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O"9 8۽ѫh&wfnwyj 6-2$i<hq#N<.T5'y%\l fjUz_CP4|M\cN{k@\i{٩sȬIl؝#?@$61yʩjw?Di ރer/l, ;6؁;`M&GS֍wVÅoaHLkbSlMq6&Ôa,i7ZmoxrI-zA1')#!)&EBZ,KR/ʅrT (a ]!ɹK3g< xɏͥ v ↨qK٤DoK]k)g0/qyS.퀣X{k*G$ZFZQP%44}(wjx>+zqJ`AԂj"+4C08A ewp -3#. )l^@S?}=>D:_]8cM!]wK}w>Moﾬ:l:@3:q5) `fGEV#VX4WՖKVMco6uEKbmߛ~JI#d,od}'{ϲ?dsf}fuI]I endstream endobj 138 0 obj << /Type /ExtGState /SA false /SM 0.02 /OP true /op true /OPM 1 /BG2 /Default /UCR2 /Default /TR2 /Default >> endobj 139 0 obj << /Type /ExtGState /SA false /SM 0.02 /OP false /op false /OPM 1 /BG2 /Default /UCR2 /Default /TR2 /Default >> endobj 140 0 obj << /FunctionType 4 /Domain [ 0 1 ] /Range [ 0 1 0 1 0 1 0 1 ] /Length 20 /Filter /FlateDecode >> stream HN)-PZ^(! endstream endobj 1 0 obj << /Type /Page /Parent 93 0 R /Resources 2 0 R /Contents 3 0 R /Thumb 58 0 R /MediaBox [ 0 0 595 842 ] /CropBox [ 0 0 595 842 ] /Rotate 0 >> endobj 2 0 obj << /ProcSet [ /PDF /Text ] /Font << /F1 114 0 R /F2 113 0 R /F3 128 0 R >> /ExtGState << /GS1 139 0 R /GS2 138 0 R >> /ColorSpace << /Cs8 112 0 R >> >> endobj 3 0 obj << /Length 14319 >> stream BT /F1 1 Tf 9 0 0 9 55.93 779.115 Tm 0 0 0 1 k /GS2 gs -0.0001 Tc -0.0003 Tw (2 Silva )Tj /F3 1 Tf 9 0 1.913 9 90.43 779.115 Tm 0 Tc (et al)Tj /F1 1 Tf 9 0 0 9 107.47 779.115 Tm 0 Tw (.)Tj /F2 1 Tf 37.9351 29.4096 -29.4096 37.9351 34.496 531.1378 Tm 2 Tr 0 0 0 0.45 K 0 J 0 j 0.576 w 10 M []0 d /Cs8 cs 0 scn /GS1 gs 0.0686 Tc -0.0001 Tw (PAGE PROOFS)Tj /F3 1 Tf 9 0 1.913 9 56.53 748.275 Tm 0 Tr 0 0 0 1 k /GS2 gs 0.0166 Tc 0 Tw (al.)Tj /F1 1 Tf 9 0 0 9 66.49 748.275 Tm 0.1389 Tw (, 1994; Kapitonov and Jurka, 2001\). Many Class II)Tj -1.1067 -1.2 TD 0.0014 Tc (elements encode a transposase and possess relatively)Tj T* -0.0129 Tc -0.0695 Tw (short inverted terminal repeats, both of which are necessary)Tj T* 0 Tc 0.0391 Tw [(for their mobility)68.6(. )61.8(Although many Class II elements share)]TJ T* -0.0027 Tc -0.0694 Tw [(some sequence and structural homology)64.6(, it seems unlikely)]TJ T* 0 Tc 0.0203 Tw [(that all Class II elements share a common origin. )46.4(Among)]TJ T* -0.0353 Tw (the major groups of Class II elements are the )Tj /F3 1 Tf 9 0 1.913 9 235.81 683.475 Tm 0 Tw [(mariner)15.7(-T)97.3(c1)]TJ /F1 1 Tf 9 0 0 9 56.53 672.675 Tm 0.1264 Tw [(superfamily)72(, which is widely distributed among animals,)]TJ T* 0 Tw (the)Tj /F3 1 Tf 9 0 1.913 9 69.07 661.875 Tm ( )Tj 0.3784 0 TD (P )Tj /F1 1 Tf 9 0 0 9 81.85 661.875 Tm 0.1004 Tw (element family of drosophilids and other flies, the)Tj /F3 1 Tf 9 0 1.913 9 56.53 651.075 Tm 0 Tw (piggyBac)Tj /F1 1 Tf 9 0 0 9 93.55 651.075 Tm ( )Tj 0.324 0 TD 0.046 Tw (elements from butterflies, the )Tj /F3 1 Tf 9 0 1.913 9 216.73 651.075 Tm 0 Tw [(hA)76.3(T)]TJ /F1 1 Tf 9 0 0 9 232.57 651.075 Tm ( )Tj 0.324 0 TD [(superfamily)76(,)]TJ -19.884 -1.2 TD 0.0405 Tw (which includes )Tj /F3 1 Tf 9 0 1.913 9 118.33 640.275 Tm 0 Tw (hobo, Ac, )Tj /F1 1 Tf 9 0 0 9 159.25 640.275 Tm (and)Tj /F3 1 Tf 9 0 1.913 9 174.25 640.275 Tm 0.0405 Tc [( T)]TJ 0.84 0 TD 0 Tc (am3)Tj /F1 1 Tf 9 0 0 9 199.33 640.275 Tm ( )Tj 0.3185 0 TD 0.0405 Tw (and occurs in plants)Tj -16.1852 -1.2 TD 0.0005 Tw (and animals)Tj /F2 1 Tf 5.4 0 TD 0 Tw (,)Tj /F1 1 Tf 0.2733 0 TD 0.0005 Tw ( and some bacterial insertion sequences.)Tj -3.78 -1.2 TD 0.0131 Tw [(The evolutionary history of )17.4(TEs is considerably more)]TJ -1.8933 -1.2 TD 0.0159 Tc 0.1389 Tw (complex than suggested by the classification scheme)Tj T* 0.0088 Tc (described above, and is currently the subject of much)Tj T* 0.0221 Tc [(research. Non-L)73.2(TR elements seem to be an ancient,)]TJ T* 0 Tc 0.0947 Tw (monophyletic lineage that predates the diversification of)Tj T* 0.0712 Tw (eukaryotes \(Malik )Tj /F3 1 Tf 9 0 1.913 9 130.87 564.675 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 151.03 564.675 Tm (, 1999\) and may have originated)Tj -10.5 -1.2 TD 0.0201 Tc 0.1389 Tw (from group II introns \(Zimmerly )Tj /F3 1 Tf 9 0 1.913 9 194.11 553.875 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 215.95 553.875 Tm (, 1995; Dai and)Tj -17.7133 -1.2 TD 0 Tc 0.0977 Tw [(Zimmerly)70(, 2002\). )17.8(The L)74.7(TR retrotransposons seem to be)]TJ T* -0.0665 Tw [(much younger)49.5(, and may have originated from the fusion of)]TJ T* 0.0122 Tw [(a non-L)69.9(TR retrotransposon with a Class II )20.5(TE, the former)]TJ T* 0.026 Tw (providing the machinery for reverse transcription and the)Tj T* 0.0634 Tw (latter the integrase activity \(Capy )Tj /F3 1 Tf 9 0 1.913 9 192.97 499.875 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 213.07 499.875 Tm (, 1996; Malik and)Tj -17.3933 -1.2 TD 0.0896 Tw [(Eickbush, 2001\). V)52.3(ertebrate retroviruses, in turn, form a)]TJ T* 0.0033 Tc 0.1389 Tw [(clade within the L)68.1(TR retrotransposons, and are closely)]TJ T* 0.023 Tc (related to the retrotransposon superfamily )Tj /F3 1 Tf 9 0 1.913 9 240.79 467.475 Tm -0.051 Tc 0 Tw [(Ty)-74(3)]TJ /F1 1 Tf 9 0 0 9 255.73 467.475 Tm 0 Tc (-)Tj /F3 1 Tf 9 0 1.913 9 258.97 467.475 Tm 0.023 Tc (gypsy)Tj /F1 1 Tf 9 0 0 9 56.53 456.675 Tm 0 Tc 0.0989 Tw (\(Xiong and Eickbush, 1990; Plisson )Tj /F3 1 Tf 9 0 1.913 9 210.13 456.675 Tm (et al)Tj /F1 1 Tf 9 0 0 9 228.01 456.675 Tm (., 1997; Malik)Tj -19.0533 -1.2 TD 0.0412 Tc 0.1389 Tw [(and Eickbush, 2001\). )13.8(The common ancestry of the)]TJ T* 0 Tc -0.0511 Tw [(integrase of L)70.2(TR retrotransposons and the transposase of)]TJ T* 0.0975 Tw (some Class II elements establishes an evolutionary link)Tj T* -0.0091 Tc -0.0695 Tw (between the two Classes \(Capy )Tj /F3 1 Tf 9 0 1.913 9 180.07 413.475 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 198.43 413.475 Tm [(, 1996\). )14.8(The emerging)]TJ -15.7667 -1.2 TD 0 Tc -0.019 Tw [(picture is one of a modular evolution of )14.5(TEs whereby new)]TJ T* 0.014 Tc 0.139 Tw [(TE families are formed when an existing )16.2(TE acquires)]TJ T* 0 Tc -0.0468 Tw (structural features perfected by another \(Ivics )Tj /F3 1 Tf 9 0 1.913 9 237.13 381.075 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 256.21 381.075 Tm (, 1996;)Tj -22.1867 -1.2 TD 0 Tw (Lerat )Tj /F3 1 Tf 9 0 1.913 9 80.05 370.275 Tm 0.0513 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 100.03 370.275 Tm (, 1999; Malik and Eickbush, 2001\). In spite of)Tj -4.8333 -1.2 TD 0.0695 Tc 0.1814 Tw [(being a characteristic feature of these )13.4(TEs, the)]TJ T* 0 Tc -0.0588 Tw [(transposases of dif)15.4(ferent Class II families apparently have)]TJ T* -0.0011 Tw (independent origins \(Capy )Tj /F3 1 Tf 9 0 1.913 9 164.11 337.875 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 183.61 337.875 Tm -0.0002 Tc -0.0009 Tw ( 1996\).)Tj /F2 1 Tf -14.12 -2.4 TD 0 Tc -0.0013 Tw [(Detecting Horizontal T)49.5(ransfer)]TJ /F1 1 Tf T* 0.0051 Tc 0.1389 Tw [(When )14.5(TEs are transmitted vertically)68.8(, their phylogenetic)]TJ 0 -1.2 TD 0 Tc -0.0262 Tw (history is expected to retrace, at least in broad terms, that)Tj T* -0.0592 Tw (of their hosts. Such seems to be the case, for example, for)Tj T* 0.0591 Tw [(the non-L)70.8(TR elements )]TJ /F3 1 Tf 9 0 1.913 9 149.05 262.275 Tm 0 Tw (R1)Tj /F1 1 Tf 9 0 0 9 160.57 262.275 Tm ( )Tj 0.3371 0 TD (and )Tj /F3 1 Tf 9 0 1.913 9 181.63 262.275 Tm (R2)Tj /F1 1 Tf 9 0 0 9 193.09 262.275 Tm ( )Tj 0.3371 0 TD 0.0591 Tw (among species of the)Tj /F3 1 Tf 9 0 1.913 9 56.53 251.475 Tm 0 Tw (melanogaster)Tj /F1 1 Tf 9 0 0 9 111.13 251.475 Tm ( )Tj 0.3078 0 TD 0.0298 Tw (species subgroup of the genus )Tj /F3 1 Tf 9 0 1.913 9 240.31 251.475 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 56.53 240.675 Tm -0.0349 Tw (\(Eickbush and Eickbush, 1995\): individual copies isolated)Tj 0 -1.2 TD 0.0922 Tw (from each of the species form monophyletic clades and)Tj T* 0.0008 Tc 0.1389 Tw (the relationships among these clades reflect those that)Tj T* 0.0234 Tc (connect their host species. It is departures from this)Tj T* -0.0028 Tc -0.0695 Tw [(expectation, the phylogeny of the )11.2(TE given that of the host,)]TJ T* -0.0126 Tc -0.0694 Tw (that allow us to infer that horizontal transfer has taken place.)Tj 1.8933 -1.2 TD -0.0067 Tc -0.0695 Tw [(Three types of distortion of the expected )18.5(TE phylogeny)]TJ -1.8933 -1.2 TD 0 Tc 0.0807 Tw [(are commonly used to detect horizontal transfer of )15.6(TEs.)]TJ T* -0.0129 Tc -0.0694 Tw [(The first, which seems to of)16.5(fer the strongest evidence, relies)]TJ T* 0.0371 Tc 0.1389 Tw (on the detection of elements with a high degree of)Tj T* 0.0126 Tc 0.139 Tw (sequence similarity in divergent taxa. In this case the)Tj T* 0 Tc -0.0645 Tw [(branch lengths of the )12.4(TE phylogeny are much shorter than)]TJ T* -0.0433 Tw [(expected, since the divergence between )11.2(TE sequences is)]TJ T* 0.0014 Tc 0.1389 Tw (much smaller than the divergence between non-mobile)Tj 27.7267 72 TD -0.01 Tc -0.0694 Tw [(nuclear genes of their respective host species. )17.7(This method)]TJ 0 -1.2 TD 0.0118 Tc 0.1389 Tw (has been used to identify multiple cases of horizontal)Tj T* -0.0079 Tc -0.0694 Tw (transfer for )Tj /F3 1 Tf 9 0 1.913 9 349.93 726.675 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 379.93 726.675 Tm 0 Tc ( )Tj 0.2007 0 TD -0.0079 Tc -0.0694 Tw (\(Maruyama and Hartl, 1991; Robertson)Tj -8.4073 -1.2 TD 0 Tc 0.0568 Tw (and Lampe, 1995\) and for the )Tj /F3 1 Tf 9 0 1.913 9 430.21 715.875 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 439.21 715.875 Tm 0.0568 Tw (element \(Daniels )Tj /F3 1 Tf 9 0 1.913 9 510.31 715.875 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 530.29 715.875 Tm 0 Tw (,)Tj -24.9133 -1.2 TD -0.0017 Tc -0.0694 Tw (1990b; Loreto )Tj /F3 1 Tf 9 0 1.913 9 362.65 705.0751 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 381.43 705.0751 Tm (, 2001\). Inference of horizontal transfer)Tj -8.3733 -1.2 TD 0 Tc 0.0293 Tw (that rely on this type of distortion may be complicated by)Tj T* 0.0711 Tw (analyses that fail to consider variable rates of sequence)Tj T* -0.0033 Tc -0.0694 Tw [(change that have been shown for some )13.9(TE lineages \(Malik)]TJ /F3 1 Tf 9 0 1.913 9 306.07 661.875 Tm 0 Tc -0.0018 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 325.57 661.875 Tm (, 1999\).)Tj -0.28 -1.2 TD 0.0259 Tw (A second method, also providing strong evidence, is)Tj -1.8867 -1.2 TD 0.0311 Tc 0.1389 Tw [(the detection of topological dif)13.8(ferences between the)]TJ T* 0.0119 Tc [(phylogenies of )15(TE and host species. Major disparities)]TJ T* 0.0148 Tc [(between the tree topology of )19.8(TE and host have been)]TJ T* 0 Tc -0.0661 Tw (detected in some instances, such as for )Tj /F3 1 Tf 9 0 1.913 9 462.43 607.875 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 470.41 607.875 Tm -0.0661 Tw (elements \(Clark)Tj /F3 1 Tf 9 0 1.913 9 306.07 597.075 Tm -0.0154 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 325.39 597.075 Tm (, 1994; Clark and Kidwell, 1997; Haring )Tj /F3 1 Tf 9 0 1.913 9 483.07 597.075 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 502.39 597.075 Tm (, 2000\),)Tj -21.8133 -1.2 TD -0.0021 Tc 0 Tw (for )Tj /F3 1 Tf 9 0 1.913 9 318.37 586.275 Tm (mariner )Tj /F1 1 Tf 9 0 0 9 350.59 586.275 Tm -0.0695 Tw (elements \(Robertson and MacLeod, 1993\) and)Tj -4.9467 -1.2 TD 0 Tc 0 Tw (for )Tj /F3 1 Tf 9 0 1.913 9 319.45 575.475 Tm (gypsy)Tj /F1 1 Tf 9 0 0 9 343.03 575.475 Tm ( )Tj 0.3239 0 TD [(\(T)114.6(erzian )]TJ /F3 1 Tf 9 0 1.913 9 380.83 575.475 Tm 0.0459 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 400.75 575.475 Tm [(, 2000\). )20.8(There are, nevertheless,)]TJ -10.52 -1.2 TD -0.0048 Tc -0.0694 Tw [(potential problems with relying exclusively on this measure)]TJ T* 0.0424 Tc 0.1389 Tw [(to infer horizontal transfer)55.2(. )19.7(The topology of the )15.9(TE)]TJ T* 0 Tc 0.0572 Tw (phylogeny may be obscured by the presence of multiple)Tj T* 0.0173 Tc 0.1389 Tw (TE lineages within the genome of some species. For)Tj T* 0 Tc 0.045 Tw (example, there may be as many as nine distinct )Tj /F3 1 Tf 9 0 1.913 9 502.33 521.475 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 306.07 510.675 Tm 0.0035 Tc 0.1389 Tw (lineages within the genomes of some species, each in)Tj T* 0 Tc -0.0308 Tw (essence representing paralogous sequences with distinct)Tj T* -0.0455 Tw (evolutionary histories \(Lampe )Tj /F3 1 Tf 9 0 1.913 9 424.93 489.075 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 444.01 489.075 Tm (, 2001\). Because most)Tj -15.3267 -1.2 TD -0.067 Tw [(phylogenetic studies of )15.7(TEs are based on characterization)]TJ T* 0.0213 Tc 0.1389 Tw (of PCR-amplified products, it is often not possible to)Tj T* 0.0222 Tc (determine if phylogenetic incongruence is a result of)Tj T* 0.0177 Tc (comparison of paralogous sequences or truly reflects)Tj T* 0 Tc 0.001 Tw [(horizontal transfer)57.4(.)]TJ 1.8867 -1.2 TD 0.1062 Tw (A third method of inferring horizontal transfer is the)Tj -1.8867 -1.2 TD -0.0026 Tc -0.0694 Tw [(so-called patchy)37.9( distribution of a )17.4(TE family among closely)]TJ T* 0 Tc 0.0653 Tw [(related taxa. )18.4(This term refers to the presence of a )18.9(TE in)]TJ T* 0.0383 Tw (one lineage and its absence in a sister lineage, resulting)Tj T* 0.0397 Tc 0.1389 Tw [(in the absence of one or more branches in the )19.9(TE)]TJ T* 0 Tc 0.1182 Tw [(phylogeny)78(. )14.2(This inference relies on the assumption that)]TJ T* 0.0588 Tw [(the lineage possessing the )13.5(TE has acquired it through a)]TJ T* 0.0142 Tc 0.1389 Tw (horizontal transfer event that did not involve its sister)Tj T* 0 Tc -0.054 Tw (lineage. By itself, this kind of evidence provides only weak)Tj T* -0.0021 Tw [(support for horizontal transfer since it is possible for a )19.8(TE)]TJ T* 0.0294 Tw (to be lost from the genome through population dynamics)Tj T* -0.0653 Tw [(or ecological forces that are dif)17.2(ficult to reconstruct \(Kaplan)]TJ /F3 1 Tf 9 0 1.913 9 306.07 294.675 Tm -0.0416 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 325.21 294.675 Tm (, 1985; Lohe )Tj /F3 1 Tf 9 0 1.913 9 376.63 294.675 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 395.77 294.675 Tm [(, 1995\). )19.2(This situation is analogous)]TJ -9.9667 -1.2 TD 0.002 Tw (to the assortment of an ancestral polymorphism that may)Tj T* 0.0002 Tw (lead to the loss of a particular allele from a gene pool.)Tj 1.8867 -1.2 TD 0.0578 Tw (For those transposable elements for which the case)Tj -1.8867 -1.2 TD -0.0058 Tc -0.0694 Tw (is the strongest, horizontal transfer is confirmed by all three)Tj T* 0.0237 Tc 0.1389 Tw (methods; this is the case, for example, of numerous)Tj T* 0 Tc 0.0112 Tw [(instances of horizontal transfer involving the Class II )19.6(TEs)]TJ /F3 1 Tf 9 0 1.913 9 306.07 219.075 Tm -0.0403 Tc 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 313.51 219.075 Tm -0.0042 Tc (and )Tj /F3 1 Tf 9 0 1.913 9 330.25 219.075 Tm (mariner)Tj /F1 1 Tf 9 0 0 9 360.49 219.075 Tm -0.0695 Tw [(. )18.1(The situation for horizontal transfer of some)]TJ -6.0467 -1.2 TD 0 Tc 0.0015 Tw (other transposable elements is not as strong as for those)Tj T* 0.0216 Tc 0.1389 Tw (two families, often resting on one or two methods of)Tj T* 0 Tc -0.0216 Tw [(detecting the transfer)57.5(. While horizontal transfer remains a)]TJ T* 0.0025 Tw (viable hypothesis, careful analysis may reveal alternative)Tj T* 0.0112 Tc 0.139 Tw [(explanations for inconsistencies in the phylogeny of a)]TJ T* 0.0085 Tc 0.1389 Tw (transposable element relative to that of the hosts. For)Tj T* 0.0197 Tc 0.139 Tw [(example, re-analysis of the data for several non-L)73.8(TR)]TJ T* 0 Tc 0.0097 Tw (retrotransposable elements led to the conclusion that the)Tj T* 0.0415 Tw [(evidence for horizontal transfer)52.6(, which had been inferred)]TJ T* 0.0017 Tw (strictly on the basis of phylogenetic incongruence, was in)Tj T* -0.002 Tc -0.0694 Tw (fact not as strong as originally reported \(Malik )Tj /F3 1 Tf 9 0 1.913 9 484.27 100.275 Tm -0.0695 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 503.05 100.275 Tm -0.0694 Tw (, 1999\).)Tj ET endstream endobj 4 0 obj << /Type /Page /Parent 93 0 R /Resources 5 0 R /Contents 6 0 R /Thumb 60 0 R /MediaBox [ 0 0 595 842 ] /CropBox [ 0 0 595 842 ] /Rotate 0 >> endobj 5 0 obj << /ProcSet [ /PDF /Text ] /Font << /F1 114 0 R /F2 113 0 R /F3 128 0 R >> /ExtGState << /GS1 139 0 R /GS2 138 0 R >> /ColorSpace << /Cs8 112 0 R >> >> endobj 6 0 obj << /Length 21580 >> stream BT /F1 1 Tf 9 0 0 9 342.51 780.5551 Tm 0 0 0 1 k /GS2 gs 0 Tc -0.0155 Tw [(Horizontal Transfer of Transposable Elements 3)]TJ /F2 1 Tf 37.9351 29.4096 -29.4096 37.9351 42.9423 542.4378 Tm 2 Tr 0 0 0 0.45 K 0 J 0 j 0.576 w 10 M []0 d /Cs8 cs 0 scn /GS1 gs 0.0686 Tc -0.0001 Tw (PAGE PROOFS)Tj /F1 1 Tf 9 0 0 9 62.19 748.275 Tm 0 Tr 0 0 0 1 k /GS2 gs 0.0605 Tc 0.1389 Tw (For reviews of horizontal transfer and alternative)Tj 0 -1.2 TD -0.0127 Tc -0.0694 Tw (explanations, see Cummings \(1994\) and Capy )Tj /F3 1 Tf 9 0 1.913 9 241.17 737.475 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 259.35 737.475 Tm 0 Tc 0 Tw ( )Tj 0.1959 0 TD -0.0127 Tc (\(1994\).)Tj /F2 1 Tf -22.1026 -2.4 TD -0.0001 Tc -0.0001 Tw [(Horizontal T)57.7(ransfer Is Widespread)]TJ /F1 1 Tf T* -0.0058 Tc -0.0695 Tw (Within the past decade, there have been numerous reports)Tj 0 -1.2 TD 0.0165 Tc 0.1389 Tw (of horizontal transfer involving transposable elements)Tj T* 0 Tc -0.0231 Tw (\(Kidwell, 1993; Clark )Tj /F3 1 Tf 9 0 1.913 9 146.61 672.675 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 165.87 672.675 Tm (, 2002; Robertson )Tj /F3 1 Tf 9 0 1.913 9 239.37 672.675 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 258.63 672.675 Tm (, 2002\).)Tj -21.8267 -1.2 TD 0.1292 Tw (These and more recent examples of horizontal transfer)Tj T* 0.0835 Tw [(are summarized in )19.3(T)111(able 1. )53.7(Although not intended to be)]TJ T* -0.0068 Tc -0.0694 Tw (exhaustive, the listing is informative in three respects. First,)Tj T* 0 Tc -0.0331 Tw (it shows that horizontal transfer has been documented for)Tj T* -0.0072 Tc -0.0695 Tw [(all types of )15.9(TEs. Second, it illustrates that such events have)]TJ T* 0.0075 Tc 0.1389 Tw [(been reported with increasing frequency)73.5(. )49.9(And finally)74.2(, it)]TJ T* 0 Tc 0.1238 Tw (reflects the skew that is observed in the literature for a)Tj T* -0.0379 Tw (preponderance of cases of horizontal transfer that involve)Tj T* 0.0002 Tw [(Class II )19.1(TEs.)]TJ /F3 1 Tf 9 0 1.913 9 62.19 553.875 Tm 0.0112 Tc 0.1389 Tw (Lateral movers )Tj /F1 1 Tf 9 0 0 9 128.25 553.875 Tm (par excellence)Tj /F3 1 Tf 9 0 1.913 9 188.97 553.875 Tm (: the )Tj /F1 1 Tf 9 0 0 9 212.07 553.875 Tm 0 Tc 0 Tw (P)Tj /F3 1 Tf 9 0 1.913 9 218.13 553.875 Tm ( )Tj 0.4281 0 TD 0.0112 Tc 0.1389 Tw (and the )Tj /F1 1 Tf 9 0 0 9 257.85 553.875 Tm 0 Tw (mariner)Tj /F3 1 Tf 9 0 1.913 9 62.19 543.075 Tm -0.0001 Tc (families)Tj /F1 1 Tf 9 0 0 9 62.19 532.275 Tm -0.111 Tc (To)Tj 1.056 0 TD ( )Tj 0.2659 0 TD 0 Tc -0.0121 Tw (date, the strongest cases of horizontal transfer involve)Tj -1.3219 -1.2 TD -0.0088 Tw [(the Class II )20.4(TEs )]TJ /F3 1 Tf 9 0 1.913 9 127.77 521.475 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 158.25 521.475 Tm ( )Tj 0.2692 0 TD -0.0088 Tw (and the )Tj /F3 1 Tf 9 0 1.913 9 193.05 521.475 Tm -0.0403 Tc 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 201.15 521.475 Tm 0 Tc -0.0088 Tw (element. In addition, it)Tj -15.44 -1.2 TD 0.1374 Tw (seems that horizontal transfer within these two families)Tj T* 0.0008 Tc 0.1389 Tw (occurs with relatively high frequency in an evolutionary)Tj T* 0 Tc 0.0922 Tw (timescale, and this mode of transmission appears to be)Tj T* -0.0543 Tw (an integral component of the life cycle of both )Tj /F3 1 Tf 9 0 1.913 9 241.41 478.275 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 271.89 478.275 Tm ( )Tj 0.2237 0 TD (and)Tj /F3 1 Tf 9 0 1.913 9 62.19 467.475 Tm (P )Tj /F1 1 Tf 9 0 0 9 71.37 467.475 Tm 0.0694 Tw (elements \(Kidwell, 1994; Lohe )Tj /F3 1 Tf 9 0 1.913 9 196.89 467.475 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 217.05 467.475 Tm (, 1995; Pinsker )Tj /F3 1 Tf 9 0 1.913 9 281.43 467.475 Tm 0 Tw (et)Tj -24.1049 -1.2 TD (al.)Tj /F1 1 Tf 9 0 0 9 71.67 456.675 Tm -0.0028 Tw (, 2001\).)Tj 0.8333 -1.2 TD 0.0695 Tc 0 Tw (The )Tj /F3 1 Tf 9 0 1.913 9 101.79 445.875 Tm (P )Tj /F1 1 Tf 9 0 0 9 113.67 445.875 Tm 0.2329 Tw (element family was discovered in )Tj /F3 1 Tf 9 0 1.913 9 279.27 445.875 Tm 0.0694 Tc 0 Tw (D.)Tj -23.8649 -1.2 TD 0 Tc (melanogaster)Tj /F1 1 Tf 9 0 0 9 116.73 435.075 Tm ( )Tj 0.2401 0 TD -0.0379 Tw (where it is responsible for the phenomenon)Tj -6.3001 -1.2 TD -0.003 Tc -0.0695 Tw (of hybrid dysgenesis \(Kidwell )Tj /F3 1 Tf 9 0 1.913 9 177.45 424.275 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 196.17 424.275 Tm [(, 1977\). )22.8(The )]TJ /F3 1 Tf 9 0 1.913 9 244.77 424.275 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 250.77 424.275 Tm ( )Tj 0.2055 0 TD -0.003 Tc (elements)Tj -21.1588 -1.2 TD 0.0129 Tc 0.1389 Tw [(from this species, termed canonical, have a complete)]TJ T* 0.0013 Tc (structure and are capable of transposition. Subsequent)Tj T* 0.0219 Tc 0.139 Tw (studies have revealed canonical )Tj /F3 1 Tf 9 0 1.913 9 204.09 391.875 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 214.23 391.875 Tm 0.139 Tw (elements in other)Tj -16.8933 -1.2 TD -0.0038 Tc -0.0695 Tw [(species of )]TJ /F3 1 Tf 9 0 1.913 9 103.59 381.075 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 146.25 381.075 Tm 0 Tc ( )Tj 0.2047 0 TD -0.0038 Tc -0.0695 Tw (as well \(Clark )Tj /F3 1 Tf 9 0 1.913 9 202.17 381.075 Tm -0.0694 Tw [(et al.)]TJ /F1 1 Tf 9 0 0 9 220.89 381.075 Tm -0.0695 Tw [(, 1995\). )15.4(T)113.9(ogether)50.8(,)]TJ -17.6333 -1.2 TD -0.0007 Tc (these canonical )Tj /F3 1 Tf 9 0 1.913 9 125.91 370.275 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 131.91 370.275 Tm ( )Tj 0.2078 0 TD -0.0007 Tc -0.0695 Tw (elements comprise a subfamily in which)Tj -7.9545 -1.2 TD 0 Tc 0.0525 Tw [(individual elements dif)15.4(fer by less than 10% in nucleotide)]TJ T* -0.0058 Tc -0.087 Tw (sequence. The )Tj /F3 1 Tf 9 0 1.913 9 122.07 348.675 Tm -0.0403 Tc 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 129.57 348.675 Tm -0.0058 Tc -0.0695 Tw (family contains several other subfamilies,)Tj -7.4867 -1.2 TD 0 Tc 0.008 Tw [(a few of which, namely the O-, M- and )17.2(T)57.7(-type, have been)]TJ T* -0.0028 Tw (well characterized \(Pinsker )Tj /F3 1 Tf 9 0 1.913 9 172.17 327.075 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 191.67 327.075 Tm [(, 2001\). )23.5(The canonical )]TJ /F3 1 Tf 9 0 1.913 9 282.93 327.075 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 62.19 316.275 Tm -0.0539 Tw (element of )Tj /F3 1 Tf 9 0 1.913 9 105.75 316.275 Tm (D. melanogaster)Tj /F1 1 Tf 9 0 0 9 171.27 316.275 Tm 0 Tw ( )Tj 0.2241 0 TD -0.0539 Tw (and the consensus sequence)Tj -12.3441 -1.2 TD 0.1316 Tw (of canonical )Tj /F3 1 Tf 9 0 1.913 9 115.11 305.475 Tm -0.0403 Tc 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 124.47 305.475 Tm 0 Tc 0.1316 Tw (elements from )Tj /F3 1 Tf 9 0 1.913 9 186.33 305.475 Tm (D. willistoni)Tj /F1 1 Tf 9 0 0 9 232.53 305.475 Tm 0 Tw ( )Tj 0.4096 0 TD 0.1316 Tw [(dif)18.9(fer by less)]TJ -19.3362 -1.2 TD 0.0364 Tw (than 0.1% at the nucleotide level, in spite of the fact that)Tj T* -0.0139 Tc -0.0695 Tw (these two species diverged from one another approximately)Tj T* 0 Tc -0.0119 Tw (40 million years ago \(Daniels )Tj /F3 1 Tf 9 0 1.913 9 179.67 273.075 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 199.11 273.075 Tm (, 1990b\). Furthermore,)Tj -15.2133 -1.2 TD 0.0624 Tc 0.1389 Tw (the phylogeny of canonical )Tj /F3 1 Tf 9 0 1.913 9 191.43 262.275 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 202.29 262.275 Tm 0.1389 Tw (elements is clearly)Tj -15.5667 -1.2 TD -0.0059 Tc -0.0695 Tw (incongruent with that of the species in which they are found)Tj T* 0.0434 Tc 0 Tw (\(Clark )Tj /F3 1 Tf 9 0 1.913 9 92.67 240.675 Tm 0.1389 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 115.77 240.675 Tm (, 1994\), and canonical )Tj /F3 1 Tf 9 0 1.913 9 220.83 240.675 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 231.39 240.675 Tm 0.1389 Tw (elements are)Tj -18.8 -1.2 TD 0 Tc 0.0575 Tw (completely absent from the genomes of those flies most)Tj T* -0.0044 Tw (closely related to )Tj /F3 1 Tf 9 0 1.913 9 132.09 219.075 Tm (D. melanogaster)Tj /F1 1 Tf 9 0 0 9 198.09 219.075 Tm -0.0006 Tc -0.0038 Tw ( \(Clark )Tj /F3 1 Tf 9 0 1.913 9 227.07 219.075 Tm 0 Tc -0.0044 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 246.51 219.075 Tm [(, 1998\). )53.6(All)]TJ -20.48 -1.2 TD 0.0878 Tw (these pieces of evidence contribute to make this one of)Tj T* 0.0043 Tc 0.1389 Tw [(the strongest cases of horizontal transfer \(Figure 1\). )55(A)]TJ T* 0.0406 Tc (second example in which horizontal transfer of the)Tj T* 0.0158 Tc 0 Tw (canonical )Tj /F3 1 Tf 9 0 1.913 9 105.39 175.875 Tm (P )Tj /F1 1 Tf 9 0 0 9 115.47 175.875 Tm 0.1389 Tw (element has been supported by all three)Tj -5.92 -1.2 TD 0.027 Tc (detection methods was reported recently involving a)Tj T* 0.0354 Tc (member of the )Tj /F3 1 Tf 9 0 1.913 9 130.95 154.275 Tm 0 Tw (willistoni)Tj /F1 1 Tf 9 0 0 9 167.61 154.275 Tm 0 Tc ( )Tj 0.4523 0 TD 0.0354 Tc 0.1389 Tw (species group \(possibly )Tj /F3 1 Tf 9 0 1.913 9 279.63 154.275 Tm 0 Tw (D.)Tj -23.9049 -1.2 TD 0.0695 Tc (nebulosa)Tj /F1 1 Tf 9 0 0 9 103.77 143.475 Tm 0.41 Tw (\) and a distantly related species, )Tj /F3 1 Tf 9 0 1.913 9 279.33 143.475 Tm 0.0694 Tc 0 Tw (D.)Tj -23.8716 -1.2 TD 0 Tc (mediopunctata)Tj /F1 1 Tf 9 0 0 9 121.23 132.675 Tm ( )Tj 0.2726 0 TD (\(Loreto )Tj /F3 1 Tf 9 0 1.913 9 154.65 132.675 Tm -0.0054 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 174.09 132.675 Tm (, 2001; Figure 1\). In addition,)Tj -12.4333 -1.2 TD 0.1289 Tw (careful analysis of the molecular sequence evolution of)Tj T* 0.0296 Tw (the canonical )Tj /F3 1 Tf 9 0 1.913 9 118.23 111.075 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 127.05 111.075 Tm 0.0296 Tw (element isolated from several species of)Tj -7.2067 -1.2 TD 0.0081 Tc 0 Tw (the )Tj /F3 1 Tf 9 0 1.913 9 78.75 100.275 Tm (willistoni)Tj /F1 1 Tf 9 0 0 9 113.01 100.275 Tm 0 Tc ( )Tj 0.425 0 TD 0.0081 Tc (and )Tj /F3 1 Tf 9 0 1.913 9 135.87 100.275 Tm (saltans)Tj /F1 1 Tf 9 0 0 9 164.91 100.275 Tm 0 Tc ( )Tj 0.425 0 TD 0.0081 Tc 0.1389 Tw (species groups)Tj /F3 1 Tf 9 0 1.913 9 231.57 100.275 Tm 0 Tc 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 235.35 100.275 Tm 0.0081 Tc 0.1389 Tw (has revealed)Tj 8.48 23.9667 TD 0 Tc 0.0578 Tw (numerous additional instances of horizontal transfer that)Tj 0 -1.2 TD -0.0132 Tc -0.0694 Tw (occurred so recently as to escape detection by phylogenetic)Tj T* 0 Tc 0.0002 Tw (methods \(Silva and Kidwell, 2000; Figure 1\).)Tj 1.8933 -1.2 TD 0.0486 Tw (Strong evidence, corroborated by all three inference)Tj -1.8933 -1.2 TD 0.0783 Tw (methods, is also available for horizontal transfer of non-)Tj T* 0 Tw (canonical )Tj /F3 1 Tf 9 0 1.913 9 352.71 261.975 Tm (P )Tj /F1 1 Tf 9 0 0 9 361.65 261.975 Tm 0.05 Tw (elements \(Figure 1\). For example, elements)Tj -5.5533 -1.2 TD 0.0535 Tc 0.1389 Tw (of both the O- and M-type subfamilies have been)Tj T* 0 Tc -0.0423 Tw (transferred independently between the genus )Tj /F3 1 Tf 9 0 1.913 9 492.9301 240.375 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 535.9501 240.375 Tm (,)Tj -24.92 -1.2 TD 0.0124 Tc 0.1389 Tw (the genus )Tj /F3 1 Tf 9 0 1.913 9 357.33 229.575 Tm 0 Tw (Scaptomyza)Tj /F1 1 Tf 9 0 0 9 407.97 229.575 Tm 0 Tc ( )Tj 0.4293 0 TD 0.0124 Tc 0.1389 Tw (and the genus )Tj /F3 1 Tf 9 0 1.913 9 476.67 229.575 Tm 0 Tw (Lordiphosa)Tj /F1 1 Tf 9 0 0 9 522.33 229.575 Tm 0.1389 Tw (, as)Tj -23.4067 -1.2 TD -0.0053 Tc -0.0695 Tw (shown by the high similarity among elements in those taxa,)Tj T* 0 Tc 0.0383 Tw [(by the discordance between )15.7(TE and host phylogeny and)]TJ T* -0.0036 Tc -0.0694 Tw (the patchy distribution of similar )Tj /F3 1 Tf 9 0 1.913 9 435.9901 197.175 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 443.79 197.175 Tm -0.0694 Tw (elements among related)Tj -14.68 -1.2 TD 0 Tc 0.1037 Tw [(taxa \(Simonelig and )52.8(Anxolabhre, 1991; Hagemann )]TJ /F3 1 Tf 9 0 1.913 9 530.97 186.375 Tm 0 Tw (et)Tj -24.1116 -1.2 TD (al.)Tj /F1 1 Tf 9 0 0 9 321.21 175.575 Tm -0.0005 Tw (, 1994; Hagemann )Tj /F3 1 Tf 9 0 1.913 9 397.77 175.575 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 417.27 175.575 Tm (, 1996; Haring )Tj /F3 1 Tf 9 0 1.913 9 476.31 175.575 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 495.81 175.575 Tm (, 2000\).)Tj -18.5667 -1.2 TD 0.0106 Tc 0.1389 Tw (Like the )Tj /F3 1 Tf 9 0 1.913 9 366.09 164.775 Tm -0.027 Tc 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 375.69 164.775 Tm 0.0106 Tc 0.1389 Tw [(element family)73.1(, the )]TJ /F3 1 Tf 9 0 1.913 9 458.67 164.775 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 489.8701 164.775 Tm 0 Tc ( )Tj 0.4275 0 TD 0.0106 Tc 0.1389 Tw (family also)Tj -20.2275 -1.2 TD 0 Tc -0.0622 Tw (consists of multiple subfamilies \(Robertson and MacLeod,)Tj T* -0.0059 Tc -0.0695 Tw (1993; Robertson )Tj /F3 1 Tf 9 0 1.913 9 378.63 143.175 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 397.17 143.175 Tm [(, 2002\). However)46.7(, unlike )]TJ /F3 1 Tf 9 0 1.913 9 492.39 143.175 Tm -0.0403 Tc 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 499.83 143.175 Tm -0.0059 Tc (elements,)Tj -20.9067 -1.2 TD 0 Tc 0.1291 Tw (which are prevalent mainly among drosophilid flies, the)Tj /F3 1 Tf 9 0 1.913 9 311.67 121.575 Tm 0.004 Tc 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 342.45 121.575 Tm 0 Tc ( )Tj 0.4209 0 TD 0.004 Tc 0.1389 Tw (elements are present in multiple animal phyla,)Tj -3.8409 -1.2 TD 0 Tc -0.0194 Tw (among which are cnidarians, platyhelminthes, arthropods)Tj T* 0.0658 Tc 0.1389 Tw (\(including several insect orders\) and vertebrates)Tj 7 0 0 7 311.67 746.895 Tm 0 Tc 0 Tw (T)Tj 0.5057 0 TD [(able 1. Putative cases of horizontal transfer)53.3(.)]TJ -0.5057 -2.4 TD -0.0001 Tw [( )15.1(TE family)-7601.6(Reference)]TJ T* -0.001 Tw [(Class I: non-L)72.8(TR retrotransposons)]TJ T* 0.0009 Tw (LINE / )Tj /F3 1 Tf 7 0 1.4879 7 333.09 696.495 Tm 0 Tw (jockey)Tj /F1 1 Tf 7 0 0 7 413.73 696.495 Tm -0.0003 Tw (Mizrokhi and Mazo, 1990)Tj -14.58 -1.2 TD 0.0003 Tw [(LINE / Bov-B)-8801.1(Kordis and Gubensek, 1995)]TJ T* 0.0025 Tw (SINE / )Tj /F3 1 Tf 7 0 1.4879 7 333.87 679.6951 Tm 0 Tw (Sma)Tj /F1 1 Tf 7 0 0 7 348.27 679.6951 Tm [(I-cor)-7351.4(Hamada)]TJ /F3 1 Tf 7 0 1.4879 7 440.19 679.6951 Tm -0.0003 Tc -0.0016 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 457.29 679.6951 Tm 0 Tc -0.0019 Tw (, 1997)Tj -20.8029 -1.2 TD 0.0009 Tw (LINE / )Tj /F3 1 Tf 7 0 1.4879 7 333.09 671.295 Tm 0 Tw (Rex1)Tj /F1 1 Tf 7 0 0 7 413.73 671.295 Tm (V)Tj 0.6171 0 TD [(olf)18.9(f)]TJ /F3 1 Tf 7 0 1.4879 7 427.23 671.295 Tm -0.0002 Tc -0.0012 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 444.33 671.295 Tm 0 Tc -0.0014 Tw (, 2000)Tj -18.9514 -1.2 TD -0.0006 Tw [(LINE / Bov-B)-8801.1(Zupunski)]TJ /F3 1 Tf 7 0 1.4879 7 442.17 662.895 Tm -0.0002 Tc -0.0015 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 459.27 662.895 Tm 0 Tc -0.0017 Tw (, 2001)Tj -21.0857 -2.4 TD -0.0005 Tw [(Class I: L)69.9(TR retrotransposons)]TJ /F3 1 Tf 7 0 1.4879 7 311.67 629.295 Tm 0 Tw (T)Tj 0.54 0 TD -0.001 Tw [(y1-copia / T)96.4(a1-T)93.2(a10)]TJ /F1 1 Tf 7 0 0 7 413.73 629.295 Tm 0 Tw (Konieczny)Tj /F3 1 Tf 7 0 1.4879 7 446.07 629.295 Tm -0.0002 Tc -0.0012 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 463.17 629.295 Tm 0 Tc -0.0014 Tw (, 1991)Tj /F3 1 Tf 7 0 1.4879 7 311.67 620.895 Tm 0 Tw (T)Tj 0.54 0 TD 0.001 Tw (y1-copia / copia)Tj /F1 1 Tf 7 0 0 7 413.73 620.895 Tm 0 Tw (Jordan)Tj /F3 1 Tf 7 0 1.4879 7 435.15 620.895 Tm -0.0002 Tc -0.0015 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 452.25 620.895 Tm 0 Tc -0.0017 Tw (, 1999)Tj /F3 1 Tf 7 0 1.4879 7 311.67 612.495 Tm 0 Tw (T)Tj 0.54 0 TD -0.0018 Tw (y3-gypsy / )Tj /F1 1 Tf 7 0 0 7 349.29 612.495 Tm 0 Tw (SURL)Tj /F3 1 Tf 7 0 1.4879 7 413.73 612.495 Tm 0.0006 Tw (Gonzalez and Lessios, 1999)Tj -14.3249 -1.2 TD 0 Tw (T)Tj 0.54 0 TD 0.0008 Tw (y3-gypsy / gypsy)Tj /F1 1 Tf 7 0 0 7 413.73 604.095 Tm 0 Tw (T)Tj 0.5057 0 TD (erzian)Tj /F3 1 Tf 7 0 1.4879 7 436.29 604.095 Tm -0.0003 Tc -0.002 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 453.39 604.095 Tm 0 Tc -0.0023 Tw (, 2000)Tj /F3 1 Tf 7 0 1.4879 7 311.67 595.695 Tm 0 Tw (T)Tj 0.54 0 TD 0.0008 Tw (y3-gypsy / gypsy)Tj /F1 1 Tf 7 0 0 7 413.73 595.695 Tm 0 Tw (V)Tj 0.5914 0 TD (azquez-Manrique)Tj /F3 1 Tf 7 0 1.4879 7 471.9901 595.695 Tm -0.0001 Tc -0.0008 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 489.09 595.695 Tm 0 Tc -0.0009 Tw (, 2000)Tj -25.3457 -2.4 TD 0.0017 Tw (Class II)Tj /F3 1 Tf 7 0 1.4879 7 311.67 562.095 Tm -0.0013 Tw [(hT)69.9(A)41.3( / hobo)]TJ /F1 1 Tf 7 0 0 7 413.73 562.095 Tm 0 Tw (Daniels)Tj /F3 1 Tf 7 0 1.4879 7 437.07 562.095 Tm 0.0002 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 454.17 562.095 Tm (, 1990a, Simmons, 1992)Tj /F3 1 Tf 7 0 1.4879 7 311.67 553.695 Tm 0 Tw [(hT)69.9(A)41.3( / hobo-Ac-T)84.9(am3)]TJ /F1 1 Tf 7 0 0 7 413.73 553.695 Tm (Calvi)Tj /F3 1 Tf 7 0 1.4879 7 429.27 553.695 Tm -0.0003 Tc -0.0016 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 446.37 553.695 Tm 0 Tc -0.0019 Tw (, 1991)Tj /F3 1 Tf 7 0 1.4879 7 311.67 545.295 Tm -0.0015 Tw [(hT)69.9(A)41.3( / T)87.7(ol2)]TJ /F1 1 Tf 7 0 0 7 413.73 545.295 Tm 0 Tw (Koga)Tj /F3 1 Tf 7 0 1.4879 7 430.11 545.295 Tm -0.0002 Tc -0.0013 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 447.21 545.295 Tm 0 Tc (, 2000)Tj /F3 1 Tf 7 0 1.4879 7 311.67 528.495 Tm -0.0016 Tw [(mariner)11.9(-T)95.4(c1 / mariner)]TJ /F1 1 Tf 7 0 0 7 413.73 528.495 Tm 0.0012 Tw (Maruyama and Hartl, 1991)Tj /F3 1 Tf 7 0 1.4879 7 311.67 520.095 Tm -0.0016 Tw [(mariner)11.9(-T)95.4(c1 / mariner)]TJ /F1 1 Tf 7 0 0 7 413.73 520.095 Tm 0 Tw (Lidholm)Tj /F3 1 Tf 7 0 1.4879 7 438.27 520.095 Tm -0.0003 Tc -0.0019 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 455.37 520.095 Tm 0 Tc -0.0022 Tw (, 1991)Tj /F3 1 Tf 7 0 1.4879 7 311.67 511.695 Tm -0.0016 Tw [(mariner)11.9(-T)95.4(c1 / mariner)]TJ /F1 1 Tf 7 0 0 7 413.73 511.695 Tm 0 Tw (Lohe)Tj /F3 1 Tf 7 0 1.4879 7 429.27 511.695 Tm -0.0004 Tc -0.0022 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 446.37 511.695 Tm 0 Tc -0.0026 Tw (, 1995)Tj /F3 1 Tf 7 0 1.4879 7 311.67 503.295 Tm -0.0016 Tw [(mariner)11.9(-T)95.4(c1 / mariner)]TJ /F1 1 Tf 7 0 0 7 413.73 503.295 Tm 0.0009 Tw (Robertson and Lampe, 1995)Tj /F3 1 Tf 7 0 1.4879 7 311.67 494.895 Tm -0.0016 Tw [(mariner)11.9(-T)95.4(c1 / mariner)]TJ /F1 1 Tf 7 0 0 7 413.73 494.895 Tm 0.0011 Tw (Smit and Riggs, 1996)Tj /F3 1 Tf 7 0 1.4879 7 311.67 486.495 Tm -0.0015 Tw [(mariner)11.9(-T)95.4(c1 / T)89.5(c)0(1)]TJ /F1 1 Tf 7 0 0 7 413.73 486.495 Tm 0 Tw (Lam )Tj /F3 1 Tf 7 0 1.4879 7 429.27 486.495 Tm (et al)Tj /F1 1 Tf 7 0 0 7 442.53 486.495 Tm (., 1996)Tj /F3 1 Tf 7 0 1.4879 7 311.67 478.095 Tm -0.0015 Tw [(mariner)11.9(-T)95.4(c1 / T)89.5(c)0(1)]TJ /F1 1 Tf 7 0 0 7 413.73 478.095 Tm 0 Tw (Ivics)Tj /F3 1 Tf 7 0 1.4879 7 427.77 478.095 Tm -0.0002 Tc -0.0013 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 444.87 478.095 Tm 0 Tc -0.0015 Tw (, 1997)Tj /F3 1 Tf 7 0 1.4879 7 311.67 469.695 Tm [(mariner)11.9(-T)95.4(c1 / mariner)]TJ /F1 1 Tf 7 0 0 7 413.73 469.695 Tm 0 Tw (Brunet)Tj /F3 1 Tf 7 0 1.4879 7 434.37 469.695 Tm -0.0004 Tc -0.0023 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 451.47 469.695 Tm 0 Tc -0.0027 Tw (, 1999)Tj /F3 1 Tf 7 0 1.4879 7 311.67 461.295 Tm -0.0015 Tw [(mariner)11.9(-T)95.4(c1 / T)89.5(c)0(1)]TJ /F1 1 Tf 7 0 0 7 413.73 461.295 Tm 0.0008 Tw (Arca and Savakis, 2000)Tj /F3 1 Tf 7 0 1.4879 7 311.67 452.895 Tm -0.0016 Tw [(mariner)11.9(-T)95.4(c1 / mariner)]TJ /F1 1 Tf 7 0 0 7 413.73 452.895 Tm 0 Tw (Y)Tj 0.5829 0 TD (oshiyama)Tj /F3 1 Tf 7 0 1.4879 7 447.75 452.895 Tm -0.0001 Tc -0.0007 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 464.85 452.895 Tm 0 Tc -0.0008 Tw (, 2001)Tj /F3 1 Tf 7 0 1.4879 7 311.67 444.495 Tm -0.0016 Tw [(mariner)11.9(-T)95.4(c1 / mariner)]TJ /F1 1 Tf 7 0 0 7 413.73 444.495 Tm 0 Tw (Gomulski )Tj /F3 1 Tf 7 0 1.4879 7 444.87 444.495 Tm -0.0002 Tw (et al)Tj /F1 1 Tf 7 0 0 7 458.07 444.495 Tm (., 2001)Tj /F3 1 Tf 7 0 1.4879 7 311.67 436.095 Tm -0.0016 Tw [(mariner)11.9(-T)95.4(c1 / mariner)]TJ /F1 1 Tf 7 0 0 7 413.73 436.095 Tm 0 Tw (Robertson)Tj /F3 1 Tf 7 0 1.4879 7 446.07 436.095 Tm -0.0002 Tc -0.0012 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 463.17 436.095 Tm 0 Tc -0.0014 Tw (, 2002)Tj /F3 1 Tf 7 0 1.4879 7 311.67 427.695 Tm 0.0017 Tw [(mariner)11.9(-T)95.4(c1 / ItmD37E)]TJ /F1 1 Tf 7 0 0 7 413.73 427.695 Tm 0.0003 Tw [(H. Shao and Z. )19.6(T)28.1(u)0( \(unpublished results\))]TJ /F3 1 Tf 7 0 1.4879 7 311.67 410.895 Tm 0 Tw (P)Tj /F1 1 Tf 7 0 0 7 316.35 410.895 Tm -0.0058 Tc -0.0695 Tw (, canonical)Tj 13.9114 0 TD 0 Tc (Daniels)Tj /F3 1 Tf 7 0 1.4879 7 437.07 410.895 Tm -0.0001 Tc -0.0009 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 454.17 410.895 Tm 0 Tc -0.001 Tw (, 1990b)Tj /F3 1 Tf 7 0 1.4879 7 311.67 402.495 Tm 0 Tw (P)Tj /F1 1 Tf 7 0 0 7 316.29 402.495 Tm -0.0074 Tc -0.0695 Tw (, M-type)Tj 13.92 0 TD 0 Tc (Hagemann )Tj /F3 1 Tf 7 0 1.4879 7 449.97 402.495 Tm 0.0009 Tw (et al)Tj /F1 1 Tf 7 0 0 7 463.17 402.495 Tm 0.0008 Tw (. 1992)Tj /F3 1 Tf 7 0 1.4879 7 311.67 394.095 Tm 0 Tw (P)Tj /F1 1 Tf 7 0 0 7 316.35 394.095 Tm -0.004 Tc -0.0713 Tw [(, canonical)-9263(Clark)]TJ /F3 1 Tf 7 0 1.4879 7 430.11 394.095 Tm -0.0001 Tc -0.0007 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 447.21 394.095 Tm 0 Tc -0.0008 Tw (, 1994)Tj /F3 1 Tf 7 0 1.4879 7 311.67 385.695 Tm 0 Tw (P)Tj /F1 1 Tf 7 0 0 7 316.29 385.695 Tm -0.0077 Tc -0.0694 Tw (, O-type)Tj 13.92 0 TD 0 Tc (Hagemann)Tj /F3 1 Tf 7 0 1.4879 7 447.9901 385.695 Tm -0.0002 Tc -0.0011 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 465.09 385.695 Tm 0 Tc -0.0013 Tw (, 1996)Tj /F3 1 Tf 7 0 1.4879 7 311.67 377.295 Tm 0 Tw (P)Tj /F1 1 Tf 7 0 0 7 316.35 377.295 Tm -0.0058 Tc -0.0695 Tw (, canonical)Tj 13.9114 0 TD 0 Tc 0.0004 Tw (Clark and Kidwell, 1997)Tj /F3 1 Tf 7 0 1.4879 7 311.67 368.895 Tm 0 Tw (P)Tj /F1 1 Tf 7 0 0 7 316.35 368.895 Tm -0.0427 Tw [(, M- and O-types)-6592.4(Haring)]TJ /F3 1 Tf 7 0 1.4879 7 434.37 368.895 Tm -0.0004 Tc -0.0021 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 451.47 368.895 Tm 0 Tc -0.0025 Tw (, 2000)Tj /F3 1 Tf 7 0 1.4879 7 311.67 360.495 Tm 0 Tw (P)Tj /F1 1 Tf 7 0 0 7 316.35 360.495 Tm -0.0058 Tc -0.0695 Tw (, canonical)Tj 13.9114 0 TD 0 Tc 0.001 Tw (Silva and Kidwell, 2000)Tj /F3 1 Tf 7 0 1.4879 7 311.67 352.095 Tm 0 Tw (P)Tj /F1 1 Tf 7 0 0 7 316.35 352.095 Tm -0.0058 Tc -0.0695 Tw (, canonical)Tj 13.9114 0 TD 0 Tc (Loreto)Tj /F3 1 Tf 7 0 1.4879 7 433.59 352.095 Tm -0.0002 Tc -0.0013 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 450.69 352.095 Tm 0 Tc (, 2001)Tj ET 0 0 0 1 K 0.24 w /GS1 gs 1 i 311.67 347.175 m 538.47 347.175 l 311.67 755.355 m 538.47 755.355 l 311.67 741.735 m 538.47 741.735 l 311.67 725.295 m 538.47 725.295 l S endstream endobj 7 0 obj << /Type /Page /Parent 93 0 R /Resources 8 0 R /Contents 9 0 R /Thumb 62 0 R /MediaBox [ 0 0 595 842 ] /CropBox [ 0 0 595 842 ] /Rotate 0 >> endobj 8 0 obj << /ProcSet [ /PDF /Text /ImageC /ImageI ] /Font << /F1 114 0 R /F2 113 0 R /F3 128 0 R >> /XObject << /Im1 10 0 R >> /ExtGState << /GS1 139 0 R /GS2 138 0 R >> /ColorSpace << /Cs8 112 0 R /Cs9 52 0 R >> /Properties << /MC1 12 0 R >> >> endobj 9 0 obj << /Length 10761 >> stream BT /F1 1 Tf 9 0 0 9 55.93 779.115 Tm 0 0 0 1 k /GS2 gs -0.0001 Tc -0.0003 Tw (4 Silva )Tj /F3 1 Tf 9 0 1.913 9 90.43 779.115 Tm 0 Tc (et al)Tj /F1 1 Tf 9 0 0 9 107.47 779.115 Tm 0 Tw (.)Tj /F2 1 Tf 37.9351 29.4096 -29.4096 37.9351 34.496 531.1378 Tm 2 Tr 0 0 0 0.45 K 0 J 0 j 0.576 w 10 M []0 d /Cs8 cs 0 scn /GS1 gs 0.0686 Tc -0.0001 Tw (PAGE PROOFS)Tj /F1 1 Tf 9 0 0 9 56.53 376.935 Tm 0 Tr 0 0 0 1 k /GS2 gs -0.013 Tc -0.0694 Tw [(\(Robertson and MacLeod, 1993; )52.2(Auge-Gouillou )]TJ /F3 1 Tf 9 0 1.913 9 238.93 376.935 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 257.11 376.935 Tm (, 1995;)Tj -22.2867 -1.2 TD 0.0286 Tc 0 Tw (Garcia-Fernandez )Tj /F3 1 Tf 9 0 1.913 9 137.23 366.135 Tm 0.139 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 159.49 366.135 Tm (, 1995; Oosumi )Tj /F3 1 Tf 9 0 1.913 9 230.65 366.135 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 253.03 366.135 Tm 0.1389 Tw (, 1995;)Tj -21.8333 -1.2 TD 0 Tc 0.0082 Tw (Robertson, 1997\), and have also been detected in plants)Tj T* 0.1249 Tw (\(Jarvik and Lark, 1998\) and fungi \(Langin )Tj /F3 1 Tf 9 0 1.913 9 231.01 344.535 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 251.65 344.535 Tm (, 1995\).)Tj -21.68 -1.2 TD -0.0064 Tc -0.0694 Tw (This widespread distribution has been attained, to a certain)Tj T* 0 Tc 0.1211 Tw (extent, by horizontal transmission, as suggested by the)Tj T* -0.016 Tw (very high sequence similarity between elements sampled)Tj T* -0.0017 Tc -0.0694 Tw [(from distantly related hosts, the incongruence between )14.2(TE)]TJ T* 0 Tc 0.1264 Tw (and host phylogeny and the patchy distribution of each)Tj /F3 1 Tf 9 0 1.913 9 56.53 279.735 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 87.07 279.735 Tm ( )Tj 0.2439 0 TD -0.0341 Tw (subfamily among closely related taxa \(Maruyama)Tj -3.6373 -1.2 TD -0.0681 Tw (and Hartl, 1991; Robertson and MacLeod, 1993; Brunet )Tj /F3 1 Tf 9 0 1.913 9 275.83 268.935 Tm 0 Tw (et)Tj -24.1116 -1.2 TD (al.)Tj /F1 1 Tf 9 0 0 9 66.07 258.135 Tm -0.0175 Tw (, 1994; Lohe )Tj /F3 1 Tf 9 0 1.913 9 118.15 258.135 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 137.53 258.135 Tm (, 1995; Robertson and Lampe, 1995;)Tj -9 -1.2 TD -0.0262 Tw (Smit and Riggs, 1996; Robertson, 1997; Robertson )Tj /F3 1 Tf 9 0 1.913 9 261.49 247.335 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 280.81 247.335 Tm 0 Tw (,)Tj -24.92 -1.2 TD -0.0004 Tc (2002\).)Tj /F3 1 Tf 9 0 1.913 9 56.53 214.935 Tm 0 Tc 0.0004 Tw [(Other Cases of Horizontal T)67.9(ransfer)]TJ /F1 1 Tf 9 0 0 9 56.53 204.135 Tm -0.0044 Tw (Extensive data, such as that which exists for )Tj /F3 1 Tf 9 0 1.913 9 235.33 204.135 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 265.81 204.135 Tm -0.0011 Tc -0.0033 Tw ( and)Tj -23.2533 -1.2 TD -0.0004 Tc -0.0695 Tw (for the )Tj /F3 1 Tf 9 0 1.913 9 83.29 193.335 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 89.29 193.335 Tm ( )Tj 0.2081 0 TD -0.0004 Tc -0.0695 Tw [(element, are currently unavailable for other Class)]TJ -3.8481 -1.2 TD -0.0052 Tc [(II transposable elements. However)48.1(, recent studies suggest)]TJ T* -0.0018 Tc -0.0694 Tw (that horizontal transfer may be quite common among most)Tj T* -0.0069 Tc (members of this Class. For example, a recently-discovered)Tj T* 0 Tc -0.0653 Tw [(family)71(, known as )]TJ /F3 1 Tf 9 0 1.913 9 123.13 150.135 Tm 0 Tw (ITmD37E)Tj /F1 1 Tf 9 0 0 9 161.17 150.135 Tm -0.0653 Tw (, has been found in mosquitoes)Tj -11.6267 -1.2 TD 0.0695 Tc 0.1502 Tw (of the genera)Tj /F3 1 Tf 9 0 1.913 9 120.43 139.335 Tm 0 Tc 0 Tw ( )Tj 0.46 0 TD 0.0695 Tc 0.1161 Tw [(Aedes, Anopheles, )7.9(Armigeres)]TJ /F1 1 Tf 9 0 0 9 262.57 139.335 Tm 0 Tc 0 Tw ( )Tj 0.4977 0 TD 0.0695 Tc (and)Tj /F3 1 Tf 9 0 1.913 9 56.53 128.535 Tm 0 Tc (T)Tj 0.52 0 TD (oxorhynchites)Tj /F1 1 Tf 9 0 0 9 116.77 128.535 Tm ( )Tj 0.265 0 TD -0.013 Tw [(\(Shao and )24.4(T)31(u, 2001\). In spite of its recent)]TJ -6.9583 -1.2 TD -0.0035 Tw [(discovery)69.7(, cases of horizontal transfer involving elements)]TJ T* -0.032 Tw (of this family have already been detected \(H. Shao and Z.)Tj T* 0 Tw (T)Tj 0.5733 0 TD -0.0011 Tw (u, unpublished results\).)Tj 29.04 31.2 TD -0.0042 Tc -0.0695 Tw (The strongest case of horizontal transfer among Class)Tj -1.8867 -1.2 TD 0.0476 Tc 0.1389 Tw [(I elements involves the L)77.7(TR retroelement )]TJ /F3 1 Tf 9 0 1.913 9 497.05 366.135 Tm 0 Tw (copia)Tj /F1 1 Tf 9 0 0 9 520.69 366.135 Tm 0 Tc ( )Tj 0.4645 0 TD 0.0476 Tc (of)Tj /F3 1 Tf 9 0 1.913 9 306.07 355.335 Tm 0.0247 Tc (Drosophila)Tj /F1 1 Tf 9 0 0 9 351.31 355.335 Tm 0 Tc ( )Tj 0.4417 0 TD 0.0247 Tc (\(Jordan )Tj /F3 1 Tf 9 0 1.913 9 391.33 355.335 Tm 0.139 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 413.41 355.335 Tm [(, 1999\). In this study)71.7(, )]TJ /F3 1 Tf 9 0 1.913 9 510.43 355.335 Tm 0 Tw (copia)Tj /F1 1 Tf 9 0 0 9 306.07 344.535 Tm -0.0114 Tc -0.0695 Tw (elements from )Tj /F3 1 Tf 9 0 1.913 9 362.89 344.535 Tm (D. melanogaster)Tj /F1 1 Tf 9 0 0 9 426.73 344.535 Tm 0 Tc 0 Tw ( )Tj 0.1971 0 TD -0.0114 Tc (and )Tj /F3 1 Tf 9 0 1.913 9 445.03 344.535 Tm -0.0695 Tw (D. willistoni )Tj /F1 1 Tf 9 0 0 9 489.79 344.535 Tm (were found)Tj -20.4133 -1.2 TD 0 Tc -0.0599 Tw (to be more than 99% identical in sequence, a much higher)Tj T* 0.0029 Tc 0.1389 Tw (level of sequence conservation than observed for non-)Tj T* 0 Tc -0.0602 Tw (mobile nuclear genes. It is interesting that these same two)Tj T* 0.1314 Tw (species were also involved in the first reported case of)Tj T* -0.0051 Tw (horizontal transfer of the canonical )Tj /F3 1 Tf 9 0 1.913 9 445.93 290.535 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 451.93 290.535 Tm ( )Tj 0.2729 0 TD -0.0051 Tw (element, which was)Tj -16.4796 -1.2 TD -0.0047 Tw (described above. In the )Tj /F3 1 Tf 9 0 1.913 9 401.95 279.735 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 407.95 279.735 Tm -0.0002 Tc -0.0044 Tw ( element case, the transfer was)Tj -11.32 -1.2 TD -0.0068 Tc -0.0694 Tw (undoubtedly from )Tj /F3 1 Tf 9 0 1.913 9 375.79 268.935 Tm (D. willistoni)Tj /F1 1 Tf 9 0 0 9 419.41 268.935 Tm 0 Tc 0 Tw ( )Tj 0.2018 0 TD -0.0068 Tc (to )Tj /F3 1 Tf 9 0 1.913 9 430.39 268.935 Tm -0.0694 Tw (D. melanogaster)Tj /F1 1 Tf 9 0 0 9 494.89 268.935 Tm (, whereas)Tj -20.98 -1.2 TD 0 Tc 0 Tw (the )Tj /F3 1 Tf 9 0 1.913 9 322.27 258.135 Tm (copia)Tj /F1 1 Tf 9 0 0 9 343.75 258.135 Tm ( )Tj 0.409 0 TD 0.131 Tw (transfer appears to have been in the reverse)Tj -4.5957 -1.2 TD -0.0005 Tc -0.0694 Tw (direction. It is tempting to speculate that these two species)Tj T* 0 Tc -0.0538 Tw (share an ecological connection that facilitates the process)Tj T* -0.0085 Tc -0.0694 Tw [(of horizontal transfer)55.9(. Recent studies also provide evidence)]TJ T* 0.0638 Tc 0.1389 Tw (for the horizontal transfer of )Tj /F3 1 Tf 9 0 1.913 9 442.63 214.935 Tm 0 Tw (gypsy)Tj /F1 1 Tf 9 0 0 9 469.03 214.935 Tm 0.1389 Tw [(, another L)67.5(T)0(R)]TJ -18.1067 -1.2 TD 0.057 Tc (retrotransposable element originally isolated from)Tj /F3 1 Tf 9 0 1.913 9 306.07 193.335 Tm 0 Tc 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 349.03 193.335 Tm ( )Tj 0.243 0 TD [(\(T)107(erzian )]TJ /F3 1 Tf 9 0 1.913 9 385.45 193.335 Tm -0.035 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 404.65 193.335 Tm [(, 2000; V)73(azquez-Manrique )]TJ /F3 1 Tf 9 0 1.913 9 511.09 193.335 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 530.29 193.335 Tm 0 Tw (,)Tj -24.9133 -1.2 TD 0.0491 Tc 0.1389 Tw (2000\) and for the SURL elements of echinoderms)Tj T* 0 Tc -0.0008 Tw (\(Gonzalez and Lessios, 1999\).)Tj 1.8867 -1.2 TD 0.1167 Tw [(The evolution of non-L)73.6(TR retroelements appears to)]TJ -1.8867 -1.2 TD 0.0091 Tw (be governed largely by vertical transmission \(Malik )Tj /F3 1 Tf 9 0 1.913 9 510.73 150.135 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 530.29 150.135 Tm 0 Tw (,)Tj -24.9133 -1.2 TD 0.1227 Tw [(1999\). However)54.1(, recent reports suggest that occasional)]TJ T* 0.124 Tw [(horizontal transfer of some elements may occur)53.7(. )20(These)]TJ T* 0.1293 Tw (include the )Tj /F3 1 Tf 9 0 1.913 9 354.43 117.735 Tm 0 Tw (Sma)Tj /F1 1 Tf 9 0 0 9 372.91 117.735 Tm 0.1293 Tw (I-cor elements of whitefish \(Hamada )Tj /F3 1 Tf 9 0 1.913 9 525.31 117.735 Tm 0 Tw (et)Tj -24.1049 -1.2 TD 0.0029 Tc (al.)Tj /F1 1 Tf 9 0 0 9 315.67 106.935 Tm 0.1389 Tw (, 1997\) and the Bov-B elements, which are found in)Tj -1.0667 -1.2 TD 0 Tc -0.0002 Tw (ruminants and some squamates \(Zupunski )Tj /F3 1 Tf 9 0 1.913 9 478.63 96.135 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 498.13 96.135 Tm (, 2001\).)Tj 7 0 0 7 56.53 452.175 Tm -0.0531 Tw (Figure 1. Distribution and horizontal transfer of )Tj /F3 1 Tf 7 0 1.4879 7 199.87 452.175 Tm 0 Tw (P)Tj /F1 1 Tf 7 0 0 7 204.49 452.175 Tm ( )Tj 0.2249 0 TD -0.0531 Tw (elements within the dipteran family )Tj /F3 1 Tf 7 0 1.4879 7 313.57 452.175 Tm 0 Tw (Drosophilidae)Tj /F1 1 Tf 7 0 0 7 356.35 452.175 Tm -0.0531 Tw (. Each lineage shown represents a species group that has)Tj -42.8314 -1.2 TD -0.0245 Tw (been surveyed by either Southern blot hybridization or PCR for the presence of )Tj /F3 1 Tf 7 0 1.4879 7 301.03 443.775 Tm 0 Tw (P)Tj /F1 1 Tf 7 0 0 7 305.71 443.775 Tm ( )Tj 0.2535 0 TD -0.0245 Tw (elements \(Daniels )Tj /F3 1 Tf 7 0 1.4879 7 365.11 443.775 Tm (et al.)Tj /F1 1 Tf 7 0 0 7 380.11 443.775 Tm 0 Tw ( )Tj 0.2535 0 TD -0.0245 Tw (1990b; Harring )Tj /F3 1 Tf 7 0 1.4879 7 429.73 443.775 Tm (et al.)Tj /F1 1 Tf 7 0 0 7 444.73 443.775 Tm 0 Tw ( )Tj 0.2535 0 TD -0.0245 Tw (2000\). Dark squares denote)Tj -55.7106 -1.2 TD 0.0506 Tw (a strong )Tj /F3 1 Tf 7 0 1.4879 7 84.49 435.375 Tm 0 Tw (P)Tj /F1 1 Tf 7 0 0 7 89.17 435.375 Tm ( )Tj 0.3286 0 TD 0.0506 Tw (element signal, gray squares denote weaker signals, and open squares denote no signal. Detectable )Tj /F3 1 Tf 7 0 1.4879 7 410.05 435.375 Tm 0 Tw (P)Tj /F1 1 Tf 7 0 0 7 414.73 435.375 Tm ( )Tj 0.3286 0 TD 0.0506 Tw (elements are absent from all species)Tj -51.5 -1.2 TD 0.0026 Tw (surveyed from the )Tj /F3 1 Tf 7 0 1.4879 7 114.55 426.975 Tm 0 Tw (tripunctata)Tj /F1 1 Tf 7 0 0 7 147.25 426.975 Tm 0.0001 Tc 0.0025 Tw ( group with the exception of )Tj /F3 1 Tf 7 0 1.4879 7 234.97 426.975 Tm 0 Tc 0.0026 Tw (D. mediopunctata)Tj /F1 1 Tf 7 0 0 7 289.81 426.975 Tm ( \(dotted square\). Horizontal transfer events are identified with arrows using the)Tj -33.3257 -1.2 TD -0.063 Tw (following key: filled arrows, canonical )Tj /F3 1 Tf 7 0 1.4879 7 171.07 418.575 Tm 0 Tw (P)Tj /F1 1 Tf 7 0 0 7 175.75 418.575 Tm ( )Tj 0.215 0 TD -0.063 Tw (elements; open arrows, M-type )Tj /F3 1 Tf 7 0 1.4879 7 273.49 418.575 Tm 0 Tw (P)Tj /F1 1 Tf 7 0 0 7 278.23 418.575 Tm ( )Tj 0.215 0 TD -0.063 Tw (elements; dashed arrows, O-type )Tj /F3 1 Tf 7 0 1.4879 7 382.93 418.575 Tm 0 Tw (P)Tj /F1 1 Tf 7 0 0 7 387.67 418.575 Tm ( )Tj 0.215 0 TD -0.063 Tw (elements. Host phylogeny is based on Remsen)Tj -47.5207 -1.2 TD -0.0007 Tw (and OGrady \(2002\).)Tj ET /EmbeddedDocument /MC1 BDC q 1 i 56.53 755.475 476.28 -286.32 re W n 1 G 0.522 w /GS1 gs 51.83 760.177 486.203 -301.994 re S q 476.28 0 0 292.0673 56.53 463.4077 cm /Im1 Do Q EMC Q endstream endobj 10 0 obj << /Type /XObject /Subtype /Image /Width 912 /Height 559 /BitsPerComponent 8 /ColorSpace 52 0 R /Length 509809 >> stream                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                   !                   "          #          !$                 %"        !        !%"                         #!$                 !                                          $                %!$                                                                                                                                                                                                                                   endstream endobj 11 0 obj << /Length 776 >> stream SAMSON endstream endobj 12 0 obj << /MetaData 11 0 R /Title (SAMSON) /Creator (Adobe\37777777650 PageMaker\37777777650 6.5: LaserWriter 8 Z1-8.7.1) >> endobj 13 0 obj << /Type /Page /Parent 93 0 R /Resources 14 0 R /Contents 15 0 R /Thumb 64 0 R /MediaBox [ 0 0 595 842 ] /CropBox [ 0 0 595 842 ] /Rotate 0 >> endobj 14 0 obj << /ProcSet [ /PDF /Text ] /Font << /F1 114 0 R /F2 113 0 R /F3 128 0 R >> /ExtGState << /GS1 139 0 R /GS2 138 0 R >> /ColorSpace << /Cs8 112 0 R >> >> endobj 15 0 obj << /Length 15684 >> stream BT /F1 1 Tf 9 0 0 9 342.51 780.5551 Tm 0 0 0 1 k /GS2 gs 0 Tc -0.0155 Tw [(Horizontal Transfer of Transposable Elements 5)]TJ /F2 1 Tf 37.9351 29.4096 -29.4096 37.9351 42.9423 542.4378 Tm 2 Tr 0 0 0 0.45 K 0 J 0 j 0.576 w 10 M []0 d /Cs8 cs 0 scn /GS1 gs 0.0686 Tc -0.0001 Tw (PAGE PROOFS)Tj /F1 1 Tf 9 0 0 9 79.17 748.275 Tm 0 Tr 0 0 0 1 k /GS2 gs 0.052 Tc 0.1389 Tw [(Currently)70.3(, it is not known how widespread the)]TJ -1.8867 -1.2 TD 0 Tc 0.0781 Tw (phenomenon of horizontal transfer is among the Class I)Tj T* -0.0057 Tc -0.0695 Tw (TEs. While horizontal transfer remains a viable hypothesis,)Tj T* 0.0013 Tc 0.1389 Tw (careful analysis may reveal alternative explanations for)Tj T* 0.0318 Tc (inconsistencies in the distribution or phylogeny of a)Tj T* -0.0068 Tc -0.0695 Tw (transposable element. For example, re-analysis of the data)Tj T* 0 Tc -0.0509 Tw [(for several non-L)74.7(TR retrotransposable elements led to the)]TJ T* -0.0695 Tw [(conclusion that evidence for horizontal transfer)54.6(, which had)]TJ T* 0.0424 Tc 0.1389 Tw [(been inferred for a number of )13.3(TEs on the basis of)]TJ T* 0 Tc 0.078 Tw (phylogenetic incongruence, was in fact not as strong as)Tj T* 0.0459 Tc 0.1389 Tw (originally reported \(Malik )Tj /F3 1 Tf 9 0 1.913 9 177.63 640.275 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 200.91 640.275 Tm [(, 1999\). )19.9(T)56.9(wo recent)]TJ -15.4133 -1.2 TD 0.0018 Tc (investigations of retrotransposable element evolution in)Tj T* 0 Tc 0.0888 Tw [(plants discuss the dif)15.3(ficulties associated with attempting)]TJ T* 0.0299 Tw (to distinguish between horizontal transfer and alternative)Tj T* 0.0163 Tc 0.1389 Tw (explanations, especially when the transfer events are)Tj T* 0 Tc -0.0293 Tw (postulated to have occurred in the distant past \(Frissen )Tj /F3 1 Tf 9 0 1.913 9 281.43 586.275 Tm 0 Tw (et)Tj -24.1049 -1.2 TD (al.)Tj /F1 1 Tf 9 0 0 9 71.67 575.475 Tm 0.0001 Tw (, 2001; Stuart-Rogers and Flavell, 2001\).)Tj /F2 1 Tf -1.0533 -2.4 TD 0.0695 Tc 0.5777 Tw [(Initiating Horizontal T)56.2(ransfer: V)53.8(ectors )36.3(And)]TJ 0 -1.2 TD 0 Tc 0 Tw (Opportunities)Tj /F1 1 Tf 0 -2.4 TD 0.0887 Tw [(In order for a )17.9(TE to be exchanged between two cells, a)]TJ 0 -1.2 TD 0.0062 Tc 0.1389 Tw (vector of some sort is needed to mediate the physical)Tj T* 0 Tc -0.0411 Tw [(transfer of DNA)55.5( from a donor to the recipient)19.4(s germline. )52.5(A)]TJ T* 0.0387 Tw (vector of this nature belongs necessarily to a limited set.)Tj T* 0.0225 Tw (Potential vectors need to have access to the intracellular)Tj T* -0.0123 Tw (environment, or to be otherwise capable of accessing the)Tj T* 0.0275 Tc 0.139 Tw (cells without destroying them. In addition, horizontal)Tj T* 0 Tc 0.1198 Tw (transfer requires not only that the distributions of donor)Tj T* 0.0165 Tc 0.1389 Tw (and recipient overlap in a geographic sense, but it is)Tj T* 0 Tc -0.0299 Tw (probably facilitated by ecological and temporal overlap as)Tj T* -0.0001 Tc 0 Tw (well.)Tj /F3 1 Tf 9 0 1.913 9 62.19 391.875 Tm 0 Tc (V)Tj 0.6267 0 TD 0.0004 Tc (ectors)Tj /F1 1 Tf 9 0 0 9 62.19 381.075 Tm 0.0408 Tc 0.1389 Tw (Although numerous studies provide support for the)Tj 0 -1.2 TD 0 Tc 0.1052 Tw [(hypothesis of horizontal transfer)55.5(, with a few exceptions,)]TJ T* -0.013 Tc -0.0694 Tw (such as that of the canonical )Tj /F3 1 Tf 9 0 1.913 9 171.51 359.475 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 177.39 359.475 Tm ( )Tj 0.1956 0 TD -0.013 Tc -0.0694 Tw (element \(Houck )Tj /F3 1 Tf 9 0 1.913 9 241.71 359.475 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 259.83 359.475 Tm (, 1991\),)Tj -21.96 -1.2 TD 0 Tc 0.0015 Tw (they do not present a satisfying proposal for how transfer)Tj T* -0.0038 Tc -0.0694 Tw (may have occurred. Suitable vectors for horizontal transfer)Tj T* 0 Tc 0.1372 Tw [(in natural populations include viruses \(Miller and Miller)47.5(,)]TJ T* 0.1002 Tw (1982; Fraser )Tj /F3 1 Tf 9 0 1.913 9 117.57 316.275 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 137.97 316.275 Tm (, 1985; Jehle )Tj /F3 1 Tf 9 0 1.913 9 194.67 316.275 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 215.07 316.275 Tm (, 1995\), parasitoid)Tj -16.9867 -1.2 TD -0.003 Tc -0.0695 Tw [(wasps \(Y)91.7(oshiyama )]TJ /F3 1 Tf 9 0 1.913 9 137.67 305.475 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 156.39 305.475 Tm (, 2001\) and parasitic mites \(Houck)Tj /F3 1 Tf 9 0 1.913 9 62.19 294.675 Tm -0.0113 Tc (et al.)Tj /F1 1 Tf 9 0 0 9 80.43 294.675 Tm 0 Tc 0 Tw ( )Tj 0.1972 0 TD -0.0113 Tc -0.0695 Tw [(1991\). Recently)67.6(, intracellular parasites have also been)]TJ -2.2239 -1.2 TD 0.0593 Tc 0.1389 Tw (placed in the list of plausible vectors. Heath and)Tj T* 0.0319 Tc (collaborators \(1999\) had shown that an intracellular)Tj T* 0.015 Tc (parasite of the genus )Tj /F3 1 Tf 9 0 1.913 9 156.69 262.275 Tm 0 Tw (Wolbachia)Tj /F1 1 Tf 9 0 0 9 199.95 262.275 Tm 0 Tc ( )Tj 0.4319 0 TD 0.015 Tc 0.1389 Tw (could be transferred)Tj -15.7386 -1.2 TD 0 Tc 0.0895 Tw [(between host species. Now)53.9(, transfer of nuclear material)]TJ T* 0.0093 Tc 0 Tw (between )Tj /F3 1 Tf 9 0 1.913 9 100.65 240.675 Tm (Wolbachia)Tj /F1 1 Tf 9 0 0 9 143.37 240.675 Tm 0 Tc ( )Tj 0.4262 0 TD 0.0093 Tc 0.1389 Tw (and an insect host has also been)Tj -9.4462 -1.2 TD 0 Tc 0.1227 Tw (documented, although the mechanism for such transfer)Tj T* 0.0055 Tc 0.1389 Tw (remains elusive \(Kondo )Tj /F3 1 Tf 9 0 1.913 9 163.65 219.075 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 184.71 219.075 Tm [(, 2002\). )22.6(These promising)]TJ -13.6133 -1.2 TD 0 Tc -0.0263 Tw (hypotheses notwithstanding, it is worth noting that to date)Tj T* 0.0161 Tc 0.1389 Tw (none of these vectors has been observed to mediate)Tj T* 0 Tc 0.1148 Tw [(horizontal transfer of a )19.6(TE between two hosts, either in)]TJ T* 0.0004 Tw (natural or in laboratory populations.)Tj 1.8867 -1.2 TD -0.0674 Tw (An interesting situation exists with the Class I element)Tj /F3 1 Tf 9 0 1.913 9 62.19 154.275 Tm -0.0018 Tc 0 Tw (gypsy)Tj /F1 1 Tf 9 0 0 9 85.65 154.275 Tm -0.0695 Tw [(. )22.9(This )15.1(TE can act as an endogenous retrovirus, since)]TJ -2.6067 -1.2 TD 0 Tc 0.0175 Tw (it encodes an envelope protein and possesses infectious)Tj T* -0.0673 Tw (properties \(Kim )Tj /F3 1 Tf 9 0 1.913 9 124.47 132.675 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 143.37 132.675 Tm (, 1994\). )Tj /F3 1 Tf 9 0 1.913 9 175.17 132.675 Tm 0 Tw (Gypsy)Tj /F1 1 Tf 9 0 0 9 200.73 132.675 Tm ( )Tj 0.2107 0 TD -0.0673 Tw (can produce virus-like)Tj -15.604 -1.2 TD 0.0902 Tw (particles \(Lcher )Tj /F3 1 Tf 9 0 1.913 9 132.87 121.875 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 153.21 121.875 Tm (, 1997\) and it has been shown in)Tj -10.1133 -1.2 TD 0.0076 Tc 0.1389 Tw (experimental conditions that )Tj /F3 1 Tf 9 0 1.913 9 182.49 111.075 Tm 0 Tw (gypsy)Tj /F1 1 Tf 9 0 0 9 206.37 111.075 Tm 0 Tc ( )Tj 0.4245 0 TD 0.0076 Tc 0.1389 Tw (can be horizontally)Tj -16.4445 -1.2 TD -0.0002 Tc -0.0695 Tw (transmitted between )Tj /F3 1 Tf 9 0 1.913 9 144.45 100.275 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 187.47 100.275 Tm 0 Tc ( )Tj 0.2083 0 TD -0.0002 Tc -0.0695 Tw (species \(Mejlumian )Tj /F3 1 Tf 9 0 1.913 9 267.57 100.275 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 286.47 100.275 Tm 0 Tc 0 Tw (,)Tj 2.8 72 TD -0.0059 Tc -0.0695 Tw [(2002\). )15.3(Thus, in principle, horizontal transfer of )]TJ /F3 1 Tf 9 0 1.913 9 490.11 748.275 Tm 0 Tw (gypsy)Tj /F1 1 Tf 9 0 0 9 513.33 748.275 Tm 0 Tc ( )Tj 0.2026 0 TD -0.0059 Tc (would)Tj -22.6093 -1.2 TD 0 Tc 0.0001 Tw [(not require a vector)48.7(.)]TJ /F3 1 Tf 9 0 1.913 9 311.67 715.875 Tm (Overlap Between Donor and Recipient Hosts)Tj /F1 1 Tf 9 0 0 9 311.67 705.0751 Tm -0.04 Tw (A fascinating example of horizontal transfer of an element)Tj T* 0.0881 Tw (of the )Tj /F3 1 Tf 9 0 1.913 9 338.31 694.275 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 368.79 694.275 Tm ( )Tj 0.3661 0 TD 0.0881 Tw (family between a parasitoid wasp and its)Tj -6.7128 -1.2 TD 0.0493 Tc 0.1389 Tw (lepidopteran host provides a good example of the)Tj T* 0 Tc -0.0141 Tw (ecological overlap between donor and recipient that must)Tj T* 0.0827 Tw [(accompany horizontal transfer \(Y)86.9(oshiyama )]TJ /F3 1 Tf 9 0 1.913 9 486.9301 661.875 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 507.21 661.875 Tm (, 2001\).)Tj -21.7267 -1.2 TD 0.0297 Tw (In this case, the parasitoid possesses a )Tj /F3 1 Tf 9 0 1.913 9 473.13 651.075 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 503.67 651.075 Tm ( )Tj 0.3077 0 TD (element)Tj -21.641 -1.2 TD -0.0115 Tc -0.0695 Tw [(with 97% sequence identity to that of its moth host, whereas)]TJ T* 0.0177 Tc 0.139 Tw (related wasps species do not possess )Tj /F3 1 Tf 9 0 1.913 9 479.19 629.475 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 510.81 629.475 Tm 0 Tc ( )Tj 0.4347 0 TD 0.0177 Tc 0.139 Tw (at all.)Tj -22.5613 -1.2 TD -0.0111 Tc -0.0694 Tw [(However)54.2(, in spite of this close physical association between)]TJ T* 0 Tc -0.0087 Tw [(parasitoid and host, a vector)53.9(, such as a virus, may still be)]TJ T* 0.0257 Tc 0.1389 Tw [(necessary to mediate the actual transfer of )10.9(TE DNA)]TJ T* 0 Tc 0.002 Tw (between cells.)Tj 1.8933 -1.2 TD 0.0155 Tc 0.1389 Tw [(The transfer of canonical )]TJ /F3 1 Tf 9 0 1.913 9 438.87 575.475 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 444.9901 575.475 Tm ( )Tj 0.4324 0 TD 0.0155 Tc 0.1389 Tw (elements between )Tj /F3 1 Tf 9 0 1.913 9 529.29 575.475 Tm 0 Tw (D.)Tj -23.9249 -1.2 TD 0 Tc (willistoni)Tj /F1 1 Tf 9 0 0 9 345.21 564.675 Tm ( )Tj 0.2341 0 TD (and )Tj /F3 1 Tf 9 0 1.913 9 364.47 564.675 Tm -0.0439 Tw (D. melanogaster)Tj /F1 1 Tf 9 0 0 9 430.05 564.675 Tm 0 Tw ( )Tj 0.2341 0 TD -0.0439 Tw (illustrates the geographical)Tj -13.3874 -1.2 TD -0.0098 Tc -0.0694 Tw (overlap that is a prerequisite for horizontal transfer between)Tj T* 0.0105 Tc 0.1389 Tw (donor and recipient. )Tj /F3 1 Tf 9 0 1.913 9 399.51 543.075 Tm (D. melanogaster)Tj /F1 1 Tf 9 0 0 9 468.21 543.075 Tm 0 Tc 0 Tw ( )Tj 0.4274 0 TD 0.0105 Tc 0.1389 Tw [(is an Old W)20.6(orld)]TJ -17.8207 -1.2 TD 0 Tc -0.0061 Tw (species whose distribution has only recently expanded to)Tj T* 0.0066 Tc 0.1389 Tw [(the New W)13.5(o)0(rld, probably as a result of human activity)]TJ T* -0.0053 Tc -0.0695 Tw [(\(Kidwell, 1983; Engels, 1992\). )18.4(Thus, the horizontal transfer)]TJ T* 0 Tc 0.1311 Tw (of the canonical )Tj /F3 1 Tf 9 0 1.913 9 380.79 499.875 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 386.79 499.875 Tm ( )Tj 0.4091 0 TD 0.1311 Tw (element was only possible once the)Tj -8.7558 -1.2 TD 0.0262 Tw (distributions of )Tj /F3 1 Tf 9 0 1.913 9 372.69 489.075 Tm (D. melanogaster)Tj /F1 1 Tf 9 0 0 9 438.9301 489.075 Tm 0 Tw ( )Tj 0.3042 0 TD (and )Tj /F3 1 Tf 9 0 1.913 9 459.45 489.075 Tm 0.0262 Tw (D. willistoni)Tj /F1 1 Tf 9 0 0 9 504.69 489.075 Tm 0 Tw ( )Tj 0.3042 0 TD (partially)Tj -21.7508 -1.2 TD 0.0171 Tc 0.1389 Tw [(overlapped, an event that occurred relatively recently)]TJ T* 0 Tc 0 Tw (\(Daniels )Tj /F3 1 Tf 9 0 1.913 9 347.19 467.475 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 366.69 467.475 Tm (, 1990b\).)Tj -6.1133 -1.2 TD 0.0096 Tw (Bimont and colleagues \(1999\) extend the importance of)Tj T* -0.0127 Tc -0.0695 Tw (the overlap between donor and recipient when they suggest)Tj T* -0.0045 Tc -0.0694 Tw (that the expansion of a species range may be concomitant)Tj T* 0 Tc 0.0798 Tw [(with the genomic invasion by )13.3(TEs. )20.5(This invasion can be)]TJ T* -0.0458 Tw [(explained both by the horizontal transfer of alien )13.7(TEs from)]TJ T* 0.0435 Tw (new species with which the invading species comes into)Tj T* 0.0047 Tc 0.1389 Tw [(contact, or by the activation of long-time resident )15.1(TEs,)]TJ T* 0 Tc -0.0692 Tw [(leading to an increase in genomic copy number)49.6(. )13.5(Thus, to a)]TJ T* 0.0109 Tc 0.1389 Tw (certain extent, a newly acquired overlap of donor and)Tj T* 0 Tc 0.0435 Tw (recipient may lead to horizontal transfer by providing the)Tj T* 0.0011 Tc 0.1389 Tw [(opportunity for such an exchange. )8.4(This hypothesis has)]TJ T* 0.0524 Tc (been invoked to explain the recent invasion of )Tj /F3 1 Tf 9 0 1.913 9 528.9901 337.875 Tm 0 Tw (D.)Tj -23.8916 -1.2 TD -0.0136 Tc (melanogaster)Tj /F1 1 Tf 9 0 0 9 364.77 327.075 Tm 0 Tc ( )Tj 0.195 0 TD -0.0136 Tc (by )Tj /F3 1 Tf 9 0 1.913 9 377.55 327.075 Tm -0.147 Tc 0.0639 Tw [(P,)-133.4( I)]TJ /F1 1 Tf 9 0 0 9 388.77 327.075 Tm 0 Tc 0 Tw ( )Tj 0.195 0 TD -0.0136 Tc (and )Tj /F3 1 Tf 9 0 1.913 9 406.89 327.075 Tm (hobo)Tj /F1 1 Tf 9 0 0 9 426.45 327.075 Tm 0 Tc ( )Tj 0.195 0 TD -0.0136 Tc -0.0694 Tw (elements, and of )Tj /F3 1 Tf 9 0 1.913 9 493.29 327.075 Tm -0.0695 Tw (D. simulans)Tj /F1 1 Tf 9 0 0 9 311.67 316.275 Tm 0 Tc -0.0008 Tw (by the retrotransposon )Tj /F3 1 Tf 9 0 1.913 9 404.25 316.275 Tm 0 Tw (412)Tj /F1 1 Tf 9 0 0 9 419.25 316.275 Tm -0.0001 Tc -0.0007 Tw [( \(V)17.1(ieira )]TJ /F3 1 Tf 9 0 1.913 9 450.09 316.275 Tm 0 Tc -0.0008 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 469.59 316.275 Tm (, 1999\).)Tj -15.6533 -1.2 TD -0.0122 Tw (A similar coincidence of expansion of host range and)Tj -1.8933 -1.2 TD 0.0361 Tc 0.1389 Tw [(invasion of )15.5(TEs by horizontal transfer is apparently)]TJ T* -0.0002 Tc -0.0695 Tw (occurring with the drosophilid species )Tj /F3 1 Tf 9 0 1.913 9 460.53 283.875 Tm (Zaprionus indianus)Tj /F1 1 Tf 9 0 0 9 535.9501 283.875 Tm 0 Tc 0 Tw (.)Tj -24.92 -1.2 TD -0.0479 Tw (Flies of the genus )Tj /F3 1 Tf 9 0 1.913 9 383.49 273.075 Tm 0 Tw (Zaprionus)Tj /F1 1 Tf 9 0 0 9 423.57 273.075 Tm ( )Tj 0.2301 0 TD -0.0479 Tw (are of afrotropical origin, and)Tj -12.6634 -1.2 TD -0.0052 Tw (are closely related to the genus )Tj /F3 1 Tf 9 0 1.913 9 438.9901 262.275 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 482.01 262.275 Tm ( )Tj 0.2728 0 TD -0.0052 Tw (\(Remsen and)Tj -19.1994 -1.2 TD 0.0145 Tw [(OGrady)69.7(, 2002; Figure 1\). )]TJ /F3 1 Tf 9 0 1.913 9 416.07 251.475 Tm (Z. indianus)Tj /F1 1 Tf 9 0 0 9 460.23 251.475 Tm 0 Tw ( )Tj 0.2925 0 TD 0.0145 Tw (is in the process of)Tj -16.7992 -1.2 TD 0.0343 Tc 0.1389 Tw (expanding its range, having recently invaded South)Tj T* 0 Tc 0.0893 Tw (America, and is rapidly spreading throughout Brazil and)Tj T* 0.0289 Tc 0.1389 Tw [(neighboring countries \(V)9.1(ilela, 1999\). In addition, the)]TJ T* 0 Tc 0.0015 Tw (ecological niche of )Tj /F3 1 Tf 9 0 1.913 9 388.29 208.275 Tm (Z. indianus)Tj /F1 1 Tf 9 0 0 9 432.33 208.275 Tm 0.0001 Tc 0.0014 Tw ( is very similar to that of )Tj /F3 1 Tf 9 0 1.913 9 529.47 208.275 Tm 0 Tc 0 Tw (D.)Tj -23.9449 -1.2 TD (simulans)Tj /F1 1 Tf 9 0 0 9 347.19 197.475 Tm ( )Tj 0.2586 0 TD -0.0194 Tw (and these species are now often found together)Tj -4.2053 -1.2 TD -0.0006 Tc -0.0695 Tw (in Brazil \(E. L., unpublished results\). Concomitant with this)Tj T* 0 Tc -0.014 Tw (range expansion, horizontal transfer of )Tj /F3 1 Tf 9 0 1.913 9 467.13 175.875 Tm 0 Tw (gypsy)Tj /F1 1 Tf 9 0 0 9 490.65 175.875 Tm ( )Tj 0.264 0 TD (between )Tj /F3 1 Tf 9 0 1.913 9 529.47 175.875 Tm (D.)Tj -23.9449 -1.2 TD (simulans)Tj /F1 1 Tf 9 0 0 9 347.19 165.075 Tm 0.0003 Tc 0.0013 Tw ( and )Tj /F3 1 Tf 9 0 1.913 9 367.23 165.075 Tm 0 Tc 0.0016 Tw (Z. indianus )Tj /F1 1 Tf 9 0 0 9 413.79 165.075 Tm (has been described by Herdia)Tj -11.3467 -1.2 TD 0.0139 Tc 0.1389 Tw [(\(2002\), based on incongruence between )13.9(TE and host)]TJ T* -0.0132 Tc -0.0694 Tw (phylogenies. Comparison of sequence divergence of a non-)Tj T* 0 Tc -0.0617 Tw (mobile nuclear gene \(superoxide dismutase, )Tj /F3 1 Tf 9 0 1.913 9 488.01 132.675 Tm 0 Tw (Sod)Tj /F1 1 Tf 9 0 0 9 504.03 132.675 Tm -0.0617 Tw (\) and the)Tj /F3 1 Tf 9 0 1.913 9 311.67 121.875 Tm 0.0164 Tc 0 Tw (gypsy)Tj /F1 1 Tf 9 0 0 9 335.97 121.875 Tm 0 Tc ( )Tj 0.4333 0 TD 0.0164 Tc 0.1389 Tw [(sequences \(T)113.8(able 2\) in these species provides)]TJ -3.1333 -1.2 TD 0.0063 Tc (additional corroboration of the hypothesis of horizontal)Tj T* 0.0657 Tc [(transfer)49.2(. )26.3(The divergence of the )11.7(TE sequences is)]TJ ET endstream endobj 16 0 obj << /Type /Page /Parent 93 0 R /Resources 17 0 R /Contents 18 0 R /Thumb 66 0 R /MediaBox [ 0 0 595 842 ] /CropBox [ 0 0 595 842 ] /Rotate 0 >> endobj 17 0 obj << /ProcSet [ /PDF /Text ] /Font << /F1 114 0 R /F2 113 0 R /F3 128 0 R >> /ExtGState << /GS1 139 0 R /GS2 138 0 R >> /ColorSpace << /Cs8 112 0 R >> >> endobj 18 0 obj << /Length 14248 >> stream BT /F1 1 Tf 9 0 0 9 55.93 779.115 Tm 0 0 0 1 k /GS2 gs -0.0001 Tc -0.0003 Tw (6 Silva )Tj /F3 1 Tf 9 0 1.913 9 90.43 779.115 Tm 0 Tc (et al)Tj /F1 1 Tf 9 0 0 9 107.47 779.115 Tm 0 Tw (.)Tj /F2 1 Tf 37.9351 29.4096 -29.4096 37.9351 34.496 531.1378 Tm 2 Tr 0 0 0 0.45 K 0 J 0 j 0.576 w 10 M []0 d /Cs8 cs 0 scn /GS1 gs 0.0686 Tc -0.0001 Tw (PAGE PROOFS)Tj /F1 1 Tf 9 0 0 9 56.53 748.275 Tm 0 Tr 0 0 0 1 k /GS2 gs 0.0056 Tc 0.1389 Tw (considerably less than that of a host gene supposedly)Tj 0 -1.2 TD -0.0141 Tc -0.0695 Tw (evolving under purifying selection. In order to further explore)Tj T* 0 Tc 0.1016 Tw [(this possibility)69.7(, PCR was used to screen 35 Neotropical)]TJ T* 0.0224 Tw (species \(14 from the subgenus )Tj /F3 1 Tf 9 0 1.913 9 183.13 715.875 Tm 0 Tw (Sophophora)Tj /F1 1 Tf 9 0 0 9 232.15 715.875 Tm 0.0224 Tw (, 19 from the)Tj -19.5133 -1.2 TD 0 Tw (subgenus )Tj /F3 1 Tf 9 0 1.913 9 98.89 705.0751 Tm (Drosophila)Tj /F1 1 Tf 9 0 0 9 141.91 705.0751 Tm 0.0889 Tw (, and one each from the subgenus)Tj /F3 1 Tf 9 0 1.913 9 56.53 694.275 Tm 0 Tw (Dorsilopha)Tj /F1 1 Tf 9 0 0 9 99.55 694.275 Tm ( )Tj 0.391 0 TD (and )Tj /F3 1 Tf 9 0 1.913 9 121.63 694.275 Tm 0.113 Tw (Zaprionus indianus)Tj /F1 1 Tf 9 0 0 9 198.73 694.275 Tm (\) for the presence of)Tj -15.8 -1.2 TD 0.0576 Tw (elements from the )Tj /F3 1 Tf 9 0 1.913 9 132.67 683.475 Tm 0 Tw (297)Tj /F1 1 Tf 9 0 0 9 147.67 683.475 Tm (/)Tj /F3 1 Tf 9 0 1.913 9 150.13 683.475 Tm (tom)Tj /F1 1 Tf 9 0 0 9 165.13 683.475 Tm ( )Tj 0.3356 0 TD 0.0576 Tw [(group of retrotransposons. )47.8(A)]TJ -12.4023 -1.2 TD 0.0507 Tw (fragment with the expected size was amplified only from)Tj T* 0.0573 Tw (species of the )Tj /F3 1 Tf 9 0 1.913 9 116.17 661.875 Tm 0 Tw (melanogaster)Tj /F1 1 Tf 9 0 0 9 170.65 661.875 Tm ( )Tj 0.3353 0 TD 0.0573 Tw (group and from )Tj /F3 1 Tf 9 0 1.913 9 238.75 661.875 Tm (Z. indianus)Tj /F1 1 Tf 9 0 0 9 56.53 651.075 Tm 0.113 Tw (\(E. L., unpublished results\). Once again the divergence)Tj 0 -1.2 TD 0.0308 Tw [(between the )14.3(TE sequences is lower than that of the host)]TJ T* 0.0098 Tw [(gene, suggesting a possible horizontal transfer \(T)104.6(able 2\).)]TJ T* -0.0071 Tc -0.0695 Tw (A careful examination of alternative possibilities is currently)Tj T* 0 Tc 0.0825 Tw [(underway)68.4(. Finally)74.2(, Brunet and collaborators \(1999\) have)]TJ T* 0.0576 Tw (detected the presence of )Tj /F3 1 Tf 9 0 1.913 9 160.21 597.075 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 190.69 597.075 Tm ( )Tj 0.3356 0 TD 0.0576 Tw (elements in )Tj /F3 1 Tf 9 0 1.913 9 243.31 597.075 Tm 0 Tw (Zaprionus)Tj -20.4983 -1.2 TD (indianus.)Tj /F1 1 Tf 9 0 0 9 92.59 586.275 Tm ( )Tj 0.2867 0 TD 0.0249 Tw (These elements are very closely related to that)Tj -4.2933 -1.2 TD 0 Tw (of )Tj /F3 1 Tf 9 0 1.913 9 67.39 575.475 Tm 0.0944 Tw (D. simulans)Tj /F1 1 Tf 9 0 0 9 115.27 575.475 Tm 0 Tw ( )Tj 0.3724 0 TD 0.0944 Tw [(\(T)109.8(able 2\). Whether the introduction of all)]TJ -6.8991 -1.2 TD 0.0714 Tw [(these )15.4(TE families into the )]TJ /F3 1 Tf 9 0 1.913 9 163.15 564.675 Tm (Z. indianus)Tj /F1 1 Tf 9 0 0 9 207.79 564.675 Tm 0 Tw ( )Tj 0.3494 0 TD 0.0714 Tw (genome is indeed)Tj -17.1561 -1.2 TD -0.0082 Tc -0.0694 Tw (recent, and results from the expansion of the species range)Tj T* 0 Tc -0.0293 Tw (remains to be shown, but the possibility is fascinating and)Tj T* -0.0007 Tw (certainly warrants further investigation.)Tj /F2 1 Tf 0 -2.4 TD -0.0057 Tc -0.0694 Tw [(Factors )35.3(Affecting the Frequency of Horizontal T)55.8(ransfer)]TJ /F1 1 Tf 0 -1.2 TD 0.0144 Tc 0.1389 Tw (A horizontal transfer event, defined as the successful)Tj T* 0 Tc -0.053 Tw [(invasion of a new species by a )15.9(TE, can be divided into two)]TJ T* -0.0054 Tw (phases: the transfer of DNA between donor and recipient)Tj T* -0.0121 Tc -0.0695 Tw [(and a subsequent increase in )8.4(TE frequency \(Kidwell, 1992\).)]TJ T* 0 Tc 0.1273 Tw [(The first phase requires the physical transfer of the )13.6(TE)]TJ T* 0.0502 Tw [(from one organism to another)51.1(, the stable integration into)]TJ T* 0.0013 Tc 0.1389 Tw [(the recipient)12.4(s germline and the expression of the )16.3(TE)17.2(s)]TJ T* 0 Tc -0.061 Tw (coding sequences. Once a stable transfer is achieved, the)Tj T* -0.023 Tw (element must increase in copy number within the cell and)Tj T* 0.045 Tc 0.1389 Tw (then spread throughout the population in order for)Tj T* 0 Tc -0.0005 Tw (horizontal transfer to succeed.)Tj 1.8933 -1.2 TD 0.0188 Tc 0.139 Tw [(Any one of these events seems unlikely)73.7(, and yet)]TJ -1.8933 -1.2 TD 0 Tc 0.0503 Tw [(horizontal transfer requires that they occur together)45.9(, in a)]TJ T* 0.039 Tw [(coordinated manner)52.1(. Because the presence and mobility)]TJ T* 0.0014 Tw (of a transposable element may be deleterious to the host)Tj T* 0.019 Tc 0.1389 Tw (\(for a review see Nuzhdin, 1999\), this spread will be)Tj T* 0 Tc 0.0598 Tw [(counteracted by selection and by host- and )16.3(TE-encoded)]TJ T* -0.0077 Tc -0.0695 Tw [(mechanisms that repress transposition. )15(Therefore, not only)]TJ T* 0 Tc 0.0755 Tw [(will an inherent predisposition of a )13.3(TE to perform any of)]TJ T* 0.0158 Tw (the required steps increase its probability of a successful)Tj T* 0.0105 Tc 0.1389 Tw [(transfer but, ultimately)67.6(, the structure and transposition)]TJ T* 0.0268 Tc [(mechanism of a )14.6(TEs are actively molded by the co-)]TJ T* 0 Tc -0.0006 Tw [(evolution between host and )16.3(TE.)]TJ /F3 1 Tf 9 0 1.913 9 56.53 240.675 Tm 0.0002 Tw (Stable Integration into a New Genome)Tj /F1 1 Tf 9 0 0 9 56.53 229.875 Tm -0.0167 Tw (The physical transfer of nuclear DNA from one species to)Tj T* 0.1128 Tw (another must occur in such a manner that the recipient)Tj T* 0.062 Tw (cell is able to take up the donor DNA. Once this occurs,)Tj T* -0.0096 Tc -0.0695 Tw (the DNA must enter the nucleus of a germline cell, integrate)Tj T* -0.0052 Tc [(into a chromosome and be expressed. )15.1(This process is best)]TJ T* 0 Tc -0.0045 Tw (understood for the )Tj /F3 1 Tf 9 0 1.913 9 132.01 175.875 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 138.01 175.875 Tm -0.0001 Tc -0.0044 Tw ( element, which for twenty years has)Tj -9.0533 -1.2 TD 0 Tc 0.0893 Tw (been used routinely to transform )Tj /F3 1 Tf 9 0 1.913 9 192.13 165.075 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 235.15 165.075 Tm ( )Tj 0.3673 0 TD 0.0893 Tw (\(Rubin and)Tj -20.214 -1.2 TD 0.0118 Tc 0.1389 Tw (Spradling, 1982\). In this case, a )Tj /F3 1 Tf 9 0 1.913 9 196.75 154.275 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 202.87 154.275 Tm ( )Tj 0.4287 0 TD 0.0118 Tc 0.1389 Tw (element construct,)Tj -16.6887 -1.2 TD -0.0027 Tc -0.0695 Tw (carrying the terminal inverted repeats, is microinjected into)Tj T* 0 Tc 0.1132 Tw (the syncytial blastoderm of early embryos, where the )Tj /F3 1 Tf 9 0 1.913 9 277.33 132.675 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 56.53 121.875 Tm 0.0251 Tc 0.139 Tw [(elements subsequently transpose into the recipient)9.6(s)]TJ T* 0 Tc 0.0389 Tw [(genome. )14.6(This requires a supply of transposase, which is)]TJ T* -0.0122 Tw (produced from transcription of a transposase gene that is)Tj 27.7267 46.4867 TD 0.134 Tw (co-injected with the )Tj /F3 1 Tf 9 0 1.913 9 389.23 518.655 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 395.23 518.655 Tm ( )Tj 0.412 0 TD 0.134 Tw (element construct. Only some of)Tj -10.3187 -1.2 TD 0.0128 Tw (these insertions will occur in nuclei of those cells that will)Tj T* 0.0054 Tc 0.1389 Tw [(ultimately form germline tissue, potentially producing a)]TJ T* 0.0003 Tc (stable fly lineage that is genetically transformed. In the)Tj T* 0.029 Tc [(laboratory)75.9(, a strong promoter is used to provide the)]TJ T* 0 Tc -0.0515 Tw [(necessary level of transposase that is needed for mobility)75.2(.)]TJ T* 0.0017 Tw [(However)58.4(, in nature the integration and proper expression)]TJ T* 0.0203 Tc 0.139 Tw [(of a )18.9(TE would seem to be a daunting impediment to)]TJ T* 0 Tc -0.0004 Tw [(successful horizontal transfer)54.9(.)]TJ 1.8867 -1.2 TD 0.0022 Tw (Some of the complex features required for horizontal)Tj -1.8867 -1.2 TD -0.004 Tc -0.0694 Tw (transfer are integral characteristics of the life cycle of some)Tj T* 0.0154 Tc 0.1389 Tw [(TE families, and reflect selection on )20.5(TEs for ef)20.1(fective)]TJ T* 0.0049 Tc (transposition. For example, the transposase of class II)Tj T* 0 Tc 0.0578 Tw (elements belonging to the )Tj /F3 1 Tf 9 0 1.913 9 413.71 378.255 Tm 0 Tw [(mariner)15.7(-T)97.3(c1)]TJ /F1 1 Tf 9 0 0 9 461.23 378.255 Tm ( )Tj 0.3358 0 TD 0.0578 Tw (superfamily have)Tj -17.5758 -1.2 TD -0.0055 Tw (been found to contain a nuclear localization signal, which)Tj T* 0.1059 Tw (explains the migration of these enzymes to the nucleus)Tj T* 0.1225 Tw [(\(either upon translation or after co-transfer with its )15.8(TE\),)]TJ T* 0.0529 Tw [(where they facilitate )20.5(TE transposition \(Ivics )]TJ /F3 1 Tf 9 0 1.913 9 481.81 335.055 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 501.79 335.055 Tm (, 1996\).)Tj -21.7467 -1.2 TD 0.0113 Tc 0.1389 Tw [(Since Class II )17(TEs transpose predominantly via DNA-)]TJ T* 0 Tc -0.0677 Tw (mediated processes \(Kaufman and Rio, 1992; see Hartl )Tj /F3 1 Tf 9 0 1.913 9 525.31 313.455 Tm 0 Tw (et)Tj -24.1049 -1.2 TD 0.0213 Tc (al.)Tj /F1 1 Tf 9 0 0 9 316.15 302.655 Tm 0.1389 Tw (, 1997 for a review of transposition in )Tj /F3 1 Tf 9 0 1.913 9 483.37 302.655 Tm 0 Tw [(mariner)17.9(-T)93.2(c1)]TJ /F1 1 Tf 9 0 0 9 306.07 291.855 Tm -0.0012 Tc -0.0695 Tw (elements\), extra-chromosomal DNA copies of the element)Tj T* 0 Tc -0.0375 Tw (are a necessary feature of their transposition mechanism.)Tj T* 0.0008 Tc 0.1389 Tw (Class I elements, on the other hand, depend on extra-)Tj T* 0 Tc 0.1007 Tw (chromosomal copies of the element in the form of RNA)Tj T* -0.0056 Tc -0.0694 Tw (intermediates, which are necessarily not as stable as DNA.)Tj T* 0 Tc 0.0794 Tw (As is the case for the transposase of Class II elements,)Tj T* 0.0031 Tc 0.1389 Tw (the enzymes encoded by Class I elements need to be)Tj T* 0 Tc 0 Tw (produced at levels high enough to ensure transposition.)Tj 1.8867 -1.2 TD -0.0089 Tc -0.0694 Tw [(Dif)15.4(ferences in the transposition mechanism of L)74.8(TR and)]TJ -1.8867 -1.2 TD 0.013 Tc 0.1389 Tw [(non-L)75.2(TR Class I elements provide useful insights into)]TJ T* 0.0053 Tc [(varying rates of horizontal transfer among )17.5(TE families.)]TJ T* 0.011 Tc 0.139 Tw [(During the transposition of L)75.3(T)0(R retroelements, a DNA)]TJ T* -0.0066 Tc -0.0694 Tw (intermediate is produced, which can insert into the genome)Tj T* 0 Tc 0.0402 Tw (in a manner analogous to Class II elements \(Luan )Tj /F3 1 Tf 9 0 1.913 9 510.43 151.455 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 530.29 151.455 Tm 0 Tw (,)Tj -24.9133 -1.2 TD 0.0196 Tc 0.1389 Tw [(1993, and references therein\). However)51.2(, for non-L)77.7(TR)]TJ T* 0 Tc 0.0583 Tw (transposable elements, an RNA intermediate is reverse-)Tj T* 0.042 Tw (transcribed directly into a chromosomal target site \(Luan)Tj /F3 1 Tf 9 0 1.913 9 306.07 108.255 Tm -0.0064 Tc -0.0694 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 324.61 108.255 Tm (, 1993\). Malik and collaborators \(1999\) suggested that)Tj -2.06 -1.2 TD 0 Tc 0.0921 Tw [(this dif)15.4(ference might be suf)23(ficient to explain the rarity of)]TJ 7 0 0 7 306.07 745.095 Tm 0 Tw (T)Tj 0.4971 0 TD -0.0011 Tw (able 2. Divergence \(substitutions per nucleotide\) of a host marker \()Tj /F3 1 Tf 7 0 1.4879 7 517.99 745.095 Tm 0 Tw (Sod)Tj /F1 1 Tf 7 0 0 7 530.47 745.095 Tm (\))Tj -32.0571 -1.2 TD 0.0009 Tw [(and of )17(TEs between )]TJ /F3 1 Tf 7 0 1.4879 7 370.15 736.6951 Tm (Z. indianus)Tj /F1 1 Tf 7 0 0 7 404.41 736.6951 Tm 0.0001 Tc 0.0008 Tw ( and two )Tj /F3 1 Tf 7 0 1.4879 7 432.85 736.6951 Tm 0 Tc 0 Tw (Drosophila)Tj /F1 1 Tf 7 0 0 7 466.33 736.6951 Tm 0.0001 Tc 0.0008 Tw ( species)Tj 4.9 0 0 4.9 492.01 739.095 Tm 0 Tc 0 Tw (1)Tj 7 0 0 7 494.71 736.6951 Tm (.)Tj /F3 1 Tf 7 0 1.4879 7 371.05 719.895 Tm -0.0017 Tw [(D. melanogaster)-3233.7(D. simulans)]TJ -8.7727 -2.4 TD -0.0022 Tw (D. simulans)Tj 2.4065 -1.2 TD 0.001 Tc 0 Tw (Sod)Tj 4.9 0 1.0415 4.9 333.61 697.095 Tm 0 Tc (2)Tj /F1 1 Tf 7 0 0 7 389.95 694.6951 Tm -0.0011 Tc (0.03)Tj /F3 1 Tf 7 0 1.4879 7 318.19 686.295 Tm 0.0011 Tc (gypsy)Tj 4.9 0 1.0415 4.9 336.49 688.6951 Tm 0 Tc (2)Tj /F1 1 Tf 7 0 0 7 389.95 686.295 Tm -0.0011 Tc (0.12)Tj /F3 1 Tf 7 0 1.4879 7 306.07 661.095 Tm 0 Tc 0.0013 Tw (Z. indianus)Tj 2.4065 -1.2 TD 0.001 Tc 0 Tw (Sod)Tj 4.9 0 1.0415 4.9 333.61 655.095 Tm 0 Tc (2)Tj /F1 1 Tf 7 0 0 7 389.95 652.695 Tm -0.0011 Tc [(0.32)-7572.8(0.33)]TJ /F3 1 Tf 7 0 1.4879 7 318.19 644.295 Tm 0.0011 Tc (gypsy)Tj 4.9 0 1.0415 4.9 336.49 646.695 Tm 0 Tc (2)Tj /F1 1 Tf 7 0 0 7 390.19 644.295 Tm -0.0008 Tc [(0.1)68.6(1)-7604.8(0.07)]TJ /F3 1 Tf 7 0 1.4879 7 313.75 635.895 Tm 0.0007 Tc (17.6/tom)Tj 4.9 0 1.0415 4.9 340.99 638.295 Tm 0 Tc (3)Tj /F1 1 Tf 7 0 0 7 456.55 635.895 Tm -0.0011 Tc (0.17)Tj /F3 1 Tf 7 0 1.4879 7 315.49 627.495 Tm 0.001 Tc (mariner)Tj 4.9 0 1.0415 4.9 339.25 629.895 Tm 0 Tc (4)Tj /F1 1 Tf 7 0 0 7 456.55 627.495 Tm -0.0011 Tc (0.06)Tj 4.9 0 0 4.9 306.13 613.095 Tm 0 Tc (1)Tj 7 0 0 7 308.83 610.695 Tm ( )Tj 0.2078 0 TD -0.0008 Tc -0.0694 Tw [(Divergence estimated according to Kimura)17.3(s 2-parameter model \(Kimura,)]TJ -0.6021 -1.2 TD 0.0001 Tc 0 Tw (1980\).)Tj 4.9 0 0 4.9 306.07 596.295 Tm 0 Tc (2)Tj 7 0 0 7 308.83 593.895 Tm 0.0002 Tc 0.0028 Tw ( Herdia \(2002\).)Tj 4.9 0 0 4.9 306.07 587.895 Tm 0 Tc 0 Tw (3)Tj 7 0 0 7 308.83 585.495 Tm 0.0008 Tw ( E. Loreto, unpublished results.)Tj 4.9 0 0 4.9 306.07 579.495 Tm 0 Tw (4)Tj 7 0 0 7 308.77 577.095 Tm ( )Tj 0.2063 0 TD -0.0022 Tc -0.0695 Tw (Genbank accession numbers for )Tj /F3 1 Tf 7 0 1.4879 7 410.53 577.095 Tm (D. simulans)Tj /F1 1 Tf 7 0 0 7 446.47 577.095 Tm 0 Tc 0 Tw ( )Tj 0.2063 0 TD -0.0022 Tc -0.0695 Tw (\(AF037052\) and )Tj /F3 1 Tf 7 0 1.4879 7 499.21 577.095 Tm (Z. indianus)Tj /F1 1 Tf 7 0 0 7 306.07 568.695 Tm 0 Tc 0 Tw (\(AF034700\) )Tj /F3 1 Tf 7 0 1.4879 7 344.95 568.695 Tm (mariner)Tj /F1 1 Tf 7 0 0 7 368.65 568.695 Tm -0.0002 Tc -0.002 Tw ( elements.)Tj ET 0 0 0 1 K 0.24 w /GS1 gs 1 i 306.19 755.475 m 306.19 755.295 l 306.07 754.575 m 532.81 754.575 l 306.07 731.355 m 532.81 731.355 l 306.07 622.815 m 532.81 622.815 l 306.07 562.695 m 532.81 562.695 l 306.07 714.915 m 532.81 714.915 l S endstream endobj 19 0 obj << /Type /Page /Parent 93 0 R /Resources 20 0 R /Contents 21 0 R /Thumb 68 0 R /MediaBox [ 0 0 595 842 ] /CropBox [ 0 0 595 842 ] /Rotate 0 >> endobj 20 0 obj << /ProcSet [ /PDF /Text ] /Font << /F1 114 0 R /F2 113 0 R /F3 128 0 R >> /ExtGState << /GS1 139 0 R /GS2 138 0 R >> /ColorSpace << /Cs8 112 0 R >> >> endobj 21 0 obj << /Length 14984 >> stream BT /F1 1 Tf 9 0 0 9 342.51 780.5551 Tm 0 0 0 1 k /GS2 gs 0 Tc -0.0155 Tw [(Horizontal Transfer of Transposable Elements 7)]TJ /F2 1 Tf 37.9351 29.4096 -29.4096 37.9351 42.9423 542.4378 Tm 2 Tr 0 0 0 0.45 K 0 J 0 j 0.576 w 10 M []0 d /Cs8 cs 0 scn /GS1 gs 0.0686 Tc -0.0001 Tw (PAGE PROOFS)Tj /F1 1 Tf 9 0 0 9 62.19 748.275 Tm 0 Tr 0 0 0 1 k /GS2 gs 0 Tc 0.0843 Tw [(non-L)70.4(TR elements)41( horizontal transfer)49.7(. )20.3(The transposition)]TJ 0 -1.2 TD 0.0341 Tw [(mechanism used by non-L)76.2(TR )11.8(TEs does not preclude the)]TJ T* 0.1271 Tw [(possibility of horizontal transfer using a virus as vector)49.7(,)]TJ T* -0.0025 Tc -0.0695 Tw (but reverse transcription probably has to occur directly into)Tj T* 0 Tc -0.0002 Tw (the viral DNA, an event that might be exceedingly rare.)Tj 1.8867 -1.2 TD 0.006 Tc 0.1389 Tw [(Thus, there appears to be a gradient of horizontal)]TJ -1.8867 -1.2 TD 0 Tc 0.0737 Tw (transfer that reflects the