αριστοτελειο πανεπιστημιο θεσσαλονικης σχολη γεωτεχνικων επιστημω

breathobeseΒιοτεχνολογία

19 Φεβ 2013 (πριν από 4 χρόνια και 6 μήνες)

246 εμφανίσεις

ǹȇǿȈȉȅȉǼȁǼǿȅ ȆǹȃǼȆǿȈȉǾȂǿȅ ĬǼȈȈǹȁȅȃǿȀǾȈ
ȈȋȅȁǾ īǼȍȉǼȋȃǿȀȍȃ ǼȆǿȈȉǾȂȍȃ –ȉȂǾȂǹ īǼȍȆȅȃǿǹȈ
ȆȇȅīȇǹȂȂǹ ȂǼȉǹȆȉȊȋǿǹȀȍȃ ȈȆȅȊǻȍȃ
ǼǿǻǿȀǼȊȈǾ ǼȆǿȈȉǾȂǾȈ Ȁǹǿ ȉǼȋȃȅȁȅīǿǹȈ ȉȇȅĭǿȂȍȃ

ȆǹȇǹīȍīǾ ȕ-ȀǹȇȅȉǼȃǿȅȊ ǹȆȅ ȈȊȃĬǼȉǿȀȅ
ȊȆȅȈȉȇȍȂǹ ȂǼ ȉȅ ȂȊȀǾȉǹ BLAKESLEA TRISPORA
ȈǼ ǽȊȂȍȈǾ ǺȊĬȅȊ

ǻǿǻǹȀȉȅȇǿȀǾ ǻǿǹȉȇǿǺǾ

ȂǹȃȉǽȅȊȇǿǻȅȊ ĭǹȃǾ
ȋǾȂǿȀȅȈ-M.Sc. in Food Biotechnology


ǼȆǿǺȁǼȆȍȃ:
ȉȡȚĮȞIJȐijȣȜȜȠȢ ȇȠȣțȐȢ


ǼȄǼȉǹȈȉǿȀǾ ǼȆǿȉȇȅȆǾ
ȉȡȚĮȞIJȐijȣȜȜȠȢ ȇȠȣțȐȢ, ȀĮșȘȖȘIJȒȢ
ȆĮȡșȑȞĮ ȀȠIJȗİțȓįȠȣ-ȇȠȣțȐ, ȀĮșȘȖȒIJȡȚĮ
ȂĮȡȓĮ ȁȚĮțȠʌȠȪȜȠȣ-ȀȣȡȚĮțȓįȠȣ, ȀĮșȘȖȒIJȡȚĮ
ǹȚțĮIJİȡȓȞȘ ȉȗĮȕȑȜȜĮ-ȀȜȦȞȐȡȘ, ȀĮșȘȖȒIJȡȚĮ
ǻȘµȒIJȡȚȠȢ ȀȣȡȚĮțȓįȘȢ, ȀĮșȘȖȘIJȒȢ
ȋȡȒıIJȠȢ ȆĮȞĮȖȚȦIJȓįȘȢ, ǹȞĮʌȜ. ȀĮșȘȖȘIJȒȢ
ȂĮȡȓĮ īĮȜȚȫIJȠȣ, ǹȞĮʌȜ. ȀĮșȘȖȒIJȡȚĮ


ĬǼȈȈǹȁȅȃǿȀǾ 2003

ȆǼȇǿȁǾȌǾ

ȈIJȘȞ ʌĮȡȠȪıĮ İȡȖĮıȓĮ įȚİȡİȣȞȒșȘțİ Ș įȣȞĮIJȩIJȘIJĮ ȕİȜIJȓȦıȘȢ IJȘȢ ĮʌȩįȠıȘȢ
țĮȚ IJȘȢ ʌĮȡĮȖȦȖȚțȩIJȘIJĮȢ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ Įʌȩ ıȣȞșİIJȚțȩ ȣʌȩıIJȡȦµĮ µİ IJȠ µȪțȘIJĮ
Blakeslea trispora. īȚĮ IJȠ ıțȠʌȩ ĮȣIJȩ, ȠȚ ȗȣµȦIJȚțȑȢ įȚİȡȖĮıȓİȢ ȑȖȚȞĮȞ ıİ
ĮȞĮțȚȞȠȪµİȞİȢ ijȚȐȜİȢ țĮȚ ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ µİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ țĮȚ ıİ
ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ µİ ĮıȣȞİȤȒ ȗȪµȦıȘ µİ ıȣȞİȤȒ ʌȡȠıșȒțȘ ȣʌȠıIJȡȫµĮIJȠȢ.
ȈIJȠ ĮȡȤȚțȩ ıIJȐįȚȠ IJȘȢ µİȜȑIJȘȢ ĮȞĮʌIJȪȤșȘțİ µȓĮ ȕİȜIJȚȦµȑȞȘ µȑșȠįȠȢ
ĮȞȐțIJȘıȘȢ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ Įʌȩ IJĮ țȪIJIJĮȡĮ IJȠȣ B. trispora. ȉȠ ȣȖȡȩ IJȘȢ ȗȪµȦıȘȢ
șİȡµĮȓȞȠȞIJĮȞ ıIJȠȣȢ 121
0
C ȖȚĮ 15 ȜİʌIJȐ țĮȚ ıIJȘ ıȣȞȑȤİȚĮ ijȣȖȠțİȞIJȡȠȪȞIJĮȞ ıİ
5.000g ȖȚĮ 20 ȜİʌIJȐ. ǹțȠȜȠȣșȠȪıİ ȑțʌȜȣıȘ IJȠȣ µȣțȘȜȓȠȣ µİ ĮʌȠıIJĮȖµȑȞȠ Ȟİȡȩ țĮȚ
ijȣȖȠțȑȞIJȡȘıȘ IJȠȣ ȣȖȡȠȪ ıİ 5.000g ȖȚĮ 20 ȜİʌIJȐ. Ǿ İțȤȪȜȚıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ
ȑȖȚȞİ µİ ʌȡȠıșȒțȘ 100 ml ĮȜțȠȩȜȘȢ/g ȟȘȡȠȪ ȕȐȡȠȣȢ ȕȚȠµȐȗĮȢ ıİ șİȡµȠıIJĮIJȠȪµİȞȠ
ʌĮȜȚȞįȡȠµȘIJȒ ĮȞĮțȓȞȘıȘȢ, ıIJĮ 300 rpm, ıİ șİȡµȠțȡĮıȓĮ 30
0
C. ȉȠ ȕ-țĮȡȠIJȑȞȚȠ
ĮʌȠµĮțȡȪȞȠȞIJĮȞ ʌȜȒȡȦȢ Įʌȩ IJĮ țȪIJIJĮȡĮ IJȠȣ µȚțȡȠȠȡȖĮȞȚıµȠȪ µİIJȐ Įʌȩ 3
įȚĮįȠȤȚțȑȢ İțȤȣȜȓıİȚȢ. ȉȠ µİȖĮȜȪIJİȡȠ ʌȠıȠıIJȩ IJȠȣ İțȤȣȜȚıµȑȞȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ
ʌĮȡĮȜȒijșȘțİ µİIJȐ IJȠ IJȑȜȠȢ IJȘȢ ʌȡȫIJȘȢ İțȤȪȜȚıȘȢ (75,94%), İȞȫ Ș ĮʌȩįȠıȘ IJȘȢ
įİȪIJİȡȘȢ țĮȚ IJȡȓIJȘȢ İțȤȪȜȚıȘȢ ȒIJĮȞ ıȘµĮȞIJȚțȐ µȚțȡȩIJİȡİȢ (16,62% țĮȚ 7,44%,
ĮȞIJȓıIJȠȚȤĮ).
ȀĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ İʌȓʌİįȠ țȦȞȚțȫȞ ijȚĮȜȫȞ µİȜİIJȒșȘțİ Ș
İʌȓįȡĮıȘ IJȠȣ ݵȕȠȜȓȠȣ, IJȘȢ șİȡµȠțȡĮıȓĮȢ, IJȠȣ pH, IJȠȣ ijȣıȚțȠȪ ijȦIJȩȢ, IJȘȢ ʌȘȖȒȢ
ȐȞșȡĮțĮ țĮȚ ĮȗȫIJȠȣ, IJȦȞ İʌȚijĮȞİȚȠįȡĮıIJȚțȫȞ ȠȣıȚȫȞ, IJȦȞ ijȣIJȚțȫȞ İȜĮȓȦȞ, IJȦȞ
ĮțȩȡİıIJȦȞ ȜȚʌĮȡȫȞ ȠȟȑȦȞ, IJȠȣ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ țĮȚ įȚĮijȩȡȦȞ ʌȡȩįȡȠµȦȞ ȠȣıȚȫȞ
țĮȚ İȞİȡȖȠʌȠȚȘIJȫȞ ıIJȚȢ țȚȞȘIJȚțȑȢ ʌĮȡĮµȑIJȡȠȣȢ IJȘȢ ȗȣµȦIJȚțȒȢ įȚİȡȖĮıȓĮȢ. ȅȚ
ʌĮȡȐµİIJȡȠȚ ĮȣIJȠȓ ȒIJĮȞ Ș ıȣȖțȑȞIJȡȦıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ, IJȘȢ ȕȚȠµȐȗĮȢ, IJȦȞ
ȣʌȠȜİȚµµĮIJȚțȫȞ ıĮțȤȐȡȦȞ țĮȚ IJȠ pH IJȠȣ ȣȖȡȠȪ IJȘȢ ȗȪµȦıȘȢ. Ǿ µİȖĮȜȪIJİȡȘ
ĮʌȩįȠıȘ IJȘȢ ȤȡȦıIJȚțȒȢ ʌĮȡĮIJȘȡȒșȘțİ ıIJȠ ȣʌȩıIJȡȦµĮ ʌĮȡĮȖȦȖȒȢ (pH 7,0) ʌȠȣ
ݵȕȠȜȚȐıIJȘțİ µİ ȑȞĮ țȠµµȐIJȚ ȐȖĮȡ Įʌȩ țȐșİ ȑȞĮ IJȡȣȕȜȓȠ țĮȜȜȚȑȡȖİȚĮȢ IJȦȞ
ıIJİȜİȤȫȞ ĮȞIJȓșİIJȠȣ ijȪȜȠȣ IJȠȣ B. trispora, IJȘȞ 8
Ș
ȘµȑȡĮ IJȘȢ ȗȪµȦıȘȢ ıIJȠȣȢ 26
0
C
țĮȚ ʌĮȡȠȣıȓĮ ijȣıȚțȠȪ ijȦIJȩȢ ıİ ȩȜȘ IJȘ įȚȐȡțİȚĮ IJȘȢ ȗȣµȦIJȚțȒȢ įȚİȡȖĮıȓĮȢ. ȉĮ
ĮʌȠIJİȜȑıµĮIJĮ ȑįİȚȟĮȞ ȩIJȚ Ș ĮȞIJȚțĮIJȐıIJĮıȘ IJȘȢ ȖȜȣțȩȗȘȢ µİ ıĮțȤĮȡȩȗȘ, ȐµȣȜȠ țĮȚ
ȖȜȣțİȡȩȜȘ µİȓȦıĮȞ IJȘȞ ʌĮȡĮȖȦȖȒ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ, İȞȫ Ƞ ݵʌȜȠȣIJȚıµȩȢ IJȠȣ
ȣʌȠıIJȡȫµĮIJȠȢ µİ țĮȜĮµʌȠțȐȜİȣȡȠ, ȣįȡȠȜȣµȑȞȘ ʌȡȦIJİȓȞȘ ıȩȖȚĮȢ Ȓ ıIJİȡİȐ
ȣʌȠȜİȓµµĮIJĮ ĮʌȩıIJĮȟȘȢ ĮȜțȠȩȜȘȢ ȠįȒȖȘıİ ıİ µȚțȡȒ ĮȪȟȘıȘ IJȘȢ ȤȡȦıIJȚțȒȢ.
ǹȞIJȓșİIJĮ, Ș ʌȡȠıșȒțȘ ȖȜȣțȩȗȘȢ, İțȤȣȜȓıµĮIJȠȢ țĮȜĮµʌȠțȚȠȪ, Span 20, Tween 80,
İȜĮȚȩȜĮįȠȣ, ȕĮµȕĮțȑȜĮȚȠȣ, ıȠȖȚȑȜĮȚȠȣ, İȜĮȧțȠȪ țĮȚ ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ, țȘȡȠȗȓȞȘȢ
țĮȚ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ (ȕȠȣIJȣȜȚȦµȑȞȠ ȣįȡȠȟȣIJȠȜȠȣȩȜȚȠ, BHT) ĮȪȟȘıĮȞ ıȘµĮȞIJȚțȐ
IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ. ȈIJȘ ıȣȞȑȤİȚĮ, įȚİȡİȣȞȒșȘțİ Ș µİIJĮȕȠȜȒ IJȘȢ
µȠȡijȠȜȠȖȓĮȢ IJȠȣ µȚțȡȠȠȡȖĮȞȚıµȠȪ µȑıȦ İȚįȚțȠȪ ʌȡȠȖȡȐµµĮIJȠȢ µȠȡijȠµİIJȡȓĮȢ ıİ
ıȤȑıȘ µİ IJȘ ıȪıIJĮıȘ IJȠȣ ȣʌȠıIJȡȫµĮIJȠȢ. ȀĮIJȐ IJȘ įȚȐȡțİȚĮ IJȠȣ ȕȚȠȜȠȖȚțȠȪ țȪțȜȠȣ
IJȠȣ B. trispora Ƞ șĮȜȜȩȢ IJȠȣ µȪțȘIJĮ įȚĮijȠȡȠʌȠȚİȓIJĮȚ ıİ ȕȜĮıIJȚțȩ țĮȚ
ĮȞĮʌĮȡĮȖȦȖȚțȩ IJµȒµĮ. ȉȠ ȕȜĮıIJȚțȩ IJµȒµĮ ȕȡȓıțİIJĮȚ µİ IJȘ µȠȡijȒ ȐșȚțIJȦȞ
µȣțȘȜȚĮțȫȞ ȣijȫȞ, µȣțȘȜȚĮțȫȞ ȣijȫȞ µİ țİȞȠIJȩʌȚĮ, țİȞȫȞ țȣIJIJȐȡȦȞ țĮȚ
İțijȣȜȚıµȑȞȦȞ țȣIJIJȐȡȦȞ, İȞȫ İʌȓ IJȠȣ ĮȞĮʌĮȡĮȖȦȖȚțȠȪ IJµȒµĮIJȠȢ ݵijĮȞȓȗȠȞIJĮȚ IJĮ
ȖĮµİIJȐȖİȚĮ ʌȠȣ ijȑȡȠȣȞ IJĮ ȩȡȖĮȞĮ ĮȞĮʌĮȡĮȖȦȖȒȢ (ȖĮµȑIJİȢ), Įʌȩ IJĮ ȠʌȠȓĮ µİ
ıȣȖȤȫȞİȣıȘ ȖİȞİIJȚțȒȢ ȠȣıȓĮȢ IJȦȞ ʌȣȡȒȞȦȞ IJȠȣȢ ʌȡȠțȪʌIJȠȣȞ IJĮ ȗȣȖȠıʌȩȡȚĮ. ȉĮ
ĮʌȠIJİȜȑıµĮIJĮ IJȘȢ µȠȡijȠµİIJȡȚțȒȢ ĮȞȐȜȣıȘȢ ȑįİȚȟĮȞ ȩIJȚ ıIJȠ ȐȡȚıIJȠ ȣʌȩıIJȡȦµĮ
ʌĮȡĮȖȦȖȒȢ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ IJȠ µİȖĮȜȪIJİȡȠ ʌȠıȠıIJȩ IJȘȢ ȟȘȡȒȢ ȕȚȠµȐȗĮȢ IJȠȣ
µȚțȡȠȠȡȖĮȞȚıµȠȪ ĮȞIJȚıIJȠȚȤȠȪıİ ıİ ȗȣȖȠıʌȩȡȚĮ (79,84%), İȞȫ IJȠ ʌȠıȠıIJȩ IJȦȞ
ȐșȚțIJȦȞ µȣțȘȜȚĮțȫȞ ȣijȫȞ, IJȦȞ µȣțȘȜȚĮțȫȞ ȣijȫȞ µİ țİȞȠIJȩʌȚĮ, IJȦȞ țİȞȫȞ
țȣIJIJȐȡȦȞ țĮȚ IJȦȞ İțijȣȜȚıµȑȞȦȞ țȣIJIJȐȡȦȞ ȒIJĮȞ ʌȠȜȪ ȤĮµȘȜȩ. Ǿ ȠȜȠțȜȒȡȦıȘ IJȘȢ
ȑȡİȣȞĮȢ ȖȚĮ IJȘ ȕİȜIJȚıIJȠʌȠȓȘıȘ IJȦȞ ʌĮȡĮµȑIJȡȦȞ IJȘȢ ȗȣµȦIJȚțȒȢ įȚİȡȖĮıȓĮȢ ıİ
İʌȓʌİįȠ țȦȞȚțȫȞ ijȚĮȜȫȞ, ʌȡĮȖµĮIJȠʌȠȚȒșȘțİ µİ IJȘ ȤȡȒıȘ ıIJĮIJȚıIJȚțȠȪ µȠȞIJȑȜȠȣ
ʌȡȩȕȜİȥȘȢ, IJȠ ȠʌȠȓȠ ȕĮıȓȗİIJĮȚ ıIJȘ µİșȠįȠȜȠȖȓĮ IJȘȢ İʌȚijĮȞİȚĮțȒȢ ĮʌȩțȡȚıȘȢ. Ȃİ IJȠ
ʌĮȡĮʌȐȞȦ µȠȞIJȑȜȠ İȡµȘȞİȪIJȘțĮȞ ıİ ȚțĮȞȠʌȠȚȘIJȚțȩ ȕĮșµȩ ȠȚ İʌȚįȡȐıİȚȢ IJȠȣ
ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ, IJȘȢ țȘȡȠȗȓȞȘȢ țĮȚ IJȠȣ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ ıIJȘ ıȣȖțȑȞIJȡȦıȘ IJȠȣ
ȕ-țĮȡȠIJȑȞȚȠȣ țĮȚ țĮșȠȡȓıIJȘțİ Ƞ ıȣȞįȣĮıµȩȢ IJȦȞ IJȚµȫȞ IJȘȢ ıȣȖțȑȞIJȡȦıȘȢ IJȠȣ
ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ (17,15 g/l), IJȘȢ țȘȡȠȗȓȞȘȢ (39,25 g/l) țĮȚ IJȠȣ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ
(9,04 g/l) IJȠȣ ȣʌȠıIJȡȫµĮIJȠȢ ʌĮȡĮȖȦȖȒȢ, Ƞ ȠʌȠȓȠȢ İʌȚIJȣȖȤȐȞİȚ IJȘ µȑȖȚıIJȘ
ıȣȖțȑȞIJȡȦıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ (2,82g/l).
ȅȚ ʌȜȘȡȠijȠȡȓİȢ ʌȠȣ ıȣȖțİȞIJȡȫșȘțĮȞ Įʌȩ IJȚȢ ʌȡȠĮȞĮijİȡșİȓıİȢ ʌİȚȡĮµĮIJȚțȑȢ
įȠțȚµȑȢ ȤȡȘıȚµȠʌȠȚȒșȘțĮȞ ȖȚĮ IJȘȞ ʌȡĮȖµĮIJȠʌȠȓȘıȘ ȗȣµȫıİȦȞ ıIJȠ İʌȓʌİįȠ IJȠȣ
İȡȖĮıIJȘȡȚĮțȠȪ ȕȚȠĮȞIJȚįȡĮıIJȒȡĮ. ȂİȜİIJȒșȘțİ Ș İʌȓįȡĮıȘ IJȠȣ ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ, IJȘȢ
țȘȡȠȗȓȞȘȢ, IJȠȣ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ țĮȚ IJȠȣ ȚıȠµȠȡȚĮțȠȪ µȓȖµĮIJȠȢ İȜĮȚȩȜĮįȠȣ,
ıȠȖȚȑȜĮȚȠȣ țĮȚ ȕĮµȕĮțȑȜĮȚȠȣ ıIJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora țĮȚ
ıIJȘ µȠȡijȠȜȠȖȓĮ IJȠȣ µȚțȡȠȠȡȖĮȞȚıµȠȪ ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ. ȉĮ ĮʌȠIJİȜȑıµĮIJĮ
ȑįİȚȟĮȞ ȩIJȚ Ș ʌȡȠıșȒțȘ ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ, țȘȡȠȗȓȞȘȢ Ȓ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ įİȞ
ȕİȜIJȓȦıĮȞ IJȘȞ ĮʌȩįȠıȘ IJȘȢ ȤȡȦıIJȚțȒȢ, İȞȫ İȣȞȩȘıĮȞ IJȠ ıȤȘµĮIJȚıµȩ µȣțȘȜȚĮțȫȞ
ȣijȫȞ µİ țİȞȠIJȩʌȚĮ, țİȞȫȞ țȣIJIJȐȡȦȞ țĮȚ İțijȣȜȚıµȑȞȦȞ țȣIJIJȐȡȦȞ. ǼʌȚʌȜȑȠȞ, Ș
ıȣȞİȡȖȚıIJȚțȒ įȡȐıȘ IJȠȣ ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ țĮȚ IJȠȣ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ İʌȑijİȡİ µȚțȡȒ
ĮȪȟȘıȘ ıIJȘȞ ʌĮȡĮȖȦȖȒ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ. Ǿ µİȖĮȜȪIJİȡȘ ʌĮȡĮȖȦȖȒ IJȠȣ
ȕ-țĮȡȠIJȑȞȚȠȣ (1,57 g/l) ʌĮȡĮIJȘȡȒșȘțİ µİ IJȠȞ ݵʌȜȠȣIJȚıµȩ IJȠȣ ȣʌȠıIJȡȫµĮIJȠȢ µİ
30,0 g/l ȚıȠµȠȡȚĮțȩ µȓȖµĮ İȜĮȚȩȜĮįȠȣ, ıȠȖȚȑȜĮȚȠȣ, ȕĮµȕĮțȑȜĮȚȠȣ țĮȚ 2,5 g/l
ĮȞIJȚȠȟİȚįȦIJȚțȩ (BHT). ȈIJȠ ȐȡȚıIJȠ ȣʌȩıIJȡȦµĮ ʌĮȡĮȖȦȖȒȢ ȕ-țĮȡȠIJȑȞȚȠȣ IJȠ ʌȠıȠıIJȩ
IJȦȞ ȐșȚțIJȦȞ µȣțȘȜȚĮțȫȞ ȣijȫȞ, IJȦȞ țİȞȫȞ țȣIJIJȐȡȦȞ, IJȦȞ İțijȣȜȚıµȑȞȦȞ țȣIJIJȐȡȦȞ
țĮȚ IJȦȞ ȗȣȖȠıʌȠȡȓȦȞ ıIJȠ ıȣȞȠȜȚțȩ ȟȘȡȩ ȕȐȡȠȢ IJȘȢ ȕȚȠµȐȗĮȢ ȒIJĮȞ 16,0%, 13,52%,
1,60% țĮȚ 51,46%, ĮȞIJȓıIJȠȚȤĮ.
Ǿ µİȜȑIJȘ ȖȚĮ IJȘ ȕİȜIJȓȦıȘ IJȘȢ ĮʌȠIJİȜİıµĮIJȚțȩIJȘIJĮȢ IJȘȢ ĮıȣȞİȤȠȪȢ ȗȪµȦıȘȢ
ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ ıȣȞİȤȓıIJȘțİ µİ IJȘ įȚİȡİȪȞȘıȘ IJȘȢ İʌȓįȡĮıȘȢ IJȘȢ IJĮȤȪIJȘIJĮȢ
ĮȞȐįİȣıȘȢ IJȠȣ ȣʌȠıIJȡȫµĮIJȠȢ țĮȚ IJȘȢ ʌĮȡȠȤȒȢ ĮȑȡĮ ıIJȠ ȗȣµȦIJȒȡĮ ıIJȘȞ ʌĮȡĮȖȦȖȒ
IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ țĮȚ ıIJȘ µȠȡijȠȜȠȖȓĮ IJȠȣ µȚțȡȠȠȡȖĮȞȚıµȠȪ. ǺȡȑșȘțİ ȩIJȚ ȠȚ
ʌĮȡĮʌȐȞȦ ʌĮȡȐȖȠȞIJİȢ İʌȚįȡȠȪȞ ıȘµĮȞIJȚțȐ ıIJȘ ıȣȖțȑȞIJȡȦıȘ țĮȚ ıIJȘȞ
ʌĮȡĮȖȦȖȚțȩIJȘIJĮ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ, ıIJȘ ȕȚȠµȐȗĮ țĮȚ ıIJȠȞ ȠȖțȠµİIJȡȚțȩ ıȣȞIJİȜİıIJȒ
µİIJĮijȠȡȐȢ µȐȗĮȢ. Ȃİ IJȘȞ ʌȡȠıĮȡµȠȖȒ ıIJĮIJȚıIJȚțȠȪ µȠȞIJȑȜȠȣ ʌȡȩȕȜİȥȘȢ
İȡµȘȞİȪIJȘțĮȞ ıİ ȚțĮȞȠʌȠȚȘIJȚțȩ ȕĮșµȩ ȠȚ İʌȚįȡȐıİȚȢ IJȘȢ IJĮȤȪIJȘIJĮȢ ĮȞȐįİȣıȘȢ țĮȚ
IJȘȢ ʌĮȡȠȤȒȢ ĮȑȡĮ ıIJȚȢ ʌĮȡĮʌȐȞȦ ʌĮȡĮµȑIJȡȠȣȢ IJȘȢ ȗȣµȦIJȚțȒȢ įȚİȡȖĮıȓĮȢ țĮȚ
țĮșȠȡȓıIJȘțİ Ƞ ıȣȞįȣĮıµȩȢ IJȦȞ ȐȡȚıIJȦȞ IJȚµȫȞ IJȠȣȢ, Ƞ ȠʌȠȓȠȢ IJȚȢ İʌȚIJȣȖȤȐȞİȚ. Ǿ
µȑȖȚıIJȘ IJȚµȒ IJȘȢ ıȣȖțȑȞIJȡȦıȘȢ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ (1,57 g/l) İʌȚIJİȪȤșȘțİ ıİ ȤĮµȘȜȑȢ
IJȚµȑȢ IJĮȤȪIJȘIJĮȢ ĮȞȐįİȣıȘȢ (150 rpm) țĮȚ ȣȥȘȜȑȢ IJȚµȑȢ ʌĮȡȠȤȒȢ ĮȑȡĮ (1,50 vvm),
İȞȫ ȠȚ µȑȖȚıIJİȢ IJȚµȑȢ IJȘȢ ʌĮȡĮȖȦȖȚțȩIJȘIJĮȢ IJȘȢ ȤȡȦıIJȚțȒȢ (0,079 g/l.d) țĮȚ IJȘȢ
ıȣȖțȑȞIJȡȦıȘȢ IJȘȢ ȕȚȠµȐȗĮȢ (24,7 g/l) ʌĮȡĮIJȘȡȒșȘțĮȞ ıİ ȣȥȘȜȑȢ IJȚµȑȢ IJĮȤȪIJȘIJĮȢ
ĮȞȐįİȣıȘȢ țĮȚ ʌĮȡȠȤȒȢ ĮȑȡĮ (500 rpm, 1,29 vvm țĮȚ 500 rpm, 1,43 vvm,
ĮȞIJȓıIJȠȚȤĮ). ǼʌȚʌȜȑȠȞ, Ș µȑȖȚıIJȘ IJȚµȒ IJȠȣ ȠȖțȠµİIJȡȚțȠȪ ıȣȞIJİȜİıIJȒ µİIJĮijȠȡȐȢ
µȐȗĮȢ (0,33 s
-1
) ʌĮȡĮIJȘȡȒșȘțİ ıİ IJĮȤȪIJȘIJĮ ĮȞȐįİȣıȘȢ 500 rpm țĮȚ ʌĮȡȠȤȒ ĮȑȡĮ
1,47 vvm. Ǿ µȠȡijȠȜȠȖȓĮ IJȠȣ µȚțȡȠȠȡȖĮȞȚıµȠȪ İʌȘȡİȐȗȠȞIJĮȞ ıȘµĮȞIJȚțȐ Įʌȩ IJȘȞ
IJĮȤȪIJȘIJĮ ĮȞȐįİȣıȘȢ țĮȚ IJȘȞ ʌĮȡȠȤȒ ĮȑȡĮ. ǵIJĮȞ Ș țĮȜȜȚȑȡȖİȚĮ ĮȞĮʌIJȣııȩIJĮȞ µİ
ȣȥȘȜȒ ʌĮȡȠȤȒ ĮȑȡĮ țĮȚ ȤĮµȘȜȒ IJĮȤȪIJȘIJĮ ĮȞȐįİȣıȘȢ İȣȞȠȠȪȞIJĮȞ Ƞ ıȤȘµĮIJȚıµȩȢ
µİȖȐȜȠȣ ĮȡȚșµȠȪ ȗȣȖȠıʌȠȡȓȦȞ µİ µȑIJȡȚȠ µȑȖİșȠȢ, İȞȫ ıİ ȑȞIJȠȞİȢ ıȣȞșȒțİȢ
ĮİȡȚıµȠȪ țĮȚ ĮȞȐįİȣıȘȢ IJȠȣ ȣʌȠıIJȡȫµĮIJȠȢ İȣȞȠȠȪȞIJĮȞ Ƞ ıȤȘµĮIJȚıµȩȢ µȚțȡȠȪ
ĮȡȚșµȠȪ ȗȣȖȠıʌȠȡȓȦȞ µİ µİȖȐȜȠ µȑȖİșȠȢ. ǼʌȓıȘȢ, Ș ĮȞȐʌIJȣȟȘ IJȠȣ µȚțȡȠȠȡȖĮȞȚıµȠȪ
ıİ ȑȞIJȠȞİȢ ıȣȞșȒțİȢ ĮİȡȚıµȠȪ țĮȚ ĮȞȐįİȣıȘȢ İȣȞȠȠȪıİ IJȠ ıȤȘµĮIJȚıµȩ µȣțȘȜȚĮțȫȞ
ȣijȫȞ µİ țİȞȠIJȩʌȚĮ, țİȞȫȞ țȣIJIJȐȡȦȞ țĮȚ İțijȣȜȚıµȑȞȦȞ țȣIJIJȐȡȦȞ.
ȈIJȠ IJİȜȚțȩ ıIJȐįȚȠ IJȘȢ İȡȖĮıȓĮȢ įȚİȡİȣȞȒșȘțİ Ș įȣȞĮIJȩIJȘIJĮ ʌĮȡĮȖȦȖȒȢ IJȠȣ
ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȘ µȑșȠįȠ IJȘȢ ĮıȣȞİȤȠȪȢ ȗȪµȦıȘȢ ȕȣșȠȪ µİ ıȣȞİȤȒ ʌȡȠıșȒțȘ
ȣʌȠıIJȡȫµĮIJȠȢ. ȂİȜİIJȒșȘțİ Ș İʌȓįȡĮıȘ IJȠȣ ȤȡȩȞȠȣ ȑȞĮȡȟȘȢ, IJȘȢ µİșȩįȠȣ țĮȚ IJȠȣ
ȡȣșµȠȪ IJȡȠijȠįȠıȓĮȢ IJȠȣ ȗȣµȦIJȒȡĮ µİ ȞȑȠ ȣʌȩıIJȡȦµĮ țĮșȫȢ țĮȚ IJȘȢ ıȪıIJĮıȘȢ IJȦȞ
ȣʌȠıIJȡȦµȐIJȦȞ ĮȞȐʌIJȣȟȘȢ țĮȚ IJȡȠijȠįȠıȓĮȢ ıIJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ. Ǿ
µȑȖȚıIJȘ ıȣȖțȑȞIJȡȦıȘ IJȘȢ ȤȡȦıIJȚțȒȢ (82,5 mg/l) İʌȚIJİȪȤșȘțİ ȩIJĮȞ ʌȡȠıIJȓșİȞIJĮȞ
ıȣȞİȤȫȢ ıIJȠ ȗȣµȦIJȒȡĮ ȞȑȠ ȣʌȩıIJȡȦµĮ Įʌȩ IJȘȞ ĮȡȤȒ IJȘȢ ȗȪµȦıȘȢ, ʌȠȣ ʌİȡȚİȓȤİ
30,0 g/l ȚıȠµȠȡȚĮțȩ µȓȖµĮ ijȣIJȚțȫȞ İȜĮȓȦȞ țĮȚ ȤĮµȘȜȑȢ ıȣȖțİȞIJȡȫıİȚȢ ĮȣȟȘIJȚțȫȞ
ʌĮȡĮȖȩȞIJȦȞ (ʌȡȦIJİȧȞȚțȒ ʌȘȖȒ ĮȗȫIJȠȣ, ȕȚIJĮµȓȞİȢ țĮȚ ȚȤȞȠıIJȠȚȤİȓĮ) µİ ȡȣșµȩ
ʌĮȡȠȤȒȢ 4,2 ml/h. Ǿ µȠȡijȠµİIJȡȚțȒ ĮȞȐȜȣıȘ IJȦȞ țȣIJIJȐȡȦȞ IJȠȣ µȚțȡȠȠȡȖĮȞȚıµȠȪ
ȑįİȚȟİ ȩIJȚ ıIJȠ ʌĮȡĮʌȐȞȦ ıȪıIJȘµĮ ȗȪµȦıȘȢ İȣȞȠİȓIJĮȚ Ƞ ıȤȘµĮIJȚıµȩȢ ȣȥȘȜȫȞ
ʌȠıȠıIJȫȞ µȣțȘȜȚĮțȫȞ ȣijȫȞ µİ țİȞȠIJȩʌȚĮ, țİȞȫȞ țȣIJIJȐȡȦȞ țĮȚ İțijȣȜȚıµȑȞȦȞ
țȣIJIJȐȡȦȞ ıIJȠ ıȣȞȠȜȚțȩ ȟȘȡȩ ȕȐȡȠȢ IJȘȢ ȕȚȠµȐȗĮȢ, ȜȩȖȦ IJȘȢ ȣȥȘȜȒȢ ıȣȖțȑȞIJȡȦıȘȢ
IJȠȣ Ǿ
2
O
2
țĮȚ IJȘȢ İʌĮȖȦȖȒȢ IJȘȢ įȡȐıȘȢ IJȦȞ İȜİȣșȑȡȦȞ ȡȚȗȫȞ ıIJȚȢ ʌĮȡĮʌȐȞȦ
ıȣȞșȒțİȢ ʌȠȣ ʌȚșĮȞȩȞ ȞĮ ʌȡȠțĮȜȠȪȞ ĮȞĮıIJȠȜȒ IJȘȢ įȡȐıȘȢ İȞȩȢ Ȓ ʌİȡȚııȠIJȑȡȦȞ
İȞȗȪµȦȞ IJȘȢ ȕȚȠıȪȞșİıȘȢ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ, țĮIJĮıIJȡȠijȒ IJȘȢ įȠµȒȢ IJȠȣ țȣIJIJĮȡȚțȠȪ
DNA țĮȚ ȐȜȜİȢ țȣIJIJĮȡȠIJȠȟȚțȑȢ įȡȐıİȚȢ.
ȉĮ ʌĮȡĮʌȐȞȦ įİįȠµȑȞĮ įİȓȤȞȠȣȞ ȩIJȚ IJȠ ıȪıIJȘµĮ IJȦȞ ĮȞĮțȚȞȠȪµİȞȦȞ ijȚĮȜȫȞ
İȓȞĮȚ IJȠ ʌȜȑȠȞ ĮʌȠIJİȜİıµĮIJȚțȩ ıȪıIJȘµĮ ȖȚĮ IJȘȞ ʌĮȡĮȖȦȖȒ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ Įʌȩ
ıȣȞșİIJȚțȩ ȣʌȩıIJȡȦµĮ µİ IJȠ B. trispora. ȉȠ ıȪıIJȘµĮ IJȘȢ ĮıȣȞİȤȠȪȢ ȗȪµȦıȘȢ ȑįȦıİ
țĮȜȪIJİȡĮ ĮʌȠIJİȜȑıµĮIJĮ ȦȢ ʌȡȠȢ IJȚȢ ȕĮıȚțȑȢ ʌĮȡĮµȑIJȡȠȣȢ IJȘȢ ȗȪµȦıȘȢ, ıİ ıȤȑıȘ
µİ IJȠ ıȪıIJȘµĮ IJȘȢ ĮıȣȞİȤȠȪȢ ȗȪµȦıȘȢ µİ ıȣȞİȤȒ ʌȡȠıșȒțȘ ȣʌȠıIJȡȫµĮIJȠȢ.


