Добавил:
Sekretar
kiopkiopkiop18@yandex.ru
t.me/Prokururor I Вовсе не секретарь, но почту проверяю
Опубликованный материал нарушает ваши авторские права? Сообщите нам.
Вуз:
Предмет:
Файл:Ординатура / Хирургия / Библиотека им академика М.И. Перельмана / Книга_5182_Библиотеки_им_академика_М_И_Перельмана
.pdf
396 Wild Edible Plants
Wang, K., Bao, L., Qi, Q/ , Zhao, F., Ma, K., Pei, Y., Liu, H. (2015). Erinacerins C- L, isoindolin-
1- ones with α- glucosidase inhibitory activity from cultures of the medicinal mushroom
Hericium erinaceus. Journal of Natural Products, 78(1), 146– 154. https:// doi.org/ 10.1021/
np5004 388
Wang, S.J., Li, Y.X., Bao, L., Han, J.J., Yang, X.L., Li, H.R., Wang, Y.Q., Li, S.J., & Liu,
H.W. (2012). Eryngiolide A, a cytotoxic macrocyclic diterpenoid with an unusual
cyclododecane core skeleton produced by the edible mushroom Pleurotus eryngii.
Organic Letters, 14(14), 3672– 3675. https:// doi.org/ 10.1021/ ol3015 19m
Wang, Y., Huang, X., & Nie, S. (2018). Novel prospective of wild mushroom polysaccharides
as potential prebiotics. In B. Singh, Lallawmsanga, & A. Passari (Eds.), Biology of
macrofungi. Fungal biology, (pp. 211– 226). Springer, Cham. https:// doi.org/ 10.1007/ 9783- 030- 02622- 6_ 10
Wang, Z., Qu, K., Zhou, L., Ren, L., Ren, B., Meng, F., Yu, W., Wang, H., & Fan, H. (2021).
Apaf1 nanoLuc biosensors identied lentinan as a potent synergizer of cisplatin in targeting
hepatocellular carcinoma cells. Biochemical and Biophysical Research Communications,
577, 45– 51. https:// doi.org/ 10.1016/ j.bbrc.2021.08.030
Wei, L., Wang, W., Hou, Y., Wang, X., Liu, Y., Xie, X., Li, X., Wang, T., & Jing, B. (2023).
Assessment of the quality characteristics of Stropharia rugosoannulata subjected to ve
different drying methods. Journal of Food Processing and Preservation, article 2518620,
1– 12. https:// doi.org/ 10.1155/ 2023/ 2518 620
Wu, F., Tang, J., Pei, F., Wang, S., Chen, G., Hu, Q., & Zhao, L. (2014). The inuence of
four drying methods on nonvolatile taste components of White Hypsizygus marmoreus.
European Food Research and Technology, 240(4), 823– 830. https:// doi.org/ 10.1007/ s00
217- 014- 2388- 4
Wu, J., Kobori, H., Kawaide, M., Suzuki, T., Choi, J. H., Yasuda, N., Noguchi, K., Matsumoto,
T., Hirai, H., & Kawagishi, H. (2013). Isolation of bioactive steroids from the Stropharia
rugosoannulata mushroom and absolute conguration of Strophasterol B. Bioscience,
Biotechnology, and Biochemistry, 77(8), 1779– 1781. https:// doi.org/ 10.1271/ bbb.130 216
Wu, J., Tokuyama, S., Nagai, K., Yasuda, N., Noguchi, K., Matsumoto, T., Hirai, H., & Kawagishi,
H. (2012). Strophasterols A to D with an unprecedented steroid skeleton: From the mushroom Stropharia rugosoannulata. Angewandte Chemie, 124(43), 10978– 10980. https:// doi.
