tion by wh ich ind iv id u als a re d iff erent ly end o wed and
, ,

by wh ich so me few a re exc ept io na lly r ich in t he elem ent s o u t o f

wh ich ra c ial charact ers a re d ev elop ed Wh en t o th ese fact s is .

add ed t he d iffi c u lty o f s elect ing an im als by o u t ward appearanc e ,

o n a cc o u nt o f the relat io n o f d o m i nant and rec es s iv e charact er s ,

we n eed feel n o s u rpr is e at t he relat i v ely s m a ll n u m ber o f

high ly prepo t ent ind iv id u als a n d t he large n u m ber of rev ers io n s
enc o u nt ered i n act u a l breed i ng .

SEC TI O N I — D ATA FR OM T H E T R O IT I NG R ECOR DS

'

I L LU ST R AT I N G PR E POT E N C Y
Seek ing m at eria l wh ich wo u ld i llu s trat e acc u rat ely , and with
su fli c ien tly large n u m b ers , t he d ifferenc es i n t he breed ing p o wers
of d ifferent ind iv id u als the writ er m ad e s o me s t u d ies in the
,

rec o rd s of tro tt ing bred h o rs es T h es e s t u d ies c o v ered all

-
.

an im als reg is t ered a n d that had m ad e track o r breed ing rec o rd s

“

fro m the o pen ing o f t he R eg is t er and t he Yearboo k d own to

and inc lu d ing t he y ea r
I n t he c o n s id erat io n o f th is m at erial and in t he c o m pari so n ,

o f i nd iv id u a ls f o u r fact s m u s t b e bo rn e in m ind : fi rs t s o me
, ,

i nd iv id u als were t o o y o u ng fo r th eir f u ll breed ing rec o rd t o b e

all i n ; s ec o nd s o m e had enj o y ed less o pp o rt u n i ty than o th er s
, ,

o wing t o th ei r rac ing engagem ent s ; th i rd s o m e s ta llio n s had ,

acc es s t o b ett er m ares and m o re of th em than had o th ers ;

, ,

1It i
s n ee d l t y th t th i p d l b i t k
es s o sa a s i g m y w k
ro v e a a o r o us as , c o v er n an ee s ,

wi th t w l lt
o c a c u a o rs .
 T R AN SM I SS I O N
55 2

fo u rth fas h io n has m u ch to d o ev en a mo ng rac e h o rses in

, , ,

infl u enc ing s elect io n es pec ially aft er an ind iv id u a l o r a fa mily

,

has acq u ired a rep u tat io n .

A llo wing a s f u lly a s po s s ib le fo r th es e fact s t he rec o rd s a re ,

wo rth s t u dy fo r t he light th ey thro w u po n t he q u es t io n o f

inh erent d ifferenc es b et ween i nd iv id u al s in res p ect t o breed in g
—
p o wers d ifferenc es so great that as we pro c eed it will b e
perfect ly ev id ent that t he line o f d es c ent ru n s thro u gh few
i n d iv id u als and q u it e in d epend ent o f the m as s :

T he t o ta l n u m b er o f p erf o r mer s lis t ed — that is that had ,

m ad e track rec o rd s go o d en o u gh t o ad m it th em t o the lis t

at th is dat e ( 1 90 1 ) was o f wh ich o r a lm os t

exact ly t wo th i rd s were t ro tt er s and 8 7 0 2 were pac ers

, ,
.

T he R eg is t er S h o wed th a t in all s ta llio n s had b een , ,

rec o rd ed at th is t ime b u t t he breed ing rec o rd s h o wed that o nly

,

6 2 7 8 o r les s than o n e in fi v e had pro d u c ed anyth ing in t he lis t ;

, ,
1

that is to s ay rou gh ly s peak ing 6 2 7 8 s ires had pro d u c ed

, ,

perfo r mers o r a n av erage o f,

each .

Great sir es Of th es e 6 2 7 8 s ires o n ly 2 0 7 had prod u c ed ten

.

o r m or e s i r es or da m s o f p
s eed ; that is o n ly 2 0 7 had br ed w e ll ,

en ou gh t o pr od u c e ei th er t en s ta llio n s each o r ten m ares each , ,

capable th em s elv es o f pro d u c ing s peed I n o th er wo rd s o f t he

2
.
,

wh o le s ta llio n s and 6 2 7 8 s ires o n ly 2 0 7 br ed s peed well ,

en o u gh t o s end it int o t he s ec o nd gen erat io n t o t he ext ent o f

eith er t en s ires o r ten da m s pr o d u c i ng s peed .

N o w th es e 2 0 7 great S ires thems elv es p r od uc ed d i rec t ly 5 3 7 7

—
perf o rm ers ( 4 2 2 6 1 1 5 1 wh ich is m or e t ha n on e fif th of .

the en t i re l is t of perf o rm ers a n d an a v erage o f 2 6 ap iec e o r , ,

s ix t i m es t he breed ing rec o rd o f t he a v erag e s ta llio n .

A ga in th es e 2 0 7 great s i res pro d u c ed 3 1 5 5 s ires of p erfo r m ers

, ,

and th ey in t u rn prod u c ed tro tt ers and pac ers

4 7 99 T h is is o v e r ha lf o f a ll t he S i re s and o v e r 6 2 p e r c e nt
o f a ll t he p erf o r mer s o f t he br eed .

1 h
T is wa s pa rt ly , as ha s b een l
a rea dy n o te d , b ec a u s e so m e in i v i d d l ua s were t oo
yo un g to av e ma h de a fu ll b re edi g nd rec o r .

1
’

T is , h o f c o u rs e, d o es n o t i l d th
nc u e o s e S i res w h ose p ro d uc e in s ires an d d ams

t o get h e r eq u a e l dt en .

3 N o te t h at 4 2 26 1 1 5 1 p m ean s 4 2 2 6
. t ro t te rs and 1 1
5 1 p a c e rs .
 T R A N SM I SS I O N

BRE ED I NG R ECOR D OF T HR E E C LASS ES OF ST A L LIONS : I , SIR ES OF

SIR Es ; 2 , SIR ES OF D A Ms ; 3 , SI R ES OF BOT H SI RES

A ND D A MS O F S PEED

C LA SS 1 C L A SS 2 C L
A SS 3

N u m b er o f s i res

To ta l p erfo rm ers go t t en direc t l y b y th es e s ires

R a t io per s ire
S ires of perf o rm ers
go tt en by ea c h c lass

R a tio pe r o rigin a l s ire

Perfo rm ers go t t en b y t h es e s ires
( l
i ne 4)
R a t io p er s ire ( in e l
R a t io t o o rigin a l s i res ( lin e I )
D a ms of p erfo rm ers go t t en b y o rigin a l s ires
R a t io t o o rigi n a l s ires

Perf o rm ers p ro d uc e d b y th es e d a ms ( li ne 9)
R a t io per d a m .

R a t io to o rigin a l s ires l( i ne 1 )
h
Perfo rm ers ( lin e 4 ) t a t were a ls o S ires o f s p ee d
R a t io t o o rigin a l s ires ( in e 1 ) l

4. On e t d
ou s i e c i rc u m s tanc e h elp s t o reli ev e t he b u rd en o f
i nf erio rity res t ing o n C la s s 2 A s ta llio n belo nging to an u n
.

fas h io nable lin e wou ld b e u s ed b u t litt le o r n o t at a ll in the s t u d ,

wh ile a mare belo ngi ng t o an eq u ally u nfas hio nable s tra i n wo ul d

n o t b e eq u a lly barred T he res u lt o f th is d is c ri m in at io n in t he
.

lo ng ru n and u nd er o u r m eth o d s o f s t u dy wo u ld appear in t he

, ,

fo r m of sires of da ms T o so me ext ent it m ean s n o t that these

.
,

S ires d id n o t p r od uc e m a les b u t that b ei ng u nfa s h io nab le th es e

, , ,

ma les had litt le o pp o rt u n ity T h is pro bab ly d oes no t acc ou nt .

fo r all d iffer enc es ev en t ho u g h t he t u rn s and cap ric es o f

,

fa s h io n a re hard er on S ires tha n o n da ms .

I t m u s t n o t b e f o rgo tt en in th is c o n n ect io n that th es e I I 3 S i res

c o n s t it u t e m o re than o n e half o f the 2 07 great es t S ires o f the rac e .

T h ey c ou ld n o t th erefo re ha v e b een s o v ery u nfas h io nab le

—
.
, ,

5
. C la ss I m u s t b e larg ely w hat i t s eem s t o b e br e ed e r s ,

o f S ir es rath er tha n o f da m s beca u s e th ere is n o rea s o n wh it

, y s

fem a le o ffs pring s h ou ld hav e been s u ppres s ed I t is c learly .

su per io r t o C la ss 2 b u t i nferio r t o C la s s 3
, .
 PR E POT EN C Y 555

6 C las s 3 ev id ent ly
. represent s t he cream o f the rac e ,

exc eed in gly pro lific and h igh ly fa s h io nab le — t he m o s t s u cc es s ,

fu l gett ers bo t h o f s peed and o f breed ers .

7 .T h ese 8 5 s i res th em s elv es pro d u c ed d i rect ly 2 5 8 1 S i res o f

perf o rmers ( 30 apiec e) th is n u m ber being o v er 4 0 per c ent of

,

all t he S i res o f t he breed T h ey pro d u c ed d i rect ly 3 7 4 6 p er

.

fo rmers or 1 4 per c ent o f all in t he lis t The 2 5 8 1 s ires d i rect ly

, .

go tt en by th em pro d u c ed perf o r m ers o r o v er 5 6 per c ent ,

o f a ll in t he lis t T h is m ean s that a li tt le o v er one p er c en t of the

.

s ir es a r e ra nd s i r es to ov er ha lf the b r eed
g .

The b ig t en h ighes t relat iv e exc ellenc e is well with in

. B u t t he
th ese 8 5 great breed ers T h eir av erage get o f perfo rm ers was .

44 ,
o r o v er t en t im es t he a v erag e o f t he ra c e b u t o u t o f t he ,

regis t ered s ta llio n s ten and ten o n ly pro d u c ed d irect ly , ,

as m any as a h u nd red o r m o re p erf o r m er s each T he tab u lat io n

1
.

o f t he br eed ing r ec o rd o f th es e t en great es t s ta llio n s is go o d

read ing fo r t he s t u d ent o f prepo t ency as it sh o ws a breed ing ,

power wh ich fu lly j u s t ifi ed th eir fa me as great pro d u c ers n o t

o n ly o f S peed b u t o f s ir es and d am s o f s peed .

TA B LE S HOWIN G T HE R ECOR D O F T HE T EN G RE AT EST PR OD U C ER S

OF S PEED 2
U P T o AN D I NC L U D I NG 1 90 1

SI R D E BY
jT R O TT E Rs P A R
CE S TO T AL
N u two o d 600 B elm o n t 64
E l ec t io n eer 1 2 5 Ha mb l eto n ian 1 0

O n wa r d 14 1 1 G eo . W il k es 5 9
1

R ed W il k es 1 7 49 G eo . W ilk es 5 19

A l c an ta ra 729 G eo . W il k es 5 19

Pilo t Me iu m d 1 5 79 Ha p py M ed iu m 400
S immo n s 2 7 44 G eo . W il k es 5 19

W il to n 598 2 G eo . W ilk es 5 19

G a mb etta W ilk es 46 5 1 G eo . W ilk es 5 19

B a ro n W ilk es 4 7 5 8 G eo . W il k es 1
5 9
T o tal

A v era ge

1 On e of t h es e o es
g in at 99 .
2 T ro tt ers an d p a c ers .
55 6 T RA N S M I SS I O N

T h is is 3 2 t im es t he breed ing rec o rd o f t he av erag e S ire an d ,

n ea r ly fi v e t im es t he rec o rd o f t he 2 07 grea t s i res t h em s elv es ,

inc lu d ed o r o v e r s ix t im es th eir rec o rd exc lu s i v e o f t hes e t en

,
.

I t is wo rt h wh ile t o n o t e t he s ires o f t h ese great breed ers O f

s peed N u m b er 2 is by Ha m b let o n ia n 1 0 ; N o 6 is by Happy
. .

Med iu m he by Ha m b let o n ia n 1 0 ; N o I is by Belm o nt by

,
.
,

A bd a llah he by Ha m b let o n ia n 1 0 ; and t he rem ain ing sev en

, ,

N O S 3 4 5 7 8 9 1 0 a re by G eo “ilk es by Ha m b le
'
.
, , , ,
.
, , , ,

t o n ia n 1 0 T h u s o f th is rema rk a b le lis t o f t en s t a llio n s all

.
, ,

exc ept o n e a re b u t t wo rem o v es fro m Ha m b let o ni a n 1 0 Of th is .

n u m b e r th o se t ha t were s ired by Geo Wilk es prod u c ed 6 4 0

,
.

tro tt er s and 2 3 2 pac ers — in a ll 8 7 2 pe rfo r mers o r m o re than , ,

66 pe r c ent o f t he wh o le .

The f amou s gran ds ires Eight s ta llio ns o f th is lis t ha v e t he .

d is t inct io n o f being g ra nd s ire t o o v er 5 00 pe rfo rm ers as fo llo ws ,

T A BLE OF FA MO I G RA N DS I R FS HA V I N G M O RE PER FO RM ERS

'

S 5 00 OR
‘
IN T HE S EC O N D G EN E R AT IO N

Per T o ta l Per Pe r fo r m in g
Dam s
fo rm ers form er : S ires and G et

G eo . W il k es
Ha m b l et o n ia n 1 0

E l ec t io n ee r

N u t wo o d
l
Be m o n t
l
A mo n t
R ed W il k es
O n wa r d

1 Co l
m n 1 n a m e o f g ra n s ire ; c o u m n
u . d
n u m b er o f p erfo rm ers o
f h i s o wn l 2

get l
c o u m n s 3 a n d 4 n u m b er o f s i res h e go t Wi t ,t ei r get in t h e is t ; c o lu m n s , h h l
5 a n d 6 n u mb e r o f a m s h e go t wit t h d
e p e rfo rm ers ro p p e b y t em ; c o u m n 7h d d h l ,

a m s — t h e s ec o n
, ,

t o t a l p er fo rm ers go t t en b y s ires an d g en e ra t io n ; co u mn 8 d , d l ,

l
n u m b e r o f pe rf o rm e rs ( c o u m n 2 ) t a t were a ls o s i r es a n d t ei r perfo rm i n g g et h , h .

NB y
Ma n o f t he s ires ( c o u m n 3 ) were n ot perf o rm ers a n d man o f t he
. l y
p e r f o rm e rs ( c o u m n 2 ) were n o t s i res l N u m b ers in
p a ren t es es refer to ea r of . h y
b irt h an d d ea th .
 8 T R A N SM I SSI ON
55

generat io n than in t he fi rs t b u t als o that th eir ,

age is again s t
th ei r sec o nd gen era tio n rec o rd -
.

R el ati on b et ween per f orman ce an d b reedi ng powers . An at

t em pt was m ad e to learn wh ether perfo rm ers a re bett er breed ers

t ha n n o rfo rm ers T h ere were at that t ime 4 9
n -
pe . s tal lio n s i n

t he lis t On ly 2 1 o f th es e had get in t he

. lis t and ,

o n ly f o u r had pro d u c ed s ires o f s peed .

T he breed ing rec o rd o f th is c la s s o f s tallio n s loo k s p it if u lly

S lim a s c o m pared wi t h that o f t he gr eat breed er s T he bes t .

breed ing rec o rd m ad e by a h o rs e in t he lis t u p t o t he ,

t ime t hes e s t u d ies were m ad e was that o f N elso n 4 2 09 who , ,

—
had prod u c ed 2 8 1 2 p eight s ires ( 5 7 and three da ms
.
,
-

Th e wh o le 4 9 in t he li s t had pro d u c ed o n ly
( 1 2-

1 94 6 5 p - and o n ly I 3 sires o f s peed 8 o f wh ich ha v e j u s t been

.
, ,

cred it ed to N elso n .

W e m ight c o nc lu d e that perfo rm anc e is no t a v ery goo d in d ex

o f breed i ng po wer b u t i t wo u ld b e a ha s ty c o nc lu s io n if m ad e
,

on th is ba s is T wo c irc u ms tanc es c o n s p ire t o k eep d o wn t he

.

breed ing rec o rd of s t al lio n s o f extreme s peed One o f th es e is .

t he fact t hat m any o f th em a re y o u ng and t he o th er is t he fact ,

that a hors e capa b le of ma k ing low rec or d s is wor th m ore f or

ra c i n th a n f or b r eed i ng p u rpos es a n d wh i le rac ing enga gemen t s
g ,

d o n o t ab s o lu t ely prev ent breed ing a m o ng s ta llio n s as it d oes ,

a mo ng m ares u nt il th eir rac ing day s are o v er yet it o pera tes

1
,

t o gr eat ly red u c e it E v id ent ly we S ha ll get litt le light o n o u r

.

q u es t io n fro m th is s o u rc e .

T u rn ing t o ind iv id u als we find that N u twoo d 600 the g reat , ,

es t s ire o f s peed ( s ee tab le o n page had a rec o rd of 2 ,

b u t that Elect io n eer t he n ext great es t S ire o f s peed had n o , ,

rec o rd Of t he b ig ten b u t o n e has a rec o rd as good as

.
“
,
”

and his breed ing rec o rd is t he lo wes t o f t he lo t .

T u rn ing t o t he great es t grand si res o f s peed Geo W ilk es ,

.

h ead s the lis t with a rec o rd of b u t Ha m b leto n ian 1 0

c o mes next hav ing pro d u c ed mo re S ires o f s peed than a n y h o rse
,

1Th w ere l 49 m i th
e re a s o l i t — t g i id a res — t n e s , a s ra n e co nc en c e, no

one o f wh m h d p d o d y th i g i t h l i t Th i f t i f
a ro uce an t t b n n e s . s ac s, o c o u rs e, n o o e

t
c o n s ru ed t m t h t th y l d t p d
o ea n ap d b t
e t h t
cou h t t h y h no ro uce s ee , u ra er a e av e

no t , l h d th pp t ity W h t k i d f b d m th y w ld m k
as a c ass , a e O o r un . a n o roo a res e ou a e

wh en t i d i r e th t
s an o
q i er u es on .
 PR EPOT E N CY 5 59

liv i n d ead and he has n o rec o rd ; Elect io neer 1 2 5 c o m es 1

g or ,

next als o wi t h n o rec o rd ; th en N u t woo d

, Belmo nt n o , ,

rec o rd follo wed by A lmo nt 2 :3 9 2 R ed W ilk es and , ,

On ward ,

F ro m th is s h o wing o f ind iv id u a ls we can argu e eith er that t he

great breed ers were too b u sy to m ak e rac ing rec o rd s o r that
breed ing po wer is i nd epend ent o f t he ab ility t o perfo rm .

N ei th er c o nc lu s io n is war ra n t ed I n t he fi rs t plac e t he breed .

,

in g rec o rd o f a s ta llio n wit h a track rec o rd is inj u red by the

fact that he has litt le oppo rt u n ity u nt il his rac ing day s a re
o v er ; b u t it is h elped by t he fact that wh en he is s ent t o t he

s t u d he ha s a s up eri or c la s s of ma r es .

A ga in it d oes n o t f o llo w if a h o rs e ha s n o t m ad e a track

, ,

rec o rd that he s h o u ld b e c o n s id ered as u na b le to d o it I t m ay

, .

b e fr o m s o m e s light defect o r fro m lack o f pro per trai n ing fro m ,

s o m e m i n o r inj u ry o r fr o m o n e o r m o re o f a h u ndred o th er
,

ca u s es ha v ing n o c o nnect io n what ev er with his inh erit ed ab ility

t o go .

E v id en t ly , if we any light u po n th is q u es t io n fro m

a re t o get
th is s o u rc e — and it o u ght t o b e o n e o f t he bes t o f s o u rc es for
,

inf o r m at io n of th is c la s s — th en we m u s t o bta i n it fro m large ,

n u m bers in which the breed ing rec o rd o f th o s e kn o wn t o b e

,

perfo r m ers is c o m pared d irect ly with that o f th os e that hav e n o

perfo r m a n c e rec o rd .

A cc o rd ing ly a tab le was prepared exh ibit ing t he breed ing

rec o rd o f 1 6 5 o f t he pr inc ipal s ta llio n s Th ey were C h o sen fro m .

t he lis t o f 2 0 7 that had pro d u c ed t en o r m o r e s ires o f s p eed o r

t en o r m o re da m s o f s peed and i nc lu d ed a ll i n d i v id ua ls tha t ha d ,

p r od uc ed b oth p erf or m i ng a nd no n p erfor m i ng s i r es

2
e xc ept a v e-
ry ,

few t he data fo r wh ich were inc o m p let e T he tab le s h o ws fi rs t

,
.
,

( c o lu m n I ) t he t o ta l n u m b er o f p erf o r m er s pro d u c ed by t he
,

It w
1
p p l l y b
as li d t h t
o H m b l
u ar t i e ev e ld g i a b ta b t
e on an 1 0 co u o n a ou u

h w e d d l gb f
as ea y b dy k w hi l
on e o re a nb d f p d
o ne s v a u e as a ree er o s ee .

By p f m
3 i m t th
er o r th t h ers t
s k ean d f 3 ose b tt
a av e a rac rec o r o 2 : 0, o r e er .

Th t m e er p f m“
no n l l w
-
i t h
er o rt ers d ;
”
i t m if
c o v ers l
a y i l d ou a rec o r an es t nc u es

tw l
o c a ss es ,th t h t m igh t h m d ose ad d i bl av ei m ta e a rec o r un er s u t a e c rc u s a n c es

an d th th t oseld t h a d cou d y
no i m t
ave A t h
on e so un i er a n c rc u s a n c es . s ere s no

w y f d i ti g i h i g b t w
a o s nth tw
u s th y
n e ll ll deen pes ef m o, d e a re a ca e n o n - er o r ers , a n

th t b l m k
e a e mp i a b tw th
es a c o th t h m d
ar son e d d ll th
een os e a av e a e a rec o r an a o ers ,

a bl bl
e or u na th t h t d e, a av e no o n e so .
 60 T R A N SM I SSI O N
5

va rio u s s t allio n s with o u t regard to their breed ing po wers ;

s ec o nd ( c o lu m n s 2 and t he n u m b er o f p erf or m i ng s ires ( that

is S ires with s peed rec o rd s 2 3 0 o r b ett er) with th eir get ;

, ,

th ird (c olu m n s 4 and t he n u m b er o f n o n perfo r m ing s i res -

,

with t heir get Fo r c o n v en ienc e th ere is add ed ( c o lu m n 6 ) t he

.

t rack rec o rd o f th ose s tallio n s (o f t he 1 6 5 ) wh ich were th em s elv es

perfo r mer s ”
fo r exa m p le lin e 6 : th is S ire m ad e a rec o rd o f
,

o n t he t rack He pro d u c ed 1 4 9 perfo r m ers 1 9 perfo r m

.
,

in g s ires that go t 1 1 1 perf o r m ers and 2 4 S i res that n ev er m ad e

,

a rec o rd on the track b u t that pro d u c ed 8 9 o ffs pring that were

perfo r m ers The en t ire table c o v ering 1 6 5 ind iv id u als is giv en
.
, ,

in o rd er that the read er may ha v e at hand t he m ean s o f mak ing

i nd iv id u a l c o m pariso n s ma n y o f wh ich a re to say the leas t
, , ,

remarkab le N a mes ha v e been o mitt ed b u t the read er may b e

.
,

int eres t ed t o kn ow that lin e 6 5 is G eo W ilk es 7 0 is Ha m b le .

,

t o nia n 1 0 1 2 0 is N u t woo d and 1 2 1 is On ward

, , .

BR EED I NG R ECOR D OF 1 6 5 L EA D ING ST A LL IONS T o S Ho w T HE R ELAT IO N

BET W EEN PER FOR M A N C E BR EED I NG POW ERS
”
AN D

Perfo rm i n g I Sires also Perfo r m in g Sires n ot Perfo r m in g

R ec o rd
G“ Perfo rm ers G et Perfo r m ers G et
 PR EPOT E N C Y 6
5 3

Perfo r min g i S ires 3 15 0 Perfo r min g S ires no t Perfo rm in g

R ec o rd
G et Perfo rmers G et Perfo rmers G et

K
) )
n v
i i a

o
t
w p.

N Nf :
‘
 6
5 4 TRA N S M I S S IO N

Perf o rm in g S ires a lso Pe rfo rm in g S ires no t Perfo rmi n g

R ec o rd
G et Perfo r me rs G et Perfo rm ers G et

W ha t light , d oes th is thro w u po n o u r q u ery ? We re

n o w,

m em b er t he d is ad v antag e o cc a s io n ed to t he perf o r m ing s ire by t he

i n t erferenc e o f track en ga gem ent s d u ring his rac ing y ears and ,

we rem em b er o n t he o th er hand t he ad v antage that c o mes fro m

, ,

his enha n c ed rep u tat io n and t he s u p er io r c la ss o f m ares o ffered .

T h is tab le tak en a s a wh o le s h o ws that th es e 1 6 5 lead ing

, ,

s tallio n s b ing h n f h igh s t n o t e that pr o d u c ed bo th p er

( e t e o e s o e

f or m i n
g and no n p f
er o
-
r mi ng s i r es ) pr o d u c ed 5 6 8 8 o ffs pri ng and

grand o ffs pri n g with t rack rec o rd s o f 2 30 o r bett er .

 66 T R A NS M ISS IO N
5

pro d u c ed o nly 4 0 perf o rm ers and 8 pe rfo r m ing s ires b u t G eo ,

.

W ilk es was o ne o f th es e a n d the a v erage o f , ( 74

1 8 ) t el ls

b u t a s m a ll part o f t he s t o ry o f th es e 8 pe rf o r m ing s ires o f th is

rem arkab le pro gen it o r o f s peed His 1 42 n on p erf or m i ng s ires .
-

prod u c ed s peed at a n av erage o f each I t wo u ld b e an .

int e res t ing s t u dy t o d et e rm in e ho w t he d es c endant s o f th ese

1 4 2 n o n pe rf o r m ing s ires c o m pared with th o s e o f t he 8 pe rf o rm
-

in g s i res — a s t u dy that t he wr it er ha s left t o o th er s

,
1
o r to a

f u t u r e t im e .

A c o n s erv at iv e c o nc lu s io n fro m t hes e data wo u ld b e that per

for ma n c e is n o t an in v ariable ind ex o f breed ing p o wer s b u t t hat
o n t he a v era g e t he p erfo r m ers a re m u ch m o re lik ely t o get s peed

tha n a re n o n p erfo r mers o f the s am e br eed ing

-
.

T h is d ifferenc e if it rea lly ex is t s is with o u t d o u bt inh erent ;

, ,

ind eed it is n o t d iffic u lt to find in s ta n c es in wh ic h that wh ich

,

s eem s to b e t he gen e ra l p ri nc ip le is rev er s ed s o that t he n on ,

rf or m er s a r e t/l e b etter b r eed er s ( s ee lin es 1 6

p e 4 4 2 79 1 3 0 , , , , ,

a nd 1 5 5 ) a ll o f wh ich s h o ws that wh ile g oo d n egat iv e t es t i mo ny

m ay b e fo u nd in a s ingle i ns t a nc e po s i t iv e s tat em ent s m u s t b e ,

bas ed u p o n a c o m preh en s iv e s t u dy o f large n u m bers A n d so .

we n eed t o go thro u gh o u r r ec o rd s and o u r exper ienc e caref u lly ,

h u nt ing fo r t he th ings that c o n s t it u t e gro u nd o n which p re

po t ency may s afely b e pred ic a t ed W ith o u t d o u bt p u r i ty of .

blood , i n tko tb c l / s t p os s i b le h
s ens e o f gu t e pe g f
r c en ta e o c a r a c ters

f a v or a b le to the p ur
p os e d es i r ed , n na [loy ed by d is tu r b i ng f ac tors ,
will b e f o u n d t o c o n s t it u t e the real bas is of prepo ten cy When .

d is c u ss ing the m ath em at ic s o f breed ing it was fo u nd that n o ,

m atter what t he c o m b inat io n s a f ew ind iv id u a ls will alway s ,

rem a in p u re By t he s a m e proc es s o f reas o n ing wh en we mix

.
,

t he elem ent s o f d es irab le character s d ilu t ing th em as litt le as ,

po s s ib le with wild b loo d we s hall by t he s a me la w o f pro b

“
, ,

abilit ies o nc e in a wh ile effect a ph eno men al c o m b in at io n S u ch

,
.

a o n e is pro d u c ed by m eth od s n o t u nd er o u r c o ntro l exc ept as ,

we i n c r ea s e t/t e p r oba b i li ty by i nc r ea s i ng Me i n ten s i ty f

o b r eed ing .

T h is is t he v ery h eart and so u l o f li n e breed ing

“
an d m ea n s
”

that t he b t bred an im a l is t he m o s t lik ely to b e prepo t ent

es -
. In
1 Wo r k f th i k i d
o i t d y w k
s n ru n s d m th
n o t a ps,i i g ee s, an on s, a a sur r s n ra te
.

T h e d a ta gi h
v en
p t m m
e re re
y t h f l b
re s en i w k
an on s o a o r o us or .
 PR EPOT E N C Y 6
5 7

t he m eant ime i t will b e well to remem ber that to t he b es t o f ou r ,

pres ent kno wledge s o m e ind iv id u als seem to b e preeminent ly

,

breed ers of perf or m ers o thers breed ers o f s ires and s t ill o th ers, ,

breed er s o f da ms ; wh ile a f ew are breed ers o f a ll three c la s ses .

I mportan ce of t he ac t u al t es t T he s t u d ent cann o t fail to n o t e

.

t hat the bu lk o f t he bu s ines s of real impro v ement is d o n e by a

v ery few rea lly great an i m a ls and that t he wo rk o f mo s t o f t he
,

s o ca lled breed ing s t ock is m er ely that o f repro d u ct io n in the

-

s en s e o f increa s e o f n u m b er s .

I t is p erf ec t ly c lear that he who is b ent u p o n acc o m p lis hing

real res u lt s will seek fo r the oc cas io na l pheno menal breed er ,

and hav ing f o u nd him will mak e t he mo s t o f him wh ile he is

, ,

ab le to repro d u c e I t is matt er o f d eep regret that so many of

.

o u r ph en o m ena l an im a ls lik e Ha m b let o n ian 1 0 were n ev er

, ,

rec o gn ized as s u ch u nt il lo ng aft er th ey were d ead and the

o ppo rt u n ity to u t ilize th em to t he bes t ad v antage had pa s sed

fo rev er lea v ing u s t o d o t he bes t we can and m ak e t he m os t of

,

t he acc id ent s th a t a re left beh ind .

I n s eek ing th es e ph en o m ena l breed er s t o o m u ch cann o t b e

s a id c o nc ern i ng t he im po rtanc e o f t he act u al breed i ng t es t as

t he la s t and fi na l cr it erio n o f breed ing po wer s — a s u bj ect o n ,

wh ich m o re will b e s a id lat er .

SECT I O N II — PR EPOT E N C Y I N SEX

T h ere is a trad it io nal belief that in gen era l the s ire is p re

po t ent o v er t he d a m I n act u al pract ic e th is is lik ely to b e t he
.

cas e fo r the s ire is in mos t cas es the bett er bred o f t he two

, , ,

parent s I f a breed er s tart s o u t to breed half blo o d s an d to giv e

.
,

his s t o ck t he m o s t ben efit p o s s ib le o f goo d b lo o d at t he lea s t

expen s e he will o f c o u r s e pr o v id e it throu gh t h e m a le s id e ; fo r
,

with o n e ma le he can infl u enc e t he b loo d o f m any o ffs pri ng ,

wh ile with t he fem a le he can infl u enc e b u t o n e in h o rs es o r

catt le and b u t few in swin e S o it c o mes a b o u t fo r p u rely
,
.
,

ec o n o m ic r ea so n s a lo n e that in gen era l s ires a re b ett er br ed

,

tha n da m s a n d o n th is acc o u nt s h o u ld b e prepo t ent

,
.

B u t t he q u es t io n o f breed ing as id e a re th ey prepo t ent b e

, ,

cau se o f sex On th is po int s pec u lat io n has lo ng b een at wo rk ,

 5 6 8 T R A N SM I SS I ON
res u lt ing in a ch o ic e c o llect io n o f beliefs c o v ering abo u t all ,

t he c o m b in at io n s p o s s ib le I t is h eld :
.

1 .T hat the m a le is prep o t ent o n gen era l p rinc ip les beca u s e ,

m a les a re s tro nger and m o re v irile than fem a les .

2 .T hat the fem a le is prep o t ent es pec ially a m o ng m a m m a ls

, ,

beca u s e her as s oc ia t io n s with t he o ff s pring a re s o m u ch m o re

int im at e bo th phy s io lo g ica lly and s o c ia lly
,
.

3 .T hat that par e nt i s pr e p o t e nt w h ich h as t h e s tr o ng e r

ner v o u s and s ex u al o rga n izat io n ,
what ev er that m ay m ean .

4 T
. hat t h e m a le i s pr e p o t e nt o v e r t h e f o r ward and u pp e r
part s of t he body and t he menta l q u a lit ies .

5 T
. hat t h e exact r e v e r s e o f t h e la s t s tat e m e nt is t h e tr u th .

6 T hat t he m a le go v ern s t he ext e rna l a n d t he f em ale t he

.

int erna l o rga n s and part s .

I n s tanc es a re n o t wa nt ing t o pro v e any o f th es e beliefs ;

ind eed pro o f by t he m eth o d o f i n s tanc e is t he f a v o ri t e fo r m o f
,

argu m ent fo r o r again s t any o ne o f th em and it is n o t too m u ch ,

t o s ay that by th is m eth o d th es e o r any o th er a s s u m pt io n s m ay

b e read ily s u b s tant iat ed .

We hav e lea rned lo ng ago t he u nreliab ility o f c o nc lu s io n s of

th is k ind a n d it is wo rth wh ile to d is t ingu is h c learly s o far as
, ,

we a re ab le b et ween what is kn o wn and what has n o t yet been

,

learn ed t o u ch in g th is im p o rtant m att er .

I n gen eral t h e sex es ar e eq u i pot en t S o far a s is n o w kn o wn

.
,

n o part o f t he ger m c ell is nat u ra lly pred es t in ed t o pr o v id e any

par t ic u lar part o f the bo dy T he ger m c ells fro m b o th parent s

.

a re b eare rs o f t he h ered itary s u b s tanc e in t he pro p o rt io n in

wh ich th ey p o s s es s it and eith er s ex m ay and d o es t ran s m it

,

any and all t he charact ers o f t he rac e to its o ffs pring o f eith er
s ex . W e m ay say th en in gen era l th a t th a t parent will b e
, ,

prepo t ent wh o s e h ered itary s u b s tanc e is leas t m ix ed and there ,

fo re m o s t int en s ifi ed a lo n g t he lin e o f es t a b lis h ed charact ers

, .

T he o n ly way we can go f a rth er than t he gen era l pr inc iple j u s t

s tat ed is by ext en s iv e s t u d ies s o lv ing t he c oe c ien t o [t er ed i t
fi f y
bet ween each p a rent and it s off s pring o f bo th sex es fo r d ifferent
ch a ract ers s eparat ely .

T h is has b een d o n e fo r a n u m b er o f charact ers b o th in m en ,

and in an ima ls th o u gh t he lis t is too s m a ll to d o m o re than t o

,
5 7 0 T R A N S M I SS I ON
2 . q u o t e Pear so n t he m a le s eems t o inheri t m o re than
To ,
“

t he f em a le beca u s e his c oe ffi c ient o f h ered ity is h igh er with ,

which ev er parent t he c o m paris o n is m ad e ( c o m pare lin es 1 3 5 , , ,

with lin es 2 4 T h is c o nc lu s io n Pear s o n d ec la res is c o n

, ,

firm ed by d a ta in eye c o lo r as well as in s tat u re c o at c o lo r and

, , ,

h ead ind ex l
.

I t is a s ign ificant fact th a t fo r t he ra c es and charact ers h ere

in v o lv ed t he c o rrelat io n bet ween bro t he r and bro th er is les s tha n
t he c o rrelat io n o r s im ila r ity bet ween s is t er and s is t er ( c o m pare
lin es 1 1 and 1 2 with lin es 1 3 and wh ich m ean s a lso that
s is t er s res em b le ea ch o th er m o re c los ely th a n d o bro th ers .

3 .T h e r es em b la nc e b et wee n m e m b er s o f t he s a m e s ex is

c lo s er than th a t be t ween m em bers o f o ppo s it e s ex es ( c o m pare

lin es 1 1 and 1 3 with lin e 1 5 ; a lso 1 2 and 1 4 with Pearso n
a ls o d ec la res th a t t he s a me princ ip le h o ld s fo r eye c olo r and
h ead ind ex and he is in c lined to believ e it general
,
2
.

T h is a u th o r po int s o u t th a t th is princ ip le if gen era l m ean s , ,

th a t i nh eritanc e in a line thro u gh o n e s ex is prepo t ent ov er

“

i nh er itanc e in t he s a m e d egree with a chang e o f s ex that is ,

that in heri tanc e t end s to ru n in s ex lines wh ich m ean s to , ,

q u o t e Pearso n (ita lic s and pa renth es es m in e) that a m a n in eye ,

c o lo r ( fo r exam p le) m o re c learly res em b les his pa terna l than his

ma ter na l grandfath er ( o r o th e r m a le anc es t o rs ) a wo m an m o re
c lo s ely res em bles her m at ernal g rand mo th er than her pa t erna l
g rand mo th er A gain a n ephew is m o re lik e his pat ernal u nc le
.
,

than his pat erna l a u nt ; a n iec e lik e her m a t erna l au nt than her ,

m a t ern a l u nc le 3
.

F u t u re in v es t igat io n s will a d d t o o u r k n o wledge in th es e

m a tt ers and p e rh a p s m o d ify s o m e gen eral s tat em ent s n o w c o n
,

s id e red s a f e b u t t he m att er a s s tat ed a b o v e r epres ent s t he bes t

,

c o n c lu s io n s o f th o s e who ha v e giv en m o s t caref u l att ent io n to

the s u bj ect u p t o t he p res en t t ime .

Com pa rat iv e v ar iab il i ty of t h e sex es T h ere ha s b een a .

gen eral tend en cy to as s ert that ma les a re m o re v ariab le t han

f em a les T h is a s s ert io n has n o t been bas ed o n act u al s t u d ies
4
.

1 Pea rs o n , G ra mm a r S c ien c e, p 4 5 9
of . .
2 Ib i d .
p 4 59 . .
3 Ib i d .
p 4 60 . .

4 S ee G e dd es h l
a n d T o m s o n , E v o u t io n S ex pp 1 2 - 1 3 ; D a rwin , A n i ma ls
of

—
, .

an d P l d
a n ts u n er D o m es t ic a t io n , I , 4 5 7 h
Pe a rs o n , C an c es o f D ea t , pp 2 56 2 60 h . .
 PR EPOT EN C Y 57 1

but po n t he th eo ret ical gro u nd that m ales lead a mo re act iv e

u

life and tak e t he lead in s ex u a l s elect io n The data j u s t c i t ed .

s eem t o s u b s tant iat e th is a s s ert io n Fo r t he s pec ies a n d charac .

t ers inv olv ed it appears that m a le o ff s pri ng fo llo w m o re c lo s ely

t he parenta l typ e than d o t he fem a le a n d ( which a m o u nt s t o ,

t he s a m e th i ng ) f em a le o ff s pri ng o r s is t er s a re m o re n ear ly a lik e , ,

than a re male o ff s pring o r bro th ers t end ing to the c o nc lu s io n

, ,

tha t m a les are m o re v ariab le than f em a les .

Pear s o n h o wev er rec o rd s data o f an exc eed ingly exha u s t iv e

l
, ,

s eries o f i n v es t iga t io n s o f v ar iab ili ty in m en and wo m en not ,

ab solu t ely b u t relat iv ely as expres sed in t he c oeffi c ient o f ,

v ar iab ility Wh ile he find s men m o re v a riab le at c ert a in ages

2
.

and in c ertain charact ers yet he d o es n o t fi nd pro no u nced and ,

d ec id ed d ifferenc es n o r a re th ese t he s ame fo r d ifferent rac es

,

o f m en He c o nc lu d es o n t he wh o le that fo r a ll rac es s t u d ied

.
, , ,

anc ient and m o d ern and fo r a ll charact ers c o v ered by t he ,

s t u d ies th ere is,

n o ev id enc e o f great er m a le v ar iab ility b ut ,

rath er of a s light ly great er female v a riab ility .

He find s fo r exa m p le that E nglis h m en a re s light ly m o re

, ,

v ariab le as t o h eight than a re E ng lis h wo m en b ut

that a m o ng Ger m a n s wo m en a re c o n s id erably m o re v ariable
, ,

than men a s th ey a re a ls o a m o ng the F rench

(4 3 5 3 8 8 )
I n t he lo ng bo nes
t imes o n e is m o re v ariable s o m et im es
som e ,

t he o th er ; th u s a s t o t he f em u r m en a re m o re v a riab le
, L ibyan ,

F rench A in o and N eo
lith ic m a n b u t with t he anc ient inhab itant s o f the
C anary I s land s t he rev ers e is t ru e ( m en wo m en , ,

I n a ll c ases exa m in ed exc ept t he F rench ( m en wo m en

, , ,

t he t ib ia is m o re v ar iab le in men ; b u t in m os t cas es t he

h u mer u s and rad iu s a re mo re v ariable in wo men .

1 Pea rs o n , C h an c es o f D ea t h, 61,
5 377 2 — .

9 M an if es t ly t h e c o effi c ien t o f v a ri a b i it is l y t he o n ly c o rrec t es t i ma t e o f c o m

p a ra t i v e v ar a i b i l it y , b ec a u s e in it s l
c a c u la t io n en en i n s ta n ce is c omp a r ed wi l l: i ts
own type as a ba se . h
T is is n ec es s a r y , b ec a u s e t h e s ta t u re of wo m en , fo r ex a m p e, l
is d if er en t fro m t h at of h
m en ; en c e t he t wo c a n n o t b e c o m p a re o n a ny c om m on d
b as is . h
T is is t he on ly way, fo r in s t an c e, as Pea rs o n p o in ts o u t in w ic we c an , h h
c o m pa re t h e v a ri a b il it y o f ma n wi t h h
t a t o f t he elep an t ; i n a n y o t er wa y t h e h h
e epl h an t wo u ld a ppea r l
m o re v a ria b e, beca u s e li e is bigg er .
 7 T R A N SM I SSI O N
5 2

av era ges
T he o f c oeffic ien t s of v ariab ility fo r all lo ng -
bo n e
d et er minat io n s a re as fo llo ws 1

Fe u c n T ra m Hl '
ll ER LS
'

R Aou s
'

T h o u gh thes e are hu m an n o t an im al data yet th ey inv o lv e , ,

s k eletal m ea s u rement s wh ich a re a m o ng t he m o s t f u nda m en ta l

o f a ll o rgan ic part s and th ey s carc ely wa rrant the s weep ing

,

as sert io n that m a les a re d ec id ed ly m o re v ariable t han females .

Pears o n is ent i rely j u s t i fi ed in h is pro t es t again s t what he ca lls

th is ps eu d o s c ien tific s u pers t it io n and t he s weeping c o nc lu
“ -

s io n s i n v o lv ing s o c ial and pra ct ica l c o n s eq u enc es affect ing

“

t he wh o le o f o u r c iv ilizat io n
“ 2 "
.

I n bo dy weight b o th a m o ng E ng lis h ( m en , wo m en , ,

an d a m o ng G er ma n s ( m en wo m en wo men , ,

a re d ec id ed ly m o re v a r iab le than m en
3
.

I n weight at b i rt h b o t h a m o ng E n g lis h ( b o y s ,
gi rls , ,

and a m o ng Ger man s ( bo ys girls t he , ,

m ales a re m o re v ar iab le ; b u t a m o ng t he B elgia n s t he rev ers e

is t ru e ( boy s girls , ,

D ata o f th is s o rt a re f u ll b u t u nf o rt u n a t ely c o nfined m os t ly ,

to h u m an s F ro m al l s o u rc es it s eem s th a t m en a re m o re v ari
.

ab le in h eight wh en s i tting and in s wiftn es s o f b lo w b u t "

,
”

that wo men a re m o re v ariab le in s tat u re Gi eight wh en s ta n d

s pan bo dy weight brea th ing capac ity s trengt h
” “ “
i ng )
”

, , ,

o f p u ll s q u eeze o f hand and keenn es s o f s ight

”

, , .

A s Pear so n po int s o u t s o me o f th es e v ar ia nt s wo u ld d isa ppear ,

if wo m en were s u bj ect ed t o t he s a m e c o nd it io n s o f life as a re

men and we n eed t o b e ca u t io u s wh en app ly ing th es e data t o
,

rac es in gen era l — fo r wh ich f u t u re r es earch es a re s o rely need ed

, .

We c ert a i n ly a re n o t warrant ed in a s s u min g s weep in g and f u nda

m ental d ifferenc es in v ariab ility bet ween t he s ex es Here agai n .

1 Pea rs o n , C h a n c es o f D ea t h , I 30 5
, .
2 Ib id . I, 2 5 6 and 3 76.

8 Inc i d en t a ll y , the sa me d a ta wa rra n t o ur l

c o nc u s io n t h at t he G erman ra c e,

bot h m en and wo m en , a re m o re v a ri a b e l th an t he E n g is l h a s to bo dy w igh t

e .
 T R A N S M I SS I O N
5 74

he fo u nd its d es c endant s h igh ly res is tant fl o u ris h ing well wh ere ,

o rd ina ry wh eat t o ta lly f a iled I t was apparent ly a tru e m u ta nt

.
, , ,

with extra s tro ng res is tanc e t o m agn es iu m .

I nflu ence of dev elopm en t u pon p repot ency M any b io lo g is t s .

c o ntend that t ra n s m is s io n d epend s to a la rge extent u po n t he

d ev elop men t o f t he parent ; th a t a s ta llio n tro tt ing bred fo r ex ,

a m p le wo u ld get s p eed m u c h m o r e s u cc es s f u lly if he h im s elf

,

were d ev elo ped or wo rk ed u po n t he track than wo u ld t he

s a m e s tallio n k ept eq u a lly h ea lthf u l a n d v igo ro u s b u t n o t d ev el

o ped a s t o act u a l s p eed A nat u ra l c o nc lu s io n o f th is c o nt ent io n

.

is o f c o u rs e that t he sa m e s ire will get m o re s p eed in his m idd le

, ,

and lat er y ears than wo u ld b e po s s ib le befo re he wa s d ev elo ped .

T his is t he v ery po int o f C a s per L R ed field s rec ent art ic les 1 '
.

s ett ing f o rth wh a t he ca lls his dyna m ic th eo ry o f h ered ity “

.

He bri ng s m any in s tanc es and m u ch argu m ent in s u ppo rt o f t he

as s u m pt io n b u t in the o p in io n o f t he wr it er t he m eth o d o f proo f
,

ad o pted is n o t c o m pet ent to s ett le t he q u es t io n n o r is any ,

m eth o d ab le t o d o s o that is b a s ed u po n t he s im p le en u m erat io n

o f i n s t anc es .

A s with any o t her q u es t io n in v o lv in g great v ar iab ility t he ,

o n ly wa y we can s ett le th is is by e m p lo y ing large n u m ber s o n

b ot/z s id es o f t he pro p o s i t io n ; in o ther wo rd s by c o m pari ng t he ,

s peed o f a ll t he h o rs es g o tt en by p erf o r m ing s ires lat e in lif e

with t he rec o rd s o f t he get o f t he sa m e s ires b ef o re d ev elo p ment ,

o r at lea s t b efo re lo ng s er v ic e o n t he tra ck E v en th en we m u s t .

learn what d ed u ct io n s to m a k e if an y o n acc o u nt o f d ifferenc es , ,

in age ; aft er wh ich we m a y h o p e t o lea rn t he real effect o f

,

d ev elo p m ent u po n prepo t en cy .

A s t he m att er s t a nd s n o w t he f ac t o f prepo t en cy is pat ent t o ,

bo th t he c as u a l o b s er v er and to t he c a ref u l s t u d ent ; b u t the

reas o n s fo r th is d ifferenc e in breed ing po wer s a re n o t yet at all
well u nd e rs t o o d Here as in m any o ther d irect io n s we awai t
.
, ,

f u t u re s t u d ies .

S u mm ary I nd iv id u a ls o f t he s a m e anc es try d iffer m a rv el

.

ousl i n th ei r breed ing p o wer s S o m e can pro d u c e exc ellenc e

y .

d irect ly in th eir o wn d es c endan t s and o th er s ind i rect ly throu gh ,

1S Th H
ee e m X XV N
or s e —
9 4
an ,
B d i g t h T tt
, os . l Tb
1 1, on ree n e ro er a so e

Am F i ld L X I I N
er zc a n e , 5 d L X,
I I I oN .
9 2 E , l t
ani f t h S t t , o .
, on vo u on o e e er
.
”
 PR EPOT EN C Y 5 75

r and da m s th ey a re a b le to get T he line o f d esc ent r u ns

s i es .

o nly thr o u gh t he few that can pro d u c e b r eed er s o f b r eed er s n o t ,

s im p ly pe rfo r m ers .

I nd iv id u a l exc ellenc e is n o t a c erta i n g u id e to breed ing p o wers ,

and many o rd ina ry ind iv id u a ls a re a mo ng t he grea t es t breed ers .

T h is is n eith er a m y s t ery n o r a fa u lt in h ered ity ; it aris es fro m

the fact that ind iv id u a l exc ellenc e is part ly a m att er o f ind iv id u a l
d ev elop men t and n o t a s u re ind ex o f real anc es tra l po s ses s io n s .

T he s pec im en m ay b e o n ly fa irly well bo rn th o u gh fa u lt less ly ,

d ev elo ped ,
i n wh ich ca s e he will pro bab ly b e a d is appo int m ent
as a breed er ; o r he m ay b e exc ellen t ly b o rn b u t o n ly fa ir ly well ,

d ev elo ped ,
in wh ich c as e he will breed b ett er than he is h im
s elf s t ill aga in he m a b e well b o rn and perfect ly d ev elo ped
y
, ,

wh ich is b es t o f all .

A ll s t u d ies yet m ad e S ho w that o n t he a v erage p erfo r mers , ,

( th o s e ind iv id u a ls p o s s es s ing h igh in d iv id u a l m e ri t ) a re b ett er

breed ers than n o n perfo rmers ; that is than t ho s e wh ich d o n o t

-
,

s h o w in th eir p er so na l d ev elo p m ent a h igh d egree o f exc ellenc e ,

th ou gh as we s ho u ld at o nc e s u r m ise th ere a re n u m erou s ex

, ,

c ep t io n s largely ar is in g fro m o u r inab ility to acc u rat ely j u dge

,

ind iv id u a ls by ext erna l appearanc es .

S PE C I A L E x e R C I S E s

W or k o u t s p ec i a l c as es o f re
p p o ten c y in th e b reed i n g a n d s peed rec o rds

o f t ro ttin g a n d ru n n i n g ho r s es , i n the a d v a n c ed reg is t r y o f c ow s, and in t h e

fa m o u s fa m ilies of b eef c a t t le a n d o f s wi n e . Pa y s pec i a l a tten tio n to rela
tiv e p repo ten c y o f o wn b ro t hers a n d t o t he c o r rela t io n b e tween i n d iv id u a l

per fo rm a n c e an d b reed i n g p o wers .

A DDI TION AL RE F EREN CES

A MEA S U RE OF I NT E N S IT v OF TR A S M I SS I ON
N . B y F ra n c is G a lto n , 1 8 99 .

N a tu re, L X , 29 .

D I ST R I B T IU ON OF PR E P O T E N C Y .
( T ro t ti n g -
ho rs e rec o rd s .
) B y F r a n c is
G alto n . N a tu re, L V I I I , 2 46 —2 4 7 .

I N F L U EN C E OF S Ex ON SI ZE OF OF F S P R I N G . By F . B . M u m fo r d .

E x per i m en t S ta t io n R ec o rd , X V , 5 4 2 .

PR E P OT E N C Y A N D X E N I A B y C C o rr en s . . . E x p er im en t S t a t io n R ec o rd ,
X I, 10 16 .

PR E POT E N C Y OF DI FF E R EN T PL A N T S . By W W . . T ra c y . E x perim en t
S ta t io n R ec o r d , X I I I , 3 2 4 .
 8 PR A C TI C A L PR O B L E M S
57

be d ne
o wit h
referenc e b o th to what is d es irab le and to what
will pay ; h en c e t he n ec es s ity o f c o n s id ering a ll pha s es o f t he
s elect io n pro b lem fro m a d o u b le s t a ndp o int Each c o n s id e rat io n .

o u t lin ed is s elf ev id ent et each is o f s u ffi c ient im po rta nc e t o

y
-
,

m eri t f u rth er c o n s id erat io n .

SECT I O N I — I D EA LS I N S ELECTI ON

A m o ng t he m u lt it u d e ariat io n s wh ich ev ery breed and

of v

ev ery v ar iety will pres ent t he breed er m u s t kn o w wh ich a re

,

u s ef u l . T he great m as s m u s t b e d is c a rd ed fro m t he m ere po int ,

o f n u m b ers and n o o n e ca u s e o f f a ilu re is m o re c o mm o n than

,

a v ac illat ing po licy rega rd ing s tand a rd s o f select io n .

T h is u nc erta inty is d u e t o n o o th er f a ct than that t he breed er

d oes n o t kn o w q u it e wh a t he want s He is in t he m ark et .
“ ”

fo r “
any go o d th in g th a t m ay t u rn u p I n t he c o u rs e o f his .

breed ing o perat io n s a great m a ny n ew and mo re o r les s pro mis

i n g th ings will appear U n les s he ha s u n lim it ed m ean s and
.

bo u nd les s s pac e fo r h is o perat io n s th es e m u s t b e d is card ed

,

wi th seem ing ru th les s n es s o r h e will s peed ily ha v e an a s s o rt

,

m ent of n o v elt ies wh ich if bred a m o ng th em s elv es will o v err u n

his prem is es and if bred int o h is pe r m an ent s t o ck will pro d u c e
,

a v er itab le j u m b le ou t o f wh ic h n o go o d th ing can c o me I n

,
.

th is way anc es try and ped igree can bec o m e s o h o peles s ly m ixed
as t o b e wo rth les s T h is m a y h a ppen with any breed and ev en
.
,

wi t h in t he lim it s o f p u ri ty o f b lo o d ; i nd eed it ha s happen ed

o v er and o v er ag a in in a ll breed s t h ro u gh the m is g u id ed en t hu
, ,

s ia s m o f br eed e rs wo rk ing wi th o u t well d efi n ed s tandard s -

.

S ta n da r d s wi sely fi x ed S t a n d a rd s m u s t n o t b e left to chanc e

. .

T h ey m u s t n o t b e warped o r a lt ered by n o v elt ies n o m atter ho w ,

c u r io u s o r attract iv e Th ey m u s t b e fix ed in ad v anc e lik e b u ild

.
,

in g plan s a n d s p ec ificat io n s a n d s h o u ld b e fix ed in t he light of

,

wlta t is n eed ed and rel/a t t/ze b r eed is li kely to af or d I nd eed .

,

t he s tandard s s h o u ld b e ro u g h ly fix ed b ef o re t he breed is ch o se n .

On ce c h osen , s ta n da r ds s h ou l d b e preserv ed u n ch an ged As .

t he art is t s ees his p ict u re befo re he m ix es his c o lo rs , and as t he

s c u lpt o r ch ip s a wa y at h is m a r b le t o br ing o u t t he part ic u lar

fi g u re t h a t s t a nd s i n his m in d u nd is t u rb ed and u n d is s u ad ed
,
 SELECT I O N
5 79

from p u rpo s e by the many af/zer exc ellent figu res t hat m tg/ct
his
b e c u t fro m t he s a me materia l s o th e breed er s h ou ld adh e re t o ,

his s tandard s d o gged ly Th ey s h o u ld b e wis ely ch o s en it is

.
,

t ru e b u t o nc e s u re o f that fact and with the law o f anc es tra l

, , ,

h ered it y in m ind no th ing s h ou ld warp t he j u dgment as to chan ge

, .

