Encountering a host
6 Walking away
Rejection 1 Ascending the host
and turning
drumming the host with antennae
2 Probing to insert ovipositor under the host
3 Laying an egg under the host
4 Withdrawing the ovipositor and drumming the dorsal part of the host with hind legs
5 Preening the antennae and legs
(Drumming time)
(Oviposition time) (Time spent searching for an oviposition site )
(Marking time) Total
handling time
Fig. 23. The sequence of behaviors shown by an Eretmocerus mundus female while searching for and after encountering whitefly nymphs.
72
ෆ࡛⾜ࡗࡓࠋᐤ⏕⻏1㢌ᑐࡋ࡚ࢥࢼࢪ࣑ࣛ㢮ᗂ1㢌ࢆ࠼㸪ࢥࢼࢪ࣑ࣛ㢮 ᗂࡢ㱋ู5㢌ࡽ30㢌ࡢᐤ⏕⻏ࢆ౪ヨࡋࡓࠋ
㸲㸬⤫ィศᯒ
ᐤࡢ㱋ูཷࡅධࢀ⋡ࢻ࣑ࣛࣥࢢࣉ࣮ࣟࣅࣥࢢᚋࡢᐤࡢᣄྰ⋡ࡣࣟࢪ ࢫࢸࢵࢡᅇᖐศᯒࢆ⏝࠸ࡓࠋࣟࢪࢫࢸࢵࢡᅇᖐศᯒࡢ⤖ᯝ㸪ᐤࡢཷࡅධ
ࢀ⋡ࡲࡓࡣᣄྰ⋡ᐤ㱋ࡢ㛫᭷ព࡞ᙳ㡪ࡀࡳࡽࢀࡓࡁࡣ㸪᳨࢝ᐃ
ࢆ⾜ࡗࡓᚋ㸪Ryan’s test ࡼࡿከ㔜ẚ㍑ࢆ⾜ࡗࡓࠋᐤ㱋✀ࢆせᅉࡋ࡚
ྛ⾜ື㛫ࢆ2せᅉࡢศᩓศᯒࡼࡾẚ㍑ࡋࡓࠋྛᐤࡢྠ୍㱋㛫㸪␗࡞ࡿ㱋 㛫ࡢ⾜ື㛫ࡢẚ㍑ࡣTukey - Kramer HSD testࡲࡓࡣStudent’s t-test᳨ᐃࢆ⏝࠸
ࡓࠋㄪᰝ⤖ᯝࡢ⤫ィゎᯒࡣJMP5㸦SAS Institute, 2002㸧ࢆ⏝࠸ࡓࠋ
⤖
⤖ᯝ
ࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳࡢᐤ⏕ࡢཷࡅධࢀ⋡ࡣᐤࡢ✀㛫࡛᭷ព࡞ᕪࡣ࡞
ࡗࡓࡀ㸦ࣟࢪࢫࢸࢵࢡᅇᖐศᯒ㸸Ȯ2 = 0.313㸪df = 2㸪p = 0.855㸹Fig. 24㸧㸪ᐤ
ࡢ㱋㛫࡛᭷ព␗࡞ࡗࡓ㸦ࣟࢪࢫࢸࢵࢡᅇᖐศᯒ㸸Ȯ2 = 22.599㸪df = 4㸪p
= 0.0002㸧ࠋࣂ࢜ࢱࣉB㸦᳨࢝ᐃ㸪Ȯ2 = 4.674㸪df = 4㸪p = 0.322㸧
࢜ࣥࢩࢶࢥࢼࢪ࣑ࣛ㸦᳨࢝ᐃ㸪Ȯ2 = 2.353㸪df = 3㸪p = 0.503㸧ࡢᐤࡢ 㱋㛫࡛ࡣ᭷ព࡞ᕪࡣ࡞ࡗࡓࡀ㸪ࣂ࢜ࢱࣉQ࡛ࡣᐤࡢ㱋㛫࡛᭷ព␗࡞
ࡾ㸦᳨࢝ᐃ㸪Ȯ2 = 13.514㸪df = 4㸪p = 0.009㸧㸪4㱋ᚋᮇࡢᗂࡢཷࡅධ
ࢀ⋡ࡣⱝ㱋ẚ᭷ពపࡗࡓ㸦Ryan’s test㸪p < 0.05㸧㸦Fig. 24㸧ࠋࡲࡓ㸪࢜
ࣥࢩࢶࢥࢼࢪ࣑ࣛࡢ4㱋ᚋᮇࡢᗂࡣ⏘༸ࡋ࡞ࡗࡓ㸦Fig. 24㸧ࠋ࡞࠾㸪࡚
ࡢᐤ⏕⻏ࡢෆ 14 ಶయ㸦4.7%㸧ࡀ࣍ࢫࢺࣇ࣮ࢹࣥࢢ㸦ᐤయᾮᦤྲྀ㸧ࢆ⾜
ࡗࡓࠋ
ࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳࡀᐤࢆᣄྰࡍࡿ⾜ືࡣࢻ࣑ࣛࣥࢢࡲࡓࡣࣉ࣮ࣟࣅ
73
㻜 㻞㻜 㻠㻜 㻢㻜 㻤㻜 㻝㻜㻜
㻮 㻽 㼀
㻴 㼛 㼟㼠 㻌㼍 㼏㼏㼑㼜 㼠㼍 㼚㼏㼑㻌 㼞㼍 㼠㼑 㻌㻔 㻑 㻕
䠍㻌㻌䠎㻌㻌䠏㻌㻌㻱㻠㻌㻌㻸㻠 㻌㻌㻌㻌
㻮㻚㻌㼠㼍㼎㼍㼏㼕
㻌㻌㻌㻌㻌㻌㻌㻮㻌㼎㼕㼛㼠㼥㼜㼑䠍㻌㻌䠎㻌㻌䠏㻌㻌㻱㻠㻌㻌㻸㻠 㻌
㼀㻚㻌㼢㼍㼜㼛㼞㼍㼞㼕㼛㼞㼡㼙
䠍㻌㻌䠎㻌㻌䠏㻌㻌㻱㻠㻌㻌㻸㻠㻌㻌㻌䚷
㻮㻚㻌㼠㼍㼎㼍㼏㼕
䚷䚷㻌䚷㻽㻌㼎㼕㼛㼠㼥㼜㼑㻌㻴㼛㼟㼠㻌㼟㼠㼍㼓㼑 㼍
㼎 㼍
㼍 㼍
Fig. 24. The acceptance rates with Wald 95 percent confidence interval of Eretmocerus mundus for different nymphal instars of the Bemisia tabaci B and Q biotypes and of Trialeurodes vaporariorum. Bars labeled with different letters differ significantly among nymphal instars of the same host species (Ryan’s test for multiple comparisons, p < 0.05). There were no significant differences in B. tabaci B biotypes and T. vaporariorum (Ryan’s test for multiple comparisons, p > 0.05). The numbers of parasitoids tested are shown in Table 7. E4 and L4 indicate the early and late fourth instar stages, respectively.
74
Table 8. The host rejection rates by Eretmocerus mundus for different nymphal instars of the Bemisia tabaci B and Q biotypes and of Trialeurodes vaporariorum after completion of the drumming and probing behaviors
Host species Host stage No. of parasitoids
tested
Host rejection rate after drumming
㸦%㸧
No. of probing parasitoids
Host rejection rate after probing
㸦%㸧
Bemisia tabaci B biotype
first 22 9.1 a 20 15.0 a
second 24 33.3 a 16 6.3 a
third 22 18.2 a 18 22.2 a
early fourth 20 30.0 a 14 21.4 a
late fourth 8 50.0 a 4 25.0 a
Bemisai tabaci Q biotype
first 27 3.7 b 26 26.9 a
second 29 24.1 ab 22 13.6 a
third 23 13.0 b 20 5.0 a
early fourth 17 17.6 ab 14 14.3 a
late fourth 15 53.3 a 7 42.9 a
Trialeurodes vaporariorum
first 16 12.5 b 14 14.3 a
second 13 46.2 ab 7 0 a
third 30 46.7 ab 16 0 a
early fourth 13 38.5 ab 8 12.5 a
late fourth 5 100 a 0 —
Values followed by different letters differ significantly among the host stages within a host species or biotype (logistic regression analysis followed by Ryan’s test, p < 0.05).
75
ࣥࢢࡢᚋほᐹࡉࢀࡓࠋࢻ࣑ࣛࣥࢢᚋࡢᐤᣄྰ⋡ࡣᐤࡢ㱋㛫࡛᭷ព␗࡞
ࡗࡓࡀ㸦ࣟࢪࢫࢸࢵࢡᅇᖐศᯒ㸸Ȯ2 = 34.664㸪df = 4㸪p < 0.0001㸧㸪ᐤࡢ✀
㛫࡛ࡣ᭷ព࡞ᕪࡀ࡞ࡗࡓ㸦ࣟࢪࢫࢸࢵࢡᅇᖐศᯒ㸸Ȯ2 = 1.240㸪df = 2㸪p =
0.538㸧㸦Table 8㸧ࠋ࢜ࣥࢩࢶࢥࢼࢪ࣑ࣛ4㱋ᗂᑐࡍࡿᣄྰࡣ࡚ࢻ࣑ࣛࣥࢢ
ᚋࡳࡽࢀࡓ㸦Table 8㸧ࠋ࡚ࡢ✀ࡢ2㱋௨ୖࡢᗂᑐࡍࡿᐤᣄྰ⋡ࡣࢻࣛ
࣑ࣥࢢᚋẚࣉ࣮ࣟࣅࣥࢢᚋࡢ᪉ࡀపࡗࡓ㸦Table 8㸧ࠋࣉ࣮ࣟࣅࣥࢢᚋࡢᐤ
ᣄྰ⋡ᐤࡢ㱋✀ࡣ୧᪉ࡶᙳ㡪ࡋ࡞ࡗࡓ㸦ࣟࢪࢫࢸࢵࢡᅇᖐศ ᯒ:ᐤ㱋㸸Ȯ2 = 6.656㸪df = 4㸪p = 0.155㸪ᐤ㸸X2 = 1.367㸪df = 2㸪p = 0.505㸹 Table 8㸧ࠋ
ࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳࡣ࢜ࣥࢩࢶࢥࢼࢪ࣑ࣛࡢ4㱋ᚋᮇࡢᗂᑐࡋ࡚ࢻ
࣑ࣛࣥࢢࡣ⾜ࢃ࡞ࡗࡓ㸦Fig. 25 a㸧ࠋࡑࢀ௨እࡢ✀ᑐࡍࡿࢻ࣑ࣛࣥࢢ㛫ࡣ ᐤࡢ㱋㛵ࢃࡽࡎᐤࡢ✀㛫࡛ᕪࡀ࡞ࡗࡓࡀ㸦ANOVA㸸F 2, 161 = 0.270㸪 p = 0.764㸧㸪ᐤࡢ㱋㛫࡛ࡣ᭷ព␗࡞ࡗࡓ㸦F 4, 161 = 5.009㸪p = 0.0024㸹Fig. 25 a㸧ࠋࣂ࢜ࢱࣉQࡢ4㱋๓ᮇࡢᗂᑐࡍࡿࢻ࣑ࣛࣥࢢ㛫ࡣ㸪1㱋3㱋 ᗂẚ᭷ព㛗ࡗࡓ㸦Tukey - Kramer HSD test㸪p < 0.05㸹ANOVA㸸F 4, 68
= 3.337㸪p = 0.015㸹Fig. 25 a㸧ࠋ
ࣉ࣮ࣟࣅࣥࢢ㛫ࡣᐤࡢ✀㛫࡛᭷ពᕪࡣ࡞ࡃ㸦ANOVA㸸F2, 161 = 0.051㸪p
= 0.950㸧㸪ᐤࡢ㱋㛫࡛᭷ព␗࡞ࡗࡓ㸦F 3, 161 = 14.145, p < 0.001㸹Fig. 25 b㸧ࠋ
ࣂ࢜ࢱࣉBࡢ4 㱋ᚋᮇᗂᑐࡍࡿࣉ࣮ࣟࣅࣥࢢ㛫ࡣ1㱋㸪2 㱋࠾ࡼࡧ 3㱋ᗂẚ᭷ព㛗ࡗࡓ㸦Tukey - Kramer HSD test㸸ANOVA㸸F 4, 55 = 6.511㸪 p = 0.0002㸹Fig. 25 b㸧ࠋࣂ࢜ࢱࣉQࡢ4㱋ࡢ๓ᮇᚋᮇᗂᑐࡋ࡚ࡣ㸪1 㱋㸪2㱋㸪3㱋ᗂẚ᭷ព㛗ࡗࡓ㸦Tukey - Kramer HSD test㸪P < 0.05㸹 ANOVA㸸F 4, 68 = 13.886㸪p < 0.001㸹Fig. 25 b㸧ࠋ
⏘༸㛫ࡣᐤࡢ✀㛫࡛᭷ព࡞ᕪࡣࡳࡽࢀ࡞ࡗࡓࡀ㸦ANOVA㸸F 2, 161 = 1.548㸪p = 0.216㸧㸪ᐤࡢ㱋㛫࡛ࡣ᭷ព࡞ᕪࡀࡳࡽࢀࡓ㸦F 3, 161 = 2.7626㸪
76
Fig. 25. Time required for behavioral events. (a) drumming time, (b) probing time, (c) oviposition time, (d) marking time, and (e) total handling time of Eretmocerus mundus on different nymphal instars of the Bemisia tabaci B and Q biotypes and ofTrialeurodes vaporariorum. Values are means ± S.E. Bars labeled with different capital letters differ significantly among host species or biotypes; bars labeled with different lowercase letters differ significantly among host stages, except for the late fourth instars, within a species or biotype (two-factor ANOVA followed by the Tukey-Kramer HSD, or the Student’s t-test for L4, p < 0.05). E4 and L4 indicate the early and late fourth instar stages, respectively.
