Ultra-Weak Photon Emission in Plants

High-Sensitivity Imaging and Modeling of Ultra-Weak

Photon Emission in Plants Under Stress
Yan-Xia Liu1, Hai-Yu Fan1, Yu-Hao Wang1, Yan-Liang Wang2, Sheng-Wen Li1 *, Shi-Jian
Li3, Xu-Ri Yao1, *, Qing Zhao1, *

1
Center for Quantum Technology Research, School of Physics, Beijing Institute of Technology, Beijing 100081, China

2
The Germplasm Bank of Wild Species & Yunnan Key Laboratory for Fungal Diversity and Green Development, Kunming Institute of

Botany, Chinese Academy of Sciences, Kunming, Yunnan 650201, China

3
School of Integrated Circuits and Electronics, Beijing Institute of Technology, 100081 Beijing, China

*Author for correspondence: lishengwen@bit.edu.cn (Sheng-Wen Li), yaoxuri@bit.edu.cn (Xu-Ri Yao), qzhaoyuping@bit.edu.cn (Qing

Zhao)

Short title: Ultra-Weak Photon Emission in Plants

Responsible Author for Material Distribution:

The author responsible for distribution of materials integral to the findings presented in this article in accordance with the policy

described in the Instructions for Authors (https://academic.oup.com/plphys/pages/General-Instructions) is Qing Zhao.

Abstract

Ultra-weak photon emission (UPE) is a noninvasive diagnostic tool that effectively

reflects the function and health status of plant cells. However, current UPE measurement
techniques are limited by resolution and sensitivity, particularly when monitoring different
plant species and stress types. This study analyzes the delayed luminescence (DL) properties
of Hydrocotyle vulgaris, Arabidopsis leaves, and Ginkgo leaves under both stress and control
conditions using an independently developed UPE imaging system. The results showed a
significant increase in initial DL intensity and an accelerated oxidative metabolic rate under
mechanical injury and oxidative stress. DL decay characteristics were significantly correlated
with the plant’s physiological state, with stress conditions exhibiting decay curves that
closely matched theoretical models. These findings confirm the established correlation
between DL and plant stress responses. The high-resolution, low-noise imaging system
significantly improves the accuracy of plant physiological state monitoring and provides new
insights into the potential of optical signals for non-chemical communication research and
agricultural applications. This technology has great potential for monitoring plant growth,
assessing environmental stress, and supporting precision agriculture.
Keywords ultra-weak photon emission, delayed luminescence, oxidative stress, reactive
oxygen species, plant stress monitoring

Introduction

Ultra-weak photon emission (UPE), an inherent property of all living systems, has been
observed across a diverse range of organisms, including animals, plants, and microorganisms.
It is defined as the spontaneous emission of a weak photon stream by organisms during
metabolic processes. The intensity of this emission typically ranges from a few photons to a
few hundred per square centimeter and wavelengths between 180−800 nm. This emission
can be considered a reflection of the organism’s local energy state[1].
In eukaryotic cells, the primary sources of UPE are the excited-state carbonyl (3R=O·)
or singlet oxygen (1O2) generated during oxidation, which serves as the ultimate photon
emitters. The mechanism of their formation is well known[2]. During normal oxidative
1
 Ultra-Weak Photon Emission in Plants

metabolism, organisms produce small amounts of reactive oxygen species (ROS), which are
essential precursors of UPE. NADPH oxidase is a multi-subunit membrane protein complex
typically found in cellular membranes, including the inner membranes of mitochondria and
chloroplasts. The enzyme acts as an electron donor and facilitates the electron transfer
reaction between a substrate and molecular oxygen (O₂), thus forming the superoxide anion
(O₂⁻). Subsequently, the action of enzymes, such as superoxide dismutase, facilitates the rapid
conversion of O₂⁻ into hydrogen peroxide (H₂O₂). H₂O₂ can undergo further reactions to
produce the highly reactive hydroxyl radical (HO·). HO· then reacts with proteins, lipids,
[3]
nucleic acids, and other biomolecules in organisms to form alkyl radicals (R·) , which
subsequently combine with O₂ to form organic peroxyl radicals (ROO·). ROO· then reacts
with other organic substances to form organic peroxides (ROOH) and R·. The cyclization or
recombination of ROO· can produce unstable, energetic intermediates, such as dioxetane
(ROOR) and tetroxide (ROOOOR) [4-5]. These unstable intermediates decompose into triplet-
excited carbonyls (3R=O·) and ground state carbonyls (R=O). The energy transfer from
3
R=O· to pigments or O₂ results in the formation of the excited-state pigments (3P· or 1P·)
and 1O2. These excited states release excess energy in the form of photons when they return
to a lower energy state[6]. The energy of the photons emitted by various radiators corresponds
to certain wavelength ranges in the electromagnetic spectrum. The emission wavelengths of
3
R=O· are between 350−550 nm, while those of the pigments are between 550−750 nm (1P·)
and 750−1000 nm (3P·). A simplified diagram of the above reaction mechanism is presented
in Figure 1.

Figure 1. (a) Schematic representation of the UPE mechanism. NADPH oxidase is located within the

2
 Ultra-Weak Photon Emission in Plants

mitochondrial or chloroplast membranes, where it catalyzes the reaction between a substrate and molecular

oxygen, resulting in the production of the O₂⁻, which is subsequently converted to H₂O₂ and HO·. HO· can

then oxidize biomolecules. These unstable intermediates have high energy levels, as exemplified by

