OVERVIEW: THE ABIOTIC FACTORS ‘STRESS AND LIGHTING’

HAVE AN INFLUENCE ON THE BIOACTIVE COMPOUND

FLAVONOID IN STRAWBERRY PLANT

Enceng Sobari 1,2)

1) Graduate Doctoral Student, Faculty of Agriculture, Universitas Padjadjaran; 2)

Department of Agriculture, State Polytechnic of Subang

1)Bandung Sumedang Highway KM.21, Hegarmanah, Jatinangor District,

Sumedang Regency, West Java 45363, Indonesia.
2) Brigadier General Katamso Street No. 37 (Behind Subang General Hospital),

Dangdeur, Subang District, Subang Regency, West Java 41211, Indonesia

1,2) [email protected]

ABSTRACT

The increasing demand for strawberries is driving advancements in technology that

support strawberry production, which is expected to become far more effective and
efficient. Strawberries are one of the plants whose fruits contain substances that are
highly beneficial for human health. Abiotic factors such as stress and light often play
a decisive role in the production of secondary metabolites like flavonoids, which are
important compounds in strawberry plants. This article aims to provide information
related to the impact of abiotic factors ‘stress and lighting” that can affect these
compounds. The studies used to support the information in this article include
reputable scientific journals published and indexed in ScienceDirect, Springer,
articles indexed in Scopus, and relevant literature on GoogleScholar. All literature
providing insights on the importance of biological activity and plant physiology,
secondary metabolites, flavonoid biosynthesis, and the phytochemistry of plants.
The influence of abiotic stress and lighting on plants can affect various aspects of
plant growth, which will eventually impact the response in terms of quality produced
and also the compounds contained within the plant itself. Based on the study of
abiotic factors such as stress and lighting, these have a crucial role in the
biosynthesis of flavonoid compounds that affect production in strawberry plants.

Keywords: Abiotic stress, Bioactive, Flavonoid, Secondary metabolites, Lighting,

Strawberry.

1. INRODUCTION

Strawberries are among the most popular fruits worldwide, not only consumed fresh
but also used in various types of processed food products such as snacks and
beverages due to their excellent nutritional content. Strawberries contain vitamin C
(39-86 mg per 100 g of fruit) and vitamin A (60 IU per 100g), as well as minerals,
pectin, potassium, calcium, and phosphorus. (Singh et al., 2018). Additionally,
strawberries are one of the plants whose fruits contain substances that are highly

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beneficial for human health. These include antioxidant agents such as anthocyanins,
which are flavonoid compounds, vitamins, sugars, and other phenolic compounds
(Martinsson et al., 2006). Flavonoids are a class of secondary metabolites produced
by plants, including phenolic compounds that belong to the flavonoid group. These
can produce natural antioxidants (Saija et al., 1995). Flavonoids, as part of the
metabolites, play a significant role in plant growth and stress response (Li et al.,
2021). These metabolites function as compounds used in plant growth and
development, divided into two types: primary metabolites, which are produced in
limited quantities and used by plants for growth, and secondary metabolites, which
serve as a defense mechanism for plants against both biotic and abiotic stress
(Angin et al., 2019). The compounds found in strawberries, such as antioxidants,
offer significant health benefits.

Secondary metabolites are compounds possessed by plants that function in the

processes of growth and development. They are small molecular organic
compounds that serve as bioactive substances, whose formation depends on the
plant's stimulation by abiotic factors such as environmental stress (T. Wang et al.,
2019). Secondary metabolites are produced by plants to enhance their
competitiveness in their environment, offering various benefits to the plant itself as
well as other living organisms, and acting as antimicrobials. More than 50,000 types
of plants have been identified as producers of secondary metabolites, with many
modern medicines beginning to rely on these plant-derived compounds (Teoh,
2016). There are over 4,000 flavonoids widely distributed across plants and
prokaryotes, which belong to a large group of secondary metabolites that can be
categorized as phenolic compounds (Woo et al., 2002; Middleton, 1998). Research
findings suggest that flavonoids can perform various functions as antioxidants and
regulators of development in photoprotection (Agati & Tattini, 2010).

Flavonoids are known to increase tolerance to various abiotic stresses, serving as a

defense mechanism for plants, and as a basis for allelopathic interactions with other
plant species. Species across all orders in the plant kingdom, from liverworts to
angiosperms, invest a significant amount of metabolic energy in producing these
compounds (Kevin S. Gould & Lister, 2005). The environmental conditions for
growing strawberries are not always ideal, abiotic factors present challenges that
must be navigated during the growth and development of strawberry plants.
Secondary metabolite compounds such as flavonoids can serve as defense
mechanisms and signals for strawberry plants in conditions of both biotic and abiotic
stress. (Goyal et al., 2012). Therefore, this study is expected to provide deeper
insights into abiotic factors such as stress and lighting that can affect these
compounds in the physiological aspects of strawberry plants. It aims to serve as an
important piece of information in the development of strawberry plant research.