SUMMARY

In the present study the production of ȕ-carotene from synthetic medium by
mated cultures of B. trispora in submerged fermentation was investigated. For this
purpose different fermentation systems were used such as shake flasks and stirred
tank reactor (batch and fed-batch culture).
At the beginning stage of this study an improved method for the extraction of
ȕ-carotene from B. trispora was developed. The fermentation broth was steamed at
121
0
C for 20 min. ȕ-Carotene was removed from the biomass by extraction with
absolute ethanol at a ratio of 1:100 at 30
0
C for 2 h in a rotary shaker incubator at 300
rpm. The carotenoid pigment was completely removed from the cells after three
repeated extractions. The removal of ȕ-carotene from B. trispora was higher during
the first stage (75,94%), whereas in the second and third stage it was very low
(16,62% țĮȚ 7,44%, respectively).
The effect of inoculum, temperature, pH, light, carbon and nitrogen source,
nonionic surfactants, natural oils, fatty acids, inducers, precursors and antioxidant on
ȕ-carotene production by B. trispora in shake flask culture was investigated. The
highest concentration of ȕ-carotene was obtained in the medium (pH 7.0) inoculated
with one loop of each culture after 8 days of fermentation, at 26
0
C, in the presence
of daylight. Sucrose, starch and glycerol did not improve the production of ȕ-
carotene, while corn meal, soy protein acid hydrolysate and distiller’s solubles
slightly increased the production of the pigment. On the other hand, glucose, corn
steep liquor, Span 20, Tween 80, olive oil, soybean oil, cottonseed oil, oleic and
linoleic acid, kerosene and antioxidant (butylated hydroxytoluene, BHT),
significantly increased the production of ȕ-carotene. Also, a semi-automated image
analysis system was used to examine the relationship between morphological
changes of the fungus and the composition of the production medium. B. trispora is a
microfungus with a life-cycle involving hyphae (intact hyphae, vacuolated and
evacuated cells and degenerated hyphae), zygophores and zygospores in submerged
culture. As indicated from the results of the morphometric analysis, in the best
production medium the growing mycelium was composed primarily of zygospores
(79,84%), whereas the concentration of intact hyphae, vacuolated and evacuated cells
and degenerated hyphae were very low. A central composite design was employed to
determine the maximum ȕ-carotene production at optimum values for the process
variables (linoleic acid, kerosene and antioxidant). The fit of the model was found to
be good. Linoleic acid, kerosene and antioxidant had a strong linear effect on ȕ-
carotene production. The concentration of ȕ-carotene was significantly affected by
linoleic acid-kerosene and linoleic acid-antioxidant interactions as well as by the
negative quadratic effects of these variables. The interaction among kerosene-
antioxidant had no significant linear effect. The maximum ȕ-carotene concentration
(2.82 g/l) was obtained at concentrations of linoleic acid (17.15 g/l), kerosene (39.25
g/l) and antioxidant (9.04 g/l).
The effect of linoleic acid, kerosene, antioxidant and

a mixture of natural oils
(olive oil, soybean oil, and cottonseed oil) on the production of ȕ-carotene by B.
trispora and the morphological characteristics of the fungus in a stirred tank reactor
were investigated. The mixture of linoleic acid and antioxidant improved slightly the
production of ȕ-carotene, while the addition of linoleic acid, kerosene or antioxidant
alone to the medium had an inhibiting effect on ȕ-carotene production. The addition
to the medium of a mixture of natural oils and antioxidant increased significantly the
concentration of the pigment. B. trispora formed hyphae, zygophores and zygospores
during ȕ-carotene production. Morphology of the fungus was studied during growth
in submerged culture in a stirred tank reactor under different compositions of the
production medium using an image analysis system. Zygospores are the
morphological forms, which are responsible for pigment production. Maximum ȕ-
carotene concentration (1,57 g/l) was obtained in culture grown in medium
supplemented with 10,0 g/l each one of olive oil, soybean oil, cottonseed oil and 2,5
g/l of antioxidant. In this case, the morphological characteristics of B. trispora
recorded showed intact hyphae, evacuated cells, degenerated hyphae and zygospores
in percentages of about 16,0%, 13,52%, 1,60% țĮȚ 51,46%, respectively, in the
biomass dry weight.
The effect of aeration rate and agitation speed on ȕ-carotene production and
morphology of B. trispora in a stirred tank reactor was investigated. Both aeration
and agitation significantly affected ȕ-carotene concentration, productivity, biomass
and the volumetric mass transfer coefficient (K
L
a). The highest ȕ-carotene
concentration (1.57 g/l) was obtained at low impeller speed (150 rpm) and high
aeration rate (1.5 vvm). Also, maximum productivity (0.079 g/ld) and biomass dry
weight (24.7 g/l) were achieved at high agitation speed and aeration rate (500 rpm,
1.29 vvm and 500 rpm, 1,43 vvm, respectively). The highest value of K
L
a (0.33 s
-1
)
was observed at high agitation speed (500 rpm) and high aeration rate (1.47 vvm).
The experiments were arranged according to a central composite statistical design.
Response surface methodology was used to describe the effect of impeller speed and
aeration rate on the most important fermentation parameters. In all cases, the fit of
the model was found to be good. All fermentation parameters (except biomass
concentration) were strongly affected by the interactions among the operation
variables. ȕ-Carotene concentration and productivity were significantly influenced by
the aeration, agitation, and by the positive or negative quadratic effect of the aeration
rate. Biomass concentration was principally related to the aeration rate, agitation
speed, and the positive or negative quadratic effect of the impeller speed and aeration
rate, respectively. Finally, the volumetric mass transfer coefficient was characterized
by the significant effect of the agitation speed, while the aeration rate had a small
effect on K
L
a. Microscopic examination showed that at high aeration rate (1.5 vvm)
and low agitation speed (150 rpm) was observed a great number of zygospores with
long diameter, while at high aeration rate and agitation speed (1.5 vvm, 500 rpm) a
small number of zygospores with extremely wide diameter were observed. At
intermediate values of aeration rate and impeller speed (1.0 vvm, 325 rpm) was
observed a common number of zygospores with average diameter. The maximum
concentration of ȕ-carotene (1.57 g/l) was observed at the same conditions where the
highest percentage of zygospores in the biomass dry weight were observed.
At the final stage of this study the potential of fed-batch culture as
fermentation system for ȕ-carotene production from a chemically defined medium by
B. trispora was investigated. The effect of the feeding medium composition, time of
addition of feeding medium, feeding rate and fermentation system on the production
of ȕ-carotene was studied. Also, a semi-automatic image analysis system was used to
determine the morphology of the microorganism, and a correlation between
morphology of the fungi and the concentration of the pigment was proposed. The
maximum ȕ-carotene concentration (82,5 mg/l) was obtained using the best batch
medium found in previous experiments and the feeding medium containing natural
oils at concentration 1% of each one and low concentration of growth factors (amino
acids, vitamins and minerals) at feeding rate of 4,2 ml/h from the beginning of the
fermentation. High percentages of vacuolated hyphae, evacuated cells and
degenerated hyphae of the microorganism were observed during the fermentation.
This was due to the formation of high amount of reactive oxygen species (e.g.
superoxide, hydrogen peroxide, hydroxyl radical) under conditions of exogenous
oxidative stress, by exposure of the microorganism to high dissolved oxygen
concentration. These oxidants have a detrimental function in the microorganism
mediating damage of cellular membranes and enzymes, DNA damage, mutagenesis
and inhibition of membrane transport processes.
The above results showed that shake flask culture was the best fermentation
system for the production of ȕ-carotene from synthetic medium by B. trispora.
Further investigation should focus on the improvement of ȕ-carotene production
under batch and fed-batch conditions in a larger fermentation scale.




ǼȊȋǹȇǿȈȉǿǼȈ

Ǿ ʌĮȡȠȪıĮ įȚįĮțIJȠȡȚțȒ įȚĮIJȡȚȕȒ İțʌȠȞȒșȘțİ ıIJȠ ǼȡȖĮıIJȒȡȚȠ ǼʌİȟİȡȖĮıȓĮȢ
țĮȚ ȂȘȤĮȞȚțȒȢ ȉȡȠijȓµȦȞ IJȠȣ ȉȠµȑĮ ǼʌȚıIJȒµȘȢ țĮȚ ȉİȤȞȠȜȠȖȓĮȢ ȉȡȠijȓµȦȞ IJȠȣ
ȉµȒµĮIJȠȢ īİȦʌȠȞȓĮȢ IJȠȣ ǹȡȚıIJȠIJİȜİȓȠȣ ȆĮȞİʌȚıIJȘµȓȠȣ ĬİııĮȜȠȞȓțȘȢ.
ȉȘȞ İʌȓȕȜİȥȘ IJȘȢ įȚĮIJȡȚȕȒȢ İȓȤİ Ƞ ȀĮșȘȖȘIJȒȢ IJȠȣ ȉµȒµĮIJȠȢ īİȦʌȠȞȓĮȢ IJȠȣ
ǹȡȚıIJȠIJİȜİȓȠȣ ȆĮȞİʌȚıIJȘµȓȠȣ ĬİııĮȜȠȞȓțȘȢ ț. ȉȡȚĮȞIJȐijȣȜȜȠȢ ȇȠȣțȐȢ, IJȠȞ ȠʌȠȓȠ
İȣȤĮȡȚıIJȫ șİȡµȐ ȖȚĮ IJȘȞ țĮșȠįȒȖȘıȘ țĮȚ IJȘȞ İȞșȐȡȡȣȞıȘ ʌȠȣ µȠȣ ʌĮȡİȓȤİ țĮșȫȢ
țĮȚ ȖȚĮ IJȚȢ ʌȠȜȪIJȚµİȢ ȚįȑİȢ țĮȚ ıȣµȕȠȣȜȑȢ IJȠȣ țĮIJȐ IJȘȞ İțʌȩȞȘıȘ țĮȚ IJȘ ıȣȖȖȡĮijȒ
IJȘȢ İȡȖĮıȓĮȢ ĮȣIJȒȢ. Ǿ ıȣµµİIJȠȤȒ IJȠȣ ıIJȘȞ ʌȡĮȖµĮIJȠʌȠȓȘıȘ IJȘȢ įȚĮIJȡȚȕȒȢ ȒIJĮȞ
țĮIJĮȜȣIJȚțȒ ĮijȠȪ ıȣȞȑȕĮȜȜİ ıȘµĮȞIJȚțȐ ıIJȘȞ İʌȓȜȣıȘ IJȩıȠ İʌȚıIJȘµȠȞȚțȫȞ ȩıȠ țĮȚ
ȐȜȜȦȞ ʌȡȠȕȜȘµȐIJȦȞ µİ ıțȠʌȩ IJȘȞ İʌȚțȑȞIJȡȦıȘ µȠȣ ıIJȘȞ ȑȡİȣȞĮ.
ǿįȚĮȓIJİȡĮ İȣȤĮȡȚıIJȫ IJȠ µȑȜȠȢ IJȘȢ IJȡȚµİȜȠȪȢ ıȣµȕȠȣȜİȣIJȚțȒȢ µȠȣ İʌȚIJȡȠʌȒȢ
ț. ȆĮȡșȑȞĮ ȀȠIJȗİțȓįȠȣ-ȇȠȣțȐ, ȀĮșȘȖȒIJȡȚĮ IJȠȣ ȉµȒµĮIJȠȢ īİȦʌȠȞȓĮȢ IJȠȣ
ǹȡȚıIJȠIJİȜİȓȠȣ ȆĮȞİʌȚıIJȘµȓȠȣ ĬİııĮȜȠȞȓțȘȢ, ȖȚĮ IJȘȞ İȞİȡȖȒ, ıȣȞİȤȒ țĮȚ
ĮȞİțIJȓµȘIJȘ ıȣµµİIJȠȤȒ IJȘȢ ıİ ȩȜİȢ IJȚȢ ijȐıİȚȢ įȚİȟĮȖȦȖȒȢ ĮȣIJȒȢ IJȘȢ İȡȖĮıȓĮȢ țĮșȫȢ
țĮȚ ȖȚĮ IJȠ ȑȞIJȠȞȠ İȞįȚĮijȑȡȠȞ IJȘȢ, IJȠ ȠʌȠȓȠ ĮȞĮȕȐșµȚıİ țĮȚ įȚİȣțȩȜȣȞİ IJȘȞ ʌĮȡȠȪıĮ
İȡȖĮıȓĮ. ǼʌȓıȘȢ, İȣȤĮȡȚıIJȫ IJȠ µȑȜȠȢ IJȘȢ IJȡȚµİȜȠȪȢ ıȣµȕȠȣȜİȣIJȚțȒȢ µȠȣ İʌȚIJȡȠʌȒȢ
ț. ȂĮȡȓĮ ȁȚĮțȠʌȠȪȜȠȣ-ȀȣȡȚĮțȓįȠȣ, ȀĮșȘȖȒIJȡȚĮ IJȠȣ ȉµȒµĮIJȠȢ ȋȘµȚțȫȞ
ȂȘȤĮȞȚțȫȞ IJȠȣ ǹȡȚıIJȠIJİȜİȓȠȣ ȆĮȞİʌȚıIJȘµȓȠȣ ĬİııĮȜȠȞȓțȘȢ, ȖȚĮ IJȚȢ ʌȠȜȪIJȚµİȢ
ıȣµȕȠȣȜȑȢ țĮȚ ʌĮȡĮIJȘȡȒıİȚȢ țĮIJȐ IJȘ įȚİȟĮȖȦȖȒ ĮȣIJȒȢ IJȘȢ İȡȖĮıȓĮȢ.
ȉȚȢ İȣȤĮȡȚıIJȓİȢ µȠȣ șȑȜȦ ȞĮ İțijȡȐıȦ ıIJȠȞ ț. ǻȘµȒIJȡȘ ȀȣȡȚĮțȓįȘ,
ȀĮșȘȖȘIJȒ IJȠȣ ȉµȒµĮIJȠȢ ȋȘµİȓĮȢ IJȠȣ ǹȡȚıIJȠIJİȜİȓȠȣ ȆĮȞİʌȚıIJȘµȓȠȣ ĬİııĮȜȠȞȓțȘȢ,
ȖȚĮ IJȚȢ ʌȠȜȪIJȚµİȢ ıȣµȕȠȣȜȑȢ IJȠȣ.
ǼʌȓıȘȢ, İȣȤĮȡȚıIJȫ ȩȜȠȣȢ IJȠȣȢ µİIJĮʌIJȣȤȚĮțȠȪȢ ijȠȚIJȘIJȑȢ IJȠȣ ǼȡȖĮıIJȘȡȓȠȣ
ǼʌİȟİȡȖĮıȓĮȢ țĮȚ ȂȘȤĮȞȚțȒȢ ȉȡȠijȓµȦȞ țĮȚ IJȠȣ ǼȡȖĮıIJȘȡȓȠȣ ȊȖȚİȚȞȒȢ țĮȚ
ȂȚțȡȠȕȚȠȜȠȖȓĮȢ ȉȡȠijȓµȦȞ IJȠȣ ȉȠµȑĮ ǼʌȚıIJȒµȘȢ țĮȚ ȉİȤȞȠȜȠȖȓĮȢ ȉȡȠijȓµȦȞ IJȠȣ
ȉµȒµĮIJȠȢ īİȦʌȠȞȓĮȢ IJȠȣ ǹȡȚıIJȠIJİȜİȓȠȣ ȆĮȞİʌȚıIJȘµȓȠȣ ĬİııĮȜȠȞȓțȘȢ, ȖȚĮ IJȘȞ
ȣʌȠıIJȒȡȚȟȘ IJȠȣȢ țĮIJȐ IJȘ įȚȐȡțİȚĮ IJȘȢ įȚĮIJȡȚȕȒȢ.



I
ȆǼȇǿǼȋȅȂǼȃǹ


ıİȜ.
ǼȊȇǼȉǾȇǿȅ ȆǿȃǹȀȍȃ..................................................................................

VIII
ǼȊȇǼȉǾȇǿȅ ȈȋǾȂǹȉȍȃ..............................................................................

XII
ǼȊȇǼȉǾȇǿȅ ǼǿȀȅȃȍȃ...................................................................................

XX
ȀǹȉǹȁȅīȅȈ ǻǾȂȅȈǿǼȊȂǼȃȍȃ ǼȇīǹȈǿȍȃ..........................................

XXIV
ȀǼĭǹȁǹǿȅ 1: ǼǿȈǹīȍīǾ...........................................................................

