AN APPLICATION OF INDEX SELECTION TO THE IMPROVEMENT OF SELF.

POLLINATED SPECIES
J. PESEK' and R. J. BAKER Research Statiort, Canada Deportment of Agriculture, Winnipeg 19, Canada. Contribution No. 399, received November 7, 1969'

ABSTRACT
Results of a genetic study of four quantitative desired genetic gains rather than relative eco' characters in-a cross of two cultivirs of nomic w;ights o=f traits, is explained in detail ticunt aestivum L. em Thell. indicated and applied to selection for maturity, height heritability of yield was lower than the herita- and yield from hybrid population of bilities of maturity and height and that inter- wheat. The methods and problems of using actions between genotypic- effects and year index selection in self-polfinated species are environmental effects were nonsignificant. discussed. modified seleclion index method. based uoon

Trithat

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The

INTR.ODUCTION

Smith (16) first suggested the use of the concept of a "discriminant function" as a logical and systematic manner of selecting plant lines to improve several quantitative characters simultaneously. He applied this technique to wheat. Since that time, this method has become known as index selection and has been used extensively in the improvement of various animal species. The application of index selection to the improvement of self-pollinated plant species, for which it was

first suggested, has been rare. The object of index selection is to maximize the average "genetic worth" of a population. Genetic worth is the sum of products of the genotypic values of the measured characters and their respective "economic weights". Thus, genetic worth reflects the overall value of a particular line or individual. Economic weights express the relative importance of one trait to another in making up the overall value of a line. Since genotypic values cannot be observed or measured directly, it is necessary to develop a linear function of the observable phenotypic values which will best discriminate among genotypes of different genetic worth. Flence, the original term for index selection was "a discriminant function for plant selection". Once a selection index has been constructed, it is possible to predict the difference between the mean of a proportion selected by truncation and the mean of the original population for each character. This "expected response" may be used to evaluate beforehand the consequences of several alternative selection procedures. "Observed response" implies that the selection has already been done. Selection indexes may be used as a basis for the simultaneous improvement of more than one character by selection, or for enhancing the effectiveness of selection for one character by incorporating information on one or more secondary characters. The latter use has been most common in plant breeding (I,'7,9, II, 13,15) because there is no need to assign economic values to all traits in this application. To improve the primary trait, such as yield, as much as possible, the economic values of the secondary traits are all set al zeto and that of the primary trait as unity. This method has generally permitted good results in selectionforyield (7, 11). Exceptions were noted by Panse and Khangonker (13) in cotton and Abraham ( 1) in rice. For improvement of several characters simultaneously, however, it is necessary to assign economic weights to the characters
lPostdoctorate Fellow (permanent address: Research Institute Hrusovany, Czechoslovakia).
Can. J. Platrt Sci, 50: 267-276 (May 1970)

for Basic Agrotechnique, Brno-

267

268

CANADIAN JOURNAL OF PLANT SCIENCE

and this is often very elaborate and in some cases almost impossible. For instance, the economic value of a character may depend upon its reaching a certain threshold

value. Protein content in wheat must be above a certain minimum level before a line is acceptable. Above that level, protein content becomes relatively less important than yield. Since Smith's (16) paper, two other approaches to the discriminant problem in selection have been proposed but both incorporate the economic weights concept. Kempthorne and Nordskog (10) developed the theory of restricted selection indexes and applied it to selection in poultry. Tallis (17) derived a further modification in the terms of an "optimum genotype". These
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modifications allow for the maintenance of one or more characters at the level of the population mean or at some pre-chosen optimum level. Difficulties in assignment of relative economic weights led us to the problem of ranking measures of several quantitative attributes when nothing is to be assumed about economic weights. The concept of "desired genetic gains" (14) which removes all needs for assigning relative economic weights provides one answer. In this study we report the results of a quantitative genetic study in a cross of two cultivars of hard red spring wheat, Triticum aestivum L. em Thell., and apply the modified selection index method to selection in this cross.

