1. Early Generation Testing, Mass Selection and Bulk Selection
PLS 664
February 21, 2011

2. Early Generation Testing
Objective: identify those populations that are likely to contain superior lines
Strategy: eliminate populations with low potential from the inbreeding process
Goal: maintain and develop lines from populations with high genetic potential

4. Jenkins, 1935
Usual method of estimating combining ability in maize was to inbreed lines, then mate them to a common tester
Jenkins saved seed from S0:1 lines through many selfing generations, then crossed them to common tester
Found that combining ability was already determined in S0:1 lines

5. Self-Pollinated Crops
Determine the generation for testing
If it is to be the F2, you will have to grow the F1 in an environment which favors seed production
A more common choice would be F2:3 lines

6. Self-Pollinated Crops
Harvest seed from individual F2 plants
Plant seeds in F2:3 progeny rows
Identify the superior rows
Harvest all seed in each selected row in bulk
Grow replicated tests of F2:4 lines
Grow replicated tests of F2:5 lines

7. Self-Pollinated Crops
Harvest selected F5 plants individually
Grow F5:6 lines in headrows
Test F5 - derived lines extensively

8. Breeder’s Decisions Generation to test
Number of reps, locations and years - tradeoff between early and late generation testing
Separate program for inbreeding or not
Selected lines can be advanced by pedigree, bulk, or SSD
Number of plants chosen from each hetergeneous line may vary

9. Genetic Considerations
Recall that there is all of the additive variance
among F2:3 lines and one-half of the additive
variance within F2:3 lines
In later generations of F2 derived lines, there is
still all of the additive variance among lines,
and considerable variance within lines, as
inbreeding progresses

10. Genetic Considerations
Therefore, one may need to take a large number of heads to adequately sample the variation within the F2 - derived line
Now one must decide how to allocate resources
Should you sample more lines or more selections within lines?

11. Pros
Inferior individuals and crosses are discarded early in the process
One hetergeneous line may yield more than one cultivar

12. Cons
When you commit a lot of resources to early generation testing, you cannot devote as much to thorough evaluation of more inbred material
If you spend a lot of time testing the early generations, cultivar release may be delayed

13. From Bernardo, 2003 CROP SCIENCE 43 : 1558-60

14. Bernardo Simulation paper; no actual data
Demonstrated that even though the genetic correlation may be high, it was more strongly affected by non-genetic (ie environmental) effects than by the generation of selfing

15. Mass Selection
Selective harvest of individuals from a heterogenous population.
Oldest method of plant improvement. Humans have been
Selecting desirable seeds for years, even since Neolithic times.
Quickest way to make progress with undeveloped populations
Reviews heavily from Fehr and Allard…also check our cited papers.

16. Examples of selection criteria height weight color
texture chemical content shippability (cookies)
storability timing of maturity taste!!
# tillers shape disease resistance
“vigor” clean leaves stay green
Selection success is related to the single plant
heritability estimate of each trait.
So it means we’re forming a population by selecting superior individuals from the group you’re working with.
They simply would look at a population (or later a production field) and choose sets of grain that were larger, disease free, appealing to the eye, and plant them back next year.
As Dr.. Murphy said in my class, if you take a job as a new prof, start using mass selection on a landrace and you’ll make great progress. On the other hand….
It’s used for cultivar purification.

17. Mass Selection – How To
Stage #1 = Selection of individuals within a heterogeneous
population. Choose favorable/desirable plants and harvest
the seed.
Stage #2 = Sampling equal numbers of seed from selected
individuals to plant the next generation.
Some of these we’ve already discussed. But now we’re actually implementing selection.
Color: fruit crops, edibles
Shippability: watermelon drop test with Dr. Wehner
Taste: strawberry breeder in CA. Wehner and watermelon
Storability : food crops
Kick test: corn breeding…trying not to hit coworker on the back of the head.
Low h2 = low selection response

18. Mass Selection in Oat : Romero & Frey, 1966
1958: Plant out heterogeneous
population of F3 oat seed. Using
a lawn clipper, cut oat plants to
uniform height (determined by check cultivar).
Harvest only top 10 cm in order to select against
short plants.
10 cm
Collected seed composed
of complete panicles, partial
panicles, or no panicle.

19. 1959 (F4), 1960 (F5), and 1961 (F6): Repeated the procedure.
Evaluated 75 plants from the unselected group of each generation,
75 plants from the selected group of each generation, and 75 plants
of a pure line check.
Results
Significant reduction plant height = 0.47” generation-1
Associated shift towards earliness = 0.23 days cycle-1
Shift towards higher yields = 0.41 grams plot-1. Non-significant
At a later date…they perform evaluations. Evals were performed in field trials.
We will look at graphs of this data.

20. Height: reduction as expected…
Height: reduction as expected….they were selecting on basis of medium height.
Earliness : not directly selected, but apparently correlated with height.
Increase of weight…must also be correlated with height.