2. RESEARCH METODOLOGY
The materials used in compiling the database for the creation of this article were
obtained from primary scientific article sources such as ScienceDirect, Springer,
articles indexed in Scopus, and relevant literature on GoogleScholar. The method
used in creating this review article involves utilizing a literature study to search for
information related to the necessary aspects as follows:

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i. Related to the topic of metabolites: All literature providing insights on the

importance of biological activity and plant physiology, secondary metabolites,
flavonoid biosynthesis, and the phytochemistry of plants.
ii. Abiotic factors in plants: Literature searches focusing on topics related to abiotic
stress such as drought stress, lighting, and other aspects related to enhancing
flavonoid compounds, considered beneficial regardless of the journal impact
factor, popularity, or its source.

3. RESULTS AND DISCUSSION

A. The Impact of Light on the Quality and Biosynthesis of Flavonoids in

Strawberries
Strawberry plants often face many issues during the cultivation process, especially
in the field, such as low productivity and the quality of the fruit produced,
necessitating a good system to anticipate these problems. Adverse and extreme
weather conditions, which can result in strawberry production losses, the concept of
controlled environment agriculture accompanied by technology application is a good
alternative for the agricultural sector (Hernández-Martínez et al., 2023;Wai et al.,
2023). Strawberries face numerous challenges during the cultivation process,
resulting in yield losses of about 30-70%. Proper management is necessary to
mitigate these issues (Mahmud et al., 2019). Cultivating strawberries in controlled
indoor environments has become the most modern alternative for producing high-
quality strawberry fruits. The majority of indoor strawberry cultivation uses
greenhouse technology, widely adopted in response to unpredictable weather
conditions that can affect the yield and quality of cultivated strawberries, such as a
decrease in the sweetness and texture of the strawberry fruit (Gavilán et al.,
2015;Chaichana et al., 2020). A decline in the quality of strawberry fruits can also
lead to consumers deciding against purchasing them (Zhou et al., 2022).

Lighting is a crucial factor in the physiological aspect of plants that can influence the
growth and development of strawberry plants due to its connection with the
photosynthesis process, which impacts the formation of strawberry fruits. Lighting
issues are particularly significant for the leaves and can greatly affect the quality and
characteristics of the produce (Pydipati et al., 2006). Supplemental lighting has been
widely successful in enhancing vegetable production in greenhouses, providing
significant differences in growth and yield. The rate of photosynthesis in leaves of
plants exposed to additional LED lighting is much higher, as indicated by increases
in leaf dry weight, leaf area, and specific leaf dry weight. These improvements
significantly impact the average fruit weight, fruit count, and overall yield (Hidaka et
al., 2013).

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(Chaichana et al., 2020) (Wai et al., 2023)

Figure 1. Addition of LED lighting for the growth of strawberry plants

In addition to nutrients and water, another factor such as light becomes an

important factor that affects the quality of fruit from strawberry plants cultivated in
greenhouses. Figure 1 shows the use of LED lighting on strawberry plants.
Supplemental lighting is recommended for producing high quality strawberry fruits
(Xu et al., 2023). Exposure to light can trigger a series of physiological processes in
strawberries and affect their commercial value. Light is one of the most important
environmental factors influencing the biosynthesis of anthocyanins in plants and can
enhance the red coloring of strawberry fruits after harvest. This is combined with a
doubling of the anthocyanin content, which is a compound derived from flavonoids
(Miao et al., 2016; L. Wang et al., 2022).

B. The Impact of Drought Stress on Plant Physiological Mechanisms

One-third of the Earth's surface is classified as arid or semi-arid, while areas

that are humid, where the world's food is produced, experience periods of severe
drought. Consequently, the impact of drought has become a primary focus on the
influence of plant productivity worldwide (Díaz et al., 2005). Drought stress is a factor
that significantly affects agricultural productivity in developing countries, reaching 50-
70%. Among abiotic stresses, it is considered the most detrimental, thereby
hindering maximum agricultural productivity worldwide (Verma & Deepti, 2016). The
harmful impact of drought conditions can have detrimental effects on growth,
chlorophyll content, cell integrity, photosynthesis, leaf water potential, and water
deficiency (Hussain et al., 2020). Drought stress causes a significant inhibition of
growth and the rate of photosynthesis, as well as inducing a number of major
changes in the morphological, physiological, and biochemical responses when
plants are exposed to drought stress. These changes can enhance the plant's ability
to survive and grow during periods of drought (Yin et al., 2005). Abiotic stress can
result from atmospheric pollutants, the presence of heavy metals, low temperatures,
or soil acidity, which can also affect the productivity and quality of plants (Figure 2).