presence of DNA intermediates)Tj T* -0.0623 Tw (during the transposition events. Horizontal transfer seems)Tj T* 0.0022 Tw (much more common for Class II elements for which DNA)Tj T* -0.0139 Tw (intermediates are a persistent feature of the transposition)Tj T* -0.0464 Tw (process. Horizontal transfer seems to be less common for)Tj T* 0.0545 Tc 0.139 Tw [(Class I L)72(TR retroelements, which produce a DNA)]TJ T* 0.0019 Tc 0.1389 Tw (intermediate only after reverse transcription of an RNA)Tj T* -0.002 Tc -0.0695 Tw [(copy)77.2(, and horizontal transfer is least common for the Class)]TJ T* 0.0061 Tc 0.139 Tw [(I non-L)74.9(TR elements, for which no DNA)50( intermediate is)]TJ T* 0.0003 Tc 0 Tw (produced.)Tj /F3 1 Tf 9 0 1.913 9 62.19 564.675 Tm 0 Tc -0.0006 Tw [(Spread W)5.5(ithin the New Host)]TJ /F1 1 Tf 9 0 0 9 62.19 553.875 Tm 0.0086 Tw (After stable integration and expression, there must be an)Tj T* 0.0028 Tc 0.1389 Tw [(increase in copy number in order for the )17.4(TE to spread)]TJ T* 0 Tc 0.0916 Tw [(reliably throughout the population. )15.1(The increase in copy)]TJ T* -0.0137 Tc -0.0694 Tw (number within the cell results in a decrease in the probability)Tj T* 0 Tc 0.1236 Tw (of a gamete being formed that carries no copies of the)Tj T* -0.0309 Tw [(element. If fact, if all copies segregate independently)73.9(, that)]TJ T* 0.0656 Tw (probability decreases exponentially as \(1/2\))Tj /F3 1 Tf 6.3 0 1.3391 6.3 237.63 492.195 Tm 0 Tw (n)Tj /F1 1 Tf 9 0 0 9 241.17 489.075 Tm 0.0656 Tw (, where )Tj /F3 1 Tf 9 0 1.913 9 274.35 489.075 Tm 0 Tw (n)Tj /F1 1 Tf 9 0 0 9 279.33 489.075 Tm ( )Tj 0.3436 0 TD (is)Tj -24.4702 -1.2 TD 0.0654 Tw [(the total number of copies per genome. )13.7(This scenario is)]TJ T* -0.0095 Tc -0.0695 Tw [(simplistic, as )16.3(TE spread may be influenced by other factors,)]TJ T* -0.0041 Tc [(including biases in )17.1(TE insertion sites. For example, several)]TJ T* 0 Tc 0.0087 Tw (TEs have been shown to transpose preferentially to sites)Tj T* 0.0849 Tw (close to the parental copy \(Engels, 1989\), meaning that)Tj T* -0.0081 Tc -0.0694 Tw [(these copies would not segregate independently)69.6(. However)52.3(,)]TJ T* 0 Tc 0.032 Tw [(this model clearly exemplifies the powerful ef)12.4(fect of copy)]TJ T* -0.0117 Tc -0.0694 Tw [(number increase on )18.5(TE transmission; if an individual carries)]TJ T* 0 Tc -0.038 Tw [(in its genome ten )21(TE copies that segregate independently)]TJ T* -0.0012 Tc -0.0695 Tw [(of each other)55.8(, the probability of it generating a gamete with)]TJ T* 0 Tc 0.0002 Tw [(no )16.9(TEs is less than one in a thousand.)]TJ 1.8867 -1.2 TD 0.0072 Tc 0.1389 Tw [(Several studies have addressed the subject of )10(TE)]TJ -1.8867 -1.2 TD 0 Tc 0.0343 Tw [(spread, both by quantifying the number of )19.8(TE copies per)]TJ T* 0.1143 Tw [(genome and by estimating the rate of spread of )16.2(TEs in)]TJ T* -0.0127 Tc 0 Tw (populations. )Tj /F3 1 Tf 9 0 1.913 9 111.15 327.075 Tm (P )Tj /F1 1 Tf 9 0 0 9 118.77 327.075 Tm -0.0695 Tw (elements have been shown to spread rapidly)Tj -6.2867 -1.2 TD 0.0007 Tc 0.139 Tw (when first introduced into naive populations \(those that)Tj T* 0 Tc 0.0311 Tw [(lacks the )19.6(TE in question\) in the laboratory \(Kidwell )]TJ /F3 1 Tf 9 0 1.913 9 266.67 305.475 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 286.47 305.475 Tm 0 Tw (,)Tj -24.92 -1.2 TD -0.0026 Tw (1988; Good )Tj /F3 1 Tf 9 0 1.913 9 111.69 294.675 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 131.19 294.675 Tm [(, 1989\). )17.2(These results mimic the spread)]TJ -7.6667 -1.2 TD 0.0394 Tc 0 Tw (of )Tj /F3 1 Tf 9 0 1.913 9 74.49 283.875 Tm (P )Tj /F1 1 Tf 9 0 0 9 84.99 283.875 Tm 0.1389 Tw (elements throughout natural populations of )Tj /F3 1 Tf 9 0 1.913 9 279.57 283.875 Tm 0 Tw (D.)Tj -23.8983 -1.2 TD 0.0695 Tc (melanogaster)Tj /F1 1 Tf 9 0 0 9 124.23 273.075 Tm 0 Tc ( )Tj 0.677 0 TD 0.0695 Tc 0.3295 Tw (within the past 50 to 100 years)Tj -7.5703 -1.2 TD 0.0137 Tc 0 Tw (\(Anxolabhre )Tj /F3 1 Tf 9 0 1.913 9 126.21 262.275 Tm 0.1389 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 147.69 262.275 Tm (, 1988\), with most flies examined)Tj -9.5 -1.2 TD 0 Tc -0.0539 Tw (recently carrying up to 60 )Tj /F3 1 Tf 9 0 1.913 9 163.35 251.475 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 171.33 251.475 Tm -0.0539 Tw (element copies \(Ronsseray )Tj /F3 1 Tf 9 0 1.913 9 281.43 251.475 Tm 0 Tw (et)Tj -24.1049 -1.2 TD (al.)Tj /F1 1 Tf 9 0 0 9 71.67 240.675 Tm -0.0608 Tw (, 1989\). Canonical )Tj /F3 1 Tf 9 0 1.913 9 145.59 240.675 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 153.57 240.675 Tm -0.0608 Tw (elements have also colonized very)Tj -10.1533 -1.2 TD -0.0004 Tc -0.0694 Tw [(recently the New W)14.1(orld species )]TJ /F3 1 Tf 9 0 1.913 9 187.35 229.875 Tm (D. willistoni)Tj /F1 1 Tf 9 0 0 9 231.69 229.875 Tm (, as suggested)Tj -18.8333 -1.2 TD 0 Tc -0.017 Tw [(by an average pairwise dif)14.8(ference between copies of less)]TJ T* 0.0679 Tw (than 1% \(Silva, 2000\) and by the low frequency of each)Tj T* -0.0019 Tc -0.0694 Tw (insertion \(A. Holyoke and M. Kidwell, unpublished results\).)Tj T* 0 Tc 0.0503 Tw (In addition, these elements were shown to spread faster)Tj T* 0.0308 Tc 0.1389 Tw (than neutral nuclear markers in natural populations,)Tj T* -0.004 Tc -0.0694 Tw (overcoming barriers such as moderate levels of population)Tj T* 0.0045 Tc 0.1389 Tw (subdivision \(Silva, 2000\), and are now present in 5-14)Tj T* 0.0695 Tc 0.1611 Tw (copies per genome \(A. Holyoke and M. Kidwell,)Tj T* 0 Tc 0.0002 Tw (unpublished results\).)Tj 1.8867 -1.2 TD -0.0535 Tw (A more dramatic example of the potential for increase)Tj -1.8867 -1.2 TD 0.0092 Tc 0.1389 Tw (in copy number accompanying horizontal transfer was)Tj T* 0 Tc 0.045 Tw (found for the )Tj /F3 1 Tf 9 0 1.913 9 116.43 100.275 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 146.97 100.275 Tm ( )Tj 0.323 0 TD 0.045 Tw (element in some host species. For)Tj 17.977 72 TD 0.1288 Tw (example, the genome of the planarian, )Tj /F3 1 Tf 9 0 1.913 9 474.75 748.275 Tm (Dugesia tigrina)Tj /F1 1 Tf 9 0 0 9 535.9501 748.275 Tm 0 Tw (,)Tj -24.92 -1.2 TD 0.004 Tc 0.1389 Tw (contains approximately 8000 copies of )Tj /F3 1 Tf 9 0 1.913 9 474.87 737.475 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 505.65 737.475 Tm 0 Tc ( )Tj 0.4209 0 TD 0.004 Tc 0.1389 Tw (and, of)Tj -21.9743 -1.2 TD -0.0116 Tc -0.0694 Tw (those sequenced, all are quite similar to each other \(Garcia-)Tj T* 0.0325 Tc 0 Tw (Fernandez )Tj /F3 1 Tf 9 0 1.913 9 361.41 715.875 Tm 0.1389 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 383.91 715.875 Tm (, 1995\). Furthermore, the planarian)Tj /F3 1 Tf 9 0 1.913 9 311.67 705.0751 Tm 0.0038 Tc 0 Tw (mariners)Tj /F1 1 Tf 9 0 0 9 347.01 705.0751 Tm 0 Tc ( )Tj 0.4207 0 TD 0.0038 Tc 0.1389 Tw (are characterized by full-length, uninterrupted)Tj -4.3474 -1.2 TD -0.0031 Tc -0.0695 Tw (reading frames and are dispersed throughout the genome.)Tj T* 0 Tc 0 Tw (T)Tj 0.5 0 TD -0.0005 Tc -0.0695 Tw [(ogether)54(, these observations suggest recent transposition)]TJ -0.5 -1.2 TD 0 Tc 0.0001 Tw [(and spread following horizontal transfer)52.5(.)]TJ /F3 1 Tf 9 0 1.913 9 311.67 651.075 Tm 0.0002 Tw [(Factors Limiting the Spread of T)68.9(ransposable Elements)]TJ /F1 1 Tf 9 0 0 9 311.67 640.275 Tm 0.0385 Tw (The invasion of a new species after the initial integration)Tj T* -0.0031 Tc -0.0695 Tw (into the host genome is not always quick, or even possible.)Tj T* -0.0027 Tc [(Several factors might play a crucial role in )13.1(TE spread, such)]TJ T* 0.0355 Tc 0.1389 Tw [(as the ef)13.2(fective population size of the host species)]TJ T* 0.0085 Tc (\(Charlesworth and Charlesworth, 1983; Brookfield and)Tj T* 0.0274 Tc [(Badge, 1997; Quesneville and )52.2(Anxolabhre, 1997\),)]TJ T* 0.0474 Tc (selection \(Nuzhdin, 1999, and references therein\),)Tj T* 0 Tc -0.0062 Tw (repression of transposition, and the presence or absence)Tj T* 0.0097 Tc 0.1389 Tw [(of host factors required for transposition. )15.9(The last two)]TJ T* 0.0044 Tc [(factors vary considerably between )13.6(TE families and are)]TJ T* 0 Tc -0.0007 Tw [(discussed more extensively below)47.3(.)]TJ 0 -2.4 TD -0.0016 Tw (Host factors)Tj 0 -1.2 TD -0.0128 Tc -0.0694 Tw [(The ef)15.5(fect of host factors on )18.1(TE spread is well demonstrated)]TJ T* 0 Tc 0.0528 Tw [(by the dif)17(ference in the taxonomic range of host species)]TJ T* -0.0075 Tc -0.0695 Tw (for the )Tj /F3 1 Tf 9 0 1.913 9 337.89 478.275 Tm -0.0403 Tc 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 345.33 478.275 Tm -0.0075 Tc -0.0695 Tw (element and )Tj /F3 1 Tf 9 0 1.913 9 395.31 478.275 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 425.37 478.275 Tm 0 Tc ( )Tj 0.201 0 TD -0.0075 Tc -0.0695 Tw [(families. )61.3(Although both these)]TJ -12.8344 -1.2 TD 0 Tc -0.0257 Tw [(families are Class II )12.5(TEs, )]TJ /F3 1 Tf 9 0 1.913 9 412.41 467.475 Tm -0.0337 Tc 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 420.33 467.475 Tm 0 Tc -0.0257 Tw (elements are phylogenetically)Tj -12.0733 -1.2 TD -0.0124 Tc -0.0694 Tw (restricted to Diptera \(mostly Drosophilids\), whereas )Tj /F3 1 Tf 9 0 1.913 9 508.59 456.675 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 311.67 445.875 Tm 0 Tc 0.0755 Tw (has been found in many animal phyla \(Robertson )Tj /F3 1 Tf 9 0 1.913 9 515.73 445.875 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 535.9501 445.875 Tm 0 Tw (,)Tj -24.92 -1.2 TD 0.0787 Tw (2002\). Despite their very similar structure and life cycle,)Tj T* -0.0065 Tc -0.0695 Tw [(these two families dif)16.1(fer in a major aspect: the transposition)]TJ T* 0.0047 Tc 0 Tw (of )Tj /F3 1 Tf 9 0 1.913 9 323.07 413.475 Tm (P )Tj /F1 1 Tf 9 0 0 9 332.91 413.475 Tm 0.1389 Tw (elements requires a host enzyme, IRBP)Tj /F3 1 Tf 9 0 1.913 9 499.29 413.475 Tm 0 Tc 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 503.07 413.475 Tm 0.0047 Tc (\(inverted)Tj -21.2667 -1.2 TD -0.0026 Tc -0.0694 Tw [(repeat binding protein\), which binds the element)13.3(s inverted)]TJ T* 0 Tc -0.0654 Tw [(terminal repeats and is responsible for the element)11.9(s initial)]TJ T* 0.0994 Tw (excision from its chromosomal locus \(Beall )Tj /F3 1 Tf 9 0 1.913 9 489.63 381.075 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 510.03 381.075 Tm (, 1994;)Tj -22.04 -1.2 TD -0.0137 Tc -0.0694 Tw (Beall and Rio, 1997\). In contrast, purified transposase alone)Tj T* -0.0122 Tc -0.0695 Tw [(is suf)17.8(ficient to support the mobility of )]TJ /F3 1 Tf 9 0 1.913 9 450.21 359.475 Tm (mariner in vitro)Tj /F1 1 Tf 9 0 0 9 506.67 359.475 Tm 0 Tc 0 Tw ( )Tj 0.1963 0 TD -0.0122 Tc (\(Lampe)Tj /F3 1 Tf 9 0 1.913 9 311.67 348.675 Tm 0 Tc 0.0875 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 332.01 348.675 Tm 0.0874 Tw [(, 1996\). )17.2(This striking distinction between these two)]TJ -2.26 -1.2 TD 0.0088 Tw (Class II elements may alone explain the wide distribution)Tj T* -0.0019 Tc 0 Tw (of )Tj /F3 1 Tf 9 0 1.913 9 321.03 327.075 Tm (mariner)Tj /F1 1 Tf 9 0 0 9 351.39 327.075 Tm 0 Tc ( )Tj 0.2067 0 TD -0.0019 Tc -0.0694 Tw (in contrast to that of )Tj /F3 1 Tf 9 0 1.913 9 431.31 327.075 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 437.31 327.075 Tm -0.0019 Tc -0.0694 Tw (. Indeed, if )Tj /F3 1 Tf 9 0 1.913 9 479.67 327.075 Tm -0.0337 Tc 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 487.23 327.075 Tm -0.0019 Tc -0.0694 Tw (elements are)Tj -19.5067 -1.2 TD 0.039 Tc 0.1389 Tw (transferred into the germ cells of a new host, their)Tj T* 0.0218 Tc (transposition and concomitant spread depend on the)Tj T* 0 Tc -0.0292 Tw (existence of a host protein with properties similar to those)Tj T* 0.0134 Tc 0.1389 Tw [(of IRBP)125.5(. )15.1(The failure of the canonical )]TJ /F3 1 Tf 9 0 1.913 9 470.31 283.875 Tm -0.027 Tc 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 479.97 283.875 Tm 0.0134 Tc 0.1389 Tw (element to be)Tj -18.7 -1.2 TD 0 Tc 0.1364 Tw (mobilized in non-drosophilids following microinjection in)Tj T* 0.1126 Tw (the laboratory is possibly due to the lack of such factor)Tj T* -0.0098 Tc -0.0695 Tw [(\(OBrochta and Handler)49(, 1988\). Interestingly)71.7(, )]TJ /F3 1 Tf 9 0 1.913 9 484.47 251.475 Tm 0 Tc 0 Tw (T)Tj 0.5 0 TD -0.0098 Tc (c1 )Tj /F1 1 Tf 9 0 0 9 500.13 251.475 Tm (elements,)Tj -20.94 -1.2 TD 0 Tc 0.0019 Tw (which belong to Class II )Tj /F3 1 Tf 9 0 1.913 9 409.35 240.675 Tm 0 Tw [(mariner)15.7(-T)97.3(c1)]TJ /F1 1 Tf 9 0 0 9 456.87 240.675 Tm 0.0001 Tc 0.0018 Tw ( superfamily and are)Tj -16.1333 -1.2 TD 0.0142 Tc 0.1389 Tw (widely distributed among animals and fungi \(Plasterk,)Tj T* 0.0365 Tc (1996\), much like their )Tj /F3 1 Tf 9 0 1.913 9 412.77 219.075 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 445.59 219.075 Tm 0 Tc ( )Tj 0.4534 0 TD 0.0365 Tc 0.1389 Tw (cousins require only)Tj -15.3334 -1.2 TD 0 Tc 0.1202 Tw [(transposase activity for their mobility \(V)55(os )]TJ /F3 1 Tf 9 0 1.913 9 486.27 208.275 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 506.85 208.275 Tm (, 1996\).)Tj -21.6867 -1.2 TD -0.0087 Tc -0.0695 Tw (This apparent minimal requirement for mobility may explain)Tj T* 0 Tc 0.0894 Tw (why horizontal transfer between distantly related taxa is)Tj T* -0.0106 Tw (relatively common for )Tj /F3 1 Tf 9 0 1.913 9 399.45 175.875 Tm -0.0977 Tc 0 Tw [(Tc)-97.7(1)]TJ /F1 1 Tf 9 0 0 9 413.61 175.875 Tm 0 Tc ( )Tj 0.2674 0 TD (and )Tj /F3 1 Tf 9 0 1.913 9 433.47 175.875 Tm (mariner)Tj /F1 1 Tf 9 0 0 9 463.95 175.875 Tm (.)Tj -15.0267 -1.2 TD 0.0209 Tc 0.1389 Tw [(The influence of host factors on )18.5(TE mobility was)]TJ -1.8933 -1.2 TD 0 Tc 0.0448 Tw (illustrated by comparing the dynamics of the spread of a)Tj T* 0.1314 Tw (TE family that was introduced into the genomes of two)Tj T* -0.004 Tc -0.0695 Tw (closely related species. When )Tj /F3 1 Tf 9 0 1.913 9 429.63 132.675 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 437.4901 132.675 Tm -0.0695 Tw (elements were introduced)Tj -13.98 -1.2 TD 0.0102 Tc 0.1389 Tw (into laboratory populations of )Tj /F3 1 Tf 9 0 1.913 9 437.67 121.875 Tm (D. melanogaster)Tj /F1 1 Tf 9 0 0 9 506.31 121.875 Tm 0 Tc 0 Tw ( )Tj 0.4271 0 TD 0.0102 Tc 0.1389 Tw (and its)Tj -22.0538 -1.2 TD 0.0057 Tc (sibling species )Tj /F3 1 Tf 9 0 1.913 9 376.05 111.075 Tm (D. simulans)Tj /F1 1 Tf 9 0 0 9 424.83 111.075 Tm (, there was a dramatic and)Tj -12.5733 -1.2 TD 0.0129 Tc [(repeatable dif)11.5(ference in the population dynamics of )]TJ /F3 1 Tf 9 0 1.913 9 532.47 100.275 Tm 0 Tc 0 Tw (P)Tj ET endstream endobj 22 0 obj << /Type /Page /Parent 93 0 R /Resources 23 0 R /Contents 24 0 R /Thumb 70 0 R /MediaBox [ 0 0 595 842 ] /CropBox [ 0 0 595 842 ] /Rotate 0 >> endobj 23 0 obj << /ProcSet [ /PDF /Text ] /Font << /F1 114 0 R /F2 113 0 R /F3 128 0 R >> /ExtGState << /GS1 139 0 R /GS2 138 0 R >> /ColorSpace << /Cs8 112 0 R >> >> endobj 24 0 obj << /Length 14863 >> stream BT /F1 1 Tf 9 0 0 9 55.93 779.115 Tm 0 0 0 1 k /GS2 gs -0.0001 Tc -0.0003 Tw (8 Silva )Tj /F3 1 Tf 9 0 1.913 9 90.43 779.115 Tm 0 Tc (et al)Tj /F1 1 Tf 9 0 0 9 107.47 779.115 Tm 0 Tw (.)Tj /F2 1 Tf 37.9351 29.4096 -29.4096 37.9351 34.496 531.1378 Tm 2 Tr 0 0 0 0.45 K 0 J 0 j 0.576 w 10 M []0 d /Cs8 cs 0 scn /GS1 gs 0.0686 Tc -0.0001 Tw (PAGE PROOFS)Tj /F1 1 Tf 9 0 0 9 56.53 748.275 Tm 0 Tr 0 0 0 1 k /GS2 gs 0 Tc -0.0602 Tw (element \(Kimura and Kidwell, 1994; Higuet )Tj /F3 1 Tf 9 0 1.913 9 226.39 748.275 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 245.35 748.275 Tm (, 1996\). )Tj /F3 1 Tf 9 0 1.913 9 277.33 748.275 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 56.53 737.475 Tm -0.0612 Tw (elements are significantly more active in )Tj /F3 1 Tf 9 0 1.913 9 215.29 737.475 Tm (D. melanogaster)Tj /F1 1 Tf 9 0 0 9 280.81 737.475 Tm 0 Tw (,)Tj -24.92 -1.2 TD -0.0079 Tc -0.0695 Tw (reaching a higher copy number per cell than in )Tj /F3 1 Tf 9 0 1.913 9 235.21 726.675 Tm (D. simulans)Tj /F1 1 Tf 9 0 0 9 280.81 726.675 Tm 0 Tc 0 Tw (.)Tj -24.92 -1.2 TD -0.054 Tw (These results suggest that, in addition to the transposase,)Tj T* 0.042 Tw (host-specific factors are necessary to support )Tj /F3 1 Tf 9 0 1.913 9 242.41 705.0751 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 251.29 705.0751 Tm (element)Tj -21.64 -1.2 TD 0.0197 Tc 0.1389 Tw [(mobility at a suf)17.9(ficient level to ensure its spread and)]TJ T* 0.0013 Tc (subsequent persistence in a species, once it has been)Tj T* 0 Tc 0.0047 Tw [(introduced by horizontal transfer)51.5(. )14(These host factors may)]TJ T* -0.0009 Tc -0.0694 Tw (be in the form of a facilitator of transposition, such as IRBP)Tj T* 0 Tc 0.094 Tw (\(Badge and Brookfield, 1997\) or may be related to host)Tj T* -0.0026 Tc -0.0694 Tw (systems not directly related to transposition, such as those)Tj T* 0.0191 Tc 0.1389 Tw [(related to DNA)50.3( repair)60.1(. Quesneville and )51.8(Anxolabhre)]TJ T* 0 Tc 0.0137 Tw (\(1997\) have suggested that a species ability to deal with)Tj T* -0.061 Tw (the damage induced by )Tj /F3 1 Tf 9 0 1.913 9 150.43 607.875 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 158.41 607.875 Tm -0.061 Tw (element excision can determine)Tj -11.32 -1.2 TD 0.0002 Tw [(the success or failure of horizontal transfer)49.3(.)]TJ 0 -2.4 TD -0.0019 Tc -0.0695 Tw (Regulation of copy number: repression of transposition )Tj /F3 1 Tf 9 0 1.913 9 271.87 575.475 Tm 0 Tw (vs)Tj /F1 1 Tf 9 0 0 9 280.81 575.475 Tm 0 Tc (.)Tj -24.92 -1.2 TD -0.0003 Tc (selection)Tj T* 0.0105 Tc 0.1389 Tw (Experimental data suggest that the initial stage of the)Tj T* -0.0045 Tc -0.0694 Tw (invasion of a naive genome by )Tj /F3 1 Tf 9 0 1.913 9 175.63 543.075 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 183.43 543.075 Tm -0.0694 Tw (elements is characterized)Tj -14.1 -1.2 TD 0 Tc 0.0204 Tw (by a very high transposition rate, on the order of 10)Tj 6.3 0 0 6.3 262.03 535.395 Tm 0 Tw (-2)Tj 9 0 0 9 267.61 532.275 Tm ( )Tj 0.2984 0 TD (per)Tj -23.7517 -1.2 TD 0.0079 Tc 0.1389 Tw (element per generation \(Engels, 1989, and references)Tj T* 0 Tc 0.0609 Tw [(therein\). )13.6(That this may be the case for other )17(TE families)]TJ T* -0.0085 Tc -0.0694 Tw (\(even if not to the extreme seen in )Tj /F3 1 Tf 9 0 1.913 9 187.39 499.875 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 195.13 499.875 Tm -0.0694 Tw (elements\) is supported)Tj -15.4 -1.2 TD -0.0059 Tc [(by the very high copy number attained by some )13.7(TE families)]TJ T* 0 Tc 0.1337 Tw (soon after invasion, as mentioned above for )Tj /F3 1 Tf 9 0 1.913 9 242.11 478.275 Tm 0 Tw (mariner )Tj /F1 1 Tf 9 0 0 9 276.31 478.275 Tm (in)Tj /F3 1 Tf 9 0 1.913 9 56.53 467.475 Tm 0.0084 Tc (Dugesia)Tj /F1 1 Tf 9 0 0 9 90.13 467.475 Tm 0.1389 Tw [(. Eventually)75.8(, however)49.8(, copy number ceases to)]TJ -3.7333 -1.2 TD 0.0006 Tc [(increase. )13.3(This stabilization in copy number can be due)]TJ T* -0.0059 Tc -0.0694 Tw [(either to selection at the host level or to a )11.2(TE self-regulation)]TJ T* -0.0054 Tc (mechanism \(Charlesworth and Charlesworth, 1983\). In the)Tj T* 0 Tc 0.0571 Tw (case of selection host fitness is a decreasing function of)Tj T* 0.137 Tw [(TE number)49.7(. In the case of self-regulation, transposition)]TJ T* 0.0002 Tw [(rate is a decreasing function of )10.9(TE number)59.6(.)]TJ 1.8933 -1.2 TD -0.0048 Tw (Regulation of transposition of Class I elements is still)Tj -1.8933 -1.2 TD 0.092 Tw [(poorly understood. However)55.7(, there is a growing body of)]TJ T* 0.0389 Tc 0.139 Tw [(evidence that suggests that self-regulation is not a)]TJ T* -0.005 Tc -0.0695 Tw (significant force in the stabilization of copy number in these)Tj T* 0 Tc 0.0296 Tw (families, since there seems to be no negative correlation)Tj T* 0.0099 Tw (between transposition rate and copy number \(Bimont )Tj /F3 1 Tf 9 0 1.913 9 275.83 337.875 Tm 0 Tw (et)Tj -24.1116 -1.2 TD 0.0058 Tc (al.)Tj /F1 1 Tf 9 0 0 9 66.19 327.075 Tm 0.1389 Tw [(, 1997; V)13.2(ieira and Bimont, 1997; Pasyukova )]TJ /F3 1 Tf 9 0 1.913 9 259.69 327.075 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 280.81 327.075 Tm 0 Tc 0 Tw (,)Tj -24.92 -1.2 TD -0.0541 Tw [(1998\). Rather)53.2(, selection has been suggested as the major)]TJ T* 0.0022 Tw [(force controlling Class I element copy number)51.8(, especially)]TJ T* 0.0259 Tc 0.1389 Tw (because of the fitness costs associated with ectopic)Tj T* 0 Tc 0.0115 Tw (recombination \(Charlesworth and Lapid, 1989; Maside )Tj /F3 1 Tf 9 0 1.913 9 275.83 283.875 Tm 0 Tw (et)Tj -24.1116 -1.2 TD (al.)Tj /F1 1 Tf 9 0 0 9 66.07 273.075 Tm -0.0537 Tw (, 2001\). Repression of transposition of Class I elements)Tj -1.06 -1.2 TD -0.0008 Tc -0.0695 Tw (can also be achieved by means of host-encoded peptides.)Tj T* 0 Tc 0.1072 Tw (This is the case for )Tj /F3 1 Tf 9 0 1.913 9 139.39 251.475 Tm 0 Tw (gypsy)Tj /F1 1 Tf 9 0 0 9 162.91 251.475 Tm 0.1072 Tw (, the transposition of which is)Tj -11.82 -1.2 TD 0.0247 Tc 0.1389 Tw (regulated by an X-linked gene called )Tj /F3 1 Tf 9 0 1.913 9 220.39 240.675 Tm 0 Tw (flamenco)Tj /F1 1 Tf 9 0 0 9 258.73 240.675 Tm 0 Tc ( )Tj 0.4416 0 TD 0.0247 Tc (in )Tj /F3 1 Tf 9 0 1.913 9 274.09 240.675 Tm (D.)Tj -23.9183 -1.2 TD 0 Tc (melanogaster)Tj /F1 1 Tf 9 0 0 9 111.13 229.875 Tm ( )Tj 0.2511 0 TD (\(Prudhomme )Tj /F3 1 Tf 9 0 1.913 9 169.63 229.875 Tm -0.0269 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 188.89 229.875 Tm [(, 1995\). V)75.5(arying rates of)]TJ -14.7067 -1.2 TD -0.0052 Tc -0.0695 Tw (transposition of these elements among natural populations)Tj T* 0 Tc 0.0141 Tw (may be due to the presence of permissive and restrictive)Tj T* -0.008 Tc -0.0695 Tw (alleles, which are kept in balance by mutation and selection)Tj T* -0.0069 Tc -0.0694 Tw [(\(Nuzhdin, 1999\). )14.1(This is probably a tight balance, delicately)]TJ T* 0 Tc -0.0608 Tw (modulated by host TE interactions and hence specific for)Tj T* 0.1124 Tw [(each such pair)54.5(, as suggested by the recent finding that)]TJ /F3 1 Tf 9 0 1.913 9 56.53 154.275 Tm 0 Tw (flamenco)Tj /F1 1 Tf 9 0 0 9 93.07 154.275 Tm ( )Tj 0.2712 0 TD -0.0068 Tw (cannot repress transposition of )Tj /F3 1 Tf 9 0 1.913 9 220.87 154.275 Tm 0 Tw (gypsy)Tj /F1 1 Tf 9 0 0 9 244.33 154.275 Tm ( )Tj 0.2712 0 TD (elements)Tj -21.1379 -1.2 TD -0.0218 Tw (from other )Tj /F3 1 Tf 9 0 1.913 9 99.67 143.475 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 142.69 143.475 Tm ( )Tj 0.2562 0 TD -0.0218 Tw (species when these are introduced)Tj -9.8295 -1.2 TD 0.0279 Tc 0 Tw (in )Tj /F3 1 Tf 9 0 1.913 9 68.05 132.675 Tm 0.139 Tw (D. melanogaster )Tj /F1 1 Tf 9 0 0 9 143.11 132.675 Tm 0 Tw (\(Mejlumian )Tj /F3 1 Tf 9 0 1.913 9 193.63 132.675 Tm 0.139 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 215.89 132.675 Tm [(, 2002\). Finally)69.5(,)]TJ -17.7067 -1.2 TD 0.0223 Tc 0.1389 Tw [(methylation of )15.6(TE sequences can also play a role in)]TJ T* 0 Tc 0.0304 Tw (repression of transposition in some organisms \(Labrador)Tj T* -0.0007 Tw (and Corces, 1997; Matzke )Tj /F3 1 Tf 9 0 1.913 9 164.11 100.275 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 183.61 100.275 Tm (, 1999\).)Tj 15.4933 72 TD -0.0108 Tc -0.0695 Tw (Self-regulation depends on repression of transposition,)Tj -1.8867 -1.2 TD 0.0618 Tc 0.1389 Tw (which encompasses many processes that reduce)Tj T* 0.0032 Tc 0.139 Tw (transposition rate to varying degrees. Self-regulation is)Tj T* 0 Tc 0.0143 Tw (known for Class II families. )Tj /F3 1 Tf 9 0 1.913 9 416.23 715.875 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 424.87 715.875 Tm 0.0143 Tw (elements in particular have)Tj -13.2 -1.2 TD 0.0417 Tw (been intensively studied and several types of repressors)Tj T* 0.0071 Tc 0.1389 Tw [(have been defined. )16.3(T)58.1(ype I repressors correspond to a)]TJ T* 0 Tc -0.0063 Tw (truncated version of the )Tj /F3 1 Tf 9 0 1.913 9 402.43 683.475 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 408.43 683.475 Tm -0.0063 Tw (-encoded transposase, and are)Tj -11.3733 -1.2 TD 0.0524 Tw [(produced by alternative splicing of the element)17.4(s mRNA;)]TJ T* 0.0206 Tc 0.1389 Tw (this yields a 66-kD repressor protein that prevents )Tj /F3 1 Tf 9 0 1.913 9 526.81 661.875 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 306.07 651.075 Tm 0.0784 Tw (element transcription \(Misra and Rio, 1990; Gloor )Tj /F3 1 Tf 9 0 1.913 9 510.13 651.075 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 530.29 651.075 Tm 0 Tw (,)Tj -24.9133 -1.2 TD -0.0079 Tc -0.0695 Tw (1993; Siebel )Tj /F3 1 Tf 9 0 1.913 9 356.41 640.275 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 374.83 640.275 Tm (, 1994; Roche )Tj /F3 1 Tf 9 0 1.913 9 430.57 640.275 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 448.99 640.275 Tm [(, 1995\). )18.7(T)36.4(ransposition)]TJ -15.88 -1.2 TD 0 Tc 0.0347 Tw (can also be repressed by a series of internally deleted )Tj /F3 1 Tf 9 0 1.913 9 526.81 629.475 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 306.07 618.675 Tm 0.0106 Tc 0.1389 Tw [(elements, called type II repressors. )17.8(These are usually)]TJ T* -0.0037 Tc -0.0695 Tw (byproducts of the repair of the double-stranded DNA break)Tj T* 0 Tc 0.0948 Tw (that occurs during )Tj /F3 1 Tf 9 0 1.913 9 382.63 597.075 Tm -0.0337 Tc 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 391.63 597.075 Tm 0 Tc 0.0948 Tw [(element transposition. Supposedly)68.1(,)]TJ -9.5067 -1.2 TD 0.0033 Tc 0.1389 Tw (type II repressors act by binding the transposase itself)Tj T* 0 Tc 0 Tw (\(Rasmusson )Tj /F3 1 Tf 9 0 1.913 9 359.77 575.475 Tm 0.0781 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 379.99 575.475 Tm [(, 1993; )58.9(Andrews and Gloor)55.2(, 1995\), by)]TJ -8.2133 -1.2 TD 0.0113 Tc 0.139 Tw [(binding the element sequence thus out-competing the)]TJ T* 0 Tc 0.0015 Tw (transposase enzyme \(Lee )Tj /F3 1 Tf 9 0 1.913 9 412.15 553.875 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 431.65 553.875 Tm (, 1996\), and by antisense)Tj -13.9533 -1.2 TD -0.0537 Tw [(RNA)57.7( interference \(Simmons )]TJ /F3 1 Tf 9 0 1.913 9 418.63 543.075 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 437.65 543.075 Tm [(, 1996\). )15(The )]TJ /F3 1 Tf 9 0 1.913 9 487.09 543.075 Tm 0 Tw [(KP)40.7( )]TJ /F1 1 Tf 9 0 0 9 500.83 543.075 Tm (element)Tj -21.64 -1.2 TD -0.0125 Tc -0.0695 Tw (is the most abundant type II repressor in natural populations)Tj T* 0 Tc 0 Tw (of )Tj /F3 1 Tf 9 0 1.913 9 316.03 521.475 Tm -0.0028 Tw (D. melanogaster)Tj /F1 1 Tf 9 0 0 9 382.03 521.475 Tm -0.0001 Tc [( \(Engels, 1989\). Finally)71(, the titration of)]TJ -8.44 -1.2 TD 0.0695 Tc 0.7041 Tw (transposase by its binding to defective or)Tj T* 0.0247 Tc 0 Tw (extrachromosomal )Tj /F3 1 Tf 9 0 1.913 9 387.61 499.875 Tm (P )Tj /F1 1 Tf 9 0 0 9 397.81 499.875 Tm 0.1389 Tw (element copies might also help)Tj -10.1933 -1.2 TD 0 Tc -0.0438 Tw (reduce transposition rates \(Simmons and Bucholz, 1985\).)Tj 1.8867 -1.2 TD -0.003 Tc -0.0695 Tw (Repression of transposition in other Class II elements,)Tj -1.8867 -1.2 TD 0 Tc 0.1126 Tw (even though not as well studied, seems to rely on self-)Tj T* 0.0926 Tw (regulation mechanisms as well \(for reviews see Hartl )Tj /F3 1 Tf 9 0 1.913 9 525.31 456.675 Tm 0 Tw (et)Tj -24.1049 -1.2 TD 0.011 Tc (al.)Tj /F1 1 Tf 9 0 0 9 315.85 445.875 Tm 0.1389 Tw [(, 1997; Labrador and Corces, 1997\). )12.4(These include)]TJ -1.0867 -1.2 TD 0.0001 Tc 0.004 Tw [(alternative splicing of the elements)18.7(s mRNA)52.9( that can give)]TJ T* 0 Tc -0.0606 Tw (rise to either repressor peptides or transposase \(Mason )Tj /F3 1 Tf 9 0 1.913 9 525.31 424.275 Tm 0 Tw (et)Tj -24.1049 -1.2 TD 0.0695 Tc (al.)Tj /F1 1 Tf 9 0 0 9 317.47 413.475 Tm 0.1939 Tw (, 1991\), negative feedback dependent on the)Tj -1.2667 -1.2 TD 0.0258 Tc 0.1389 Tw (concentration of transposase \(Lohe and Hartl, 1996;)Tj T* 0.0188 Tc (Labrador and Corces, 1997\), transposase titration by)Tj T* 0.0142 Tc (methods such as dominant-negative complementation)Tj T* 0 Tc 0 Tw (\(Lohe )Tj /F3 1 Tf 9 0 1.913 9 332.47 370.275 Tm 0.099 Tw (et al.)Tj /F1 1 Tf 9 0 0 9 352.87 370.275 Tm (, 1997\), and/or RNA