E v eryth ing that h elps t o s ec u re t he id ea l s h o u ld b e acc ept ed ,

and ev eryth ing else n o matt er ho w attract iv e in it s elf s h o u ld

—
, ,

b e p u s h ed a d e
s i u n les s ind eed
,
t he breed er ha v e u n lim it ed
, ,

m ean s and is m ind ed to d o n o t o n e th ing b u t m any th ings .

Keep b l ood lin es pu re B u t if t he breed er is m ind ed t o ind u lge

.

in experiment s o u t s id e t he ch os en s tandard th ese experim ent s ,

m u s t b e carr ied o n sepa rat ely B lood l in es m u s t b e kep t pu r e .

,

n o t p u re wi th in b r eed lin es s im p ly b u t r emem beri ng t he law o f , ,

anc es t ral h ered ity and the p u ll o f t he a n c es t o rs back o f t he

i mmed iat e parent th ey s h o u ld b e k ept a s p u re as select io n can
,

ma k e th em .

Ob j ect s of sel ec t i on I nd eed wh ile o n e o bj ect o f s elect io n is

.
,

to red u c e n u m be rs by far the large r o bj ect is to p u r ify the

,

a nc es try to t he en d that i nh er itanc e fr o m a ll t he a n c es t o r s s ha ll

,

b e alik e so that t he p u ll o f t he rac e s hall n o t b e d ifferent

,
“

fro m the t ran s m is s io n o f the imm ed iat e parent Th is being so .

,

s elec ti o n accor d i n
g to 7 uc i lla ti ng
t
s ta n d a r d s is n o s elec ti on a t a ll,

and he who ret u rn s fro m ea ch s t a t e fa ir o r expo s it io n with n ew

rath er than i mprov ed s tandard s canno t h o pe to m eet the h igh es t
s u cc es s a s a breed er o r c o ntr ib u t e rea l exc ellenc e to t he breed

he ha s ch o s en .

SECT I ON II —HISTOR I CA I . K N OWL ED G E OF T HE BR EED

N EC ESSA R Y
T h is is a lm o s t
f ev id ent and yet the n u m ber o f breed ers
s el -
,

who d o n o t po s s es s it and the read in es s with wh ich large m o n ey

,

is inv es t ed and p lan s m ad e wh ich will req u ire a lifet im e t o carry

out — a ll with t he mo s t m ea ger kn o wledge o f t he breed th a t

—
,

is ch o sen s h o w ho w light ly th is m a t t er res t s in t he m ind s o f

,

m a ny o th erwis e int elligent and ca u t io u s b u s in es s m en .

T h es e m en pro c eed as if t he m at eria l t hey were to wo rk wit h

were n ew m a t eria l ready t o b e m o ld ed fo r t he fi rs t t ime int o
,
5 8 0 PR A CT I C A L PR OB L EMS
any d es ired fo rm wh ile i n t ru t h it i v e rv ld m a t eria l with s o

wh ich m a ny m en h a v e wo rk ed befo re — s o m et im es fo r pro fit

, ,

, ,

o ften fo r a m u s em ent m e rely .

A n d t he breed has inh erit ed t he res u lt s o f a ll th es e ex per i

m ent s bo th had and g oo d s o th a t t h is m a t eria l is in so m e
, ,

res pect s t he bett e r and in s o m e t he wo rs e fo r wh a t o thers hav e

tr ied t o d o wi th i t T he m o s t o rd ina ry b u s in es s s en s e a n d t he
.

c o m m o n es t b io lo gic a l pri nc ip les ind ic a t e t hat befo re t he breed er

b egi n s s erio u s o perat io n s he s h o u ld kn o w a ll that can b e kn o wn
o f t he breed o r t he v ariety he pro po s es t o wo rk wi th In no .

o t her wa can h e m ak e int elligent s elect io n

y .

A few c ru d e ex a m p les will s u ffi c e t o illu s trat e M any breed ers .

o f E ng lis h s t ra in s o f catt le will n o t o n ly d es t ro y a wh it e ca lf ,

b u t will c o n s id er it s appea ranc e an ev id enc e o f i m p u ri ty o f b loo d ,

n o t kn o wing that th es e breed s in g en era l h a v e d es c end ed fr o m

t he wild wh it e catt le o f Great B r it a in I t is o n ly rec ent ly that.

wh it e has b een res t o red to fa v o r as a go o d S ho rt ho m c o lo r .

T he B erk s h ire s win e a re t he res u lt o f a cro s s o f t he large

E ng lis h ho g with t he s ma ll th in ha i red p lu m c o lo red N ea po litan
,
-
,
-
,

and m o re tha n o n e B erk s h ire h erd ha s been r u in ed by s elect ing

fo r breed i n g p u rpo s es t he p lu m p q u ick m a t u ring fi n e h aired
,
-
,
-
,

and m s t attract iv e p igs

o .

A fa m o u s breed er o f K ent u cky was rem arkab ly s u cc es s f u l

with S h o rth o rn s yet in his effo rt s t o s ec u re a h igh h ead an d a lo w
,

bris k et he fo rgo t t he nat u ra l wild type and s peed ily his breed ing ,

c a me to b e kno wn by it s s lo p in g r u m p and s p lit q u art ers .

Th e b reed er sh ou l d k n ow t h e pecu liar i ti es of hi s b l ood . Th e

firs t in fo r m a t io n n eed ed by t he pro s pect iv e breed er is a goo d
kn o wledge o f t he inh erent f a u lts o f t he breed He n eed s t o .

k n o w fo r ex a m p le that t he Berk s h ires a re nat u ra lly d efic ient i n

, ,

t he b a rn s and t he Po land C h ina s in t he s h o u ld er s ; that t he

,
-

D u ro c J ers ey s a re u n ev en in typ e a n d t he C h es t er Wh it es a ,

b it c o ar s e in th e b o n e He n eed s t o rea lize that t he J ers ey is

.

s o m et im es extrem ely d elicat e ; th a t m a n y o f t he Ho ls t ein F rie -

s ian s a re ro u gh and t h a t t h e breed is preem in en t ly s h o rt ta iled

‘

-
, ,

— henc e t he pro v is io n in t he s ca le o f po int s th a t t he bo ne o f

t he t a il s ho u ld rea ch t he h o ck which wa s b u t rarely t he cas e in
,

t he f o u ndat io n s t o ck o f the breed .

 8 PR A C I I C A I
’

PR OB LE MS
'

5 2 .

aft er birth with its attendant lo s s es I n all th eo ry we wou ld

,
.

prev ent t he b irth o f u npro fit a b le ind iv id u a ls and we s u cc eed ,

n early in pro po rt io n a s we a re s k illf u l in s elect io n .

Sel ecti on res u l t s in ab sol u t e in c rease in q u al i t y , n ot m erel y in

an el ev ati on of t h e av erage b y el imin at in g t h e l ower v al u e
s .

We ha v e been t o ld that ct io n res u lt s o nly in ra is ing the

s ele

a v erage by c u tt ing o ff the lo we r v a lu es b u t that t he u pper ,

v a lu es a re n o t i n flu enc ed th er eby T h is is C learly an e rro r as .

,

will b e seen by a refe renc e t o any sy s tema t ic breed ing experi

m ent s and es pec ia lly to the tab les giv ing t he res u lt s o f s elect ing
co rn fo r h igh o r lo w pro t ein o r h igh o r lo w o il Here it is s een
1
.

that in t he pro gress o f select io n by t he u s e o f s u cc es s i v ely

, ,

i nc r ea s ing s ta nd a r d s n ew a n d mg/1 7 v a l u es c on s ta n tly app ea r ed

,
r . .

N o t o n ly tha t b u t t he pri nc ip le is s t ill o p era t iv e aft er ten y ears

,

o f s elect io n and the c oe fi,

c i en t of v a r ia b i li ty is no t i n m os t c as es , ,

n gen era l i t m a s a id that t he res u lt o f

2
g r o wi n
g less I .
y b e

s y s t em at ic s elect io n is to S h ift t he type b u t n o t great ly to

red u c e v ariab ility and wh en app lied t o a n u m ber o f ch a racters

,

at t he s am e t ime it v ery c lea rly and v ery rapid ly d efin es the

type o f t he s tra in o r breed .

I n breed ing t he beet fo r s u gar t he c o w fo r m ilk t he h o rse , ,

fo r s peed o r any a n imal o r p la nt fo r any d efi n it e q u a lity th ere

, ,

is ev ery r eas o n fo r b eliev ing that we ha v e s u cc eed ed in p ro

d u c in g a h igh er o rd er o f exc ellenc e than ev er a ro s e s po ntan eo u s ly
in t he rac e wh ile in a s tat e o f n a t u re ; th a t is to s ay we ha v e ,

d o n e m o re tha n to ra is e the a v erag e we hav e elev at ed t he ,

Th e u pper l i mit s th is inc rea se o f

of i m prov em en t . M an ifes t ly
q u ality cann o t go o n in d efi n itely W e cann o t breed t he h o rs e .

to b e a s large a s t he elephant ; o r if we c o u ld th ere will b e an , ,

u pp er limit s o m ewh e re What will set th ese lim it s is an int eres t

.

in g q u es t io n I n s o m e ca s es n o d o u bt t he lim it wo u ld b e fix ed
.
, ,

by p u rely mech a n ic a l pri nc ip les in o th ers by phy s io lo gical

3
,
.

res trict io ns s u ch as the S ize o f the h ea rt and t he labo r o f

,

4 96
1 S ee pa ges 494 and .

11 Ib i d .

3 Fo r ex a m p e, t l h ere is a m ec h an ic a l l im it to t he len g t h o f leg, or t o th e s ize

of u dd er .
 S EL ECT I ON 8
5 3

c irc u lat ing the b lood ; b u t apparent ly we ha v e n o t yet in any ,

lin e appro ach ed a lim it s o h igh that v a riat io n is n o t ab u ndant ly

,

ab le t o pres ent s t ill h igh er v alu es Ho w lo ng th is m ay go o n is .

a q u es t io n bo th o f s c ient ific and o f u t ilitarian int eres t b u t it will ,

b e rem em b ered that v ariat io n is s u ppo s ed n o t to b e red u c ib le

belo w so me 8 5 o r 8 9 per c ent o f it s o riginal a m o u nt 1
.

Sel ec t i on f or d efi ni t e pur poses oft en again st v al u ab l e q u ali ties ,

es peciall y f ertility , 2
v i gor , an d po n l on gev ity . S o int ent a re we u

s ec u ri ng s o m e c o v et ed charact er a s ear ly m at u r i ty s ize m ilk i ng , , ,

o r feed i ng q u ality that we o v e r loo k o th e r les s v is ib le b u t n o ne

,

the les s es sent ia l q u a lit ies T h is is bes t s een a m o ng o u r m eat

.

pro d u c ing an imals Fo r exa m p le i t is t he h eav y fles hed early

’

.
,
-
,

m at u ring so w p ig that find s her way t o t he p rize r ing and u lt i

m a t ely t o t he fa s h io nab le breed ing pen N o w th is is n o t t he .

mo s t pro lific type o f s win e and u nd er t his po licy o f s elect io n , ,

prim arily fo r fl es h and fat it is n o t s u rpris ing that o f all o u r , ,

an im a ls th o s e bred fo r m eat prod u ct io n are lo wes t in fert ility

,
.

We kn o w o f n o fu nda m ental rea so n why it m u s t b e s o ; it S im p ly

is so b ecau s e fert ility ha s b een so gen era lly n eg lect ed in t he
exc lu s iv e s tandard s and m eth o d s o f s elect io n em p lo y ed .

Fert ility “
v igo r and lo ngev ity a re a ll rela t iv e t erm s
“
”
.
, ,

A ll an ima ls and p lant s ha v e s o m e d egree o f v igo r and n early ,

a ll a re ab le to repro d u c e a t lea s t t o s o m e ext ent T he ev ils o n

,
.

th is s c o re aris e no t fro m t he n o n breed er o r t he ind iv id u al that -

,

s u cc u m b s in ear ly life b u t fro m th o s e ind iv id u a ls wh ich th o u gh

, ,

n o t en t ir ely want ing a re et d efic ient in th o s e f u nda menta l

y ,

q u a lit ies that a re o f n ec es s ity c o rrelat ed with pro pagat io n o f a

v igo ro u s la s t ing and pro s pe ro u s rac e
, ,
I t is t he s hy breed er .

that c o m es to n o th ing and that is the ro o t o f m any o f t he ev ils

,

o f t he breed ing h erd .

T he relat iv e v a lu es o f pro lific and o f s hy breed er s may b e

bro u ght o u t by c o m pa ring three c o ws fo r exam p le o n e o f wh ich , ,

will prod u c e t wo ca lv es bef o re s he s t o p s breed ing an o th er fo u r , ,

and an o th er s ix A ft er fiv e generat io n s the f er tile f ema le

.

d es c endant s o f each would b e as fo llows ass u m ing that o n e ha lf ,

1 Pea rs o n , G ra m m a r of S c ien c e, p 48 3 . .

2 The wo r d “
l y
fert i it d
is u s e in p referen c e to “
d y
fec u n it b ec a u s e t he la t t er
t erm refers es p ec ia ll y f
to em a es l .
 8 PR ACTI CA L PR OB L EM S
5 4

t he calv es are ma les and on e ha lf fema les and that a ll d esc en d a n ts ,

a r e p r olific in tb c s a me p r op or tion as t he o rig ina ls .

N U M BER OF LI V ING AND PR OD U C IN G FE M A LES AT T II E EN D OF

V AR IOU S G EN E R AT IONS , F R OM Cows OF D I FFE R ENT

D EG R E ES OF FER T ILIT Y

Co w N o .

Firs t Sec o n d Third Fo u rth Fifth

G en era tio n G en era tio n G en era ti o n G en era tio n G en erat io n

I t is a y t o s ee that n o m att er what t he ind iv id u al exc ellenc e

e s

of c ow N o I and her d es c endant s th ey c o u ld n ev er b u ild u p a

.
,

h erd T h eir rat e o f reprod u ct io n is so lo w as on ly to k eep goo d

.

t he o r igina l n u m b er C aref u l s earch will d is c o v er a s u rpris ing

.

n u m ber o f fem ales o f th is C lass in t he herd s o f o therwise s u c

c es s fu l s t ock m en — u s eles s fro m any s ta ndp oi nt exc ept t he
,

S h o w ring .

On the o th er h a nd c o w N o 2 and her d es c endant s prod u c e

,
.

at a rat e that will n o t o n ly k eep th eir n u m bers goo d b u t will

ad mit o f s elect io n and th is is t he ca se to a great er ext ent with
,

N o 3 wh o s e d es c endant s in t he fifth gen erat io n wo u ld b e no

.
,

fewer than 2 4 3 a s c o m pared with 3 2 fo r N o 2 and 1 fo r N o 1 . . .

I t is ea s y to s ee th a t o n e s u ch c o w a s N o 3 in a h erd o f 2 0 .

lik e N o I wo u ld in a few y ear s by v ery breed ing p o wers

.
, ,

d om inat e the h erd a t the s a m e t ime a ffo rd ing gen erou s n u m

,

bers fo r select io n wh erea s t he d es c endant s o f N o 1 wou ld

, .

a ffo rd n o o ppo rt u n ity what ev er fo r select io n I t is c lear to t he .

mo s t ca s u al s t u d ent that w/t en ou r s ta n d a r d s a r e d ec id ed ly

ag a i ns t tb e l u
g/t es t
f er til i ty t/zey a r e d a ng er ou s , if n ot f a ta l, to
Me ra c e.

Need of comparat iv ely l a rge n u mb er s in b reedi n g operati ons .

Ob v io u s ly c o m pa rat iv ely large n u m bers a re n ec es sary in o rd er to

,

p ro v id e s elect io n with mat eria l s u fli c ien t fo r s ec u rin g u n if o rmity

 8 6 PRA C T I C A L PR OB L EM S
5

ha v e c o nt rib u t ed m os t to th eir breed s ha v e o ft en been in c on

s p ic u o u s a s ind iv id u a ls and u nf o rt u na t ely o ft en ha v e been d ea d , ,

lo ng b ef o re th eir real s erv ic e to t he breed was kn own and

rec o gn ized .

Need of t h e act u al b reedin g t es t S o v alu ab le is t he exc ep .

t io n a l breed er a n d s o i m po s s ib le is it t o kn o w him ( o r her) in

,

ad v a nc e by t he o rd ina ry m eth od s o f j u dg ment that o n ly t he ,

act u al breed in g t es t is reliab le T he o n ly s afe m eth o d is t o .

s elect t he h erd o f f em a les o f h igh f ert ility a n d u n ifo r m ly exc el

lent breed ing rec o rd and th en kno wing t he f em a le S id e by lo ng

, ,

and int im at e exp er ienc e s elec t the s i r e f or leis perf or ma nc e

1
,

i n g etti ng y ou ng
rd .

T he t es t s S h o u ld firs t b e m ad e with a few well -

kn o wn and ,

th erefo re fa ir ly aged f em ales ,

. Too m u c/z ca n n ot be s a i d a a i ns t
g
t/ze rac tic e of p u t ti n n ew s i r e a t onc e i n t o f u ll s er v ic e
p g a y ou ng
in tb c ner d , no ma tter w/za t lt is i n di v id ua li ty or h is p ed ig r ee .

Ho wev er prom is ing he m u s t b e s u bj ect ed to , t he act u a l t es t ,

and aft er ha v ing pro v ed his breed ing po wers with kno wn fem ales
lt e s /t ou ld b e u s ed to t/ze u t mos t a s long as b e w i ll b r eed s u cc es s

f l ul
y and n,o t d is card ed b eca u s e o f lo s s o f b lo o m d ec li n e i n ,

fo rm o r ev en fo r t he acq u irem ent o f a n ev il d is po s it io n I t is

, .

fro m t he pro v ed pat riarch s and fro m t he grand mo thers o f the

h erd that rea l exc ellenc e will c o m e and t he rea l v alu e o f pro v ed ,

breed ers ma le o r fema le is beyo nd c o m p u tat io n

,
2
,
.

In stal l t h e su ccessor early I t is n ev er t o o ear ly to s eek a n ew .

h ead to a n es tab lis h ed h erd Prov ed S ires a re s eld o m fo r s ale .

,

and the o n ly rec o u rs e fo r the breed er is t o pro v e his o wn ;

ind eed what he n eed s is a S ire that will pro d u c e well wi th Iris
,

tak es m u ch t im e and o ft en m any trials to find a wo rthy

It
s u cc es s o r t o t h e h ead o f t he h erd Pu tt ing it o ff t o o lo ng and .
,

a feeling o f fanc ied s ec u r ity a re t he t wo cau s es o f lea v ing a ,

1 A l
b ree in g rec o r
c o m p et e d d h ld s ou be k ep t o f eac h fe m a l e s ep a ra t el y . S ee
C h a p ter o n A n ima B ree in g
"
l d .

h
3 T h e a u t o r h as a m a s s o f d a ta c o ll ec t e df ro m h d d fb d f m
un re s o ree ers , ro

w ich h it a pp ea rs t h ty a oung bu ll s a re c o m mo n ly p refe rre db th y h p

ec a u s e e a re c ea er,

b ec a u s e t h ei r p i d
er o f o s erv ic e is l o n g er , a n d b ec a u s e th y
e ma re m g b l o re an a ea e.

It a p p ea rs to o t h t wh
a en a t es t is m a d e it is c o m mo n ly t p ld d k w
no u on o an no n

c o ws , but u po n h if
e e rs .
 SELECT I ON 8
5 7

herd with o u t a h ead and ,

of t he e nfo rc ed ev il pract ic e o f usi ng
an u nt ried s i re .

Comparati v e v al u e of mal e an d f emal e . I n t he m att er of p re

p o t ency as we hav e already s een neith er parent has any part ie

, ,

u lar ad v a ntag e o v er the o th e r B u t th is refers t o a s ingle .

o ff s pr ing a n d is o n ly a part o f t he q u es t io n
,
T he rea l d iffer .

enc e is o n e o f n u m be r s A m o ng an im a ls the s i re m ay pro d u c e

.

perhap s a h u ndred in a s eas o n wh ile t he d a m is lim it ed to on e ,

ind iv id u a l o r at m o s t ( a m o ng h o g s ) to t wo litt er s T he tro tt ing .

rec o rd s S h o w that c ertain S ires pro d u c ed lit era lly h u ndred s o f off
s p ring in t he lis t b u t t he gr eat es t rec o rd e v er m ad e by any m are
,

was that o f Green Mo u ntai n M a id who pro d u c ed n ine liv i ng fo a ls , .

F o r p u rely m ath em at ica l reas o n s therefo r e t he fem a le is o f , ,

v a s t ly les s c o n s eq u enc e in h erd o r breed i m pro v ement ind eed , ,

wh erev er po lyga m o u s m at ing o cc u rs I t is h ere a q u es t io n o f .

n u m bers and o ppo rt u n ity A s regard s th ese t he u pper limit o f.

,

t he m a le is v ery h igh and o f t he fem a le v e ry lo w wh ich fact ,

t eaches t he nec es sity o f extrem e care in the select io n o f t he

s i re n o t so m u ch fo r b io lo gi ca l as fo r n u merica l rea so n s
,
T he .

S i ng le fem a le i s th erefo re compa ra ti v ely i ns ign ifica n t U n less

, ,
.

S he b e o n e o f t he few ph en o m ena l br eed ers her i nd iv id u a l p o wer

fo r goo d is exc eed ing ly low and t he read in es s o f m any b u yers ,

t o pay extrem e pric es fo r f em ales es pec ia lly o f catt le is wh o lly , ,

u nacc o u ntab le .

Th e sire mor e t han h al f t h e h erd I t has bec o m e a pro v erb .

that t he s ire is half t he h erd He is fa r m o re than that He . .

is h alf o f t he fi rs t gen erat io n three q u arters o f t he n ext s ev en , ,

eighth s o f t he th ird and s o o n u nt il if ju d ic io u s s elec ti on b e ma i n

, ,

ta in ed f or a
f e w g en era tions , t/t e d i a m eter of tb c lt er d wi ll b e
fi x ed by Me s i r e b ei ng tr
a lon e
u e t he.f o llyT h is
o f m a int a in ing ,

a S ire with b u t t wo o r three h igh c las s fem ales is ev id ent ; he -

s h o u ld ha v e larger o pp o rt u n ity A ll th is mean s th a t as a beg in .

,

n ing n u m bers a re o f mo re c o n seq u enc e relat iv ely than q u al ity

,

o n the S id e o f t he d a m and that if t he breed er m u s t ch oo s e

,

bet ween the t wo it is bett er to p u t a g iv en a m o u nt o f mo ney

int o a goo d n u m ber o f p la in fem a les than int o a s ma ller n u m ber
o f h igh q u a lity b u t that in a ll ca s es t he S ire S h o u ld ha v e q u a lity
,

a n d p lenty o f i t b eca u s e o f t he p r inc ip le h ere s tat ed

, .
 8 8 PR A C TI CA L PR OB LEM S
5

S ize i n t h e da m ; q u a li ty in t h e sir e I n m any lin es o f breed .

in g , s iz e i n t he S i re is c o n s id ered by m any breed ers a s o f firs t

im po rtanc e . T h is is
again s t reas o n a n d b io logic a l pri nc ip les .

W e n eed i n t he s ire all t he d es i ra b le cha ract ers p o s s ib le and ,

th ese a re m o s t rea d ily fo u nd in an im als o f med i u m no t extreme , ,

s ize I t is c o m para t iv ely ea s y t o get S ize a lo n e a n d th is can b e

.
,

go tt en o n the s id e o f t he d a m T he herd m u s t d epend fo r u n i .

fo rm ity la rgely u po n the S ire and he s ho u ld b e freed a s m u c h ,

as po s s ib le fro m t he req u irem ent o f s ize

1
.

Natu r al sel ecti on al ways a t wor k N at u ra l s elect io n is a lway s .

at wo rk in fi eld and fl ock and h erd Of th is we m ay b e well .

as s u red N o m att er wh a t we d es ire to acc o m plis h o u r s u cc es s

.
,

o r o u r fa ilu re will t u rn in t he las t ana ly s is u po n the fitn es s of

, ,

t he pro d u ct t o liv e a n d t o repro d u c e a m id t he c o nd i t io n s by

wh ic h i t m u s t b e s u rro u nd ed .

So me o f t he b es t th ing s a m o n g b o th plants and an im als are

weak o r c o m para t i v ely u n pro li fic N at u ral s elect io n is d ec id ed ly .

again s t th eir s u rv iv a l no m att er ho w v alu able they m ay b e t o

,

us . L ack o f c o n s t it u t io n o r v igo r is ea s ily seen b u t lack o f ,

breed ing po wers is n o t s o ea s ily d et ect ed and h ere is wh ere ,

t he great es t a m o u nt o f tro u b le a ris es .

We hav e a lready s een that in nat u re t he po p u latio n t ha t is

b or in to Me wor ld is p r opo r ti on ed a cc or d i ng to r ela ti v e f er til ity ,
n

wh ile t he p o p u la t io n that is pe rm itt ed t o r ema i n is c o nd it io ned

u po n rela t iv e po wers t o res is t ad v erse c o nd it ion s and to fi t int o
t he c o nd it io n s o f life .

T h is s a m e relat io n o bta in s in o u r h erd s with th is d iff erenc e , ,

that we with o u r s elect io n d ec id e arb it ra rily what s hall liv e

T hat is right and acc o rd ing t o ec o n o m ic n ec es s ity — o n ly in
.
, ,

d o ing s o we m u s t n o t a s s u m e th a t a ll ind iv id u als an d types a re

eq u a lly fert ile and eq u a lly ab le t o pro paga t e th em s elv es .

I t m ay b e in c erta in in s tanc es that i n o rd er to s ec u re what

we d es ire we s ha ll o f n ec es s ity proc eed t em p o rar ily with so me

1 A m a n ifes t e xc ep t io n to t hi
s gen era l p ri n c ip e l is in t h e b ree in g o f raft d d
h o rs es fro m fa rm m a res . Here s ize is an job ec t io n o n t he d
a m s s i e, b es i es
’
d d
b ein g d iffi l t t
cu o
g e t . W e ig h t is th e l if
c z e d ides er a t u m j ust now ( it wi n o t ll a lw ya s

be so ) ni d f t
ra h -
o rs e d
b ree in g , an d u n d er p r es en t c irc u m s t a n c es it m us t b e so u g h t
es p ec a i ll y i t h
n e S i re .
590 PR A C T IC A L PR O BL E M S

In tru th we ha v e litt le exact info rm at io n o n wh ic h t o rely b u t ,

t he wr it e r s er io u s ly q u es t io n s t he c o rrectn es s o f th is c o nc lu s io n

f r o m t/t e s ta n d o i n t o tb e oflirp r i u
p f g T h a t br ee d ing at an. i m m a
t u re age Ch eck s t he gro wth o f fem a les is n ext t o c ertain b u t it is ,

a lso tru e that t he h eif er will m ak e a b ett er m ilk e r and a m o re c er

ta in breed er if bred b efo re m at u r ity and befo re f u nct io n s o th er

th a n m ilk pro d u ct io n ha v e b ec o me t he prev a iling hab it o f life .

T hat the pro geny o f imm at u re an im a ls is n ec es sarily fa u lty

is d o u btf u l I n nat u re ev ery wh ere r ep r od u c t ion b eg i ns bef or e
.

ma tu r i ty and in m a n at lea s t it ha s been s h o wn that t he lengt h

,

o f lif e o f fi r s t ch i ldr en is o n t he a v erag e f o u r y ears m o re than

, ,

th a t o f t he lat es t bo rn .

C o n s id erat io n s already ad v anc ed in c o nn ect io n with t es t ing

breed ers n ec es s itat e t he breed in g at a c o m parat iv ely ad v anc ed
age and a ll th ing s p o int t o t he c o nc lu s io n that i n pract ic e breed
,

in g m a y begin early and c o nt in u e as lo ng a s p o s s ib le M erely t o .

ga in t i me if fo r n o o th er rea s o n ea rly breed ing is to b e ad v o cat ed

, ,
.

Bl emish es an d accid en ta l in j u r i es N o t with s t a nd ing p o p u lar

.

o p in io n t he breed ing an im a l is n o n e the wo r s e fo r acc id ent al

,

inj u ry ; that is s o fa r as his o r h er b r eed i ng v a lu e is c o nc e rn ed

, .

The q u es t io n is n o t w/zet/t c r t/ze ma r e is sp a v i n ed b u t wh a t k ind ,

o f a h o ck had s he na tu r a lly a n d had S he s u ffi c ient occa s ion to ,

b e s pa v in ed I t is ea s y t o m ak e a bad s h o win g and to say b itt er

.

th ing s abo u t the p ract ic e o f breed ing inj u red an im a ls b u t t he ,

ev id enc e o n inh eri t a nc e a ll s h o ws that inj u ries a s s u ch a re n o t

tran s m itt ed T h is s h o u ld n o t free t he m ind fro m t he o bligat io n

.

t o j u dge acc u rat ely a s t o wh eth er t he p a rt wa s n a t u r a lly perfect

o r na tu r a lly d ef ect i v e .

Dimcu lt ies in sel ec ti on i n crea se r apidl y wi t h t he n u mb er of

p o in t s o n w hi c h s el ect i on i s t o b e b as ed T h is p u re ly m ath em at ic
. al
c o n s id erat io n s eem s n o t to arr es t the att ent io n o f breed ers as it
s h o u ld . If we s elect fo r o n e p o int o n ly we get ah ead ra p id ly .

T hat has b een t he a d v antage o f t he tro tt er S p eed was t he o n ly .

req u i rem ent and wh ile it in v o lv ed m a n y s u bo rd inate c o nd it io n s

, ,

s u ch a s a p erf ect bo dy v ig o r e nd u r a nc e m enta l c o u ra ge and

, , , ,

d et er m inat io n yet n o o th er req u irem ent ha s been add ed C o lo r

,
.
,

s ize ,
s t le
y a ct i on
,
c o nf o r m a t io
,
n — a ll ha v e b een d is regard ed
,

fo r t he o n e o bj ect s p eed , .
 S E LEC T I O N 59 1

Ov er again s t th is the D u tch B elt ed C att le fo r exa m p le hav e , ,

an ab s o lu t e c o lo r req u i rem ent E v ery c o w m u s t firs t hav e a .

wh it e b elt aro u nd her bod y T h is c ert ai n ly has n o th ing to d o .

with her m ilk ing abilit ies yet t he ab s u rd s pec i ficat io n has go n e
,

into t he v ery na m e o f t he breed — a fact that will k eep t he ,

breed m at eria lly beh ind it s c om pet it o rs in m att ers fo r wh ich we

breed c ows 1
.

The m o re c learly t o S h ow t he ext ent o f t he ha n d ic ap of '

s tr iv ing aft er m any p o int s in s elect io n let t he s t u d ent wo rk it ,

o u t m ath em at ica lly I f b u t o ne po int is req u ired and it can b e

.
,

sa t is fi ed s a in o n e t enth o f t he ind iv id u a ls th e n t h e chanc es

y , ,

o f gett i ng i t a re o n e in t en and o n e t enth o f the breed is ,

a v ailab le .

N o w t o th is let u s add a s ec o nd req u i rem ent that can b e

fo u nd in b u t o n e th i rd o f t he ind iv id u als The pro bab ility o f .

find ing th ese two po int s in t he sa me ind iv id u a l will th en bec o me

no t 1
1 6
or
} b ut 1
36 x
a
g,
o r 1
35
and o n ly ab o u t three an im a ls
, ,

in o n e h u ndred will m eet t he req u irem ent s .

Need of r edu cin g th e r eq u iremen t s t o t h e u t il i ty b a sis . In

fa s h io nable an ima l breed ing we ha v e so m u lt ip lied o u r po int s
that we a re n o lo nger ab le to find any v ery large pro po rt io n o f
th em in any o n e ind iv id u a l and we are o ft en o b liged to t o lerat e ,

po s it iv e ev ils in o rd er to get t he req u irement s ev en with in t he

lim it s o f t he h erd T h is is v irt u ally m ix ed breed ing
. .

What is n ed ed i a ret u rn t o fir s t p rin c p e

e s i l S — t o s elect a ,

v ery li m i ted n u m b er o f t he po int s mo s t im p o rtant fro m t he

u t ility s t a ndpo int L et th es e he so few and s o pro n o u nc ed that

.

they m ay a ll b e fo u nd in ev ery ind iv id u al o f t he breed ing h erd .

T h en la ter a s n u mb er s m u lt ip ly o th er po int s can b e add ed a

, , ,

few at a t ime u p o n a pract ica lly p u re anc es try s o f a r a s p r ev iou s

,

p o i n ts a r e c on c ern ed “
T h is o n e th i ng I d o
. s h o u ld b e w o rn i n

t he hat o f ev ery breed er A litt le c o u rage h ere wo u ld s o o n

.

wo rk wo nd ers b u t po int s hav e bec o m e so m u lt ip lied in s o m e

”

o f o u r breed s th a t a ll po s s ib ility o f find ing any ind iv id u a ls that

po sses s th em a ll has lo ng a go been pass ed leav ing u s in a ,

1 T he writ er h es ita t es t o u s e as fo rc efu l la n gu a ge a s t h e ab o v e rega r di gn an

y
b ree d , l
b ec a u s e in gen era a ll br eed s a re go o , b u t t is is a s tep s o c ea r d h l ly d
a v ers e

to l k h l
iv e s t o c in teres ts t a t n o a n g u a ge is t o o s t ro n g in c o n em n a t io n
-
d .
 PR A C TI C A L PR O B L EM S
59 2

welln igh h o peles s j u m b le with ped igrees m ean ing next to n o th ing ,

s o far a s d efi n i t e inf o rm at io n goes .

Im port an c e of pedi gree E n o u gh has b een s h o wn t o po int .

c lear ly to the fact that the s im ply goo d ind iv id u al is wo rth “

les s a s a breed er 1
He m u s t b e t he pro d u ct o f a goo d anc es try
.
,

and m o reo v er o f t he n g/zt kin d o f go o d anc es t ry I t is n o t .

en o u gh that t he an im a l o r p lant is n o t m ix ed in i t s b loo d We .

o u ght t o kn o w and o u r p ed igrees o u ght t o s h o w what wer e t he

, ,

s p ec ia l cha ra ct er s o f t he anc es t o r s T h ere is yet s o m u ch .

v a riab ility in a ll o u r breed s that a s i m p le g u ara n ty o f n o n

i nf u s io n o f o u t s id e b loo d is no t en o u gh S o m eth ing po s it iv e is .

n eed ed and great s u cc es s a wait s t he breed wh os e breed ers will

,

t ak e a few po int s at a t im e and es tab lis h a d o u b le regi s try o n e o f ,

wh ich S h al l b e a rec o rd o f til e d eg r ee in w/t ic /t tlt e in d i v id ua l

a c tu a lly p oss ess ed t/t e d o m i na n t c lea ra c t er s of tlze b r eed I f t h e .

“
ad v anc ed regis try o f so me o f t he dai ry breed s can b e s afe
”

gu ard ed again s t ab u s es and th en b e u s ed as a bas is fo r select io n

, ,

it will b e o f u nt o ld benefi t to t he breed s and t o t he c o u ntry at

larg e .

S EC T I ON I V R AT I O N A L S E LECT I O N

Wh en and to wh a t extent t o d epart fro m s a fe general p rin

c i ples o n acc o u n t o f ec o n o mic o r o th er c o n s id era t io n s is a m att er

calling fo r the m os t d is cri m ina t ing j u d gm ent .

Fan cy poin t s I t is perf ect ly ea s y t o s h o w that if t he breed er

.

su cc eed s in fix ing r ea lly u s ef u l charact ers he will hav e his hand s

m o re than f u ll and yet d es p i t e a ll th is fa s h io n c o n s tant ly s et s

, ,

c erta in fancy po int s and in s is t s u po n th eir o bs er v anc e T he

, .

t ro u b le is n o t o n ly that m o s t o f th es e fancy point s ha v e litt le o r

'

n o u t ility b u t a lso that lik e any o th er c a pr ic e o f fas h io n th ey

, , ,

a re li k ely t o change freq u ent ly and with o u t warn ing wh ereas ,

a ll c o n s id era t io n s o f s elec t io n r eq u ire c o n s tancy and S im p li c i ty .

W hat th en s hall t he breed er d o ? He is b ent u p o n b u ild ing

, ,

u p a h erd o f the h igh es t pra ct ic a l v a lu e and he has caref u lly ,

weigh ed t he relat iv e v a lu e o f a ll u t ilita rian chara ct er s A ll o f a .

1 T he reg ress io n t ab e c l l ea r ly h s o ws t h at an d d lf
in fe rio r i n iv i ua ro m a
go o d
y
a n c es t r is in e v er y wa y s u p e rio r to a p e rf e ct in d i id l f m h t
v ua ro a e ero gen eo u s

t y
a n c es r . Bo t h a re ev i l s, b ut of t he t wo t h e l a t te r is b y fa r t he wo rs e .
5 94 PR A CT I C A L PR O B LEM S

h o ping t he c raze m ay pas s befo re the o ld s t o ck o f fem ales S hall

pas s away I f it d oes n o t th en at a ll hazard s s o me rem nant s
.
,

at leas t m u s t b e pres erv ed p u re and u nalloyed as a n u c leu s

aga in s t t he day wh en t he pend u lu m will s wing back t o t he
no rm a l o r perhap s to t he o ther extrem e
,
.

Br eeders f ad s T he ab o v e ha s referenc e to req u i rem ent s

’
.

im po sed by t he b u yer B u t t he fact is breed ers th em s elv es

.
,

hav e m u lt ip lied th eir n at u ral d ifli c u lties eno rm o u s ly and u se

les s ly by fad s o f th eir o wn i n v ent io n t he tyra nny o f wh ich is ,

ev en wo rs e than that ex erc is ed by t he a lien b u y er A gai n s t all .

th is t he s tro nges t pro t es t is far to o weak .

Why fo r exa m p le s h o u ld a few c u r ly ha irs o n t he back o f a

, ,

hog d is q u alify him a s a breed er ? Why a re c o ws and b u lls

s elect ed by t he S ize o r s hape o f t he es c u tch eo n ? Why mu s t
a J ersey hav e a b lack t o ngu e ? Why m u s t t he ta il bo n e o f a
Ho ls t ein F ries ian c o w reach to t he h ock Why d id t he S ho rt ho m
-

breed ers t wenty fi v e y ears a go caref u lly k ill ev ery roan o r wh it e

-

ca lf and s o y ield th em selv es t o t he c o lo r craze that fo r a d ecad e

or m
,

o re t he breed m ad e pro gres s back ward Why su ch frant ic

h o rro r at t he sev ent een s 1
req u iring that a boo k b e wri tten
,

black lis t ing lit erally th o u s a nd s o f the bes t animals of t he breed

A b s u rd s tandard s o f th is cha ract er S h o u ld b e res is t ed to t he
u t m o s t by ev ery rep u tab le breed er and by ev e ry rea l f riend o f

t he breed wh eth er a ris i ng ign o rant ly o r fro m a m a lic io u s d esi re

,

t o narro w t he range o f p o s s ib le s a les t o d is cred it t he ani m a ls ,

o f c o m pet i t o rs o r t o d es tro y th eir h erd s

,
T he wri t er is well .

aware that breed ers as a c la ss a re s ec o nd to n o ne either in ,

int elligenc e o r i n h o n o r He is a ware t o o that m any o f t hes e

.
, ,

foo lis h req u irement s o r o bj ect io n s lik e t he q u erl o n t he pig , ,

g e t s tart e d n o o n e kn o ws ho w and ga in s t rength by r epet it io n ;

,

b u t t he wh o les a le d es tr u ct io n o f rep u tat io n s and ev en o f h erd s

and f o rt u n es by t he wa r o n t he s ev ent een s is c o n v inc ing proo f
,
“

that ind iv id u a ls ev en in th is h o n o rab le c o mpany a re n o t abo v e

, ,

t he m o s t da s tard ly m eth o d s o f red u c ing c o m pe t it io n I nd iv id .

u a ls o f th is s o rt a re n o t bon a i d e breed er s th y re c o mm erc ial

f e a

p ira tes who u s e ped igreed liv e s t o ck as m at erial fo r s pec u lat io n .

T h ey ha v e a dd ed n o th ing t o t he exc ellenc e of any breed o r t o ,

1 R f i h
e e re n c em d t t h S h th m im p t t i
s e re a e of 8 7 e or o or a on o 1 1 .
 S ELECT I O N
5 95

the h o n o r of breed ers who as a c las s a re the so u l of h o no r and

, , , ,

wo u ld n o m o re f als ify a p ed igr ee than th ey wo u ld ro b a bank .

S u c h metho d s m u s t b e m et and breed ers fad s gen era lly

’

s h o u ld rec eiv e t hei r ev er las t i ng q u iet u s wi th in t he c o nfin es o f

o u r breed ers as s oc iat io n s I f th is m ay b e then t he ind iv id u a l

'
.
,

will b e reas o nab ly s af e ; if n o t th en he m u s t t ak e his chanc es

,

with t he res t b u t aga in s t th is i n s id io u s enem y o f all goo d breed

,

in g h is v o ic e and his pen s h o u ld b e in s tant a nd act iv e th is in ,

t he int eres t n o t o n ly o f his o wn b u s i nes s b u t of t he breed he lo v es .

Fas hiona b l e pedigrees A ll agree an d t he law o f anc es tra l

.
,

h ered ity prov es that t he ances try back o f t he ind iv id u al is

,

extrem ely po t ent ; et th is p o t ency larg ely res id es in t he n ear b

y y
-

m em ber s and t o see a breed er p o ring o v er a ped igree ru n n i ng

,

t en o r t welv e gen erat io n s ba ck to an appro v ed ind iv id u a l o n

t he f em a le s id e and th en gra v ely n o dd ing appro bat io n o f a
,

goo d fo u ndat io n th is is ind eed bo th h u m o ro u s and pat het ic

, .

A cc o rd ing to t he law o f anc es t ral h ered ity as s tat ed by Ga lt o n

and f u lly n o t ed in a prev iou s chapt er each gen erat io n a n d each ,

in di v id u a l o f t he v ariou s generat io n s ha s an infl u enc e r epres ent ed

by t he fo llo wing fract io n s waiv ing a ll q u es t io n s of prepo t ency
,

R E LAT I V E I NT ENSITV OF BLOOD LI NES A ND A PPR o x I M AT ELV R ELA T I V E

I N FLU E NC E or D I FFERENT G E NERA TIONS AND I N D IV I D U A LS
FO R T EN G EN E R A T IO NS BA C KWA R D

IN FL U E N CE or G ENE RA F LU E EA
IN EN C or CH

T I ON —PE R C EN T IN D D U A L —P C E
IVI ER N T

T o ta l

1 h
T is wi ll b e 1 00 if c a rri e dt o in fin it y .
 6 PR A C T I C A L PR O BLEMS
59

By th is we s ee t ha t t he ind iv id u al inh erit s fro m n o les s than

2 04 6 ind iv id u als with in t en g en erat io n s o f anc es try and t ha t , ,

on tice a v erag e C haract e rs p o s s es s ed by a S ing le i nd i v id u al o f

,

t he t enth gen e rat io n back ha v e an infl u enc e a m o u nt ing t o n o t

o v er o n e t en th ou s andth o f o n e pe r c ent o f t he t o ta l h eritage ,

repres ent ing a pro babi ity f ab t o e in a m illio n c ertai n t o

l o o u n —
b e h ea rd fro m b u t o f litt le c o n s eq u enc e as a f ou n d a tion .

We m u s t remem ber that b es id es t he fo u nd a t io n th ere a re “ ”

1 0 2 3 otlzer anc es t o r s o f t he t enth g en era t io n and s o me 1 02 2 ,

int erv en ing anc es t o rs each m o re po werf u l by far than t he

,

s o called f o u ndat io n
-
S i x gen erat io n s back t he infl u enc e is b u t
.

1 5 per c ent fo r t he s i xty fo u r i nd iv id u a ls i n v o lv ed o r abo u t

.
-
,

1 o f o n e p er c ent fo r each T h is is an a m o u nt o f infl u enc e

46
.

wh ic h fo r pract ica l p u rp o s es m ay b e c o n s id ered a s a n egligible

q u ant ity and it is fo r th is rea so n that in m any lin es t he d ict u m
,

ha s go n e o u t The S ixth cross is p u re

“
th is m ean ing that
”

, ,

n early 99 per c en t is c o v ered by t he t op W e ha v e seen

“
.

already that th is agrees perfect ly with m ath em at ica l theo ry .

T he s o ca lled f o u n dat io n is th eref o re n o t a f o u ndat io n b u t

-
,

o n ly a b eg i nn ing and i t is t he top and n o t t he b o tt o m that giv es

, , ,

charact er t o the ped igree .

T h is is n o t int end ed t o d is pa rage p u rity o f p ed igree ev en to

the t enth gen era t io n and b ey o nd b u t it is int end ed a s a pro t es t
,

again s t t he b lind f o llo wing o f c ertain p ed igrees b eca u se o f the

f o u n dat io n .

N o r is it t o b e c o n s tr u ed a s a crit ic is m d irect ed agai ns t breed

in g a lo ng appro v ed lin es ; fa r fro m i t ; b u t it is a p lea fo r the
caref u l s t u dy and ra t io nal v a lu at io n o f ped igrees .

Wh a t d ec eiv es the breed er is t h e fa ct that t he sh o rt fo rm

ped igree as it is o ft en pres ent ed ru n s o n ly o n t hef ema le S id e so
, , ,

that o f the 2 04 6 a nc es t o rs o f t he fir s t t en gen era t io ns o nly eight

, ,

o r t en f em a le s a n d i s i s w —
th e r re o u ld a ppear t he o th er 2 0 2 6
n o t being n o t ed T hey ex is t h o wev er and th eir infl u enc e is t o
.
, ,

b e reck o n ed wit h Of c o u r s e i t is t ru e that i n C lo s e breed ing

.

t he s a m e ind iv id u a l app ear s m any t im es in a p ed igree and th us ,

his o r her in flu enc e is m u lt ip lied ; b u t t he po int h ere m ad e is

that a s ingle ind iv id u al ten o r ev en s ix gen erat io n s back c o u nts
fo r lit t le s o far a s it s per s o na l infl u enc e is c o nc ern ed .
 8 PR ACT I C A L PR OB L EM S
59

t he rec o rd s d o no t u nd ertak e to s u pply b u t wh ic h breed h is t o ries

,

S h o u ld a ffo rd fo r t he en light en m ent o f y o u ng br eed er s and t o

t he en d that u nfo rt u nat e c o m b inat io n s m ay n o t b e u n witt i ng ly

m ad e T he s t u d ent will reca ll t he d ifferenc e bet ween bro th ers
.
,

which s ho ws t he n eed fo r s elect io n ev en with in t he p ed igree .

Wh en a fa m ily bec o m es fa m o u s it is bred with les s ab ility and

care than bef o re wh ereas it is d eser v ing o f great er and m o re
,

caref u l att ent io n B u t c o nd it io n s a re aga ins t it t he ind iv id u als

.

ha v e a h igh c o m m erc ia l v a lu e and ev eryth in g goes at a s tro ng

,

pric e i nd ifferent and bad as well as go od A S lo ng as men

,
.

will pay fo r breed ing regard les s o f q u a lity breed ers are lik ely
, ,

t o s ell ev eryth ing t h at will find a b u y er A larg e n u m ber o f

.

breed er s hav e repo rt ed to t he wri t er that th ey s ell 1 00 per c ent

o f th ei r an im als fo r breed ing p u rpo s es U nd er c o nd it io n s s u ch
.

a s th es e all s elect io n is pr a ct ica lly aband o n ed and we s h o u ld n o t ,

exp ect lo ng er t o m a inta i n exc ellenc e T h u s it is that t he v ery

.

s u cc es s o f a p o p u lar s tra in is lik ely t o pro v e i t s u nd o ing .

T his s tat e o f a ffa ir s s u fli c ien t ly acc o u nt s fo r m o s t d is as t e rs that

ha v e o v ertak en fa s h io nable fa m ilies in t he pa s t and we a re n o t ,

et warra nt ed in ass u m ing th a t a fa v o ri t e s tra in is bo u nd q u ick ly

y
t o wear o u t o r o th erwis e c o m e t o a n en d m ak ing it n ec es sa r y ,

that we S h o u ld be fo rev er effect ing n ew c o m binat io n s I nd eed .

,

th ere is t he b es t o f gro u nd fo r t he c o nfid ent belief that if a

t ith e o f t he lab o r were bes t o wed u po n the pres er v at io n of a u se
fu l s tr a i n that wa s exp end ed t o o rigi nat e it we m ight ha v e i t wi th
u s i nd e fin it ely t o t he infin it e go o d o f t he breed and t he las t ing
,

s er v ic e o f m a n .I t is n o t a c o nt in u a lly rec u rring s tra in o f n ew

creat io n s that is m o s t need ed b u t rath er well pro t ect ed and
,
-

so lid ly bred lin es o f lo ng es tablis h ed exc ellenc e and u nq u es t io n ed

-

anc es try Here in t he Op in io n o f t he wr it er lies the fi eld fo r

.
, ,

f u t u re effo rt o f A m erican breed ers .

 CH A PT E R XV I I
SY STEM S OF BR E EDING

Of t he ar io u s p o s s ible sy s t em s o f breed ing s o m e are bett er

v

adapt ed to o n e p u rpo se o thers t o an o th er ; s o me again a re

,

pec u liarly adapt ed to an ima ls a n d o th ers to p lant s T he prac ti

,
.

c al breed er S h ou ld fi r s t o f all ha v e a c lear id ea o f what he is

try ing to d o and th en an acc u rat e kn o wledge o f t he v ario u s

,

s y s t em s that can b e em p lo y ed t o ach iev e h is p u rpo s e .

SEC TI O N I — PU R POS ES I N BR E E D I N G

T he gen eral pr inc ip le that s h o u ld d ec id e t he sy s t em t o b e

c hos en and adh ered to d epend s u po n t he an s wer to the fo llo wing
q u es t io n I s t he p u rpos e of t he breed ing t o im pro v e t he lt erd ,

that is the h o me s t o ck of the far m er ; is it to im pro v e t he b reed

,

or v a r i ety a s a wlt ole ; o r is it t o or ig i na te n ew v ariet ies

T he an s wer to th is q u es t io n S h o u ld d et er m in e t he sy s t em o f
breed ing to b e ad o pt ed T h es e p u rpo ses a re s epa rat e an d d is
.

t inct T he fi rs t is herd im pro v ement t he ackno wledged o bj ect

.
,

o f wh ich is t o b u ild u p t he h o m e s t o ck u nt il i t appro ach es in

exc ellenc e t h e appr o v ed br eed s o r s tra in s T h is is p u rely c o m

.

m erc ial and p u r ely s el fi s h i n t he b es t s en s e o f t he t er ms in that

, ,

the breed er is no t o pe rat ing fo r t he go o d o f anyb o dy o r anyth ing

b u t h im self and h is o wn and is n o t a iming to o u td o anybo dy
,

els e ; h is p u rp o s e is rath er t o s ec u r e fo r h im self t h e im pr o v e

, ,

m ent that o th er s ha v e o rigi nat ed I t is the ch eapes t and ea s ies t

.

o f all f o r m s o f breed ing and pr o d u ct iv e o f t he m o s t ra p id res u lt s .

T he sec o nd p u rp o se is o n t he o th er hand c hiefly t he i m pro v e

, ,

m ent o f a rec o gn ized breed o r v a riety — a h i m pro v em ent in

,

t end ed to end o w t he rac e m o re rich ly t han ev er b efo re T h is is .

t he v ery h igh es t s ty le o f fin is h ed breed ing and ca lls fo r t he mo s t

,

intelligent and expen s iv e meth o d s beca u s e in th is ca s e t he

,

breed er is a lead er n o t a fo llower and an im itat o r

,
.

5 99
600 PR A CT I C A L PR O BLEM S

T he th ird p u rpo se in breed ing is n o t t o im pro v e anyth ing ,

b u t to s ec u re s o m eth ing ent irely n ew d iff erent fro m and p re , ,

s u m ab ly bett er than any prev io u s ly ex is t ing s t ra in s

,
I n carry ing .

ou t th is p u rpo s e t he breed er proc eed s u po n t he a ss u m pt io n t h at

o u r v ariet ies a re t oo few that gap s ex is t which m ay b e fi lled u p ;

and that it is bett er t o pro d u c e so m eth ing n ew than to d e
pend u po n im pro v ing the o ld T h is is the m os t a m b it iou s o f all .

fo rm s o f breed ing and appea ls t o creat iv e gen iu s rath er th a n

,

t o c o n s er v at iv e b u s in es s in s t inct s wh ich inc lin e t o im pro v e ,

m ent o f ex is t ing ra c es ra t her t ha n t o t he prod u ct io n o f n ew o n es .

N a t u ra lly it is m o s t c o m mo n in p lant breed ing in wh ich n u m bers ,

a re n o t s o s eriou s a m att er and in wh ich breed ing o perat io n s a re

,

les s ex pen s iv e .

A S has been r em a rk ed th es e three p u r po s es i n breed ing a re

,

ent i rely d is t inct T h es e d is t inct io n s s ho u ld b e c lea r ly in t he

.

m ind o f t he s t u d ent when s t u dy ing s y s tem s o f breed ing and t he ,

breed er h im self m u s t b e in n o s en s e u nc erta in as to wh ich o n e

is rea lly in his m ind wh en he b egi n s h is breed ing o pe rat io n s .

If his p u rpo s e is t o im pro v e h is o wn let him f rank ly ad m i t it ,

t o h im s elf and pro c eed acc o rd ing ly lea v ing h igh p ric es and ,

hazard o u s ent erpris es to o t hers I f o n t he o th er hand he .

, ,

h o pes to d o So meth ing d is t inct iv e fo r h is breed and is s at is fied ,

t hat he has t he m o n ey and t he pat ienc e to d o it then again his ,

p u rpo se is c lear c u t and his metho d s a re well ind icat ed .

A ll b reedi n g ex pen s iv e ex cept h erd i mprov emen t l

A ll fo r m s .

o f breed ing a re c o s t ly w/zen ev c r t/zc p u rpos e is to p rod uc e s om e

tlzi ug b etter tita n ev er b e f o r e I f t he p u rp o s e is o n l y .t o m u lt i

l x c ellc n c e th en it is c o m para t iv ely ch eap b u t t he o rigi nal
py e , ,

pro d u ct io n o f exc ellenc e wh ich is breed ing in the h igh es t ,

s en s e o f t he t er m is r elat iv ely expen s iv e b ecau se s o few

, ,

ind iv id u a ls plant o r an i m a l exc el ei th er th eir pred ec es s o r s o r

, ,

th eir c o nt em po rar ies and so few o f t hes e can pro paga t e th ei r

,

o wn exc ellenc e .

Wit h th es e c o n s id erat io n s in m ind it is wo rth wh ile to get a

c lear id ea o f t he d ifferent s y s t em s o f breed ing a v a ilab le fo r the
v ar io u s p u rpo s es .

1 H d im p
er m t i
ro v e p ien d i f t
s a n ext h h m t k
ress o n u s e n re eren c e o e o e s oc ,

wh het er p l t an i m l
o r an a .
60 2 PRA CT I C A L PR O BL EM S

SECT I O N I I G RA D I N G

By g rad i ng is meant t he m at ing of a c o m mo n o r r ela t iv ely

”

u n im pro v ed par ent wi th o n e t hat is mo re h ig hly i m pro v ed , that

is , a p u re bred T he m at ing might b e mad e eith er way b u t

.
”

in pract ic e t he m a le is tak en for the p u re bred parent fo r -

ec o n o m ic reas o n s On e p u re bred b u ll wi th a h erd o f t wenty

.
-

c o ws can gi v e all t he calv es i n t he h erd a pu re bred Sire ( t ha t is -

,

m ak e th em ha lf b loo d s ) wh erea s if t he m ak ing o f ha lf b lo od s

,

were att em pt ed in t he o th er way it wo u ld req u ire t wenty p u re

bred ind iv id u als and the cro p o f ca lv es wo u ld hav e n o mo re
,

im prov em ent ; bes id es wh ich t he i m pro v em ent m ad e wo u ld b e ,

n o t i n o n e b u t in t wenty lin es each with it s s had e o f d i ff er enc e ,

.

