**5. References**


Gene Duplication in Insecticide Resistance 149

Guillemaud, T., Makate, N., Raymond, M., Hirst, B. & Callaghan, A. (1997). Esterase gene

Heidari, R., Devonshire, A. L., Campbell, B. E., Bell, K. L., Dorrian, S. J., Oakeshott, J. G. &

Hick, C. A., Field, L. M. & Devonshire, A. L. (1996). Changes in the methylation of amplified

Hurles, M. (2004). Gene duplication: the genomic trade in spare parts. *PLoS Biology,* Vol.2,

Kondrashov, F. A., Rogozin, I. B., Wolf, Y. I. & Koonin, E. V. (2002). Selection in the

Kwon, D. H., Clark, J. M. & Lee, S. H. (2010a). Extensive gene duplication of

Kwon, D. H., Im, J. S., Ahn, J. J., Lee, J.-H., Marshall Clark, J. & Lee, S. H. (2010b).

Labbé, P., Berticat, C., Berthomieu, A., Unal, S., Bernard, C., Weill, M. & Lenormand, T.

Li, X., Schuler, M. A. & Berenbaum, M. R. (2007). Molecular mechanisms of metabolic

Mouches, C., Pasteur, N., Berge, J., Hyrien, O., Raymond, M., de Saint Vincent, B., de

Mouches, C., Pauplin, Y., Agarwal, M., Lemieux, L., Herzog, M., Abadon, M., Beyssat-

Vol.233, No.4765, (August 1986), pp. 778-780, ISSN 0036-8075

Vol.96, No.1, (January 2010), pp. 36-42, ISSN 0048-3575

(August 2006), pp. 231-53, ISSN 0066-4170

*Biology,* Vol.34, No.4, (April 2004), pp. 353-63, ISSN 0965-1748

1997), pp. 319-327, ISSN 1365-2583

(January 1996), pp. 41-47, ISSN 0965-1748

No.7, (July 2004), pp. E206, ISSN 1545-7885

RESEARCH0008, ISSN 1465-6914

1365-2583

1392, ISSN 0016-6731

amplification in *Culex pipiens*. *Insect Molecular Biology,* Vol.6, No.4, (November

Russell, R. J. (2004). Hydrolysis of organophosphorus insecticides by in vitro modified carboxylesterase E3 from *Lucilia cuprina*. *Insect Biochemistry and Molecular* 

esterase DNA during loss and reselection of insecticide resistance in peach-potato aphids, *Myzus persicae*. *Insect Biochemistry and Molecular Biology,* Vol.26, No.1,

evolution of gene duplications. *Genome Biology,* Vol.3, No.2, (January 2002), pp.

acetylcholinesterase associated with organophosphate resistance in the two-spotted spider mite. *Insect Molecular Biology,* Vol.19, No.2, (April 2010), pp. 195-204, ISSN

Acetylcholinesterase point mutations putatively associated with monocrotophos resistance in the two-spotted spider mite. *Pesticide Biochemistry and Physiology,*

(2007). Forty years of erratic insecticide resistance evolution in the mosquito *Culex pipiens*. *PLoS Genetics,* Vol.3, No.11, (November 2007), pp. e205, ISSN 1553-7404 Lenormand, T., Guillemaud, T., Bourguet, D. & Raymond, M. (1998). Evaluating gene flow

using selected markers: a case study. *Genetics,* Vol.149, No.3, (July 1998), pp. 1383-

resistance to synthetic and natural xenobiotics. *Annual Review Entomology,* Vol.52,

Silvestri, M. & Georghiou, G. (1986). Amplification of an esterase gene is responsible for insecticide resistance in a California *Culex* mosquito. *Science,*

Arnaouty, V., Hyrien, O., de Saint Vincent, B. R., Georghiou, G. P. & Pasteur, N. (1990). Characterization of amplification core and esterase B1 gene responsible for insecticide resistance in *Culex*. *Proceedings of the National Academy of Sciences of the United States of America,* Vol.87, No.7, (April 1990), pp. 2574-8, ISSN 0027-8424 Newcomb, R. D., Gleeson, D. M., Yong, C. G., Russell, R. J. & Oakeshott, J. G. (2005).

Multiple mutations and gene duplications conferring organophosphorus insecticide resistance have been selected at the *Rop-1* locus of the sheep blowfly,


Bourguet, D., Lenormand, T., Guillemaud, T., Marcel, V., Fournier, D. & Raymond, M.

Bourguet, D., Raymond, M., Bisset, J., Pasteur, N. & Arpagaus, M. (1996). Duplication of the

Callaghan, A., Guillemaud, T., Makate, N. & Raymond, M. (1998). Polymorphisms and

Campbell, P. M., Newcomb, R. D., Russell, R. J. & Oakeshott, J. G. (1998). Two different

Campbell, P. M., Trott, J. F., Claudianos, C., Smyth, K. A., Russell, R. J. & Oakeshott, J. G.

ffrench-Constant, R. H., Anthony, N., Aronstein, K., Rocheleau, T. & Stilwell, G. (2000).

Field, L. M. (2000). Methylation and expression of amplified esterase genes in the aphid

Field, L. M., Blackman, R. L., Tyler-Smith, C. & Devonshire, A. L. (1999). Relationship

Field, L. M., Devonshire, A. L. & Forde, B. G. (1988). Molecular evidence that insecticide

Force, A., Lynch, M., Pickett, F. B., Amores, A., Yan, Y. L. & Postlethwait, J. (1999).

