Efficiency of the Use of Doubled-Haploids in Recurrent Selection
for Combining Ability
A. Bouchez* and A. Gallais
ABSTRACT
In population improvement for combining ability, the use of selfed
progenies increases genetic advance per cycle, but it can unduly in-
crease the cycle length. Haplodiploidization (HD) can be very efficient
because it induces complete homozygosity in a short period. We com-
pared, from a theoretical approach, the potential of recurrent selection
with a tester using doubled haploids (SDHT), with selection with
testcrosses of S0, S1, or S2 plants, with or without the use of off-season
nurseries, for an annual plant like maize (Zea mays L.). With the
same selection intensity, and without off-season nurseries, SDHT with
a 4-yr cycle is the most efficient method. Efficiency increases with
lower heritability, with an advantage in comparison to the test of S0
plants (S0T) of 40 to 50% at low heritability (h2 ⬍ 0.15) and 12% at
high heritability (h2 ⫽ 0.8). Use of off-season nurseries reduces the
advantage of SDHT. With a 3-yr cycle, SDHT remains the best at
low heritability with a gain of 27% (for h2 ⫽ 0.1) in comparison to
S0T with a 2-yr cycle. When using constant effective size, the advantage
of SDHT is further reduced or suppressed at the benefit of S0T in 2
yr. The use of HD in recurrent selection for combining ability has its
biggest advantage when heritability is low. Consideration of variety
development will give more advantage to HD.
F or the improvement by recurrent selection of the
value of lines that can be derived from a population,
haplodiploidization (HD) appears to be a very efficient
process (Griffing, 1975; Gallais, 1989, 1990a, 1993b;
Goldringer et al., 1996). This is due to the fact that it
allows direct selection on line value, whereas by the use
of partially inbred progenies, selection can be biased by
heterozygosity. However, considering the improvement
of combining ability of a population, the usefulness of
HD may be questioned. Indeed, in the absence of epista-
sis, the combining ability of lines derived from an S0
plant can be predicted from the combining ability of
the S0 plant (Gallais, 1990b). Nevertheless, HD can be
efficient in increasing the variance among progenies
from testcrosses and thus the heritability. HD also has
the advantage of giving more uniform progenies. If the
cycle length is increased in comparison with combining
ability tests at the S0 level, the genetic advance per unit
of time will not necessarily be greater. As already shown
by Griffing (1975), the length of HD process is a critical
parameter to consider. Thus, the advantage of HD in
recurrent selection for combining ability with a tester is
not obvious. However, an advantage of HD in recurrent
selection is to reduce the time required for varietal de-
velopment by pedigree selection. Indeed, with the use
of HD, if the tester is used as a parent of the new
hybrids, recurrent selection is more than a method of
A. Bouchez, INRA-UPS-INA.PG, Station de Génétique Végétale,
Ferme du Moulon, 91190 Gif Sur Yvette , France; A. Gallais, INA.PG,
16 rue Claude Bernard, 75321 Paris Cedex 05, France, and INRA-
UPS-INA.PG, Station de Génétique Végétale, Ferme du Moulon,
91190 Gif Sur Yvette, France. Received 13 Jan. 1999. *Corresponding
author (bouchez@mons.inra.fr).
Published in Crop Sci. 40:23–29 (2000).
23
population improvement, it becomes a method of recur-
rent varietal development.
Griffing (1975) has already considered some aspects
of the use of HD in recurrent selection for improving
combining ability; however, he considered neither the
use of S1 or S2 progeny testcrosses, nor the possible use
of off-season nurseries. Strahwald and Geiger (1988)
have shown that with the use of off-season nurseries,
the advantage of HD, for the development of lines tends
to disappear. In a theoretical study on optimizing sugar
beet breeding plans based on selection among testcross
progenies, Borchardt and Geiger (1997) showed that
testing S3 lines maximizes genetic gain compared with
S1 lines or doubled haploid lines.
In our paper, we consider only the use of recurrent
selection with a tester (which can be a population), with
or without the use of an off-season nursery. Recurrent
selection with HD is compared with recurrent selection
at the S0 plant level (i.e., without inbreeding) and with
selection at the S1 and S2 plant level (i.e., with one or
two generations of self-pollination before crossing to
a tester).
