3-1 ⥴ゝ
20 ୡ⣖ࡢ㎰ᴗࡣ⛉Ꮫᢏ⾡ࡢⓎᒎࡼࡗ࡚⏕ࡲࢀࡓᏛ〇ရࡀᵝࠎ࡞ኚࢆࡶࡓࡽࡋࡓ㸬
Ꮫ㎰⸆ࢆ⏝࠸ࡓ㜵㝖᪉ἲࡣᐖ⿕ᐖࢆࡁࡃపୗࡉࡏࡿ㧗࠸㜵㝖ຠᯝࢆⓎࡋ㎰ᐙࡢ ປാ㈇ᢸࢆ㍍ῶࡍࡿࡇࡽᗈࡃ⏝ࡉࢀ࡚࠸ࡿ㸬2007ᖺ࠾ࡅࡿ᪥ᮏࡢᏛ㎰⸆ࡢฟⲴ 㔞ࡣ⣙30ࢺࣥ㸪ฟⲴ㔠㢠ࡣ⣙3,600൨ࡉࢀ࡚࠾ࡾ㸪௬Ꮫ㎰⸆ࢆ⏝ࡋ࡞࠸ሙྜ
࡛ࡣỈ✄ࡢ㔞ࡀ 20㹼50%పୗࡍࡿ᥎ ࡉࢀ࡚࠸ࡿ㸬ࡑࡢ㠃㸪Ꮫ㎰⸆ࡣ⎔ቃ㈇ᢸࡀ
ࡁࡃᜏᖖⓗ࡞⏝ࡼࡗ࡚㎰ᆅࡢᅵተ࠾ࡼࡧỈ㉁ởᰁࢆᘬࡁ㉳ࡇࡋ࡚࠸ࡿ㸬ࡉࡽᏛ
㎰⸆౫Ꮡࡋࡓ㜵㝖ࡣṧ␃㎰⸆ၥ㢟ࡸ⸆⪏ᛶᐖࡢⓎ⏕㸪ᚤ⏕≀ᒙࡢ༢⣧క࠺ከ ᵝᛶࡢపୗ㸪ࣜࢧ࣮ࢪ࢙ࣥࢫࡢⓎ⏕࠸ࡗࡓၥ㢟ࡀሗ࿌ࡉࢀ࡚࠸ࡿ (ᮃ᭶, 1997; ⏥ᩫࡽ㸪 1986; ఀ⸨ࡽ㸪1999; ⸨ࡽ㸪2000)㸬
ࡇࡢࡼ࠺࡞≧ἣࢆ⫼ᬒᏛ㎰⸆௦᭰ᢏ⾡ᑐࡍࡿ♫ⓗࢽ࣮ࢬࡀ㧗ࡲࡗ࡚࠸ࡿ㸬⎔ቃ ప㈇Ⲵ㎰ᴗࡋ࡚㸪㍯సయ⣔ࡸᐖᢠᛶရ✀ࡢ⏝㸪⇕ࡼࡿᾘẘࡸᶵᲔ࡞ࢆ⏝࠸ࡓ
≀⌮ⓗ࡞㜵㝖㸪ኳᩛࡸࣇ࢙ࣟࣔࣥࡢ⏝࡞ࡶ⤌ࡳྜࢃࡏࡓ⥲ྜⓗᐖ⟶⌮ (IPM㸸 Integrated Pest Management) ࡀ㔜せ࡞ᙺࢆᯝࡓࡍ⪃࠼ࡽࢀࡿ㸬࡞࡛ࡶᚤ⏕≀⏝
ࡢ◊✲ࡣ┒ࢇ࡛࠶ࡾ㸪ᚤ⏕≀ࡢᵝࠎ࡞᭷⏝ຠᯝࡀゎ᫂ࡉࢀ࡚࠸ࡿ㸬௦⾲ⓗ࡞ࡋ࡚᰿⢏
⳦ࡼࡿ❅⣲ᅛᐃࡸ࣮ࣂࢫ࣮࢟ࣗࣛ⳦᰿⳦ࡼࡿࣜࣥ㓟྾ࡢ⿵ຓస⏝ࡼࡾ⫧ᩱᡂศ ࡢ྾ࡀຠ⋡ⓗ࡞ࡾᡂ㛗ࡀಁ㐍ࡉࢀࡿ (Ho and Trappe, 1975; George et al., 1995) 㸬ࡲ
ࡓ⎔ቃࢫࢺࣞࢫࡢᢠᛶࡸ⦆ຠᯝࡀุ᫂ࡋ࡚࠾ࡾ㸪⇱ࡸሷ㸪㔜㔠ᒓ࡞ࡢ㠀⏕≀ࢫࢺ
ࣞࢫᑐࡍࡿᢠᛶࡀୖ᪼ࡍࡿሗ࿌ࡉࢀ࡚࠸ࡿ (Dimkpa et al., 2009)㸬ࡲࡓᅵተᚤ⏕≀
ᒙࡀከᵝࡍࡿࡇ࡛㐃స㞀ᐖࡢᅇ㑊㸪㍍ῶຠᯝࡀ࠶ࡿࡉࢀ࡚࠸ࡿ㸬ࡇࡢࡼ࠺࡞ᚤ⏕≀
ࡀ᳜≀ࡶࡓࡽࡍከᵝ࡞᭷⏝࡞ຠᯝࡢࡦࡘࡋ࡚ᐖ㜵㝖ࡀ࠶ࡾ㸪ᐖ⪏ᛶࢆಁ㐍ࡍࡿ
78
ᚤ⏕≀ࢆ⏝࠸ࡿ⏕≀ⓗ㜵㝖ࡣ⎔ቃ㈇Ⲵࡀᑡ࡞ࡃ㸪⸆⪏ᛶᐖࡀⓎ⏕ࡋࡃ࠸ࡉࢀ࡚
࠸ࡿ㸬ᚤ⏕≀ࡢᐖ㜵㝖ຠᯝࡋ࡚㸪᳜≀ࡢᐖᢠᛶࢆㄏᑟࡍࡿ㌟⋓ᚓᢠᛶ (Systemic Acquired Resistance: SAR) ࡸㄏᑟ㌟ᢠᛶ (Induced Systemic Resistance:
ISR) ࡀ▱ࡽࢀ࡚࠸ࡿ (Lawton et al., 1996)㸬ࡲࡓ᳜≀ࡢ⪏ᛶࡸ⏕⫱ࢆಁ㐍ࡍࡿ᳜≀⏕⫱
ಁ㐍᰿ᅪ⣽⳦ (Plant Growth-Promoting Rhizobacteria: PGPR) ࡸ᳜≀⏕⫱ಁ㐍⳦㢮 (Plant Growth-Promoting Fungi: PGPF) ࡀ≉ᐃࡉࢀ㸪㏆ᖺ㸪ࡇࡢࡼ࠺࡞ᚤ⏕≀ࡀ᳜≀ࡢ චᶵᵓࢆάᛶࡋ࡚ᐖᢠᛶࢆ㧗ࡵࡿ᳜≀ࣉࣟࣂ࢜ࢸࢡࢫࡋ࡚ὀ┠ࡉࢀ࡚࠾ࡾ㸪 ᵝࠎ࡞ᚤ⏕≀ࡀ㜵㝖㈨ᮦ࡛ၟᶆⓏ㘓㸪ᐇ⏝ࡀ㐍ࡵࡽࢀ࡚࠸ࡿ㸬ᐇ㝿B. subtilis ࡸ㠀
ཎᛶ Fusarium ⳦࡞ࡢᚤ⏕≀ࡀ㎰⸆Ⓩ㘓ࡉࢀ㸪ࡑࢀࡒࢀ࣎ࢺ࣮࢟ࣛỈ㸪࣐ࣝ࢝ࣛ
ࢺࡋ࡚ၟᴗⓗ⏝ࡉࢀ࡚࠸ࡿ (⏣ཱྀࡽ㸸2003)㸬ᐖ⪏ᛶࢆㄏᑟࡍࡿᚤ⏕≀ࡢ≉ᐃࡣ᳜