1
ȀǼĭǹȁǹǿȅ 2: ǹȃǹȈȀȅȆǾȈǾ ǺǿǺȁǿȅīȇǹĭǿǹȈ……………………. 6
2.1 ȂȚțȡȠȠȡȖĮȞȚıµȠȓ ʌȠȣ ȋȡȘıȚµȠʌȠȚȠȪȞIJĮȚ ȖȚĮ IJȘȞ ȆĮȡĮȖȦȖȒ
ȕ-ȀĮȡȠIJȑȞȚȠȣ…………………………………………………………

6
2.1.1 ȂȪțȘIJİȢ…………………………………………………………….. 6
2.1.2 ǽȪµİȢ………………………………………………………………... 16
2.1.3 ǺĮțIJȒȡȚĮ……………………………………………………………. 17
2.1.4 DZȜȖȘ………………………………………………………………… 18
2.1.5 ǿıIJȠțĮȜȜȚȑȡȖİȚİȢ……………………………………………………. 19
2.2 ǺȚȠıȪȞșİıȘ ȕ-ȀĮȡȠIJȑȞȚȠȣ……………………………………………… 20
2.3 īİȞİIJȚțȒ ȉȡȠʌȠʌȠȓȘıȘ ȂȚțȡȠȠȡȖĮȞȚıµȫȞ ʌȠȣ ȋȡȘıȚµȠʌȠȚȠȪȞIJĮȚ ȖȚĮ
IJȘȞ ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ.................................................................
24
2.4
a
ȆĮȡȐȖȠȞIJİȢ ʌȠȣ ǼʌȘȡİȐȗȠȣȞ IJȘȞ ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ..........….. 28
2.4.1 ĬȡİʌIJȚțȐ ȈȣıIJĮIJȚțȐ țĮȚ ǿȤȞȠıIJȠȚȤİȓĮ................................................ 28
2.4.2 ǼʌȚijĮȞİȚȠįȡĮıIJȚțȑȢ ȅȣıȓİȢ țĮȚ ǹȞIJȚȠȟİȚįȦIJȚțȐ............................... 37
2.4.3 ǼȞİȡȖȠʌȠȚȘIJȑȢ..................................................................................... 38
2.4.4 ǹȞĮıIJȠȜİȓȢ.......................................................................................... 40
2.4.5 pH........................................................................................................ 41 2.4.6 ĬİȡµȠțȡĮıȓĮ....................................................................................... 42
2.4.7 ǹțIJȚȞȠȕȠȜȓĮ........................................................................................ 43
II
2.4.8 ȋȡȩȞȠȢ ǽȪµȦıȘȢ................................................................................ 44
2.4.9 ǹİȡȚıµȩȢ țĮȚ ǹȞȐįİȣıȘ..................................................................... 45
2.5 ȂȑșȠįȠȚ ǽȪµȦıȘȢ ʌȠȣ ȋȡȘıȚµȠʌȠȚȠȪȞIJĮȚ ȖȚĮ IJȘȞ ȆĮȡĮȖȦȖȒ
ȕ-ȀĮȡȠIJȑȞȚȠȣ............................................................................................
46
2.5.1 ǹıȣȞİȤȒȢ ǽȪµȦıȘ ǺȣșȠȪ...................................................................
2.5.1.1 ǹȞĮțȚȞȠȪµİȞİȢ ĭȚȐȜİȢ...............................................................
46 46
2.5.1.2 ǽȣµȦIJȒȡĮȢ µİ ȈȪıIJȘµĮ ǹȞȐįİȣıȘȢ.......................................... 47
2.5.1.3 ǺȚȠĮȞIJȚįȡĮıIJȒȡĮȢ ȈIJȒȜȘȢ......................................................... 48
2.5.2 ǹıȣȞİȤȒȢ ǽȪµȦıȘ ǺȣșȠȪ µİ ȈȣȞİȤȒ ȆȡȠıșȒțȘ ȊʌȠıIJȡȫµĮIJȠȢ..... 48
2.6
a
ǹȞȐțIJȘıȘ țĮȚ ȀĮșĮȡȚıµȩȢ IJȠȣ ȕ-ȀĮȡȠIJȑȞȚȠȣ....................................… 50
2.7
a
ǼµʌȠȡȚțȒ ǹȟȚȠȜȩȖȘıȘ IJȘȢ ȆĮȡĮȖȦȖȒȢ ȕ-ȀĮȡȠIJȑȞȚȠȣ µİ
ĮĮĮ
ȂȚțȡȠȠȡȖĮȞȚıµȠȪȢ...................................................................................
53

ȀǼĭǹȁǹǿȅ 3: ȊȁǿȀǹ Ȁǹǿ ȂǼĬȅǻȅǿ.....................................................
54
3.1 ȈIJİȜȑȤȘ ȂȚțȡȠȠȡȖĮȞȚıµȫȞ ʌȠȣ ȋȡȘıȚµȠʌȠȚȒșȘțĮȞ ȖȚĮ IJȘȞ ȆĮȡĮȖȦȖȒ
ȕ-ȀĮȡȠIJȑȞȚȠȣ...........................................................................................
54
3.1.1 ȆİȡȚȖȡĮijȒ ȈIJİȜİȤȫȞ.......................................................................... 54
3.1.2 ǻȚĮIJȒȡȘıȘ ȈIJİȜİȤȫȞ.......................................................................... 54
3.2 ǺİȜIJȚıIJȠʌȠȓȘıȘ IJȘȢ ǼțȤȪȜȚıȘȢ IJȠȣ ȕ-ȀĮȡȠIJȑȞȚȠȣ Įʌȩ IJĮ ȀȪIJIJĮȡĮ IJȠȣ
B. trispora.................................................................................................
55
3.2.1 ǹȞȐʌIJȣȟȘ ǼµȕȠȜȓȠȣ............................................................................ 55
3.2.2 ȊʌȩıIJȡȦµĮ ȆĮȡĮȖȦȖȒȢ țĮȚ ȈȣȞșȒțİȢ ǽȪµȦıȘȢ............................... 55
3.2.3 ȂȑșȠįȠȚ ǼțȤȪȜȚıȘȢ............................................................................. 55
3.2.4 ȀĮIJİȡȖĮıȓĮ IJȠȣ ȊȖȡȠȪ ǽȪµȦıȘȢ ʌȡȚȞ IJȘȞ ǼțȤȪȜȚıȘ IJȠȣ
ȕ-ȀĮȡȠIJȑȞȚȠȣ...................................................................................
58
3.3 ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ Įʌo ȈȣȞșİIJȚțȩ ȊʌȩıIJȡȦµĮ µİ IJȠ B. trispora
ıİ ǹȞĮțȚȞȠȪµİȞİȢ ȀȦȞȚțȑȢ ĭȚȐȜİȢ..........................................................
59
3.3.1 ǹȞȐʌIJȣȟȘ ǼµȕȠȜȓȠȣ............................................................................ 59
3.3.2 ǼʌȓįȡĮıȘ IJȦȞ ȆĮȡĮȖȩȞIJȦȞ

IJȘȢ ǽȪµȦıȘȢ ıIJȘȞ ȆĮȡĮȖȦȖȒ
ȕ-ȀĮȡȠIJȑȞȚȠȣ...................................................................................
59
3.3.2.1 ȈȣȖțȑȞIJȡȦıȘ ǼµȕȠȜȓȠȣ............................................................. 59
3.3.2.2 ǹȡȤȚțȩ pH.................................................................................. 60
3.3.2.3 ĬİȡµȠțȡĮıȓĮ.............................................................................. 60
III
3.3.2.4 ȋȡȩȞȠȢ....................................................................................... 60
3.3.2.5 ĭȦIJȚıµȩȢ.................................................................................... 60
3.3.3 ȈȪıIJĮıȘ IJȠȣ ȊʌȠıIJȡȫµĮIJȠȢ.............................................................. 61
3.3.3.1 ȆȘȖȒ DZȞșȡĮțĮ........................................................................... 61
3.3.3.2 ȆȘȖȒ ǹȗȫIJȠȣ............................................................................. 61
3.3.3.3 ǼʌȚijĮȞİȚȠįȡĮıIJȚțȑȢ ȅȣıȓİȢ țĮȚ ǹȞIJȚȠȟİȚįȦIJȚțȐ...................... 61
3.3.3.4 ĭȣIJȚțȐ DzȜĮȚĮ țĮȚ ǹțȩȡİıIJĮ ȁȚʌĮȡȐ ȅȟȑĮ............................... 62
3.3.3.5 ȆȡȩįȡȠµİȢ ȅȣıȓİȢ țĮȚ ǼȞİȡȖȠʌȠȚȘIJȑȢ...................................... 62
3.3.4 ǹȞĮȜȣIJȚțȠȓ ȆȡȠıįȚȠȡȚıµȠȓ................................................................. 62
3.3.4.1 ȆȡȠıįȚȠȡȚıµȩȢ ǺȚȠµȐȗĮȢ.......................................................... 62
3.3.4.2 ȆȡȠıįȚȠȡȚıµȩȢ ȈĮțȤȐȡȦȞ......................................................... 63
3.3.4.3 ȆȡȠıįȚȠȡȚıµȩȢ pH..................................................................... 64
3.3.4.4 ȆȡȠıįȚȠȡȚıµȩȢ ȕ-ȀĮȡȠIJȑȞȚȠȣ................................................... 64
3.3.4.5 ȆȡȠıįȚȠȡȚıµȩȢ ȊʌİȡȠȟİȚįȓȠȣ IJȠȣ ȊįȡȠȖȩȞȠȣ.......................... 65
3.3.5 ǹȞȐʌIJȣȟȘ ȂĮșȘµĮIJȚțȠȪ ȂȠȞIJȑȜȠȣ ȆȠȜȜĮʌȜȒȢ ȆĮȜȚȞįȡȩµȘıȘȢ µİ
IJȘ ȂİșȠįȠȜȠȖȓĮ IJȘȢ ǼʌȚijĮȞİȚĮțȒȢ ǹʌȩțȡȚıȘȢ..................................
70
3.3.5.1 ȆİȚȡĮµĮIJȚțȩȢ ȈȤİįȚĮıµȩȢ.......................................................... 70
3.3.5.2 ȈȣȜȜȠȖȒ ǻİįȠµȑȞȦȞ.................................................................. 72
3.3.5.3 ǹȞȐȜȣıȘ ǻİįȠµȑȞȦȞ țĮȚ ȂĮșȘµĮIJȚțȒ ȆİȡȚȖȡĮijȒ................... 72
3.3.5.4 ǹȟȚȠȜȩȖȘıȘ IJȠȣ ȂȠȞIJȑȜȠȣ........................................................ 73
3.3.6 ȂİȜȑIJȘ IJȦȞ ȂȠȡijȠȜȠȖȚțȫȞ ȋĮȡĮțIJȘȡȚıIJȚțȫȞ IJȠȣ B. trispora µİ
ȌȘijȚĮțȒ ǹȞȐȜȣıȘ ǼȚțȩȞĮȢ................................................................
77
3.3.6.1 ȆİȡȚȖȡĮijȒ IJȠȣ ȈȣıIJȒµĮIJȠȢ ʌȠȣ ȋȡȘıȚµȠʌȠȚȒșȘțİ ȖȚĮ IJȘ
ȌȘijȚĮțȒ ǹʌİȚțȩȞȘıȘ țĮȚ ǼʌİȟİȡȖĮıȓĮ IJȘȢ ǼȚțȩȞĮȢ...........
77
3.3.6.2 ǺĮıȚțȑȢ ȆĮȡȐµİIJȡȠȚ IJȘȢ ȂȠȡijȠµİIJȡȚțȒȢ ǹȞȐȜȣıȘȢ IJȘȢ
ȌȘijȚĮțȒȢ ǼȚțȩȞĮȢ..................................................................
78
3.4 ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ Įʌȩ ȈȣȞșİIJȚțȩ ȊʌȩıIJȡȦµĮ µİ IJȠ B. trispora
ıİ ǹıȣȞİȤȒ ǽȪµȦıȘ ǺȣșȠȪ ıİ ǽȣµȦIJȒȡĮ ǹȞȐįİȣıȘȢ.........................
81
3.4.1 ȆİȡȚȖȡĮijȒ IJȘȢ ȈȣıțİȣȒȢ.................................................................... 81
3.4.2 ȆȡȠİIJȠȚµĮıȓĮ IJȠȣ ȊʌȠıIJȡȫµĮIJȠȢ țĮȚ ȈȣȞșȒțİȢ ȀĮȜȜȚȑȡȖİȚĮȢ IJȠȣ
ȂȚțȡȠȠȡȖĮȞȚıµȠȪ...............................................................................
86
3.4.3 ǼʌȓįȡĮıȘ IJȘȢ ȈȪıIJĮıȘȢ IJȠȣ ȊʌȠıIJȡȫµĮIJȠȢ ıIJȘȞ ȆĮȡĮȖȦȖȒ
ȕ-ȀĮȡȠIJȑȞȚȠȣ...................................................................................
86
IV
3.4.4 ǼʌȓįȡĮıȘ IJȠȣ ǹİȡȚıµȠȪ țĮȚ IJȘȢ ǹȞȐįİȣıȘȢ ıIJȘȞ ȆĮȡĮȖȦȖȒ
ȕ-ȀĮȡȠIJȑȞȚȠȣ...................................................................................
87
3.4.5
ȥ
ȆȡȠıįȚȠȡȚıµȩȢ ȈȣȖțȑȞIJȡȦıȘȢ țĮȚ ȆĮȡĮȖȦȖȚțȩIJȘIJĮȢ
ȕ-ȀĮȡȠIJȑȞȚȠȣ, ǺȚȠµȐȗĮȢ, ȈĮțȤȐȡȦȞ, Ǿ
2
ȅ
2
, pH, ǻȚĮȜȣIJȠȪ
ȅȟȣȖȩȞȠȣ țĮȚ ȀĮIJĮȞȐȜȦıȘȢ ȈĮțȤȐȡȦȞ..........................................
87
3.4.6 ȂİȜȑIJȘ IJȦȞ ȂȠȡijȠȜȠȖȚțȫȞ ȋĮȡĮțIJȘȡȚıIJȚțȫȞ IJȠȣ B. trispora µİ
Į
ȌȘijȚĮțȒ ǹȞȐȜȣıȘ ǼȚțȩȞĮȢ..............................................................
88
3.4.7 ǹȞȐʌIJȣȟȘ ȂĮșȘµĮIJȚțȫȞ ȂȠȞIJȑȜȦȞ ȆȠȜȜĮʌȜȒȢ ȆĮȜȚȞįȡȩµȘıȘ µİ
IJȘ ȂİșȠįȠȜȠȖȓĮ IJȘȢ ǼʌȚijĮȞİȚĮțȒȢ ǹʌȩțȡȚıȘȢ...............................
91
3.4.7.1 ȆİȚȡĮµĮIJȚțȩȢ ȈȤİįȚĮıµȩȢ…...................................................... 91
3.4.7.2 ȈȣȜȜȠȖȒ ǻİįȠµȑȞȦȞ.................................................................. 92
3.4.7.3 ǹȞȐȜȣıȘ ǻİįȠµȑȞȦȞ țĮȚ ȂĮșȘµĮIJȚțȒ ȆİȡȚȖȡĮijȒ................... 94
3.4.7.4 ǹȟȚȠȜȩȖȘıȘ IJȠȣ ȂȠȞIJȑȜȠȣ........................................................ 94
3.5 ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ Įʌȩ ȈȣȞșİIJȚțȩ ȊʌȩıIJȡȦµĮ µİ IJȠ B. trispora
ıİ ǹıȣȞİȤȒ ǽȪµȦıȘ ǺȣșȠȪ µİ ȈȣȞİȤȒ ȆȡȠıșȒțȘ ȊʌȠıIJȡȫµĮIJȠȢ ıİ
ǽȣµȦIJȒȡĮ ǹȞĮįİȪıİȦȢ............................................................................
100
3.5.1 ȆİȡȚȖȡĮijȒ IJȘȢ ȈȣıțİȣȒȢ.................................................................... 100
3.5.2 ȈȣȞșȒțİȢ ǹȞȐʌIJȣȟȘȢ ȂȚțȡȠȠȡȖĮȞȚıµȠȪ țĮȚ ǹȡȤȒ ȁİȚIJȠȣȡȖȓĮȢ IJȘȢ
ǹıȣȞİȤȠȪȢ ǽȪµȦıȘȢ ǺȣșȠȪ µİ ȈȣȞİȤȒ ȆȡȠıșȒțȘ
ȊʌȠıIJȡȫµĮIJȠȢ....................................................................................
100
3.5.3
ĮĮ
ȆȡȠıįȚȠȡȚıµȩȢ ȈȣȖțȑȞIJȡȦıȘȢ țĮȚ ȆĮȡĮȖȦȖȚțȩIJȘIJĮȢ
ȕ-ȀĮȡȠIJȑȞȚȠȣ, ǺȚȠµȐȗĮȢ, ȈĮțȤȐȡȦȞ, Ǿ
2
ȅ
2
, pH, ǻȚĮȜȣIJȠȪ
ȅȟȣȖȩȞȠȣ țĮȚ ȀĮIJĮȞȐȜȦıȘȢ ȈĮțȤȐȡȦȞ..........................................
101
3.5.4 ȂİȜȑIJȘ IJȦȞ ȂȠȡijȠȜȠȖȚțȫȞ ȋĮȡĮțIJȘȡȚıIJȚțȫȞ IJȠȣ B. trispora µİ
ȌȘijȚĮțȒ ǹȞȐȜȣıȘ ǼȚțȩȞĮȢ................................................................
101

ȀǼĭǹȁǹǿȅ 4: ǹȆȅȉǼȁǼȈȂǹȉǹ Ȁǹǿ ȈȊǽǾȉǾȈǾ.............................
103
4.1 ǺİȜIJȚıIJȠʌȠȓȘıȘ IJȘȢ ǼțȤȪȜȚıȘȢ IJȠȣ ȕ-ȀĮȡȠIJȑȞȚȠȣ Įʌȩ IJĮ ȀȪIJIJĮȡĮ IJȠȣ
B. trispora.................................................................................................
103
4.1.1 ǼʌȓįȡĮıȘ IJȠȣ ȋȡȩȞȠȣ......................................................................... 103
4.1.2 ǼʌȓįȡĮıȘ IJȘȢ ĬİȡµȠțȡĮıȓĮȢ.............................................................. 107


V
4.1.3 ǼʌȓįȡĮıȘ IJȘȢ ȈȣȖțȑȞIJȡȦıȘȢ IJȘȢ ǹȚșĮȞȩȜȘȢ țĮȚ IJȘȢ ǹȞĮȜȠȖȓĮȢ
ǵȖțȠȣ ǹȚșĮȞȩȜȘȢ/ȄȘȡȠȪ ǺȐȡȠȣȢ ǺȚȠµȐȗĮȢ......................................
110
4.1.4 ǼʌȓįȡĮıȘ IJȦȞ ǻȚĮįȠȤȚțȫȞ ǼțȤȣȜȓıİȦȞ............................................. 113
4.1.5 ǼʌȓįȡĮıȘ IJȘȢ ȀĮIJİȡȖĮıȓĮȢ IJȠȣ ȊȖȡȠȪ IJȘȢ ǽȪµȦıȘȢ........................ 115
4.2 ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ Įʌo ȈȣȞșİIJȚțȩ ȊʌȩıIJȡȦµĮ µİ IJȠ B. trispora
ıİ ǹȞĮțȚȞȠȪµİȞİȢ ȀȦȞȚțȑȢ ĭȚȐȜİȢ..........................................................
118
4.2.1 ǼʌȓįȡĮıȘ IJȠȣ ǼµȕȠȜȓȠȣ ıIJȘȞ ȆĮȡĮȖȦȖȒ IJȠȣ ȕ-ȀĮȡȠIJȑȞȚȠȣ țĮȚ ıIJȘ
ȂȠȡijȠȜȠȖȓĮ IJȠȣ B. trispora...............................................................
118
4.2.2 ȂİIJĮȕȠȜȒ IJȦȞ ȀȚȞȘIJȚțȫȞ ȆĮȡĮµȑIJȡȦȞ IJȘȢ ǽȪµȦıȘȢ µİ IJȠ
ȋȡȩȞȠ...............................................................................................
125
4.2.3 ǼʌȓįȡĮıȘ IJȘȢ ĬİȡµȠțȡĮıȓĮȢ ıIJȚȢ ȀȚȞȘIJȚțȑȢ ȆĮȡĮµȑIJȡȠȣȢ IJȘȢ
ǽȪµȦıȘȢ...........................................................................................
128
4.2.4 ǼʌȓįȡĮıȘ IJȠȣ ǹȡȤȚțȠȪ pH IJȠȣ ȊʌȠıIJȡȫµĮIJȠȢ ıIJȚȢ ȀȚȞȘIJȚțȑȢ
ȆĮȡĮµȑIJȡȠȣȢ IJȘȢ ǽȪµȦıȘȢ.............................................................
131
4.2.5 ǼʌȓįȡĮıȘ IJȠȣ ĭȣıȚțȠȪ ĭȦIJȩȢ ıIJȘȞ ǹȞȐʌIJȣȟȘ IJȠȣ B. trispora țĮȚ
ıIJȘȞ ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ...........................................................
134
4.2.6 ǼʌȓįȡĮıȘ IJȘȢ ȆȘȖȒȢ DZȞșȡĮțĮ ıIJȘȞ ǹȞȐʌIJȣȟȘ IJȠȣ B. trispora țĮȚ
ıIJȘȞ ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ...........................................................
137
4.2.7 ǼʌȓįȡĮıȘ IJȘȢ ȆȘȖȒȢ ǹȗȫIJȠȣ ıIJȘȞ ǹȞȐʌIJȣȟȘ IJȠȣ B. trispora țĮȚ
ıIJȘȞ ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ...........................................................
140
4.2.8 ǼʌȓįȡĮıȘ IJȦȞ ǼʌȚijĮȞİȚȠįȡĮıIJȚțȫȞ ȅȣıȚȫȞ ıIJȘ ȂȠȡijȠȜȠȖȓĮ IJȠȣ
B. trispora țĮȚ ıIJȘȞ ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ...................................
145
4.2.9 ǼʌȓįȡĮıȘ IJȦȞ ĭȣIJȚțȫȞ ǼȜĮȓȦȞ ıIJȘȞ ǹȞȐʌIJȣȟȘ IJȠȣ B. trispora țĮȚ
ıIJȘȞ ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ...........................................................
148
4.2.10 ǼʌȓįȡĮıȘ IJȦȞ ǹțȩȡİıIJȦȞ ȁȚʌĮȡȫȞ ȅȟȑȦȞ ıIJȘȞ ǹȞȐʌIJȣȟȘ IJȠȣ
B. trispora țĮȚ ıIJȘȞ ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ................................
152
4.2.11 ǼʌȓįȡĮıȘ IJȠȣ ǹȞIJȚȠȟİȚįȦIJȚțȠȪ ıIJȘȞ ȆĮȡĮȖȦȖȒ IJȠȣ
ȕ-ȀĮȡȠIJȑȞȚȠȣ................................................................................
155
4.2.12 ǼʌȓįȡĮıȘ IJȦȞ ǼȞİȡȖȠʌȠȚȘIJȫȞ ıIJȘȞ ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ....... 159
4.2.13 ǼʌȓįȡĮıȘ IJȘȢ ȈȪıIJĮıȘȢ IJȠȣ ȊʌȠıIJȡȫµĮIJȠȢ ıIJȘ ȂȠȡijȠȜȠȖȓĮ IJȠȣ
B. trispora........................................................................................
165
4.2.14 ǺİȜIJȚıIJȠʌȠȓȘıȘ IJȘȢ ȆĮȡĮȖȦȖȒȢ ȕ-ȀĮȡȠIJȑȞȚȠȣ ıİ ǹȞĮțȚȞȠȪµİȞİȢ
ȀȦȞȚțȑȢ ĭȚȐȜİȢ................................................................................
171
VI
4.3 ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ Įʌo ȈȣȞșİIJȚțȩ ȊʌȩıIJȡȦµĮ µİ IJȠ B. trispora
ıİ ǹıȣȞİȤȒ ǽȪµȦıȘ ǺȣșȠȪ ıİ ǽȣµȦIJȒȡĮ ǹȞȐįİȣıȘȢ...........................
183
4.3.1 ǼʌȓįȡĮıȘ IJȠȣ ȁȚȞİȜĮȧțȠȪ ȅȟȑȠȢ țĮȚ IJȠȣ ǹȞIJȚȠȟİȚįȦIJȚțȠȪ ıIJȘ
ȈȣȖțȑȞIJȡȦıȘ ȕ-ȀĮȡȠIJȑȞȚȠȣ, ǺȚȠµȐȗĮȢ, ȈĮțȤȐȡȦȞ, ǻȚĮȜȣIJȠȪ
ȅȟȣȖȩȞȠȣ țĮȚ ıIJȠ pH IJȠȣ ȊȖȡȠȪ IJȘȢ ǽȪµȦıȘȢ..............................
183
4.3.2 ǼʌȓįȡĮıȘ IJȘȢ ȀȘȡȠȗȓȞȘȢ ıIJȘ ȈȣȖțȑȞIJȡȦıȘ ȕ-ȀĮȡȠIJȑȞȚȠȣ,
ǺȚȠµȐȗĮȢ, ȈĮțȤȐȡȦȞ, ǻȚĮȜȣIJȠȪ ȅȟȣȖȩȞȠȣ, țĮȚ ıIJȠ pH IJȠȣ
ȊȖȡȠȪ IJȘȢ ǽȪµȦıȘȢ.........................................................................
191
4.3.3 ǼʌȓįȡĮıȘ IJȦȞ ĭȣIJȚțȫȞ ǼȜĮȓȦȞ, IJȠȣ ǹȞIJȚȠȟİȚįȦIJȚțȠȪ țĮȚ IJȘȢ
ȀȘȡȠȗȓȞȘȢ ıIJȘ ȈȣȖțȑȞIJȡȦıȘ ȕ-ȀĮȡȠIJȑȞȚȠȣ, ǺȚȠµȐȗĮȢ,
ȈĮțȤȐȡȦȞ, ǻȚĮȜȣIJȠȪ ȅȟȣȖȩȞȠȣ, țĮȚ ıIJȠ pH IJȠȣ ȊȖȡȠȪ IJȘȢ
ǽȪµȦıȘȢ...........................................................................................
196
4.3.4 ȂİȜȑIJȘ IJȦȞ ȂȠȡijȠȜȠȖȚțȫȞ ȋĮȡĮțIJȘȡȚıIJȚțȫȞ IJȠȣ B. trispora țĮȚ
IJȘȢ ȆĮȡĮȖȦȖȒȢ ȕ-ȀĮȡȠIJȑȞȚȠȣ ıİ ǹıȣȞİȤȒ ǽȪµȦıȘ ǺȣșȠȪ ıİ
ǽȣµȦIJȒȡĮ ǹȞȐįİȣıȘȢ µİ ǻȚĮijȠȡİIJȚțȐ ȊʌȠıIJȡȫµĮIJĮ......................
206
4.3.5 ǼʌȓįȡĮıȘ IJȠȣ ȇȣșµȠȪ ȆĮȡȠȤȒȢ ǹȑȡĮ țĮȚ IJȘȢ ȉĮȤȪIJȘIJĮȢ
ǹȞȐįİȣıȘȢ ıIJȘ ȈȣȖțȑȞIJȡȦıȘ ȕ-ȀĮȡȠIJȑȞȚȠȣ, ǺȚȠµȐȗĮȢ,
ȈĮțȤȐȡȦȞ, ǻȚĮȜȣIJȠȪ ȅȟȣȖȩȞȠȣ, țĮȚ ıIJȠ pH IJȠȣ ȊȖȡȠȪ IJȘȢ
ǽȪµȦıȘȢ...........................................................................................
221
4.3.6 ȂİȜȑIJȘ IJȦȞ ȂȠȡijȠȜȠȖȚțȫȞ ȋĮȡĮțIJȘȡȚıIJȚțȫȞ IJȠȣ B. trispora
țĮIJȐ IJȘȞ ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ ıİ ǹıȣȞİȤȒ ǽȪµȦıȘ ǺȣșȠȪ
ıİ ǽȣµȦIJȒȡĮ ǹȞȐįİȣıȘȢ µİ ǻȚĮijȠȡİIJȚțȑȢ ȈȣȞșȒțİȢ ǹİȡȚıµȠȪ
țĮȚ ǹȞȐįİȣıȘȢ.................................................................................
230
4.3.7 ǺİȜIJȚıIJȠʌȠȓȘıȘ IJȦȞ ȈȣȞșȘțȫȞ ǹİȡȚıµȠȪ țĮȚ ǹȞȐįİȣıȘȢ ȖȚĮ IJȘȞ
ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ǹıȣȞİȤȒ ǽȪµȦıȘ
ǺȣșȠȪ ıİ ǽȣµȦIJȒȡĮ ǹȞȐįİȣıȘȢ........................................................
237
4.4 ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ Įʌȩ ȈȣȞșİIJȚțȩ ȊʌȩıIJȡȦµĮ µİ IJȠ B. trispora
ıİ ǹıȣȞİȤȒ ǽȪµȦıȘ ǺȣșȠȪ µİ ȈȣȞİȤȒ ȆȡȠıșȒțȘ ȊʌȠıIJȡȫµĮIJȠȢ ıİ
ǽȣµȦIJȒȡĮ ǹȞȐįİȣıȘȢ...............................................................................
255
4.4.1 ǼʌȓįȡĮıȘ IJȠȣ ȋȡȩȞȠȣ DzȞĮȡȟȘȢ ȉȡȠijȠįȠıȓĮȢ ıIJȘȞ ȆĮȡĮȖȦȖȒ
ȕ-ȀĮȡȠIJȑȞȚȠȣ, ıIJȘ ȈȣȖțȑȞIJȡȦıȘ IJȘȢ ǺȚȠµȐȗĮȢ, IJȦȞ ȈĮțȤȐȡȦȞ,
IJȠȣ ǻȚĮȜȣIJȠȪ ȅȟȣȖȩȞȠȣ țĮȚ ıIJȠ pH IJȠȣ ȊȖȡȠȪ IJȘȢ
ǽȪµȦıȘȢ..............................................................................................
255
VII
4.4.2 ǼʌȓįȡĮıȘ IJȠȣ ȇȣșµȠȪ ȇȠȒȢ IJȠȣ ȊʌȠıIJȡȫµĮIJȠȢ ȉȡȠijȠįȠıȓĮȢ ıIJȘȞ
ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ, ıIJȘ ȈȣȖțȑȞIJȡȦıȘ IJȘȢ ǺȚȠµȐȗĮȢ, IJȦȞ
ȈĮțȤȐȡȦȞ, IJȠȣ įȚĮȜȣIJȠȪ ȅȟȣȖȩȞȠȣ țĮȚ ıIJȠ pH IJȠȣ ȊȖȡȠȪ IJȘȢ
ǽȪµȦıȘȢ..............................................................................................
261
4.4.3 ǼʌȓįȡĮıȘ IJȘȢ ȂİșȩįȠȣ ȉȡȠijȠįȠıȓĮȢ IJȠȣ ǽȣµȦIJȒȡĮ ıIJȘȞ
ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ, ıIJȘ ȈȣȖțȑȞIJȡȦıȘ IJȘȢ ǺȚȠµȐȗĮȢ, IJȦȞ
ȈĮțȤȐȡȦȞ, IJȠȣ ǻȚĮȜȣIJȠȪ ȅȟȣȖȩȞȠȣ țĮȚ ıIJȠ pH IJȠȣ ȊȖȡȠȪ IJȘȢ
ǽȪµȦıȘȢ...........................................................................................
262
4.4.4 ǼʌȓįȡĮıȘ IJȘȢ ȈȪıIJĮıȘȢ IJȠȣ ȊʌȠıIJȡȫµĮIJȠȢ ȉȡȠijȠįȠıȓĮȢ ıIJȘȞ
ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ, ıIJȘ ȈȣȖțȑȞIJȡȦıȘ IJȘȢ ǺȚȠµȐȗĮȢ, IJȦȞ
ȈĮțȤȐȡȦȞ, IJȠȣ ǻȚĮȜȣIJȠȪ ȅȟȣȖȩȞȠȣ țĮȚ ıIJȠ pH IJȠȣ ȊȖȡȠȪ IJȘȢ
ǽȪµȦıȘȢ...........................................................................................
267
4.4.5 ȂİȜȑIJȘ IJȦȞ ȂȠȡijȠȜȠȖȚțȫȞ ȋĮȡĮțIJȘȡȚıIJȚțȫȞ IJȠȣ B. trispora
țĮIJȐ IJȘȞ ȆĮȡĮȖȦȖȒ ȕ-ȀĮȡȠIJȑȞȚȠȣ ıİ ǹıȣȞİȤȒ ǽȪµȦıȘ ǺȣșȠȪ
µİ ȈȣȞİȤȒ ȆȡȠıșȒțȘ ȊʌȠıIJȡȫµĮIJȠȢ.............................................
273
4.5 ȈȊīȀȇǿȈǾ ȉȍȃ ȂǼĬȅǻȍȃ ȆǹȇǹīȍīǾȈ Ǻ-ȀǹȇȅȉǼȃǿȅȊ.......... 282

ȈȊȂȆǼȇǹȈȂǹȉǹ.........................................................................................