21. Mass selection in soybean for maturity and
calcareous soil tolerance : Fehr, unpublished
Yellowing of soybean indicates lack of ability to utilize available
iron.
Cultivar with desirable levels of tolerance and maturity were planted
as standards.
Plants with more yellowing than standards were removed before
flowering.
One pod/plant was harvest and bulked from selected individuals
** These are two examples of selections based on individual
plants within the population.
rg DI
h2D h2I
It makes sense to assume that height and yield are correlated b/c of the limited resources of Pn. You only have so many products that either must go into growing upwards, or filling grain head.
Height and earliness are an unexpected correlation. But a positive one that the breeder can take advantage of.
h2D = heritability of directly selected trait
h2I = heritability of indirectly selected trait
rg DI = genetic correlation between D&I traits

22. Mass selection of seed size in soybeans : Fehr & Weber, 1968
1963: F6 plants were planted and 400 plants of early maturity
were selected.
Top 1/4 of main stem and all branches were removed to discard
poor seed; remainder of plant was threshed together.
Seed was passed over different sized sieves and 25% of the largest
and smallest seeds were retained.

23. Specific gravity tests in glycerol-water solutions were performed
for each size group. 25% of the seeds with a high density and
25% of the seed with a low density were selected.
1964 : 2300 F7 seeds from each group were planted: 400 selected
for harvest in same fashion.
Repeat sizing and specific gravity tests.

24. 1965 : Process repeated with 1000 F8 seeds from each group.
Results
Linear change in seed size
Progress for high protein- low oil was best in large seed/high specific
gravity set.
Progress for high oil - low protein was best in small seed/low specific
This is an example of selections and sampling being performed
simultaneously on the harvested seed.
Obviously protein and oil are correlated with seed size and density.

25. Mass Selection
Selection can be applied to 1) individual plants, 2) seeds.
Selection involved is 1) artificial selection, 2) natural selection,
3) both.
Selection for 1) one trait, 2) multiple traits
Sampling methods :
1) random sample of selected seeds bulked
2) equal quantities of seeds harvested & bulked from selected
individuals
3) selection performed on seeds which are used to plant out the
next generation. (Selection & sampling done simultaneously).

26. Cultivar Purification : mass selection is used routinely
in the maintenance of purity for self-pollinated cultivars
or inbreds of cross-pollinating species. It involves roguing of
off-types (removal of individuals that do not conform to the
normal types).
Genetic Considerations:
Leads to higher percentages of desired genotypes.
Effectiveness is a function of h2 of trait on a single plant basis.
Improving h2 will improve gain.
Also a function of correlation if selecting for opposite trait.

27. Advantages
Rapid, inexpensive procedure for increasing frequency of desired
genes.
Can repeat over years until no more progress seems apparent.
Disadvantages
Can only be used in environment where character is expressed.
Prevents use of off-season nurseries.
Limited value for low h2 traits (like yield).

28. Mass Selection
Additional thoughts Mass selection is distinguished from other forms of selection by two criteria: 1. Mass selection is the selection of individuals rather than families. 2. It is based on the performance of the individual in that generation, not on a progeny test.

29. History: 1. Natural selection: removes less fit individuals. 2
History: 1. Natural selection: removes less fit individuals. 2. Leaming, 1825, began selecting the best ears in a field of OP corn and improved the Leaming variety. 3. Pearl, 1907, conducted mass selection for egg production in chickens. Overall, the results were mixed. This led to the conclusion that mass selection was not very successful with low heritability traits, i.e. those in which the environmental effect was large.

30. In a plant breeding context, we think of two forms of mass selection: 1. Selection of pre-flowering traits, followed by intermating the selected individuals that same season. 2. Selection on an individual plant basis of top performing females (in an cross pollinated crop) Selection on an individual plant basis, no intermating (eg in self pollinated crops)

31. The best-known example of mass selection is the work of C. O
The best-known example of mass selection is the work of C. O. Gardner (Crop Sci. 1: ). He began with the OP variety Hays Golden. To begin with, Gardner determined yield from each plant at 10% moisture. He then bulked 20 seeds from each selected plant to provide seed for the next cycle of selection.

32. In 1956 he imposed a grid system on his selection scheme: what was the motivation for doing this? Micro environmental variation was impeding his selection response. He planted 40 plants per cell and selected the top 4 yield plants in each cell, to reduce problems associated with soil variability, etc. Equal numbers of seeds from each cell were bulked for the next cycle of selection.

33. Selection Response Population Kg/ha % of check Irradiated pops C0 5040
100
4696
92.2 C1
5206
103.3
5384
106.8 C2
5364
106.4
5428
107.7 C3
5708
113.2
6057
120 C4
6991
127.8

34. The C0 was Hays Golden. In order to evaluate selection response, new seed from each cycle of selection would have to be generated and the experiment would be grown in replicated plots at several locations. To determine actual selection response, the usual approach is to regress cycle means on cycle numbers, and the linear regression coefficient is used as the estimate of selection response.

35. Selection Response over 15 Cycles of Selection
% of ck.
150
C0 C C15

36. Also known as mass selection Grow the segregating bulk
Modified Bulk System
Also known as mass selection
Grow the segregating bulk
Harvest seed from superior individuals
Combine or “bulk” the harvested seed
This bulk constitutes the population for the next year, or next cycle of selection

37. Modified Bulk System Very simple to implement
Has an excellent track record (dating back to Neolithic times!)
Combines selection and inbreeding
May be difficult to perceive superior individuals in a heterogeneous population

38. Modified Bulk System Does not work well with low heritabililty traits
Does allow the breeder to shape the population while inbreeding

39. UK Modified Bulk System
F2: select bright clean heads from early, short disease free plants; thresh in bulk
F3: same procedure
F4: select bright clean heads from early, short disease free plants; keep each head separate and thresh into headrow trays
F5: plant F4:5 headrows; harvest superior headrows separately
F6: test F4:6 lines in single rep trials at 2-3 locations