COMPARISON OF TANDEM AND INDEX SELECTION IN THE

MODIFIED PEDIGREE METHO.D OF BREEDING

SELF.POLLINATED SPECIES'
J. Pnstr" eNr R. J. Baxon
Researcb Station, Canadu Departtnent of Agricztlture, Winnipeg 19, Manitoba
Received Mry 22, 1969
ABSTRACT
Two-staee tandem selection and index selection were simulated for the modified
pedigree-method.of. breeding self-pollinated crops.,-We assumed that two
tralts we-re negatrvely correlited due to repulsion linkage;, and considered
correlations r"igi.rg from 0.00 to -0.95 correiponding to Iinkage intensities of
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0.50 to 0.05. Eath"trait was controlled b1' five'different loci with equal genetic
efiects" Three levels of environmental variability were assumed. Economic
values were set at 0.5, 1.0 and 1.5 for one trait and 1.0 for the other.
Index selection was suDeriot to tandem selection for all combinations of para-
meters simulated. The efficiency of tandem selection was increased substan-
tially by selecting the most v-aluable trait first. The eficiency.of index
seleitiori can be increased by more frequent esrimation of selection index
coefrcients,
We did not consider positively correlated traits.
INTRODUCTION
Many problems in the biometrical genetics of self-pollinated croPs exist and
a soluti,on to them would aid in the improvement of these important economic'
sDecies. Few studies have been made on Senetic covaliation between two of
rirore characters under selection, even thorigh the recombination of correlated
For personal use only.

traits is an important aspect of crop improvement.

In most plant breeding progfams, it is necessary to consider several traits
simultaneously. Three miin se'iection methods are available: index selection
(6,8), independent culling (9), and tandem selection (7). Hazel and Lush
(7) consideied the efficiencies of these three methods, assuming one-stage
selection for two uncorrelated traits with equal variances and heritabilities.
They concluded that index selection was the most efficient method.
However, plant breeders have made little use of index selection, but have
concentrated mbre on various forms of tandem selection in which they select
first for one trait and then for another. In view of the report by Hazel and
Lush of greater efficiency of index selection over tandem seleciion for un-
correlatei traits, and in view of the fact that many traits are correlated, it
seems useful to investigate the relative efficiencies of these two methods of
selection as they apply io cor-related traits in self-pollinated crops. That is the
purpose of this study.
MATERIALS AND METHODS
The modified pedigree method of plant breeding, an adaptation of a scheme
proposed by Gouliien (5) and desCribed by Brim (4), was chosen for study.
l*sentially, this method consists of advancing each F, plant in the population
by single seed descent. In F, and succeeding generations, one seed is used
from each plant in the population to propagate the next generation' When

'Contrib rtion No. 376, Research Station, Canada Department of Agriculrure, Winnipeg 19,
A4anitoba.
?ermanent address: Research Institute for Basic Agrotechnique, Brno-Hrusovany, Czecho-
slovakia.
Can. J. Plant Sci, Vol. 49,773-781 (1969)
773
 C{NADIAN JOURNAL OF PLANT SCIENCE [Vol" 49

the desired level of inbreeding is attained, each progeny, which traces to a

single F, plant, is bulked. The advantages of this method compared with the
pedigree system of breeding were discussed by Brim (4).
Consider selfing the F, generation for four generations to produce an F"
population. Selection is practiced on this generation and on the F" generation.
Now, suppose that the phenotvpes of the two correlated trairs are designated
X and I' and the corresponding genotypes are r and y. In tandem selection,
a proportion of the F. individuals is selected on the basis of trait X. Those
individuals are then selfed to produce the F, population. Selection for trait
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Z is then practiced on the F, population. For index selection, an appropriate

index, 1: b"X + b,Y, is calculated from the phenotypic, genetic and economic
parameters of the unselected Fo population ( 3 ). This one index is used to score
individuals in crch gencration.
To compare tandcm and index seiection, assume that two identical F.
generations lvere subjected to thc two methods of selection and that surviving
individuals in F, u'cre further subjected to the two methods of selectiori.
Selection intcnsities wcrc assumed to be constant for both methods of selection
and for both gencrations.
The expected responsc to selection can be prcdicted from theory for the
cwo methods. Let I desienatc the selcction differential expressed in standard
deviations rnd lct tlre geirorvpic i'arirnces and plrenotl pic variances of thc
For personal use only.