287
ȆȇȅȉǹȈǼǿȈ īǿǹ ȂǼȁȁȅȃȉǿȀǾ ǼȇǼȊȃǹ.............................................

291
ǺǿǺȁǿȅīȇǹĭǿǹ.............................................................................................. 293

VIII
ǼȊȇǼȉǾȇǿȅ ȆǿȃǹȀȍȃ

ıİȜ.
ȆȓȞĮțĮȢ 1.1 ȆİȡȚİțIJȚțȩIJȘIJĮ ȠȡȚıµȑȞȦȞ ijȡȠȪIJȦȞ țĮȚ ȜĮȤĮȞȚțȫȞ ıİ
ȕ-țĮȡȠIJȑȞȚȠ........................................................................................................

2
ȆȓȞĮțĮȢ 3.3.1 ǼʌȓʌİįĮ IJȚµȫȞ IJȦȞ ʌĮȡĮȖȩȞIJȦȞ ʌȠȣ ȤȡȘıȚµȠʌȠȚȒșȘțĮȞ ıIJȠȞ
ʌİȚȡĮµĮIJȚțȩ ıȤİįȚĮıµȩ IJȠȣ µȠȞIJȑȜȠȣ ʌȠȜȜĮʌȜȒȢ ʌĮȜȚȞįȡȩµȘıȘȢ ȖȚĮ IJȘ
ıȣȖțȑȞIJȡȦıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ.........................................................................
75
ȆȓȞĮțĮȢ 3.3.2 ȆİȚȡĮµĮIJȚțȩȢ ıȤİįȚĮıµȩȢ țĮȚ IJȚµȑȢ IJȘȢ µİIJĮȕȜȘIJȒȢ
ĮʌȩțȡȚıȘȢ (ıȣȖțȑȞIJȡȦıȘ ȕ-țĮȡȠIJȑȞȚȠȣ) ʌȠȣ µİIJȡȒșȘțĮȞ ıIJȚȢ įȚĮijȠȡİIJȚțȑȢ
įȠțȚµȑȢ IJȠȣ ʌİȚȡĮµĮIJȚțȠȪ ıȤİįȚĮıµȠȪ..............................................................
76
ȆȓȞĮțĮȢ 3.3.3 īİȦµİIJȡȚțȑȢ ʌĮȡȐµİIJȡȠȚ IJȘȢ µİȜȑIJȘȢ IJȦȞ µȠȡijȠȜȠȖȚțȫȞ
ȤĮȡĮțIJȘȡȚıIJȚțȫȞ IJȠȣ B. trispora ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ țȦȞȚțȑȢ
ijȚȐȜİȢ µİ ȥȘijȚĮțȒ ĮȞȐȜȣıȘ İȚțȩȞĮȢ.................................................................
80
ȆȓȞĮțĮȢ 3.4.1 īİȦµİIJȡȚțȑȢ ʌĮȡȐµİIJȡȠȚ IJȘȢ µİȜȑIJȘȢ IJȦȞ µȠȡijȠȜȠȖȚțȫȞ
ȤĮȡĮțIJȘȡȚıIJȚțȫȞ IJȠȣ B. trispora ıİ ȗȣµȦIJȒȡĮ ĮȞĮįİȪıİȦȢ µİ ȥȘijȚĮțȒ
ĮȞȐȜȣıȘ İȚțȩȞĮȢ................................................................................................
90
ȆȓȞĮțĮȢ 3.4.2 ǼʌȓʌİįĮ IJȚµȫȞ IJȦȞ ʌĮȡĮȖȩȞIJȦȞ ʌȠȣ ȤȡȘıȚµȠʌȠȚȒșȘțĮȞ ıIJȠ
ʌİȚȡĮµĮIJȚțȩ ıȤİįȚĮıµȩ IJȠȣ µȠȞIJȑȜȠȣ ʌȠȜȜĮʌȜȒȢ ʌĮȜȚȞįȡȩµȘıȘȢ ȖȚĮ IJȘ
ıȣȖțȑȞIJȡȦıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ, IJȘȞ ʌĮȡĮȖȦȖȚțȩIJȘIJĮ, IJȘ ȕȚȠµȐȗĮ țĮȚ IJȠȞ
ȠȖțȠµİIJȡȚțȩ ıȣȞIJİȜİıIJȒ µİIJĮijȠȡȐȢ µȐȗĮȢ.......................................................
96
ȆȓȞĮțĮȢ 3.4.3 ȆİȚȡĮµĮIJȚțȩȢ ıȤİįȚĮıµȩȢ țĮȚ IJȚµȑȢ IJȦȞ µİIJĮȕȜȘIJȫȞ
ĮʌȩțȡȚıȘȢ ʌȠȣ µİIJȡȒșȘțĮȞ ıIJȚȢ įȚĮijȠȡİIJȚțȑȢ įȠțȚµȑȢ IJȠȣ ʌİȚȡĮµĮIJȚțȠȪ
ıȤİįȚĮıµȠȪ.........................................................................................................
97
ȆȓȞĮțĮȢ 4.1.1 ǼʌȓįȡĮıȘ IJȘȢ țĮIJİȡȖĮıȓĮȢ IJȠȣ ȣȖȡȠȪ IJȘȢ ȗȪµȦıȘȢ ıIJȘȞ
ĮʌȠµȐțȡȣȞıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ Įʌȩ IJĮ țȪIJIJĮȡĮ IJȠȣ B. trisporĮ (ȋȡȩȞȠȢ
ȗȪµȦıȘȢ 8 ȘµȑȡİȢ).............................................................................................
117
ȆȓȞĮțĮȢ 4.2.1 ȂȠȡijȠȜȠȖȚțȐ ȤĮȡĮțIJȘȡȚıIJȚțȐ IJȠȣ B. trispora ıIJȘ µȑȖȚıIJȘ
ıȣȖțȑȞIJȡȦıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ țȦȞȚțȑȢ
ijȚȐȜİȢ (ȆȠıȠıIJȩ İʌȓ IJȠȚȢ İțĮIJȩ IJȦȞ įȚĮijȠȡİIJȚțȫȞ µȠȡijȫȞ IJȦȞ µȣțȘȜȚĮțȫȞ
ȣijȫȞ, ȗȣȖȠıʌȠȡȓȦȞ țĮȚ IJȠȣ İȞįȠțȣIJIJĮȡȚțȠȪ ȕ-țĮȡȠIJȑȞȚȠȣ ıIJȠ ıȣȞȠȜȚțȩ
ȟȘȡȩ ȕȐȡȠȢ IJȘȢ ȕȚȠµȐȗĮȢ)..................................................................................
123
IX
ȆȓȞĮțĮȢ 4.2.2 ǼʌȓįȡĮıȘ IJȘȢ ʌȘȖȒȢ ȐȞșȡĮțĮ ıIJȚȢ țȚȞȘIJȚțȑȢ ʌĮȡĮµȑIJȡȠȣȢ IJȘȢ
ȗȪµȦıȘȢ ȖȚĮ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ Įʌȩ ıȣȞșİIJȚțȩ ȣʌȩıIJȡȦµĮ µİ IJȠ
B. trispora ıİ țȦȞȚțȑȢ ijȚȐȜİȢ (ȋȡȩȞȠȢ ȗȪµȦıȘȢ 8 ȘµȑȡİȢ)..............................
139
ȆȓȞĮțĮȢ 4.2.3 ǼʌȓįȡĮıȘ IJȘȢ ʌȘȖȒȢ ĮȗȫIJȠȣ ıIJȚȢ țȚȞȘIJȚțȑȢ ʌĮȡĮµȑIJȡȠȣȢ IJȘȢ
ȗȪµȦıȘȢ ȖȚĮ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ Įʌȩ ıȣȞșİIJȚțȩ ȣʌȩıIJȡȦµĮ µİ IJȠ
B. trispora ıİ țȦȞȚțȑȢ ijȚȐȜİȢ (ȋȡȩȞȠȢ ȗȪµȦıȘȢ 8 ȘµȑȡİȢ)..............................
143
ȆȓȞĮțĮȢ 4.2.4 ȈȪıIJĮıȘ İțȤȣȜȓıµĮIJȠȢ țĮȜĮµʌȠțȚȠȪ....................................... 144
ȆȓȞĮțĮȢ 4.2.5 ǼʌȓįȡĮıȘ IJȦȞ İʌȚijĮȞİȚȠįȡĮıIJȚțȫȞ ȠȣıȚȫȞ ıIJȘ µȠȡijȠȜȠȖȓĮ
IJȠȣ B. trispora țĮȚ ıIJȚȢ țȚȞȘIJȚțȑȢ ʌĮȡĮµȑIJȡȠȣȢ IJȘȢ ȗȪµȦıȘȢ ȖȚĮ IJȘȞ
ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ țȦȞȚțȑȢ ijȚȐȜİȢ (ȋȡȩȞȠȢ ȗȪµȦıȘȢ 8 ȘµȑȡİȢ)........
147
ȆȓȞĮțĮȢ 4.2.6 ǼʌȓįȡĮıȘ IJȦȞ ijȣIJȚțȫȞ İȜĮȓȦȞ ıIJȚȢ țȚȞȘIJȚțȑȢ ʌĮȡĮµȑIJȡȠȣȢ
IJȘȢ ȗȪµȦıȘȢ ȖȚĮ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ țȦȞȚțȑȢ
ijȚȐȜİȢ (ȋȡȩȞȠȢ ȗȪµȦıȘȢ 8 ȘµȑȡİȢ)...................................................................
150
ȆȓȞĮțĮȢ 4.2.7 ǼʌȓįȡĮıȘ IJȠȣ µȓȖµĮIJȠȢ IJȦȞ ijȣIJȚțȫȞ İȜĮȓȦȞ ıIJȚȢ țȚȞȘIJȚțȑȢ
ʌĮȡĮµȑIJȡȠȣȢ IJȘȢ ȗȪµȦıȘȢ ȖȚĮ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora
ıİ țȦȞȚțȑȢ ijȚȐȜİȢ (ȋȡȩȞȠȢ ȗȪµȦıȘȢ 8 ȘµȑȡİȢ)................................................
151
ȆȓȞĮțĮȢ 4.2.8 ǼʌȓįȡĮıȘ IJȦȞ ĮțȩȡİıIJȦȞ ȜȚʌĮȡȫȞ ȠȟȑȦȞ ıIJȚȢ țȚȞȘIJȚțȑȢ
ʌĮȡĮµȑIJȡȠȣȢ IJȘȢ ȗȪµȦıȘȢ ȖȚĮ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora
ıİ țȦȞȚțȑȢ ijȚȐȜİȢ (ȋȡȩȞȠȢ ȗȪµȦıȘȢ 8 ȘµȑȡİȢ)................................................
154
ȆȓȞĮțĮȢ 4.2.9 ǼʌȓįȡĮıȘ IJȠȣ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ ıIJȚȢ țȚȞȘIJȚțȑȢ ʌĮȡĮµȑIJȡȠȣȢ
IJȘȢ ȗȪµȦıȘȢ ȖȚĮ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ țȦȞȚțȑȢ
ijȚȐȜİȢ (ȋȡȩȞȠȢ ȗȪµȦıȘȢ 8 ȘµȑȡİȢ)...................................................................
157
ȆȓȞĮțĮȢ 4.2.10 ǼʌȓįȡĮıȘ įȚĮijȩȡȦȞ ȠȣıȚȫȞ ıIJȚȢ țȚȞȘIJȚțȑȢ ʌĮȡĮµȑIJȡȠȣȢ IJȘȢ
ȗȪµȦıȘȢ ȖȚĮ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ țȦȞȚțȑȢ
ijȚȐȜİȢ (ȋȡȩȞȠȢ ȗȪµȦıȘȢ 8 ȘµȑȡİȢ)...................................................................
163
ȆȓȞĮțĮȢ 4.2.11 ȂȠȡijȠȜȠȖȚțȐ ȤĮȡĮțIJȘȡȚıIJȚțȐ IJȠȣ B. trispora ıIJȘ µȑȖȚıIJȘ
ıȣȖțȑȞIJȡȦıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ țȦȞȚțȑȢ
ijȚȐȜİȢ ȤȡȘıȚµȠʌȠȚȫȞIJĮȢ ȣʌȠıIJȡȫµĮIJĮ µİ įȚĮijȠȡİIJȚțȒ ıȪıIJĮıȘ (ȆȠıȠıIJȩ
İʌȓ IJȠȚȢ İțĮIJȩ IJȦȞ įȚĮijȠȡİIJȚțȫȞ µȠȡijȫȞ IJȦȞ µȣțȘȜȚĮțȫȞ ȣijȫȞ,
ȗȣȖȠıʌȠȡȓȦȞ țĮȚ IJȠȣ İȞįȠțȣIJIJĮȡȚțȠȪ ȕ-țĮȡȠIJȑȞȚȠȣ ıIJȠ ıȣȞȠȜȚțȩ ȟȘȡȩ
ȕȐȡȠȢ IJȘȢ ȕȚȠµȐȗĮȢ)...........................................................................................
169
ȆȓȞĮțĮ 4.2.12 ǹȞȐȜȣıȘ įȚĮțȪµĮȞıȘȢ ȖȚĮ IJȠ µȠȞIJȑȜȠ ʌȠȜȜĮʌȜȒȢ
ʌĮȜȚȞįȡȩµȘıȘȢ įİȣIJȑȡȠȣ ȕĮșµȠȪ IJȘȢ ıȣȖțȑȞIJȡȦıȘȢ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ..........
176
X
ȆȓȞĮțĮȢ 4.2.13 ǼțIJȓµȘıȘ IJȦȞ µİȡȚțȫȞ ıȣȞIJİȜİıIJȫȞ IJȠȣ µȠȞIJȑȜȠȣ
ʌȠȜȜĮʌȜȒȢ ʌĮȜȚȞįȡȩµȘıȘȢ įİȣIJȑȡȠȣ ȕĮșµȠȪ ȖȚĮ IJȘ ıȣȖțȑȞIJȡȦıȘ IJȠȣ
ȕ-țĮȡȠIJȑȞȚȠȣ......................................................................................................
177
ȆȓȞĮțĮȢ 4.2.14 ȆȡȠȕȜİʌȩµİȞȘ (prd) țĮȚ ʌĮȡĮIJȘȡȠȪµİȞȘ (obs) µȑȖȚıIJȘ
ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ țȦȞȚțȑȢ ijȚȐȜİȢ
ȤȡȘıȚµȠʌȠȚȫȞIJĮȢ ıȣȞșİIJȚțȩ ȣʌȩıIJȡȦµĮ ʌȠȣ ȑȤİȚ ݵʌȜȠȣIJȚıIJİȓ µİ IJȠȞ
ȐȡȚıIJȠ ʌȡȠȕȜİʌȩµİȞȠ ıȣȞįȣĮıµȩ IJȠȣ ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ, IJȘȢ țȘȡȠȗȓȞȘȢ țĮȚ
IJȠȣ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ...........................................................................................
182
ȆȓȞĮțĮȢ 4.3.1 ȂİȜȑIJȘ țȚȞȘIJȚțȫȞ ʌĮȡĮµȑIJȡȦȞ IJȘȢ ĮıȣȞİȤȠȪȢ ȗȪµȦıȘȢ
ȕȣșȠȪ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ȗȣµȦIJȒȡĮ
ĮȞȐįİȣıȘȢ µİ įȚĮijȠȡİIJȚțȐ ȣʌȠıIJȡȫµĮIJĮ ʌĮȡĮȖȦȖȒȢ......................................
212
ȆȓȞĮțĮȢ 4.3.2 ȂȠȡijȠȜȠȖȚțȐ ȤĮȡĮțIJȘȡȚıIJȚțȐ IJȠȣ B. trispora țĮIJȐ IJȘȞ
ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ
µİ įȚĮijȠȡİIJȚțȐ ȣʌȠıIJȡȫµĮIJĮ ʌĮȡĮȖȦȖȒȢ (ȆȠıȠıIJȩ İʌȓ IJȠȚȢ İțĮIJȩ IJȦȞ
įȚĮijȠȡİIJȚțȫȞ µȠȡijȫȞ IJȦȞ µȣțȘȜȚĮțȫȞ ȣijȫȞ, µȠȡijȠȜȠȖȚțȫȞ įȠµȫȞ țĮȚ IJȠȣ
İȞįȠțȣIJIJĮȡȚțȠȪ ȕ-țĮȡȠIJȑȞȚȠȣ ıIJȠ ıȣȞȠȜȚțȩ ȟȘȡȩ ȕȐȡȠȢ IJȘȢ ȕȚȠµȐȗĮȢ)..........
213
ȆȓȞĮțĮȢ 4.3.3 ȂİȜȑIJȘ țȚȞȘIJȚțȫȞ ʌĮȡĮµȑIJȡȦȞ IJȘȢ ĮıȣȞİȤȠȪȢ ȗȪµȦıȘȢ
ȕȣșȠȪ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ȗȣµȦIJȒȡĮ
ĮȞȐįİȣıȘȢ µİ įȚĮijȠȡİIJȚțȑȢ ıȣȞșȒțİȢ ĮİȡȚıµȠȪ țĮȚ ĮȞȐįİȣıȘȢ.......................
229
ȆȓȞĮțĮȢ 4.3.4 ȂȠȡijȠȜȠȖȚțȐ ȤĮȡĮțIJȘȡȚıIJȚțȐ IJȠȣ B. trispora ıIJȘ µȑȖȚıIJȘ
ıȣȖțȑȞIJȡȦıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ ȗȣµȦIJȒȡĮ
ĮȞȐįİȣıȘȢ µİ įȚĮijȠȡİIJȚțȑȢ ıȣȞșȒțİȢ ĮİȡȚıµȠȪ țĮȚ ĮȞȐįİȣıȘȢ (ȆȠıȠıIJȩ İʌȓ
IJȠȚȢ İțĮIJȩ IJȦȞ įȚĮijȠȡİIJȚțȫȞ µȠȡijȫȞ IJȦȞ µȣțȘȜȚĮțȫȞ ȣijȫȞ, ȗȣȖȠıʌȠȡȓȦȞ
țĮȚ IJȠȣ İȞįȠțȣIJIJĮȡȚțȠȪ ȕ-țĮȡȠIJȑȞȚȠȣ ıIJȠ ıȣȞȠȜȚțȩ ȟȘȡȩ ȕȐȡȠȢ IJȘȢ
ȕȚȠµȐȗĮȢ)............................................................................................................
234
ȆȓȞĮțĮ 4.3.5 ǹȞȐȜȣıȘ įȚĮțȪµĮȞıȘȢ ȖȚĮ IJȠ µȠȞIJȑȜȠ ʌȠȜȜĮʌȜȒȢ
ʌĮȜȚȞįȡȩµȘıȘȢ įİȣIJȑȡȠȣ ȕĮșµȠȪ IJȘȢ ıȣȖțȑȞIJȡȦıȘȢ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ (A),
IJȘȢ ʌĮȡĮȖȦȖȚțȩIJȘIJĮȢ (B), IJȘȢ ȕȚȠµȐȗĮȢ ȟȘȡȩ ȕȐȡȠȢ (C) țĮȚ IJȠȣ
ȠȖțȠµİIJȡȚțȠȪ ıȣȞIJİȜİıIJȒ µİIJĮijȠȡȐȢ µȐȗĮȢ (D)..............................................
244
ȆȓȞĮțĮȢ 4.3.6 ǼțIJȓµȘıȘ IJȦȞ µİȡȚțȫȞ ıȣȞIJİȜİıIJȫȞ IJȠȣ µȠȞIJȑȜȠȣ
ʌȠȜȜĮʌȜȒȢ ʌĮȜȚȞįȡȩµȘıȘȢ įİȣIJȑȡȠȣ ȕĮșµȠȪ ȖȚĮ IJȘ ıȣȖțȑȞIJȡȦıȘ IJȠȣ ȕ-
țĮȡȠIJȑȞȚȠȣ (A), IJȘȞ ʌĮȡĮȖȦȖȚțȩIJȘIJĮ (B), IJȘ ȕȚȠµȐȗĮ ȟȘȡȩ ȕȐȡȠȢ (C) țĮȚ
IJȠȞ ȠȖțȠµİIJȡȚțȩ ıȣȞIJİȜİıIJȒ µİIJĮijȠȡȐȢ µȐȗĮȢ (D)................................……..
245
XI
ȆȓȞĮțĮȢ 4.3.7 ȆȡȠȕȜİʌȩµİȞİȢ (prd) țĮȚ ʌĮȡĮIJȘȡȠȪµİȞİȢ (obs) µȑȖȚıIJİȢ
IJȚµȑȢ IJȘȢ ıȣȖțȑȞIJȡȦıȘȢ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ, IJȘȢ ʌĮȡĮȖȦȖȚțȩIJȘIJĮȢ, IJȘȢ
ıȣȖțȑȞIJȡȦıȘȢ IJȘȢ ȕȚȠµȐȗĮȢ țĮȚ IJȠȣ ȠȖțȠµİIJȡȚțȠȪ ıȣȞIJİȜİıIJȒ µİIJĮijȠȡȐȢ
µȐȗĮȢ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıIJȠȣȢ ȐȡȚıIJȠȣȢ
ʌȡȠȕȜİʌȩµİȞȠȣȢ ıȣȞįȣĮıµȠȪȢ IJȘȢ IJĮȤȪIJȘIJĮȢ ĮȞȐįİȣıȘȢ țĮȚ IJȠȣ ȡȣșµȠȪ
ʌĮȡȠȤȒȢ ĮȑȡĮ ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ ȗȣµȦIJȒȡĮ
ĮȞȐįİȣıȘȢ..........................................................................................................
254
ȆȓȞĮțĮȢ 4.4.1 ȈȪıIJĮıȘ IJȦȞ ȣʌȠıIJȡȦµȐIJȦȞ ĮȞȐʌIJȣȟȘȢ țĮȚ IJȡȠijȠįȠıȓĮȢ ȖȚĮ
IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ µİ
ıȣȞİȤȒ ʌȡȠıșȒțȘ ȣʌȠıIJȡȫµĮIJȠȢ ıİ ȗȣµȦIJȒȡĮ ĮȞĮįİȪıİȦȢ............................
278
ȆȓȞĮțĮȢ 4.4.2 ȂȠȡijȠȜȠȖȚțȐ ȤĮȡĮțIJȘȡȚıIJȚțȐ IJȠȣ B. trispora țĮIJȐ IJȘȞ
ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ µİ ıȣȞİȤȒ ʌȡȠıșȒțȘ
ȣʌȠıIJȡȫµĮIJȠȢ ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ µİ įȚĮijȠȡİIJȚțȐ ȣʌȠıIJȡȫµĮIJĮ
ĮȞȐʌIJȣȟȘȢ țĮȚ IJȡȠijȠįȠıȓĮȢ țĮȚ įȚĮijȠȡİIJȚțȑȢ µİșȩįȠȣȢ țĮȚ ȡȣșµȠȪȢ
IJȡȠijȠįȠıȓĮȢ (ȆȠıȠıIJȩ İʌȓ IJȠȚȢ İțĮIJȩ IJȦȞ įȚĮijȠȡİIJȚțȫȞ µȠȡijȫȞ IJȦȞ
µȣțȘȜȚĮțȫȞ ȣijȫȞ, µȠȡijȠȜȠȖȚțȫȞ įȠµȫȞ țĮȚ IJȠȣ İȞįȠțȣIJIJĮȡȚțȠȪ
ȕ-țĮȡȠIJȑȞȚȠȣ ıIJȠ ıȣȞȠȜȚțȩ ȟȘȡȩ ȕȐȡȠȢ IJȘȢ ȕȚȠµȐȗĮȢ).....................................
279
ȆȓȞĮțĮȢ 4.4.3 ȂİȜȑIJȘ țȚȞȘIJȚțȫȞ ʌĮȡĮµȑIJȡȦȞ IJȘȢ ĮıȣȞİȤȠȪȢ ȗȪµȦıȘȢ
ȕȣșȠȪ µİ ıȣȞİȤȒ ʌȡȠıșȒțȘ ȣʌȠıIJȡȫµĮIJȠȢ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ
ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ µİ įȚĮijȠȡİIJȚțȐ
ȣʌȠıIJȡȫµĮIJĮ ĮȞȐʌIJȣȟȘȢ țĮȚ IJȡȠijȠįȠıȓĮȢ țĮȚ įȚĮijȠȡİIJȚțȑȢ µİșȩįȠȣȢ țĮȚ
ȡȣșµȠȪȢ IJȡȠijȠįȠıȓĮȢ........................................................................................
281
ȆȓȞĮțĮȢ 4.5.1 ǺĮıȚțȑȢ ʌĮȡȐµİIJȡȠȚ IJȘȢ ȗȣµȦIJȚțȒȢ įȚİȡȖĮıȓĮȢ ȖȚĮ IJȘȞ
ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora Įʌȩ ıȣȞșİIJȚțȩ ȣʌȩıIJȡȦµĮ ıİ
įȚĮijȠȡİIJȚțȐ ıȣıIJȒµĮIJĮ ȗȪµȦıȘȢ......................................................................
285

XII
ǼȊȇǼȉǾȇǿȅ ȈȋǾȂǹȉȍȃ

ıİȜ.
ȈȤȒµĮ 2.1.1 ǹʌİȚțȩȞȘıȘ IJȦȞ ıIJĮįȓȦȞ ıȤȘµĮIJȚıµȠȪ IJȠȣ ȗȣȖȠıʌȠȡȓȠȣ țĮIJȐ IJȘȞ İȖȖİȞȒ
ĮȞĮʌĮȡĮȖȦȖȒ IJȠȣ B. trispora. a) ǻȪȠ µȣțȘȜȚĮțȠȓ ȕȡĮȤȓȠȞİȢ ʌȠȣ ʌȡȠȑȡȤȠȞIJĮȚ Įʌȩ IJĮ
(+) țĮȚ (-) ıIJİȜȑȤȘ IJȠȣ µȚțȡȠȠȡȖĮȞȚıµȠȪ ʌȜȘıȚȐȗȠȣȞ țĮȚ İijȐʌIJȠȞIJĮȚ, b) ʌȣțȞȩ
ʌȡȦIJȩʌȜĮıµĮ țĮȚ ʌȣȡȒȞİȢ ıȣȖțİȞIJȡȫȞȠȞIJĮȚ ıİ ıȘµİȓĮ İʌĮijȒȢ țĮȚ IJĮ ȖĮµİIJȐȖȖİȚĮ ıIJĮ
ȐțȡĮ IJȦȞ ʌȡȠȖĮµİIJĮȖȖİȓȦȞ ȤȦȡȓȗȠȞIJĮȚ Įʌȩ IJȠȞ ȣʌȩȜȠȚʌȠ șĮȜȜȩ µİ įȚĮijȡȐȖµĮIJĮ, c)
ıȤȘµĮIJȚıµȩȢ IJȠȣ ȗȣȖȦIJȠȪ țĮIJȐ IJȘ ıȣȖȤȫȞİȣıȘ ʌȣȡȒȞȦȞ IJȦȞ įȪȠ ȖĮµİIJĮȖȖİȓȦȞ, d) Ƞ
ȗȣȖȦIJȩȢ ĮʌȠțIJȐ ʌĮȤȚȐ IJȠȚȤȫµĮIJĮ țĮȚ µİIJĮIJȡȑʌİIJĮȚ ıİ ȗȣȖȠıʌȩȡȚȠ ..............................