The following theoretical development assumes no
epistasis and no genotype ⫻ environment (G ⫻ E) inter-
action. Gallais (1991) has shown that epistasis is ex-
pected to have a low effect on relative efficiency of the
breeding methods for combining ability. Furthermore,
in a large range of realistic situations, the presence of
genotype ⫻ location interaction has only a small effect
on the ratio of the two phenotypic standard deviations
associated to any pair of methods when selection is on
average performance across locations. Interaction with
an uncontrolled factor, such as years, decreases the ac-
tual heritability. Thus, here such an effect is considered
through heritability. Methods are compared on the basis
of the genetic advance per unit of time, at the level of
the breeding population. The cost of the methods is not
considered. However, it is assumed that for a given year
of testing, the number of plots is fixed. Furthermore,
time is considered through the cycle length. The consid-
eration of the cost would need to assess the whole breed-
ing plan, including varietal development (Geiger and
Tomerius, 1997). For such a development, to be realistic,
it would be necessary to solve the problem of response
to recurrent selection on several cycles, and response
to pedigree selection, which are not well resolved. Our
study is valid for choosing, at a given cycle of the breed-
ing population, the recurrent selection scheme having
the greatest genetic advance per unit of time.
Abbreviations: HD, haplodiploidization; SDHT, single-doubled-hap-
loid descent recurrent selection for combining ability with a tester;
S0T, S1T, S2T, recurrent selection for combining ability with a tester
with respectively S0, S1, S2 plants.
24 CROP SCIENCE, VOL. 40, JANUARY–FEBRUARY 2000

MATERIALS AND METHODS
Description of the Methods
From a practical point of view, we will consider the case
of an annual crop such as maize, for which it is possible to
grow off-season nurseries. Basically, a cycle of recurrent selec-
tion has four stages: (i) production of plants to be tested, i.e.,
the derivation of selection units: S1, S2 plants or doubled-
haploid lines; (ii) crossing of these units to the tester, and
simultaneous selfing in order to maintain tested plants; (iii)
progeny testing; and (iv) intercrossing of selected units. Note
that with a plant like maize, with no prolificacy, it may be
difficult to self and cross simultaneously (S⫹TC) with the
tester used as male parent. The tester can be used as a female.
However, for early types, it may be impossible to produce
enough seeds for trials. In this situation, many breeders prefer
to produce S1, S2, or S3 progenies in order to select at the S0,
S1, or S2 plant level, respectively, and thereafter to cross these
progenies in isolation with the tester used as male parent. We
assume that field testing has to take place in a normal growing
season to get a representative agronomic evaluation.
The four selection stages of a cycle are scheduled differently
for each selection method, according to the use of off-season
nurseries and S⫹TC (Table 1). As a result, each method can
be applied with varying cycle lengths. With selection among
S0 plants (S0T) and use of S⫹TC, the length of the cycle will
be three generations, i.e., 3 yr with an annual plant without
access to an off-season nursery and only 2 yr with an off-
season intercrossing. It cannot be shorter, because we assumed
agronomic tests will occur in a normal growing season. If S1
progenies of the S0 plants are used for crossing to the tester,
the cycle length will be four generations long, and could last
from 2 to 4 yr depending on the use of off-season nurseries;
a realistic length is 3 yr. With selection among S1 plants (S1T),
or S2 plants (S2T), the cycle needs one or two more generations
for selfing. Depending on the use of off-season nurseries, and
on the kind of progenies used to evaluate combining ability,
the cycle length can vary from 2 to 5 yr for S1T and from 3
to 6 yr for S2T.
In the application of recurrent selection with HD, several
lines can be derived from an S0 plant. Increasing the number
of lines per S0 decreases the number of S0 plants studied, and,
on average, it is at the expense of the total variance and
effective population size (Gallais, 1989). The best approach
is to derive only one line per S0 plant, which is equivalent to
the classical single-seed descent method (Gallais, 1988). This
leads to the single-doubled-haploid descent recurrent selection
method (SDHT). Its cycle length depends mostly on the length
of the HD process. We only consider cases in which the whole
HD process is well controlled, and does not take more than
2 yr. With an annual plant without off-season generation, and
assuming that the whole HD process takes 2 yr, the complete
SDHT cycle will be 2 yr longer than S0T, or 1 yr longer than
S1T. If doubled-haploid lines can be developed from S0 plants
within 1 yr, then SDHT becomes comparable to S1T for cycle
length. With maximum use of off-season resources, all four
stages of S0T or S1T can be realized within 2 yr, while SDHT
as S2T will take at least one more year. Then, with DH lines, the
cycle length can vary from 3 to 5 yr, and cannot be shortened
because of field testing constraints.