㻜
㻡㻝㻜
㻝㻡
㻞㻜 Marking time (s)
Marking time Aa Aa Ba Aa
Aa Ab
AaAa
AaAa
ABa AaAa
Aa
(d) 㻜
㻞㻜㻜
㻠㻜㻜
㻢㻜㻜
㻤㻜㻜㻝㻜㻜㻜 㻮㻽㼀
Total handling time (s)
Total handling time 䠍䠎䠏E4 L4 B. tabaci B biotype
䠍䠎䠏E4 L4 B. tabaci Q biotype
䠍䠎䠏E4 L4 T. vaporariorum
Ab Ab
Ab ABb
Aa Bb Ab Bb
Aa
Aa Aa Aab AB
b Aa
b Host stage
(e)
㻜㻞㻜
㻠㻜
㻢㻜
㻤㻜㻝㻜㻜
Drumm ing tim e (s)
Drumming time Aa Aa Aa AaAa Aa
Aa Aa
Aa
(a) Ab
Aab Ab
Aa Aab 㻜
㻞㻜㻜
㻠㻜㻜
㻢㻜㻜
㻤㻜㻜㻝㻜㻜㻜Searching time Ab Ab
Aa Aab Ab Ab
Aa
Aa
(b) Aa Aa
Aa Aa 㻜
㻞㻜㻜
㻠㻜㻜
㻢㻜㻜
㻤㻜㻜㻝㻜㻜㻜 㻮㻽㼀
Oviposition tim e (s)
Oviposition time 䠍䠎䠏E4 L4 B. tabaci B biotype
䠍䠎䠏E4 L4 B. tabaci Q biotype
䠍䠎䠏E4 L4 T. vaporariorum
Aab Ab
AabAa Aa AaAa Aa Aa Aa AaAa Aa
(c) Host stage
Probing time (s)
77
p = 0.044㸹Fig. 25 c㸧ࠋࣂ࢜ࢱࣉBࡢ⏘༸㛫ࡣ㸪2㱋3㱋ᗂẚ㸪4 㱋ᚋᮇᗂ࡛᭷ព㛗ࡗࡓ㸦✀ࡈᐤࡢ㱋㛫࡛Tukey - Kramer HSD test
࡛ẚ㍑㸪p < 0.05㸹ANOVA㸸F 4, 55 = 3.381㸪P = 0.015㸹Fig. 25 c㸧ࠋ
࣐࣮࢟ࣥࢢ⾜ືࡣ࢜ࣥࢩࢶࢥࢼࢪ࣑࡛ࣛࡣほᐹࡉࢀ࡞ࡗࡓ㸦Fig. 25 d㸧ࠋ࣐
࣮࢟ࣥࢢ㛫ࡣᐤࡢ✀㛫࡛᭷ព࡞ᕪࡣ࡞ࡗࡓࡀ㸦ANOVA㸸F 2, 161 = 0.009㸪 p =0.991㸧㸪ᐤࡢ㱋㛫࡛᭷ព␗࡞ࡗࡓ㸦F 4, 161 = 5.006㸪p = 0.0024㸧ࠋࣂ
࢜ࢱࣉBᑐࡍࡿ࣐࣮࢟ࣥࢢ㛫ࡣ 3㱋ᗂࡀ᭱ࡶ㛗ࡃ㸪1㱋ᗂࡀ᭱ࡶ▷
ࡗࡓ㸦Tukey - Kramer’s HSD test, p < 0.05㸹ANOVA㸸F 4, 55 = 3.243㸪p = 0.0185㸸
Fig. 25 d㸧ࠋࣂ࢜ࢱࣉQᑐࡍࡿ࣐࣮࢟ࣥࢢ㛫ࡣᐤࡢ㱋㛫࡛᭷ព࡞ᕪࡀ
࡞ࡗࡓ㸦ANOVA㸸F 4, 68 = 1.615㸪p = 0.181㸧ࠋ
ࣂ࢜ࢱࣉQ ࣂ࢜ࢱࣉ B ࡢ1 㱋࠾ࡼࡧࣂ࢜ࢱࣉ Q࢜ࣥࢩࢶ ࢥࢼࢪ࣑ࣛࡢ3㱋ࡢ⥲⏘༸㛫ࢆẚࡿ㸪࠸ࡎࢀࡶࣂ࢜ࢱࣉQࡀ᭷ព
▷ ࡃ 㸪 ࡑ ࡢ ࡛ ࡣ ⏘ ༸ ࡋ ࡓ ᐤ ⏕ ⻏ ࡢ ⥲ ⏘ ༸ 㛫 ࡣ ᐤ ࡢ ✀ 㛫 ࡛ ᕪ ࡣ ࡞ ࡃ 㸦ANOVA㸸F 2, 161 = 0.472㸪p = 0.625㸧㸪ᐤࡢ㱋㛫࡛ࡣ᭷ពᕪࡀ࠶ࡗࡓ 㸦ANOVA㸸F 2, 161 = 17.297㸪p < 0.0001㸧ࠋ࡚ࡢ✀ࡢ⪁㱋ᗂᑐࡍࡿ⥲⏘༸
㛫ࡀ㛗ࡃ࡞ࡗࡓ㸦Tukey-Kramer HSD test㸪p < 0.05㸹ࣂ࢜ࢱࣉB F 4, 55 = 7.820㸪p < 0.001㸪ࣂ࢜ࢱࣉQ F 4, 68 = 17.556㸪p < 0.001㸪࢜ࣥࢩࢶࢥࢼࢪ
࣑ࣛF 3, 38 = 3.005㸪p = 0.042㸹Fig. 25 e㸧ࠋ
⪃
⪃ᐹ
ᐤ⏕⻏ࡢ⏘༸⾜ືࡣỴࡲࡗࡓ㡰ᗎ࡛⾜ࢃࢀࡿ㸦Vinson㸪1998㸧ࠋࢳࢳࣗ࢘࢝
ࢶࣖࢥࣂࢳࡢ⏘༸⾜ືࡣ㸪ᚑ᮶㸪3ࡘศࡅࡽࢀ࡚࠸ࡓࡀ㸦Foltyn and Gerling㸪 1985㸹Headrick et al.㸪1996㸧㸪Ardeh et al.㸦2005㸧ࡣ5ࡘ⣽ศࡋ࡚࠸ࡿࠋᮏ◊
✲࡛ࡣArdeh et al.㸦2005㸧ࡼࡿࠕゐゅࡼࡿᐤࡢࣉ࣮ࣟࣅࣥࢢࠖࠕᐤ
ୖࡢࡰࡗ࡚㸪ࡑࡢ࿘㎶ࢆゐゅࡼࡾ᥈⣴ࠖࡢ⾜ືࡣ᫂☜ศࡅࡿࡇࡀᅔ
78
㞴࡛࠶ࡗࡓࡇࡽ㸪ࠕࣉ࣮ࣟࣅࣥࢢࠖࡢࡦࡘࡢ⾜ືࡋࡓࠋࡲࡓ㸪ࠕ⏘༸⟶
ࡢᤄධ⏘༸ࠖࡘ࠸࡚ࡣ㸪ᐤ⏕⻏ࡀ⏘༸⟶ࢆᤄධࡋ࡚࠸ࡿࡣ⭡㒊ࢆఱᗘࡶ
ືࡋ㸪⏘༸ࡣ⭡㒊ࢆືࡉ࡞࠸ᵝᏊࡀほᐹࡉࢀࡓࡓࡵ㸪ࠕ⏘༸⟶ࡢᤄධࠖ
ࠕ⏘༸ࠖศࡅ㸪ࠕ⏘༸⟶ࡢᤄධࠖࡣࠕࣉ࣮ࣟࣅࣥࢢࠖྵࡵࡓࠋᚑࡗ࡚㸪ᮏ
◊✲࡛ࡣ㸪ࠕࢻ࣑ࣛࣥࢢࠖ㸪ࠕࣉ࣮ࣟࣅࣥࢢࠖ㸪ࠕ⏘༸ࠖ࠾ࡼࡧࠕ࣐࣮࢟ࣥࢢࠖࡢ4 ࡘศ㢮ࡋ࡚ㄪᰝࡋࡓࠋ
ࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳࡢᐤࡋ࡚ࢱࣂࢥࢥࢼࢪ࣑ࣛࡣ࢜ࣥࢩࢶࢥࢼࢪࣛ
࣑ࡼࡾ㐺ࡋ࡚࠸ࡿ⪃࠼ࡽࢀ࡚࠸ࡿࠋࡓ࠼ࡤ㸪ࢱࣂࢥࢥࢼࢪ࣑ࣛᑐࡍࡿᐤ
⏕⋡ࡣ࢜ࣥࢩࢶࢥࢼࢪ࣑ࣛࡼࡾ㧗ࡃ㸪Ꮴ⋡ࡶ㧗࠸࠺࠼㸪ᗂࡢⓎ⫱ᮇ㛫ࡶ▷
ࡃ㸪ᡂࡢᑑࡶ㛗࠸ሗ࿌ࡉࢀ࡚࠸ࡿ㸦Heinz and Parrella㸪1994㸹Greenberg et al.㸪2002㸧ࠋGreenberg et al.㸦2002㸧ࡶࢱࣂࢥࢥࢼࢪ࣑ࣛᑐࡍࡿᐤ⏕⋡ࡣ࢜ࣥ
ࢩࢶࢥࢼࢪ࣑ࣛࡢ 3.4 ಸ࡛㸪Ꮴ⋡ࡶࢱࣂࢥࢥࢼࢪ࣑ࣛࡢ 84.9%ᑐࡋ࡚࢜ࣥ
ࢩࢶࢥࢼࢪ࣑࡛ࣛࡣ 19.9%࡛࠶ࡗࡓሗ࿌ࡋ࡚࠸ࡿࠋ୍᪉㸪ᐤ⏕⻏⏘༸⤒㦂 ࡀ࠶ࡿሙྜࡸᐤ⏕⻏ࡀ」ᩘ✀ࢆ㑅ᢥ࡛ࡁࡿሙྜࡣᐤ⏕⻏ࡢᐤ⏕⋡ࡀᐤࡼ
ࡾ␗࡞ࡿࡇࡀሗ࿌ࡉࢀ࡚࠸ࡿࡀ㸦Cornell and Pimentel㸪1978㸹van Alphen and Vet㸪
1986㸧㸪ᮏヨ㦂࡛ࡣࢱࣂࢥࢥࢼࢪ࣑ࣛ࢜ࣥࢩࢶࢥࢼࢪ࣑ࣛࡢ㛫࡛ᐤ⏕⋡᭷ព
࡞ᕪࡣㄆࡵࡽࢀ࡞ࡗࡓ㸦Fig. 24㸧ࠋࡇࡢཎᅉࡣ㸪ᮏヨ㦂࡛ࡣ⏘༸ᮍ⤒㦂ࡢᡂ
1 㢌 1 ಶయࡢࢥࢼࢪ࣑ࣛᗂࢆ౪ヨࡋࡓࡓࡵ㸪㑅ᢥࡢవᆅࡀ࡞ࡃ㸪ࡇࢀࡲ࡛
ࡢሗ࿌␗࡞ࡿ⤖ᯝࡀᚓࡽࢀࡓࡢࡶࡋࢀ࡞࠸ࠋ
ࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳࡣࢱࣂࢥࢥࢼࢪ࣑ࣛࡢ4㱋ᗂᑐࡍࡿᐤ⏕ࡣᑡ࡞
࠸ሗ࿌ࡉࢀ࡚࠸ࡿࡀ㸦Urbaneja and Stansly㸪2004㸹Qiu et al.㸪2004㸧㸪ᮏヨ㦂
࡛ࡶ4㱋ᚋᮇࡢᐤࡢᐤ⏕⋡ࡣⱝ㱋ᗂẚ᭷ពపࡃ㸦Fig. 24㸧㸪ࡇࢀࡲ࡛
ࡢሗ࿌ྠࡌ⤖ᯝ࡛࠶ࡗࡓࠋ
Table 8♧ࡋࡓᐤᣄྰ⋡ࢆࡳࡿ㸪ࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳࡣࢻ࣑ࣛ