MATERIALS AND METHODS The parental cultivars for this study were 'Garnet' and 'Kenya 360.H'. With respect to the four characters measured in this study, Gamet headed earlier, ripened earlier, had shorter straw, and was lower yielding than Kenya 360.H. Lines were advanced from single seeds in the F, generation up to single seeds in the F" generation by single seed descent, as suggested by Goulden (6). Seed from random F. plants was bulked and grown as plots to produce the F, generation. Bulk seed from 48 F, plots and the two parental cultivars was sown in replicated yield trials (two replicates) at Brandon and Regina in 1961 . Seed harvested from the yield plots was again sown at these two places in 1968. Days to heading (DH)' days to ripening (DR), height in centimeters (HT) and yield in decagrams per plot (Y) were recorded for all plots in all trials. A combined analysis of variance and covariance was performed on the data. Mean squares for each character and mean cross products for each pair of characters were calculated for sources of variation relating to genotypes, years, places, the interactions, and error. Components of variance and covariance were then estimated as outlined by Comstock and Moll (4). Phenotypic and genotypic correlations were estimated by the methods outlined by Baker et al. (3). Pesek and Baker's (14) modification of the theory of index selection involves calculation of index coefficients accordins to the following matrix formulation:

b:G-th.
where

?
z

b is the vector of index coefficients, G-' is the inverse of the genotypic variance-covariance matrix, /z is the vector of desired genetic gains and p/zis lhe reciprocal of the selection differential in standard units. Since p/z is constant for any experiment, an equivalent solution can be obtained from:
b:G-lh.

PESEK AND BAKER-INDEX SELECTION

269

The above expression represents the following set of simultaneous equations for the specific example of four characters: b*n I bzgtz I ba*z * bqu -- ht hgrz I bzgzz -f 6:9:: -f bqzt : hz bqn I hzgzz * b;Brr I bqzq : ht b$rt I bz7z+ I b*tr * b4++ : h+ where the bo's are the index coemcients to be solved for, g,,, is the genotypic covariance between characters i and j, and ho is the desired genetic gain for the i'n
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character.

After estimating the genetic components of covariance, the b,'s were evaluated for two different sets of desired genetic gains. The resulting b,'s were then combined with the character means of each of the 48 lines to sive a selection index for each line of the form: Ie : brXLn I brXre f- btXre I brXt*
values of the four traits for that particular line. The five lines having the highest index values were chosen as the selected population, and their means were compared with those of the original population and with the means of the parental cultivars.
RESULTS Estimates of components of variance and heritabilities of each of the four characters are given in Table 1. Heritability has been expressed on a line mean basis, the mean being taken over two replicates at each of two places in each of two years. An approximate test of significance for a difference from zero is given by comparing an estimate to twice its standard deviation. By this criterion, it is apparent

/" being the index value of the k'" line and X,,,,

X",,, X",, a;fld X,* being the mean

that the genotype x year interaction component of variance is unimportant for all characters. For yield, the genotype x place interaction component is nonsignificant and the genotype component of variance would be judged as just approaching significance. The heritability estimates indicate that relative yields of lines are
more dependent on environmental influence than the other three traits.

The genotypic and phenotypic correlations among the four characters are listed in -Iable 2. All phenotypic correlations are positive and significant at the 77o leveI of probability. The genotypic correlations between yield and the other characters are larger than the corresponding phenotypic correlations.
Table

1.

Estimates of variance comporlents and heritabilitl. (n'rean basis)

Componentx I)a1s to head

62,
62,^,

Dill's to ripen
13.01+3.07
0.

Height (cm) 5+.91+ 5.03

3

Yield (dkg)
7 .94+1 3.26+3

7 2

0.18+0.15 0.93+0.09
8+.2

.03+t.67
18

6'ctr
n2 o2"

0.41+0

.01+0.,19

28+0.56 2.+1+0.92 1 .35+0. 78 4.78+0.48

85.
1

3 3

9.93+ 1 .03
91

81+3 .41 .03+ 1 .68

.

.03+3.

.O2

29

2.10+3.6r
12 21

-79

.74++
48.
1

.7t+2.18

-80

h'(%)

.0

1t2s : component of variance due to genotypic difierences among lines, 6t I : component of variance due to the interaction of genotypic and year effects, dtr, : component of variance due t; the inteiacLion of genotvpic and place efiects, dzrrJ : component of variance due to the second order interaction of genotypes, places and years, t2e : the variance due to plot-to-plot environmental variations.