Expression of Genetic Advance
Consider the case of a breeding population and its tester.
To simplify the notation, the combining ability of a S0 genotype
will be denoted AT , i.e., the additive effect of the S0 genotype
crossed to a specific tester (Gallais, 1989; 1990b). The testcross
value with a given tester is equivalent to a quantitative charac-
ter without dominance. Note that, with this notation, if the
tester is the population itself, AT is one-half A, the classical
additive effect for per se value. A general expression of the
expected genetic advance per cycle has been given by Gallais
(1991)
␪ covPTOT
⌬G ⫽ i , [1]
√var PT
where i is the selection intensity, ␪ the degree of selection
control on both sexes, cov PTOT is the parent-offspring covari-
ance for testcross value, and var PT the phenotypic variance

Table 1. Cycle description for four methods of recurrent selection for combining ability with a tester, in the case of an annual crop such
as maize. Plants are grown in summer (SUM) or winter in off-season nurseries (WIN).
Year 1 Year 2 Year 3 Year 4 Year 5 Year 6
Selection Cycle length
method† SUM WIN SUM WIN SUM WIN SUM WIN SUM WIN SUM
years
S0 T 4 S‡ – TC – T – IC
3 S⫹TC – T – IC
3 S TC T – IC
2 S⫹TC – T IC
2 S TC T IC
S1T 5 S – S – TC – T – IC
4 S – S⫹TC – T – IC
4 S S TC – T – IC
3 S S TC – T IC
2 S S⫹TC T IC
S2T 6 S – S – S – TC – T – IC
5 S – S – S⫹TC – T – IC
5 S – S S TC – T – IC
4 S – S – S⫹TC – T IC
4 S – S S TC – T IC
3 S S S⫹TC – T IC
3 S S S TC T IC
SDHT 5 ⇐ HD ⇒ – TC – T – IC
4 ⇐ HD ⇒ – TC – T IC
4 HD ⇒ TC – T – IC
3 HD ⇒ TC – T IC
3 ⇐ HD ⇒ TC T IC
† Methods of recurrent selection for combining ability with testcrosses of S0 plants: S0T, of S1 plants: S1T, of S2 plants: S2T, or of doubled haploids: SDHT.
‡ S: selfing, TC: crossing with a tester, T: agronomical test, IC: intercrossing, S⫹TC: selfing and testcrossing on the same plant, HD: complete haplodiploidiza-
tion process.
 BOUCHEZ & GALLAIS: DOUBLED-HAPLOIDS IN RECURRENT SELECTION FOR COMBINING ABILITY 25

among selection units. cov PTOT is equal to (1 ⫹ F)/2 ␴A2 T, ␴A2 T of replications and of the plot-heritability h2S01 at the S0 level:
being the additive variance for testcross value, F the coefficient
of inbreeding of selection units: F ⫽ 0 for S0 plants or S1 (1 ⫹ F ) hS01 ␴AT 1
⌬G ⫽ i . [5]
families, 1/2 for S1 plants or S2 families, 3/4 for S2 plants or S3 √(1 ⫹ F ) h2S01 ⫹ (1 ⫺ h2S01)/b t
families and 1 for doubled haploid lines. As in our situation
␪ ⫽ 2, it results It can be noted that when heritability is low, i.e., there
is a great environmental variance in comparison to genetic
(1 ⫹ F ) ␴2AT variance, phenotypic variances under different methods can
⌬G ⫽ i . [2] be considered as approximately equal. Then, differences in
√var PT genetic advance only depend on the genetic variance among
With an experimental design with n individuals per plot progenies multiplied by (1 ⫹ F ) and divided by cycle length.
and b replications, the phenotypic variance among selection At the other extreme, when heritability is high, differences in
units is genetic advance are due to the square root of genetic variance
among progenies multiplied by (1 ⫹ F )1/2, and divided by the
var PT ⫽ ␴2GBT ⫹ ␴2p/b ⫹ (␴2WGT ⫹ ␴2e)/bn, cycle length. These two extremes for the genetic advance in
where ␴2GBT is the genetic variance between progenies, ␴2p the standard units of the genetic variance (␴AT ⫽ 1) are easy to
environmental variance between plots, ␴2e the environmental compute (Table 2).