traits X and ]'be designatedbv V,, V,, V,, and V', respectively. C,,, and
C,' will denote the genotypic and phenotypic covariance betrveen the two
traits. To distinguisfi the F. p^m-.t.rs froir those of the F" generation, we
will use the integers 6 and 7 as subscripts preceding the parameter designations.
Thus, oC", refers to the genctic covariance betrveen traits X and I' in the F"
generation, while
"2" t.I.tr to the phenotypic variance of trait I in the F,
generation. Using this terminology, the following formulae represent the ex-
pected response of the two traits to the two methods of selection.
(i) Expected response in trait X due to tandem selection:

^G.:il:r,''""
/x \ lV c t vv

(il) Erpected response in trait f due to tandem selection of Y after X:

LG":;lt'', + i',v"
!u/x lzVv
(lli) Expected response in trait X due to inder selection:

:'i b,.6v, + b, 6c,, b" tV, I bttC'"

LG, It
aD

(io) Expected response in trait I' due to index selection:

.b,'eC," I b"'eV,, .b^'rC,, L br'tV,
-.l'v:/ -'
,D ,
-where
,D : rlb"".;v"+br"rvo+2Lt\jc, h : 6or7.
 November 19691 PEsEK AND BAKER-TANDEM AND INDEx sELECTIoN 775

If the changes in genetic parameters from Fu to F" could be derived from

theory, it wouli be p"ossible to evaluate AG" and AG,. Because this is not
possi6le with the theo'ry we know, we find it necessary to use the methods of
ittont" Carlo simulatioi to study the efficiencies of the two methods and to
determine the effects of selection on the genetic and phenotypic parameters.
In our simulation of the modified pedigree method of plant breeding,
the F, of two inbred lines was selfed to prod"uce 600 F" individuals. Each F,
individual produced progeny by selfing-, and one Pfo.geny was selected at
random to ptop"g"t" th""tt.it generatioi and so on until production of an Fu
population'ot oo"o individuals.- selection intensity wa-s s:t ^r 40"A in both
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g.ner"tions to oLrtain a final selected proportion oi rc"1.. According to Baker

\t1 , rc"1" is in the range that minimizes fhe coefficient of variation of resPonse
to selection.
A simulation program was dcsigned for the IBM 360-65 computer. . Ten
loci in a self-polHhate"d species werE simulated. The five odd-numbered loci
controlled trait X, and the five remaining controlled trait f' All loci were as-
signed equal genetic effecrs. If a locis was homozygous and dominant, it
c6nributed a"genetic value of 3.0 to the crait. A heterozygous locus had a
value of 2.5 and a homozygous recessive locus a value of 1'5.
A negative correlation was simulated b1' choosing the- genotvpe of the
F, such th-at the loci governing the tu,'o traits rvere in repulsion. A negative
For personal use only.

correlation was chose-n to approximate the common occulfence of negative

correlations between such economically important traits as yield and quality
(2). The degree of negative correlation rvas adiusted u1 aJ1g1in9 the prob-
abilitl' of rec6mbination=betrveen ad]acent loci. The probabilities of fecom-
bination considered were 0.50,0.35,0.20 and 0.05, and rvere the same betrveen
all pairs of loci. These recombination values resulted in genetic correlation
ranging from 0.0 to -0.95.
Three environmental variances we1e simulated: Vu: 9.0, 16.0, and 25'0'
These levels gave F. heritabilities ranging from 0.80 to 0.30' Three economic
values of trai-t X were considered 1a,: O.S,l.0 and 1.5) while the economic
value of trait Y remained constant (4, : 1.0).
For each combination of linkage intensity, environmental variance and
selection method including economic values in index selection, the following
information was obtained:- (i) genotypic and phenotypic means, valiance and
correlations in the F. and F" populations; (il) genotypes, genotypic and pheno-
typic values, and index scores bf tne selected individuals in the F" generation'
The experiment was replicated but, because no significant diffelences
existed between replicates, the results are Presented as means over two repli-
cates.
To check the cornputer program fol systematic error, the simulated
genetic palameters were substituted into the formulae for expected lesPonse
and the expected response was then compared with the lesPonse observed in
the simulaied populations. The correlations between the two measures of
response wefe greater than 0.90 for all comparisons made, indicating that the
simulation program was behaving properly.
 776 CANADIAN JOURNAL OF PLANT SCIENCE lVol. 49