≀ࡢᐖᢠࢆಁࡍ⏕≀㈨ᮦࡢ㛤Ⓨࡘ࡞ࡀࡾ㸪ప⎔ቃ㈇Ⲵᆺ㎰ᴗ࠾࠸࡚᭷ព⩏࡛࠶ࡿ㸬 ୡ⏺ࡢᚤ⏕≀㈨ᮦᕷሙ㢠ࡣẖᖺ12㸣ୖ᪼ࡋ࡚࠾ࡾ㸪2004ᖺ4൨6800ࢻࣝࡢつᶍ࡛࠶ࡗ ࡓࡀ㸪2010ᖺࡣ10൨7500ࢻࣝ2ಸ௨ୖࡢᡂ㛗ࢆ㐙ࡆ࡚࠾ࡾ㸪ࡉࡽ2018ᖺࡲ࡛
2൨2000ࢻࣝ㐩ࡍࡿண ࡉࢀࡿ᭷ᮃ࡞ᕷሙ࡞ࡿぢ㎸ࡲࢀ࡚࠸ࡿ(Thakore 2006)㸬
᭱㏆㸪ᥬᩫᕝᕤᴗᰴᘧ♫ࡀࢿᐖᢠᛶࢆㄏᑟࡍࡿ᭷⏝ᚤ⏕≀ Penicillium pinophilum YS-31ᰴ (௨ୗ㸪YS-31ᰴ) ࢆ≉チⓏ㘓ࡋࡓ㸬ࡇࡢ⳦ࡣỈ✄ᇵᅵࡼࡾ༢㞳ࡉ
ࢀࡓࡀᐖᢠᛶࡢヲ⣽࡞ㄏᑟ࠾ࡼࡧ⪏ᛶᶵᵓࡣゎ᫂ࡉࢀ࡚࠸࡞࠸㸬ᮏ❶࡛ࡣYS-31ᰴࡢ
ࢿᐖᑐࡍࡿⓎᢚṆᛶࡢཎᅉ࡛࠶ࡿࢆ᫂ࡽࡍࡿࡓࡵྠᰴࡢࢿᐖᑐࡍࡿ
㜵㝖ຠᯝ࠾ࡼࡧᐖᢠᛶㄏᑟ⬟ࢆホ౯ࡋࡓ㸬
79
33-2 ᐇ㦂ᮦᩱ࠾ࡼࡧ᪉ἲ
౪ヨ᳜≀
౪ヨ᳜≀ࡋ࡚ࢿရ✀ࢥࢩࣄ࢝ࣜ㸪ࢩࣟࢾࢼࢬࢼရ✀Col-0ࢆ⏝࠸ࡓ㸬
ᇵᆅసᡂ
⣧ࡉࢀࡓศ㞳⳦ᰴࡢᇵ㣴ࡣPDA (potato dextrose agar) ᇵᆅ10 mg/lࢼࢫࢱࢳࣥ㸪
50 mg/lࣥࣆࢩࣜࣥ㸪10 mg/lࣜࣇࣥࣆࢩࣥ㸪100 μg/l࣑ࢥࢼࢰ࣮ࣝ࡞ࡿࡼ࠺ῧຍࡋ㸪
ᐮኳࡀ⤊⃰ᗘ0.8㸣࡞ࡿࡼ࠺ຍ࠼ࡓᇵᆅ (௨ୗ㸪ᢠ⏕≀㉁ධࡾPDAᇵᆅ) ⏝ࡋࡓ㸬ࡲ ࡓᢠ⏕≀㉁ධࡾPDB (potato dextrose broth) ᇵᆅࡣୖグྠ⃰ᗘ࡞ࡿࡼ࠺ㄪᩚࡋࡓ㸬
ࢿᐖⓎᢚṆヨ㦂
ᮏᐇ㦂ࡣᥬᩫᕝᕤᴗᰴᘧ♫ࢢࣟࣂ࢜ㄢࡢ ᐊ࡛⾜ࡗࡓ㸬㯏⢏࡛ᇵ㣴ࡋࡓࢿ⣠
ᯤ⳦ Rhizoctonia solani࠾ࡼࡧࢿࡶࡳᯤ⣽⳦⳦ Burkholderia glumae ࢆࢿദⱆ
⢄ឤᰁࡉࡏ㸪Ỉ✄⏝⫱ⱑᇵ㣴ᅵ400 g✀ࡋࡓ㸬YS-31ᰴࡢ⳦యᠱ⃮ᾮ (1107 cfu/ml)
ࢆᄇ㟝᥋✀ࡋࡓฎ⌮༊ྠᰴࢆ᥋✀ࡋ࡞࠸ᑐ↷༊ࢆタࡅࡓ㸬⫱ⱑჾ࡛30Υ㸪100 㛫ฟⱆ ฎ⌮ࢆ⾜ࡗࡓᚋ㸪ࢿ⣠ᯤ⳦ฎ⌮ヨ㦂ࡣ20᪥㛫㸪ࢿࡶࡳᯤ⣽⳦⳦ヨ㦂ࡣ 25᪥㛫 ᐊ࡛⫱ⱑࡋ㸪ࢿᗂⱑࡢⓎࢆホ౯ࡋࡓ㸬
RNAᢳฟ㸪㏫㌿ᛂ࠾ࡼࡧᐃ㔞ⓗRT-PCRゎᯒ
YS-31ᰴࢆᢠ⏕≀㉁ධࡾPDBᾮయᇵᆅ500 ml࡛㸪⣙25ºC㸪ᬯᡤ࡛10᪥㛫㟼ᆅᇵ㣴ࡋ ࡓᚋ㸪ᇵ㣴ᾮࢆMILLEX GV 0.22 μm (MILLIPORE) ࡛ࣇࣝࢱ࣮ℐ㐣⁛⳦ࡋ࡚ℐᾮࢆᅇ
ࡋࡓ㸬
➨ 1 ❶ྠᵝࡢ᪉ἲ࡛Ⓨⱆฎ⌮ࢆ⾜ࡗࡓࢿ࠾ࡼࡧࢩࣟࢾࢼࢬࢼࢆỈ⪔⁐ᾮ (2%
MGRL㸪pH5.6) ࡛3㐌㛫Ỉ⪔᱂ᇵࡋ㸪YS-31ᰴฎ⌮༊࡛ࡣ᳜≀యࡢ᰿㒊ࢆYS-31ᰴࡢℐ
ᾮ 24 㛫ᾐₕᚋ㸪ࡧỈ⪔⁐ᾮ࡛ 24 㛫ᇵ㣴ࡋࡓࡢࡕᆅୖ㒊ࢆᅇࡋࡓ㸬ᑐ↷༊ࡣ
80
YS-31ᰴࡢℐᾮฎ⌮ࢆ⾜ࢃࡎ㸪48㛫ᚋᆅୖ㒊ࢆᅇࡋࡓ㸬࡞࠾᱂ᇵ⎔ቃࡣ➨1❶ྠ
᮲௳࡛࠶ࡿ㸬
RNAᢳฟ࠾ࡼࡧ㏫㌿ᛂ㸪ᐃ㔞ⓗRT-PCRࡣ➨1❶ྠᵝࡢ᪉ἲ࡛⾜ࡗࡓ㸬ࢿᐖ ᛂ⟅ᛶ㛵㐃㑇ఏᏊࡣ㸪PR-1a (pathogenesis-related gene 1a ; Os07g0129200)㸪PR-1b (Os01g0382000)㸪PBZ1 (probenazole-inducible protein ; Os12g0555200)㸪OsGLN1 (endo-1,3-glucanase ; Os07g0539100) 㸪 