variance at the plant level within plot, and ␴2WGT the within
progeny genetic variance. It can be shown (see appendix) that Comparison of the Methods
␴2GBT ⫽ (1 ⫹ F ) ␴2AT The different methods will be compared according to their
genetic advance per unit of time with or without the use of
and off-season nurseries (Table 1). Methods can be compared at
the same selection intensity or on the basis of the same effec-
␴2WGT ⫽ (1 ⫺ F ) ␴2AT ⫹ ␴2WT, tive size of the population of intercrossed plants. At the same
␴2WT being the genetic variance due to heterogeneity of the selection intensity, their relative efficiencies depend only on
tester. When selection units are doubled-haploid lines, the three parameters: heritability, cycle length, and inbreeding
within progeny genetic variance depends only on the heteroge- coefficient. Such a comparison favors the schemes with the
neity of the tester, ␴2WT being then zero when the tester is a most inbred selection units. Ignoring the other parameters,
the selection of N completely homozygous individuals leads
homozygous line.
to an effective size that is half that with the selection of N
The general expression [1] of genetic advance per cycle can
non-inbred individuals. This is equivalent to having a greater
be transformed to get explicit heritabilities of the associated
selection intensity. To ensure comparable effective population
testing systems. Then
size for the various methods, if pS0 is the selection rate for
(1 ⫹ F ) ␴2AT S0T, pS1, pS2 and pDH for S1T, S2T and SDHT, respectively, it
⌬G ⫽ i√1 ⫹ F hF ␴AT with h2F ⫽ . [3] is necessary to set pS1 ⫽ 1.5 pS0, pS2 ⫽ 1.75 pS0 and pDH ⫽
var PT
2 pS0. More generally, with an inbreeding coefficient F among
Assuming that the number of plants per plot is sufficiently selection units, the selection rate pF applied to the method
great to neglect the contribution of the within-plot genetic must equal (1 ⫹ F ) pS0. For high values of F, the intensity
variance in comparison to the between-plot environmental may become unrealistic. For example, to compare SDHT and
variance and to the genetic variance among progenies, i.e., S0T with the same effective genetic size, if pS0 is 0.15 then
n ⬎ 30 (Gallais, 1993a), the phenotypic variance becomes pDH becomes 0.30. Considering the cost of SDHT, such a low
var PT ⫽ (1 ⫹ F ) ␴2AT ⫹ ␴2p/b. selection intensity is quite unrealistic. Thus, both types of
comparisons were developed, with pS0 ⫽ 0.05 and 0.10 for the
Then, the design heritability at the S0 level is studies with the same effective size.
Methods are compared first on the basis of the same number
␴2AT of replications, and second at their optimum number. To deter-
h2S0 ⫽ .
␴2AT ⫹ ␴2p/b mine such an optimum, we consider a fixed total number of
plots for a given year of testing. Then, for a given number of
Heritabilities for designs with the same number of replications intercrossed plants, the rate of selection with b replications is
but different F values can be expressed in terms of h2S0 pb ⫽ b p1. Increasing the number of replications increases the
design-heritability but decreases the selection intensity. The
(1 ⫹ F ) h2S0
h2F ⫽ . first effect is favorable, the second unfavorable. Then an opti-
(1 ⫹ F ) h2S0 ⫹ (1 ⫺ h2S0) mum number may result. At this optimum, comparisons have
been developed on the basis of the same number of inter-
When expressed per unit of Time t, genetic advance becomes crossed plants and on the basis of the same effective size.
⌬G ⫽ (i√1 ⫹ F hF ␴AT)/t
or RESULTS
(1 ⫹ F ) hS0 ␴AT 1 Selection with Same Selection Intensity
⌬G ⫽ i . [4]
√(1 ⫹ F ) h ⫹ (1 ⫺ h ) t
2
S0
2
S0
Without Off-Season Nurseries
The breeding methods can be easily compared on the basis Without off-season nurseries, considering the shortest
of the same number of replications. However, the optimum possible schemes with same selection intensity (Fig. 1),
number of replications can vary according to the level of SDHT in 4 yr is the best method whatever the heritabil-
inbreeding. To determine this optimum, Formula [4] was trans- ity. Its superiority is all the greater when the heritability
formed to express the genetic advance in terms of the number is low. With low S0-heritability (h2S0 ⫽ 0.10), the gain in
26 CROP SCIENCE, VOL. 40, JANUARY–FEBRUARY 2000

Fig. 2. Genetic advance per year (in genetic standard deviations) as a

Fig. 1. Genetic advance per year (in genetic standard deviations) as a function of S0 heritability under different recurrent selection methods,
function of S0 heritability under different recurrent selection methods, with use of off-season (with same selection rate pS0 ⫽ 0.05).
when no off-season is used (with same selection rate pS0 ⫽ 0.05).