interference \(Jensen )Tj /F3 1 Tf 9 0 1.913 9 525.31 370.275 Tm 0 Tw (et)Tj -24.1049 -1.2 TD (al.)Tj /F1 1 Tf 9 0 0 9 315.55 359.475 Tm 0.0592 Tw [(, 1999\). )14.2(These studies support the contention that the)]TJ -1.0533 -1.2 TD 0.0032 Tc 0.139 Tw (mobility of Class II elements is, to a large extent, self-)Tj T* 0.0003 Tc 0 Tw (regulated.)Tj /F2 1 Tf 0 -2.4 TD 0 Tc 0.0006 Tw [(Correlates of Horizontal T)52.6(ransfer)]TJ /F1 1 Tf T* -0.0029 Tc -0.0695 Tw (As described above, horizontal transfer is apparently more)Tj 0 -1.2 TD 0 Tc -0.0467 Tw [(frequent among Class II than Class I )18.3(TEs, and this pattern)]TJ T* 0.0168 Tc 0.139 Tw [(seems to be due in part to the dif)14.8(ferent transposition)]TJ T* 0 Tc -0.012 Tw [(mechanisms used by the two classes. )56.6(Another reason for)]TJ T* -0.0457 Tw [(this dif)17.6(ference may be that while Class II elements appear)]TJ T* 0.115 Tw [(to be mostly self-regulating, Class I )22.5(TEs are not. Could)]TJ T* 0.0387 Tw [(this dif)22.1(ference in regulatory mechanism be related to the)]TJ T* -0.0169 Tw [(incidence of horizontal transfer? Possibly)73.4(, if the degree to)]TJ T* 0.0957 Tw [(which self-regulation represses transposition is ef)12.9(fective)]TJ T* 0.0003 Tc 0 Tw (enough.)Tj 1.8867 -1.2 TD 0 Tc 0.0423 Tw [(In order for a )18.8(TE lineage to persist through time, the)]TJ -1.8867 -1.2 TD -0.019 Tw (rate at which new functional elements arise must balance)Tj T* 0.0536 Tc 0.1389 Tw (the rate at which they are lost \(Charlesworth and)Tj T* 0 Tc -0.0258 Tw [(Charlesworth, 1983\). )54.2(A)53.7( functional element \(those capable)]TJ T* -0.0139 Tw (of transposition and which encode functional enzymes\) is)Tj T* 0.0387 Tw (lost due to a variety of processes, which include random)Tj T* 0.0713 Tw (loss \()Tj /F3 1 Tf 9 0 1.913 9 328.21 121.875 Tm 0 Tw (e.g.)Tj /F1 1 Tf 9 0 0 9 343.21 121.875 Tm 0.0713 Tw (, failure of the host to reproduce\), substitutions)Tj -4.1267 -1.2 TD 0.0291 Tc 0.1389 Tw (and insertions/deletions that render the transposase)Tj T* 0.0072 Tc (inactive, excision \()Tj /F3 1 Tf 9 0 1.913 9 383.41 100.275 Tm 0 Tw (e.g.)Tj /F1 1 Tf 9 0 0 9 398.65 100.275 Tm 0.1389 Tw (, a Class II element that, during)Tj ET endstream endobj 25 0 obj << /Type /Page /Parent 93 0 R /Resources 26 0 R /Contents 27 0 R /Thumb 72 0 R /MediaBox [ 0 0 595 842 ] /CropBox [ 0 0 595 842 ] /Rotate 0 >> endobj 26 0 obj << /ProcSet [ /PDF /Text /ImageC /ImageI ] /Font << /F1 114 0 R /F2 113 0 R /F3 128 0 R >> /XObject << /Im2 28 0 R >> /ExtGState << /GS1 139 0 R /GS2 138 0 R >> /ColorSpace << /Cs8 112 0 R /Cs10 54 0 R >> /Properties << /MC2 30 0 R >> >> endobj 27 0 obj << /Length 11200 >> stream BT /F1 1 Tf 9 0 0 9 342.51 780.5551 Tm 0 0 0 1 k /GS2 gs 0 Tc -0.0155 Tw [(Horizontal Transfer of Transposable Elements 9)]TJ /F2 1 Tf 37.9351 29.4096 -29.4096 37.9351 42.9423 542.4378 Tm 2 Tr 0 0 0 0.45 K 0 J 0 j 0.576 w 10 M []0 d /Cs8 cs 0 scn /GS1 gs 0.0686 Tc -0.0001 Tw (PAGE PROOFS)Tj /F1 1 Tf 9 0 0 9 62.19 748.275 Tm 0 Tr 0 0 0 1 k /GS2 gs 0.0235 Tc 0.1389 Tw (transposition, fails to re-insert\) and selection against)Tj 0 -1.2 TD 0 Tc -0.0374 Tw (individual hosts with specific deleterious insertions or with)Tj T* -0.019 Tw [(too many )19.8(TE copies. When the rate of transposition is not)]TJ T* 0.0384 Tw [(high enough to counteract the ef)15(fect of these processes,)]TJ T* 0.0839 Tw [(the )18.6(TE lineage will eventually go extinct. )56.8(Among Class I)]TJ T* 0.0851 Tw (elements, the rate of transposition is apparently at least)Tj T* 0.125 Tw (one order of magnitude larger that the rate of excision,)Tj T* 0.0695 Tc 0.2707 Tw [(and selection keeps )17.4(TE copy number in check)]TJ T* 0 Tc -0.0005 Tw [(\(Charlesworth and Langley)68(, 1991\).)]TJ 1.8867 -1.2 TD -0.0111 Tc -0.0694 Tw (A relevant question is whether the self-regulating Class)Tj -1.8867 -1.2 TD 0.0046 Tc 0.139 Tw [(II )20.2(TEs can maintain a rate of transposition that is high)]TJ T* 0 Tc 0.0652 Tw (enough to prevent lineage extinction, once repression is)Tj T* -0.0139 Tw (established. Not much is known about transposition rates)Tj T* -0.0028 Tw [(of Class II )15.5(TEs in natural population. However)57.4(, laboratory)]TJ T* -0.0123 Tw (studies show that a few generations after the introduction)Tj T* 0.0281 Tc 0 Tw (of )Tj /F3 1 Tf 9 0 1.913 9 74.19 586.275 Tm (P )Tj /F1 1 Tf 9 0 0 9 84.45 586.275 Tm 0.1389 Tw (elements into naive populations the )Tj /F3 1 Tf 9 0 1.913 9 243.39 586.275 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 253.65 586.275 Tm (cytotype)Tj -21.2733 -1.2 TD 0.0056 Tc 0.1389 Tw (becomes established, a condition characterized by the)Tj T* 0.0077 Tc 0.139 Tw (suppression of )Tj /F3 1 Tf 9 0 1.913 9 126.81 564.675 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 136.65 564.675 Tm 0.139 Tw (element transposition \(Engels, 1979;)Tj -8.2733 -1.2 TD 0.0163 Tc 0.1389 Tw (Kidwell, 1985; Engels, 1989, and references therein\).)Tj T* 0.0331 Tc (Although not thoroughly understood, the )Tj /F3 1 Tf 9 0 1.913 9 242.97 543.075 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 253.35 543.075 Tm (cytotype)Tj -21.24 -1.2 TD 0 Tc -0.0195 Tw (condition is maternally inherited and depends on both the)Tj T* 0.0076 Tw (presence and the location of )Tj /F3 1 Tf 9 0 1.913 9 178.11 521.475 Tm 0 Tw (P )Tj /F1 1 Tf 9 0 0 9 186.69 521.475 Tm 0.0076 Tw (element insertions, and is)Tj -13.8333 -1.2 TD 0.0506 Tw (related to the transmission of repressor peptides or their)Tj T* -0.0306 Tw [(mRNA)57.4( to the zygote, through the oocyte)14.7(s cytoplasm \(see)]TJ T* -0.054 Tw (Labrador and Corces, 1997 for a review; Ronsseray )Tj /F3 1 Tf 9 0 1.913 9 267.39 489.075 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 286.47 489.075 Tm 0 Tw (,)Tj -24.92 -1.2 TD 0.0203 Tc 0.1389 Tw (1998; Ronsseray )Tj /F3 1 Tf 9 0 1.913 9 138.15 478.275 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 159.99 478.275 Tm (, 2001, Simmons )Tj /F3 1 Tf 9 0 1.913 9 236.73 478.275 Tm (et al)Tj /F1 1 Tf 9 0 0 9 255.87 478.275 Tm (, 2002\).)Tj -21.52 -1.2 TD 0 Tc 0 Tw (T)Tj 0.5867 0 TD 0.0076 Tc 0.139 Tw [(runcated peptides with putative repressor capabilities)]TJ -0.5867 -1.2 TD 0 Tc -0.0234 Tw (have also been found in )Tj /F3 1 Tf 9 0 1.913 9 159.21 456.675 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 202.23 456.675 Tm ( )Tj 0.2546 0 TD -0.0234 Tw (species other than )Tj /F3 1 Tf 9 0 1.913 9 279.93 456.675 Tm 0 Tw (D.)Tj -23.9383 -1.2 TD 0.0159 Tc (melanogaster)Tj /F1 1 Tf 9 0 0 9 118.47 445.875 Tm 0.1389 Tw (, which shows that repressors can arise)Tj -6.2533 -1.2 TD 0 Tc -0.0091 Tw [(repeatedly \(Nouaud and )52.2(Anxolabhre, 1997\). Moreover)50.7(,)]TJ T* 0.068 Tw (the presence of the )Tj /F3 1 Tf 9 0 1.913 9 144.21 424.275 Tm 0 Tw (KP)Tj /F1 1 Tf 9 0 0 9 156.21 424.275 Tm ( )Tj 0.346 0 TD 0.068 Tw (repressor at a high frequency in)Tj 16.9273 36 TD -0.0023 Tc -0.0694 Tw (populations of )Tj /F3 1 Tf 9 0 1.913 9 368.67 748.275 Tm (D. melanogaster)Tj /F1 1 Tf 9 0 0 9 433.77 748.275 Tm 0 Tc 0 Tw ( )Tj 0.2063 0 TD -0.0023 Tc -0.0694 Tw (worldwide provide support)Tj -13.7729 -1.2 TD 0 Tc 0.0695 Tw (for the idea that new repressors can quickly spread and)Tj T* -0.0294 Tw (be maintained by selection \(Black )Tj /F3 1 Tf 9 0 1.913 9 446.97 726.675 Tm (et al.)Tj /F1 1 Tf 9 0 0 9 466.17 726.675 Tm (, 1987; Jackson )Tj /F3 1 Tf 9 0 1.913 9 530.97 726.675 Tm 0 Tw (et)Tj -24.1116 -1.2 TD (al.)Tj /F1 1 Tf 9 0 0 9 321.21 715.875 Tm -0.0272 Tw [(, 1988\). )14.6(These studies strongly suggest that repression)]TJ -1.06 -1.2 TD 0 Tw (of )Tj /F3 1 Tf 9 0 1.913 9 321.27 705.0751 Tm (P)Tj /F1 1 Tf 9 0 0 9 327.27 705.0751 Tm ( )Tj 0.2264 0 TD -0.0516 Tw [(element transposition can be quite ef)16.9(fective. Whether)]TJ -1.9597 -1.2 TD -0.054 Tw (or not repression of transposition is as strong in other self-)Tj T* -0.0002 Tw (regulated Class II elements remains to be determined.)Tj 1.8933 -1.2 TD -0.0121 Tw (Finally it should be noted that complete repression of)Tj -1.8933 -1.2 TD 0.1024 Tw [(transposition may not be required for extinction of a )13(TE)]TJ T* -0.054 Tw (lineage. Kaplan and collaborators \(1985\) have shown that)Tj T* -0.0081 Tc -0.0695 Tw [(self-regulating )12.3(TE lineages are expected to go extinct when)]TJ T* 0.0682 Tc 0.1389 Tw (non-functional elements can be transposed by a)Tj T* -0.0108 Tc -0.0694 Tw (transposase produced in )Tj /F3 1 Tf 9 0 1.913 9 408.57 618.675 Tm 0 Tw (trans)Tj /F1 1 Tf 9 0 0 9 428.07 618.675 Tm (, )Tj /F3 1 Tf 9 0 1.913 9 432.27 618.675 Tm (i.e.)Tj /F1 1 Tf 9 0 0 9 443.91 618.675 Tm -0.0694 Tw (, a transposase encoded)Tj -14.6933 -1.2 TD 0 Tc 0.0199 Tw [(by another element. )9.1(This is the case for )]TJ /F3 1 Tf 9 0 1.913 9 473.07 607.875 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 479.07 607.875 Tm ( )Tj 0.2979 0 TD 0.0199 Tw (elements and,)Tj -18.8979 -1.2 TD 0.0001 Tw [(probably)68.4(, for other Class II )17.4(TE families as well.)]TJ 1.8933 -1.2 TD -0.0141 Tc -0.0695 Tw (Assuming, for the sake of argument, that self-regulation)Tj -1.8933 -1.2 TD -0.0019 Tc (is indeed conducive to extinction, then continual horizontal)Tj T* 0 Tc -0.0454 Tw [(transfer becomes necessary for the survival of Class II )13.9(TE)]TJ T* 0.0365 Tc 0.1389 Tw (families. While Class I elements persist through an)Tj T* 0 Tc 0.1053 Tw (equilibrium between transposition and loss, Class II are)Tj T* 0.0137 Tc 0.1389 Tw [(always on the run \(Figure 2\). )48.9(As long as repression)]TJ T* -0.0022 Tc -0.0695 Tw [(persists, at least one )20.3(TE copy must find its way into a naive)]TJ T* 0 Tc 0.0387 Tw (genome, where repression is still absent, giving rise to a)Tj T* 0.0047 Tc 0.1389 Tw (new pool of functional copies to re-initiate the cycle of)Tj T* -0.0087 Tc -0.0694 Tw [(transposition and spread. )12.5(This scenario leads to a few quite)]TJ T* 0 Tc 0.0337 Tw (interesting predictions. First, because it depends on how)Tj T* -0.0067 Tc -0.0694 Tw (quickly repression of transposition arises, copy number per)Tj T* 0 Tc 0.0989 Tw (genome is likely to vary among populations and among)Tj T* -0.0467 Tw [(species. )12.7(There is already evidence that this is the case for)]TJ /F3 1 Tf 9 0 1.913 9 311.67 435.075 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 317.67 435.075 Tm ( )Tj 0.3258 0 TD 0.0478 Tw [(elements, the copy number of which is known to dif)10.5(fer)]TJ -0.9925 -1.2 TD -0.0027 Tc -0.0694 Tw (among species of the )Tj /F3 1 Tf 9 0 1.913 9 396.75 424.275 Tm 0 Tw (willistoni)Tj /F1 1 Tf 9 0 0 9 429.9901 424.275 Tm 0 Tc ( )Tj 0.2059 0 TD -0.0027 Tc -0.0694 Tw (species group \(Daniels and)Tj 7 0 0 7 62.19 164.955 Tm 0 Tc -0.0067 Tw (Figure 2. Stages of the life cycle of a hypothetical Class II, self-regulated, transposable element lineage. \()Tj /F3 1 Tf 7 0 1.4879 7 387.15 164.955 Tm 0 Tw (i)Tj /F1 1 Tf 7 0 0 7 388.71 164.955 Tm -0.0067 Tw [(\) )55.7(An element is transferred into a germline cell of)]TJ -46.6457 -1.2 TD -0.128 Tw [(host A. )-59.1(\()]TJ /F3 1 Tf 7 0 1.4879 7 86.91 156.555 Tm 0 Tw (ii)Tj /F1 1 Tf 7 0 0 7 90.03 156.555 Tm -0.0689 Tw [(\) Once the element has successfully integrated into the host DNA)56.7( and is expressed, transposition will start, with a concomitant)]TJ 54.3258 0 TD 0 Tw ( )Tj 0.2091 0 TD -0.0689 Tw (rapid increase in copy)Tj -58.5121 -1.2 TD -0.0622 Tw [(number)55.8(. \()]TJ /F3 1 Tf 7 0 1.4879 7 91.35 148.155 Tm 0 Tw (iii)Tj /F1 1 Tf 7 0 0 7 95.97 148.155 Tm -0.0622 Tw [(\) Repression of transposition arises and spreads throughout the host population. )54.9(As a result, the growth rate in copy number sl)]TJ 54.5614 0 TD (ows. \()Tj /F3 1 Tf 7 0 1.4879 7 496.17 148.155 Tm 0 Tw (iv)Tj /F1 1 Tf 7 0 0 7 501.27 148.155 Tm -0.0622 Tw [(\) )60.2(At this time)]TJ -62.7257 -1.2 TD -0.0516 Tw (the rate of loss of functional elements is higher than the rate at which they are created by transposition, and the number of f)Tj 53.2861 0 TD (unctional elements in the genome)Tj -53.2861 -1.2 TD -0.0636 Tw [(slowly decreases. )19.1(This process that can take many millions of years \(abbreviated period represented by a dashed line\). \()]TJ /F3 1 Tf 7 0 1.4879 7 427.17 131.355 Tm 0 Tw (v)Tj /F1 1 Tf 7 0 0 7 430.71 131.355 Tm -0.0636 Tw [(\) Finally)76.5(, no functional elements are)]TJ -52.6457 -1.2 TD -0.0148 Tw [(left in the genome of host )52.9(A, and this )13(TE lineage becomes extinct. Sometime between \()]TJ /F3 1 Tf 7 0 1.4879 7 329.79 122.955 Tm 0 Tw (ii)Tj /F1 1 Tf 7 0 0 7 332.91 122.955 Tm -0.0148 Tw (\) and \()Tj /F3 1 Tf 7 0 1.4879 7 352.89 122.955 Tm 0 Tw (v)Tj /F1 1 Tf 7 0 0 7 356.37 122.955 Tm -0.0148 Tw (\), a functional element may be transferred horizontally \(HT\))Tj -42.0257 -1.2 TD -0.0214 Tw [(to a new host and the process begins anew)50.8(. So, the death of the lineage in host )59.7(A)49.9( does not necessarily imply the death of the )15(T)]TJ 55.5796 0 TD [(E family)74.3(. Eventually)82(, another)]TJ -55.5796 -1.2 TD -0.0337 Tw [(functional element may escape to yet another host. If, at this time, repression of transposition has subsided in host )55.2(A, it can)]TJ 53.6543 0 TD 0 Tw ( )Tj 0.2443 0 TD -0.0337 Tw [(be invaded again. )15.3(The presence)]TJ -53.8986 -1.2 TD -0.0271 Tw [(of multiple, quite divergent )21.8(TE subfamilies of )]TJ /F3 1 Tf 7 0 1.4879 7 200.07 97.755 Tm 0 Tw (mariner)Tj /F1 1 Tf 7 0 0 7 223.77 97.755 Tm ( )Tj 0.2509 0 TD (and )Tj /F3 1 Tf 7 0 1.4879 7 238.95 97.755 Tm (P)Tj /F1 1 Tf 7 0 0 7 243.57 97.755 Tm ( )Tj 0.2509 0 TD -0.0271 Tw (elements in the same host is probably the signature of independent waves of horizontal transfer)Tj -26.1623 -1.2 TD 0 Tw (\(Clark)Tj /F3 1 Tf 7 0 1.4879 7 80.85 89.355 Tm -0.0001 Tc -0.0009 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 97.95 89.355 Tm 0 Tc -0.001 Tw (, 1995; Lampe)Tj /F3 1 Tf 7 0 1.4879 7 142.71 89.355 Tm -0.0001 Tc -0.0009 Tw ( et al.)Tj /F1 1 Tf 7 0 0 7 159.81 89.355 Tm 0 Tc -0.001 Tw (, 2001\).)Tj ET /EmbeddedDocument /MC2 BDC q 1 i 62.19 391.215 476.28 -208.32 re W n 1 G 0.456 w /GS1 gs 58.092 420.845 484.931 -247.067 re S q 476.28 0 0 209.6875 62.19 181.9833 cm /Im2 Do Q EMC Q endstream endobj 28 0 obj << /Type /XObject /Subtype /Image /Width 1046 /Height 460 /BitsPerComponent 8 /ColorSpace 54 0 R /Length 481161 >> stream                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                              endstream endobj 29 0 obj << /Length 776 >> stream SAMSON endstream endobj 30 0 obj << /MetaData 29 0 R /Title (SAMSON) /Creator (Adobe\37777777650 PageMaker\37777777650 6.5: LaserWriter 8 Z1-8.7.1) >> endobj 31 0 obj << /Type /Page /Parent 93 0 R /Resources 32 0 R /Contents 33 0 R /Thumb 74 0 R /MediaBox [ 0 0 595 842 ] /CropBox [ 0 0 595 842 ] /Rotate 0 >> endobj 32 0 obj << /ProcSet [ /PDF /Text ] /Font << /F1 114 0 R /F2 113 0 R /F3 128 0 R >> /ExtGState << /GS1 139 0 R /GS2 138 0 R >> /ColorSpace << /Cs8 112 0 R >> >> endobj 33 0 obj << /Length 12862 >> stream BT /F1 1 Tf 9 0 0 9 55.93 779.115 Tm 0 0 0 1 k /GS2 gs -0.0001 Tc -0.0003 Tw (10 Silva )Tj /F3 1 Tf 9 0 1.913 9 95.47 779.115 Tm 0 Tc (et al)Tj /F1 1 Tf 9 0 0 9 112.45 779.115 Tm 0 Tw (.)Tj /F2 1 Tf 37.9351 29.4096 -29.4096 37.9351 34.496 531.1378 Tm 2 Tr 0 0 0 0.45 K 0 J 0 j 0.576 w 10 M []0 d /Cs8 cs 0 scn /GS1 gs 0.0686 Tc -0.0001 Tw (PAGE PROOFS)Tj /F1 1 Tf 9 0 0 9 56.53 748.275 Tm 0 Tr 0 0 0 1 k /GS2 gs 0 Tc -0.0673 Tw [(Strausbaugh, 1986\). Second, in populations where the )12(TE)]TJ 0 -1.2 TD -0.0041 Tc -0.0695 Tw (copy number is stable, an increase in the number of copies)Tj T* 0 Tc 0.057 Tw (should be possible through the elimination of the source)Tj T* 0.0877 Tw [(of repression. Finally)67.1(, unless they are a recently formed)]TJ T* 0.0099 Tc 0.1389 Tw [(family)70.4(, self-regulated )18.1(TEs should all show evidence of)]TJ T* 0 Tc 0.0181 Tw [(horizontal transfer)54.5(. )14.1(The maximum length of time before a)]TJ T* 0.0042 Tc 0.1389 Tw (horizontal transfer is required \(time to extinction of the)Tj T* 0 Tc -0.0326 Tw (family\) will depend on several characteristics, such as the)Tj T* 0.0125 Tc 0.1389 Tw (number of copies per genome and the rate of loss of)Tj T* 0 Tc 0.0819 Tw (functional elements. In this context, the study of class II)Tj T* 0.004 Tc 0.1389 Tw (families in which transmission seems to be exclusively)Tj T* 0 Tc 0.0001 Tw (vertical, if any is found, would be extremely fruitful.)Tj /F2 1 Tf 0 -2.4 TD 0 Tw (Conclusions)Tj /F1 1 Tf 0 -2.8 TD 0.0099 Tc 0.1389 Tw [(Successful )14(TE horizontal transfer events depend on a)]TJ 0 -1.2 TD 0 Tc 0.1195 Tw (stable transfer between donor and recipient and on the)Tj T* 0.0016 Tw (subsequent spread throughout the new host populations.)Tj T* 0.004 Tw [(Although detected for all types of )12.9(TEs, horizontal transfer)]TJ T* 0.0975 Tw (seems considerably more frequent among Class II than)Tj T* -0.0531 Tw (Class I families. Evidence summarized here indicates that)Tj T* -0.0589 Tw [(this pattern is due to fundamental dif)13.7(ferences between the)]TJ T* -0.0013 Tc -0.0695 Tw [(two )19.2(TE Classes. )12.9(The major transposition mechanism used)]TJ T* 0 Tc -0.0352 Tw (by Class II elements is better suited for horizontal transfer)Tj T* 0.1088 Tw [(than those used by Class I )15(TE families. In addition, the)]TJ T* -0.0067 Tc -0.0695 Tw [(type of copy number regulation used by Class II )17.7(TEs, which)]TJ T* 0 Tc -0.0256 Tw (seem to be mostly self-regulated, may make it impossible)Tj T* -0.0431 Tw (for these elements to sustain a rate of transposition that is)Tj T* 0.0069 Tc 0.1389 Tw (compatible with their long-term survival within a single)Tj T* -0.0113 Tc -0.0695 Tw [(genome. )15.4(Thus, horizontal transfer may be the chance event)]TJ T* 0 Tc 0.008 Tw (that dictates which Class II families survive and which do)Tj T* -0.0004 Tc 0 Tw (not.)Tj 1.8933 -1.2 TD 0 Tc -0.0676 Tw (Extensive genomic data, often in the form of complete)Tj -1.8933 -1.2 TD 0.1288 Tw (genomes, is currently being gathered at an astonishing)Tj T* 0.0145 Tc 0.1389 Tw [(rate. Mining this type of data for )12.8(TEs is sure to bring)]TJ T* 0.0084 Tc (advances on many fronts. For example, it will make it)Tj T* -0.0124 Tc -0.0695 Tw [(possible to objectively assess the distribution of known )11.6(TEs,)]TJ T* -0.0116 Tc -0.0694 Tw (to estimate more reliably the incidence of horizontal transfer)Tj T* 0 Tc 0.0592 Tw [(among them, and to identify new )18.8(TE families. Studies of)]TJ T* -0.0129 Tc -0.0695 Tw (horizontal transfer have practical implications as well. Some)Tj T* 0.0184 Tc 0.139 Tw (TEs, such as )Tj /F3 1 Tf 9 0 1.913 9 117.01 312.675 Tm 0 Tw (piggyBac)Tj /F1 1 Tf 9 0 0 9 155.35 312.675 Tm 0 Tc ( )Tj 0.4354 0 TD 0.0184 Tc 0.139 Tw (and those of the )Tj /F3 1 Tf 9 0 1.913 9 234.13 312.675 Tm 0 Tw [(mariner)18.1(-T)87.5(c1)]TJ /F1 1 Tf 9 0 0 9 56.53 301.875 Tm 0.0113 Tc 0.1389 Tw [(superfamily)70(, are already being used as transformation)]TJ 0 -1.2 TD 0 Tc 0.1071 Tw (vectors in a wide variety of taxa \(Sherman et al., 1998;)Tj T* -0.0076 Tc -0.0694 Tw (Rubin et al., 1999; Mamoun et al., 2000; Zhang et al., 2000;)Tj T* 0 Tc 0.0859 Tw [(Handler)54.4(, 2002\). )14.1(The usefulness of )15.4(TEs in this context is)]TJ T* -0.0198 Tw (phenomenal since, through insertional mutagenesis, they)Tj T* -0.021 Tw (provide a tool to rapidly identify and recover of genes that)Tj T* 0.0415 Tw (contribute of specific phenotypes \(Spradling et al., 1999;)Tj T* 0.1292 Tw (Fischer et al., 2001; Horn et al., 2003\). In addition, the)Tj T* 0.0037 Tc 0.139 Tw [(inherent ability of )15.3(TEs to spread can be explored as a)]TJ T* 0 Tc -0.0029 Tw [(means to transform, and thus, ultimately)70.4(, control, species)]TJ T* 0.0881 Tw (that are involved in the transmission of diseases or that)Tj T* -0.0097 Tc -0.0695 Tw (are, themselves, considered to be pests \(Gueiros-Filho and)Tj T* 0 Tc 0.0004 Tw [(Beverley)70(, 1997; )56.9(Atkinson et al., 2001\).)]TJ 1.8933 -1.2 TD 0.0149 Tc 0.1389 Tw (The success of these elements as transformation)Tj -1.8933 -1.2 TD 0.0092 Tc (vectors rests mainly on two bases: on studies of their)Tj T* -0.0061 Tc -0.0694 Tw (transposition mechanisms that show transposase to be the)Tj T* 0 Tc 0.0841 Tw [(only factor required for transposition. )14.3(This suggests that)]TJ T* -0.0047 Tw (the success of such experiments does not hinge on host-)Tj T* -0.0036 Tc -0.0695 Tw (encoded factors, which might be absent in distantly related)Tj T* 0 Tc 0.0551 Tw [(taxa. )14.5(They also rely on studies of the occurrence of )17.1(TEs)]TJ 27.7267 72.4 TD -0.0062 Tc -0.0695 Tw (that reveal the presence of specific elements across a wide)Tj 0 -1.2 TD 0 Tc -0.0691 Tw (variety of taxa, since this provides circumstantial evidence)Tj T* -0.0033 Tc -0.0695 Tw (for the success of such elements as tools in transformation)Tj T* 0.0359 Tc 0.1389 Tw [(studies. )52.7(Additional studies of horizontal transfer)57.9(, by)]TJ T* 0 Tc 0.0259 Tw (providing further insights into the features of interspecific)Tj T* 0.0573 Tw [(transfer)56.7(, may prove invaluable for the fields of functional)]TJ T* 0.0001 Tw (and medical genomics.)Tj /F2 1 Tf 0 -2.4 TD -0.0001 Tc 0 Tw (Acknowledgements)Tj /F1 1 Tf T* -0.0307 Tc (We)Tj 1.4599 0 TD ( )Tj 0.1991 0 TD -0.0094 Tc -0.0695 Tw (would like to thank Margaret G. Kidwell, David G. Harris)Tj -1.659 -1.2 TD 0 Tc -0.0352 Tw (and Shelby L. Bidwell for critical review of the manuscript,)Tj T* 0.0594 Tc 0.1389 Tw (and for suggestions and lengthy discussions that)Tj T* 0 Tc 0.1293 Tw (significantly contributed to the clarity of the manuscript.)Tj T* 0.0293 Tc 0 Tw (We)Tj 1.5748 0 TD ( )Tj 0.4624 0 TD 0.0455 Tc 0.1389 Tw [(also thank Margaret G. Kidwell and )61.4(Andrew J.)]TJ -2.0371 -1.2 TD 0 Tc -0.0404 Tw [(Holyoake, and Jake )18.8(T)37.7(u)0( and Hongguang Shao for allowing)]TJ T* 0.0004 Tw (us to use their unpublished results.)Tj /F2 1 Tf 0 -2.4 TD 0.0003 Tc 0 Tw (References)Tj /F1 1 Tf T* 0 Tc 0.1237 Tw [(Andrews, J.D., and Gloor)55.4(, G.B. 1995. )59.4(A)53.7( role for the KP)]TJ 0.6267 -1.2 TD 0.1266 Tw (leucine zipper in regulating )Tj /F3 1 Tf 9 0 1.913 9 425.83 521.475 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 431.83 521.475 Tm ( )Tj 0.4046 0 TD 0.1266 Tw (element transposition in)Tj /F3 1 Tf 9 0 1.913 9 311.71 510.675 Tm -0.001 Tw [(Drosophila melanogaster)34.1(.)]TJ /F1 1 Tf 9 0 0 9 413.89 510.675 Tm -0.0001 Tc -0.0009 Tw ( Genetics)Tj /F3 1 Tf 9 0 1.913 9 451.93 510.675 Tm 0 Tc 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 454.39 510.675 Tm -0.001 Tw (141: 587-594.)Tj -16.48 -1.2 TD 0.0576 Tw (Anxolabhre, D., Kidwell, M.G., and Periquet, G. 1988.)Tj 0.6267 -1.2 TD 0.0223 Tc 0.139 Tw (Molecular characteristics of diverse populations are)Tj T* 0.0021 Tc 0.1389 Tw (consistent with the hypothesis of a recent invasion of)Tj /F3 1 Tf 9 0 1.913 9 311.71 467.475 Tm -0.0104 Tc -0.0694 Tw (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 409.03 467.475 Tm 0 Tc 0 Tw ( )Tj 0.1982 0 TD -0.0104 Tc -0.0694 Tw (by mobile )Tj /F3 1 Tf 9 0 1.913 9 449.59 467.475 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 455.53 467.475 Tm ( )Tj 0.1982 0 TD -0.0104 Tc -0.0694 Tw (elements. Mol. Biol.)Tj -16.1782 -1.2 TD 0 Tc 0 Tw (Evol.)Tj /F3 1 Tf 9 0 1.913 9 331.69 456.675 Tm ( )Tj /F1 1 Tf 9 0 0 9 334.21 456.675 Tm 0.0003 Tw (5: 252-269.)Tj -3.1267 -1.2 TD 0.0426 Tc 0.1389 Tw (Arca, B., and Savakis, C. 2000. Distribution of the)Tj 0.6267 -1.2 TD 0.0033 Tc (transposable element )Tj /F3 1 Tf 9 0 1.913 9 403.33 435.075 Tm 0 Tw (Minos)Tj /F1 1 Tf 9 0 0 9 427.57 435.075 Tm 0 Tc ( )Tj 0.4202 0 TD 0.0033 Tc 0.1389 Tw (in the genus )Tj /F3 1 Tf 9 0 1.913 9 487.03 435.075 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 530.29 435.075 Tm 0 Tc (.)Tj -24.2867 -1.2 TD -0.0011 Tw (Genetica 108: 263-267.)Tj -0.6267 -1.2 TD -0.0026 Tc -0.0694 Tw [(Atkinson, P)126.7(.)0(W)56.8(., Pinkerton, )58.9(A.C., and OBrochta, D.A. 2001.)]TJ 0.6267 -1.2 TD 0 Tc 0.1143 Tw [(Genetic transformation systems in insects. )53.7(Annu. Rev)76.7(.)]TJ T* -0.0005 Tw (Entomol. 46: 317-346.)Tj -0.6267 -1.2 TD 0.0065 Tc 0.139 Tw [(Auge-Gouillou, C., Bigot, )23.2(Y)126.8(., Pollet, N., Hamelin, M.H.,)]TJ 0.6267 -1.2 TD 0 Tc -0.024 Tw (Meunier-Rotival, M., and Periquet, G. 1995. Human and)Tj T* 0.0737 Tw (other mammalian genomes contain transposons of the)Tj /F3 1 Tf 9 0 1.913 9 311.71 348.675 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 342.19 348.675 Tm -0.0002 Tw [( family)68.8(. FEBS Lett.)]TJ /F3 1 Tf 9 0 1.913 9 416.05 348.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 418.57 348.675 Tm -0.0002 Tw (368: 541-546.)Tj -12.5 -1.2 TD 0.0653 Tw [(Badge, R.M., and Brookfield, J.F)112.4(. 1997. )20(The role of host)]TJ 0.6267 -1.2 TD -0.0052 Tc -0.0694 Tw (factors in the population dynamics of selfish transposable)Tj T* 0 Tc 0.0004 Tw [(elements. J. )23.2(Theor)45.4(. Biol. 187: 261-271.)]TJ -0.6267 -1.2 TD 0.0322 Tw [(Beall, E.L., )58.5(Admon, )56.2(A., and Rio, D.C. 1994. )48.9(A)53.7( )]TJ /F3 1 Tf 9 0 1.913 9 489.79 305.475 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 311.71 294.675 Tm 0.0628 Tw (protein homologous to the human p70 Ku autoimmune)Tj T* -0.011 Tc -0.0695 Tw (antigen interacts with the )Tj /F3 1 Tf 9 0 1.913 9 408.61 283.875 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 414.49 283.875 Tm ( )Tj 0.1975 0 TD -0.011 Tc -0.0695 Tw (transposable element inverted)Tj -11.6175 -1.2 TD 0 Tc 0.0004 Tw [(repeats. Proc. Natl. )58.3(Acad. Sci. USA. 91: 12681-12685.)]TJ -0.6267 -1.2 TD 0.0083 Tc 0.1389 Tw (Beall, E.L., and Rio, D.C. 1997. )Tj /F3 1 Tf 9 0 1.913 9 443.59 262.275 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 487.33 262.275 Tm 0 Tc ( )Tj /F3 1 Tf 9 0 1.913 9 491.23 262.275 Tm (P)Tj /F1 1 Tf 9 0 0 9 497.23 262.275 Tm 0.0083 Tc (-element)Tj -20.6133 -1.2 TD -0.0086 Tc -0.0695 Tw (transposase is a novel site-specific endonuclease. Genes)Tj T* 0 Tc 0 Tw [(Dev)71.3(.)]TJ /F3 1 Tf 9 0 1.913 9 329.53 240.675 Tm ( )Tj /F1 1 Tf 9 0 0 9 332.05 240.675 Tm (1)Tj 0.4867 0 TD 0.0013 Tw (1: 2137-2151.)Tj -3.3733 -1.2 TD 0.0106 Tc 0.1389 Tw (Berg, D.E., and Howe, M.M., eds. 1989. Mobile DNA.)Tj 0.6267 -1.2 TD 0 Tc 0.0003 Tw [(American Society of Microbiology)69.8(, W)40.4(ashington, D.C.)]TJ -0.6267 -1.2 TD 0.1149 Tw [(Bimont, C., V)16.8(ieira, C., Borie, N., and Lepetit, D. 1999.)]TJ 0.6267 -1.2 TD 0 Tw (T)Tj 0.5733 0 TD -0.0409 Tw (ransposable elements and genome evolution: the case)Tj -0.5733 -1.2 TD 0 Tw (of )Tj /F3 1 Tf 9 0 1.913 9 321.73 186.675 Tm 0.0001 Tw (Drosophila simulans)Tj /F1 1 Tf 9 0 0 9 402.73 186.675 Tm (. Genetica)Tj /F3 1 Tf 9 0 1.913 9 443.77 186.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 446.23 186.675 Tm 0.0001 Tw [(107: 1)66.8(13-120.)]TJ -15.5733 -1.2 TD 0.0413 Tc 0.1389 Tw [(Bimont, C., V)23(ieira, C., Hoogland, C., Cizeron, G.,)]TJ 0.6267 -1.2 TD 0.0019 Tc [(Loevenbruck, C., )53.7(Arnault, C., and Carante, J.P)134.7(. 1997.)]TJ T* 0 Tc 0.1291 Tw (Maintenance of transposable element copy number in)Tj T* 0.0642 Tw (natural populations of )Tj /F3 1 Tf 9 0 1.913 9 402.01 143.475 Tm (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 502.63 143.475 Tm 0 Tw ( )Tj 0.3422 0 TD (and )Tj /F3 1 Tf 9 0 1.913 9 523.81 143.475 Tm (D.)Tj -23.3116 -1.2 TD (simulans)Tj /F1 1 Tf 9 0 0 9 347.23 132.675 Tm -0.0012 Tw (. Genetica)Tj /F3 1 Tf 9 0 1.913 9 388.21 132.