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RECURRENT SELECTION

Overview
The goal of recurrent selection is to improve the mean performance of a population of plants; a seconda ry
goal, but nevertheless very im portant, is to ma intain the gen etic va riability present in the population to the
extent possible.

Progress in selection is based on the heritability of the trait and the types of genetic variation controlling the
trait in the particu lar population under selection and on the selectio n differentia l.

W e will initially discuss various recurrent selection m ethods, and then discuss the ge netic param eters that are
associated with them, as a means to differentiate between them.

The generalized method consists of three parts: (1) De velopm ent of a base population with which to begin
selection, (2) Evaluation of individuals from the population, and (3) Selection of superior individuals and
intercrossing to form a new population.

Evaluation of individuals in the population


Individuals can be evaluated in one of two ways:

Ph enoty pic evaluation considers the individual plant per se, so that no evaluatio n of the actu al breeding value
of the individual is possible.

In con trast, genotypic evaluation is based on the performance of the progeny of the plant under evaluation,
and therefore, provides information on the individual’s breeding value. Three types of progeny may be
evaluated: (1) half-s ib progeny, (2) full-sib progeny, and (3) selfed progeny. Half-sibs are formed by crossing
the individuals to be evaluated to a com m on parent (wh ich can be a population, an inbred line, or a specific
genotype), called a tester; thu s, a ll half-s ib fam ilies have the tester in comm on. To produced full-sib families,
the individuals to be evaluated are crossed in pairwise combinations–thus, full-sib families do not have any
pare nts in com m on, although all the parents c om e from the sam e population. Finally selfed progeny are
produced by self-fertilizing the individuals to be evaluated for one or two generations. These partially inbred
progeny are evaluated.

Methods of intrapopulation improvement


Recurrent phenotypic selection

Recurrent phenotypic selection (RPS), also termed m ass selection (because this term is used in other
situations, I prefer to not use it here), is the simplest form of recurrent selection. Typically, RPS has three
stages: (1) a population of p lants is plante d in a nursery in a manner that allows notes to be taken on individual
plants, (2) the population is evaluated for the trait/traits of interest and the best individuals are identified, and
(3) seed is harvested from the selected individuals to constitute a new population from which the next c ycle
will begin.

A m ajor d eterm inant in rate of progress is parental control; that is, is the seed that is harvested from the
selected plants the result of pollination from only those selection s (i.e., both parents a re controlled), or is it
formed from pollen derived from all the plants in the nursery, both selected and unselected (i.e., only the
fem ale pare nt is co ntrolled ). The form er sh ould theoretically double the rate of pro gres s (i.e., the genetic gain
from selection). However, in some instances, control of both parents is not possible. Consider selection for
grain yield in m aize . Evaluatio n can only occur after pollination , obviously, so when the best ears are selected
from individual plants, clearly the pollen has derived from the entire population, and only the female parent
is controlled. Perennial plants, in contrast, can be dug up from their location and all selections can be
transplanted to a isolated crossing nu rsery (called a polycross nursery) where the plants are typically
replicated to ensure random intercrossing among all the selections, or to the greenhouse to enable hand
crossing among them.

Selection of plants spaced widely apart suffers from two major problems. First, the spaced planting may not
reflect the density of comm ercial production, and if the correlation between performance in space planting and
performance under comm ercial density is low, then selection will be ineffective and progress slow or non-
existent.

Second, single plant performan ce can b e significantly affected by variation in the field. Thus, if for some
environmental reason plants perform better on one side of the nursery than the other, phenotype-based
selection will be ineffective because the phenotype is undually affected by the environment, not the genotype.
One way around this problem is gridding, in which the field is divided into a num ber of equally sized
subdivisions. Selections are then m ade within each grid, such that an equa l numb er of individuals are
identified in each subdivision, for example, five plants within each 5 x 5 grid.

As sugge sted by the exam ple of gridding, R PS can be im prov ed b y making sm all adjustm ents that enable
better evaluation of phe notype s. Further improvements to the RPS system can be envisioned, and Burton
{, 1982 #829} summ arizes a number of m odificatio ns that he has adapte d to selection of forage yield in
bahiagrass. He term s his prog ram , Recurrent Restricted Phenotypic Selection (RRPS), because he has
placed restrictions on how he handles his material to ensure the greatest gain. Gridding is just one of many
such res triction s. H is m eth od is a classic example of practical breeding–attention to the smallest detail and
hard work, as his last sentence says, have resulted in the most effective forage breeding program in the world.

Selection will undoubtedly result in diminishing genetic variability, particularly if few genes are involved in the
trait, because as they become fixed, genetic variability goes down. Selection is most effective when allele
frequencies are near 0.5; if the desirable alleles are below 0.5, increasing their frequency will lead to increased
genetic variation for the trait they control, until their frequency passes 0.5.