length (Table 1). A 3-yr cycle has also been considered

efficiency in comparison to S0T in 3 yr is 43%, and tends
towards 50% as heritability approaches zero (Table 2).
With h2S0 ⫽ 0.40, the gain is still 27%, and for h2S0 ⫽ 0.80,
the gain is only 12%. Gains for SDHT in 5 yr are 25%
less. S2T in 5 yr is clearly the worst whatever the herita-
bility and at most equal to S0T at low heritability. Other
methods are relatively close to S2T when S0-heritability
is low. When S0-heritability increases, S0T in 3 yr and
S1T in 4 yr are significantly better than S2T and SDHT
in 5 yr.
With the Use of Off-Season Nurseries
To restrict the number of curves on Fig. 2, each selec-
tion method was studied at its shortest realistic cycle
for S1T, because the 2-yr cycle is quite complicated.
Whatever the heritability, S1T in 2 yr obviously becomes
more competitive than all other methods. It is followed
by SDHT in 3 yr, S0T in 2 yr and S1T and S2T in 3 yr.
In 4 yr, SDHT is the least efficient, especially for high
heritabilities. In 3 yr, it is still a competitive method for
medium to low heritability. The relative efficiencies with
same selection intensity (SI) can also be expressed in
reference to S0T in 2 yr (Table 3). These results show
the strong effect of the cycle length on the genetic ad-
vance per unit of time. The rank of the methods is
approximately the rank of the cycle length. This illus-
trates the strong effect of off-season nurseries. Notations
for selection method will be followed by the cycle length

Table 2. Relative efficiency of four recurrent selection methods for combining ability with a tester, for various heritability values (h2S0),
and for realistic cycle length using off-season generation or not (italics). Relative efficiency is expressed in reference to the genetic
advance for S0T in three years, for 3 situations: same Selection Intensity (SI), same Effective Size (ES) with selection rate ps0 ⫽ 0.05,
and same Effective Size (ES) with ps0 ⫽ 0.10.
Cycle length
2 years 3 years 4 years 5 years
Heritability Selection
h2S0 method SI ES (0.05) ES (0.10) SI ES (0.05) ES (0.10) SI ES (0.05) ES (0.10) SI ES (0.05) ES (0.10)
S0T 150 150 150 100 100 100 75 75 75 – –
0.0† S 1T 225 206 199 150 137 133 113 103 100 – – –
S2T –‡ – – 175 154 147 131 116 110 105 93 88
SDHT – – – 200 170 160 150 128 120 120 102 96
0.1 S1T 220 201 194 146 134 130 110 100 97 – – –
S2T – – – 169 149 142 127 112 106 101 89 85
SDHT – – – 191 162 152 143 122 114 114 97 91
0.4 S1T 205 188 182 137 125 121 103 94 91 – – –
S2T – – – 153 135 129 115 101 97 92 81 77
SDHT – – – 169 144 135 127 108 101 101 86 81
0.8 S1T 190 174 168 127 116 112 95 87 84 – – –
S2T – – – 138 122 116 104 91 87 83 73 70
SDHT – – – 149 127 119 112 95 89 89 76 71
1.0 S1T 184 168 163 122 112 108 92 84 81 – – –
S2T – – – 132 117 111 99 87 83 79 70 67
SDHT – – – 141 120 113 106 90 85 85 72 68
† Limit when heritability decreases towards zero.
‡ Unrealistic situation for maize.
 BOUCHEZ & GALLAIS: DOUBLED-HAPLOIDS IN RECURRENT SELECTION FOR COMBINING ABILITY 27

Table 3. Relative efficiency of five recurrent selection methods

for combining ability with a tester, for various heritability val-
ues (h2S0) with same selection intensity, and according to their
cycle length (in parentheses). Relative efficiency is expressed
in reference to the genetic advance for S0T in 2 yr.
