Behavioural features of Bursaphelenchus xylophilus in the mating process

in Nematology
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The pine wood nematode (PWN), Bursaphelenchus xylophilus, is able to produce hundreds of thousands of offspring in a short time. Its mating behaviour might play a basic role in this marked fecundity. We investigated the features of the mating behaviour of B. xylophilus using long-time microscope video and repeated observations. The mating behaviour of B. xylophilus could be separated into a series of sequential sub-behaviours, including cruising, approaching, encountering, touching, hooping, locating, attaching, ejaculating, separating, quiescence and roaming. Overall, the process of mating behaviour could be divided into four different phases, searching, contacting, copulating and lingering; the mean times for these different phases varied significantly with 21.8 ± 2.0 min, 28.0 ± 1.9 min, 23.6 ± 0.7 min and 7.2 ± 0.5 min for each of the four phases, respectively. Attraction between the sexes was observed, indicating that sex pheromone(s) might be involved in mating behaviour of B. xylophilus. In addition, when one female was placed with three males, male-male competition was observed, which might be caused by mating-choice pressure from the female. Intra-sexual competition of females was also occasionally observed.


International Journal of Fundamental and Applied Nematological Research



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  • Sub-behaviours in mating process of Bursaphelenchus xylophilus. These photographs of the typical features are taken from the videos of mating process. The mating process includes searching, contacting, copulating and lingering phases. Female and male search everywhere (cruising), attract, approach each other (approaching) until encountering each other (encountering). They touch each other with their heads (touching). The male curls his tail, hoops the female’s body (hooping), and slides along the female’s body and positions at the vulva of the female (positioning). The male clamps and attaches to the vulva of the female (attaching), inserts his spicule into the vulva and ejaculates sperm (ejaculating). The male loosens the curled tail and slides down from the female’s body (separating). They appear transiently quiescent (quiescence). The male subsequently roams around the female and occasionally touches the female (roaming).

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  • Duration percentage of four phases in the mating process of Bursaphelenchus xlophilus. This figure is published in colour in the online edition of this journal, which can be accessed via

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  • Intra-sexual competition and mating choice during the mating process of Bursaphelenchus xylophilus. A, B, C: Males hooped the female’s bodies and competed for the female’s vulva; D: Males swung their body vigorously to hit the mating male or female; E, F: The males even dragged one of them by trapping its body; B, G; The females swung their bodies violently to hit the mating male or female, or waited nearby for the mating male; H, I; The female dragged the male away in order to avoid interference from other males.

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  • A: Contacting frequency between male and female of Bursaphelenchus xylophilus; B: Mating success for the first contact. Calculated mean values are from 30 replicates ±SE. Means with the same lowercase letter within each column are not significantly different at P<0.05 when compared by Tukey’s Honest Significant Difference test.

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