org/ 10.1002/ ange.201205 351
Wu, L., Liu, X., Hu, R., Chen, Y., Xiao, M., Liu, B., & Zeng, F. (2022). Prebiotic Agrocybe
cylindracea crude polysaccharides combined with Lactobacillus rhamnosus GG postpone
aging- related oxidative stress in mice. Food & Function, 13(3), 1218– 1231. https:// doi.org/
10.1039/ D1F O020 79J
Xu, Q., Wang, H., Li, T., Chen, L., Zheng, B., & Liu, R. H. (2020). Comparison of phenolics,
antioxidant, and antiproliferative activities of two Hypsizygus marmoreus varieties. Journal
of Food Science, 85(7), 2227– 2235. https:// doi.org/ 10.1111/ 1750- 3841.15173
Xue, Z., Li, J., Cheng, A., Yu, W., Zhang, Z., Kou, X., & Zhou, F. (2015). Structure identi-
cation of triterpene from the mushroom Pleurotus eryngii with inhibitory effects against
breast cancer. Plant Foods for Human Nutrition, 70(3), 291– 296. https:// doi.org/ 10.1007/
s11 130- 015- 0492- 7
Xue, Z., Ma, Q., Chen, Y., Lu, Y., Wang, Y., Jia, Y., Zhang, M., & Chen, H. (2020). Structure
characterization of soluble dietary ber fractions from mushroom Lentinula edodes (Berk.)
Pegler and the effects on fermentation and human gut microbiota in vitro. Food Research
International, 129, 108870. https:// doi.org/ 10.1016/ j.food res.2019.108 870
Yan, Q.- X., Huang, M.- X., Sun, P., Cheng, S., Zhang, Q., & Dai, H. (2020). Steroids, fatty acids and
ceramide from the mushroom Stropharia rugosoannulata Farlow apud Murrill. Biochemical
Systematics and Ecology, 88, 103963. https:// doi.org/ 10.1016/ j.bse.2019.103 963
Edible and Medicinal Mushrooms as an Eco-Friendly Source of Food 397
Yang, Z., Xu, J., Fu, Q., Fu, X., Shu, T., Bi, Y., & Song, B. (2013). Antitumor activity of a poly-
saccharide from Pleurotus eryngii on mice bearing renal cancer. Carbohydrate Polymers,
95(2), 615– 620. https:// doi.org/ 10.1016/ j.carb pol.2013.03.024
Yao, F., Gao, H., Yin, C. M., Shi, D. F., & Fan, X. Z. (2023). Effect of different cooking methods
on the bioactive components, color, texture, microstructure, and volatiles of Shiitake
mushrooms. Foods, 12(13), 2573. https:// doi.org/ 10.3390/ foods1 2132 573
Ye, K., Fu, C., Ma, S., Du, H., Chen, S., Liu, D., Ma, G., & Xiao, H. (2023). Comprehensive
assessment of Hypsizygus marmoreus polysaccharides through simulated digestion and
gut microbiota fermentation in vitro. Food Hydrocolloids, 144, 108989. https:// doi.org/
10.1016/ j.food hyd.2023.108 989
Yu, Q., Guo, M., Zhang, B., Wu, H., Zhang, Y., & Zhang, L. (2020). Analysis of nutritional com-
position in 23 kinds of edible fungi. Journal of Food Quality, 2020, 1– 9. https:// doi.org/
10.1155/ 2020/ 8821 315
Zan, X., Cui, F., Li, Y., Yang, Y., Wu, D., Sun, W., & Ping, L. (2015). Hericium erinaceus
polysaccharide- protein HEG- 5 inhibits SGC- 7901 cell growth via cell cycle arrest and
apoptosis. International Journal of Biological Macromolecules, 76, 242−253. https:// doi.