Selection method
Heritability
h2S0 S1T(2) S1T(3) S2T(3) SDHT(3) SDHT(4)
0.1 146 97 112 127 95
0.4 136 91 102 112 84
0.8 126 84 92 100 75

in parentheses; S1T(2) means S1T in 2 yr. To get the

advantage of S1T(2) (46% for h2S0 ⫽ 0.10 and still 26%
for h2S0 ⫽ 0.80), it is necessary to conduct simultaneously,
in off-season, selfing and crossing to the tester. If we
look (in Table 3) at the four most comparable schemes,
with the maximum realistic use of off-season nurseries
for a species like maize, i.e., S0T(2), S1T(3), S2T(3), and
SDHT(3), then SDHT(3) gives a gain of 27% for h2S0 ⫽
0.10, and is equal to S0T(2) for h2S0 ⫽ 0.80. S2T(3) is only
about 10% less than SDHT(3) whatever the heritability.
SDHT in 4 yr is at most equal to S0T in 2 yr at low
heritability, and is 25% less at high heritability.
Selection with Same Effective Size
As expected, the relative efficiencies for SDHT, S2T,
and S1T are reduced relatively to selection at the same
selection intensity, and all the more as the tested plants
are more inbred (Table 2). With a 5% rate of selection
in S0T, the reduction in relative efficiency is about 10%
for S1T, 12% for S2T, and 15% for SDHT. With this
selection intensity, it is clear that the results are not
drastically changed. When a method has a strong advan-
tage on the basis of the same selection intensity for all
methods, it also has a great advantage on the basis of
the same effective size. Without off-season nurseries,
this increases the inferiority of S2T and SDHT in 5 yr.
S1T in 4 yr also becomes significantly inferior to S0T.
The advantage of SDHT in 4 yr is still 22% at low
heritability but disappears at high heritability. With the
use of off-season nurseries, the advantage of S1T in 2 yr
in comparison to S0T in 2 yr is 34% for h2S0 ⫽ 0.10 and
16% for h2S0 ⫽ 0.80. All other methods are inferior to
S0T in 2 yr, except SDHT in 3 yr at low heritabilities.
In particular, SDHT in 3 yr remains better than S0T in
3 yr, even at high heritabilities. With a lower selection
intensity (pS0⫽ 0.10), the reduction in relative efficiency
of SDHT in 3 or 4 yr is about 6% greater (Table 2).
Fig. 3. Domains of efficiency for SDHT compared with S0T, S1T and
S2T, according to relative cycle length (ratio of the SDHT length
to the length of the compared method), and to S0 plot-heritability
with (same selection intensity).
are nearly vertical. This indicates a low effect of herita-
bilities on these boundaries, mainly for the boundary
S2T-SDHT. The boundary S0T-SDHT clearly depends
on S0-heritability. This means that the relative cycle
length of SDHT can be longer at low heritabilities than
at high heritabilities. For example, with SDHT in 4 yr
and S0T in 2 yr, the ratio is 2, and then whatever the
heritabilities, S0T will be better than SDHT (approxi-
mately equal at very low heritabilities). With a shorter
SDHT cycle (3 yr) and S0T in 2 yr, with S0-heritability
less than 0.80, SDHT will be the best, whereas if h2S0 ⬎
0.80, it will be S0T.
Because S1T can be a competitive method, domains
of efficiency were calculated for this method (Fig. 4).
This method appears more efficient than SDHT only
when its cycle is approximately 20% shorter. The
boundary between S1T and S2T is nearly vertical at a
relative cycle length of 0.9. This means that for the same
cycle length, as expected, S2T is more efficient than S1T.
If the cycle length for S1T is shorter than that for S2T,

Effect of the Cycle Length

As long as the cycle for SDHT is not longer than for
other methods, SDHT is obviously the most efficient
whatever the heritability (Fig. 3). When its cycle is about
25% longer than that for S1T or about 10% in compari-
son to S2T, it becomes less efficient. For example, with
SDHT in 4 yr and S1T in 3 yr, the ratio of cycle length
is 1.33, and then S1T will be better than SDHT. If SDHT
Fig. 4. Domains of efficiency for S1T compared with S0T, S2T and
is in 5 yr and S2T in 4 yr, the cycle length ratio is 1.25, SDHT, according to relative cycle length (ratio of the S1T length
and then S2T will be better than SDHT. Note that the to the length of the compared method), and to S0 plot-heritability
separations of the domains S2T-SDHT and S1T-SDHT (same selection intensity). (a) S1T ⬎ S2T and S1T ⬍ SDHT.
