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Principles of Plant Genetics and Breeding

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the presence <strong>of</strong> β-carotene, but no lycopene. This<br />

finding suggests that enzyme 4 may be present in rice<br />

endosperm naturally, or that it could be induced by<br />

lycopene to turn lycopene into β-carotene. Analysis also<br />

showed the presence <strong>of</strong> lutein <strong>and</strong> zeaxanthin, both<br />

products derived from lycopene. None <strong>of</strong> the above was<br />

found in the control (non-engineered) plants.<br />

Matters arising from the development <strong>of</strong><br />

“Golden Rice”<br />

The initial golden rice lines produced 1.6–2.0 µg <strong>of</strong><br />

β-carotene per gram <strong>of</strong> grain. The recommended daily<br />

allowance (RDA) set by health agencies for children<br />

is 0.3 mg/day. Estimates <strong>of</strong> the bioavailability <strong>of</strong> βcarotene<br />

have been put at less than 10% in some cases.<br />

The scientists intend to refine their invention to make it<br />

produce 3–5-fold its present level <strong>of</strong> β-carotene.<br />

Upon an international intellectual property rights (IPR)<br />

audit commissioned by the Rockefeller Foundation<br />

through the International Service for the Acquisition<br />

<strong>of</strong> Agri-Biotech Applications, Potrykus <strong>and</strong> his team<br />

realized their invention utilized 70 IPRs <strong>and</strong> TPRs<br />

(technical property rights) owned by 32 different companies<br />

<strong>and</strong> universities. Because <strong>of</strong> the humanitarian<br />

goal <strong>of</strong> the project, the development team negotiated<br />

with owners <strong>of</strong> these patents to allow the use <strong>of</strong> their<br />

inventions under the “freedom to operate” clause.<br />

Further, because <strong>of</strong> public pressure <strong>and</strong> the need for big<br />

business to tone down their pr<strong>of</strong>it-oriented public<br />

image, the key companies (e.g., Monsanto) <strong>of</strong>fered free<br />

licenses for their IPRs involved with “Golden Rice”.<br />

It might take an estimated additional 5–10 years<br />

before the ordinary person for whom it is intended to<br />

benefit can produce golden rice. The characters <strong>of</strong> the<br />

present genotype must be bred into as many locally<br />

adapted varieties <strong>and</strong> ecotypes in as many rice-growing<br />

countries as quickly as possible. It is also important that<br />

such breeding efforts be organized such that all rules<br />

<strong>and</strong> regulations concerning the h<strong>and</strong>ling <strong>and</strong> use <strong>of</strong><br />

genetically modified organisms be strictly followed to<br />

avoid stirring up additional controversies. Agronomic<br />

<strong>and</strong> other studies will have to be conducted to determine<br />

how well such golden rice yields, its palatability<br />

<strong>and</strong> digestibility, <strong>and</strong> public acceptance.<br />

The next phase requires additional funds. Because<br />

<strong>of</strong> public protests against the product for a variety <strong>of</strong><br />

reasons, public funding began to quickly dwindle. In<br />

January <strong>of</strong> 2001, a new effort was launched in the<br />

Philippines for a comprehensive set <strong>of</strong> tests to determine<br />

the efficiency, safety, <strong>and</strong> usefulness <strong>of</strong> “Golden Rice”<br />

BREEDING COMPOSITIONAL TRAITS AND ADDED VALUE 411<br />

for people in the developing world. The joint effort<br />

includes the Philippines-based IRRI, Syngenta, <strong>and</strong> the<br />

Rockefeller Foundation. Further, the IRRI has set up<br />

a humanitarian board to oversee this project <strong>and</strong> to<br />

ensure that the highest st<strong>and</strong>ards for testing, safety, <strong>and</strong><br />

support are achieved. The board includes several public<br />

<strong>and</strong> private organizations, such as the World Bank,<br />

Cornell University, the Indo-Swiss Collaboration in<br />

Biotechnology, <strong>and</strong> the Rockefeller Foundation.<br />

<strong>Breeding</strong> improved oil quality<br />

Oil quality improvement is a major breeding objective<br />

for major oil crops such as soybean <strong>and</strong> rape. Soybean oil<br />

accounts for about 22% <strong>of</strong> the world’s total edible oil<br />

production.<br />

Chemical composition <strong>of</strong> seed oil<br />

By chemical composition, soybean oil consists primarily<br />

<strong>of</strong> palmitic (C16:0), stearic (C18:0), oleic (C18:1),<br />

linoleic (C18:2), <strong>and</strong> linolenic (C18:3) acids, <strong>of</strong> which<br />

palmitic acid is one <strong>of</strong> the two major components<br />

<strong>of</strong> saturated fatty acids. Consequently, the physical,<br />

chemical, <strong>and</strong> nutritional quality <strong>of</strong> soybean depends<br />

significantly on its palmitic acid content. Breeders are<br />

interested in lowering the palmitic acid content <strong>of</strong><br />

soybean oil so as to lower its total saturated fatty acid<br />

content for higher desirability for human consumption.<br />

On average, soybean seed oil contains about 120 g/kg<br />

<strong>of</strong> palmitic acid.<br />

On the other h<strong>and</strong>, seed oil <strong>of</strong> modern oil seed rape<br />

contains about 60% oleate (C18:1), 20% linoleate<br />

(C18:2), 10% linolenate (C18:3), <strong>and</strong> only 4% palmitate<br />

(C16:0) <strong>and</strong> 2% stearate (C18:0). The major breeding<br />

goal for oil seed rape quality improvement is to increase<br />

the oleate content, which decreases the amounts <strong>of</strong><br />

polyunsaturated fatty acids, linoleate <strong>and</strong> linolenate.<br />

This would enhance its nutritional quality <strong>and</strong> increase<br />

its potential value as an industrial oil.<br />

Generally, food products with less than 7% <strong>of</strong> total<br />

saturated fatty acids (C16:0 + C18:0 + C20:0 + C22:0)<br />

qualifies to be labeled “low in saturated fatty acids” in<br />

the USA.<br />

Approaches to breeding oil quality<br />

Conventional breeding approaches have been successfully<br />

applied to improve the seed quality <strong>of</strong> various oil<br />

crops. Soybean lines with reduced palmitic acid content

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