270

CANADIAN JOURNAL OF PLANT SCIENCE

Table 2. Genot-vpic and phenotypic correlations among four traitsJ

DH
Da1's to head (DH) Da.vs to ripen (DR)

DR
0.91+*

HTY
0..48*x

0.26xx

Height (HT) Yield (Y)

0.98 0.57
0.6+

0.52*x 0.7+

0.32**
0. 55**

0.57
0. 74

t*Significant at th.e 17o lcvel ni probrbilitt. fPhenotypic correlarjons abor e the djagonal. gPnot] l ic belo\'.

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The flrst example of application of the modified method (14) of index selection was based on desired gains of: (1) a decrease of 2.75 days (5% ) in days to heading, (2) a decrease of 4.83 d,ays (5%) in days to ripen, (3) a decrease of 1.63 cJntimeterc (2Vo ) in height, and (4) an increase of 3.25 decagrams (about 3.2 bu/acre, i.e., to%) in yield. By using these values and the estimates of genetic covariances and variances in the four linear equations, the following index ioefficients were obtained: b,: 15.70, b,: -14.14, b": -1.97, and bn: 6.84. Thus, the index value of a line is obtained by multiplying its average days to head by 15.70, subtracting 14.14 times the number of days to ripen, subtracting 1.97 times the height in centimetels, and adding 6.84 times the yield psr plot i1 dega; grams. For the index value of line 1, which averaged 52.8 days to head, 95.1 days to ripen,82.6 centimetels in height and 39.92 decagrams per plot in yield, one obtains:
1s.70(52.s)

Desired genetic gains for the second example allowed for some delay in maturity in hope of a greater increase in yield. Thus, the desired genetic gains were a 5Vo inuease in days to heading (2.1 5), a 5Vo increase in days to ripel. (4.83), a IVo decrease in height (-0.81) and a 2o7o increase in yield (6.50). The index coefficients werei br : 5.20, b": -4.42, b": -I.37, and b' = 3'58' The index value of the first line was:

14.14(95.1)

1.97(82.6)

6 81(39.92)

-405

+ 3.58(39.92) : -116 0 The mean values for each character and the two index values for each line
5,20(s2.8)

1.42(95.1)

r.37(82.6)

are

presented in Table 3. The index values have all been made positive by subtracting their minimum value. The five lines with the highest index value were then selected, and the means of the various characte$ were compared with what would have been expected on

summarized in Table 4. The ranking of selected lines indicates that use of a selection index often leads to the selection of lines which do not excal in any

a theoretical basis and with the original parents of the cross. These data are

one character but tend to be more intermediate. Yield, which has the most weight in the desired gains, is represented by lines which tend to rank high. Expected gains, calculated by prediction equations (5, 8), are distributed in proportion to the desired gains. However, most of the realized gains depart from those expected both in sign and magnitude.
DISCUSSION

Application of index selection to self-pollinated species may meet with the unique situation of selection among homozygous lines where the selected line or lines will

PESEK AND BAKER-INDEX SELECTION

271

-fable 3. 'fable of means and index values of lines and parents

Da.vs

Line No

to head
52

Da.vs to

ripen 95.
1

Height (cm, 82.6

Yield

(dkg)
39 .92

Index A
126.+ 133.8 t01 .7 80.2 62.3

Index B
57

i
2 3

.8

4
5

55. 8 51 .2 54. 8
.)o . .')

100. t) 97 .4

86.4
85 .3
tJ.l

41 .68

.0

95.2

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98.4
97 .6
9+

85.

6
7

8 9 10
11 72 l.)

.)+.r 54.4
.)/.6
52

-r

80.8
73.7
85. 1 83 .3

34.98 35.08 33.51

.)r.lt

60.3
40.9 40.2

9'1.1 101 .9

29.20 31 .88
29.41

96.6
105. 1 105. 7
35

26.3
38.2
41 .9
37

.8

94.0
95 .,1 92 .5

80.8
.0 7+.2 82.8 82.8
81
6.)

3t.61
29.75
29 .89

.4
8

.+ 1+.2

88.

54. 1 52 .6

76.0
107 .9 85 .0

34.6
28 .1

65.2
58

7+
1.)