28 CROP SCIENCE, VOL. 40, JANUARY–FEBRUARY 2000
Fig. 5. Genetic advance per cycle (in genetic standard deviations) as
a function of the number of replications of selection units, for
the same number of intercrossed plants, with the same selection
intensity (pS0 ⫽ 0.05 for design without replication) and with S0 plot-
heritability ⫽ 0.2.
for example, S1T in 3 yr and S2T in 4 yr, the relative
cycle length is 0.75, and then, S1T is better than S2T.
The boundary between S1T and S0T depends more on
S0-heritability. For high heritabilities, h2S0 ⫽ 0.80, with a
cycle length ratio greater than 1.30 (for example, S1T
in 4 yr and S0T in 3 yr) S0T is more efficient than S1T.
When heritability is lower S1T remains the best method.
For low heritability, with a ratio of 1.50 (S1T in 3 yr and
S0T in 2 yr), S0T tends to be the best.
It is obvious from Fig. 3 and 4 that the cycle length
ratio has a stronger impact than the heritability value.
The relative cycle length determines most of the relative
efficiency, while heritability modulates this relative effi-
ciency to a lesser extent. Consideration of the same
effective size still decreases the effect of heritability,
even for the comparison between S0T and SDHT.
Comparison at the Optimum
The optimum number of replications depends on the
selection method, i.e. on the inbreeding coefficient (F)
of selection units. Obviously, the cycle length has no
influence on such an optimum. With SDHT, two replica-
tions are optimum only when h2S0 is lower or equal to
0.2 (Fig. 5). As the selection units have a smaller in-
breeding coefficient, the optimum number of replica-
tions increases up to three for S0T at h2S0 ⫽ 0.2. The
optimum is relatively flat, and thus the differences in
genetic advance with 1, 2, or 3 replications are small.
When heritability is greater or equal to 0.5 , an evalua-
tion based on single replicate trials ensures the best
genetic advance whatever the method. For a given heri-
tability, the optimum is approximately the same for each
method. With a constant effective size, conclusions
about the optimum number of replications are not
changed.
DISCUSSION
When off-season nurseries are not used, it appears
that SDHT can increase genetic advance per unit of
time in comparison to the other methods when a cycle
can be achieved in 4 yr. With the use of off-season
nurseries, SDHT in a 3-yr cycle has an advantage only
with low heritability values. This advantage is reduced,
and even suppressed if comparisons are made on the
basis of the same effective genetic size. In this case,
excluding S1T in 2 yr, which can be difficult to apply in
a species like maize, S0T in 2 yr appears to be the most
efficient except at very low heritability (Table 2). This
is due to the shortness of its cycle. As previously men-
tioned, the cycle length has a greater effect than the
coefficient of inbreeding and the rank of the methods
corresponds approximately to their cycle length, the
worst having the longest cycle. Indeed, reducing the
cycle length from three to 2 yr increases genetic advance
per year by 50%. One self from S0 to S1 can increase
the expected genetic advance per cycle by 50% at low
heritability, and the gain per year is only 22%. With
high heritability, the genetic advance per year is reduced
to 8% (Table 2). This illustrates that the additional gain
expected by the use of inbreeding can be suppressed by
the associated longer cycle. The use of inbreeding is
efficient, mainly at low heritability, if it can be accom-
plished with the use of off-season nurseries. Even with
SDHT, it is necessary to have a well controlled HD
process to have SDHT among the best methods for the
genetic advance per unit of time. It now appears possible
in maize to complete the HD process in 12 or 18 mo
by the in situ gynogenesis process (Bordes et al., 1997).
The development in off-season nurseries for simultane-
ous pollination and crossing to the tester (S⫹TC) can
decrease the cycle length for several schemes (Table 1)
and then increase genetic advance per unit of time. In
this case, S1T in 2 yr will be one of the best schemes.
The difficulty with such a scheme, for a plant like maize,
will be the low quantity of seeds produced, even by
crossing the candidate plant to several plants of the
tester, a solution which implies the use of a pure line
or a single-cross as tester.