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 November 1969] IESEK AND BAKER-TANDEM AND rNDEx sELEcrroN 777

RESULTS AND DISCUSSION

In a plant breeding program, most selection for quantitative traits is based on
the performance of inbred lines grown in replicated experiments. In this
simulation study, we considered only individual plants. We believe, however,
that inbreeding in Fu and F, is sufficiently high for the genotypic value of a
single plant to be close to that of its sibs. Although we have considered only
single plants, the results are directly applicable to practical plant breeding.
Theoreticallv. one would exDect selection for one or more traits to alter
the genetic varjance of traits and also the generic covariance among traits.
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Selection would affect these generic parameters directly and would alter the
phenotypic parameters .orir.qo.n... Table I shows'the results of selection
"r " selection resulted in decreases in the genetic variance
for trait X in the Fu. 'fhis
of both traits and a reduction in the genetic covariance betwEen these traits,
except where the initial genetic correla-tion between the traits was zero. Simi-
larly, index selection in the F. reduced the magnitude of these genetic para-
meters to an extent dependent upon the economic values of the two traits and
the magnitude of the initial sen-etic correlation. The reduction in masnitude
of the -genetic paramet.r, #", less when heritabilitv was lower. Li index
selectiori, the trait with the hiehest economic value wis subiect ro rhe greatest
changes in genetic parameters: This is because the most valuable trait-is sub-
For personal use only.

iected to th-e mosr intensive selection.

From these observations on changes in genetic parameters under selection,
we are able to draw conclusions aborit the efiect of selection on heritabilities.
When the environmental variance is constant, as in this study, selection reduces
heritability by decreasing the genetic variance. With index selection and
equal economic values oiboth iraits, heritability would be reduced at about
the samE rate for both traits. With tandem seiection, rhe heritability of the
first trait selected would be reduced most. This would occur whether or not
the first trait selected was the most valuable from an economic standpoint and
it emphasizes that, in tandem selection with negative genetic correlations among
traits, it is very important to select for the miost economically important traic
ln the hfst steD.
Whether ot ,rot negative genetic correlation resulting from linkage can
be reduced in magnitude by se-Iection is of interest both iheoreticallv -and in
practice. Changei in geneiic correlations are of major concern in applying
ielection index irocedrires to breeding programs. If'the correlations'ihingE
during a breeding program, it may b-e nec-essary to calculate new index c"o-
efficie-nts. Althoigi r"eductions ii genetic .r^rirrrces would be expecred to
incleas.e the magnitude of the g.n"-ti. correlation, there is a corrisponding
reduction in thJ magnitude of ihe senetic covariance. This results in tittG
net change in the ginetic correlatiJn coefficient except for index selection,
where the relative economic weights are different. The results are shown in
Table 2.
Changes in genetic parameters other than the genetic correlation can also
influence ihe caiculation of index coefficients in iidex selection. Therefore,
we calculated new index coefficients for the F, generation and compared them
with those calculated for the Fu generation. If-we calculate the,ratios B, and
 778 CANADIAN JOURNAL OF PLANT SCIENCE lVoI. 49

Table 2. Genetic correlation coefficients before and after tandem and index selection*

Index

V" Initial Tandern a, : !,a, : I o, : 1,a" - | a"- 11,4, : I

0.05 9.0 .85
- .89 -76 - .95 -79
16.0 - .85 -.81 - .96 -.83
25.0 - - .82 - .95 - .82
11
0.24 9.0 - .69 - .55 - .59 n1 -67
16.0 -58 - .63 - .70
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25.0 - .63 - .70 - .67

0.35 90 - .32 - .28
.lt
-.49 - .45
160 -_28
.33 -38 - .41 - .38
250 -.Jt - .41 -
J/

0.50 9.0 00 .02 .00 .03 - .02

16.0 06 - .06 -22
250 06 - to -07 -16
tp : plolabilitv of recotnbination, 11, : enrironmental vatiance

B, of ,b. to ,b" and,,bo t:o ,b,, rcspcctiyely, the resulting ProPortions should
have a value of unin'if thcrc have^been no changcs in the values of the inder
coefficicnts due to sciection. These results are pre;cnted in Table 3. Substantial
For personal use only.