E xpres s ed in t er m s o f mo n ey it is p oss ib le to gi v e a ll t he ,

i —
ca lv es in a h erd a p u re bred S re that is m ak e th em all half
-
,

b loo d s at a t o ta l c os t of appro x im at ely t wo d ollars per calf

-
,

as s u m ing o f c o u rse a reas o nable n u m ber of c o ws in the h erd

, ,

a n d a b u ll at a m od erat e pr ic e b u t goo d en o u gh fo r gr ad in g If .

t he m ak ing of half b loo d ca lv es wer e acc o m p lis h ed in t he o th er

-

way h o wev er that is by pro v id ing the p u re bred pa rent o n t he -

—
, , ,

dam s S id e
’
it wo u ld c o s t at t he s a me relat iv e rat e C lose to
, , ,

fo rty d ollars as a m in im u m T h is s ho ws the n eces sarily extreme

.

c o s t o f p u re bred s as c o m pared with grad es .

D ISA PPEA R A NC E OF U N IM PR OV ED BLOOD BY T HE CONT IN U OU S U SE

OF PU RE BRED SIR ES
-

SI RE S D AM S O F FS PR I N G

G ENE RAT I ONS

Per Cen t of Un
im pro v ed

I m pro v ement by grad ing is o f c o u rse lim it ed to h erd im pro v e

m ent . It add s n o th ing t o the breed b u t it d is trib u t es b reed ,
60 4 PRAC T IC A L PR O B LE M S

e xc ellenc e ra p id ly and with extrem e c ertainty S u ch a s ire is .

al mo s t s u rely prep o t ent o v er t he da m s what ev er th ey m a b e

y , ,

and the m ath em at ic s o f m at ing S h o ws t hat if t he pract ic e is

c o nt in u ed fo r S ix gen erat io n s b u t o n e and a h alf per c ent o f t he
,

o rig i nal u n im pro v ed b lo o d will r em a in a s is s h o wn i n t he tab le ,

at t he bo tt o m of page 6 02 .

By th is we s ee th a t t he u n i m pro v ed blo o d s oo n b ec o m es in s ig
n ifi c an t and rap id ly d is app ear s T h is is why it is th at in t he
.

ea r ly day s o f a breed t he s ixth o r s ev enth cro s s is d ec lared

eligib le t o r ec o rd .

I t s h o u ld b e n o t ed that if any o n e o f th es e gen erat io n s b e

bred wit h it s elf ( grad es with grad es ) n o progres s is m ad e Thu s .

ind iv id u a ls o f the s ec o nd gen erat io n a re o n e fo u rth u n im

pro v ed and bred to a generat io n o f th eir o wn k ind th ey will
, , ,

s t ill r em a in o n e f o u rth u n im pro v ed By t he s a m e pr inc ip le

.
,

ha lf b lo o d s bred t o ha lf b loo d s will pro d u c e h a lf b lo o d s in d efi

n i tely . T he eff ect s o f grad ing c ea s e t he m o m ent we d is c o nt in u e
t he p u r e bred S ire
-
.

Ab u se of gr adin g T he ch ief dra wback in grad ing is that it is

.

lik ely n o t t o b e f o llowed u p T he breed er is alm o s t c ertain to

.

ch oo s e so me pro m is ing ha lf o r three q u art er b lo o d for a S ire -

beca u s e he looks a s g ood as a p u re bred and th en by t he ,

la w o f a n c es tra l h ered ity a ll i mp r ov em en t s tops exc ept t he litt le

that can b e acc o m plis h ed by t he S lo w pro c es s o f s elect io n .

A dv an t ages of gr adi n g . For ec on om ic p p

ur os es g r a d es may b e
eq ua l to p u r e wor t/ des s f or b r eed ing p u rp os es
b r ed s , b u t tlt ey a re

th is is t he p la in c o nc lu s io n o f what is well kn o wn of the prin

c i ples o f breed ing Grad ing is ch eap By t he u se o f a s ingle
. .

ind iv id u a l it c r
at o nc e s o meth ing m o r e than half o f the
s e u es

t o tal exc ellenc e o f the breed and if f o llo wed u p it will sec u re in
,

t ime thro u gh S ires a lo n e p ra c tica lly a ll of it

, ,
.

T h is is t he sy s t em o f breed ing t o b e rec o m m end ed t o the

grea t m a s s o f s t o ck m en and if it c o u ld b e gen era lly ad o pted
,

and f ollowed up it wo u ld add m illio n s to A m eric a n ag ric u ltu re .

E v ery s t o ck m an kn o ws that t he great b u lk o f t he b es t catt le in

t he m ark et s a re h igh grad e S h o rth o rn s and Her efo rd s
-
T he .

acc o m pany ing figu res s u rely S h o w that the les s kn own A ngus -

a n d i t s c lo s e relat iv e t he G a llo way

,
a re eq u a lly s u cc es s f u l fo r
,
6 06 PRA C TI C A L PR O B LE M S

grad ing is t he breed ing fo r begi nners He c a n eas ily s ho w t he .

no v ic e t hat if he wi ll k eep his o ld fema les o r if n o t get p lenty , , ,

o f s u ch as a r e ea s ily a v a ilab le he can ha v e a s m any g rad es

,

with in a y ear a s he can pro v id e fem a les n o w a n d that s peed ily ,

he will o wn a h erd that fo r a ll pract ical p u rpo s es exc ept b reed

in g will b e a s go od as anybo dy s all at a c os t o f o n ly t wo o r
’

three d o llar s per calf and c o rres po nd ing ly les s o r m o re for o t her
,

an im a ls S u ch a c o u r se will d emo n s trat e at o nc e t he exc ellenc e

.

o f t he br eed and mak e fr iend s n o t enemies of the m an and

, , ,

his n eighbo rs .

T he b u rd en is u p o n t he breed ers and o wn ers of p u re bred -

fl o ck s and herd s to lead in a cr u sad e fo r gra d ing T hey need .

t he m ark et fo r th ei r exc es s o f m a les and if th is m ark et were ,

f u lly d ev elo ped and t he m as s o f s tock men fu lly a liv e to t he

,

ad v antages o f grad ing th is m ark et alon e wo u ld ab so rb at good

—
,

pric es a ll the ma le o u tp u t fro u breed ing herd s

m o r a c o n su m ,

mat io n they s tand s o rely in need o f attaini ng .

T he f em ale o u tp u t of o u r breed ing herd s S h ou ld b e u sed fi rs t , ,

to reé n fo rc e the h o m e h erd s and aft er that to s u pply d efic ienc ies
,

i n o th er rep u tab le h erd s A n y f u rth er s u rp lu s an im als s h o u ld go

.

t o t he o pen m ark et exc ept i n s o m e rare cas es in wh ich th ey a re

,

need ed fo r t he real fo u nd ing o f n ew h erd s .

T he m a i n difli c u lty is t hat t h e breed ers as a ru le a re too , ,

int ent u p o n s elling fem a les and s ett ing u p a m u lt it u d e o f li tt le

breed er s in a s m a ll b u s ines s ; wh erea s th ey s h o u ld b e n ot only
int ent b u t p ers is t ent in s elling m a les fo r grad ing p u rposes
, , .

T h is is t heir great m a rk et t hei r nat u ral o u t let and it s exp loita

, ,

t io n is th eir o ppo rt u n ity T he a u th o r has replies fro m h u ndred s

.

o f breed er s o n t h is p o int A large S h are o f th em pro fes s to ex

.

pend as m u ch effo rt t o s ell fem a les as to s ell m a les and a few ,

ev en m o re A s so c iat io n s ha v e m u ch to d o a lo ng th is lin e
. .

Begin ani mal b reedin g b y gr adi n g Grad ing is t he s afes t b egin.

n ing ev en for t he pro s pect iv e breed er o f pu re bred s t ock N o t

,
-
.

o n ly is it ch eap and S af e b u t it will b ri ng o u t C lear an d s tro ng i n

,

t he grad es t he m a in breed po int s and a few generat io n s o f gra d es

,

fro m lo w to h igh will s pread o u t befo re the eyes of the breed er

s u ch a pan o ra m a o f br eed c ha ract er s a s he wo u ld n o t see in

years o f p u re breed ing o n a s m a ll s ca le ; ind eed th ere is n o ,

608 PR AC T I C A L PR OBLE MS

q u ick er cheaper o r m o re th o ro u gh way o f bec o m ing acq u aint ed

, ,

with a breed than thro u gh it s grad es .

Disadv an tage of gr adin g T he o n ly d is ad v a n tag e that c a n b e

—
.

m ent io ned is th is that the fir s t res u lt s are s o em in ently S at is

,

fact o ry that s o m e prom is ing grad e is lik ely t o b e select ed as a

s ire rega rd les s o f t he la w o f a n c es t ra l h ered ity wh ereu po n all
, ,

f u rth er im pro v ement s t o p s T his is so likely to b e the case that

.

i t m ay b e s ai d in gen era l t hat t he v ery s u cc es s o f grad ing is the

great es t gu aranty o f it s failu re .

S ECT I ON III —C R OSS I NG OR H Y B R I DIZ I N G

A lm o s t t he e xact
ppos it e o f g rad ing cro s s ing c o m b ines
o ,

anc es tral lines of t wo d is t inct ra c es breed s o r v ariet ies in the , , ,

h o pe eith er o f s ec u ring a b lend o r else of gett ing a fo rt u itou s

c o m b inat io n o f C haracters .

T h is f o rm o f breed ing is adapt ed o n ly t o t he pro d u ct io n o f

n ew s trai n s i n wh ich it exc els
, Of c o u rs e i t s o m ix es bloo d lin es
.

a s t o eff ect u a lly d es tro y the i n flu enc e o f t he anc es try and a ll

m ean i ng and v alu e o f ped igree I t s h o pe is in s tart ing a n ew

.

s t ra in wh ich m a p erchanc e breed p u r e

, y .

T he o peratio n o f Mend el s law t eaches ho w s m a ll is th is

’

chanc e I f th is law alway s h eld with all r ac es a n d c haracters

.
'

it wo u ld o f c o u rs e b e i mposs i b le t o s ec u re p er m an ent s t ra in s
by cro s sing b u t the fact rem ain s that perm an ent hybrid s [ra v e
,

en tly b een s ec u r ed by th is met ho d es pec ia lly a m o ng p lant s

f q
r e u , ,

wh ich is a n o t ewo rt hy fact in breed ing .

Adv an t ages of c rossin g N o t with s t a nd ing t he o perat io n o f

.

M end el s la w a s a gen era l pri nc ip le cro s s ing is a fr u itf u l so u rc e

’

o f n ew s t ra i n s Hyb rid izat io n is bett er adapt ed to p lant s than

.

t o a n im a ls b eca u s e o f t he n eed o f V igo ro u s s elect io n aft erward

and th eref o re o f relat iv ely large n u m bers I t was a fa v o rit e
, ,
.

m eth od o f p lant im prov ement t wenty y ears ago b u t it h as fa llen ,

largely int o d is u s e b eca u se o f t he inc o n s tancy o f M end el s

’

midd le t erm ( t he 5 0 pe r c ent appa r en t hyb rid s ) and beca u s e as

go o d o r bett er res u lt s can o ft en b e s ec u red by s elect io n alo n e ,

wit/t ou t d es t ruc tion f

o tlzc p ed i
g
ree a nd t/re i nflu enc e o f til e
a nc es t ry .
6 10 PRA CT I C A L PR OBLE MS

The reci procal cross . S t range it may at fi rs t appea r the

as ,

t wo p os s ib le cro ss es by int e rchange o f t he

sex es o ft en th o u gh ,

n o t al way s d iffer s u b s ta nt ially I t is s aid t hat the c o m mo n

,
.

mu le m o re n early r es em b les t he as s and t he h inny t he h o rse , .

Ot her in s tanc es ha v e been n o t ed and t he po int ha s been u rged

,

that rec ipro ca l cros ses a re in genera l d is s im ilar I t is the .

writ er s o pin io n that t he r u le app lies o n ly to t ho s e part ic u la r

’

cha ract ers in wh ich t he o n e parent ( eit her m ale o r female) is -

prepo t ent o v er the o th er beca u s e of sex Ho wev er s ta t is tical .

,

ev id enc e o n r ec ipro cal cro s s es is al mo s t t o ta lly lack ing .

T he wh o le s u bj ect o f hybrid izat io n s eem s at present t o

pro m is e litt le of int eres t t o an im al breed ers b eyo nd the prod u c
t io n o f t he c o m mo n mu le b u t if we may p lac e a s hrewd gu es s
, ,

it will yet b e fo u nd a fr u itf u l s o u rc e o f n ew v ariet ies in c er ta i n

ra c es o f plant s in wh ich pro pagat io n is so easily effect ed by
,

b u dd ing g raft ing o r o ther fo r m o f asex u a l m u lt iplicat io n th u s

, , ,

a v o id ing t he effect s of M en d el s law in a way q u ite impos s ib le

’

with ani m a ls

SECT I O N I V — LI N E BR EED I NG

By “
lin e
breed i ng is m eant t he res tr ict io n o f select io n an d
mat ing t o t he ind iv id u a ls o f a S ing le lin e o f d es c ent The p u r .

po s e o f th is s y s t em o f breed ing is real breed im pro v emen t ,

t o get t he b es t that can b e go tt e n o u t o f t he rac e an d b etter ,

th a n ev er befo re if po s s ib le .

E xper ienc e has S h o wn th a t if t he p u rp o s e b e breed im pro v e

ment o r ev en h erd im pro v em ent carried to it s lim it s i t is n o t
, ,

en o u gh t o c o nfin e s elect io n t o t he lim it s o f t he breed A ll .

breed s a re exc eed ingly v ariab le and real res u lt s ai mi ng at any

,

th ing m o re than mere m u lt ip licat io n can fo llo w o n ly c los ely

dra wn lin es ith n t e breed
w i h — breed ing in lin e o r li ne breed ing
, , .

L in e breed ing exc lu d es ev eryth ing o u t s id e t he appro v ed and

C h o s en lin e o f breed ing I t n o t o n ly c o m b in es a n i mals v ery
.

s im ila r i n th ei r charact er s b u t it narro ws t he ped igree t o few

,

and c lo s ely related lin es o f d es c ent Th is p u ri fi es t he ped i

“
.
”

gree rap id ly and gi v es t he anc es try t he larges t p o s s ible o ppo r

t u n i ty . T he s y s t em is em in ent ly c o n s er v at iv e I t d is c ou rages .
 SY ST EM S O F BR E ED I N G 61 I

variability and rap id ly red u c es it to a min i m u m Mo r eo v er

,
.
,

what ev er v ar iat io n s d o occ u r will b e i n li n e wi t/t t/ze p r o m i n en t

c lza r a c ter s o f tire eli asen
b ra nd s o
f tlze b r eed .

A dv a n t a g es of line
b reedin g . at u re o f
The n

res u lt s sec u red by th is

s y s t em c an a lm os t c er
ta in ly b e pred ict ed ; and
wh en th ey d o appear and ,

im pro v em ent is at hand ,

i t is back ed u p by t he
m o s t po werf u l h ered itary
infl u enc e o bta inab le b e F 5 B D ,k 6 3 d li IG b . d 0 B . k h i
a ro n u e , a n e- re er s re.

ca u s e o f the S im plic ity P p t y f A J L j y R ro I

er ll i io . . ov e o , o sc o , no s .

F ig 5 d 5 h w g t f t h i b
u res 1 an 2 s o e o s oar
and s tren gth of t he an
c es t r
y w
,
h ich if t he s elect io n ha s b een g oo d a ll
,
p u lls i n t he ,

s a m e d irect io n The rec o rd s o f a ll breed s will S h o w the p ro

.

n o u n c ed res u lt s that ha v e f o llo wed j u d ic io u s lin e breed i ng A .

v o lu m e c o u ld b e fi lled with p ict u res o f fa m o u s an im a ls s o

p r o

FIG .
5 1 . L in e b re
-
d B erk hi
s re i
p gs . G et of B a ro n D u k e 6 3d

d u c ed . h o wn are of s win e
T ho s e S reaso n that t he p ig is ,
for t he
po p u larly s u ppo sed to b e t he m o s t
s en s it iv e t o c lo s e br eed ing .

Disadv an t ages of lin e b reedin g T he ch ief danger in lin e .

breed ing is that t he breed er will select by ped igree aband o n ing ,
612 PR A CT I C A L PR O B LEM S

rea l in d iv id u al select io n A line bred ped igree is v alu able o r

.
-

d a ngero u s in exact pro po rt io n as t he ind iv id u als ha v e been kept

u p t o grad e I t will n o t replac e s elect io n b u t o n t he c o nt rary
.
, , ,

calls fo r t he mo s t d iscri m inat ing care wi t/t in t he line .

I f the breed er s elect s by pap er and n o t in the yard s an d a , ,

few gen erat io n s o f in f erio r a n im a ls creep in th en li ne breed ing ,

will c o ns ig n t he wh o le b u nch to t he lim bo s q u ick er and mo re

c ert a in ly than will any o th er kn n y t o w s s em o f br d ng
ee i — a ,

F IG .
5 2 . L in e b re
-
d y li
ea r n
g B er k his res . G et of Ba ro n D u k e 63 d

fat e that has o v ertak en m o re than one lin e that u nfo rt u nat ely
beca me prem at u rely fas h io nab le .

Lin e b reedi ng t h e b est sy st em ther for improv emen t . No o

s y s t em o f br eed ing ha s ev er s ec u red t he r es u lt s that lin e breed

in g ha s s ec u red and if t he pres en t s t a te o f kn o wledge is reas o n

,

ab ly s o u nd n o o t h er sy s t em will ev er b e S O p o werf u l in getti ng

,

t he m o s t po s s ib le o u t o f a giv en breed o r v a riety es pec ia lly o f ,

an im a ls and th is with t he grea t es t c ert a inty as we go alo ng

, .

T he o n ly req u irem ent is n ot to a ba n d on i n d i v id ua l s elec tion

, A .

ped igree is n o t a cr u tch o n wh ich inc o m pet enc e can lean ; it is

a gu ara nty o f b loo d lin eS — a fi eld i n s id e o f which breed ing
,

o perat io n s and s elect io n m ay wi th c o nfid enc e b e c o nfin ed .

T he wo rd c o nfin ed is u s ed ad v is ed ly fo r aft er lin e breed

”

, ,

in g has been p ract ic ed fo r a few generat io n s t he anc es t ry ,

bec o mes a k nd i o f p r br d
u e ee o f it o wn
s — a breed with in a ,

breed so t s peak
,
o — and a ny att em pt to intro d u c e b lood fro m
,
6 14 PR A C TI C A L PR O B LEMS

of t he dam . T h is fo rm o f breed ing pract ic ed wh en it is de

is
s i red t o s ec u re a ll that is po s s ib le o f
t he b loo d o f t he S i re .

2 B reed ing t he d a m t o her o wn s o n o r s o n s s u cc es s iv ely

.
,

th u s increas ing t he b loo d lin es o f t he fem ale s id e Th is fo rm is .

p ract ic ed wh en it is t he dam s b loo d lines that are t o b e ’

pres erv ed and c o nd en s ed B o th sy s tem s a re n ec es sarily limit ed

.

t o t he lifet im e o f t he ind iv id u a ls in v o lv ed Eith er s y s t em can

.

o f c o u r s e b e appro x im at ed by t he u s e o f g ra ndda u ght er o r

g r and s o n w
, h ich w o u ld by c o m m o n c o n s ent b e called inbreed

in g b u t relat io n s h ip mo re remo t e wo u ld g en era lly b e regard ed

,

m erely as lin e breed ing .

3 B
. re ed i ng t o ge th e r o f br o th er and S is t e r — a f o r m o f i n ,

breed ing wh ich pres erv es the b lo o d lin es fro m bo th s ire and d am
in eq u a l pro p o rt io n s I t is inferio r to eith er o f t he o th ers as a
.

m ean s o f s trength en ing prev io u s ly ex is t ing b lood lin es b u t it is ,

freely em p loy ed wh en the c om b ina tion ha s pro v ed ex c ep tiona lly

s ucc es sf u l v i rt u a lly es t a b lis h i ng a n ew typ e
,
I t has all t he da n gers
.

o f t he o th er t wo and in a la rger d egree b eca u s e we ha v e p rae

, ,

t ically no acq u a intanc e with the n ew c o m b inat io n wh ereas in ,

s tr ength en i ng t he pr o p o rt io n o f o n e lin e o f anc es try o v er an o th er ,

wh eth er it b e that o f t he S ire o r that o f t he d am we a re d eali ng ,

with prev io u s ly ex is t ing b loo d lin es kn o wn t o b e ha rm o n io u s .

A m o ng p lant s th ere are t wo f o r m s o f inbr eed ing na mely ,

1 T hat in wh ich the fert ilizat io n is with po llen fro m a n o th er

.

fl o wer o n t he s a me plant .

2 T hat in wh ich fert ilizat io n is by p o llen o f the s a m e fl o wer

. .

T h is being h erm aphro d it ic is t he c lo s es t i maginab le inbreed ing

, , ,

a n d exc eed s anyth ing that is p o s s ib le wi th a n im a ls .

Adv an ta ges of in b reedin g N o b o dy C la im s ad v antages in in

.

breed ing per s e b u t it is t he ac m e o f line breed ing and wh en

, ,

s u per io r ind iv id u a ls a re at hand it is t h e m o s t po werf u l m eth o d

kn o wn o f m ak ing t he m o s t o f th eir exc ellenc e I t is the metho d .

by wh ich the h igh es t po s s ib le perc entage o f t he blood of an

exc ept io n a l ind iv id u a l o r o f a p a rt ic u lar ly f o rt u nat e n ick c an
b e pres erv ed f u s ed int o and u lt i m at ely m ad e t o C ha ract erize a n
,

ent i r e lin e o f d es c ent o n b o th s id es .

I f pers is t ed in t h e o u t s id e b lo o d d isapp ea rs by the s ame la w

,

that go v ern s gra d ing and t he p ed igree is s peed ily enriched to

,
 SY S TEM S OF B R EE D I N G 1 5

an almos t u n lim it ed extent by the b lo od Of a single an im a l ,

in pra ct ic e gen era lly that o f t he s ire I t is a m eth od n o t so

,
.

m u ch o f o riginat i ng exc ellenc e as o f m ak ing t he m o s t of it wh en

it d o es appear and it is n o t t o o m u ch t o s ay that a large p ro
,

po rtio n of the really great s ires hav e been s tro ngly inbred .

A n i nbred an im a l is o f c o u rse en o r mo u s ly prepo t ent o v er

ev eryth ing els e I t s ha lf o f t he anc es try being largely o f id en
.
,

t ical b lood is alm o s t c erta in to d o m inat e t he Offs pri ng I nbreed

,
.

in g is th eref o re rec o gn ized a s t he s tro nges t o f all breed ing

, , ,

giv ing ris e to t he S im p les t o f ped igrees an ad v antage q u ick ly

,

reco gnized wh en we recall the law of anc es tra l hered ity I n th is .

res pect it is all that line breed ing is and m o re .

A s ec o nd ad v antage is that s u cc es s f u l a s s o c iat io n s o f C har

ac t er s a re pres er v ed intact and n o t S hatt ered by t he i nf u s io n o f

n ew s t ra in s I f t he breed er were d ea ling with b u t a S ingle char

.

acter he c o u ld read ily fi nd it s eq u al ; and th ere wo u ld b e litt le

need fo r inbreed ing ; b u t ev en if breed ing fo r b u t a s ingle u t ili
tarian charact er he a lway s has at lea s t t wo o th ers V igo r and
, ,

fertilit y whic h m u s t b e inc lu d ed in select io n I n pra ct ic e he has

'

.
,

many m o re and a S ingle ind iv id u a l that c o nta in s all or m os t o f

,

th em in a h igh d egree is a v eritab le b o nanza ; nat u rally the

t em ptat io n is to m ak e t he m os t o f an o ppo rt u n ity wh ich is no ne
t oo freq u ent in t he breed ing b u s in es s .

A ll th ings c o n s id ered n o o th er kn o wn m eth o d o f breed ing

,

eq u a ls t his fo r int en s ify ing b loo d lin es d o u b ling u p ex is t ing

,

c om b inat io n s and mak ing t he m o s t o f exc ept io nal ind iv id u als

,

o r o f u n u s u a lly v a lu ab le s tra in s .

Di sad v an ta ges of in b reedin g C learly h o wev er th is is no t a

.
, ,

g u n t o h i t t h e b e ar and m iss t he ca lf T h is d o u b.li ng

”
u p
proc ess th is int en sifying of Chara ct ers increas ing th eir pro s pect s
, ,

fro m po ss ib ility t o pro bability and aft erward to c erta inty wo rk s ,

exact ly t he sa m e fo r o n e charact er a s fo r an o th er ; i t afi ec ts a ll
'

d i a m eter s of tlze i n d i v i d u a ls in v olv ed ba d a s well a s g ood ; and

,

s o it is that th is m eth o d wh ich is app licab le to b o th p lant and

,

an im al breed ing and wh ich aim s at m ak ing t he great es t u se

,

po ss ible of o u r m o s t v a lu able po ssess io n s has been follo wed ,

alik e by t he m o s t s tr ik ingly s u cc ess f u l res u lt s and by the m os t

s t u pend o u s d is a s t er s that ev er o v ert o o k t he breed ing b u s in ess .
616 PRA C T I CA L PR OB LE M S

Plenty xa m p les o f s u cc ess es c a n b e in s tanc ed and ev ery

of e ,

breed er is fa m iliar with th em T he fa ilu res hav e been m any .

,

b u t th ey a re n o t t o b e c o u nt ed h ere fo r t he b lo o d lines in ,

v o lv ed a re lo ng S inc e ext inct .

Special dan gers f rom in b reedi n g T rad it io n ev erywh er e has i t

.

th a t inbreed ing if lo ng c o nt in u ed is pract ically c erta in to en d

, ,

in lo s s o f V igo r and o f f ert ility and p lenty o f in s t a nc es a re giv en

,

“
to p ro v e it .

N ow a ra t io na l c o n s id erat io n o f the princ ip les o f tran s m iss io n

h as a lready led u s t o exp ect that bad charact er s a s well a s good
will b e int en s ifi ed W e c o u ld n o t expect S O p o werf u l a m eth o d
.

t o wo rk o n ly to o u r a d v antage and t o gra nt im m u n ity fro m d is

ad v a ntage in a ll ca ses .

What we wa nt to kn ow is wh eth er in res pect to tro u b le , ,

we a re t o lo o k o u t fo r lib eli/zood o r fo r c er ta in ty ; wh eth er d isas

t er is i n ev itab le o r o n ly ext rem ely pro ba ble
,
T h is q u es t io n has .

been m u ch b efogged by c erta in catchy s t a t ement s s u ch as ,

N at u re a bh o rs inc es t u o u s breed ing a ll o f wh ich c o nf u s e an

”

eth ic a l and s oc ia l q u es t io n with t h e b io lo gica l o n e in wh ich o nl y

we a re int eres t ed .

I n b reedi n g n ot n ec es sari l y disa st r ou s Ou r att ent io n is c o n .

s ta n t l
y ca lled to nat u re s pr o v is io n s fo r prev ent i ng i nbreed in g
’

,
”

a n d to ingen io u s d ev ic es fo r ind u c ing cros s p o llinat io n by

“

in s ect a id ; b u t we a re n o t r em ind ed that m any s pec ies o f

”

plant s a re S elf p o llinat ed n o r is o u r att ent io n ca lled to t he m any

-
,

fa m o u s si res that were s tro ng ly inbred n o r to t he fact that in ,

nat u re a m o ng grega rio u s a n im a ls the h ea d o f t he h erd is S ire o f

pra ct ica lly a ll the yo u ng ( s o lo ng as he rem a in s m as t er) m any o f ,

o f wh o m ar e th u s d o u b ly h is N o r d o we h a v e i t ca lled to o u r
.

a tt en t io n th a t wh ile c o rn S eem s p ec u lia r ly s en s it iv e t o i nbreed

,

in g wh eat is s elf f ert ilizing to t he c lo s es t po s s ib le d egree and

,
-
,

that it is perhap s t he m o s t v igo rou s pro lific and all ro u nd c o s , ,

-

m o po lita n s u cc es s a m o ng o u r d o m es t ic p lant s .

Lack of v i gor an d low f ert ili t y t h e t wo mos t common def ec ts .

I f what ha s b een s a id and S h o wn has any m ean ing i t is that a ny ,

charact er can b e bred u p o r d own s trength en ed or weak ened ,

by th is m eth o d o f b reed ip g Why th en its ev il rep u tat io n with

.

res pect t o v igo r a n d fert ility ? I s th ere s o m e inh erent in j u ry

6 18 PRA C TI C A L PR OB LEM S

Darwi n ’
far as plants are c o n
s ex per imen t s .
1
F o rt u nat ely so

c ern ed we a re n o t with o u t s o m e acc u rat e data t end ing t o S h ow

t he act u a l eff ect o f inbreed ing u po n t he t wo m o s t im po rta nt
charact ers h ere u nd er d is c u s s io n — na m ely v igo r and fert ility , , ,

— and fo r a great v ariety o f s pec ies T he experi ment s are t oo .

ext en s iv e to f u lly d is c u s s ev en by ab s t ract c o v eri ng as th ey d o ,

s o me fifty s ev en s p ec ies belo nging to fifty two gen era ; b u t

-
11 -
,

th eir res u lt s m ay b e briefly s tat ed .

T he caref u l s t u dy o f th es e exp erim ent s s h o ws t he f o llowi ng

facts : ( I ) that in g en era l and with o u t a d o u bt cro s sed fo rm s , ,

( b o th th ey and th ei r o ff s pri ng ) a r e o n the a v erag e m u ch mo r e , ,

f ert ile and far m o re v igo ro u s than are the s elf fert ilized ( 2 ) b u t -

tlza t t/zis is a ll s ec ies , n o r is i t tr u e f a ll in d i v id ua ls ,

n ot tr ue o f p o

ev en withi n thos e spec ies mos t sens i ti v e to i n b r eed ing .

T h u s o f t he 8 3 S p ec ies t es t ed fo r h eight , 2 6 o r n early o n e

, ,

th ird were eith er with in 5 per c ent o f t he h eight o f th eir cros s

,

bred c o m pan io n s , o r els e exc eed ed th em in h eight Of th ese 2 6 .

ca ses h o wev er he c o nc lu d es th a t 1 4 were act u ally inferio r

, ,
if ,

no t in h eight at le s t in o th er respect s
a ,
— lea v ing 1 2 o r o n e , ,

s ev enth o f a ll that q u it e C lea r ly were n o t i nf er io r wh en i nbred

, ,

and in s o m e ca ses were d ec id ed ly the b ett er fo r it Co nc lu d ing 3

.
,

D arwin remark s

The f if w
re o re l d th e exc u
p ie w h i h ea e pp o
e s i m a t lec
y e q as l t h e r e c r a r x e u ,

ar e th i t y r -
sev en
p i i w
s h i hec est h e m n f t h e m e c h e i g h ts o f t h e ea n o an

Ch l
1 D wi
a r es C ar d S l f F t ili t i
n, ro s s i t h V g t b l K i gd m
an e er za on n e e e a e n o ,

p .
4 8 [D2 A pp l. t 8 C me on
p y ] I t
1 i of t t an t h t w d. t p s u n o r u na e a e o no oss es s

e q ll
ua y f ll d
u tand t t
ex a c i b d i
a a as
g m g im o l b t n t t h
ree i p i
n t a on an a s, u a s o n our

k wl dg i l imit d t g
no e e s l e l t d t i d i id l p i
o en era res u Th m k d
s an o n v ua ex er en c es . e ar e
s u c c es s o f l b di g d
c o se ree i b n di g i
an h d i tt ib t d t t h
ev en n ree n n our er s s a r u e o e

s p iec al k i ll S f m ot b
a
“
d Th t th i i
as e r ree t th f ll er .
p l
"
t i a i h w s s no e u ex an a on s s o n

by h t e ex
p i f t h
e r en c e oU it d S t t e B n f A im l I d t y S m y
e a es u rea u o n a n us r . o e ears

a
g i
o t b m ec a y t m
e n ec es sa r t h t ko f re
g i p iove
g t w e s oc
q t o u n ea s o ne ua r ers a

c o ns id b l di t
e ra e w y A s anc e a t a m w . t d
s ev ere s o r t d ly a s en c o u n ere en ro u e a n on a

f w p ig w
e s d F m t h f w d wi th i f i f t id b l d th
ere sa v e . ro es e e , an no n us on o ou s e oo , e

p res en t t k i
s oc d d d d th w it i
s es c en e , d ib l y i f m d th t th t k i
an e r er s c re n or e a e s oc s

e xc e p t i ll y ona ig v d f t il
o ro u s a n er e.

9
Th t d e s u t d i i g th d t
en es r n
p t h fl t e f a a u l f f t ili t i
on e e i ec s o c ro s s o r se -
er za on n

g en era l h lsd d
ou h p ii
r ea
pp 3c8 a 8 4 . fv D w i, C . 2 d S -
lf
2 F t ili
, o ar
’
n s ro s s a n e er za

ti on, e c t ; f .d t or i g th ff t p
a a c o n c ern n d p d ti h h ld d e e ec u on s ee ro uc on e s ou rea

c h p i pp 3 —3 5 5 d f d t
a . x, . 12 an i g thor ff t h p iii pp 8 5 3
a a c o n c ern n o er e ec s , c a . v , . 2 - 1 1

for d t il d p t f d iff t p i
e a e re or s o h p ii i p i ll y ii
eren s ec es s e e c a s -v
es ec a
—
. .
,

D wi
3 ar C n, d S lf F t il i t i
ro s s a n e t pp 7 9 83
er za on, e c .
, . 2 2 .
 Y
S STE M S O F BR EED I N G 619

c rossed p la n ts ex c eed s h
t a t o f the s e f fert ili zed b y 2 2 per
l -
c en t , wh ereas
h
t ere a re onl y fiv e s pec ies in w i c h h the m ea n o f the m ean h eig h ts o f t he

self -
fertilized plan ts ex c eed s t h a t o f th e c ross ed , an d h
t is o n l y b y 9 pe r c en t .
1

T he writ er again calls att ent io n to the fact that wh ile a v erag es
a re o f p rim e c o n s eq u enc e i n c o m m erc ia l tran s act io n s th ey d o ,

no t d ec id e pr i nc ip les o f breed ing and it is extrem ely s u gges t iv e ,

that ev en fi v e s pec ies were d ec id ed ly m o re v igo ro u s when inbred .

I t d et er m ines d efi n it ely that th ere is n ot/t i ng in /zer en tly a nd

n ec essa r i ly ev i l i n i n b r eed i ng p er s e fo r if s u ch were t he cas e it , ,

wo u ld m ak e i t s elf ev id ent i n ev ery in s tanc e .

S p eak ing o f t he fertility o f s elf f er t ilized flo wers D arwin s ays in -

, ,

T h eir fert ility ranges fro m zero t o f ert ility eq u aling that of t he
cros sed fl owers ; and of th is fact n o exp lanat ion can b e o ff ered .
"

N o t o n ly was t h is tr u e b u t t he s elf fert ilized f o r m s were so m e

,
-

t imes actu a lly m o re fert ile than the cro s sed 3

.

T h is m y s t ery fo r wh ich n o exp lanat io n can b e o ffer ed

, is ,

largely c leared u p by o u r mo d ern k n o wledge o f h ered ity as ,

is s h o wn by wh a t fo llo ws .

Th e t ot al effect s of in b reedin g A ll charact ers b o th goo d and .

,

bad ex is t in v ario u s d egrees in d ifferent ind iv id u als T he pro b

,
.

lem in breed ing is to s ec u re t he s tro nges t c o m b inat io n s o f d es i r

ab le characters and it is eas y to s h ow that th is is acc o m p lish ed
,

by inbreed ing N o t o n ly that b u t it is als o ea sy to s h o w that

.
,

the s a m e m eth od s will s ec u re t he lo wes t atta inab le int en s ity ,

a c o n s u m mat io n d es irab le with u n welc o me charact ers and goo d ,

to kn o w ab o u t a s a gen eral p o s s ib ility .

T ak e fo r exa m p le t h ree int en s it ies o f any s ing le charact er

, , ,

d is regard ing fo r t he m o ment all q u es t io n s o f c o rrelat io n L et .

th ese three inten s it ies b e repres ent ed by 3 2 and I res pec , , ,

tiv ely 2 b eing t he m ea n

,
.

I f n o w we exc lu d e i nbreed ing we find three u n io n s p os s ib le

, , , ,

— nam ely 3 2 3 I and 2 I b u t if we reso rt to inbreed

, , ,

in g we m ak e a lso the m at ings 3

, 3 2 2 I I Wh i ch u n io n s , ,
.

a re rich es t in res u lt s ? I n wh ich ha v e b loo d lin es been m o s t

int en s ifi ed

1 D a rwin , C ro ss and l
S e f F er t i iza t io n , l et c .
, p . 2 8 3.
2 Ib i d p 3 2 6
—
. . .

3 I b id .
pp .
3 2 2
3 5
2 .
620 PR A C T I C A L PR OB L EM S

R E LA T I V E EFFE C T OF OU T B R EE D I NG AND or I N BR EE D I NG

M A rm t '
M m - Px x s x r s On s p nm c

O u tb ree in gd

d
I n b ree in g 2 2 o

l 1

F ro m th is we s ee that b o th s y s tem s pro d u c e t he s a m e mea n ,

b u t that inbreed ing pro d u c es t he wid er extrem es ( 3 and I ) .

Henc e t he great es t r a ng e o f po s s ib ilit ies lies with inbr eed ing so ,

far as i mmed ia te pa ren tag e a lon e is c o nc ern ed a n d t he ad v antage ,

is o f c o u r s e s t ill great er in t he anc es t ry f a rth er back .

A ga in t he tab le s h o ws what will h a pp en o n t he av erage u nd er

,

t he law o f regres s io n b u t in exc ept io n a l cas es t he la w of p ro

,

g r e s s io n w i ll app ly fr
,
o m wh ich we s ee th a t t he ad v a ntage fo r

inbreed ing is s t ill great er ; in o th er wo rd s i t is by inbreed ing ,

that t he h ig hes t and the lo wes t a tt ain a ble res u lt s can b e p ro

d u c ed and th is is bec a u s e n o o th er s y s t em can pro d u c e s o h igh
,

l o w) a m id p a rent o r in th e en d s o p u re an anc es try A l l

( o r -
,
.

o f th is ind ic a t es a pr inc ip le th a t is a b u nd a nt ly p o werf u l fo r

int en sify ing go o d ch a ract er s o r fo r breed ing o u t ev il o n es T he .

fact that it is th u s po werf u l arg u es again s t its u s e with any b u t

s u p er io r ind iv id u a ls .F u rth er m o re inbreed ing is a s u preme t es t
,

o f exc ellenc e and if a fa m ily lin e o r an i nd iv id u a l end u r es it it s

, ,

characters a re a b o v e reproach .

Not al l in b red in di v i dual s in f eri or t o t h e cr os s b red ,

-
ev en in
spec i es especi al ly sen si t iv e t o in b reedin g On e o f D arwin s m o s t
.
’

t n
ex e s iv e s er ies o f exp erim ent s was carr ied o n with t he c o m m o n
62 2 PR A CTI C A ] PR OB L E MS .

On t his po int c o m pare t he fact s repo rt ed in t he f ollowi ng

tab le fo r the fo u rth generat io n ‘
.

Here a gain each inbred p la nt is inf er io r t o it s part ic u lar m at e ,

b u t on ly t/zr ee o f t he cro s s bred s eq u a led t he b es t inbred p lant

-

o f t he s eri es
( 8 0 i nch es ) and ,

I a ll b u t on e o f t he inbred s were
” W
E:
“ 0"
C l xo ss x u RsD

m o re v igo ro u s than the poo r

es t cros s bred -
.

T he s a m e g eneral fact is
n o t ic eab le in t he next ( fifth )
gen era t io n th o u gh n o t q u ite ,

so pro n o u nc ed exc ept that in ,

one c as e t he inbred p lant

eq u a led it s o wn m at e Ev i .

d en t ly s o m eth ing was prepar

in g to h a pp en .

Th e appear an ce of Hero I n t he n ext (s ixth ) gen erat ion

.
”

there appear ed a s p ec ia lly v igo ro u s p lant th a t o v ert o pped its o wn

c o m pet it o r by ha lf an inch and exc eed ed in h eight all b u t three
o f t he s eries D ar win na m ed th is p lant
.
“
Hero and rem ark s , ,

I wa s s o m u ch s u rpr is ed at th is f a ct that I res o lv ed to a sc erta in

wh eth er th is p lant wo u ld tran s m it it s po wers o f growth t o it s
s eed ling s .

A cc o rd ingly he f ert ilized a n u m ber o f fl owers of Hero wit h

th eir o wn p o llen and p lant ed the s eed lings in c o m pet it io n with
,

o th er inbr ed p lant s and with cro s s bred as well T he two tab les -
.

o n t he n ext pag e s h o w ho w t he d es c endant s o f Hero acq u itt ed

th em s elv es .

Here th en o u t o f a s p ec ies s en s it iv e t o i nbreed ing has a risen

, , ,

a p lant that is s tro ng v igo ro u s and pro lific and its o wn inbred
, , ,

s eed ling s at o nc e d em o n s trat e th ei r s u p e rio rity n o t o n ly t o o th er

inbred s t o ck b u t a ls o t o th eir cro s s ed c o m pet it o rs A s D a rwi n .

rem ark s 2
,
Hero tran s m itt ed t o it s o ff s pring a pec u liar c o ns ti
t u t io n ad a pt ed fo r s elf f ert ilizat io n and again
- 8 “
I t appear s , ,

th eref o re that Hero and it s d es c endant s ha v e v ar ied fro m t he

,

c o m mo n type n o t o n ly in acq u i ring great p ower o f growth an d

D wi 1 C ar d S l f F t ili t i
n, t
ro s s a n p 34 e er za on , e c .
, . .

I b id p 5
2
. . I b id 5
0 .
3 . 1 .
 SYST E MS OF B R EE D I N G 62 3

increas ed fertility when s u b OFFSPRING o r H ERO CO MPA RE D WIT H

d s elf f ert ilizat io n b u t O RD IN A R Y I N B RE D SEE DL I NGS
je c t e t o -
, ,

i n n o t pro fit ing fro m a cro s s SEV E NT H GE N E R AT ION o r

with a d is t inct s t o ck . I N BR EE D I N G
”

Here is exc ellenc e thro u gh

inbreed ing u nd er what may N w m 0?

b e called t he hard es t c o nd i
t io n s and it gi v es great en
, i n .

c o u ragem en t t o t he belief that

if it is nec ess ary t o s ec u re a

s t ra i n o f p lant o r an im a l that
8 i 2 n
will pro s per u nd er inbreed i ng
.

that s train can b e prod u c ed ,

and that it s pro d u ct io n is a

q u e s t i o n o n ly o f t im e pa ,

t ien c e a n d expe n se Hero will OFFS PR I NG OF H E RO COM PA RE D WIT H

,
.

u nd o u bt ed ly b e ca lled a m u C R OSS FER T I LIZ A T I ON SEED LINGS -

,

t a nt in th es e d ays b u t m u tant s ,
S EV E N T H G E NE RA TI ON o r
a re welc o m e I t m u s t b e b o rn e
.
I N BR EED I N G
in m i nd that Hero was n o t t he
On ly i nd iv i d u a l that d em o n “ “ UM 0"

s t ra t ed it s s u p erio r i ty to cr os s

bred p lant s b u t that th is was

, i n .

a c o m m o n ci r c u m s t a n c e
throu gh o u t t he experiment s .

N or was t he m o rn ing g lo ry -

t he o n ly cas e o f t he k i nd C o n .

c ern ing his experim ent s with M im u lu s ( t he mo nk ey flo wer of n o -

,

c o n seq u enc e t o u s exc ept a s s h o wing a princ ip le) he say s 1

I the th i d
n d f
r t h ge era t i
an t ll
ou r i ty ft ll d ed t ha
n o ns a a v ar e , o en a u o, v

i g l ge wh ite fl we
n ar b l t h d wi th c ri m
o rs pp eo c d m o ge t b t/ th e so n , a a re a n s o z

the sel f fert ilized p lan t s

i n ter c ro ss ed a n d I t prev a iled in a ll the la ter self
-
.

fer t ilized gen era tio n s , to the ex c lu s io n o f ev ery o t er v ariet , a n d tran s h y

h h
m i tted i ts c a ra c ters fait fu lly, b u t d is a ppea red fro m th e i n terc rossed
p l a n t s .T he s el f fer ti li zed p la n ts
-
b elo n i
g gn to t is v a r iet
y were n o t o n l
y h
ta ller b u t mo re fert ile t ha n the i n terc ro ss ed pla n ts , t ou g h h th ese la tte i r n

the ea rlier gen era tio n s were m u c h ta ller a nd m ore fer tile th a the s elf
n

fertilized plan ts .

1 D a rwin , C ro ss an d S elf Fert iliza tio n , etc .,

p 3
.
4 8 .
( I ta lics a re m in e ) .
624 PR ACT I C A ] PR . OB LE MS
He add s ,
1
Th is v a r iety s eem s to lea v e b ec ome spec ia lly a da
p ted

to p ro
fi t i n ev ery way by se lff er tilia a ti on , a l t/zong /z t/zis p r oc es s

wa s
'

iry u mon s to flee pa r en t p la n ts d u r i ng t/ze fi r s t f ou r g en era

'

so

D ar win s wh o le d is c u s s io n o f h igh ly s elf f ert ile v arie

’
t i on s .
-

t ies is exc eed ingly v a lu ab le n o t o n ly b eca u s e t he a u th o r s eem s

2
,

t o c o n s id er t he p hen o m ena in exp lic a b le b u t m o re es p ec ia lly b e ,

c a u s e it es tab lis h es t he f act that t he c lo s es t inbreed ing is n o t

n ec es s arily fatal .

I t s h o u ld b e n o t ed h o wev er that th es e a re exc ept io n al in , ,

s tanc es c o n s t it u t ing n o arg u m ent fo r i nd is c ri m inat e inbr eeding

, ,

b u t th ey d o S h o w that inbreed in g is n o t n ec es sa ri ly h ead ed

s trai ght fo r d is a s t er and with a f u ll h ead o f s t ea m .

T h e breed ing b u s in es s d ea ls n o t with a v era ges b u t with po s

s ib ilities and it is h igh t im e th a t t he f o o lis h h o rr o r Of inbreed
,

in g b e d is s ipat ed I f breed er s had b een a s caref u l in c erta in

.

o th er r es pect s a s th ey ha v e b een t o a v o id t he s light es t f o r m o f

inbreed ing o u r fl o ck s a n d h erd s wo u ld ha v e pro gres s ed farth er

,

alo ng the road o f im pro v em ent .

Ex peri en ce i n an imal b reedin g A n y o n e who will tak e t he .

tro u b le to s t u dy the ped igrees o f fam o u s fa m ilies in almos t any

lin e Of s t o ck breed ing will find that t he f o u ndat io n b lo o d is
mo s t int en s ely bred I nd eed t he pract ica l breed e r wo rk ing wi th
.
,

m at erial that is rea lly o f d is t i nct iv e a n d p ec u liar merit c o mes

soo n t o t he po int at wh ich c lo s e breed ing is i n ev itab le and he ,

m u s t fac e t he is s u e s oo n er o r lat er if he is to m a k e any real u se

o f his v a lu ab le cr eat io n s T o breed th em o u t is b u t t o d is s ipat e
.

th eir exc ellenc e and the o n ly p ract ical c o u r s e is c lo s e breed ing

,
.

A m o ng catt le breed er s th is pract ic e is t o o well kn own t o

n eed m o re than a pas sing m ent io n b u t t he fo llo wing ext ract s ,

fro m pers o nal lett ers rec ent ly rec eiv ed will S h o w ho w it wo rk s
u po n t he h igh ly o rgan ized h o r s e and t he q u ick breed i ng h ea v y -
,

flc s hed s wi ne .

T he v et eran breed er o f A rab h o r s es R and olph Hu nt ingt o n , ,

o f R o ch es t er N ew Y o rk writ es as f o llo ws
, ,

W i th me lo e b eed i g h p c d s t t f
r p i t y a
n d m y b e
ast ro v e a s u re es or ur , n s ,

m t
os i fo m e l t h
un r b e i br su d i g th d m t he
s av e d t he
e n n ree n e a o r so n an o r

D1 wi arC d S l f F t il i t i
n, t p 34
ro s s a n8 ( t li
I mie ) er za on, e c .
, . . a c s a re ne .

I b id p p 3 4 7 3 5
2 . .
-
2 .
626 PRA C TI C A L PR OBLEM S

a n y s tu d
y wh a tev er ha sd ec id ed o tio th t i b eed i ng is d a gero u s I
a n n a n r n .

p re s m ue o u fa t h e r tell s t h i rsi m p ly b ec a s eut h ei fa tsh e S t o ld t h em s o u r rs

an d th ei g rd fa t h e
ra n b efo e th em an d o t o e in m a y t h o u sa n ds h as
rs r , n n n

e e gi e
v r v the m t ter a y t i l
n a r s e io s t h o g h t
n E en with a t ial i t
r a o r u u . v r

d oe o t fo ll w t h t e e y c a
s n o a wi ll b e a s c c e
v r a ym se e t h a n th e m a ti g u s s, n or n

o f im a ls o t rela ted will b e a

an n cc in e e y c e T he a n im als m a ted
su ess v r as .
,

wh et h er kin o ot m st b er n ited to p rod u c e goo d e lts ; t h a t is hav e

, u su r su ,

n o weak es i c o m mo
n s na d as m u c h go o d n,s p o s ib le
n a s .

How t T h ere a re t wo s it u at io n s es pe
t o prac ice in b reedi n g .

c ia lly ind ic a t ing th is m et h o d o f breed i ng O ne is g rad ing in .

,

wh ich it m ay o rd inarily b e pract ic ed with im p u ni ty The o t her .

a rises i n t he v ery b es t h erd s wh en t he breed er fi nd s h im self in

po ss es s io n o f a s m a ll a m o u nt o f v ery s u perio r b loo d and is d ebat
i n g ho w t o h a nd le i t I f he in s is t s u p o n breed ing
.
“
out he
will lo s e it by d is s ipat io n He ha s go n e t o th e lim it s o f lin e
.

breed ing ; what s hall he d o ?

I n a c a s e o f th is k ind t he o n ly c ou r se th a t pro m ises anythi ng
is i nbreed in g I t p u t s t he line t o t he s ev eres t pos s ib le t es t o f
.
,

c o u rs e and t he hazard is grea t b u t t he po s sible res u lt s a re

, ,

ph en o m enal T he rea lly goo d br eed er s h ou ld a lways b e rea dy

.

t o acc ept what ev er hazard is i n v o lv ed .

I f it i s t o b e d o n e at a ll t he b es t way is t o “
d o i t a n d be
,

d on e w i l l: i t and kn o w t he wo r s t a t o nc e M any breed ers

, .
,

fearing t he c o n s eq u enc es go at t h e j ob ginger ly breed ing

, ,

a litt le m o re c lo sely with each s u cc es s iv e trial as if t o t es t the ,

s it u at io n b efo re m ak ing t he b o ld a n d fi n a l s tro k e T h is if n o t .

,

s u cc es s f u l is t o u nd erm in e t he s it u at io n and acc u m u lat e n u m

,

b ers o f u nd es i ra b le ind iv id u a ls ; in any ev ent it c o n s u m es t im e

th a t is v alu a b le fo r a n im als gro w Old q u ick ly
, .

T he pro per way is t o m a k e the b o ld es t s tro k e a t onc e s o that , ,

if t he wo r s t happen s t h e o r igina l s t oc k is left fo r o th er t rials

,

and t he breed er is n o t p o s s es sed Of a h erd that is d es troyed by

u n s u cc es s f u l ha lf h ear t ed att em pt s at i nbr eed ing
,
-
.

SECT I O N V I — B R EED I N G F R O M T H E B EST

T h is ha s referenc e t o the p ract ic e o f s elect ing and breed ing

fro m the b es t ind iv id u a ls b u t with o u t ref erenc e to b lo o d lin es .

I t is pro bable ind eed it is c ert ain th a t in pro c ess o f t ime

, ,
 SY STEM S OF BR E EDI N G 62 7

e xc eed ingly v alu ab le rac es c o u ld b e es tablis h ed in th is way ,

es p ec ia lly o n res t rict ed area s and m o re part ic u lar ly wi th fi eld

cro p s.

B u t in act u al p ract ic e t he breed er fo llo wing th is meth o d

a mo ng an im als s u cc eed s in gett ing togeth er a c o n f u sed j u m ble ,

o u t o f wh ich n o th i ng o f n o t e can b e es tab lis h ed I t is the .

pract ic e fo llo wed by prim it iv e ra c es and by careless fa rm ers ,

and as soo n as so me att ent io n is paid to s trains to fa m ilies , ,

and to bloo d lines it passes at o nc e int o so me o n e o f t he o th er

,

fo rm s o f breed ing already d isc u s sed .

I n p lant breed ing t he pri nc ip le o perat es d ifferent ly Here .

n u m bers m ay b e em p loyed so ext en s iv ely that aft er ha v ing

ch osen t he s t o ck we can lit era lly h u nt thro u gh th o u s and s fo r
t he th ing we want T h is wh en f o u n d is s trict ly s peak ing a
.
, ,

m u tant and ha v ing fo u n d it t he p lant breed er may proc eed a t

,

onc e to m u lti ly i t by c u t ti ng s or to b r eed i t p u r e

p po ss i b ly by ,

inbreed ing c ertain ly wi th as litt le cro s s ing as po s s ib le T h is is

,
.

t he s y s t em f o llo wed by L u th er B u rbank and by p lant breed ers ,

generally who are lo o k ing fo r n ew th ings th o u gh it is o ft en ,

c o m b ined with cross ing .

N eith er in an im al n o r I n p lant breed ing h o wev er a re we to , ,

expect m u ch s u cc ess exc ept by regard i ng a nces tra l li n es and

liv ing and wo rki ng in f u ll realizat io n that the la w o f anc es tral

h ered ity is a fact .

S u mmar y T he s y s t em o f breed ing to b e f o llowed d epend s

.

u po n t h e p u rpo s e t o b e acc o m p lis h ed Grad ing is t he pract ical

.

m eth o d of im pro v ing c o mm o n s t o ck and o f q u ick ly and ch eap ly

gett ing acq u aint ed with t he es s ent ia l charact er s of a breed .

I f t he p u rpo s e is br eed im pr o v em ent thro u gh t he p erf ec t io n

o f fa m ily lin es th en line breed ing and ev en inbreed ing will b e
,

the s y s t em s f o u nd m o s t effect iv e .

I f n ew typ es n ew s t rai n s and n ew creat io n s gen era lly a re

, ,

s u ght t
o ,
w o c o u r s es a re o pe n — ei th er t o watch fo r acc id enta l
,

m u tat io n s o r to ha s t en th eir appearanc e by cro s s ing a f o r m

, ,

o f breed i ng that pro d u c es i nd iv id u als wh ich a re goo d b u t wh ich , ,

u nd er t he c o mm o n la w o f anc es tra l h ered ity a re t oo bad m ix ,

t u r es t o pro d u c e a u n if o rm type and u nd er M en d el s la w a re t oo

’

u n s tab le t o pro d u c e a c o n s t a nt typ e o f any k i nd T he sy s t em .

6 28 PR A C TI C A L PR OB LEMS

of cro s s ing is th erefo re bes t adapt ed to p lant s wh ich can b e

, , ,

pro pagat ed a sex u a lly and th erefo re free fro m t he lim itat io n s
,

j u s t ment io ned .

S PE C I A L E x e R C I S E s

M ake c a l c u la t i o n s s ho wi n g the rela tiv e ex pen se o f gr a d i n g as c o m p a red

wit h b reed in g p u re fo r d iff eren t c las ses o f a n ima ls .

A lso ma k e c ri tic a l s tud y o f m a n y ped ig rees o f fa mo u s a n i ma ls i n o rd er

to t ra c e the s ys tems o f b reed in g a c t u a lly em plo yed , es pec ia lly a s to lin e
b reed ing an d in b reed i n g .

ADDITIONAL REFER ENCES

Lo s s O P V I G O R P RO M I N B R EE D I N G . By H .
J . W eb b er . S c ien c e, 1 90 l ,

No .
3 20, p . 257 .

POL L I N A T I O N OP A PP L E S AND PE A S . E xperi men t S ta tio n R ec o rd ,

X I I I, 62 0 .

R E C I PR OC A L C R O S S ES ( wi th ex ten d ed b ib liog r a p y) h . M a in e S ta t io n
R epo rt , 1 90 4 , pp S I
.
- 8
9 .
6 30 PRAC TI C A L PR O BLEMS

s pec imen s than a re da ms wh ich s h o u ld giv e a h eav y p repo n

,

d eranc e o f males ,
— a fact n o t s u b s ta nt ia t ed .

