Лигниназы базидиомицетов тема диссертации и автореферата по ВАК РФ 03.00.04, доктор биологических наук Леонтьевский, Алексей Аркадьевич
Леонтьевский, Алексей Аркадьевич
доктор биологических наук
2002
- Специальность ВАК РФ03.00.04
- Количество страниц 267
Оглавление диссертации доктор биологических наук Леонтьевский, Алексей Аркадьевич
Список сокращений.
I. Введение.
1.1. Актуальность проблемы.
1.2. Состояние вопроса.
1.3. Цель и задачи исследования.
1.4. Научная новизна.
1.5. Практическая значимость.
1.6. Связь работы с крупными программами.
1.7. Апробация работы.
1.8. Публикации.
1.9. Благодарность.
II. Обзор литературы «Биодеградация лигнина грибами белой гнили»
II. 1. Лигнин: определение, функции, распространение.
II. 1.1. Общие сведения.
II. 1.2. Биосинтез лигнина.
II.1.3. Строение лигнина.
II. 1.4. Препараты лигнина.
11.2. Микробиология разложения лигнина.
11.2.1. Анаэробное разложение лигнина.
11.2.2. Разложение лигнина бактериями.
- Актиномицеты.
- Эубактерии.
- Туннелирующие бактерии.
11.2.3. Разложение лигнина грибами.
- Грибы «мягкой» гнили древесины.
- Грибы «бурой» гнили древесины.
- Грибы «белой» гнили древесины.
- Другие грибы.
11.2.4. Микробные сообщества.
11.3. Физиология разложения лигнина грибами белой гнили.
11.3.1. Субстратная адаптация грибов белой гнили.
11.3.2. Условия разложения лигнина грибами белой гнили.
- Лигнинолитическая активность как функция вторичного метаболизма.
- Индуцибельность лигнинолитической системы.
- Необходимость косубстрата.
- Необходимость аэробных условий.
11.4. Энзимология разложения лигнина грибами белой гнили.
11.4.1. Лигнинолитические ферментные комплексы.
11.4.2. Лигнин пероксидаза.
- Определение.
- История изучения и распространение.
- Строение лигнин пероксидазы.
- Каталитические свойства.
- Лигнинолитические свойства.
- Функции лигнин пероксидазы.
- Особенности экспрессии и идентификации.
11.4.3. Мп-пероксидаза.
- Определение.
- История изучения и распространение.
- Строение Мп-пероксидазы.
- Каталитические свойства.
- Лигнинолитические свойства.
- Функции Мп-пероксидазы.
- Особенности экспрессии и идентификации.
11.4.4. Другие пероксидазы.
- Mn-зависимая лигнин пероксидаза.
- Классическая пероксидаза
- «Новые» пероксидазы.
11.4.5. Лакказа.
- Определение.
- История изучения.
- Распространение.
- Локализация.
- Строение лакказы.
- Каталитические свойства.
- Лигнинолитические свойства.
- Функции лакказы.
- Особенности экспрессии и идентификации.
11.4.6. Перекись-генерирующие ферменты.
- Глюкозо-оксидаза.
- Метанол-оксидаза.
- Глиоксаль-оксидаза.
- Арил-алкоголь оксидаза.
11.4.7. Редуктазы.
- Арил-алкоголь дегидрогеназа.
- Хинон-редуктаза.
- Целлобиозо-дегидрогеназа.
11.5. Неэнзиматические реакции при биодеградации лигнина.
11.6. Прикладное значение грибов белой гнили.
11.6.1. Обработка природных лигнин-содержащих материалов.
11.6.2. Разложение ксенобиотиков.
III. Методы и материалы.
111.1. Микроорганизмы.
111.2. Методы культивирования микроорганизмов.
111.3. Методы определения активности ферментов.
111.4. Очистка ферментов.
111.5. Электрофорез и изоэлектрофокусировка.
111.6. Спектральные методы.
111.7. Высокоэффективная жидкостная хроматография.
111.8. Лигнинолитические реакции культур грибов и очищенных ферментов.
111.9. Биоремедиация почвы.
111.10. Идентификация продуктов разложения хлорфенолов. 117 III. 11. Аналитические методы.
III. 12. Материалы и реактивы.
IV. Результаты.
IV.1. Лигнинолитические свойства гриба Panus tigrinus 8/18 при разных способах культивирования.
