В статье изложена история изучения численности и разнообразия мицелиальных грибов в образцах глубинных горизонтов и активного слоя Арктики и Антарктиды, установленных с помощью традиционных микробиологических методов. На обширном экспериментальном материале показано, что общая численность грибов в глубинных горизонтах антарктических отложений в целом ниже, чем в Арктике. Подтверждается микроочаговое распределение грибов в многолетнемерзлых отложениях, а также обсуждается их таксономический состав. Анализируется видовое разнообразие и выявляются таксоны с высокой частотой встречаемости, среди которых находятся грибы родов Penicillium и Geomyces (tm Pseudogymnoascus), адаптированные к существованию в экстремальных экотопах. Обсуждаются особенности их метаболизма при действии разных стессоров и реакции грибов рода Geomyces на изменение абиотических факторов на генетическом уровне. The article sets out the history of the study the number of mycelial fungi established using traditional microbiological methods in deep horizons and in the samples of the active layers of the Arctic and Antarctica. An extensive experimental data shows that the total number of fungi in the deep horizons of the Antarctic sediments as a whole is lower than in the Arctic. The microfocal distribution of fungi in permafrost deposits is confirmed, and their taxonomic composition is also discussed. Biodiversity is analyzed and the taxa with a high frequency of occurrence are detected. Among them there are fungi from genera Penicillium and Geomyces (tm Pseudogymnoascus) adapted to life in extreme ecotopes. The features of their metabolism under the influence of different stressors and the reaction of Geomyces on the changing of abiotic factors at the genetic level are discussed.

Secondary metabolites of 25 Penicillium strains isolated from high-latitude ecosystems (upper layer of Antarctic soils and Arctic permafrost deposits) were analyzed. Out of the studied strains, 80% were found to produce secondary metabolites belonging to benzodiazepine alkaloids (anacin, cyclopenin, and cyclopeptin), quinoline alkaloids (viridicatin and 3-methoxyviridicatin), diketopiperazine alkaloids (aurantiamine, 3,12-dihydroroquefortine, roquefortine and rugulosuvin B), polycyclic indole alkaloids (communesin B and chаetoglobosine A), clavine ergot alkaloids (rugulovasins A and B, festuclavine, fumigaclavines A and B, and cyclopiazonic acid), polyketides (mycophenolic acid and citreoviridin), terpenes (andrastins A and B and phomenone), and N-acetyltriptamine. Most strains of the Penicillium subgenus isolated from anthropogenically impaired upper layers of Antarctic deposits and from subsurface Arctic deposits exhibited more complete spectra of secondary metabolites compared to three strains isolated from permafrost 15 000 to 600 000 years old. This is the first report on andrastins formation by a P. restrictum. Wide occurrence of rugulovasins in P. variabile strains was shown.

Secondary metabolites of 22 fungal strains (genus Aspergillus, section Usti) isolated at diverse geographic regions, including the Arctic permafrost deposits, were studied. The studied strains were found to synthesize a variety of biologically active compounds, structurally identified as drimane sesqueterpenoids, isoquinoline alkaloids (TMC-120 A−C, derivative 1), meroterpenoids (austalides О and J), and anthraquinone pigments (averufin, versicolorin C). Desferritriacetylfusigen production by A. calidoustus isolates is reported for the first time. The individual spectra of secondary metabolites were used for reidentification of 17 strains, of which 15 were identified as A. calidoustus and two, as A. pseudodeflectus.

The effect of potassium sorbate, sodium benzoate, and sodium nitrite used as preservatives in the food industry in the production of such mytotoxins as citrinin cyclopiazonic and mycophenolic acids by the contaminating fungi Penicillium citrinum, P. commune, and P. brevicompactum, respectively, was investigated. It was shown that the effect of preservatives used at concentrations relevant to the food industry on the synthesis of mycotoxins depended on the species-specific biochemical and physiological features of the cultures. The growth of P. brevicompactum was inhibited to the highest degree by sodium nitrite and potassium sorbate, and the growth of P. commune was so inhibited by sodium benzoate. It was established that the introduction of 0.015% sodium nitrite into the medium resulted in 1.3- and 1.4-fold reductions of the production of citrinin and mycophenolic acid, respectively, while the production of cyclopiazonic acid did not change in comparison with the control. The introduction of 0.015% sodium benzoate caused a more than 1.5-fold increase of the concentration of citrinin, cyclopiazonic, and mycophenolic acids, and the addition of 0.02% potassium sorbate increased the production of cyclopiazonic and mycophenolic acids by 1.7 and 2.6 times, respectively.

Exometabolites of 22 strains of the genus Penicillium, section Chrysogena isolated from low-temperature ecotopes of various geographical regions were analyzed. The ecotopes included permafrost deposits, frozen volcanic ash, a fossil horse, cryopeg, and water from an Antarctic lake. The studied strains were found to contain exometabolites belonging to the groups of penicillins (penicillin G), chrysogines (chrysogine, 3-acetyl-quinazolin-4(3H)-one, 2-pyruvoylaminobenzamide, 2-(2-hydroxypropionylamino)-benzamide, and questiomycin A), roquefortines (3,12-dihydroroquefortine, roquefortine, glandicolines A and B, and meleagrine), xanthocillins (xanthocillin X), and simple tryptophan derivatives (N-acetyltriptamine and indoleacetic acid). In five P. chrysogenum strains and three P. nalgiovense strains, a correlation was found between exometabolite spectra and morphological characteristics of the cultures isolated from modern ecotopes. For other strains species, identification was based on morphological features due to the absence of biosynthesis of penicillin G on the major chemotaxonomic markers for these species.