org/ 10.1016/ j.ijbio mac.2015.01.060
Zhai, X., Zhao, A., Geng, L., & Xu, C. (2013). Fermentation characteristics and hypogly-
cemic activity of an exopolysaccharide produced by submerged culture of Stropharia
rugosoannulata. Annals of Microbiology, 63(3), 1013– 1020. https:// doi.org/ 10.1007/ s13
213- 012- 0555- z
Zhang, B., Li, Y., Zhang, F., Linhardt, R. J., Zeng, G., & Zhang, A. (2020). Extraction, structure
and bioactivities of the polysaccharides from Pleurotus eryngii: A review. International
Journal of Biological Macromolecules, 150, 1342– 1347. https:// doi.org/ 10.1016/ j.ijbio
mac.2019.10.144
Zhang, J., Feng, N., Liu, Y., Zhang, H., Yang, Y., Liu, L., & Feng, J. (2022). Bioactive compounds
from medicinal mushrooms. Advances in Biochemical Engineering/ Biotechnology, 184,
219– 268. https:// doi.org/ 10.1007/ 10_ 2 022_ 202
Zhang, M., Zhang, Y., Zhang, L., & Tian, Q. (2019). Mushroom polysaccharide lentinan for
treating different types of cancers: A review of 12 years clinical studies in China. Progress
in Molecular Biology and Translational Science, 163, 297– 328. https:// doi.org/ 10.1016/
BS.PMBTS.2019.02.013
Zhang, W., Tian, G., Geng, X., Zhao, Y., Ng, T., Zhao, L., & Wang, H. (2014). Isolation and
characterization of a novel lectin from the edible mushroom Stropharia rugosoannulata.
Molecules, 19(12), 19880– 19891. https:// doi.org/ 10.3390/ mol ecul es19 1219 880
Zhou, G., Liu, H., Yuan, Y., Wang, Q., Wang, L., & Wu, J. (2024). Lentinan progress in inamma-
tory diseases and tumor diseases. European Journal of Medical Research, 29(1), 8. https://
doi.org/ 10.1186/ s40 001- 023- 01585- 7
Regulation of the
Biological Activity
14
of Medicinal
Macromycetes Using
Low- Intense QuasiMonochromatic and
Laser Light
Oksana Mykchaylova, Margarita Lomberg,
Alexander Galkin, and Natalia Poyedinok
14.1 INTRODUCTION
Searching for new sources of physiologically active compounds to obtain functional
and safe products is one of the most critical tasks of modern biotechnology (Badalyan
& Rapior, 2020; Deshmukh et al., 2022; Hyde et al., 2019). Edible and medicinal
macromycetes, in addition to their nutritional value and bioavailability, represent a rich
but largely underutilized resource of useful natural compounds with diverse pharmacological activities (Hobbs, 2023; Lu et al., 2020; Martinez- Medina et al., 2021; RangelVargas et al., 2021; Song et al., 2020; Stabnikova et al., 2024; Venturella et al., 2021).
Considering that edible mushrooms are producers of valuable metabolites, prospects are
opening up for their use in the development of new functional foods and nutraceuticals
398
DOI: 10.1201/9781003486794-14
Regulation of the Biological Activity of Medicinal Macromycetes 399
without serious problems regarding the safety of their use (Bell et al., 2022; Cohen
et al., 2014; Giavasis, 2014; Ivanov et al., 2021; Rangsinth et al., 2023; Valverde
et al., 2015). Depending on the purpose of use, various products based on edible and
medicinal mushrooms have been developed, which can be broadly divided into categories: functional foods, dietary supplements, nutraceuticals, and designer foods,
including probiotics and prebiotics, which can provide benecial effects on health especially by frequent human consumption (Kour et al., 2022; Morris et al., 2016; Niego
et al., 2021, 2023; Rangel- Vargas et al., 2021) (Figure 14.1).
Although edible and medicinal mushrooms have been used by humans for thousands
of years, it is only in recent decades that mushroom consumption has increased signicantly, mainly due to the growing awareness that a stable and balanced diet plays a
key role in the normal functioning and the maintenance of health (Reis et al., 2017).
According to Food and Agriculture Organization Statistical (FAOSTAT) data, the
current global mushroom output trend is rising, with a global mushroom production
of 43 million metric tons in 2018– 2019. Mushroom consumption is expected to rise,
FIGURE 14.1 Graphical representation of the use of edible and medicinal mushroombased products.
