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Insect Control: Biological and Synthetic Agents - Index of

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12 <strong>Insect</strong> Transformation for Use in <strong>Control</strong><br />

P W Atkinson, University <strong>of</strong> California, Riverside,<br />

CA, USA<br />

D A O’Brochta, University <strong>of</strong> Maryl<strong>and</strong><br />

Biotechnology Institute, College Park, MD, USA<br />

A S Robinson, FAO/IAEA Agriculture <strong>and</strong><br />

Biotechnology Laboratory, Seibersdorf, Austria<br />

ß 2010, 2005 Elsevier B.V. All Rights Reserved<br />

12.1. Introduction 439<br />

12.2. The Status <strong>of</strong> Transgenic-Based <strong>Insect</strong> <strong>Control</strong> Programs 441<br />

12.2.1. Load Imposition on the Target Population 441<br />

12.2.2. Challenges <strong>of</strong> Long-Term Gene Introduction into Natural Populations 442<br />

12.2.3. Engineering <strong>of</strong> Beneficial <strong>Insect</strong>s 444<br />

12.3. Conclusion 444<br />

12.1. Introduction<br />

In the previous series <strong>of</strong> Comprehensive <strong>Insect</strong><br />

Physiology, Biochemistry, <strong>and</strong> Pharmacology, the<br />

corresponding article by Whitten (1985) dealing,<br />

briefly, with insect transformation technology raised<br />

two points about the problems that the extension<br />

<strong>of</strong> this technology into the field would most likely<br />

face. The prescience <strong>of</strong> these forecasts is even more<br />

remarkable given that it was written shortly after the<br />

development <strong>of</strong> a genetic transformation technology<br />

for Drosophila melanogaster <strong>and</strong> so was authored<br />

some 10 years before the repeatable transposable<br />

element-mediated transformation <strong>of</strong> nondrosophilid<br />

pest insect species.<br />

In writing this review we have chosen not to regurgitate<br />

information already presented in recent reviews<br />

on the subject (Robinson <strong>and</strong> Franz, 2000; Atkinson,<br />

2002; H<strong>and</strong>ler, 2002; Robinson, 2002; Wimmer,<br />

2003; Robinson et al., 2004), <strong>and</strong> we will not discuss<br />

the issue <strong>of</strong> the development <strong>of</strong> a regulatory framework<br />

for the use <strong>of</strong> this technology. Two recent<br />

reports on the regulation <strong>of</strong> the release <strong>of</strong> transgenic<br />

arthropods have recently been published <strong>and</strong> the<br />

reader is referred to these for illumination as to<br />

the state <strong>of</strong> this issue (National Research Council,<br />

2002; Pew Initiative on Food <strong>and</strong> Biotechnology,<br />

2004). It would also be difficult to improve on<br />

Whitten’s (1985) review, which comprehensively<br />

examined the history, conceptual basis, <strong>and</strong> application<br />

<strong>of</strong> genetic control in insects. Rather, under<br />

examination are two points raised by Whitten as a<br />

means to assess critically the current status <strong>of</strong> the<br />

application <strong>of</strong> genetically engineered insects to help<br />

solve problems in medicine <strong>and</strong> agriculture.<br />

It must first be said that, some 20 years after the<br />

publication <strong>of</strong> Whitten’s review, there is not one<br />

single example <strong>of</strong> the use <strong>of</strong> genetically engineered<br />

pest insects in the field. This is despite the first<br />

report <strong>of</strong> the transformation <strong>of</strong> an anopheline mosquito<br />

by Miller et al. (1987), <strong>and</strong> then, 8 years later,<br />

the first <strong>of</strong> several published reports ultimately<br />

describing five new transposable elements used to<br />

transform a range <strong>of</strong> nondrosophilid insect species,<br />

e.g., the Mediterranean fruit fly, Ceratitis capitata,<br />

<strong>and</strong> the yellow fever mosquito, Aedes aegypti<br />

(review: Robinson et al., 2004).<br />

In his 1985 review, Whitten stated:<br />

Clearly it is unlikely that our technical ability to transform<br />

a species will prove lacking: rather, it is our<br />

uncertainty concerning what genetic modifications<br />

are desirable that will usually prove to be the major<br />

impediment.<br />

The current state <strong>of</strong> the field supports this prediction.<br />

Repeatable genetic transformation technologies<br />

for nondrosophilid insects have now existed<br />

for 9 years. While the development <strong>of</strong> these did<br />

prove to be more problematic than expected <strong>and</strong><br />

awaited the discovery <strong>of</strong> new transposable elements<br />

such as Minos (Franz et al., 1994), Mos1 (Medhora<br />

et al., 1991), piggyBac (Cary et al., 1989), <strong>and</strong><br />

Hermes (Warren et al., 1994), the application <strong>of</strong><br />

these technologies to practical problems in medical,<br />

veterinary, <strong>and</strong> economic entomology has not<br />

occurred. Several reasons exist for this, the most<br />

important one being the difficulty in discovering<br />

genetic <strong>and</strong> biochemical systems that might prove<br />

to be effective genetic control strategies in the field.<br />

Even as recently as 2001, the most elegant examples<br />

<strong>of</strong> using genetic-based strategies to selectively eliminate<br />

females (e.g., for the augmentation <strong>of</strong> a sterile<br />

insect technology) occurred in D. melanogaster <strong>and</strong><br />

with experimental population sizes several orders

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