ࣥࢢ୰㐼㐝ࡋࡓᗂࢆᐤࡋ࡚ཷࡅධࢀࡿ࠺ุ᩿ࡋ࡚࠸ࡿࡼ࠺ࡳ
79
࠼ࡿࠋࡲࡓ㸪ᐤ⏕⻏ࡀᐤࢆ㐺ุ᩿ࡋࡓሙྜࡣ⏘༸⾜ືࢆ┤ࡕ୰Ṇࡋ
࡚࠸ࡿࠋᐤ⏕⻏ࡢ⏘༸⾜ືࡣᐤࡢࡁࡉ㸦Salt㸪1958㸧ࡸయ⾲ࡢ࣡ࢵࢡࢫᡂ ศࡢ㐪࠸㸦Muratori et al.㸪2006㸧ࡀᙳ㡪ࡋ࡚࠸ࡿ⪃࠼ࡽࢀ࡚࠸ࡿࡀ㸪ࡑࡢヲ
⣽ࡣ࡛᫂࠶ࡿࠋ
ᐤ㐼㐝ᚋࡢࢻ࣑ࣛࣥࢢࡽ⏘༸⤊ࡲ࡛ࡢ⥲⏘༸㛫ࡣ࠸ࡎࢀ࠾࠸࡚
ࡶ4㱋ᗂ㸦288㹼522 ⛊㸧ࡀ᭱ࡶ㛗ࡃ㸪1㹼3㱋ᗂ㸦96㹼231⛊㸧ẚ࡚⣙
2㹼4ಸࡶ㛗ࡗࡓ㸦Fig. 25 e㸧ࠋࣂ࢜ࢱࣉBQ࡛ࡣ4㱋ࡢࣉ࣮ࣟࣅࣥࢢ
㛫ࡀ1㹼3㱋ᗂẚ㛗࠸㸦Fig. 25 c㸧ࠋⱝ㱋ᗂẚ㸪4㱋ᗂࡣⴥᅛ
╔ࡋ࡚࠸ࡿࡇࡽ㸪ᐤⴥࡢ㝽㛫ࡀ⊃࠸ࠋࡲࡓ㸪ࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳ ࡢ⏘༸⟶ࡣ‴᭤ࡋ㸪ⷧ⫗࡛࠶ࡿࡓࡵ㸦Gerling et al.㸪1998㸧㸪ࡇ࠺ࡋࡓ⊃࠸㝽㛫
ᤄධࡍࡿࡇࡀᅔ㞴ࡳࡽࢀࡿ㸦Ardeh et al.㸪2005㸧ࠋࡑࡢࡓࡵ㸪ࣉ࣮ࣟࣅࣥ
ࢢ㛫ࡣቑຍࡋ㸪⥲⏘༸㛫ࡣ㛗ࡃ࡞ࡗࡓ⪃࠼ࡽࢀࡿࠋ
ከࡃࡢᐤ⏕⻏ࡣ㐣ᐤ⏕ࢆ㑊ࡅࡿࡓࡵ⏘༸ᚋࡢᐤ࣐࣮ࢡࢆࡘࡅࡿ㸦Vinson㸪 1998㸹Nufio and Papaj㸪2001㸧ࠋࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳࡢሙྜࡣ⏘༸ᚋࡢᐤ
ࢆᚋ⬮࡛྇ࡃࢻ࣑ࣛࣥࢢࡢ㝿࣐࣮ࢡࢆࡘࡅࡿ 㸦Ardeh et al.㸪2005㸧ࠋBuckner and Jones㸦2005㸧ࡣ㸪ࡇࡢᏛ≀㉁ࡣᐤ⏕⻏ࡢ⭡㒊ࡲࡓࡣᚋ⬮࠶ࡿ≉ูࡢ⭢
ࡽศἪࡉࢀࡓ࣓ࢳࣝศᯞࣝ࢝ࣥ⬡㉁㸦additional lipids㸧ࡢΰྜ≀࡛࠶ࡿ
ሗ࿌ࡋ࡚࠸ࡿࠋᮏ◊✲࡛ࡣ㸪࣐࣮࢟ࣥࢢ⾜ືࡣࣂ࢜ࢱࣉBࣂ࢜ࢱࣉ
Q࡛ほᐹࡉࢀࡓࡀ㸪࢜ࣥࢩࢶࢥࢼࢪ࣑࡛ࣛࡣほᐹࡉࢀ࡞ࡗࡓ㸦Fig. 25 d㸧ࠋࡇ ࡢ⯆῝࠸⌧㇟ࡢヲ⣽ࡘ࠸࡚ࡣᚋࡢ᳨ウࢆᚅࡕࡓ࠸ࠋ
௨ୖࡢ࠾ࡾ㸪ࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳࡣࢱࣂࢥࢥࢼࢪ࣑ࣛࣂ࢜ࢱࣉB㸪
ࣂ࢜ࢱࣉQ࠾ࡼࡧ࢜ࣥࢩࢶࢥࢼࢪ࣑ࣛࡢ࠸ࡎࢀࡶᐤ⏕ࡍࡿࡀ㸪⪁㱋ᗂࡼ
ࡾⱝ㱋ᗂࢆዲࡴࡇࡀ᫂ࡽ࡞ࡗࡓࠋ
80
➨
➨㸱⠇ '1$ ࣐࣮࣮࢝ࢆ⏝࠸ࡓᐤ⏕⻏ࡢ㆑ูἲ
ᮦᩱ࠾ࡼࡧ᪉ἲ
⏕≀㎰⸆ࡋ࡚㈍ࡉࢀ࡚࠸ࡿ࢜ࣥࢩࢶࢶࣖࢥࣂࢳ㸪ࢧࣂࢡࢶࣖࢥࣂࢳ࠾ࡼ
ࡧࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳࢆ⏝࠸ࡓ㸦Table 9㸧ࠋᐤ⏕⻏ࡢ࣐࣑࣮࣮࢝ࢻࢆ࣐࣑࣮
ࡢ㒊ศࡢࡳࢆษࡗ࡚࢞ࣛࢫ⟶㸦ཱྀෆᚄ10 mm㸪⬗ᚄ21 mm㸪㧗ࡉ45 mm㸧ෆ
1࣮࢝ࢻࢆධࢀ㸪ᜏ ᐊෆ㸦25 °C㸪47% RH㸪16L8D㸧࡛ಖᏑࡋ㸪⩚ࡋࡓᡂ
ࢆ 1 ಶయࡎࡘ 0.5mL ࣐ࢡࣟࢳ࣮ࣗࣈ୰ධࢀ࡚Ѹ80Υ࡛ಖᏑࡋࡓࠋ࡞࠾㸪ࢧ
ࣂࢡࢶࣖࢥࣂࢳ࠾ࡼࡧࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳࡣ㸪ಖᏑඛ❧ࡕᙧែࡼࡾ㞤 㞝㆑ูࢆ⾜ࡗࡓࠋࡲࡓ㸪ᑐ↷ࡋ࡚㸪࢚ࢱࣀ࣮ࣝ୰ಖᏑࡋࡓࢱࣂࢥࢥࢼࢪࣛ
࣑㸦ࣂ࢜ࢱࣉB࠾ࡼࡧQ㸧࠾ࡼࡧ࢜ࣥࢩࢶࢥࢼࢪ࣑ࣛࡶ⏝࠸ࡓ㸦Table 9㸧ࠋ ࣉ࣐࣮ࣛࡢタィࡣᅜ㝿ሷᇶ㓄ิࢹ࣮ࢱ࣮࣋ࢫⓏ㘓ࡉࢀ࡚࠸ࡿࣜ࣎ࢰ࣮࣒
RNA 㑇ఏᏊ㡿ᇦ㸦rDNA㸧ࡢሷᇶ㓄ิࢆඖ⾜ࡗࡓࠋ⏝࠸ࡓࢹ࣮ࢱࡣࢳࢳࣗ࢘
࢝ࢶࣖࢥࣂࢳࡢ18SrDNAࡽ5.8SrDNAࡢ㡿ᇦ㸦AF273631-AF273636㸧࠾ࡼ
ࡧ࢜ࣥࢩࢶࢶࣖࢥࣂࢳࡢ Internal Transcribed Spacer 1㸦ITS1㸧㡿ᇦࢆྵࡴ㸪 18SrDNA ࡢ 3'ᮎ ➃ ࡽ 5.8SrDNA ࡢ 5'ᮎ ➃ ࡅ ࡚ ࡢ 㡿 ᇦ 㸦AY615782- AY615783㸧࡛࠶ࡿࠋ
ಶయࡈWizard® Genomic DNA Purification Kit㸦ࣉ࣓ࣟ࢞㸧ࡼࡾDNAᢳ ฟࢆ⾜࠸㸪᭱⤊ⓗᚓࡽࢀࡓDNAࡣTris-EDTA㸦10mM-0.1mM㸪pH 8.0㸧⁐ᾮ
୰⁐ゎࡋ࡚㸪௨ᚋࡢPCRᛂ౪ヨࡋࡓࠋ
PCRᛂࡣ㸪1ᛂ࠶ࡓࡾdNTP 0.2μmol/mL㸪ྛࣉ࣐࣮ࣛ 0.2μmol/mL㸪ࢤ
ࣀ࣒DNA0.5μL㸪ExTaq (ࢱ࢝ࣛ) 0.2unit㸪࠾ࡼࡧῧࣂࢵࣇ࣮10ศࡢ1㔞ࢆ
ྵࡴ㸪㔞10μLࡢỈ⁐ᾮ୰࡛⾜ࡗࡓࠋPCRᛂࡣiCyclerࢧ࣮࣐ࣝࢧࢡࣛ
࣮㸦ࣂ࢜ࣛࢵࢻ㸧ࢆ⏝࠸㸪94Υ3ศࡢᚋ㸪94Υ30⛊㸪55Υ30⛊㸪72Υ60⛊ࡢ ࢧࢡࣝࢆ35ᅇ⧞ࡾ㏉ࡋࡓᚋ㸪72Υ࡛10ศ㛫ಖᣢࡋࡓࠋቑᖜࡋࡓDNA᩿∦
81
Table 9. Strains of whitefly parasitoids and whiteflies used for this study
Species Strain* Supplier No. of
females No. of males Encarsia formosa Tsuya-Top Agrisect 16
En-Strip Arysta
LifeScience 16 Tsuyakobachi
EF30 Syngenta 16
Eretmocerus remicus Sabaku-Top Agrisect 12 12 Er-Card Arysta
LifeScience 8 8
Eretmocerus mundus Bemipar Arysta
LifeScience 8 8
Trialeurodes
vaporariorum 8**
Bemisia tabaci B-type 8**
Q-type 8**
* For parasitoid wasps, strain names are commercial names.
** Sex were not identified.
82
18S 5.8S
EmF
EmR EfF
EfR
Fig. 26. Schematic diagram of primers designed on the sequences of ribosomal RNA gene in this study. White and black arrows indicate the primers designed on the sequences of Encarsia formosa and Eretmocerus mundus, respectively.
83
Table 10. Primers of whitefly parasitoids used in this study
Name Sequence (5’–3’)
EfF CCTTCTCGTTTTGCACTTGG
EfR TCGACGAGTTCATTGATGTGA
EmF AACGTATGCGGATAACAACG
EmR GCAATTAGCTGCGTTCTTCA
D3-4046F * GACCCGTCTTGAAACACGGA
D3-4413R * TCGGAAGGAACCAGCTACTA
* Gillespie et al. (2005)
84
ࡣTAE⦆⾪ᾮ୰ࡢ1.5%࣮࢞ࣟࢫࢤࣝୖ࡚100V࡛40ศ㛫㟁ẼὋືࢆ⾜ࡗ
࡚ศ㞳ࡋ㸪࢚ࢳࢪ࣒࢘ࣈ࣐ࣟࢻ⁐ᾮ࡛ᰁⰍᚋ㸪UV ↷ᑕୗ࡛ࣂࣥࢻࣃࢱ࣮ࣥ
ࢆ☜ㄆࡋࡓࠋ࡞࠾㸪సᡂࡋࡓࣉ࣐࣮࡛ࣛࡢቑᖜ☜ㄆඛ❧ࡕ㸪rDNA ࡢ 28S 㒊ศ㡿ᇦࢆቑᖜࡍࡿỗ⏝ⓗ࡞ࣉ࣐࣮ࣛ㸦D3-4046F࠾ࡼࡧD3-4413R㸹 Gillespie et al.㸪2005㸪Table 10㸧ࢆ⏝࠸ࡓPCRࡼࡗ࡚DNAᢳฟࢆ☜ㄆࡋࡓࠋࡲࡓ㸪ᚓ
ࡽࢀࡓቑᖜDNA᩿∦ࡀITS1㡿ᇦ࠺☜ㄆࡍࡿࡓࡵ㸪࢜ࣥࢩࢶࢶࣖࢥࣂࢳ 㸦࢚ࣥࢫࢺࣜࢵࣉ®࠾ࡼࡧࢶࣖࢺࢵࣉ®㸧㸪ࢧࣂࢡࢶࣖࢥࣂࢳ㸦࢚࣮ࣝ࢝ࢻ®࠾ࡼ
ࡧࢧࣂࢡࢺࢵࣉ®㸧ྛ⣔⤫1ಶయࡎࡘ㸪ࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳ㸦࣑࣋ࣃ࣮ࣝ®㸧 ࡢ2ಶయࡢቑᖜDNA᩿∦ࡢሷᇶ㓄ิࢆḟ♧ࡍ᪉ἲ࡛ỴᐃࡋࡓࠋୖグPCR᮲
௳࡚㔞30μLࡢᛂᾮ୰࡛ቑᖜࢆ⾜ࡗࡓᚋ㸪MinElute PCR purification kit㸦࢟
ࢤࣥ㸧ࢆ⏝࠸࡚⢭〇ࡋࡓࡶࡢࢆ㗪ᆺࡋ㸪PCR࡛⏝࠸ࡓࣉ࣐࣮ࣛࢆ⏝ࡋ
࡚BigDye® Terminator v3.1 Cycle Sequencing Kit 㸦ࣉࣛࢻࣂ࢜ࢩࢫࢸ࣒ࢬ㸧
ࡼࡾࢧࢡࣝࢩ࣮ࢡ࢚ࣥࢫᛂࢆ⾜ࡗࡓࠋሷᇶ㓄ิࡢỴᐃࡣ ABI PRISM®
3100 ࢪ࢙ࢿࢸࢵࢡࢼࣛࢨ࣮㸦ࣉࣛࢻࣂ࢜ࢩࢫࢸ࣒ࢬ㸧ࢆ⏝࠸ࡓࠋ
ᚓࡽࢀࡓሷᇶ㓄ิࡣᅜ㝿ሷᇶ㓄ิࢹ࣮ࢱ࣮࣋ࢫ࠾࠸࡚ᥦ౪ࡉࢀࡿ BLAST
ࡼࡾ㢮ఝ㓄ิࡢ᳨⣴ࢆ⾜ࡗࡓࠋ
⤖
⤖ᯝ
DNAࢆᢳฟࡋࡓࡍ࡚ࡢಶయ࠾࠸࡚㸪D3-4046F࠾ࡼࡧD3-4413Rࢆ⏝࠸ࡓ PCR ࡼࡗ࡚ቑᖜ⏘≀ࡀ☜ㄆ࡛ࡁࡓࠋࡇࡢࡇࡽ㸪DNA ࡣᢳฟ࡛ࡁࡓ⪃
࠼ࡽࢀࡓࠋ࢜ࣥࢩࢶࢶࣖࢥࣂࢳࡢ235bp ࡢ㡿ᇦࢆቑᖜࡉࡏࡿࡼ࠺タィࡋࡓࣉ
࣐࣮ࣛ㸦EfF࠾ࡼࡧEfR㸹Fig. 26㸪Table 10㸧ࡼࡿPCR ࡢ⤖ᯝ㸪࢜ࣥࢩࢶࢶ
ࣖࢥࣂࢳࡽࡣᮇᚅ㏻ࡾ⣙250bpࡢቑᖜ᩿∦ࡀ☜ㄆࡉࢀࡓࡀ㸪ࢳࢳࣗ࢘࢝ࢶ
ࣖࢥࣂࢳ㸪ࢧࣂࢡࢶࣖࢥࣂࢳ࠾ࡼࡧࢥࢼࢪ࣑ࣛ㢮࡛ࡣࡃቑᖜࡀ☜ㄆ࡛ࡁ࡞
ࡗࡓࠋࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳࡢ527bp ࢆቑᖜࡍࡿࡼ࠺タィࡋࡓࣉ࣐࣮ࣛ
85
㸦EmF࠾ࡼࡧEmR㸹Fig. 26㸪Table 10㸧ࡼࡿPCRࡼࡗ࡚㸪ࢳࢳࣗ࢘࢝ࢶ
ࣖࢥࣂࢳ࠾ࡼࡧࢧࣂࢡࢶࣖࢥࣂࢳࡢ୧✀࡛⣙500bpࡢPCR⏘≀ࡢቑᖜࡀ☜ㄆ࡛
ࡁ㸪ࡲࡓቑᖜ᩿∦㛗ࡣࢧࣂࢡࢶࣖࢥࣂࢳࡢ࠺ࡀࡸࡸ㛗ࡗࡓࠋ୍᪉㸪࢜ࣥࢩ ࢶࢶࣖࢥࣂࢳ࠾ࡼࡧࢥࢼࢪ࣑ࣛ㢮࡛ࡣቑᖜ⏘≀ࡣᚓࡽࢀ࡞ࡗࡓࠋ
࢜ࣥࢩࢶࢶࣖࢥࣂࢳ࡛ࡣ EfF EfR ࠾ࡼࡧ EfF EmR ࡼࡗ࡚ᚓࡽࢀࡓ 2
✀㢮ࡢቑᖜDNA᩿∦ࢆ㸪ࢧࣂࢡࢶࣖࢥࣂࢳࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳ࡛ࡣEmF
EfRࡼࡗ࡚ᚓࡽࢀࡓ᩿∦ࡢሷᇶ㓄ิࡢゎᯒࢆ⾜ࡗࡓࠋࡑࡢ⤖ᯝ㸪࢜ࣥࢩࢶ ࢶࣖࢥࣂࢳࡽᚓࡽࢀࡓ㓄ิ㸦AB662969–AB662972㸧ࡣ㸪ࡶ᪤▱ࡢ࢜ࣥࢩ ࢶࢶࣖࢥࣂࢳࡢ㓄ิ㸦AY615782ࡶࡋࡃࡣAY615783㸧99%ࡽ100%୍⮴ࡋ ࡓࠋࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳࡢ㓄ิ㸦AB662975㸪AB662976㸧ࡣ᪤▱ࡢࢳࢳࣗ
࢘࢝ࢶࣖࢥࣂࢳࡢ㓄ิ㸦AY878190㸧99%୍⮴ࡋࡓࠋࡲࡓ㸪ࢧࣂࢡࢶࣖࢥࣂ
ࢳࡢ㓄ิࡣࡇࢀࡲ࡛ᅜ㝿ሷᇶ㓄ิࢹ࣮ࢱ࣮࣋ࢫⓏ㘓ࡉࢀ࡚࠸࡞ࡗࡓࡀ㸪
ᅇᚓࡽࢀࡓ㓄ิ㸦AB662975㸪AB662976㸧ࡣEr. warrae Naumann and Schmidt 98%୍⮴ࡋࡓࠋࡇࡢࡇࡽ㸪ᚓࡽࢀࡓቑᖜDNA᩿∦ࡣࣉ࣐࣮ࣛࢆタィࡋࡓ ITS1㡿ᇦࡢࡶࡢ࡛࠶ࡿ☜ㄆ࡛ࡁࡓࠋ
タィࡋࡓ4ࡘࡢࣉ࣐࣮ࣛࡢ࠺ࡕ3ࡘ㸦EfF㸪EmF࠾ࡼࡧ EmR㸧ࢆ⏝࠸ࡓቑ ᖜ࡛ࡣ㸪3✀ࢶࣖࢥࣂࢳ࡚ࡽቑᖜ᩿∦ࡀᚓࡽࢀࡓ㸦Fig. 27㸧ࠋࡲࡓ㸪3✀㛫
࡛ቑᖜ᩿∦㛗ࡀ␗࡞ࡾ㸦࢜ࣥࢩࢶࢶࣖࢥࣂࢳ⣙300ࠥ400bp㸪ࢧࣂࢡࢶࣖࢥࣂࢳ
⣙600ࠥ700bp㸪ࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳ⣙500bp㸧㸪ศᏊ㔞࣐࣮࣮࢝ࡋ࡚100bp
ࣛࢲ࣮࣐࣮࣮࢝ࢆ⏝࠸ࡿࡇࡼࡾ㸪᫂☜༊ูྍ⬟࡛࠶ࡗࡓࠋࡲࡓ㸪ࢥࢼࢪ
࣑ࣛ㢮ࡽࡣቑᖜ᩿∦ࡀ☜ㄆࡉࢀ࡞ࡗࡓ㸦Fig. 27㸧ࠋ
⪃
⪃ᐹ
rDNA 㡿ᇦࡣሷᇶ㓄ิࡢኚ␗ࡀᑡ࡞ࡃሷᇶ㓄ิࡀࡼࡃಖᏑࡉࢀ࡚࠸ࡿ㡿ᇦ
ኚ␗ࡀከࡃከᆺ᳨ฟࡀᐜ᫆࡞㡿ᇦࡀྵࡲࢀ࡚࠸ࡿࡇࡽ㸪᪻ࡢ✀㛫࣭✀
86
M 1 2 3 4 5 6 7 8
Fig. 27. Electrophoregram of amplified fragments for three parasitoid wasps of whitefly by multiplex polymerase chain reaction using three primers EfF, EmF and EmR. Lanes are: M: 100bp ladder molecular weight marker; 1-2: Encarsia formosa; 3-4: Eretmocerus eremicus; 5-6:
Eretmocerus mundus; 7: Trialeurodes vaporariorum; 8: no DNA template.
