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416 Wild Edible Plants
TABLE 14.3 Influence of the cultivation method on photo- induced changes in the
growth of macromycetes
632.8 NM
SPECIES
Cordyceps
militaris
LASER
Linear growth on agar medium, % increase in growth rate
10.3±0.1a5.1±0.2 0 0 13.8±0.6a9.5±0.2
Surface growth in liquid medium, % increase in biomass
6.1±0.3 5.9±0.2
Growth during deep cultivation, % increase in biomass
19.5±0.5a11.6±0.2
Flammulina
velutipes
Linear growth on agar medium, % increase in growth rate
13.8±0.2a9.5±0.3
Surface growth in liquid medium, % increase in biomass
25.3±0.29 19.6±1.6
Growth during submerged cultivation, % increase in biomass
56.2±1.8a36.3±0.7
Ganoderma
lucidum
Linear growth on agar medium, % increase in growth rate
7.8±0.7
Surface growth in liquid medium, % increase in biomass
25.8±1.2a17.9±0.6
Growth during submerged cultivation, % increase in biomass
66.7±1.4a41.7±1.1
Hericium
erinaceus
Linear growth on agar medium, % increase in growth rate
37.5±1.1a 18.7±0.9
Surface growth in liquid medium, % increase in biomass
39.2±1.8a29.7±0.18 32.2±1.1a20.3±0.8a45,5±5.2a33.3±1.8
Growth during submerged cultivation, % increase in biomass
51.9±3.3a36.5±4.2
Inonotus
obliquus
Linear growth on agar medium, % increase in growth rate
11.3±0.2a10.7±0.1
Surface growth in liquid medium, % increase in biomass
12.6±0.8a7.9±0.3
Growth during submerged cultivation, % increase in biomass
17.2±0.8a9.7±0.4
Lentinula
edodes
Linear growth on agar medium, % increase in growth rate
0 0 3.5±0.6 2.2±0.6 12.5±1.0a1.2±0.2
Surface growth in liquid medium, % increase in biomass
75.0±1.7 28.9±1.4
Growth during submerged cultivation, % increase in biomass
139.9±2.5a93.4±0.8
Source: Poyedinok et al. (2013).
Note: «a» – statistically significant differences relative to control are indicated (p ≤ 0.05). Data are
the mean±standard error for three replications.
a
625.0 NM
INCOHERENT
a
a
a
a
a
a
6.7±0.7
a
a
a
a
a
a
a
a
a
514.5 NM
LASER
1.6±0.2 0 15.3±0.3a9.3±1.1
7.8±0.2
522.0 NM
INCOHERENT
a
1.6±0.4 25.9±1.21a11.9±1.18
488.0 NM
LASER
463.0 NM
INCOHERENT
a
a
0 0 12.6±0.3a10.3±0.3
1.1±0.2 0 34.2±1.9a21.0±2.5
– 11.6±1.0a0 78.1±1.1a44.2±4.4
– 1.9±0.1 1.6±0.1 10.4±1.0a9.6±0.7
10.8±0.8a6,0±0,3
a
36.5±1.4a8.5±0,8
a
a
34.9±0.8a26.0±0.8a75.3±0.6a38.7±0.6
32.9±0.7a20.0±1.3a29,6±2,4a21.1±2.8
43.1±1.8a31.0±2.3a68.9±2.8a43,5±1.8
1.1±0.2 – 0.7±0.1 26.6±1.4a22.3±0.78
1.3±0.1 1.8±0.1 36.6±1.8a21.03±0.7
2.2±0.1
a
5.3±0.2
a
56.9±2.0a23.6±0.9
66.8±2.0a20.3±1.3a76.8±1.9a45.8±1.3
117.5±2.3a87.7±1.7a153.9±3.0a104.2±3.3
a
a
a
a
a
a
a
a
a
a
a
a