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 390.73 132.675 Tm -0.0012 Tw (100: 161-166.)Tj -9.4067 -1.2 TD -0.0055 Tc -0.0695 Tw [(Black, D.M., Jackson, M.S., Kidwell, M.G., and Dover)57.6(, G.A.)]TJ 0.6267 -1.2 TD -0.0027 Tc 0 Tw (1987. )Tj /F3 1 Tf 9 0 1.913 9 335.95 111.075 Tm (KP)Tj /F1 1 Tf 9 0 0 9 347.89 111.075 Tm 0 Tc ( )Tj 0.2058 0 TD -0.0027 Tc -0.0695 Tw (elements repress )Tj /F3 1 Tf 9 0 1.913 9 419.59 111.075 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 425.59 111.075 Tm -0.0027 Tc -0.0695 Tw (-induced hybrid dysgenesis)Tj -12.6533 -1.2 TD 0 Tc 0 Tw (in )Tj /F3 1 Tf 9 0 1.913 9 321.13 100.275 Tm -0.0095 Tw (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 421.09 100.275 Tm (. EMBO J. 6: 4125-4135.)Tj ET endstream endobj 34 0 obj << /Type /Page /Parent 95 0 R /Resources 35 0 R /Contents 36 0 R /Thumb 76 0 R /MediaBox [ 0 0 595 842 ] /CropBox [ 0 0 595 842 ] /Rotate 0 >> endobj 35 0 obj << /ProcSet [ /PDF /Text ] /Font << /F1 114 0 R /F2 113 0 R /F3 128 0 R >> /ExtGState << /GS1 139 0 R /GS2 138 0 R >> /ColorSpace << /Cs8 112 0 R >> >> endobj 36 0 obj << /Length 17018 >> stream BT /F1 1 Tf 9 0 0 9 337.47 780.5551 Tm 0 0 0 1 k /GS2 gs 0 Tc -0.0155 Tw [(Horizontal Transfer of Transposable Elements 11)]TJ /F2 1 Tf 37.9351 29.4096 -29.4096 37.9351 42.9423 542.4378 Tm 2 Tr 0 0 0 0.45 K 0 J 0 j 0.576 w 10 M []0 d /Cs8 cs 0 scn /GS1 gs 0.0686 Tc -0.0001 Tw (PAGE PROOFS)Tj /F1 1 Tf 9 0 0 9 62.19 748.275 Tm 0 Tr 0 0 0 1 k /GS2 gs -0.0125 Tc -0.0694 Tw [(Brookfield, J.F)114.3(., and Badge, R.M. 1997. Population genetics)]TJ 0.6267 -1.2 TD 0 Tc 0.1384 Tw (models of transposable elements. Genetica)Tj /F3 1 Tf 9 0 1.913 9 245.97 737.475 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 249.63 737.475 Tm 0.1384 Tw (100: 281-)Tj -20.2 -1.2 TD -0.0006 Tc 0 Tw (294.)Tj -0.6267 -1.2 TD 0.031 Tc 0.1389 Tw [(Brunet, F)110.7(., Godin, F)114.3(., Bazin, C., and Capy)70.9(, P)128.2(.)0( 1999.)]TJ 0.6267 -1.2 TD -0.0129 Tc -0.0695 Tw (Phylogenetic analysis of )Tj /F3 1 Tf 9 0 1.913 9 162.09 705.0751 Tm 0 Tw (Mos1)Tj /F1 1 Tf 9 0 0 9 183.63 705.0751 Tm -0.0695 Tw (-like transposable elements)Tj -12.8667 -1.2 TD 0 Tc -0.0001 Tw (in the )Tj /F3 1 Tf 9 0 1.913 9 92.37 694.275 Tm 0 Tw (Drosophilidae)Tj /F1 1 Tf 9 0 0 9 147.39 694.275 Tm -0.0001 Tw (. J. Mol. Evol. 49: 760-768.)Tj -9.4667 -1.2 TD 0.0014 Tw [(Brunet, F)107.7(., Godin, F)118.5(., David, J.R., and Capy)72.3(, P)124.4(.)0( 1994. )18.8(The)]TJ /F3 1 Tf 9 0 1.913 9 67.83 672.675 Tm -0.0081 Tc 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 97.83 672.675 Tm 0 Tc ( )Tj 0.2004 0 TD -0.0081 Tc -0.0695 Tw (transposable element in the )Tj /F3 1 Tf 9 0 1.913 9 208.17 672.675 Tm 0 Tw (Drosophilidae)Tj /F1 1 Tf 9 0 0 9 262.23 672.675 Tm 0 Tc ( )Tj 0.2004 0 TD -0.0081 Tc [(family)69.3(.)]TJ -21.8004 -1.2 TD 0 Tc (Heredity)Tj /F3 1 Tf 9 0 1.913 9 101.37 661.875 Tm ( )Tj /F1 1 Tf 9 0 0 9 103.83 661.875 Tm -0.0003 Tw (73: 377-385.)Tj -4.6267 -1.2 TD 0.0764 Tw [(Calvi, B.R., Hong, )14.3(T)111(.J., Findley)78.1(, S.D., and Gelbart, W)54.4(.M.)]TJ 0.6267 -1.2 TD 0.0164 Tc 0.1389 Tw (1991. Evidence for a common evolutionary origin of)Tj T* 0.001 Tc (inverted repeat transposons in )Tj /F3 1 Tf 9 0 1.913 9 196.71 629.475 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 239.79 629.475 Tm 0 Tc ( )Tj 0.4179 0 TD 0.001 Tc 0.1389 Tw (and plants:)Tj /F3 1 Tf 9 0 1.913 9 67.83 618.675 Tm 0 Tc 0 Tw (hobo)Tj /F1 1 Tf 9 0 0 9 87.87 618.675 Tm (, )Tj /F3 1 Tf 9 0 1.913 9 92.73 618.675 Tm (Activator)Tj /F1 1 Tf 9 0 0 9 127.77 618.675 Tm -0.0153 Tw (, and )Tj /F3 1 Tf 9 0 1.913 9 149.97 618.675 Tm 0 Tw (T)Tj 0.52 0 TD (am3)Tj /F1 1 Tf 9 0 0 9 172.17 618.675 Tm -0.0153 Tw (. Cell)Tj /F3 1 Tf 9 0 1.913 9 192.57 618.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 194.91 618.675 Tm -0.0153 Tw (66: 465-471.)Tj -14.7467 -1.2 TD -0.0003 Tc -0.0695 Tw [(Capy)72.7(, P)125.9(.)0( 1998. Classification of transposable elements. In:)]TJ 0.6267 -1.2 TD 0 Tc 0.0583 Tw [(Dynamics )52(And Evolution Of )16.9(T)37.7(ransposable Elements. P)126(.)]TJ T* 0.0778 Tw [(Capy)74(, C. Bazin, D. Higuet, and )14.4(T)111(.)0( Langin, eds. Landes)]TJ T* -0.0007 Tw [(Bioscience, )57(Austin. p. 37-52.)]TJ -0.6267 -1.2 TD 0.0125 Tc 0.139 Tw [(Capy)70.8(, P)126.2(., )63.9(Anxolabhre, D., and Langin, )11.2(T)110.2(.)0( 1994. )20.9(The)]TJ 0.6267 -1.2 TD 0.0306 Tc 0.1389 Tw (strange phylogenies of transposable elements: are)Tj T* 0 Tc -0.047 Tw [(horizontal transfers the only explanation? )14.2(T)37.7(rends Genet.)]TJ T* 0.0027 Tw (10: 7-12.)Tj -0.6267 -1.2 TD 0.0366 Tw [(Capy)74(, P)132.9(., V)10.9(italis, R., Langin, )19.9(T)111(., Higuet, D., and Bazin, C.)]TJ 0.6267 -1.2 TD 0.0173 Tc 0.1389 Tw (1996. Relationships between transposable elements)Tj T* 0 Tc -0.0655 Tw (based upon the integrase-transposase domains: is there)Tj T* 0.0001 Tw (a common ancestor? J. Mol. Evol. 42: 359-368.)Tj -0.6267 -1.2 TD 0.048 Tc 0.1389 Tw [(Charlesworth, B., and Charlesworth, D. 1983. )11.8(The)]TJ 0.6267 -1.2 TD 0 Tc 0.0882 Tw (population dynamics of transposable elements. Genet.)Tj T* 0.001 Tw (Res. 42: 1-27.)Tj -0.6267 -1.2 TD 0.0137 Tc 0.1389 Tw [(Charlesworth, B., and Langley)69.1(, C.H. 1991. Population)]TJ 0.6267 -1.2 TD 0.0152 Tc (genetics of transposable elements in )Tj /F3 1 Tf 9 0 1.913 9 227.73 435.075 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 272.13 435.075 Tm 0.1389 Tw (. In:)Tj -22.7 -1.2 TD 0 Tc 0.1133 Tw [(Evolution )51(At )23(The Molecular Level. R.K. Selander)50.8(, )56(A.G.)]TJ T* 0.0582 Tw [(Clark, and )18.1(T)111(.S. Whittam, eds. Sinauer )55.6(Associates, Inc.,)]TJ T* 0.0001 Tc 0 Tw (Sunderland.)Tj -0.6267 -1.2 TD -0.0105 Tc -0.0694 Tw [(Charlesworth, B., and Lapid, )54.1(A. 1989. )50(A)56.5( study of ten families)]TJ 0.6267 -1.2 TD 0.0089 Tc 0.1389 Tw (of transposable elements on X chromosomes from a)Tj T* 0 Tc -0.0192 Tw (population of )Tj /F3 1 Tf 9 0 1.913 9 121.53 370.275 Tm (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 221.43 370.275 Tm (. Genet. Res. 54:)Tj -17.0667 -1.2 TD 0 Tw (1)Tj 0.4867 0 TD -0.0002 Tc (13-125.)Tj -1.1133 -1.2 TD 0 Tc 0.132 Tw [(Clark, J.B., )52.1(Altheide, )21(T)111(.K., Schlosser)54.4(, M.J., and Kidwell,)]TJ 0.6267 -1.2 TD 0.039 Tc 0.1389 Tw (M.G. 1995. Molecular evolution of )Tj /F3 1 Tf 9 0 1.913 9 223.11 337.875 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 229.47 337.875 Tm ( )Tj 0.4559 0 TD 0.039 Tc (transposable)Tj -18.4159 -1.2 TD 0.0025 Tc 0.1389 Tw [(elements in the genus Drosophila. I. )15.1(The )]TJ /F3 1 Tf 9 0 1.913 9 241.47 327.075 Tm 0 Tw (saltans)Tj /F1 1 Tf 9 0 0 9 270.09 327.075 Tm 0 Tc ( )Tj 0.4194 0 TD 0.0025 Tc (and)Tj /F3 1 Tf 9 0 1.913 9 67.83 316.275 Tm 0 Tc (willistoni)Tj /F1 1 Tf 9 0 0 9 101.37 316.275 Tm -0.0001 Tw ( species groups. Mol. Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 228.87 316.275 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 231.39 316.275 Tm -0.0001 Tw (12: 902-913.)Tj -18.8 -1.2 TD 0.0342 Tc 0.1389 Tw [(Clark, J.B., and Kidwell, M.G. 1997. )54.2(A)54.5( phylogenetic)]TJ 0.6267 -1.2 TD 0.0226 Tc (perspective on )Tj /F3 1 Tf 9 0 1.913 9 134.43 294.675 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 140.67 294.675 Tm ( )Tj 0.4395 0 TD 0.0226 Tc 0.1389 Tw (transposable element evolution in)Tj -8.5329 -1.2 TD -0.0047 Tc -0.0694 Tw [(Drosophila. Proc. Natl. )53.6(Acad. Sci. USA. 94: 1)73.1(1428-1)71.5(1433.)]TJ -0.6267 -1.2 TD 0 Tc 0.0185 Tw [(Clark, J.B., Kim, P)128.6(.C., and Kidwell, M.G. 1998. Molecular)]TJ 0.6267 -1.2 TD 0.0306 Tc 0.1389 Tw (evolution of )Tj /F3 1 Tf 9 0 1.913 9 122.43 262.275 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 128.73 262.275 Tm ( )Tj 0.4475 0 TD 0.0306 Tc 0.1389 Tw (transposable elements in the genus)Tj -7.2142 -1.2 TD 0.0003 Tc [(Drosophila. III. )15.1(The )]TJ /F3 1 Tf 9 0 1.913 9 150.03 251.475 Tm 0 Tw (melanogaster)Tj /F1 1 Tf 9 0 0 9 204.57 251.475 Tm 0 Tc ( )Tj 0.4172 0 TD 0.0003 Tc 0.1389 Tw (species group. Mol.)Tj -15.6106 -1.2 TD 0 Tc -0.0003 Tw (Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 107.85 240.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 110.37 240.675 Tm -0.0003 Tw (15: 746-755.)Tj -5.3533 -1.2 TD 0.0198 Tc 0.1389 Tw [(Clark, J.B., Maddison, W)54.2(.)0(P)131.2(., and Kidwell, M.G. 1994.)]TJ 0.6267 -1.2 TD 0 Tc 0.0831 Tw (Phylogenetic analysis supports horizontal transfer of )Tj /F3 1 Tf 9 0 1.913 9 282.93 219.075 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 67.83 208.275 Tm -0.0007 Tw (transposable elements. Mol. Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 222.87 208.275 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 225.39 208.275 Tm (1)Tj 0.4867 0 TD -0.0007 Tw (1: 40-50.)Tj -18.62 -1.2 TD -0.0047 Tw (Clark, J.B., Silva, J.C., and Kidwell, M.G. 2002. Evidence)Tj 0.6267 -1.2 TD 0.0078 Tc 0.1389 Tw (of horizontal transfer of )Tj /F3 1 Tf 9 0 1.913 9 169.23 186.675 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 175.29 186.675 Tm ( )Tj 0.4247 0 TD 0.0078 Tc 0.1389 Tw (transposable elements. In:)Tj -12.3647 -1.2 TD 0 Tc 0.0949 Tw [(Horizontal Gene )12.9(T)37.7(ransfer)58.7(. M. Syvanen, and C.I. Kado,)]TJ T* -0.0001 Tw [(eds. )54.6(Academic Press, San Diego. p. 161-171.)]TJ -0.6267 -1.2 TD -0.008 Tc -0.0694 Tw [(Cummings, M.P)125.8(. 1994. )20.1(T)36.3(ransmission patterns of eukaryotic)]TJ 0.6267 -1.2 TD 0.0348 Tc 0.1389 Tw (transposable elements: arguments for and against)Tj T* 0 Tc -0.0003 Tw [(horizontal transfer)49.5(. )15.7(T)44.3(rends Ecol. Evol. 9: 141-145.)]TJ -0.6267 -1.2 TD 0.0113 Tw [(Dai, L., and Zimmerly)75.9(, S. 2002. Compilation and analysis)]TJ 0.6267 -1.2 TD -0.0132 Tc -0.0695 Tw (of group II intron insertions in bacterial genomes: evidence)Tj T* 0 Tc 0.0156 Tw [(for retroelement behavior)49(. Nucleic )56.6(Acids Res. 30: 1091-)]TJ T* 0 Tw (1)Tj 0.4867 0 TD -0.0002 Tc (102.)Tj 26.6067 73.2 TD 0.0241 Tc 0.1389 Tw [(Daniels, S.B., Chovnick, )56.1(A., and Boussy)74.3(, I.A. 1990a.)]TJ 0.6333 -1.2 TD -0.0128 Tc -0.0694 Tw (Distribution of the )Tj /F3 1 Tf 9 0 1.913 9 385.71 737.475 Tm 0 Tw (hobo)Tj /F1 1 Tf 9 0 0 9 405.27 737.475 Tm 0 Tc ( )Tj 0.1958 0 TD -0.0128 Tc -0.0694 Tw (transposable element in the genus)Tj /F3 1 Tf 9 0 1.913 9 317.37 726.675 Tm 0 Tc 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 360.33 726.675 Tm -0.0007 Tw (. Mol. Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 424.89 726.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 427.35 726.675 Tm -0.0007 Tw (7: 589-606.)Tj -12.8533 -2.4 TD -0.0437 Tw (Daniels, S.B., Peterson, K.R., Strausbaugh, L.D., Kidwell,)Tj 0.6333 -1.2 TD 0.079 Tw [(M.G., and Chovnick, )60.8(A. 1990b. Evidence for horizontal)]TJ T* 0.0102 Tc 0.1389 Tw (transmission of the)Tj /F3 1 Tf 9 0 1.913 9 397.11 683.475 Tm 0.1491 Tc 0 Tw ( P)Tj /F1 1 Tf 9 0 0 9 407.07 683.475 Tm 0 Tc ( )Tj 0.4271 0 TD 0.0102 Tc 0.1389 Tw (transposable element between)Tj /F3 1 Tf 9 0 1.913 9 317.37 672.675 Tm 0 Tc 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 360.33 672.675 Tm -0.0001 Tw ( species. Genetics)Tj /F3 1 Tf 9 0 1.913 9 433.89 672.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 436.41 672.675 Tm -0.0001 Tw (124: 339-355.)Tj -13.86 -1.2 TD -0.0192 Tw [(Doolittle, W)52.8(.)0(F)109(., and Sapienza, C. 1980. Selfish genes, the)]TJ 0.6333 -1.2 TD -0.0007 Tc -0.0694 Tw (phenotype paradigm and genome evolution. Nature)Tj /F3 1 Tf 9 0 1.913 9 519.09 651.075 Tm 0 Tc 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 520.95 651.075 Tm -0.0007 Tc (284:)Tj -22.62 -1.2 TD 0.0003 Tc (601-603.)Tj -0.6333 -1.2 TD 0.0517 Tc 0.1389 Tw [(Eickbush, D.G., and Eickbush, )15.9(T)109.4(.H. 1995. V)55.7(ertical)]TJ 0.6333 -1.2 TD 0 Tc 0.046 Tw (transmission of the retrotransposable elements )Tj /F3 1 Tf 9 0 1.913 9 509.01 618.675 Tm 0 Tw (R1)Tj /F1 1 Tf 9 0 0 9 520.47 618.675 Tm ( )Tj 0.324 0 TD (and)Tj /F3 1 Tf 9 0 1.913 9 317.37 607.875 Tm (R2)Tj /F1 1 Tf 9 0 0 9 328.83 607.875 Tm ( )Tj 0.2933 0 TD 0.0153 Tw (during the evolution of the )Tj /F3 1 Tf 9 0 1.913 9 438.27 607.875 Tm (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 317.37 597.075 Tm -0.0003 Tw (species subgroup. Genetics)Tj /F3 1 Tf 9 0 1.913 9 428.37 597.075 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 430.89 597.075 Tm -0.0003 Tw (139: 671-684.)Tj -13.2467 -1.2 TD 0.0476 Tc 0.1389 Tw [(Engels, W)51.2(.)0( 1979. Hybrid dysgenesis in )]TJ /F3 1 Tf 9 0 1.913 9 491.55 586.275 Tm 0 Tw (Drosophila)Tj -19.0983 -1.2 TD 0.0127 Tc (melanogaster)Tj /F1 1 Tf 9 0 0 9 373.23 575.475 Tm 0.1389 Tw [(: rules of inheritance of female sterility)70.5(.)]TJ -6.2067 -1.2 TD 0 Tc -0.0006 Tw (Genet. Res. 33: 219-236.)Tj -0.6333 -1.2 TD -0.0536 Tw [(Engels, W)50.1(.)0( 1989. )]TJ /F3 1 Tf 9 0 1.913 9 380.79 553.875 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 386.79 553.875 Tm ( )Tj 0.2244 0 TD -0.0536 Tw (elements in )Tj /F3 1 Tf 9 0 1.913 9 436.41 553.875 Tm (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 535.9501 553.875 Tm 0 Tw (.)Tj -24.2867 -1.2 TD 0.0408 Tc 0.1389 Tw (In: Mobile DNA. D.E. Berg, and M.M. Howe, eds.)Tj T* 0 Tc 0.065 Tw [(American Society of Microbiology)77.1(, W)31.7(ashington, D.C. p.)]TJ T* 0.0003 Tc 0 Tw (437-483.)Tj -0.6333 -1.2 TD 0 Tc -0.0643 Tw [(Engels, W)52.7(.R. 1992. )14.1(The origin of )]TJ /F3 1 Tf 9 0 1.913 9 440.13 510.675 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 446.13 510.675 Tm ( )Tj 0.2137 0 TD -0.0643 Tw (elements in )Tj /F3 1 Tf 9 0 1.913 9 495.45 510.675 Tm 0 Tw (Drosophila)Tj -19.5316 -1.2 TD (melanogaster)Tj /F1 1 Tf 9 0 0 9 371.91 499.875 Tm -0.0164 Tw (. Bioessays)Tj /F3 1 Tf 9 0 1.913 9 417.75 499.875 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 420.09 499.875 Tm -0.0164 Tw (14: 681-686.)Tj -12.0467 -1.2 TD 0.0331 Tc 0.1389 Tw [(Finnegan, D. 1992. )14.6(T)37.4(ransposable elements. In: )17.7(The)]TJ 0.6333 -1.2 TD 0 Tc -0.0437 Tw (Genome of )Tj /F3 1 Tf 9 0 1.913 9 363.57 478.275 Tm (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 463.23 478.275 Tm [(. D.L. Lindsley)68.7(, and)]TJ -16.2067 -1.2 TD -0.0007 Tc -0.0695 Tw [(G. Zimm, eds. )58.6(Academic Press, New )15.3(Y)93(ork. p. 1092-1)74.4(107.)]TJ -0.6333 -1.2 TD 0 Tc 0.125 Tw (Finnegan, D.J. 1989. Eukaryotic transposable elements)Tj 0.6333 -1.2 TD 0.0002 Tw [(and genome evolution. )22.3(T)31(rends Genet. 5: 103-107.)]TJ -0.6333 -1.2 TD 0.0071 Tc 0.1389 Tw [(Fischer)54.6(, S.E., W)-5.3(ienholds, E., and Plasterk, R.H. 2001.)]TJ 0.6333 -1.2 TD -0.007 Tc -0.0695 Tw (Regulated transposition of a fish transposon in the mouse)Tj T* 0 Tc -0.0001 Tw [(germ line. Proc. Natl. )53.6(Acad. Sci. USA. 98: 6759-6764.)]TJ -0.6333 -1.2 TD -0.0103 Tc -0.0695 Tw [(Fraser)53.7(, M.J., Brusca, J.S., Smith, G.E., and Summers, M.D.)]TJ 0.6333 -1.2 TD 0.0695 Tc 0.343 Tw [(1985. )19.8(T)33.8(ransposon-mediated mutagenesis of a)]TJ T* 0 Tc 0.0009 Tw [(baculovirus. V)20.9(irology)]TJ /F3 1 Tf 9 0 1.913 9 400.23 381.075 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 402.75 381.075 Tm 0.0009 Tw (145: 356-361.)Tj -10.12 -1.2 TD -0.0215 Tw [(Frissen, N., Brandes, )52.3(A., and Helsop-Harrison, J.S. 2001.)]TJ 0.6333 -1.2 TD 0.0069 Tc 0.1389 Tw [(Diversity)75.2(, origin, and distribution of retrotranspososns)]TJ T* 0 Tc 0 Tw (\()Tj /F3 1 Tf 9 0 1.913 9 320.37 348.675 Tm (gypsy)Tj /F1 1 Tf 9 0 0 9 343.83 348.675 Tm ( )Tj 0.3046 0 TD (and )Tj /F3 1 Tf 9 0 1.913 9 364.35 348.675 Tm (copia)Tj /F1 1 Tf 9 0 0 9 385.83 348.675 Tm 0.0266 Tw (\) in conifers. Mol. Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 498.09 348.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 500.85 348.675 Tm 0.0266 Tw [(18: 1)70.7(176-)]TJ -20.3867 -1.2 TD 0 Tw (1)Tj 0.48 0 TD -0.0002 Tc (188.)Tj -1.1133 -1.2 TD 0 Tc 0.0374 Tw [(Garcia-Fernandez, J., Bayascas-Ramirez, J.R., Marfany)70(,)]TJ 0.6333 -1.2 TD -0.0109 Tc -0.0695 Tw [(G., Munoz-Marmol, )59.6(A.M., Casali, )54.2(A., Baguna, J., and Salo,)]TJ T* 0 Tc -0.0254 Tw (E. 1995. High copy number of highly similar )Tj /F3 1 Tf 9 0 1.913 9 491.61 305.475 Tm 0 Tw [(mariner)15.7(-)]TJ /F1 1 Tf 9 0 0 9 524.97 305.475 Tm (like)Tj -23.0667 -1.2 TD 0.0329 Tw (transposons in planarian \()Tj /F3 1 Tf 9 0 1.913 9 421.77 294.675 Tm 0 Tw (Platyhelminthe)Tj /F1 1 Tf 9 0 0 9 480.87 294.675 Tm 0.0329 Tw (\): evidence for)Tj -18.1667 -1.2 TD -0.0695 Tw [(a trans-phyla horizontal transfer)52(. Mol. Biol. Evol.)]TJ /F3 1 Tf 9 0 1.913 9 504.21 283.875 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 506.07 283.875 Tm -0.0695 Tw (12: 421-)Tj -20.9667 -1.2 TD -0.0006 Tc 0 Tw (431.)Tj -0.6333 -1.2 TD 0 Tc -0.0338 Tw [(Gloor)51.7(, G.B., Preston, C.R., Johnson-Schlitz, D.M., Nassif,)]TJ 0.6333 -1.2 TD 0.0038 Tc 0.1389 Tw [(N.A., Phillis, R.W)58.2(., Benz, W)55.5(.K., Robertson, H.M., and)]TJ T* 0.0212 Tc [(Engels, W)53.4(.R. 1993. )25.9(T)52.2(ype I repressors of )]TJ /F3 1 Tf 9 0 1.913 9 495.15 240.675 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 501.33 240.675 Tm ( )Tj 0.4381 0 TD 0.0212 Tc (element)Tj -20.8781 -1.2 TD 0 Tc 0.0002 Tw [(mobility)75.7(. Genetics)]TJ /F3 1 Tf 9 0 1.913 9 387.69 229.875 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 390.21 229.875 Tm 0.0002 Tw (135: 81-95.)Tj -8.7267 -1.2 TD 0.0317 Tc 0.1389 Tw [(Gomulski, L.M., )15.5(T)109.4(orti, C., Bonizzoni, M., Moralli, D.,)]TJ 0.6333 -1.2 TD 0 Tc -0.0371 Tw [(Raimondi, E., Capy)73.3(, P)132.6(., Gasperi, G., and Malacrida, )53.7(A.R.)]TJ T* 0.0136 Tc 0.1389 Tw [(2001. )60.4(A)53.9( new basal subfamily of )]TJ /F3 1 Tf 9 0 1.913 9 454.71 197.475 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 486.09 197.475 Tm 0 Tc ( )Tj 0.4305 0 TD 0.0136 Tc 0.1389 Tw (elements in)Tj /F3 1 Tf 9 0 1.913 9 317.37 186.675 Tm 0 Tc 0.068 Tw (Ceratitis rosa)Tj /F1 1 Tf 9 0 0 9 371.01 186.675 Tm 0 Tw ( )Tj 0.346 0 TD 0.068 Tw (and other tephritid flies. J. Mol. Evol. 53:)Tj -6.306 -1.2 TD 0.0003 Tc 0 Tw (597-606.)Tj -0.6333 -1.2 TD 0 Tc 0.1289 Tw [(Gonzalez, P)122.6(., and Lessios, H.A. 1999. Evolution of sea)]TJ 0.6333 -1.2 TD -0.0552 Tw (urchin retroviral-like \(SURL\) elements: evidence from 40)Tj T* -0.0001 Tw (echinoid species. Mol. Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 448.41 143.475 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 450.87 143.475 Tm -0.0001 Tw (16: 938-952.)Tj -15.4667 -1.2 TD -0.0012 Tc -0.0695 Tw [(Good, )51.8(A.G., Meister)56.1(, G.A., Brock, H.W)58.4(., Grigliatti, )12.8(T)109.8(.A., and)]TJ 0.6333 -1.2 TD 0.0229 Tc 0.1389 Tw [(Hickey)70.9(, D.A. 1989. Rapid spread of transposable )]TJ /F3 1 Tf 9 0 1.913 9 532.47 121.875 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 317.37 111.075 Tm 0.0243 Tc 0.1389 Tw (elements in experimental populations of )Tj /F3 1 Tf 9 0 1.913 9 493.47 111.075 Tm 0 Tw (Drosophila)Tj -19.3116 -1.2 TD 0 Tc (melanogaster)Tj /F1 1 Tf 9 0 0 9 371.91 100.275 Tm -0.0184 Tw (. Genetics)Tj /F3 1 Tf 9 0 1.913 9 412.23 100.275 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 414.57 100.275 Tm -0.0184 Tw (122: 387-396.)Tj ET endstream endobj 37 0 obj << /Type /Page /Parent 95 0 R /Resources 38 0 R /Contents 39 0 R /Thumb 78 0 R /MediaBox [ 0 0 595 842 ] /CropBox [ 0 0 595 842 ] /Rotate 0 >> endobj 38 0 obj << /ProcSet [ /PDF /Text ] /Font << /F1 114 0 R /F2 113 0 R /F3 128 0 R >> /ExtGState << /GS1 139 0 R /GS2 138 0 R >> /ColorSpace << /Cs8 112 0 R >> >> endobj 39 0 obj << /Length 16999 >> stream BT /F1 1 Tf 9 0 0 9 55.93 779.115 Tm 0 0 0 1 k /GS2 gs -0.0001 Tc -0.0003 Tw (12 Silva )Tj /F3 1 Tf 9 0 1.913 9 95.47 779.115 Tm 0 Tc (et al)Tj /F1 1 Tf 9 0 0 9 112.45 779.115 Tm 0 Tw (.)Tj /F2 1 Tf 37.9351 29.4096 -29.4096 37.9351 34.496 531.1378 Tm 2 Tr 0 0 0 0.45 K 0 J 0 j 0.576 w 10 M []0 d /Cs8 cs 0 scn /GS1 gs 0.0686 Tc -0.0001 Tw (PAGE PROOFS)Tj /F1 1 Tf 9 0 0 9 56.53 748.275 Tm 0 Tr 0 0 0 1 k /GS2 gs 0.0279 Tc 0.139 Tw [(Gueiros-Filho, F)107.4(.J., and Beverley)74.2(, S.M. 1997. )23.4(T)32.2(r)0(ans-)]TJ 0.6333 -1.2 TD -0.0016 Tc -0.0694 Tw (kingdom transposition of the Drosophila element )Tj /F3 1 Tf 9 0 1.913 9 252.91 737.475 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 62.23 726.675 Tm 0.0044 Tc 0.1389 Tw (within the protozoan )Tj /F3 1 Tf 9 0 1.913 9 149.83 726.675 Tm 0 Tw (Leishmania)Tj /F1 1 Tf 9 0 0 9 196.27 726.675 Tm 0.1389 Tw (. Science)Tj /F3 1 Tf 9 0 1.913 9 234.85 726.675 Tm 0 Tc 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 238.63 726.675 Tm 0.0044 Tc 0.1389 Tw (276: 1716-)Tj -19.6 -1.2 TD -0.0006 Tc 0 Tw (1719.)Tj -0.6333 -1.2 TD -0.0099 Tc -0.0694 Tw [(Hagemann, S., Haring, E., and Pinsker)50.7(, W)54.2(.)0( 1996. Repeated)]TJ 0.6333 -1.2 TD -0.0119 Tc (horizontal transfer of )Tj /F3 1 Tf 9 0 1.913 9 142.39 694.275 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 148.33 694.275 Tm ( )Tj 0.1967 0 TD -0.0119 Tc -0.0694 Tw (transposons between )Tj /F3 1 Tf 9 0 1.913 9 234.73 694.275 Tm 0 Tw (Scaptomyza)Tj -18.9116 -1.2 TD 0 Tc (pallida)Tj /F1 1 Tf 9 0 0 9 88.21 683.475 Tm -0.0001 Tc -0.0006 Tw ( and )Tj /F3 1 Tf 9 0 1.913 9 108.25 683.475 Tm 0 Tc (Drosophila bifasciata)Tj /F1 1 Tf 9 0 0 9 191.77 683.475 Tm (. Genetica)Tj /F3 1 Tf 9 0 1.913 9 232.81 683.475 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 235.33 683.475 Tm -0.0007 Tw (98: 43-51.)Tj -19.8667 -1.2 TD 0.0449 Tc 0.1389 Tw [(Hagemann, S., Miller)52.1(, W)53.6(.J., and Pinsker)55.1(, W)53.6(.)0( 1992.)]TJ 0.6333 -1.2 TD 0 Tc -0.049 Tw (Identification of a complete )Tj /F3 1 Tf 9 0 1.913 9 169.99 661.875 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 175.99 661.875 Tm -0.049 Tw (-elements in the genome of)Tj /F3 1 Tf 9 0 1.913 9 62.23 651.075 Tm -0.0003 Tw (Drosophila bifasciata)Tj /F1 1 Tf 9 0 0 9 145.75 651.075 Tm [(. Nucleic )58.1(Acids Res. 20: 409-413.)]TJ -9.9133 -1.2 TD 0.1292 Tw [(Hagemann, S., Miller)54.5(, W)49.2(.J., and Pinsker)57.2(, W)55.9(.)0( 1994. )14.5(T)57.7(w)0(o)]TJ 0.6333 -1.2 TD 0.0586 Tc 0 Tw (distinct )Tj /F3 1 Tf 9 0 1.913 9 98.71 629.475 Tm 0 Tc (P)Tj /F1 1 Tf 9 0 0 9 105.25 629.475 Tm ( )Tj 0.4755 0 TD 0.0586 Tc 0.1389 Tw (element subfamilies in the genome of)Tj /F3 1 Tf 9 0 1.913 9 62.23 618.675 Tm 0 Tc 0.0005 Tw (Drosophila bifasciata)Tj /F1 1 Tf 9 0 0 9 145.75 618.675 Tm (. Mol. Gen. Genet. 244: 168-175.)Tj -9.9133 -1.2 TD 0.037 Tw [(Hamada, M., Kido, )16.7(Y)127(., Himberg, M., Reist, J.D., )14.7(Y)40.3(ing, C.,)]TJ 0.6333 -1.2 TD 0.1246 Tw [(Hasegawa, M., and Okada, N. 1997. )60.8(A)47( newly isolated)]TJ T* 0.0188 Tw (family of short interspersed repetitive elements \(SINEs\))Tj T* 0.0791 Tw (in coregonid fishes \(whitefish\) with sequences that are)Tj T* 0.0617 Tw (almost identical to those of the )Tj /F3 1 Tf 9 0 1.913 9 189.61 564.675 Tm 0 Tw (Sma)Tj /F1 1 Tf 9 0 0 9 208.09 564.675 Tm 0.0617 Tw (I family of repeats:)Tj -16.2067 -1.2 TD 0.103 Tw (possible evidence for the horizontal transfer of SINEs.)Tj T* 0 Tw (Genetics)Tj /F3 1 Tf 9 0 1.913 9 97.69 543.075 Tm ( )Tj /F1 1 Tf 9 0 0 9 100.21 543.075 Tm -0.0008 Tw (146: 355-367.)Tj -4.8533 -1.2 TD -0.0112 Tc -0.0695 Tw [(Handler)55.8(, )50.7(A. 2002. Use of the )]TJ /F3 1 Tf 9 0 1.913 9 164.95 532.275 Tm 0 Tw (piggyBac)Tj /F1 1 Tf 9 0 0 9 201.13 532.275 Tm 0 Tc ( )Tj 0.1973 0 TD -0.0112 Tc -0.0695 Tw (transposon for germ-)Tj -15.6306 -1.2 TD 0 Tc -0.0316 Tw (line transformation of insects. Insect Biochem. Mol. Biol.)Tj T* -0.002 Tw [(32: 121)74(1.)]TJ -0.6333 -1.2 TD -0.0053 Tw [(Haring, E., Hagemann, S., and Pinsker)50.2(, W)54.7(.)0( 2000. )52.1(Ancient)]TJ 0.6333 -1.2 TD 0.0158 Tc 0.1389 Tw (and recent horizontal invasions of drosophilids by )Tj /F3 1 Tf 9 0 1.913 9 277.33 489.075 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 62.23 478.275 Tm -0.0005 Tw (elements. J. Mol. Evol. 51: 577-586.)Tj -0.6333 -1.2 TD -0.0124 Tc -0.0695 Tw [(Hartl, D.L., Lohe, )52.5(A.R., and Lozovskaya, E.R. 1997. Modern)]TJ 0.6333 -1.2 TD 0.0042 Tc 0.1389 Tw (thoughts on an )Tj /F3 1 Tf 9 0 1.913 9 128.53 456.675 Tm 0 Tw (ancyent)Tj /F1 1 Tf 9 0 0 9 160.33 456.675 Tm 0 Tc ( )Tj /F3 1 Tf 9 0 1.913 9 164.17 456.675 Tm 0.0042 Tc (marinere)Tj /F1 1 Tf 9 0 0 9 199.93 456.675 Tm 0.1389 Tw (: function, evolution,)Tj -15.3 -1.2 TD 0 Tc -0.0001 Tw [(regulation. )53.6(Annu. Rev)75.6(. Genet. 31: 337-358.)]TJ -0.6333 -1.2 TD -0.0373 Tw [(Heath, B.D., Butcher)56.5(, R.D., Whitfield, W)47.5(.G., and Hubbard,)]TJ 0.6333 -1.2 TD 0.0133 Tc 0.1389 Tw [(S.F)115.9(. 1999. Horizontal transfer of )]TJ /F3 1 Tf 9 0 1.913 9 201.61 424.275 Tm 0 Tw (Wolbachia)Tj /F1 1 Tf 9 0 0 9 244.69 424.275 Tm 0 Tc ( )Tj 0.4302 0 TD 0.0133 Tc (between)Tj -20.7036 -1.2 TD 0.0115 Tc 0.1389 Tw (phylogenetically distant insect species by a naturally)Tj T* 0 Tc 0.0004 Tw [(occurring mechanism. Curr)56.3(. Biol. 9: 313-316.)]TJ -0.6333 -1.2 TD 0.002 Tc 0.1389 Tw [(Herdia, F)106.5(.)0( 2002 Evoluo do retroelemento )]TJ /F3 1 Tf 9 0 1.913 9 243.43 391.875 Tm 0 Tw (gypsy)Tj /F1 1 Tf 9 0 0 9 267.01 391.875 Tm 0 Tc ( )Tj 0.4189 0 TD 0.002 Tc (em)Tj -23.1722 -1.2 TD 0.0307 Tc 0.1389 Tw (espcies de )Tj /F3 1 Tf 9 0 1.913 9 118.57 381.075 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 164.35 381.075 Tm 0 Tc ( )Tj 0.4476 0 TD 0.0307 Tc (e )Tj /F3 1 Tf 9 0 1.913 9 177.67 381.075 Tm 0.1389 Tw (Zaprionis indianus)Tj /F1 1 Tf 9 0 0 9 256.93 381.075 Tm (. Uma)Tj -21.6333 -1.2 TD 0 Tc 0.0735 Tw [(abordagem filogentica, Doctoral )18.3(Thesis, Universidade)]TJ T* -0.0001 Tw [(Federal do Rio Grande do Sul, Porto )53.8(Alegre, Brazil.)]TJ -0.6333 -1.2 TD 0.0038 Tc 0.139 Tw [(Hickey)69.7(, D.A. 1982. Selfish DNA: a sexually-transmitted)]TJ 0.6333 -1.2 TD 0 Tc -0.0001 Tw (nuclear parasite. Genetics)Tj /F3 1 Tf 9 0 1.913 9 166.75 337.875 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 169.27 337.875 Tm -0.0001 Tw (101: 519-531.)Tj -12.5267 -1.2 TD -0.0107 Tc -0.0694 Tw [(Higuet, D., Merot, H., )49.6(Allouis, S., and Montchamp-Moreau,)]TJ 0.6333 -1.2 TD -0.0097 Tc -0.0695 Tw [(C. 1996. )24.8(The relationship between structural variation and)]TJ T* 0 Tc -0.0559 Tw (dysgenic properties of )Tj /F3 1 Tf 9 0 1.913 9 151.27 305.475 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 157.27 305.475 Tm ( )Tj 0.2221 0 TD -0.0559 Tw (elements in long-established P-)Tj -10.7821 -1.2 TD 0.0927 Tw (transformed lines of )Tj /F3 1 Tf 9 0 1.913 9 146.23 294.675 Tm (Drosophila simulans)Tj /F1 1 Tf 9 0 0 9 228.13 294.675 Tm (. Heredity)Tj /F3 1 Tf 9 0 1.913 9 267.43 294.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 270.79 294.675 Tm (77:)Tj -23.1733 -1.2 TD -0.0004 Tc (9-15.)Tj -0.6333 -1.2 TD 0 Tc -0.0205 Tw [(Horn, C., Of)14.1(fen, N., Nystedt, S., Hacker)54.2(, U., and Wimmer)57(,)]TJ 0.6333 -1.2 TD -0.0565 Tw (E.A. 2003. )Tj /F3 1 Tf 9 0 1.913 9 105.73 262.275 Tm 0 Tw (piggyBac)Tj /F1 1 Tf 9 0 0 9 142.75 262.275 Tm -0.0565 Tw (-based insertional mutagenesis and)Tj -8.9467 -1.2 TD 0.0436 Tc 0.1389 Tw (enhancer detection as a tool for functional insect)Tj T* 0 Tc 0.0006 Tw (genomics. Genetics)Tj /F3 1 Tf 9 0 1.913 9 141.25 240.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 143.77 240.675 Tm 0.0006 Tw (163: 647-661.)Tj -9.6933 -1.2 TD -0.0054 Tc -0.0695 Tw (Houck, M.A., Clark, J.B., Peterson, K.R., and Kidwell, M.G.)Tj 0.6333 -1.2 TD 0 Tc 0.0921 Tw (1991. Possible horizontal transfer of )Tj /F3 1 Tf 9 0 1.913 9 212.47 219.075 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 255.43 219.075 Tm ( )Tj 0.3701 0 TD (genes)Tj -21.8368 -1.2 TD 0.0008 Tc 0.139 Tw (by the mite )Tj /F3 1 Tf 9 0 1.913 9 112.57 208.275 Tm (Proctolaelaps regalis)Tj /F1 1 Tf 9 0 0 9 197.53 208.275 Tm (. Science)Tj /F3 1 Tf 9 0 1.913 9 235.87 208.275 Tm 0 Tc 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 239.59 208.275 Tm 0.0008 Tc 0.139 Tw [(253: 1)70.6(125-)]TJ -19.7067 -1.2 TD 0 Tc 0 Tw (1)Tj 0.48 0 TD -0.0002 Tc (128.)Tj -1.1133 -1.2 TD -0.0028 Tc -0.0695 Tw [(Ivics, Z., Hackett, P)126.8(.B., Plasterk, R.H., and Izsvk, Z. 1997.)]TJ 0.6333 -1.2 TD 0 Tc 0.0337 Tw (Molecular reconstruction of )Tj /F3 1 Tf 9 0 1.913 9 173.65 175.875 Tm (Sleeping Beauty)Tj /F1 1 Tf 9 0 0 9 239.53 175.875 Tm (, a )Tj /F3 1 Tf 9 0 1.913 9 252.61 175.875 Tm -0.091 Tc 0 Tw [(Tc)-91(1)]TJ /F1 1 Tf 9 0 0 9 266.83 175.875 Tm 0 Tc (-like)Tj -22.7333 -1.2 TD -0.0058 Tc -0.0694 Tw (transposon from fish, and its transposition in human cells.)Tj T* 0 Tc 0 Tw (Cell)Tj /F3 1 Tf 9 0 1.913 9 77.71 154.275 Tm ( )Tj /F1 1 Tf 9 0 0 9 80.23 154.275 Tm 0.0002 Tw (91: 501-510.)Tj -2.6333 -1.2 TD 0.1336 Tw [(Ivics, Z., Izsvk, Z., Minter)48.7(, )56.3(A., and Hackett, P)134.3(.B. 1996.)]TJ 0.6333 -1.2 TD -0.0064 Tc -0.0694 Tw (Indentification of functional domains and evolution of )Tj /F3 1 Tf 9 0 1.913 9 266.29 132.675 Tm -0.0977 Tc 0 Tw [(Tc)-91.3(1)]TJ /F1 1 Tf 9 0 0 9 280.33 132.675 Tm 0 Tc (-)Tj -24.2333 -1.2 TD 0.0238 Tw [(like transposable elements. Proc. Natl. )59.1(Acad. Sci. USA.)]TJ T* -0.0002 Tw (93: 5008-5013.)Tj -0.6333 -1.2 TD 0.038 Tc 0.1389 Tw [(Jackson, M.S., Black, D.M., and Dover)50.9(, G.A. 1988.)]TJ 28.3533 72 TD 0.0473 Tc (Amplification of KP elements associated with the)Tj 0 -1.2 TD 0.0695 Tc 0.2666 Tw (repression of hybrid dysgenesis in )Tj /F3 1 Tf 9 0 1.913 9 484.15 737.475 Tm 0 Tw (Drosophila)Tj -18.9049 -1.2 TD 0 Tc (melanogaster)Tj /F1 1 Tf 9 0 0 9 366.25 726.675 Tm -0.0188 Tw (. Genetics)Tj /F3 1 Tf 9 0 1.913 9 406.63 726.