ROOR and ROOOOR. They decompose into 3R=O· and R=O. 3R=O· then transfers energy to pigments

or O2 leading to the production of excited-state pigments (3P· or 1P·) and 1O2. The reaction scheme is

based on[2]. (b) UPE electronically excited species. 3R=O· emission wavelengths are located in the near-

ultraviolet and blue-green regions, while the excited state pigments are located in the green-red (1P·) and

near-infrared (3P·) regions. 1O2 is in the red and near-infrared regions. The solid arrows indicate the energy

level transitions from the excited state to the ground state, while the dashed arrows indicate the
3R=O
· transitions and the energy transfer between the pigments and O2.
ROS concentrations during plant growth and metabolism are closely associated with
UPE. Low ROS levels signal external stimuli, regulate plant growth and development, and
work in synergy with other defense mechanisms to enhance plant resistance[7]. However,
when ROS levels remain persistently elevated, an oxidative stress response is triggered,
impairing the plant’s antioxidant system and causing oxidative damage to tissues[8]. ROS
accumulation disrupts the balance between production and scavenging, resulting in increased
UPE intensity. The existing literature indicates that different stress conditions significantly
affect plant UPE in varying ways. For example, in Arabidopsis, UPE increases and persists
for several hours after mechanical damage, accompanied by changes in the spectral pattern
of photon emission[9]. The endogenous photon emission intensity of Arabidopsis seeds
significantly increases during the early stages of seed germination, and this increase depends
on the dose of locally applied H₂O₂[10]. In sunflower plants subjected to biotic and abiotic
stresses, a significant decrease in photon emission was observed in healthy plants under
controlled growth conditions. In contrast, a marked increase was observed in stressed plants
under water stress[11]. Peanut and rice seedlings exhibited the most pronounced UPE when
exposed to white light, followed by red and yellow lights. Conversely, the lowest
luminescence intensity was observed under blue light. Under natural light conditions, UPE
exhibited the slowest decay, while all other monochromatic lights exhibited faster decay rates
than that of natural light[12]. These results indicate that UPE, as a macroscopic manifestation
of microscopic life activities, is highly sensitive to internal changes in plants and external

3
 Ultra-Weak Photon Emission in Plants

factors. Compared to traditional destructive detection methods (e.g., analysis of physiological

and biochemical indicators), UPE is a noninvasive diagnostic tool that can be used to assess
the physiological status of plants.
UPE can be divided into two main categories: spontaneous luminescence and
exogenously induced luminescence. In the latter category, light-induced UPE is referred to
as delayed luminescence (DL). DL refers to the emission of weak light radiation following
the transition of a biological sample from a photoexcited state to a ground state after
irradiation with white or monochromatic light. Compared to spontaneous luminescence, DL
has a higher signal-to-noise ratio and luminescence intensity, with peaks reaching hundreds
of thousands of photons per square centimeter. The DL phenomenon is closely related to
various fundamental biological processes, including metabolic activities, growth and
development, genetic variation, and intercellular information transfer. Consequently, it is a
crucial and sensitive physical indicator for assessing the functionality and health status of
biological cells[13-19]. DL is closely associated with the excitation of coherent electron and
exciton states in biological macromolecules, a conclusion that is supported by experimental
evidence[20-22]. The advantages of DL—high sensitivity, convenient operation, rapid response,
and nondestructive detection—have led to its widespread use in various fields. These fields
include the differentiation of normal cells from tumor cells[23], the determination of seed
germination[24,25], the identification of herbal medicines[26-29], food safety assessment[30], and
fruit ripening[31]. The results of these studies demonstrate that changes in DL parameters can
effectively reflect the physiological state of organisms, thus providing new insights and
avenues for the advancement of bioanalytical techniques based on changes in DL. Despite
some progress in the study of DL in plants, the field still presents significant challenges.
Current techniques do not overcome the problems of weak plant DL signals, interference
from background noise, and variability across different plant species and environmental
conditions. Additionally, the underlying mechanism of DL remains poorly understood, and
the relationship between light signal attenuation and physiological changes in plants in
response to environmental stress has not been fully elucidated. There is considerable scope
for improving the existing technology, particularly in terms of imaging resolution, signal
sensitivity, and data processing accuracy.

4
 Ultra-Weak Photon Emission in Plants

In light of the above considerations, this study proposes the implementation of a light-
induced methodology for the systematic observation of DL in Hydrocotyle vulgaris,
Arabidopsis and Ginkgo leaves under natural growth conditions, considering different stress
factors. The methodology involves using a highly sensitive two-dimensional photon imaging
technique, which enables effective monitoring of oxidative metabolic processes in plants and
their responses to oxidative stress. Additionally, a comprehensive investigation of the DL
decay kinetics in plant leaves will be conducted to elucidate the relationship between changes
in DL parameters and plant physiological states. This study aims to provide a new basis for
understanding the quantum effects and the physical mechanisms of UPE.

Results

DL of three plant species in their natural state

Arabidopsis plants and leaves, H. vulgaris leaves, and Ginkgo leaves were imaged in
their natural state, as shown in Figure 2(a). Figure 2(b) presents the trend of DL intensity over
time. After multiple exposures, the DL intensity of all three plants exhibited a nonlinear decay
trend, characterized by an initial high intensity followed by a rapid decline and subsequent
stabilization. This observation suggests that by the end of the measurement period, the plants’
DL had almost ceased and had entered a stable spontaneous luminescence phase.
The DL decay differed significantly between the Arabidopsis plant and its leaves. The
plant had a higher initial intensity (0.2736) than the leaves (0.2258) and stabilized earlier. In
contrast, the leaves showed greater fluctuation, with a notable decrease at t = 120s, slight
recovery, and then stabilization, reflecting different metabolic and physiological responses.
The DL ability also varied between plant species. Arabidopsis leaves had the weakest
and shortest luminescence, with a rapid decay. H. vulgaris had a higher initial intensity
(0.3037) but decayed quickly, with a brief luminescence duration. Ginkgo leaves had the
strongest and most persistent delayed luminescence, with the highest initial intensity (0.5353)
and a slower decay rate, maintaining a high intensity for a longer period.

5
 Ultra-Weak Photon Emission in Plants

a Sample 3 s s s 12 s 1 s1 s b

Arabidopsis
plant

.2 3 . 3 . 3 . . 2 . 2

Arabidopsis
leaf

.22 . . 3 . 23 . 3 3 . 32

ydrocotyle
ulgarist

.3 3 . . 3 . 2 1 . 1 . 1

in go leaf

. 3 3 .1 2 . 3 . . 3 . 2

Figure 2. (a) DL imaging results of Arabidopsis (plants and leaves), H. vulgaris leaves, and Ginkgo leaves

in the natural state. Sample sizes are presented in front of the qCMOS camera. The exposure time was 30

s, and the total measurement time was 180 seconds. The red data represents the DL intensity. The image

size was 230 × 230 pixels. Subsequent figures follow a similar pattern. (b) The trend of DL intensity over

time.

