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PBG 650 Advanced Plant Breeding

Module 10: Recurrent Selection


Recurrent selection

Cyclical selection of populations


form families
evaluate in trials
recombine selections
Pedigree selection and improvement of elite lines
are also cyclical processes, but the population
structure is not so clearly defined
Selfing and introgression of new germplasm are
common features of both selection systems
Recurrent selection and development of lines can be
integrated into a comprehensive system

Bernardo, Chapt. 10
Rationale for recurrent selection

Selfing systems:
Fixation of alleles is so rapid that the impact of selection is limited
Probability of obtaining segregants with all of the favorable alleles
controlling a quantitative trait is small
Example: with 5 loci, all alleles have p=0.5
1/32 chance to get all of the good alleles
Recurrent selection:
systematically increases the frequency of favorable alleles

Example: with 5 loci, all alleles have p=0.6

1/13 chance to get all of the good alleles


maintains the genetic variation within a population to permit continual
progress from selection
Recurrent selection in practice

Why is it not used more often? (Bernardo)


Easy to apply in cross-pollinated crops; difficult in self-
pollinating crops
male sterility systems can be used
Objectives are long-term
several generations needed to complete a cycle
Immediate output is an improved open-pollinated variety, not
a line or hybrid
Need to choose one or a few populations for selection
not as much opportunity for speculation in use of germplasm
Nonetheless, there are many examples of widescale use of
varieties developed from recurrent selection schemes
Expected selection response

Progeny
Phenotype






b OP
h2


Expected





selection



response



R = h2S







Mid-parent



Phenotype


Mean of






selected


parents

Selection Selected parents


differential
S
Truncation point

Mean of all possible parents


(selection candidates)

Source: Lecture by Jean-Luc Jannink at Iowa State, 2004


Response to selection

R=h2S X0

S
Selection differential
S XS X0 Xs Select best
10% of C0

60 70 80 90 100 110 120 130 140 150


Response to selection
Recombine to form C1
R X1 X0 X1

Realized heritability
R
R 113 100
h
2
0.75
S 117 100
60 70 80 90 100 110 120 130 140 150

Falconer and Mackay, Chapt. 11


Predicting response to selection

R=h2S
Need estimates of h2 and the selection
differential

In theory, h2 is only applicable for a single


generation, because heritability depends on
gene frequencies. In practice, predictions
seem to work for 5-10 generations.
Selection differential

S can be predicted if we can assume:


normal distribution of phenotypes
truncation selection
Standardized selection differential (i)
S = iP Derivation of selection intensity from
the standard Normal distribution

i = S/P = z/p i = z/p


=1

p = proportion selected
z = height of curve at z
p

truncation point
i = standard deviations 0 x i

from the mean


Values of standardized selection differential

p i p i
0.90 0.20 0.09 1.80
0.80 0.35 0.08 1.86
0.70 0.50 0.07 1.92
0.60 0.64 0.06 1.99
0.50 0.80 0.05 2.06
0.40 0.97 0.04 2.15
0.30 1.16 0.03 2.27
0.25 1.27 0.02 2.42
0.20 1.40 0.01 2.67
0.15 1.55 0.005 2.89
0.10 1.76 0.001 3.37

Becker, 1984 Appendix Tables 2 and 3 (infinite population size)


Response to selection

R=h2S

S = iP
i
2
A 2

R=ih2P h 2
2
R A

P P
Applies to individual plants in a population
Selections made before flowering + controlled matings among
selected individuals
Mass selection + selfing of selected plants
Family selection

(O) (X)
Parental plant Selection Unit
in reference (progeny mean)
population

(W)
(R) Individual in
Recombination improved
unit population

Cov(X,W) determines expected gain from selection

Hallauer, Carena and Miranda (2010) Chapt. 6


Intrapopulation Improvement

Method Progenies tested Recombination


unit
Mass selection (both parents) Individual plants Individual plants
Mass selection (one parent) Individual plants outcrossed seed

Half-sib (progeny selected) Half-sib families Half-sib families

Half-sib (parent is selfed) Half-sib families S1 family

Modified ear-to-row Half-sib families outcrossed seed

Full-sib Full-sib families Full-sib families

S1 family S1 family S1 family

S2 family S2 family S2 family


Intrapopulation Improvement
Method Expected Gain Generations/Cycle
Mass selection (both parents) i P
2
A
1
Mass selection (one parent) i (1/2) A P
2
1
Half-sib (progeny selected) i (1/4) A2 Phs 2

Half-sib (parent is selfed) i (1/2) A2 Phs 3

S1/Testcross i (1/2) A
2
Phs 4

Modified ear-to-row i (1/8) A2


Phs 1

Full-sib i (1/2) A2 Pfs 2

S1 family* i A2 PS1
3

i (3/2) A P
2
S2 family* 4
S2

P is the square root of variance; pertains to selection units


*additive variance for inbred progeny includes an additional
component that is a function of the degree of dominance
Phenotypic variance of families

Half-sibs P
HS
1
re + +
2
e
1
e
2
GE
1
4
2
A

Full-sibs P
FS
1
re + + +
2
e
1
e
2
GE
1
2
2
A
1
4
2
D

S1 families P
S1
1
re + + +
2
e
1
e
2
GE
2
A
1
4
2
D

S2 families
P
S2
1
re + + +
2
e
1
e
2
GE
3
2
2
A
3
16 2
D


2
Error variance r = # replications
e e = # environments
2

GE Variance due to genotype x environment interactions
Interpopulation Improvement

