International Journal of Horticultural Science and Technology (2025) Vol. 12, No. 1, pp.

83-100

International Journal of Horticultural Science and Technology

Journal homepage: http://ijhst.ut.ac.ir

A Preliminary Experimental Protocol for Enhanced Tomato Callus

Formation and Growth via Several Medicinal Plant Extracts
Omid Sohrabi1*, Abbdollah Hatamzadeh1, Azim Ghasemnezhad2, Habibollah Samizadeh1, Vahid Erfani-moghadam3
1 Plant Production Department, Guilan University, Rasht, Iran
2 Agricultural Sciences and Natural Resources, Gorgan University, Gorgan, Iran
3 Department of New Technologies in Medicine, Golestan University, Gorgan, Iran

ARTICLE INFO ABSTRACT

Article history: Medicinal plants have been used historically for various treatments and
in human nutrition. Due to their natural occurrence and
Received: 4 August 2023,
biodegradability, medicinal plants can be an eco-friendly alternative to
Received in revised form: 30 December 2023,
Accepted: 5 January 2024 toxic chemicals such as pesticides, fungicides, and herbicides. An in
vitro experiment was conducted to test the impact of six medicinal
plant extracts on tomato growth and callus induction. Two
Article type: concentrations of each extract (50 and 100 mg L-1) were added to the
Murashige and Skoog (MS) medium culture. Results showed that
Research paper Juniperus sabina extract (50 mg L-1) accelerated tomato seed
germination by 100% in vitro, compared to the control group in MS
Keywords: basal medium. Callus growth index (CGI) and callus weight (CW)
increased by 50% and by more than 200% when using Taraxacum
Allelopathy,
officinale (100 mg L-1) and Conocarpus erectus (50 mg L-1) extracts,
Kermanshah, respectively. Biochemical analysis revealed that the extracts were rich
Plant growth regulators, in phenolic compounds (348 mg g-1 of total phenol), flavonoids (162
Plant for plant, mg g-1), antioxidants (61%), and auxin. A rise in antioxidant activity,
Tissue culture plant growth regulators (PGR), and plant defense elicitation probably
contributed to these outcomes. Plant extracts also affected the
biochemical content of calluses, except for their total phenol. We
COPYRIGHT recommend using these plant extracts to increase growth, accelerate
seed germination, and promote callus induction. Optimizing
© 2023 The author(s). This is an open-
concentrations and combinations of medicinal plant extracts require
access article distributed under the further research to maximize their benefits to different plant species
terms of the Creative Commons and their growth-related values.
Attribution License (CC BY). The use,
distribution or reproduction in other Abbreviation: Auxin content (AC), Callus growth index (CGI), Callus
medium is permitted, provided the weight (CW), 2,2-diphenyl-1-picrylhydrazyl (DPPH), Indole acetic acid
original author(s) and source are cited, (IAA), Murashige and Skoog culture medium (MS), Seedling fresh
weight (SFW), Stem length (SL), Total flavonoid (TF), Total phenol (TP)
in accordance with accepted academic
practice. No permission is required from
the authors or the publishers.

Introduction1 amino acids, and algae extracts can become a

In recent years, an increase in global populations fundamental approach to agricultural safety and
has been concomitant with the prevalence of sustainability (De Saeger et al., 2020). For
diseases caused by unintentional consumption of centuries, common knowledge was that plants
synthetic chemicals by humans, thus leading to a can affect the growth and development of their
greater emphasis on natural materials in surrounding plants through the production and
agriculture. As a result, using biological secretion of specific substances (Pan et al., 2015).
stimulants such as microorganisms, humic acid, This phenomenon is called allelopathy, and it

*Corresponding author’s email: [email protected]