9
ȈȤȒµĮ 2.1.2 ǻȠµȒ IJȦȞ įȚĮijȠȡİIJȚțȫȞ µȠȡijȫȞ IJȡȚıʌȠȡȚțȠȪ ȠȟȑȠȢ, ʌȡȠȠȡµȠȞȫȞ,
ĮʌȠIJȡȚıʌȠȡȩȜȘȢ țĮȚ ĮʌȠIJȡȚıʌȠȡȓȞȘȢ. ȉȡȚıʌȠȡȚțȩ ȠȟȪ-A: X = H, H (1), ȉȡȚıʌȠȡȚțȩ
ȠȟȪ-B: X = O (2), ȉȡȚıʌȠȡȚțȩ ȠȟȪ-C: X = H, OH (3), 4-ǻȚȣįȡȠ-µİșȣȜİıIJȑȡĮȢ IJȠȣ
IJȡȚıʌȠȡȚțȠȪ ȠȟȑȠȢ (4), ȂİșȣȜİıIJȑȡĮȢ IJȠȣ IJȡȚıʌȠȡȚțȠȪ ȠȟȑȠȢ (5), ȉȡȚıʌȠȡȩȜȘ (6),
ȉȡȚıʌȠȡȓȞȘ (7), ǹʌȠIJȡȚıʌȠȡȩȜȘ (8), ǹʌȠIJȡȚıʌȠȡȓȞȘ (9) (ıIJȚȢ İȞȫıİȚȢ 4, 5, 6, 7, 8 țĮȚ 9
ȋ: ȅ Ȓ Ǿ țĮȚ ȅǾ).................................................................................................................
12
ȈȤȒµĮ 2.1.3 ȂİIJĮȕȠȜȚıµȩȢ IJȦȞ ʌȡȠȠȡµȠȞȫȞ țĮȚ ıȤȘµĮIJȚıµȩȢ IJȠȣ IJȡȚıʌȠȡȚțȠȪ ȠȟȑȠȢ
țĮIJȐ IJȘ ıȪȗİȣȟȘ IJȦȞ (+) țĮȚ (-) ıIJİȜİȤȫȞ IJȠȣ B. trispora. Ǿ ıȪȞșİıȘ įȚĮijȠȡİIJȚțȫȞ
ʌȡȠȠȡµȠȞȫȞ ıIJĮ ĮȞIJȓșİIJĮ ijȪȜĮ IJȠȣ µȚțȡȠȠȡȖĮȞȚıµȠȪ ȠijİȓȜİIJĮȚ ıIJȠ įȚĮijȠȡİIJȚțȩ İȓįȠȢ
µİIJĮIJȡȠʌȒȢ IJȘȢ pro-S µİșȣȜȚțȒȢ ȠµȐįĮȢ IJȠȣ C-1 IJȘȢ 4-įȚȣįȡȠ-IJȡȚıʌȠȡȓȞȘȢ ıİ țȐșİ
ıIJȑȜİȤȠȢ. Ǿ µİșȣȜȠµȐįĮ İȓIJİ įİȞ µİIJĮȕȐȜȜİIJĮȚ (IJȡȚıʌȠȡȓȞȘ) İȓIJİ ȠȟİȚįȫȞİIJĮȚ ıİ
ʌȡȦIJȠIJĮȖȒ ĮȜțȠȩȜȘ (IJȡȚıʌȠȡȩȜȘ) ıIJȠ (-) ıIJȑȜİȤȠȢ țĮȚ ȠȟİȚįȫȞİIJĮȚ ıİ țĮȡȕȠȟȣȜȠµȐįĮ
ʌȠȣ ıIJȘ ıȣȞȑȤİȚĮ İıIJİȡȠʌȠȚİȓIJĮȚ ıIJȠ (+) ıIJİȜȑȤȠȢ.  İȞȗȣµȚțȒ ĮȞIJȓįȡĮıȘ, įȚȐȤȣıȘ
IJȦȞ ʌȡȠȠȡµȠȞȫȞ IJȠȣ İȞȩȢ ıIJİȜȑȤȠȣȢ ıIJȠ İıȦIJİȡȚțȩ IJȦȞ țȣIJIJȐȡȦȞ ıIJİȜȑȤȠȣȢ IJȠȣ
ĮȞIJȓșİIJȠȣ ijȪȜȠȣ...................................................................................................................

13
ȈȤȒµĮ 2.2.1 ǺȚȠıȪȞșİıȘ ȕ-țĮȡȠIJȑȞȚȠȣ.............................................................................. 23
ȈȤȒµĮ 2.6.1 ȈIJȐįȚĮ ʌĮȡĮȖȦȖȒȢ ȕ-țĮȡȠIJȑȞȚȠȣ Įʌȩ ıȣȞșİIJȚțȩ ȣʌȩıIJȡȦµĮ µİ IJȠ
B. trispora. ȊʌȩıIJȡȦµĮ 1: Potato dextrose agar (PDA). ȊʌȩıIJȡȦµĮ 2 (g/l): ǼțȤȪȜȚıµĮ
țĮȜĮµʌȠțȚȠȪ 70,0, ȐµȣȜȠ țĮȜĮµʌȠțȚȠȪ 50,0, ȣįȡȠȤȜȦȡȚțȒ șİȚĮµȓȞȘ 0,01, KH
2
PO
4
0,5,
MgSO
4
. H
2
0 0,1. ȊʌȩıIJȡȦµĮ 3 (g/l): ǻȚĮȜȣIJȩ ȣʌȩȜİȚµµĮ ĮʌȩıIJĮȟȘȢ ĮȜțȠȩȜȘȢ 70,0,
ȐµȣȜȠ țĮȜĮµʌȠțȚȠȪ 60,0, ıȠȖȚȐȜİȣȡȠ 30,0, ȕĮµȕĮțȑȜĮȚȠ 30,0, ĮȞIJȚȠȟİȚįȦIJȚțȩ 0,3,
MgSO
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. H
2
0 0,2, ȣįȡȠȤȜȦȡȚțȒ șİȚĮµȓȞȘ 0,5, ȣįȡĮȗȓįȚȠ IJȠȣ 4-ʌȣȡȚįȚȞȠțĮȡȕȠȟȣȜȚțȠȪ

XIII
ȠȟȑȠȢ 0,6, țȘȡȠȗȓȞȘ 20,0, pH 6,3. ȉȠ ȣʌȩıIJȡȦµĮ ĮʌȠıIJİȚȡȫȞİIJĮȚ ıIJȠȣȢ 121
Ƞ
C ȖȚĮ 55
ȜİʌIJȐ (Ș țȘȡȠȗȓȞȘ țĮȚ IJȠ ȣįȡĮȗȓįȚȠ IJȠȣ 4-ʌȣȡȚįȚȞȠțĮȡȕȠȟȣȜȚțȠȪ ȠȟȑȠȢ
ĮʌȠıIJİȚȡȫȞȠȞIJĮȚ ȟİȤȦȡȚıIJȐ µİ įȚȒșȘıȘ). ȂİIJȐ 48 ȫȡİȢ Įʌȩ IJȘȞ ĮȡȤȒ IJȘȢ ȗȪµȦıȘȢ
ʌȡȠıIJȓșİIJĮȚ ıIJȠ ȗȪµȦIJȒȡĮ 1,0 g/l ȕ-ȚȠȞȩȞȘ țĮȚ 5,0 g/l țȘȡȠȗȓȞȘ. Ǿ ȖȜȣțȩȗȘ
ʌȡȠıIJȓșİIJĮȚ ıȣȞİȤȫȢ ıIJȠ ȗȣµȦIJȒȡĮ țĮȚ Ș ıȣȞȠȜȚțȒ ʌȠıȩIJȘIJĮ IJȘȢ ȖȜȣțȩȗȘȢ ʌȠȣ
ȤȡȘıȚµȠʌȠȚİȓIJĮȚ ıİ ȩȜȘ IJȘ įȚȐȡțİȚĮ IJȘȢ ȗȪµȦıȘȢ İȓȞĮȚ 42,0 g/l ........................................

50
ȈȤȒµĮ 2.6.2 ǹȞȐțIJȘıȘ țĮȚ țĮșĮȡȚıµȩȢ ȕ-țĮȡȠIJȑȞȚȠȣ Įʌȩ IJȠ B. trispora......................... 52
ȈȤȒµĮ 3.3.1 ȆȡȩIJȣʌȘ țĮµʌȪȜȘ ȖȚĮ IJȠȞ ʌȡȠıįȚȠȡȚıµȩ IJȦȞ ıĮțȤȐȡȦȞ.............................. 67
ȈȤȒµĮ 3.3.2 ȆȡȩIJȣʌȘ țĮµʌȪȜȘ ȖȚĮ IJȠȞ ʌȡȠıįȚȠȡȚıµȩ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ.......................... 68
ȈȤȒµĮ 3.3.3 ȆȡȩIJȣʌȘ țĮµʌȪȜȘ ȖȚĮ IJȠȞ ʌȡȠıįȚȠȡȚıµȩ IJȠȣ Ǿ
2
ȅ
2
....................................... 69
ȈȤȒµĮ 3.4.1 ǽȣµȦIJȒȡĮȢ ĮȞȐįİȣıȘȢ ʌȠȣ ȤȡȘıȚµȠʌȠȚȒșȘțİ ȖȚĮ IJȘȞ ʌĮȡĮȖȦȖȒ
ȕ-țĮȡȠIJȑȞȚȠȣ Įʌȩ ıȣȞșİIJȚțȩ ȣʌȩıIJȡȦµĮ µİ IJȠ µȪțȘIJĮ B. trispora...................................

85
ȈȤȒµĮ 3.4.2 ȆȡȠıįȚȠȡȚıµȩȢ IJȠȣ K
L
a µİ IJȘ įȣȞĮµȚțȒ µȑșȠįȠ........................................... 98
ȈȤȒµĮ 3.4.3 ȆȡȠıįȚȠȡȚıµȩȢ IJȠȣ K
L
a µİ IJȘ įȣȞĮµȚțȒ µȑșȠįȠ (Ǿ țȜȓıȘ IJȘȢ İȣșİȓĮȢ
ȚıȠȪIJĮȚ µİ –1/K
L
a)...............................................................................................................
99
ȈȤȒµĮ 4.1.1 ǼʌȓįȡĮıȘ IJȠȣ ȤȡȩȞȠȣ ıIJȘȞ İțȤȪȜȚıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ Įʌȩ IJĮ țȪIJIJĮȡĮ
IJȠȣ B. trispora µİ įȚĮijȠȡİIJȚțȠȪȢ įȚĮȜȪIJİȢ İțȤȪȜȚıȘȢ. -- ĮȚșĮȞȩȜȘ, - - µİșĮȞȩȜȘ,
- - ĮțİIJȩȞȘ, - - ʌİIJȡİȜĮȧțȩȢ ĮȚșȑȡĮȢ, -- İȟȐȞȚȠ, - - µİșĮȞȩȜȘİȟȐȞȚȠ (11), - -
ȚıȠʌȡȠʌȣȜȚțȒ ĮȜțȠȩȜȘ, - - ȠȟȚțȩȢ ĮȚșȣȜİıIJȑȡĮȢ (ĬİȡµȠțȡĮıȓĮ İțȤȪȜȚıȘȢ 30
0
C)........
106
ȈȤȒµĮ 4.1.2 ǼʌȓįȡĮıȘ IJȘȢ șİȡµȠțȡĮıȓĮȢ ıIJȘȞ İțȤȪȜȚıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ Įʌȩ IJĮ
țȪIJIJĮȡĮ IJȠȣ B. trispora µİ įȚĮijȠȡİIJȚțȠȪȢ įȚĮȜȪIJİȢ İțȤȪȜȚıȘȢ. -- ĮȚșĮȞȩȜȘ, - -
µİșĮȞȩȜȘ, - - ĮțİIJȩȞȘ, - - ʌİIJȡİȜĮȧțȩȢ ĮȚșȑȡĮȢ, -- İȟȐȞȚȠ, - - µİșĮȞȩȜȘİȟȐȞȚȠ
(11) - - ȚıȠʌȡȠʌȣȜȚțȒ ĮȜțȠȩȜȘ, - - ȠȟȚțȩȢ ĮȚșȣȜİıIJȑȡĮȢ (ȋȡȩȞȠȢ İțȤȪȜȚıȘȢ 2
ȫȡİȢ).....................................................................................................................................
109
ȈȤȒµĮ 4.1.3 ǼʌȓįȡĮıȘ IJȘȢ ıȣȖțȑȞIJȡȦıȘȢ IJȘȢ ĮȚșĮȞȩȜȘȢ ıIJȘȞ İțȤȪȜȚıȘ IJȠȣ
ȕ-țĮȡȠIJȑȞȚȠȣ Įʌȩ IJĮ țȪIJIJĮȡĮ IJȠȣ B. trispora (ȋȡȩȞȠȢ İțȤȪȜȚıȘȢ 2 ȫȡİȢ, șİȡµȠțȡĮıȓĮ
İțȤȪȜȚıȘȢ 30
0
C, ĮȡȚșµȩȢ įȚĮįȠȤȚțȫȞ İțȤȣȜȓıİȦȞ 3, 100 ml įȚĮȜȪIJȘ/g ȟȘȡȠȪ ȕȐȡȠȣȢ
ȕȚȠµȐȗĮȢ)..............................................................................................................................

111
ȈȤȒµĮ 4.1.4 ǼʌȓįȡĮıȘ IJȘȢ ĮȞĮȜȠȖȓĮȢ IJȠȣ ȩȖțȠȣ IJȘȢ ĮȚșĮȞȩȜȘȢ ʌȡȠȢ IJȠ ȟȘȡȩ ȕȐȡȠȢ
IJȘȢ ȕȚȠµȐȗĮȢ IJȠȣ B. trispora ıIJȘȞ İțȤȪȜȚıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ (ȋȡȩȞȠȢ İțȤȪȜȚıȘȢ 2
ȫȡİȢ, șİȡµȠțȡĮıȓĮ İțȤȪȜȚıȘȢ 30
0
C, ĮȡȚșµȩȢ įȚĮįȠȤȚțȫȞ İțȤȣȜȓıİȦȞ 3).........................

112
ȈȤȒµĮ 4.1.5 ǼʌȓįȡĮıȘ IJȦȞ įȚĮįȠȤȚțȫȞ İțȤȣȜȓıİȦȞ ıIJȘȞ ĮʌȠµȐțȡȣȞıȘ IJȠȣ
ȕ-țĮȡȠIJȑȞȚȠȣ Įʌȩ IJĮ țȪIJIJĮȡĮ IJȠȣ B. trispora µİ įȚĮijȠȡİIJȚțȠȪȢ įȚĮȜȪIJİȢ İțȤȪȜȚıȘȢ.

XIV
-- ĮȚșĮȞȩȜȘ, - - µİșĮȞȩȜȘ, - - ĮțİIJȩȞȘ, - - ʌİIJȡİȜĮȧțȩȢ ĮȚșȑȡĮȢ, -- İȟȐȞȚȠ, - -
µİșĮȞȩȜȘİȟȐȞȚȠ (11), - - ȚıȠʌȡȠʌȣȜȚțȒ ĮȜțȠȩȜȘ, - - ȠȟȚțȩȢ ĮȚșȣȜİıIJȑȡĮȢ (ȋȡȩȞȠȢ
İțȤȪȜȚıȘȢ 2 ȫȡİȢ, șİȡµȠțȡĮıȓĮ İțȤȪȜȚıȘȢ Ș ȐȡȚıIJȘ ȖȚĮ țȐșİ İțȤȣȜȚıIJȚțȩ µȑıȠ ȩʌȦȢ
ijĮȓȞİIJĮȚ ıIJȠ ıȤȒµĮ 4.1.2)....................................................................................................
114
ȈȤȒµĮ 4.2.1 ǹȞȐʌIJȣȟȘ IJȠȣ B. trispora țĮȚ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ țȦȞȚțȑȢ ijȚȐȜİȢ.
--, - -, - - : IJȠ ȣʌȩıIJȡȦµĮ ݵȕȠȜȚȐıIJȘțİ µİ țȪIJIJĮȡĮ IJȠȣ ıIJİȜȑȤȠȣȢ 14271, 14272
țĮȚ µȓȖµĮ IJȦȞ ʌĮȡĮʌȐȞȦ ıIJİȜİȤȫȞ, ĮȞIJȓıIJȠȚȤĮ. - -, --, - - : IJȠ ȣʌȩıIJȡȦµĮ
ݵȕȠȜȚȐıIJȘțİ µİ 1,0 %, 2,5 %, țĮȚ 5,0 % (v/v) Įʌȩ IJȠ țȐșİ ȑȞĮ ıIJȑȜİȤȠȢ ʌȠȣ
ĮȞĮʌIJȪȤșȘțĮȞ ȤȦȡȚıIJȐ ıİ ȣȖȡȩ ȣʌȩıIJȡȦµĮ ȖȚĮ 2 ȘµȑȡİȢ, ĮȞIJȓıIJȠȚȤĮ. - -: IJȠ
ȣʌȩıIJȡȦµĮ ݵȕȠȜȚȐıIJȘțİ µİ 5,0 % (v/v) µȓȖµĮ IJȦȞ 14271 țĮȚ 14272 ʌȠȣ ȑȤȠȣȞ
ĮȞĮʌIJȣȤșİȓ µĮȗȓ ıİ ıȣȞșİIJȚțȩ ȣʌȩıIJȡȦµĮ ȖȚĮ 2 ȘµȑȡİȢ. (ȊʌȩıIJȡȦµĮ ʌĮȡĮȖȦȖȒȢ, g/l:
ȖȜȣțȩȗȘ 50,0, İțȤȪȜȚıµĮ țĮȜĮµʌȠțȚȠȪ 80,0, ȣįȡȠȜȣµȑȞȘ țĮȗİȓȞȘ 2,0, İțȤȪȜȚıµĮ ȗȪµȘȢ
1,0, L-ĮıʌĮȡĮȖȓȞȘ 2,0, KH
2
PO
4
1,5, MgSO
4
. 7H
2
0 0,5, Span 20 10,0, Tween 80 1,0,
ȣįȡȠȤȜȦȡȚțȒ șİȚĮµȓȞȘ 5,0 mg/l. ĬİȡµȠțȡĮıȓĮ 26
0
C, ĮȡȤȚțȩ pH ȣʌȠıIJȡȫµĮIJȠȢ 7,
IJĮȤȪIJȘIJĮ ĮȞĮțȓȞȘıȘȢ 200 rpm)...........................................................................................
122
ȈȤȒµĮ 4.2.2 ȂİIJĮȕȠȜȒ IJȦȞ țȚȞȘIJȚțȫȞ ʌĮȡĮµȑIJȡȦȞ IJȘȢ ȗȪµȦıȘȢ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ
ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ țȦȞȚțȑȢ ijȚȐȜİȢ. -- ȕ-țĮȡȠIJȑȞȚȠ, - - ȕȚȠµȐȗĮ ȟȘȡȩ
ȕȐȡȠȢ, - -ȣʌȠȜİȚµµĮIJȚțȐ ıȐțȤĮȡĮ, - - pH (ȊʌȩıIJȡȦµĮ ʌĮȡĮȖȦȖȒȢ țĮȚ ıȣȞșȒțİȢ IJȘȢ
ȗȪµȦıȘȢ ȩʌȦȢ ıIJȠ ȈȤȒµĮ 4.2.1)..........................................................................................
127
ȈȤȒµĮ 4.2.3 ǼʌȓįȡĮıȘ IJȘȢ șİȡµȠțȡĮıȓĮȢ ıIJȚȢ țȚȞȘIJȚțȑȢ ʌĮȡĮµȑIJȡȠȣȢ IJȘȢ ȗȪµȦıȘȢ
țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ țȦȞȚțȑȢ ijȚȐȜİȢ. --, - -, - -, -
-  24, 26, 28 țĮȚ 30
0
C, ĮȞIJȓıIJȠȚȤĮ. (ȊʌȩıIJȡȦµĮ ʌĮȡĮȖȦȖȒȢ ȩʌȦȢ ıIJȠ ıȤȒµĮ 4.2.1.
ǹȡȤȚțȩ pH ȣʌȠıIJȡȫµĮIJȠȢ 7, IJĮȤȪIJȘIJĮ ĮȞĮțȓȞȘıȘȢ 200 rpm)............................................

130
ȈȤȒµĮ 4.2.4 ǼʌȓįȡĮıȘ IJȠȣ ĮȡȤȚțȠȪ pH IJȠȣ ȣʌȠıIJȡȫµĮIJȠȢ ıIJȚȢ țȚȞȘIJȚțȑȢ ʌĮȡĮµȑIJȡȠȣȢ
IJȘȢ ȗȪµȦıȘȢ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ țȦȞȚțȑȢ ijȚȐȜİȢ.
--, - -, - -, - -, --, - -, - -  ĮȡȤȚțȩ pH 5,0, 6,0, 7,0, 8,0, 9,0, 10,0 țĮȚ 11,0,
ĮȞIJȓıIJȠȚȤĮ. (ȊʌȩıIJȡȦµĮ ʌĮȡĮȖȦȖȒȢ ȩʌȦȢ ıIJȠ ıȤȒµĮ 4.2.1. ĬİȡµȠțȡĮıȓĮ 26
0
C,
IJĮȤȪIJȘIJĮ ĮȞĮțȓȞȘıȘȢ 200 rpm)...........................................................................................
133
ȈȤȒµĮ 4.2.5 ǼʌȓįȡĮıȘ IJȠȣ ijȦIJȚıµȠȪ ıIJȘȞ ĮȞȐʌIJȣȟȘ IJȠȣ B. trispora țĮȚ IJȘȞ ʌĮȡĮȖȦȖȒ
ȕ-țĮȡȠIJȑȞȚȠȣ ıİ țȦȞȚțȑȢ ijȚȐȜİȢ. --, - -, - -, - - țĮȚ - -, - -, - -, - - :
ȕ-țĮȡȠIJȑȞȚȠ țĮȚ ȕȚȠµȐȗĮ ȟȘȡȩ ȕȐȡȠȢ, ĮȞIJȓıIJȠȚȤĮ. Ǿ ȗȪµȦıȘ ȑȖȚȞİ : -- țĮȚ - -
ʌĮȡȠȣıȓĮ ijȦIJȩȢ, - - țĮȚ - - ĮʌȠȣıȓĮ ijȦIJȩȢ, - - țĮȚ - - IJȚȢ 3 ʌȡȫIJİȢ ȘµȑȡİȢ

XV
ĮʌȠȣıȓĮ ijȦIJȩȢ țĮȚ IJȚȢ ȣʌȩȜȠȚʌİȢ 5 ȘµȑȡİȢ ʌĮȡȠȣıȓĮ ijȦIJȩȢ, - - țĮȚ - - IJȚȢ 3 ʌȡȫIJİȢ
ȘµȑȡİȢ ʌĮȡȠȣıȓĮ ijȦIJȩȢ țĮȚ IJȚȢ ȣʌȩȜȠȚʌİȢ 5 ȘµȑȡİȢ ĮʌȠȣıȓĮ ijȦIJȩȢ. (ȊʌȩıIJȡȦµĮ
ʌĮȡĮȖȦȖȒȢ țĮȚ ıȣȞșȒțİȢ IJȘȢ ȗȪµȦıȘȢ ȩʌȦȢ ıIJȠ ȈȤȒµĮ 4.2.1).........................................
136
ȈȤȒµĮ 4.2.6 ǼʌȓįȡĮıȘ IJȘȢ ıȣȞİȡȖȚıIJȚțȒȢ įȡȐıȘȢ IJȠȣ ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ țĮȚ IJȠȣ
ĮȞIJȚȠȟİȚįȦIJȚțȠȪ ıIJȘȞ ĮȞȐʌIJȣȟȘ IJȠȣ B. trispora țĮȚ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ
țȦȞȚțȑȢ ijȚȐȜİȢ. --, - - țĮȚ - -, - -: ȕ-țĮȡȠIJȑȞȚȠ țĮȚ ȕȚȠµȐȗĮ ȟȘȡȩ ȕȐȡȠȢ,
ĮȞIJȓıIJȠȚȤĮ. ȈIJȠ ȕĮıȚțȩ ȣʌȩıIJȡȦµĮ ʌȠȣ ʌİȡȚȖȡȐijİIJĮȚ ıIJȠȞ ȆȓȞĮțĮ 4.2.6 ʌȡȠıIJȑșȘțİ:
-- țĮȚ - - ĮȞIJȚȠȟİȚįȦIJȚțȩ 10,0 g/l, - - țĮȚ - - µȓȖµĮ ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ țĮȚ
ĮȞIJȚȠȟİȚįȦIJȚțȠȪ (20,0 țĮȚ 10,0 g/l, ĮȞIJȓıIJȠȚȤĮ)..................................................................
158
ȈȤȒµĮ 4.2.7 ǼʌȓįȡĮıȘ IJȘȢ ıȣȞİȡȖȚıIJȚțȒȢ įȡȐıȘȢ IJȠȣ ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ țĮȚ IJȘȢ
țȘȡȠȗȓȞȘȢ ıIJȘȞ ĮȞȐʌIJȣȟȘ IJȠȣ B. trispora țĮȚ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ țȦȞȚțȑȢ
ijȚȐȜİȢ. --, - -, - - țĮȚ - -, - -, - - : ȕ-țĮȡȠIJȑȞȚȠ țĮȚ ȕȚȠµȐȗĮ ȟȘȡȩ ȕȐȡȠȢ,
ĮȞIJȓıIJȠȚȤĮ. ȈIJȠ ȕĮıȚțȩ ȣʌȩıIJȡȦµĮ ʌȠȣ ʌİȡȚȖȡȐijİIJĮȚ ıIJȠȞ ȆȓȞĮțĮ 4.2.6 ʌȡȠıIJȑșȘțİ:
-- țĮȚ - - µȓȖµĮ ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ 20,0 g/l țĮȚ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ 10,0 g/l, - - țĮȚ - -
țȘȡȠȗȓȞȘ 40,0 g/l, - - țĮȚ - - µȓȖµĮ ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ 20,0 g/l, ĮȞIJȚȠȟİȚįȦIJȚțȠȪ 10,0
g/l țĮȚ țȘȡȠȗȓȞȘȢ 40,0 g/l................................................................................................
164
ȈȤȒµĮ 4.2.8 ȉȣʌȠʌȠȚȘµȑȞİȢ İʌȚįȡȐıİȚȢ IJȠȣ ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ (ȋ
1
), IJȘȢ țȘȡȠȗȓȞȘȢ (ȋ
2
)
țĮȚ IJȠȣ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ (ȋ
3
) ıIJȘ ıȣȖțȑȞIJȡȦıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ....................................
178
ȈȤȒµĮ 4.2.9 ȉȡȚıįȚȐıIJĮIJȘ ĮʌİȚțȩȞȚıȘ IJȘȢ İʌȚijȐȞİȚĮȢ ĮʌȩțȡȚıȘȢ IJȘȢ ıȣȖțȑȞIJȡȦıȘȢ
IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ įȚĮijȠȡİIJȚțȑȢ ıȣȖțİȞIJȡȫıİȚȢ ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ țĮȚ țȘȡȠȗȓȞȘȢ
(ȈȣȖțȑȞIJȡȦıȘ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ 9,04