changes occurred when economic values of thi tl-o traits were unequal and the
geneiic correlation rvas highlv ncgative. \,Ve conclude that a single index-cal-
Eulated on the initial popuiation sl-r"ould bc used for onlv a few- generations if the
genetic correlation large and the tlaits have unequal economic values. A
ii
ieduction in the magnitude of the genetic correlation may be a good indicator
of the need to calculate a new index.
The effectiveness of the two selection methods was assessed by comparing
genetic gains. Expected genetic gains ale often not realized because of such

-lable 3. Measures of chanqe in index coefficients B, and Bn+

w: - 2, ur
^ -L - \
^ -1
R R R R R
V"

0.05 9 .0 959 97 03 0.98 1.70 -0.65

16.0 224 90 01 098 1.80
25.0 338 87 05 .20 2.23 +.+6

0.20 9.0 050 49 12 .01 .30 091

16.0 0.62 07 05 54 0. 86
250 38 25 23 0. 81
-o 22

0.35 9.0 r 23 16 1 .33 .t+ 1 .14

16.0 1 29 ZJ 48 1 49 .29 1.15
25 0 0.98 18 14 1 .06 .14 1 .84

0.50 9 0 1.18 .)L 25 1 26 .39 1 .40

16.0 110 1l 11 1 04 .05 0.93
25 .0 1 .09 23 12 1 28 .35 |.3+
*P : probability of recombination, V, : environmental variance. See text for definition of B: and Br.
**Division by zero.
 November 1959] pEsEri AND BAKER-TANDEM AND rNDEX sELEcrroN 779

disturbing influences as skewness in the selected population, epistasis and

genotype-environment interaction. However, in computer simulation of a
genetic system we are able to compare simulated gains rather than expected
genetic gains for the particular model under consideration.
The genetic gains computed in this study are presenred in Table 4. In
tandem selection, the trait selected in the first qeneration r,vas improved most
when the negative genetic correlation between the tr,vo traits was large. Wirh
extremely Iarge negative genetic correlation (r,u :
-0.95), randem selection
resulted in a einetiiloss fo-r the second trait. This asain shows that in tandem
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selection onJ should select for rhe most imoortarit trait first. With zero
glnetic correlation between the trvo traits, approximately the same amount
of improvement was made in both traits by tandem selecrion.
With index selection, a genetic loss for the least valuable trait occurred
when the economic values wire unecual. When the economic values of the
t\'vo traits were equal, there rvas verv little response in either trait when the
negative correlation was high.
Genetic worth of the selected population rvas calculated for both methods
of selection by weighting the geneiic gain for each trait according to its
economic value. llhe differences bctr.vecn the changes in genetic worrh for
index and tandem selection was then erpressed as a pioportion of the absolute
For personal use only.

value of rhe change in genetic \\,orrh for tandem seleciion. This proportion,
S, was used as --.mu?" of the superiority of index selection over'tandem
" values of S indicate thit index selcction is suoerior and
selection. Positive
negative values of S indicate that tandem selection is superior. S : 0 would
indicate that there was no difference in thc eificiencies of the trvo methods of
selection. Estimates of S are presented in Tabie 5. Because all values of S
are greatef than zero, we conciude that index selection is superior to tandem
selection under all combinations of parameters considered in this studv. The

Table 4. Realized genetic gains under tandem and index selection*

lndex selection
Tandem
selection a,: |,a": I a":1,6":l a":1+,ar:1
AC AG, LG" A,G, AG, AG. AUr

0.05 9 .0 +.06 - s 47 706 0.58 0 65 7.27

160 3.40 -2.70 -7.C0
-6 93 / .1.) -0.28 1.24 7.r5 -6.95
-7.03
25.0 3t7 -25+ -7.09 7.00 -051 1.21 6.59 -672
o.20 9.0 .87 0. 59 -3 36 529 1.78 1 63 + 14 -1 56
16.0 49 0.61 -2 87 1.24 1 .09 1 81 4.08 -2 15
25.O 25 0 ,32 -3.+7 435 1.04 1.39 3.57 -1.56
0.35 9.0 .81 161 0 .12 3.62 2 .24 1 .9+ 3 .63 0.54
160 38 12+ 087 295 1 92 1 .62 3 .07 0.90
25 .0 .04 1++ 015 29A 1 .56 1 .78 2 64 0.74
0.50 9.0 252 2.03 1 .65 3.60 3.06 2.55 3.24 1.85
16.0 2 .16 r 71 |.49 3 19 2.67 2 41 2 90 1.63
25.0 1 .88 1 .65 1 .39 2.74 2.40 1.90 271 1.33
+P : probability of recombination and V" : environmental varrance.
 780 CANADIAN JOURNAL OF PLANT SCIENCE [VoI. 49