Recurrent half-sib selection


The gene ral schem e for recu rrent half-sib selection (RH SS) is to cross individuals be ing evaluated to a
comm on tester, evaluate the half-sib progeny of each plant, select the best individuals, and intercross them.
RHSS can be co nducted for either general or sp ec ific com bining ability. General combining ability (GCA)
refers to the average performance of an individual’s progeny, which are developed by crossing that individual
to a variety of other individuals . If the tester being used is a population of plants, then the progeny
performance represents the pa rental plant’s GC A. If the tester were an inbred line, then the progeny
performance would represent specific combining ability (SCA), becau se it wo uld represent the ability of a
particular plant to combine with a specific inbred rather than with a broad selection of genotypes.

The earliest RHSS program used the ear-to-row method (Hopk ins, 1899). For this metho d, he produce d half-
sib seed by interpollinating a number of individuals from a population; thus, the population per se is the tester,
and selection will be for GCA. The ears thus represent half-sib families, with the pollen coming from the entire
population. The ears were then planted into a single row in an unreplicated trial in one environment (hence,
literally ear-to-row), and selection was based on this trial. (Note that this evaluatio n is not unlike that for RPS,
except that here we’re evaluatin g progeny perform ance rather than single plants). The ears of selected plants,
which received pollen from the population (a ltho ugh this was skewed toward more pollen from the neighboring
half-s ib fam ilies rather than a random sample of the population’s alleles), were then planted into rows next
year. Note that if two seasons of maize could be grown in a single year, two cycles of selection could be
com pleted per year. This was a sim ple, tho ugh not particularly effec tive, selection m etho d.

The modified ear-to-row method (Lonnquist, 19 64, and F ig 15.5 in Fehr, 1987) made a number of alterations
to Hopk ins’ method to avo id som e of its shortcom ings. First, testing the half-sib families is done at three
locations with one replication per location. At two of these locations, data is c ollected an d plan ts are harvested
as norm al done. At the third location, the half-sib families are plante d in isolation. Between half-sib families,
rows consistin g of a b ulk of all half-sib families are planted. The half-sib families are detasseled and the bulk
rows are use d as m ales to effe ct pollination.

Selections are based on the data from the first two locations. From the selected families in the isolation, the
five best ears are selected and bulked to form the next cycle; these will be the half-sib family rows next
sea son . These m odificatio ns still only control the fem ale parent, but at least the pollination occurs with the
entire population ra the r than jus t neighboring half-sib fa m ilies. The with in ro w s electio n is like ly only
moderately effective at best, but it may add a bit to the progress.

A better approach to th ose discussed above would be to control both parents (Jenk ins, 1940 and F ig 15.6 in
Fehr, 1987). In order to effect this, however, would require an extra season–that is, evaluations need to be
conducted, and then, bas ed o n those e valua tions, the parents wou ld be intercross ed. Unless parents can be
maintained in the field (as with perennial forage or turf species), the intercrossing nee ds to be done with
remnant seed; if half-sib seed is to be used for evaluation, then some can be held in reserve to be used for
crossing if that parent is selec ted. Intercrossing of selections is done as described for the modified ear-to-row
method above, except that the male rows consist of a bulk of only the selected half-sib families. The
intercrossing can be done in an off-season nursery, so that a cycle of selectio n can still be completed in one
year.

Selection with in half-sib fa m ilies during the intercrossing phase cannot be done if an off-season nursery is
used, simp ly becau se that nursery is in an environmen t outside the target region of the crop. (So, to be mo re
specific, selec tion could be done, but m ost like ly, it wo uld not be advanta geous to advancing the genetic
potential of your populations.) However, if intercrossing is done in the target environment, either because two
seasons per year can be grow n there or b ecause no winter nursery is used, then selection could be practiced
with in half-sib families. Of course, this is essentially RPS within families, and selection based on unreplicated
single plants is not the most effective type of selection, but some additional gain might be possible, depending
on the trait being selected. A further discussion of this topic with forage crops as the primary focus is Vogel
and P eders en {, 199 3 #38 }.

Now, even though intercrossing based on rem nant half-sib seed of selecte d plants is better than only
controlling the female parents, the half-sib seed being intercrossed only contains half of its alleles from the
selected parent; the rem ainder com es from the population as a whole, which includes a preponderance of
nonselected plants. Thus, an even more effective intercrossing method would be to use selfed seed rather
than half-sib seed. (Again, if the plants themselves can be maintained indefinitely, or can be cloned, then
using them is clearly the best strategy of all.)