DL of three plant species under stress

Effects of mechanical injury

The DL Imaging System monitored the changes in DL intensity of H. vulgaris,

Arabidopsis, and Ginkgo leaves after mechanical injury. Figure 3(a) illustrates the imaging
results of leaves from the three plants under control and injury conditions, with the red line
denoting the injury site. The data for the first 30 seconds are shown in the figure, while the
data for the first 90 seconds can be found in the Supplementary Figure S1. 3D waterfall
plots of DL intensity and its decay for three plants at different injury times are shown in
Supplementary Figure S2. Figure 3(b) presents a histogram of the difference in DL intensity
between the mechanically injured group and the control groups over time, demonstrating the
effect of mechanical injury on the DL. The fitting results (Figure 3(c-e)) indicated that the DL
decay of H. vulgaris and Ginkgo leaves followed the exponential model (Eq.(5)) and
Arabidopsis conformed to the polynomial model (Eq.(6)). The detailed regression
coefficients (a), slope parameters (b), and fitting coefficients (R²) are given in Table 1 and
Table 2, respectively.

6
 Ultra-Weak Photon Emission in Plants

In monitoring DL intensity imaging in different plants, we observed significant

differences in their sensitivity and response time to mechanical injury. H. vulgaris showed a
rapid and strong response within 5 min, with a marked increase in DL intensity to 0.6716,
approximately 1.95 times that of the control group. Although the response diminished over
time, it remained elevated, indicating a sensitive and prolonged response to mechanical injury.
In contrast, Arabidopsis showed a more moderate response, with a smaller increase in DL
intensity and less fluctuation, although the intensity gradually increased after injury. Ginkgo
leaves showed a more stable response, with DL intensity increasing to 0.6693 within 5 min,
approximately 1.56 times that of the control, and remaining at a high level for the remainder
of the time period. These results highlight differences in the adaptive mechanisms of different
plants in response to mechanical injury, with H. vulgaris and Ginkgo leaves showing more
intense and sustained responses, whereas Arabidopsis showed relatively milder and less
fluctuating responses.
The fitting results also indicated that the injury groups generally had higher initial
intensities and faster decay rates, reflecting the rapid response mechanism that plants employ
following mechanical damage. The R² values for all groups were close to 1, indicating an
excellent model fit. Notably, the injury group exhibited a DL decay pattern that was highly
consistent with the fitted model, indicating that the response of plants to mechanical injury
follows a predictable pattern. In Arabidopsis, the injury group had a larger coefficient for the
quadratic term, suggesting that its nonlinear decay component was more pronounced. The
higher-order terms (A3 and A4) were negative in both the control and injury groups, with
larger absolute values in the injury group, indicating greater volatility and more substantial
decay in the higher stages of decay. In summary, the injury groups of all three plants showed
greater consistency in initial intensity, decay rate, and model fit, underscoring the significant
influence of mechanical injury on the DL decay pattern in plants.

7
 Ultra-Weak Photon Emission in Plants

a Sample min min 1 min 1 min 2 min 2 min 3 min b

ydrocotyle
ulgarist

.3 . 1 . 2 . 3 . 2 2 . 1 . 22

Arabidopsis
leaf

.2 .2 .2 .2 3 .2 13 .321 .2 2

in go leaf

. 2 . 3 . . 1 2 . . . 3

c d ydrocotyle
e
Arabidopsis
ulgarist in go leaf
leaf

Figure 3. Effect of mechanical injury on plant DL. (a) Initial intensity (t = 30 s) imaging results of DL

after mechanical injury in three plants. 0 min indicates the control. Time represents the duration of

mechanical injury, which was monitored every 5 min for 30 min Samples present the true state of the

leaves before the experiment, with the red line indicating the location of the mechanical injury. (b) D-value

in DL intensity in the stress and control groups. The error bar comes from the average of three

measurements. (c-e) Examples of the fit of DL control and stress groups decaying over time for H. vulgaris,

Arabidopsis, and Ginkgo leaves. The data comes from the average of three measurements.

Effects of oxidati e stress

The leaves of H. vulgaris, Arabidopsis, and Ginkgo were treated with a 3% H2O2
solution, and the response was monitored using a DL imaging system. The resulting images
are presented in Figure (a), with 0 min representing the untreated control. Data for the first
30 seconds are shown in the figure, while the data for the first 90 seconds can be found in the
Supplementary Figure S3. 3D waterfall plots of DL intensity and its decay for three plants
at different oxidative stress times are shown in Supplementary Figure S . Bar graphs were
constructed to demonstrate the changes in DL intensity between the oxidative stress group
and the control group (Figure (b)). Figure (c-e) presents the fitting of the intensity decay for
the three plants during DL. The detailed regression coefficients (a), slope parameters (b), and

