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Principles of Plant Genetics and Breeding

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BREEDING FOR RESISTANCE TO DISEASES AND INSECT PESTS 381<br />

variety (or range <strong>of</strong> varieties) suitable for deployment to farmers. As described above, this is not a good option for cassava <strong>and</strong><br />

the other heterozygous, vegetatively propagated crops. Here the desire is to directly enhance the performance <strong>of</strong> existing farmerpreferred<br />

varieties, l<strong>and</strong>races, <strong>and</strong> elite breeding lines, without changing their other beneficial characteristics, which may be lost<br />

due to unpredictable segregation when sexual crossing is carried out. It is thus necessary to genetically transform each desired<br />

cassava variety individually. This presents important challenges, as one has to correctly identify which varieties to target for such<br />

investment <strong>and</strong> then develop the technical capacity to manipulate this germplasm within the tissue culture <strong>and</strong> genetic transformation<br />

systems established for the crop. Greenhouse <strong>and</strong> field testing must then follow for each new transgenic variety. With support<br />

from the United States Agency for International Development, the DDPSC is currently engaged in this process for preferred<br />

cassava varieties from East Africa.<br />

Future directions for transgenic cassava<br />

Biotechnology can have a significant impact on the genetic improvement <strong>of</strong> vegetatively propagated crops, including cassava, if<br />

sufficient resources are committed to such efforts. However, with only five research institutes capable <strong>of</strong> producing transgenic cassava<br />

plants, it is obvious that the funding being committed to this <strong>and</strong> other crops important in developing countries such as plantains,<br />

sorghum, millet, <strong>and</strong> sweet potato are not in proportion to their importance as sources <strong>of</strong> food security <strong>and</strong> economic well-being<br />

for people in the tropical regions. Nevertheless, important progress has been made over the last 5–10 years in developing transgenic<br />

programs for cassava. Genes <strong>of</strong> agronomic interest have been integrated into the crop <strong>and</strong> field trials – a critical step in the<br />

process towards product development – that are being initiated in Africa <strong>and</strong> at CIAT, Colombia. Future programs should benefit<br />

from mapping <strong>and</strong> other genomic-based research. BAC (bacterial artificial chromosome) libraries are being developed for the<br />

crop <strong>and</strong> being used to identify <strong>and</strong> isolate genes responsible for resistance to CMD <strong>and</strong> bacterial blight disease. Once available<br />

these can be integrated into farmer-preferred varieties using the transgenic technologies described above. It is also hoped to<br />

access beneficial genes present in the wild relatives <strong>of</strong> cassava. Within this germplasm can be found traits for enhanced protein<br />

accumulation in the roots, longer shelf-life <strong>of</strong> harvested roots, <strong>and</strong> disease <strong>and</strong> pest resistance. Cassava has an inherent capacity<br />

for high rates <strong>of</strong> photosynthesis <strong>and</strong> the ability to accumulate large amounts <strong>of</strong> starch within its storage roots. It has already been<br />

shown possible to shut down starch accumulation in cassava through the application <strong>of</strong> transgenic technologies. Can this unused<br />

energy be diverted towards the accumulation <strong>of</strong> biosynthetic plastics? Could genes for apomixis be introduced to cassava to<br />

enable true seeds to be produced, thereby revolutionizing the propagation <strong>of</strong> disease-free propagules? Reaching such goals<br />

though traditional breeding alone is not possible. They are only feasible if robust biotechnology programs are developed for cassava.<br />

References<br />

Ceballos, H., C.A. Iglesias, J.C. Perez, <strong>and</strong> A.G.O. Dixon. 2004. Cassava breeding: Opportunities <strong>and</strong> challenges. <strong>Plant</strong> Mol.<br />

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viruses associated with transgene-induced gene silencing in cassava. <strong>Plant</strong> Mol. Biology (available at http://www.kluweronline.com/issn/0167-4412/current).<br />

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Gonzalez-de Schöpke, A.E., C. Schöpke, N.J. Taylor, R.N. Beachy, <strong>and</strong> C.M. Fauquet. 1998. Regeneration <strong>of</strong> transgenic plants<br />

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Li, H.Q., C. Sautter, I. Potrykus, <strong>and</strong> J. Pounti-Kaerlas. 1996. Genetic transformation <strong>of</strong> cassava (Manihot esculenta Crantz).<br />

Nature Biotechnol. 14:736–740.<br />

Sangaré, A., D. Deng, C.M. Fauquet, <strong>and</strong> R.N. Beachy. 1999. Resistance to African cassava mosaic virus conferred by mutant <strong>of</strong><br />

the putative NTP-binding domain <strong>of</strong> the Rep gene (AC1) in Nicotiana bethamiana. Mol. Biol. Rep. 5:95–102.<br />

Schöpke, C., N.J. Taylor, R. Carcamo, et al. 1996. Regeneration <strong>of</strong> transgenic cassava plants (Manihot esculenta Crantz) from<br />

microbombarded embryogenic suspension cultures. Nature Biotechnol. 14:731–735.<br />

Siritunga, D., <strong>and</strong> R.T. Sayre. 2003. Generation <strong>of</strong> cyanogen-free transgenic cassava. <strong>Plant</strong>a 217:367–373.<br />

Taylor, N., P. Chavarriaga, K. Raemakers, D. Siritunga, <strong>and</strong> P. Zhang. 2004. Development <strong>and</strong> application <strong>of</strong> transgenic technologies<br />

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Taylor, N.J., M. Edwards, R.J. Kiernan, C. Davey, D. Blakesley, <strong>and</strong> G.G. Henshaw. 1996. Development <strong>of</strong> friable embryogenic<br />

callus <strong>and</strong> embryogenic suspension cultures in cassava (Manihot esculenta Crantz). Nature Biotechnol. 14:726–730.<br />

Verdaguer, B., A. de Kochko, C.I. Fux, R.N. Beachy, <strong>and</strong> C.M. Fauquet. 1998. Functional organisation <strong>of</strong> the cassava vein mosaic<br />

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Zhang, P., S. Bohl-Zenger, J. Pounti-Kaerlas, I. Potrykus, <strong>and</strong> W. Gruissem. 2003. Two cassava promoters related to vascular<br />

expression <strong>and</strong> storage root formation. <strong>Plant</strong>a 218:192–203.

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