4 . T h a t s e r v ic e e ar ly i n h ea t w ill pr o d u c e a m a le ( s o m e s a
y a
female) and that s erv ic e lat e in h eat ( with o v u m s ta le) will
,

prod u c e t he o ppo s it e s ex —. Dis pro v ed by t he f act that in

nat u re females es pec ia lly in h erd s a re s erv ed early in h eat
, , ,

a fact that s h o u ld mak e the o ffs pring pra ct ically all o f o n e s ex .

5 . T hat t he o ld e r par e nt w ill d e t er m i n e t h e s ex

,
— s o m e s a
y
the parent neares t t he p rime o f life ; n o t s u b s tant iated .

6 T hat extreme s ex u a l exc it em ent o n t he part o f t he f ema le

.

is a lmo s t c erta in to res u lt in m a le ( so m e say fem a le) o ff s pri ng .

T h is is a d iffi c u lt as s u m pt io n t o pro v e o r to d is pro v e b eca u s e ,

ev eryt hi ng t u rn s u po n wha t wo u ld b e called extrem e exc i t ement ,

and t he s ingu lar fact is that t he believ ers in th is t heo ry th em

selv es app ear q u it e u nab le t o d ec id e wh ich s ex is ind icat ed by

t he v io lent d is t u rbanc es o f the fem ale ; so me s ay o n e s o me say ,

t he o th er u nt il it loo k s lik e a cas e o f t he ind igo t es t o v er agai n

, .

I t is inc o nc eiv ab le that t he g en era l d is t u rbanc e o f t he bo dy

att end ing h eat s h o u ld ha v e t he slight es t infl uenc e u po n the
chara ct er o f t he u n io n o f t he n u c lear m att er of two germ c ells ,

wh ich is all tha t we no w kn o w t o b e in v o lv ed in fert iliza t io n .

I t is no t ic eab le tha t n early ev ery th eo ry o n d et er m inat io n o f

sex c o ntai n s s o me tra c e o f ma le su perio ri ty ,
m any go ing so
far as to s tat e that fema les a re u nd ev elo ped m ales T h is c o n .

c c it is ev id enc ed wherev er an ad v antage is s u ppo sed to exis t as ,

by exc es s o f fert ilizat io n , s u ch ad v antage b ei ng a lway s gi v en

t o t he m a le .

A y n th e o ry n o t i n v o lv i ng o b s c u r e d is t i nct io sn — as d oes thi s

o ne c an b e ea s ily pro v ed o r d is pr o v ed by t he s tat is ti c al

m eth od alway s rem em bering that a c o rrelat io n u p to 5 0 per
,

c ent is in ev itable ind icat ing n o ca u se at wo rk b u t ch a nc e I t

, .

is far m o re pro fitab le t o lea v e s p ec u lat io n and inq u i re wha t is

a ct u a lly kno wn ab o u t t he ca u s es that d et er m in e s ex .

Sex differen ces sl ight . F i rs t o f all t he id ea o f f u nda m en t al

,

s ex d i ff erenc es is great ly e x aggera t ed . A bo u t t he o n ly att ri b u t e

that can b e as cribed to ma leness in gen eral thro u gh o u t the
wh o le range o f life is a litt le h igh er s tat e o f act iv ity u s u a lly b u t ,

n o t a lway s acc o m pan ied by so m ewhat d ecrea s ed s ize .Typ ically

 T HE D ETER M I N ATI ON OF SM 63 1

t he ov u m is large, well
pplied with no u ris h ment and n o t
su ,

i v en to act iv i ty ; wh ile t he s p er m p la sm s perm a tozod n

g o r , ,

po llen gra in is s ma ll and po o r in foo d m at eria l b u t charact er

, , ,

ized by great act iv ity .

A s id e fro m t h is m a les and fem ales d iffer fa r les s than is

,

po pu larly s u ppo sed T he art ific ial c o n v ent io na lit ies and t he es
.

t ab lis hed d i v is io n s o f lab o r exaggerat e d ifferenc es o f s ex in m a n ,

and o v er enth u s ias t ic writ ers ha v e f o rm ed o u t o f thes e exaggera

-

t io n s c o nc lu sio ns a s far reach in g as th ey a re gro tes q u e

-
.

S ex d ifferenc es a re few and s light and m o s t ly c o nn ect ed with,

t he s erio u s b u s in es s o f repro d u ct io n W e n eed n o t th erefo re

.
, ,

in seek ing ca u s es for th eir d et er minat io n loo k fo r s u ch a s s tri k e ,

at the v ery fo u ndat io n of rac ial cha racters Sex is so meth ing .

s u per i m po s ed u p o n a ll o th er c o n s id erat io n s n o t a f u nda m enta l

d iv is io n halv ing the po p u lat io n int o o n e s ect io n that m ay and ,

an o th er that m ay n ot ent er int o the f u ll p os ses s io n o f a ll the

,

end o wm ent s o f t he rac e .

SECTI O N II — I NFLU EN C E OF N U T R I T I ON

In ta d poles . A cc o rd ing to Pflu ger ,

three fo r m s d ev elo p 1

a ) d is t in ct m ales b
( ) d i t i nct f m al and ( ) h r m aphr d it
( s ,
e es c e o, es .

I n the las t ca se t he ma le o rgan s “

d ev elo p ro u nd p rimit iv e
o v aries ,
and if the tadp o les are t o b ec om e m a les the inc lo sed
f emale o rgan s are ab s o rbed .

A cc o rd ing t o Y o u ng s ex in tadpo les rem a in s a lo ng t im e

2
,

ind et er m in at e and d u ring th is t im e t he a m o u nt o f f oo d ex ert s

,

a c o ntrolling infl u enc e u po n the sex He had three broo d s o f .

tadpo les .

B roo d I u nd er nat u ral c o nd it io n s d ev elo ped 5 4 per c ent

, ,

fem a les b u t wh en fed freely with beef i t d ev eloped fem a les in

,

t he pro p o rt io n o f 7 8 per c ent ; the pro po rt io n o f f em a les fro m

brood 2 was increas ed by a gen ero u s d iet o f fis h fro m 6 1 per ,

c ent t o 8 1 per c ent ; an d in the s a me way a d iet o f frogs fl es h ’

ra ised the pro po rt io n in broo d 3 fro m 5 6 per c ent wh en left “

a o ne o 9 pe c ent h en fed
l t 2 r w — a ll o f wh ich loo k s a s th o u gh
,

n u t rit io n ha s so me infl u enc e u po n sex in frogs at leas t .

1 G dd d Th m
e T h E l ti
es an o f S
son , p e
4 5 vo u I
on ob i d ex , . .
2
.
 PRA CT IC A L PR OB L EM S
3 2

In pl an t lice gen eral it m ay b e sa id that in s u mmer

.
l
In ,

wh en fa v o rab le c o nd it io n s o f life a re at t he max im u m th ese ,

c reat u res prod u c e parth en ogenet ica lly generat io n af ter gen era
t io n and o n ly females b u t with the c oo l of a u t u m n an d it s
, ,

les s en ed foo d s u pp ly m ales appear and s ex u al repro d u ctio n is

, ,

res u m ed ; ind eed to q u o t e fro m G edd es and T h o mso n

,
2 “
in ,

t he a rt i fic ia l en v i ro n ment o f a greenh o u s e eq u i v alent to a per ,

p et u a l s u m m e r o f w ar m th and ab u ndant f oo d t he pa rt hen o ,

gen et ic s u cc ess io n o f fema les has been expe rim entally o b serv ed
fo r f o u r y ears I t s eem s in fact to c o nt in u e u nt il lo weri ng o f
.

t he t em p erat u re and d im in u t io n o f fo o d r eintrod u c e m a les and

s ex u a l reprod u ct io n Oth ers ha v e s tat ed t ha t m ales may b e
.

prod u c ed at any t ime m erely by lett ing t he p lant s o n whi c h t he

lic e a re f eed ing bec o me s o mewhat d ried u p .

SECT I ON III — I N F LU E N C E O F F E R T I LI ZATI ON

I n b ees .
3
A s is kno wn bees are of three fo rm s as
no w well ,

regard s sex dro n es ( males ) pro d u c ed fro m u nfert ilized eggs ;

, ,

wo rk er s ( fema les g en era lly b u t n o t alway s s t erile) pro d u c ed

, ,

fro m f ert ilized eggs ; and q u een s ( fert ile fem ales ) also p ro ,

d u c ed fro m f ert ilized egg s b u t q u art ered i n s pec ial c ells and
i v en large a m o u nt s o f s pec ia l f oo d A s r em ark ed by Gedd es
g .

and T h o m s o n ro yal d iet and p lenty o f it d ev elo ps the repro

,

d u c tiv e o rga n s o f t he f u t u re q u een s and at any t im e wi th in "

th e fi r s t eight day s o f lar v a l life t he add i t io n o f a litt le food will

d et er m in e t he s trik ing s tr uct u ral and f u nct io nal d ifferen c e
bet ween wo rk er and q u een 4
T h is fact is o ft en tak en a d v a n
”
.

t age o f by the n u rs e b ees wh en d is a s t er threat en s thro u gh the

lo s s o f the q u een c ells Ha s t ily extract ing s o m e wo rker larv a
.

fro m th ei r o rd inary c ells th ey d epo s it th em in t he q u een c ells , ,

giv ing th em royal f ood and S peed ily th ey acq u ire all t he cha rac
,

ters o f any o th er f ert ile q u een a res u lt p lain ly d u e eit her to ,

t he charact er o r t o t he q u ant i ty o f the f oo d o r t o bo th ,

.

I n t he ca s e o f b ees th erefo re f ert ilizat io n s eems t o b e t he

, ,

d ec id ing facto r as to d ifferenc es b et ween m ale and f em ale an d ,

G dd
1 e d Th m Th E
es a n l t i f S p 49o so n , e vo u on o ex,

I b id p p 4 6—
. .

I b id p 5
2 . . 0
.
48 I b id p 4 7 3
. . .
2 . . .
6 34 PR A CT I C A L PR OB L EM S

SECT I ON IV — SEX I N MA M M A L S

T em peratu re and n u tri t io n S eem to b e c o nt rolling facto rs in

m any o f t he lo wer fo r ms and wh en s u ch is t he cas e it is rem ark ,

ab le that in ev ery in s ta nc e the pro d u ct io n o f females s eem s t o

acc o m pan y t he m o re fa v o rab le c o nd it io n s and that o f males ,

t he hard er o r less fa v o rab le .

A m o ng m a mm a ls th ere is litt le exp erim ent a l ev id enc e b u t ,

that litt le po int s to t he sa me gen eral fact as fo u nd a mo ng lo wer

ani m als th o u gh the larger an imals a re m an ifes t ly les s d irec t ly
,

infl u enc ed by s u rr o u nd ing c o nd it io n s G irou d iv id ed a fl o ck of .

three h u ndred ewes int o t wo eq u al lo t s o n e o f wh ich was ex ,

t r em ely well fed T h is lo t was s erv ed by two y ou ng ra m s ; t he

.

o th er s cant ily fed by t wo m at u r e o n es

,
T he well fed lo t ( serv ed
,
.
-

by yo u ng ram s) pro d u c ed 60 per c ent fem ales the o th er lo t o nly ,

0
4 p e r c ent f e m a les
1
T h e d i ff e r enc e in ag e o f t he ra m s i ntro .

d u c es a s ec o nd elem ent b u t t he fact s if tr u e a re s ign ificant , , ,

.

T h is is ab o u t all that is kn o wn o f th is pha s e o f the q u es t io n .

E xperim ental ev id enc e s eem s to ind icat e that ab u ndant f ood ,

o pt im u m t em p erat u res and gen era lly fa v o ra b le c o nd iti o n s t end ,

t o t he pro d u ct io n o f fem a les ; b u t wh en we c o m e to pred i ct t he

li m it s t o wh ich t he ru le will app ly we m u s t pro c eed with great
cau t io n c o nfes s ing that o u r exact kn owledge is exc eed ingly
,

lim ited .

SECT I ON V — THE “
A C C ESSOR Y C H R OM OSOM E AND

SEX D ET ER M I N AT I ON

C erta in in eq u alit iesb et ween germ c ells in res pect to the d is

t rib u tion o f chro mat i n m att er ha v e lo ng b een kn o wn 3
.

G dd
1 e d Th m Th E l ti
es a n f S p 5o so n , e vo u on o ex , . 1 .

W il
2 S t di
so n Ch m m III j
“
l f E pu i m t
es l on ro o so es ,
”
ou r na o x er en a
—
,

Z ol gy 1 1 1 N
o o , pp 39 A
, l t
o h .f ll w i
I g, B d Th D. t 1 m i t i . f so e o o n : ea r , e e er na on o

S ex ; C tl asTh H e d it y f S B ll ti f tb“
Al m f e C m ere o ex ,
”
u e n o e u se n o o

p t Z
ara i ve ol gy X L 4 ; M C l
o
g ( 9 )
o 0 , T h A y C
, h m m Sc un I 2 ,
“
e c c ess o r ro o so e . ex

D et errn in an t B i ol ogi ca l B u ll eti n , 1 1 1 , I , 2 ; M o rga nS elf Fert iliza t io n -

In d uce d b y A rt ifi c ial M ean s , ”

j ou r n a l o
f E xper i m en ta l Z ool ogy , I , 1 S t u ies in d
S perm a t o gen es is wit h S pec ia R eferen c el t o t h e A c c ess o ry C ro m o s o m e, Pu b ica h l
t io n N o .
3 6 , C a rn egi e I n s t it u t e, W as h in gto n , D C . .
, S ep t em b er, 1 9 5;
0 W ils o n ,
“
Th e
C h ro m o so m es in R e l a tio n t o t he D et erm in a t io n o f S ex in I n s ec t s , S c ien ce, X X I I ,

5 6 4 , O c t o b er, 1 90
5 .
3 W ils o n , T h e C ell, pp . 271- 272 .
 T H E D ET E R M I N AT I O N OF SM 635

Fo r xa mp le W ilso n repo rt s that as lo ng ago as 1 8 9 1 Hen

e ,

king “
d is c o v ered that in t he s econ d s perm at o cyt e d iv isio n of
Pyr rhoc o ris o n e o f t he chro m o s o mes pas s es u nd iv id ed i nt o o n e
o f t he da u ght er c ells s perm at id s ) wh ich r ec eiv es t welv e eb ro
( ,

m ati n elem ent s wh ile it s s is t er rec eiv es b u t elev en ; s o that

”

, ,

o f t he fo u r resu lt ing s per m at o zo a t wo po s s ess an add i t io na l ,

chro m o so me as c o m pared with t he o th er two 1

.

Oth er d is c o v eries were repo rt ed a n d Pa u lmier ( 1 8 98 , ,

wo rk i ng with A na s a i n W ils o n s lab o ra t o ry f ou nd that in t he

’

firs t s perm at ocyte d iv is io n elev en t et rad s appeared o n e of wh ich ,

was m u ch s m a lle r than t he o th ers

“
and s eem ed to aris e from
a s ingle n u c leo lu s lik e b ody and by a pro c es s d iff er ing c o n
-

s id era b ly fro m t he o th er s He add s I n the s ec o nd (and la s t )

.

s pe rm at o cyt e d iv is io n t he larg r dyad d iv id e t o f o rm

( )
t en e s

chro m o s o mes in t he u s u al mann er ; t/ze s m a l l dy a d b owev er , ,

f a i ls to d i v i d e, a s s i ng bod i ly i n to til e erm a tid s

p o v er on e o f s p .

In th is ca se th eref o re half the s pe rm at id s rec eiv e t en s ing le

, ,

chro m osom es wh ile the rem aind er rec eiv e in add it io n a s m all
,

dyad 2
.
”

T he fact was g rad u ally es tab lis h ed that at lea s t in Hem ipt era ,

and in c erta in o th er in s ect s o n e o f t he C hro m at in m as s es o f the

,

m a le m at u rat io n c ell d iffer s fr o m its f ello ws and u nd ergoes o n e ,

les s d iv is io n than th ey so that o f t he gro u p o f f o u r s perm at ozo a

, ,

aris ing o u t o f the d o u ble d iv is io n o f t he s p er m at o cyt e t wo will ,

po s s es s an add it io nal chro m osom e as c o m pared with t he o th er

t wo .T h is add itio na l m em ber has been v ariou s ly na m ed by d if
feren t expe riment ers t he t erm s “
acc esso ry chrom oso m e and
,

h et ero tro p ic chrom o so m e being the m os t c o mm o n ”

.

Here is abo u t wh ere t he m a tt er res t ed t ill M c C lu n g ( 1 90 2 )

ad v anc ed the th eo ry that t he acc es so ry chro m os o m e is the s ex
d eter m i na n t a ss u m i ng ( e rro n eo u s ly as we n o w believ e) that if
,

the o v u m s h ou ld b e f ert ilized by o n e o f t he s p er m at ozo a c o nta in

in g t he acc es s o ry ch ro m o s o m e the o ff s p ring wo u ld th en b e
pro v id ed with t he acc es so ry and its sex wo u ld th erefo re b e
m a le ; wh ile if t he f ert ilizat io n s h o u ld b e by o n e o f t he s p er m a
t o zoa d es t it u t e o f t he acc es s o ry t he o ff s p ri ng wo u ld o f n ec es s ity ,

b e o f the o pp os i t e s ex .

W il 1 Th C ll p s on,
7 e I b
e id p 7, . 2 1 .
2 . . 2 2 .
6 36 PR A C TI C A L PR OB LE MS

T h is v iew o f c ase mad e it appea r that t he fem ale c ells c o r

t he
res po nd m os t c lo s ely with th o s e s per mat ozo a wh ich a re d es t it u t e
o f t he ac c es s o r and h e r e t he m att er s t o o d u nt il M o ntg o m er
y ,
y
Gro s s and Wallac e ( 1 90 5 ) d is c o v ered that in c er ,

ta in s pec ies at lea s t the o v u m has t he s a me n u m ber o f chro m o

,

s o mes as t he s pe r m at o zo a wi th Me a cc es s ory T h ese expe rimen t e rs .

ca me t o t he c o nc lu s io n that o nly o n e o f t he t wo c las s es Of S pe r

mat o zoa is f u nct io na l na m ely that in wh ich t he h et e ro tr o p ic (a c
, ,

c es s o ry) C hr o m o s o m e is pres ent T h o s e o f t he o th er c las s were

.

as s u med t o d egenerat e aft er t he fas h io n Of po lar bod ies

,
1
.
”

Wilso n s h o ws that t he s ex es in h em ipt e r s o f t his

“

type d o in fact S h o w a c o n s tant d ifferenc e in t he n u m ber Of

chro mo so mes ?
He ha s d et er m in ed in at lea s t f o u r gen era
”

, ,

that t he n u m ber of c/zr omos om es i n til e f em a le cell c or r espond s

wi t/c tlze la rg er n u m ber i n tlze m a le c ell ; in o th er wo rd s that t he ,

“
acc es so ry chro mo so me tho u gh pres ent in b u t half t he s per ,

m atozoa is fo u nd in a ll f em a le germ c ells

,
Th is b eing tr u e t he .
,

remarkab le s perm at ozo a a re n o t th o s e wi t]: the acc es so ry ,

b u t on t he c o ntrary th o s e with o u t it and th ey a re to b e

, , ,

rega rd ed as in s o me sen se d efic ient .

Wils o n s h o ws c o nc lu s iv ely t he oppos i te o f Gro s s and Wa llac e s ’ ’

hypo th esis na mely that wh en a female c ell u n it es with a s per

, ,

m a to zo On d es t i t u te o f t he acc es s o ry chro m o s o me th en t he ,

acc es s o ry o f the o v u m fi nd s n o m at e and a m a le d ev elop s ; and

that o n t he o th er hand if t he o v u m happen s to b e fert ilized by
, ,

o n e o f t he s pe r m at o zo a pr o v id ed with an acc ess o ry th en eac h ,

acc es so ry find s its m at e th ere is th en n o s olita ry acc esso ry and

, ,

a fem a le res u lt s .

E xt en d ing his exp er im ent s W ils o n find s t wo k ind s o f acc es ,

s o ry chro m o s o m es t he o n e already d es cri bed wh ich is s maller

, ,

than the o rd ina ry ch ro m o s o m es and an o th er wh ich is larger I n ,

.

th is c o nnect io n it s h o u ld b e rem ark ed t hat his in v es t iga t ion s

s h o w great d iff erenc es i n s ize a m o ng t he chro m oso mes gen era lly ,

b u t that t he acc es so ry can read ily b e d et ect ed wh eth er ,

larg er o r s ma ller than the o th ers and that all C h ro m o s o m es , ,

large or s ma ll exc ept t he acc es s o ry

,
can read ily b e as signed ,

in pai r s u nd er t he m icro s c o pe .

j 1 l of
ou r n a E p i m t lxZ ol
er gy I I I N en a p o I b iod , , o . I , . 2 .
2 .
6 38 PR A CT I C A L PR O B LE M S

I NFL U EN CE OF N U T R IT I O N ON SE X . E x peri m en t S ta t io n R ec ord , X V I ,

228 .

PA R T H E N O G E N E T I C FERTI LI Z A T ION IN T HE HO N E Y B E E . E xper imen t

S ta tio n R ec o rd , X V , 7 92 .

R EC E N T T H E O R I E S I N R E G A R D To D ET E R M I N A T I O N OF S E x . By T .

H . M o rg a n . Po p u la r S c ien c e M o n t l hy , LX I V , 97 1 16 -
.

S E X CO N T R O L . B y Pr o fes s o r S c h en c k o f A u s t r ia S c ien c e, V I I , 7 3 6
.

738
S EX D ET E R M I N A T I O N IN B E ES .
( A d is c u s s io n of t he D zier zo n D i c -
k el
c o n tro v ers y) . By B . S po rr er . E x peri m en t S ta tio n R ec o rd , X I , 5 6 1 ,

SEx D E T E R M I N A T I O N , — W H ET H E R B U D S HA L L BE L E A F O R FL O W E R .

By E . S G o ff
. . A meri c a n G a rd en , 1 90 1 , pp .
3 3 3 , 3 46 3 4 7
33 0 - -
.

S Ex IN M IC E . B y Pa rs o ns and C o pem a n . Pro c eed i n gs o f the R o yal

y
S o c iet , Lo n d o n , L X X I I I , 3 2 4 8 -
.

S Ex I N PL A N T S A M A TT E R O F N T R IT I O N U . By T . M eh a n . R ep o rt ,
D ep a rt m en t of Ag r i c u ltu re, 1 8 98 , pp 5 36 5 4 8 ; E x per imen t S ta ti o n
.
-

R ec o rd , X 1 , 9 1 0 .

W I S C O N S I N E x P E R I M E N T S T A T I O N R E PO R T , 1 900 , pp . 2 66 - 2 8 5 ; 1 90 1 ,

PP -
3 04 —3 1 6 ~

Z I EG L E R ’
S T HE O R Y OP S Ex D ET E R M I N A T I O N . By T . H . Mo rga n .

S c ien c e, XX I I, 8 3 9- 8 4 1 .
 CHA PTER XI X
PLANT B R EEDING

Wh en t he princ iples of breed ing are o nc e u nd ers t oo d th eir ,

app licat io n to s pec ial ca ses either in p lant o r an im al breed ing

, ,

m
is largely a m att er o f c o m o n s en se and n o ext end ed d is c us s io n
,

o f part ic u lar o pera t io n s is n ec ess ary .

I t has a lready b een rem ark ed that t he breed er n eed s t he

u t m o s t po s s ib le fa mi liari ty wi th t he part ic u lar lin e he h o pes t o

i m pro v e T h is fa m iliarity he will get la rgely thro u gh exp erien c e

.
,

b u t he cann o t a ff o rd t o n eglec t any so u rc e o f i nfo m I a tio n that

will enlarge his acq u aintanc e wi th t he breed o r t he v ariety fo r ,

ev ery i t em o f kn o wledg e will c o n s t i t u t e a v a lu ab le a s set in his

bu s in ess wh en the t ime c o mes as it su rely will fo r weigh ing

, ,

s light d iff erenc es in t he ba lanc e in o rd er t o d et er m in e q u es t io n s

o f s elect io n . T h is in v o lv es d etail wh ich o n ly the pract ica l

breed er can acq u ire and u po n wh ich att em pt ed in s t ru ct io n
,

a m ou nt s t o litt le m o re than acad emic d is s ertat io n Certain

~
.

s pec ial fact s and pri nc ip les h o wev er ru n th ro u gh plant breed

, ,

in g as d is t inct fro m an i mal breed i ng and th ese it is well to

, ,

c lear ly u nd ers tand in ad v anc e of act u al o pera t io n s .

SECT I ON I — A D V A NTAG ES A N D L I M I T AT I ON S

A dv an ta ges in plan t b reedi n g T he p lant breed er po s s es s es n o

.

les s than s ix d is t inct ad v antages as c o m pared wi th t he breed er

of an imals
1 L arge n u m bers giv i n g exc ellent o ppo rt u n ity fo r s elect io n
.
, .

2 R ap id repro d u c ing po wers resu lt i ng i n a m a rv elo u s s a v ing

.
,

o f t im e a s c o m par ed wi th that n ec ess ar ily c o n s u m ed in t he

s lo wer pr oc es s o f an im a l br eed i ng .

3 T
. h e relat i v e ch eapn es s o f i nd i v i d u a ls m,
ak ing wh o les a le

d es t ru ct io n ec o n o mically po s s ib le .

63 9
6 4o PR A C TI C A L PR O B L E M S

4. T he gre at e r l ik eli h o o d o f m u tat io n s ar i s ing fr o m m e r e

po int o f n u m bers if fro m n o o th er ca u s e and the grea t er ea s e

, ,

wi th wh ich th es e m ay b e d et ected if th ey d o a ris e .

5. T h e gr eat e r C h anc e o f pr e s e r v ing m u t a t io n s o w i ng t o ,

rap id po wers o f reprod u ct io n .

6 T he po s s ib ility o f r epro d u c ing asex u a lly by b u dd ing c u t

.
,

t ing etc wh ich o v erc o m es t o a large ext ent t he d is as t ers o f

,
.
,

s t erility and a v o id s t he o p era t io n o f M end el s la w in t he p ro pa

’

g a t io n o f hybr i d s .

T he p lant breed er th erefo re n o t o n ly enj oy s s u perio r ad v a n

, ,

tages in s elect io n b u t he is f ree t o m ak e f u ll u s e o f m u tat io n s

,

and t he princ iple of cro ss ing t wo f o r m s o f i m pro v em ent a ll b u t

,

c lo sed t o the an imal breed er N at u rally th erefo re his o perat io n s

.
, ,

as s u m e o n e o f three well d efin ed fo r m s o r s y s t em s o f breed ing :

-

I. S t ra ight s elect io n o r breed ing fro m t he bes t the pu r

, ,

po se being t he im pro v ement o f ex is t ing v ariet ies rath er than the

pro d u ct io n o f n ew s tra in s .

2. M ainta in ing ext en s iv e p lant ing s in t he h o pe o f d et ecti ng

s p o ntan eo u s m u tant s t he o bj ect b ei n g t he pro d u ct io n o f n ew
,

v ariet ies .

3. C r o s s ing o r,
hybr id iz i ng w i th t h e p u r
,
p o s e o f pr od u c in g
n ew s tra in s .

N ew s tra in s pro d u c ed by eith e r t he s ec o n d o r t he th ird meth od

a re o f c o u r s e v a riab le a nd capab le o f im pro v em ent by s tra ight

s elect io n . Each s y s t em o f breed ing req u i res i t s o wn m eth o d s ,

s u it ed t o t he m at e ria l in h a nd a n d t h e C h a r a ct er o f t he im pro v e

m e n t s o u ght S o m e s pec ies d o bes t with o n e s y s t em o th ers

.
,

with an o th er and o n ly exper ienc e can d ec id e wh ich is m o s t

,

pro lific o f res u lt s in a part ic u la r cas e T he fir s t s tra ight selec .

,

t io n is the safes t and is a lway s c ertain o f res u lt s ; b u t mo s t

, ,

s p ec ies r es p o nd well t o t he S ec o nd and t he th i rd wh ich wi th , ,

s u itab le m at er ia l a re capab le o f t h e rich es t res u l t s kn o wn t o all

,

breed ing and a re the sy s t ems pa r ex cellenc e fo r t he pro d u ct io n

,

o f n ew s tr a in s .

C ro s s ing ha s la tt erly fa llen i nt o so m e d is rep u t e beca u s e o f

t h e em pha s is la id o n M end el s la w and t h e pri nc ip le o f m u tat io n
’

is b u t rec ent ly rec o g n ized ; b u t t he pred ict io n is v ent u red that

t he fo r m er will b e res t o red to fav o r i n p lant breed in g and that ,
64 2 PRA CT I C A L PR O B L EM S

st ra in will pro s per ev en b ett er in t he hand s o f t he fa rmer an d ,

th ereby s t and m o re ch a nc es o f p lea s in g when p u t to t he ac t u a l

t es t o n t he fa rm o r in t he o rchard S pec io u s b u t fa u lty is t he .
,

o n ly c o rrect v erd ict a s t t h is po s i t io n th o u g h it m a b e m a i

o
y, n

ta i n ed p e rh,
a ps a s t o so
,
m e s pe c ia l s t rai n s p a rt ic u la r ly s e ns i
t iv e t o h igh fert ility I m pro v ement is what is aimed at by t he
.

breed er — t hc pro d u c t io n o f be tt e r s t rai ns than befo re

, On .

th is there a re t wo S igni fic a nt po int s

I. T he breed e r wi ll n o t kn o w wh en he ha s s u cc eed ed in p r o
d u c in g an i m pro v em ent u n less t he s o il c o nd it io n s a re g oo d
en o u gh t o pe rm it f u ll and c o m p let e d ev elo p m ent .

2. A ll ex perien c e g oes t o S h o w that p lant s a re m o re v a riab le

i n s o ils o f h igh f e rt ility tha n i n s o ils o f lo w fert ili ty T h is is t h e .

expe rienc e o f D e V ries o f D arw in and so far as is k n o wn to t he

, , ,

writ er o f ev ery p lant breed er u po n rec o rd

,
.

T he o bj ect in a ll p lant b reed ing is t he pro d u c t io n Of i mpro v e

m en t T h is is pa rt ly d epend ent u p o n f ert ili ty and i n gen eral
.
, , ,

p lant breed ing o pera t io n s will b e mo s t s u cc es s fu l o n land s o f

-

m ax im u m fe rt ility S o m e acc lim at iza t io n m ay aft erward b e

.

nec es s a ry as to s o il as well as to C lim at e b u t t he la tt er is ,

in v o lv ed in a ll p lant breed ing — and the f o rm er too fo r th a t

, ,

m att er — fo r n o s o il can b e fa ir ly repres enta t iv e o f an y v ery

,

great ext ent o f t errit o ry .

Th e b al an ce Of ferti li ty M u ch rem ain s to b e lea rn ed as t o

.

t he elem ent s that s h o u ld pred o m inat e in a fert ile so il In .

genera l n itro g en fav o rs t he growth o f leaf and s t em b u t th ere

, ,

is m u ch reas o n t o b eliev e th a t s eed fo rm at io n is int im ately

relat ed to t he s u pp ly o f ph o s ph o r u s T he bo tan is t will t ell u s .

that S apro legn ia fo r exa m ple gro ws lu x u ria nt ly in beef extra c t

, ,

o r pept o n e b u t pr o d u c es n o repro d u ct iv e o rga n s ; gro wn in

,

n u trien t s o lu t io n s c o nta in in g ab u nd a nc e o f ph o s ph o ru s ho w ,

ev er repro d u ct iv e o rgan s f o rm r ead ily es pec ia lly in t he female

, , .

With o u t d o u bt m u ch rem ain s t o b e lear n ed in t he matter of

m ak ing u p o f s o ils m o s t fa v o rab le fo r t he pro d u ct io n o f d es i ra b le
v ariati o n s i n d i ff erent c la s s es o f p lant s I n th is m att er o f s o il
.

fert ility and c u lt u ral req u i rem ent s let t he p la n t breed er pro v i d e
m ax imu m c o nd it io n s with n o f ea r o f ev il c o n seq u enc es
,
I f he .

c a n s u c c eed in pro d u c i ng a d es i rab le v ar iety by b oo k o r by

 PLA NT BR EED I NG 64 3

croo k acc lim at izat io n will c o m e to his a id and h elp him to

,

preserv e it .

Of o n e fact we m ay b e well as s u red na m ely that ev ery plant

, ,

is at it s bes t u nd er op t im u m c o nd i t io n s T hat is t he t im e when

.

fav o rab le v ariat io n s m ay b e mo s t c o nfid ent ly expec ted in o ther

wo rd s that is t he t im e wh en i t will res po nd m o s t fa v o ra b ly t o
,

s elect io n T he wri t er believ es th is to b e t he gen eral princ iple

.

f ro m wh ich t o wo rk o u t t he c o nd it io n s u nd er which t he pa rtic u lar

s tr ai n o r s t ra in s y ield bes t res u lt s b u t t he p lant breed er m u s t
,

n o t d ec eiv e h i m s elf int o th ink ing that he will get v a lu ab le d ev i

a t io n s fro m type wh ile t he p lant i s end u ri ng hard c o nd it io n s t he ,

eff ect o f wh ich is t o br i ng ev erythi ng t o t he d ead lev el o f med i

oc ri t y wh e re li tt le im pro v em ent is po ss ib le by breed ing wi th ou t

,

fi rs t impro v ing t he c o nd it io n s o f gro wth .

SECT I O N I I I Y
S ST EM S OF PLA NT I N G

In general three sy s t em s of p lant ing a re in u s e a m o ng p lant

,

b reed er s
I
. T he n u rs ery s y s t em in wh ich t he p lant s a re t reat ed a s
,

ind iv id u a ls each being g iv en ab u ndanc e o f roo m an d each m ad e

, ,

t he ba s is o f s elect io n at t he c lo s e o f t he sea so n .

2. T he fi eld s y s t em i n wh ich ind iv id u a l p lant s a re not giv en

,

s pec ia l o pp o rt u n i t ies .S eed is s a v ed fro m t he bes t p lant s b u t ,

n o att em pt is m ad e t o id ent ify and i so lat e part ic u lar parentag e .

T h is is im pro v ement rath er than breed ing and is in c o m mo n

“
,

u s e a m o ng g en era l far m er s .

3. What W e bb er ca lls t h e B u rbank m e th o d o f cr o wd ing

th ou s and s of s eed lings c lo s e t ogeth er o n goo d so il and then
s elect ing t he few that a re ab le t o end u r e t h e batt le and s u rv iv e .

Ea ch s y s t em has it s ad v antages es pec ia lly t he fi rs t and t h i rd

, ,

b u t t he t h ird m erges int o t he fir s t t he m o m ent that rea lly c lo se

wo rk b egin s .

A ll th ing s c o n s id ered it is alt o geth er lik ely that t he great er

,

part of o u r res u lt s will b e o btain ed by the so called n u r sery -

m eth o d . I n any ev ent t h is is t he m eth od that lend s it s elf to

t he b es t grad e o f wo rk and to t he m os t c o m p let e r ec o rd s I t is .

t he o n e th eref o re that will rec eiv e f u rth er c o n sid erat io n h ere .

6 44 PR A C TIC A L PR OB LEM S

Pl ot or row in t h e n u rser y s yst em . S hall t he i nd iv id u als o f a

si ng le s elect io n b e plant ed t o geth er in recta ngles o r gro wn in
s epa rat e r o ws ? Ea ch s y s t em has it s ad v ocat es and t he m att er ,

is t o b e d ec id ed fi rs t o f all by c o n v en ien c e i n c u lt iv at io n and

harv es t in g and sec o nd by c o n v en ienc e in k eeping rec o rd s The
,
.

t wo s y s t em s a re well illu s t rat ed by t he m eth o d s em p lo yed at

t he t wo experim ent s ta t io n s o f M inn es o ta and Illino is .

Th e pl ot syst em T h is s y s t em is bes t d es c rib ed by i t s u s e a t

.

t he M inneso ta s t at io n wh ere i t ha s been f u lly elab o ra t ed a n d is

,

c o ns tant ly emplo y ed for all breed ing wo rk U nd er th is sy s tem

1
.

as th ere em p loyed t he pro c ed u re is s u b s tant ia lly as fo llo ws ,

ta k ing wh ea t as an illu s tra t io n Wh en a n ew v ariety is rec eiv ed

.

o r a p ro m is ing p lant is select ed i ts h is t o ry is rec o rd ed o n a

rec o rd s h eet 5 5 x 8 5 inch es and p u nch ed at t he en d fo r fi ling

. .

I t is giv en a c lass na me and a n u m ber o f i t s o wn N u rs ery

S to ck N o . and if it s pra n g fro m a n u m bered s t ock that ,

n u m ber is a lso rec o rd ed as Parent S to ck N O T h is s h eet .

is kn o wn as the I ntro d u ct o ry S h eet

“
and is repro d u c ed h ere ”

o n a r ed u c ed s ca le .

Form 6 1 S EL EC TED S TOC K —INTR OD U CTOR Y S HEET

C lass Name of Mi nn No
WHEAT Parent S tock
. .

Par ent S tock

of

Q Q Q Q Q Q Q Q Q Q Q Q Q Q Q Q Q Q

Ori gi n an d His tory of Paren t S toc k

The d is th en p lant ed by it s elf in a rectangu lar p lo t ten

s ee ,

p la n ts s q u are if p o s s ib le — h enc e kn o wn as a “
, c entg en e r
p lo t N o tes a re t ak en bo th o n t he c en tgen er p lo t as a whole
.

and o n ind iv id u al plant s and rec o rd ed o n S h eet s (s ee tab le o n

,

o ppo s it e pag e red u c ed s ize)

,
.

1 S ee B u l leti n A o 62 , M in n es o t a A gric u lt u ra l E xperim en t S ta t io n , fo r f u ll

'

. a

d es c rip t io n o f t he
p o t l y
s s t em as us e d
a t t is s t a t io n h .
64 6 PRA C TI CA L PR OB LEMS

M an ifes t ly all the p lant s

o n th is c entgen er p lo t ha v e

t he s a m e c entg en er n u m
b er,
b u t any pro m is ing
p lant in t he p lo t may b e
g iv en a n u r s ery n u m ber ”

at any t ime so that after

,

the s y s t em is well s t a rt ed
ev e ry p lant ha s t wo n u m

bers it s c entgener n u mber

,

and its n u rs ery n u m ber .

F ro m th ese o ri g ina l rec o rd s

three o th er s h eet s a re mad e
u p a s a k ind o f ledg er ac

c o u nt with th is part ic u lar

s e l e c t i o n ext end i ng per
,

hap s o v er sev era l y ear s .

S ee fo r m s 6 5 6 7 and 6 8
, ,
.

T he s y s t em s eem s so m e
wh a t c o m p l i c a t e d t o a
no v ic e b u t it is eas ily and
,

q u ick ly m as t ered by th o s e
engag ed i n t he wo rk a n d it ,

is app lied t o a ll f o rm s o f
b reed ing carried o n at the
M inn es o ta s t a t io n .

Th e row syst em T h is .

s y s t em is b es t illu s trat ed

by its u s e a t the U n iv er
s it o f I llin o is wh ere i t is
y ,

em p lo y ed n o t o n ly fo r c o rn

b reed ing b u t a lso fo r wh eat ,

o a t s and o th er p lant s
, T he .

s a m e s y s t e m h a s b een

ad o pt ed b y t h e I l l i n o i s
S eed C o rn B reed er s A S S O
’

c ia t io n and is in u s e by a ll
,

it s m em b ers .
64 8 PR A CT I C A L PR OB L EM S

U nd er th is s ys tem ea ch ea r Of c o rn fo r exa m ple is plant ed , ,

in a s eparat e ro w and as m any ro ws will b e u s ed as th ere a re

,

ear s in t he fir s t s elect io n T he ea r in t he boo k a n d t he ro w i n

.

t he fi eld th en ha v e t he s a m e n u m ber .

S u ppo se we a re s tart ing an experiment o n breed ing fo r h igh

o il I t is t he fi rs t y ear and we ha v e t he t wenty h igh es t Oil
”
.
,

ear s that c o u ld b e f o u nd o u t o f t he s eed at hand T h ese ea rs .

will b e n u m bered n o t fro m I t o 2 0 b u t fro m 1 0 1 t o 1 2 0

, ,
.

N ext y ear th ey will b e n u m b ered 2 0 1 to 2 2 0 o r 2 2 5 o r t o wh a t , ,

ev er n u m b er o f c ar s ma y b e a v a ilab le T he h u ndred s a lway s .

S h o w t he n u m b e r o f y ear s o r gen erat io n s o f i m pro v em ent and ,

t he res t o f t he n u m b e r s h o ws t he fi eld n u m be r o f t he ea r and o f

t he ro w in wh ich it is p la nt ed T h u s if we find c ar N o 6 1 2
.
,
.
,

we kn o w that it is t he S ixth y ear o f t he expe rim en t o r t he ,

s i xth s elect io n fro m t he o r ig i n a l s t o ck a n d th a t i t is p lant ed in ,

1 2 o f that y ear s fi eld T h is c a r lik e a ll o t h ers that

’
ro w N o . .
,

ha v e been a na ly zed has it s la bo rat o ry n u m ber o r an n u al ear

, ,

n u m ber by wh ich it s c o m po s it io n m ay b e t rac ed ; b u t it s

,

“
pe d igr ee n u m b er 6 1 2 f o r e x,
a m p le i s,
t h e o n e fro m wh ich ,

it s breed ing is tra c ed A s a m ple p a ge fro m s u ch a regis ter

.

boo k s h o ws t he s ys t em in f u ll th ere being n o o th er rec o rd s ,

exc ept t he ch em ica l ana ly s es T h is is t he ent i re rec o rd o f t he

.

h igh o il c o rn fo r 1 90 2 ( S ee tab le o n o ppo s it e page )

-
. .

S elect ing 6 0 7 o f th is t a b le fo r exa m p le we s ee by t he rec o rd, ,

th a t it s d a m o f the y ear bef o re was N o 5 04 ; that it s n u m ber .

i n t he ch em ic a l la b o ra t o ry wa s 3 9 2 3 ; that t he c a r was
inch es lo ng it s t ip c i rc u m fe renc e
,
in ch es and i t s b u tt c i r ,

c u m feren c e inch es ; th a t it had 1 2 ro ws and that each ro w ,

ha d an a v era ge o f 4 8 k ern els ; th a t t he ea r weigh ed o u nc es ,

and had p e r c e nt o f o il W e lea rn t o o that

. i t wa s p lant ed , ,

in ro w N o 7 wh ic h wa s .
,
h ills lo ng and pro d u c ed 8 5 po u n d s
o f c o rn o r at t he rat e o f
,
b u s hels per acre ; that there
were in all 1 4 0 ea rs o f c o rn in the ro w t he a v erag e Oil c o nt ent ,

o f wh ich wa s

A S a pract ical d et a il in c o rn breed ing it m a y he remark ed ,

t h a t t he b es t ea rs a re a lway s p la nt ed i n t he m idd le ro ws t o g iv e

t h em t he ad v ant a ge in t he m att er o f p o llin at io n .

A d t mi d b y
1 s e er mp it
ne mpl f tw t y
a co osg e sa e o en a v era e ears .
650 PR A CT I C A L PR OB LEM S

As bet ween t he t wo s y s t em s
nd iv id u a l m u s t tak e hi s t he i
cho ic e T he ro w s y s t em s eem s to hav e t he ad v antage o f S im
.

lic ity es p ec ially fo r p lant s s tand i ng in c u lt iv at ed ro ws I t i s

p , .

t he o n e mo s t r ead ily u nd er s t oo d and m o s t ea s ily m a n aged b y

t he far m er b u t eith e r is ea s y o f app lic a t io n
, .

Th e perf orm an ce record On e o f t he s u rpris es o f plant breed

.

in g is t he v ery d iff erent app eara nc e Of t he pro geny fro m eq u a lly

pro m is ing ind iv id u al ears h ead s o r o th er select io n A d iff er , , .

enc e Of t wo t o o n e is n o t at a ll u nc o m m o n and n o t i nf req u ent ly ,

a ro w o r c entgener p lo t fro m pro mis ing s eed pro v es almos t

wo rt h less Henc e t he n ec ess ity o f acc u ra t e rec o rd s o f ent ire
.

ro ws and c entgen er p lo t s Th ere is litt le u se in wa s t ing t ime

l
.

o n i n f e r io r m at eria l fo r t he b es t i nd iv id u a l p lant fro m s u ch a

,

frat ern ity wo u ld b e b u t p o o r s to ck fo r breed ing p u rpo ses A ll .

i nd iv id u a l s elect io n s fo r f u t u re p lant ing th en s h o u ld b e m ad e , ,

fro m ro ws (o r c entgener p lo t s ) with a h igh av erage perfo rm

anc e rec o rd .

M u l t ipl yin g plot s an d fi elds A ft er a n ew s tra in has pro v ed .

s at is fact o ry i n t he n u r s ery ro w o r p lo t it m u s t n eed s b e mu l ,

t ip lied in o rd er t o s ec u re a s u ffi c ient q u ant ity fo r s a le I t is .

u s u ally c u s t o m ary t o s elect fo r at lea s t three g en era t io n s a s i n ,

t he prod u ct io n o f s u ga r b eet s eed th en m u lt ip ly by p lant ing

-
,

o u t in t he o p en fi eld and s elling t he t o tal cro p fo r s eed Th u s .

t he beet s eed o f t he m ark et is gen era lly t wo rem o v es fro m

t he m o th er b eet wh o s e s elect io n wa s m ad e by s u gar c o nt ent
, .

S eed produ ct i on a b u s in ess I t is fa s h io nable t o ad v ise t he .

fa r m er t o pro d u c e his o wn s eed F ro m the s tandpo int Of .

acc limat izat io n it is goo d ad v ic e b u t fro m any o th er s tandpo int ,

it is bad c o u n s el pro v id ing o f c o u r s e th a t p lant breed ers and

, , ,

s eed s men liv e u p t o th eir r es p o n s ib ilit ies .

I t is u nth inkab le that t he far m er who is p rim a rily engaged in

pro d u ct io n can a lso b e a s k illed im pro v er in a ll that he pro d u c es .

He m ay breed o n e o r t wo lin es o f an i mals o r p la n t s h im self ,

a n d if h e b e a breed er by nat u re and t ra in i ng and if he hav e ,

t he leis u re that is well ; b u t by t he s a m e t o k en he sh o u ld

,

1 Th e ro w of th e ro y d
w s s t em c o rres p o n s o f c o u rs e, t o , th e c en tgen er p lo t
of the p l ot s y s t em , an d b o t h l d
in c u e t he en t ire p ro gen y of an
“
in i v id d u al
”
s elec t io n .
 PLA N T BR EE D I N G 65 I

e xpect s o me o th er s pec ialis t to p rov id e him with his fou ndat io n

s t oc k in o th er lin es N o m a n liv ing o r that will ev er b e bo rn
.
, ,

will s u cc eed in breed ing all lin es o f an im als and p lant s exc ept ,

t o th eir c o nf u s io n and gen era l d egra dat io n .

S O wh ile it is p o p u lar t o ad v is e t he farm er to pro d u c e his

o wn s eed s it is b ett er b u s i n ess p o licy fo r h im t o b u and s p end
, y ,

a litt le t im e and pat ienc e in acc lim at ing t he s tra in s if n eed b e

fo r a y ear o r t wo befo re p u tt in g the n ew s t o ck i nt o c o m m o n u s e
, .

T h is will genera lly s u flic e and if in a s p ec ia l cas e a s p erhap s

‘
, , ,

in c o rn n o im pro v ed v a riety will s u cc ess f u lly acc lim at e th en

, ,

th ere is no th ing to b e d o ne b u t to s et abo u t t he pro d u ct io n o f

an im pro v ed s tra in o f the lo ca l v ariety b u t th is can b e aec o m
p lis h ed by a fe w p e r s o n s o r ev e n by o n e p e rs o n as well as it

wo u ld b e d o n e if ev ery n eighbo r u nd ert oo k t he tas k .

I m pro v em ent ev en in p lant s is c o s t ly b u s ines s and wh en

, , ,

o n c e i t is eff ect ed a r ea l i n v es t m ent ha s b een m ad e that can b e

m u lt ip lied as o ft en a s m en and land s a re a v a ilab le .

The int eres t s o f agr ic u lt u re d ema n d that so m e m en g iv e th ei r

t ime and gen iu s to t he im pro v ement o f anim als and p lant s .

T h eir n u m ber h o wev er will n ev er b e relat iv ely la rge and it

, , ,

n eed n o t b e I t is exped ient t ha t a ll farm ers addres s th em selv es

.

t o t he s erio u s b u s in es s o f learn ing ho w t o h and le and pro d u c e

o n a c o mmerc ial s ca le t he r ea lly exc ellent cr eat io n s i n p lant and

an imal life that t he geniu s o f breed ers is ab le t o o riginat e .

A DDITI ON A L REF EREN CES

A B ST R A C T OF PA PE R S R E A D A T T HE N EW YOR K C ON FE R EN CE OF PL A N T
AN D A NI MA L B R EED E R S S
( pe tem b er 3 0 O c t o b e r -
2, E x peri
m en t S ta tio n R ec o rd , X I V , 2 0 8 —2 22 .

B I B L I O G R A P HY .
( A referen c e to fo rt y eigh t
-
a r ti c les on p la n t b reed i n
g .
)
E x pe ri men t S ta tio n R ec o rd , X V , 7 7 0 ; a ls o i n E x peri m en t S t a tio n
h y
R ec o rd , X V I , 3 5 4 , t ir t o n e a rti c les -
.

BR EEDI NG A NI MA LS AND PL A N T S . By W M Ha y s
. . .
’
B reed ers G a zet te,
X L I , 8 92 , 94 4 .

B R E ED ING COR N By C . . P Har tle

. y . Y ea r B oo k , U n i ted S ta tes D ep a r t
men t o f A gri c u ltu re, 1 90 2 , p 5 39. .

B R E E D I N G C O T T ON . By H .
J . W eb b er . Y ea r B oo k , U n i ted S ta tes D ep ar t
m en t of A g ri c u ltu re, 1 90 2 , p 3 5
. 6 .

B R EED I N G F OR EA R LI N E S S . E x peri m en t S ta tio n R ec o rd , X , 3 5 2 .

65 2 PRA C TI C A L PR OB LE M S

B R E E D I N G PE A N UTS . N o rt h C a ro li n a E x peri me n t S ta t io n , B u lleti n N O .

168, pp .
4 2 1 -
4 34 ; a ls o i n E x p e ri m en t S ta tio n R ec o rd , X I , 1 0 3 2 .

B R EEDI NG W HE A T . By W illia m Saun de rs . A g r i c u lt u r a l S c ien c e, 1 8 99,

74 8 7 ;
-
i n E x pe r i m en t S ta tio n R ec o rd , X I I , 3 3 9
als o .

B R EEDING W O R K O P T H E M I N N E S O T A E x PE R I M E N T S T A T I O N . By
W M Ha y s
. . . B reed e rs
’
G a zette, X L I V , 1 187 .

C O F F E E HY B R I D . G a rd en er
’
s C h ro n ic le, 1 8 99 , p . 2 40 .

C o oPE R A T I v E B R E E D I N G . By W M . Ha y s B reed ers G azet te X L V , I 4

. .
’
, .

C R OS S , M A I Z E T E OS I N T E -
. By J . W Ha rs h b e rg er G a rd e n a n d F o res t
. .
,

1 8 96 , p .
5 2 2 ; a lsoi n E x pe ri m en t S ta tio n R ec o rd , V I I I , 5 6 3 .

C R OS S B R E E D I N G O F FR I IT S S u m mar f s e ri es o f e x pe ri m e n ts c o v e r
(
'
-
y o .

in g a n u mb e r of y
ea rs ) By L B ud d . I o wa Ho rt ic u lt u ra l S o c ie ty,
J . . .

1 900 , pp 7
. 8 ; al
1 s 7
o i
6 n
- E x pe r i m1e n t S t a t io n R ec o rd , X I I I, 4 54 .

C R OSS I NG O F PE A S , B E A N S , E TC E x per im en t S ta t io n R ec o rd , X V I , 2 6 3 . .

C R O S S I NG S T R A wB E R R I E S B y F W C a rd a n d G E A d a m s . R od e . . . . . h
E x per im en t S ta tio n R epo r t,
I s la n d 1 90 0 , pp . 247 —2 6 7 ; a l so i n E x pe ri
m en t S ta tio n R ec o rd , X I I , 94 4 .

C R O S S I N G V A R I ET I E S . By B . I) . Ha ls ted . N ew J erse y E x perimen t S t a

t io n R epo r t , 1 90 1 , 8 —
pp 3 9 4 1 . 1 ; a ls o in E x pe ri m en t S t a tio n R ec o rd ,
X I V , 56 8 .

D I F FE R ENC E IN PL A N T AND A NIMAL B R E E D I NG . By W . M . Ha ys .

B ree d ers
’
G a zet te, X L I V , 1 1 32 .

E F F EC T OP S OI L ON D E V E LOP M E N T . E x per im en t S ta tio n R ec o rd , X V I , 22 .

E x PE R I M E NT s IN PL A N T B R EE D I N G O N T HE D O M I N I O N E x PE R I M E N T A L
FA R MS . By W illia m S a u n d e rs . T ra n sa c ti o n s o f the R o yal S oc iety
of C a n ad a , 1 90 2 , p . 1 1 5 .

E X PE R I M E N T S U T H E R B U R B A N R B y D id S t rr J o d a
OF L . av a r n . Po p u lar
S c ien ce M t h ly L X V I 2 1 2 2 5
on , , 0 -
.

G ER MA N M ET HO D O P B R E E D I N G S U G A R B E ET S E pe ime . x r nt S ta t io n
R ec o rd , X I I I , 6 4 2 9 4 8 - .

K
HY B R I D , B L A C B E R R Y R A S P B E R R Y -
. E x per im en t S t a t io n R ec o rd , V I I ,

HY B R I D C O R N . By C . P . Ha r t le y . Y ea r B oo k , U n i ted S ta tes D ep a rt m en t
of A g r ic u ltu re , pp 5 9 5 5
3 0 1 90 2 , -
. .

HY B R I D TO MATO ES E x p er i m en t S t a tio n R ec o rd , X I I I , 3 4 8
. .

HY B R I D I Z ATIO N ( L i s ts o f b.rid s a n d g en e r a l la ws o f ered i t hy h y .

) By
F A
. . W a u gh , A m e r ic a n G a r d en , 1 8 99 , No
34, p 43 1
. 2 . .

HY B R I D I Z A T I O N . Pa pers b y B a tes o n , D e V ries , B a iley, a nd Web b er .

S c ien c e, X , 1 1 3 -
1 16 .

HY B R I D I Z A T ION IN B EA NS . By R . A . E m ers o n . N eb ras ka E x pe r imen t

S ta tio n R epo rt , 1 90 3 , pp 3 3 6 8 .
-
a ls o i n E x peri m en t S ta t io n R ec o rd ,
X V I , 56 3 5 64 -
.

HY B R I D I Z A T I O N OP B A EL EV, W HE A T , OA T S , AND FR U IT S . By W il liam

S a u n d ers . T ra n s a c tio n s of th e Ro y al S o c iety of C an ad a, 1 894
,

pp . 1 39 -
142 ; a ls o i n E xp er imen t S t a tio n R ec o rd , V I I , 2 73 -
2 75 .
 CH A PTER XX
AIm IAL BR EEDING

SEC TI O N I A D V A NT AG ES A ND D I SA D V A NTAG ES

Adv antages . A n i ma l breed ing po ss es ses three s u b s tan t ial

ad v antages o v er p la n t breed ing
I
. M o re freed o m fro m c lim at ic and o th er loca l c o n d it io n s S O ,

that the prod u ct is fitt ed fo r a wid er range o f s erv ic e .

2. R elat iv ely go o d pric es beca u s e t he in d i v id u a l has a h ig h

,

v a lu e a n d is n o t S O eas ily o r s o rap id ly m u lt ip lied .

3
. I n v o lv ing s u p er io r beings an im a l breed ing bec o m es o n e O f
,

t he h igh es t f o rm s Of art I n th is we a re d ea lin g n o t o nly wi t h

.