8
 Ultra-Weak Photon Emission in Plants

fitting coefficients (R²) are given in Supplementary Table 1 and Supplementary Table 2,
respectively.
Monitoring and imaging the initial DL intensity of the three plants’ leaves under
oxidative stress over time revealed that different plants exhibited different response
mechanisms. During the initial 5 min, the DL intensity of H. vulgaris decreased by 13.0%
compared to the control group. However, after 30 min, it returned to a higher level, indicating
a robust antioxidant capacity. Arabidopsis displayed a more sensitive response to oxidative
stress, with a sustained decrease in DL intensity. At the 5-minute mark, its DL intensity
decreased to 0.2375, representing a 24.3% reduction. Subsequently, the DL intensity
decreased further to 0.2486 after 30 min, representing a total change of 27.4%. In contrast,
the Ginkgo leaves displayed a remarkable ability to adapt to oxidative stress. Within the first
5 min, the DL intensity increased from 0.4555 to 0.4979, a change of 9.3%. After 30 min, the
DL intensity continued to increase and reached 0.6546, an increase of 43.7%. These results
indicate that the Ginkgo leaves could rapidly activate their antioxidant response mechanism,
resulting in increased DL intensity. The results revealed that the three plants had different
response mechanisms when exposed to oxidative stress, and applying H2O2 facilitated
changes in the DL intensity of the plants.
Data fitting results indicated that the decay of DL in the three plants under oxidative
stress was more consistent with the fitted model, suggesting that oxidative stress had a
significant impact on the plants' DL characteristics. Oxidative stress significantly increased
the initial DL intensity in both H. vulgaris and Ginkgo leaves, with Ginkgo leaves exhibiting
an initial intensity nearly three times higher than that of the control group, indicating a
stronger response to oxidative stress. Regarding the decay rate, H. vulgaris showed a slight
increase with no significant change, while Ginkgo leaves showed a significantly faster decay
rate, reflecting their stronger adaptive capacity. In contrast, Arabidopsis showed a faster
decay rate, indicating a weaker ability to cope with oxidative stress. These results highlight
the different response mechanisms of the three plants under oxidative stress and underline
the high sensitivity of DL imaging as an effective tool for monitoring oxidative responses in
plants.

9
 Ultra-Weak Photon Emission in Plants

a Sample min min 1 min 1 min 2 min 2 min 3 min b

ydrocotyle
ulgarist

.3 .3 2 .331 .3 2 .3 . 1 . 22

Arabidopsis
leaf

.313 .23 .2 .2 .2 .233 .2

in go leaf

. . . . 11 .3 . .

c d ydrocotyle
e
Arabidopsis
ulgarist in go leaf
leaf

Figure . Effect of oxidative stress on plant DL. (a) Initial intensity (t = 30 s) imaging results of DL after

oxidative stress in three plants. 0 min indicates the control. Time represents the duration of oxidative stress,

which was monitored every 5 min for 30 min. Samples present the true state of the leaves before the

experiment. (b) D-value in DL intensity in the stress and control groups. The error bar comes from the

average of three measurements. (c-e) Examples of the fit of DL control and stress groups decaying over

time for H. vulgaris, Arabidopsis, and Ginkgo leaves. The data comes from the average of three

measurements.

Effect of different light qualities

Figure (a) presents the results of the DL imaging of plant leaves under white, red, and

blue lights. Figure (b) and Figure (c) compare the effects of different light qualities on the
DL of the same plant and an analysis of the DL intensity of the three plants under different
light qualities at the same exposure time. Figure (d-f) presents an example of the fitting of
the DL decay with time for the three plants. The fitting results demonstrate that the decay
patterns of H. vulgaris and Ginkgo leaves were consistent with the exponential model (Eq.
(5)), whereas that of Arabidopsis displayed a polynomial decay pattern (Eq. (6)). The detailed
regression coefficients (a), slope parameters (b), and fit coefficients (R²) are presented in
Supplementary Table 3 and Supplementary Table .

10
 Ultra-Weak Photon Emission in Plants

Different light qualities significantly influenced the initial intensity and decay rate of
DL in the leaves of the three plant species, with both common patterns and individual
differences in their responses. Under white light, the initial luminescence intensity was
highest and the decay rate was slow for all three plants. In contrast, under blue light, the
intensity was the lowest and the decay rate was the fastest, particularly in H. vulgaris and
Ginkgo leaves. Under red light, the DL intensity fell between that of white and blue light,
and the decay rate was more moderate. These findings suggest that plant responses to
different wavelengths of light vary significantly, particularly under white light conditions.
The observed differences among plants' responses were related to their
photoacclimation mechanisms. Under white light, Ginkgo leaves exhibited the highest initial
luminescence intensity, followed by H. vulgaris, with Arabidopsis showing the lowest
intensity. The initial intensity of H. vulgaris was similar to and significantly higher than, that
under blue light in both white and red light conditions, with DL persistence being
significantly higher, especially under red light. In contrast, the luminescence intensity and
decay rate of Arabidopsis showed less variation, particularly under red and blue light, and its
response was more consistent. Under blue light, the R² value for Arabidopsis was 0.9998,
indicating that its DL decay followed a polynomial model. Under red light, Ginkgo leaves
displayed a higher initial DL intensity and a slower decay, exhibiting an exponential decay
pattern similar to that of H. vulgaris.

11
 Ultra-Weak Photon Emission in Plants

a ydrocotyle Arabidopsis
in go le af
ulgarist le af
3 s s s 3 s s s 3 s s s

white light
0.4671 0.0973 0.0420 0.2857 0.0612 0.0557 0.4819 0.0815 0.0355

red light

0.4202 0.1350 0.0613 0.1614 0.0464 0.0464 0.2814 0.0501 0.0188

blue light

0.1897 0.0345 0.0155 0.1540 0.0390 0.0334 0.2256 0.0299 0.0272

b c

d e f

Figure . The effect of different light qualities on plants’ DL. (a) DL imaging results of the three plants

under white, red, and blue light irradiation presented for a single measurement with an exposure of 30 s,

for a total of 180 s. The results for the first 90 s are presented. (b) Decreasing trend of DL with increasing

exposure time for the same plant under different light qualities. (c) Decreasing trend of DL intensity of

three plants with the same exposure time under different light qualities. (d-f) Fitted curves of DL decay

with time for H. vulgaris, Arabidopsis, and Ginkgo leaves under different light qualities. The data represent

the mean of three measurements.

Physical mechanisms and quantum modeling of DL in plants

12
 Ultra-Weak Photon Emission in Plants

The observed DL in plants and the luminescence process of glow sticks have similarities.
Specifically, the luminescence mechanism of glow sticks comprises a series of chemical
reactions triggered by physical stimuli (e.g., bending). In this process, H2O2 inside the glow
stick reacts with the fluorescent dye, decomposing and producing water and high-energy 1O2.
The 1O2 then exchanges energy with the fluorescent dye, causing the dye molecule to jump
to the excited state. When the dye molecule returns to its ground state, energy is released in
the form of photons, producing visible light. This luminescence mechanism is analogous to
the DL phenomenon in plants, particularly in chemical reactions involving H2O2.
a b

Figure . Physical mechanisms and quantum modeling of DL in plants. (a) Theoretical model of quantum

dot luminescence. The left and right sides indicate two electrodes, and the top line indicates the

electrochemical potential of the two electrodes. (b) Theoretical model of DL in plants. e and g denote the

two energy levels of the final luminescence, and the left side denotes the chemical potential provided by

the complex chemical reaction process. The graph represents the trend of the gain intensity with the course

of the chemical reaction.

