should be viewed as high estimates because the weather conditions, pathogen, and host susceptibility were optimal. The average annual US losses are probably more similar to those estimated by Green and Campbell for Canada (1979). The three rusts combined reduce the annual US wheat crop by about 2% (table 3.1); most of the losses are caused by leaf rust. The low level of rust losses is attributable mainly to the use of resistant cultivars.

Health and Environmental Impacts

No formal assessment of the health or environmental impact of conventionally breeding wheat for resistance to rust has been undertaken by regulatory agencies, inasmuch as the products of conventional plant breeding have generally not required their oversight. Rust-resistant wheat cultivars, regardless of the source of their resistance genes, have been widely grown and consumed in food products with no history of causing health problems.

Little is known about the biochemical basis or gene products for plant protection against rust, but these genes are likely to be similar to other genes in the large class of race-specific resistance genes isolated from other plants. The presence of pest-resistance genes can affect end-use quality by affecting the grain protein content. For example, leaf rust detrimentally affects leaves reducing their potential for nitrogen remobilization to the grain and reducing grain protein content (Cox et al. 1997). Lower grain protein content is generally considered a detrimental effect in hard wheats but a beneficial effect in soft wheats (Finney et al. 1987). Stem rust tends to reduce the flow of photosynthate and nitrogen to the grain; but because nitrogen is mobilized early in grain development, the overall result of stem rust is generally an increase in protein content in the grain, possibly including shriveled kernels.

Environmentally, the use of rust-resistant wheat cultivars has reduced the use of fungicides, but the extent of this reduction is not well documented, because effective pest-protection has been widely deployed for many years and the economics of wheat production often preclude the widespread use of fungicides that are effective against rust.

3.1.2 Bt Crops

The most widely used transgenic pest-protected plants are cultivars that express insecticidal proteins derived from the bacterium Bacillus thuringiensis (Bt). Cotton and corn are protected from some of their lepidopteran pests by Bt proteins in the Cry1A and Cry9C groups. The po-



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