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 408.91 726.675 Tm -0.0188 Tw (120: 1003-1013.)Tj -11.4267 -1.2 TD 0.0361 Tc 0.1389 Tw [(Jarvik, )21.3(T)113.8(., and Lark, K.G. 1998. Characterization of)]TJ /F3 1 Tf 9 0 1.913 9 311.71 705.0751 Tm 0 Tc 0 Tw (Soymar1)Tj /F1 1 Tf 9 0 0 9 347.71 705.0751 Tm 0.0157 Tw (, a )Tj /F3 1 Tf 9 0 1.913 9 360.49 705.0751 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 391.03 705.0751 Tm ( )Tj 0.2937 0 TD 0.0157 Tw (element in soybean. Genetics)Tj /F3 1 Tf 9 0 1.913 9 512.65 705.0751 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 515.29 705.0751 Tm (149:)Tj -22.62 -1.2 TD 0.0001 Tc (1569-1574.)Tj -0.6267 -1.2 TD -0.0059 Tc -0.0694 Tw [(Jehle, J.A., Fritsch, E., Nickel, )59.6(A., Huber)48.3(, J., and Backhaus,)]TJ 0.6267 -1.2 TD 0 Tc 0.0018 Tw [(H. 1995. )15(TCl4.7: a novel lepidopteran transposon found)]TJ T* 0 Tw (in )Tj /F3 1 Tf 9 0 1.913 9 321.43 661.875 Tm 0.0261 Tw (Cydia pomonella)Tj /F1 1 Tf 9 0 0 9 388.69 661.875 Tm 0 Tw ( )Tj 0.3041 0 TD 0.0261 Tw [(granulosis virus. V)12(irology)]TJ /F3 1 Tf 9 0 1.913 9 491.83 661.875 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 494.53 661.875 Tm 0.0261 Tw (207: 369-)Tj -20.3133 -1.2 TD -0.0006 Tc 0 Tw (379.)Tj -0.6267 -1.2 TD 0.0213 Tc 0.1389 Tw [(Jensen, S., Gassama, M.P)129.3(., and Heidmann, )18.9(T)105.6(.)0( 1999.)]TJ 0.6267 -1.2 TD 0 Tc 0 Tw (T)Tj 0.5667 0 TD 0.0678 Tc 0.1389 Tw (aming of transposable elements by homology-)Tj -0.5667 -1.2 TD 0 Tc 0 Tw (dependent gene silencing. Nat. Genet. 21: 209-212.)Tj -0.6267 -1.2 TD 0.1104 Tw [(Jordan, I.K., Matyunina, L.V)98.1(., and McDonald, J.F)104.1(. 1999.)]TJ 0.6267 -1.2 TD -0.0094 Tc -0.0695 Tw [(Evidence for the recent horizontal transfer of long terminal)]TJ T* 0 Tc 0.0316 Tw [(repeat retrotransposon. Proc. Natl. )57.9(Acad. Sci. USA. 96:)]TJ T* 0 Tw (12621-12625.)Tj -0.6267 -2.4 TD -0.0377 Tw [(Jordan, I.K., Rogozin, I.B., Glazko, G.V)93.6(., and Koonin, E.V)94.6(.)]TJ 0.6267 -1.2 TD -0.001 Tc -0.0695 Tw (2003. Origin of a substantial fraction of human regulatory)Tj T* 0 Tc 0.0439 Tw [(sequences from transposable elements. )17.1(T)37.7(rends Genet.)]TJ T* 0.0021 Tw (19: 68-72.)Tj -0.6267 -1.2 TD 0.0533 Tc 0.1389 Tw [(Kapitonov)73.1(, V)95.1(.)0(V)91.6(., and Jurka, J. 2001. Rolling-circle)]TJ 0.6267 -1.2 TD 0 Tc 0.0476 Tw [(transposons in eukaryotes. Proc. Natl. )58.7(Acad. Sci. USA.)]TJ T* -0.0002 Tw (98: 8714-8719.)Tj -0.6267 -1.2 TD 0.0894 Tw [(Kaplan, N., Darden, )21.7(T)111(., and Langley)67.6(, C. 1985. Evolution)]TJ 0.6267 -1.2 TD 0.0032 Tc 0.1389 Tw (and extinction of transposable elements in Mendelian)Tj T* 0 Tc -0.0001 Tw (populations. Genetics)Tj /F3 1 Tf 9 0 1.913 9 398.23 456.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 400.75 456.675 Tm -0.0001 Tw (109: 459-480.)Tj -10.52 -1.2 TD -0.0121 Tc -0.0695 Tw [(Kaufman, P)127.8(.D., and Rio, D.C. 1992. )]TJ /F3 1 Tf 9 0 1.913 9 442.39 445.875 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 448.33 445.875 Tm ( )Tj 0.1964 0 TD -0.0121 Tc -0.0695 Tw (element transposition)Tj /F3 1 Tf 9 0 1.913 9 311.71 435.075 Tm -0.0093 Tc (in vitro)Tj /F1 1 Tf 9 0 0 9 336.91 435.075 Tm 0 Tc 0 Tw ( )Tj 0.1992 0 TD -0.0093 Tc -0.0695 Tw (proceeds by a cut-and-paste mechanism and uses)Tj -2.9992 -1.2 TD 0 Tc 0.0008 Tw [(GTP)16( as a cofactor)58.7(. Cell)]TJ /F3 1 Tf 9 0 1.913 9 404.11 424.275 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 406.63 424.275 Tm 0.0008 Tw (69: 27-39.)Tj -11.1733 -1.2 TD 0.0389 Tc 0.1389 Tw (Kidwell, M.G. 1983. Evolution of hybrid dysgenesis)Tj 0.6267 -1.2 TD 0.0082 Tc (determinants in )Tj /F3 1 Tf 9 0 1.913 9 379.39 402.675 Tm (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 482.41 402.675 Tm (. Proc. Natl.)Tj -18.9667 -1.2 TD 0 Tc 0.0007 Tw (Acad. Sci. USA. 80: 1655-1659.)Tj -0.6267 -1.2 TD 0.0226 Tc 0.1389 Tw (Kidwell, M.G. 1985. Hybrid dysgenesis in )Tj /F3 1 Tf 9 0 1.913 9 487.93 381.075 Tm 0 Tw (Drosophila)Tj -19.3249 -1.2 TD 0.0266 Tc (melanogaster)Tj /F1 1 Tf 9 0 0 9 369.13 370.275 Tm 0.1389 Tw (: nature and inheritance of )Tj /F3 1 Tf 9 0 1.913 9 489.13 370.275 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 495.37 370.275 Tm ( )Tj 0.4435 0 TD 0.0266 Tc (element)Tj -20.8501 -1.2 TD 0 Tc -0.0004 Tw (regulation. Genetics)Tj /F3 1 Tf 9 0 1.913 9 391.75 359.475 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 394.27 359.475 Tm -0.076 Tc (11)Tj 0.96 0 TD 0 Tc -0.0005 Tw (1: 337-350.)Tj -10.76 -1.2 TD -0.0328 Tw (Kidwell, M.G. 1992. Horizontal transfer of )Tj /F3 1 Tf 9 0 1.913 9 470.83 348.675 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 476.83 348.675 Tm ( )Tj 0.2452 0 TD -0.0328 Tw (elements and)Tj -18.5919 -1.2 TD 0.001 Tc 0.1389 Tw (other short inverted repeat transposons. Genetica)Tj /F3 1 Tf 9 0 1.913 9 516.49 337.875 Tm 0 Tc 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 520.27 337.875 Tm 0.001 Tc (86:)Tj -23.1733 -1.2 TD 0.0003 Tc (275-286.)Tj -0.6267 -1.2 TD 0 Tc -0.0377 Tw (Kidwell, M.G. 1993. Lateral transfer in natural populations)Tj 0.6267 -1.2 TD -0.0001 Tw [(of eukaryotes. )52.2(Annu. Rev)75.6(. Genet. 27: 235-256.)]TJ -0.6267 -1.2 TD -0.0058 Tc -0.0694 Tw [(Kidwell, M.G. 1994. )17.5(The evolutionary history of the )]TJ /F3 1 Tf 9 0 1.913 9 501.79 294.675 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 507.73 294.675 Tm ( )Tj 0.2028 0 TD -0.0058 Tc (family)Tj -21.9828 -1.2 TD 0 Tc -0.0003 Tw (of transposable elements. J. Hered. 85: 339-346.)Tj -0.6267 -1.2 TD 0.0578 Tw [(Kidwell, M.G., Kidwell, J.F)111.4(., and Sved, J.A. 1977. Hybrid)]TJ 0.6267 -1.2 TD 0.0368 Tw (dysgenesis in )Tj /F3 1 Tf 9 0 1.913 9 369.43 262.275 Tm 0.0367 Tw (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 469.81 262.275 Tm (: a syndrome of)Tj -17.5667 -1.2 TD 0.022 Tc 0.1389 Tw (aberrant traits including mutation, sterility and male)Tj T* 0 Tc 0.0003 Tw (recombination. Genetics)Tj /F3 1 Tf 9 0 1.913 9 408.73 240.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 411.25 240.675 Tm 0.0003 Tw (36: 813-833.)Tj -11.6867 -1.2 TD -0.0026 Tc -0.0695 Tw (Kidwell, M.G., Kimura, K., and Black, D.M. 1988. Evolution)Tj 0.6267 -1.2 TD 0.0328 Tc 0.139 Tw (of hybrid dysgenesis potential following)Tj /F3 1 Tf 9 0 1.913 9 484.63 219.075 Tm 0.1718 Tc 0 Tw ( P)Tj /F1 1 Tf 9 0 0 9 494.95 219.075 Tm 0 Tc ( )Tj 0.4498 0 TD 0.0328 Tc (element)Tj -20.8098 -1.2 TD -0.0077 Tc -0.0694 Tw (contamination in )Tj /F3 1 Tf 9 0 1.913 9 377.29 208.275 Tm (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 475.15 208.275 Tm (. Genetics)Tj /F3 1 Tf 9 0 1.913 9 514.33 208.275 Tm 0 Tc 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 516.13 208.275 Tm (1)Tj 0.48 0 TD -0.0077 Tc (19:)Tj -23.1933 -1.2 TD 0.0003 Tc (815-828.)Tj -0.6267 -1.2 TD 0 Tc -0.0694 Tw [(Kidwell, M.G., and Lisch, D. 2000. )17.1(T)44.3(ransposable elements)]TJ 0.6267 -1.2 TD 0.0689 Tw [(and host genome evolution. )18.8(T)37.7(rends Ecol. Evol. 15: 95-)]TJ T* -0.0006 Tc 0 Tw (99.)Tj -0.6267 -1.2 TD 0.0527 Tc 0.139 Tw [(Kim, )54(A., )24.3(T)110.4(erzian, C., Santamaria, P)126.5(., Plisson, )52.6(A.,)]TJ 0.6267 -1.2 TD 0 Tc 0.083 Tw [(Prudhomme, N., and Bucheton, )59.2(A. 1994. Retroviruses)]TJ T* -0.0559 Tw (in invertebrates: the )Tj /F3 1 Tf 9 0 1.913 9 391.75 132.675 Tm 0 Tw (gypsy)Tj /F1 1 Tf 9 0 0 9 415.27 132.675 Tm ( )Tj 0.2221 0 TD -0.0559 Tw (retrotransposon is apparently)Tj -11.7287 -1.2 TD -0.0083 Tc -0.0695 Tw (an infectious retrovirus of )Tj /F3 1 Tf 9 0 1.913 9 410.17 121.875 Tm (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 507.85 121.875 Tm (. Proc.)Tj -21.7933 -1.2 TD 0 Tc 0.0003 Tw [(Natl. )54.3(Acad. Sci. USA. 91: 1285-1289.)]TJ -0.6267 -1.2 TD 0.0181 Tc 0.1389 Tw [(Kimura, K., and Kidwell, M.G. 1994. Dif)20.4(ferences in )]TJ /F3 1 Tf 9 0 1.913 9 526.81 100.275 Tm 0 Tc 0 Tw (P)Tj ET endstream endobj 40 0 obj << /Type /Page /Parent 95 0 R /Resources 41 0 R /Contents 42 0 R /Thumb 80 0 R /MediaBox [ 0 0 595 842 ] /CropBox [ 0 0 595 842 ] /Rotate 0 >> endobj 41 0 obj << /ProcSet [ /PDF /Text ] /Font << /F1 114 0 R /F2 113 0 R /F3 128 0 R >> /ExtGState << /GS1 139 0 R /GS2 138 0 R >> /ColorSpace << /Cs8 112 0 R >> >> endobj 42 0 obj << /Length 16976 >> stream BT /F1 1 Tf 9 0 0 9 337.47 780.5551 Tm 0 0 0 1 k /GS2 gs 0 Tc -0.0155 Tw [(Horizontal Transfer of Transposable Elements 13)]TJ /F2 1 Tf 37.9351 29.4096 -29.4096 37.9351 42.9423 542.4378 Tm 2 Tr 0 0 0 0.45 K 0 J 0 j 0.576 w 10 M []0 d /Cs8 cs 0 scn /GS1 gs 0.0686 Tc -0.0001 Tw (PAGE PROOFS)Tj /F1 1 Tf 9 0 0 9 67.83 748.275 Tm 0 Tr 0 0 0 1 k /GS2 gs -0.0088 Tc -0.0695 Tw (element population dynamics between the sibling species)Tj /F3 1 Tf 9 0 1.913 9 67.83 737.475 Tm 0.0295 Tc 0.1389 Tw (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 175.23 737.475 Tm 0 Tc 0 Tw ( )Tj 0.4465 0 TD 0.0295 Tc (and )Tj /F3 1 Tf 9 0 1.913 9 199.11 737.475 Tm 0.1389 Tw (Drosophila simulans)Tj /F1 1 Tf 9 0 0 9 286.47 737.475 Tm 0 Tc 0 Tw (.)Tj -24.2933 -1.2 TD -0.0002 Tw (Genet. Res. 63: 27-38.)Tj -0.6267 -1.2 TD 0.059 Tc 0.1389 Tw [(Kimura, M. 1980. )57.7(A)52.7( simple method for estimating)]TJ 0.6267 -1.2 TD 0.0695 Tc 0.1829 Tw (evolutionary rate of base substitutions through)Tj T* 0.0054 Tc 0.1389 Tw (comparative studies of nucleotide sequences. J. Mol.)Tj T* 0 Tc 0 Tw (Evol.)Tj /F3 1 Tf 9 0 1.913 9 87.87 683.475 Tm ( )Tj /F1 1 Tf 9 0 0 9 90.33 683.475 Tm -0.001 Tw [(16: 1)69.6(1)76(1-120.)]TJ -3.1267 -1.2 TD -0.0585 Tw [(Koga, )52.5(A., Shimada, )52.8(A., Shima, )60.8(A., Sakaizumi, M., )14(T)111(achida,)]TJ 0.6267 -1.2 TD 0.1048 Tw (H., and Hori, H. 2000. Evidence for recent invasion of)Tj T* 0.0327 Tc 0.139 Tw (the medaka fish genome by the )Tj /F3 1 Tf 9 0 1.913 9 212.25 651.075 Tm 0 Tc 0 Tw (T)Tj 0.5533 0 TD 0.0327 Tc (ol2)Tj /F1 1 Tf 9 0 0 9 230.13 651.075 Tm 0 Tc ( )Tj 0.4496 0 TD 0.0327 Tc (transposable)Tj -18.483 -1.2 TD 0 Tc 0.0006 Tw (element. Genetics)Tj /F3 1 Tf 9 0 1.913 9 140.37 640.275 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 142.89 640.275 Tm 0.0006 Tw (155: 273-281.)Tj -8.9667 -1.2 TD -0.0598 Tw (Kondo, N., Nikoh, N., Ijichi, N., Shimada, M., and Fukatsu,)Tj 0.6267 -1.2 TD -0.111 Tc 0 Tw (T.)Tj 0.778 0 TD ( )Tj 0.2961 0 TD 0 Tc 0.0181 Tw (2002. Genome fragment of )Tj /F3 1 Tf 9 0 1.913 9 188.25 618.675 Tm 0 Tw (Wolbachia)Tj /F1 1 Tf 9 0 0 9 230.25 618.675 Tm ( )Tj 0.2961 0 TD (endosymbiont)Tj -18.3428 -1.2 TD 0.0279 Tw (transferred to X chromosome of host insect. Proc. Natl.)Tj T* 0.0004 Tw (Acad. Sci. USA. 99: 14280-14285.)Tj -0.6267 -1.2 TD -0.0063 Tc -0.0694 Tw [(Konieczny)69.8(, )60.7(A., V)53.8(oytas, D.F)111.4(., Cummings, M.P)126.7(., and )57.2(Ausubel,)]TJ 0.6267 -1.2 TD 0 Tc 0 Tw (F)Tj 0.5533 0 TD 0.052 Tc 0.1389 Tw [(.M. 1991. )58(A)52.3( superfamily of )]TJ /F3 1 Tf 9 0 1.913 9 197.73 575.475 Tm (Arabidopsis thaliana)Tj /F1 1 Tf 9 0 0 9 67.83 564.675 Tm 0 Tc 0 Tw (retrotransposons. Genetics)Tj /F3 1 Tf 9 0 1.913 9 175.89 564.675 Tm ( )Tj /F1 1 Tf 9 0 0 9 178.41 564.675 Tm (127: 801-809.)Tj -12.9133 -1.2 TD 0.0258 Tc 0.139 Tw [(Kordis, D., and Gubensek, F)110.3(.)0( 1995. Horizontal SINE)]TJ 0.6267 -1.2 TD -0.0072 Tc -0.0695 Tw (transfer between vertebrate classes. Nat. Genet.)Tj /F3 1 Tf 9 0 1.913 9 255.21 543.075 Tm 0 Tc 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 257.01 543.075 Tm -0.0072 Tc -0.0695 Tw (10: 131-)Tj -21.02 -1.2 TD -0.0006 Tc 0 Tw (132.)Tj -0.6267 -1.2 TD 0.0323 Tc 0.1389 Tw [(Labrador)53.7(, M., and Corces, V)88.3(.G. 1997. )19.5(T)43.3(ransposable)]TJ 0.6267 -1.2 TD 0.012 Tc (element-host interactions: regulation of insertion and)Tj T* 0 Tc 0.0002 Tw [(excision. )54.9(Annu. Rev)69.2(. Genet. 31: 381-404.)]TJ -0.6267 -1.2 TD 0.0331 Tw [(Lam, W)58.1(.L., Seo, P)129.6(., Robison, K., V)12(irk, S., and Gilbert, W)55.5(.)]TJ 0.6267 -1.2 TD 0.038 Tc 0.1389 Tw (1996. Discovery of amphibian )Tj /F3 1 Tf 9 0 1.913 9 203.85 478.275 Tm -0.0577 Tc 0 Tw [(Tc)-95.7(1)]TJ /F1 1 Tf 9 0 0 9 219.03 478.275 Tm 0.038 Tc 0.1389 Tw (-like transposon)Tj -16.8 -1.2 TD 0 Tc -0.0005 Tw (families. J. Mol. Biol. 257:359-366.)Tj -0.6267 -1.2 TD 0.0527 Tw [(Lambert, M.E., McDonald, J.F)113.6(., and W)16.1(einstein, I.B., eds.)]TJ 0.6267 -1.2 TD 0.0328 Tw [(1988. Eukaryotic )18.3(T)31(ransposable Elements as Mutagenic)]TJ T* 0.0002 Tw (Agents. Cold Spring Harbor Laboratory Press.)Tj -0.6267 -1.2 TD 0.0262 Tw (Lampe, D.J., Churchill, M.E., and Robertson, H.M. 1996.)Tj 0.6267 -1.2 TD -0.0537 Tc 0 Tw (A )Tj 1.0128 0 TD 0 Tc (purified )Tj /F3 1 Tf 9 0 1.913 9 110.07 413.475 Tm (mariner)Tj /F1 1 Tf 9 0 0 9 140.55 413.475 Tm ( )Tj 0.3995 0 TD 0.1215 Tw [(transposase is suf)14.8(ficient to mediate)]TJ -8.4795 -1.2 TD 0 Tw (transposition )Tj /F3 1 Tf 9 0 1.913 9 121.41 402.675 Tm 0.0001 Tw (in vitro)Tj /F1 1 Tf 9 0 0 9 147.87 402.675 Tm (. EMBO J. 15: 5470-5479.)Tj -9.52 -1.2 TD 0.1266 Tw [(Lampe, D.J., W)37.6(alden, K.K., and Robertson, H.M. 2001.)]TJ 0.6267 -1.2 TD 0.1017 Tw (Loss of transposase-DNA interaction may underlie the)Tj T* -0.0466 Tw (divergence of )Tj /F3 1 Tf 9 0 1.913 9 123.57 370.275 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 154.05 370.275 Tm ( )Tj 0.2314 0 TD -0.0466 Tw (family transposable elements and)Tj -9.8114 -1.2 TD -0.0113 Tw (the ability of more than one )Tj /F3 1 Tf 9 0 1.913 9 178.29 359.475 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 208.77 359.475 Tm ( )Tj 0.2667 0 TD -0.0113 Tw (to occupy the same)Tj -15.9267 -1.2 TD 0.0003 Tw (genome. Mol. Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 164.91 348.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 167.37 348.675 Tm 0.0003 Tw (18: 954-961.)Tj -11.6867 -1.2 TD 0.0347 Tc 0.1389 Tw [(Langin, )20.9(T)112.4(., Capy)69.6(, P)125.9(., and Daboussi, M.J. 1995. )22.7(The)]TJ 0.6267 -1.2 TD 0 Tc 0.1346 Tw (transposable element )Tj /F3 1 Tf 9 0 1.913 9 158.85 327.075 Tm 0 Tw (impala)Tj /F1 1 Tf 9 0 0 9 185.37 327.075 Tm 0.1346 Tw (, a fungal member of the)Tj /F3 1 Tf 9 0 1.913 9 67.83 316.275 Tm 0 Tw (T)Tj 0.52 0 TD (c1-mariner)Tj /F1 1 Tf 9 0 0 9 115.53 316.275 Tm [( superfamily)70(. Mol. Gen. Genet. 246: 19-28.)]TJ -5.9267 -1.2 TD -0.01 Tc -0.0695 Tw [(Lcher)50.1(, P)130.1(., Bucheton, )58.8(A., and Plisson, )52.7(A. 1997. Expression)]TJ 0.6267 -1.2 TD 0.0083 Tc 0.1389 Tw (of the )Tj /F3 1 Tf 9 0 1.913 9 95.91 294.675 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 139.65 294.675 Tm 0 Tc ( )Tj 0.4252 0 TD 0.0083 Tc (retrovirus )Tj /F3 1 Tf 9 0 1.913 9 185.55 294.675 Tm (gypsy)Tj /F1 1 Tf 9 0 0 9 209.43 294.675 Tm 0 Tc ( )Tj 0.4252 0 TD 0.0083 Tc 0.1389 Tw (as ultrastructurally)Tj -16.1585 -1.2 TD 0.0144 Tc (detectable particles in the ovaries of flies carrying a)Tj T* 0 Tc 0 Tw (permissive )Tj /F3 1 Tf 9 0 1.913 9 112.77 273.075 Tm (flamenco)Tj /F1 1 Tf 9 0 0 9 149.25 273.075 Tm ( )Tj 0.2089 0 TD -0.0691 Tw [(allele. J. Gen. V)13.9(irol. 78: 2379-2388.)]TJ -9.8822 -1.2 TD -0.0045 Tw [(Lee, C.C., Mul, )13.5(Y)133.7(.M., and Rio, D.C. 1996. )15.8(The )]TJ /F3 1 Tf 9 0 1.913 9 245.97 262.275 Tm 0 Tw (Drosophila)Tj -19.5383 -1.2 TD (P)Tj /F1 1 Tf 9 0 0 9 73.83 251.475 Tm (-element KP repressor protein dimerizes and interacts)Tj -0.6667 -1.2 TD -0.0554 Tw (with multiple sites on )Tj /F3 1 Tf 9 0 1.913 9 151.41 240.675 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 157.41 240.675 Tm -0.0554 Tw (-element DNA. Mol. Cell. Biol. 16:)Tj -9.9533 -1.2 TD 0.0001 Tc 0 Tw (5616-5622.)Tj -0.6267 -1.2 TD 0 Tc 0.0299 Tw [(Lerat, E., Brunet, F)112(., Bazin, C., and Capy)73.2(, P)132.9(.)0( 1999. Is the)]TJ 0.6267 -1.2 TD 0.0771 Tw (evolution of transposable elements modular? Genetica)Tj T* 0.0015 Tw (107: 15-25.)Tj -0.6267 -1.2 TD 0.0085 Tc 0.1389 Tw (Lidholm, D.A., Gudmundsson, G.H., and Boman, H.G.)Tj 0.6267 -1.2 TD 0.0131 Tc [(1991. )50.7(A)60.1( highly repetitive, )]TJ /F3 1 Tf 9 0 1.913 9 176.07 175.875 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 207.45 175.875 Tm 0.1389 Tw (-like element in the)Tj -15.5133 -1.2 TD 0.0085 Tc (genome of )Tj /F3 1 Tf 9 0 1.913 9 116.13 165.075 Tm (Hyalophora cecropia.)Tj /F1 1 Tf 9 0 0 9 203.97 165.075 Tm 0 Tc 0 Tw ( )Tj 0.4254 0 TD 0.0085 Tc 0.1389 Tw (J. Biol. Chem.)Tj /F3 1 Tf 9 0 1.913 9 267.39 165.075 Tm 0 Tc 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 271.17 165.075 Tm 0.0085 Tc (266:)Tj -22.5933 -1.2 TD 0 Tc (1)Tj 0.4867 0 TD -0.0002 Tc [(1518-1)71.8(1521.)]TJ -1.1133 -1.2 TD 0 Tc -0.0356 Tw [(Lohe, )57.8(A.R., De )52.5(Aguiar)56.7(, D., and Hartl, D.L. 1997. Mutations)]TJ 0.6267 -1.2 TD 0.0091 Tc 0.1389 Tw (in the )Tj /F3 1 Tf 9 0 1.913 9 95.43 132.675 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 126.51 132.675 Tm 0 Tc ( )Tj 0.426 0 TD 0.0091 Tc 0.1389 Tw (transposase: the D,D\(35\)E consensus)Tj -6.946 -1.2 TD 0 Tc 0.0632 Tw [(sequence is nonfunctional. Proc. Natl. )53.5(Acad. Sci. USA.)]TJ T* -0.0002 Tw (94: 1293-1297.)Tj -0.6267 -1.2 TD -0.0027 Tc -0.0694 Tw [(Lohe, )54.6(A.R., and Hartl, D.L. 1996. )57.5(Autoregulation of )]TJ /F3 1 Tf 9 0 1.913 9 258.57 100.275 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 317.37 748.275 Tm 0.0038 Tc 0.1389 Tw (transposase activity by overproduction and dominant-)Tj T* 0 Tc -0.0283 Tw (negative complementation. Mol. Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 485.91 737.475 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 488.13 737.475 Tm -0.0283 Tw (13: 549-555.)Tj -19.6067 -1.2 TD -0.0603 Tw [(Lohe, )53.1(A.R., Moriyama, E.N., Lidholm, D.A., and Hartl, D.L.)]TJ 0.6333 -1.2 TD 0.0332 Tw (1995. Horizontal transmission, vertical inactivation, and)Tj T* 0.0026 Tc 0.139 Tw (stochastic loss of )Tj /F3 1 Tf 9 0 1.913 9 392.61 705.0751 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 423.27 705.0751 Tm 0.1389 Tw (-like transposable elements.)Tj -11.7667 -1.2 TD 0 Tc 0 Tw (Mol. Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 376.83 694.275 Tm ( )Tj /F1 1 Tf 9 0 0 9 379.35 694.275 Tm (12: 62-72.)Tj -7.52 -1.2 TD -0.0119 Tc -0.0695 Tw [(Loreto, E.L., V)75.3(alente, V)90.3(.L., Zaha, )52.9(A., Silva, J.C., and Kidwell,)]TJ 0.6333 -1.2 TD -0.0057 Tc -0.0694 Tw (M.G. 2001. )Tj /F3 1 Tf 9 0 1.913 9 362.55 672.675 Tm (Drosophila mediopunctata)Tj /F1 1 Tf 9 0 0 9 465.27 672.675 Tm 0 Tc 0 Tw ( )Tj /F3 1 Tf 9 0 1.913 9 467.07 672.675 Tm (P)Tj /F1 1 Tf 9 0 0 9 473.07 672.675 Tm ( )Tj 0.2029 0 TD -0.0057 Tc -0.0694 Tw (elements: a new)Tj -17.5029 -1.2 TD 0 Tc -0.0004 Tw [(example of horizontal transfer)57.7(. J. Hered. 92: 375-381.)]TJ -0.6333 -1.2 TD -0.002 Tc -0.0694 Tw (Luan, D.D., Korman, M.H., Jakubczak, J.L., and Eickbush,)Tj 0.6333 -1.2 TD 0 Tc 0 Tw (T)Tj 0.5 0 TD -0.0693 Tw (.H. 1993. Reverse transcription of )Tj /F3 1 Tf 9 0 1.913 9 455.31 640.275 Tm 0 Tw (R2Bm)Tj /F1 1 Tf 9 0 0 9 480.27 640.275 Tm ( )Tj 0.2087 0 TD -0.0693 Tw [(RNA)53.1( is primed)]TJ -18.3087 -1.2 TD 0.0433 Tw (by a nick at the chromosomal target site: a mechanism)Tj T* 0.0003 Tw [(for non-L)75.7(TR retrotransposition. Cell)]TJ /F3 1 Tf 9 0 1.913 9 457.23 618.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 459.75 618.675 Tm 0.0003 Tw (72: 595-605.)Tj -16.4533 -1.2 TD 0.023 Tw [(Mackay)71.7(, )19(T)111(.)0(F)109(.)0( 1989. )22.1(T)31(ransposable elements and fitness in)]TJ /F3 1 Tf 9 0 1.913 9 317.37 597.075 Tm -0.013 Tw (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 417.27 597.075 Tm (. Genome)Tj /F3 1 Tf 9 0 1.913 9 456.69 597.075 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 459.09 597.075 Tm -0.013 Tw (31: 284-295.)Tj -16.38 -2.4 TD -0.0691 Tw [(Makalowski, W)51.3(.)0( 2000. Genomic scrap yard: how genomes)]TJ 0.6333 -1.2 TD 0.0002 Tw (utilize all that junk. Gene)Tj /F3 1 Tf 9 0 1.913 9 415.41 564.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 417.87 564.675 Tm 0.0002 Tw (259: 61-67.)Tj -11.8 -1.2 TD 0.1039 Tw [(Malik, H.S., Burke, W)53.9(.D., and Eickbush, )18.2(T)111(.H. 1999. )17.2(The)]TJ 0.6333 -1.2 TD 0.0602 Tc 0.1389 Tw [(age and evolution of non-L)76(TR retrotransposable)]TJ T* 0 Tc 0.0004 Tw (elements. Mol. Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 418.41 532.275 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 420.87 532.275 Tm 0.0004 Tw (16: 793-805.)Tj -12.1333 -1.2 TD -0.0121 Tc -0.0695 Tw [(Malik, H.S., and Eickbush, )10.5(T)112.2(.H. 2001. Phylogenetic analysis)]TJ 0.6333 -1.2 TD -0.0127 Tc (of ribonuclease H domains suggests a late, chimeric origin)Tj T* 0.0165 Tc 0.1389 Tw [(of L)72.9(TR retrotransposable elements and retroviruses.)]TJ T* 0 Tc -0.0011 Tw (Genome Res.)Tj /F3 1 Tf 9 0 1.913 9 372.87 489.075 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 375.33 489.075 Tm (1)Tj 0.4867 0 TD -0.0011 Tw [(1: 1)73.6(187-1)77(197.)]TJ -7.56 -1.2 TD 0.0485 Tc 0.1389 Tw [(Mamoun, C.B., Gluzman, I.Y)130.6(., Beverley)69.4(, S.M., and)]TJ 0.6333 -1.2 TD 0.0054 Tc [(Goldberg, D.E. 2000. )21.9(T)36.4(ransposition of the )]TJ /F3 1 Tf 9 0 1.913 9 494.97 467.475 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 317.37 456.675 Tm 0.0266 Tc (element )Tj /F3 1 Tf 9 0 1.913 9 355.05 456.675 Tm (mariner)Tj /F1 1 Tf 9 0 0 9 387.21 456.675 Tm 0 Tc ( )Tj 0.4435 0 TD 0.0266 Tc 0.1389 Tw (within the human malaria parasite)Tj /F3 1 Tf 9 0 1.913 9 317.37 445.875 Tm 0 Tc 0.1373 Tw (Plasmodium falciparum)Tj /F1 1 Tf 9 0 0 9 412.11 445.875 Tm (. Mol. Biochem. Parasitol.)Tj /F3 1 Tf 9 0 1.913 9 517.83 445.875 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 521.61 445.875 Tm (1)Tj 0.48 0 TD (10:)Tj -23.1733 -1.2 TD 0.0003 Tc (405-407.)Tj -0.6333 -1.2 TD 0.0506 Tc 0.1389 Tw (Maruyama, K., and Hartl, D.L. 1991. Evidence for)Tj 0.6333 -1.2 TD -0.0044 Tc -0.0695 Tw (interspecific transfer of the transposable element )Tj /F3 1 Tf 9 0 1.913 9 508.17 413.475 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 317.37 402.675 Tm -0.0061 Tc (between )Tj /F3 1 Tf 9 0 1.913 9 352.77 402.675 Tm (Drosophila)Tj /F1 1 Tf 9 0 0 9 395.31 402.675 Tm 0 Tc ( )Tj 0.2024 0 TD -0.0061 Tc (and )Tj /F3 1 Tf 9 0 1.913 9 413.79 402.675 Tm (Zaprionus)Tj /F1 1 Tf 9 0 0 9 453.27 402.675 Tm -0.0695 Tw (. J. Mol. Evol. 33: 514-)Tj -15.1 -1.2 TD -0.0006 Tc 0 Tw (524.)Tj -0.6333 -1.2 TD 0.0298 Tc 0.1389 Tw [(Maside, X., Bartolome, C., )49.9(Assimacopoulos, S., and)]TJ 0.6333 -1.2 TD 0.062 Tc (Charlesworth, B. 2001. Rates of movement and)Tj T* 0.0294 Tc (distribution of transposable elements in )Tj /F3 1 Tf 9 0 1.913 9 493.05 359.475 Tm 0 Tw (Drosophila)Tj -19.2649 -1.2 TD 0 Tc (melanogaster)Tj /F1 1 Tf 9 0 0 9 371.91 348.675 Tm (: )Tj /F3 1 Tf 9 0 1.913 9 377.37 348.675 Tm 0.0551 Tw (in situ)Tj /F1 1 Tf 9 0 0 9 401.37 348.675 Tm 0 Tw ( )Tj 0.3331 0 TD (hybridization )Tj /F3 1 Tf 9 0 1.913 9 457.89 348.675 Tm (vs)Tj /F1 1 Tf 9 0 0 9 466.89 348.675 Tm ( )Tj 0.3331 0 TD 0.0551 Tw (Southern blotting)Tj -16.9465 -1.2 TD 0.0005 Tw (data. Genet. Res. 78: 121-136.)Tj -0.6333 -1.2 TD 0.1189 Tw [(Mason, P)122.9(., Strem, M., and Fedorof)18.3(f, N. 1991. )22(The )]TJ /F3 1 Tf 9 0 1.913 9 519.9301 327.075 Tm 0 Tw (tnpA)Tj /F1 1 Tf 9 0 0 9 317.37 316.275 Tm -0.0019 Tc (and )Tj /F3 1 Tf 9 0 1.913 9 334.17 316.275 Tm (tnpD)Tj /F1 1 Tf 9 0 0 9 353.07 316.275 Tm 0 Tc ( )Tj 0.2066 0 TD -0.0019 Tc -0.0695 Tw (gene products of the )Tj /F3 1 Tf 9 0 1.913 9 436.65 316.275 Tm 0 Tw (Spm)Tj /F1 1 Tf 9 0 0 9 455.07 316.275 Tm 0 Tc ( )Tj 0.2066 0 TD -0.0019 Tc -0.0695 Tw (element are required)Tj -15.5066 -1.2 TD 0 Tc 0.0007 Tw (for transposition in tobacco. Plant Cell)Tj /F3 1 Tf 9 0 1.913 9 468.45 305.475 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 470.97 305.475 Tm 0.0007 Tw (3: 73-85.)Tj -17.7 -1.2 TD 0.1275 Tw [(Matzke, M., Mette, M., )53.3(Aufsatz, W)55.8(., Jakowitsch, J., and)]TJ 0.6333 -1.2 TD 0.0313 Tw [(Matzke, )57(A. 1999. Host defenses to parasitic sequences)]TJ T* 0.0144 Tc 0.1389 Tw (and the evolution of epigenetic control mechanisms.)Tj T* 0 Tc 0 Tw (Genetica)Tj /F3 1 Tf 9 0 1.913 9 353.37 262.275 Tm ( )Tj /F1 1 Tf 9 0 0 9 355.83 262.275 Tm -0.0011 Tw (107: 271-287.)Tj -4.9067 -1.2 TD 0.0161 Tc 0.1389 Tw [(McDonald, J.F)108(. 1993. Evolution and consequences of)]TJ 0.6333 -1.2 TD 0 Tc 0.0262 Tw [(transposable elements. Curr)52.5(. Opin. Genet. Dev)80.7(. 3: 855-)]TJ T* -0.0006 Tc 0 Tw (864.)Tj -0.6333 -1.2 TD 0.0348 Tc 0.1389 Tw (McFadden, J., and Knowles, G. 1997. Escape from)Tj 0.6333 -1.2 TD 0 Tc 0.0411 Tw (evolutionary stasis by transposon-mediated deleterious)Tj T* 0.0004 Tw [(mutations. J. )21.2(Theor)52(. Biol. 186: 441-447.)]TJ -0.6333 -1.2 TD -0.0029 Tw [(Mejlumian, L., Plisson, )55(A., Bucheton, )49.5(A., and )21.2(T)111(erzian, C.)]TJ 0.6333 -1.2 TD -0.0006 Tw (2002. Comparative and functional studies of )Tj /F3 1 Tf 9 0 1.913 9 495.45 175.875 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 317.37 165.075 Tm 0.1385 Tw (species invasion by the )Tj /F3 1 Tf 9 0 1.913 9 417.87 165.075 Tm 0 Tw (gypsy)Tj /F1 1 Tf 9 0 0 9 441.39 165.075 Tm ( )Tj 0.4165 0 TD 0.1385 Tw (endogenous retrovirus.)Tj -14.1965 -1.2 TD 0 Tw (Genetics)Tj /F3 1 Tf 9 0 1.913 9 352.83 154.275 Tm ( )Tj /F1 1 Tf 9 0 0 9 355.35 154.275 Tm -0.0008 Tw (160: 201-209.)Tj -4.8533 -1.2 TD 0.0109 Tc 0.1389 Tw [(Mendiola, M.V)85.4(., Bernales, I., and de la Cruz, F)115.6(.)0( 1994.)]TJ 0.6333 -1.2 TD 0.0336 Tc [(Dif)22.8(ferential roles of the transposon termini in )]TJ /F3 1 Tf 9 0 1.913 9 519.03 132.675 Tm 0 Tw (IS91)Tj /F1 1 Tf 9 0 0 9 317.37 121.875 Tm -0.0033 Tc -0.0694 Tw [(transposition. Proc. Natl. )54.4(Acad. Sci. USA. 91: 1922-1926.)]TJ -0.6333 -1.2 TD 0 Tc 0.0182 Tw [(Miller)54.7(, D.W)51.6(., and Miller)57.9(, L.K. 1982. )54.4(A)53.7( virus mutant with an)]TJ 0.6333 -1.2 TD -0.0137 Tc -0.0695 Tw (insertion of a )Tj /F3 1 Tf 9 0 1.913 9 367.65 100.275 Tm 0 Tw (copia)Tj /F1 1 Tf 9 0 0 9 388.53 100.275 Tm -0.0695 Tw (-like transposable element. Nature)Tj /F3 1 Tf 9 0 1.913 9 519.5701 100.275 Tm 0 Tc 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 521.3101 100.275 Tm -0.0137 Tc (299:)Tj ET endstream endobj 43 0 obj << /Type /Page /Parent 95 0 R /Resources 44 0 R /Contents 45 0 R /Thumb 82 0 R /MediaBox [ 0 0 595 842 ] /CropBox [ 0 0 595 842 ] /Rotate 0 >> endobj 44 0 obj << /ProcSet [ /PDF /Text ] /Font << /F1 114 0 R /F2 113 0 R /F3 128 0 R >> /ExtGState << /GS1 139 0 R /GS2 138 0 R >> /ColorSpace << /Cs8 112 0 R >> >> endobj 45 0 obj << /Length 17289 >> stream BT /F1 1 Tf 9 0 0 9 55.93 779.115 Tm 0 0 0 1 k /GS2 gs -0.0001 Tc -0.0003 Tw (14 Silva )Tj /F3 1 Tf 9 0 1.913 9 95.47 779.115 Tm 0 Tc (et al)Tj /F1 1 Tf 9 0 0 9 112.45 779.115 Tm 0 Tw (.)Tj /F2 1 Tf 37.9351 29.4096 -29.4096 37.9351 34.496 531.1378 Tm 2 Tr 0 0 0 0.45 K 0 J 0 j 0.576 w 10 M []0 d /Cs8 cs 0 scn /GS1 gs 0.0686 Tc -0.0001 Tw (PAGE PROOFS)Tj /F1 1 Tf 9 0 0 9 62.23 748.275 Tm 0 Tr 0 0 0 1 k /GS2 gs 0.0003 Tc 0 Tw (562-564.)Tj -0.6333 -1.2 TD 0.0446 Tc 0.1389 Tw (Misra, S., and Rio, D.C. 1990. Cytotype control of)Tj /F3 1 Tf 9 0 1.913 9 62.23 726.675 Tm 0 Tc 0.0066 Tw (Drosophila P)Tj /F1 1 Tf 9 0 0 9 113.83 726.675 Tm 0 Tw ( )Tj 0.2846 0 TD 0.0066 Tw (element transposition: the 66 kd protein is)Tj -6.018 -1.2 TD 0.0001 Tw [(a repressor of transposase activity)75.3(. Cell)]TJ /F3 1 Tf 9 0 1.913 9 219.13 715.875 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 221.59 715.875 Tm 0.0001 Tw (62: 269-284.)Tj -18.34 -1.2 TD 0.0461 Tc 0.1389 Tw [(Mizrokhi, L.J., and Mazo, )46.7(A.M. 1990. Evidence for)]TJ 0.6333 -1.2 TD 0.0109 Tc (horizontal transmission of the mobile element )Tj /F3 1 Tf 9 0 1.913 9 257.29 694.275 Tm 0 Tw (jockey)Tj /F1 1 Tf 9 0 0 9 62.23 683.475 Tm 0 Tc 0.1386 Tw (between distant )Tj /F3 1 Tf 9 0 1.913 9 130.27 683.475 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 173.23 683.475 Tm ( )Tj 0.4166 0 TD 0.1386 Tw [(species. Proc. Natl. )51.1(Acad.)]TJ -12.7499 -1.2 TD -0.0004 Tw (Sci. USA.)Tj /F3 1 Tf 9 0 1.913 9 100.69 672.