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 SHORTER ARTICLES AND DISCUSSION

STUDIES ON INBREEDING. VI. SOME FURTHER CON-

SIDERATIONS REGARDING COUSIN AND
RELATED KINDS OF MATING'

IN the firstof these studies2the writerdealt with the results,

in so far as concernedcoefficients of inbreeding,which would
followcontinuedbrotherX sister,parent X offspring, and cousin
X cousin mating. Regardingmatingsof the lattertype it is de-
sired now to record certainfurtherfacts.
PEDIGREE TABLE I (HYPOTHETICAL)
To ILLUSTRATE THE CONTINUED BREEDING OF FIRST-COUSIN X FIRST-COUSIN
SINGLE COUSINS

A ~ ~b~~~~~~~~
l SF {A {4
_~~~~~~~~
Kf~~~~~~~~~
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 6__ _ _ 1
Generaion
nmber a 2I
V~ln~be1
Genec1tiOI1 1 3 4 .5

1 Papers fromthe Biological Laboratory of the Maine Agricultural Experi-

ment Station No. 85.
2 AMER. NAT., Vol. XLVIII, 1913, pp. 577-614.
570
No. 5S5] SHORTER ARTICLES AND DISCUSSION 571

There are, of course,two possible sorts of firstcousins,single

and double. In thefirstcase one of the parentsof any individual
is a brother(or sister) to the one of the parentsof the otherindi-
vidual in the mating. In the second case, both the parents
occupythis relationto the parentsof the otherindividual in the
mating.
These twosortsof firstcousinshipare shownin Pedigree Tables
I and II.
PEDIGREE TABLE II (HYPOTHETICAL)
To ILLUSTRATE THE CONTINUED BREEDING OF FIRST-COUSIN X FIRST-COUSIN
DOUBLE COUSINS

_0

(
~0_

ehiell {rere
l op
b

page 09 n 9 fteerirlae.Tepeetvle

Generation number 12 3 4 5

Thevaluesofthecoefficients -ofinbreedingfor continuedsingle

and double cousin matingare shown in Table I.
It will be seen that Pedigree Table I and the third columnof
Table I are differentfrom the correspondingvalues givell on
pages 591 and 592 of the earlier paper. The present values
should be substitutedfortheearlierones,w11hich w-erebased"Lmon
572 THE AMERICAN NATURALIST [VOL. XLIX

the erroneous assunimtipnthat half the double-cousinvalues

would give single-cousinvalues.

TABLE I
VALUES OF THE SUCCESSIVE COEFFICIENTS OF INBREEDING IN THE CASE OF
CONTINUED COUSIN MATING

of
Coefficient AncestralGeneration Coefficient
forSingle Coefficient
forI)ouble
Inbreeding Inclulded Cousins Cousins

zo 1 0 0
Z1 2 0 0
Z2 3 1 _D2.00 50.00
Z 4 50.00 75.00
Z4 5 68.715 87.50
Z5 6 81.25 93.775
z6 7 839.06 96.98
Z7 8 93.75 98.44
Z 9 96.48 99.22
z9 I
10 98.05 99.61
ZIO 11 98.93 99.8O
Z11 912 '9.41 99.90
Z12 13 99.68 99.95
Z1.s 14 99.83 99.98
1i4f 1D5 99.91 99.99
Z15, 1B 99.95 99.994

2/0 6 8 /0 /=' /.r

C f 5
1160G./. -t

8)lothel x-itlllednuhlinteslcsiecefcenso gh

K /~~~~~~ECA ~~
FIG.1. C /ve /fibed hwn a helmtn aeo otne

brohe
xsite/reig hri h ucsiecofiinsonrei-hv h
maxmumvales /b otne aetxofpigmtn;()cniudfrt
cosnxfrs-osnmai-weeth osnhi
tinedirs-cosisdula,2XC)
K'he x ~s-osnmtn
on hr h cuisii
/0 le
ad()cn
i-e('xC)
14

FIG.co.tCures aio ofinbreed ing, shoing( the of

limitingv case xconine
No. 5S5] SHORTER ARTICLES AND DISCUSSION 573

The data of Table I are given graphicallyin Fig. 1, together

withthe curve for brotherX sister and parentX offspring.
From the table and figureit is seen that with continuedin-
breedingaccordingto any one of these four types the coefficient
approaches the value 100. The rate of approach is different,
however,in the different cases. The curves fall into two pairs.
The brotherX sister and the double cousin curves are precisely
alike so far as concernstheir curvatureor shape at any given
point. Similarly,the parent X offspringand singlecousincurves
are of the same shape. The essentialpoint of clifferenceis that
the cousin curves lag a generation behind tlMeothers.
Let us now considerthe question of the degree of inbreeding
followingO'continuedmatings of the avuncular type of relation
ship. Pedigree Table III gives a pedigreein which each mating
is of uncle X niece.