Su p erb phy s ica l fo rm b u t with m enta l q u a lit ies as well and h e re

, ,

t he breed er may c o m e i nt o s y m path et ic pe rso na l relat io n s wi th t h e

pro d u ct s o f his h a nd and o f his gen iu s T he s a me s y m pa th et ic .

relat io n will b e c laimed by t he lo v er o f plant s and es pec ially by ,

t he lo v er o f fl o wers b u t t he f a ct rem ai n s that o n ly wi th a n im al s

d o we fi nd c o n s c io u s n es s a n d int elligent res po n se to o u r m oo d s
and pas s io n s What f eeling o n eart h o u t s id e of h u man a fi ec t io n
.
, ,

can appro ach t he attach m ent ex is t ing bet ween a man an d his
h o rse o r bet ween a d o g and his mas t er ?
,

Disadv an tages B u t t he an ima l breed er m u s t fac e a lo ng li ne

.

o f li m itat io n s and d is ad v antag es S o m e o f th es e can b e eas ily

.

s ing led o u t and s tat ed o th er s a re t o o s u bt le fo r p u tt i ng in t o

,

wo rd s :
1
. N u m b ers a re n ec es s arily few a n d reprod u ct io n is relat iv ely
,

s lo w m ak ing s elect io n d iflic u lt fr o m m ere s carc ity o f m at eri a l

‘
.
,

2
. I nd iv id u a ls a re c o s t ly and th o s e o f h igh b reed i ng po wers
,

extrem ely s o . T h is m ak es rea lly go o d br eed ing n o t o n ly ex

pen s iv e b u t in a m eas u re haza rd o u s fo r pric es hav e a way ,

of tak ing a s u dd en d ro p o ften with litt le o r n o warn ing

,

wh en n o th ing b ett er than t he o pen m ark et affo rd s an o u t let

fo r s u rp lu s s t o ck .
 A N I MA L BR EED I N G
’
65 5

3. c
T he h aract e r s t o b e s e le ct e d and br e d fo r a re genera lly

n o t few b u t m any and d i ffi c u lt ies in select io n n ec es s arily i ncrea se

,

o u t o f a ll pro p o rt io n t o t he n u m b er o f p o i nt s to b e atta in ed .

4 . A S if t h e br e e d e r did n o t ha v e tr o u b les e n o u gh o f h i s o w n ,

fas h io n is c o nt in u ally add ing po int s that d emand his attent io n

mo s t im perio u s ly ev en at t he expen s e o f b ett er th ings
,
.

5. A n i m a l s pr o pagat e b u t s lo w ly and ,
br ee d i ng o pe r at io n s

nec es sarily extend o v er m any years and sev eral generat io ns the
po p u lat io n cann o t th erefo re b e s pread o u t to v iew and th ere is
, , ,

m o re o r les s u nc ert ai nty as t o act u al fa m ily h is t o ry and i nd iv id

u al m e ri t — ,
a ll Of wh ich m ak es select io n m o re o r les s d iffi c u lt

and u nc ertain .

6 T he y o u ng of m o s t an im a ls are pro m is ing b u t s elect io n

.
,

canno t s afely b e m ad e at extrem e immat u ri ty fo r t he d iff er ,

en c es bet ween in f erio rity and s u p erio r ity a re bro u ght o u t o n ly in

t he d ev elo p m ent that c o mes wi th f u ll m at u rity TIce ex c ellenc e of

.

b r eed ing is m a i n ly s /zown in Me capa c i ty f or d ev elop m en t and ,

th is canno t b e fo reto ld exc ept as it may b e pred ict ed by a

genera l kn o wledge o f t he part ic u lar fam ily line and t he s pe
c ial b loo d c o m b inat io n i n v o lv ed S o me o f t he mo s t pro m is ing
.

“
y ou ng th ings a re t he bitteres t d is appoint m ent .

7 . A n i m a ls a re d iffi c u lt O f d e v e lo p m ent and ,

m any o f t h e

bes t bred th ings a re nev er pro perly d ev eloped

-
.

8 A n im a ls d o n o t rep rod u c e a sex u a lly and th eir s u cc es s f u l

.
,

pro d u ct io n is c o nd it io ned u po n h igh s ex u al fert ility N o w im per .

,

fec t sex u a l d ev elo p ment is o n e Of t he m o s t c o m m o n if no t t/ze ,

m os t com m on d efect in b o th plant s and an im a ls

,
Plant s m ay b e
.

pro pagated by b u d s o r c u tt ing s b u t an im als a re at the m ercy Of

,

s ex u a l r epro d u ct io n I n nat u re t he ex is t in g lin es a re k ept at

.

lea s t fa i rly f ert ile by nat u ra l s elect io n b u t in d o m es t icat io n n o

,

s u ch c o ntro lli ng i nfl u enc e ex is t s u n les s s u pplied by t he breed er

h im self Th is lack he m u s t pro v id e fo r if he h o pes to s u cc eed

.
,

b u t it c o n s t it u t es o n e o f his p rinc ip al d iffi c u lt ies .

9 . F a s h io n and c u s t o m d e c r ee that an i m a ls s h o w n at t h e fa i r s

S ha ll b e p u t i nt o extrem e c o nd i t io n and th is is a c o n s tant m enac e

,

t o th e effi c iency Of a breed in g h e rd .

10 A s tro ng v ein Of s pec u lat io n ha s ent ered int o m any lines

.

Of an i m al breed ing t he t end en cy of wh ic h is to u n s ettle r ic e

, p s
656 PRA C TI C A L PR OBLEMS

and c o nd it io n s generally in what o u ght t o b e o ne Of t he m o s t

s t ead ily c o nd u ct ed o f al l t he i nd u s tr ies .

T h es e in br ief a re s o m e o f t he lim itat io n s Of t he an im a l

, ,

breed er He canno t a ffo rd t o c lo s e his ey es to th eir ex is t en c e

.
,

fo r th ey a re rea lit ies He will get o n bes t t o frank ly c o n fes s th ei r

.

ex is t enc e a n d t o m eet th em t o t he b es t o f his ab ility I t rem a i n s .

t o exa m in e a li tt le m o re c lo s ely i n t o so m e o f th ese a n d o th e r

d etailed c o n sid erat io n s that m u s t engage t he breed er s attentio n '
.

SECT I O N I I F EW E R C H A RA C TER S FO R SEL EC T I O N

The great es t s ing le im pro v em ent po s s ib le in p res ent d ay -

a n im al breed ing in m o s t lines wo u ld b e to free the S it u at io n

fro m u n im po rtant charact ers A t bes t t he breed er m u s t pay .

rega rd to a large n u m b er o f c o n s id era t io n s i n m ak ing select io n s .

C o n s t it u t io n a l v igo r h igh pro d u ct iv e po wers a n d u t ility fo r t h e

, ,

p u rpo s e in m ind a re f u nda m ent a l c o n s id erat io n s and the las t ,

( u t i li ty ) is m o re t h a n lik ely t o c o v er m a n y po i nt s .

N o w t he d iffi c u lt ies O f s elect io n increa s e at a s u r pr is ing rat e

a s req u i rem en t s m u lt ip ly I f a pro p er d egr ee o f c o n s t it u ti o na l
.

v igo r is f o u nd in b u t o n e an i m a l o ut o f t wo — and it is n o t ,

h igh er than that th en t he chanc es o f a pa rt ic u lar an im a l pro v

,

in g s at is f a ct o ry in th is res p ect a re b u t o n e o u t o f t wo o r as we , ,

sa
y h is pro b a b ility is
i I f aga i n b u t o n e an im a l o u t o f three
.
, ,

1 —
,

is f u lly f ert ile and it is d o u btf u l if t he pro po rt io n is h ighe r

, ,

in c erta in lin es a t lea s t th en t he c ha nc es o f a s tro ng c o n s t it u

,

t io n and f ull fert ility b eing fo u nd in t he s a me ani mal a re b u t

I f n o w we add t o th is a th ird req u ir em ent f ou nd
3
2
x or
a .
, , ,

s ay in b u t o n e an im a l o u t o f t en th en o u r chanc es ha v e b een
, ,

red u c ed to 5 x x 1 15 1
wh ich is eq u iv a lent t o saying that
“

o n ly o n e an im a l i n 6 0 tak en at rand o m that is b u t o n e an im al ,

in 60 f o a ll that a b r nt t e breed will f u lly meet o u r

re o n i o h —
d em and s and sa t is fy o u r req u irem ent s ex c ept i n s o f a r a s fire ,

c ha r a c tersi n q u es t i o n m ay b e r ela ted by ca u s a t i o n a nd to ” ra t

ex ten t ov er lap .

1 B y fu ll fert i itl y is not m ea n t t he p o wer t o rep ro d u c e as a gai n s t l

ab s o u t e

b a rren n es s , b u t ra th er fu ll m a x im u m p o we rs
an d o f rep ro d u c t io n , t h a t is , regu lar

d
b ree in g t h ro u g h out a rea s o n a b ly
lo n g ife l .
658 PR A CT I C A L PR OB LEM S

at bes t a m ixt u re Of goo d and e v il — ,

a m ixt u re m o reo v er ,
,
that
will n ev er p u rify it s elf and th a t
,
c an be p u ri fi ed o n ly by a retu rn
t o fi rs t p ri nc ip les
.

S ECT IO N I I I FA SHI O N

As fa s hi o n d ecrees t he c u t o f o u r c lo th es S O it a lso d ecrees

,

the length o f t he ta il Of a c o w o r t he s h a pe o f her h o rn and t he ,

h eight a t wh ich a ho rs e s h o u ld ra is e his feet fro m the gro u nd .

I f f a s h io n wo u ld b e rea so nab le and c o n s is t ent and s tab le it

, , ,

wo u ld n o t b e S O bad fo r breed ers c o u ld finally adj u s t th em s elv es

,

t o it s d em and s ; b u t it is n o t s tab le and Oft en i t is n eith e r

,

reas o nab le n o r c o n s is t ent.

N o w it is n o t s o eas y t o cha n g e t he c o nfo r m at io n o f an an im al

as it is t o a lt er t he c u t o f a gar m ent wh ich m ean s at t he mo s t
,

o n ly a t u rn o f t he s h ear s th i s way o r th a t Ev ery o n e o f th es e

.

d ecrees o f fa s h io n ind icat es a n add it io na l req u irem ent fo r s elec

t io n a n d we kn o w what tha t m ean s to t he breed er ; n o t on ly
,

th a t b u t s u ch d ecrees a re c erta in t o b e s h o rt liv ed changi ng fo r

,
-
,

o th er s m o r e o r les s tro u b les o m e . Wo rs t o f a ll m any o f th ese ,

req u irem ent s o f fas h io n a re to t he d is t inct and per man ent d is

ad v antage o f the breed — th a t is per m an ent u nt il bred o u t
, ,

wh ich we ha v e learn ed req u ires appro x im at ely s ix generatio ns

O f s u cc ess f u l s elect io n
.

B u t t he m a nd a t es o f fa s h io n a re t o b e reck o n ed with errat ic ,

and tro u b leso m e th o u gh th ey may b e fo r in a v ery large meas u re

,

th ey deter m in e s a les a n d fi x p r ic es N o w t he breed er is in t he

.

b u s in es s fo r m o n ey a n d he m u s t s ell s t ock o r aband o n all h o pe

,

o f p ro fi t
,
— wh ich in t he en d m ea n s to a band o n t he ent erp rise
ent i rely ; and n o pha s e o f pract ica l breed ing ca lls fo r m o re wis

d o m and s hre dn w e s s than th s part ic ar pro b lem

i u l — ho w t o ,

m eet t he chan ging d em and s o f t he m ark et and k eep t he h erd o r

s t u d intact and if p o s s ib le im pro v ing .

I n s o far a s th es e fas h io n s em anat e fro m t he Open m ark e t

th eir c o ntro l is p ract ica lly beyo nd the breed er B u t s o m e o f t he .

wo r s t o f th em em anat e fro m a m o n g t he breed ers th em s elv es who ,

s o met im es s eem bent o n in v ent ing art i fic ia l is s u es o n wh i ch t o

m a k e s a les. S o m et im es th ere c o mes a f eeling apparent ly t hat , ,

 A N I M A L B R E ED I N G 6 59

th ere a re too m any an im a ls o f a giv en breed a v a ilable and that , ,

in S elf pro t ect io n n ew s tandard s m u s t b e s et u p

-
,
.

T h is is c o nf u s ing t o t he b u y er and o n ly h u rtf u l t o t he breed

A s a m att er o f fact th ere n ev er were and n ev er can b e t o o m any
,

really exc ellent an im als o f any breed The q u es t io n O f d ev elo p

.

in g t he m ark et fo r p u r e bred s t u ff will b e d is c u s s ed la t er b u t h ere

-
,
.

it is en o u gh t o s ay that n o art ific ia l s tandard s s h o u ld b e to lerat ed

in any breed m erely t o creat e s ales T h is m att er can b e c o n
.

tro lled by the breed ers them selv es in th eir o wn as s o c iat io n s and ,

wh en it is c o ntro lled a large s hare o f t he sen s eless a n d d is t u rb ing

“
d ecrees o f fas h io n will ha v e d is appeared and t he rem ain ing
o n es will ha v e b een m os t ly m o d i fi ed i nt o c o m parat iv e har m less

nes s T h ere is to o m u ch h o m em ad e law pa ssing fro m mo u th to

.

m o u th a m o ng breed ers with o u t t he s anct io n o f a s s o c iat io n s and

, ,

m u ch o f it wo u ld n ev er b e s erio u s ly s u pp o rt ed o n any fl oo r i f t he
ad v o cat es were rea lly req u ired t o serio u s ly d efend it Here is a .

d u ty that ev ery as soc iat io n o wes t he breed it ad v ocat es and

wh o se int eres t s it m ainta in s .

B u t wh en a ll is s aid and d o n e ho w s ha ll t he ind iv id u a l proc eed to

,

m eet t he d ecrees o f a craze that in his j u dg ment will s peed ily pa s s ?

T h ere is n o bett er way than by t he u s e Of s i res that s tro ngly
poss es s the po int s d em and ed in th e m ark et a lway s being care ,

fu l t o pres erv e a goo d ly n u m b er o f the bes t fem a les u n co n ta m i

n o ted fro m t h e i nf ect io n T h es e will f o r m t he n u c leu s o f t he
.

n ew h erd o r s t u d aft er t he c raze h a s pa s s ed and t he pend u lu m

has s wu ng back t o t he n o rm a l .

Here t he breed er m u s t b e wis e in his j u dg m ent a s t o wh eth er

a n ew th ing is o n ly a pa ss ing craze o r is rea lly a per m an ent
im pr o v em ent in t he breed and h er e his acc u m u lat ed kn o wledge
,

o f an im a ls will s er v e him well ; b u t h e S h o u ld b e well ad v is ed

that by t he pro per u s e o f t he S ire a h erd may b e m ad e to t u rn

o u t a n ew s ty le o f a n im a l fo r a c o n s id erab le t im e w i t/t ou t i n a ny

way afi ec ti ng t/ze r ea l c/za ra c ter of t/zef on n da ti on and th is c a n b e

,

c o nt in u ed as lo ng as t he old s t o ck o f fem ales la s t s A s t he t ime .

o f th eir en d appro ach es h o wev er so m eth ing m u s t b e d o n e t o

, ,

res t o re th eir n u m ber or els e the n ew point m u s t b e acc ept ed and

,
.

bred int o t he fem a les wh ich really c o n s t it u te the backbo n e o f

ev ery pr od u c ing h erd .
6 60 PR A CT I C A L PR O BLE MS

S ECT IO N I V — S HOW R I N G C O N SEQ U EN C ES

-

A n i m als that h a v e m ad e th ei r rec o rd in t he S h o w ring a re

no ne t he wo r se fo r that fact and thi s s u cc es s add s grea t ly t o
,

th eir cred it as ind iv id u a ls and to the c o mmerc ial v alu e of th eir

g e t aft er ward .A l th ou gh t he exc es s iv e fitt i ng req u ired i n t he

ri ng is o ft en i nj u rio us it is n o t n ec es s arily s o
,
I t is o f c o u rs e .

tr u e that n o an im a l will rem ain lo ng in fo rm n o r can t he

“
,

proc es s o f fitt ing b e repea t ed m any t imes S h ow r ing anim als .

-

a re th u s o ft en a d is a pp o int ment t o t he eye lat er on b u t th is is ,

n o d etri m ent t o th eir breed i ng p o wer s T he o n ly da ng er f ro m

.

exc ess iv e fitt ing is i t s eff ect u po n f ert ility and if th is has been ,

im p a i red it will v ery s oo n b ec o m e ev id ent .

I t is a s eri o u s q u es t io n a s to w/zen a breed er c an a ffo rd t o

tak e t he ris k o f p u tt ing int o t he ring a v alu ab le breed ing an im al .

A s a m att er Of fact if n o t o f n ec es s ity m o s t S h ow a ni mals a re

, ,

yo u ng The writ er d oes n o t s hare t he o p in io n that s h o w rin g

.
-

an im a ls ha v e n ec es s arily been inj u red fo r breed ing p u rposes any ,

m o re th a n he s hares t he Opin io n that S h o w ring s u cc es s is a -

g u aranty Of breed ing po wer s U po n th is po in t n o thi ng is reli

.

able b u t the act u al t es t .

SECT I O N V — T ESTI N G O F S I R ES A N D D A M S

Wh en werem em b er that v ariability ca n no t b e red u c ed belo w

8 9 o r 90 pe r c en t o f t he v ariability o f t he rac e o r in extrem e , ,

cases perh a ps to 8 5 per c ent and wh en we apprec iat e the fac t

, ,

that n o m att er ho w m u ch t he type (o r m ean ) h as been im pro v ed

t h e v ar iab ility r em ai n s th en we a re n o lo nger s u rpris ed at t he la rge
,

n u m ber o f m ed io cre ind iv id u a ls that appear ev en in b lood lin es

t he m o s t ar is t oc rat ic and th a t ha v e b een lo nges t in the p u rp le “ ”
.

T he n ec ess i ty fo r select io n th erefo re wi ll a lway s exis t an d

, , ,

wh en we add t o th is that o th er fact that m any med io cre look ing

,
-

an imals a re after all great breed ers and m any exc ept io nal ind iv i d
u a ls b itt er d is app o int m ent s th ere is an add it io nal reas o n fo r
,

t he a ct u a l t es t .

A n d s t ill aga in n o o n e kn o ws p o s it iv ely what will b e th e

,

res u lt o f a n ew c o m b in at io n o f b loo d lines u nt il it has been

66 2 PR A CT I CA L PR O BLE MS

T he t es t ing o f yo u ng females is a d iffi c u lt b u s ines s T h er e .

is litt le u s e in breed ing th em to u nkn o wn s ires wh o s e o wn breed ,

ing p o wers a re pro b lem a t ica l T o d o that is t o m eas u re o n e .

yard s t ick with an o th er wh o s e s tandard o f acc u racy is u nkno wn .

T he y o u ng fem a les S h o u ld b e bred wh en ev er po ss ib le to tes ted

s i r es ,
and then the breed er will ha v e an acc u rat e mea su re o f
wha t th ey S h o u ld b e expect ed to d o u nd er h erd c o nd it io n s .

U n ity of th e h erd T he o ld p la n o f ha v ing repres ent ed in t he

.

s a me h erd all fa s h io nab le fa m ilies in t he pers o n o f it s f em ales is

fo rt u nat ely pa s s ing S u ch a h erd n o m att er ho w exc ellent o r

.
,

well bred it s ind iv id u a ls wa s aft er a ll b u t a m o t ley c o llect io n

,

o f s tro ng ly bred d iff erenc es o n wh ich n o S ire ev er b o rn c o u ld ,

b e expect ed t o s u cc eed S u ch att em pt s ha v e in v o lv ed m any

.

breed er s in a h o peles s tangle o f M end elis m fo r thes e v io lent ,

ad m ixt u res o f f a m ily lin es a m o u nt to litt le els e than cro s s ing ,

ei th er in th eo ry o r in pract ic e .

T he i n d iv id u a l breed er s u cc eed s b es t who att em pt s t o d o a

d is t inct iv e th ing and who preserv es o n e type thro u gh o u t his
,

c o llect io n of f ema les wh ich is his h erd He will fi nd t his d iffi

,
.

c u lt eno u gh to a cc o m p lis h with o u t s eek ing t he m u lt ip licat io n

o f typ es ; and he will if he i s wis e d is card m any fema les in t he
, ,

t es t ing beca u s e if wo rk ing with well kn o wn and t es t ed s t ra in s

, ,
-

o f lin e bred s t o ck o n e or two tes ts of a p a r t ic u la r c om b i na ti on a r e

-
,

as
g ood a s a d oz en T h e re is n o n eed that t he breed er s h o u l d
.

was te t ime and mo n ey and liv e in u nc erta inty if o n ly he will k eep

his type d is t inct h is b loo d lin es p u re and wil l t es t ev ery an im al
, ,

that is t o ha v e a per m anent p lac e in the h erd I f he will n o t d o .

th is he will ind eed b e tread ing a m aze o f u nc erta inty and will ,

b e ready t o say at t he en d o f a lo ng lif e and as t he fru it o f his

expe r ienc e V e rily breed ing is a lo tt ery an h o n es t ly u tt ered
“ ”

, , ,

b u t g ro s s lib el o n o n e o f t he great es t pro f es s io n s on earth .

Tes t in g si res A well es t a b lis h ed h erd has al way s in serv ic e a

.
-

m at u re and well t es t ed S ire who ha s pro v ed his breed ing po wer

-

o n so m e o f t he b es t kn o wn f em a les o f t he h erd N o t o n ly that

- 1
.
,

b u t t he o wn er o f s u ch a h erd is a lway s lo o k ing fo r h is s u cc es s o r .

1 A m o n g the m a n y an s wers to q u es t io ns to u c hi ng th is p o in t a s u rp ris in gl y

la rge p ro p o rt io n o f b ree ers d c o n fess t o t es t in g b u ls , l no t on co ws of k no wn b reed~
in g po wers b u t o n lzei/er s
.
.
 A N I MA L B R EE D I N G 66 3

If an Old and pro v ed s ire can b e had that is t he s ire to b uy , ,

b u t o rd inari ly S ires o f th is k ind a re n o t o b ta inab le fo r if th ey , ,

a re rea lly t es t ed s i res th ey a re us u a lly h eld in t he h erd that

,

t es t ed th em u nt il th eir p erio d o f u sef u ln es s is o v er I f h o wev er .

, ,

o n e is a v a ilab le it is a treas u re that S h o u ld no t go begging

, ,

as it o ft en d o es I f t he y o u n g breed er wo u ld m ak e i t a r u le to
.

b u y o n ly o ld t e t ed ire
,
s s s — th o u gh th ere is n o v irt u e in o ld
,

s ir es h w u ld d o b ett er breed ing than m any an o th er

p e r se
,
e o

with lo ng exp er ienc e beh ind h im who is c o n s t a nt ly acc u m u la

t ing exc ellenc e and as c o n s tant ly d iss ipat ing it by the u se of
u nt es t ed S i res .

The wri t er has c o nd u ct ed an ext en s iv e c o rres po nd enc e with

breed ers o f catt le o n th is po int and has fo u nd that the a lmo s t
,

u n iv e rs a l pract ic e i s t o b u a y u ng b u ll pro bab ly a yearli ng

y o , ,

and p u t lzi m a t onc e i n to s er v ic e on Me en ti r e lter d Th is is .

bu s ines s s u ic id e fo r it c o n s t it u t es a bar to any v ery h igh d egree

,

o f s u cc es s and is bes id es extrem ely dangero u s I t is h ead ed

, ,
.

s t ra ight fo r m ed io c rity — with in t he breed o f c o u r s e b u t i t is ,

med io cr ity n ev erth eles s .

Tes t in g y ou n g S ir es T h is t es t t o b e m o s t v a lu ab le s h o u ld
.
, ,

b e m ad e o n so m e o f t he be tt er fem a les o f t he h erd wh o s e ,

breed ing po wers a re kno wn I t wo u ld b e fo lly to u se the v ery

.

ch o ic es t ind iv id u als fo r th ey a re need ed fo r mo re Certa in wo rk

,

with t he t es t ed s ire at t he h ead b u t s o m eth ing m u s t b e kn o wn

abo u t t he fem ales o n wh ich ev en the prelim ina ry t es t is mad e ,

o r i t is o f litt le v alu e .

A s a ll y o u ng th ing s loo k pro m is ing thi s t es t will n o t b e ,

wo rth m u ch u nt il the y o u ng ha v e n ea red m at u rity To b e s u re .

,

s o me i nd iv id u a ls will b e s o u npro m is ing a s t o S h o w it at b irth ,

o r so o n aft e r b u t v ery m any m ed io cre an i m a ls will n o t m ak e

,

thei r med ioc ri ty m an if es t u nt il m at u rity appro ach es Th ey th en .

exh ib it th eir inab ili ty t o tak e o n t he fin er fin is h and t he b ett er

t o u ch es that go with the breed .

On th is po int expe rienc e is f u ll On e o f t he fin es t s u ck ing c olt s

.

ev er kn o wn t o t he wr it er wa s exc eed ing ly pe rf ect a s a y ear ling ,

ga v e good pro m is e as a t wo year Old c o m m enc ed to f a ll a way as

- -
,

a three year o ld and befo re he was fi v e had d ev elo ped int o a

- -
,

v eri tab le pelt er with ewe neck and sway back .

6 64 PR ACT I CA L PR OB LE MS

I f the yo u ng that are to S h ow t he breed ing powers of p ro spec

t iv e s ires m u s t reach p ract ical mat u rity m u ch t im e will b e c o n
s u med in t he pro c ess I n catt le fo r exa m p le t he y o u ng m u s t
.
, ,

b e n o t les s than o n e and preferab ly th ey s h o u ld b e at lea s t two

, ,

years old Pract ically a year was c o n s u m ed in pregnancy a n d

.
,

t he hu ll wa s a y ear o ld at t he t i m e o f ser v ic e T h is will mak e t he .

pro s p ect iv e s i re at leas t fo u r y ears o ld by t he t im e his breed ing

po wers a re act u ally known T h is is t he age at wh ich m os t b u lls
.

a re s o ld as u g ly and dang er o u s a p ra ct ic e wh ich is d ep lo rable

“ ”

, .

A ll b u lls a re u g ly o r dang ero u s o r bo th that is alway s t o b e

, ,

as s u med ,
b u t at fo u r years o f age th ey a re j u s t ready t o ent er
u po n t he p erio d o f real u s ef u ln es s and t he rec o rd s will s h o w
,

that all great s ires ha v e d o ne th eir wo rk no t as y earlings b u t

lat er aft er th eir p o wers were kn o wn
,
.

Hav ing b een t es t ed and pro v ed o n a p o rt io n o f t he h erd a S i re ,

is o f c ou rs e p lac ed in f u ll s erv ic e and it is b u t b u si ness s ens e

, , ,

to m ak e t he m o s t o f h im a s lo ng as he is able t o c o nt in u e at its
h ead ; it is ev en mo re b u sin es s sen se to begin at o nce to loo k
fo r his s u cc es so r .

A h erd wi t h ou t a head Herd s pa s s q u ick ly and a herd with

.
,

o u t a h ead is d oo m ed t o s peedy ext inct io n u n less a s u i tab le o n e

can b e fou nd A h erd in s u ch a c o nd it io n presents a hard

.

prob lem to t he breed er and o wn er He ha s a lo t o f acc u m u lat ed

.

exc ellenc e b u t it is liab le t o d ie b efo re he can u se i t u n les s he

,

p u t s a pro p er s ire at the h ead with o u t d elay I f he cann o t do .

th at in all pro bab ility d is s ipat io n will f o llo w wh ich is b u t

, ,

an o th er fo rm o f ext inct io n I n th is ev ent t he o nly p ract ical

.

r emedy seem s to b e d is pers io n and th is is t he real reaso n fo r

,

m o re than o n e o f t he d is p ers io n s ales that c o m e alo ng each yea r

t o ar o u s e o u r wo nd er .

The d is p ers io n s ale a ff o rd s an exc ept io nal o ppo rt u n ity to

s ec u re real exc ellenc e in br eed er s fo r an i ma ls a re th ere o ff ered
,

that o rd inarily n o p u rchas er c o u ld b u y b u t th ese sales a re no t ,

nec es sarily fo r the bes t int eres t o f t he breed ; in m any cases

it wo u ld ha v e been bett er if t he herd c o u ld ha v e been k ept
t ogeth er .

T es t ed i nd i v id u als m ale and female a re t he backbo ne of t he

, ,

h erd and th es e a r e what t he wis e breed er will p res erv e thro u gh

,
6 66 PR A CT I C A L PR O B LE M S

at lo ng pric es fro m the h erd s of th o s e who were wise enou gh ,

to pr o t ect th em s elv es and m ak e ev eryth ing ta u t wh ile t he s t o rm

m ight la s t b u t who c o nt in u ed in b u s in es s agains t t he day wh en
,

t he m ark et wo u ld a ga i n want t he pro d u c e o f th ei r h erd s .

Wh en t he h erd is r ed u c ed at s u ch a t im e and it m u s t b e ,

red u c ed it is the yo u ng a n d u npro v ed s t u ff that S h o u ld b e

,

s ac ri fic ed fi rs t and i t i s m arv elo u s ho w m u ch can b e s o ld o ff

,

w i t/to u t d is t u r b i ng tlze r ea l n u c leu s o r

p r od u c i ng p a rt o f the lzer d .

I f the h erd is n o t a ll t he breed er c o u ld d es ire s u ch a t ime is ,

t he m o s t fa v o rab le o pp o rt u n ity that will ev er appear t o perf ec t

tlce lt er d by p u re/za s e T h is is t he t im e to gath er in t he mo th er s
.

and the g rand m o th er s o f t he breed fro m o th er herd s k eep th em ,

o u t o f t he b u tch er s hand , and s et th em at wo rk ; and abo u t

’

t he t im e th ey ha v e pro d u c ed an o th er gen er at io n th eir fo rm er ,

o wn ers o r o th er eq u a lly anx io u s p u rch as er s will b e ready t o pay

, ,

m o re fo r a c a lf tha n t he t es t ed d am a n d s ire b o t h c o s t Ev ery .

o n e wh o has liv ed lo ng a m o ng breed er s ha s s een th is s ta m ped e

o u t o f the b u s in ess f o llo wed by an eq u a lly i n san e t u m b le i nt o i t .

T he s o lid breed er will h o ld h im s elf well t o geth er at s u ch a t im e

and av a il h im s elf o f t he oppo rt u n ity b o th to impro v e his h erd
and to reap an as s u red ha rv es t lat er o n .

S EC TI O N V I I R EC O R D S

On e of req u irem ent s o f all goo d breed ing o perat io n s is

t he
an acc u rate sy s t em o f rec o rd s c o v eri ng ev ery im po rtant d et ail
, ,

lea v ing n o th ing to m em o ry Mo reo v er t he rec o rd S ho u ld b e

.
,

m ad e o n t he s po t .

Herd rec or d s Sim p le rec o rd s o f all p u rchas es sa les b irth s

.
, , ,

and d eath s are matt ers o f o rd inary b u s ines s acc o u nt s a n d in v en

t o ry b u t in add it io n to th es e th ere s h o u ld b e k ep t what m ay b e
,

kn o wn as the perfo rm a nc e rec o rd of the h erd T h is c o n s is t s o f .

th ree d is t inct ly separa t e feat u res :

I. A n acc u ra t e d es cript io n in writ ing of ev ery ind iv id u al a n i
m al o f t he h erd tak en n o t o n ly a t m at u r ity b u t at b irth o r ti me
,

o f p u rcha s e and as o ft en th ereaft er as changes in d ev elo p m ent

,

o cc u r S u ch a rec o rd S h o u ld b e a d es cript iv e h is t o ry o f the in

.

d iv id u al fro m b irt h to d ea th o r at lea s t to d is pos al acc o mpa n ied

, ,
 A N I M A L BR EED I N G 66 7

by ph o t ograph s if po ssib le and by any meas u rement s weight s

, , , ,

ach iev ement s o r fact s of any k ind that may later as s is t the j u dg
,

ment in b as ing c o nc lu s io n s as to selec t io n T he rec o rd o f t he .

s erv ic e m a l es s h ou ld b e es p ec ia lly f u ll and c o m p let e .

2. A f u ll d es cript io n o f ev ery ind iv id u a l o ff s p ring o f ev ery s ire

o f t he h erd a s c o m pared with his o wn d es c ript io n and with that

o f t he d a m — s u ch a series o f d es cr ipt io n s t o c o n s t it u t e t he
,

breed in g rec o rd o f the s ires .

3
. A br e e d ing r ec o rd s h o u ld b e o pen ed w ith ev ery fem a le of

t he h erd s h o wing dat e o f s er v ic e o r s erv ic es dat e o f b irth o f all

, ,

o ffs pring and s o m e na me o r n u m b e r whi ch may s erv e as an

,

id ent ificat io n m ark fo r ev ery ind iv id u a l pro d u c ed wh eth er liv ing ,

o r d ead .

I f th es e three lin es o f data a re k ept t he breed er will ha v e n o t

,

o n ly an acc u rat e pers o na l d es c ript io n o f ev e ry an im a l c o n s tit u t

in g t he h erd and ev ery o n e pro d u c ed by t he h erd b u t he will ,

also ha v e a c o m plet e breed in g rec o rd of ev ery an im a l bo th male ,

a n d f em a le and t he t wo t o g eth er will S h o w n o t o nly how m u ch

,

each an im a l ha s pro d u c ed b u t als o what wa s it s q u a lity When .

breed ing o p erat io n s ha v e been carried o n aft er th is p lan fo r a

n u m ber o f y ear s s u ch rec o rd s will c o ns t it u t e a m in e o f info rm a
,

t io n wh ich n o m emo ry c a n s u pply with s u fli c ien t acc u racy to b e ‘

d ependab le T ru s t ing to rec o llect io n is dangero u s and t he fl eet

.
,

in g i m pres s io n s f o r m ed o f c erta in ind iv id u als bec o m e ra pid ly d is

t o r ted and u nreliab le a s t im e pa s s es ho w rap id ly n o o n e kn o ws
,

u nt il he ha s been c o nfro nt ed a few t im es wi th his o wn rec o rd

m ad e fi rs t hand and u po n t he s po t T h ese rec o rd s a re es s ent ia l

.

t o t he b es t breed ing and ab s o lu t ely ind is pen sab le to t he m a n who

ma y s u cc eed t o t he m anagem ent o f t he h erd lat er o n .

B reed ers too o ft en pro c eed as if th ey expec t ed to liv e fo rev er ,

o r at lea s t a s lo ng a s t he h erd ex is t s wh erea s we mu s t loo k

,

fo r ward to t he t im e when an es tab lis h ed h erd s ha ll o u t la s t the

lifet im e o f it s f o u nd er and perhap s t wo three o r mo re gen era
, , ,

t io n s o f men s hall b e in v o lv ed in S hap ing the po lic ies of it s

breed ng i — ,
a ll o f wh ich is po s s ib le o n ly wh en t he m o s t perf ect

rec o rd s a re k ept .

I n c o nnect io n wi t h pr iv at e h erd rec o rd s t he fo llo wing is

append ed as th e p lan in act u al use by t he late Hon orab le
66 8 PR ACT I CA L PR OBLE MS

L . H . Kerrick ,
of B loo m i ngt o n ,
a v ery s u cc ess f u l I llino is ,
breed er of t he A berd een A ngu s T hes e rec o rd s are k ept o n -
.

cardboa rd 5 5 x 8 } inches and c o ntain ing n o print ing wha tev er ,

.

T h ey a re k ept o n edge in a lphab et ica l o rd er in a s pec ial b o x an d ,

it is b u t a m o m ent s wo rk t o n o t e t he pres ent c o nd i t io n o f a n y

’

female in the herd ; mo reo v er ev ery t im e the o wner c o n su lt s a ,

card he is a ffo rd ed a t a g la nce a n d in v olu n ta r i ly a pic tu re of

tire c omp lete b r eed i ng r ec or d of tb c c ow f o r lzer en ti re l ifet i m e .

T wo rec o rd s a re pres ent ed o n e o f F a ir La dy o f V eru la m and ,

t he o th er o f her fir s t o ff s pring F ancy F a i r ,

.

Ermi ne Bea rer

March 21 , 1 88 8 Fair La dy of V erula m 91 2 2 Fair La d y of Chil
lioot he
Ju n e 2 0, 1 8 90 Fan c y F ai r C . 1
5 21 5 E l l e n r ea g h of

K i n n o u l Par k 1 02 0 3

Ju n e 1 8, 1 8 91 W a p ell a Bo y B . 1 5094 C C A llen

. . 1 1 2 66

A p ril 1 8 , 1 8 92 E b o n L izzie C . 1 6988 E b o n is t 5 2 66

M a rc h 2
5 , 1 8 93 Bun t y B . 1 9 1 55 Eb o n is t 5 66
2

M a rc h 1 2 , 1 8 94 I re n e o f t he W ells C . 2 1 2 43 Eb o n is t 5 2 66
Ju n e 1 5 , 1 8 95 Lu c y M . of t h e W ell s . C . 2
3 5
4 1 C ra igo of Es t i ll 19
5 1 8
M ay 2 , 1 8 96 Pa irie D . C . 2 4 96 1 C ra igo of Es t il l 1 95 1 8

M arc h 1 5 ,
897 1 y
L gia o f th e W e ls l C . 2 7 94 5 C ra igo Of Es t ill 1 95 1 8

Ja n u a ry 2 7 , 1 8 98 F lo ren c e F o f the W el . l s C 3 2 444

. W ill is E G ra
. y 2 47 5 1

A p ri 6,l 1 8 99 E ri e of t he W ell s C 3 4 1 96
. C ra ft o f th e W ells 2 3 4 50

Ma rc h 3 0 , 1 900 S en a t o r I I o a r of t he W e ll s B 4 1 94 9
. C ra ft o f th e W ells 2
3 5
4 0

F eb ru a r y 1 2, 1 90 1 S t a t es m a n of t he W ell s B 47 8 1 3
. C ra f t o f t h e W ells 2 34 5 0

Jan u ar y 1 7 , 1 90 2 Bun k er o f t he W ells B 5 7 7 03

. Pa i n s t a k er o f
A b erlo u r 3 4 2 2 0

Ma rc h 1 8, 1 904 F l id
or a of the W ell s C 7 57 5 1
. Pa in s t a k er o f
A b erlo u r 3 4 2 2 0

Ellenrea gh of

K inn onl Park I 3

ju ne 2 3 , 1 8 90 Fa n cy Fair 1 52 15
Fai r Lady of

Verula m
A u gu s t 2 0 , 1 8 92 W ap el la La dy H C Al
. . l en

S ep t emb er 1 0, 1 893 F a ir Fa n c y E b o n is t
Ju l y 2 , 1 8 94 R ose F . of th e W ell s E b o n is t
Feb ru a r y6 , 1 89 7 L it t l e B en C ra igo of Es till
D ec em b er 2 8, 1 89 7 Op era to r W ells o f th e W il lis E G ra
. y
Ja n u ary 2 1, 1 8 99 G ra ym on t o f t he W el ls W il lis E G ra
. y
F eb ru a r y 2 4 , 1 900 e o f th e W ell s
C ro u j Ro y l
a Ju dge
Jan u a r y 1
5 , 1 90 1 Miss W es t o f th e W ells Pa i n s t a ke r o f
A b erlo u r 34 2 2 0
6 70 PR A CT I C A L PR O B LEM S

T h is latt er point is c o v ered in the s y s t em in u se by A J Lo v e . .

f R s c o e I llin o is a w ell k n o wn breed e r o f B erk s h i res

jy
o ,
o o , ,a -
,

s a m p le card fr o m wh o s e h erd is h ere r epro d u c ed

I ndex No . 1 6. I mport ed Bes s ie II 55 1 0 1 , farrowed A pril 1 0 to Mast er

piece 77 000 Farro w ed 5 , sa v ed 5 : b
. oar s 4 , sow 1

So l db oar to J W Ma rt in G o t h a m W is c o n s in
. .
, ,

L . W ll . B ro wn , Ber i n , I in o is l
J R L g
. S w d I ll i
. o an , e ar , n o is

Hib b d 8 B w M i h ig
ar 1 ro n, c an

so w N b ek I H p it l
ra s a n s an e os a , L in c o l n, N eb ras k a

To ta l fo r lit t er of 1 9 5
0 54

In c o nn ect io n with Offi c e rec o rd s o f th is k ind the breed ing “

b oo k k ept at t he barn s h o u ld rec o rd the d a te o f ev ery s er v ic e ,

t h e dat e s ex a n d any d is t ing u is h ing fact c o nc e rni ng t he b irt h

, ,

o f ev ery ind iv id u a l liv ing o r d ead a n d any o th er fact that wo u ld

, ,

prov e of t he s light es t v a lu e in es t im at ing what t he h erd is d o ing

o r ha s d o n e .

I f th en in add it io n th ere were k ept an acc u rat e d es c ript iv e

, , ,

rec o rd o f ev ery in d iv id u a l that is c o n sid ered wo rthy to ent er

th e h erd o r t o b e s o ld a s a br eed er and b es id es th is a ls o a s a c
, ,

c u ra te a r ec or d o b c u n wor t/ty p r od u c ts o f t he h erd t he breed e r

f t ,

wo u ld ha v e — n o t o n ly fo r his o wn s at is fact io n and pro fit wh en

m emo ry fa ils o r b ec o m es c o nf u s ed b u t fo r that o f o th er s who ,

may hand e l h i s bre d ng

e i — a rec o rd o f q u al it ies goo d and bad

o n wh ich a s k illf u l br eed er c a n s afely ba s e h is s elect io n With .

o u t a rec o rd s u ch as th i s a ll r ea l s elect io n is li m i t ed t o what is

d o n e with the eyes o n an im als s t ill liv ing exc ept a s a ma n may ,

b e g u id ed by an u nc erta in m em o ry Ho w u nc er ta in t hat is he .

will rea lize wh en he rev is it s in f u ll prim e o f lif e t h e h ills an d

v a lley s o f h is b o yh oo d .

Pedi gr ee records T hat wh ich is n eed ed with in t he h erd is

.

eq u a lly i m p o rtant wi th i n t he breed T he fact s of h ered ity go .

t o S h o w that all goo d breed ing req u ir es that t he type S ha ll b e

u nchanged fo r at lea s t s i x gen e rat io n s if we h o pe to get any ,

th ing lik e u n if o r m ity o f o ffs pring I f th is b e t ru e we need .

,

acc u ra t e rec o rd s c o v ering all im p o rtant d eta ils and all v alu able
c ha ract ers fo r at lea s t t he six gen erat io n s requ ired to prod u ce
a s table type .
 A N I MA L B R EE D I N G 67 1

N ow o u r ped igree rec o rd s f u rn is h litt le info r m at io n o u t s id e o f

bloo d lin es and th ey a re t o tally s ilent as to what t he ind iv id u als
,

act u al ly were in thei r o wn per s o nalit ies T h is info r mat io n t he

.

breed er n eed s and m u s t ha v e if he is to s u cc eed S o m e s light .

beginni ng has been m ad e in t he way of track rec o rd s am o ng

rac ing h o rs es and in ad v anc ed r egis t ry a m o ng dairy c o ws
,
.

T h en t oo breed ers a im to s u pp ly in th eir pri v at e cata logu es

, ,

s o m e d eta iled inf o rm at io n a b o u t part ic u lar an im a ls b u t in many

cases s u ch d escript io n c o ntain s so large an element o f ad v er
t is in g a s t o thro w d o u bt u p o n i t s acc u racy .

A S t he ca s e s tand s t o d ay th ere is n o way in wh ich an im par

,

t ial and t ru s t wo rthy rec o rd can b e as s u red ev en o f o u r m os t

fa m ou s and v alu ab le an im als T h is being t he ca se the in div id u a l
.
,

breed er c o m ing int o t he b u s in es s m u s t d ev o t e y ears o f his life

t o t he acc u m u lat io n o f a m a ss Of data m o re o r less reliab le ,

gath ered i rreg u lar ly and o ft en s u rrept itio u s ly fro m the u nd er

c u rrent o f s id e ta lk in which o ld and pro m in ent breed ers lik e ,

o th er m o rt a ls so m et im es ind u lge
,
.

N o w th is o u ght n o t t o b e S u ch inf o rm at io n belo ngs t o the

.

breed and to f u t u re breed er s who ha v e a right to kn o w t he fact s

,

abo u t t he a n im als wh o s e b lo o d lin es th ey a re o b liged to u s e and

s o m et im e i n s o m e wa w h en c o m m erc ia l int eres t s a re n o lo nger
, y ,

su pr e m e
,
— if n o t b ef o re an acc u rat e and im part ia l d es c rip
,

t io n o f ev ery great a n im al will b e m ad e a m att er o f perm an ent

rec o rd and will find its way into t he h is t o ry o f the breed .

I t is t o t he i nt eres t o f the breed that th is s h ou ld b e s o and it ,

is a ls o to t he int eres t o f t he y o u ng breed er that he m ay pro c eed

,

at o nc e and int elligent ly with his breed ing o pera t io n s and n o t

s pend t wenty o f t he b es t y ear s o f his lif e in c o llect ing inf o r m a

t io n by ind irect and o ft en d ev io u s m eth od s — inf o r m at io n that

,

is by go o d right s p u b lic pro pert y and a s s u ch is t he rightf u l

,

h er itage o f ev ery man fro m the mo m ent he bec o m es a breed er

o f that part ic u lar breed .

A brilliant f u t u re a wa it s t he breed that will s ec u re and pu t

int o it s h is t o ry a n a cc u ra te and cri t ical d es cr ipt io n at lea s t o f ,

ev ery f a m o u s an im al s a id d es cr ipt io n c o v er ing a ll d is t ing u is h ing

,

o r u n u s u a l tra it s b o th d es ira b le and u nd es i rab le and n o t c o nfin ed

,

t o extra v agant pra is e .

672 PRA CT I CA L PR OB LE MS

If th is is ev er t o b e d o n e s o me fea s ible p lan m u s t b e d ev ised

, .

N o w in th is m att er t wo th ing s a re self ev id ent : fi rs t s u c h tr u tlt

-
,

u l and cri t ica l d es cr ipt io n c o u ld n o t b e m ad e o r at lea s t m ad e

f ,

pu blic d u ring t he lifet ime o f t he an imal wh ile large c o mmerc ial

, ,

int eres t s were i n v o lv ed and s ec o nd m o re than on e m an s j u dg

’

m ent s h o u ld b e c o n s u lt ed in m ak ing t he act u a l rec o rd .

T he wri t er v ent u res t o s u gges t that wh ile the a n im al liv es ,

and is in his p rim e and wh ile his charact er and ach iev ement s
,

a re well kn o wn a f u ll s tat em ent o f his ach iev ement s b e m ad e

,

and t wo critical d es cript io n s b e prepared on e by t he o wn er , ,

t he o th er by a c o m m itt ee o f t he a s so c ati n
i o — all t o fi nd a per
,

m anent plac e in t he p u b lis h ed rec o rd s o f t he breed .

Ho w th is Obj ect can b e ach iev ed is pro b lem at ical b u t u ntil it ,

can b e ac h iev ed the bes t resu lts in an im al breed ing will nev er
,

b e po s s ib le On e th ing is c ertain
.
,
t he p u b lic at large and t he
a s s o c ia t io n of breed ers in part ic u lar hav e an inh erent if n ot a ,

v es t ed right in ev ery an im al that c o m es pro m in ent ly befo re the

,

p u b lic and s o m et im e in s o m e way th is larger righ t of p u blic

, , ,

o wn ers h ip will b e c o nc ed ed great ly to t he g enera l int eres t s o f

t he breed and to t he c o n v en ienc e o f f u t u r e breed ers ; in o t her

wo rd s n o t ev en priv at e c o m m erc ial int eres t s will alway s int er
,

v en e t o prev ent a rec o rd o f t he fact s u nt il by t he d eath of all

, ,

part ies po s ses sing act u al kn o wledge t he rea l perso nality of a

,

famo u s an imal has bec o m e lo s t bey o nd the p o wer of res to rat io n .

T he b lo od of n o t o n ly o n e fa mo u s an im a l b u t o f m any fam ou s
an im als ru n s thro u gh the ped igrees of a ll o u r herd s T he fame .

of s o m e o f th em res t ed o n real exc ellenc e and well earn ed meri t ; -

that o f o th ers was d ue ch iefly to s k illf u l m anagement oft en to ,

s hrewd ad v ert is ing A ni mals o f bo th c la s ses p osses sed poin ts

.

of h igh exc ellenc e and bo th c la s ses als o p o s s es sed d efect s

,
The .

p u blic has a right to the f ac ts wh ich a re n o les s th an a v es t ed

,

int eres t t o ev ery m a n wh o o wn s a breed ing h erd .

SECTI O N V II I —D I SPOSAL OF SU R PLU S FEMALES

T he h erd it s elf mu s t ma ke t he firs t draft u po n it s f emale

o u tp u t in o rd er t o s ec u re m at eria l t o reenf o rc e it s n u m b ers .

S o m e f ema les will b e need ed by o th er breed ers o f s tand ing to

6 74 PR A C TI C A L PR OB LE M S

t ck n eed s t he s ires fo r t he s erv ic e a n d t he breed ers need

s o ,

t he m ark et .

B reed ers a re s elling to o m u ch back and f o rth a mo ng th em

s elv es. T he breed ing b u s in es s is too m u ch o f a m u t u a l ben efi t
ass o c iat io n wh ile th ere is an u nd ev elo ped p u blic wit h a lm o s t
,

u n li m i t ed b u y ing p o wer s that n eed s to b e ed u cat ed and it s

,

b u y ing po wers d ev elo ped Ma ny a breed er wo rk s ind u s trio u s ly

.

to s ell t wo o r three f em a les and a s ire to a n o v ic e part ly fo r t he ,

m o n ey that is in t he s a le and part ly t o s pread t he go s pel o f

bett er breed ing as he th ink s,
.

I t d oes n o t wo rk that way A n o v ic e has b een s ta rt ed in a

.

s m a ll b u s in es s T he chanc es a re great that he will n o t s u cc eed

. .

He will eith er fail and c u rs e t he breed o r s u cc eed o nly in d if ,

feren tly well and m ak e an u nd es irable c o m pet it o r who is willing

t o s ell s t ock o f t he s a m e breed ing at pric es m u ch belo w
“

what th ey m u s t c o s t wh en pro d u c ed by c a ref u l breed ing .

I f t he s a me m a n had b o u ght a s i re he wo u ld ha v e b een s at is

fi ed with the n ew breed a n d he wo u ld b e o n t he ro ad to a p er m a
,

nent hab it o f k eep ing b ett er liv e s t o ck He wo u ld then bec o m e a .

c u s t o mer agai n and again F ro m any p o int o f v iew the breed ers
.

m u s t d ev elo p the m ark et fo r s i r es fo r g rad ing p u rpo s es .

SECT I ON X —C O M M U N IT Y B R EE D I N G

M any ad v antages will fo llo w if an ent ire c o m m u n ity will go

int o t he pro d u ct io n o f a p a rt ic u lar c la s s o f an i ma ls as fo r , ,

exa m p le dr i v ing h o r s es
,
T h ere a re a th o u s and litt le d etails in
.

t he s u cc es s f u l m anagem ent o f any b u si n es s and fo r t he b es t ,

s u cc ess ,
m ind m u s t react u po n m ind I f a who le c o m m u n ity .

wo u ld go int o the pro d u ct io n o f dr iv ing h o rs es and d is c u s s t he

driv er his breed in g care d ev elo p ment and ed u cat io n as c o m
, , , , ,

m u n it ies n o w d is c u s s c o rn rais ing i n t he c o rn belt in a few ,

years ev ery man wo man and ch ild in that part ic u lar locali ty
, ,

wo u ld “
kn o w all abo u t h o r s es T h ey wo u ld s oo n b ec o me
”
.

s k illf u l driv er s and as is n o w t he ca s e in t he fa m ou s b lu e g rass

, ,
-

regio n o f K ent u c ky the c o m m u n ity wo u ld ha v e a rep u ta tion

,

that wo u ld attract b u yers and a h o rs e wo u ld bring m o re m o ney

,

than he c o u ld bring if he were t he o n ly o n e in t he neighbo rh oo d .

 A N I M A L BR E E D I N G 6 75

L et the wh o le c o mm u ni ty as far as po s s ib le breed the sa m e

, ,

ki n d o f h o rs e o r o th er ani m al so that it m ay win a rep u tat io n

,

fo r a d is t inct iv e pro d u ct a n d it will n o t o n ly d o bett er breed ing

,

than it wo u ld if it were t o breed m any types b u t the b u s in ess ,

will b e v a s t ly mo re pro fi tab le Pract ica lly a ll t he canaries o f

.

t he wo rld a re bred in t wo v illages in G er m any and n o b i rd with ,

a false n o t e is ev er allo wed to liv e s o s k illed hav e t he ent ire

,

v illag e bec o m e i n what m ight b e ca lled canary exc ellenc e No .

i nd iv id u a l breed er can ev e r eq u a l t he d egree o f s u cc es s wh ich

is h ere ev o lv ed wh ere pract ica lly no o th er int eres t enga ges
,

t he att ent io n of t he c o m m u n ity .

SE CT I ON X I — T HE Y O U N G B R EEDER

T h ere is no rea so n why t he you ng breed er s h o u ld n o t pos sess

h im self th o ro u gh ly and q u ick ly with t he p rinc ip les o f breed ing .

What he lack s is experienc e with an im a ls and real inf o r m at io n

ab o u t breed s He s h o u ld get his experienc e eith er by grad ing
.

o r by a s s o c iat io n with a g oo d h erd b u t he m u s t n eed s p ick u p

,

his i n f o r m at io n m u c h o f i t at lea s t fro m int im a t e as so c iat io n

, ,

with m en who a re in t he act iv e b u s i n es s o f br eed ing .

Th e su ck er in t h e sal es r in g A b o v e a ll t he yo u ng breed er
”
.
,

m u s t k eep his h ead He d o es n o t n eed and b e cann o t a ffo rd t o

.

pay large pric es fo r females I f he s ees s o me o ld and es tablis h ed

.

breed er b idd ing h igh o n a y o u ng female he m u s t n o t a s s u me that ,

he can a ff o rd t o b id eq u a lly h igh T h er e a re a d ozen rea s o n s

.

why t he o ld er breed er m ay want that part ic u lar an im al n on e of ,

wh ich wo u ld app ly t o t he y o u ng breed e r Fo r exa m p le it may .

,

b e t he o n ly o n e o f that part ic u lar breed ing o u t s id e t he h erd o f

t he Old er breed er and h e can perhap s o r th ink s he can affo rd
, , ,

t o pay ev en m o re th a n t he an i m a l is wo rth fo r t he S ak e o f c o n
tro lling that part ic u lar c o m binat io n ; Or he m ay want it t o pu t
int o t he S h o w r ing o r t o m at e with a part ic u lar S ire N o ne of
,
.

th ese reas o n s wo u ld app ly to t he y o u ng breed er who if he bu y s , ,

it tak es it fo r it s meri t
,
.

T he b es t way fo r a y o u ng breed er to get h is s tart in p u re bred -

an im als is to get it fro m a rep u t a b le br eed er who can b e pers u ad ed

t o S ell s o m e rea lly exc ellent and t es t ed o r part ially t es t ed f em a les
, ,
.
6 76 PR ACT I C A L PR OBLE MS

I t is d o ing n o inj u s t ic e t o any breed t o remi nd the yo u ng

breed e r that t he b u lk o f yo u ng th ing s c o m e to v ery little He .

will b e s af er with o ld an ima ls — ev en th os e o f c o ns id era b le ag e ,

pro v id ing th ey a re s t ill fert ile .

W ith him m u ch d epend s u po n pric e He has n o c all t o pa y .

extrem e p ric es He cann o t s ell h is s t u ff a t m axi m u m p ric es t ill

.

he has been in t he b u s i n es s lo ng en o u gh t o acq ui re so m et hi ng

o f a rep u tat io n and o n e o f his bes t ea rly rep u tat io ns is as a
,

caref u l j u d ic io u s b uyer
,
.

I f t he y o u ng breed er ev e r los es his h ead in t he sa les ring let ,

him n o t d o i t o n a f em a le that can at bes t pro d u c e b u t few o r , ,

u po n extremely y o u ng th ing s which s tand abo u t as m an y chan c es ,

o f c o m ing o u t wro ng as th ey d o o f c o mi ng o u t right .