To further quantify the plant luminescence process, we borrowed the model of quantum
dot luminescence to describe its luminescence mechanism. Figure (a) illustrates the
luminescence process of quantum dots, in which electrons jump from the electrodes to the
upper energy level of the quantum dots and from the upper energy level to the lower energy
level, emitting photons in the process. The energy of the luminescence process comes from
the “electrochemical potential” of the electrons in the electrode. The DL of plants, Conversely,
is an energy transfer process driven by the chemical potential provided by a chemical reaction.
Therefore, there exists a similarity in the nature of the energy transfer between the two
phenomena.
The process of plant luminescence is triggered by a complex chemical reaction (Figure

13
 Ultra-Weak Photon Emission in Plants

(b)). In this process, the jump between the excited state and the ground state determines the
final luminescence intensity. Assuming that the electron jumps in the process of plant
luminescence follow an energy transfer mechanism similar to that of quantum dot
luminescence, we can simplify it into two energy levels: the ground state (g) and the excited
state (e). A chemical reaction causes the electrons in the excited state to occupy the number
Ne, which decreases with the decay term, providing sufficient chemical potential and
releasing energy to produce light, as per the following equation:
𝜕𝑡 𝑁𝑒 = −𝛾𝑁𝑒 + 𝜅𝑁𝑐 (1)
𝜕𝑡 𝑁𝑐 = −Γ𝑡− 𝑁𝑐 + Γ𝑡+ 𝑁𝑔 (2)
Eq. (1) describes the excited state generation process, where the first term represents the
decay of the occupation number Ne of the excited state with time and is the rate constant for
the jump from the excited state (e) to the ground state (g). The second term represents the
increase in Ne due to the transition from the intermediate state (c) to the excited state (e) and
is the rate constant for the jump. Eq. (2) describes the evolution of the occupancy number Nc
of the intermediate state with time, where Γ𝑡− is the rate constant for the transition from the
intermediate state (c) to the excited state (e), and Γ + corresponds to the rate constant for a
chemical reaction to excite an electron to a higher energy state. As a result of the chemical
reaction, the system’s energy changes from the lowest to a higher state. By analogy with the
quantum dot luminescence process, we propose to write these two jump coefficients as Γ + ≡
𝛾0 𝑛̅𝑡 , Γ − ≡ 𝛾0 （1 − 𝑛̅𝑡 ）, where nt denotes a quantity analogous to the chemical potential
and varies with the chemical reaction process.
The model’s predictions regarding the kinetics of the chemical reactions were
effectively verified in the experiments. The experimental results revealed that mechanical
injury and oxidative stress significantly increased the DL intensity of plants, a phenomenon
consistent with the description of the electron jump rate and energy transfer in the model.
Mechanical injury disrupted plant cellular structures and induced ROS accumulation. This
process accelerated the electron jump from the excited state to the ground state and increased
the DL intensity. Conversely, oxidative stress increased the luminescence intensity by
increasing the H2O2 concentration, which increased the chemical potential of the reaction and
the number of electrons occupying the excited state and facilitated the electron jump. In a

14
 Ultra-Weak Photon Emission in Plants

study of the effect of different light qualities on DL in plants, the photosynthesis efficiency
was higher under white light irradiation. Compared with monochromatic light irradiation,
white light activated stronger chemical reactions and promoted the conversion of electrons
from the intermediate to the excited states. Additionally, white light irradiation may promote
the production of photosynthetic intermediates (e.g., ATP and NADPH), which is directly
related to the chemical potential provided by the chemical reactions in the model. According
to the model, the metabolites provided additional energy for electron jumps under light
irradiation, further facilitating the electron transfer from the excited state to the ground state
and increasing the DL intensity. This result is consistent with the experimental observations,
suggesting that the wavelength of the light source during photosynthesis directly affects the
DL intensity and decay characteristics.
Furthermore, for the simplified two-energy system, we can derive the following
evolution equations:
𝜕𝑡 𝑁𝑒 = −𝛾𝑁𝑒 + 𝐺(𝑡)𝑁𝑒 (3)
where G(t) denotes the gain of the chemical reaction for Ne. Although these equations are
time-inclusive, considering the slower rate of the chemical reaction relative to the
luminescence process, we can assume that G(t) changes gradually with time and stabilizes
on longer time scales. In the solution of the model, we can approximate the process to obtain
the Ne, the time evolution of the plant’s luminescence: 𝑁𝑒 (𝑡) ≅ 𝐶exp[𝛾𝑡 + 𝐺(𝑡)𝑡], and thus
derive the decay properties of plant luminescence. Additionally, combining the polynomial
and exponential decay terms, a functional model for fitting the DL data was built:

𝐷(𝑡) = ∑ 𝑎𝑖 𝑡 𝑖 · e−𝛾𝑡 (4)

𝑖
where D(t) is the luminous intensity, ai is the polynomial coefficient, γ is the decay rate, and
t is the time. The model can capture the time-dependent characteristics of the DL process in
plants well, especially the decay trend on long-time scales.
Through the above model, we can further explore the quantitative features of the DL
process, especially by analyzing the second-order autocorrelation function of DL, 𝑔(2) (𝑡),
to reveal the intrinsic law of the plant luminescence mechanism. Additionally, we can
accurately characterize the temporal and spatial properties of the luminescence process in

15
 Ultra-Weak Photon Emission in Plants

plants by adjusting the parameters in the model, which can provide a theoretical basis for
further studies on the effects of plant physiology and chemical reactions on the luminescence
phenomenon.