PEDIGREE TABLE III (HYPOTHETICAL)

To ILLUSTRATE THE MATING OF UNCLE X NIECE

VI

h~~~~~~
In)
So~~~~~~~~~~(
IJ

IL1
f2

~lb

a_ ~~~~~~I _ (1A
(1 ~~~~ h

Generation number 12 3 4 5
574 THE AMIERICAN NATURALIST [VTOL. XLIX

of
From this table it appears that the values of the coefficients
inbreedingAwrillbe exactlythe same for this type of mating as in
the ease of single cousin mating. Or, in other words,Z's form
series.
the followi-ing
TABLE II

VALUES OF COEFFICIENTS OF INBREEDING FOR CONTINUED

UNCLE X NIECE AMATING

Coefficient Number of Ancestral Generations Value of Coefficient

0 1 0
Zi 2 0
Z. 3 25.00
z/ 4 50.00
Z/ 5 68.75
Z. 6 81.25
etc. etc. etc. ns in Table I

From the data presentedin this and formerpapers it is clear

that inbreedingcontinmcdl for about ten generations,quite re-
gardless of the type of mating. providedonly it 'be conti'auously,
followed,leads to within one or two per cent. of complete"con-
"
centrationof blood. The bearinuo of thisresult-uponthe general
questionof the degreeof inbreedingwhichexistsin the ancestry
of our domesticanimals to-day is obvious. To considerbut a
single case: In 17893 a law was passed prohibitingthe importa-
tionof cattleinto the Island of Jersey. Hence it followsthat all
pure-bred Jersey cattle of the present time must be of the
descendantsof the relativelyfew animals on the Island in 1790.
Taking threeyears as about the average generationinterval in
cattle,this means about fortygenerationssince the Island was
closed to importation. The concentrationof lines of descent
Whichmusthave occ-urred in this tiue merelyby the droppingof
lines and quite regardlessof the type of matingis obvious. This
is not the place to go in detail intothe discussionof inbreedingin
Jerseys,especiallyas I hope shortlyto publish the resultsof an
extensivestudyof thismatter,but it seemsdesirableto emphasize
the bearing of sluchhypotheticalpedigrees for particular types
of matingas are given in this and earlier papers,on the general
problemof inbreeding.
It is possible to extend now somewhatthe table of general
equations oiven by Jenninz.s4for coefficients of inbreedingafter
3 Teste Rees's Encyclopedia and H. S. Redfield, Nat!. Stockman amd
Fanner, December 15, 1892.
4 AMER. NAT., Vol. XLIII, p. 695, 1914.
No. 5805] SHORTER ARTICLES AND DISCUSSION 575

9ngenerationsof each particular type of mating. We have the

followingvalues, where n denotesthe numberof ancestral gen-
erationseoiieerned,or, as Jenningsputs it, the numberof suc-
whichhave takenplace.
cessiveinbreeclings

Type of Mating Coefficient of Inbreeding

Self-fertilization ...... .................... 21-1

On.
*1 22
Brother X sister ... .......................
- 2n
G'ousiii X cousin , single............2
-2 22 (from n== 2
Cousin X cousin, double *.-.-.-.-.... 0tto mt - CO
Parent X offspring. ........................ 211
-

Uncle X jiece*e..................------- ' - 2n

RAYMOND PEARL

AN ATTEMPT TO PRODUCE MUTATIONS THROUGH

HYBRIDIZATION
THERE is no more interestingprobleni to the experimental
evolutionistthan the one relating to the cause or causes of the
originof mutations. Until we are able to solve this problemAwe
can only accept what the gods give in our breedingexperiments.
When a mutationarises it is usually a simple processto prod-uce
a pure stock. By mutation is meant any deviation from the
normal type which reappears in some of the descendants. In
the followingexperimentmost of the abnormalitiesthat were
foundneverreappeared in the offspring.
My experimentshave been confinedto the fruitfly,Drosophila
, a species kept for years "'under cultivation'" at
aminpelophcila
Columbia University. This species has proved to be very plas-
tic, throwingoffgreat numbersof mnutant forms. At the sug-
gestion of Dr. T. H. MIorganI crossed sone of these mutants
with wild stockof the same species fromwidelyseparated locali-
ties in order to test whetherthroughhybridlization mutations
arise in greaternumbersthan in inbred stock.
The idea that new forms arise from crossing more or less
closelyrelated species is an old one. One findsmanyreferences
in Darwiin's works to this conception. For instance, in the
" Origin of Species " Darwin says: