西瓜銀斑病毒對南黃薊馬幼蟲發育速率及存活率之影響

比較健康及感染 WSMoV 的南黃薊馬飼養於花豆上的存活率 (圖四)，

兩者間並無顯著差異 (² = 2.84, d. f. = 1, P = 0.09)。健康薊馬與感染 WSMoV 的薊馬於花豆上發育至前蛹期的時間分別為 119.4 ± 31.6 小時及 122.9 ± 18.4 小時，兩者間無顯著差異 (表二，Log-likelihood: -339.09, P = 0.73)。以 上結果顯示 WSMoV 的感染對薊馬幼蟲的存活率及發育至前蛹期的時間無直接影響。

表二、西瓜銀斑病毒 (Watermelon silver mottle virus) 的感染對南黃薊馬 (Thrips palmi) 於花豆苗上的幼蟲存活率與幼蟲發育至前蛹期時間的 直接效應

Table 2. Direct effects of Watermelon silver mottle virus infection on larval survival rate and developmental time from newly larvae hatched to prepupae of Thrips palmi on kidney bean seedlings

Test statistic P-value

Survival rate 2.81^a 0.09

Developmental time -339.09^b 0.73

a The log-rank test was used for planned comparison. d. f. = 1.

b The negative binomial distribution was used for planned co mparision.

圖四、幼蟲存活率 (Kaplan–Meier)。健康南黃薊馬 (Thrips palmi) 幼蟲飼育於花豆 苗上的存活率 (n = 65, 黑線) 與感染西瓜銀斑病毒 (Watermelon silver mottle virus) 的南黃薊馬幼蟲於花豆苗上存活率 (n = 70, 虛線)。

Fig. 4. Cumulative survival rate of Thrips palmi larvae (Kaplan–Meier). Healthy thrips larvae were raised on kidney bean seedlings (n=65, drawn line).

Watermelon silver mottle virus-infected thrips larvae were raised on

kidney bean seedlings (n=70, dashed line).

3.2 間接效應

健康南黃薊馬幼蟲飼育於健康植株、經薊馬取食植株及薊馬接種感染 WSMoV 的植株之幼蟲存活率，三者之間沒有顯著差異 (表二、圖五 A，²= 4.6, d. f. = 2, P = 0.31)。結果顯示薊馬取食與 WSMoV 並不會透過薊馬的寄主 植物間接影響薊馬的存活率。發育至前蛹期時間方面，健康南黃薊馬幼蟲於健康植株及薊馬取食植株發育時間有顯著差異 (表三、圖六，Log-likelihood:

-235.10, P < 0.05)；但健康南黃薊馬幼蟲於薊馬接種感染 WSMoV 之植株與 經薊馬取食之植株的發育時間，沒有顯著差異 (Log-likelihood: -163.44, P = 0.52)；健康南黃薊馬幼蟲於健康植株與薊馬接種感染 WSMoV 的植株發育

Time (hours)

20 40 60 80 100

0.0 0.2 0.4 0.6 0.8 1.0

Cummulative survival rate

時間亦無顯著差異 (Log-likelihood: -220.80, P = 0.53)。結果顯示 WSMoV 並 不會透過薊馬的寄主植物間接影響，薊馬的存活率與發育時間。

感染 WSMoV 之南黃薊馬幼蟲於健康植株、經薊馬取食植株及薊馬接種感染 WSMoV 的植株之存活率，三者之間沒有顯著差異 (圖五B，²= 3.5, d. f. = 2, P = 0.17)。經薊馬取食與 WSMoV 對感染 WSMoV 之薊馬的存活率 沒有間接效應，此結果與健康薊馬的結果相同。

21 Healthy thrips larvae were raised on healthy watermelon seedlings (n=47, drawn line), on thrips-damaged watermelon seedlings (n=32, dashed line) and on thrips-inoculated watermelon seedlings (n=32, grey line). (B) Watermelon silver mottle virus-infected thrips larvae were raised on healthy watermelon seedlings (n=46, drawn line), on thrips-damaged watermelon seedlings (n=26, dashed line) and on thrips-inoculated watermelon seedlings (n=40, grey line).

Time (hours)

發育至前蛹期時間方面，感染 WSMoV 的薊馬幼蟲於健康植株與薊馬 取食植株之發育速率有顯著差異 (表三、圖六，log-likelihood: 193.58, P <

0.01)；感染 WSMoV 的薊馬幼蟲於薊馬接種感染 WSMoV 的植株及健康 植株之發育速率有顯著差異 (log-likelihood: -266.85, P < 0.01) ；感染 WSMoV 的薊馬幼蟲於薊馬接種感染 WSMoV 的植株及經薊馬取食植株之 發育速率有顯著差異 (log-likelihood: -193.58, P < 0.01)。感染 WSMoV 的 薊馬幼蟲於健康植株上發育至前蛹期的時間最長，其次為於薊馬接種感染 WSMoV 者，最短者為於經薊馬取食植株發育者。結果顯示不論薊馬有無感染 WSMoV，當薊馬幼蟲飼育於經薊馬取食的植株上會縮短薊馬幼蟲發育至前蛹的發育時間，但 WSMoV 的感染會減弱薊馬取食植株造成的影響。

表三、西瓜銀斑病毒 (Watermelon silver mottle virus, WSMoV) 對南黃薊馬 (Thrips palmi) 於西瓜苗上的幼蟲發育至前蛹期時間的間接效應 Table 3. Indirect effects of Watermelon silver mottle virus (WSMoV) on larva

developmental time from newly hatched to prepupa of Thrips palmi on watermelon seedlings

Comparision

Healthy larvae WSMoV-infected larvae Test statistic^a P Test statistic^a P

Healthy vs. thrips-damaged -235.10 < 0.05 -203.92 < 0.05

Thrips-inoculated vs.

thrips-damaged

-163.44 0.52 -193.58 < 0.01

Healthy vs. thrips-inoculated -220.80 0.53 -266.85 < 0.01

a. The negative binomial distribution was used for planned comparision.

23 watermelon leaf. Black columns: healthy thrips on thrips-inoculated plants (number of prepupa = 16); Watermelon silver mottle virus (WSMoV)-infected thrips on thrips-inoculated plants (number of prepupa = 28); Grey columns: healthy thrips on thrips-damaged plants (number of prepupa = 20); WSMoV-infected thrips on thrips-damaged plants (number of prepupa = 15); White columns: healthy thrips on healthy plants (number of prepupa = 32); WSMoV-infected thrips on healthy plants (number of prepupa = 28). Different letters indicate statistical differences between means at P < 0.05 analyzed by negative binomial distribution.

0 50 100 150

Developmental time (hours)

Healthy thrips WSMoV-infected thrips ab

a a

c b

24 4. 南黃薊馬成蟲之取食偏好性

將南黃薊馬成蟲釋放於健康植株及經薊馬取食的植株之間，雌薊馬在 2、8、12、24 或雄薊馬在釋放 2、12、24 小時均對薊馬取食植物有較高的偏好性 (圖七)。當南黃薊馬釋放於經薊馬取食及薊馬接種感染 WSMoV 的植株間，雌薊馬在任何時間點對兩種植物的偏好性沒有差異，雄薊馬於釋放 2、 4 小時對薊馬接種感染 WSMoV 的植株有較高的偏好性 (圖八)。這指出 WSMoV 的感染可增加雄薊馬對植株的偏好度。當南黃薊馬釋放於健康植株及薊馬接種感染 WSMoV 的植株間，雄薊馬在釋放後 4、8、12、24 小時或雌薊馬在任何時間點對薊馬接種感染 WSMoV 的植株有較高的偏好性 (圖九)。結果顯示南黃薊馬偏好取食感染 WSMoV 的植株。

25 thrips-damaged (grey columns) and healthy seedlings (white columns) in three trials. (B) Number of female thrips adults on thrips-damaged (grey columns) and healthy seedlings (white columns) in three trials.

Ten adults were released between the two seedlings at the beginning of the experiment. * and ** indicate significant differences (P < 0.05 and P < 0.01) between numbers of thrips adults on thrips-damaged and healthy seedlings by sign test. Error bars indicate the standard error of the mean.

Time after thrips release (hours)

(A) (B)

Time after thrips release (hours)

26 Watermelon silver mottle virus (WSMoV)-infected watermelon seedlings. (A) Number of male thrips adults on thrips-damaged (grey columns) and WSMoV-infected seedlings (black columns) in three trails. (B) Number of female thrips adults on thrips-damaged (grey columns) and WSMoV-infected seedlings (black columns) in three trials. Ten adults were released between the two plants at the beginning of the experiment. * and

** indicate significant differences (P < 0.05 and P < 0.01) between numbers of thrips adults on thrips-damaged and WSMoV-infected seedlings by sign test.