400 Wild Edible Plants
indicating a continuous upward trend in mushroom production. The global mushroom
market was valued at USD 45.3 billion in 2020 and is predicted to increase at a compound annual growth rate of 7% from 2022 to 2027 (Hamza et al., 2024; Niego et al.,
2021, 2023).
Currently, nutrition is considered a fundamental basis for maintaining human
health and development throughout life. Moreover, it is very important to have a proper
diet and sufcient nutrients for physical growth, mental development, performance
and productivity, health, and well- being. From this point of view, edible and medicinal
mushrooms can be considered functional and safe food products.
Functional foods are regular or everyday foods consumed as part of a normal
daily diet. They must consist of naturally occurring components (perhaps in unnatural
concentrations or found in foods that do not normally contain them). Functional foods
have a positive impact on target body functions beyond their nutritional value, improving
well- being and quality of life and/ or reducing the risk of diseases (El- Sohaimy, 2012).
Fresh edible and medicinal mushrooms and their dried powders, aqueous extracts,
concentrates, and bioactive compounds can be incorporated into foods to develop fortied functional foods with the aim to substitute meat, fat, our, minerals, phosphates,
and antioxidants in addition to contributing long shelf- life and low- cost production.
Bakery, meat, dairy, fermented, fruit and vegetable, traditional products, and beverages
have already been studied in vitro to present value- added and mushroom- based func-
tional foods. In these studies, some changes in the nutritional, sensorial, textural, and
pharmacological properties of novel value- added products were observed (Bulam et al.,
2022; Das et al., 2021; Dorin and Melinda, 2021; Gonzales et al., 2020; Kumar et al.,
2021; Kumaresan et al., 2022; Reis et al., 2017; Rangel- Vargas et al., 2021; Simakhina
et al., 2022; Stabnikova et al., 2024; Yurchak & Sharkova, 2022).
Due to the high bioactive potential of edible and medicinal mushrooms, as well as
the growing interest in natural medicines for the “treatment” of various physiological
disorders, a new and specic term is now recognized: “mushroom nutraceuticals”,
which should not be confused with general nutraceuticals, functional foods’ nutrition,
and/ or general purpose food products or pharmaceuticals (Reis et al., 2017). The word
“mushroom nutraceuticals” has been devised by Chang and Buswell (1996) and can be
dened as rened or partially dened extractives from both mycelium or fruit bodies,
frequently ingested in the form of capsules as a dietary supplement and symbolize an
important constituent of the expanding mushroom biotechnology industry (Kour et al.,
2022). Mushroom nutraceuticals include puried extracts from fruiting bodies, puried
polysaccharides, and dried mycelium biomass or mushrooms, which are consumed in
the form of capsules, tablets, powder, syrup, and solution (Kour et al., 2022; Hozova
et al., 2004; Reis et al., 2017). Mushroom nutraceuticals contain the following:
• lipids, especially unsaturated fatty acids;
• vitamins such as vitamins E, D, and vitamin C;
• proteins, peptides, and amino acids, including lectins, leucine, and valine;
• carbohydrates, especially polysaccharides such as lentinan.
Due to the presence of a wide range of bioactive compounds in mushrooms, they have
great potential for the production of new nutraceutical formulations. However, despite
the recognition of the enormous potential of edible and medicinal mushrooms as the
Regulation of the Biological Activity of Medicinal Macromycetes 401
basis of such formulations, there remain some unresolved issues affecting their preparation and subsequent marketing, such as issues of safety, standardization, regulation,
efcacy, and mechanism of action (Kour et al., 2022).
Thus, the role of functional foods is primarily related to disease risk reduction
rather than disease prevention, while nutraceuticals are typically consumed to promote
well- being through the prevention and/ or treatment of diseases and/ or disorders (Reis
et al., 2017; El- Sohaimy, 2012). Additionally, nutraceuticals should not be confused
with medications, which are administered in precise doses under medical supervision.