87
ෆኚ␗ࡢゎᯒࡼࡃ⏝࠸ࡽࢀ࡚࠸ࡿ㸦Ardeh et al.㸪2005㸹Heraty et al.㸪2007㸹 Ji et al. 2003㸹Vickerman et al.㸪2004㸹Yara and Kunimi㸪2009㸧ࠋᮏ◊✲࡛ࡣ㸪ࡇ ࡢ࠺ࡕኚ␗ࡢከ࠸ࡉࢀࡿInternal Transcribed Spacer 1㸦ITS1㸧㡿ᇦࡢሷᇶ㓄ิ
ࢆゎᯒࡍࡿࡇࡼࡗ࡚㸪✀㛫ኚ␗ࢆ᳨ฟࡍࡿࡇึࡵ࡚ᡂຌࡋࡓࠋᮏἲ
ࡼࡾ㸪࢜ࣥࢩࢶࢶࣖࢥࣂࢳ㸪ࢧࣂࢡࢶࣖࢥࣂࢳ࠾ࡼࡧࢳࢳࣗ࢘࢝ࢳࣖࢥࣂࢳ
ࢆᐜ᫆㆑ูࡍࡿࡇࡀྍ⬟࡞ࡗࡓࠋᮏ◊✲࡛ࡣᡃࡀᅜ࡛⏕≀㎰⸆ࡋ࡚Ⓩ
㘓ࡢ࠶ࡿ3✀ࡢࡳࢆᑐ㇟ࡋࡓࡀ㸪ᅜෆ࡛ࡣࢥࢼࢪ࣑ࣛ㢮ࡢᅵ╔ᐤ⏕⻏ࡀከࡃ
▱ࡽࢀ࡚࠾ࡾ㸦Ლ⏣㸪1980㸹Kajita㸪2000㸹ᑠ⃝ࡽ㸪1992㸧㸪ᅵ╔ኳᩛࢆ⏝ࡋ ࡓᐖ㜵㝖ࡀồࡵࡽࢀ࡚࠸ࡿࠋᚋࡣ㸪ᮏ᪉ἲࢆࡼࡾⓎᒎࡉࡏ㸪ᅵ╔ᐤ⏕⻏ࡶ
ྵࡵࡓ㆑ูἲࡢ㛤Ⓨࡀᮇᚅࡉࢀࡿࠋ
ᾏእ࡛ࡣ࣑ࢺࢥࣥࢻࣜDNAࡢPCR-RFLPࡼࡾ Encarsiaᒓࡢ㆑ูἲࡀ㛤
Ⓨࡉࢀ࡚࠸ࡿࡀ㸦Monti et al.㸪2005㸧㸪ࡇࡢᡭἲࡣไ㝈㓝⣲ฎ⌮ࡢࡓࡵ㛫
ࢥࢫࢺࡀࡾ㸪ษ᩿㠃ࡀ༑ศ࡞ሙྜࡣゎᯒࡀᅔ㞴࡞ࡿࡇࡀ࠶ࡿࠋ୍
᪉㸪ᮏ◊✲࡛㛤Ⓨࡋࡓ࣐ࣝࢳࣉࣞࢵࢡࢫPCR ἲࡣ 1ᅇࡢPCR ᛂ࡛3 ✀ࢆྠ
㆑ู࡛ࡁࡿࡇࡽMonti et al.㸪㸦2005㸧ࡢᡭἲࡼࡾඃࢀ࡚࠸ࡿࠋ
88
➨
➨㸲⠇ ẅᚤ⏕≀㎰⸆ࡢᙳ㡪
ᮦᩱ࠾ࡼࡧ᪉ἲ
㸯㸬ᐤ⏕⻏ᑐࡍࡿẅࡢᙳ㡪 㸦㸯㸧ẅ
ࢺ࣐ࢺⓏ㘓ࡢ࠶ࡿẅࡢ࠺ࡕ24ࢆ౪ヨࡋࡓ㸦Table 11, 12㸧ࠋ
㸦㸰㸧ᐤ⏕⻏
⏕≀㎰⸆ࡋ࡚㈍ࡉࢀ࡚࠸ࡿࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳ㸦࣑࣋ࣃ࣮ࣝ®㸧㸪ࢧ
ࣂࢡࢶࣖࢥࣂࢳ㸦࢚࣮ࣝ࢝ࣝ®㸧࠾ࡼࡧ࢜ࣥࢩࢶࢶࣖࢥࣂࢳ㸦࢚ࣥࢫࢺࣜࢵࣉ®㸧 ࡢ࣐࣑࣮࣮࢝ࢻࢆ౪ヨࡋࡓࠋ࣐࣑࣮࣮࢝ࢻࡣヨ㦂ࡲ࡛෭ⶶᗜ㸦8Υ㸧ෆ࡛ಖ⟶ࡋ ࡓࠋᡂࡣ࣐࣑࣮࣮࢝ࢻࢆᜏ ᐊෆ㸦25Υ㸪16L8D㸧⛣ືࡋ㸪⩚ࡋࡓ2㹼3
᪥㱋ࢆ౪ヨࡋࡓࠋ
㸦㸱㸧ᐤ⏕⻏ࡢ࣐࣑࣮ᑐࡍࡿẅࡢᙳ㡪
ᯇ㸦2006㸧ࡢ᪉ἲ‽ࡌ࡚ㄪᰝࡋࡓࠋࡍ࡞ࢃࡕ㸪ྛẅࢆᖖ⏝⃰ᗘ㸦400 㹼5000 ಸ㸧␃Ỉ࡛ᕼ㔘ࡋ㸪࣐࣑࣮࣮࢝ࢻ㸦⻈ᩘ80㹼100 ⛬ᗘ㸧ࢆ10 ⛊㛫 ᾐₕࡋࡓࠋྛᐤ⏕⻏ࡶ3ᯛࡢ࣐࣑࣮࣮࢝ࢻࢆ౪ヨࡋࡓࠋᑐ↷ࡣ␃Ỉࡋࡓࠋ
࣐࣑࣮࣮࢝ࢻࡣᾐₕᚋ㸪࣮࣌ࣃ࣮ࢱ࢜ࣝࡢୖ࡛⇱ࡋ㸪ࢩ࣮ࣕࣞ㸦┤ᚄ 9 cm㸪
῝ࡉ2 cm㸧ධࢀ㸪ᜏ ჾෆ㸦25Υ㸪16L8D㸧2㐌㛫㟼⨨ࡋࡓࠋ⩚ᩘ㠀
⩚ᩘࢆ║ᐇయ㢧ᚤ㙾㸦SHZ-U㸸ࢽࢥࣥ㸧ୗ࡛ᩘ࠼㸪⸆ࡈAbbott㸦1925㸧 ࡢ⿵ṇᘧ࡛Ṛ⋡ࢆィ⟬ࡋ㸪ḟࡢIOBCࡢホ౯ᇶ‽㸦Hassan㸪1992㸹Amano and
Haseeb㸪2001㸧ᇶ࡙ࡁ 4ẁ㝵ศࡅࡓ㸹ࣞ࣋ࣝ 1㸦Ṛஸ⋡ 30%௨ୗ㸪ᙳ㡪ࡀ
࡞࠸㸧㸪ࣞ࣋ࣝ2㸦30㹼79%㸪ഹᙳ㡪ࡀ࠶ࡿ㸧㸪ࣞ࣋ࣝ3㸦80㹼99%㸪୰⛬ᗘ
ᙳ㡪ࡀ࠶ࡿ㸧㸪ࣞ࣋ࣝ4㸦99%௨ୖ㸪㠀ᖖᙳ㡪ࡀ࠶ࡿ㸧ࠋ
89
Mortalityb)Evalution CategorriesC)Mortalityb)Evalution CategorriesC)Mortalityb)Evalution CategorriesC) Neonicotinoids Acetamiprid20S.P.×2,00099.3487.931004 Clothianidin16S.P.×2,000100496.1399.64 Dinotefuran20S.G.×3,00098.5379.7299.14 Imidacloprid10W.P.×2,00090.1353.0261.02 Nitenpyram10S.P.×1,00095.5355.1292.23 Thiacloprid30W.D.G.×4,00092.7362.2276.82 Synthetic pyrethroids Etofenprox20E.C.×1,00025.0133.7298.23 Permethrin20E.C.×2,00053.5254.3295.73 Organophosphates Acephate50W.P.×1,00052.0215.4147.92 Fenitrothion15E.C.×2,00093.7393.1397.43 Insect growth regulators Buprofezin40W.P.×1,00023.3113.8101 Cyromazine8.3S.L.×1,00010.1113.911.71 Flufenoxuron10E.C.×2,00018.4120.1117.61 Lufenuron5E.C.×3,000010110.81 Acaricides Chinomethionat25W.P.×1,50023.019.6113.11 Milbemectin1E.C.×1,50072.0267.4242.52 Pyridaben20F.L.×1,000100489.2390.93 Otheres Bt10W.D.G.×1,000013.816.91 Chlorphenapyr10F.L.×2,00051.8223.513.21 Emamectin-benzoate1E.C.×2,00086.7349.8246.82 Pymetrozine25W.P.×3,00023.4120.0115.61 Spinosad25W.D.G.×5,000100497.5399.14 Tolfenpyrad15E.C.×1,000100410041004 Sulfur52F.L.×40013.2114.6120.41 C) Evalution categories:The results of expressed as safe(1), slightly harmful(2), moderately harmful(3), harmful(4) whereby 1=<30% mortality, 2=30-79% mortality, 3=80-98% mortality, 4=99-100% mortality.
b) Corrected mortality accrding to Abbott's formula based on check value
a)S.P.:water soluble powder,W.P.:wettable powder,W.D.G.:water dispension concentrate,S.G.:soluble granule,E.C.:emulsifiable concentrate,S.L.:soluble concentrate,F.L.:flowable
Table 11. The side effects of pesticides on pupae of parasitoids in pupae of whitefly by direct dipping method Encarsia Formosa pesticidesFormulation (%a.i. & type)a)Dilution
Eretmocerus mundusEretmocerus eremicus
90
Mortalityb)Evalution CategorriesC)Mortalityb)Evalution CategorriesC)Mortalityb)Evalution CategorriesC) Neonicotinoids Acetamiprid20S.P.×2,000100410041004 Clothianidin16S.P.×2,000100410041004 Dinotefuran20S.G.×3,000100410041004 Imidacloprid10W.P.×2,000100410041004 Nitenpyram10S.P.×1,000100410041004 Thiacloprid30W.D.G.×4,00037.3298.3385.33 Synthetic pyrethroids Etofenprox20E.C.×1,000100410041004 Permethrin20E.C.×2,000㸫㸫10041004 Organophosphates Acephate50W.P.×1,000100410041004 Fenitrothion15E.C.×2,000100410041004 Insect growth regulators Buprofezin40W.P.×1,00025.413211.21 Cyromazine8.3S.L.×1,00019151.324.41 Flufenoxuron10E.C.×2,000010101 Lufenuron5E.C.×3,000018.8101 Acaricides Chinomethionat25W.P.×1,50095.6310041004 Milbemectin1.0E.C.×1,500㸫㸫92.631004 Pyridaben20F.L.×1,000100498.3398.53 Otheres Bt10W.D.G.×1,000㸫㸫014.91 Chlorphenapyr10F.L.×2,0001004㸫㸫㸫㸫 Emamectin-benzoate1.0E.C.×2,000100410041004 Pymetrozine25W.P.×3,00011.7114.8101 Spinosad25W.D.G.×5,000100410041004 Tolfenpyrad15E.C.×1,000100410041004 Sulfur52F.L.×400㸫㸫28.311.61
Dilution a)S.P.:water soluble powder,W.P.:wettable powder,W.D.G.:water dispension concentrate,S.G.:soluble granule,E.C.:emulsifiable concentrate,S.L.:soluble concentrate,F.L.:flowable b)Corrected mortality accrding to Abbott's formula based on check value C)Evalution categories:The results of expressed as safe(1), slightly harmful(2), moderately harmful(3), harmful(4) whereby 1=<30% mortality, 2=30-79% mortality, 3=80-98% mortality, 4=99-100% mortality.