Method Progenies tested Recombination unit

Reciprocal recurrent Half-sib families S1 families

Reciprocal full-sib Full-sib families S1 families

Testcross Testcrosses S1 families


Reciprocal recurrent selection
Half-sibs evaluated (Design I matings)
A0 S1 recombined A1
A0 females
HS yield trials
A1 x B1 (improved cross)
HS yield trials
B0 females
Full-sib RRS
B0 S1 recombined B1 plants must have two ears
Full-sibs evaluated twice the number of plants
can be evaluated
A0 S1 recombined A1 continue to inbreed and
evaluate specific crosses

FS yield trials
A1 x B1 (improved cross)

B0 S1 recombined B1
Interpopulation Improvement

Method Expected Gain Generations


per cycle
Reciprocal recurrent

(i (1/2) 2
A ( P1) P(P1hs ) ) + (i (1/2) 2
A (P 2 ) P( P 2hs ) ) 3

Reciprocal full-sib

3
(i (1/2)( 2
A ( P1) + 2
A (P 2 ) ) P(P1xP2 fs)
Testcross

Depends on choice of tester, but typically 3


Cross P1 plants to inbred line from P2
Cross P2 plants to inbred line from P1
Phenotypic variance of families for RRS

r = # replications
e = # environments

P(P1hs) 1
re 2e(P1) + 1e GE(P1)
2
+ 12 2A(P1)

P(P1xP2fs) 1
re 2e + 1e 2GE + 12 (2A(P1) + 2A(P2) ) + 14 D(P1P2)
2
Comprehensive breeding program

Development of breeding populations from diverse


sources such that the performance of the
population cross is maximized while maintaining
high levels of genetic variance within each
population
Application of an effective recurrent selection
procedure
Development of inbreds from each population with
good combining ability and recycling of superior
inbreds back into the base populations

Eberhart et al., 1967


Increasing selection response

Increase the selection differential (reduce


proportion selected)
Increase the coefficient of A2
i
2

Increase A2
R A

P
Reduce nongenetic effects
Reduce generations/cycle or increase
generations/year
Choice of selection method

I. Breeding Objectives
Open-pollinated varieties, synthetics or hybrids
Status of commercial seed sector
Strategy for distribution of seed
Elite variety or genetic resource
Target production environments
Low or high inputs?
Narrow or broad adaptation?
Number of traits, relative importance of traits
Choice of selection method

II. Genetic, Environmental, External Factors


Heritability of the trait(s)
Extent of GXE
Type of gene action
Effects of inbreeding on the trait
Expected gain per cycle
Number of seasons per cycle
Growing seasons per year and availability of off-
season nurseries
Seed quantities required for screening
Costs and resources available
Maize families seed quantity issues
Family Crosses Seed quantity Comments
Half-sibs 1. Collect pollen in bulk 1. One ear 1. Controlled pollinations or
and cross to a female by detasseling
plant
2. Take pollen from one 2. ~4 ears 2. Full-sib families within
male and cross to half-sibs
several females

Full-sibs Cross two plants One or two ears With or without reciprocals
S1, S2, etc. Self pollination One ear Seed quantities decrease
with inbreeding
Can increase a line by
selfing or by sib-mating
Testcrosses 1. Cross one male plant 1. ~4 ears
to a female tester and
self
2. Cross an S1 line to a 2. many ears Controlled pollinations or
tester (population or by detasseling (if S1 line is
inbred line) female)

one ear at least four single-row plots


Maintenance of Maize Streak Virus Resistance

Modified full-sib family selection


Year 1 Evaluate full-sib families in target environments
Main season,
savanna zone Off-season: data entry and analysis

Year 2 Recombine selected full-sib families by making


First season, plant to plant crosses between families
forest zone

Year 2 1) Plant F1 families ear-to-row under MSV


Second season infestation
(high disease 2) Remove susceptible plants and offtypes
pressure) before flowering
3) Make reciprocal crosses between best plants
in good rows to generate new full-sib families
Reciprocal S1 Testcross Selection (modified)
Year 1 Self
First season

Year 1 Evaluate ~500 S1 families (2 reps, 2 loc)


Second season Select for disease resistance and other
(high disease highly heritable traits
pressure) Testcross to the reciprocal population

Year 2 Evaluate ~ 200 testcrosses (3 reps, 4 loc) in


Main season the target environments
Select for yield and other agronomic traits

Year 2 Recombine selected S1 families


Off-season
Can stagger populations so that one is at the S1 stage and the other is
at the testcross stage each year
Meadowfoam - use of blue bottle flies as pollinators
S1 testcross selection in meadowfoam
Year 1 (spring)
Self ~300 plants in the greenhouse with blue bottle flies
Year 1 (fall) + Year 2 (spring)
Plant rows of ~5 seeds per family in isolation with bees, 2 blocks
Reject S1 families with poor agronomic characteristics (disease,
insect damage, small seeds, etc)
Harvest ~6000 seeds per family in bulk
Year 2 (fall) + Year 3 (spring)
Evaluate ~150 testcross families in yield trials, select ~30
Year 3 (fall)
Recombine S1 seed of selected families in greenhouse

Further selfing of selected S1 lines


Evaluation of experimental varieties in yield trials
Balancing resources for recurrent selection

*Daylength can be controlled in the greenhouse to complete


a generation in four months
Makes efficient use of greenhouse space
New experimental varieties can be evaluated every year