.'i

108.8

33.79

.50. (,

98.8 98.0
96.1.

..i

30.88 32.95
33 .05

16 1n

53.2 53 .4
52 .0

18

t9
20
21 22

53.4 52.0
JI. I

93.9 92.8 92.6

91 .4 92 .1

80.0 76.4 78.5

tr.l

30.32 32.81 26.06

33 .65

99.7 82.5 76.6 99.1 105.7

97 .9

43.0 38.6
37 .0

32.0
33 .5

39 .3

43.t

23

nt

52.+ 5,1 .8

95.0
95 .4

77.7 76.7 80.3 88.6

82 .3

132.8

36.5
53 .3

33.35
33 .06

t2+.4
79 .1 140 .4

25 26 27 28 29

55.8 53.4
.)J. 53. 53.
.).).
Z

96.9
93 .6

39.16 36.99
.)l . Jr

51.0 33.9
5,/. t .)+.+ 43.9

128.5

77.2
81 .5

110.0

J.). I

100.5 99 .9
91 .2

53 .5
1 1
I

96.4
95 .0

80.8
81

26.79 30.76
.39 .05 .48 .92 .95 30. 75
33 32 33 29 39

0.0

i09.0
58.4

41 .5

zr.t

0.0

30
JI

32
33

58.

95.8 t02.2
95.9

82.8 .0 87.9

43.6

91.5 95.2
65.9

21 .2

5+.2

93.0
86 .9 81 .0

38.0 22.7
41 .0
21 .9

34 35 36

56.4 54.6
.).) . o

96.0
98. 95.
8

32.3+
37 .96

38
39 40
41
A'
+.1

62.6 s3.5
51 .1

98.6
105.6
8

92.7
86.6
83 .6 76 .5

12t.4
63.3

30.79 30. 19

68.9 50.4

46.6

29.0
17.8 25.3 15.2 26.1
JJ. J
41 .9 49 .7

44 4J 46 48

54.5 58.0 56.4 53.0 52.9 54.5

52.9

93.2 95.2
99 .2
97 .2

70.9
80.
8

25.76 26.95 29.06

.tl -.).)

42.7
60. 9 90 .4

93.2

7+.9 75.2
81 .3

32.35
29 .52

103.2 116.4

94.4
93.
91
1

99.8
125.8 153 .7
774.2 105 .6

39.2
.)+. /

80.8
81 .5 89 .4

37.79
35 .36

59.4
57

i01.1

.9

60.7

31.00
36.
11

4t.7
l.l
.

.9

i00.8
96.6
91

85.

28.90 32.51
27 .9+ 45. 58

45.5

+r .7
.t

Mean Garnet

.).). t,

81 .3 81 87

Kenya 360.H

51.9
61 .1

.6

107.4

.8 .6

81 .3 111 .5

25.9

59.0

2'72

CANADIAN JOURNAL OF PLANT SCIENCE

=
G

; 111A OEN

'o

reior cj c;*\O4DD<,

c4N<.

- N4+r

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/.
a

= .j

z
A

\O4O Ar\O\Cc}, 4O.ee\O so+eo

C'O\ =!-C' rC<i a

;6FoF
4=\Yc;t

c+o' o@9
Co e

+&c a.q
r4N N<r

'o
c

tr

o
= c

I I rc-c . :-+=J]:e

*'**.c- --<+9^l-

+
N

.U

J
a c
O

F =

:
t

.i
tr

4 : =6CrN :jN

@ +,r e

-CoO

JCt

a-r o+o

*t

-6d656rco

t t =Fr

&*

a o
d

d a/

Ilrcee*N!i1 ;":
ll

E c
N 4
ll

@ooo,r +Nee

NO Y

*,

/,
F
G

..- ,; ;
'o
o =
\OOle+n

-.;

;c;i

ONC]

;:etP:
--=-e-t

C^)C

li

oco ,;: ::-

.O+C

, =t9 =_

t
G

:acr,i N+N

>al

(a

4=3C
=r

+- a;

q-?
4\On

z
4CN .+N O+ NN

+'^i-+ _*i,b 'N-ts+ .[1* Eyi, 7;= *?f E;.i i.",n. oYr ;'3i

.=.=.