The recent data on exometabolite biosynthesis in fungi of the genus Penicillium is summarized. The study of creative species, as well as those isolated from extreme ecotopes, resulted in the identification of a number of novel, biologically active compounds. Alkaloid biosynthesis has been shown to begin on the first day of fungus cultivation and to proceed throughout the cultivation period. Idiophase kinetics was observed for the biosynthesis of polyketide metabolites. The mechanisms of regulation of biosynthesis of promising bioactive compounds are discussed.

Fungi of the genus Penicillium isolated from little studied habitats are able to synthesize both previously known and new physiologically active compounds with diverse structures. They include secondary metabolites of alkaloid nature, i.e., ergot alkaloids, diketopiperazines, quinolines, quinazolines, benzodiazepines, and polyketides. We discuss the use of profiles of secondary metabolites for taxonomy purposes. Studying the physicochemical characteristics of producers of biologically active compounds showed that the biosynthesis of alkaloids is initiated on the first days of cultivation and proceeds simultaneously with growth. The cyclic character of alkaloid accumulation was recorded related to the processes of alkaloid biosynthesis, excretion from cells, degradation in culture fluid, and consumption by cells. Synchronic variations in the concentrations of intracellular tryptophan and alkaloids are necessary for the regulation of the optimal quantity of tryptophan necessary for the culture.

Fungi of the genus Penicillium isolated from little studied habitats are able to synthesize both previously known and new physiologically active compounds with diverse structures. They include secondary metabolites of alkaloid nature, i.e., ergot alkaloids, diketopiperazines, quinolines, quinazolines, benzodiazepines, and polyketides. We discuss the use of profiles of secondary metabolites for taxonomy purposes. Studying the physicochemical characteristics of producers of biologically active compounds showed that the biosynthesis of alkaloids is initiated on the first days of cultivation and proceeds simultaneously with growth. The cyclic character of alkaloid accumulation was recorded related to the processes of alkaloid biosynthesis, excretion from cells, degradation in culture fluid, and consumption by cells. Synchronic variations in the concentrations of intracellular tryptophan and alkaloids are necessary for the regulation of the optimal quantity of tryptophan necessary for the culture.

Биосинтез фумихиназолинов F и G (ФХ), РС-2 и пигментов грибом P. thymicola ВКМ FW-869 напрямую зависит от количества углеродного субстрата (маннит) в среде. При всех изученных концентрациях маннита преобладала продукция пигментов. Условием преимущественного биосинтеза ФХ грибом P. thymicola является лимитирование источником углерода (маннитом) и присутствие NaCl в среде культивирования гриба. NaCl оказывает регуляторное влияние на образование вторичных метаболитов, увеличивая биосинтез ФХ и снижая образование пигментов. Максимальные показатели биосинтеза ФХ и подавление продукции пигментов достигнуты при концентрации маннита в среде 20 г/л и 2.5% NaCl.

Biosynthesis of fumiquinazolines F and G (FQs), PC-2, and pigments by the fungus P. thymicola VKM FW-869 is directly dependent on the content of carbon substrate (mannitol) in the medium. Pigment production prevailed at all of the tested mannitol concentrations. The necessary conditions for predominant FQ biosynthesis by the fungus P. thymicola are carbon source (mannitol) limitation and presence of NaCl in the cultivation medium. NaCl has a regulatory effect on the formation of secondary metabolites by enhancing FQ biosynthesis and reducing pigment formation. The maximum values of FQ biosynthesis and inhibition of pigment production are obtained at a mannitol concentration of 20 g/l and 2.5% NaCl in the medium.

Проведен скрининг продуцентов вторичных метаболитов среди 25 штаммов грибов рода Penicillium, выделенных из отложений вечной мерзлоты Арктики, Антарктиды и из мерзлого вулканического пепла Камчатки. Обнаружено, что половина из исследованных штаммов синтезирует биологически активные вещества алкалоидной природы: эргоалкалоиды, дикетопиперазины и производные хинолина. Большая часть из идентифицированных метаболитов относится к микотоксинам. Найден штамм Penicillium waksmanii – продуцент эпоксиагроклавина-I и хиноцитрининов, исследованы его основные физиолого-биохимические характеристики.

Четыре культуры грибов рода Penicillium, относящихся к различным видам подрода Furcatum Pitt: P. citrinum Thom 1910, P. corylophiium Dierckx 1901, P. fellutanum Biourge 1923 и P. waksmanii Zaleski 1927, продуцировали эргоалкалоиды агроклавин-I, эпоксиагроклавин-I, их N–N-димеры – димер эпоксиагроклавина-I и смешанный димер эпоксиагроклавина-I и агроклавина-I, а также хинолиновые метаболиты – хиноцитринины А и Б. Изучена физиолого-биохимическая характеристика продуцентов. Определены оптимальные условия биосинтеза компонентов метаболома. Добавка цинка в среду стимулировала биосинтез эргоалкалоидов во всех случаях, продукция же хиноцитрининов возрастала только у P. citrinum, а у P. corylophiium, P. fellutanum и P. waksmanii подавлялась. Это свидетельствует о том, что гены путей биосинтеза этих метаболитов находятся у продуцентов в разных кластерах.