Because functional foods and nutraceuticals have properties found in foods and drugs,
they are considered to straddle the boundary between conventional foods and drugs
(Reis et al., 2017).
Sometimes this concept is also misused to describe another food concept the
“dietary supplement”. Dietary supplements have essentially a feed function, taking the
form of medicines (pills or capsules). Once they take the form of medicines and are not
consumed as traditional food in the daily diet, they cannot be regarded as functional
foods (Reis et al., 2017). Mushroom dietary supplements on the market are presented
in the following forms:
• powders from cultivated fruiting bodies, aqueous or alcoholic extracts of these
fruiting bodies;
• dried and crushed preparations from a combination of substrate, mycelium,
and mushrooms primordia;
• biomass or extracts from mycelium obtained by deep cultivation in fermenters
or bioreactors;
• dried fruiting bodies of wild mushrooms in the form of capsules or tablets;
spores and their extracts (Kour et al., 2022; Reis et al., 2017).
One of the advantages of using mushroom- based food additives is their safety. Most
of the mushrooms that are used in dietary supplements are cultivated on an industrial
scale under strictly controlled conditions. Industrial strains of mushrooms undergo strict
taxonomic control, are easily propagated in vitro, and are capable of growing as mycelial biomass in submerged cultures. Growing medicinal mushrooms under standard
conditions make it possible to control and intensify the synthesis of certain biologically
active substances and obtain a nal product with a reproducible chemical composition,
comparable biological effects, and greater safety for health (Chang & Wasser, 2018).
Although about 270 species of mushrooms have proven medicinal properties, only a
few are considered nutraceuticals. Species most commonly found in dietary supplements
include Agaricus bisporus (Button mushroom), Ophiocordyceps sinensis (Cordyceps),
Ganoderma lucidum (Reishi), Grifola frondosa (Maitake), Hericium erinaceus (Lion’s
mane), Lentinula edodes (Shiitake), and Trametes versicolor (Turkey tail) (Niego et al.,
2021). Further study of the biological properties of mushrooms will signicantly expand
the list of potential candidates that can be classied as nutraceuticals.
Medicinal mushrooms have been used for thousands of years in traditional oriental
medicine for the treatment and prevention of various diseases without side effects (Hobbs,
2023; Zhang et al., 2022). Using modern methods, many secondary metabolites such
as polysaccharides, proteins and their complexes, phenolic compounds, polyketides,
triterpenoids, steroids, alkaloids, and nucleotides have been isolated and identied
402 Wild Edible Plants
from fruiting bodies, mycelium, and culture liquid of fungi. Some compounds have
antidiabetic, antioxidant, antitumor, immunomodulatory, antimicrobial, and antiviral
effects (Chang & Wasser, 2018; Niego et al., 2021, 2023; Sandargo et al., 2019). The
presence of a large spectrum of biological active compounds in mushrooms can be
explained by the fact that in nature, the development of fungal organisms occurs in
direct contact with the environment. Therefore, they are constant objects of stressors
of a physical and chemical nature. External factors such as the absence or excess of a
specic food source, adverse lighting and temperature conditions, changes in the gas
composition of the environment, and mechanical damage to the mycelium trigger stress
response processes. Under natural conditions, the survival of mushrooms under stress
is the rule rather than the exception. During evolution, fungi have developed survival
strategies by creating barrier systems and synthesizing numerous bioactive metabolites
that allow them to survive in their natural environment (Hyde, 2022).