.2.=- Eg

E.

E
tr

PESEK AND BAKER-INDEX

SELECTION

273

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selection and since evaluation of the final product is already complete. The effectiveness of index selection will be useful in programs of recurrent selection. cycles of recurrent selection can be obtained by ielecting a portion of the population and recombining the genetic material of tie selected portion to form u tre* population which is to be the foundation of a new cycle o1 selection. At first look, the large discrepancies between expected and realized gains raise doubt as to the validity of the general theory of index selection. Howiver, estimates of both expected and realized, gains ire subject to error, the error of the difference thus being compounded. A minimum estimate of the standard deviation of the difference between the expected and. realized gains can be obtained by taking the standard deviation of the realized gains as (V/s + v/4g),', where it is the estimate of the phenotypic variance of line means. Having done this, the differences between realized, and expected gains all fall within thJ range of twice their standard deviations. The discrepancies between realized, and exp"ected gains . are thus well within the limits of random environmental variability und th"rJfor" should not discredit index selection methods. The sizes of the expected gains deserve comment, for they are small relative to the gains that are desired of this particular wheat cross. Aimitting some level of error.inthese expectations, it is s?iil clear that several cycles of selection would be required to obtain the desired results. The realized gain estimated for yield was greater than the expected gain. The large error associated with this measurement, however, casts doubt on comparisons oirealized with expected gains for yield. . As will always be !rye in the application of this method (14), the exp-ected gains are a constant multiple of the desired gains. Thus, one'would thut the desired gains should be obtained if ind& selection is continued foi enoush "*p""t cycles. This will not be true in actual practice if pleiotropic effecrs impose liriitations on the gain that can be realized.. If this should be the case, one would observe changes in genetic parameters during cycles of selection. The nature of these changes would indicate whether the deiired gains could be achieved. - .It tl interesting to ask what economic weights would have to be assigned to obtain the same selection results as are obtainJd by the modifled method, which bypasses this step. By using the relation pb = da of conventional theory and inserting the index coefficients of the first example (b) and the phenotypic (p) and
genotypic

be the final product of selection. We see no value for index selection in this instance, since the goals of the program will depend upon what is available for

relationships among four traits. The equivalent economic values for the second example, in spite of changes in selection aims, are similar to those of the first except for a general reduction in magnitude._ Il may also be seen that the index coefficients oflhe second example are similar both in sign and magnitude to those of the first. If we take into accor]nt the large positive correlations among all four characters and express our two goals as attempts to decrease three (or one) characters and increase the other on"- (ot

fhat would have resulted in identical expectations. For the first exthe equivalent economic values would have been 2.0 for days to head,, ?-pl-"' -2.2 for days to ripen, -0.3 for height in centimeters and 1.0 for yleld per plot in decagrams. These values appear to be nonsense from un e.orro-i" point oi view, but do serve to indicate our inability to consider as a whole the genetic inter-

values

(a)

(G)

