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Principles of Plant Genetics and Breeding

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428 CHAPTER 23<br />

unit: in the field, these are plots (<strong>of</strong> rows <strong>of</strong> plants); in<br />

the greenhouse, a unit could be a pot. For example,<br />

applying different levels <strong>of</strong> fertilizer, planting at different<br />

spacing, or evaluating different genotypes (as in breeding),<br />

are all treatments that elicit variable responses from<br />

plants. However, in addition to the expected variation<br />

from a treatment, there is always some variation that is<br />

unintended <strong>and</strong> cannot be accounted for, <strong>and</strong> is known<br />

in research as experimental error. By definition, this is<br />

the variation among plots that are treated alike. Major<br />

sources <strong>of</strong> error are those due to soil heterogeneity,<br />

the operator’s inability to conduct the experiment uniformly<br />

as intended, <strong>and</strong> interplant competition.<br />

Soil (site) variability<br />

Soils are naturally heterogeneous, some more so than<br />

others. Natural variation may originate from differences<br />

in soil minerals, soil moisture, organic matter, or topography.<br />

The tops <strong>of</strong> slopes are drier than the bottom<br />

parts; nutrients wash down <strong>and</strong> accumulate at the<br />

bottom; depressions may drain poorly. In addition to<br />

these natural sources <strong>of</strong> soil variation, humans create<br />

additional variation through how they manage the soil<br />

in production. Before selecting <strong>and</strong> using a site for<br />

genotype evaluation, the breeder should know about<br />

the previous use <strong>of</strong> the l<strong>and</strong> (history <strong>of</strong> use). The parcel<br />

<strong>of</strong> l<strong>and</strong> <strong>of</strong> interest may have been differentially managed<br />

(e.g., different plant species were grown, or different<br />

tillage practices were imposed). The breeder should<br />

use all available data (e.g., yield records, management<br />

records) <strong>and</strong> visual observations to identify general patterns<br />

<strong>of</strong> variation at the proposed site. Experimental<br />

design techniques <strong>and</strong> other tactics can be used to minimize<br />

the effects <strong>of</strong> soil heterogeneity in field trials.<br />

As previously indicated, breeders usually have selected<br />

sites at which they conduct their yield trials (e.g., experimental<br />

stations <strong>of</strong> universities). Are locations then a<br />

fixed variable in ANOVA? Some argue that locations<br />

are r<strong>and</strong>om effects since the breeder has no control over<br />

the meteorological factors that occur at locations. Most<br />

breeders also consider years <strong>of</strong> testing as r<strong>and</strong>om effects.<br />

It is important to have at least two replications <strong>of</strong> each<br />

treatment (genotypes) in each trial for estimating error<br />

variance.<br />

Tactics for reducing experimental error<br />

In order to correctly <strong>and</strong> effectively evaluate the desired<br />

variation, the researcher should eliminate or, more real-<br />

istically, minimize extraneous variation. Some errors<br />

come from natural soil variability whereas others are<br />

human in origin. The plant breeder may use certain field<br />

plot techniques to minimize experimental errors.<br />

Use <strong>of</strong> border rows Different genotypes differ in various<br />

plant characteristics to varying extents (e.g., growth<br />

rate, size, height, nutrient <strong>and</strong> moisture uptake). When<br />

planted next to each other, interplot competition may<br />

cause the performance <strong>of</strong> one genotype to be influenced<br />

by another in the adjacent plot. In early generation or<br />

preliminary trials, which usually include large numbers<br />

<strong>of</strong> genotypes, breeders <strong>of</strong>ten use fewer rows (e.g., two<br />

rows) in planting a plot in order to save resources. In<br />

advanced yield trails, four-row plots are customarily<br />

used. Data are collected on the middle rows only, because<br />

they are protected from border effects. Some breeders<br />

minimize border effects by increasing interplot spacing<br />

or using a common genotype for planting the border <strong>of</strong><br />

all plots in a test in which row plots are few (one or two<br />

rows). To reduce interplot competition, the materials<br />

may be grouped according to competitive abilities.<br />

Proper choice <strong>of</strong> plot size <strong>and</strong> shape Several factors<br />

affect the optimum plot size to use in field evaluation<br />

<strong>of</strong> genotypes (breeding objectives, stage <strong>of</strong> breeding,<br />

resources available, equipment). Evaluation <strong>of</strong> an F 2<br />

segregating population is <strong>of</strong>ten based on individual<br />

plant performance (i.e., individual plants are essentially<br />

plots). Consequently, the plants are adequately spaced<br />

to allow the breeder enough room to examine each<br />

plant.<br />

Some breeders use what are called microplots, consisting<br />

<strong>of</strong> planting hills or short rows <strong>of</strong> test plants. This<br />

tactic is used in the early stages <strong>of</strong> genotype evaluation<br />

as a quick <strong>and</strong> inexpensive way <strong>of</strong> eliminating inferior<br />

genotypes.<br />

Row plots are commonly used by breeders for genotype<br />

evaluation. The size <strong>and</strong> shape depends on the plant<br />

species, the l<strong>and</strong> available, <strong>and</strong> the method <strong>of</strong> harvesting.<br />

Generally, row plots are rectangular in shape.<br />

Adequate number <strong>of</strong> replications Replication or repetition<br />

<strong>of</strong> treatment in a test is critical to statistical<br />

analysis, providing a means <strong>of</strong> estimating statistical<br />

error. The number <strong>of</strong> replications used usually varies<br />

between two <strong>and</strong> four; fewer replications may be used in<br />

early evaluation <strong>of</strong> genotypes while advanced yield tests<br />

usually have four replications. The number <strong>of</strong> replications<br />

depends on the accuracy desired in the analysis, the

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