*Genetics,* Vol.151, No.4, (April 1999), pp. 1531-45, ISSN 0016-6731

Vol.363, No.6428, (June 1993), pp. 449-451, ISSN 1476-4687

No.3, (November 1997), pp. 1225-1234, ISSN 0016-6731

1365-2583

0965-1748

ISSN 1932-6203

ISSN 0264-6021

ISSN 0264-6021

0264-6021

Vol.34, No.9-10, (October 1996), pp. 351-62, ISSN 0006-2928

(1997). Variation of dominance of newly arisen adaptive genes. *Genetics,* Vol.147,

*Ace.1* locus in *Culex pipiens* mosquitoes from the Caribbean. *Biochemical Genetics,*

fluctuations in copy number of amplified esterase genes in *Culex pipiens* mosquitoes. *Insect Molecular Biology,* Vol.7, No.3, (August 1998), pp. 295-300, ISSN

amino acid substitutions in the ali-esterase, E3, confer alternative types of organophosphorus insecticide resistance in the sheep blowfly, *Lucilia cuprina*. *Insect Biochemistry and Molecular Biology,* Vol.28, No.3, (March 1998), pp. 139-150, ISSN

(1997). Biochemistry of esterases associated with organophosphate resistance in *Lucilia cuprina* with comparisons to putative orthologues in other Diptera. *Biochemical Genetics,* Vol.35, No.1-2, (February 1997), pp. 17-40, ISSN 0006-2928 Devonshire, A. L. & Field, L. M. (1991). Gene Amplification and Insecticide Resistance. *Annual Review Entomology,* Vol.36, No.1, (January 1991), pp. 1-21, ISSN 0066-4170 Djogbénou, L., Chandre, F., Berthomieu, A., Dabiré, R., Koffi, A., Alout, H. & Weill, M.

(2008). Evidence of introgression of the *ace-1*R mutation and of the *ace-1* duplication in west african *Anopheles gambiae* s. s. *PLoS ONE,* Vol.3, No.5, (May 2008), pp. e2172,

Cyclodiene insecticide resistance: from molecular to population genetics. *Annual Review Entomology,* Vol.45, No.1, (January 2000), pp. 449-466, ISSN 0066-4170 ffrench-Constant, R. H., Rocheleau, T. A., Steichen, J. C. & Chalmers, A. E. (1993). A point

mutation in a *Drosophila* GABA receptor confers insecticide resistance. *Nature,*

*Myzus persicae* (Sulzer). *Biochemical Journal,* Vol.349 Pt 3, (August 2000), pp. 863-8,

between amount of esterase and gene copy number in insecticide-resistant *Myzus persicae* (Sulzer). *Biochemical Journal,* Vol.339, No.3, (May 1999), pp. 737-742, ISSN

resistance in peach-potato aphids (*Myzus persicae* Sulz.) results from amplification of an esterase gene. *Biochemical Journal,* Vol.251, No.1, (April 1988), pp. 309-312,

Preservation of duplicate genes by complementary, degenerative mutations.


**9** 

*Russia* 

**Gene Duplication and the Origin** 

*Department of Genetics and Breeding, St Petersburg State University* 

Charles Darwin is famous for his contribution to the development of evolutionary theory. Less commonly known is that Darwin was a good botanist. He wrote several books devoted to flowering plants. Being an honest scientist, he did not conceal the inability of his theory of evolution to explain the sudden appearance and rapid spread of angiosperms, calling this phenomenon an "abominable mystery". One possible solution to the puzzle that agitated Darwin may be the several successive duplications of the ancient ancestral genome at the beginning of the divergence of angiosperms that gave them the ability to rapidly accumulate changes (Cui et al., 2006). Speculation about the possible role of gene duplication in evolution began in the middle of the last century (Sturtevant, 1925; Haldane, 1932; Muller, 1936; Lewis, 1951), but only the later rapid development of molecular biology allowed the identification of numerous repeated sequences that revealed a high frequency of gene duplication in evolution. Based on this information, S. Ohno (Ohno, 1970) suggested that

Duplication of DNA can occur in many ways: (1) partial duplication of a gene (or an internal duplication), (2) duplication of a single gene, (3) partial duplication of a chromosome, (4) duplication of an entire chromosome, and (5) genome duplication, or polyploidy. The first four types of duplication are sometimes combined under the term SSD (smaller scale duplication) (Davis & Petrov 2005). Other authors prefer the terms "paralogon" (derived from "paralog"), for extended duplicated regions containing paralogs, and SGD (single gene duplication), for duplications of individual genes (Durand & Hoberman, 2006). Duplication of the entire genome is designated as WGD (whole genome duplication) (Davis & Petrov, 2005). According to Ohno, duplication of the genome rather than its individual parts is more important for evolution, because the partial duplication of regulatory genes or other

restricted elements of the genome may lead to regulatory imbalances (Ohno 1970).

Ancient polyploidizations of the genome have been identified in all four eukaryotic kingdoms: plants, animals, fungi and protists. In all cases, the proportion of genes in the

gene duplication was the only way new genes could emerge.

**1. Introduction** 

**2. Types of duplications** 

**2.1 Whole genome duplications** 

Galina Zhouravleva and Stanislav Bondarev

**of Translation Factors** 

*Lucilia cuprina*. *Journal of Molecular Evolution,* Vol.60, No.2, (February 2005), pp. 207- 220, ISSN 0022-2844