Regulation of the Biological Activity of Medicinal Macromycetes 417
light. There is no signicant difference in the growth rate of irradiated mycelium on agar
medium from the control was not found in L. edodes, under all irradiation modes. In
C. militaris, F. velutipes, G. lucidum, and I. obliquus there were no differences in linear
growth rate about to control when irradiated with green light. In this case, the absence
of signicant differences between experiment and control for some macromycetes
when growing on agar media can be explained by the difference in their morphological
changes in response to light inuences. Therefore, to obtain a reliable picture, in our
opinion, it is advisable to compare growth rates in liquid media at different cultivation
methods.
Analysis of surface growth of macromycete mycelium on the liquid medium by
determining the biomass of the grown mycelium showed reliable differences in this
indicator under the same light exposure regimes according to attitude towards control.
To obtain the greatest stimulating effect from irradiation submerged cultivation turned
out to be the most preferable. Increased biomass accumulation during dynamic cultivation mode varied depending on the species and irradiation mode from 12 to more
than 154% compared to cultivation on the same medium in stationary mode. Thus,
we have established that the method of cultivating irradiated strains of macromycetes
to a signicant extent inuences the degree of photostimulation of growth processes.
Also, the period preceding fruiting is reduced, and fruiting time during solid- phase
cultivation. At the same time, there is a signicant increase in the yield of fruiting
bodies and an improvement in their quality. Low- intensity laser light in the visible
part of the spectrum was used in submerged cultivation biotechnology not only as
a growth stimulator but also for the synthesis of biologically active substances ‒
polysaccharides, melanins, and antibiotics (Poyedinok, 2013; Poyedinok et al., 2013,
Poyedinok et al., 2015b).
Our results conrm that low- intensity light in the visible part of the spectrum can
be used in the biotechnology of submerged cultivation of Inonotus obliquus not only as
a growth stimulator but also as an inducer of melanin synthesis (Figure 14.2).
FIGURE 14.2 Effect of low- intensity light on the synthesis of melanin by medicinal
mushroom Inonotus obliquus. A ‒ control without irradiation, B ‒ irradiation with blue laser
light (488 nm).

418 Wild Edible Plants
In our research, the irradiation of mycelial with blue laser light (488 nm)
increased the amount of melanin synthesized by the culture of I. obliquus during
submerged cultivation. The maximum stimulation of growth, which was expressed in
an increase in the accumulation of mycelium biomass, blue coherent light ‒ 56.9%.
At the same time, the amount of endomelanin increased by 250.0%. Analysis of
the dynamics of melanin accumulation by the culture of I. obliquus showed that
its synthesis reaches the stationary phase after exposure to laser light with a high
intensity of 488 nm on the 9th day of cultivation, whereas irradiation red laser light
(632.8 nm) on the 12th day of cultivation, the transition to the stationary phase was
not noted. Thus, the cultivation of the I. obliquus under the indicated regime allowed
us to signicantly increase the yield of melanin, and to speed up the cultivation time
(Poyedinok et al., 2019).
An analysis of the experience of using articial light to stimulate biological
processes in plant growing and mushroom growing showed that it is limited to methods
based on constant illumination of crops at different stages of morphogenesis (Arjona
et al., 2009; Huang et al., 2017; Namba et al., 2002). However, this requires the instal lation of special lighting over large areas, additional energy costs, and maintenance.
Whereas studies conducted on cells of various biological objects have shown that shortterm irradiation (from fractions of a second to tens of minutes) with low- intensity light
in relatively small doses (102‒103 J/ m2) promotes a macro- effect that lasts for a long
time (Karu, 1996). Conducting similar studies on photointensication of technological
stages of surface cultivation of macromycetes on solid substrates by short- term irradiation of seed material with low- intensity light using the example of Pleurotus ostreatus,
Lentinula edodes, Flammulina velutipes, Hericium erinaceus, and Cordyceps militaris,
which have a number of advantages compared to previously known (Miyazaki et al.,
2011; Myoung- Jun, 2011; Namba et al., 2002), showed the promise of using new
methods in biotechnology (Figures 14.3 and 14.4).
Using seed material activated by short- term irradiation with low- intensity laser light
made it possible to reduce the fouling of L. edodes substrate blocks by 20 days when
using low- intensity light in the red and green wavelength ranges and by 30 days when
using blue light. Activation of the seed mycelium of P. ostreatus in all irradiation modes,
and F. velutipes in the blue wavelength range and red (laser), made it possible to reduce
the incubation time until complete fouling of the substrate by 5 days (Poyedinok, 2013).
Irradiation of H. erinaceus seed material with red and blue light of different coherence made it possible to shorten the specied cultivation stage by 10 days (Poyedinok
et al., 2000). While studying the formation process of fruiting in these macromycetes,
an earlier onset of fruiting was established on substrates inoculated with grain mycelium activated in various irradiation modes. In L. edodes, fruiting began 35‒40 days
earlier when blue light was used to stimulate growth processes, 30 days earlier when
using red light, and 10‒15 days earlier when using green light. In P. ostreatus, the
period before the onset of fruiting, depending on the mode of activation of the seed, was
reduced by 4‒6 days, in H. erinaceus ‒ by 11‒15 days, and in F. velutipes by 4‒12 days
(Poyedinok, 2013).
Thus, literature data and our research prove the promise of using articial laser
light in biotechnologies for cultivating edible and medicinal mushrooms.

