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UNIVERSITE DE BOURGOGNE THÈSE Yongbo LIU - Université de ...

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2002), but certain studies showed no e ffects because of compensatory growth (Hawkes and<br />

Sullivan 2001; Boalt and Lehtila 2007). In an experiment in which we simulated herbivory by<br />

clipping l eaves o f B. juncea we also observed compensatory growth at low herbivory level<br />

(Liu et al., 2009; 2010b). The effect of herbivory on pl ant growth and reproduction <strong>de</strong>pends<br />

on lots of factors such as plant species (Rogers and Siemann 2002), resource level (Hawkes<br />

and S ullivan 2001; R ogers a nd S iemann 2002) , i nduced i nsect-resistance an d d amage t ime<br />

(Agrawal 1998; 1999).<br />

In t he current s tudy, t rBC2 a nd nt rBC2 w ere exposed t o i nsects onl y at pl ant l ategrowth<br />

stage so that it resulted in poor effects. The timing of the insect attack is important.<br />

Agrawal (1999) found that insect-resistance was induced by caterpillar herbivory at the fourth<br />

true l eave o f w ild r adish, an d t his r esistance p ersisted i n n ewly f ormed l eaves o f d amaged<br />

plants, reducing the mass of caterpillars feeding on induced plants compared to un-induced<br />

controls. T hus, t he h erbivory at e arly-growth s tage of pl ants could i nduce t he r esistance t o<br />

herbivory for i nsect-susceptible pl ants, which could un<strong>de</strong> restimate t he differences b etween<br />

resistant and susceptible plants. In this study, the late flowering plants showed higher fitness<br />

than the early flowering ones at the time of insect <strong>de</strong>posit. Most studies involving Bt insect-<br />

resistant f ocused on t he e ffects of i nsects f rom t he ear ly-growth t o m ature, a nd t hey f ound<br />

insect-resistant transgenic plants showed higher fitness only un<strong>de</strong>r mo<strong>de</strong>rate or higher insect<br />

pressure (e.g. Ramachandran et al. 2000; Letourneau and Hagen 2009).<br />

The impact of the insects on the seed weight and number <strong>de</strong>creased with the increased<br />

proportion of t ransgenic pl ants. In c ontrast, w hen pl ants w ere not s ubjected t o i nsects, t he<br />

growth and reproduction of both trBC2 and ntrBC2 showed apparently erratic variations, with<br />

a maximum in populations composed by one half of resistant plants. This genotype proportion<br />

in a l imited r esource e nvironment c ould c orrespond t o t he s patial pl ant di stribution t hat<br />

maximizes an eventual fitness difference between the transgenic and non-transgenic plants. In<br />

other words, in years without insect attack, the spread of transgenic plants would be stopped<br />

by a l ow r eproduction va lue r elative t o t he na tive s usceptible pl ants i f t he t ransgenic<br />

proportion r eaches 50% . O nce ove r t his t hreshold, t he t ransgenes w ould not c onfer a ny<br />

reproduction c ost, w hich ope n t he w ay f or i ts f ixation i n t he popu lations. U n<strong>de</strong>r i nsect<br />

pressure, the growth and reproduction in both trBC2 and ntrBC2 showed linearly increasing<br />

trends w ith t he i ncreased pr oportion of t ransgenic pl ants. T his c ould b e i nterpreted b y a<br />

protection effect applied to all the plants in a cag e, perhaps because the insects leaved from<br />

132

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