g/l)............................................................................
179
ȈȤȒµĮ 4.2.10 ȉȡȚıįȚȐıIJĮIJȘ ĮʌİȚțȩȞȚıȘ IJȘȢ İʌȚijȐȞİȚĮȢ ĮʌȩțȡȚıȘȢ IJȘȢ ıȣȖțȑȞIJȡȦıȘ
IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ įȚĮijȠȡİIJȚțȑȢ ıȣȖțİȞIJȡȫıİȚȢ ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ țĮȚ
ĮȞIJȚȠȟİȚįȦIJȚțȠȪ (ȈȣȖțȑȞIJȡȦıȘ țȘȡȠȗȓȞȘȢ 39,25 g/l).........................................................
180
ȈȤȒµĮ 4.2.11 ȉȡȚıįȚȐıIJĮIJȘ ĮʌİȚțȩȞȚıȘ IJȘȢ İʌȚijȐȞİȚĮȢ ĮʌȩțȡȚıȘȢ IJȘȢ ıȣȖțȑȞIJȡȦıȘȢ
IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ įȚĮijȠȡİIJȚțȑȢ ıȣȖțİȞIJȡȫıİȚȢ țȘȡȠȗȓȞȘȢ țĮȚ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ
(ȈȣȖțȑȞIJȡȦıȘ ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ 17,15 g/l).....................................................................
181
ȈȤȒµĮ 4.3.1 ǼʌȓįȡĮıȘ IJȘȢ ıȣȖțȑȞIJȡȦıȘȢ IJȠȣ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ țĮȚ IJȠȣ ȜȚȞİȜĮȧțȠȪ
ȠȟȑȠȢ ıIJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ
ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ. a, b, c : ǺĮıȚțȩ ȣʌȩıIJȡȦµĮ ʌȠȣ ʌİȡȚȑȤİȚ 0, 2,5 țĮȚ 5,0 g/l
ĮȞIJȚȠȟİȚįȦIJȚțȩ, ĮȞIJȓıIJȠȚȤĮ. --, - -, - -, --  ȜȚȞİȜĮȧțȩ ȠȟȪ 0, 5,0, 10,0 țĮȚ 15,0
g/l, ĮȞIJȓıIJȠȚȤĮ......................................................................................................................
188
ȈȤȒµĮ 4.3.2 ǼʌȓįȡĮıȘ IJȘȢ ıȣȖțȑȞIJȡȦıȘȢ IJȠȣ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ țĮȚ IJȠȣ ȜȚȞİȜĮȧțȠȪ
XVI
ȠȟȑȠȢ ıIJȚȢ țȚȞȘIJȚțȑȢ ʌĮȡĮµȑIJȡȠȣȢ IJȘȢ ĮıȣȞİȤȠȪȢ ȗȪµȦıȘȢ ȕȣșȠȪ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ
ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ. a, b, c : ǺĮıȚțȩ ȣʌȩıIJȡȦµĮ
ʌȠȣ ʌİȡȚȑȤİȚ 0, 2,5 țĮȚ 5,0 g/l ĮȞIJȚȠȟİȚįȦIJȚțȩ, ĮȞIJȓıIJȠȚȤĮ. --, - -, - -, -- țĮȚ - -,
- -, - -, -- ȕȚȠµȐȗĮ ȟȘȡȩ ȕȐȡȠȢ țĮȚ ȣʌȠȜİȚµµĮIJȚțȐ ıȐțȤĮȡĮ ıİ ıȣȖțİȞIJȡȫıİȚȢ
ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ 0, 5,0, 10,0 țĮȚ 15,0 g/l, ĮȞIJȓıIJȠȚȤĮ......................................................
189
ȈȤȒµĮ 4.3.3 ǼʌȓįȡĮıȘ IJȠȣ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ țĮȚ IJȠȣ ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ ıIJȘ ıȣȖțȑȞIJȡȦıȘ
IJȠȣ įȚĮȜȣIJȠȪ ȠȟȣȖȩȞȠȣ țĮȚ ıIJȠ pH IJȠȣ ȣȖȡȠȪ IJȘȢ ȗȪµȦıȘȢ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ
ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ. a,
b, c : ǺĮıȚțȩ ȣʌȩıIJȡȦµĮ ʌȠȣ ʌİȡȚȑȤİȚ 0, 2,5 țĮȚ 5,0 g/l ĮȞIJȚȠȟİȚįȦIJȚțȩ, ĮȞIJȓıIJȠȚȤĮ.
--, - -, - -, -- țĮȚ - -, - -, - -, --  įȚĮȜȣIJȩ ȠȟȣȖȩȞȠ țĮȚ pH ıİ ıȣȖțİȞIJȡȫıİȚȢ
ȜȚȞİȜĮȧțȠȪ ȠȟȑȠȢ 0, 5,0, 10,0 țĮȚ 15,0 g/l, ĮȞIJȓıIJȠȚȤĮ.............................
190
ȈȤȒµĮ 4.3.4 ǼʌȓįȡĮıȘ IJȘȢ țȘȡȠȗȓȞȘȢ ıIJȚȢ țȚȞȘIJȚțȑȢ ʌĮȡĮµȑIJȡȠȣȢ IJȘȢ ĮıȣȞİȤȠȪȢ
ȗȪµȦıȘȢ ȕȣșȠȪ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ȗȣµȦIJȒȡĮ
ĮȞȐįİȣıȘȢ. ȉȠ ȕĮıȚțȩ ȣʌȩıIJȡȦµĮ ʌȠȣ ʌİȡȚİȓȤİ 5,0 g/l ȜȚȞİȜĮȧțȩ ȠȟȪ țĮȚ 2,5 g/l
ĮȞIJȚȠȟİȚįȦIJȚțȩ ݵʌȜȠȣIJȓıIJȘțİ µİ țȘȡȠȗȓȞȘ 5,0 g/l (a) țĮȚ 10,0 g/l (b) IJȘȞ 3Ș ȘµȑȡĮ IJȘȢ
ȗȪµȦıȘȢ țĮȚ 5,0 g/l IJȘȞ 6Ș ȘµȑȡĮ IJȘȢ ȗȪµȦıȘȢ (c). -- ȕ-țĮȡȠIJȑȞȚȠ, - - ȕȚȠµȐȗĮ ȟȘȡȩ
ȕȐȡȠȢ, - - ȣʌȠȜİȚµµĮIJȚțȐ ıȐțȤĮȡĮ. ʌȡȠıșȒțȘ țȘȡȠȗȓȞȘȢ ıIJȠ
ȣʌȩıIJȡȦµĮ...........................................................................................................................
194
ȈȤȒµĮ 4.3.5 ǼʌȓįȡĮıȘ IJȘȢ țȘȡȠȗȓȞȘȢ ıIJȘ ıȣȖțȑȞIJȡȦıȘ IJȠȣ įȚĮȜȣIJȠȪ ȠȟȣȖȩȞȠȣ țĮȚ ıIJȠ
pH IJȠȣ ȣȖȡȠȪ IJȘȢ ȗȪµȦıȘȢ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ
ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ. ȉȠ ȕĮıȚțȩ ȣʌȩıIJȡȦµĮ ʌȠȣ ʌİȡȚİȓȤİ
5,0 g/l ȜȚȞİȜĮȧțȩ ȠȟȪ țĮȚ 2,5 g/l ĮȞIJȚȠȟİȚįȦIJȚțȩ ݵʌȜȠȣIJȓıIJȘțİ µİ țȘȡȠȗȓȞȘ 5,0 g/l (a)
țĮȚ 10,0 g/l (b) IJȘȞ 3Ș ȘµȑȡĮ IJȘȢ ȗȪµȦıȘȢ țĮȚ 5,0 g/l IJȘȞ 6Ș ȘµȑȡĮ IJȘȢ ȗȪµȦıȘȢ (c).
-- įȚĮȜȣIJȩ ȠȟȣȖȩȞȠ, - - pH. ʌȡȠıșȒțȘ țȘȡȠȗȓȞȘȢ ıIJȠ ȣʌȩıIJȡȦµĮ…………...........
195
ȈȤȒµĮ 4.3.6 ǼʌȓįȡĮıȘ IJȘȢ ıȣȖțȑȞIJȡȦıȘȢ IJȠȣ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ țĮȚ IJȦȞ ijȣIJȚțȫȞ İȜĮȓȦȞ
ıIJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ ȗȣµȦIJȒȡĮ
ĮȞȐįİȣıȘȢ. a, b, c : ǺĮıȚțȩ ȣʌȩıIJȡȦµĮ ʌȠȣ ʌİȡȚȑȤİȚ 0, 2,5 țĮȚ 5,0 g/l ĮȞIJȚȠȟİȚįȦIJȚțȩ,
ĮȞIJȓıIJȠȚȤĮ. --, - -, - - ȚıȠµȠȡȚĮțȩ µȓȖµĮ İȜĮȚȩȜĮįȠȣ, ıȠȖȚȑȜĮȚȠȣ țĮȚ
ȕĮµȕĮțȑȜĮȚȠȣ 15,0, 30,0 țĮȚ 45,0 g/l, ĮȞIJȓıIJȠȚȤĮ...............................................................
202
ȈȤȒµĮ 4.3.7 ǼʌȓįȡĮıȘ IJȘȢ ıȣȖțȑȞIJȡȦıȘȢ IJȦȞ ijȣIJȚțȫȞ İȜĮȓȦȞ țĮȚ IJȠȣ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ
ıIJȚȢ țȚȞȘIJȚțȑȢ ʌĮȡĮµȑIJȡȠȣȢ IJȘȢ ĮıȣȞİȤȠȪȢ ȗȪµȦıȘȢ ȕȣșȠȪ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ
ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ. a, b, c: ǺĮıȚțȩ ȣʌȩıIJȡȦµĮ ʌȠȣ

XVII
ʌİȡȚȑȤİȚ 0, 2,5 țĮȚ 5,0 g/l ĮȞIJȚȠȟİȚįȦIJȚțȩ, ĮȞIJȓıIJȠȚȤĮ. --, - -, - - țĮȚ - -, - -, - 
ȕȚȠµȐȗĮ ȟȘȡȩ ȕȐȡȠȢ țĮȚ ȣʌȠȜİȚµµĮIJȚțȐ ıȐțȤĮȡĮ ıİ ıȣȖțİȞIJȡȫıİȚȢ ȚıȠµȠȡȚĮțȠȪ
µȓȖµĮIJȠȢ İȜĮȚȩȜĮįȠȣ, ıȠȖȚȑȜĮȚȠȣ țĮȚ ȕĮµȕĮțȑȜĮȚȠȣ 15,0, 30,0 țĮȚ 45,0 g/l,
ĮȞIJȓıIJȠȚȤĮ............................................................................................................................
203
ȈȤȒµĮ 4.3.8 ǼʌȓįȡĮıȘ IJȠȣ ĮȞIJȚȠȟİȚįȦIJȚțȠȪ țĮȚ IJȦȞ ijȣIJȚțȫȞ İȜĮȓȦȞ ıIJȘ ıȣȖțȑȞIJȡȦıȘ
IJȠȣ įȚĮȜȣIJȠȪ ȠȟȣȖȩȞȠȣ țĮȚ ıIJȠ pH IJȠȣ ȣȖȡȠȪ IJȘȢ ȗȪµȦıȘȢ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ
ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ. a,
b, c : ǺĮıȚțȩ ȣʌȩıIJȡȦµĮ ʌȠȣ ʌİȡȚȑȤİȚ 0, 2,5 țĮȚ 5,0 g/l ĮȞIJȚȠȟİȚįȦIJȚțȩ, ĮȞIJȓıIJȠȚȤĮ.
--, - -, - - țĮȚ - -, - -, - -  įȚĮȜȣIJȩ ȠȟȣȖȩȞȠ țĮȚ pH ıİ ıȣȖțİȞIJȡȫıİȚȢ
ȚıȠµȠȡȚĮțȠȪ µȓȖµĮIJȠȢ İȜĮȚȩȜĮįȠȣ, ıȠȖȚȑȜĮȚȠȣ țĮȚ ȕĮµȕĮțȑȜĮȚȠȣ 0, 30,0 țĮȚ 45,0 g/l,
ĮȞIJȓıIJȠȚȤĮ............................................................................................................................

204
ȈȤȒµĮ 4.3.9 ǼʌȓįȡĮıȘ IJȘȢ țȘȡȠȗȓȞȘȢ ıIJȚȢ țȚȞȘIJȚțȑȢ ʌĮȡĮµȑIJȡȠȣȢ IJȘȢ ĮıȣȞİȤȠȪȢ
ȗȪµȦıȘȢ ȕȣșȠȪ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ȗȣµȦIJȒȡĮ
ĮȞȐįİȣıȘȢ. To ȕĮıȚțȩ ȣʌȩıIJȡȦµĮ ʌȠȣ ʌİȡȚİȓȤİ 2,5 g/l ĮȞIJȚȠȟİȚįȦIJȚțȩ țĮȚ 30,0 g/l
ȚıȠµȠȡȚĮțȩ µȓȖµĮ İȜĮȚȩȜĮįȠȣ, ıȠȖȚȑȜĮȚȠȣ țĮȚ ȕĮµȕĮțȑȜĮȚȠȣ ݵʌȜȠȣIJȓıIJȘțİ µİ 5,0 g/l
țȘȡȠȗȓȞȘ IJȘȞ 3
Ș
ȘµȑȡĮ IJȘȢ ȗȪµȦıȘȢ. -- ȕ-țĮȡȠIJȑȞȚȠ, - - ȕȚȠµȐȗĮ ȟȘȡȩ ȕȐȡȠȢ, - -
ȣʌȠȜİȚµµĮIJȚțȐ ıȐțȤĮȡĮ -- įȚĮȜȣIJȩ ȠȟȣȖȩȞȠ, -- pH......................................................
205
ȈȤȒµĮ 4.3.10 ǼʌȓįȡĮıȘ IJȘȢ ĮȞȐįİȣıȘȢ țĮȚ IJȠȣ ĮİȡȚıµȠȪ ıIJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ
µİ IJȠ B. trispora ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ. a, b, c, d, e
0, 150, 325, 500 țĮȚ 700 rpm, ĮȞIJȓıIJȠȚȤĮ. --, - -, - -, - -, --  0, 0,5, 1,0, 1,5
țĮȚ 2,0 vvm, ĮȞIJȓıIJȠȚȤĮ.......................................................................................................
226
ȈȤȒµĮ 4.3.11 ǼʌȓįȡĮıȘ IJȘȢ ĮȞȐįİȣıȘȢ țĮȚ IJȠȣ ĮİȡȚıµȠȪ ıIJȚȢ țȚȞȘIJȚțȑȢ ʌĮȡĮµȑIJȡȠȣȢ
IJȘȢ ĮıȣȞİȤȠȪȢ ȗȪµȦıȘȢ ȕȣșȠȪ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ
ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ. a, b, c, d, e  0, 150, 325, 500 țĮȚ 700 rpm, ĮȞIJȓıIJȠȚȤĮ. --, - -,
- -, - -, -- țĮȚ - -, - -, - -, - -, -- ȕȚȠµȐȗĮ ȟȘȡȩ ȕȐȡȠȢ țĮȚ ȣʌȠȜİȚµµĮIJȚțȐ
ıȐțȤĮȡĮ ıİ ıȣȞșȒțİȢ ĮİȡȚıµȠȪ 0, 0,5, 1,0, 1,5 țĮȚ 2,0 vvm, ĮȞIJȓıIJȠȚȤĮ...........................
227
ȈȤȒµĮ 4.3.12 ǼʌȓįȡĮıȘ IJȘȢ ĮȞȐįİȣıȘȢ țĮȚ IJȠȣ ĮİȡȚıµȠȪ ıIJȘ ıȣȖțȑȞIJȡȦıȘ IJȠȣ
įȚĮȜȣIJȠȪ ȠȟȣȖȩȞȠȣ țĮȚ ıIJȠ pH IJȠȣ ȣȖȡȠȪ IJȘȢ ȗȪµȦıȘȢ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-
țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ. a, b,
c, d, e  0, 150, 325, 500 țĮȚ 700 rpm, ĮȞIJȓıIJȠȚȤĮ. - -, - -, - -, - -, -- țĮȚ --,
- -, - -, - -, --  pH țĮȚ ıȣȖțȑȞIJȡȦıȘ įȚĮȜȣIJȠȪ ȠȟȣȖȩȞȠȣ ıİ ıȣȞșȒțİȢ ĮİȡȚıµȠȪ
0 0 5 1 0 1 5 2 0 ȓ

XVIII
0, 0,5, 1,0, 1,5 țĮȚ 2,0 vvm, ĮȞIJȓıIJȠȚȤĮ.............................................................................. 228
ȈȤȒµĮ 4.3.13 ȉȣʌȠʌȠȚȘµȑȞİȢ İʌȚįȡȐıİȚȢ IJȘȢ IJĮȤȪIJȘIJĮȢ ĮȞȐįİȣıȘȢ (ȋ
1
) țĮȚ IJȠȣ ȡȣșµȠȪ
ʌĮȡȠȤȒȢ ĮȑȡĮ (ȋ
2
) ıIJȘ ıȣȖțȑȞIJȡȦıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ....................................................
246
ȈȤȒµĮ 4.3.14 ȉȣʌȠʌȠȚȘµȑȞİȢ İʌȚįȡȐıİȚȢ IJȘȢ IJĮȤȪIJȘIJĮȢ ĮȞȐįİȣıȘȢ (ȋ
1
) țĮȚ IJȠȣ ȡȣșµȠȪ
ʌĮȡȠȤȒȢ ĮȑȡĮ (ȋ
2
) ıIJȘȞ ʌĮȡĮȖȦȖȚțȩIJȘIJĮ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ..............................................
247
ȈȤȒµĮ 4.3.15 ȉȣʌȠʌȠȚȘµȑȞİȢ İʌȚįȡȐıİȚȢ IJȘȢ IJĮȤȪIJȘIJĮȢ ĮȞȐįİȣıȘȢ (ȋ
1
) țĮȚ IJȠȣ ȡȣșµȠȪ
ʌĮȡȠȤȒȢ ĮȑȡĮ (ȋ
2
) ıIJȘ ıȣȖțȑȞIJȡȦıȘ IJȘȢ ȕȚȠµȐȗĮȢ............................................................
248
ȈȤȒµĮ 4.3.16 ȉȣʌȠʌȠȚȘµȑȞİȢ İʌȚįȡȐıİȚȢ IJȘȢ IJĮȤȪIJȘIJĮȢ ĮȞȐįİȣıȘȢ (ȋ
1
) țĮȚ IJȠȣ ȡȣșµȠȪ
ʌĮȡȠȤȒȢ ĮȑȡĮ (ȋ
2
) ıIJȠȞ ȠȖțȠµİIJȡȚțȩ ıȣȞIJİȜİıIJȒ µİIJĮijȠȡȐȢ
µȐȗĮȢ...................................................................................................................................
249
ȈȤȒµĮ 4.3.17 ȉȡȚıįȚȐıIJĮIJȘ ĮʌİȚțȩȞȚıȘ IJȘȢ İʌȚijȐȞİȚĮȢ ĮʌȩțȡȚıȘȢ IJȘȢ ıȣȖțȑȞIJȡȦıȘȢ
IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ įȚĮijȠȡİIJȚțȑȢ ıȣȞșȒțİȢ ĮȞȐįİȣıȘȢ țĮȚ ĮİȡȚıµȠȪ..............................
250
ȈȤȒµĮ 4.3.18 ȉȡȚıįȚȐıIJĮIJȘ ĮʌİȚțȩȞȚıȘ IJȘȢ İʌȚijȐȞİȚĮȢ ĮʌȩțȡȚıȘȢ IJȘȢ
ʌĮȡĮȖȦȖȚțȩIJȘIJĮȢ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ įȚĮijȠȡİIJȚțȑȢ ıȣȞșȒțİȢ ĮȞȐįİȣıȘȢ țĮȚ
ĮİȡȚıµȠȪ…………………………………………………………………………………...
251
ȈȤȒµĮ 4.3.19 ȉȡȚıįȚȐıIJĮIJȘ ĮʌİȚțȩȞȚıȘ IJȘȢ İʌȚijȐȞİȚĮȢ ĮʌȩțȡȚıȘȢ IJȘȢ ȕȚȠµȐȗĮȢ ȟȘȡȩ
ȕȐȡȠȢ ıİ įȚĮijȠȡİIJȚțȑȢ ıȣȞșȒțİȢ ĮȞȐįİȣıȘȢ țĮȚ ĮİȡȚıµȠȪ................................................
252
ȈȤȒµĮ 4.3.20 ȉȡȚıįȚȐıIJĮIJȘ ĮʌİȚțȩȞȚıȘ IJȘȢ İʌȚijȐȞİȚĮȢ ĮʌȩțȡȚıȘȢ IJȠȣ ȠȖțȠµİIJȡȚțȠȪ
ıȣȞIJİȜİıIJȒ µİIJĮijȠȡȐȢ µȐȗĮȢ ıİ įȚĮijȠȡİIJȚțȑȢ ıȣȞșȒțİȢ ĮȞȐįİȣıȘȢ țĮȚ
ĮİȡȚıµȠȪ...............................................................................................................................
253
ȈȤȒµĮ 4.4.1 ǼʌȓįȡĮıȘ IJȠȣ ȤȡȩȞȠȣ ȑȞĮȡȟȘȢ IJȘȢ IJȡȠijȠįȠıȓĮȢ ıIJȚȢ țȚȞȘIJȚțȑȢ
ʌĮȡĮµȑIJȡȠȣȢ IJȘȢ ĮıȣȞİȤȠȪȢ ȗȪµȦıȘȢ ȕȣșȠȪ µİ ıȣȞİȤȒ ʌȡȠıșȒțȘ ȣʌȠıIJȡȫµĮIJȠȢ țĮIJȐ
IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ. a, b: ʌȡȠıșȒțȘ
1l ȣʌȠıIJȡȫµĮIJȠȢ IJȡȠijȠįȠıȓĮȢ 1 µİ ȡȣșµȩ 3,5 ml/h µİIJȐ IJȘȞ 2Ș ȘµȑȡĮ IJȘȢ ȗȪµȦıȘȢ țĮȚ
Įʌȩ IJȘȞ ĮȡȤȒ IJȘȢ ȗȪµȦıȘȢ, ĮȞIJȓıIJȠȚȤĮ. c: ʌȡȠıșȒțȘ 1l ȣʌȠıIJȡȫµĮIJȠȢ IJȡȠijȠįȠıȓĮȢ 2
µİ ȡȣșµȩ 3,5 ml/h Įʌȩ IJȘȞ ĮȡȤȒ IJȘȢ ȗȪµȦıȘȢ. -- ȕ-țĮȡȠIJȑȞȚȠ, - - ȕȚȠµȐȗĮ ȟȘȡȩ
ȕȐȡȠȢ, - - ȣʌȠȜİȚµµĮIJȚțȐ ıȐțȤĮȡĮ (ȊʌȩıIJȡȦµĮ ĮȞȐʌIJȣȟȘȢ 1,  ȑȞĮȡȟȘ
IJȡȠijȠįȠıȓĮȢ)........................................................................................................................
259
ȈȤȒµĮ 4.4.2 ǼʌȓįȡĮıȘ IJȠȣ ȤȡȩȞȠȣ ȑȞĮȡȟȘȢ IJȘȢ IJȡȠijȠįȠıȓĮȢ ıIJȘ ıȣȖțȑȞIJȡȦıȘ IJȠȣ
įȚĮȜȣIJȠȪ ȠȟȣȖȩȞȠȣ țĮȚ ıIJȠ pH IJȠȣ ȣȖȡȠȪ IJȘȢ ȗȪµȦıȘȢ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ
ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ µİ ıȣȞİȤȒ ʌȡȠıșȒțȘ
ȣʌȠıIJȡȫµĮIJȠȢ ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ. a, b: ʌȡȠıșȒțȘ 1l ȣʌȠıIJȡȫµĮIJȠȢ IJȡȠijȠįȠıȓĮȢ
1 µİ ȡȣșµȩ 3,5 ml/h µİIJȐ IJȘȞ 2Ș ȘµȑȡĮ IJȘȢ ȗȪµȦıȘȢ țĮȚ Įʌȩ IJȘȞ ĮȡȤȒ IJȘȢ ȗȪµȦıȘȢ,

XIX
ĮȞIJȓıIJȠȚȤĮ. c: ʌȡȠıșȒțȘ 1l ȣʌȠıIJȡȫµĮIJȠȢ IJȡȠijȠįȠıȓĮȢ 2 µİ ȡȣșµȩ 3,5 ml/h Įʌȩ IJȘȞ
ĮȡȤȒ IJȘȢ ȗȪµȦıȘȢ. -- įȚĮȜȣIJȩ ȠȟȣȖȩȞȠ, - - pH (ȊʌȩıIJȡȦµĮ ĮȞȐʌIJȣȟȘȢ 1,  ȑȞĮȡȟȘ
IJȡȠijȠįȠıȓĮȢ)........................................................................................................................
260
ȈȤȒµĮ 4.4.3 ǼʌȓįȡĮıȘ IJȘȢ µİșȩįȠȣ țĮȚ IJȠȣ ȡȣșµȠȪ IJȡȠijȠįȠıȓĮȢ (ȣʌȩıIJȡȦµĮ
IJȡȠijȠįȠıȓĮȢ 3) ıIJȚȢ țȚȞȘIJȚțȑȢ ʌĮȡĮµȑIJȡȠȣȢ IJȘȢ ȗȪµȦıȘȢ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-
țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ. a, b: ȡȣșµȩȢ IJȡȠijȠįȠıȓĮȢ 4,2 țĮȚ
8,4 ml/h Įʌȩ IJȘȞ ĮȡȤȒ IJȘȢ ȗȪµȦıȘȢ, ĮȞIJȓıIJȠȚȤĮ. c: ʌȡȠıșȒțȘ 667 ml ȣʌȠıIJȡȫµĮIJȠȢ
IJȡȠijȠįȠıȓĮȢ 3 IJȘȞ 2Ș, 8Ș țĮȚ 14Ș ȘµȑȡĮ IJȘȢ ȗȪµȦıȘȢ. -- ȕ-țĮȡȠIJȑȞȚȠ, - - ȕȚȠµȐȗĮ
ȟȘȡȩ ȕȐȡȠȢ, - - ȣʌȠȜİȚµµĮIJȚțȐ ıȐțȤĮȡĮ (ȊʌȩıIJȡȦµĮ ĮȞȐʌIJȣȟȘȢ 2,  ʌȡȠıșȒțȘ
ȣʌȠıIJȡȫµĮIJȠȢ IJȡȠijȠįȠıȓĮȢ 3)..........................................................................................
265
ȈȤȒµĮ 4.4.4 ǼʌȓįȡĮıȘ IJȘȢ µİșȩįȠȣ țĮȚ IJȠȣ ȡȣșµȠȪ IJȡȠijȠįȠıȓĮȢ (ȣʌȩıIJȡȦµĮ
IJȡȠijȠįȠıȓĮȢ 3) ıIJȘ ıȣȖțȑȞIJȡȦıȘ IJȠȣ įȚĮȜȣIJȠȪ ȠȟȣȖȩȞȠȣ țĮȚ ıIJȠ pH IJȠȣ ȣȖȡȠȪ IJȘȢ
ȗȪµȦıȘȢ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ.
a, b: ȡȣșµȩȢ IJȡȠijȠįȠıȓĮȢ 4,2 țĮȚ 8,4 ml/h Įʌȩ IJȘȞ ĮȡȤȒ IJȘȢ ȗȪµȦıȘȢ, ĮȞIJȓıIJȠȚȤĮ. c:
ʌȡȠıșȒțȘ 667 ml ȣʌȠıIJȡȫµĮIJȠȢ IJȡȠijȠįȠıȓĮȢ 3 IJȘȞ 2Ș, 8Ș țĮȚ 14Ș ȘµȑȡĮ IJȘȢ
ȗȪµȦıȘȢ. -- įȚĮȜȣIJȩ ȠȟȣȖȩȞȠ, - - pH (ȊʌȩıIJȡȦµĮ ĮȞȐʌIJȣȟȘȢ 2,  ʌȡȠıșȒțȘ
ȣʌȠıIJȡȫµĮIJȠȢ IJȡȠijȠįȠıȓĮȢ 3)............................................................................................
266
ȈȤȒµĮ 4.4.5 ǼʌȓįȡĮıȘ IJȘȢ ıȪıIJĮıȘȢ IJȠȣ ȣʌȠıIJȡȫµĮIJȠȢ IJȡȠijȠįȠıȓĮȢ ıIJȚȢ țȚȞȘIJȚțȑȢ
ʌĮȡĮµȑIJȡȠȣȢ IJȘȢ ȗȪµȦıȘȢ ȕȣșȠȪ țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ
ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ. a, b, c, d: ȣʌȩıIJȡȦµĮ IJȡȠijȠįȠıȓĮȢ 4, 5, 6 țĮȚ 7, ĮȞIJȓıIJȠȚȤĮ.
ȇȣșµȩȢ IJȡȠijȠįȠıȓĮȢ 4,2 ml/h Įʌȩ IJȘȞ ĮȡȤȒ IJȘȢ ȗȪµȦıȘȢ. -- ȕ-țĮȡȠIJȑȞȚȠ, - -
ȕȚȠµȐȗĮ ȟȘȡȩ ȕȐȡȠȢ, - - ȣʌȠȜİȚµµĮIJȚțȐ ıȐțȤĮȡĮ (ȊʌȩıIJȡȦµĮ ĮȞȐʌIJȣȟȘȢ 2)................
271
ȈȤȒµĮ 4.4.6 ǼʌȓįȡĮıȘ IJȘȢ ıȪıIJĮıȘȢ IJȠȣ ȣʌȠıIJȡȫµĮIJȠȢ IJȡȠijȠįȠıȓĮȢ ıIJȘ
ıȣȖțȑȞIJȡȦıȘ IJȠȣ įȚĮȜȣIJȠȪ ȠȟȣȖȩȞȠȣ țĮȚ ıIJȠ pH IJȠȣ ȣȖȡȠȪ IJȘȢ ȗȪµȦıȘȢ țĮIJȐ IJȘȞ
ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ µİ ıȣȞİȤȒ
ʌȡȠıșȒțȘ ȣʌȠıIJȡȫµĮIJȠȢ ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ. a, b, c, d: ȣʌȩıIJȡȦµĮ IJȡȠijȠįȠıȓĮȢ
4, 5, 6 țĮȚ 7, ĮȞIJȓıIJȠȚȤĮ. ȇȣșµȩȢ IJȡȠijȠįȠıȓĮȢ 4,2 ml/h Įʌȩ IJȘȞ ĮȡȤȒ IJȘȢ ȗȪµȦıȘȢ. --
įȚĮȜȣIJȩ ȠȟȣȖȩȞȠ, - - pH (ȊʌȩıIJȡȦµĮ ĮȞȐʌIJȣȟȘȢ 2)........................................................
272
ȈȤȒµĮ 4.5.1 ǺȚȠıȪȞșİıȘ IJȠȣ H
2
O
2
țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ B. trispora
ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ ĮȞĮțȚȞȠȪµİȞİȢ țȦȞȚțȑȢ ijȚȐȜİȢ (a), ıİ ȗȣµȦIJȒȡĮ
ĮȞȐįİȣıȘȢ (b) țĮȚ ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ µİ ıȣȞİȤȒ ʌȡȠıșȒțȘ ȣʌȠıIJȡȫµĮIJȠȢ ıİ
ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ (c). -- and - - ıȣȖțȑȞIJȡȦıȘ H
2
O
2
ıIJȘ ȕȚȠµȐȗĮ țĮȚ ıIJȠ ȣȖȡȩ
IJȘȢ ȗȪµȦıȘȢ, ĮȞIJȓıIJȠȚȤĮ

.....................................................................................................
286

XX
ǼȊȇǼȉǾȇǿȅ ǼǿȀȅȃȍȃ

ıİȜ.
ǼȚțȩȞĮ 3.4.1 ĭȦIJȠȖȡĮijȓĮ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ ʌȠȣ ȤȡȘıȚµȠʌȠȚȒșȘțİ ȖȚĮ IJȘȞ
ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ IJȠ µȪțȘIJĮ B. trispora...............................................