IV.1.1. Скрининг лигнинолитических грибов.
IV. 1.2. Стационарная погруженная культура Panus tigrinus 8/18.
IV. 1.3. Твердофазная культура/*, tigrinus 8/18.
IV. 1.4. Определение ключевого лигнинолитического фермента.
IV.2. Новый способ погруженного культивирования Panus tigrinus 8/18.
IV.2.1. Индукция лигнинолитических ферментов.
IV.2.2. Подбор компонентов культуральной среды.
IV.2.3. Новые приемы культивирования.
IV.2.4. Варианты нового способа погруженного культивирования.
IV.3. Mn-зависимая лигнин пероксидаза Panus tigrinus 8/18.
IV.3.1. Физико-химические свойства очищенной МпЛП.
IV.3.2. Каталитический цикл.
IV.3.3. Лигнинолитические свойства.
IV.4. Лакказа Panus tigrinus 8/18 и феномен желтых лакказ.
IV.4.1. Общая характеристика.
IV.4.2. Спектральные свойства.
IV.4.3. Каталитические свойства.
IV.4.4. Множественные формы.
IV.4.5. Феномен желтых лакказ.
IV.4.6. Лигнинолитические свойства голубых и желтых лакказ.
IV.5. Прикладное значение лигнинолитических грибов и ферментов.
IV.5.1. Обработка лигнин-содержащих материалов.
IV.5.2. Разложение хлорфенолов грибами белой гнили.
V. Обсуждение результатов. 181 V.I. Лигнинолитические свойства Panus tigrinus 8/18 при разных способах культивирования. 181 V.2. Новый способ погруженного культивирования Panus tigrinus 8/18. 183 V.3. Mn-зависимая лигнин пероксидаза (МпЛП) P. tigrinus. 185 V.4. Лакказа P. tigrinus 8/18 и феномен желтых лакказ.
V.5. Прикладное значение лигнинолитических грибов и ферментов.
VI. Выводы.
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Введение диссертации (часть автореферата) на тему «Лигниназы базидиомицетов»
1.1. Актуальность проблемы.
Лигнин - ароматический биополимер, обязательный компонент всех наземных растений, крайне широко распространен на Земле, составляя 25% сухого веса фотосинтезирующей биомассы. В отличие от других биополимеров, - белков, нуклеиновых кислот и полисахаридов, лигнин не имеет регулярной структуры и содержит более десяти разных типов межмономерных связей, что определяет его устойчивость к химическому и микробиологическому разложению. Считается, что лигнин - самое устойчивое природное соединение.
Значимость лигнина для биосферы и хозяйственной деятельности человека трудно переоценить: сам факт существования лигнина определяет возможность существования растительного мира; изобилие лигнина и высокое удельное содержание углерода (вдвое выше чем у целлюлозы) делают этот материал одним из главных участников глобального круговорота углерода; продукты биотрансформации лигнина - основной источник гумуса, определяющего плодородие почв; лигноцеллюлоза - один из основных сырьевых ресурсов современной цивилизации. Однако, определение места лигнина в природе и решение технологических и экологических проблем при использовании лигнин-содержащих материалов ограничены недостаточным пониманием процессов биодеградации лигнина. Прогресс в этой области знания заметно уступает таким смежным вопросам как биодеградация целлюлозы или биосинтез лигнина. Причины - в сложности строения и вариабельности структуры лигнина у разных видов и даже в разных частях растений, в недостатке надежных и адекватных методов его изучения.
К настоящему времени в качестве эффективного биодеструктора лигнина охарактеризована лишь одна группа микроорганизмов, это - базидиомицеты, вызывающие т.н. белую гниль древесины. Пока неясно, является ли это следствием упомянутых технических трудностей, так что открытие новых микроорганизмов-лигнинолитиков из других таксонов лишь вопрос времени, или же это, - отражение реальной картины ограниченности природных лигнинолитических ресурсов вследствие сложности и устойчивости структуры лигнина.
В качестве лигнинолитических агентов грибов белой гнили определены внеклеточные ферменты класса оксидоредуктаз: лигнин пероксидаза, Мп-пероксидаза и лакказа. Стало очевидным, что эти ферменты работают в составе сложно организованных ферментных комплектов. Однако, до настоящего времени нет общепринятых представлений о деполимеризации лигнина отдельными ферментами и о взаимодействиях внутри лигнинолитических ферментных комплексов.