The primary method of growing macromycetes currently remains solid- phase cultivation for fruiting bodies production, but this is a long and labor- intensive process
(Arya & Rusevska, 2022; Dedousi et al., 2023; Nikšić et al., 2022; Stabnikova et al.,
2010). The most promising technology for rapidly and efciently producing mushroom
mycelium and valuable metabolites from edible and medicinal mushrooms is submerged
cultivation in bioreactors (Bakratsas et al., 2021; Berovic & Zhong, 2022a). Cultivation
of mushrooms in a submerged state has signicant industrial potential. However, its
success on a commercial scale depends on the development of technological stages for
intensifying the process of obtaining the nal product, including the use of environmentally friendly regulators of the biosynthetic activity of the producing fungus (Berovic
& Zhong, 2022a, 2022b, 2023). The creation of new technologies involves an in- depth
study of the factors regulating the functions of the fungal organism, which will make it
possible to use their natural potential with the most excellent efciency and ensure the
obtaining of products of the required quality in the required quantity.
Light is one of the environmentally friendly regulators of morphogenesis, biosynthetic, and biological activity of producing macromycetes (Kojima et al., 2015; Nakano
et al., 2010; Yue et al., 2022). Understanding the inuence of light on the physiology and
metabolism of macromycetes may positively impact the development of bioprocesses
that focus on the targeted synthesis of specic bioactive compounds.
Studying the mechanisms of fungal photoreactions is a complex task, both from
a fundamental and practical point of view. Knowledge of a producer’s photoreception
mechanisms is an integral part of the targeted photoregulation of its activity in biotechnological processes.
For most macromycetes, light is an essential source of temporal and spatial information regarding environmental changes (Wang et al., 2016). In addition, it can indicate
various environmental stresses such as exposure to genotoxic ultraviolet radiation (UV),
oxidative stress, increased temperature, or decreased humidity (Fuller et al., 2016; Yu
& Fischer, 2019).
For decades, reactions to light have been studied in at least 100 species of fungi
from different systematic groups. Particular attention was paid to the study of the perception of light signals by fungi and their transformation into morphological, physiological, and metabolic reactions. The mechanisms by which fungi perceive blue,
near- UV, green, and red light have been studied (Corrochano, 2007, 2019; Corrochano
Regulation of the Biological Activity of Medicinal Macromycetes 403
& Galland, 2006). It has been established that the processes of spore germination, development and growth of mycelium, and formation of fruiting bodies can be controlled by
lighting conditions (Corrochano, 2007, 2019; Corrochano & Galland, 2006; Fischer
et al., 2016; Idnurm et al., 2010; Igbalajobi et al., 2019; Yu & Fischer, 2019; Yu et al.,
2021). Moreover, although a massive amount of information has been accumulated
over the last century on various aspects of photoreception and signal transduction by
fungi, issues of photoregulation of specic physiological reactions, such as changes in
metabolism, are multidimensional processes that require further research. Once basic
science has identied the signaling pathways that control, for example, the biosynthetic
activity of a producing fungus, applied science can develop physical or genetic means
by which these pathways can be regulated and the yield of the desired end product can
be increased.
14.2 MECHANISMS OF FUNGAL
PHOTORECEPTION
During evolution, fungi formed the photoregulatory system mycochrome. A distinctive
feature of this system is the dependence of some stages of morphogenesis and physiology on the duration and intensity of exposure to light (Rodriguez- Romero et al., 2010).
The reactions of various types of fungi to light began to be studied more than 150 years
ago. They can be rapid, such as phototropism, or take a certain time. Light- induced
reactions in fungi can be triggered by very brief exposure (nanoseconds to minutes)
to low- intensity light (Corrochano, 2007). The long- term effects of light depend primarily on genetic reprogramming of the fungal genome and typically involve signicant
changes in gene expression patterns. The main question is how much- coordinated activation and repression of hundreds of genes is achieved (Fischer et al., 2016).