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Sohrabi et al., Int. J. Hort. Sci. Technol. 2025 12 (1): 83-100

refers to the ability of plants to stimulate or develop more efficient and sustainable
inhibit the growth of other plants (Mahboobi and agricultural practices.
Heidarian, 2016). Plants produce allelopathic or Tomato belongs to the Solanaceae family and is
allelochemical substances, both known and cultivated worldwide as a fruit. It is a source of
unknown, but primarily specific compounds such bioactive, antioxidant, and antimicrobial
as tannins, phenolic acids, lignins, alkaloids, compounds (Silva-Beltran et al., 2015). Given the
flavonoids, coumarins, and terpenoids (Da Silva et benefits of medicinal plants, we aimed to apply six
al., 2015). Allelopathy can be negative (Santonja medicinal plant extracts on tomato seeds in vitro
et al., 2019) or positive when considered a and assess callus formation. The six medicinal
phenomenon and as hormesis or positive plants were Juniperus sabina (Adams and
allelopathy (Da Silva et al., 2015). Previous Wingate, 2008; Dahmane et al., 2015), Rheum
research showed that seaweed extract application ribes (Keser et al., 2019), Allium
through foliar spraying can increase anthocyanins jesdianum (Amiri, 2007), Conocarpus
(hydrocinnamic acid) and flavonols (Salvi et al., erectus (Nascimento et al., 2016), Taraxacum
2019). Today, plant growth control is a leading officinale (Faria et al., 2019), and Dorama
topic of discussion in crop production, especially aucheri (Ajani and Claβen–Bockhoff, 2021;
in marketing management and ripening Mianabadi et al., 2015). Thus, the present work
uniformity (Rietveld, 1983). The effect of aimed to investigate the impact of six medicinal
allelopathic compounds on plant growth is plant extracts on tomato growth and callus
diverse and depends on many factors. For induction compared to a regular protocol for
instance, juglone, the dominant allelopathic tomato culture that uses MS with plant hormones.
compound of walnut, reportedly limited the
growth of cucumber by stimulating the synthesis Materials and Methods
of abscisic acid (ABA) in cucumber as a growth Plant materials and extraction
inhibitor or by preventing the production of All medicinal plants were collected from the
growth-promoting hormones such as auxin, Kermanshah region, except for Conocarpus
cytokinin, and gibberellin (Terzi et al., 2003). erectus and Juniperus sabina. All plants were
Parallel to allelopathy, an alkaloid compound, identified based on available sources (Nemati and
cylindrospermopsin, reportedly halted mitosis in Jalilian, 2011; Sohrabi, 2022). The plant material
bean plants when used at high concentrations was washed with water and dried in the shade.
(Garda et al., 2015). Ziziphus mauritiana plant Then, the plants were extracted in a ratio of 1:10
extract decreased mitotic indices in onion (plant material to 85% methanol solvent) using a
(Owolarafe et al., 2020). Ricinus cold maceration method for two periods of 24 h
communis methanolic extract significantly (Sohrabi, 2022). After preparing the powder (Fig.
affected corn seed (Zea mays L.) germination at 1), the methanolic extract was obtained. Extracts
different levels; the extract concentration at 10 from the two periods were mixed and used for the
mg mL-1 resulted in zero germination (Saddiqe et next steps. The extracts were concentrated using
al., 2020). an IKA HB10 rotary evaporator (Fig. 1B). The
Some allelopathic compounds can have a positive concentrated extracts were then placed in a
effect on plant growth. For example, a low freeze-dryer (Christ Beta 2-8 LB plus, made in
concentration (2.5-5%) of chia (Salvia Germany) for 90 h. This was followed by
hispanica L.) and wormwood (Artemisia preparing the pure and dry powder of the plant
absinthium L.) extract had a positive effect on extracts (Fig. 1C).
lettuce (Lactuca sativa L.) germination (Erhatic et
al., 2023). Priming safflower seeds with fennel,
Preparing tissue culture medium and
peppermint, and cumin extracts changed GA3
content in safflower (Carthamus tinctorius L.)
samples
and had a positive effect on seed germination To prepare plant samples in vitro, seeds of the CH
(Alivand and Farajzadeh, 2017). It is also worth cultivar (Fallat Company) were initially placed in
70% ethanol for 1 min. Then, they were
noting that the allelopathic properties of a plant
can vary depending on extract concentration. For disinfected by immersing them in 5% sodium
example, garlic extract (Allium sativum L.) hypochlorite for 10 min and rinsing them with
sterile distilled water.
showed both stimulatory and inhibitory effects on
plant growth, with low concentrations having a The seeds were then grown under a laminar hood
stimulatory effect and high concentrations on MS culture medium containing 8% agar and
3% sucrose, 2 mg L-1 BA and 0.1 mg L-1 IAA
completely stopping growth (Cheng et al., 2016).
Thus, understanding how allelopathic (Gerszberg et al., 2016), with ten seeds in each
compounds affect plant growth can help us glass. To produce callus, 1 cm stem explants were