Discussion

DL imaging is an emerging biomonitoring tool that offers higher sensitivity, rapid

response, and noninvasive assessment of oxidative stress levels in plants compared with
traditional biochemical and molecular assays. In this study, we systematically analyzed the
DL properties of three plants under natural growth and stress-induced conditions. The results
revealed that the DL intensity was significantly correlated with the stress level of the
environment to which the plants were exposed. The physiological status of plants under
different types of stress changed significantly, and these changes could be effectively
monitored by the DL intensity fluctuations. In an environment of high oxidative stress, the
plants displayed stronger DL intensity, reflecting their ability to respond to environmental
stress. Additionally, by combining this tool with image pattern recognition and statistical
analysis, we could accurately indicate the health status of plants, providing a new
technological tool for future plant health monitoring and management. DL imaging enables
us to assess and intervene in plant growth status in a timely manner before visual symptoms
become apparent, thus improving the sustainability of agricultural production.
Measuring DL intensity under natural growing conditions provides a valuable basis for
assessing plant health status and normal metabolic processes. Analyzing the decay pattern of
DL intensity over time gave us insight into the oxidative metabolic state of plants and
revealed the dynamic changes in intracellular ROS. These findings provide new perspectives
for understanding the metabolic activities of plants under natural conditions.
Plants undergo oxidative metabolism and produce a certain amount of ROS during
normal growth; however, when they are subjected to mechanical injury, ROS production
increases significantly, leading to oxidative stress. In this case, the plant’s antioxidant system
cannot effectively scavenge the excess ROS, leading to their accumulation. This imbalance
between generation and scavenging exacerbates oxidative stress, further damaging
biomolecules like proteins, lipids, and nucleic acids in the leaves. At the same time, the ROS

16
 Ultra-Weak Photon Emission in Plants

accumulation may contribute to the production of more high-energy states in plant cells,
which release more photons when jumping to the ground state. As a result, the initial DL
intensity of the damaged group was usually higher than that of the control group.
The differences in the responses of different plants to mechanical injury reflect their
different mechanisms of response to oxidative stress. Our results revealed that the DL
intensity significantly increased in mechanically injured plants, which is consistent with the
findings of Prasad et al[9]. Additionally, H. vulgaris displayed a strong oxidative stress
response, whereas Arabidopsis was more stable, and Ginkgo leaves responded significantly
in terms of initial DL intensity but declined more significantly over time. Future studies could
further explore how the DL property can enhance the stress tolerance performance of plants.
Endogenous metal ions in plants (e.g., Fe2 or Cu2 ) react with exogenous H2O2
solutions in a Fenton reaction to produce HO·, which accelerates oxidative reactions in
plants[2]. To test this hypothesis, we treated the leaves of three plants (H. vulgaris, Arabidopsis,
and Ginkgo leaves) with a 3% H2O2 solution and monitored their responses using a DL
imaging system. The results revealed that the initial DL intensity significantly increased in
plants under oxidative stress. Exogenous H2O2 induced oxidative stress and reacted with
intracellular metal ions to generate hydroxyl radicals and hydroxide ions, which increased
the intracellular ROS concentration and significantly increased the DL intensity. However,
with H2O2 depletion and the excited-state substances, the DL intensity gradually decreased
and eventually tended toward the steady state. The differences in the responses of different
plants to oxidative stress may be due to their differences in the rate of the Fenton reaction or
the primary oxidation reaction, which directly affects the DL intensity and its decay time.
Our results indicated that H. vulgaris and Ginkgo leaves could effectively regulate the
intracellular ROS levels, resulting in an increase in DL intensity, whereas Arabidopsis
exhibited higher sensitivity, resulting in a decrease in DL intensity. Related studies have also
demonstrated the critical role of H2O2 and oxidative processes in UPE; for example, the UPE
intensity in potato tubers was significantly correlated with endogenous H2O2[32].
Photobiology is an essential field that studies the effects of light on plant physiology,
development, and ecology. The health status of plants under different light qualities is usually
assessed by indicators like the photosynthetic rate, respiration rate, and chlorophyll content.

17
 Ultra-Weak Photon Emission in Plants

Compared with static bioindicators, DL imaging provides more detailed dynamic information
and can effectively integrate the complex relationship between light quality, time, and plant
physiological status. The results revealed that the DL intensity in plants under white light was
significantly higher than under red and blue lights, further confirming the importance of the
light source wavelength in plant physiological responses. Different light qualities had
different effects on the DL decay rate in plants: white light had the highest initial DL intensity
and a slow decay rate, while blue light had the lowest initial DL intensity and fastest decay
rate. Ginkgo leaves had the highest initial intensity in all experimental conditions, followed
by H. vulgaris, while Arabidopsis had a relatively low DL intensity. Notably, the DL
properties of H. vulgaris exhibited unique variations under red light conditions, suggesting a
higher sensitivity to that wavelength. Related studies have demonstrated that different light
wavelengths interact with each other through relevant pigments to affect the hormonal
balance in plants, which in turn triggers physiological and ecological changes in plants[33,34].
We hypothesize that the differences in the response mechanisms of plants under different
light conditions may be due to their different internal pigment systems, which have different
absorption and utilization capabilities for different light wavelengths, thus affecting plant
growth and development.
Additionally, we analyzed the DL decay properties and verified the consistency of the
quantum model with experimental data. By simplifying the theoretical DL model (Eq. (4)),
several fitting forms were derived by combining different experimental data and limiting
cases. By adjusting key parameters in the model (e.g., the decay rate γ), we investigated the
main DL decay modes and the differences between plant species. The experimental data
revealed that the DL intensity of H. vulgaris and Ginkgo leaves decayed exponentially with
time, indicating that the DL of these plants is mainly driven by fast and direct processes,
which is consistent with the assumption of an exponential decay term in the theoretical model.
When the decay rate γ is negative (i.e., γ = −b, where b is a positive decay constant), the
model exhibits an exponential decay property (Eq. (5)). The exponential decay term describes
the rapid decrease in the luminescence intensity with time. However, the DL decay in
Arabidopsis does not strictly follow the exponential decay law and displays a more complex
time dependence. This deviation could be related to intermediate reactions, complex