As far as the influence of heritability is concerned,
we must note that in maize breeding for grain yield,
plot heritability at the S0 level is often about 0.40 to
0.50, leading to design–heritability greater than 0.50 with
two or three replicates (Moreno-Gonzalez and Hal-
lauer, 1982; Gallais, 1996; Sampoux and Gallais, 1996).
With design heritabilities of 0.60 to 0.70 and the same
selection intensity, the relative genetic gain obtained by
the use of SDHT, in comparison to S0T, is between 0
and 25%. With the same effective size, there will be no
significant gain, and worse, SDHT can be lower than
S0T. Note that a gain of 10% is low in absolute value.
Thus, considering HD as a costly process, its interest
for the improvement of combining ability in recurrent
selection is not as clear as for the improvement of line
value. However, genotype ⫻ year interaction will fur-
ther reduce heritability. If genotype ⫻ year interaction
represents 50% of the total genetic variance, operational
 BOUCHEZ & GALLAIS: DOUBLED-HAPLOIDS IN RECURRENT SELECTION FOR COMBINING ABILITY
heritability will not be 0.60 to 0.70 but 0.30 to 0.35. Then,
the advantage of SDHT will be increased. Whatever
the situation, there still remains an interest in SDHT
because it allows rapid development of new hybrids,
particularly when off-season nurseries are not used.
Moreover, HD produces completely homozygous lines,
while some residual heterozygosity is possible when
lines are derived by single-seed descent or pedigree
selection. Then, if the HD process is expensive, it could
be used every second cycle of recurrent selection to
quickly develop hybrids, in alternation with the use of
S0T. S0T in 2 yr is the shortest selection scheme available
that allows separation of self-pollination and crossing
to the tester. Note also that, from the point of view of
the existence of genotype ⫻ year interaction, with the
same expected genetic advance, a short cycle will be
better than a long cycle. In 6 yr, three complete S0T
cycles can be developed, whereas only two cycles will
be developed for SDHT in 3 yr. Obviously, the breeder
has to consider the cost of genetic advance. However,
it is sometimes efficient to invest more, even if this
increases the cost of genetic advance, in order to achieve
quicker varietal development, allowing success on the
market, which will make the investment profitable. Use
of doubled-haploids, if well controlled, is a tool to satisfy
this goal.
APPENDIX
Derivation of Between and Within
Progenies Variance
Consider, for one locus, a random mating population with
alleles A1p, A2p, . . . Aip, Ajp and a tester population with alleles
A1t , A2t , . . . Ait, Ajt.
The value of a genotype Aip, Ajt from the population x tester
cross can be written
Yijpt ⫽ ␮T ⫹ ␣ip ⫹ ␣tj ⫹ ␤ijpt [1]
where ␮T is the mean of all testcross progenies, ␣ip the additive
effect of allele Aip in combination with the tester, ␣jt the additive
effect of allele Ajt in combination with the population, and
␤ijpt the dominance effect, i.e. the interaction between two al-
leles, one from the population and the other from the tester.
Now consider the combining ability of a genotype Aip Ajp
from the S0 population. It is
T(ij) ⫽ ␮T ⫹ 1/2(␣ip ⫹ ␣jp )
or
T(ij) ⫽ ␮T ⫹ T ␣i ⫹ T ␣j
with
T ␣i ⫽ 1/2 ␣ip. [2]
The variance among progenies will be
␴2GBT ⫽ ␴2AT
with
␴2AT ⫽ 2E (T ␣2i )
and more generally with inbred plants
␴2GBT ⫽ (1 ⫹ F )␴2AT. [3]
The within progeny variance will be
␴2GWT ⫽ ␴2Gtot ⫺ ␴2GBT. [4]
29
From [1] :
␴2Gtot ⫽ E (␣ip)2 ⫹ E (␣ti)2 ⫹ E (␤ijpt )2.
According to [2], the first term is equal to 2␴2AT and the last
two represent variance due to the heterogeneity of the tester
(␴2WT). Then,
␴2Gtot ⫽ 2␴2AT ⫹ ␴2WT
and from [4]
␴2GWT ⫽ (1 ⫺ F )␴2AT ⫹ ␴2WT
For a set of loci, with a population in linkage equilibrium and
without epistasis, the formulae for variances remain valid,
except that variance components represent summation of one-
locus components over the set of loci.
ACKNOWLEDGMENTS
The authors are very grateful to the reviewers and to Prof.
Kendall R. Lamkey, Technical Editor, for their helpful sugges-
tions and revision of the English.
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