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Figure 2. The impact of abiotic stress on plants that can affect growth and plant
productivity (Díaz et al., 2005)

Under stress conditions, plants are capable of initiating defense responses such as
physiological, biochemical, and molecular mechanisms under drought to enhance
their survival ability in conditions of water scarcity during growth and development
periods (Amoah et al., 2019). Drought in plants elicits a complex response, ranging
from wilting processes, reduced photosynthesis rates, causing stomatal closure, and
increased leaf temperature at physiological, metabolic, and developmental response
levels, dependent on the degree of drought stress (Guimarães-Dias et al., 2012).
Once stress occurs, complex biochemical and physiological mechanisms are
activated to protect crucial processes such as cell respiration, photosynthetic activity,
and nutrient transport. Additionally, stress tolerance responses are induced to
preserve organelles and tissue structures, such as the meristem (Kramer, 1983;
Kramer & Boyer, 1995).

C. The Role of Secondary Metabolites in Plants

Bioactive compounds are substances found in nature and are part of the food chain.
Possessing the ability to interact with one or more living tissue compounds,
demonstrating effects on human health (Biesalski et al., 2009). Possessing an
unlimited combination of functional groups, including hydroxyl, alcohol, aldehyde,
alkyl, benzyl rings, and steroids, each can produce a diversity of plant compounds,
with unique characteristics (Beckles & Roessner, 2012). More than 8,000 different
phenolics have been identified within the plant kingdom, representing one of the
most abundant and widespread classes of plant secondary metabolites (Cheynier et
al., 2013). Through complex metabolic pathways, these are referred to as secondary
metabolites, whose compounds have the capability for synthesis used by plants for
various functions, especially in responding to biotic and abiotic stress encountered
(Michel, 2011). Various research findings indicate that these metabolites fall into
three classes: polyphenols, alkaloids, and terpenoids, demonstrating the bioactivity
of these molecules in antitumor, antiviral, antimicrobial, antioxidant, and anti-
inflammatory activities. Therefore, plants containing these metabolites can be

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utilized in the fields of therapy, pharmacology, cosmetology, and food (Rispail et al.,
2005; Sompila et al., 2021).

D. The Role of Flavonoids in Plants

Flavonoid content is abundantly stored within plant bodies, aiding in the

response to growth and development throughout their lifespan. Flavonoids are the
most plentiful secondary metabolite compounds in plants (Han et al., 2023). These
compounds protect against UV rays, pests, and diseases, and also play a crucial
role in the color, taste quality, and nutritional value of the fruits produced (Gouot et
al., 2019). Flavonoids primarily consist of five subclasses: anthocyanins, flavanones,
flavanonols, flavonols, and flavanols (Durazzo et al., 2019). The accumulation of
flavonoids can be a key defense for plants against various environmental stresses,
as flavonoids belong to the polyphenol compounds with antioxidant activity,
comprising flavones, flavanones, flavonols, isoflavones, and anthocyanins (P. Wang
et al., 2018;Sun et al., 2020). Flavonoids, as bioactive polyphenolic compounds in
plants with structures like Kaempferol-3-O-rutinoside, quercetin-3-O-glucoside, and
kaempferol-3-O-glucoside, can play a significant role in the prevention and mitigation
of the impact of several diseases in humans. These flavonoid glycosides are
responsible for antioxidant activities and form the scientific basis for their use in
traditional medicine. (Badmus et al., 2016). The flavonoid fraction from each plant
possesses an effective hepatoprotective effect as flavones, which clinically reduces
symptoms in liver damage (Gupta et al., 2015). The flavonoid content has a close
relationship between plant parts and the phenological development of the plant,
which can affect the content of compounds such as rutin, amentoflavone,
hyperoside, isoquercitrin, quercitrin, quercetin, avicularin, (+)-catechin, and (–)-
epicatechin (Çirak et al., 2014).

E. Flavonoid Biosynthesis Induced by Stress

The role of flavonoids can offer new insights into enhancing plant production
potential by utilizing plant varieties rich in flavonoids as crops tolerant to biotic and
abiotic stress (Nakabayashi et al., 2014). High flavonoid content is found in the early
development stages of young fruit tissue, evidenced by high gene expression related
to the biosynthesis and accumulation of flavonoids (Moriguchi et al., 2001). The
enhancement of anthocyanin biosynthesis enzymes from the pathway involved in
phenylpropanoid metabolism, which converts phenylalanine and related substrates
through phenylalanine ammonia-lyase (PAL) into flavonoids including anthocyanins.
The proteins whose synthesis is concurrently increased include phenylalanine
ammonia-lyase (PAL), chalcone isomerase (CHI), flavanone 3-hydroxylase (F3H),
dihydroflavonol 4-reductase (DFR), and UDP-glucose:flavonoid-3-O-
glucosyltransferase. PAL has been identified as an enzyme positively associated
with the ripening of strawberry fruits (Song et al., 2015).

4. CONCLUSIONS

Abiotic factors such as stress and light play a crucial role in the biosynthesis
of flavonoid compounds in strawberry plants. These factors influence the production
of flavonoids and can optimize growth. The production of strawberry plants
containing bioactive compounds is beneficial for human health.

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