PAL (Phenylalanine ammonia-lyase ; Os02g0626600)㸪Peptidase S8 (Os01g0653800)㸪GSTU4 (Os10g0528300)㸪Peroxidase (Os07g0638400) ࡢ8㑇ఏᏊ㸪ࢩࣟࢾࢼࢬࢼࡣPR1㸪PR2㸪PR3㸪PR4㸪PR5㸪PDF1-2㸪 Peroxidase (At5g64120)㸪WRKY41ࡢ8㑇ఏᏊࡢⓎ⌧㔞ࢆ ᐃࡋࡓ㸬ࢿࡢෆ㒊ᶆ‽㑇ఏ Ꮚࡋ࡚Rubq1ࢆ㸪ࢩࣟࢾࢼࢬࢼࡢෆ㒊ᶆ‽㑇ఏᏊࡣUBQ1ࢆ⏝࠸ࡓ㸬ᮏᐇ㦂࡛⏝࠸ࡓ
ࣉ࣐࣮ࣛ㓄ิࢆTable 8⾲グࡋࡓ㸬
81
Name Sequence (5'-)
F TTATCCTGCTGCTTGCTGG
R GGTCGTACCACTGCTTCTCC
F ACGGGCGTACGTACTGGCTA
R CTCGGTATGGACCGTGAAG
F CGTCCTCGAGCTCATACTCC
R GAGAGGCTGTGGAAGGTCTG
F GAGCCGAACACCTTAGTTGC
R TTCCCCGACACTATCTTTCG
F TCCCATTGTTTGGGAGAAAA
R GAATATTCCTTGCATTATTCAGACA
F AGTGGCCAGGAAGACTCTGA
R GCTTCTTCATCAGCGGGTAG
F TGGAGAGCTTACCAGGCTG
R CCATATTCCCCAAAATGTGC
F TCAGGGGCAGGATCTACAAC
R AGTTCCTCACCGTGTTGTCC
F GTGGTGGCCAGTAAGTCCTC
R GGACACAATGATTAGGGATCA
F TTCTTCCCTCGAAAGCTCAA
R CGTTCACATAATTCCCACGA
F GTCTGAATCAAGGAGCTTAGCC R GATGGACTTGGCAAGGTATCG F GACGCCGACCGTGCCGCCGGG R CGGCGACTCTCCCGTCTTGGCC F GGACCAATGCAGCAACGGAGGC R GGCTGCCCAATGAGCTCATTGCC F ATGTGAGCCTCGTAGATGGTTAC R GATCCATGACCTTAAGCATGTCG
F TTTGCTGCTTTCGACGCAC
R CGCAAACCCCTGACCATG
F TAGGGTCGGGTAAACCTGTG
R GTTGCTCTCTCCGGTGGTC
F CAGCTTCAGGGTTCGTCTTC
R GATCAGGCGCAAGCTCTATC
F TCGTAAGTACAATCAGGATAAGATG R CACTGAAACAAGAAAAACAAACCCT UBQ1 (internal std.)
OsRubq1 Os07g0677500 Os01g0653800
Os02g0626600
PR1
PR2
PR3
PR4
PR5
PDF1-2
peroxidase
WRKY41
Table 8. List of primers used in quantitative RT-PCR experiments.