Тем не менее, во всем мире ведется интенсивная разработка биотехнологий на основе лигнинолитических грибов и ферментов как для обработки лигноцеллюлозных материалов (производство бумаги, хлопкового и льняного волокна и т.п.), так и для утилизации лигнин-содержагцих отходов. Эффективность лигнинолитических систем грибов при разложении устойчивого лигнина и неспецифичность лигнинолитических ферментов послужили поводом для использования интактных культур грибов и отдельных ферментов для разложения устойчивых ксенобиотиков. Описаны многочисленные примеры трансформации или минерализации разных типов устойчивых ксенобиотиков, что также активно применяется для разработки технологий биоремедиации различных природных сред.
1.2. Состояние вопроса.
К началу настоящей работы было известно, что базидиомицеты белой гнили древесины являются единственной группой микроорганизмов, способных разлагать лигнин [Kirk, 1984; Leisola and Fiechter, 1985]. Были изучены также физиологические условия проявления лигнинолитической активности этими грибами. Лигнинолитическая активность проявлялась как функция вторичного метаболизма вне зависимости от наличия или отсутствия лигнина, лигнин не разлагался без легко усваиваемого косубстрата, эффективность разложения лигнина стимулировалась повышенным парциальным давлением кислорода. Для соблюдения этих условий был разработан метод погруженного культивирования грибов в виде мицелиального мата в тонком слое жидкой среды, в 100% кислородной атмосфере, без перемешивания[Клгк et al., 1978]. Этот метод позволял получать культуры грибов с активированной лигнинолитической системой, однако, был слишком трудоемким, дорогим, и, главное, давал крайне низкий выход лигнинолитических ферментов. Кроме того, искусственная жидкая среда отличалась по физическим свойствам и составу от природных ростовых субстратов грибов, - древесины, травяного или лиственного опада. Это оставляло сомнения в идентичности лигнинолитических ферментных систем, получаемых в лабораторных условиях, и существующих в природе.
Характерным признаком лигнинолитических грибов белой гнили, отличающим их от других дереворазрушающих грибов, была способность продуцировать внеклеточные фенолоксидазы, - лакказу и пероксидазу. Однако, выделенные фенолоксидазы не окисляли нефенольные подструктуры, преобладающие в полимерном лигнине и определяющие его устойчивость к разложению. Этот парадокс был разрешен в 1984 г., когда у гриба белой гнили Phanerochaete chrysosporium были открыты новые лигнинолитические пероксидазы - лигнин пероксидаза и Mn-пероксидаза [Tien and Kirk, 1984; Kuwahara et al., 1984a], Надежды на прорыв в изучении энзимологии разложения лигнина связывали только с лигнин пероксидазой: это единственный фермент, для которого надежно доказали способность разрушать нефенольные подструктуры лигнина.
Другие ферменты, - лакказа и Mn-пероксидаза, - воспринимались как вспомогательные, способные лишь детоксифицировать фенольные продукты деполимеризации лигнина лигнин пероксидазой и вызывать некоторые изменения в полимерной структуре лигнина. Позднее, в 1991 г. был обнаружен эффект медиации у лакказ: в присутствии синтетических соединений определенной структуры голубая лакказа приобретала способность разлагать устойчивые соединения с редокс-потенциалом, превосходящим собственный [Bourbonnais and Paice, 1990]. Аналогичные реакции были найдены и для Mn-пероксидазы. Этот фермент разлагал нефенольные подструктуры лигнина в присутствии медиаторов радикальной природы, образующихся при сопряженном окислении тиолов или ненасыщенных липидов [Gold et al., 1990; Jensen et al., 1996]. Эти находки привели к размыванию идеи ключевого фермента лигнинолитического ферментного комплекса, каковым считали лигнин пероксидазу. Но строгих доказательств эффективности феномена медиации для лакказы и Mn-пероксидазы при разложении лигнина в природе не получено до сих пор.
Подавляющее количество исследований биодеградации лигнина проводили на примере базидиомицета белой гнили Phanerochaete chrysosporium. Интенсивное изучение этого организма дало свои результаты: основные сведения по физиологии и энзимологии разложения лигнина грибами были получены в экспериментах с Р. chrysosporium. Однако, со временем выяснилось, что P. chrysosporium не продуцирует лакказу в обычных условиях, чем отличается от типичных лигнинолитических грибов. Кроме того, концентрирование исследовательского интереса на единственном, хотя и хорошо изученном во всех аспектах организме, оставляло за рамками научного поиска разнообразие ферментных и метаболических систем других грибов.