18
 Ultra-Weak Photon Emission in Plants

molecular interactions, or multistep energy transfer processes. Therefore, the DL properties

of Arabidopsis must be described by a more complex polynomial model, especially for longer
decay times, where the polynomial part can better capture the complexity of the luminescence
process. As the decay rate γ tends to zero, the exponential decay term exp(−γt) becomes 1,
and the model simplifies to the polynomial form presented in Eq. (6), at which point the
luminescence intensity is no longer affected by the exponential decay and represents a purely
polynomial process.
From a theoretical modeling perspective, exponential decay, which assumes that the
system follows a single decay pathway, is more effective for the DL decay in H. vulgaris and
Ginkgo leaves, but it is difficult to fully capture the complex interactions in biological
processes. For example, the system may contain multiple reactants, different decay channels,
or nonlinear interactions between molecules, which can lead to more complex decay
properties, as observed in Arabidopsis. For all plant samples, the polynomial part of the
theoretical model can, in some cases, better describe the complexity of the luminescence
process.
Although the theoretical model better fits the DL process in the experimental data, it
still has some limitations. For example, the model parameters depend on the experimental
conditions and can be influenced by plant species, chemical reaction rates, and external
environmental factors (e.g., temperature and light). Therefore, the parameters must be
adjusted to improve the fitting accuracy under different experimental conditions. Future
studies should consider more complex physical models that introduce higher-order decay
functions and biological factors to improve the understanding of the DL process in plants
further.
The present study reveals the differences in the oxidative metabolic capacity of different
plants by comparing the DL characteristics of three plants. The results demonstrate that DL
intensity and decay rate can effectively reflect the physiological status of plants. DL, as a
sensitive physiological detection method, provides valuable information for understanding
plant response mechanisms in complex environments. This finding provides a new
perspective for the study of plant adaptation mechanisms to adversity and provides a
scientific basis for future plant health monitoring and management.

19
 Ultra-Weak Photon Emission in Plants

Conclusion

Using a high-resolution and low-noise imaging system, this study provides a highly
accurate noninvasive plant monitoring method that can effectively reflect plant cells’
physiological status and health level. This method provides a new idea for the study of plant
stress response and offers a wide range of application potentials, especially in plant health
monitoring, environmental stress assessment, and precision agriculture management, with
important practical significance. In the future, by combining DL technology with quantum
physics modeling, we expect to further elucidate the response mechanism of plants to their
external environment and provide new technical approaches and theoretical support for the
realization of precision agriculture and the enhancement of plant resistance.

Material and Methods

Plant material and growing conditions

Hydrocotyle vulgaris is a perennial herbaceous wet plant of the genus Hydrocotyle. It is

native to Europe, south North America, and Central America. Due to its rapid reproduction,
large population, and strong adaptability, H. vulgaris is classified as an invasive plant in
China[35]. Our research team purchased H. vulgaris from the flower market of the Chinese
Academy of Agricultural Sciences and grew it in a well-lit environment at a suitable room
temperature. During the experiment, the plants were watered regularly to prevent wilting,
and new leaves of similar age and size were selected for each measurement to ensure
comparable data.
Arabidopsis thaliana is an annual herbaceous plant belonging to the Cruciferae family
and is an important model organism in molecular genetics research. The Columbia ecotype
(Columbia, Col-0) was used in this study. Seeds were soaked in distilled water at 4°C for 4
days before being planted in growing pots containing peat substrate (Klasmann, Potground
H). Arabidopsis plants were grown under a 16-h light/8-h dark photoperiod with a photon
flux density of 100 μmol·m−2·s−1, with the ambient temperature maintained at 25°C and 60%
relative humidity. The growth cycle was completed in 6−7 weeks. New plants or leaves of
approximately the same age and size were selected for each measurement to ensure

20
 Ultra-Weak Photon Emission in Plants

consistency.
The Ginkgo tree is a perennial deciduous tree belonging to the genus Ginkgo in the
family Ginkgoaceae. It is rich in various biologically active components, including
flavonoids and pigments, which give it a high medicinal value. The leaves were harvested on
the campus of the Beijing Institute of Technology, and the research team selected only healthy
leaves free from pests and avoided picking fallen or discolored leaves to ensure the quality
of the samples and the reliability of the experimental results.

DL imaging system

The configuration of the DL imaging system for radiation detection is presented in

Figure . The system comprises four main components: a sample chamber, a qCMOS camera,

a water-cooling system, and a computer. Due to the extremely low intensity of the DL photon
flux emitted by the organism, a highly sensitive qCMOS camera (Hamamatsu ORCA-Quest)
was selected. The camera’s qCMOS imaging sensor has single photon sensitivity and
subpiston readout noise characteristics, minimal dark current signal loss, and is equipped
with a two-dimensional photon counting function that allows simultaneous acquisition of
temporal and spatial information on DL intensity. The camera operates in the single photon
number resolved mode with a scan rate of 17.6 frames per second and has a spectral
sensitivity of 400−1000 nm with a quantum efficiency of up to 85%. The accompanying
Computar objective (M2514-MP2, f = 25 mm) allows efficient DL measurement in
organisms. A water-cooling system (LX-150 cooling circulator) was used to maintain the
temperature of the camera at −40°C and minimize the occurrence of dark counts. The
experiment was conducted in an opaque darkroom, with similarly sized intact plant samples
(plants or leaves) positioned on a completely sealed black dark box sample stage, which was
located within the camera’s field of view. For each measurement, the same image acquisition
parameters were maintained, with a single imaging exposure time of 30 seconds and six
consecutive measurements with a cumulative imaging time of 180 seconds. The distance
between the plant and the detector was 25 cm, and the distance between the leaf and the
detector was 5 cm. After acquiring the detector signal, the data were transferred to a computer
outside the darkroom. HCImage software was used for image acquisition and analysis, while

21
 Ultra-Weak Photon Emission in Plants

data correction was achieved by subtracting the background signal before each measurement.
Image processing techniques were also integrated to improve the measurement system’s
image quality.

Figure . Schematic of DL imaging.