Os10g0528300 Os07g0539100 Os12g0555200 Os01g0382000 Os07g0129200
82
33-3 ⤖ᯝ࠾ࡼࡧ⪃ᐹ
YS31⳦ᰴ᥋✀ࡼࡿᐖᢚṆᛶࡢホ౯
ࢿ⣠ᯤ⳦࠾ࡼࡧࢿࡶࡳᯤ⣽⳦⳦ᑐࡍࡿ YS-31ᰴࡢᐖᢚṆຠᯝヨ㦂ࡢ⤖ᯝ㸪 ᑐ↷༊࡛ࡣᐖࡢⓎ⏕࡛⏕㛗ࡀᢚไࡲࡓࡣᯤṚࡋࡓࢿᗂⱑࡀከࡃᏑᅾࡋࡓࡀ㸪YS-31 ᰴ ฎ⌮༊࡛ࡣⓎࡀᢚไࡉࢀࢿᗂⱑࡀᖖ࡞⏕㛗ࡋ࡚࠸ࡿࡇࡀ☜ㄆ࡛ࡁࡓ (Fig. 21)㸬
YS-31 ᰴฎ⌮༊ࡢࢿࡣⱑࡀ⨯ⱑࡼࡾࡶከࡃᏑᅾࡋ࡚࠸ࡿࡇࡽ㸪ࢿ⣠ᯤ࠾
ࡼࡧࢿࡶࡳᯤ⣽⳦ᑐࡋ࡚Ⓨᢚไຠᯝࡀ࠶ࡿࡇࡀุ᫂ࡋࢿࡢ⫱ⱑ㜵㝖ຠᯝ ࡀ㧗࠸ᚤ⏕≀㈨ᮦ࡞ࡿࡇࡀ♧၀ࡉࢀࡓ㸬ࡋࡋᮏヨ㦂࡛ࡣ⫱ⱑᅵ࡛ࡢᐖᢚṆࡋ᳨
ドࡋ࡚࠾ࡽࡎ㸪Ỉ⏣⛣᳜ࡋࡓᚋࡶ㜵㝖ຠᯝࡀ⥅⥆ࡍࡿࡢ᳨ドࡀᚲせ࡛࠶ࡿ㸬
YS-31ᰴࡣࢿ࠾ࡼࡧࢩࣟࢾࢼࢬࢼࡢᐖᛂ⟅ᛶ㑇ఏᏊࢆⓎ⌧ㄏᑟࡍࡿ
YS-31 ᰴࡀᐖᛂ⟅ᛶ࠾ࡼࡧᢠᛶ㑇ఏᏊࢆⓎ⌧ㄏᑟࡍࡿ㸪YS-31 ᰴࢆ᥋✀ࡋࡓࢿ
࠾ࡼࡧࢩࣟࢾࢼࢬࢼᆅୖ㒊࡛ᐃ㔞ⓗRT-PCRࡼࡗ࡚㑇ఏᏊⓎ⌧㔞ࢆ ᐃࡋࡓ (Fig. 22)㸬
YS-31ᰴࡢᇵ㣴ℐᾮฎ⌮༊ࡢࢿ࡛ࡣ㸪ឤᰁ≉␗ⓗࢱࣥࣃࢡ㉁PR-1aࡸPR-1b㸪ࣇࢺ
ࣞ ࢟ ࢩ ࣥ ྜ ᡂ 㑇 ఏ Ꮚ bPAL (phenylalanine ammonia-lyase) ࡸ glucan endo-1,3- beta-glucosidase㸪άᛶ㓟⣲㝖ཤ⣔㑇ఏᏊGSTU4ࡸ࣌ࣝ࢜࢟ࢩࢲ࣮ࢮࡢⓎ⌧ࡀᑐ↷༊ẚ
㍑ࡋ࡚᭷ពㄏᑟࡉࢀ࡚࠸ࡓ (Fig. 22A)㸬ḟYS-31ᰴฎ⌮༊ࡢࢩࣟࢾࢼࢬࢼ࡛ࡣ㸪PR1
ࡸPR2㸪PR5㸪ᢠ⳦άᛶࢱࣥࣃࢡ㉁ࢹࣇ࢙ࣥࢩࣥ㑇ఏᏊ PDF1.2 ࡸάᛶ㓟⣲㝖ཤ⣔㑇ఏ ᏊࡢⓎ⌧ࡀㄏᑟࡉࢀ࡚࠸ࡓ୍᪉࡛㸪PR3ࡸPR4ࡢⓎ⌧ኚືࡣᑐ↷༊ẚ㍑ࡋ࡚᭷ព࡞ᕪࡣ
࡞ࡗࡓ (Fig. 22B)㸬ࡇࢀࡲ࡛ࡢ᳜≀ࡢᐖᢠࢩࢢࢼࣝఏ㐩ᶵᵓࡢ◊✲ࡽᐖᢠᛶ 㑇ఏᏊࡣࢧࣜࢳࣝ㓟ࢪࣕࢫࣔࣥ㓟/࢚ࢳࣞࣥࡼࡿࢩࢢࢼࣝఏ㐩⤒㊰࡛Ⓨ⌧ࡀไᚚࡉࢀ࡚
࠾ࡾ㸪ࡉࡽࡇࢀࡽࡢ᳜≀࣍ࣝࣔࣥࡼࡿࢩࢢࢼࣝఏ㐩ࡣᣕᢠⓗస⏝ࡍࡿࡇࡀ᫂ࡽ
࡞ࡗ࡚࠸ࡿ㸬᳜≀ᐖᢠࡢ◊✲ࡽ㸪SARࡣࢧࣜࢳࣝ㓟ࢩࢢࢼࣝఏ㐩⤒㊰㸪ISRࡣࢪ
ࣕࢫࣔࣥ㓟/࢚ࢳࣞࣥࢩࢢࢼࣝఏ㐩⤒㊰ࡀࡁࡃ㛵ࡋ࡚࠾ࡾ㸪ࡲࡓPR-1 ࡸPDF1.2 ࡢⓎ
⌧ㄏᑟࡶࡑࢀࡒࢀ␗࡞ࡿࡇࡀุ᫂ࡋ࡚࠸ࡿ㸬ࡋࡋᐃ㔞ⓗRT-PCRࡢ⤖ᯝࡽYS-31ᰴ
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ࡣࢩࣟࢾࢼࢬࢼࡢᐖᛂ⟅ᛶࢩࢢࢼࣝఏ㐩ࡲ࡛▱ࡽࢀ࡚࠸࡞࠸ᮍ▱ࡢ⤒㊰࡛ㄏᑟࡋ
࡚࠸ࡿࡇࡀ♧၀ࡉࢀࡓ㸬
⏕≀㜵㝖㈨ᮦࡣᗈࡃㄆ▱ࡉࢀࡣࡌࡵ㸪⏕ែᏛⓗࡘศᏊ⏕≀Ꮫⓗࣉ࣮ࣟࢳࡼࡾᅵተ
୰ࡸ᳜≀ࡢ᰿ᅪ㸪᰿㠃㸪᳜≀యෆ⏕ᜥࡍࡿ⭾࡞ᩘࡢᚤ⏕≀ࢆከゅⓗࡘ⥙⨶ⓗᢅ࠺
ゎᯒࡀྍ⬟࡞ࡾᚤ⏕≀ࡢ᭷⏝ຠᯝࡀゎ᫂ࡉࢀࡘࡘ࠶ࡿ (ⓒ⏫ࡽ 2014)㸬ࡇࢀక࠸㸪ཎ
⳦ࡢ㑇ఏⓗ㸪⏕Ꮫⓗ≉ᛶࡘ࠸࡚ࡢ㧗ᗘ࡞ᇶ┙ⓗ◊✲࡞ࡽࡧࢹ࣮ࢱ࣮࣋ࢫࡢタ❧ࡀ㐍