84
ǼȚțȩȞĮ 4.2.1 ǹʌİȚțȩȞȚıȘ IJȦȞ µȠȡijȠȜȠȖȚțȫȞ ȤĮȡĮțIJȘȡȚıIJȚțȫȞ IJȠȣ B. trispora
ıIJȘ µȑȖȚıIJȘ ıȣȖțȑȞIJȡȦıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ țȦȞȚțȑȢ ijȚȐȜİȢ (µȠȞȐįĮ
µȒțȠȣȢ 50 µm). a, b, c: IJȠ ȣʌȩıIJȡȦµĮ ݵȕȠȜȚȐıIJȘțİ µİ țȪIJIJĮȡĮ IJȠȣ
ıIJİȜȑȤȠȣȢ 14271, 14272 țĮȚ µȓȖµĮ IJȦȞ ʌĮȡĮʌȐȞȦ ıIJİȜİȤȫȞ, ĮȞIJȓıIJȠȚȤĮ. d, e:
IJȠ ȣʌȩıIJȡȦµĮ ݵȕȠȜȚȐıIJȘțİ µİ 2,5% (v/v) Įʌȩ IJȠ țȐșİ ȑȞĮ ıIJȑȜİȤȠȢ ʌȠȣ
ĮȞĮʌIJȪȤșȘțĮȞ ȤȦȡȚıIJȐ ıİ ȣȖȡȩ ȣʌȩıIJȡȦµĮ ȖȚĮ 2 ȘµȑȡİȢ țĮȚ 5% (v/v) µȓȖµĮ
IJȦȞ 14271 țĮȚ 14272 ʌȠȣ ĮȞĮʌIJȪȤșȘțĮȞ µĮȗȓ ıİ ıȣȞșİIJȚțȩ ȣʌȩıIJȡȦµĮ ȖȚĮ 2
ȘµȑȡİȢ, ĮȞIJȓıIJȠȚȤĮ................................................................................................
124
ǼȚțȩȞĮ 4.2.2 ǹʌİȚțȩȞȚıȘ IJȦȞ µȠȡijȠȜȠȖȚțȫȞ ȤĮȡĮțIJȘȡȚıIJȚțȫȞ IJȠȣ B. trispora
ıIJȘ µȑȖȚıIJȘ ıȣȖțȑȞIJȡȦıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ țȦȞȚțȑȢ ijȚȐȜİȢ
ȤȡȘıȚµȠʌȠȚȫȞIJĮȢ ȣʌȠıIJȡȫµĮIJĮ µİ įȚĮijȠȡİIJȚțȒ ıȪıIJĮıȘ (µȠȞȐįĮ µȒțȠȣȢ 50
µm). a, b: IJȠ ȕĮıȚțȩ ȣʌȩıIJȡȦµĮ ݵʌȜȠȣIJȓıIJȘțİ µİ 3,0% țĮȚ 5,0% ȖȜȣțȩȗȘ,
ĮȞIJȓıIJȠȚȤĮ. c: ȉȠ ȕĮıȚțȩ ȣʌȩıIJȡȦµĮ (ȤȦȡȓȢ Span 20 țĮȚ Tween 80)
ݵʌȜȠȣIJȓıIJȘțİ µİ 5,0% ȖȜȣțȩȗȘ țĮȚ 8,0% İțȤȪȜȚıµĮ țĮȜĮµʌȠțȚȠȪ. ȉȠ ȕĮıȚțȩ
ȣʌȩıIJȡȦµĮ ʌȠȣ ʌİȡȚİȓȤİ 5,0% ȖȜȣțȩȗȘ țĮȚ 8,0% İțȤȪȜȚıµĮ țĮȜĮµʌȠțȚȠȪ
ݵʌȜȠȣIJȓıIJȘțİ µİ 0,25% ĮȞIJȚȠȟİȚįȦIJȚțȩ (d), 3,0% ȚıȠµȠȡȚĮțȩ µȓȖµĮ
İȜĮȚȩȜĮįȠȣ, ıȠȖȚȑȜĮȚȠȣ țĮȚ ȕĮµȕĮțȑȜĮȚȠȣ țĮȚ 0,25% ĮȞIJȚȠȟİȚįȦIJȚțȩ (e),
2,0% ȜȚȞİȜĮȧțȩ ȠȟȪ țĮȚ 0,25% ĮȞIJȚȠȟİȚįȦIJȚțȩ (f), 4,0% țȘȡȠȗȓȞȘ (g) țĮȚ 2,0%
ȜȚȞİȜĮȓțȩ ȠȟȪ, 4,0% țȘȡȠȗȓȞȘ țĮȚ 1,0% ĮȞIJȚȠȟİȚįȦIJȚțȩ (h). (ǺĮıȚțȩ
ȣʌȩıIJȡȦµĮ ȩʌȦȢ ʌİȡȚȖȡȐijİIJĮȚ ıIJȠȞ ȆȓȞĮțĮ 4.2.11)...........................................
170
ǼȚțȩȞĮ 4.3.1 ǹʌİȚțȩȞȚıȘ IJȦȞ µȠȡijȠȜȠȖȚțȫȞ ȤĮȡĮțIJȘȡȚıIJȚțȫȞ IJȠȣ B. trispora
ıIJȘ µȑȖȚıIJȘ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ ȗȣµȦIJȒȡĮ
ĮȞȐįİȣıȘȢ (µȠȞȐįĮ µȒțȠȣȢ 50 µm). a: b, c, d: ȊʌȩıIJȡȦµĮ ʌĮȡĮȖȦȖȒȢ 1, 2, 3
țĮȚ 4, ĮȞIJȓıIJȠȚȤĮ (ȩʌȦȢ ʌİȡȚȖȡȐijȠȞIJĮȚ ıIJȠȞ ȆȓȞĮțĮ
4.3.1)......................................................................................................................

215
XXI
ǼȚțȩȞĮ 4.3.2 ǹʌİȚțȩȞȚıȘ IJȦȞ µȣțȘȜȚĮțȫȞ ȣijȫȞ µİ țİȞȠIJȩʌȚĮ, țİȞȫȞ
țȣIJIJȐȡȦȞ țĮȚ İțijȣȜȚıµȑȞȦȞ țȣIJIJȐȡȦȞ IJȠȣ B. trispora ʌȠȣ ĮȞĮʌIJȪııİIJĮȚ ıIJȠ
ȣʌȩıIJȡȦµĮ ʌĮȡĮȖȦȖȒȢ 6 µİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ
IJȘȞ 4
Ș
ȘµȑȡĮ IJȘȢ ȗȪµȦıȘȢ. (ȊʌȩıIJȡȦµĮ ʌĮȡĮȖȦȖȒȢ 6 ȩʌȦȢ ʌİȡȚȖȡȐijİIJĮȚ
ıIJȠȞ ȆȓȞĮțĮ 4.3.1, µȠȞȐįĮ µȒțȠȣȢ 50 µm)..........................................................
216
ǼȚțȩȞĮ 4.3.3 ǹʌİȚțȩȞȚıȘ IJȦȞ µȠȡijȠȜȠȖȚțȫȞ ȤĮȡĮțIJȘȡȚıIJȚțȫȞ IJȠȣ B. trispora
ıIJȘ µȑȖȚıIJȘ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ ȗȣµȦIJȒȡĮ
ĮȞȐįİȣıȘȢ (µȠȞȐįĮ µȒțȠȣȢ 50 µm). a: ȂȣțȘȜȚĮțȑȢ ȣijȑȢ țĮȚ ȗȣȖȠıʌȩȡȚĮ IJȠȣ
B. trispora ʌȠȣ ĮȞĮʌIJȪııİIJĮȚ ıIJȠ ȣʌȩıIJȡȦµĮ ʌĮȡĮȖȦȖȒȢ 7. b: ȂȣțȘȜȚĮțȑȢ
ȣijȑȢ µİ țİȞȠIJȩʌȚĮ, țİȞȐ țȪIJIJĮȡĮ țĮȚ İțijȣȜȚıµȑȞĮ țȪIJIJĮȡĮ IJȠȣ B. trispora
ʌȠȣ ĮȞĮʌIJȪııİIJĮȚ ıIJĮ ȣʌȠıIJȡȫµĮIJĮ 8, 9, 10, 11 țĮȚ 12. (ȊʌȠıIJȡȫµĮIJĮ 7, 8,
9, 10, 11 țĮȚ 12 ȩʌȦȢ ʌİȡȚȖȡȐijȠȞIJĮȚ ıIJȠȞ ȆȓȞĮțĮ
4.3.1)......................................................................................................................
217
ǼȚțȩȞĮ 4.3.4 ǹʌİȚțȩȞȚıȘ İțijȣȜȚıµȑȞȦȞ țȣIJIJȐȡȦȞ (Į) țĮȚ µȣțȘȜȚĮțȫȞ ȣijȫȞ
µİ țİȞȠIJȩʌȚĮ, țİȞȫȞ țȣIJIJȐȡȦȞ țĮȚ İțijȣȜȚıµȑȞȦȞ țȣIJIJȐȡȦȞ (b) IJȠȣ
B. trispora ʌȠȣ ĮȞĮʌIJȪııİIJĮȚ ıIJĮ ȣʌȠıIJȡȫµĮIJĮ 14 (4
Ș
ȘµȑȡĮ ȗȪµȦıȘȢ) țĮȚ
15 (16
Ș
ȘµȑȡĮ ȗȪµȦıȘȢ), ĮȞIJȓıIJȠȚȤĮ, µİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ ȗȣµȦIJȒȡĮ
ĮȞȐįİȣıȘȢ. (ȊʌȠıIJȡȫµĮIJĮ 14 țĮȚ 15 ȩʌȦȢ ʌİȡȚȖȡȐijȠȞIJĮȚ ıIJȠȞ ȆȓȞĮțĮ 4.3.1,
µȠȞȐįĮ µȒțȠȣȢ 50 µm).........................................................................................
218
ǼȚțȩȞĮ 4.3.5 ǹʌİȚțȩȞȚıȘ IJȦȞ µȠȡijȠȜȠȖȚțȫȞ ȤĮȡĮțIJȘȡȚıIJȚțȫȞ IJȠȣ B. trispora
țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ ȗȣµȦIJȒȡĮ
ĮȞȐįİȣıȘȢ (µȠȞȐįĮ µȒțȠȣȢ 50 µm). a, b: ǽȣȖȠijȩȡȠȚ (2
Ș
ȘµȑȡĮ ȗȪµȦıȘȢ) țĮȚ
ȗȣȖȠıʌȩȡȚĮ (20
Ș
ȘµȑȡĮ ȗȪµȦıȘȢ), ĮȞIJȓıIJȠȚȤĮ, IJȠȣ B. trispora ʌȠȣ
ĮȞĮʌIJȪııİIJĮȚ ıIJȠ ȣʌȩıIJȡȦµĮ ʌĮȡĮȖȦȖȒȢ 20. c, d: ȂȣțȘȜȚĮțȑȢ ȣijȑȢ țĮȚ
ȗȣȖȠıʌȩȡȚĮ IJȠȣ B. trispora ʌȠȣ ĮȞĮʌIJȪııİIJĮȚ ıIJĮ ȣʌȠıIJȡȫµĮIJĮ 17, 23,
ĮȞIJȓıIJȠȚȤĮ, IJȘȞ 20
Ș
ȘµȑȡĮ IJȘȢ ȗȪµȦıȘȢ. e, f: ȂȣțȘȜȚĮțȑȢ ȣijȑȢ µİ țİȞȠIJȩʌȚĮ,
țİȞȐ țȪIJIJĮȡĮ, İțijȣȜȚıµȑȞĮ țȪIJIJĮȡĮ țĮȚ ȗȣȖȠıʌȩȡȚĮ IJȠȣ B. trispora ʌȠȣ
ĮȞĮʌIJȪııİIJĮȚ ıIJĮ ȣʌȠıIJȡȫµĮIJĮ 22 țĮȚ 24 ĮȞIJȓıIJȠȚȤĮ, IJȘȞ 20
Ș
ȘµȑȡĮ IJȘȢ
ȗȪµȦıȘȢ. (ȊʌȠıIJȡȫµĮIJĮ 17, 20, 22, 23 țĮȚ 24 ȩʌȦȢ ʌİȡȚȖȡȐijȠȞIJĮȚ ıIJȠȞ
ȆȓȞĮțĮ 4.3.1).........................................................................................................
219


XXII
ǼȚțȩȞĮ 4.3.6 ǹʌİȚțȩȞȚıȘ IJȦȞ IJȦȞ ȗȣȖȠıʌȠȡȓȦȞ, IJȦȞ țİȞȫȞ țȣIJIJȐȡȦȞ țĮȚ IJȦȞ
İțijȣȜȚıµȑȞȦȞ țȣIJIJȐȡȦȞ IJȠȣ B. trispora ʌȠȣ ĮȞĮʌIJȪııİIJĮȚ ıIJĮ ȣʌȠıIJȡȫµĮIJĮ
ʌĮȡĮȖȦȖȒȢ 2, 3, 7, 13 țĮȚ 25 ıIJȠ IJȑȜȠȢ IJȘȢ ȗȪµȦıȘȢ. (ȊʌȠıIJȡȫµĮIJĮ 2, 3, 7,
13 țĮȚ 25 ȩʌȦȢ ʌİȡȚȖȡȐijȠȞIJĮȚ ıIJȠȞ ȆȓȞĮțĮ
4.3.1)......................................................................................................................
220
ǼȚțȩȞĮ 4.3.7 ǹʌİȚțȩȞȚıȘ IJȦȞ µȠȡijȠȜȠȖȚțȫȞ ȤĮȡĮțIJȘȡȚıIJȚțȫȞ IJȠȣ B. trispora
ıIJȘ µȑȖȚıIJȘ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ µİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ ıİ ȗȣµȦIJȒȡĮ
ĮȞȐįİȣıȘȢ µİ įȚĮijȠȡİIJȚțȑȢ ıȣȞșȒțİȢ ĮİȡȚıµȠȪ țĮȚ ĮȞȐįİȣıȘȢ (µȠȞȐįĮ
µȒțȠȣȢ 50 µm). a: 1,0 vvm-150 rpm, b: 1,0 vvm-500 rpm, c: 2,0 vvm-150 rpm,
d: 2,0 vvm-500 rpm, e: 1,0 vvm-700 rpm, f: 2,0 rpm-700 rpm țĮȚ g: 0,5 vvm-
150 rpm.................................................................................................................
235
ǼȚțȩȞĮ 4.3.8 ǹʌİȚțȩȞȚıȘ IJȦȞ ȐșȚțIJȦȞ µȣțȘȜȚĮțȫȞ ȣijȫȞ, IJȦȞ µȣțȘȜȚĮțȫȞ
ȣijȫȞ µİ țİȞȠIJȩʌȚĮ, IJȦȞ țİȞȫȞ țȣIJIJȐȡȦȞ țĮȚ IJȦȞ İțijȣȜȚıµȑȞȦȞ țȣIJIJȐȡȦȞ
IJȠȣ B. trispora ıIJȘ µȑȖȚıIJȘ ıȣȖțȑȞIJȡȦıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ ĮıȣȞİȤȒ
ȗȪµȦıȘ ȕȣșȠȪ ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ. a: 0,5 vvm- 0 rpm Ȓ 0 vvm-150 rpm,
b: 2,0 vvm-0 rpm, c: 0 vvm-700 rpm (µȠȞȐįĮ µȒțȠȣȢ 50
µm)........................................................................................................................
236
ǼȚțȩȞĮ 4.4.1 ǹʌİȚțȩȞȚıȘ IJȦȞ µȠȡijȠȜȠȖȚțȫȞ ȤĮȡĮțIJȘȡȚıIJȚțȫȞ IJȠȣ B. trispora
țĮIJȐ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ µİ ıȣȞİȤȒ
ʌȡȠıșȒțȘ ȣʌȠıIJȡȫµĮIJȠȢ ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ (µȠȞȐįĮ µȒțȠȣȢ 50 µm). a:
ȂȣțȘȜȚĮțȑȢ ȣijȑȢ µİ țİȞȠIJȩʌȚĮ, țİȞȐ țȪIJIJĮȡĮ țĮȚ İțijȣȜȚıµȑȞĮ țȪIJIJĮȡĮ IJȠȣ
B. trispora (12
Ș
ȘµȑȡĮ ȗȪµȦıȘȢ) ʌȠȣ ĮȞĮʌIJȪııİIJĮȚ µİ ıȣȞİȤȒ ʌȡȠıșȒțȘ IJȠȣ
ȣʌȠıIJȡȫµĮIJȠȢ IJȡȠijȠįȠıȓĮȢ 1 µİ ȡȣșµȩ 3,5 ml/h Įʌȩ IJȘȞ ĮȡȤȒ IJȘȢ ȗȪµȦıȘȢ
(ȊʌȩıIJȡȦµĮ ĮȞȐʌIJȣȟȘȢ 1). b, c: ȂȣțȘȜȚĮțȑȢ ȣijȑȢ țĮȚ ȗȣȖȠıʌȩȡȚĮ IJȠȣ
B. trispora (4
Ș
ȘµȑȡĮ ȗȪµȦıȘȢ) ʌȠȣ ĮȞĮʌIJȪııİIJĮȚ µİ ıȣȞİȤȒ ʌȡȠıșȒțȘ IJȠȣ
ȣʌȠıIJȡȫµĮIJȠȢ IJȡȠijȠįȠıȓĮȢ 2 µİ ȡȣșµȩ 3,5 ml/h țĮȚ IJȠȣ ȣʌȠıIJȡȫµĮIJȠȢ
IJȡȠijȠįȠıȓĮȢ 3 µİ ȡȣșµȩ 4,2 ml/h, ĮȞIJȓıIJȠȚȤĮ, Įʌȩ IJȘȞ ĮȡȤȒ IJȘȢ ȗȪµȦıȘȢ
(ȊʌȩıIJȡȦµĮ ĮȞȐʌIJȣȟȘȢ 1 țĮȚ 2, ĮȞIJȓıIJȠȚȤĮ). d: ȂȣțȘȜȚĮțȑȢ ȣijȑȢ µİ
țİȞȠIJȩʌȚĮ, țİȞȐ țȪIJIJĮȡĮ țĮȚ İțijȣȜȚıµȑȞĮ țȪIJIJĮȡĮ IJȠȣ B. trispora (4
Ș
ȘµȑȡĮ
ȗȪµȦıȘȢ) ʌȠȣ ĮȞĮʌIJȪııİIJĮȚ µİ ıȣȞİȤȒ ʌȡȠıșȒțȘ IJȠȣ ȣʌȠıIJȡȫµĮIJȠȢ
IJȡȠijȠįȠıȓĮȢ 4 µİ ȡȣșµȩ 4,2 ml/h Įʌȩ IJȘȞ ĮȡȤȒ IJȘȢ ȗȪµȦıȘȢ (ȊʌȩıIJȡȦµĮ
ĮȞȐʌIJȣȟȘȢ 2). e: ȂȣțȘȜȚĮțȑȢ ȣijȑȢ țĮȚ ȗȣȖȠıʌȩȡȚĮ IJȠȣ B. trispora (10
Ș
ȘµȑȡĮ
ȗȪµȦıȘȢ) ʌȠȣ ĮȞĮʌIJȪııİIJĮȚ µİ ıȣȞİȤȒ ʌȡȠıșȒțȘ

XXIII
IJȠȣ ȣʌȠıIJȡȫµĮIJȠȢ IJȡȠijȠįȠıȓĮȢ 5 Ȓ 6 µİ ȡȣșµȩ 4,2 ml/h Įʌȩ IJȘȞ ĮȡȤȒ IJȘȢ
ȗȪµȦıȘȢ (ȊʌȩıIJȡȦµĮ ĮȞȐʌIJȣȟȘȢ 2). f: ȂȣțȘȜȚĮțȑȢ ȣijȑȢ µİ țİȞȠIJȩʌȚĮ, țİȞȐ
țȪIJIJĮȡĮ țĮȚ İțijȣȜȚıµȑȞĮ țȪIJIJĮȡĮ IJȠȣ B. trispora (20
Ș
ȘµȑȡĮ ȗȪµȦıȘȢ) ʌȠȣ
ĮȞĮʌIJȪııİIJĮȚ µİ IJȘȞ ʌȡȠıșȒțȘ 667 ml ȣʌȠıIJȡȫµĮIJȠȢ IJȡȠijȠįȠıȓĮȢ 3 IJȘȞ 2Ș,
8Ș țĮȚ 14Ș ȘµȑȡĮ IJȘȢ ȗȪµȦıȘȢ (ȊʌȩıIJȡȦµĮ ĮȞȐʌIJȣȟȘȢ 2). (ȊʌȠıIJȡȫµĮIJĮ
ĮȞȐʌIJȣȟȘȢ țĮȚ IJȡȠijȠįȠıȓĮȢ ȩʌȦȢ ʌİȡȚȖȡȐijȠȞIJĮȚ ıIJȠȞ ȆȓȞĮțĮ 4.4.1).
...............................................................................................................................

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XXIV
ȀǹȉǹȁȅīȅȈ ǻǾȂȅȈǿǼȊȂǼȃȍȃ ǼȇīǹȈǿȍȃ

1. Roukas, T., Mantzouridou, F. (2001) An improved method for extraction of
ȕ-carotene from Blakeslea trispora. Applied Biochemistry and Biotechnology
90: 37-45.

2. Mantzouridou, F., Roukas, T., Kotzekidou, P. (2002) Effect of the aeration
rate and agitation speed on ȕ-carotene production and morphology of
Blakeslea trispora in a stirred tank reactor: mathematical modeling.
Biochemical Engineering Journal 10: 123-135.

3. Mantzouridou, F., Roukas, T., Kotzekidou, P. (2002) Optimization of
ȕ-carotene production from synthetic medium by Blakeslea trispora in a
stirred tank reactor and relationship between morphological changes and
pigment formation. Food Biotechnology 16: 167-187.

4. Mantzouridou, F., Roukas, T., Kotzekidou, P., Liakopoulou, M. (2002)
Optimization of ȕ-carotene production from synthetic medium by Blakeslea
trispora: A mathematical modeling. Applied Biochemistry and Biotechnology
101: 153-175.



1

ȀǼĭǹȁǹǿȅ 1
ǼǿȈǹīȍīǾ

ȉȠ ȕ-țĮȡȠIJȑȞȚȠ Ȓ ʌȡȠȕȚIJĮµȓȞȘ ǹ İȓȞĮȚ µȓĮ ȜȚʌȠįȚĮȜȣIJȒ ȤȡȦıIJȚțȒ (țȓIJȡȚȞȠ
ȦȢ ʌȠȡIJȠțĮȜȓ ȤȡȫµĮ) ʌȠȜȪ įȚĮįİįȠµȑȞȘ ıIJȘ ijȪıȘ. ǺȡȓıțİIJĮȚ țȣȡȓȦȢ ıIJȠȣȢ
ijȣIJȚțȠȪȢ țĮȚ ȗȦȧțȠȪȢ ȚıIJȠȪȢ țĮȚ ȖȚĮ ʌȡȫIJȘ ijȠȡȐ ĮʌȠµȠȞȫșȘțİ Įʌȩ IJȠ țĮȡȩIJȠ
(Daucus carota) ıİ țȡȣıIJĮȜȜȚțȒ µȠȡijȒ (Wackenroder 1831). ǹʌĮȞIJȐIJĮȚ ıİ
µİȖȐȜȠ ĮȡȚșµȩ ijȡȑıțȦȞ ijȡȠȪIJȦȞ țĮȚ ȜĮȤĮȞȚțȫȞ (țĮȡȩIJĮ, ȖȜȣțȠʌĮIJȐIJİȢ,
ȡȠįȐțȚȞĮ, IJȠµȐIJİȢ, ț.Ȝ.ʌ.) µİ IJȘ µȠȡijȒ trans-ȚıȠµİȡȠȪȢ, İȞȫ įİȞ ıȣȞIJȓșİIJĮȚ ıIJĮ
ȗȫĮ. ǹʌȩ IJĮ ijȣIJȚțȐ µȑȡȘ IJȘȢ IJȡȠijȒȢ IJȦȞ ȗȫȦȞ µİIJĮijȑȡİIJĮȚ ıIJĮ ȜȚʌȓįȚĮ IJȠȣ
ĮȓµĮIJȠȢ, IJȠȣ ȖȐȜĮțIJȠȢ țĮȚ IJȦȞ ĮȣȖȫȞ. Ǿ ʌİȡȚİțIJȚțȩIJȘIJĮ ȠȡȚıµȑȞȦȞ IJȡȠijȓµȦȞ
ijȣIJȚțȒȢ ʌȡȠȑȜİȣıȘȢ ıİ ȕ-țĮȡȠIJȑȞȚȠ ĮȞĮijȑȡİIJĮȚ ıIJȠȞ ȆȓȞĮțĮ 1.1.
Ǿ ȤȘµȚțȒ įȠµȒ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ ȑȖȚȞİ ȖȞȦıIJȒ Įʌȩ IJȠȞ Karrer IJȠ 1930
(Malmström țĮȚ Anderson 2001). ǼȓȞĮȚ ȑȞĮȢ ʌȠȜȣĮțȩȡİıIJȠȢ ȣįȡȠȖȠȞȐȞșȡĮțĮȢ
(ʌȠȜȣȑȞȚȠ) ʌȠȣ ʌİȡȚȑȤİȚ 40 ȐIJȠµĮ ȐȞșȡĮțĮ țĮȚ ĮʌȠIJİȜİȓIJĮȚ Įʌȩ µȓĮ ʌȠȜȣİȞȚțȒ
ĮȜȣıȓįĮ µİ İȞȞȚȐ trans-(E-) įȚʌȜȠȪȢ įİıµȠȪȢ, ıIJĮ ȐțȡĮ IJȘȢ ȠʌȠȓĮȢ ıȣȞįȑȠȞIJĮȚ įȪȠ
įĮțIJȪȜȚȠȚ ȕ-ȚȠȞȩȞȘȢ (Mathews țĮȚ Van Holde 1996).


2

ȆȓȞĮțĮȢ 1.1 ȆİȡȚİțIJȚțȩIJȘIJĮ ȠȡȚıµȑȞȦȞ ijȡȠȪIJȦȞ țĮȚ ȜĮȤĮȞȚțȫȞ ıİ ȕ-țĮȡȠIJȑȞȚȠ
(Mangels țĮȚ ıȣȞ. 1993).