A D DITION AL REF EREN CES

A N I MA L B R E ED ING . By W . M . Ha y s . B reed ers

’
G aze tt e, X LV , 1 99,

B R E E D I N G B E ES TO I NC R EA S E LE NG T H OP TONG UE . By J . M R an k i n
. .

M ic h ig a n E xperi m en t S ta t io n R epo rt , 1 8 9 7 p , . 12 7 a lso i n E x perimen t

S ta tio n R ec o rd , X I , 6 1 , 1 06 2 .

B R EEDI NG E x P E R I M E N T s w1 T 11 S H E E P . M is s o u ri E xpe ri m en t S ta tio n ,

B u lleti n N o .
53 , pp . 1 6 7- 1 88 ; a ls o i n E x per i m en t S ta tio n R ec o rd ,
X I V, 383 .

B R E E D I N G PO U LT R Y . E x pe ri men t S ta t io n R e c o rd , X l l l , 1 76 .

B R E EDI NG PO U LT R Y P OR E G G PR O D U CTI O N By G M G o well M ai ne

—
. . . .

E x pe ri men t S ta t io n , B u llet i n N o .
79 ; No .
93 , pp . 69 9 2 ; al so in
E x peri men t S ta tio n R ec o rd , X V , 3 94 .

h aw
'

B R E E D I N G S HE E P T O C HA N G E B R E E D I N G S E A S O N By T S M in
—
. . .

n es o ta E x perimen t S ta tio n , B u l leti n N o .

78, pp 7 1 8 7 . .

C R O S S B R E E D I N G C HI C K E N S
-
. B y E P M iles . . . V i rg in i a E x peri men t S ta tio n ,
B u lle t i n N O 96 , p 6 ;
. . i n E x pe ri men t S ta tio n R epo rt , X 1 , 1 0 7 4
a ls o .

C R O S S -
B R E E D I N G S HE E P . B y F W in ter . A g ri c u ltu ra l G a ze tte, Lo n d o n ,
.

1 900 , p . 2 46 .

C R OS S B R E E DI N G S WI N E
-
. E x peri men t S ta tio n R ec o rd , X I , 1077 .

C R O S S I N G C A TT L E . E x per im en t S ta tio n R epo r t, V I I I , 7 2 0 .

HY B R I D , G A M E C OC K U -
G I N E FOW L
A -
By T V i la ro B u lleti n
. . . of the
A m er i c a n M u seu m of N a tu ra l H is to r y , 1 8 97 , p . 22 5 a ls o i n E x pe ri
m en t S t a tio n R ec o rd , I X , 1 03 1 .

PE D I G R E E S T O C K R EC O R D S .
( R epo r t of C o m m i ttee on P h o togr aph ic
M e th o d s of Pres er v i n g .
) B y Fr a n c is G a lto n . R epo rt of the B ri ti sh
A s s o c ia t io n fo r the A d v an c e men t of S c ien c e, 1 8 99, pp 4 2 4 4 2 9 .
— .
6 78 PR ACT I C A L PR O B LEM S

anybod y ev er s eriou s ly d o u bt ed th is fact Th is is alt ogether .

o u t s id e t he q u es t io n o f t he d u a l p u rp o se an ima l fo r n o ,

att em pt has b een m ad e to breed the J ers ey fo r o th er p u rpo ses

than m ilk pro d u ct io n N o r is i t a repro ach t o J ers ey s that th ey
.

d o n o t exc el i n s o m eth ing fo r wh ich th ey ha v e n ev er bee n bred .

A S well expect th em t o tak e rec o rd s u p o n t he ra c e t rack o r t o d o ,

any o th er th ing fo r wh ich th ey ha v e n o t been fi tt ed by select io n .

T he infl u enc e o f t he en v i ro n m ent is therefo re perm is s i v e

rath er than a s sert iv e . I t a ff o rd s the ma t er ial and t he o p po r t u
n ity fo r d ev elo p m ent o f what was b o rn int o t he ind iv id u a l and ,

what was n o t b o rn int o i t cann o t d ev elo p n o m att er ho w fa v o r,

able the en v iro n m ent ,

— a s witn es s t he v ery d i fferent d ev elo p

ment o f t wo i nd iv id u a ls d ifferent ly b o rn b u t liv i ng u nd er t he

s am e c o nd it io n s o f life . I f an i nd iv id u a l is exc ept io nal we may ,

s ay o f him with c o nfid enc e that he wa s bo th well b o rn and well

c o nd it io n ed d u ring d ev elo p ment I f o n t he o ther hand he is

.
, ,

i nf erio r we a re u nc erta in wh eth er t o attrib u t e it t o n o n inh eri t

,
-

anc e o f v alu able ch a ra ct ers o r to th ei r failu re to d ev elo p o wing

,

t o u nf o rt u nat e c o nd i t io n s o r t o b o th
,
.

Well b red in div i du al s shou l d h av e good con diti on s I t is m an i

-
.

fes t ly u n wis e to expend t im e m o n ey and th o u ght o n t he p ro

, ,

d u ct io n o f ind iv id u a ls h igh ly end o wed wit h the rich es t po s s ib ili t ies

o f t h e rac e and th en fa il to pro v id e t he n ec es s a ry c o n d it io n s fo r

th eir d ev elo p m ent Ord ina ry b u s in es s sen s e th eref o re d icta t es

.
, ,

that the breed er s h o u ld s ec u re id ea l c o nd it io n s fo r t he fu ll and

pro per d ev elop ment o f t he charact ers wh o se im pro v ement he
aim s to sec u re thro u gh f o rt u nat e c o m b inat io n s of b lood lines .

To s ee h erd s o f t he b es t bred an im a ls s u ff eri ng fo r feed is at

-

o nc e path et ic fr o m t he h u m an itarian s tandp o int and u na c c o u n t

ab le fro m a b u s ines s s tandpo int .

Ha v ing s pent t im e and m o n ey fo r the inf u s io n o f the h igh es t

po ss ibilit ies int o t he h erd c e rt ain ly b u s iness fo resight d em and s
,

that th eir f u ll realizat io n s ha ll b e prev ent ed by n o o rd inary c i r

c u m s ta n c e ;
y e t w hat s hare o f t he b es t br ed an im a ls and what
-

pro po rt io n Of o u r i mpro v ed seed s a re giv en f u ll o ppo rt u ni ty t o

S h o w what i s rea lly i n th em ?

T he breed er is int eres t ed fo r an o th e r rea s o n i n s ec u ring t h e

f u ll d ev elo p ment o f a ll that is bo rn i nt o ind iv id u als and fa m ily
 D EV ELOPM ENT 6 79

lin es . In th er way can he j u dge of their real exc ellenc e and

no o

i n n o o th er way is his s elec tio n s af e The p ract ic e of fitt ing for .

t he s ho w ring is Oft en d ep lo red and n o t wi th o u t goo d reas o n

,

—
,

b u t o f o e th ng
n i w e m a y b e we ll as s u red we can n ev er b e ,

c ertain o f t he capac it ies o f an ind iv id u al u ntil th ey hav e been

p u t t o t he t es t by d ev elo p m ent .

On e o f t he hard f a ct s o f an im al breed ing is that t he d ev elop

m en t o f y o u n g th ing s is v ery o ft en left t o men who are n o t
S k illed in an i m a l pr o d u ct io n T h ey s eem to a s s u me that t he
.

well bred an im al in s o m e way can get a lo ng u nd er les s fa v o rable

-

c o nd it io n s than can t he u n im prov ed — a k i nd of o ff s ett ing o f ,

breed ing aga in s t feed ing and care .

I mpr ov emen t con sist s in prod u cin g an im al s an d pl an t s ab l e t o

mak e good r et u rn s f or goo d c on di t i on s , n ot m er el y t o ex i s t u n d er

hard c on di t i on s . T h is
fact o u ght t o b e pa s t ed in t he hat of
ev ery far m er T he p u rpo s e o f br eed ers is n o t t o pro d u c e s train s
.

that can liv e o n next to n o th ing and that a re ab le t o end u re ,

hard c o nd it io n s and n o t d ie o u tright ; it is to pro d u c e s t rain s of

an im als and p lant s that a re able to m ak e g ood retu r n s fo r t he
f u ller f eed and bett er c a re wh ich the c iv ilized and e d u cat ed
farm er can giv e a s c o m pared with nat u re wh ich is capric iou s , ,

o r wi th t he u n s k illf u l sem i s a v age who is i m pro v id ent

-
,
.

On e o f the m o s t s erio u s fa u lt s o f u n im pro v ed s t ra in s is that

th ey d o n o t respo nd to bett er f oo d o r to m o re o f it T h ey hav e .

been s elect ed fo r gen erat io n s fo r th eir powers o f res is tanc e to

hard c o nd it io n s and that is wh ere th eir s trengt h lies N ow that
,
.

we can pro v id e b ett er fo o d we n eed an im a ls o f h igh er effi c iency

,

ind eed th a t is o u r arg u m ent fo r b ett er an im a ls T h en aga in

,
.
, ,

hav ing a n im als o f h igh er effic iency we need better f eed and ,

m o re o f i t and th a t is o u r arg um ent fo r b etter bred c o rn and

,
-

o th er c ro p s S o t he t wo — a n im a l breed ing and b ett er feed ing

.

react the o n e u po n t he o th er and bo th go with bett er farm i ng

,

and with the greater n eed s of an ad v anc ing c iv ilizat io n .

The well b red an i mal i s a h i gh c lass mac hi n e T h is fact can

- - .

n o t b e to o well u nd er s t o o d by ev ery m a n who c o m es i nto relat io n s

with the well bred an im al and i t is tru e wh eth er we c on s id er

-
, ,

anim a ls a s m ach in es fo r the prod u c ing of m ilk o r m eat of labo r , ,

o r o f b o dy c o v eri ng ; wh eth er we c o n s id er that th ey are to

680 PRA C TI C A ] PR OB L E M S

m in is t er to n ec es s it ies o r to cat er t o o u r enj oy ment b y

ou r

perso nal s erv ice a s with t he s add ler and the d riv i ng h o rse
, .

E xc ellenc e is n o t to b e m eas u red by the po wer t o wi th s ta n d

d ep riv at io n b u t rath er by effi c iency to d o wo rk u nd er f ull feed
,

and u nd er goo d c o nd it io ns ; and to th is en d it is b u t good bu s i

nes s sen s e t o s ec u re fo r each ind iv id u al t he f u ll d ev elo p ment o f
all t he u sef u l q u a lit ies wi th wh ich h e is n at u ra lly end o wed .

Dev el opm en t is a stu dy b y i t sel f Here is an ent ire fi eld .

a lmo s t u nexp lo red We kn o w in a genera l way that t he

.
, ,

en ergy o f em bry o n ic d ev elo p m ent is n ev er att ai n ed lat er in

“

life and that if we wo u ld sec u re f u ll d ev elo p ment in gr o wth we

,

mu st k eep the y o u ng t h ing gro wi ng I n s o m e way o r o th er .

th is b u sines s of bo dy d ev elo p m ent if o nc e ch eck ed is nev er, ,

agai n f u lly res u med I t is tru e that a few expert s hav e learned
.

fairly well ho w t o d ev elo p s peed in h o rses and o th ers ho w t o ,

train s add lers and driv ers b u t wlzet/zer we cons id er tlze g r owtlz
f
o tlze b ody , t/t e d ev elop m en t of i ts f unc tion s , o r tite ed uc a ti on of
tice m en ta l f ac u l t i es , we do n ot y et p os s es s ev en tlze ru d i m en ts f
o

tire kno wledg e of tb c f m os t W

s ucc es s u l i th u s o n ly
d ev elop m en t .

an occa s io nal ind iv id u al enj o y s o pt im u m c o nd it io n s thro u gh o u t

his lif e and o n ly a few exh ibit in th ei r o wn perso na lity t he really
,

wo nd erf u l capac ity o f the breed to wh ich th ey belo ng I f one .

were t o say that t he s c ienc e and pra ct ic e of breed ing is far bett er
kn own than that o f d ev elo p m ent aft erward he wo u ld b e well ,

wi th in t he tru th and in t he o p in io n o f t he wr it er h ere wi ll lie

,

s o m e o f o u r great es t i m pr o v em ent s o f the n ear f u t u re T he .

exc ess iv e fitt ing o f an o ccas io na l i nd iv id u a l fo r t he S h o w ri ng ,

regard les s of c o n s eq u enc es aft erward is n o t what is h ere mea nt , ,

b u t rath er t he o rd erly and f u ll d ev elo p m ent in s u b s t a nt ial ly al l ,

i nd iv id u als o f th os e q u alit ies wh ich we d eem v alu ab le s o t hat

, ,

we may f u lly realize in ou r an imal s the q u alit ies we select and

breed fo r in o u r yard s .
68 2 A PPEND I X

a ll ? ( )
3 W h at a re the ob ec ts j of hag d
v in
i fferen t k in d s o f a v era g es ?
Suc h q es tio s as th ese a re
u n apt to b e o v erloo k ed b y t o se wh o av e fo r med h h
the h b i t o f a e a g i g all k i
a v r n n d s o f r es u l ts wi t h o u t c a ref ul t h o u gh t .

I n po p u lar la n gu age, we s pea o f the av er age d a ily tem pera t u re, th e k

a v er a ge len g t h
o f ea rs o f c o r n , the a v er age s tu d en t , the a v er age c i t izen ;

an d we s h o u ld k now th e ex a c t mea n i n g to be c o n v e ed y b y th ese a n d S im i

la r ex p ress io n s b efo re u s in g h
t em in s c ien ti fi c d is c o u rse .

k
T he ta in g o f a n a v er age p res u ppo ses a po p u la tio n w o se v a ria tes ha v e h
a c erta in m ea s u ra b le c a ra c ter a b o u t w i c h
we a re c o n c ern ed , an d t a t h h h
th e m eas u rem en t of t h is c h a r a c ter d ifi ers i n d ifferen t in d iv id ua ls . We
a t tem p t to des cr i be t hi s p p
o u l a t io n b y pu ttin g as id e the meas u rem en ts of
i n d i v id u als a n d c o n s tr u c ti n g a s i n g le i n ter m ed ia te n u m b er w i c h s all b e h h
d es c r ipti v e o f the to ta l po p u la t io n , i n so far a s o n e n u mb er c an d es c rib e
a po p ula tio n .

T he S in gle i n term ed ia te n u m b er wh i c h a n s wers h

t is p u rpos e is , in the

g en er al s e n se , s o m e k in d o f a n a v era
ge . We t h u s u se a v er a
ges fo r d esc r ip
t iv e p u r p o ses i n th e in teres t of b rev i t y; b u t , t a k en a lo n e, a n a v era ge c a n

not c o m pletel y d es c r ib e a p p
o u la t i o n an y m o re h
t an the m o tio n o f th e

c en ter o f g ra v ity of a s y s tem o f m a terial p a r t ic les c a n c o m pletely d es c ri b e

t he m o t io n of the s ep a ra te p ar tic les .

In s ta t i n g w h at a n a v er age is we ha v e a ls o s ta ted i ts fu n c tio n ; b u t, as

ju st i n d ic a ted , it m u s t not be ass u m ed t h a t an a v erag e
gi v es a ll t he i n fo r

h h is to b e sec ed fro m the me reme t o f a pop la ti on I t

m a tio n w ic ur as u n u .

c a n o ly ta k e th e pla c e o f the m ass o f fi g

n es fo c e tai p ec i l p poses
r ur r r n s a u .

I n fa c t th e e h , b ee a te d e c y b y o m ewha t c a eless wo kers wi th

r as n n n , s r r

st a ti tic l d a ta to a tt c h too m c h i m po r ta c e to a e ges a nd n o t eno g h

s a a u n v ra u

—
,

t d e i tio
o v af o m the a e ge ns a po in t t h a t will b e d eal t wi th i
r v a later
ra , n

s ec t io n .

Th e e are fi e d iffe e t k i
r v r n n ds o f a v er a ges in c o m m o n u se for d iff eren t
pu r po e sT h e es e . 1
) the s ar a ri t h m etic m ea n , ( )2 the weighted a ri t h metic
mea n , ( 3) the g eo m et ri c m ean , (4) th e m od e, ( 5) t h e m ed ia n .

Wh ile s o m e of h
t es e a v erages h av e b een d efi n ed , a n d u sed freely in the
tex t , i t s eem s well t o res ta te h
t ese d efi n itio n s toget er wit the o t ers , the h h h
h ei e pec ti e
b et ter to d is c u ss t r r s v a d v a n t ages a nd d is ad v a n tages , a n d so me

o f the p r po e to wh ic h e c h i a d pt ed
u s s a s a .

T he ari thmetic mean T he ri th m eti c m ean o f a po pu la tio n. a o f n v a r ia tes

m a y b e d efi n ed as fo llo ws :
su m o m ea s ur em en t qf n v a r ia tes
a r i thm etI c m ea n
f

T h at is , to f i n d t/t e a r i t/u u eti c m ea n qf n v a r i a tes , we d i v id e tlte su m o

f th e
m ea s u r em en ts of ” us e v a r ia tes by th e n u m ber o f v a r ia tes .

h
T u s , in the c as e o f a h
t o u s a n d ears o f c o rn , th e a ri t h m etic mean o f the

len g th s of the ea rs is the s u m o f the len g t s o f 1 00 0 ea rs h d iv id ed b y 1 000 .

T he u se o f th is k i n d o f a n av era e
g ha s a lwa y s b een ta k en b y ob se rv ers a s the
b es t m et h od o f c o m b i n in g d i rec t m eas u remen ts o f the s a me q u an t i t y. Th is is
 A PPEN D I X 68 3

the av era ge mos t c om mo n ly em plo ed , y a n d o ne o f the s tro n

ges t a rgu m en ts

ad v a n c ed to j u s ti fy t hi s m et h od is its u n iv ers a l a c c ep ta n c e . 1 1 is wo rth

wh ile, h owev er , to c a ll a tten tio n to o n e o f the ab u ses o f the a rit h metic
m ean . F o r in s ta n c e, i f a v ery few ( s a y fo u r ) meas u rem en ts h av e b een
m ad e o f a c er ta i n c h a ra c ter , th e a rith m etic m ea n h as o ften b een ta k en as
a good i n d ex o f t h eir m ean i n g ; b u t i f t h es e few meas u remen ts d ifi er
wid ely, to repo rt th eir a ri th metic m ea n is to fu rn is h a v ery m is lead in g a nd
u n tru s two r th y p iec e o f in fo r m a tio n T h is h as o ften b een d o n e b y tho se .

who h a e gi en n o t h o gh t to ta tis tic al meth ods

v v u s .

T h ere i a so t o f c o mm e c ial po in t i v olv ed in the a ri th meti c m ea

s r r n n

wh ic h s ho ld n o t b e o e loo k ed F o r i s ta n c e i f a real es ta te d ea ler S ell

u v r . n , s

a hu nd red lo t a t a rio s p i c es o f wh ic h the a ri th m et i c al a era ge i

s v u r $800 ,
v s ,

t h is a s u es u s t h a t i f th e eller h ad so ld ea c h o f the lo ts fo r $
s r s 800 ins tead ,

of s elli n g a t d iffere t p ic es he wo u ld h av e rea li ed p rec is ely the s a m e

n r , z

f o m the s ale o f the wh o le u m b e o f lo ts as he has reali ed fro m sellin g

r n r z

at va r y in g ra tes e e tho g h we h a e n o i n fo rm a t io n as to wh a t a n y in d i
, v n u v

has pa id fo r a lo t
v id u a l .

Weighted arithmeti c mean . A s li

g h t m od ifi c atio n o f the ab o v e meth od
is o ften u s ed . T o ill u s t r a te, the t ho u sa n d m eas u rem en ts of len g t hs of ea rs

o f c o rn m ay b e a r ra n ged , let us s a y, i n h alf in c h gro

-
u ps as fo llo ws

Freq u en c ies .

I
wh ere, fo r in s tan c e, the 6 in c -
h gro up i n c lu d es a ll ea rs wh os e len g t hs are

b etween an d I n g en er a l , if v 1 , v 2 , v , rep resen t the c las s m a r ks ,

an d f f,
,, , f ,
rep res en t the c o rres po n d i n
g fr eq u en c ies , t en h

S ta ted in wo rd s , t is m ea n is h
by m u ltip lyi ng ea ch m a r k of a ob ta i n ed

c las s by the cor r espon di ng f r eq u en cy , a n d d i v i d i ng the s u m o

f th e p r od u c ts

by the tota l p op u la tion .

h ki
T is nd of a v era ge is u s ed a g rea t d ea l i n ou r wo rk an d is a p p ro x i

m a tely eq u al to the ord in a r y a ri t h m et i c a l i f the gro u ps a re fa i rl

a v erage y
n arr o w. I ts a d v a n ta ge o v er the h
o rd in a ry a ri t m et i c m ea n l ies in the fa c t

h
t a t it is m o re eas il y c o m p u ted . I n repo r ti n g the m ea n d ai l tem pera tu re, y
the a v er age len g t h o f c a rs o f c o rn , the a v era ge h eigh t o f a c er ta in C la s s o f

m en , k in ds o f a erages is m ea n t W e u e th es e a erages
o n e o f t he a b o v e v . s v

so m u c h in th is wo r k t h a t we s pea k o f th em as the m ea n fo r b re ity

”
, v ,

s o t h a t wh en t he term m ea n is u s ed with o u t a li m i tin g a d j ec ti e i t is to

”
v ,

b e u n d er s to od th a t a n a ri th m et ic m ean is m ea n t .

The geometri c mean The g eom etr i c m ea n of n n u m ber s is f ou n d by m u l

.

tip lyi ng the n u m ber s tog ether a n d ex t r a cti ng the n th r oot o

f th e p r od u c t .
684 A PPE N D I X

Le t us as s u m e h
t a t d u ri n g a d ec a d e the a tten d a n c e a t a un iv ers i t
y in
c reased 1 00 per c en t , a n d let us p p
ro o s e t he pr o b lem o f fi n d i n g th e av era g e
an n u al r a te of in c rea se . W ill i t d o to reso r t to the a ri thm eti c mean in
h
t is c ase a n d sa y tha t th e av era ge ra te o f in c rease is 10 per c en t ? N o : an
in c r eas e of 10 per c en t a n nu a ll y wo u ld gi v e a n a tten d an c e

ti m es the a t ten d a n c e a t the b eg in n i n g o f t he d ec a d e W h at we really wa n t

W
.

is : h
t at is , an i n c reas e of a li ttle mo re t ha n 7 per c en t ea c h

yea r will d o u b le the po p u la tio n in a d ec ad e .

T he geo m etri c a l av er age i s b u t li ttle u sed i n o u r wo r , b u t i t is b ro u gh t k

fo rwa rd h
ere to rem i n d u s t a t a n a v er a ge c a n , i n gen er al , b e d e en d ed
p h
u po n o n ly to serv e a d efi n ite p u rp ose a n d , w en the p u r pos e is k n o wn , we h
a re so m et im es c on fi n ed t o o n e k i n d o f a v era ge, o r a t leas t a b le to see th e

ad v a n t ag e o f o n e k in d o f a v era ge o v er an o t he r . S u pp ose th a t we kn o w the

p ro tei n to h a v e b een i n c reas ed 5 0 p er c en t in ten years ’
c o n ten t o f c o rn

h
b reed i n g C a n we sa y t at the a v era ge a n n u al ra te o f in c reas e was 5 per
.

c en t ? C lea rly we c a n n o t W a t we S o uld d o is to ta e . h h k

an d sa
y t h at the a v era ge a n n u a l r a te o f in c rease is a p p r o x im a tel y 4 pe r c en t .

T he mod e . W hen we s p ea k o f the a v era ge c o llege s t u d en t o r th e a v er

a ge c i ti zen we c erta in ly do not hav e referen c e to the a r i t h metic or geo

m etri c a v er a ge o f an y th i ng . W h en we sa
y a m a n is an a v era ge c i ti zen

we m ean t h at he r ep resen ts a wh ic h
t pey is me t o ften er t h an any o t h er .

If a c o m mu n i ty h as ten mil lio n a ires , b u t all h

the o t er c i ti zen s are i n po v

er ty, a n a r i t h m etic a l a v era

ge h
m ig t g iv e the im press io n t a t the peo p le o f h
the c o m m u n it y a re in go o d fi n a n c ia l c o n d i tio n , w ile r ea ll
y th e a v era ge h
c i ti zen is i n po v er t y
T he a v era ges th u s fa r d is c u ssed a re in n o wa y
.

h
s u i ted to d es c r ib e t is po p u la tio n , b u t th e a v erage c alled the
“
m od e is
”

u se fu l fo r t is pu r po s e h .

If a p p
o u la ti o n be a r r a n g ed in ser ia te o rd er wi t h res pec t to so me c ha r
a c ter, a m o d e is a v a lu e to w ic h h th ere c o r res p o n d s a
g rea ter fr eq u en c y
h
t a n to v alu es j us t p r e c ed i n
g a n d im m ed ia tely fo llo wi n g i t in the a rr an
g e

m en t .A po p u la tio n m ay h a e m re t h a
v o n one mod e, b u t the po p u la ti o n s
wi th h h h
w ic we s all d ea l o ly o n e h a e in ge e al
v , n r , n .

T h is k i d o f a e age seem s to b e a b ou t the s a me as th a t o f the n ew

n v r s

e w h e t h ey pea k o f t he a e g e c i t i e n I a d em o w e o f te ra
p p
a rs n s n c a c y v n z . r

h e the c y o f the g re tes t good fo the greates t u mb er a d i n s i t tha t

ar r a r n ,
”
n s

legi l t i
s a h ll b en efi t the a e age m a n the m a n a t the mo d e
on s a v r , .

R e e t i g aga i to the t h o a d ears o f c o n ar r a ged i n h al f in c h gro u ps

v r n n us n r n -
,

it s h o ld b e n o ted th a t the freq e c y i c e e up to the c lass o f m a r k

u u n n r as s

9 i n c h e a dst h e d e c ea se
n W e m ig h
n t c o c l rd e t h a t 9 si .c h e is e xa c t ly n u n s

the mod e fo t h i p p l t io
r I t m t
s b e o
r em e m b
u a e r e d t h a t na l.l m e a u e us s r

me t f o m
n s r to i ch we e p la c ed i t he 9 in c h g o p a n d t h at
n es r n - r u ,

a d ifi eren t g o p i g m ig h t c h th e f e i m w h t H e i
r g e u n q e c e o e a an n c e 9 s r u n s s .

s a id b e the em i r i ca l m od e, a nd the theor etic a l m od e is d efin ed

to p as a
68 6 A PPEN D I X

S ECT I O N III — G R A PH I C R E P R E S E N T A T I O N OF
S T A T I S T IC S

A m ere tab u la t ion o f a n y c o n s id era b le n u m b er o f figu res d oes n o t m a e k

i t pos s ib le, i n gen era l , fo r the m i n d to g r as p the m a i n fa c ts wh ic the h
fig u res repres en t in fa c t , s u c h a ta b u la t io n o f, s a y a t ho u s an d
fig u res , m a y
ma ke no i m p res s io n on the m in d wh i c h is a t a ll wor th men ti o n in g B y the .

gr ap h ic m et h od h o we
, v er, the c h ief c h a r a c teris tics o f a m ass of fig u r es a re
p resen ted to th e eye b y m ea n s of a p i c t u re o r c u rv e . T he g p g
ra i vhes ,

at a
g la n c e, im po r ta n t fa c ts w ic h m a
y b e o v er loo h ked , or wh ic h c an be
o b ta i n ed f ro m the fig u res o n l b y c o n s id er ab le la b o r y .

T he u se o f g p
r a h ic m et h od s in S ta t i s t ic s is v er ye x ten s iv e, an d has p ro v ed
to be of g rea t se r v i c e . I n fa c t , ev er yo ne wh o h as to d eal wi t h c o mp lic a ted

g ro u
p s o f figu res c o mes to a p prec ia te the g p
r a h ic m et h od , as it en a b les o n e

to p erc eiv e h o gh the eye

rela t io n s t r u .

I t is the o b j ec t o f t h i ec tio to h o w ho w gs S n s ra p hs a re fo rm ed f ro m giv en

d a ta .

h eq u en cy cu rv es . Let u s c o n s id er th e g r a p h o f the fo llo wi n g freq u e n c y

d is trib u t io n , in w ic h h the fi rs t l i n e o f th e ta b le g i v es the ma r ks of t he
c lass es , a n d t he s ec o n d
gi v es the n u m b er o f v a r ia tes i n the c lass es :

W h a t we p p
ro o s e to do he e r is to p re se n t a si
g n ifi c an t p i c t u re of th is

p p
o u la t io n .

D ra w two li n es , 0X an d O V, at r ig ht a n g les to ea c h o t er h F
( gi . I ) .

T he e s referen c e lin es a re c a lled c o ord i n a te a x es . T he l in e 0X is c alled

the x a x is , a n d
-
the lin e O l the y
’
-
a x is . T he po i n t 0, fro m w ic we m eas h h
u re, is c a l led the o ri g in , o r zero po i n t . B eg in n i n g a t t is po in t m ar o ff h k
the ax i s e u a l in terv a ls a s c a le c o n v e n ien t to the p rob lem a t
on x q u po n
-

ha nd . F ro m the s a m e zero po i n t la y O ff eq u al i n ter v a ls a ls o o n th e y -a x is .

Th es e n eed no t be on t he sa me s c a le a s t h o s e o n the x -a x is , b ut s ho u ld

be s u i ted to b es t b ri n g o u t th e fa c ts to b e S ho wn b y t he g r a p N ot h . all

g ra p h s a re d ra wn u po n the s a m e s c a le h
t erefo re, n o r a re the two axes of

the sa m e g p
ra h a li ke as to s pa c i n g o r s c a le .

I n th e pa r ti c u la r c a s e in a n d , let ea c h
in terv al a lo n g 0X rep resen t a h
h al f i n c h
T he q u es tio n as to w a t ea c i n ter v a l s a ll represen t is a ma tter
. h h h
of the s c a le us ed a nd t he s c a le must b e c ho s en to s u it th e p a r ti c u lar d a ta
in ha nd . N ex t, a lo n g 0X la y o ff t he c la s s m a r s C o rres po n d i n g to ea c h k .

c las s m a rk t he er is a freq u en c y
F ro m t he v a r io u s po i n ts o n the x -a x is
.

T h is d is t r ib u t io n is t a ken a s
1 a rep rese n t at iv e o f a n y freq u en c y di s t rib u t io n I t is . no t ,

ho wev er, m a d e u p a rt i fic ia y, b u t ll a ct ua ll y represen ts t he d is t ri b u t ion o f eight h und red ea rs

of corn wit h res pec t to l en gt h .
 A PPEN DI X 68 7

wh ic h rep resen t m ar k s la y o ff li n es p ara llel to the y a x is a n d o f

c la s s -

h
len g t s c o rres po n d i n g to the v a ri o us freq u en c ies , a c c o rd in g to the s c a le o n
th e y a x is
-
W en t is is d o n e t e re will b e a s eries o f pa rallel li nes a t
. h h h
eq u a l d is ta n c es a pa r t, a ll per pen d ic u la r to th e x a x is a n d pa r allel to the -

y
-
g h
ax is , b u t o f len t s c o r res po n d i n
g to th e v a r io u s fre u en c ies a n d therefo re
q
u ne u a l
q o
. i n i J
n g t he to p s o f t h e li n e s s o c o n s t r u c ted b y s t r a ig t l i n es h
g iv es th e f r eq u en cy p oly
g on s h own
i n F ig I T he to ps o f the lin es t u s
. . h
j o i ned g i ve an o rd erl y a r ra n gem en t o f po i n ts , t ro u
g w ic h i t m a y bhe h h
po ss ib le td ra w a s m o o th c u rv e I f i t is im pos s ib le to d ra w a s mo o t c u rv e
o . h
h h h h
t ro u g a ll o f t em , d ra w a s m oo t c u r v e a s n ea r as po ss i b le to a ll o f t em h
T he c u rv e so d ra wn is c alled a f r eq u en cy c u rv e ( n o t s own in figu re) h .

4 5 as 1 a 9

FIG . I

A n y po in t P i n the pl a n e r epr es en ts two n u m b ers : the o n e n u mb er is

rep rese n ted b y the d i t
s a n ce o f th e p o i n t f r o m t e y a x is , a n d the o t er b y
h -
h
i ts d is ta n c e fro m the x -ax is . The n u mb er w ich h gi v es the d is ta n c e o f P fro m
they -ax is is c a lled the a b s c is s a o f P , a nd the n u m b er w ich h gi v es i ts d is ta n c e
fro m th e x -ax is is c a l led i ts o r d i n a te . T h e two n u m b ers h
to g et er are o ften
s po k en o f as the c o ord i n a tes o f the po in t P .

S i gn ifi ca n ce of ar ea un d er curv e. C o n s t ru c t rec ta n g les su c h as A B CD

an d B C EF
’
on the o rd i n a tes a t c lass m a r s as m id li n es , m a k -
ki n
g th e s id es
A I ) , B C, etc .
, b is ec t the c lass i n terv a ls a lo n g the x a x is -
. S u p po s e, n o w,

h
t at we d efi n e u n it area as a rec ta n gle b o un d ed b y A B , A D , B C, a n d a
‘

lin e pa ra llel to A B a n d j u s t fa r en o u g h h
fr o m i t s o t a t the d is ta n c e
b etween A B a n d t is l in e represen ts h u i t f eq e c y
n T h e the a ea o f
r u n . n r

A B CD is 1 1 0 , a n d t h e a rea o f a ll s u c h rec ta gle t k en toget h e

n is eq al
s a r u

nu mer i c a lly to the to ta l p p

o u l a tio n . I nd ra wi g the m o th c n e me s o u rv n

t i o n ed a b o v e, we s ho u ld a im to ma k e the a ea b etwee the c e the

r n u rv
688 A PPEN D I X

x a x is , a n d
-
the two en d o rd i n a tes ( i n t is h c as e o rd i n ates a t 4 an d 1 0
) eq u al

to th e s u m o f the a reas o f t es e h rec ta n les

g . T he a rea u n d er th e c u rv e
then repres en ts the to ta l po p u la tio n . h
T is is im po r ta n t po in t, b ec a use
an

i t presen ts to th e eye ho w m u c o f the h p p

o u la t io n i s i n c l u d ed b e tween a ny

t wo m eas u rem en ts F or i ns ta n c e, a t a gla n c e yo u c o u ld es tim a te a p pro x i

.

m a tely th e po r tio n o f th e po pu lati o n d isc us s ed i n Fig 1 , w o s e meas u re . h

m en ts a re b etween 5 a n d 8 T he u se o f the a rea u nd er the freq u en c y c u rv e
.

will b e fo u n d h elpf ul in our d isc uss io n of p r o b a b le err o r .

Choice of sca le I n d ra wi n g a g rap . t he q u es tio n a lways a ri ses as to h

h
w a t s c a le s h a ll b e u sed i n plo t tin g, b u t u n fo r t u n ately n o d efi n i te r u le c a n
b e la id d o wn I t ma y,
. h owe e v r, p ro v e u s efu l to c a ll a tten t io n to a few

po i n ts . Firs t , we s o u ld h c h oo s e suc h a s c a le t h at we c a n p lo t a ll the po in ts

on one p a
g e o f t h e pa per u sed ; fo r i t i s a r v a g ea t a d a n t g e to h a e th e v

w h o le g rap h o n one pa pe r , t h us ma k in g i t v is ib le to the eye i n i ts en ti r ety .

S ec o n d , i f the po i n t i n v o lv ed i n the i n v es tiga t io n is a q u es t io n o f ra te o f

i n c reas e o r d ec reas e, we s o u ld s elec t s u c a s c a le as to m a k e the c u rv e
h h
r eas o n ab l y s teep . F req u en c y c u rv es are u s ed a g r ea t d ea l in the s tu d y o f th e
so c ia l s c ien c es , as well as in n a tu r al s c ien c e . F o r i n s ta n c e, the so c io lo gi s t

p r es e n ts the po p u lat io n o f a c i t y o r c o u n tr y fo r s u c c ess iv e y ea rs by us i n g

y ea rs as the mar ks o f c las ses , -

la yi n g t h ese o ff a lo n g the x -ax i s, — a nd

th e p p
o u l a t io n fo r t h ese y e a rs a s o rd i n a tes .

N egat iv e v a lu es ea s il y represented graph i ca ll y We . o ften d es ire to p lo t

ne
g a t iv e v a lu es as well as po s i ti v e v a lu es , a n d t is is h eas ily a c c o m p lis ed h
by a S lig ht wh a t has a lread y b een d o n e i n c o n n ec t io n wi th
ex tens io n o f

F ig 1 . W i th the d a ta ex h ib ited in Fig 1 it migh t h av e b een fo u n d c o n

. .

v en ien t to u se the m ea n as the o ri i n a n d to

g p lo t t he freq u en c y wi t h res pec t
to d ev ia tio n s fro m th e m ea n S i n c e t he m ea n is i n t h is c ase
. we h av e
the fo llo wi n g s et o f d ev ia t io n s a n d c o rres po n d i n g freq u en c ies to p lo t :

We s ho u l d now la y o ff t h e p o s i t iv e d ev ia t io n s to ward the rig t fro m the h

o ri
g i n 0
( F ig . 2
) a nd the ne
g a tiv e d ev ia tio n s to wa rd the left fro m O T he .

f req u en c ies s ho u ld , o f c o u rs e, b e plo t ted u pwa rd fro m X X , j u s t as i n F ig 1

’
. .

T he r es u l t o f
p lo t t i n g t is h y
freq u en c d is trib u t io n is s own i n F ig 2 T is h . . h
h
s o u ld b ri n g h
o m e to the rea d er , who is n o t v er
y fa m i li a r wi t th e h u se o f

n eg a tiv e n u m b ers , the fa c t t a t n eg a ti v e h n u m b ers m a

y b e j u s t as real

a n d u s e fu l as p o s i t i v e n u mb e rs .

T he freq u en c y po lygo o f Fig 2 n . d o es n ot d iffer i n fo rm fro m t h at of

F ig . 1 . I t is on ly d iff e e tly ela ted

r n r to th e li n es o f referen c e OX an d OI ’
.

Grap hi ca l mea nin g of m ed ian , mean , an d mode . I f i n F ig . 1 we selec t o n

the c u r v e a po i n t wh o s e o rd i n a te d i v id es the a rea u n d er the c u rv e i n to t wo

eq u al p a r ts, t he a b s c iss a o f h
t is po i n t is t he m edia n o f the po p u la ti o n . T he
6 90 A PPE N D IX

Let us ta e as k illu s tra t io n th e p lo t t in g o f the gr a p

an h of y 2x 4 .

Here we see h
fro m the eq u a t io n t a t c o r res po n d in g to a n y v a lue ass ig n ed

to x we get a v a lu e Of
y eq u a l t o twi c e x p lu s 4 . T he c o r res po n d i n g v a l u es

are as fo llo ws :

y
I
4
l 6
j 8
I 10
j 1 2 1 4 + 2 0 l

Lo c a t i n g , in Fig 3 , the po in ts c o rres po n d i n g to t

. h e e l es a d
s va u , n d ra w
h
in g a s m o o t c u rv e t ro u g t em , we a v e the g ra ph h h h h o f the f c tio un n . h
T is

g p
ra h is a s t r a ig h t li n e.

W e lea v e as a n ex er c is e fo r the s t u d en t to fi n d the g ra p h ofy x

9
. Fo r
a pp lic a ti o n o f g r ap h of fu n c t io n , s ee Pro b ab ili ty C u rv e, S ec tion V I .

S EC T I ON IV S M OO T H I N G OF F I G U R E S

S o meti mes t he f req u en c y d is tri b u tio n of a p p

o u l a t io n a r r a n ged with
res pec t to so m e c ha ra c ter ha s m a n y s m a ll irregu la ri t ies w ic h h a r is e merely
fro m the way in w ic h h the meas ur emen ts were ta k en and gr o u ped . In
su c h a c as e a p ro c ess g c a lled “ sm
oo t hi n
”
c an o ften be em p lo y ed to

o b t a i n reg u la ri t A n o tewo r t y . hy
in s ta n c e o f s m oo t in g is to b e s een i n the h
a d j u s tin g o f the p o p u la t io n c en s u s wi t res p e c t to a ge, t ere b ei n g a gr ea t h h
m an ym ore peo p le who rep o r t t ei r a ges a s 4 0 t h h a a 39 o r 4 1 I n
n s .

fa c t, S o m etim es the u n s m o o t ed fi g u res S o w o n e h h h l f mo re peo ple o f

a

h
ag e 4 0 t a n o f ag e 3 9 o r 4 1 I t is , then , m an ifes tl . y d e i ab le to s moo t h
s r

h
t es e c en s u s ret u rn s if t h ey a re to g i ve ev en an a p p ro x i m atel
y c o rrec t

im p res s io n .

In rep res en t i n g s u c h a di s tri b u tio n

g p
ra h ic a lly we ha ve to d ra w a s m oo t h
lin e in the h b o rh o o d o f the po i t
n eig n s, b ut n o t n ec ess a ri l y th ro g h u an y o f

h
t em . T h is s moo t h li e i the es u lt n s r o f t he a ttem p t to p resen t wh a t the
d is t r ib u tio n wo uld b e i f the c a u s es of the s m a ll i rregu lar it ies c o u ld be
rem ov ed . S o metim es i t is c o n v en i en t to s m oo t h fig u res wit ho u t reso r ti n g
to a g p
r a h . T he e r a re so m e ra t he r refi n ed but c o m plic a ted a lgeb r a ic

m et h od s
1
of d o in g th is , b u t i n gen er a l a v ery s i m p le m et o d c an b e used h .

To ex p la in t is m et o d , t a k e th e fo llo wi n g freq uen c y d is tr ib u tio n ( wh i c h

h h
was o b ta in ed b y m eas u r in g the c ir c u m feren c es of 99 5 ea rs o f c o r n
) ,
i n whi c h
the g rou p i n g s i n to gi nc
-
h c la s s es a re n o t well s elec ted .

1 Da rwin , P hi los ophi c a l h l ag a zi n e and jou r n a l , Ju y, l 1 87 7 .

 A PPE N D I X 69 1

I t m a y b e well to expla i n th e c ief so u r c e o f t is i rregu la r i t T his h h y .

c an b e s e en b y o b s erv i n g two c las s es ,

s u c as t he 7-
i n c c las s a n d th e h h
i n c c la s s
-
h
A s the meas u remen ts were rec o rd ed to the n ea res t ten t
. h
inc h , the 7 i n c -
h c lass i n c l u d es the m eas u rem en ts rec o rd ed a s 7, and

wh ile the -
in c h c las s in c lud es onl y th ose rec o r d ed as and

T ish sh o ld u e v id en tl
y p r od u c e a b ias ed res u l t . I n s tead of m a k in g a n ew

freq u e y t
nc a b le wit h d iffer en t g ro u p in g , we ma y s u b s ti tu te fo r ea c
a h
freq u e cy n a n u m b er h
d er iv ed b y s m oo t in g T is s m oo thi n g c a n b e a c c o m . h
p lis h ed b y s u b s ti tu ti n
g fo r ea c h fr eq u e n c ,
y ex c ep t the two ex trem e o n es ,

th e m ea n o f the g i v en freq u en c y a n d the im med ia tely b e fo re

o ne a nd

the o n e i m m ed ia tely a fter it . T h us , fo r freq u en c y o f ea rs o f len g t h

z + 4 + u
in c he s we s h o u ld s ub s t i t u te Bu t as h
t is is o n ly a n a pp ro x i
3
m a tio n , we m a y a s well ta k e the n ea res t i n teg ra l v a l u e, o r 6 F o r a n ex t rem e .

freq u en c y , we s u b s titu te t he mea n ( to n ea res t i n teger ) o f the ex trem e fre

q u e n ey t a k en t wic e a n d th e ad a c e n t j freq u en c y ta ken once . T h us , fo r th e
2 2 4
freq u en c y c o r res po n d in g to leng t h4 .
5 i n c h es we s u b s t i t u te

or, in in tegra l n u mb ers , 3 . so m et im es d es i rab le to a pp l

I t is y t is
p r o c ess h
mo re h
t a n o n c e to a g iv en d is t r ib u t io n i n o rd er to
g i v e i t t he d es i red

reg u la ri t y .

T he res ul ts of the s c em e h fo r the g iv en fre uen c

q y d is t r ib u tio n a re a s

follo ws :

— —

4 1
2
‘
T
|6 1
I n c hes .
S0o
5 5 75 6 0 . z $6 5 75 7 25 7 5 7 75 8 0 8 2s 3 5
r

\ '
U n s mo o t hed I
Ireq u enc nes
z 4
'i 13 24 20 74 1 1 5 98 13 98 w8 1 55 67 IO m

13 1 s m oo the d
6
W

? Li
1 26 162

w
3 '4 19 30 73 I ZO " 0
freq u en c ies

1
1
2d s m o o t hed
freq u en c ies 4 l 8 13 24 43 70 10 2 1 20 14 1 1 38
_ l _ l _ L

I n gen era l al
g eb ra ic term s , i f v
"
v
z,
. v ,,
a re th e m a r k s o f c l as s es a n d

a
,,
a
2,
a t he c o r res po n d in g freq u en c ies , i n s moo t in g th e h ’
a s we s ho u ld

s u b s t i t u te fo r t h em th e fo llo win g v a lu es r es pe c t iv ely :

a l + a + a + 11
,
+ a a
0 0 .

i i

It c an be eas il
y fro m t h
s een ese a l eb ra ic ex p ress io n s t h a t the a ri t h
g
m etic m ea n o f the m eas u remen ts is sc a r c el y a ffec ted a t a ll b s mo o t h in ,
y g
b ut t hat t he m o d e is s o met i m es c o n s i d e ra b l y c ha n ged . 1 1
g e n er a l, t he
“ s an
t d a r d d ev ia t io n ”
( t o b e d is c u s s ed i n S ec tio n V I I ) is b u t 5
'i
g h t l y
a ffec ted b y s m oo t i n g h .
69 2 A PPE N D I X

S E CT I ON V — A P PL I C A T I O N OF T HE T H E O R Y OF
PR O B A B I L I T Y

T he rea d er s ho u ld u n d ers ta n d t ho ro u g h ly t h at w h at is c om mo nl
y k n o wn

as a la w o f n a t u re is a g en e r a liza t io n b as ed u po n ex per ien c e, a n d th a t ch su

a la w c a n n o t b e p ro v ed in th e s t r ic t l y lo g i c a l s en s e , b ut o nl y i n t he s en s e o f

e s tab lis hi ng a h igh

d eg ree o f p ro b a b ili t y i n i ts fa v o r T o il l u s t ra te, we m a y .

ta k e o ne o f the b e s t -
h
es ta b lis ed fa c ts o f p ys i c a l s c ie n c e, n a m el , t a t a ll free h y h
b od ies a re a tt ra c ted b y t he ea r t h . T he ev id en c e fo r t is h s ta tem en t c o n s is ts

in t he fa c t t h a t th e th o u s a n d s a n d ev e n m illio n s o f b o d ies w ic h h ha ve b een

o b se rv ed h a v e, wit ho u t e x c ep tio n , fo llo wed t is h ru le . h
T is ha s es ta b lis h ed
a v er y h ig h d egree of h c o ei a b le h o we e
p ro b a b ili t y . I t is a l t o g et er nc v , v r,

th t a m e b od y e i t wh i h w ld b e
so p
x s s
e lle d b y t h e c e t h A lt
ouh o g h re ar . u

e pe ien c e h a
x r e t b li h ed a o s wh elm i g p b b ility ag i t
s a s n h a
v er n ro a a ns suc n

o c e c e we m
c u rr n t t e l
, k the f c t th t e pe i e h p ed the
us no ov r oo a a x r en c as ro v

s ta tem e t ly i the
n e th a t i t h a
on n e t b li h d sens h igh d g f p ob s s a s e a e ree o r a

b ili f y It h. d e th i d as d on th i g m o th
s an th i It i d
c an b tf l
o no n re an s . s ou u

wh eth e a y pe r li i g h
n ee h n d ed p i t ed a t
rs o n v n as S dom n a u r e n n es oss ran ,

a ll o f wh i c h c m e h ea d p a d t i
a l l i t i p o ib le t h
s u
a t t h,i m ig h
n t h a s
pp e n s ss s .

E e if n o
v n e h e th em f ll wi th
on as ll h e d
s en
p w e a re l e l y
a n o t a a s u , c ar

j t ifi d i c o l d i g th t t h e will a lw y b e me h d p a d o m e
us e n nc u n a er a s so ea s u n s

ta i ls p Iuf a t h. d pe ie b e t d
o us a n
e t n d o m t h nn
e p b a b i
s li ty th t o ss a ra , ro a

t h ey will a ll fa ll h e d p i O s m l l t h t w
a e ms u
y fe l ys S
sa y i f t h e w h o lea a a sa ,

h m a ra c e we e to d e o te ge e ti t th e to i g f p i s th o usa n d
u n r v a n ra on o ss n o en n e , a

a t a ti m e th e e w ld till b e e y
r m a ll p b a b ili t y th a t
ou s a v e wo ld r
, y o s ro an n u

to ss a ll h e d A ll th i go e t
a s h w th a t c e ta i po ib le e e t h a e u c h
. s s o s o r n ss v n s v s

a l igh t p o b a b ili t y t h t we h
S r ld o t e pec t th em to h pp en i the l i feti m e
a s ou n x a n

O f a gi en i d i id a l
v J t o t ime g
n v u . a d ob e us s tio s a re m a d e i
, as o es o n n s rv a n n

g re a te r n u m b e r s , e x c ep t io n s m a y b e fo u n d t o a n y o f t h e s o c a l led - “ la ws of

n a tu re
”
. h
S uc ex c ep tio n s a re, o wev er, i n m a n y c a s es ex c eed in g ly h u n li k ely .

h
I t is h o ped t a t the fo rego i n g p a v es the wa y fo r th e s t a tem en t th at ,

h h
w ile i n t is s u b j ec t m a n res u lts m a y b e s ta ted i n term s y of p ro b a b i li ties ,
th e e s res u l ts do not d i ffe r in reli a b ili t y on t h t a a c c o un t fro m h
t ose o f an y

o th e r s c ien c e b as ed on e x per ien c e . If a th o us a nd p en n ies b e tossed at

ra nd o m , h
t ere is not hi ng mo re u n c ert a i n th a n h
t at g
a i v en pen n
y w ill
b e hea d s, b u t it is a m a t ter o f c o m m o n ex pe rien c e h
t a t th e ra t io o f the
n u m b er o f h ea d s to the to t a l n u m b er o f pe n n ies to s s d
is , i n gen eral, n ea rly
e

o ne h a lf . We may he re rec a ll the s ta te m en t o f S ec tio n I , t a t th e t eo r

y h h
of p ro b a b i li t y is n eed ed i n t is h s u b ec j t b ec a u se we d ea l with o c c u rren c es

a nd c h h
a ra c t ers o f s u c
t th t we wi h to m k e t teme t i n ega d
a n a u re a s a s a n s r r

to la ge
a mbe
r f t h em t k e t g t h e
nu It i
r o m a t ter o f c o m mo experi
a n o e r . s a n

e nce th a t e lt h as r su g a d a t i
s, suc
o O b t i e d f m l g e
a v e ra
a n u m b ers es n r s a n ro r

o f c e e n
as s , a r ly t t io a y W e fi d the a e age t t re o f a t h o an d
ea r s a n r . n v r s a u us

i d i id
n vl el c t d
ua s st do m f m l g e p p l t io
e e a a d a e much
ra n ro a ar o u a n, n r su r

p r i ed si f p o t k i g
, o t
u h e d n
o m m a
p le f
n a t h oan a d f ro m t h
r ra n
e a me sa o us n s
6 94 A PPEN DI X

th o us a n d peo p le t i rty t h h o usa n d a re v a c c in a ted , a n d t at h fiv e h u n d red pe r

so ns ha v e s m a llpo x . If v a c c in a t io n ha s no in fl u en c e o n the n u m b er o f c as es

o f s m a llpo x , w h at is the p ro b ab ilit y th a t a pe rso n will b e b o t h v ac c i n a ted

an d h av e s m a llpo x ?

S in c e o ne h un d red tho us a n d is a large n u m b er, we m a y g iv e

p ro b a b ilit y th at a perso n is v a c c i n a ted

p ro b a b i l i t y th at a pers o n h as s m a llpo x ;

p ro b a b ili t y th at a p e r s on is b o t h v a c c i na te d

a nd ha s s m a llpo x .

3
F u rt h erm o re,
5000
x 1 000 00 1 5 0 , t he n u mb er o f pe rs o n s we s h o u ld e x

peet b o th to b e v ac c in a ted a nd to h a v e s m a llpo x , if v a c c i n a t io n ha s no

in fl u en c e o n the n u m b er o f c a s es o f s m a llpo x .

Ill u st rations of p rob a b il it y . L et us th r ow ou t u p on a ta ble a t r a n d om f ou r

p en n ies ; wh a t is the pr oba b i l i ty tha t ex a c t ly r c f them wi ll be hea d s and the

r es t ta i ls when r ta hes v a l u es 0, 1 2, 3 4 t , ,

( ) 1 Pro b ab ilit y th a t 0 will b e head a d 4 tail i n s s

(D
4

( )
2 P o b a b ili ty t h t 1 wi ll b e h e d a d 3 ta i l i
r a a n s s

( 3) P o b a b ili t y th a t 2 will b e h e d a d 2 t il i
r a s n a s s

(4) P b b i li t y t h t 3 will b e h d a d 1 t il i
ro a a ea s n a s

( 5) P o b b ility th a t 4 will b h e d
r a d 0 t il i e a s an a s

In ( ) 2 th e c oefli c ien t 4 a p pea rs b e fo re b ec a u se wit h fo u r c o in s h

t ere

a re fo u r d iff eren t c o mb in a tio n s po s s ib le 1

, ea c h c o n s is tin g o f 1 h ea d and 3
ta ils . y
S im ila rl in ( 3 ) the c oe ffi c ie n t 6 a ppea rs b e c a u se w i t h fo u r c o ins

h
t ere a re po s s ib le s ix c o m b i n a tio n s , ea c h c o n s is t in g o f 2 he ad s a n d 2 t a ils .

T he ab ov e illu s t rat io n m a y b e gen era l ized a nd the res u l t

p u t in to the
fo llo win g fo rm
If n c o i n s a re h ow
t r n u po n a ta b le a t ran d o m , the p ro b a b ilit y t h a t ex ac tl
y
r of t h em will b e he d a s and the res t ta ils is g iv en b y the r + I s t ter m o f

the

the
b i n o mia l

m b ol
"
ex pa n s io n

C in d ic a tes th e
(2 a )
n um b er
t h at i s, in ot

o f c o m b in a tio n s o f n
h er s b o ls ,

h
t in gs ta en r
wh ere

k
s
y ,

at a t im e .

In or d er th a t the read er m a y mo re fu ll y a pp rec ia te the g rea ter p ro b

ab il it y Of ge t t i n
g a n a lm o s t eq u a l n u m b er of he da s a nd ta ils i n to ss i n g

pen n ies t h an of g etwid ely d i fferen t n u mb ers , we p resen t the fo llo win g
t i n
g
tab le fo r Ob ta in ed fro m Q u e telet S u r la t h eo r ie d es p ro b a b il i tés
n 999 , .

1 Fo r d efi n itio n o f a co m b in at io n , see text, p .

511 .
 A PPE N D I X 69 5

C o lu m n s 1 a n d 2 gi v e the n u mb er o f eads an d ta i ls whose p ro b ab ili ty h

is i n q u es tio n C o lu m n 3 g iv es the prob a b ili t o f exa c tly th e n u m b er o f
. y
h ea d s a n d t ai ls i n d i c a ted i n c o lu m n s 1 a n d 2 .

HE AD S T AL
I S P R BAB L T
O I I Y

”
mp
Ca ” : 0 02 s 4 50
“ C 0 0 2 1 069
QP
”
m :
4 40
0-0 1 1 7 94 4 3°
00 0 44 2 3 420
1 10

A gla n c e a t the ta b le s o ws t h h at , in the lo n g ru n , o ne S ho u ld ex pec t 4 99

hea d s a n d 5 00 t a ils mo re t a n h t imes as o ften as 4 2 0 h ead s a n d 5 79
ta ils . h
I n t is c o n n ec tio n it is i n teres t in g to i n q u i re in to a c as e men tion ed
on p g
a e 69 2 , n a mel , t he y p ro b a b i li t o f ge t y
t i n g a ll ea d s in a S i n le th ro w
g h
Of a h
t o u sa n d pe n n ies h
T is p ro b a b ili t
. y is

1
—

( an i n teg e r c o n ta i n in g 3 0 2 fi g u res )

and th e s t a te m e n t m ad e on p g
a e 6 92 as to the h u ma n r ac e
( p p
o u l atio n
1, d ev o tin g i tse l f to to ss i n g pen n ies is c lea r l a s a fe s ta temen t y '
.