ࡵࡽࢀ࡚࠸ࡿ㸬᭷⏝ᚤ⏕≀ࡢ≉ᐃ᪉ἲࡸᐖ㜵㝖ࡢ≉ᛶࡸ㜵㝖స⏝ᶵᵓࡢゎ᫂࡞㜵㝖
⏝ྥࡅࡓ◊✲ࡶ⢭ຊⓗ⾜࡞ࢃࢀ࡚࠾ࡾ㸪ᚤ⏕≀㜵㝖ࡢ✀㢮ࡀቑࡋ㸪ᐖࡢ✀㢮ࡈ
᭷ຠᶵ⬟ࡍࡿࡢ≉ᐃࢆ⾜࠸ᚤ⏕≀㜵㝖ࡢ⏝ࢆࡍࡿᑐᛂࡀྍ⬟࡞ࢀࡤ㸪IPM ࡢⓎᒎࡁࡃ㈉⊩ࡍࡿ࠸࠼ࡿ㸬⏕≀㈨ᮦ㸪≉᭷⏝ᚤᚤ⏕≀ࡀ㜵㝖ࡶࡓࡽࡍ᭷┈ᛶ ࡣࡁࡃ㸪᳜≀ఏᰁᛶᐖ㜵㝖⬟ࢆ᭷ࡍࡿᚤ⏕≀ࢆ᭷ຠᡂศࡋ࡚⏝࠸ࡿ㜵㝖ࡑࢀࡽ
ࡢ㐺ษ࡞⏝᪉ἲࡀồࡵࡽࢀ࡚࠸ࡿ㸬
ᮏ❶࡛ࡣ㸪YS-31 ᰴࡣࢿᐖࡢⓎᢚṆࡶ᳜≀✀ࢆ㉸࠼ࡓᐖᢠᛶㄏᑟࡢᶵ
⬟ࢆ᭷ࡍࡿ⣒≧⳦࡛࠶ࡿࡇࢆ♧ࡋࡓ㸬ᚋ㸪YS-31 ᰴࡢ⪏ᛶᶵᵓࡘ࠸࡚ࡉࡽ࡞ࡿ◊✲
ࡍࡿࡶ㎰ᆅ࡛ࡢ⳦ࡢⓎ⏕⋡ࡸᐖ㜵㝖ຠᯝ࡞ࡢᐇ⏝ᛶࢆホ౯ࡋ㸪ࣇ࣮ࣝࢻࡢ ᛂ⏝ྥࡅ࡚ࡢㄪᰝࡀᚲせ࡛࠶ࡿ㸬ᅇࡢᐇ㦂࡛YS-31ᰴࡀస≀✀ᐖᢠᛶㄏᑟࢆಁ
㐍ࡍࡿᚤ⏕≀㈨ᮦ࡞ࡾ࠺ࡿྍ⬟ᛶࡀ࠶ࡿࡇࢆド᫂ࡋࡓ㸬ࡑࡢ୍᪉࡛ YS-31ᰴࡀ᳜≀య ࡢ⏕⫱ಁ㐍ࡸࣂ࣐࢜ࢫቑຍ࡞ࡢຠᯝࡣ᳨ドࡋ࡚࠾ࡽࡎ㸪YS-31 ᰴࡢᚤ⏕≀㈨ᮦࡋ࡚
ࡢ᭦࡞ࡿᶵ⬟ゎᯒࡀᮃࡲࢀࡿ㸬
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Figure 21. Effects of P. pinophilum sp. YS-31 inoculation in rice seedlings. Disease symptoms caused by (A) Rhizoctonia solani and (B) Burkholderia glumae infection on rice seedlings. Left shows control plants and right shows plants inoculated by P. pinophilum sp YS-31.
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Figure 22. Expression change of pathogen resistance-related genes in rice leaf and Arabidopsis shoots treated with P.pinophilum. (A) rice and (B) Arabidopsis treated percolate of P.pinophilum transferred seedlings were grown for 3 weeks. Error bars represent SE of the averages of three technical replication. Transcript levels were normalized to the Rubq1 in rice and UBQ1 in Arabidopsis as the internal control. Asterisks show a significant difference by Student’s t test (P <