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obtained from the resulting seedlings.

Fig. 1. (A) Medicinal plants as a natural compound source. (B) Concentration of herbal extracts. (C) Pure extract
powders of the medicinal plants.

Applying treatments in culture medium standard curve for total flavonoid measurement,
Before adding plant extracts to the culture which was expressed as mg g-1 using a relevant
medium, the culture medium was cooled and the equation:
extracts were sterilized using a 45-micron filter.
Then, they were added to Murashige and Skoog 𝑦 = 0.002𝑥 – 0.0282 (𝑅2 = 0.99)
(MS) culture medium at two concentrations (50
and 100 mg L-1) (Sohrabi, 2022). Each replication Plant growth hormone measurement (auxin)
consisted of three explants, and each sample was Auxin content in the plants was measured using a
a mixture of all three explants in each replication method described by Gang et al. (2019). In this
to obtain a final sample. method, a Salkowski reagent was used for
measuring auxin levels. For this purpose, one
Morphological measurement gram of callus tissue was boiled in 10 mL of 80%
The effect of plant extracts on seed germination ethanol. The callus tissue was then ground and
was investigated by measuring traits such as stem the solution was filtered using Whatman filter
length (SL) and seedling fresh weight (SFW). paper. Then, 1 mL of the resulting extract was
Callus growth index (CGI) and callus weight (CW) mixed with 2 mL of the Salkowski reagent. To
were evaluated in the cultured samples under the prepare the Salkowski reagent, a 0.5 M solution of
influence of plant extract treatment (Alizadeh, FeCl3 was prepared. One mL of it was mixed with
2013). 50 mL of 35% perchloric acid. The resulting
mixture was placed in a steam bath at 40-50 °C for
Biochemical measurement 15 min to complete the reaction. The presence of
Total phenols (TP) and total flavonoids (TF) were auxin was indicated by a pink color. The
measured using a method described by Chang et absorbance of each sample was read at a
al. (2002). The percentage of inhibition of free wavelength of 530 nm using a spectrophotometer.
radicals (DPPH) was determined using a method
described by Brand-Williams et al. (1995). Statistical analysis
Different concentrations of gallic acid were used The experiment comprised a completely
as a standard for the standard curve. The total randomized design with three replications under
phenol was expressed as mg g-1 of gallic acid using in vitro conditions in laboratories of the
the equation 𝑦 = 0.0005𝑥 + 0.0257 (𝑅2 = Agricultural Sciences and Natural Resources
0.9925). Quercetin was used for determining the Department at Guilan University and Gorgan

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University (2020-2021). Data analysis was length, whereas this relationship had a linear
performed using SAS version 9.4 software, and pattern in other cases (Fig. 2). The results largely
mean values were compared using the least depended on plant species and extract
significant difference (LSD) (p≤0.01). Figures concentration. For example, as seen in Figure 2,
were illustrated using Microsoft Excel. Juniperus sabina, Rheum ribes, and Taraxacum
officinale treatments led to a decrease in growth
Results when using them at high extract concentrations.
Stem length However, Allium jesdianum, Dorema aucheri, and
Table 1 indicates that the effect of treatments on Conocarpus erectus treatments resulted in
stem length was significant (p≤0.01). In the case enhanced growth when using them at high extract
of some extracts, an increase in extract concentrations (Fig. 2).
concentration resulted in a decrease in seedling