T he res u l ts i n th e a b o v e ta b le m ay well b e exh ib ited g ra p h ic a lly ( F ig .

FIG .
4
SC A L E : V er tica l. 1 in c h
Ho rizo n ta l 3 i nc h
, 10
6 96 A PPE N D I X

I f we h ad ta k en a ll the in term ed ia te in tege rs fro m 4 99 wit 5 00 to h

wi t h 5 59 , we s h o u ld h av e h ad ten tim es as man p o iyn ts w i c wo u ld h h
a rr an ge th em selv es a lo n g the c u rv e i n Fig 4 B y i n c r eas i n g the n u mb er
. .

o f c o i ns and d ec reas i n g the h o r i zo n ta l s c a le , we c a n get the po i n ts a s

c lo s e to get e r as h
w e p leas e T is c u rv e i n. F ig 4 ih
s t he so -
c a lled p ro b .

a b ili ty c u rv e a n d i t a ppro a c h es v er n ea r l y y
the c u rv e o f er ro r , o r n o r m al
h
freq uen c y c u rv e, w ic h we are n o w p rep a red to d is c u ss .

S EC T I O N V I — N OR MA L PR O B A B I L I T Y C U R V E

I t has b een fo u n d t h at t he freq u en c y e

c u rv s of a g r ea t ma n y b io lo g i c a l
meas u remen ts fo llo w a c u r v e v a r io u s ly k n o w n as th e p ro b a b i l i t yc u rv e ,
”

n o rm a l p ro b a b i l i t y c u rv e,
” “
c u rv e of er ro r ,
”
or n o r m al freq u en c y
c u rv e .
”
I n p a r tic u la r i t is k n o wn as the c u rv e o f erro r,
”
b ec a use i f
er ro rs whi c h a n Ob serv er ma k es in a refi n ed s et o f d irec t meas u reme n ts
on g
a i v en
q u an ti t b e p lo t tey
d as a b s c iss as , th e c o rr es p o n d i n g o rd i n a tes

h
o f po i n ts o n t is c u rv e r ep resen t the freq u en c i es o r p r ob a b i li ti es o f the

erro rs .

T h e gen era l fo r m o f the c u rv e is s o wn i n F ig 5 h

The o rig i n is ta en . . k
a t th e m ea n h k
T en , i f a n y m a r o f a c la ss is a b o v e the mea n , i ts d ev ia
.

tio n is p o s i tiv e, a n d i t wo u ld b e p lo tted to the r i

g t fro m th e o ri in 0, whi le
g h
i f the m a r k o f a c la s s ha
th e m ea n , i ts d ev i a ti o n is n eg a tiv e, a n d
is les s t n

i t wo u ld b e p lo tted to the left fro m 0 Fo r t he b en efi t o f t ose who ar e . h

6 98 A PPEN D IX

Ho wev er , the n o rm a l c u rv es gi v e, i n g e n era l , at le a s t a v a lu a b le fi rs t

a pp ro x im a tio n , an d we s h a ll fo llo w the u s u a l m e t h od o f em p lo y in g s t a t is

ti c a l c o n s t an ts d er iv ed fro m t his c u rv e : fo r h
t es e c o n s t a n ts a r e si
g n i fi c an t,

1
ev en i f t h e d is tri b u t io n is no t n o rmal .

A rea u nd er the prob a b ili t y curv e has a n im port a nt m eani n g I f we se lec t .

u n i t a rea as ex pla i ned in S ec t io n I l l t he a rea r ep rese n ts t he to t a l po p u la t io n ,

,

a n d th e a rea b etwee n th e t wo or di n a tes . t he c u rv e, a n d r -

a x is r e p res e n ts the

n u m b er o f v a r ia tes b e t wee n t h e se o rd i n a te s . I f we loo k u po n th e c u rv e as

rep res e n t i n g p ro b a b i li t ie s in s tea d of freq u e n c ies our h o r izo n ta l sc a le is

u n c ha n g ed but our v er t i c a l s c a le m u s t b e m u l t i p l ied b y t he to t a l po p u la
t io n . T h
i f the po p u la t io n is
us, 8 00 , as i n t h e c a s e o f F ig l , we s o u ld sa y . h
h h
t a t w a t t ere rep rese n te h
u n i ty s d h oul d b e m u l t ip lied b y 8 00 in o r er t a t d h
it s h a ll rep res e n t un i t o f p ro b a b il i t y . h
T e n t he e n ti re a rea u n er t he c u r v e d
will b e u n it y , a nd the a rea b etween t wo or di n a t es , t he c u rv e, a n d r - ax is . is
s im p l y the p r o b a b ili t y h
t a t a v a r i a te s e lec te d at ra n d o m wo u ld lie with i n
t hi s in terv a l .

S EC T I O N V I I — PR O B A B L E E R R OR A N D S TA N D A R D
D EV I A T I O N

I f we h ave es ti m a te d the o
p p u la t io n of a c i ty at a nd h av e

g o o d re a s o n to t h i k t h t th
n a e c h a n c es a re ev e n t h at t hi s is c o r re c t h
wi t i n
t 000 , we g iv e m h m e i
uc or n fo rm a t io n by s t a t in g h
t a t the p p
o u l a t io n is
t oo , oo o ziz 1 00 0 th a b y gi i
n v ng m er el y th e fig u res a nd lea v in g the
rea d er en tirel y in d o u b t a s t o the a c c u ra c y o f th e d et er m in a t io n .

I n d es c ri b i n g a fr eq u en c y d i s t r i b u t i o n t he a v era ge g i v es a b so lu tel y no

id ea as to w h e t h er d e v ia t io n s a re la rge o r s m a ll , n ot hing in re
g a r d to th e
s
p r ea d of the d is t rib u t io n . I t is th e j
o b ec t o f t he “ s ta n d a rd d ev ia t io n to

b e d es c r ip t iv e of t hi s v a r ia b i li t y , a nd i t is the o b e c t j o f t he so - c a lled

p r o b a b le err o r
”
d ic t wh t o fid
to in i t b p l c ed i
a e a c n en c e s o e a n s ta t is t ic al

res u l ts . T he us e wh i h h b ee m d e o f b t h
c t d d de i
as n a o “
s an ar v a t io n a nd

p ro b a b le er r or m ke it a e y to d w ll l g o th i
s un n c essa r e on er n s po i n t , but
it is o ur p p
u r o s e h e e to h o w h w the f
r m las d i the t t o or u s u se n ex a re d eriv ed .

Prob a b le error of a s in gle v ariate . TIre p r oba b le er r or

of a s ta
g ]
: wa n k /e

pp u la t i o n is d efi n ed tha t depa r t u r e f r om fil e m ea n , eff/ber s i de,

‘

y
r a o as on

wi t/t i n f
a r /1 iclz ex a c tly on e I /ze era r ia tes a re
f ou n d .

B y the u se of t h e p ro b ab i li t y c u rv e F
( gi . 6 ) the pro b a b le er ro r m a y
ea s i l y be exp l a i n ed g eo m et r i c a lly wh e n we loo k u p o n the en ti re a rea u n d er

the c u rv e as rep resen t i n

g the to t a l p o p u la t io n . I n F ig 6 we d ra w t wo .

o rd i na tes , ST and S T
’ ’
, eq u a ll y d is ta n t fro m t he m ea n , a n d s u c t a t on e h h
h a lf o f the en t i re a rea u n d er the c u rv e lies b et ween t h em , in o t h er wo rd s ,

is b o u n d ed b y the c u r v e th e x a x is , S T, a n d T ,
- ’
. T he n 1 OS rep rese n ts th e

p r o b a b le er r o r o f a s i n
g le v ar ia te I f we s o u ld . h u se a s i n le v a ri a te s elec te
g d
a t ra n d o m to rep res en t the po p u la tio n i t is a n ev en c h a nc e h
t a t th a t s i n gle
v a r i a te wo u ld b e les s or m o re t ha n OS fro m the b es t v alu e.

1 See Y l u e, P r oc eed i n g s o
f Me R oy a l S oc iet y L X , 4 7 7 ,
-
489 .
 A PPEN D I X 6 99

An a p p r o x i m a t e v a lu e fo r the p ro b a b le er ro r o f a s i n g le v a r ia te in an y

p p
o u la t i on may b e ea s il y o b ta i n e d
in the fo llo wi n g m a n n e r
I . A r ra n ge the v a r ia tes in the or d er o f m ag n it u d e .

2 . Cou n t o ne fo u r t h of the v a r ia tes o f lea s t m eas u rem en ts a n d n o te the

m ea su rem en t of the u p per o ne of h
t ese v a ri a tes . Let 11 rep rese n t h
t is
m ea s u rem en t .

3 . Co u n t o ne fo u r t h of t he v a ria tes o f g r ea tes t meas u re m e n ts and n o te

th e m eas u rem en t of the lo wer o f th es e v a r ia tes . Let 7; re p res en t t hi s

m ea s u rem en t .

'
7 u
4 . T he n —
a g iv es the
p ro b a b le erro r o f a s in g le v a ria te .

T he fo r m u l afo r the p ro b a b le er ro r in a s i n gle v a r i a te is

w h ere 2 x
2
m ea n s the sum o f t he s q u a r es o f th e d ev ia t io n s fro m the m ea n
and n is the n u m b e r o f v a ri a t e s . T he c o n c ep t io n o f the p ro b a b le er ro r o f

0
'
S 8

FIG . 6

a s i n gle v a r ia te is o f v a lu e b ec a u s e i t a id s in the d eri v a t io n f the p ro b a b le

o

erro r o f o t he r i m p o r ta n t res u l ts . T h e fo rm u la fo r the s ta n d a r d

d ev ia tio n

18 ( p g
a e 429 ) so t h a t t he p ro b a b le e r ro r o f a s i n le v a r ia te is o
g b ta in ed
V n

fro m th e s ta n d a rd de v ia t io n o f t he p p
o u l a t io n b y m u l t iplyin g t h e s ta n d a rd
de v i a t io n b y
700 A PP E N D I X

A s h as b een po in ted o u t i n t he tex t , the s ta n d a rd d ev ia t io n g iv es a g o o d

id ea o f th e s prea d o f the d is tr ib u tio n F ro m the a c c o m p a n yi n g fo o t n o t e 1
.

we a re n o w in a p o s i t io n t o a p p rec i a te i ts m a th em a t i c a l si
g n i fic an c e . It
. 1 2
l
is the o
'
i n the eq u a tio n y “J a
e of th e n o rm al p ro b a b i l it y c u rv e,
k
an d b ea rs a s im ilar rela t io n to the p ro b ab i li ty h c u rv e t at t he r a d i u s o f a

c i rc le b ea rs to t he c i r c le . If a is sm a ll th e p o b a b ili t y r c u rv e is c ro wd ed

to g et e r h s o as to res em b le c u rv e A in Fig 7 wh ile i f i .

, a s la rge i t is s p re a d

o u t s o as to res e m b le c u rv e B i n F ig 7 . .

Hen c e the s ta n d a rd d ev ia t io n a lo n g wi t h the m ea n c o m p letel y d es c rib es

the d is trib u tio n w h en i t is

o f v a ri a b
n o rm al

y
.
”

a n o r m al
T he ex p ress io n
“
d is trib u tio n , is
E
a
, wh i c h is th u s a

p er fec t m eas u re ili t fo r a g oo d mea s u re

o f v a ria b i li t y wh e n t he di s t r ib u t i o n is n o t n o r m a l, b u t i t is n o t c o m p letel y

d es c ri p ti v e o f the p o p u la t io n .

A not he r m eas u re o f v a r ia b i li t y is s o m e ti m es u s ed wh ic h c o n s is ts s im p ly

k
i n ta i n g th e a ri t m e tic a l a v erage h of the n d ev ia t io n s , — t h ese d ev ia ti o n s
b ein g gi v en the po s i ti v e s ig n .

1 T he ro b a b i lit t ha t a s s tem o f n v a r ia tes

p y y l
fa l in in terv a l s x
,
to x
,
Ax , x
,
to x
, Ar,
x ,. to x“ Ax is gi v en b y
r
e
a x/2 ir

T his ma y be wr itten m ore b riefly as

e
1
on ( 2 tr ) ?

F or g
a i v en set o f d evi a t i o n s whic h oc c u r, 6 sh o u ld be s elec ted so as to ma ke the
ro b a b i li t P a m ax i m u m
p y .

E q ua tin g t he firs t deri va t i v e to zero , we o b ta i n

2x 3
n o ?

N o w, b y mea n s of i n tegra l l l
c a c u u s, tab l es a re fo r med of t he a rea in c l uded b y t he cu rv e
x 2

2 03
,

a 2 r

the x -a x is , a n d a n y t wo o rd in a t es at eq ua l di t s a n c es i a fro m t he mea n . S uc h a ta b l e wit h

t he a rgu men t
x

0
'
is fo u n d i n D a v en po rt
‘

s S t a t is t ica l Met hods , seco n d edi ti on , pp . ri g ms

-
.

T his t ab l e s ho ws t ha t
0 -6 7 4
5 ( l)
when ju s t o n e ha f l of t he a rea u n der the c u r v e is i nc l u d ed a s desc rib ed ab ov e . By defi n it io n ,
t he pa rt ic u l l
a r v a ue o f t g i v en b y
( ) is
i ca ll d e l
t he pro b a b e error i n a s in g e va ri a te, a n d l we
s ha ll re p res en t it b y I f
Hen c e , E; . 6 74 5 d .

3 l
S ee G a t o n , Na t u ra lI n heri ta n c e, p 6.2 .
7 0 2 A PPE N D I X

d iff e r b u t s li
g h tl y fr o m ea c h o t he r, b u t i f th e m eas u re m en ts a re s u ffi c ien t l
y
a c c u ra te h
t e re will b e so me d iffe e r n c es . I f we s h o ld
u fi n d the m ea n of

h
t es e m ea n s ( w e m a
y c a ll it t he s e c o n d m ea n
) we c o u ld p lo t a f re q u e n cy

c u rv e o f t h e d is tr ib u t io n o f m ea n s j u s t a s we a v e fo r t he d ev ia t io n s h
o f th e o r ig in a l v a ri a tes . Of c o u r se h t is c u rv e wo u l d be mu c h c ro wd ed

to get er, h li ke .
A
I of F ig .
7 . T o ma k e th i s
gen e ra l, wi t h a v er y
la rge po p u
la tio n , and wi t h n in ea c h g ro u p i n s tea d of 1 00 0 , t he fo llo wi n g res ult is
o b ta i n ed :

I f I f , is t he p ro b a b le e rro r of a s i n g le v a r ia te, h
t a t o f th e m ea n of n

v a ri a tes is

t h at is , to fi n d the p ro b a b le e rro r o f the m ea n , d i v i d e t/ w p r oba ble er r o r

s i n le v a r i a te by f h m b of
o
f a
g s u a r e r o ot (
q t e n u er

Prob a b le err or in st a nd a rd d ev ia t ion . T a in gk u p a ga in t he m illio n c as es

o f s ta t u re d iv id ed i n to a t h o u s a nd g ro u p s a s a n ill u s t ra t io n , s u p po s in
g t h at

the s ta n d a rd d ev ia tio n of ea c h o f t es e h h
t o u s a nd g r o u ps b e fo u n d , we
s h o u ld s ee h h
t a t t ey d i ffer b u t s li
g h tly H w . o e v e r, i f the c o m p u ta tio n s a n d

m ea s u re m e n ts b e
v er y re fi n ed th e wil l b
er e d ev ia t io n s . T he e s s ta n d ard
d ev ia t io n s c o n s t it u te a freq u e n c y d is t rib u t io n w h ose s ta n d a r d d ev ia t io n
c an b e fo u n d , a nd t he p r o b a b le e r ro r of t h e s ta n d a r d d ev ia t io n c an be
ob ta in ed j ust as we h av e s h o wn in the c a s e o f a s i n g le v a r ia te .

G en er a lizi n g t hi s so as to h a v e a v er y la rge n u m b er o f gro u ps ea c c o n h

tai n i n g n v a r i a tes t a ke n as a s a m p le t o rep res en t the p p
o u la tio n , the p ob
r

a b le e rro r o f the s t a n d a rd d ev ia t io n is

t h is , to fi n d th e p r o b a b le erro r i n
at t he s ta n d a rd d ev ia tio n , d iv i de Me
fir oo
/
a b /e er r or i n M e m ea n by x o
[
Formu las for p rob a b le error in some im portant sta tist ica l con sta nt s E n o u g . h
d
h a s n o w b ee n s a i to g i v e the c o n c e p t io n o f t h e p ro b a b le erro r i n a n y
s ta t is t i c a l d e te rm i n a t io n a nd a g e n er a l n o t io n o f the m et hod s b y whic h
fo r m u las fo r the p r o b a b le e r ro r a r e d er iv ed .

I t is s c a r c el
y n ec es s a r y to r em a r k t h at th e p ro b a b le er r o r d o es not ta e k
i n to k e eith er o f b e a tio n o r c o m p u ta t io n W e
ac c o u n t ev id en t m is ta s o s rv .

ar e s u m i g th a t th e e h a e b ee
as n elim i a t ed I t h as to d o with erro rs
s v n n .

( d e i a tio
v
) d e to
ns a i d eufi n i tely l a ge u mb e r o f
n n ass i n a b le c a u se
n
g r n u s

su c h t h t the er o
a a e d i t i b t ed a c c o d i n
r rs
g t o th
r l w o f p ro b a
s rb i l i t y
u r e a s .

I t ee m s n e e s y to c o n t in e the d i c u io n o f p ro b a b le erro r in
u n c ssa r u s ss

o th e d te m i
r eti r b t i t d o e eem well to c o l lec t to ge th er fo r p rpos e
na o ns , u s s , u s

of refe r en c e , fo r m u la s fo r the p r ob a b le
the er ro r in so me of th e mo s t
i m po r t a n t s t a t is t i c a l c o n s ta n ts .
 A PP E N D I X 7 3
0

In w h at fo llo ws

a is to r ep res en t th e s ta n d a r d d ev ia tio n
n i s to r ep res en t the n u m b er o f v a r i a tes ;
e is to r epres en t t he c oe ffi c ien t o f v a ri a b ilit y;
r is to rep res en t t he c o effi c ien t o f c o r rela tio n .

n r:
p r o b a b le err o r in a s i n g le o b s erv a t io n .

E 0
p r o b a b le er ro r in the m ea n .

v” v ”

E "l O . 6 74 5 a
o b a b le err o r in s ta n d a rd d ev ia tio n
p r .

V 2 V2 n

74 5
o b b le er ro r in c o e fii c ien t o f v an a
[
t 2 p r a
7 [1
b i li t y

a p p ro x i m a tel
y, i f C i s no t g rea ter h
t an t o p er c e n t .

p r o b a b le e r ro r in c oe ffi c ien t o f c o r rela t io n .

p ro b a b l e e rro r i n the reg res s io n c o effi c ien t

’
0
r
"
2

S EC T I O N V I I I — C O R R E LA T I O N T HE O R Y

Defi n it ion . T7 00 m ea s u r a ble e/ mr a e/er s of an i n d i v id u a l, or

of r ela ted
’ ’
in d i v i d u a l s . a r e s a id to be eor r e/a l ea if to a s elee/ea s er i es of s i z es
of !b e on e

fli er e cor r espon d s i z es of ( b e oI /t er wll os e m ea n v a l u es a r e jun c t i ons o f Me

s elec ted v a l u es . T he wo rd “ s i zes "

, he e r u s ed , s h o u ld b e ta ke n to m ea n
n u m er ic a l m ea s u re ”
.

Fo r the k e o f c o n c reten ess a n d s i m plic it y we m a y th in k o f m ea u rin g

sa ,
s

t he c o rela t io o f s o
r n wi t h re pec t t th i fa th e
ns T o ren d er the b o e
s o e r rs . a v

d efi n itio i n y m b o lic la ngu age a d to d e el p the m eth o d o f d eterm in in g

n s n v o

th e fu n c ti o n m en tio n ed in the d efi n i tio n a re th e fi rs t po i n ts in the a p p lic a tio n

of m at h em a ti c s to the t eo r h y o f c o r rela tio n . h
F o r t is pu rpo s e let :r a n d y ,

represe n t v a ria b les s u c h th at y (

M ) r g iv es the m ea n v a lu e o f y c o r res po n d
in g to a s elec ted r . T he n t he p r o b lem is to d eter m i n e (
M r
) .

S u pp o s e the fo llo wi n g s y s te m of c o rres po n d in g v a lu es res u l ts fro m

meas u rem en t : wh ere n is a v er y la rge
n u m b er i n d i c a t in g t he p p
o u la t io n of fa t he rs a nd c o r res po n d i n g so n s .

T he e
s o b s erv a t io n s a re s a id to fo r m a to t al p p
o u l a t io n or u n iv ers e of

o b se rv a t io n s . A s i t will b e m o re c o n v en ien t to d ea l wi t h t he d ev ia tio n s

of the o b s erv a t io n s fro m t h ei r m ea n v a lu e t anh wi t h t he meas u rem e n ts
h
t em s elv es , let ( x l , y l ) , y ” ) rep resen t the d ev ia t ion s of
 7 4
0 A PP E N D I X

t he o b s erv a t io n s fro m t h ei r m ea n v a lu e . T h ese d ev ia t io n s m a y b e con

v en ien t ly rep res en te d wit h res pec t to c o o rd i n a t e a x es F

( g i .

B y th e ra n g e a lo n g t he x a x is we s a ll m ea n s u c a n i n terv a l t ha t o rd i
-
h h
n a tes d ra wn a t t he ex t rem i ties o f the i n ter v a l i n c lu d e b e t wee n t em t he h
to ta l po p u la t io n . T h us , i n F ig 8 , the ra n ge is t a
. ken fro m a to b . T is h
r a n ge m a y well b e d i v id ed i n to so m e n u mb e r, s a y s, of e
q u a l p a r ts , ea c h
of wid t h A x , b y o rd i n a tes a t th e p o in ts o f iv is io n l f we let x ,
g
d .
’

. r ,
’
be th e a b s c is s a s o f th e feet o f t he o rd in a tes t ro u g the m id d le po i n ts h h
of th e s c la s s es , we s h all c a ll t he e s t he mar k s of th e c las s es o f y
’
s . T he
v a l u es o f
y w i c h h b elo n g to a
g i v en c las s o f r a r e s a i . d to fo rm a
y
-
a rr a y .

FIG 8 .

L e t the c r o s s es ( ) x in 8 rep res e n t th e m ea n s of th e y

’
s in ea c h of

the s - a rr a y s . If c o rrela t io n ex is ts , t he se m ea n s d o not lie a t ra n d o m o v er

t he field , b u t a rra n g e t h em s e l v es m o re or les s in the fo rm o f a s moot h

c u r v e c a lled the c u rv e o f reg res s io n .
”
T is h c u rv e is a c ru d e
p i c t u re o f t he

f u n c ti o n w i c h h d e fi n es t he c o r rela t io n of the y - ch c te a ra r re la t i v e to th e
. r c -
h a ra c ter . E x pe r ien c e h a s s ho wn t h at, i
n m a y se tn f s o m eas u remen ts ,
t hi s l i n e is a p p r o x i m a tely a s t ra ig ht l in e . F th i
or e s r as o n , a nd fo r s im

p lic i t y, t h e l i ne j
s u b e c ted to the c o n d i tio n t h at the su m o f th e s u a res o f th e
q
d e v ia tio n s ( m eas u red p a r a llel t o the y a x is -
and weig h ted wi t h n u m b er o f
p o i n ts in a rra y) of t he m ea n s fro m i t s h a ll be a m i n i m u m , is c a lled t he
“ lin e o f reg res s io n . W h en the m ea n s lie ex a c t l y on t he l i n e t he re
g res

s io n is sai d to be t ru l y l i n ea r .
”
7 0 6 A PP EN D I X

If r I , a ll the i n d iv id u a l p o in ts of th e p p
o u la t i on wi ll lie o n t he li n e
of r eg ress io n , and we c an h
t ere fo re, w h en o ne c h a r a c ter i s g iv en , t ell
ex a c t ly w h at th e a s s o c ia ted ch is i n m a g n i tu d e
a ra c ter In . h
t is c a s e t h e

c o r re la t io n is s a id to be p e r fec t p o s i t iv e c o r r e la t io n S i m i la rl . y if
, r t,

t h e c o r rela t io n wo u ld b e p e r fec t n eg a ti v e c o r rela t io n .

T hree v ari a b les . T he t h eo r y o f c o r re la t io n is ea s il ye x ten d ed t o a pply t o

m o re t h a n t wo v a r ia b les . Fo r ex a m p le, h
we m ig t i n v es t ig a te the c o r rela t io n
o f th e s ta t u res o f s o n s wi t h res pec t to the s ta t u res o f b o th p a re n ts T is is . h
th e c a s e o f b ipa re n t a l i n h e it r ance t rea ted i n the tex t, p a ge 5 2 9, an d t he fo r
m u las th e re u s ed m u s t b e s pec ia l c a s es h
o f t o s e w ic h h
we a re a b o u t to d e riv e
fo r g i v in g th e m o s t p ro b a b le v a lue o f a v a r ia b le wh e e
3 r 3 is the n u m e ri c a l

v a lu e o f a c a ra c te r h c o r rela te d wi t h h c a ra c te rs of m ea s u rem en ts 1
.
'

and
y .

S u p po se th at t he fo llo wi n g s
y s te m o f c o rres po n d in
g d e v i a t io n s fro m the
m ea n s h a v e res u l ted fro m m e as u rem en t : ( my s,
r

R ep res en t t h es e m ea s u re m e n ts wi t h res p ec t to c o or i n a te d
ax es in t h ree d im en s io n s . Th es e a x es e to b e t k
ar a e n a t ri
g h t a n g les to

ea c h ot h er, a s is c o n v e n t io n a l in ana l y t ic g m et y eo r , and m ay b e re fer red

to as t h e x , y , a n d z a xes . It no w req u i res t wo let ters to m a rk a n a rr a y of

2 s
’
. We s h a ll c a ll th e ma r k o f a c la s s . N o w i m agin e th e m ea n s

p lo t t e d fo r e v er y z - a r ra y . If c o rre la t io n e x is ts , thes e m ea n s will n o t l ie a t

ra n d om in s p a c e, b u t wi l l a rra n ge t h em s elv es a pp ro x i m a tel y o n a s u r fa c e

c a l le d t he s u r fa c e o f reg res s io n .
”
T he eq u a t io n o f a s u r fa c e is o f t h e fo r m
We s h a ll c o n s id e r onl y t he c a s e wh e e th i f f c t io i f
r s un n s o

th e fi rs t d eg ree , fo r t he s a m e rea s o n s t a t we c h o id e d o ly th
ns c as e o f re n e

a fi rs t d eg ree fu n c t io n i n th e c a s e o f c o r rela t io n of two v a ria b les .

It res u lts t h at t he req u ired fu n c t io n is

l
‘
—l '

r 0
'

f f O
'

w h e re r
) ,
is the c o r re l a t io n c o effi c ien t b e t wee n the y a n d z -c h a ra c ters , a n d

s i m i la r m ea n i n g s a re to b e g iv en to t he o t er r s , as i n ic a ted b y t he s u b h ‘
d
s c r i ts
p .
q T h is e u a tio n g iv es t he m e a n v a l u e o f t h e z a rra s c o rres p o n d in
g to -
y
g iv en r a n d y , i f t e
. h y
c a n b e g iv en b y a li n ea r fu n c tio n I f t e
y c a nnot be . h
a c c u ra tel g i v en b
y a l iyn ea r fu n c t io n , t is eq u a t io n m u s t m erel b e loo ed h y k
iv in
g a fi rs t a pp ro x i m a tio n
u po n as g .

S ta n d a rd dev ia t ion of a rrays . I f th e e

q u a t io n
( I ) b e u s ed to es tim a te th e

v a lu e o f th e a c -
h a ra c ter c o r res p o n d i n
g t o a s e le c t e d . 1
'

and
y we, h a v e t he

sq u a re o f s ta n d a rd d ev ia t io n o f ea c h 3 -
a rra
y ab o u t t hi s es t i m a ted v a l u e

iv en b t he e x p re s s io n
g y
"
3
a r by )

a s a n a v er a g e v a lu e, w h e e the r s u m m a t io n ex ten d s to a ll th e o b s erv at io n s ;

and

r r (r r , ,
 A PP E N D I X 7 7
0

W h en ex p a n d ed a n d ex p r es se d i n terms o f r s a nd
’ ’
0 s, ( )
2 b ec o m es

)
2 rzx f rl
r y y
2
rx
y

T he fo rm u las i n the tex t i n d is c u s s in g b ipa ren t a l in eri t a n c e a re

u s ed h
s pec ia l c as es o f ( I ) a n d ( 3 ) j u s t d eriv ed T is m ay b e v er i fied b y m a i n g
. h k
th e fo llo win g s u b s ti tu t io n s

Pu t 1 [i v y fi g , 3 fi x, r2 , r ” r,, r
”
73 . a: a
s , 0,

T he n i n t he ne w n o t a t io n
( ) I b ec o m es

S i n c e i n t he c as e d is c u ss ed i n the tex t the pa ren ts were k

ta en eq u i

po t en t , r
l
r
, , a nd k
b y ma in g t is h s u b s t i tu t io n in ( )
4 we g et

7 108

wh ic h i s the fo rm u la u s ed in tex t L ik ewis e, i f we m a k e . h

t es e s u b s t i t u tio n s
in we get fo r t he v a ri ab ili t y o f a n a rr a y o f s o ns

wh ic h is the fo r m u la u sed i n the tex t .

M ore t han th ree v a ria b les I t is ea si ly . seen th a t the m et h o ds em plo yed

i n the c as e o f two a n d t h ree v ar i a b les c a n be ex ten d ed to a n y n u m b er o f

v aria b les . Ho wev er, the c o m plexi ty o f the a lgeb ra ic exp ress io n s b ec o mes s o
g rea t h
t at i t d oes n o t s eem well to p res en t a m o re ex ten d ed d is c u ss io n h ere .

Fo r the g en er a l c as e o f an y n u mb er o f v a ria b les , the r ea d er wi t c o n s id er h

h
a b le m a t em a tic a l tra i n i n g is referr ed to the t rea tm en t b y a rl Pear s o n in K
th e P lzilosop / uea l Tr a n s a c tion s of tire R oya l S ociety , A , C L X X X V I I , 1 8 96 ,
'

a nd A , CC, 1 90 3 . I n the pa pers just refer red to the gen era l ex press io n is
a ls o g i v en fo r the v a r ia b ili t
y o f an a rr a y i n the c as e o f a n y n u m b er o f

v a r ia b les . h
I t is fro m t is gen era l ex press io n t a t the fo rm u la u s ed i n th e h
tex t fo r th e v a r iab ili ty o f a n a rr a y o f o ffsp r i n g a fter 7: gen era ti o n s o f
s elec ti o n is d er iv ed .

Formula for th e correlat ion coe mcient r wh ich is b etter adapt ed to n u m er

ical ca lcu la ti on . I n the fi rs t p la c e, the c a lc u la t io n o f th e s tan d a rd d ev ia

t io n s o f b o t h y s s tem s o f v a ri a tes s o u ld b e d o n e b h y th e s ho r ter m et h od

p res en ted on p g
a e 46 5 .
7 08 A PPEN D Ix

T he v a lu e o b ta in ed fo r r on p g
a e 7 0 5 is
Er r
r

w he e r r and
.
y repres e n t d ev ia t io n s f ro m th e m ea n s , a n d t he s u mm a ti o n is
ex ten d ed to ev ery p a i r o f c o r r es po n d i n
g d ev ia t io n s . T he c a lc u la t io n of

Er r
no a
can be mu c h s h o rte n ed by a n eq u iv a len t fo rm u la w ic h h we s h a ll
, ,
no w d eri v e . Fo llo wi n g the n o ta tio n on p a
g es le t G ,
and 6 ,

r ep res e n t c lass m a rk s n ea r the m ea n s o f t he s y tem s s o f v a r i a tes in d ic a t ed

b y the s u b s c ri p ts , a n d C C, c o rrec t io n s to th e e c l
s as s ma r k s wh i c h g i v e
the c o r rec t mea n v a lu es so t h at

Le t y
’
r ep resen t d ev ia t io n s fro m G , an d G, w hic h c o rr es po n d

d ev ia t io n s .r,
y fro m t he m ea n . T hen
1 a
”
C} ,
C)
,
z
] ,

Cy )
ne w,
Ef y Q E/ EQ C;
’

Ef f C, 2 ( r
. C , ) C 2(y
, Q) . EC G ,

sa y

h
T is is a fo rm u la m u c h b et ter a d a p ted to c o m p u ta t io n h
t a n t he form u la
Er r

I ts a pp lic a t io n is s h o wn i n th e tex t, p age 4 6 5 .

S EC T I O N I X R A N D O M S A M PLE S

We k n o w fu ll well th a t we c an n o t, i n gen era l, m eas u re a ll th e in d i v id

n a ls o f a ra c e o r p p la t io n
o u w ho se c h a ra c ter is ti c s we wis to d es c ri b e hVt e
'
.

a re o b liged to ge t o u r i n fo rm a t io n a n d to c o n s tr u c t o u r s c ien c e b y th e
se le c t io n a n d ex a m i n a tio n o f s a m p les k
ta en a t ra n d o m fro m la rge gro u p
a

of in d iv id u a ls . T o i llu s t ra te, i t is no t p r a c t i c a b le to m eas u re the s ta tu re

n o r, h h
in gen eral , a n y o t er c a ra c ter o f a ll th e ad u lts in the U n ited S ta tes .

We h
m us t b e c o n ten t to d ea l wit a reas o n a b ly s m a ll n umb er tha t will m a e k
the meas u rem e n ts a feas ib le u n d er t a k in g .

A n in v es ti ga to r is a lwa y s c o n c er n ed a b o u t the n u m b er o f v a ri a tes whic h

m u s t b e m ea s u red i n o rd er t h at c o n fid e n c e may b e p la c ed in h is r es u l ts .
7 I 2 A PP E N D IX
A c o m m o n a nd i m p o r ta n t c lass o f s ta t is ti c a l d ed u c tio n s , w ic h h ho s u ld

rec eiv e v er y c r i ti c a l exa m i n a tio n , m a y b e illus tra ted as fo llo ws

S u ppose t h at o , u t of a to ta l o f ten yea rs w ic h h h av e b een o b serv ed , the
a pp le c ro p i th i
n s lo c a lit y has b een in j u red b y fro s t fo u r ea r s , a n d has y
b een j
u n i n u red s i x h y yea rs . I f t is d a ta fo r ten ea rs is a ll the ev id en c e we
h a e b ea ing o n the p rob ab ilit y o f a app le c o p the b e t es tima te we c an
v r n r , s

i e fo r the p o b b il it y t h t th e p p le rc 0 p i th
a i s lo c a li t y w i ll t b
a e a n
g v n o r

i j ed b y f o ts i a g i e y ea is l I f h o we e o r d a ta e te d o e
°

'
n ur r s n v n r r r , v r, u x n v r

t wen ty fi e y e -
i fi ftee o f wh ic h th e app le c 0p has b een
v a rs , n i j red b y
n r un n u

fro s t, we ag a i n g i ve
( 53
5 36 ) as the b es t es tim a te fo r the p rob a b il i ty
h
t a t the a p ple c ro p i n t is lo c a li ty h will not be in j u red b y fros ts in a g iv en
y ear ; a n d c er ta i n l y m o re c o n fid en c e c a n b e p la c ed in the res u lt h
t an wh en
o n ly ten y e we e ta ke a rs r n .

W e m ig h t c r y o r ill t a t io a r u us r n b ac k a hu n d red y ears in s ix t

y of wh i c h
th e a p ple c o p i th i lo c alit y h s
r n s a b een j
u n i n u red b y fro s ts and we s ho u ld

s ti ll g i v e as th e m o s t p ro b a b le v al u e o f the p ro b ab ili ty t a t a n a pp le h
c r0 p h
i n t is lo c a lit y wi ll n o t b e i n j u red b y fros t in a gi v en yea r I t s o u ld . h
be n o ted t at h we a re he er d ea li n g wit h the p r oba bi l i ty f o a p roba bil i ty , or

wh a t D e M o rga n h as c a lled the m f b b ili ty

”
p res u p t io n o a p r o a .

T he c r i tic a l ex a m i na tio n o f s uc h p ro b a b i li t ies as the a b ov e d eriv ed

fro m o b s erv a t io n s h o u ld i n c lu d e so m e c ri ter io n wh ic h will i n d ic a te the
a c c u ra c y o f the a pp r o x im a tion wh en onl y a lim i ted n u m b er o f c as es c an be
ex a m i n ed . Suc h a c ri terio n m ay b e fo u n d in the p rob a b le erro r o f the

p r o b a b i l i t y .

h
T he p rob lem in a n d m ay wel l b e s ta ted i n the fo llo win g gen era l fo r m
A b ag c o n ta i n s a n i n d efi n itel la rge n u mb er o f w ite a n d b lac b a lls i n y h k
un k no wn a tio i f l r I l n b a lls h ave b een d ra wn a s a r a n d o m sa m ple, a n d

III a re wh i te a n d a n re b la c k , we g iv e as the b es t v a lu e o f the p ro b a b i li t y

of d ra wi n g a h
w i te b al l — ” 1
W h at i s the p ro b a b le er ro r in t is h res ul t ?
7 1:5
.

11

O r , in ot he r wo rd s , i n m 1: t ria ls , a n ev e n t has h a ppe n ed III ti mes a n d

rn
fa i led to h a p pen n tim es i f we d ed u c e fro m t is t h h at
”I n
is the p rob a b i li ty

h
t a t t he ev en t will h a p pen o n a g i v en o c c as io n , wh a t is the prob a b le erro r

in t h is l
re u t
s ?

F ro m the wo r k s of La pla c e , Po iss o n , a nd D e M o rga n , i t fo llo ws th a t

fu n
the p ro b a b le er ro r I n Is g I v en b y the fo rm u la :l;
W it m n m
A pplied to ou r illu s t ra tio n o f th e a pp le c ro p wh en the d a ta c o v ered on l y
'

S
i
t 2
ten y ea rs , t hi s
p ro b a b le er ro r fo rm u la g iv es ; t
l
f—
7 2 l: 0 . I 0 4 .
:

F ro m the m a g n i t u d e o f t is p ro b a b le er ro r , i t is a t o n c e s een t h h a t th e
res u lt
( d er iv ed fro m ten o b s erv a tio ns
) fo r t h e p ro b a b i li t t a t an y h a p p le

c ro p will b e u n i n u red j b y fro s t c a n at m os t b e s a id to be b u t a c ru d e a n d

 INDEX

y h
A c c es s o r c ro m o so me, a s rela t e to d h
666 ; fas io n in , 65 8 6 59 ; b yb egin n ers ,
—
,

s ex, 63 4 6 3 7 ; G ro ss a n d W a llac e d
6 7 5 ; rec o r s in , 666- 6 7 2 ; s o w rin g
. h -

636 ; Hen i n g u po n , 63 5 ; k c o n s eq u en c es , 660 ; t es t in i d

— g
u po n , s res an

Mc C lu n g u po n , 63 5 ; Pa u lm ier u po n, ( 1 mm660 664 t es t in

g o u n g fem a es , y l
l
63 5 ; W i s o n 11 o n , 63 4 , 63 7 . 6 1 2 .

A cc lima t iza t io n , ea rin g o f, o n in s ta A n ima ls , gro wt o f, in u en c e b y ea t , h fl d h

l
b ili ty o f i v in g ma t ter, 3 08 3 1 6 ; b y -
2 58, 2 59 ; h h
ig er regen era t io n in ,
in oc u la t io n , 38 2 ; effec t o f, u po n 3 5 3
2 , 2 6 ; r e u c t io n i n , c o d
m p e
a r d
t ran s mis s io n , 3 7 4 3 8 6 ; e x t
-
e n t o f, h
wit plan ts , 1 6 5
—
.

3 5 3
7 , 7 6 ; i n ge n e ra l, 1 0 5 3 1 4 3 1 6 ; , y
A n t, p o l mo rp is m o f, 2 0 h .

d d
o f t he in i v i u a l a n d o f t he ra c e, 3 7 4 ; d
A n t en na ev e o p e as a fo o t, 4 3 l d .

o f rac es , 3 8 4 ; o f wo o s a ge o n d ig h h A p es , meris tic v a ria t io n in teet o f, 4 8 , h

l d
a n d lo w a t it u es , 3 7 8 ; p erm an enc e 49
o f, 1
5 h
t o c emic a s , 3 08 3 1 1 ; to l A p ric o t , m u ta n t o f t he p lu m , 1 1 2
d
-
; .

c o l , 3 1 3 ; to elec t ric i t , 3 1 4 ; t o y ig h h y
A rra s o f th e c o rr ela tio n ta b le, 4 5 9 .

tempera t u re, 3 1 1 3 1 3 t o h o t s p rin gs ,

- A rt em ia , exp e rim en ts wit , 2 8 3 h .

3 79 ; t o l h
ig t , 3 3
1 ; t o p o i s o n s , 3 8 1 A r tem ia s a li n a , effec t o n , b y eg ree o f d
3 8 2 t o t em p era t u res , 3 7 6, 3 8 1 t ra n s s a lin it , 1 02 y .

mis s io n o f. 3 7 4 : wit o u t s elec tio n , h h

A rt ifi c ia l pa rt en o gen es is , experim en ts
7
3 9 3 1 8 , in , 2 7 8- 2 8 2 .

d d
A c i s ec ret e b y a n im a ls an d p lan ts , A s s o rta t i v e ma t in g, 1 63 .

2 67 . A s ter, 1 4 7 .

A c q u ire c a rac t ers , 1 8 2 , 3 08 —

d h 3 1 1 a s y
A s m m et r , 70 y .

dis t in c t f ro m c o n gen ita , 3 54 3 56 ; in l -

A ta v is m , 1 92 — 1 94 .

heri ta n c e o f, 3 4 9 ; n o n exis t en c e o f, -
A t t ra c t io n o f o o rs , 2 7 5 d .

A u ric u la r a ppe n a ges in m a mm a ls , 4 5 , d

A c tI n Ia n , rea c t I o n to c h em ic a ls , 2 73, 46
2 74 ; regen era t io n in t he 3 3 4 , . A v era ge d
ev ia t io n , 4 2 7 ; illu s t ra te , d
A d a p ta t io n s , 350 ; n o t u n iv ersa l , 2 06 44 1 -
4 43 .

2 08 , 412 .

d
A lu m, Joh n, o rigin a to r of C a ta wb a B a c illu s tu ber c u los is exc it es a b n o rm a l
ra
g p e. '34 g r o wt , 98 h .

fl
A ge, in u en c e o f, u po n fu n c tio n a ac t iv l B ac teria, effec t o f c u lt u re m e iu m d
fl
ity, 94 in u en c e o f, u po n p repo ten c , y u po n , 229 .

Ba i ly exp erim en ts in a c c lim a tiza t io n ,

b
73 e
—
,

A l in ts m c o m m o n in m o s t s pec ies , 1 1 4 . 6
37 37 8 ; o n b u d v a riet ies , 1 8 1 ; o n

A m ito s is , 1 5 1 . t he go o s eb er r , I 30 y .

A moeb a , res po n s e to lig t , 2 5 3 , 2 5 4 h . B a rd eleb en , s t u ies in m a mm te , 4 7 d .

h
A mp ias t er, I 4 7 B a rren n es s t o c e rta in in iv i u a s , 2 0 1 d d l
B a t m ic in u en c es 2 0 2 —
. .

h y
A m p ic o n , 50 . h fl
2 08 , .

A n c es t ra l h d y
e re it , fo rm u a fo r, 5 3 3 , l B ea r, v a ria t io n in igits o f, 5 8 d .

5 34 la w o f , 5 5 5 34
2 -
. B ees effec t o f fo o u po n , 2 2 6
, d .

d
A n c es t ra l i io p as m , 1 7 3 l . B ego n ia , regen era tio n o f, 2 38 , 3 3 1 .

A n c es t ra l u n its 1 7 3 l d h
B e a t e in eri ta n c e, 4 7 5
—
, . .

d
A n ima l b ree in g, 65 4 6 7 6 ; a v a n t ages d B er t, ex perim en t s in g ra ft in g 1 0 7 , .

o f, 6 5 4 ; d d
is a v a n tages o f, 6 5 4 6 5 5 ; . j
B i om etr i l a , o u rn a o f s ta t is t ic a s t u
'

, l l dy
d d
is a v a n ta ge o f m a n c a ra c ters in , y h 4 7 8~
6 56 , 6 5 7 ; u rin
g ad ep r ess io n , 66 ,
5 d h
B io p o rs , I 4 , 2 08 .
7 16 I N D EX

l h
B ipa ren t a in erit an c e, 5 2 9 5 3 3 -
. wit h an d wit h out d iff eren tia t io n , 1 4
9
d
B ir s , effec t o f ea t u po n ev e o p m en t h d l 1 5 0 .

o f , 2 5 9 ; v a ria t io n in igits o f, 56 d . C en tgen er p lo ts , 64 4 .

h k
B irt m a r s , 1 8 9- 1 9 1 . C en t ro s o m e, 1 4 6 .

l
Bis ex u a rep ro u c t io n , a c a u s e o f v a ria d C erv ic a l fis t u la: in m a m ma s 4 4 l , .

t io n , 1 60 1 63 ; i n t ro u c es n o n ew
-
d h
C an c e, a c c o u n ts fo r u n u s u a o c c u r l
h t
c a ra c ers , 1 6 3 . ren c es , 1 8 7 ; as is t in c t f ro m c a u s e, d
B l kb
ac y err , ev o l u t io n o f, 1 3 1 - 1 3 3 . 6
3 5 ; la w o f , 6
3 5 3
, 66 .

Bl m i h e
e s s on b ree d ers , 590 . h
C a ra c ter, efi n e , 17; d m ean in g o f d
Bl di h
en n c 48 1
ara c ters , . t erm , 1 1 - 1 3 .
Bl d d d exc lu s iv e i n h erita n c e, 47 5 Ch ara c ters , ac q u i re d 1 82 a c q u ire df ro m
—
en e an .
, ;
B o n n et , experi men ts in r egen era t io n , t he en v iro n m en t, 3 0 2 3 4,
0 08
3 3 1 1 ;
-

3 I6 . a c q u ire h it d , in f 3 49 ;
er tanc e o , a re n o
B o n n ier, exp erim en ts o n ac c lim a t iza “
ac
q i d
u re 35 3 8 ,60
”
; b l d d 48 1 ; -
en e ,

t io n , 3 7 8 ; experim en ts wit an e h d d co mb i i d fi it p p ti
ne n e 0n
5 4 e ro or o ns,
l
io n , 2 2 3 .
5 3 1; g con i t l den q i ad 354 an ac u re ,

B o rn , exp erim en ts in graft in g, 1 08 , 3 3 6 .

35 6 ; d p d e
e en t p n94 96 ; u on s ex , 1 - 1
B ra in n o t n ec es s a r to c o or in a t e m o y d d d l pm t f h w i fl
ev e o en o d 36 , o n u enc e , 1
t io n , 4 00, 4 0 1 . 6
3 3 ; d m i o t d l
n an t t 3 1 4 ; an a en , 1 ,

d
B ree er s b u s in ess is th e p ro u c tio n o f
’
d d mi t d
o nan an i 5 4 5 5; rec es s v e, 1 , 1
s ires , 60 5 l m t y 4 ; h wb h i t
e e en a r 1 e av e
—
.
, o n ra n s
d
B ree ers fa s , 594
'
d . mi i ss o n, 4 7 3 4 78 ; l t t 474 ; t a en , no
d
B ree ers o f s pee a nd b ree ers o f d d n ec es s a r il d p ti 4 ; f d l t
,
a a v e, 12
d b y d l p m t 35 0 ; f
o a u as
d
b ree ers c o n t ra s te , 5 5 7 d . i fl n u en c e ev e o en , o
d
B ree in g p o m o rp is m in 4 7 6 , 4 7 7 ;
, ly h , t he i d i id n l e th
v ua f th ar ose o e rac e,
l
o b em s o f, o u t lin e , 3 d i i f i i d
5 p p
— 35 7
s es
p r ; u r o ; g or 4 n
3 4 5 o
; g 1 1 or n an
-
,

in , 5 99, 600 ; s s tem s o f, 5 99 6 2 7 ; y dg e en erac y f t ot d 4 5; t , c o n ras e , 1 oo

t r u e, o r s ta b ilit o f t p e, 5 4 1 - 5 44 y y . m y i an , im l b
n an d i g 6 56 6 5 7 ;
a ree n , ,

B ro wn S eq u a r , exp erim en ts o n m u t i a
-
d l i b ilit y f i
v ar a m p p l ti
o , n sa 444 e o u a on, .

t io n s , 3 6 7 . C h em i l ti
c a ac l im ti t i t 308
o n , ac c a za on o ,

d
B ru c e, s t u ies in m amm at , 4 7 .
3 1 1 ; f o t i
s ec re 8
3 3 3 4 8 o n s, , .

B u d v a ria t io n , 1 8 1 . C h m i l ff t
e ca e f l igh t 4 0
ec s o , 2 .

B u d v a riet ies rep ro u c e b y s ee s , 1 8 1 d d . C h m i l e ti

e ca r ac f p t pl m 2 64 o ns o ro o as ,

ll
B u , E \V , o rigi n a t o r o f C o n c o r d 28
é
. .

g ra pe, '3 4 C her i rc a ls , effec t o f, u po n g e rm i n a t io n ,

B u llfin c h, eff ec t o f fo o u po n 2 28 d , . 27 1 ; r hy th m i cal c o n t rac t io n s tim u

l
B u m b eb ee, an ten n a o f, ev elo p e a s a d d la t e d by , 2 76 - 2 78 .

foo t, 4 3 . C h em t i 7 7
o ax s , 2 1 , 2 2.
k y
B u rb a n s s tem o f p la n t in g, 6 4 3 . C h em t pi m 7 7 6
o ro s , 2 1, 2 .

B u rri ll
exp erimen t s in c ro ss in g s t ra w C h m ti g l 4 5
ro a n ra n u es , 1
h m ti m tt 1 4 5—5
, .

b e rr ies , 1 8 4 . C ro a n a er, 1 2.
C h m m
ro 1 46
o eres , .

l d
C a m e , ev elo p m en t o f fo o t o f, 60 . C h m m 1 46 5 ; b t h lf
ro oso es , - 1 2 u a t he
d
C a s t ra t io n , in irec t eflec ts o f, u po n t he
'

no r m l mb a f i p th
nu g er o , n ar en o en etic
bo dy fu n c t io n s , 1 00 . i d i id l
n v 1 80 ; m p it i
ua s , co os on o f,
ly
C a ta t ic p o is o n s , 2 6 6 . 1 74 ; c o n s tan t in n u m b er fo r t he
C a ts v a ria t io n in igits o f 57
, d , . s pec ies , 1 46 ; n u m b er o f, ev en in
l
C a t t le, a c c im a t iza t io n o f, 3 7
5 ev e o p d l b is ex u al rep ro u c t io n , 1 46 ; n u mb er d
m en t o f fo o t o f 5 8 ; m eris t i c v a ria t io n , o f, re u c e d d
b y m a tu ra t io n res to red ,

d
in igits o f, 6 2 6 3 ; rev ers io n s in 1 92 , , . b y fert iliza tio n 1 70 ; n u mb er o f, ,

C a v e a n im a ls , 2 4 2 . s o m et im es a ve 1 80 h l d , .

ll
C e , t he a s a s t ru c t u ra l u n it, 1 4 3 , 1 4 4 ; hy h
C r sa n t em u ms , ex p eri men ts wit , b y h
—
,

d ifferen t ia tio n in , 1 4 4 ; effec t o f gra v D e V ries , 1 1 9 1 2 1 .

i t u po n , 2 39 . l
C ea v age effec t o f o u ts i e c o n it io n s
, d d
C el d
iv is io n , as a c a u s e o f v a ria t io n , u po n , 3 40 ; geo m et ric a c a ra c t er o f, l h
8 i rregu la ri t ies in , 1 0—
1
55 1 1 5 1
5 ;
2

n
9
-
.

h
m ec a n is m o f, 1 4 5- 1 5 2 ; o u t in e o f l , ffic ien t , o f ass o rta t i v e m a t in g 5 3 2 ; ,

1 46, 1 4 7 ; v a ria t io n in ra te o f, 3 4 0 ; of c o rrelat io n , s o rt m et o fo r h h d

7 18 I N D EX

D ev ia t io n , a v era ge, 4 7 ;
2 n a t u re o f, En v i ro n m en t , a lwa s s elec tiv e, 3 5 1 y
l h l y l
—
50 n o t t ra ns m is s ib e a s s uc b ea rin g o f, o n in s ta b i it of iv i n g
8—
349 , 3 ; ,

349 ; s ta n d d ar , 42 43 1 ; s tan d d ar , m a t ter, 2 90 2 93 , 3 0 2 3 04 ; ca u s e o f -

m ea n in g o f 4 3 2 , , 433 . e v o u t io n l of t he o rs e,
3 0 2 -
0
3 4 h
D ev ia t io n a n d pro b a b e erro r i u s l ll d
irec t ac t io n o f, 30 7 ; fo od , 2 2 5 - 2 30 ,
t ra t ed 4 4 1 4 4 3 ,
-
.
g e n e ra effe c t l
o f, u po n d ev e o m e n t ,
p l
D e V ries , ex peri m en ts o f, 1 1 4 1 2 9 ; ex 2 2 1 - 2 2 5 ; h o w in u en c es t
p e o f ra c e fl y
—
-
,

p eri e nc e in p an t b ree in g 64 2 ; ex l d , 2 90 2 9 3 ; in flu en c e o f, u po n c l ea v a g e,

p eri m e n t s wi t h C h r s a n t em u m , 1 1 9 y h 3 4 0 ; in u en c e o f, flu po n
pa rt h e n o

12 1 ex perim en ts wi th p rim ro s e, g e n e s is , 1 78 i n u en c e o f, u po n v a ri a fl
1 2 1 1 2 9 ; p ro u c t io n o f n ew s p ec ies
-
d b ili ty, 2 2 0 2 93 -

b m u t a t io n , 1 2 1 1 2 9 - . y
E p i leps , t ra n s m i s s ion o f, 36 7 .

D i eren t ia tio n , b y c e iv is io n , 1 4 9, ll d d h
E v i en c e t a t is n o t ev i en c e, 3 5 3 d .

1 50 ; ca u s es o f, 3 4 3 ; f ro m in t ern a l l
E v o u t io n , n o w e ge o f, k n ee e ld
in d d
ca u s es , 1 4 4 m ec an is m o f, 1 4 2 1 5 4 ; h -
d
b ree in g, 5 ; n o t c o n fi n e t o m o rp o d h l
p o a ri t l y
an d p ro m o rp o o gy o f o v u m , h l o
gy , 7 6 ; W e is m an n
’
s t h eo r o f , 1 5 2 y .

34 1- 343 ~ E wa rt , ex p eri m en ts in t e ego n , 1 8 6 l y .

l d
D igi t a is , m e i c in a q u a i i es o f, affec te l lt d E x erc is e, effec t o f, u p o n f u n c ti o n a l
l
b y o c a it , 2 2 3 l y . a c t iv it y
95 9, 6 , .

D ig its , m eris t ic v a ria t io n in , 3 6 4 -

. hh l
E xo p t a m ia , t ra n s mis s io n o f, 3 67 .

h
D im o rp is m o f ea rwig, S o r t ho m s , l fl
E x tern a in u en c es as c a u s es o f v a ri a b il
Herefo rd s . e tc 2 0 .
, . it y, 2 2 0— 2 93 .

D is a ppea ra n c e o f pa r ts , 3 06 . E x t ra win g, 4 3 .

D isa pp ea rin g pa rts , 2 8 8 . E ye, effec t o f ig t u po n 2 4 3 l h , .

D isc o n t in u it in v a ria t io n , 1 9 y . y d
E es , egen era t io n o f, in c a v e S p ec ies ,
D is eas e, t ra n s m is s io n o f, 368 , 38 4 .
4 1 1 , 4 12 ; s u pe r nu mera ry, 5 1.