superiority of index selection over tandem selection depends in part on the

relative economic values assigned to the traits, being proportionately greater.
when the least valuable trait is selected first in tandem selection. Index selec-
tion is much more efficient when the absolute value of the negative correlation
is greater. There is also an overall trend for index seleciion to be more
efficient as the heritability of the traits decreases.
These results indicate that simultaneous improvement of two or more
negatively correlated traits can be more efficiently obtained by applying index
selection than by applying tandem selection. While index selection is more
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efficient than tandem selection. it mav also recuire more effort on the part of
the plant breeder. He must grow material in a replicated design which permits
the necessary estimates of phenotypic and genotypic parameters. Further-
more, he must assign relative economic values to those traits he is trying to
improve. Since in?er selection was found to be from ll to 47 l'L more
efficient than tandem selection (Table 5), plant breeders should be encouraged
to initiate studies which rvill result in the realistic assignment of economic
values to the important quantitative traits.

Table 5. Average superiority, S, of index selection over tandem selection*

For personal use only.

Superiority (S)
V" o, : !,a" : I a, : 7,0, : I a,: l!,4., : I Mean
1 ,lA
0. 05 9.0 3.49 1.23 0. 50
16.0 4.71 0.36 0. 54 1.87
25.0 +.62 0. 11 0.67 1.80

Mean +.27 0. s7 0. 57 1.80

0.20 9.0 t.36 0.38 0.37 0. 70

16.0 1 .08 038 0.40 0.62
25.0 r.78 0.55 0.73 1.02

Mean 1.41 0.++ 0. 50 0. 78

0.35 9.0 0.47 0.23 0.39 0.36

16.0 0.76 0.18 0.67 0.54
25 .0 0.52 0.35 0. s7 0.48
Mean 0.58 0.25 0. 54 0.46
0.50 9.0 0.34 0.23 0. 16 0. 25
16.0 0.41 0.31 0.21 0.31
25.0 0.32 0.21 0.20 0.25
Mean 036 0.25 0. 19 0.27
*See text for defrnition of S; P : probability of recombination; tr/e : environmental variance.

ACKNOWLEDGEMENTS
This work v/as caffied out while one of us (J.P.) held a Postdoctorate Fellow-
ship from the National Research Council of Canada, which we gratefully
acknowledge. We also thank NRC for making arrangements for comPuter
facilities.
 November 19691 PESEK AND BAKER-TANDEM AND INDEX SELECTION 78L

REFERENCES
1. B.r,rnn, R. J. 1956. Predicted variance of response ro selection. Ph.D. Thesis, Uni-
versity of Minnesota, St. Paul, Alinn.
2. Bexr,n, R. J., BnNlnr.o,r.v, V. AtI. and KeurueNN, M. L. 1968. Inheritance of and inter-
relationships among yield and several quality traits in common wheat. Crop Sci. B,
725-728.
3. Bocren, W.A. 1968. A'Ianual of procedures in quantitative genetics. Washington State
Universiry Piess, Pullman, Washington.
4. BnIr,r, C. A. 1966. A modified pedigree method of selection in soybeans. Crop Sci.
6, 220.
5. Gour-nax, C. H. 1939. Problems in plant selection. pp. 132-133. 1z Proceedings of
the Seventh International Genetics Congress, Cambridge University Press.
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6. Hazar-, L. N. 1943. The genetic basis of consrructing selection indices. Genetics

23,476-490.
7. Hezw, L. N. and Lusn, J. L. 1942. The efficiency of three methods of selection. J.
Hered. Il, 39)-402.
8. Surrrr, H. F" 1936. A discriminant function for plant selection. Ann. Eugen. 7,
2+O-250,
'WETLER,
9. YouNc, S. S. Y. and FI. 1960. Selection for two correlated traits by inde-
pendent culling levels. J. Genet. 57,329-338.
For personal use only.