variance-covariance matrixes,

it is possible to esiimate the economic

274

CANADIAN JOURNAL OF PLANT SCIENCE

is due to tho three), we can see that the insensitivity of the index coefficients demonstrated manifold interrelationrhip, u-ong the characters. This is further negative by noting that days to ilead unJ duy, to ripen are given .p,o,ttiiu" and on yield is not detrimental weights, respectively, so that their combined influence to the goals of the Program. high yielding Such considerations raise doubt as to whether an early maturing, from this cross. one would need several cycles of cultivar could be a.u"top"a selection selection to reach tn" !6uf. Palmer (12) has indicated that recurrent crops as far as any one holds great promise foi ttre breeders of self-pollinated improvement. of character is concerned. This is no less true for the simultaneous In our case, his procedure would involve selecting several quantitative characters. start a new the two best lines with regard to index values and intercrossing to cycle. with the method used in the present study, each.cycle.would require at time consuming least five years. Hence, the overall program would be prohibitively would have to be devised. and methods of shortening the cycle time A modification of thi"s type of recurrent selection can be suggested in the light crop of Baker's results (2), which indicate that intermating in a self-pollinated best just the such as wheat or barley is technically feasible. Rather than selecting a foundatwo lines for intercrossing, several lines could be intermated to produce This procedur-e would also help to tion population for u n"# cycle of selection. (2)' reduce the random genetic'deviations associated with small sample sizes gen-etic with- divergent One would be advised to start a recurrent selection program breeding and calculate selection indexes based on long term goals of the material could be extracted from the program. Lines excelling in one or--several traits Selecfi1l immediate needs during any iroltu- and evaluated foi ability towould, however, be on the basiscycle. index of the cycle tion of lines to start the next respect to numerous system, which tends to retain individuals whose performance in to exclude those with u'Auptiu" traits is close to the population means, and tends in Table 4. extreme expression of a single triit, ut can be seen from rank values the main requisites for using index selectiort From our examples it is clear that of the goals are quantitative data, estimates of genetic parameteis and a statement progru-. Ttre first requirerient m"itts that visual ratings such as "good" or of th" ,,bad,,'muit be replaced ty ratings on a suitable scale. If the data for a given trait are discontinuour (..g.ratin!'s of 1 to 5) or if the trait is controlled by few genes, one should anticipite sigriificant deviations from theoretical expectations' Methods of estimating g"tt.1i" parameters necessaly for construction of selection indexes fall into four" Jategories analogous to those employed in estimating of genetic variances for one trait] They may be derived -from (l) regression combinations of traits' (il) progeny means on parental values for all possible differences between genetically homogeneous and heterogeneous populations,.(lli) data from F, and bJckcross populations, and (iv) combined variance-covariance analysis. In the modified p"iig."" method of selection, as used here, it is natural to replicate essentially homozygous genotypes over several environments which slows down cycle time. Nonsignincant estimates of genotype T yeq variance in this study uod io another (3) iniicate that the cycle time could be reduced by one year by testing in one instead of two years' Intergeneration comparisons, as between F, and Fn, constitute a form of replication *oittty of investigation relative to early generation selection' These com-

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PESEK AND BAKER-INDEX SELECTION

275

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parisons afford estimates of genetic variances and covariances in much the same way as animal breeders obtain information necessary for the application of index selection. Such estimates could be obtained in early generations and thus shorten the time required for each cycle of a recurrent selection program. Once estimates of genetic parameters are available, it is merely routine to incorporate the desired gains of the selection program into a selection index that constitutes an objective basis for the simultaneous imorovement of ouantitative characters. It should be noted that bur method can also be used for simultaneous index selection of a primary trait (say yield) along with selection of a set of secondary traits (components of yield). If no information is available for the suitable assignment of desired gains in the secondary traits, the desired responses for each trait can be simply derived from the formula:
ftj : rij

!*_

li'r,

where subscripts

i, j

refer

to primary and

desired gains, g's the genetic variances between traits i and j.

secondary traits respectively, h's ate of the traits and r is the genetic correlation

Selection for quantitative traits is complicated by qualitative characters such as disease resistance. For example, can disease reaction, which is simply inherited, be incorporated into a selection index? Of course such factors cannot be incorporated, but it is possible to apply the methods of index selection to those lines that have been proven to be disease resistant. There is a distinct possibility, however, that selection for disease resistance will have an adverse effect on subsequent improvement of quantitative traits. For example, in cases of repulsion linkages between genes for resistance and genes for yield, which might be expected in a cross between a high yielding cultivar and a disease resistant cultivar, selection for disease resistance would likely reduce the mean yield. Hence, greater improvement would be necessary to attain the original goals. Research on interrelationships of qualitative and quantitative characters is needed. If one were to adopt a long term recurrent selection program for quantitative traits, it might be advantageous to select first for quantitative traits under disease-free conditions and subsequently to introduce those genes necessary for survival by backcrossing. The modified index theory introduced by Pesek and Baker (14) and presented here in terms of two worked examples should prove useful to those workers involved directly in the improvement of economic species. The method combines the goals of the breeder with the genetic restrictions of the population with which he is working, into an objective rule for selection. The necessary calculations are difficult if more than four or flve characters are involved, but no more so than the method of Smith (16) and much less so than the methods of Tallis (17) and Kempthorne and Nordskog (10). With access to electronic computers, the matrix manipulations are routine.

