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Principles of Plant Genetics and Breeding

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354 CHAPTER 19<br />

accumulation. Of course, this will have to be done<br />

within reasonable agricultural limits, as dictated by<br />

weather <strong>and</strong> cropping sequence. Genotypes can be<br />

adapted to new growing conditions (e.g., cold tolerance<br />

to allow the farmer to plant earlier than normal).<br />

2 Tolerance <strong>of</strong> adverse environmental factors.<br />

Because <strong>of</strong> the vagaries <strong>of</strong> the weather <strong>and</strong> the<br />

presence <strong>of</strong> other inconsistencies or variation in the<br />

production environment (climate, product management,<br />

etc.), biomass can be enhanced by breeding for<br />

tolerance to these factors. Such breeding efforts may<br />

be directed at developing tolerance to abiotic stresses<br />

(e.g., drought, heat, cold). This would allow the cultivar<br />

to produce acceptable yields in the face <strong>of</strong> moderate<br />

to severe adverse environmental conditions.<br />

3 Pest <strong>and</strong> disease resistance. Diseases <strong>and</strong> pests can<br />

reduce biomass by killing plant tissue (or even an<br />

entire plant in extreme cases), <strong>and</strong> stunting or reducing<br />

the photosynthetic surface <strong>of</strong> the plant. Disease<strong>and</strong><br />

pest-resistance breeding will enhance the biomass<br />

potential <strong>of</strong> the crop. <strong>Breeding</strong> to control pests is one<br />

<strong>of</strong> the major undertakings in plant breeding.<br />

Ideotype concept<br />

<strong>Plant</strong> breeders may be likened to plant structural <strong>and</strong><br />

chemical engineers who manipulate the genetics <strong>of</strong><br />

plants to create genotypes with new physical <strong>and</strong><br />

biochemical attributes for high general worth. They<br />

manipulate plant morphology (shape, size, number <strong>of</strong><br />

organs) to optimize the process <strong>of</strong> photosynthesis,<br />

which is responsible for creating the dry matter on<br />

which yield depends. Once created, dry matter is partitioned<br />

throughout the plant according to the capacity<br />

<strong>of</strong> meristems (growing points <strong>of</strong> the plant) to grow.<br />

Partitioning (pattern <strong>of</strong> carbon use) is influenced by<br />

both intrinsic (hormonal) <strong>and</strong> extrinsic (environmental)<br />

factors. Certain plant organs have the capacity to act as<br />

sinks (importers <strong>of</strong> substrates) while others are sources<br />

(exporters <strong>of</strong> substrates). However, an organ may be a<br />

source for one substrate at one point in time <strong>and</strong> then a<br />

sink at another time. For example, leaves are sinks for<br />

nutrients (e.g., nitrates) absorbed from the soil while<br />

they serve as sources for newly formed amino acids.<br />

<strong>Plant</strong> genotypes differ in patterns for partitioning <strong>of</strong><br />

dry matter. This means plant breeders can influence dry<br />

matter partitioning. Pole (indeterminate) cultivars <strong>of</strong><br />

legumes differ in patterns <strong>of</strong> the partitioning <strong>of</strong> dry<br />

matter from bush (determinate) cultivars. Similarly, in<br />

cereal crops, tall cultivars differ from dwarf cultivars in<br />

the pattern <strong>of</strong> dry matter partitioning. The concept <strong>of</strong><br />

the plant ideotype was first proposed by C. M. Donald<br />

to describe a model <strong>of</strong> an ideal phenotype that represents<br />

optimum partitioning <strong>of</strong> dry matter according to<br />

the purpose for which the cultivars will be used. For<br />

example, dwarf (short statured) cultivars are designed<br />

to channel more dry matter into grain development<br />

whereas tall cultivars produce a lot <strong>of</strong> straw. Tall cultivars<br />

are preferred in cultures where straw is <strong>of</strong> economic<br />

value (e.g., for crafts or firewood). Consequently, ideotype<br />

development should target specific cultural conditions<br />

(e.g., monoculture, high density mechanized<br />

production, or production under high agronomic<br />

inputs). All breeders, consciously or unconsciously, have<br />

an ideotype in mind when they conduct selection within<br />

a segregating population.<br />

<strong>Plant</strong> morphological <strong>and</strong> anatomical traits (e.g., plant<br />

height, leaf size) are relatively easy for the breeder<br />

to identify <strong>and</strong> quantify. They do not vary in the short<br />

term <strong>and</strong> also tend to be highly heritable. Consequently,<br />

these traits are most widely targeted for selection by<br />

breeders in these programs. The wheat ideotype defined<br />

by Donald comprised the following:<br />

1 A relatively short <strong>and</strong> strong stem.<br />

2 A single culm.<br />

3 Erect leaves (near-vertical).<br />

4 A large ear.<br />

5 An erect ear.<br />

6 Simple awns.<br />

It is not practical to specify every trait in modeling an<br />

ideotype. The degree to which a plant model is specified<br />

is left to the discretion <strong>of</strong> the breeder. A more accurate<br />

ideotype can be modeled if the breeder has adequate<br />

information about the physiological basis <strong>of</strong> these traits.<br />

The group <strong>of</strong> traits used to define the ideotype presumably<br />

are those that would contribute the most to crop<br />

economic yield under the range <strong>of</strong> environmental <strong>and</strong><br />

management conditions that the crop would encounter<br />

during its life. As previously indicated, physiological<br />

processes are common to all plants, but there is no<br />

universal ideotype in plant breeding. This is because<br />

<strong>of</strong> the vast morphological <strong>and</strong> physiological diversity<br />

<strong>of</strong> crops, <strong>and</strong> the wide range in their economic end<br />

products as well as the cultural environments.<br />

The role <strong>of</strong> partitioning in determining crop yield<br />

depends on the species <strong>and</strong> the products <strong>of</strong> interest. In<br />

forages, the total above-ground vegetative material is<br />

harvested as the end product. The importance <strong>of</strong> partition<br />

in the economic yield in this instance is small. In<br />

other crops, the desired product is enhanced at the

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