Table 1. Statistical analysis of the effect of treatments on the morphological and biochemical properties of tomato
callus.
S.O.V df SL SFW CW CGI TP TF Anti- Auxin
oxidant
Treatments 12 1.54** 0.0025** 0.0078** 1.22** 271.1 ns 6.44** 123.22** 0.0023**
Error 26 0.06 0.0004 0.0014 0.34 144.05 1.63 37.09 0.00047
CV 3.46 11.58 25.45 39.56 18.68 13.73 7.88 21.83
ns non-significant and * and ** are significant at 5 and 1 percent probability levels, respectively.

10
a
9 ab
bc bc
de cd
8 ef ef ef ef ef
f
7 g
Stem length (cm)

6
5
4
3
2
1
0

Plant extract (mg L-1)

Fig. 2. Effects of medicinal plant extracts on tomato seedling stem length in vitro. Control was MS medium without any
plant extract.

Additionally, the biochemical potential of the potential, particularly with the presence of auxin
medicinal plant extracts revealed their hormonal (Table 2).

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Table 2. Biochemical profile and amount of auxin in medicinal plant extracts.

Medicinal plant TP (mg g-1) TF (mg g-1) Antioxidant potential (%) Auxin (mg g-1)

J. sabina 31.1870 15.893 74.285 0.3838

A. jesdianum 21.0870 15.898 12.780 0.3260
T. officinale 22.595 14.757 80.483 0.2832
C. erectus 62.545 18.56 67.568 0.2553
D. aucheri 31.663 17.043 58.46 0.3567
R. ribes 40.269 15.5 80.805 0.1483

Highest growth in stem length (8.9 cm ± 0.117 2, the Taraxacum officinale plant extract at 100
cm) was observed in response to Juniperus sabina mg L-1 (6.25 ± 0.117 cm) decreased stem growth
extract at 50 mg L-1 compared to the control (7.2 compared to the control (7.21 ± 0.117 cm).
cm ± 0.117). Moreover, no significant difference
was observed between seedlings treated with Seedling fresh weight
Juniperus sabina, Dorema aucheri, Allium Seedling fresh weight was also significantly
jesdianum, and Conocarpus erectus extracts (Fig. affected by the treatments (p≤0.01) (Table 1).
2). On the other hand, in some treatments, a high Furthermore, it was evident that the plant
concentration resulted in suppression. For extracts had various effects on seedling fresh
example, using Taraxacum officinale extract at weight (Fig. 3). Some treatments had stimulatory
100 mg L-1 decreased the stem length to 6.25 ± effects, while others had inhibitory effects. As
0.117 cm compared to the control (7.2 ± 0.117 revealed in Figure 3, some treatments such as
cm). In addition to differences in morphology, the Juniperus sabina extract increased SFW through a
germination rate of seeds cultivated in the culture linear pattern of changes in extract concentration.
medium with plant extracts was much higher In contrast, a negative relation was observed in
than the control. Figure 2 shows the results of SFW by increasing the Allium jesdianum extract
using the plant extracts. For instance, the concentration.
Juniperus sabina extract (at both concentrations) The highest seedling fresh weight was observed
stimulated growth, while Taraxacum officinale at in response to 100 mg L-1 Juniperus sabina extract
100 mg L-1 reduced stem length. Based on our (0.231 ± 0.0052 g), while the lowest weight was
results, tomato seedlings showed significant observed in response to 50 mg L-1 Rheum ribes
growth differences under the influence of the (0.129 ± 0.0052 g) (Fig. 3).
medicinal treatment extracts. As shown in Figure
Seedling fresh weight (g)

0.3
0.25 a
ab ab ab
0.2 bcd bc bcd
cd cde cde
de de
0.15 e
0.1
0.05
0

Plant extract (mg L-1)

Fig. 3. Effects of medicinal plant extracts on tomato seedling fresh weight in vitro. Control was MS medium without
any plant extract.