0.05).
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⥲
⥲ྜ⪃ᐹ
ᮏ◊✲࡛ࡣࢩࣟࢾࢼࢬࢼ࠾ࡼࡧࢿ࠾ࡅࡿ」ᩘ✀ࡢ࣑ࢿࣛࣝࢫࢺࣞࢫ࠾ࡼࡧ᭷⏝ᚤ
⏕≀ฎ⌮ࡼࡗ࡚Ⓨ⌧ኚືࡍࡿ㑇ఏᏊࢆ࣐ࢡࣟࣞゎᯒࢆ㍈ࡋ࡚≉ᐃࡋ㸪㑇ఏᏊ ᶵ⬟ࡽศᏊ⏕⌮Ꮫⓗᛂࡢໟᣓⓗࣉࣟࣇࣜࣥࢢࢆ⾜ࡗࡓ㸬ࡉࡽ᰿ᅪ࣑ࢿࣛࣝࢫࢺ
ࣞࢫᅉᏊ≉␗ⓗᛂ⟅ࡍࡿ㑇ఏᏊⓎ⌧ࣂ࣐࣮࣮࢜࢝ࢆྠᐃࡍࡿ⡆᫆ⓗ᪉ἲࡢᵓ⠏ࢆヨ
ࡳࡓ㸬
ẚ㍑ࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒ゎᯒࡼࡗ࡚N㸪P㸪Kࡢᰤ㣴ḞஈࡸAl3+㸪Cd2+㸪Cu2+㸪NaCl ࡢẘᛶ࢜ࣥࢫࢺࣞࢫࢆ᰿ᅪฎ⌮ࡋࡓࣔࢹ᳜ࣝ≀ᆅୖ㒊ࡢࢫࢺࣞࢫ≉␗ⓗᛂࡲࡓࡣᬑ 㐢ⓗᛂ⟅㑇ఏᏊࡢྠᐃࢆ⾜ࡗࡓࡇࢁ᰿ᅪ㒊ࡢᰤ㣴Ḟஈࡲࡓࡣẘᛶ࢜ࣥࢫࢺࣞࢫࡼࡿ
ᆅୖ㒊ࡢ㌿ኚືࡣࢫࢺࣞࢫᅉᏊࡈ≉␗ᛶࡀ㧗࠸ࡇࡀ♧ࡉࢀࡓ㸬ᆅୖ㒊࠾ࡼࡧᆅୗ
㒊࡛ඹ㏻ࡋ࡚ᙉㄏᑟࡍࡿ㑇ఏᏊࡣᑡ࡞ࡗࡓࡀ㸪㢮ఝࡋࡓᶵ⬟ࡢ㑇ఏᏊࡀⓎ⌧ㄏᑟࡋ࡚࠸
ࡿࡇࡽ᰿ᅪ㒊ࡢࢫࢺࣞࢫ㞀ᐖࡣᆅୖ㒊ࡶྠᵝࡢ㞀ᐖࢆࡶࡓࡽࡋ࡚࠾ࡾ㸪᳜≀ࡣᆅୖ
㒊ᆅୗ㒊ࡑࢀࡒࢀ⤌⧊≉␗ⓗ࡞㑇ఏᏊࡀⓎ⌧ㄏᑟࡋ࡚⪏ᛶࢆ⋓ᚓࡋ࡚࠸ࡿྍ⬟ᛶࡀ⪃࠼
ࡽࢀࡿ㸬Alࢫࢺࣞࢫ࡛ࡣ STOP1 ୗὶ㑇ఏᏊࡀⓎ⌧ㄏᑟࡉࢀ࡚࠾ࡾ㸪ࡉࡽ࣐ࢡࣟࣞ
ゎᯒࡢ⤖ᯝ㸪ࣇࣝࢱࣜࣥࢢࡼࡾ㝖እࡉࢀ࡚࠸ࡿࡀࣜࣥࢦ㓟ᨺฟࢺࣛࣥࢫ࣏࣮ࢱ࣮㑇 ఏᏊAtALMT1ࡀᆅୖ㒊࡛ࡶⓎ⌧ㄏᑟࡋ࡚࠸ࡿࡇࡀ☜ㄆ࡛ࡁࡓ㸬Alࡣ᰿㒊✚ࡋ㸪᳜
≀యෆ⛣⾜ࡋࡃ࠸ࡇࡽAlࢫࢺࣞࢫࢩࢢࢼࣝఏ㐩ࡼࡿㄏᑟࡔ᥎ ࡉࢀࡿ㸬᰿ᅪ
࣑ࢿࣛࣝࢫࢺࣞࢫᑐࡍࡿᆅୖ㒊ࡢࢩࢢࢼࣝఏ㐩ᶵᵓࡢゎ᫂ࡣ㐍ࡵࡽࢀ࡚࠾ࡽࡎ㸪ᚋ ࡢゎᯒࡀᮃࡲࢀࡿ㸬
᰿ᅪ㒊ࡢࢫࢺࣞࢫᑐࡍࡿ᳜≀యᆅୗ㒊ࡢ㌿ゎᯒࡼࡾศᏊ⏕⌮Ꮫⓗᛂ⟅ࢆ᥎ ࡋ⪏ᛶ
㛵ࡍࡿ㑇ఏᏊࡢྠᐃࢆ⾜࠺࠶ࡓࡾ᭷ព⩏࡞ᡭἲ࡛࠶ࡿ㸬ࡇࢀࡣ᰿㒊ࡀ┤᥋ⓗ᰿ᅪ
࣑ࢿࣛࣝࢫࢺࣞࢫࡉࢀ࡚࠸ࡿࡓࡵ࡛࠶ࡿ㸬ࡑࡢ୍᪉࡛ᆅୖ㒊ࡢ㌿ᛂ⟅ࡣ᰿ᅪ㒊ࡢࢫ ࢺࣞࢫᅉᏊࡼࡿ┤᥋ⓗ࡞ᙳ㡪ࡲࡓࡣࢩࢢࢼࣝఏ㐩ࢆࡋࡓ㛫᥋ⓗ࡞ᙳ㡪ࡀ࠶ࡿࡓࡵ⪏ᛶ 㑇ఏᏊࡢ≉ᐃ࠾ࡼࡧゎᯒࡀᅔ㞴࡛࠶ࡿ㸬࡞࡛ࡶ㐣Ꮡᅾࡍࡿẘᛶ࢜ࣥ✀ࡼࡾ⏕ࡌ
ࡿάᛶ㓟⣲ࡣࢩࢢࢼࣝఏ㐩≀㉁࡛࠶ࡿྠ㓟ࢫࢺࣞࢫࡢཎᅉ≀㉁࡛ࡶ࠶ࡾ㸪ᚑ᮶㸪
87
⏕≀ⓗࢫࢺࣞࢫ㠀⏕≀ⓗࢫࢺࣞࢫࡣ␗࡞ࡿ⊂❧ࡋࡓࢩࢢࢼࣝఏ㐩ᶵᵓ࡛࠶ࡿ⪃࠼ࡽࢀ
࡚ࡁࡓࡀ㸪㏆ᖺ࡛ࡣάᛶ㓟⣲࡞ࡽࡧ᳜≀࣍ࣝࣔࣥࢆࡋࡓࢩࢢࢼࣝ⤒㊰ࡀ┦⥭ᐦ
ࡘ㔜」ࡋ࡚࠸ࡿࡇࡀሗ࿌ࡉࢀ࡚࠸ࡿ㸬Zhaoࡽ (2008) ࢆྵࡵࡓከࡃࡢ◊✲࡛ࢩࣟࢾࢼ ࢬࢼࡢ᰿㒊࠾ࡅࡿ࣑ࢿࣛࣝࢫࢺࣞࢫㄏᑟ㑇ఏᏊࡢඹ㏻㡯ࡋ࡚άᛶ㓟⣲ࢫࢺࣞࢫ㛵
ࡍࡿ㑇ఏᏊࢭࢵࢺࡋ࡚ㄏᑟࡉࢀ࡚࠾ࡾ㸪ࡇࢀࡽࡢ㑇ఏᏊᶵ⬟ࡀ᰿㒊ࡢⓎ⌧㑇ఏᏊ㧗࠸
㢮ఝᛶࢆ♧ࡋ࡚࠸ࡿ㸬
ᮏ◊✲࡛ࡣ㸪᰿ᅪ࣑ࢿࣛࣝࢫࢺࣞࢫࡋ࡚N㸪P㸪Kࡢᰤ㣴Ḟஈ࠾ࡼࡧAl㸪Cd㸪Cu㸪NaCl ࡢẘᛶ࢜ࣥࢫࢺࣞࢫฎ⌮࡛Ⓨ⌧ㄏᑟࡉࢀࡿ㑇ఏᏊ⩌╔┠ࡋẚ㍑ࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒
ゎᯒࡼࡾᚓࡽࢀࡓࢫࢺࣞࢫ≉␗ⓗᛂ⟅㑇ఏᏊࢆ⏝࠸᳜≀ࡢᰤ㣴≧ែࢆ᳨ᰝࡍࡿ㑇ఏᏊⓎ⌧
ࣂ࣐࣮࣮࢜࢝ࡢ㛤Ⓨࢆヨࡳࡓ㸬ࡑࡢ⤖ᯝ㸪ᗂ᳜≀యࡢࢫࢺࣞࢫึᮇᛂ⟅ࡢࢺࣛࣥࢫࢡࣜ
ࣉࢺ࣮࣒ࡀ㛗ᮇ㛫⏕⫱࠾ࡼࡧᅵ⪔᱂ᇵࡋࡓ᳜≀ᛂ⏝࡛ࡁࡿࡇࡀ᫂ࡽ࡞ࡗࡓ㸬ࡲࡓ
᭷⏝ᚤ⏕≀ࢆ᥋✀ࡋࡓ᳜≀యࢆ㌿ᛂ⟅࡛ホ౯ࡋ㸪ᐖᢠㄏᑟᛶࡢศᏊ⏕⌮Ꮫⓗホ౯ࡀ
ྍ⬟࡛࠶ࡿࡇࢆ♧ࡋࡓ㸬㑇ఏᏊࡢ࡞ࡣ」ᩘࡢࢫࢺࣞࢫࡼࡗ࡚ㄏᑟࡉࢀࡿࡶࡢࡀከ ࡃᏑᅾࡋ࡚࠾ࡾ㸪Kobayashiࡽ (2013) ࡣAtALMT1ࡀAl3+ࡸపpHࢫࢺࣞࢫࡢࡳ࡞ࡽࡎ
⏕≀ࢫࢺࣞࢫ࡛࠶ࡿࣇࣛࢪ࢙ࣜࣥࡸ࣮࢜࢟ࢩࣥ࡞ࡢ᳜≀࣍ࣝࣔࣥ࠸ࡗࡓᵝࠎ࡞่⃭࡛
ㄏᑟࡉࢀࡿࡇࢆ᫂ࡽࡋ࡚࠸ࡿ㸬ࡇࡢࡼ࠺࡞ഇ㝧ᛶࡢࢫࢺࣞࢫㄏᑟ㑇ఏᏊࢆ㝖ࡃࡓࡵ
ẚ㍑ࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒ゎᯒ࡛㑇ఏᏊᩘࢆ⤠ࡗࡓࡢࡕ」ᩘࡢ㑇ఏᏊⓎ⌧㔞ࢆ ᐃࡍࡿ
ࡇ࡛㌿ࣂ࣐࣮࣮࢜࢝ࡢ⢭☜ᛶ࠾ࡼࡧຠ⋡ᛶࡀྥୖࡍࡿ⪃࠼ࡽࢀࡿ㸬⎔ቃప㈇Ⲵᆺ
㎰ᴗࢆᐇ㊶ࡍࡿ࠶ࡓࡾ㸪ୡ⏺ࡢከᵝ࡞ᅵተ࡛᱂ᇵࡉࢀࡿస≀ࢆ⢭ᐦ㎰ᴗ (Precision
agriculture) ࡛ࣂ࣐࣮࣮࢜࢝ࢆྵࡵࡓ」ྜⓗ࡞ᰤ㣴≧ែ⟶⌮ࡼࡗ࡚᳜≀ࡢᰤ㣴≧ែࢆ
ᢕᥱࡍࡿࡇࡀᮇᚅࡉࢀࡿ㸬ࡲࡓᆅୖ㒊ࡢⓎ⌧㑇ఏᏊࢆ᳨ᰝࡍࡿࡇ࡛Ⰻᅵተࡢ㎰ᴗ࡛
ࡣ༊⏬ࡈࡢ⣽ࡸ࡞ᰤ㣴⟶⌮ࡀྍ⬟࡞ࡾ㸪࠼ࡤ㓟ᛶᅵተ࡛ࡣᴟ➃పpHࡢ༊⏬ᑐ ࡋ࡚▼⅊ࢆ㏣⫧ࡍࡿࡇ⏕⏘ᛶࢆྥୖࡍࡿࡇࡀྍ⬟࡞ࡿ㸬ᵝࠎ࡞≀㉁ࡀࣂ࣐࣮࢜࢝
࣮࡞ࡗ࡚࠸ࡿࡀ㸪miRNAࡢⓎ⌧㔞ࡶࢫࢺࣞࢫ≧ែࢆ♧ࡍ᭷⏝࡞㑇ఏᏊⓎ⌧ࣂ࣐࣮࢜࢝
࣮࡞ࡾ࠺ࡿࡇࡀሗ࿌ࡉࢀ࡚࠸ࡿ㸬㑇ఏᏊⓎ⌧ࣂ࣐࣮࣮࢜࢝ࡣᅵተ᳨ᰝ࡛ࡣ⢭ᐦ࡞ホ ౯ࡀᅔ㞴࡛࠶ࡿࢫࢺࣞࢫ㞀ᐖࢆ᳨ᰝࡍࡿࡇࡀྍ⬟࡛࠶ࡿࡓࡵ㸪㎰ᆅ࡛ࡢస≀ᰤ㣴⟶⌮
88
㈉⊩ࡍࡿ⪃࠼ࡽࢀࡿ㸬ࡲࡓ㓟ᛶᅵተࢆࣔࢹࣝࡋ࡚㯮࣎ࢡᅵࢆ⏝࠸㸪STOP1ୗὶ㑇ఏᏊ
ࢆAlᛂ⟅ᛶࣂ࣐࣮࣮࢜࢝㑇ఏᏊࡢⓎ⌧㔞ࢆほᐹࡋࡓࡇࢁ㸪ⴭࡋࡃⓎ⌧ㄏᑟࡉࢀ࡚࠸ࡿ
ࡇࡀุ᫂ࡋࡓ㸬ࡲࡓࢱࣂࢥ࡛ࡶNtALS3ࡀAlᛂ⟅ࡍࡿ㌿ࣂ࣐࣮࣮࡛࢜࢝࠶ࡿࡇ
ࢆド᫂ࡋࡓ㸬᳜≀ᬑ㐢ⓗᏑᅾࡍࡿࢫࢺࣞࢫᛂ⟅ᶵᵓࡢ㑇ఏᏊࢆ≉ᐃࡍࡿࡇࡀ㸪ࣂ
࣐࣮࣮࢜࢝ࡢ≉ᐃ᭷⏝࡞㑇ఏᏊ࡛࠶ࡿ࠸࠼ࡿ㸬 Yangࡽ (2011) ࡀ♧ࡋࡓࡼ࠺㸪 ᐊ
࡛᱂ᇵࡋࡓ᳜≀ࡢࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒ࢹ࣮ࢱࡼࡾᚓࡽࢀࡓࣂ࣐࣮࣮࢜࢝㑇ఏᏊࡀࣇ
࣮ࣝࢻ᱂ᇵࡋࡓ᳜≀ࡶ㐺ᛂ࡛ࡁࡿࡇࡀ♧ࡉࢀ࡚࠸ࡿ㸬ᮏᐇ㦂࡛ࡶỈ⪔᱂ᇵ⣔ࡼࡾ≉ᐃ ࡋࡓࣂ࣐࣮࣮࢜࢝㑇ఏᏊࡀᅵ⪔᮲௳ࡼࡿ㛗ᮇࢫࢺࣞࢫฎ⌮ᑐࡋ࡚ࡶ༑ศ࡞ᛶࡀ
࠶ࡿࡇࢆ♧ࡋࡓ㸬㑇ఏⓗ⫼ᬒࡀ␗࡞ࡿရ✀࡛ࡣࣂ࣐࣮࣮࢜࢝㑇ఏᏊࡢⓎ⌧㔞ࡁ࡞
ᕪࡀฟࡿࡇࡶண ࡉࢀࡿ㸬࠼ࡤ㸪❅⣲⏝ຠ⋡ (NUE; nitrogen use efficiency) ࡸAl