The issues of photoreception of light energy in the mycelium and the mechanisms
of reactions that occur after light absorption are quite complex and are still being intensively studied. It is currently known that light sensors in fungi are chromoproteins,
which are low molecular weight compounds that absorb light in certain parts of the
spectrum and initiate protein reactions. Some photoreceptive proteins with diverse
structures, mechanisms of action, and photosensitivity have been identied in fungi
(Galindo et al., 2022). Photoreceptors are specialized proteins or protein complexes that
are capable of catching light, generating a signal that travels into the cell and stimulates
a cellular response. Their light- absorbing cofactors, known as chromophores, are heterocyclic organic molecules capable of capturing photons of light (Herrera- Estrella
& Horwitz, 2007). Fungal photoreceptors often have multiple protein domains with
different functions. Typically, a photoreceptor domain is either attached to a chromophore or is near a chromophore, which, when illuminated, can establish a chemical
bond with that domain (Bayram & Bayram, 2023). The rst step in light perception is
a physical reaction, the absorption of photons by an organic chromophore molecule,
which triggers a light- dependent signaling pathway that causes protein conformational
changes and transduction into biochemical reactions affecting gene expression in the
404 Wild Edible Plants
body (Fischer et al., 2016). Different fungal photoreceptors respond to different light
signals across a wide range of wavelengths. Four types of different light receptors have
been identied: blue light, UV light, green, and red- light receptors, which are capable
of sensing short, medium, and long wavelengths of light (Bayram & Bayram, 2023;
Wang et al., 2016). Currently, the following types of blue light receptor families have
been established: White Collars, Vivid, Cryptochrome, and Photolyase (Bayram &
Bayram, 2023; Corrochano, 2019). White Collar (WC) proteins are the most studied
fungal light receptors. The second group of blue light photoreceptors is Vivid protein
(VVD), which is not widely distributed in the fungal kingdom. The third group of blue
light receptors in fungi are proteins of the cryptochrome family, photolyase, which were
discovered only in the 90s of the twentieth century. Their function is still poorly understood compared to WC and VVD.
The initial breakthrough in the molecular understanding of phototransduction in
fungi occurred when the White Collar- 1 (WC- 1) gene was cloned by chromosomal
movement and complementation of mutants (Ballario et al., 1996). WC- 1 is a zinc
nger transcription factor that binds to the promoters of light- regulated genes. The
second White Collar Protein (WC- 2) is also a GATA- like zinc nger transcription factor
(Herrera- Estrella & Horwitz, 2007).
Proteins similar to WC- 1 and WC- 2 have been identied in the genomes of asco- ,
basidio- , and zygomycetes, many of which are necessary to explain the photoresponse
of fungi to light. It has been suggested that WC complexes arose at the early stages
of fungal evolution to regulate their photoresponse, as photoreceptors and transcription factors. Isolation and characterization of photoreceptors in various macromycete
species are based on the previously identied WC- 1 and WC- 2 genes of Neurospora
crassa (Linden & Macino, 1997; Talora et al., 1999).
Ambra et al. (2004) studied the photomorphogenesis of the ascomycete Tuber
borchii. The TbWC- 1 gene, homologous to the blue light photoreceptor in Neurospora
crassa NcWC- 1 (White Collar- 1), which consists of a light- oxygen- voltage (LOV)
domain and a transcription factor fragment, was identied and cloned. The amino acid
sequence of the gene (TbWC- 1) showed high domain similarity with NcWC- 1, except
for the polyglutamine regions. It has been established that TbWC- 1 mRNA is controlled
by blue light, and its steady- state level increases upon irradiation. In silico analysis of
the TbWC- 1 sensor domain (LOV) supports the hypothesis that TbWC- 1 is a blue light
photoreceptor (Ambra et al., 2004). Also, genes responsible for blue light photoreceptors
have been identied in the basidiomycetes Coprinopsis cinereus and Lentinus edodes.
Terashima et al. (2005) studied the effect of light on the formation of fruiting bodies
of the basidiomycete Coprinopsis cinereus. As a result, the Dst1 gene was cloned, which
is a 1,175 amino acid protein containing two PAS (Per- Arnt- Sim) domains, a helical
structure, and a putative glutamine- rich transcriptional activation domain (AD). One of
the PAS domains showed signicant similarity to LOV domains, which are known blue
light receptors, suggesting that Dst1 is a blue light receptor in C. cinereus (Terashima
et al., 2005).