This new modification, besides eliminating the need for assignment of economic weights, can replace the methods of Tallis and of Kempthorne and Nordskog. To replace the restricted index method of Kempthorne and Nordskog (10), one specifies zero desired genetic gain for the traits to be restricted. By specifying a desired genetic gain which will result in the optimum being approached, the optimum genotype index method (17) can be approximated. If a breeder should

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want to realize certain restrictions in the first cycle of selection, this method would be inferior in that the restrictions are not realized at the expense of the improvement in other characters. This criticism can be overcome adequately by calculating several sets of index coefficients corresponding to difierent sets of desired gainJ, and comparing the gains to be expected by the use of each index'
ACKNOWLEDGEI\{ENTS We gratefully acknowledge a Postdoctorate Fellowship from the National Research Council of Canada held 6y the senior author during the course of this study' We are indebted to Dr. A. e. Campbell of this station for providing data and for comments on the manuscfipt. The data were collected by the cereal breeders of the Brandon and Regina Risearch Stations, whose assistance we appreciate'
REFERENCES Note on discriminant fr-rnction for varietal selection in rice. 1. ABRAHAM T. P. 1953. Vol. 18, p. 3 in Proc. 4th Meet. Working PartY on Rice Breeding. [nt. Rice Comrn..

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2. BAKER, R. J. 1968. Extent of intermating in selj-Pollinated species counteract the effect of genetic drift. Crop Sci' 8: 547-550'
725-728.

F.A.O., Rome.

necessary to

3.BAKER,R.J.,BENDELOW,V.M.andKAUFMANN,M'L'1968'Inheritanceand8: interrelationshrp u-ong li"fd and several quality traits in common wheat. Crop Sci'

4. 5. 6. 7.

COMSTOCK, R. h,. and MoLL, R. H. 1963. Genotype environmental interaction' p. 164-196 iz Statisticaf genetics and plant breeding, NAS-NRC 982' Washington' D'C' FINNEY, D. J. 1962. Genetic gains under three methods of selection. Genet. Res' Camb.3: 417-423. GOULDEN, C. H. 1939. Problems in plant selection. p. 132-133 in Proc' 7th Int'
Genet. Congr., Camb. University Press. HANSON, D. W. and JoHNSON, H. W. 195'7. Methods for calculating and evaluattng a general ielection index obtaineci by pooling information from two or more experlments'
Genetics 42: 421-432.

g. HAZEL, L. N. 1943. The genetic basis for 9.

constructing selection indexes. Genetics

28: 476-490. JOHNSON, H. W., ROBINSON, H. F. and CoMSTOCK, R. E. 1955. Genotypic and phenotypic correlation in soybeans and their implication in selection' Agrot 1.47:
417-483.

10. KEMPTHORNE, O. and NORDSKOG,

Biometrics 15: 10-19.

A. W. 1959. Restricted selection indexes.

from a selection index technique in

11. MANNING,
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cotton. Heredity l0: 303-322. 12. PALMER, T. P. 1953. Progressive improvement in self-fertilized crops. Heredity 7: 13. PANSE, V. G. and KHANGONKER, S. A. 1949. A discriminant function for of yield in cotton. Indian Cotton Grow. Rev. 3: 356-375.
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H. L. 1957. Yield

improvement

14. PESEK, J. and BAKER, R. J. 1969. Desired improvement in relation to selection indices. Can. J. Plant Sci. 49: 803-804. 15. ROBINSON. H. F., COMSTOCK, R' E. and HARVEY, P. H' 1951. Genotvpic and
phenotypic correlations in cotn and their implication to selection. Agron. J. 43: 283-287.

16. SMITH, H. F. 1936. A discriminant function for plant selection. Ann. Eng. 7:240-250. 17. TALLIS, G. M. 1962. A selection index for optimurn genotype. Biometricis 18:
1.20-122.

This article has been cited by: 1. M. Ndoumbe, D. Bieysse, C. Cilas. 2001. Multi-trait selection in a diallel crossing scheme of cocoa. Plant Breeding 120:4, 365-367. [CrossRef] 2. Fu-Hua Liu, S. M. Smith. 2000. Measurement and Selection of Parasitoid Quality for MassReared Trichogramma minutum Riley Used in Inundative Release. Biocontrol Science and Technology 10:1, 3-13. [CrossRef]

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