В настоящее время, при изучение энзимологии биодеградации лигнина большее внимание уделяется новым видам грибов, распространяется практика использования твердофазного культивирования на природных ростовых субстратах, общий характер исследований переориентируется от изучения отдельных ферментов к выяснению взаимосвязей внутри ферментных ансамблей, включая неэнзиматические реакции.
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Заключение диссертации по теме «Биохимия», Леонтьевский, Алексей Аркадьевич
VI. Выводы.
1. Впервые изучены физиологические и лигнинолитические характеристики гриба Panus tigrinus в условиях погруженного и твердофазного культивирования.
P. tigrinus проявлял лигнинолитическую активность при переходе культуры ко вторичному метаболизму. При сопоставлении с другими грибами, P. tigrinus демонстрировал наибольшую скорость разложения лигнина. В отличие от типичных лигнинолитических грибов, лигнинолитическая активность P. tigrinus стимулировалась марганцем.
2. Изучены состав и организация лигнинолитического ферментного комплекса гриба Panus tigrinus.
P. tigrinus не продуцирует лигнин пероксидазу при любых условиях культивирования. В состав лигнинолитического ферментного комплекса Р. tigrinus входят новая Мп-зависимая лигнин пероксидаза, как ключевой фермент, и семейство лакказ.
3. Разработан новый способ погруженного культивирования грибов, повышающий выход лигнинолитических ферментов.
Способ включает использование минеральной среды, лимитированной по азоту, специфических органических и неорганических индукторов ферментов, детергента твина-80, иммобилизацию мицелия и температурный сдвиг. Выход ферментов, по сравнению со стандартными условиями культивирования увеличен в 50 раз.
4. Впервые выделен и охарактеризован новый лигнинолитический фермент -Mn-зависимая лигнин пероксидаза (МпЛП) Panus tigrinus.
Новый фермент обладает уникальным каталитическим циклом: в его заключительной реакции Соединение II восстанавливается до нативно й1 формы как марганцем, так и органическим субстратом. Вследствие этого, каталитические свойства фермента сочетают характеристики лигнин пероксидазы и Mn-пероксидазы. Физико-химические свойства нового фермента аналогичны свойствам других лигнинолитических пероксидаз.
5. Впервые обнаружен и изучен феномен желтых лакказ у лигнинолитических грибов.
Желтые лакказы грибов Partus tigrinus, Phlebia radiata, Phlebia tremellosa и Agaricus bisporus, выделенные из твердофазных культур, не имели типичных спектральных и каталитических свойств, в отличие от голубых лакказ из погруженных культур. Предполагается, что желтые лакказы образуются в результате модификации обычных голубых лакказ продуктами разложения лигнина. При этом изменяются вторичная структура и микроокружение атомов меди в активном центре, желтая лакказа приобретает способность окислять устойчивые подструктуры лигнина. Выделены и охарактеризованы множественные формы желтой лакказы гриба Panus tigrinus.
6. Разработаны способы практического применения свободных и иммобилизованных голубых лакказ грибов для делигнификации лигноцеллюлозных материалов и трансформации хлорфенолов.
При использовании грибных лакказ для отбеливания лигноцеллюлозного волокна достигали 70-75% делигнификации.
Впервые изучена реакция трансформации 2,4,6-трихлорфенола лакказами. Предложен способ удаления хлорфенолов из водной среды иммобилизованными лакказой (за 10-15 мин. при 400 мг/л токсиканта).
7. Разработаны способы практического применения культуры Panus tigrinus для биоремедиации почвы и воды.
В полевых условиях P. tigrinus снижал содержание полихлорфенолов до 48% за 4 месяца при исходной концентрации до 500 мг на кг почвы.
При трансформации 2,4,6-трихлорфенола в воде культурой P. tigrinus и очищенными лакказами/*. tigrinus и С. versicolor, впервые обнаружен 3,5-дихлоркатехол как продукт о^шо-дегалогенирования хлорфенола. Впервые предложен способ адаптации лигнинолитических грибов к высоким концентрациям ксенобиотиков на примере хлорфенолов.
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