ǼȓįȠȢ ȉȡȠijȓµȠȣ
Į


ȕ-țĮȡȠIJȑȞȚȠ
(mg /100 g IJȡȠijȓµȠȣ)


ǼȓįȠȢ ȉȡȠijȓµȠȣ
Į

ȕ-țĮȡȠIJȑȞȚȠ
(mg /100 g IJȡȠijȓµȠȣ)


ȀĮȡȩIJȠ

6,6

ȂȐȖțȠ

2,9
īȜȣțȠʌĮIJȐIJĮ 5,9 ȆİʌȩȞȚ 2,0
ȁȐȤĮȞȠ 5,3 ǺİȡȓțȠțȠ 1,6
ȂĮȧȞIJĮȞȩȢ 5,1 ȀİȡȐıȚ 0,6
ȈʌĮȞȐțȚ 4,9 ȇȠįȐțȚȞȠ 0,5
ȆȡȐıȠ 3,5 ȉȠµȐIJĮ 0,5
ȇĮįȓțȚ 2,0 ȀĮȡʌȠȪȗȚ 0,4
ȂʌȡȩțȠȜȠ 1,5 ǻĮµȐıțȘȞȠ 0,2
ȈʌĮȡȐȖȖȚ 0,5 ȂȒȜȠ 0,1
ȆȐʌȡȚțĮ 0,5 ȆȠȡIJȠțȐȜȚ 0,1
Į
Ǿ ʌİȡȚİțIJȚțȩIJȘIJĮ IJȦȞ ijȡȠȪIJȦȞ țĮȚ ȜĮȤĮȞȚțȫȞ ıİ ȕ-țĮȡȠIJȑȞȚȠ İȟĮȡIJȐIJĮȚ Įʌȩ IJȘȞ
ʌȠȚțȚȜȓĮ țĮȚ IJȠ ȕĮșµȩ ȦȡȓµĮȞıȘȢ IJȠȣȢ.

ȉȠ ȕ-țĮȡȠIJȑȞȚȠ, ȜȩȖȦ IJȘȢ ȕȚIJĮµȚȞȚțȒȢ IJȠȣ įȡȐıȘȢ, ȤȡȘıȚµȠʌȠȚİȓIJĮȚ ȦȢ
ıȣµʌȜȒȡȦµĮ IJȘȢ ĮȞșȡȫʌȚȞȘȢ įȚĮIJȡȠijȒȢ țĮȚ ȦȢ ʌȡȩıșİIJȠ ıIJȚȢ ȗȦȠIJȡȠijȑȢ.
ǼʌȚʌȜȑȠȞ, ȕȡȓıțİȚ İijĮȡµȠȖȒ ȦȢ ȤȡȦıIJȚțȒ ȖȚĮ IJȘȞ İȞȓıȤȣıȘ IJȠȣ ȤȡȫµĮIJȠȢ
IJȡȠijȓµȦȞ țĮȚ ʌȠIJȫȞ (ȕȠȪIJȣȡȠ, µĮȡȖĮȡȓȞȘ, µĮȖİȚȡȚțȐ ȜȓʌȘ, ĮȞĮȥȣțIJȚțȐ ț.Ȝ.ʌ.), ȦȢ
ĮȞIJȚȠȟİȚįȦIJȚțȩ ıIJĮ țȠȞıİȡȕȠʌȠȚȘµȑȞĮ IJȡȩijȚµĮ (Ciegler țĮȚ ıȣȞ. 1963a, Boskou
1989, Hui 1992, Kim țĮȚ ıȣȞ. 1997a) țĮȚ ȦȢ țȪȡȚȠ ıȣıIJĮIJȚțȩ ıIJȘȞ ʌĮȡĮıțİȣȒ
țĮȜȜȣȞIJȚțȫȞ (Leung țĮȚ Foster 1996). ȉĮ IJİȜİȣIJĮȓĮ ȤȡȩȞȚĮ ȤȡȘıȚµȠʌȠȚİȓIJĮȚ țĮȚ
ıIJȘ ijĮȡµĮțİȣIJȚțȒ ȕȚȠµȘȤĮȞȓĮ. ȈIJȚȢ ĮȡȤȑȢ IJȘȢ įİțĮİIJȓĮȢ IJȠȣ `80 įȚĮʌȚıIJȫșȘțİ Ș
įȡȐıȘ IJȠȣ ıĮȞ ȕȚȠȜȠȖȚțȩ ĮȞIJȚȠȟİȚįȦIJȚțȩ µȑıȦ IJȠȣ µȘȤĮȞȚıµȠȪ IJȦȞ İȜİȣșȑȡȦȞ
ȡȚȗȫȞ (Burton țĮȚ Ingold 1984, Di Mascio țĮȚ ıȣȞ. 1990, Packer 1992, Edge țĮȚ
ıȣȞ. 1997). ǻȚȐijȠȡİȢ ȑȡİȣȞİȢ ȑįİȚȟĮȞ ȩIJȚ µʌȠȡİȓ ȞĮ µİȚȫıİȚ IJȠȞ țȓȞįȣȞȠ ݵijȐȞȚıȘȢ
țĮȡțȓȞȠȣ, țĮȡįȚĮțȫȞ ʌĮșȒıİȦȞ, țĮIJĮȡȡȐțIJȘ, ĮȡIJȘȡȚȠıțȜȒȡȦıȘȢ țĮȚ ȐȜȜȦȞ
ıȠȕĮȡȫȞ ʌĮșȒıİȦȞ (Byers țĮȚ Perry 1992, Lepage țĮȚ ıȣȞ. 1996, Burri 1997,
Omaye țĮȚ ıȣȞ. 1997). ȉȑȜȠȢ, İȞȚıȤȪİȚ IJȘ ȜİȚIJȠȣȡȖȓĮ IJȠȣ ĮȞȠıȠʌȠȚȘIJȚțȠȪ
ıȣıIJȒµĮIJȠȢ țĮȚ țĮșȣıIJİȡİȓ IJȘ įȚĮįȚțĮıȓĮ ȖȒȡĮȞıȘȢ IJȦȞ țȣIJIJȐȡȦȞ (Chew 1993,
Snodderly 1995).

3
ȉȠ ȕ-țĮȡȠIJȑȞȚȠ ʌĮȡĮȜĮµȕȐȞİIJĮȚ µİ İțȤȪȜȚıȘ Įʌȩ ijȣıȚțȐ ʌȡȠȧȩȞIJĮ, İȞȫ
ʌĮȡȐȖİIJĮȚ IJȩıȠ µİ ȤȘµȚțȑȢ µİșȩįȠȣȢ ȩıȠ țĮȚ µİ IJȘ ȤȡȘıȚµȠʌȠȓȘıȘ
µȚțȡȠȠȡȖĮȞȚıµȫȞ. ȉȠ ıȣȞșİIJȚțȩ ȕ-țĮȡȠIJȑȞȚȠ ʌĮȡĮıțİȣȐıIJȘțİ ȖȚĮ ʌȡȫIJȘ ijȠȡȐ IJo
1950 Įʌȩ IJȠȣȢ Karrer țĮȚ Milas, İȞȫ Įʌȩ IJȠ 1954 ȐȡȤȚıİ Ș ȕȚȠµȘȤĮȞȚțȒ ʌĮȡĮȖȦȖȒ
IJȘȢ ȤȡȦıIJȚțȒȢ țĮȚ Ș ݵʌȠȡȚțȒ IJȘȢ İțµİIJȐȜȜİȣıȘ (Hui 1992). ȈIJĮ IJȑȜȘ IJȠȣ 1970 Ș
İIJȒıȚĮ ʌĮȖțȩıµȚĮ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ ĮȞİȡȤȩIJĮȞ ʌİȡȓʌȠȣ ıIJȠȣȢ 27 IJȩȞȠȣȢ,
İȞȫ ıȒµİȡĮ IJȠ ȪȥȠȢ ʌĮȡĮȖȦȖȒȢ ȟİʌİȡȞȐİȚ IJȠȣȢ 100 IJȩȞȠȣȢ İIJȘıȓȦȢ țĮȚ µȩȞȠ IJȠ
10,0% ʌĮȡĮȜĮµȕȐȞİIJĮȚ Įʌȩ ijȣıȚțȐ ʌȡȠȧȩȞIJĮ (Baets țĮȚ ıȣȞ. 2002). ȈȪµijȦȞĮ µİ
IJĮ ıIJȠȚȤİȓĮ IJȘȢ ǼșȞȚțȒȢ ȈIJĮIJȚıIJȚțȒȢ ȊʌȘȡİıȓĮȢ ȖȚĮ IJȠ 2002, ıIJȘȞ ǼȜȜȐįĮ ȠȚ
İȚıĮȖȦȖȑȢ țĮȡȠIJİȞȠİȚįȫȞ ʌȠȣ ȤȡȘıȚµȠʌȠȚȠȪȞIJĮȚ țȣȡȓȦȢ ȦȢ ȤȡȦıIJȚțȑȢ ıIJȘ
ȕȚȠµȘȤĮȞȓĮ IJȡȠijȓµȦȞ ĮȞȑȡȤȠȞIJĮȚ ıİ 1 IJȩȞȠ ʌİȡȓʌȠȣ.
ȉĮ IJİȜİȣIJĮȓĮ ȤȡȩȞȚĮ, ȜȩȖȦ IJȘȢ ĮȣȟĮȞȩµİȞȘȢ ȗȒIJȘıȘȢ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ țĮȚ
IJȠȣ ȣȥȘȜȠȪ țȩıIJȠȣȢ IJȦȞ ȣȜȚțȫȞ ʌȠȣ ȤȡȘıȚµȠʌȠȚȠȪȞIJĮȚ ȖȚĮ IJȘ ȤȘµȚțȒ IJȠȣ ıȪȞșİıȘ,
Ș ʌĮȡĮȖȦȖȒ IJȠȣ µİ IJȘ ȤȡȒıȘ µȚțȡȠȠȡȖĮȞȚıµȫȞ ĮʌȠIJİȜİȓ ȕĮıȚțȩ ĮȞIJȚțİȓµİȞȠ
ȑȡİȣȞĮȢ ıİ ʌĮȖțȩıµȚȠ İʌȓʌİįȠ. ȍıIJȩıȠ, Ș ȕȚȠIJİȤȞȠȜȠȖȚțȒ ʌĮȡĮȖȦȖȒ IJȘȢ
ȤȡȦıIJȚțȒȢ ıİ ȕȚȠµȘȤĮȞȚțȒ țȜȓµĮțĮ ĮʌĮȚIJİȓ IJȘ ȤȡȒıȘ µȚțȡȠȠȡȖĮȞȚıµȫȞ µİ
ȚțĮȞȩIJȘIJĮ ʌĮȡĮȖȦȖȒȢ ȣȥȘȜȒȢ ıȣȖțȑȞIJȡȦıȘȢ ʌȡȠȧȩȞIJȠȢ, ȤĮµȘȜȠȪ țȩıIJȠȣȢ
ȣʌȠıIJȡȦµȐIJȦȞ țĮȚ ĮʌȠIJİȜİıµĮIJȚțȫȞ țĮȚ ȠȚțȠȞȠµȚțȫȞ IJİȤȞȚțȫȞ ĮȞȐțIJȘıȘȢ țĮȚ
țĮșĮȡȚıµȠȪ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ.
īȚĮ IJȘȞ ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ ıİ ʌİȚȡĮµĮIJȚțȒ țȜȓµĮțĮ
ȤȡȘıȚµȠʌȠȚȒșȘțĮȞ įȚȐijȠȡĮ ıIJİȜȑȤȘ ȕĮțIJȘȡȓȦȞ, ȗȣµȫȞ, µȣțȒIJȦȞ țĮșȫȢ țĮȚ ijȪțȘ
(Ciegler 1965, Cerda-Olmedo 1985). ȅȚ ʌİȡȚııȩIJİȡȠȚ Įʌȩ IJȠȣȢ µȚțȡȠȠȡȖĮȞȚıµȠȪȢ
ʌȠȣ µİȜİIJȒșȘțĮȞ ʌĮȡȒȖĮȞ ȤĮµȘȜȑȢ ıȣȖțİȞIJȡȫıİȚȢ IJȘȢ ȤȡȦıIJȚțȒȢ (4,0-50,0 mg/l)
ȩIJĮȞ țĮȜȜȚİȡȖȠȪȞIJĮȞ IJȩıȠ ıİ ıȣȞșİIJȚțȐ ȣʌȠıIJȡȫµĮIJĮ ȩıȠ țĮȚ ıİ ȣʌȠʌȡȠȧȩȞIJĮ
ȖİȦȡȖȚțȫȞ ȕȚȠµȘȤĮȞȚȫȞ (Ciegler 1965). ȅȚ Barnett țĮȚ ıȣȞ. (1956) ʌĮȡĮIJȒȡȘıĮȞ
ȩIJȚ ȩIJĮȞ ıIJİȜȑȤȘ ĮȞIJȓșİIJȠȣ ijȪȜȠȣ IJȠȣ µȪțȘIJĮ Choanephora cucurbitarum
țĮȜȜȚİȡȖȠȪȞIJĮȞ µĮȗȓ ıİ ıȣȞșİIJȚțȩ ȣʌȩıIJȡȦµĮ Ș ʌĮȡĮȖȦȖȚțȩIJȘIJĮ ıİ ʌȡȠȕȚIJĮµȓȞȘ
ǹ ȒIJĮȞ ĮȣȟȘµȑȞȘ (15-20 ijȠȡȑȢ µİȖĮȜȪIJİȡȘ), ȑȞĮȞIJȚ IJȠȣ țȐșİ ıIJİȜȑȤȠȣȢ ȤȦȡȚıIJȐ.
ȅȚ Hesseltine țĮȚ Anderson (1957) țĮȚ ȠȚ Prieto țĮȚ ıȣȞ. (1964) ʌȠȣ µİȜȑIJȘıĮȞ IJȘȞ
ʌĮȡĮȖȦȖȚțȩIJȘIJĮ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ Įʌȩ ıIJİȜȑȤȘ IJȘȢ ȠȚțȠȖȑȞİȚĮȢ Choanephoraceae
ȩʌȦȢ Choanephora conjuncta, Blakeslea circinans țĮȚ Blakeslea trispora ȕȡȒțĮȞ
ȩIJȚ ıİ ȩȜİȢ IJȚȢ ʌİȡȚʌIJȫıİȚȢ ȠȚ țĮȜȜȚȑȡȖİȚİȢ țĮIJȐ IJȘȞ İȖȖİȞȒ ĮȞĮʌĮȡĮȖȦȖȒ ʌĮȡȒȖĮȞ
ʌİȡȚııȩIJİȡȠ ȕ-țĮȡȠIJȑȞȚȠ Įʌȩ țȐșİ ȑȞĮ ıIJȑȜİȤȠȢ ȤȦȡȚıIJȐ. Ǿ µİȖĮȜȪIJİȡȘ ʌȠıȩIJȘIJĮ

4
IJȘȢ ȤȡȦıIJȚțȒȢ (1000,0 mg/l) ʌĮȡȒȤșȘțİ µİ µȓȖµĮ IJȦȞ (+) țĮȚ (-) ıIJİȜİȤȫȞ IJȠȣ
µȪțȘIJĮ Blakeslea trispora (Atkinson țĮȚ Mavituna 1985).
ȈȒµİȡĮ, Ș ȕȚȠµȘȤĮȞȚțȒ ʌĮȡĮȖȦȖȒ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ ȖȓȞİIJĮȚ µİ IJȠ µȪțȘIJĮ
B. trispora ʌȠȣ ĮȞĮʌIJȪııİIJĮȚ ıİ ıȣȞșİIJȚțȩ ȣʌȩıIJȡȦµĮ ıİ ȗȣµȦIJȒȡİȢ ĮȞȐįİȣıȘȢ
țĮȚ µİ IJĮ ijȪțȘ Dunaliella saline țĮȚ Dunaliella bardawil, IJĮ ȠʌȠȓĮ țĮȜȜȚİȡȖȠȪȞIJĮȚ
ıİ ȜȓµȞİȢ µİ ĮȜµȣȡȩ Ȟİȡȩ (Ninet țĮȚ Renaut 1979, Borowitzka țĮȚ Borowitzka
1988, Cerda-Olmedo 1989, Benemann 1990, Vandamme 1992, Baets țĮȚ ıȣȞ.
2002). ȉȠ µİȖĮȜȪIJİȡȠ ʌȠıȠıIJȩ IJȠȣ ıȣȞȠȜȚțȠȪ ȕ-țĮȡȠIJȑȞȚȠȣ Įʌȩ IJȠ B. trispora
ĮʌĮȞIJȐIJĮȚ µİ IJȘ µȠȡijȒ all-trans-ȚıȠµİȡȠȪȢ (75,0-95,0%) (Anderson țĮȚ ıȣȞ. 1958,
Ciegler țĮȚ ıȣȞ. 1963b, Dholakia țĮȚ Modi 1984), ıİ ĮȞIJȓșİıȘ µİ IJĮ ijȪțȘ ʌȠȣ
ʌĮȡȐȖȠȣȞ ȚıȠµȠȡȚĮțȩ µȓȖµĮ all-trans- țĮȚ 9-cis-ȕ-țĮȡȠIJȑȞȚȠȣ (Ben-Amotz țĮȚ
Avron 1983, 1990, Ben-Amotz țĮȚ ıȣȞ. 1988a,b, 1989). ȈIJȘ įİȪIJİȡȘ ʌİȡȓʌIJȦıȘ, ĮȞ
țĮȚ Ș ıȣȞȠȜȚțȒ ʌȠıȩIJȘIJĮ IJȘȢ ʌĮȡĮȖȩµİȞȘȢ ʌȡȠȕȚIJĮµȓȞȘȢ ǹ İȓȞĮȚ µİȖȐȜȘ, Ș
ȕȚȠȜȠȖȚțȒ IJȘȢ įȡȐıȘ İȓȞĮȚ µİȚȦµȑȞȘ ȜȩȖȦ IJȠȣ 9-cis- ȚıȠµİȡȠȪȢ ʌȠȣ İȓȞĮȚ ȜȚȖȩIJİȡȠ
İȞİȡȖȩ (Zechmeister 1962, ȆĮȞȑȡĮȢ 1995). ǼʌȓıȘȢ, Ș İijĮȡµȠȖȒ IJȘȢ µİșȩįȠȣ
ʌĮȡĮȖȦȖȒȢ IJȘȢ ȤȡȦıIJȚțȒȢ Įʌȩ ijȪțȘ ʌİȡȚȠȡȓȗİIJĮȚ ıİ ȤȫȡİȢ ȩʌȠȣ İʌȚțȡĮIJȠȪȞ
țĮIJȐȜȜȘȜİȢ țȜȚµĮIJȠȜȠȖȚțȑȢ ıȣȞșȒțİȢ țĮȚ ȣʌȐȡȤȠȣȞ µİȖȐȜİȢ İțIJȐıİȚȢ ĮȜµȣȡȫȞ
ȜȚµȞȫȞ ȩʌȦȢ ıIJȘȞ ǹȣıIJĮȜȓĮ, ıIJȘ ȇȦıȓĮ, ıIJȠ ǿıȡĮȒȜ, ıIJȘȞ ǿıʌĮȞȓĮ, ıIJȘ ȋȚȜȒ țĮȚ
ıIJȘȞ ȀȓȞĮ (Shlomai țĮȚ ıȣȞ. 1991, Vandamme 1992, Baets țĮȚ ıȣȞ. 2002).
ȉĮ ȣʌȠıIJȡȫµĮIJĮ ʌȠȣ ȤȡȘıȚµȠʌȠȚȒșȘțĮȞ ȖȚĮ IJȘȞ ʌĮȡĮȖȦȖȒ IJȠȣ ȕ-
țĮȡȠIJȑȞȚȠȣ İȓȞĮȚ ıȣȞȒșȦȢ ıȣȞșİIJȚțȐ ȣʌȠıIJȡȫµĮIJĮ ʌȠȣ ʌİȡȚȜĮµȕȐȞȠȣȞ ȖȜȣțȩȗȘ,
ıĮțȤĮȡȩȗȘ, ȐµȣȜȠ (Ciegler țĮȚ ıȣȞ. 1959a, Costa țĮȚ ıȣȞ. 1987, Yun țĮȚ ıȣȞ.
1990), țİȜȜȠȕȚȩȗȘ (Dholakia țĮȚ Modi 1982), ȖȜȣțİȡȩȜȘ (Martelli țĮȚ ıȣȞ. 1992),
įȚĮijȠȡİIJȚțȐ İȓįȘ įȘµȘIJȡȚĮțȫȞ (ȕȡȫµȘ, ıȚIJȐȡȚ, țȡȚșȐȡȚ, țĮȜĮµʌȩțȚ, ȡȪȗȚ, ıȓțĮȜȘ
ț.Ȝ.ʌ) țĮȚ ʌȡȠȧȩȞIJĮ ıȩȖȚĮȢ (ıȠȖȚȐȜİȣȡȠ ț.Ȝ.ʌ) (Ciegler țĮȚ ıȣȞ. 1959a), ijȣIJȚțȐ
ȑȜĮȚĮ, ȜȓʌȘ, ĮțȩȡİıIJĮ ȜȚʌĮȡȐ ȠȟȑĮ, țȘȡȠȗȓȞȘ, İʌȚijĮȞİȚȠįȡĮıIJȚțȑȢ ȠȣıȓİȢ,
ĮȞIJȚȠȟİȚįȦIJȚțȐ, ʌȡȩįȡȠµİȢ ȠȣıȓİȢ țĮȚ İȞİȡȖȠʌȠȚȘIJȑȢ (Mackinney țĮȚ ıȣȞ. 1953,
Anderson țĮȚ ıȣȞ. 1958, Ciegler țĮȚ ıȣȞ. 1959b, 1961, 1962, 1963a,b, Ciegler
1965, Ninet țĮȚ ıȣȞ. 1969, Ninet țĮȚ Renaut 1979, Elsava țĮȚ ıȣȞ. 1974, Feofilova
țĮȚ Kudryavtseva 1974, Desai țĮȚ ıȣȞ. 1975, Feofilova țĮȚ Arbuzov 1975,
Feofilova țĮȚ Bekhtereva 1976, Galasko țĮȚ ıȣȞ. 1976, Desai țĮȚ Modi 1977, Rao
țĮȚ Modi 1977, Dandekar țĮȚ ıȣȞ. 1980, Yakovleva țĮȚ ıȣȞ. 1980, Mehta țĮȚ ıȣȞ.
1981, Feofilova țĮȚ ıȣȞ. 1982, Govind țĮȚ ıȣȞ. 1982, Dholakia țĮȚ Modi 1983,
1984, Atkinson țĮȚ Mavituna 1985, Cerda-Olmedo țĮȚ Hüttermann 1986, Kim țĮȚ

5
ıȣȞ. 1997a,b, Jeong țĮȚ ıȣȞ. 2001, Mantzouridou țĮȚ ıȣȞ. 2002a). ǼʌȓıȘȢ, ȑȤȠȣȞ
ȤȡȘıȚµȠʌȠȚȘșİȓ ʌİȚȡĮµĮIJȚțȐ țĮȚ ȣʌȠʌȡȠȧȩȞIJĮ įȚĮijȩȡȦȞ ȖİȦȡȖȚțȫȞ ȕȚȠµȘȤĮȞȚȫȞ,
ȩʌȦȢ IJȣȡȩȖĮȜĮ (Lampila țĮȚ ıȣȞ. 1985a,b, Fregova 1994), ıȣµʌȣțȞȦµȑȞȠ ȖȜİȪțȠȢ
ıIJĮijȣȜȚȫȞ (Buzzini țĮȚ Martini 1999, Buzzini 2000), İțȤȪȜȚıµĮ ıĮțȤȐȡȦȞ Įʌȩ
ıĮțȤĮȡȠțȐȜĮµȠ (Martelli țĮȚ ıȣȞ. 1990) țĮȚ µİȜȐııĮ (Bhosale țĮȚ Gadre 2001a,
Goksungur țĮȚ ıȣȞ. 2002).
Ȉİ ȩȜİȢ IJȚȢ ʌĮȡĮʌȐȞȦ µİȜȑIJİȢ Ș µȑȖȚıIJȘ ıȣȖțȑȞIJȡȦıȘ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ
ȒIJĮȞ ıȤİIJȚțȐ ȤĮµȘȜȒ (4,0-1000,0 mg/l). ǼʌȚʌȜȑȠȞ, ıIJȚȢ ʌİȡȚııȩIJİȡİȢ Įʌȩ IJȚȢ
ʌȡȠȘȖȠȪµİȞİȢ İȡİȣȞȘIJȚțȑȢ İȡȖĮıȓİȢ Ș ʌĮȡĮȖȦȖȒ ȕ-țĮȡȠIJȑȞȚȠȣ ȑȖȚȞİ ıİ
ĮȞĮțȚȞȠȪµİȞİȢ ijȚȐȜİȢ, İȞȫ İȜȐȤȚıIJİȢ µİȜȑIJİȢ ĮȞĮijȑȡȠȞIJĮȚ ıIJȘ ȤȡȒıȘ
ȕȚȠĮȞIJȚįȡĮıIJȒȡȦȞ IJȩıȠ ıİ İȡȖĮıIJȘȡȚĮțȩ, ȩıȠ țĮȚ ıİ ʌȚȜȠIJȚțȩ İʌȓʌİįȠ (Ciegler țĮȚ
ıȣȞ. 1963b, Ninet țĮȚ Renaut 1979, Costa țĮȚ ıȣȞ. 1987, Shlomai țĮȚ ıȣȞ. 1991,
Kim țĮȚ ıȣȞ. 1999, Bhosale țĮȚ Gadre 2001a, Mantzouridou țĮȚ ıȣȞ. 2002b,c).
ȆĮȡȩȜȠ ʌȠȣ Ș ĮʌȩįȠıȘ țĮȚ Ș ʌĮȡĮȖȦȖȚțȩIJȘIJĮ IJȦȞ įȚĮijȩȡȦȞ µİșȩįȦȞ
ȗȪµȦıȘȢ İȟĮȡIJȐIJĮȚ ıİ µİȖȐȜȠ ȕĮșµȩ Įʌȩ IJȘ µȠȡijȠȜȠȖȓĮ IJȘȢ țĮȜȜȚȑȡȖİȚĮȢ (Cox
țĮȚ ıȣȞ. 1998), įİȞ ȑȤİȚ ȖȓȞİȚ µȑȤȡȚ ıȒµİȡĮ İʌĮȡțȒȢ įȚİȡİȪȞȘıȘ IJȘȢ
ĮȜȜȘȜİʌȓįȡĮıȘȢ IJȦȞ µȠȡijȠȜȠȖȚțȫȞ ȤĮȡĮțIJȘȡȚıIJȚțȫȞ IJȦȞ µȚțȡȠȠȡȖĮȞȚıµȫȞ µİ
IJȘȞ ʌĮȡĮȖȦȖȒ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ.
ȈțȠʌȩȢ IJȘȢ İȡȖĮıȓĮȢ ȒIJĮȞ Ș ĮȪȟȘıȘ IJȘȢ ıȣȖțȑȞIJȡȦıȘȢ IJȠȣ ȕ-țĮȡȠIJȑȞȚȠȣ
Įʌȩ ıȣȞșİIJȚțȩ ȣʌȩıIJȡȦµĮ µİ IJȘ ȤȡȘıȚµȠʌȠȓȘıȘ IJȠȣ µȪțȘIJĮ B. trispora. īȚĮ IJȘȞ
ȣȜȠʌȠȓȘıȘ IJȠȣ ʌĮȡĮʌȐȞȦ ıțȠʌȠȪ ȑȖȚȞİ ȕİȜIJȚıIJȠʌȠȓȘıȘ IJȘȢ İțȤȪȜȚıȘȢ IJȠȣ
ȕ-țĮȡȠIJȑȞȚȠȣ Įʌȩ IJĮ țȪIJIJĮȡĮ IJȠȣ µȚțȡȠȠȡȖĮȞȚıµȠȪ, įȚİȡİȪȞȘıȘ IJȦȞ ʌĮȡĮȖȩȞIJȦȞ
ʌȠȣ İʌȘȡİȐȗȠȣȞ IJȘȞ ĮȞȐʌIJȣȟȘ IJȠȣ µȚțȡȠȠȡȖĮȞȚıµȠȪ țĮȚ IJȘȞ ʌĮȡĮȖȦȖȒ
ȕ-țĮȡȠIJȑȞȚȠȣ ıİ ĮȞĮțȚȞȠȪµİȞİȢ ijȚȐȜİȢ țĮȚ ıİ ȗȣµȦIJȒȡĮ ĮȞȐįİȣıȘȢ,
ȕİȜIJȚıIJȠʌȠȓȘıȘ IJȦȞ įȚİȡȖĮıȚȫȞ ʌĮȡĮȖȦȖȒȢ IJȘȢ ȤȡȦıIJȚțȒȢ ıIJĮ ʌĮȡĮʌȐȞȦ
ıȣıIJȒµĮIJĮ ȗȪµȦıȘȢ µİ ĮȞȐʌIJȣȟȘ µĮșȘµĮIJȚțȫȞ µȠȞIJȑȜȦȞ ıİ İȡȖĮıIJȘȡȚĮțȒ
țȜȓµĮțĮ, µİȜȑIJȘ IJȘȢ ʌĮȡĮȖȦȖȒȢ IJȘȢ ȤȡȦıIJȚțȒȢ ıİ ĮıȣȞİȤȒ ȗȪµȦıȘ ȕȣșȠȪ µİ