Later, Sano et al. (2007) cloned and sequenced the photoreceptor gene (Le.phrA) of
the basidiomycete Lentinula edodes. The blue light photoreceptor Le.phrA was found
to have 924 amino acid residues, and contain a serine- rich region, a LOV domain, and
two PAS (Per- Arnt- Sim) domains. It was smaller than other LOV domain- containing
Regulation of the Biological Activity of Medicinal Macromycetes 405
fungal blue light photoreceptors such as Coprinopsis cinerea Dst1, and Neurospora
crassa WC- 1 (1,167 aa). It was found that the Le.phrA gene is transcribed at all stages
of forming the fruiting body of L. edodes, but it was most actively transcribed in the
immature fruiting body (Sano et al., 2007).
Kanda et al. (2007) identied photoregulated genes that control enzymatic activity,
in particular tyrosinases of such macromycete species as Polyporus arcularius, Lentinula
edodes, and Agaricus bisporus (Kanda et al., 2007). Nakano et al. (2010) rst identi -
ed 15 genes with increased and 13 with decreased regulation of the photoresponse to
blue light in the mycelium of the fungus Pleurotus ostreatus (Nakano et al., 2010). In
addi tion, most fungal genomes contain genes for other photoreceptors. Sensitivity to red
light is realized in fungi by phytochrome, a molecule that until recently was considered
unique to plants (Rodriguez- Romero et al., 2010; Wang et al., 2016). This type of
photoreceptor, which is responsible for red and far- red light, was rst discovered in two
model objects Aspergillus nidulans and Neurospora crassa. Phytochromes are large
proteins containing several domains for light reception and signal transmission. One of
the striking features of phytochromes is that they are not found in vertebrates, but are
specic to plants, bacteria, and fungi (Bayram & Bayram, 2023; Navarro et al., 2013).
Retinal- based opsin systems have recently been discovered, the biological functions
of which still require study. Opsins are membrane- bound proteins with seven transmembrane helices that act as green light receptors (Bayram & Bayram, 2023; Yu &
Fischer, 2019). They are the main components of vision in vertebrates, including the
human eye. However, their role in fungal photoreception has not been fully studied.
Based on spectral sensitivity analysis, it was suggested that the photoreceptors of
Schizochyllum commune (Yli- Mattila, 1985) and Coprinellus congregatus contain a
compound related to avin or a pteridine (Durand & Furuya, 1985), which were also
found in Pleurotus ostreatus (Richartz & Maclellan, 1987). It has been suggested that
in Favolus arcularius the possible photoreceptor pigments may be neither carotenoids
nor avonoids (Kitamoto et al., 1972). However, in macromycete species other than
Coprinopsis cinerea, biochemical and/ or molecular studies of putative photoreceptors
have yet to be performed.
Opsin systems at fungal have been described in ascomycetes and basidiomycetes,
and sequence similarities have allowed three groups of opsins to be distinguished: the
rst group of opsin family proteins act as light- dependent proton pumps (Leptosphaeria
maculans); a second group similar to the N. crassa sensory opsin, which is expected
to be slow- cycling photosensors; the third group (opsin- like proteins) includes the socalled accessory forms of opsin, present in many fungal species (Bayram & Bayram,
2023; Rodriguez- Romero et al., 2010). However, the issue of the light- regulating role of
opsins in fungi and the interaction of opsins with other light- sensitive systems remains
insufciently studied.
It is now established that fungi can sense near- ultraviolet, blue, green, red, and
far- red light using up to 11 photoreceptors and signaling cascades to control a large
portion of the genome and thereby adapt to environmental conditions (Yu & Fischer,
2019). Regarding the time of manifestation of photosensitivity of vegetative mycelium,
two hypotheses were put forward and tested: photoinduction manifests itself when the
mycelium becomes physiologically mature; photoinduction begins when space and
nutrition resources are exhausted (Poyedinok, 2013). As a result of the research, the
Соседние файлы в папке Библиотека им академика М.И. Перельмана