D is eas es d u e t o ab s en c e o f s ec ret io n s ,
2 69 . d
F a s o f the b ree er, 5 94 d .

D is t rib u t io n , o ffs p rin g c o n s t it u t e a, 4 1 9 . F a n to p t rees , 1 1 2

-
.

Do dd
, ll
W i ia m , o rigi n a to r o f p u m , l h
Fas io n in a n im a b ree in g, 6 58 , 6 5 9 l d .

1 33 . Fa t ten in g, u n s u c c ess f u l in a r n es s , d k
l
D o g, a c c im a tiza t io n o f, 3 7 6 ; b e a v io r h 2 46 .

o f , w en ep ri v eh d
o f b ra in 4 00, 4 0 1 d , F ea r, n o t p res en t a t b irt , 403 h .

m eris t ic v a ria t io n in teet o f,

5 ;
0 h l fl
Fem a e, in u en c e u po n , b y p rev io u s
v a ria t io n i n ig its o f , 5 7 d . m a t in g, 1 8 5 - 1 8 9 ; ma t u ra tio n a n d re
fl
D o gs , in u en c e o f oc a it u p o n , 2 2 4 l l y d uc t io n i n , 1 6 5
-
1 6 9 .

t e ego n l in , 1 86 y l d
F em a es , is p o s al o f s u r p u s , 6 7 2 l .

D o rfm eis t e r, ex peri m en ts wi t b u t te r h '

Fe ré , ex peri m en ts o f , o n c ic s , 2 5 9 h k .

fl
ies 2 6 2 , . l y
Fe rt i it , c a rac t ers c o rrela t e h wit , d h
l l y l
re a ti v e, 1 96- 2 00 ; effec t o f f oo d
—
D o u b e pers o n a it , 1 06 . 1 97
l
D o u b in g o f pa rt s , as ea , 6 7 69 h d . u po n , 2 2 6 ; effec t o f, u p o n t pe, 1 98 , y
k l
D u c s , re a t i v e weig t o f b o n es o f t a m e h 1 99 ; effec t o f, u p o n t p e an d v a ria y
a n d wi , 95 ld . b ility, 4 49- 4 5 1 ; impo rt an c e o f, 1 99,
D u go n g, v a ri a t io n in igi ts o f, 5 7 d .
5 8 4 , 5 8 9 ; l es s w i t ex t rem e s o f a h
D wa rfs , reas o n s fo r, 2 7 . ra c e t a n wi t h h
t he m eans , 4 9 1 ; o ft en
yd
D a s , 1 66 . o
pp o s e b y s e e c tdion, 8
5 3 l .

l
F ert i iza t io n , b y th e po a r b o , 1 79, l dy
E th w
ar o rm, m eris tic v a riatio n in gen era 1 80 ; c o n n ec t io n o f, wit m u ta t io n , h
t i v e o pe n in g o f, 4 4 d
1 80 ; o f en o s p erm , 1 8 3 , 1 8 4 ; ma n n er
—
.

Ea rth wo n n s , regen era tio n in , 3 1 7 3 2 0 . o f, 1 6 1 s ign ifi c a n c e o f, 1 7 0 ; in u enc e fl

d
E a rwi g, i m o rp is m o f, 2 0 h . o f, u po n s ex , 6 3 2 , 6 3 3 .

Eggs , regen era t io n in , 3 2 4 , 3 2 5 . h

F is , u p s t rea m m o v em en ts , 2 3 5 .

hl h
E r ic , ex p eri m en ts wit m ic e, 309 h h
F is er s t u ies in fi s t u lae , 4 5 d .

E i m er, o n a a p t a t io n , 2 06 —
. ,

d
2 0 8 ; o n th e l ll
F a ge a ta, ac c im a t iza t io n to ig t em l h h
or
y o f o rt o gen es is 2 04 2 08 h ,
-
.
p t era u res ,
3 9
7 3 8 1 , .

l
E ec t ri c it y
a c c im a t i za t io n t o ,
,
3 4
1 l . Fl mm
a a rio n , experim en ts wit h ligh t ,

E m b ryo s , regen era t io n in , 3 2 4 , 3 2 5 .

d
E n o s p erm , fert ili za t io n o f, 1 8 3 , 1 84 . F la t fis hes , 4 1 5 .
 I N D EX 19

F oo d , effec t o f c o tto n s ee o n po r , 2 2 8 ; d k G eo tax is , 2 3 6

—
.

eff ec t o f , u p o n b u ll fi n c hes , 2 2 8 eff ec t G eo t ro p is m , 2 3 6 2 3 9 ; in a n im a s , 2 38 l

o f, u po n bo dy
t em p era t u re, 2 30 ; o f s p ro u t a n d ro o t , 1 1 1 .

effec t o f, u p o n c o n s t i t u t io n a v igo r, l G erm , in fec t io n o f , 1 8 5 ; in u en c e o f fl

3 7 2 , 3 3
7 ; e ff ec t o f , p
u on e ve o
p d l l
a ge o r s ta en es s u p o n , 1 8 2 ; in i v i u d d
m en t, 3 7 0- 3 7 4 ; effec t o f , u po n fer a it l y
o f, 1 8 2 ; v aria t io ns in , a re t ra n s

t ility, 2 2 6 ; effec t o f, u p o n f u n c t io n a l m itt ed , 3 4 8 .

y
a c t iv it , 96, 97 ; en erg o f, 2 2 9 ; ex y l
G erm p a s m an d t ra n s m iss io n , 3 5 5 .

c ess o f , 2 2 7 ; h o w re u c e b y iv in g d d l l
G erm in al s e ec t io n , 1 62 , 2 1 3 - 2 1 5 .

b ein gs , 2 2 8 ; in u en c e o f, u p o n regen fl G erm in a v es ic e, 1 66 l l .

c ra tio n , 3 2 7 , 3 2 8 ; in u en c e o f, u p o n fl G er m in a t io n , eff ec t o f c em ic a s u po n , h l
l y
v a ri a b i it , 2 2 - 2 3 0 ;
5 p r o po rt io n o f, 271 .

u se d
in gro wt , 2 2 5 q u a i t a ht iv e l G ian ts , reas o n fo r, 2 7 .

effec ts o f, 2 2 8- 2 30 ; q u a n t it a t iv e l d
G a n s , s p ec ifi c s ec ret io n s o f, 2 69 .

effec t s o f, 2 2 5 2 2 7 ; we l b re -
rac es l -
d l
G o t z, exp eri m en ts o n th e d o g, 4 00,
req u ire m o re, 2 27 .
40 1 .

d
F o u n a t io n n o t in a rem o t e fema e, l y
G o o seb err , ev o u tio n o f, 1 30 l .

5 95 5 . 9 6 . ll
G o ri a , ex t ra in c is o r in , 4 9 .

Fras er o n effi c ien cy o f c o ws , 7 8- 8 1 d

G ra in g 60 2 - 608 ; a v an t a ges o f, 604 d
—
. ,

l
F ra terna v aria b ili t , 5 00 504 y . ab u s e o f 604 ; b egi n b y t o get ex pe
, ,

Fr ee m a rt in s , 1 7 6
-
. ric u c e, 606 ; is a v a n tage o f, 608d d
F req u en c yd
is t r ib u t io n , t p ic a , 4 2 1 y l . dis a ppea ra n c e o f u n im p ro v e b o o d l d
Freq u en c yd
is t rib u t io n a n d t h e b in o m ia l in , 60 2 .

h
t eo rem , 509 . G ra f t in g fro g m a e u p o f p iec es o f
, d
F ro g, in flu en c e o f ea t u po n g ro wt h o f, h t wo d d l
in iv i u a s , 1 08 ; i u s t ra t in g ll
2 s8 . s tab i it l y l of iv in g m a t ter, 3 3 5 , 33 6 ;
F u n c t io n a l ac t iv it , b y s u gg es t io n , 2 4 3 y m a m ma r g a n y l d in to ea r o f gu in ea
effec t s o f l h
ig t u p o n , 2 4 1 2 4 6 ; h o w -
i
p g , 1 0 7 ; s p u r o f c o c in t o c o mb , k
d d l h
ep en en t u p o n ig t , 2 5 1 , 2 5 4 . 1 07 two s p ec ies o f a n im a t o get er, l h
l
F u n c t io n a v aria tio n 7 5 1 09 ; b e t ween ,
-
33 6 ; t w o p iec es o f t a po e, 33 6 d l .

d d d l
ifferen t in i v i u a s o f t he s a m e s p e G ra p es , ev o u t io n o f, 1 3 4 l
— —
.

c ies , 7 7 9 1 l h
d u e t o ig t , 2 3 9 2 5 4 ; o f y
G ra v it , effec t o f u p o n ev e o p men t , , d l
d d l
s a m e in iv i u a fro m d a
y t o da , 91
y 2 39 ; effec t o f, u po n iv in g m a t ter, l
9 4 ; id d
n u c e b y c as t ra t io n 1 00 ; i n 2 36 — 2 3 9 ; eff ec t o f, u po n p ro to p as m , l
d u c ed b y c o n ta c t , 2 3 3—
,

d d
2 3 6 ; in u c e 2 3 9 ; effec t o f, regen era t io n ,
—
upo n
l fl
b y ex t ern a in u en c es , 98 , 1 0 1 1 0 5 ; -
2
3 9 33 2 ; in s t ru
gg e w i t p o la ri t , l h y
fl d d
in u en c e b y fo o , 96 , 9 7 , 2 2 5 2 3 0 ;
—
2 7
z
-
.

fl
in u en c e b d g ra y
v i t , 2 3 6 2 3 9 ; in G re i s res 5 5 2 ; t he ten grea tes t , 5 5 5
t i , .

d
flu en c ed b y a rd c o n itio n s , 97 ; in G ro s s o n t h e ac c es s o r c ro m o s o m e, y h
d
flu en c ed b y t he rep ro u c t iv e fa c u ties , l 6 6
i
.

1 00 G ron , A P , 71 d e A n gu s s t eers b e
£
. . .

F un c ti di h g d ly i p
o ns , sc ar e on n res en c e lo n in
g g to 5 , .

of l igh t 4 43 ;
, 2 i d
2, d 2 ex e rc s e un er G ro wt h
as in u en c e
, fl
b y t em p era t u re, d
ab m l
no r d it i
a co n7 9 o ns , 10 -
10 . 2 54— 262 ; d
irec tio n o f, d u e t o ig t , l h
F un k D , N ea n e l d l t fpi .
, c a r- o a o o r ze 2 49 d
irec t io n o f, i n u en c e b y ea t , fl d h
i g g d
r n rat tl b l e cai
g g t 6 7 e e on n o, 0 . 2 5 9 ; geo m et ric a c a ra c t e r o f c lea v l h
F ii b i g
r r n t di i bi d 4
er , s u es n r s, 2. age i n , 3 3 9 ; reta r e b y ig t, 2 4 5 , dd l h
2 46 .

G ll 98
a s, 7 , 2 0 . h
G ro wt fo rc e, 2 03 , 2 04 .

G lt
a on, h d it y 4 7 8 ;
on ere l w f , on a o G u in ea p ig, g raf t in g m a m m a r g a n y l d
t
a n c es ra l h d it y 5 8 ere f t l , 2 ; on ra ern a in to ea r o f, 1 0 7 ; s u pp o s e t ra n s d
i b il i t y 5
v ar a
5 , l w f
00, i h i t 01 on a o n er m is s io n o f m u t i a t io n s o f 3 6 7 l , .

a ri c e, 1 93 , 1 94 ; o n s t u dies o f s ta t u re,

4 s1 . H ab it n o t t he b as is of in s t in c t, 40 1
G am et ic p u ri t , 5 —21 y .
4 03 .

G emm u es , 2 08 l Hab it s, a re t he t ra n s m it t e d? 6
3 3,
— — y
.

G en et ic s e ec t io n 1 96 2 00l 3 86 4 03 ; f o u n c ed o n i ti t
ns nc s , no t

l d
, .

y
G en t r , N H o n in b ree in g, 6 2 5
. .
, d . the rev ers e, 40 1 4 03 ;
-
ea rn e f ro m
7 2 0 I N DEX

e ld ers , 3 53 ; n o t a c q u ir e d h c a ra c t e rs , Ho rs es , ac c l im
375 ; a t iza t io n o f, c o rre~

35 3 6
80 - l
a t io n b e t ween c o o r , s ex, a n d s ee ,
p l d
l l a rn s , B F .
g .
, 89
, exa m p e o f l l on ev i t y .
4 68 4 7 1 ; in b ree in g in 6 2 4 ; p o we r
-
d ,

H ea pe experim en ts o n rab b its , 1 90

, . o f t ra n s m iss io n a m o n
g 4 08 ; t e e on
g , l y
hy h
Hea rt , r t m ic c o n tra c t io n o f, 3 98 , 3 99 . in , 1 8 5 .

Hea t , ac c lim a t iza tio n t o , 3 1 1 3 1 3 , 3 7 9 -

h
Ho rs es o e i n e , 6 5 kd y .

3 8 1 e ff ec t o f, u p o n an im a ac t iv i l Ho t s p ri n gs , I n f u s o ria in , 3 1 1 ; if e in , l
t ies , 2 5 5 effec t o f, u p o n i rec t io n o f d 379
g roh wt , 2
59 ; effec t o f , u p o n g ro w t h Ho u g ht on, Abe l , o rigin a to r o f
go o s e
l
o f a n i m a s , 2 8 , 2 9 ; eff ec t o f, u po n
5 5 b err y , 1 30 .

g wt h f
ro p l o a n ts , 2 55
-
2 57 . Hu n ter experi m en ts in g ra f t in g, 1 0 7
, .

H dg h g m
e e o , eris tic v a ria t io n in v ert e H u n t in gt o n R an o p , o n b ree in g ,
, d l h d
b ra:o f, 39 . 62 4 .

l
He io t ro p is m , 2 4 7 ; c o n it io n s t a t d e d h y
H b ri ds , c h a ra c t er of d es c en d a n t s o f,

t erm in e 2 5 4 ; d u e t o t he u m in o u s
, l 5 1 4- 5 2 1 ; Me n d l e s
'
la w o f,
5 3 5 5 1 -
2 ;
y
ra s , 2 4 8 ; gen era p rin c ip es go v em l l s t er i l it y o f, 60 7 .

in g, 2 5 1 , 2 5 4 ; o f a m oeb a , 2 5 3 , 2 5 4 y
H p ert ro p hy , 2 8 8, 2 89 .

o f in s ec t s , 1 04 .

k
Hen in g o n t he ac c es s o ry c ro m o s o m e, h Id t an s, 1 73 .

63 5 . Id l ea s l
in s e ec t io n , 5 7 8 , 5 7 9 .

Herb s t, exp eri m en t s in regen era t io n , I di pl o as m , 1 5 2 , 2 08 .

3 2 8 . I ds , 1 46, 2 08 .

d
Her , man agem en t o f, u rin g d ep res d ll
I in o is s ta t io n , s s tem o f p an tin g a t, 6 4 6 y l .

s io n in p ric es , 66
5 666 rec o r s o f, d y
I m m u n it , n a t u ra a n d a c q u i red , 38 2 l
—
;
-
.

666 67 0 ; u n i t o f, 66 2 ; wit o u t a y h I m p ro v em en t u pp er im it s o f , 58 2 , l .

hea , 664 d . d
I n b ree in g, 6 1 3- 6 2 6 ; a v an t ag es o f , d
d y
Here it , 4 7 3 5 4 7 ; c o efi c ien t o f, 48 6 -
6 1 4 , 6 1 5 ; A J Lo v e o on, 62
5’ ; j y
—
.
, .

‘
4 88 ; c o e flic ien ts o f iffe ren t re a d l a mo n g a n im a s , 6 2 4 6 2 6 ; D a rwin s l
t io n s h ips 4 88 ; fa m o
, u s gra n s ires d , exp erim en ts 0 11 , 6 1 8 6 2 4 ; d is ad v an —
55 6 ; la w o f a n c es t ra , 5 5 53
2 4 m an l -
t ages o f, 6 1 5 ; fo rm s o f, 6 1 3 , 6 1 4 ;
n er o f, 4 2 0 4 3 1 ; m a te ria b as is o f,
-
l ho w to p ra c tic e, 6 2 6 ; o ft en m o re
h
2 09 ; m a t em a t ic a l n a t u re o f , 5 1 0 ; h
v igo ro u s t an o u tb r ee in g, 6 2 2 -6 2 4 d
m ea n o f o ffs p rin g n o t m ea n o f t he l k
ac o f v igo r a n d lo w fert i it com l y
p a re n t s , 4 9 0 ; m e a s u r e o f 4 8 6 ; m is , d
m o n efec t s , 6 1 6, 6 1 7 ; N H G en t r . . y
c o n c e t io n s o f , 4 7 3 ; o ffs p ri n g iff d o n , 62 n o t a ll in b re i n iv i u a ls d d d
5
—
p e r ;
f ro m p a ren ts , 4 8 2 , 4 8 3 ; o ffs p rin g in ferio r, 6 2 0 6 2 3 ; n o t n ec ess a ri d is ly
m o re m e io c re t a n t h e pa ren ts , d h as t ro u s , 6 1 6 6 2 6 ; R an o ph H u n t -
d l
—
4 8 4 4 86 ; o rigin o f t h e exc ep t io n a l in gto n o n , 6 2 4 ; s p ec ia a n gers in , l d
d d l
i n i v i u a , 4 99 5 00 ; p ro gress io n i n , , l
6 1 6 ; t o ta effec t s o f, 6 1 9, 6 2 0 .

4 9 2 4 98 ; p ro per c o n c ep t io n s o f 4 7 3 ;
-
, d d l
I n iv i u a , t h e, 3 5 2 ; p o ss es s es a ll t h e
l
s ta t is t ic a m et o s i n s t u h d
o f, 4 2 6 , dy h
c a ra c ters o f t he ra c e,
35 3
7 , 6 0 .

47 8 ; hw a t is t ra n s m it t e ? 5 1 1 d . d d l y
I n iv i u a i t in o ffsp ri n g fro m sa m e
d d
Her efo r s im o rp is m in 2 0 , h , .
p t 5 3
a ren s , 0 .

Hero t h e i n b re m o rn in g g o r , 6 2 2
, d -
l y . In f t i l it y
er mm d a co on efec t , 6 1 6, 6 1 7 ;
Her rin gha m s t u ies o n n erv es , 4 3 , d . eff t f 5 8 4 5 8 9
ec o , , .

H o m e o s is 3 7 ; in in s ec ts , 4 3 ; in v ert e In h it er mp l
a n c e, co ex , 5 7;
2 be l a t ed ,
b rae a n d ri b s , 4 0—
,

42 .
47 5 ; b l d d d en e an exc u s iv e, l 475 ;
y
H o n e b ee, e es u n it e 6 5 y d , . f ro m s ep a ra te a n c es t o rs , 5 2 7 ; fro m
k
H o p in s , exp eri m en ts in c o rn b ree in g, d t he ra c e, 1 93 1 94 ; n o t lim ite t o s ex , , d
4 93 498
-
'
4 7 4 ; o f a c q u ire c a rac t e rs , 2 9 2 , 3 4 9 d h
Ho rn s , m eris t ic v a ria t io n in , 2 , 5 3, 66 ; o ffs rin
p g m o re m e ioc re t a n t h e d h
r egen era t io n o f, 3 2 6 .
p a ren t s 8
4 4 4 ,8 6 ; p a rt ic u la t-
e, 4 6
7 ;
Ho rs e, b eggin g in s tin c t in , 3 5 3 ; c a u s es p g
r o re ss io n i n 4 9 498
2 -
, .

l
o f ev o u t io n o f , 3 0 2
3 0 4 ; efec t i v e -
d l
I n o c u a t io n , im m u n it b y, 3 8 2 y .

v o ic e in 3 53 ; e v e o m en t o f fo o t
, p d l I n s ec t p o is o n s , 2 7 0 .

Of 5
, 8 59 ; , e v o u t io n o f 2 8
9 3 5;0 l ,
-
I n s t a b i it l y
of , l
i v in g ma t t er, i l u s tra t e l d
ex t rem e a ge o f 8 9 ; m e ris t ic v a ri a, b y o rigi n o f ifferen t ia te t is s u e, 33 6 d d
t io n i n d
igi ts o f, 60 6 1 , .
33 8 of
p ro t o p as m 39 8 l , .
7 2 2 I N D EX

M dl
en e
’
s la w 5 1 3- 5 5;
2 exp erim en ta l M u m fo rd , experim en ts
in fee in g , 8 d
1 6— 2 1
,

id
ev en c e on, 5 5 ; ex pe rim en t s o n po r k
p o u c t io n , 2 2 8
r d .

wit h m i c e,
5 4
2 l
M u sc e fib er, eff ec t o f ig t u p o n , 2 4 7 l h
M u t ab i it o f s p ec ies , 2 98—
.
.

M d l l
en e s
’
a w a n d ga m et ic p u rit y , 5 2 1 . l y 30 5 .

M eri n os in N ew Z ea a n , 2 2 3 l d . M ll ta n ts , 2 1 ; o rig in o f, in to ad fla x ,
Meris m 3 3 , . 1 1 5 1 18 -
.

M eri s t ic v a ria t io n 3 3 7 4 -
; c erv ic a l fis l
M u ta t io n , b io o gica s i ifican c e o f, 1 3 6 , l
h d £8
,

t u la an d a u ri c u a r a p pen l d a ges , 44 1 7 ;
3 i s d
t i n g ui s e 111 o r i n ar va d y
46 ; d o u b lin g o f c o m p lic a t e p ar ts , d r ia t io n , 1 10 ; ec o n o m ic s i n ifica n c e
g
—
64 6 9 ; d u e to c e l div is io n , 7 2 , 1 5 8 ; l o f, 1
35 3 1 -
6 ; i n C r sa n t em u m , 1 1
9 hy h
ho m a o s is in , 3 7 ; in i gits , 5 3— 64 d 1 2 1 ; i n p rim ro se 1 2 1 - 1 2 9 ; la ws o f, ,

in ey e s , 5 1 ; in gen erat iv e pa rts , 4 4 1 2 7 1 29 ; -

in t o a d fla x , 1 1 5 1 1 8 ; i n -

—
,

h d
in ea , 6 7 , 68 ; i n ho rns , 5 2 , 5 3 ; i n A m eric a n n a tiv e p lan ts , 1 2 9 1 3 5 ; i n
l
egs , 64 in m a m m a , 46, 4 7 ; in ra ia l d g e n e ra l, 1 1 0 — 1 3 8 ; r ela t io n o f, t o t e
—
s eri es , 7 0 7 3 ; in s p in a n erv es , 4
2
l °
d u c t io n a n d fert i iza t io n , 1 80 l .

h
in t eet , 4 8- 5 1 in v ert eb ra an d ri b s , M u ta t io n a n d e emen ta r s pec ies, 1 2 8 l y .

39 4 2-
; in “i n gs
5 1 M u ta t io n an d pa rt en o gen es is 1 7 9 h , .

l
M ic e, ac c im at iza t io n o f , t o ric in , 309 ; l
M u t i a t io n s a re t e t ra n s m it t e ? 36 3
, h y d
v a ria t io n in igi ts of , 58 d .
3 6 8 ; e x eri m en ts u o n t ra n s m iss io n
p p
Mic ros o m es , 1 46 . o f, 3 6 7 ; res emb la n c e o f, to n a t ura l

M id -pa ren t, C a to n s , 48 1
’
l . d efo rm i t , 3 66 y .

-
d
M id p a ren t ev ia t io n fo rm u la fo r, 5 3 1 , .

M id pa ren t v a ria b i it , fo rm u a fo r , 53 1
-
l y l . l
N age i, ex perim en ts wit c o pp er, 2 68 h .

k d
Mil p ro u c t io n , v a ria t io n in , 7 8 - 8 0, h
N a rw a l, meris t ic v a ria t io n in t u s k o f,
2 , 93 t ra n s m it t e b y m a es , 360 d l .
7 0 .

lk
M i s ec re tio n b y m a es , 1 0 5 , 1 0 7 l N a t u ra l s elec t io n a wa s a t wo rk , 5 88 l
h
.
.

M in n es o ta s ta tio n , s s t em o f p la n t in g y N ec ta rin e, m u tan t o f t e pe a c , 1 1 2 h .

a t 64 4
, . N eo D a rwin ian s, 3 5 4
-
.

M itos is , as a c a u s e o f v aria tio n , 1 5 5 N ee -La ma rc ia n s, 3 4

5 k .

181 ; d l
et a i s o f, 1 4 5 1 5 2 ; irr eg u a r -
l erv es , m eris t ic v a ri a tio n in , 4 2 .

i t ies in , 1 50- 1 5 2 ; p a t o o g ic a l, 1 5 0 h l N eu geb a u er, s t u ies in ma m m a , 4 7 d .

1 s1 . l
N u c eu s , 1 4 5 1 5 2 -
.

M ixed b ree in g p u ri t in 507 d , y , .

dl
Mo a c o effi c ien t 4 2 2 , 4 2 3 , . d
O o rs , a t t ra c t io n o f, 2 7 5 .

d
M o e the 4 2 1 , 4 2 2 ; em p i ric a l
, , an d O il c o n t en t o f c o rn , 8 3- 8 5 ; e ff ec t of
th eo ret ic a l , 422 ; p rac t ic a l v a ue l o f, l
s e ec t io n u o n , 4 4 , 4 4 6 ;
p 5 p g
ro r es s io n
d
in b ree in g fo r, 496- 498
f
4 2 .

Mo d i i ca t io n s d u e to ex t e m a l in fl u en c es , y
O ld G ra n n , exa m ple o f lo n gev ity a n d
t ra n s m iss io n o f, 3 4 8 —
4 17 . ex t rem e fert ilit , 89, 90 y .

M o is t u re eff ec t o f, u p o n d ev e o l pm en t , y
O oc t e, 1 6 5
—
.
,

2 30 2 3 3 ; effec t o f, u
po n s p in y Oogo n ia, 1 6 5 .

gr o wt h , 231 . O rien ta t io n , 2 4 7 .

l
M o n t e C a r o a n d ro u et t e 3 6 5 l , . h
O ri gin o f c a rac t ers , 4 1 3 - 4 1 5 .

M o rga n o b s erv a tio n s o n ac c lim a t iza

, h
O rt o gen es is , 2 04 - 2 08 ; exp a in ed b y l
t io n to h ea t , 3 1 1 on fea r in c hi k c s, g er m i n a s e l ec t io nl, 2 1
5 .

4 03 . Os b o rn , res ea rc es o n ev o lu tio n o f t h e h
M o rga n h o rs e 2 96 , . h o rs e,
30 2 .

M o rn in g g o ry , D a rwin s
-
’
l exp eri m en t s O ss ifi c a t io n , 99 .

in in b ree in g 6 2 1 - 6 2 4 d , . y
O t o c o n , teet o f, 0 h .

h l
M o rp o o gic a l v a ri a tio n , 2 5 2 9 ; c a u s es -
d
O v erfee in g, ev i e ec ts o f, 2 2 7 l .

o f, 2 7 ; in m u b err ea v es , 2 6 l yl . O v u m , 1 6 1 , 1 6 5 ; po la ri t o f, 3 4 1 3 4 3 ; y -

M o ss ro s es a b u d v a riet , 1 8 1 , y .
p ro m o r p o o g o f , 3 h l y
4 1 s egm en t a
h fl h d
Mo t s, ig t o f, et erm in e b y lig t , d h h
t io n o f, wit o u t fert il iza t io n , 1 7 7
1 80
é m t i d d by
2 0 . .

Mo e en n uce c o n t ac t , 2
34 . y
O x gen , effec t o f, u po n p ro to p las m , 2 6 5 .

Mu lb y l
er r p l y m ea v es , o o rp hi sm in , 26 .

Mu l t ip l y i g p l t 6 5
n o s, 0 . Pa n mix ia 2 8 8 , .

Mu l t ip l m it i 5
o ar os s . 1 1 . Pa r ry, o rigin a t o r of k
b lac b e rry, 1 32 .
 I ND EX 7 2 3

Pa rt h en o gen es is , 1 62 ; b ut one p o ar l Po l a ri t y
in s t r u gg e wit g ra v it , 2 3 7 ;
, l h y
bo dy in , 1 7 9 ; in u enc e by con fl d di o f o v u m,
3 4 1 343
-
.

t io n s l
o f if e, 1 7 8 ; i n u en c e b y t em fl d Po llin a t io n , i n i rec t eff ec t o f, 1 8 5 d .

p e ra t u re, 2 8 1 L
’
o eb s ex eri m en t s i n ,
p ly h
Po m o rp is m i n p ra c t ic a l b ree in g, d
2 78 2 82 ; b u t
-
a lf t he n o rm a n um h l 47 6 , 47 7 ; w i t rega r t o s ex , 2 0h d .

h
b er o f c ro m o s o m es in , 1 80 . l l
Po p a r, ac c im a t iza tio n o f, 3 7 6 .

h
Pa rt en o gen es is a n d m u ta tio n , 1 7 9 . l
Po p u a t io n , 4 2 1 t p e o f, 4 2 2 y .

h
Pa rt en o gen etic rep ro u c t io n , v a ria t io n d k
Po r , eff ec t o f c o t to n s ee u po n , 2 2 8 d .

in , 1 7 7 - 1 80 . Prec ip it in tes t , 3 8 2 .

l
Pa rt ic u a t e in erita n c e, 4 7 6 h Preferen t ia m a t in g, 1 63 l
—
. .

Pa u lmi er o n t he a c c es s o r c ro m o s o m e, y h fl
Pren a ta l in u en c es , 1 8 9 1 9 1 .

63 5 . Prepo t en c 55 1 , y5 5
7 ; ,a s a ff ec t e b y d
Pea rs o n , o n t h e b a t m ic in u en c es , h fl age, 5 7 3 ; as in ic a t e b y p erfo rm : d d
2 03 o n c a u s es o f v a ria b i it , 2 2 0 ; o n l y an c e, 5 5 8 5 6 7 ; as re a t e -
to c o n s t i l d
l y
fert i it , 1 96 1 98 ; o n la w o f an c es t ra -
l tu t io n a l v igo r, 5 7 3 ; as s o wn by h
h y
eredit , 5 2 9 ; o n re u c t io n o f v a ria d t ro t t in g rec o r s , 5 5 1 5 66 ; in u en c e d -
fl
b ilit y, 5 3 6 ; o n te ego n , 1 88 l y . of d l
ev e o p m en t u p o n 5 7 4 : b e c ause ,

d
Pe ig ree, im po rta n c e o f, 59 2 ; rec o r s d o f s ex , 6
5 7 5 70
—
-
.

o f, 6 7 0 6 7 2 . Pri m ro s e, s ev en m u t a n ts in eig t gen h

d h
Pe igrees , fas io n a b le, 5 9 5 . e ra t io n s o f , 1 2 7 .

l fl
Pe o ric o we rs i n t o a d fla x , 1 1 5 1 1 8 -
. l
Pro b a b e err o r, o f c o effic ien t o f v a ria
Perfo rm an c e, as a gu i e t o b reed in g d b ili ty, 4 4 1 ; o f m ea n , 44 0 ; o f s ta n d
p o wers ,
55 58 6 7 ; rec o r s i n p
-
a n t d l a rd d
ev ia t io n , 44 0 ; m ea n in g o f ,

b ree in g, 6 50 d .
4 3 7 44 0
-

Pfeffer, exp erim en t s o n c em o tro p is m , h Pro b a b l e erro r an d d ev ia tio n ll

i u s t ra t e d ,

2 7 3. 44 1 -
4 43 .

Ph t y t h i 40
o os n es s , 2 . Pro g ress io n , a few o ffs p rin g ex t rem e,
Ph t t i 48
o o ax s , 2 .
4 92 4 98 ; in o il c o n t en t , 4 96 4 98 ; in
- -

Ph t t
o o o n us , 2 48 .
p ro tein c o n ten t, 4 93 4 9 ; o rigin o f
5
-

Phy i l gi l l ti
s o o ca 0 1 58 9
s e ec on, 2 , . the exc ep t ion a l in i v i u a , 4 99, 500 d d l .

Phy i l gi l it 4 7 5 08
s o o ca un s , 1 , 1 , 1 2, 2 h d
Pro o f b y m et o o f ins ta n c e, 1 8 7 , 3 66 .

21 3 ; ff t
as a d b y d ec et i 7 0 re uc on, 1 . Pro t ein c o n t en t o f c o rn , effec t o f s elec
Pig s, l im ti ti f 3 7 5 ; d l p
ac c a za on o , ev e o t io n u po n . 4 4 5 , 4 4 6 ; p ro g ress io n in
m t en f 58 ; me i ti
o , i ti i r s c v ar a on n d
b ree in g fo r, 4 94 , 4 95 .

d igi t f 63 s o , . l
Pro t o p as m , a c em ic a s u b s t an c e wit h l h
Pl i
a na r an , g t i re en eraf 3 3 on o , 2 1, 2 2, h l
c em ic a p ro pert ies , 1 4 4 ; a c t i v i t o f, y
33 4 33 5
, ; g t i f
re en era t i g on o s a rv n , d u e to ex tern a as we as t o in ter l ll
3 72 g t i
re en era f w h p l i t 335 on o en s l
n a l im p u s es , 3 98- 4 0 1 ; as in flu en c e d
d i g 63 9—
; , , .

Pl an t b ree 65 ; n d tg 1 a v an a es h l
b y c em ic a a gen t s , 2 6 4 2 8 5 ; effec t
f 63 9—
-
,

an d l im it t i a 64 ons o
pl t y , 1 o s s o f c o n ta c t u po n , 2 3 3- 2 3 6 ; effec t o f
t m f
e 64 4 64 6 ;
o r, il -
d lt so an cu u re g ra v it
p y
u o n , 2 9 ; eff ec t o f
3 e a t a n d h
con d it i f 64 6 4 3 ; y tem f
o ns o r, 1- s s s o co l d
u on, 2
p 55 ; i r rit a t io n o f,
3 98 ;

p l an t i g i n6 43
— 6 49 n, . effec t o f ex t ern a a g en ts u p o n , 3 96 , l
Pl an t li ff t
c e, f e d it i ec p o con o ns u on
3 97 p ; s ic a b hy
as is o f if e, 1 4 2 l
, 1 43 l
re p d ti ro uc 0 ; p t
on,h g 1 t i 2 ar en o en e c l
re a t iv e s t a b i it a n d m u tab i i t l yo f, l y
on l y t h igh t m p t
a 8 e era u res , 2 1 . 2 95 , 3 4 6 .

Pl t
a n s, e ff t f h t p ec o
g w t h f
ea u on ro o , Pro to zo a , b u t o n e ma t u ra t io n iv is io n d
2
55 5 7
-
; 2 g t i re i 3 5 en era on n, 2 . in , 1 6 s .

Pl to y t m f p l ti g i b di g
s s e o an n n ree n y l d
Pu rit o f b o o lin es ess en t ia , 5 7 9 l .

w k 64 4 6 4 6
or ,
-
. yh
P t o n , m eris t ic v a ria tio n in v ert eb ra
Pl mu l ti f 3 3
s , ev o u on o , 1 . a n d rib s o f, 3 9, 4 0 .

P io son i y im m v it y t 3 0 8 un o,

l im t i ti t 308 —
, .

P io s o ns, ac c a
3 za
; on o, 1 1 Q ua l it a t iv e effec t s o f fo o , d 2 2 8- 2 3 0 .

t lyt i
ca a 66 67 f m i
c, 2 , t 2 70 ; ro n s ec s , 2 Q u an t i t a t iv e effec t s o f fo o d , 22 5
-
2 28 .

t i
ox c, 2 68 .

P lo ar b di o 64 ; f m ti
es , 1 f 67 ; f or a on o , 1 or R a c e, a ffec t e b y in t ern a in d l fl u en c es ,

m ti f i ll t t d 6 8 f t il i t i 1 9 6- 2 1 7 ; ex t in c t io n o f, 4 1
a on o , us ra e , 1 er za on
5 .

b y 79 80 ; formation of second 67
, l , 1
,
1 . d
R a ia l s m m et r , 4,
y y 3
 I N D EX
7 4
2

d
R an o m s a m p e 4 2 0 l , . R ev ers io n a n d a t a v is m , 3 0 5 .

l
R a t , ta i o f, g ra ft e in t o b ac , 1 0 7 d k . h
R eo ta x is , 2 3 5 .

l l
R a t io n a s e ec t io n , 5 9 2 , 5 93 . h d
R in o c ero s , ev e o p m en t o f fo o t o f , 6 0 l .

R a ts , v a ria t io n in igits o f, 5 8 d . R ib b ert , gra ft in g m a m ma r g a n o n e a r y l d

R ea u m u r, exp erim en ts in regen era tio n , of u in ea p i , 6
'

g g 33 .

16 k
R o eb u c , m e ris ti c v a ri a tio n in h
i
. o rn s

R e ip ro c a l c ro s s 5 2 5 , 6 1 0 o f, 6 6 6
— 53 5
.
, , , .

d
R ec o r s , in a n im a b ree in g, 666 6 7 2 ; l d R o ma n es , o n in s tin c t , 3 89 ; o n t ra n s
l
in p an t b ree in g, 64 5 - 6 4 9 ; o f t he d m iss io n , 3 54 ; o n t ra n s m iss io n o f
er , 66 6—
h d 6 70 ; o f p e igree, 67 0— 67 2 d . l
m u ti a t io n s , 367 .

R ed fi eld , C a s p er L , o n p rep o t en c , 5 7 4 . y R o o ts , d
ev e o p m en t o f, lb y ex t ern a l
o n t ra n s m iss io n , 4 07 , 4 08 . d
c o n it io n s , 1 04 ; g ro wt o f, i n r u n h
d
R e u c t io n , a c a u s e o f v a ria b ilit , 1 63 y n in
g wa t e r, 235 .

1 8 1 , 1 7 5 , 1 7 6 ; a p p a ren t p u rp o s e o f , d
R o s e, rec o r o f, Wi t N o ra , 7 8- 8 0 h .

1 6 7 ; c o n n ec t io n wit m u ta t io n , 1 8 0 ; h h
R o u let te w eel, ho w mad e u p , 3 6 5 .

d
en d p ro u c ts o f, 1 6 7 , 1 68 ; h o w a c R o u x , exp eri m en ts u po n s egm en t in g
l hd
c o m p is e , 1 69 ; i l u s t ra t e , 1 68 ; in l d f ro gs eggs , 343 ; o n p repa ra t io n o f
’

l
t h e m a e, 1 69, 1 7 0 ; in th e fem a e, l a n t i to x in ,
3 1 0 .

1 6 5 1 6 9 ; in m a e a n d fem a e c o m
-
l l y
R o w s s t em o f p a n tin g in b ree in g l d
p a d
re , 1 64 ; i n p a n ts , 1 7 1 1 7 3 ; i n l wo r k 64 6
,
-
64 9 .

l
a n im a s a n d p a n t s c o m p a re , 1 6 5 ; l d
l
o s s es s u s t a in e d
b y, 1 7 0 ; o p p o rt u n i S ac h s , ex peri m en t s o n
gro wt h of p l a n ts ,
t ies fo r ac c i en t d
u ri n g 1 7 1 ; W eis d , 2 56 .

’
d
ma n n s p re ic t io n c o n c em in g 1 7 3 , , l
S a a man d er, regen era t io n in , 3 1 6 .

1 74 ; res u ts in o ssl o f c ro m a t in l h l
S a in e s o l
u t io n s , effec t o f, u o n
p e v e d l
m a t ter, 1 7 0 , 1 7 3 ; s ign ifi c a n c e o f 1 7 0 o p m en t , 2 8 2 - 2 8
, .
5 .

R e flex a c t io n t h e b as is o f i ns t in c t i v e l
S a m p e, ra n o m , 4 2 0 d .

a c ts , fly
S a w , a n t en n a o f, ev elo p e in t o a d d
R egen e ra t io n , b ea rin g o f, o n s t a b i it l y f o o t, 4 3
—
.

of l
iv in g m a t t er, 3 1 6 3 3 2 ; b y t ra n s S c h m a n k ewit s c h , exp erimen ts wit A r h
fo rm a tio n , 2 2 ; c a ra c ter o f r es t o re h d t em ia , 1 0 2 , 2 8 3 .

p a r t in , 3 2 6 , 3 2 7 ; e ff e c t o f a g e u p o n , h
S ea u rc in , L o eb s exp erim en ts wi t
’
h ,

33 1 e ffec t o f o we rin flg p e r io u p o n, d 2 78, 2 8 2 .

33 1 ; e f fec t o f fo o u po n , d 3 7 3
2 2 8 ; l
S ea , v a ria tio n in igits o f, 5 7 d .

—
,

effec t of
g ra v it u p on, y 3 2 9 33 ; 2 , h
S ec retio n s c em ic a ac t io n o f, 3S3 , 3 8 4 l .

l
effec t o f ight u po n , 3 2 8 ; effec t o f d d
S ee p ro u c tio n a b u s in ess , 6 50 .

t em p era t u re u p o n , 3 2 7 ; fi rs t i n fo rm , d
S ee lin gs , res po n s e o f, t o gra v it , 2 3 6 y .

af terwa r d
in s ize, 3 1 7 ; g ro wt in , h d
S ee s , e ff ec o f m is t u re u po n , 2 3
t o 2

n o t u n ifo rm , 3 1 7 , 3 1 8 ; fro m o b iq u e l S egm en ta t io n , e en en c e

p d o f, u p on d
s u rfa c e, 33 4 ; e t e r h
o m o r h o s is
p i n wa t er c o n t en t 1 78 ; g e o m e t r i c a l
—
, ,

33 2 ; in t e r n a f a c lt o rs in , 3 3 33 5
2 ; i n h
c a rac t e r o f c ea v age in , 3 3 9 ; l not

a n im a s , 3 1 6 ; l i n ea rt wo rm s 3 1 7 h , d d
ep en en t o n fert iliza t io n . 1 7 8 .

3 2 0 ; i n e m b r o s an d e y gg 3 s , 2 4 , 3 5;
2 l
S e ec t io n , 5 7 7 5 9 8 ; b
-
e m i s es a n d l h
in fis h, 3 1 8 ; i n h
ighe r a n im a s , 3 2 5 , l d
a c c i en t a l j i n u ries b ea rin g o n , 5 90 ;

3 2 6 ; i n t h e p a n a ria n , l 3 2 1 , 3 2 , 334 ,
2 c es s a t io n o f , 2 88 ; effec t o f, u p o n t pe
y
3 35 ; i n
p l
a n ts ,
3 2 5 ; in s a a m a n er , l d an d l y
v a r iab i it 4 4 5, 4 4 6 ; gen e ra
, l
3 1 6 ; i n a c t i n ia n ,
3 34 ; la t era ,
333, l p p l d
ri n c i les in v o v e m ,
5 8 1 -
5 2 h is

33 5 ; n o t a w a s l yc o m p e t e, 3 8 3 2 0,
1 — l k d d
t o ri c a l n o wle ge o f t he b ree es s en

3 3 3 5
2 . 2 : p o a r it l y i n .
3 2 0 , 3 9 33 3 :
2 — l
t ia in , 5 7 9 5 8 1 d i ea ls m ,
,
°
'

h
w a t d et em i in es c a ra c t er o f re h d
i mp o rtan c e o f p e igree in , 5 9 2 in °

s t o re dp a rt , 3 33 c reas e d n u mb er o f p o i n ts in
—
. ,

d
R eg res s io n , a v a n ta ge a n d d is a d v a n d fl
in irec t effec ts o f, 4 4 7 44 8 ; in u en c e
t a ge o f, 4 8 5 ; a s t o s t a t u re, 4 7 9 4 8 1 o f a e in
g 8
5 9ll y
; fa
, da c o f “
f o u n a t io n

d ia g ra m o f 4 8 9 ; o ffs p rin g m o re m e
, l l
fem a es in , 59 5 5 96 ; im it o f p o wer ,

h
d io ere t a n p a re n ts 4 8 4 4 86 , . d y
o f, t o re u c e v a ri a b ilit
5 34 5 3 7 , ,
°

R e g res s io n c o e lfi c ien t , 4 66, 4 8 7 —4 90 n a t u ra l l s e ec t io n a lwa s a t wo r k

g
.

R eg res s io n t a b e 4 7 9 4 8 2 l ,
-
.
5 8 8 ; n eed l o f a rge n u m ers fo r, 5 8 4

R e v e rs io n , 1 6 , 1 9 2 1 94 -
. n ee d o f t h e a c t u al tes t fo r, 5 86 ,
 6 I N D EX
7 2

y
T es t in g o u n g fem a es , 66 1 l . l y
fert i i t , 4 4 9- 4 5 1 c o effic ien t o f, 4 3 3
T et ra s , 1 66 d . d et erm i n e d
f ro m gro u ps , 4 2 6 ; d ev i
h
T igm o ta xis , 2 3 5 . a t io n f ro m t p e, 4 2 y effec t o f s e ec l
h l
T remm a t o o gy, c o mp a re wi t ev o lu d h t io n u po n , 4 4 5 , 4 4 erro n eo u s c o n
t io n , 2 ; d
efi n e , 1d m o re t a n a s t u h dy c ep tio n s o f, 4 2 5, 4 2 6 ; ig es t in h h
h l y
in m o rp o o g , 8 ; p ro b e m s o f, o u t l l l
f ert i e s o i s , 64 2 ; in heig ts o f b ro t h h
l d
in e , 3 5 -
. e rs , 50 1 in o il a n d p ro tein , 4 4 5 , 44 6 ;
T hy dro i , eff ec t o f e x t i rp a t io n o f, 3 8 3 . hy l h
in p s ic a c a ra c t ers o f c o rn , 4 4 7 ,

T o a d fla x , experim en t s wit , b y D e V r ies , h l

4 4 8 ; im i t a tio n s o f, 8, 9 ; im it t o t h e l
1 1 5- 1 1 8 . d
re u c t io n o f ,
53 534 7 ;
-
m e as u re o f ,

T o x ic po is o n s , 2 68 d
b y a v erage ev ia t io n , 4 2 7 ; m eas u re
—
.

T ra n s m is s io n , 34 7 4 71 e t e r o
g he n e o u s , o f, b
y d dd
s tan a r ev ia t io n , 4 2 8 - 4 1
3
4 2 6 ; h o w c a r ac t e h
rs b e a v e in h
, 473 n a t u re o f, 1 0, 1 1 ; o f ifferen t c a r d h
47 8 ; m a n n e r o f 4 2 0 -
4 3 1 ; e ff ec t o f a c t ers in s a m e po p u a t io n , 444 ; o f l
—
,

ac c i m la t i z a t io n u
p o n 3 7 4 3 8 0 ; e ff ec t s exes ,
5 7 0 -
57 3 ; u t i m a tle u n i t o f
1 5 - 1 7 ; u n i ts o f, 2 08 —
, ,

of d l
ev e o p m en t u p o n , 4 0 7 - 409 ; o f 213 .

dis eas e, 3 68 , 3 8 4 o f effec ts o f fo o , d V a ria t io n , b u d, 1 8 1 ; ca u s es o f, 1 4 1

37 3 7 4
0 -
; o f e ff e c t s o f u s e a n d d is
3 4 7 ; c a u s e b y db is e x u a l r e p r o d uc

u s e , 4 0 4 4 7; 0 -
o f a b i t s , 3 8 h
6 -
4 0 3 ; o f t io n , 1 60- 1 63 ; c a us e b y c e d iv is io n , ll d
y
imm u n it , 3 8 2 ; o f m u t i a t io n s , 3 64 l 1 5 5- 1 8 1 ; c a u s e d
b y extern al in flu
3 6 8 ; o f s ta t u re, 4 80 ; o f v a ri a t io n , en c es , 2 2 0- 2 9 ;
3 c a use b y d
r ed u c

34 8 4 1
-
7 ; n o t u n e ss g e r m l i s a ff ec t e , d t io n p roc e ss , 1 6 3 1 8 1 ,
-
1 7 5 , 1 76 ;
4 1 6 , 1
4 7; o ff s p ri n g n o t l i e p a r e n t sk , c a us es o f, m u s t b e s t u i e , 1 4 1 d d con

8 8 ff s rin m m i d d
fi n e to ra c ial c a rac ters , 3 58 ; c o n h
—
4 o p o re e o c re
4 2 , 3 ; g
h
t a n t h e p a ren ts 4 8 4 4 86 ; o rigin o f ,
-
t in u o u s a n d d
isc o n t in u o us , 1 8 2 2 ;
t h e exc ep t io n a i n i v i u a , 4 99, 5 00 ; l d d l c o rre a t e , 1 6 ;l d d
o es n o t e x ten to d
p r o g r es s io n i n, 4 9 2 -
4 9 8 ; w a t i s h -
l
n o n iv in g m a t ter, 2 3 ; d u e to ag e o r

t ra ns m it te , 5 1 1 d . l
s ta en ess o f germ, 1 8 2 ; d u e to t em

T rem b ley, ex peri m en ts o n reg en era tio n ,

’

p e ra t u re , 2 6 2 -
2 6 4 ; f u n c t io na , 75 l
3 1 6 . 1 09 ; f u n c t io nal , d u e to age, 94 fu n c
T ro t ters , c o rrela t io n b et ween c o o r, l t io n a l, b etween d
iff eren t in iv i u a ls d d
s ex , a n d s p ee , 4 68- 4 7 1 d . o f t he sa m e s pec ies , 77 9
- 1 ; f u n c

T ro t t i n g r ec o r s s o win g p repo t en c , d h y t io n a l, d u e t o u s e o r d
is u se, 9 5 , 96 ;
55 5 1 66
-
. l
f u n c t io n a , f ro m da y t o da y sa m e i n ,

T u m o rs 99, 2 7 1 d d l
iv i u a , 9 1 - 94 ; f u n c tio n a , i n u c e l d d
b y ex t ern a l in u en c es , 98, 1 0 1 -
.
,

l d
T u rt e, o u b e ea o f, 6 7 l h d . fl 10 ;
5
d l
T win s , 1 7 6 ; i en t ic a , 1 7 6 ; f ro m a s in fl
f u n c t io n a l, in u en c e b y fee , 96 97 d d
h o w fa r p o ss ib e, 2 9 —
,

g le o v u m, 1 76 . l 3 4 6 ; in u c e d d
y
T p e as d
is t in c t f ro m t he in iv i u a d d l b y fo o d , 2 2 5 2 30 ; in u en c e o f m o is
-

—
, ,

35 2 c o n c ep t io ns o f , 4 2 0 4 2 5 ; effec t -
t u re u po n , 2 30 2 3 3 ; in u en c e fl by d
—
;
o f en v i r n m o e n t u p o n , 2 9 0 -
2 93 ; e f l y
ferti it , 1 96 2 00 ; i n u en c e b y t h e fl d
feet o f fertilit u po n , 1 98 , 1 99 4 491 , d
r ep ro u c t iv e f u n c t io n s , 1 00 ; in flu
45 1 ; e ff e c t 0 e e t io n u p o n , 44 ,
5 l h h h
en c ed b y s ig t c em ica l c an ges
—
,

6 m t b i it f l y2 8
9 3 5 0 n a t u ra l 210 2 1 1 h in c em ic a l c o m pos it io n o f
—
a o ;
d —
44 ; u , , ,

4 2 2 ; p o w e r o f s e e c lt i o n t o m o if , d y s ee s , l y
c o rn , 83 86 ; in fert i it , 90

53 5 7 4 4 ; s e e c t io nl s t a n a r fo r d d
4 2 0, l dy
i n gen era b o l fa c u t ies , 8 6 ; in m ea t
—
-
,

4 52 ; s t a b i it o f l y
2 9 6 2 9 7 5 , 5 44
4 1
p d lk
ro u c t io n , 8 1 8
3 ; i n mi s e c ret io n ,
—
, , , .

y
T p e a n d v a ria b i it , 4 1 9 4 5 1 l y -
77 8 1 9 93h
1 -
; in p a rt e n o g e n e t ic re
—
.
,

p rdo u c t io n , 1 7 7 1 8 0 ; in s u a r p ro
g
U s e a fu n c t io n o f s t ru c t u re, 3 8 7 . d d
u c t io n , 86 ; i n ra c e, c a u s e b y ex
U s e an d d
is u s e, eff ec t o f, 2 8 5 2 90 ; ef -
l fl
t ern a in u en c es , 2 90- 2 93 ; in ra t e o f
fec t o f, u p o n f u n c t io n a a c t i v ity, 9 5 . l c elld l
iv is io n , 3 40 ; in v ita fu n c t io n s ,
9 6 ; e ff e c t s o f w en t ra n s m it t e 4 0
, 4 h d , 8 7 89 ; in tern a l c a u ses o f, 1 5 5 - 2 1 7
-

fl
4 0 7 ; in u en c e o n t ra n s m is s io n , 36 3 . k d h l l
in s o f 1 7 2 3 ; m o rp o o gic a , s u b
,
-

s ta n t i v e , l
m eris tic , fu n c tio n a 22 ; ,

V a ri a b il ity am o ffs
p rin g f s am e h l l
m eris t ic , 33 7 4 ; m o rp o o gic a , 2 5
—
, ong o -

p a ren t s,
5 00-
5
04 ; as a ff ec t e d by 2 9 ; s u b s t a n t iv e, 3 0 3 2 ; q u an t ita t i v e
 I N D EX 7 7 2

an d q u a l it a t iv e,7 ; 1 n a tu re o f,
35 6 173 , 1 74 ; p redic t io n c o n c ern in g l o ss

3 6 4 ; n o t n o ta b l l ess y
in p a r t h e n o .
of h ere d it y ar ma tt er, 1 73, 1 74 ; on

g e n e s is , 1 7 8 ; t r o u g h in h e r h
i tan c e o f t ra n s m iss io n , 3 54 .

d
mo ifi ca tio n s 2 92 un its o f, 2 08 2 1 3 ;
,
- W h a le , v aria t io n in d igi t s o f,
57 .

u n iv ers a it l y
o f, 7 , 8 . W h ea t l im ti ti
, f
ac c a za on o , 37 6 ; in flu
V a ria t io n s , d u e to c a u s es in t ern a l to t he f l
enc e o lity p oc a u on, 2 2 2.

germ , are t ra ns m it te , 3 48 ; d u e t o d W h it H ghe, igi t f C li t

u , or na o r o n on

ex t ern a l fl
in u en c es , t ran s m is s io n o f, g p ra
34 e, '

3 4 8 -
4 1 7 ; d u e t o c a u s es n o t a ff ec t i n g W h it y f d f
ne p l, t i un3 o r ex o ra on, 02 .

t h e germ n o t t ran s m itt e , 4 1 6 4 1 7 ; d , W ill w ff t f g it y

o , e gecw h f o ra v on ro t o ,

o c c u r a c c o r in d t o t h e b i m i l t h e 2 8
W ili
g n o a .

o rem ,
5 0 8 5 09
, .
Jh o n, igi t f b l kb
o y
n, o r na o r o ac e rr ,

d
V a riet ies , u ra tio n o f, 544, 5 4 5 1 2
W ili
. .

d
V igo r, o ft en o pp o s e b y s e ec t io n , 58 3 l . th
o n on h m m 63 4 e c ro oso e, .

ll
V ita im i ts as to ig t, 2 44 l h . Wi g n p , m su y 43 er n u era r , .

V oc h t i n g s e x perim en ts o n gra v it a n d
'
y Wi g n p m
s, su y 5 ern u era r , 1 .

p l y
o a ri t , 2 7
3 . W rit in g with the feet , 2 86 , 2 8 7 .

la c e o n t he a c c es s o r c ro mo s o m e, y h
n 6
X en ia , 1 8 3 , 1 84 .

3
W a t er, eff ec t o f, u po n gro wt , 2 30 2 3 3 h —
d
.

l
W a t t es , 4 6 V o u n g b ree ers , 67
5 .

th
.

yl d
W a a n , o rigin a to r o f p u m , 1 33 l .
Y u c c a mo , 1 0 5 ; 1n s tin c t iv e ac t s o f,
W eepin g v a ri eties , 1 1 2 3 88 .

l l
.

W eis m an n , expe rimen ts wit b u t t er h Yu ’

e s fo rm u a, 6
4 5 , 4 57 .

fl
ies , 2 6 2 - 2 64 ; o n ea t p o in t , 2 0 2 ; d h
l l
o n g e rm in a s e ec t io n , 2 1
4 o n o rig in a , ex erim en ts in
Zo j p reg e n era t io n o f
h
o f c a ra c t ers , 4 1 , 4 1 4
3 p re ic t io n o f, d l
b a s t o m eres , 3 2 5 .