Formulation (%a.i. & type)a)pesticidesEretmocerus mundusTable 12. The side effects of pesticides on adults of whitefly parasitoids using residual film method. Eretmocerus eremicusEncarsia Formosa
91
㸦㸲㸧ᐤ⏕⻏ࡢᡂᑐࡍࡿẅࡢᙳ㡪
ヨ㦂ࡣⷧ⭷ἲ㸦ᑠ⃝ࡽ㸪1998㸹ᯇ㸪2006㸧࡛⾜ࡗࡓࠋࡍ࡞ࢃࡕ㸪ྛẅ
ࢆࢭࢺࣥ㸦99.5%㸧⁐ࡋᖖ⃰ᗘࡋ㸪ࡑࡢ0.1 mlࢆ⟶⎼㸦ཱྀෆᚄ5.4mm㸪
⬗ᚄ11.7 mm㸪㧗ࡉ35 mm㸧ᢞධࡋࡓᚋ㸪⟶⎼ࢆỈᖹᅇ㌿ࡉࡏ࡚ෆ㠃ⷧ
⭷ࢆసࡗࡓࠋࡑࡢᚋ㸪⟶⎼ࡢᗏ5%ࡢ⻏⻤ࢆᰁࡳࡇࡲࡏࡓࢁ⣬㸦5 mm 5 mm㸧
ࢆ㣵ࡋ࡚ධࢀࡓࠋ⟶⎼࠶ࡓࡾ10㢌ࡢᐤ⏕⻏ᡂࢆධࢀ㸪⟶⎼ࡢཱྀࢆࢼࣟࣥ
ࢦ࣮ࢫ࡛そ࠸㸪ᜏ ᐊෆ㸦25Υ㸪16L8D㸧㟼⨨ࡋࡓࠋẅࡈ⟶⎼ࢆ 6 ᮏ ࡎࡘ౪ヨࡋࡓࠋฎ⌮ 24 㛫ᚋ㸪⏕Ꮡᩘ㸪Ṛஸᩘࢆ║ᐇయ㢧ᚤ㙾 㸦SHZ-U㸸ࢽࢥࣥ㸧ୗ࡛ᩘ࠼㸪Abbott㸦1925㸧ࡢ⿵ṇᘧ࡛Ṛ⋡ࢆィ⟬ࡋࡓࠋ
ୖグࡢIOBCࡢホ౯ᇶ‽ᇶ࡙ࡁ4ẁ㝵ศࡅࡓࠋ
㸰㸬ᐤ⏕⻏ᑐࡍࡿᚤ⏕≀㎰⸆ࡢᙳ㡪 㸦㸯㸧ᚤ⏕≀㎰⸆
3✀㢮ࡢᚤ⏕≀㎰⸆㸦࣐ࢥࢱ࣮ࣝ®㸪ࣂ࣮ࢱࣞࢵࢡ®࠾ࡼࡧ࣎ࢱࢽ࣮࢞ࢻES®㸧
ࢆ౪ヨࡋࡓࠋ᭷ຠᡂศ㔞ࡣࡑࢀࡒࢀ㸪3.0 × 109 spores / g㸪5.0 × 108 spores / g㸪
࠾ࡼࡧ 2.1 × 1010 spores / ml࡛࠶ࡿࠋྛᚤ⏕≀㎰⸆ࡣつᐃࡢ⃰ᗘ␃Ỉ࡛ᕼ㔘 ࡋ࡚⏝ࡋࡓ㸦࣐ࢥࢱ࣮ࣝ®࠾ࡼࡧࣂ࣮ࢱࣞࢵࢡ®ࡣ 1000 ಸ, ࣎ࢱࢽ࣮࢞ࢻ ES®ࡣ500ಸ㸧ࠋ
㸦㸰㸧ᐤ⏕⻏
࢜ࣥࢩࢶࢶࣖࢥࣂࢳ㸦࢚ࣥࢫࢺࣜࢵࣉ®㸧ࢧࣂࢡࢶࣖࢥࣂࢳ㸦࢚࣮ࣝ࢝ࢻ®㸧 ࡢ࣐࣑࣮࣮࢝ࢻ㸦࣮࢝ࢻᙜࡓࡾࡢ࣐࣑࣮ᩘ80㹼100 㢌㸧ࢆ෭ⶶᗜෆ㸦8Υ㸧࡛ᩘ
᪥㛫ಖ⟶ࡋ⏝ࡋࡓࠋ
92
㸦㸱㸧ᐤ⏕⻏ࡢ࣐࣑࣮ᑐࡍࡿᚤ⏕≀㎰⸆ࡢᙳ㡪
࣐࣑࣮࣮࢝ࢻࢆྛᚤ⏕≀㎰⸆ࡢ⁐ᾮ⣙10⛊㛫ᾐₕࡋ㸪వศ࡞Ỉࢆᡶ࠸ⴠ
ࡋࡓᚋ㸪ࢩ࣮ࣕࣞ㸦┤ᚄ9 cm㸪῝ࡉ2 cm㸧ෆ㟼⨨ࡋࡓࠋᑐ↷༊ࡋ࡚␃Ỉ
ࢆ⏝࠸ࡓࠋ1 1ᅇࡘࡁ 1ᯛࡢྠ୍ࡢ⟽ࡢ࣐࣑࣮࣮࢝ࢻࢆ⏝ࡋࡓࠋࢩ࣮ࣕ
ࣞࡣࣃࣛࣇ࣒࡛ࣝᐦᑒࡋ㸪6 ᪥㛫ᜏ ჾෆ㸦25°C㸪16L8D㸧㟼⨨ࡋࡓࠋ⩚
ᩘ㠀⩚ᩘࢆ║ᐇែ㢧ᚤ㙾㸦SHZ-U㸸ࢽࢥࣥ㸧ୗ࡛ᩘ࠼㸪Ṛஸ⋡ࢆ⟬ฟ ࡋࡓࠋྛฎ⌮ࡶ3࡛⾜ࡗࡓࠋ
㸦㸲㸧ᐤ⏕⻏ࡢᡂᑐࡍࡿᚤ⏕≀㎰⸆ࡢᙳ㡪
ࢁ⣬㸦┤ᚄ9 cm㸧ࢆྛᚤ⏕≀㎰⸆ࡢ⁐ᾮ⣙ 10⛊㛫ᾐₕࡋ㸪వศ࡞Ỉࢆᡶ
࠸ⴠࡋࡓᚋࢩ࣮ࣕࣞෆᩜ࠸ࡓࠋ࣐࣑࣮࣮࢝ࢻ1ᯛࢆࢁ⣬ࡢୖ⨨ࡁࣃࣛࣇ
࣒࡛ࣝࢩ࣮ࣕࣞࡢ⦕ࢆᐦᑒࡋ㸪7᪥㛫ᜏ ჾෆ㸦25°C㸪16L8D㸧㟼⨨ࡋࡓࠋ ᑐ↷༊ࡋ࡚␃Ỉࢆ⏝࠸ࡓࠋ⩚ᩘ㠀⩚ᩘ㸪Ṛஸᡂᩘࢆ║ᐇែ㢧ᚤ
㙾㸦SHZ-U㸸ࢽࢥࣥ㸧ୗ࡛ᩘ࠼ࡓࠋṚஸᡂࡢయୖࡢ⣒≧⳦ࡢ᭷↓ࢆ☜ㄆࡋࡓࠋ
࣐࣑࣮࣮࢝ࢻࡣྠ୍ࡢ⟽ࡢࡶࡢࢆ⏝ࡋ㸪ྛฎ⌮ࡶ3࡛⾜ࡗࡓࠋ
㸦㸳㸧⤫ィศᯒ
ᐤ⏕⻏ࡢ⩚⋡Ṛஸ⋡ࡣࡈゅኚࡋ㸪ᐤ⏕⻏ࡢ✀ᚤ⏕≀㎰⸆ࡢ
✀㢮ࡢ㸰せᅉࡢศᩓศᯒ㸦2 ඖ㓄⨨ศᩓศᯒ㸧ࢆ⾜ࡗࡓࠋస⏝ࡀㄆࡵࡽࢀ
ࡓሙྜࡣ㸪ከ㔜ẚ㍑᳨ᐃ㸦Tukey-Kramer test㸧ࢆ⾜࠸㸪స⏝ࡀㄆࡵࡽࢀ࡞
ࡗࡓሙྜࡣྛせᅉࡢỈ‽㛫ࡢከ㔜ẚ㍑㸦Tukey-Kramer test ࠾ࡼࡧ Student’s
t test㸧ࢆ⾜ࡗࡓࠋ࡞࠾㸪ࢹ࣮ࢱࡢゎᯒࡣ࢚ࢡࢭࣝ⤫ィ2012㸦SSRI㸧ࡲࡓࡣJMP5
㸦SAS Institute, 2002㸧ࢆ⏝࠸ࡓࠋ
93
⤖
⤖ᯝ
㸯㸬ᐤ⏕⻏ࡢ࣐࣑࣮ᑐࡍࡿẅࡢᙳ㡪
ẅᾐₕฎ⌮ࡋࡓ 3 ✀ࡢᐤ⏕⻏࣐࣑࣮ࡢṚஸ⋡ࢆ Table 11 ♧ࡋࡓࠋ IGRࡢ4㸦ࣈࣉࣟࣇ࢙ࢪࣥ㸪ࢩ࣐ࣟࢪࣥ㸪ࣇࣝࣇ࢙ࣀࢡࢫࣟࣥ࠾ࡼ
ࡧࣝࣇ࢙ࢾࣟࣥ㸧㸪ࢳࣀ࣓ࢳ࢜ࢿ࣮ࢺ㸪Bt 㸪ࣆ࣓ࢺࣟࢪࣥ࠾ࡼࡧ࢜
࢘ࡣ3✀ᐤ⏕⻏ᑐࡋ࡚࡚ࣞ࣋ࣝ 1࡛࠶ࡗࡓࠋ࢚ࢺࣇ࢙ࣥࣉࣟࢵࢡࢫ㸪
ࢭࣇ࢙࣮ࢺ࠾ࡼࡧࢡࣟࣝࣇ࢙ࢼࣆࣝࡣ3✀ࡢᐤ⏕⻏ࡢෆ࠸ࡎࢀᑐࡋ
࡚ࣞ࣋ࣝ㸯࡛࠶ࡗࡓࠋࢿ࢜ࢽࢥࢳࣀࢻ⣔ࡢࢭࢱ࣑ࣉࣜࢻ㸪ࢡࣟࢳࢽ ࢪࣥ㸪ࢪࣀࢸࣇࣛࣥ㸪ࢲࢽࡢࣆࣜࢲ࣋ࣥ㸪ࡑࡢࡢ⣔⤫ࡢࢫࣆࣀࢧࢻ
࠾ࡼࡧࢺࣝࣇ࢙ࣥࣆࣛࢻࡣ 3 ✀ᐤ⏕⻏ࡢෆ࠸ࡎࢀᑐࡋ࡚ࣞ࣋ࣝ 4 ࡛
࠶ࡗࡓࠋ
㸰㸬ᐤ⏕⻏ࡢᡂᑐࡍࡿẅࡢᙳ㡪
ẅࡢⷧ⭷ฎ⌮ࡋࡓ3✀ࡢᐤ⏕⻏ᡂࡢṚஸ⋡ࢆTable 12 ♧ࡋࡓࠋIGR
ࡢ3㸦ࣈࣉࣟࣇ࢙ࢪࣥ㸪ࣇࣝࣇ࢙ࣀࢡࢫࣟࣥ࠾ࡼࡧࣝࣇ࢙ࢾࣟࣥ㸧
࠾ࡼࡧࣆ࣓ࢺࣟࢪࣥࡣ3✀ᐤ⏕⻏ᑐࡋ࡚࡚ࣞ࣋ࣝ 1࡛࠶ࡗࡓࠋBt࠾ࡼ
ࡧ࢜࢘ࡣࢧࣂࢡࢶࣖࢥࣂࢳ࢜ࣥࢩࢶࢶࣖࢥࣂࢳᑐࡋ࡚ࣞ࣋ࣝ 1 ࡛
࠶ࡗࡓࠋIGR ࡢࢩ࣐ࣟࢪࣥࡣࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳ࢜ࣥࢩࢶࢶࣖࢥࣂ
ࢳᑐࡋ࡚ࣞ࣋ࣝ1࡛࠶ࡗࡓࡀ㸪ࢧࣂࢡࢶࣖࢥࣂࢳᑐࡋ࡚ࣞ࣋ࣝ2࡛࠶ࡗࡓࠋ
ࢿ࢜ࢽࢥࢳࣀࢻ⣔㸪ྜᡂࣆࣞࢫࣟࢻ⣔㸪᭷ᶵࣜࣥ⣔㸪ࢲࢽ࠾ࡼࡧ ࡑࡢࡢ⣔⤫㸦ࢡࣟࣝࣇ࢙ࢼࣆࣝ㸪࢚࣐࣓ࢡࢳࣥᏳᜥ㤶㓟ሷ㸪ࢫࣆࣀࢧ ࢻ࠾ࡼࡧࢺࣝࣇ࢙ࣥࣆࣛࢻ㸧ࡣ࠸ࡎࢀࡶ3✀ᐤ⏕⻏ᑐࡋ࡚ࣞ࣋ࣝ3ࡲࡓࡣ
ࣞ࣋ࣝ4࡛࠶ࡗࡓࠋ
94
㸱㸬ᐤ⏕⻏ࡢ࣐࣑࣮ᑐࡍࡿᚤ⏕≀㎰⸆ࡢᙳ㡪
ᚤ⏕≀㎰⸆ᾐₕฎ⌮ࡋࡓᐤ⏕⻏࣐࣑࣮ࡢ⩚⋡ࢆTable 13♧ࡋࡓࠋ࣐࣑
࣮⾲㠃⳦⣒ࡀほᐹࡉࢀࡓ㸦Fig. 28㸧ࠋᐤ⏕⻏ࡢ⩚⋡ᑐࡍࡿ2せᅉࡢศᩓศ ᯒࡢ⤖ᯝ㸪ᐤ⏕⻏ࡢ㛫࡛᭷ព࡞ᕪࡣ࡞ࡗࡓࡀ㸦F1, 16 = 1.580㸪p = 0.227㸧㸪ᚤ
⏕≀㎰⸆㛫࡛ࡣ᭷ព࡞ᕪࡀ࠶ࡗࡓ㸦F3, 16 = 73.382㸪p < 0.0001㸧ࠋᐤ⏕⻏ᚤ⏕
≀㎰⸆㛫ࡢ┦స⏝ࡣㄆࡵࡽࢀ࡞ࡗࡓ㸦F3, 16 = 0.514㸪p = 0.678㸧ࠋ࣐ࢥࢱ
࣮ࣝ®ࡢ⩚⋡㸦40.5㹼40.6%㸧ࣂ࣮ࢱࣞࢵࢡ®ࡢ⩚⋡㸦55.1㹼56.2%㸧࠾ࡼ
ࡧᑐ↷㸦19.3~60.4%㸧ࡣ᭷ព࡞ᕪࡀ࡞ࡗࡓ㸦Tukey–Kramer HSD test㸪p < 0.05㸧ࠋ
࣎ࢱࢽ࣮࢞ࢻES® ࡢ⩚⋡ࡣ࣐ࢥࢱ࣮ࣝ®㸪ࣂ࣮ࢱࣞࢵࢡ®࠾ࡼࡧᑐ↷ẚ
᭷ពపࡗࡓ㸦p < 0.05㸧ࠋ
㸲㸬ᐤ⏕⻏ࡢᡂᑐࡍࡿᚤ⏕≀㎰⸆ࡢᙳ㡪
ᚤ⏕≀㎰⸆ࡢᐤ⏕⻏ࡢᡂᑐࡍࡿṚஸ⋡ࢆTable 14♧ࡋࡓࠋṚஸࡋࡓᡂ
ࡢయୖ⳦⣒ࡀほᐹࡉࢀࡓ㸦Fig. 29㸧ࠋᐤ⏕⻏ࡢ⩚⋡ࡘ࠸࡚2せᅉࡢศᩓ ศᯒࡢ⤖ᯝ㸪ᐤ⏕⻏ࡢ⩚⋡ࡣ✀ࡼࡾ᭷ព␗࡞ࡗࡓࡀ㸦F1,16 = 4.84㸪p = 0.04㸧㸪 ᚤ⏕≀㎰⸆㛫࡛ࡣ᭷ព࡞ᕪࡣ࡞ࡗࡓ㸦F3,16 = 0.31㸪p = 0.82㸧ࠋࡲࡓ㸪Ṛஸ⋡
ᐤ⏕⻏㛫࡛ࡣ᭷ព࡞ᕪࡣ࡞ࡗࡓࡀ㸦F1,16 = 1.69㸪p = 0.21㸧㸪ᚤ⏕≀㎰⸆㛫࡛ࡣ
᭷ព࡞ᕪࡀㄆࡵࡽࢀࡓ㸦F3,16 = 33.68㸪p < 0.01㸧ࠋࡲࡓ㸪ᐤ⏕⻏ᚤ⏕≀㎰⸆㛫