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 103.21 672.675 Tm -0.0004 Tw (87: 9216-9220.)Tj -5.1867 -2.4 TD -0.0051 Tc -0.0694 Tw [(Nekrutenko, )52(A., and Li, W)57.3(.H. 2001. )19.7(T)32.6(ransposable elements)]TJ 0.6333 -1.2 TD 0.0021 Tc 0.139 Tw [(are found in a large number of human protein-coding)]TJ T* 0 Tc -0.02 Tw [(genes. T)37.7(rends )-20(Genet.)]TJ /F3 1 Tf 9 0 1.913 9 148.75 629.475 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 151.27 629.475 Tm (17: 619-621.)Tj -10.5267 -1.2 TD 0.0304 Tc 0.1389 Tw [(Nouaud, D., and )49.9(Anxolabhre, D. 1997. )]TJ /F3 1 Tf 9 0 1.913 9 239.35 618.675 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 245.59 618.675 Tm ( )Tj 0.4473 0 TD 0.0304 Tc (element)Tj -20.8206 -1.2 TD 0.0089 Tc 0.1389 Tw (domestication: a stationary truncated )Tj /F3 1 Tf 9 0 1.913 9 219.79 607.875 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 225.91 607.875 Tm ( )Tj 0.4258 0 TD 0.0089 Tc 0.1389 Tw (element may)Tj -18.6125 -1.2 TD -0.0003 Tc -0.0694 Tw (encode a 66-kDa repressor-like protein in the )Tj /F3 1 Tf 9 0 1.913 9 240.31 597.075 Tm 0 Tw (Drosophila)Tj -19.5316 -1.2 TD 0.0009 Tc (montium)Tj /F1 1 Tf 9 0 0 9 96.73 586.275 Tm 0 Tc ( )Tj 0.4178 0 TD 0.0009 Tc 0.1389 Tw (species subgroup. Mol. Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 240.85 586.275 Tm 0 Tc 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 244.63 586.275 Tm 0.0009 Tc 0.1389 Tw [(14: 1)74(132-)]TJ -20.2667 -1.2 TD 0 Tc 0 Tw (1)Tj 0.48 0 TD -0.0002 Tc (144.)Tj -1.1133 -1.2 TD 0 Tc 0.1101 Tw [(Nuzhdin, S.V)86.1(. 1999. Sure facts, speculations, and open)]TJ 0.6333 -1.2 TD 0.101 Tw (questions about the evolution of transposable element)Tj T* -0.0001 Tw [(copy number)53.3(. Genetica)]TJ /F3 1 Tf 9 0 1.913 9 154.75 543.075 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 157.27 543.075 Tm -0.0001 Tw (107: 129-137.)Tj -11.1933 -1.2 TD 0.0024 Tc 0.1389 Tw [(OBrochta, D.A., and Handler)53.8(, )53(A.M. 1988. Mobility of )]TJ /F3 1 Tf 9 0 1.913 9 277.33 532.275 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 62.23 521.475 Tm -0.0035 Tc -0.0695 Tw (elements in drosophilids and nondrosophilids. Proc. Natl.)Tj T* 0 Tc 0.0007 Tw (Acad. Sci. USA. 85: 6052-6056.)Tj -0.6333 -1.2 TD -0.0025 Tw [(Oosumi, )11.9(T)111(., Belknap, W)62.1(.R., and Garlick, B. 1995. )]TJ /F3 1 Tf 9 0 1.913 9 252.79 499.875 Tm 0 Tw (Mariner)Tj /F1 1 Tf 9 0 0 9 62.23 489.075 Tm -0.0001 Tw (transposons in humans. Nature)Tj /F3 1 Tf 9 0 1.913 9 187.27 489.075 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 189.73 489.075 Tm -0.0001 Tw (378: 672.)Tj -14.8 -1.2 TD -0.0042 Tc -0.0694 Tw [(Orgel, L.E., and Crick, F)110.3(.H. 1980. Selfish DNA: the ultimate)]TJ 0.6333 -1.2 TD 0 Tc 0.0007 Tw (parasite. Nature)Tj /F3 1 Tf 9 0 1.913 9 126.25 467.475 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 128.77 467.475 Tm 0.0007 Tw (284: 604-607.)Tj -8.0267 -1.2 TD 0.0735 Tw [(Pasyukova, E.G., Nuzhdin, S.V)94(., and Filatov)72.1(, D.A. 1998.)]TJ 0.6333 -1.2 TD 0.1026 Tw (The relationship between the rate of transposition and)Tj T* 0.1187 Tw (transposable element copy number for )Tj /F3 1 Tf 9 0 1.913 9 223.63 435.075 Tm 0 Tw (copia)Tj /F1 1 Tf 9 0 0 9 245.17 435.075 Tm ( )Tj 0.3967 0 TD (and )Tj /F3 1 Tf 9 0 1.913 9 267.31 435.075 Tm (Doc)Tj /F1 1 Tf 9 0 0 9 62.23 424.275 Tm 0.0089 Tc 0.1389 Tw (retrotransposons of )Tj /F3 1 Tf 9 0 1.913 9 146.35 424.275 Tm (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 249.43 424.275 Tm (. Genet.)Tj -20.8 -1.2 TD 0 Tc -0.0009 Tw [(Res. 72: 1-1)78.7(1.)]TJ -0.6333 -1.2 TD -0.0051 Tc -0.0694 Tw [(Plisson, )48.3(A., )18(T)112.6(eysset, L., Chalvet, F)113.4(., Kim, )52.7(A., Prudhomme,)]TJ 0.6333 -1.2 TD 0 Tc 0.0076 Tw [(N., )23.6(T)111(erzian, C., and Bucheton, )52.5(A. 1997. )58.2(About the origin)]TJ T* -0.0104 Tc -0.0694 Tw (of retroviruses and the co-evolution of the )Tj /F3 1 Tf 9 0 1.913 9 221.83 381.075 Tm 0 Tw (gypsy)Tj /F1 1 Tf 9 0 0 9 244.87 381.075 Tm 0 Tc ( )Tj 0.1982 0 TD -0.0104 Tc (retrovirus)Tj -20.4915 -1.2 TD 0 Tc 0.0392 Tw (with the Drosophila)Tj /F3 1 Tf 9 0 1.913 9 139.45 370.275 Tm 0 Tw ( )Tj 0.3172 0 TD (flamenco)Tj /F1 1 Tf 9 0 0 9 178.81 370.275 Tm ( )Tj 0.3172 0 TD 0.0392 Tw (host gene. Genetica)Tj /F3 1 Tf 9 0 1.913 9 262.93 370.275 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 265.81 370.275 Tm (100:)Tj -22.62 -1.2 TD -0.0004 Tc (29-37.)Tj -0.6333 -1.2 TD 0 Tc 0.1133 Tw [(Pinsker)54(, W)53.3(., Haring, E., Hagemann, S., and Miller)51.3(, W)60(.J.)]TJ 0.6333 -1.2 TD -0.0026 Tc -0.0695 Tw [(2001. )21.4(The evolutionary life history of )]TJ /F3 1 Tf 9 0 1.913 9 204.43 337.875 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 210.43 337.875 Tm ( )Tj 0.2059 0 TD -0.0026 Tc -0.0695 Tw (transposons: from)Tj -16.6726 -1.2 TD 0.0695 Tc 0.2077 Tw (horizontal invaders to domesticated neogenes.)Tj T* 0 Tc 0 Tw (Chromosoma)Tj /F3 1 Tf 9 0 1.913 9 116.23 316.275 Tm ( )Tj /F1 1 Tf 9 0 0 9 118.69 316.275 Tm (1)Tj 0.4867 0 TD (10: 148-158.)Tj -7.3933 -1.2 TD 0.0707 Tw [(Plasterk, R.H. 1996. )11.4(The )]TJ /F3 1 Tf 9 0 1.913 9 160.03 305.475 Tm -0.0977 Tc 0 Tw [(Tc)-97.7(1)]TJ /F1 1 Tf 9 0 0 9 174.13 305.475 Tm 0 Tc (/)Tj /F3 1 Tf 9 0 1.913 9 176.65 305.475 Tm (mariner)Tj /F1 1 Tf 9 0 0 9 207.13 305.475 Tm ( )Tj 0.3487 0 TD 0.0707 Tw [(transposon family)68.7(.)]TJ -16.4487 -1.2 TD -0.0004 Tw [(Curr )21.6(T)111(op Microbiol. Immunol.)]TJ /F3 1 Tf 9 0 1.913 9 177.07 294.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 179.53 294.675 Tm -0.0004 Tw (204: 125-143.)Tj -13.6667 -1.2 TD 0.0116 Tw [(Prudhomme, N., Gans, M., Masson, M., )20(T)111(erzian, C., and)]TJ 0.6333 -1.2 TD 0.0071 Tc 0.0811 Tw [(Bucheton, A. )-57.8(1995. )]TJ /F3 1 Tf 9 0 1.913 9 145.75 273.075 Tm 0 Tw (Flamenco)Tj /F1 1 Tf 9 0 0 9 185.77 273.075 Tm 0.1389 Tw (, a gene controlling the)Tj /F3 1 Tf 9 0 1.913 9 62.23 262.275 Tm 0 Tc 0 Tw (gypsy)Tj /F1 1 Tf 9 0 0 9 85.69 262.275 Tm ( )Tj 0.3781 0 TD 0.1001 Tw (retrovirus of )Tj /F3 1 Tf 9 0 1.913 9 140.95 262.275 Tm (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 241.93 262.275 Tm (. Genetics)Tj -19.9667 -1.2 TD 0.0021 Tw [(139: 697-71)72.5(1.)]TJ -0.6333 -1.2 TD -0.0222 Tw [(Quesneville, H., and )48.8(Anxolabhre, D. 1997. )57.5(A)53.7( simulation)]TJ 0.6333 -1.2 TD 0 Tw (of )Tj /F3 1 Tf 9 0 1.913 9 72.19 229.875 Tm (P)Tj /F1 1 Tf 9 0 0 9 78.19 229.875 Tm -0.0002 Tw ( element horizontal transfer in )Tj /F3 1 Tf 9 0 1.913 9 199.27 229.875 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 242.29 229.875 Tm -0.0002 Tw (. Genetica)Tj -20.0067 -1.2 TD 0.0004 Tw (100: 295-307.)Tj -0.6333 -1.2 TD 0.0074 Tc 0.1389 Tw (Rasmusson, K.E., Raymond, J.D., and Simmons, M.J.)Tj 0.6333 -1.2 TD 0.0077 Tc (1993. Repression of hybrid dysgenesis in )Tj /F3 1 Tf 9 0 1.913 9 239.65 197.475 Tm 0 Tw (Drosophila)Tj -19.4583 -1.2 TD 0.0561 Tc (melanogaster)Tj /F1 1 Tf 9 0 0 9 122.83 186.675 Tm 0 Tc ( )Tj 0.4731 0 TD 0.0561 Tc 0.139 Tw (by individual naturally occurring )Tj /F3 1 Tf 9 0 1.913 9 277.33 186.675 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 62.23 175.875 Tm 0.0006 Tw (elements. Genetics)Tj /F3 1 Tf 9 0 1.913 9 139.27 175.875 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 141.73 175.875 Tm 0.0006 Tw (133: 605-622.)Tj -9.4667 -1.2 TD 0.0623 Tc 0.1389 Tw [(Remsen, J., and OGrady)71.2(, P)128.8(.)0( 2002. Phylogeny of)]TJ /F3 1 Tf 9 0 1.913 9 62.23 154.275 Tm 0 Tc 0 Tw (Drosophilinae)Tj /F1 1 Tf 9 0 0 9 117.25 154.275 Tm ( )Tj 0.3755 0 TD 0.0975 Tw (\(Diptera: Drosophilidae\), with comments)Tj -6.4888 -1.2 TD 0.0354 Tc 0.139 Tw (on combined analysis and character support. Mol.)Tj T* 0 Tc 0.0005 Tw (Phylogenet. Evol.)Tj /F3 1 Tf 9 0 1.913 9 132.25 132.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 134.77 132.675 Tm 0.0005 Tw (24: 249-264.)Tj -8.6933 -1.2 TD 0.0025 Tc 0.1389 Tw (Robertson, H.M. 1997. Multiple )Tj /F3 1 Tf 9 0 1.913 9 188.83 121.875 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 219.43 121.875 Tm 0 Tc ( )Tj 0.4194 0 TD 0.0025 Tc 0.1389 Tw (transposons in)Tj -17.886 -1.2 TD 0 Tc 0.037 Tw (flatworms and hydras are related to those of insects. J.)Tj T* -0.0003 Tw (Hered. 88: 195-201.)Tj 27.0933 72 TD -0.0061 Tc -0.0694 Tw (Robertson, H.M., and Lampe, D.J. 1995. Recent horizontal)Tj 0.6267 -1.2 TD 0 Tc 0.0632 Tw (transfer of a )Tj /F3 1 Tf 9 0 1.913 9 363.97 737.475 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 394.45 737.475 Tm ( )Tj 0.3412 0 TD 0.0632 Tw (transposable element among and)Tj -9.5345 -1.2 TD -0.009 Tc -0.0694 Tw (between Diptera and Neuroptera. Mol. Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 499.15 726.675 Tm 0 Tc 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 500.95 726.675 Tm -0.009 Tc -0.0694 Tw (12: 850-)Tj -21.0267 -1.2 TD -0.0006 Tc 0 Tw (862.)Tj -0.6267 -1.2 TD 0.0009 Tc 0.1389 Tw (Robertson, H.M., and MacLeod, E.G. 1993. Five major)Tj 0.6267 -1.2 TD 0 Tc -0.0463 Tw (subfamilies of )Tj /F3 1 Tf 9 0 1.913 9 368.41 694.275 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 398.89 694.275 Tm ( )Tj 0.2317 0 TD -0.0463 Tw (transposable elements in insects,)Tj -9.9183 -1.2 TD 0.0548 Tc 0.1389 Tw [(including the Mediterranean fruit fly)70.9(, and related)]TJ T* 0 Tc 0.0003 Tw (arthropods. Insect. Mol. Biol.)Tj /F3 1 Tf 9 0 1.913 9 425.77 672.675 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 428.23 672.675 Tm 0.0003 Tw (2: 125-139.)Tj -13.5733 -1.2 TD 0.0342 Tc 0.1389 Tw [(Robertson, H.M., Soto-Adames, F)110(.N., W)41.9(a)0(lden, K.K.,)]TJ 0.6267 -1.2 TD 0 Tc 0 Tw (A)Tj 0.6667 0 TD 0.0175 Tc 0.139 Tw [(vancini, R.M., and Lampe, D.J. 2002. )19.3(The )]TJ /F3 1 Tf 9 0 1.913 9 501.37 651.075 Tm 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 311.71 640.275 Tm 0 Tc 0.0367 Tw (transposons of animals: horizontally jumping genes. In:)Tj 0 -1.2 TD 0.0949 Tw [(Horizontal Gene )12.9(T)44.3(ransfer)52(. M. Syvanen, and C.I. Kado,)]TJ T* -0.0005 Tw [(eds. )54.2(Academic Press, San Diego, CA.)]TJ -0.6267 -1.2 TD 0.1224 Tw [(Roche, S.E., Schif)19.9(f, M., and Rio, D.C. 1995. )]TJ /F3 1 Tf 9 0 1.913 9 491.83 607.875 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 497.83 607.875 Tm (-element)Tj -20.68 -1.2 TD 0.0695 Tc 0.8243 Tw (repressor autoregulation involves germ-line)Tj T* 0.002 Tc 0.1389 Tw (transcriptional repression and reduction of third intron)Tj T* 0 Tc -0.0005 Tw [(splicing. Genes Dev)77.7(. 9: 1278-1288.)]TJ -0.6267 -1.2 TD 0.0039 Tw [(Ronsseray)72.4(, S., Boivin, )54.4(A., and )54.8(Anxolabhre, D. 2001. )]TJ /F3 1 Tf 9 0 1.913 9 523.81 564.675 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 529.81 564.675 Tm (-)Tj -24.2333 -1.2 TD 0.0313 Tc 0.1389 Tw (Element repression in )Tj /F3 1 Tf 9 0 1.913 9 411.19 553.875 Tm (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 519.01 553.875 Tm 0 Tc 0 Tw ( )Tj 0.4482 0 TD 0.0313 Tc (by)Tj -23.4816 -1.2 TD -0.0067 Tc -0.0695 Tw (variegating clusters of P-lacZ-white transgenes. Genetics)Tj T* 0 Tc -0.0008 Tw (159: 1631-1642.)Tj -0.6267 -1.2 TD -0.0512 Tw [(Ronsseray)72.4(, S., Lehmann, M., and )59.2(Anxolabhre, D. 1989.)]TJ 0.6267 -1.2 TD -0.0682 Tw (Copy number and distribution of )Tj /F3 1 Tf 9 0 1.913 9 438.73 510.675 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 444.73 510.675 Tm ( )Tj 0.2098 0 TD (and )Tj /F3 1 Tf 9 0 1.913 9 463.51 510.675 Tm (I)Tj /F1 1 Tf 9 0 0 9 466.03 510.675 Tm ( )Tj 0.2098 0 TD -0.0682 Tw (mobile elements)Tj -17.3564 -1.2 TD 0 Tw (in )Tj /F3 1 Tf 9 0 1.913 9 322.21 499.875 Tm 0.1107 Tw (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 423.25 499.875 Tm 0 Tw ( )Tj 0.3887 0 TD 0.1107 Tw (populations. Chromosoma)Tj -12.782 -1.2 TD 0.0009 Tw (98: 207-214.)Tj -0.6267 -1.2 TD -0.0047 Tc -0.0695 Tw [(Ronsseray)76.6(, S., Marin, L., Lehmann, M., and )54.3(Anxolabhre,)]TJ 0.6267 -1.2 TD 0 Tc -0.0083 Tw (D. 1998. Repression of hybrid dysgenesis in )Tj /F3 1 Tf 9 0 1.913 9 489.79 467.475 Tm 0 Tw (Drosophila)Tj -19.5316 -1.2 TD 0.0006 Tc (melanogaster)Tj /F1 1 Tf 9 0 0 9 366.31 456.675 Tm 0 Tc ( )Tj 0.4175 0 TD 0.0006 Tc 0.1389 Tw [(by combinations of telomeric )]TJ /F3 1 Tf 9 0 1.913 9 491.77 456.675 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 497.77 456.675 Tm 0.0006 Tc (-element)Tj -20.6733 -1.2 TD 0 Tc 0.0647 Tw (reporters and naturally occurring )Tj /F3 1 Tf 9 0 1.913 9 446.11 445.875 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 452.11 445.875 Tm ( )Tj 0.3427 0 TD 0.0647 Tw (elements. Genetics)Tj -15.9427 -1.2 TD -0.0008 Tw (149: 1857-1866.)Tj -0.6267 -1.2 TD -0.0136 Tc -0.0695 Tw [(Rubin, E.J., )57.9(Akerley)72.1(, B.J., Novik, V)92(.N., Lampe, D.J., Husson,)]TJ 0.6267 -1.2 TD 0 Tc 0.0137 Tw (R.N., and Mekalanos, J.J. 1999. )Tj /F3 1 Tf 9 0 1.913 9 442.93 413.475 Tm (In vivo)Tj /F1 1 Tf 9 0 0 9 469.03 413.475 Tm 0 Tw ( )Tj 0.2917 0 TD 0.0137 Tw (transposition of)Tj /F3 1 Tf 9 0 1.913 9 311.71 402.675 Tm 0.0661 Tc 0 Tw (mariner)Tj /F1 1 Tf 9 0 0 9 346.33 402.675 Tm 0.1389 Tw (-based elements in enteric bacteria and)Tj -3.8467 -1.2 TD -0.0084 Tc -0.0694 Tw [(mycobacteria. Proc. Natl. )55.6(Acad. Sci. USA. 96: 1645-1650.)]TJ -0.6267 -1.2 TD 0.0695 Tc 0.1761 Tw [(Rubin, G.M., and Spradling, )59.8(A.C. 1982. Genetic)]TJ 0.6267 -1.2 TD 0 Tc 0.0219 Tw (transformation of )Tj /F3 1 Tf 9 0 1.913 9 382.15 370.275 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 425.17 370.275 Tm ( )Tj 0.2999 0 TD 0.0219 Tw (with transposable element)Tj -12.9066 -1.2 TD 0.0009 Tw (vectors. Science)Tj /F3 1 Tf 9 0 1.913 9 377.77 359.475 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 380.23 359.475 Tm 0.0009 Tw (218: 348-353.)Tj -8.24 -1.2 TD 0.0345 Tw [(Shao, H. and )18.6(T)37.7(u, Z. 2001. Expanding the diversity of the)]TJ 0.6267 -1.2 TD 0.0049 Tc 0.1389 Tw [(IS630-T)111.4(c1-mariner superfamily: discovery of a unique)]TJ T* -0.0087 Tc -0.0695 Tw (DD37E transposon and reclassification of the DD37D and)Tj T* 0 Tc -0.0005 Tw [(DD39D transposons. Genetics 159: 1)74.5(103-1)70.4(1)76(15.)]TJ -0.6267 -1.2 TD -0.0293 Tw [(Sherman, )57.1(A., Dawson, )57.2(A., Mather)57.1(, C., Gilhooley)68.2(, H., Li, )18.2(Y)127(.)0(,)]TJ 0.6267 -1.2 TD 0.0695 Tc 0.1487 Tw (Mitchell, R., Finnegan, D., and Sang, H. 1998.)Tj T* 0 Tc 0 Tw (T)Tj 0.5733 0 TD -0.0426 Tw (ransposition of the )Tj /F3 1 Tf 9 0 1.913 9 391.75 283.875 Tm 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 434.83 283.875 Tm ( )Tj 0.2354 0 TD (element )Tj /F3 1 Tf 9 0 1.913 9 471.07 283.875 Tm (mariner)Tj /F1 1 Tf 9 0 0 9 501.55 283.875 Tm ( )Tj 0.2354 0 TD -0.0426 Tw (into the)Tj -21.3287 -1.2 TD -0.0004 Tw (chicken germ line. Nat. Biotechnol.)Tj /F3 1 Tf 9 0 1.913 9 450.25 273.075 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 452.77 273.075 Tm -0.0004 Tw (16: 1050-1053.)Tj -16.3 -1.2 TD 0.0017 Tw [(Siebel, C.W)54.1(., Kanaar)53.1(, R., and Rio, D.C. 1994. Regulation)]TJ 0.6267 -1.2 TD -0.0329 Tw (of tissue-specific)Tj /F3 1 Tf 9 0 1.913 9 377.95 251.475 Tm -0.0329 Tc 0 Tw ( P)Tj /F1 1 Tf 9 0 0 9 386.17 251.475 Tm 0 Tc -0.0329 Tw (-element pre-mRNA splicing requires)Tj -8.2733 -1.2 TD -0.0296 Tw [(the RNA-binding protein PSI. Genes Dev)71(. 8: 1713-1725.)]TJ -0.6267 -1.2 TD 0.0029 Tc 0.1389 Tw (Silva, J.C. 2000 Population genetics of )Tj /F3 1 Tf 9 0 1.913 9 471.19 229.875 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 477.19 229.875 Tm ( )Tj 0.4198 0 TD 0.0029 Tc (transposable)Tj -18.8064 -1.2 TD 0.026 Tc 0.1389 Tw (elements and their host species, with emphasis on)Tj /F3 1 Tf 9 0 1.913 9 311.71 208.275 Tm -0.0007 Tc -0.0695 Tw (Drosophila willistoni)Tj /F1 1 Tf 9 0 0 9 389.95 208.275 Tm 0 Tc 0 Tw ( )Tj 0.2078 0 TD -0.0007 Tc (and )Tj /F3 1 Tf 9 0 1.913 9 408.73 208.275 Tm -0.0695 Tw (Drosophila sturtevanti)Tj /F1 1 Tf 9 0 0 9 494.95 208.275 Tm (, Doctoral)Tj -20.36 -1.2 TD 0 Tc -0.0003 Tw [(Thesis, )20.7(The University of )52.4(Arizona, )19.1(T)31(ucson.)]TJ -0.6267 -1.2 TD -0.0054 Tc -0.0694 Tw (Silva, J.C., and Kidwell, M.G. 2000. Horizontal transfer and)Tj 0.6267 -1.2 TD 0 Tc 0.006 Tw (selection in the evolution of )Tj /F3 1 Tf 9 0 1.913 9 423.07 175.875 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 429.07 175.875 Tm ( )Tj 0.284 0 TD 0.006 Tw (elements. Mol. Biol. Evol.)Tj -13.324 -1.2 TD -0.0002 Tw (17: 1542-1557.)Tj -0.6267 -2.4 TD 0.1374 Tw (Silva, J.C., Shabalina, S.A., Harris, D.G., Spouge, J.L.,)Tj 0.6267 -1.2 TD 0.0084 Tc 0.1389 Tw [(and Kondrashov)72(, )58.4(A.S. 2003. Conserved fragments of)]TJ T* -0.0055 Tc -0.0695 Tw (transposable elements in intergenic regions: evidence for)Tj T* 0.0676 Tc 0.1389 Tw (widespread recruitment of MIR- and L2-derived)Tj T* 0.0264 Tc 0.139 Tw [(sequences within the mouse and human genomes.)]TJ ET endstream endobj 46 0 obj << /Type /Page /Parent 95 0 R /Resources 47 0 R /Contents 48 0 R /Thumb 84 0 R /MediaBox [ 0 0 595 842 ] /CropBox [ 0 0 595 842 ] /Rotate 0 >> endobj 47 0 obj << /ProcSet [ /PDF /Text ] /Font << /F1 114 0 R /F2 113 0 R /F3 128 0 R >> /ExtGState << /GS1 139 0 R /GS2 138 0 R >> /ColorSpace << /Cs8 112 0 R >> >> endobj 48 0 obj << /Length 11916 >> stream BT /F1 1 Tf 9 0 0 9 337.47 780.5551 Tm 0 0 0 1 k /GS2 gs 0 Tc -0.0155 Tw [(Horizontal Transfer of Transposable Elements 15)]TJ /F2 1 Tf 37.9351 29.4096 -29.4096 37.9351 42.9423 542.4378 Tm 2 Tr 0 0 0 0.45 K 0 J 0 j 0.576 w 10 M []0 d /Cs8 cs 0 scn /GS1 gs 0.0686 Tc -0.0001 Tw (PAGE PROOFS)Tj /F1 1 Tf 9 0 0 9 67.83 748.275 Tm 0 Tr 0 0 0 1 k /GS2 gs 0 Tc 0.0007 Tw (Genet. Res. \(in press\).)Tj -0.6267 -2.4 TD 0.0572 Tc 0.1389 Tw (Simmons, G.M. 1992. Horizontal transfer of )Tj /F3 1 Tf 9 0 1.913 9 267.39 726.675 Tm 0 Tw (hobo)Tj /F1 1 Tf 9 0 0 9 67.83 715.875 Tm 0.0695 Tc 0.5068 Tw (transposable elements within the )Tj /F3 1 Tf 9 0 1.913 9 240.27 715.875 Tm 0 Tw (Drosophila)Tj -18.9049 -1.2 TD 0.0199 Tc (melanogaster)Tj /F1 1 Tf 9 0 0 9 124.53 705.0751 Tm 0 Tc ( )Tj 0.4369 0 TD 0.02 Tc 0.1389 Tw (species complex: evidence from DNA)Tj -6.7369 -1.2 TD 0 Tc -0.0001 Tw (sequencing. Mol. Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 178.41 694.275 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 180.87 694.275 Tm -0.0001 Tw (9: 1050-1060.)Tj -13.1867 -1.2 TD 0.0083 Tc 0.1389 Tw [(Simmons, M.J., and Bucholz, L.M. 1985. )19.3(T)32.6(ransposase)]TJ 0.6267 -1.2 TD 0 Tc -0.0558 Tw (titration in )Tj /F3 1 Tf 9 0 1.913 9 108.33 672.675 Tm (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 207.87 672.675 Tm (: a model of cytotype)Tj -15.56 -1.2 TD -0.0037 Tc -0.0695 Tw [(in the P-M system of hybrid dysgenesis. Proc. Natl. )52.4(Acad.)]TJ T* 0 Tc 0.0001 Tw [(Sci. USA. 82: 81)77.5(19-8123.)]TJ -0.6267 -1.2 TD 0.0005 Tc 0.1389 Tw [(Simmons, M.J., Haley)72.2(, K.J., and )23.9(Thompson, S.J. 2002.)]TJ 0.6267 -1.2 TD 0 Tc -0.0277 Tw (Maternal transmission of )Tj /F3 1 Tf 9 0 1.913 9 167.67 629.475 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 173.67 629.475 Tm ( )Tj 0.2503 0 TD -0.0277 Tw (element transposase activity)Tj -12.0103 -1.2 TD -0.0011 Tc 0 Tw (in )Tj /F3 1 Tf 9 0 1.913 9 76.71 618.675 Tm -0.0694 Tw (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 175.89 618.675 Tm 0 Tc 0 Tw ( )Tj 0.2075 0 TD -0.0011 Tc -0.0694 Tw (depends on the last )Tj /F3 1 Tf 9 0 1.913 9 256.17 618.675 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 262.17 618.675 Tm ( )Tj 0.2075 0 TD -0.0011 Tc (intron.)Tj -21.8008 -1.2 TD 0 Tc 0.0001 Tw [(Proc. Natl. )52.9(Acad. Sci. USA. 99: 9306-9309.)]TJ -0.6267 -1.2 TD -0.0019 Tc -0.0694 Tw (Simmons, M.J., Raymond, J.D., Grimes, C.D., Belinco, C.,)Tj 0.6267 -1.2 TD 0.0215 Tc 0.1389 Tw [(Haake, B.C., Jordan, M., Lund, C., Ojala, )14.2(T)112.5(.A., and)]TJ T* 0 Tc -0.0282 Tw [(Papermaster)50.7(, D. 1996. Repression of hybrid dysgenesis)]TJ T* 0.0215 Tc 0 Tw (in )Tj /F3 1 Tf 9 0 1.913 9 79.17 564.675 Tm 0.1389 Tw (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 184.95 564.675 Tm 0 Tc 0 Tw ( )Tj 0.4384 0 TD 0.0215 Tc 0.1389 Tw (by heat-shock-inducible)Tj -13.4518 -1.2 TD -0.0109 Tc -0.0694 Tw (sense and antisense )Tj /F3 1 Tf 9 0 1.913 9 149.07 553.875 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 155.01 553.875 Tm -0.0109 Tc -0.0694 Tw (-element constructs. Genetics)Tj /F3 1 Tf 9 0 1.913 9 269.97 553.875 Tm 0 Tc 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 271.77 553.875 Tm -0.0109 Tc (144:)Tj -22.66 -1.2 TD 0.0001 Tc (1529-1544.)Tj -0.6267 -1.2 TD 0 Tc -0.0605 Tw [(Simonelig, M., and )57.6(Anxolabhre, D. 1991. )49.3(A)53.7( )]TJ /F3 1 Tf 9 0 1.913 9 239.49 532.275 Tm 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 245.49 532.275 Tm ( )Tj 0.2175 0 TD -0.0605 Tw (element of)Tj /F3 1 Tf 9 0 1.913 9 67.83 521.475 Tm -0.0056 Tc -0.0694 Tw (Scaptomyza pallida)Tj /F1 1 Tf 9 0 0 9 144.33 521.475 Tm 0 Tc 0 Tw ( )Tj 0.203 0 TD -0.0056 Tc -0.0694 Tw (is active in )Tj /F3 1 Tf 9 0 1.913 9 188.19 521.475 Tm (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 286.47 521.475 Tm 0 Tc 0 Tw (.)Tj -24.2933 -1.2 TD 0.0001 Tw [(Proc. Natl. )52.9(Acad. Sci. USA. 88: 6102-6106.)]TJ -0.6267 -1.2 TD 0.0341 Tc 0.1389 Tw [(Smit, )52.8(A.F)118.3(., and Riggs, )52.7(A.D. 1996. )]TJ /F3 1 Tf 9 0 1.913 9 213.21 499.875 Tm 0 Tc 0 Tw (T)Tj 0.64 0 TD 0.0341 Tc (iggers)Tj /F1 1 Tf 9 0 0 9 245.31 499.875 Tm 0 Tc ( )Tj 0.451 0 TD 0.0341 Tc 0.1389 Tw (and DNA)Tj -20.171 -1.2 TD 0.0083 Tc (transposon fossils in the human genome. Proc. Natl.)Tj T* 0 Tc 0.0006 Tw (Acad. Sci. USA. 93: 1443-1448.)Tj -0.6267 -1.2 TD -0.0022 Tc -0.0695 Tw [(Spradling, )52.5(A.C., Stern, D., Beaton, )54(A., Rhem, E.J., Laverty)75.3(,)]TJ 0.6267 -1.2 TD 0 Tc 0 Tw (T)Tj 0.5 0 TD 0.1132 Tw [(., Mozden, N., Misra, S., and Rubin, G.M. 1999. )15.6(The)]TJ -0.5 -1.2 TD 0.0062 Tc 0.1389 Tw (Berkeley Drosophila Genome Project gene disruption)Tj T* -0.0042 Tc -0.0694 Tw (project: Single )Tj /F3 1 Tf 9 0 1.913 9 125.55 435.075 Tm 0 Tc 0 Tw (P)Tj /F1 1 Tf 9 0 0 9 131.49 435.075 Tm -0.0042 Tc -0.0694 Tw (-element insertions mutating 25% of vital)Tj /F3 1 Tf 9 0 1.913 9 67.83 424.275 Tm 0 Tc 0 Tw (Drosophila)Tj /F1 1 Tf 9 0 0 9 110.85 424.275 Tm -0.0003 Tw ( genes. Genetics)Tj /F3 1 Tf 9 0 1.913 9 178.41 424.275 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 180.87 424.275 Tm -0.0003 Tw (153: 135-177.)Tj -13.1867 -1.2 TD -0.0446 Tw [(Stuart-Rogers, C., and Flavell, )55.4(A.J. 2001. )18.5(The evolution of)]TJ /F3 1 Tf 9 0 1.913 9 67.83 402.675 Tm -0.0777 Tc 0 Tw [(Ty)-76.2(1)]TJ /F1 1 Tf 9 0 0 9 82.17 402.675 Tm 0 Tc (-)Tj /F3 1 Tf 9 0 1.913 9 85.17 402.675 Tm -0.0015 Tc (copia)Tj /F1 1 Tf 9 0 0 9 106.59 402.675 Tm 0 Tc ( )Tj 0.207 0 TD -0.0015 Tc -0.0695 Tw (group retrotransposons in gymnosperms. Mol.)Tj -4.5137 -1.2 TD 0 Tc -0.0003 Tw (Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 107.85 391.875 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 110.37 391.875 Tm -0.0003 Tw (18: 155-163.)Tj -5.3533 -1.2 TD 0 Tw (T)Tj 0.4867 0 TD -0.0115 Tc -0.0695 Tw [(erzian, C., Ferraz, C., Demaille, J., and Bucheton, )52.7(A. 2000.)]TJ 0.14 -1.2 TD 0.0127 Tc 0.1389 Tw (Evolution of the )Tj /F3 1 Tf 9 0 1.913 9 138.09 370.275 Tm 0 Tw (Gypsy)Tj /F1 1 Tf 9 0 0 9 164.13 370.275 Tm 0 Tc ( )Tj 0.4296 0 TD 0.0127 Tc 0.1389 Tw (endogenous retrovirus in the)Tj /F3 1 Tf 9 0 1.913 9 67.83 359.475 Tm 0 Tc 0.0275 Tw (Drosophila melanogaster)Tj /F1 1 Tf 9 0 0 9 168.15 359.475 Tm 0 Tw ( )Tj 0.3055 0 TD 0.0275 Tw (subgroup. Mol. Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 273.69 359.475 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 276.45 359.475 Tm (17:)Tj -23.18 -1.2 TD 0.0003 Tc (908-914.)Tj -0.6267 -1.2 TD 0 Tc (V)Tj 0.64 0 TD 0.0437 Tc 0.1389 Tw [(azquez-Manrique, R.P)127.9(., Hernandez, M., Martnez-)]TJ -0.0133 -1.2 TD 0 Tc 0.1386 Tw (Sebastin, M.J., and de Frutos, R. 2000. Evolution of)Tj /F3 1 Tf 9 0 1.913 9 67.83 316.275 Tm 0 Tw (gypsy)Tj /F1 1 Tf 9 0 0 9 91.35 316.275 Tm ( )Tj 0.296 0 TD 0.018 Tw (endogenous retrovirus in the )Tj /F3 1 Tf 9 0 1.913 9 211.23 316.275 Tm (Drosophila obscura)Tj /F1 1 Tf 9 0 0 9 317.37 748.275 Tm -0.0005 Tw (species group. Mol. Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 437.91 748.275 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 440.37 748.275 Tm -0.0005 Tw [(17: 1)70.2(185-1)77(193.)]TJ -14.3 -1.2 TD 0 Tw (V)Tj 0.6533 0 TD -0.0459 Tw [(ieira, C., and Bimont, C. 1997. )20.3(T)31(ransposition rate of the)]TJ /F3 1 Tf 9 0 1.913 9 317.37 726.675 Tm 0 Tw (412)Tj /F1 1 Tf 9 0 0 9 332.37 726.675 Tm ( )Tj 0.3976 0 TD 0.1196 Tw (retrotransposable element is independent of copy)Tj -2.0643 -1.2 TD 0.12 Tw (number in natural populations of )Tj /F3 1 Tf 9 0 1.913 9 453.87 715.875 Tm (Drosophila simulans)Tj /F1 1 Tf 9 0 0 9 535.9501 715.875 Tm 0 Tw (.)Tj -24.2867 -1.2 TD -0.0003 Tw (Mol. Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 376.83 705.0751 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 379.35 705.0751 Tm -0.0003 Tw (14: 185-188.)Tj -7.52 -1.2 TD 0 Tw (V)Tj 0.6533 0 TD -0.0086 Tw (ieira, C., Lepetit, D., Dumont, S., and Bimont, C. 1999.)Tj -0.02 -1.2 TD 0 Tw (W)Tj 0.9 0 TD -0.0035 Tw (ake up of transposable elements following )Tj /F3 1 Tf 9 0 1.913 9 495.45 683.475 Tm 0 Tw (Drosophila)Tj -19.5316 -1.2 TD 0.0128 Tc (simulans)Tj /F1 1 Tf 9 0 0 9 353.79 672.675 Tm 0 Tc ( )Tj 0.4297 0 TD 0.0128 Tc 0.1389 Tw (worldwide colonization. Mol. Biol. Evol.)Tj /F3 1 Tf 9 0 1.913 9 521.85 672.675 Tm 0 Tc 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 525.6901 672.675 Tm 0.0128 Tc (16:)Tj -23.1467 -1.2 TD 0.0001 Tc (1251-1255.)Tj -0.6333 -1.2 TD 0 Tc (V)Tj 0.6667 0 TD 0.0175 Tc 0.1389 Tw (ilela, C.C. 1999. Is )Tj /F3 1 Tf 9 0 1.913 9 402.03 651.075 Tm (Zaprionus indianus)Tj /F1 1 Tf 9 0 0 9 482.19 651.075 Tm 0 Tc 0 Tw ( )Tj 0.4344 0 TD 0.0175 Tc 0.1389 Tw (Gupta, 1970)Tj -18.7478 -1.2 TD 0.0614 Tc (\(Diptera, Drosophilidea\) currently colonizing the)Tj T* 0 Tc 0.0552 Tw (Neotropical region? Drosophila Information Service)Tj /F3 1 Tf 9 0 1.913 9 522.9301 629.475 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 525.9301 629.475 Tm (82:)Tj -23.1733 -1.2 TD -0.0004 Tc (37-39.)Tj -0.6333 -1.2 TD 0 Tc (V)Tj 0.6333 0 TD 0.0138 Tc 0.1389 Tw [(olf)24(f, J.N., Korting, C., and Schartl, M. 2000. Multiple)]TJ 0 -1.2 TD 0.0247 Tc [(lineages of the non-L)73.7(TR retrotransposon )]TJ /F3 1 Tf 9 0 1.913 9 495.8701 597.075 Tm 0 Tw (Rex1)Tj /F1 1 Tf 9 0 0 9 517.77 597.075 Tm 0 Tc ( )Tj 0.4416 0 TD 0.0247 Tc (with)Tj -22.7082 -1.2 TD 0 Tc -0.0209 Tw (varying success in invading fish genomes. Mol Biol Evol)Tj T* -0.0002 Tw (17: 1673-1684.)Tj -0.6333 -1.2 TD 0 Tw (V)Tj 0.6667 0 TD 0.0571 Tc 0.1389 Tw (os, J.C., De Baere, I., and Plasterk, R.H. 1996.)Tj -0.0333 -1.2 TD 0 Tc 0 Tw (T)Tj 0.5667 0 TD -0.0583 Tw (ransposase is the only nematode protein required for )Tj /F3 1 Tf 9 0 1.913 9 531.45 553.875 Tm 0 Tw (in)Tj -23.5316 -1.2 TD (vitro)Tj /F1 1 Tf 9 0 0 9 334.35 543.075 Tm 0.0005 Tw ( transposition of )Tj /F3 1 Tf 9 0 1.913 9 400.41 543.075 Tm -0.091 Tc 0 Tw [(Tc)-91(1)]TJ /F1 1 Tf 9 0 0 9 414.57 543.075 Tm 0 Tc 0.0005 Tw [(. Genes Dev)72.5(. 10: 755-761.)]TJ -11.4333 -1.2 TD 0.0569 Tw [(Xiong, )16.6(Y)127(., and Eickbush, )17.1(T)111(.H. 1990. Origin and evolution)]TJ 0.6333 -1.2 TD -0.0003 Tw (of retroelements based upon their reverse transcriptase)Tj T* 0.0004 Tw (sequences. EMBO J. 9: 3353-3362.)Tj -0.6333 -1.2 TD 0 Tw (Y)Tj 0.58 0 TD 0.0487 Tw [(oshiyama, M., )19.5(T)37.7(u, Z., Kainoh, )18.9(Y)127(., Honda, H., Shono, )20.6(T)111(.)0(,)]TJ 0.0533 -1.2 TD 0.1109 Tw (and Kimura, K. 2001. Possible horizontal transfer of a)Tj T* 0.0237 Tw (transposable element from host to parasitoid. Mol. 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Proc. Natl.)Tj -19.4733 -1.2 TD 0.0006 Tw (Acad. Sci. USA. 97: 9665-9670.)Tj -0.6333 -1.2 TD -0.004 Tc -0.0694 Tw [(Zimmerly)71.6(, S., Guo, H., Perlman, P)132(.S., and Lambowitz, )52.4(A.M.)]TJ 0.6333 -1.2 TD 0.0098 Tc 0.1389 Tw (1995. Group II intron mobility occurs by target DNA-)Tj T* 0 Tc 0.0003 Tw (primed reverse transcription. Cell)Tj /F3 1 Tf 9 0 1.913 9 449.37 370.275 Tm 0 Tw ( )Tj /F1 1 Tf 9 0 0 9 451.89 370.275 Tm 0.0003 Tw (82: 545-554.)Tj -15.58 -1.2 TD 0.0561 Tc 0.1389 Tw [(Zupunski, V)89.2(., Gubensek, F)108.7(., and Kordis, D. 2001.)]TJ 0.6333 -1.2 TD -0.0138 Tc -0.0695 Tw [(Evolutionary dynamics and evolutionary history in the R)17.9(T)0(E)]TJ T* 0 Tc 0.0502 Tw [(clade of non-L)70.7(TR retrotransposons. Mol. Biol. 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