Experimental methods and stress treatments

The experiments were conducted in a dark room with the temperature maintained at
18℃. The air conditioning was run for 1 h before the start of the experiment, and the
apparatus was allowed to reach equilibrium for 30 min to ensure the stability of the
background signal and consistency of the ambient temperature. All biological samples were
allowed to acclimatize in an external dark room for 30 min before measurement to minimize
the effects of the environment and prevent the material’s residual luminescence from
affecting the results. Measurements were made by illuminating the sample and measuring the
intensity of photons re-emitted from the sample after the light source was switched off.
Because different leaf areas were used for each sample, the DL intensity was normalized to
the photon intensity emitted per unit time and per unit leaf area. During the measurement
process, fresh leaves of comparable growth stage were selected. They were first washed with
deionized water, and the surface moisture was removed with filter paper. The leaves were
then exposed to a 220 V and 10 W white LED light source for 5 min. Then, they were quickly

22
 Ultra-Weak Photon Emission in Plants

transferred to the sample chamber, where measurements of the leaves’ DL intensity decay
over time immediately started. The total duration of each measurement was set at 180 s, with
three consecutive measurements. Background noise measurements were conducted under the
same experimental conditions to obtain the final DL intensity measurements. These values
were then subtracted from the average of the three measurements to obtain the final result.
Mechanical injury
The plant leaves were mechanically damaged using a sharp blade in a low-light
environment. Precautions were taken to avoid additional stress or damage to other parts of
the plant samples. After 5 min of mechanical damage, the samples were exposed to a 220 V
and 10 W white LED light source for 5 min. This step ensured that the leaves could quickly
adapt to the changed environment. Following the illumination period, DL intensity
measurements were initiated.
2O2 processing
During the H2O2 treatment, a freshly prepared 3% H2O2 solution was applied evenly to
the surface of the plant samples to ensure optimum leaf infiltration. An equal amount of water
was used as a control treatment to ensure the reliability of the experimental results. After 5
min of H2O2 treatment, the samples were irradiated for 5 min under a 220 V and 10 W white
LED light source. This step was done to promote the recovery of photosynthesis and related
physiological responses. At the end of the treatment, DL measurements were started to assess
the effect of H2O2 on the plants’ DL.
Light quality
LED curing lights (emitting white, red, and blue) of 220 V and 5 W were used as
different light sources. The white light source emitted a full spectrum of light, while the red
and blue light sources had central wavelengths of 660 and 460 nm, respectively. During the
experiment, plant leaves treated with deionized water were exposed to their respective light
sources at 5 cm from the LED lamps for 5 min. After irradiation, the sample leaves were
immediately transferred to the sample chamber, and their DL intensity measurement started
immediately.

Cur e-fitting model

23
 Ultra-Weak Photon Emission in Plants

A suitable mathematical model was selected to describe the DL decay curves and make
a quantitative comparison of the differences in DL intensities exhibited by different plants.
The DL intensity data (in counts per 30 s) of H. vulgaris and Ginkgo leaves over time were
subjected to an exponential regression model as follows:
y = a × exp(b × x) (5)
where y is the response variable, x is the predictor variable, a is the regression coefficient
representing the initial DL intensity, and b is the dynamic parameter of the exponential
decay (i.e., the slope parameter) representing the DL decay rate.
For the Arabidopsis DL intensity data, a polynomial model was fitted with the
following model form:
y = A0 + A1 x + A2 x 2 + A3 x 3 + A4 x 4 (6)
where y is the response variable, x is the predictor variable, and the DL is characterized by
the parameters A0, A1, A2, A3, and A4.

Supplementary Data

Supplementary Figure S1. The dynamics of DL intensity for three plants (during the
first 90 s) at different mechanical injury times.

Supplementary Figure S2. 3D waterfall plots of DL intensity and its decay for three
plants at different mechanical injury times.

Supplementary Figure S3. The dynamics of DL intensity for three plants (during the
first 90 s) at different oxidative stress times.

Supplementary Figure S . 3D waterfall plots of DL intensity and its decay for three
plants at different oxidative stress times.

Supplementary Table 1. Exponential model fit data for the effect of oxidative stress
on DL in plants.

Supplementary Table 2. Polynomial model fit data for the effect of oxidative stress
on DL in Arabidopsis.

Supplementary Table 3. Exponential model fitting data for the effect of different light

24
 Ultra-Weak Photon Emission in Plants

qualities on the DL in plants.

Supplementary Table . Polynomial model fit data for the effect of different light
qualities on the DL in Arabidopsis.

Acknowledgments and Funding

This research did not receive any specific grant from funding agencies in the public,
commercial, or not-for-profit sectors. Thanks to Figdraw (www.figdraw.com) for help with
character cartoons.

Author Contributions

Yan-Xia Liu: Conceptualization, Methodology, Software, Investigation, Formal

Analysis, Validation, Writing−original draft. ai-Yu Fan: Methodology, Investigation,
Formal Analysis. Yu- ao Wang: Software, Data curation. Yan-Liang Wang:
Conceptualization, Methodology. Sheng-Wen Li: Conceptualization, Methodology,
Investigation. Shi-Jian Li: Conceptualization, Methodology, Writing−review & editing. Xu-
Ri Yao: Conceptualization, Formal analysis, Methodology, Resources, Writing−review &
editing. Qing Zhao: Resources, Supervision, Writing−review & editing.

Conflicts of Interest: The author declares no conflict of interest.

Table 1. Exponential model fit data for the effect of mechanical injury on DL in plants.

control wounded control wounded control wounded

Plant species
a a b b R2 R2

H. vulgaris 0.9704 2.0064 −0.0335 −0.0434 0.9826 0.9918

Ginkgo 1.5059 2.6518 −0.0406 −0.0517 0.9885 0.9933

Table 2. Polynomial model fit data for the effect of mechanical injury in Arabidopsis.

Arabidopsis A A1 A2 A3 A R2

Control 0.9158 −0.0311 3.8873E-4 −2.0743E-6 4.0124E-9 0.9937

Wounded 1.3328 −0.0476 6.2758E-4 −3.5203E-6 7.1168E-9 0.9983

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 Ultra-Weak Photon Emission in Plants

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