ᑐࡍࡿ᭷ᶵ㓟ᨺฟࡀရ✀ࡈ␗࡞ࡗ࡚࠸ࡿࡇࡀุ᫂ࡋ࡚࠸ࡿ (Gallais and Hirel, 2004; Ikka et al., 2008)㸬ᮏ◊✲࡛ࡣࣂ࣐࣮࣮࢜࢝㑇ఏᏊࡢရ✀㛫࡛ࡢᬑ㐢ⓗᛂ⏝ᛶࡣᐇ 㦂ࡋ࡚࠸࡞࠸ࡀ㸪Yangࡽࡣࣇ࣮ࣝࢻ࡛⫱࡚ࡓ」ᩘࡢࢺ࢘ࣔࣟࢥࢩ㞧ရ✀࡛NḞஈࣂ
࣐࣮࣮࢜࢝ࡀN Ḟஈᛂ⟅ࡍࡿࢆヨ㦂ࡋࡓࡇࢁከࡃࡢရ✀࡛ᰤ㣴≧ែࡢホ౯ࡀྍ⬟࡛
࠶ࡿࡇࢆ♧ࡋࡓ㸬ࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒ࢆ⏝࠸ࡓゎᯒ࡛ࡣ」ྜࢫࢺࣞࢫࢆ⏝࠸ࡓゎᯒࡸ
ᅵተ࡛⏕⫱ࡉࡏࡓ᳜≀ࡢ㌿ゎᯒࡶ⾜ࢃࢀ࡚࠾ࡾ㸪ࣇ࣮ࣝࢻ⎔ቃୗ࠾ࡼࡧࡑࢀࢆᶍࡋࡓ ࢫࢺࣞࢫᛂ⟅ᶵᵓࡢゎ᫂ࡀ⢭ຊⓗ㐍ࡵࡽࢀ㸪ࡉࡽ࣮࣑࢜ࢡࢫゎᯒࢆ⤫ྜࡋࡓᢏ⾡࡛ࣇ
࣮ࣝࢻ࠾ࡅࡿ᱂ᇵ⟶⌮ࢆ⾜࠺᪉ἲࡀ᳨ウࡉࢀ࡚࠸ࡿ (Jensen et al., 2010; Sato et al.,2011; Nagano et al., 2012; Alexandersson et al., 2014)㸬
ࣔࢹ᳜ࣝ≀࡛࠶ࡿࢩࣟࢾࢼࢬࢼࡣᵝࠎ࡞ࢫࢺࣞࢫฎ⌮࡛ࡢࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒ࢹ࣮
ࢱࡀᐇࡋ࡚࠾ࡾศᏊ⏕≀Ꮫⓗゎᯒࡢᇶ┙ࡋ࡚ࡁࡃ㈉⊩ࡋ࡚࠸ࡿ㸬ࡑࡢ୍᪉࡛᳜≀✀
ࡈ㔞ࡢ㌿ࢹ࣮ࢱࢆ✚ࡍࡿࡇࡣᅔ㞴࡛࠶ࡾ㸪ࡲࡓရ✀㸪᳜≀ࡢ⏕⫱ࢫࢸ࣮ࢪࡈ
ࡢ⣽࡞㌿ሗࢆ㞟✚ࡍࡿࡇࡶ⮳㞴࡛࠶ࡿ㸬ᚑࡗ࡚ࢩࣟࢾࢬࢼࡸࢿ࡞ࡢࣔࢹ
᳜ࣝ≀ࡼࡾᚓࡽࢀࡓ㌿ሗࡽ࣮࢜ࢯࣟࢢࢆ≉ᐃࡋ㸪」ᩘࡢ㑇ఏᏊⓎ⌧ࣂ࣐࣮࣮࢜࢝
࡛᳜≀ࡢᗣ≧ែࢆホ౯ࡍࡿࡇࡀ⌧ᐇⓗ࡛࠶ࡿ⪃࠼ࡽࢀࡿ㸬୍⯡ⓗస≀ࡣࢫࢺࣞࢫ ឤཷᛶ࡛࠶ࡿࡓࡵ㸪ࢫࢺࣞࢫᑐࡋ࡚Ⓨ⌧ᢚไࡉࢀࡿ㑇ఏᏊ⩌ࡀࣂ࣐࣮࣮࢜࢝㐺ࡋ࡚
89
࠸ࡿ⪃࠼ࡽࢀࡿࡀ㸪Alࢫࢺࣞࢫ≉␗ⓗⓎ⌧ㄏᑟࡍࡿ㑇ఏᏊࡋ࡚ALS3ࡀ᳜≀✀ᬑ㐢
ⓗᏑᅾࡍࡿ㌿ࣂ࣐࣮࣮࡛࢜࢝࠶ࡿド᫂ࡋࡓ㸬ᐇ㝿ࡢ㎰ᆅ࡛ࢫࢺࣞࢫ⪏ᛶࡢࡸ
㑇ఏᏊⓎ⌧ࣂ࣐࣮࣮࢜࢝ࡢ㛤Ⓨࡢࡓࡵࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒ࢹ࣮ࢱࡢ✚ࡀࡼࡾ㔜せ
࡞ࡿ㸬ᮏ◊✲ࡶ㸪ࡑࡢࣔࢹࣝ࡞ࡿ◊✲࡛࠶ࡾᆅୖ㒊ࡢ㌿ᛂ⟅ࡢゎ᫂ࡀࡁࡃ๓㐍ࡍ
ࡿ⪃࠼ࡽࢀࡿ㸬
௨ୖࡢࡇࢆ㋃ࡲ࠼㸪ᮏ◊✲ࡢᡂᯝࡋ࡚࣑ࢿࣛࣝࢫࢺࣞࢫᑐࡍࡿࣔࢹ᳜ࣝ≀ࡢᗂ᳜
≀యࡢྛࢫࢺࣞࢫ≉␗ⓗ㑇ఏᏊࡢ≉ᐃ࠾ࡼࡧ㸪ࡑࢀࢆ⏝࠸࡚⡆౽࡞᪉ἲ࡛㑇ఏᏊⓎ⌧ࣂ
࣐࣮࣮࢜࢝ࢆ᥈⣴㸪ྠᐃࡀྍ⬟࡛࠶ࡿࡇࢆ♧ࡋࡓ㸬Ỉ⪔ᅵ⪔ࡢ᰿ᅪ⎔ቃࡢᕪ␗ࡣ࠶ࡾ
࡞ࡀࡽࡶ㸪ᆅୖ㒊࡛ࡣྠ୍ࡢ㑇ఏᏊࡢⓎ⌧ኚ࡛࠶ࡿࡇࡀ᥎ ࡉࢀࡿ㸬ᮏ◊✲࡛ࡶỈ⪔
᱂ᇵ᳜≀యࡢ㌿ኚືࡀࢫࢺࣞࢫึᮇࡢⓎ⌧ㄏᑟ㑇ఏᏊ࡛࠶ࡗ࡚ࡶ⏕⫱୰ᮇࡢ᳜≀యࡲࡓ ࡣᅵ⪔᱂ᇵࡢ᳜≀యࡶᛂ⏝࡛ࡁࡿࡇࢆⓎぢࡋࡓ㸬࣮࣑࢜ࢡࢫ◊✲ࡼࡿࣂ࣐࣮࢜࢝
࣮ࡼࡿ᳜≀ࡢᰤ㣴⟶⌮ࡣ⌧ᐇࢆᖏࡧ࡚ࡁ࡚࠸ࡿ㸬✚ࡉࢀࡓ⭾࡞࣮࣑࢜ࢡࢫࢹ࣮ࢱ
ࡽศᏊ⏕⌮ⓗᛂࢆᫎࡋࡓ㸪㑇ఏᏊⓎ⌧ไᚚᶵᵓࢆゎ᫂ࡋࣔࢹࣝࢆᵓ⠏ࡋ㸪᳜≀ࡢ⎔
ቃࢫࢺࣞࢫᛂ⟅ᶵᵓࡢ⌮ゎ㈉⊩ࡍࡿ⪃࠼ࡽࢀࡿ㸬
90