࡛స⏝ࡀㄆࡵࡽࢀࡓࡓࡵ㸦F3,16 = 4.07, p = 0.03㸧㸪ᐤ⏕⻏ᚤ⏕≀㎰⸆ࡢ
࡚ࡢ⤌ࡳྜࢃࡏࡼࡿከ㔜ẚ㍑᳨ᐃࢆ⾜ࡗࡓࡇࢁ㸪࣐ࢥࢱ࣮ࣝ®ࡢ࢜ࣥࢩ ࢶࢶࣖࢥࣂࢳࢧࣂࢡࢶࣖࢥࣂࢳࡢṚஸ⋡㸦ࡑࢀࡒࢀ 75.8%㸪89.3%㸧㸪࣎ࢱࢽ
࣮࢞ࢻ ES®ࢆฎ⌮ࡋࡓࡁࡢ࢜ࣥࢩࢶࢶࣖࢥࣂࢳࢧࣂࢡࢶࣖࢥࣂࢳࡢṚஸ⋡
㸦ࡑࢀࡒࢀ90.3 %㸪79.0 %㸧ࡣࣂ࣮ࢱࣞࢵࢡ®ࢆฎ⌮ࡋࡓ࢜ࣥࢩࢶࢶࣖࢥࣂࢳࡢ Ṛஸ⋡㸦29.2%㸧ᑐ↷ࡢṚஸ⋡㸦30.5㸣㸧ẚ᭷ព㧗ࡗࡓ㸦Tukey-Kramer HSD test㸪p < 0.05㸧ࠋ
95
Prasitoid speciesProduct (Entomopathogenic fungus)No. of parasitoids mummy testedNo. of emerged parasitoidsEmergence rate (%) Mycotal® (L. muscarium)81.7 ± 6.132.7 ± 2.040.6 ± 4.6 aA Encarsia formosaVertalec® (L. longisporum)87.7 ± 7.147.7 ± 6.955.1 ± 8.4 aA BotaniGard ES® (B. bassiana)87.3 ± 2.62.0 ± 0.62.3 ± 0.6 bA Water (Control)79.7 ± 6.839.0 ± 2.649.3 ± 3.5 aB Mycotal® (L. muscarium)86.0 ± 5.535.3 ± 6.940.5 ± 6.1 aA Eretmoserus eremicusVertalec® (L. longisporum)81.3 ± 4.845.3 ± 0.956.2 ± 4.0 aA BotaniGard ES® (B. bassiana)92.7 ± 8.75.3 ± 3.45.2 ± 2.9 bA Water (Control)79.0 ± 1.547.7 ± 0.760.4 ± 1.2 aA
Table 13䚷Emergence rates of aphelinid species pupae follwing treatment with mycoinsecticides The different lower case letters refers to significance of the difference between entomopathogenic fungus within same parasitoid (two-factor ANOVA follwed by Tukey Kramer HSD test, p < 0.05). The different upper case letters refers to significance of the difference between parasitoids within same entomopathogenic fungu (two-factor ANOVA follwed by Student t test, p < 0.05).
96
Fig. 28. The pupae of Eretomocerus eremicus after BotaniGardES® treatment.
97
PrasitoidProduct (Entomopathogenic fungus)No. of parasitoids mummy testedNo. of emerged parasitoidsEmergence rate (%)aNo. of infected dead parasitoidsMortality (%)b Mycotal® (L. muscarium)77.0 ± 2.335.7 ± 6.645.9 ± 7.3 aA27.0 ± 5.075.8 ± 4.6 a Encarsia formosaVertalec® (L. longisporum)79.0 ± 2.337.7 ± 4.147.9 ± 5.7 aA11.0 ± 1.529.2 ± 2.5 bc BotaniGard ES® (B. bassiana)78.7 ± 2.937.7 ± 3.547.7 ± 2.9 aA34.0 ± 3.190.3 ± 1.1 a Water (Control)76.7 ± 3.842.3 ± 5.456.1 ± 9.6 aA12.0 ± 2.630.5 ± 10.2 bc Mycotal® (L. muscarium)83.0 ± 7.639.7 ± 4.847.6 ± 2.3 aA35.3 ± 3.989.3 ± 0.9 a Eretmocerus eremicusVertalec® (L. longisporum)86.0 ± 5.233.7 ± 3.039.1 ± 2.5 aA20.7 ± 2.961.9 ± 8.1 ab BotaniGard ES® (B. bassiana)80.7 ± 6.832.7 ± 4.140.3 ± 2.1 aA26.7 ± 6.679.0 ± 11.1 a Water (Control)78.0 ± 7.130.3 ± 3.438.8 ± 1.6 aA6.3 ± 1.222.1 ± 5.7 c
Table 14䚷Emergence rates and mortalities of aphelinid species adults follwing treatment with mycoinsecticides a) The different lower case letters refers to significance of the difference between entomopathogenic fungus within same parasitoid (two-factor ANOVA follwed by Tukey Kramer HSD test, p < 0.05). The different upper case letters refers to significance of the difference between parasitoids within same entomopathogenic fungu (two-factor ANOVA follwed by Studentt test, p < 0.05). b) The combination of all parasitoids and entomopathogenic fungus were compared (Tukey Kramer HSD test, p < 0.05). There is no significant difference between the same characters.