Error bars indicate the standard error of the mean.

Time after thrips release (hours) Time after thrips release (hours)

Fig. 9. Feeding preference of Thrips palmi adults to thrips-damaged versus Watermelon silver mottle virus (WSMoV)-infected watermelon seedlings. (A) Number of male thrips adults on healthy (white columns) and WSMoV-infected seedlings (black columns) in three trials. (B) Number of female thrips adults on healthy (white columns) and WSMoV-infected seedlings (black columns) in three trials. Ten adults were released between the two plants at the beginning of the experiment. * and

** indicate significant differences (P < 0.05 and P < 0.01) between numbers of thrips adults on thrips-damaged and WSMoV-infected seedlings by sign test. Error bars indicate the standard error of the mean.

Time after thrips release (hours) Time after thrips release (hours)

28 et al., 1999a; Whitfield et al., 2005)。另外，煙草花薊馬的一齡與二齡幼蟲也 可以傳播 TSWV (Sakimura, 1963)。東方番茄薊馬 (Ceratothripoides claratris) 傳播與 WSMoV 相同血清群之番椒黃化病毒 (Capsicum chlorosis virus, TSWV 雌薊馬成蟲與雄薊馬成蟲的取食電位 (electrical penetration graphs)，

de Kogel and van Deventer, 2003; Hamilton et al., 2005; Zhang et al., 2011)。南 黃薊馬形態上很可能也具有類似構造產生的聚集費洛蒙或性費洛蒙，以吸引

具有偏好性。可是 Abe et al. (2009) 以阿拉伯芥為材料，發現當西方花薊馬 取食後，有關 JA 調控植物防禦基因會大量表現。以塗抹 JA 之阿拉伯芥植株以及健康的植株對雌西方花薊馬進行偏好性測試，結果發現雌薊馬偏好 健康的阿拉伯芥植株。但有趣的是以 JA-insensitive coi1-1 阿拉伯芥突變株，進 行薊馬偏好性試驗，發現薊馬對感染 TSWV 的阿拉伯芥突變株與健康的阿拉伯芥 突變株的偏好性沒有顯著差異 (Abe et al., 2012)。這顯示 JA 與南黃薊馬的 關係仍須更深入的研究才能澄清。

在本實驗中， WSMoV 感染的西瓜對雄南黃薊馬具有較高的吸引力，

但是對雌薊馬則沒有顯示此種差異。但不論雌薊馬或雄薊馬對薊馬取食過的西瓜與健康西瓜相比較有較高的偏好性。這指出病毒的感染及薊馬的取食會影響寄主植物對薊馬的吸引力，此外雄薊馬在 WSMoV 傳播上應扮演重要的角色。

4. 結論

本研究確認南黃薊馬傳播 WSMoV 的傳播機制、試驗 WSMoV 對其病媒南黃薊馬幼蟲發育速率與存活率的直接與間接效應，此外也比較南黃薊馬對健康、薊馬取食及薊馬傳播感染 WSMoV 之植物取食偏好性進行調查。

南黃薊馬主要以持續性增殖型傳播模式傳播 WSMoV，與西方花薊馬傳播 TSWV 之傳播模式相同。南黃薊馬主要於一齡幼蟲期獲得病毒，直到羽化為成蟲期才得以傳播 WSMoV ，且病毒傳播能力沒有性別差異。感染 WSMoV 對南黃薊馬幼蟲之發育速率與存活率沒有直接影響。但是感染 WSMoV 的植物對南黃薊馬的幼蟲發育速率有間接的影響，南黃薊馬幼蟲於感染 WSMoV 的植物上之發育速率相較於健康植物上生長之南黃薊馬幼蟲快。薊馬成蟲取食偏好性方面，南黃薊馬雄成蟲對感染 WSMoV 的植物，

顯示出較大的偏好性。本研究呈現 WSMoV 與南黃薊馬之互利共生關係，

並提供南黃薊馬對 WSMoV 交互作用的概略架構，但是有關 WSMoV 與南 黃薊馬交互作用更細部的關係，尚需進一步研究才可釐清。

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在文檔中南黃薊馬 (纓翅目：薊馬科) 媒介西瓜銀斑病毒之傳播模式及病毒對薊馬生活史特徵之影響 (頁 28-0)

西瓜銀斑病毒對南黃薊馬幼蟲發育速 率 及存活率之影響