Regulation of the Biological Activity of Medicinal Macromycetes 419
FIGURE 14.3 The influence of low- intensity laser light on the growth and formation of
fruiting bodies of edible and medicinal mushrooms: Pleurotus ostreatus (A); Flammulina
velutipes (B); Fomitopsis betulina (C). 1, control without irradiation; 2, irradiation with blue
laser light (488 nm); 3, irradiation with green laser light (530 nm); 4, irradiation with red
laser light (650 nm).

420 Wild Edible Plants
FIGURE 14.4 The influence of low- intensity laser light on the growth and fruiting of
edible and medicinal mushrooms: Cordyceps militari cultivation on a nutrient medium
glucose- peptone- yeast agar (A); C. militaris cultivation on a liquid nutrient medium glucose-
peptone- yeast (B); Flammulina velutipes cultivation on a plant substrate (C). 1, control
without irradiation; 2, irradiation with blue laser light 488 nm.

Regulation of the Biological Activity of Medicinal Macromycetes 421
14.6 CONCLUSIONS
The results of the analysis of literary data conrm the signicant inuence of light
on the morphogenesis and metabolism of edible and medicinal mushrooms. Although
photoreceptors for red, green, and blue light have been identied in mushrooms, the
molecular mechanisms of response to blue light are the most studied to date. The study
of photobiological reactions, as well as the collected experimental material on the
stimulating effect of low- intensity light on the biosynthesis of edible and medicinal
mushrooms, contribute to a deeper understanding of the basic mechanisms of the effect
of light on these organisms. Particular attention should be paid to the experimental
assessment of the role of light as an environmental factor in the process of cultivating
edible and medicinal mushrooms to obtain products for both food and pharmacological needs.
Thanks to progress in the development of LED and laser technology, it has become
possible to use certain parts of the optical spectrum with precise control of the intensity and polarization of radiation when growing edible and medicinal mushrooms.
Experimental studies have established effective modes of using LED and laser light
sources to induce spore germination, synthesis of mycelial mass and various metabolites,
morphogenesis, and fruiting in the cultivation of biotechnologically important species
of mushrooms.
In the context of global sustainability efforts, edible and medicinal mushrooms are
emerging as valuable sources of food and nutraceuticals and promising producers of
natural compounds with diverse pharmacological activities. However, realizing their
full potential requires a deeper understanding of how various factors, including articial
low- intensity light, affect their morphogenesis and metabolic processes.
Studying the complex relationship between light and mushroom physiology not
only stimulates various steps in biotechnological processes but also agricultural innovation, which is important for responsible resource management and environmental protection. Continued research into light signaling mechanisms in mushrooms is essential
to promoting sustainable agricultural practices and ensuring the long- term viability of
our planet’s ecosystems.
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