98
Fig. 29. The adult of Eretomocerus eremicus after Myctal® treatment.
99
⪃
⪃ᐹ
Ἑྜ㸦1988㸧ᯘ㸦1996㸧ࡣ㸪ࣈࣉࣟࣇ࢙ࢪࣥ㸪Bt࠾ࡼࡧ࢜࢘ࡣ࢜
ࣥࢩࢶࢶࣖࢥࣂࢳࡢᡂ࣐࣑࣮ᙳ㡪ࡣ࡞࠸ሗ࿌ࡋ࡚࠾ࡾ㸪ᮏヨ㦂ࡶྠᵝ ࡢ⤖ᯝ࡛࠶ࡗࡓࠋࡲࡓ㸪ࡇࢀࡽࡢẅࡣࢧࣂࢡࢶࣖࢥࣂࢳࢳࢳࣗ࢘࢝ࢶ
ࣖࢥࣂࢳࡢᡂ࣐࣑࣮ᑐࡋ࡚ᙳ㡪ࡀᑠࡉ࠸ࡇࡀ᫂ࡽ࡞ࡗࡓ㸦Table 11, 12㸧ࠋࡇࢀࡲ࡛ᙳ㡪ࡀ࡛᫂࠶ࡗࡓ IGR ࡢࣇࣝࣇ࢙ࣀࢡࢫࣟࣥࣝࣇ࢙ࢾ
ࣟࣥࡣ3✀ᐤ⏕⻏ࡢᡂ࣐࣑࣮ᑐࡍࡿᙳ㡪ࡣᑠࡉ࠸ࡇࡀ᫂ࡽ࡞ࡗ
ࡓ㸦Table 11, 12㸧ࠋࢳࣀ࣓ࢳ࢜ࢿ࣮ࢺࡣ࢜ࣥࢩࢶࢶࣖࢥࣂࢳࡢᡂࢇ
ᙳ㡪ࡀ࡞࠸ሗ࿌ࡋ࡚࠸ࡿࡀ㸦Ἑྜ㸪1988㸹ᯘ㸪1996㸧㸪ᮏヨ㦂࡛ࡣᡂᑐࡋ
࡚ࡣᙳ㡪ࡀࡳࡵࡽࢀࡓ㸦Table 11, 12㸧ࠋࡲࡓ㸪ࢿ࢜ࢽࢥࢳࣀࢻ⣔㸪ྜᡂࣆ
ࣞࢫࣟࢻ⣔࠾ࡼࡧ᭷ᶵࣜࣥ⣔ࡣ࢜ࣥࢩࢶࢶࣖࢥࣂࢳࡢᡂ࣐࣑࣮ᑐ ࡋ࡚ᙳ㡪ࡀ࠶ࡿሗ࿌ࡉࢀ࡚࠸ࡿࡀ㸦Ἑྜ㸪1988㸹ᯘ㸪1996㸧㸪ᮏヨ㦂࡛ࡶྠᵝ ࡢ⤖ᯝࡀᚓࡽࢀࡓ㸦Table 11, 12㸧ࡇࡽ㸪ᐤ⏕⻏ే⏝࡛ࡁ࡞࠸⪃࠼ࡽࢀ
ࡿࠋ
᪻ཎᛶ⣒≧⳦ࡣ࢜ࣥࢩࢶࢶࣖࢥࣂࢳࢧࣂࢡࢶࣖࢥࣂࢳࡢ࣐࣑࣮ឤᰁ ࡋࡃ࠸ሗ࿌ࡉࢀ࡚࠸ࡿ㸦Fransen and van Lenteren㸪1993㸹Shipp et al.㸪2003㸧ࠋ ᮏヨ㦂࡛ࡶ࣐ࢥࢱ࣮ࣝ®ࣂ࣮ࢱࣞࢵࢡ®ࡣ࣐࣑࣮ឤᰁࡋ࡞ࡗࡓ㸦Table 13㸧ࠋࡋࡋ㸪࣎ࢱࢽ࣮࢞ࢻ ES® ࡢ⩚⋡ࡣ࣐ࢥࢱ࣮ࣝ®ࣂ࣮ࢱࣞࢵࢡ® ẚ᭷ពపࡃ㸦Table 13㸧㸪࣐࣑࣮ឤᰁࡋࡓࡶࡢ⪃࠼ࡽࢀࡿࠋ࢜ࣥࢩࢶࢶ
ࣖࢥࣂࢳࢧࣂࢡࢶࣖࢥࣂࢳࡢᡂᑐࡍࡿ᪻ཎᛶ⣒≧⳦ࡢឤᰁࡣ㧗‵ᗘ
⎔ቃୗ࡛㧗ࡲࡿ㸦Faria and Wraight㸪2001㸹Ludwing and Oetting㸪2001㸧ࠋᮏヨ㦂 ࡣࢩ࣮ࣕࣞෆࡢ㧗‵ᗘ⎔ቃୗ㸦95%RH㸧࡛⾜ࡗࡓࡓࡵ㸪ឤᰁࡋࡸࡍࡗࡓࡢ
ࡶࡋࢀ࡞࠸ࠋ
௨ୖࡢࡼ࠺ẅࡸ᪻ཎᛶ⣒≧⳦ࡼࡗ࡚ࡣᐤ⏕⻏ࡁ࡞ᙳ㡪ࢆ
࠼ࡿྍ⬟ᛶࡀ࠶ࡿࠋᐤ⏕⻏ࢆ⏝ࡍࡿሙྜ㸪ᐤ⏕⻏ᙳ㡪ࡢࡁ࠸ࢆࢃ࡞
100
࠸㜵㝖య⣔ࡢᵓ⠏ࡀ㔜せ࡛࠶ࡿࡀ㸪ᙳ㡪ࡢࡁ࠸ࢆ⏝ࡍࡿሙྜࡣ㸪ࡑࡢᙳ 㡪ࢆపῶࡉࡏࡿࡓࡵࡢᕤኵࢆู㏵᳨ウࡍࡿᚲせࡀ࠶ࡿࠋ
101
➨㸳❶ ⥲ྜ⪃ᐹ
ࢥࢼࢪ࣑ࣛ㢮㸪ࢨ࣑࣐࢘㢮㸪ࣁࣔࢢࣜࣂ࢚㢮࡞ᚤᑠᐖࡣ㸪ᚤᑠ㸪ᗈ㣗 ᛶ㸪㧗࠸ቑṪ⋡㸪ẅᢠᛶ࠸ࡗࡓඹ㏻ࡢ≉ᚩࢆᣢࡕ㸪ࡇࢀࡽᐖࡼࡿ
ᐊ᱂ᇵస≀ࡢ⿕ᐖࡀၥ㢟࡞ࡗ࡚࠸ࡿ㸦Stansly and Naranjo㸪2010㸧ࠋࡑࡢᑐ
⟇ࡋ࡚ẅኳᩛࢆ⤌ࡳྜࢃࡏࡓIPMࡀὶ࡞ࡗ࡚࠸ࡿ㸦Heinz et al.㸪 2004㸹Stansly and Naranjo㸪2010㸧ࠋIPM࡛ࡣ⤒῭ⓗ⿕ᐖチᐜỈ‽㸦EIL㸧ࡢᴫᛕ
✀ࠎࡢᢏ⾡ࢆ⤌ࡳྜࢃࡏࡿࡇࡢ㔜せᛶࡀᙉㄪࡉࢀ㸪せ㜵㝖Ỉ‽ࡼࡾ㜵㝖 ࡀ⾜ࢃࢀࡿࠋࡓ࠼ࡤ㸪࢜ࣥࢩࢶࢥࢼࢪ࣑ࣛࡢࢺ࣐ࢺࡢせ㜵㝖Ỉ‽ࡣᰴ࠶ࡓࡾ
ᡂ⣙ 25 㢌㸦୰ἑࡽ 1979㸧㸪ࢱࣂࢥࢥࢼࢪ࣑ࣛࡢ࣓࡛ࣟࣥࡣⴥ࠶ࡓࡾᡂ 50 㢌㸦ᗈ℩㸪1997㸧ࡉࢀ࡚࠸ࡿࠋࡋࡋ㸪ࡇ࠺ࡋࡓᚑ᮶ࡢIPMࡣࡍ࡛Ⓨ⏕ࡋ
࡚࠸ࡿᐖࢆᑐ㇟ࡋࡓࡶࡢ࡛࠶ࡾ㸪⿕ᐖࢆᮍ↛㜵Ṇࡍࡿ࠸࠺ほⅬࡽࡣ ᚲࡎࡋࡶ༑ศࡣゝ࠼࡞࠸ࠋࡓ࠼ࡤ㸪TYLCV ࢆ፹ࡍࡿࢥࢼࢪ࣑ࣛ㢮ࡢሙ
ྜ㸪ࡈࡃࢃࡎ࡞ಶయ࡛ࡶ ᐊෆධࡋ࡚ࡋࡲ࠼ࡤ㸪࢘ࣝࢫࡀᘏࡋ
ࡡ࡞࠸㸦➨2 ❶㸧ࠋࡇࡢࡓࡵ㸪 ᐊ᱂ᇵ࠾ࡅࡿIPM ࡢ⟇ᐃᙜࡓࡗ࡚ࡣ㸪ࡲ
ࡎᐖࡢධ㜵Ṇ⟇ࢆㅮࡌࡿࡇࡀඛỴ࡛࠶ࡿࠋࡑࡇ࡛㸪ᮏ◊✲࡛ࡣࢥࢼࢪࣛ
࣑㢮ࡢධ㜵Ṇ⟇ࡋ࡚ᙉไẼࢩࢫࢸ࣒ࢆ᳨ウࡋࡓ㸦➨3❶㸧ࠋࡑࡢ࠺࠼࡛㸪 ᐊෆ࡛Ⓨ⏕ࡋ࡚ࡋࡲࡗࡓࢥࢼࢪ࣑ࣛ㢮ᑐࡋ࡚ࡣᐤ⏕⻏ࢆ⏝࠸ࡓ⏕≀ⓗ㜵㝖 ἲ㸦➨4❶㸧ࡉࡽࡣᐤ⏕⻏ే⏝࡛ࡁࡿẅ᪻ཎᛶ⣒≧⳦ࢆ᳨ウࡋ ࡓ㸦➨4 ❶㸧ࠋᮏ❶࡛ࡣ㸪ࡇࢀࡽࡢ◊✲ᡂᯝࢆᇶࡋ࡚᪂ࡋ࠸IPM ࢆᥦࡋࡓ
࠸ࠋ
ᐊෆᐖࢆධࡉࡏ࡞࠸ ᐊᵓ㐀ࡢᇶᮏࡣ㛤ཱྀ㒊ࡢ㜵ࢿࢵࢺࡢᒎᙇ
࡛࠶ࡿࠋ⌧⾜ࡢ ᐊ࡛ࡣ1.0 mm ┠ྜ࠸ࡢ㜵ࢿࢵࢺࡀᒎᙇࡉࢀ࡚࠸ࡿࡀ㸪ࡇ ࡢ㜵ࢿࢵࢺ࡛ࡣ TYLCV ࢆಖẘࡋࡓࢥࢼࢪ࣑ࣛ㢮ࡀ ᐊෆධࡋ࡚ࡋࡲ࠺
ࡓࡵ㸦➨2❶㸧㸪ࢥࢼࢪ࣑ࣛ㢮ࢆධࡉࡏ࡞࠸ࡼࡾ⣽࡞┠ྜ࠸ࡢ㜵ࢿࢵࢺࡀ
102
ᚲせ࡛࠶ࡿࠋᮏ◊✲ࡼࡗ࡚0.38 mm ┠ྜ࠸ࡢ㜵ࢿࢵࢺ㧗࠸ධ㜵Ṇຠᯝ ࡀ࠶ࡿࡇࡀ᫂ࡽ࡞ࡗࡓ㸦➨3❶㸧ࠋࡋࡋ㸪ᒇእࡽࡢ㢼ᅽຊ ᐊෆእ ࡢẼ ᕪࡼࡗ࡚Ẽࡉࢀࡿ⌧⾜ࡢ⮬↛Ẽ ᐊ㸦Bournet and Boulard㸪2010㸹 బ℩㸪2015㸧࡛ࡣ㸪⣽࡞┠ྜ࠸ࡢ㜵ࢿࢵࢺẼຠ⋡ࡣⴠࡕ࡚㧗 ࣭ከ
‵࡞ࡾࡸࡍ࠸㸦Sase and Christianson㸪1990㸹Montero et al㸪1996㸹Muooz et al.㸪
1999㸧ࠋࡑࡢ⤖ᯝ㸪స≀ࡢ⏕⫱ᝏᙳ㡪ࢆཬࡰࡍࡇ࡞ࡿ㸦㧗ᶫ㸪1977㸹Díaz
and Fereres㸪2007㸧ࠋ୍᪉㸪ᙉไẼࡣ⮬↛Ẽẚ✵Ẽࡢຠ⋡ࡀ㧗࠸࠺
࠼㸦Ganguly and Ghosh㸪2011㸧㸪㝖‵ຠᯝࡶ࠶ࡿ㸦Hu et al.㸪2013㸧ࠋࡇ࠺ࡋࡓࡇ
ࡽ㸪ᙉไẼࡣ≉㜵ࢿࢵࢺࢆᒎᙇࡍࡿ ᐊ㐺ࡋ࡚࠸ࡿ㸦Kittas and Katsoulas㸪2005㸹Flores-Velazquez et al.㸪2014㸧ࠋ୍⯡ⓗ࡞㝜ᅽᘧᙉไẼࡣᐖ
ࡀ ᐊࡢ㝽㛫ࡽ ᐊෆᐜ᫆ධ࡛ࡁࡿࡀ㸪㝧ᅽᘧᙉไẼࡣ✵Ẽࡀ ᐊࡢ㝽㛫ࡽእฟࡿࡓࡵᐖࡢධࡣᑡ࡞࠸ࡉࢀ࡚࠸ࡿ㸦Berlinger and Lebiush-Mordechi㸪1996㸹㎷㸪2007㸧ࠋ㝜ᅽᘧᙉไẼ ᐊࡣ⮬↛Ẽ ᐊẚ
TYLCV ࡢⓎ⏕ࡀ㧗࠸࠸࠺ሗ࿌ࡀ࠶ࡿ㸦Berlinger and Lebiush-Mordechi㸪
1996㸧ࠋᮏ◊✲࠾࠸࡚㸪㝧ᅽᘧᙉไẼࢩࢫࢸ࣒㸦Ẽᡪ㸸┤ᚄ0.8m㸪㢼㔞
350ੑ㸭m㸦400w㸧㸪྾Ẽࢲࢡࢺ0.4 mm 0.2 mm┠ྜࡢ㜵ࢿࢵࢺ㸪ኳ❆࣭
ഃ❆1mm┠ྜࡢ㜵ࢿࢵࢺࢆᒎᙇ㸧࡛ࡣTYLCVࡢⓎ⏕ࡣㄆࡵࡽࢀࡎ㸪ࡉࡽ
ᐊෆࡢ ᗘࡣ⮬↛Ẽ㸦ኳ❆࣭ഃ❆ 1mm ┠ྜࡢ㜵ࢿࢵࢺࢆᒎᙇ㸧ࡢࡑࢀ
ẚ2㹼3Υపࡃ࡞ࡿࡇࡀࢃࡗࡓ㸦➨3❶ ➨3⠇㸧ࠋ㝧ᅽᘧᙉไẼࡣᐖ
ࡢධ㜵ṆẼຠ⋡ࡢྥୖࡢ୧㠃ࢆවࡡഛ࠼ࡓඃࢀࡓࢩࢫࢸ࣒⪃࠼ࡽࢀࡿࠋ ḟ㸪 ᐊෆධࡋ࡚ࡋࡲࡗࡓᐖᑐࡍࡿ㜵㝖ᑐ⟇ࡘ࠸࡚⪃ᐹࡍࡿࠋ
࢜ࣥࢩࢶࢶࣖࢥࣂࢳ㸪ࢧࣂࢡࢶࣖࢥࣂࢳ㸪ࢳࢳࣗ࢘࢝ࢶࣖࢥࣂࢳࡣ࠸ࡎࢀࡶ
ࢱࣂࢥࢥࢼࢪ࣑ࣛࡢᐦᗘࢆపୗࡉࡏࡿࡇࡀ᫂ࡽ࡞ࡗࡓࡀ㸪ᐤ⏕⻏ࡢ㜵㝖
ຠᯝࡣᏘ⠇ࡼࡾࡸࡸ␗࡞ࡗࡓࡇࡽ㸦➨4❶ ➨1⠇㸧㸪ḟ㏙ࡿࡼ࠺࡞
⏝ἲࡀ⪃࠼ࡽࢀࡿࠋึኟࡣ㸪࠸ࡎࢀࡢᐤ⏕⻏ࢆ⏝ࡋ࡚ࡶࣂ࢜ࢱࣉBࡢ