The difficulty here is in getting both s elfed and half-sib seed from the sam e plant. In som e species, th is would
not be a problem . Fo r m aize it is. T wo alternatives exist in m aize. First, if the m aize population re liably
produces two ears, then one can be used for selfing and other for produ cing ½ s ib see d. Many m aize
populations do not have this a bility, so in this ca se, you can self pollinate the sing le ear on the individual to
be tested and use pollen from that individual to pollinate several individuals of the tester. The ears on the
tester, then, represent th e half-sib family to be tested. Recombining selfed progeny will require three seasons:
(1) selfing and cros sing to the te ster, (2 ) evaluation, and (3) intercrossing se lfed proge ny. This will lengthen
a cycle of selection. The benefit gained from using selfed rather than half-sib seed may be offset by the longer
selection cycle. More on this subject will be discussed in the next section.

Other types of testers besides the population per se can be used. If the population per se is the tester, then
the m eth ods discussed above can be com pleted in the shortest tim e if h alf-sib seed is used for recombination.
If a different population or an inbred line is used as th e teste r, then follo wing re com binatio n, individuals need
to be crossed to the tester, necessitating three seasons to complete a cycle, similar to using selfed seed for
recombination. Thus, the tester ch ose n ca n influe nce gain p er cycle. Note that using an inbred tes ter could
pose a p roblem if the popu lation developed does no t com bine well with a different tester.

Recurrent full-sib selection

Recurrent full-sib selection (RFSS) tests paired plant crosses rather than crosses of individual plants with a
population (or inbred) tes ter. Fo r this m eth od, paired crosses are m ade betw een individuals in the population
in season one. Evaluations are conducted in the field the next season and the best families are identified.
The third season is used to recombine the best families using remnant seed from the first season. Thus, the
second selection cycle begins in the fourth season, with paired crosses between individuals. To streamline
procedures, paired crosses could be done durin g the third season, combining recombination of the best
families with full-sib fa m ily development for the next evaluation cycle. Thus, paired crosses between
individuals from the selected families can be made. The disadvantage of this is that less recombination
occurs between cycles of selection. The obvious advantage is that it allows a cycle to be completed each
year, with th e use of a single off-season nursery.

Recurrent selection amo ng selfed families


a. Evaluation is based on self-pollinated progeny, usually formed by one
generation of selfing, although two generations has also been used.

b. Season 1: S 0 plants from the popu lation are selfed to produces S 0:1
lines.

c. Season 2: S 0:1 lines evaluated, best lines selected

d. Season 3: Rem nant S 1 seed used to intercross selected lines,


producing the Cycle 1 population with new S 0 plants.

e. Season 4: Self S 0 plants as in S eas on 1 .....

f. Many m odifica tions a re po ssible. Could tes t S 1:2 lines or S 0:2 lines,
depending on the desire of the breede r. The latter (S0:2) is useful if you
cannot produce enough S 0:1 seed for testing.

Methods of interpopulation improvement


Reciprocal recurrent selection

designed to improve both the general and spe cific comb ining ability of two populations. The general idea is
to test and select in each of two populations, using the opposite population as a tester. Alleles do not m igrate
between p opulations, however!

2. Two starting populations should be genetically divergent (although selection will


probably drive them apart anywa y)–the point is that the y should co m bin e well
together.

Reciprocal half-sib selection

a. Season 1: Individual plants (e.g., 100) from each po pulation are (1)
selfed and (2) outcrossed to several (e.g., 6) random ly
chosen plants from the opposite population to generate ½
sib families

b. Season 2: 100 ½ sib families from each population are evaluated in


replicated tests; top 10 are identified

c. Season 3: G o back to selfed seed of supe rior ind ividua ls and


intercross to form Cycle 1 population within each
population. DO NO T m ix the s elections of the two
populations! Note that if the plants can be maintained
clonally, you could go back to th e original plants and
intercross, which would mean you wouldn’t have to
produce S1 seed in Season 1.

d. Season 4: Begin next cycle using Cycle 1 seed of each population.

e. Modifications:
i. Use inbred testers developed from the opposite population. The
inbred tester would be replaced by superior ones as the program
progressed.

ii. Produce half-sib seed of each population as in ear-to-row selection.


Then plant half-sib fam ilies in alternate rows with the opposite tester
population, detassel the half-sib lines and use the open pollinated
seed for testing. The best lines are then intermated using the
remnant half-sib seed of each parent. The disadvantage is that
only the female parent is controlled.

Reciprocal full-sib selection


a. Used when the desired outcome is comm ercial hybrid seed.

b. Season 1: 200 phenotypically desirable S 0 plants in Pop A paired with


200 from Pop B. Each plant is both selfed an d cro sse d to
produce S 1 and fu ll-sib seed. Selfed seed can be stored
for intercrossing of des irable p aren ts and also advanc ed to
develop inbred lines.

c. Season 2: Evaluation and selection

d. Season 3: Intercrosss selfed seed of top 20 individuals from each


population–>Cycle 1 populations for A and B.

e. Season 4: Begin again.

f. Alternative if can’t produce both selfed and crossed seed at the same time:
use S 0:1 lines to m ake c rosses. Ad ds a year.

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