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Callus growth index highest CGI (2.5 ± 0.126). In contrast, the lowest
Table 1 indicates that the effect of treatments on CGI was observed in control plants (0.5 ± 0.126).
CGI was significant (p≤0.01). Taraxacum All treatments showed significant differences
officinale extract at 100 mg L-1 resulted in the with the control group (Fig. 4).

3
a
Callus growth index

2.5 ab
abc ab
2 abc abc abc
bc
1.5 bc
c c c
1
d
0.5
0

Plant extract (mg L-1)

Fig. 4. Effects of medicinal plant extracts on tomato callus growth index in vitro. Control was MS medium without any
plant extract.

An interesting finding was the versatility of the internal auxin in callus (0.16 ± 0.0052 μg g-1),
Rheum ribes plant extract, where CGI decreased whereas the Allium jesdianum extract (50 mg L-1)
in response to the increase in extract caused the lowest auxin content in callus (0.062
concentration (Figs. 6C and D). A significant ± 0.0052 μg g-1).
difference in CGI between the control and other As shown in Figure 7, a downward trend was
treatments are evident in Figures 5 and 6. In the observed in the amount of auxin in all treatments
control group, the MS basal medium that compared to the control. An increase in Juniperus
contained some amounts of plant growth sabina and Dorema aucheri extract
regulators (auxin and cytokinin) did not yield concentrations led to a decrease in internal auxin
significant results after a two-week period. content, whereas Conocarpus erectus, Allium
However, the culture medium supplemented with jesdianum, Taraxacum officinale, and Rheum
medicinal plant extracts demonstrated a ribes extracts increased the internal auxin
remarkable outcome in this regard (Figs. 5 and 6). content. Table 2 shows that extracts of medicinal
plants contain high amounts of specific
Auxin content compounds and auxin. The sensitivity of callus
Based on the analysis of variance (Table 1), a cells to auxin may be higher compared to mature
significant difference (p≤0.01) was observed in plants, which could explain the relationship
the auxin content of callus due to the application between the lowest CGI and the highest internal
of plant extracts in vitro. As shown in Figure 7, the auxin content in each sample (Figs. 5 and 6).
control treatment led to the highest amount of

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Fig. 5. (A) Control sample on the first day. (B) Control sample after two weeks. (C) Medium with Taraxacum officinale
extract on the first day. (D) Medium with Taraxacum officinale extract after two weeks. The control was MS medium
without any plant extract.

Fig. 6. Effects of different medicinal plant extracts on tomato callus formation compared to the control group. (A)
Control, (B) Taraxacum officinale 100 mg L-1, (C) Conocarpus erectus, (D) Rheum ribes 50 mg L-1, (E) Allium
jesdianum 50 mg L-1, (F) Juniperus sabina 50 mg L-1, (G) Conocarpus erectus 100 mg L-1, (H) Rheum ribes 100 mg L-1.
The control was MS medium without any plant extract.

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0.18 a
0.16
b
Auxin (μg g-1 IAA)

0.14 bc bc bc
0.12
0.1 de cde cde bcde de de de
0.08 e
0.06
0.04
0.02
0

Plant extract (mg L-1)

Fig. 7. Effects of medicinal plant extracts on tomato callus auxin content in vitro. Control was MS medium without any
plant extract.

Callus weight (Fig. 8), which is probably due to amplified cell

Plant extracts showed significant effects growth and division. The observed increase in CW
(p≤0.01) on CW in vitro (Table 1). As shown in is consistent with the increase in other traits such
Figure 8, the control group showed the lowest CW as callus volume, growth, and internal auxin
(0.05 ± 0.0052 g). Conocarpus erectus extract at levels. As mentioned, callus samples treated with
50 mg L-1 caused the highest CW (0.24 ± 0.0052 Conocarpus erectus extract at 50 mg L-1 exhibited
g). All treatments resulted in an increase in CW the highest CW, whereas the control samples had
the lowest CW (Fig. 8).

0.3
a
Callus weight (g)

0.25
ab
0.2 bc bc bc
bcd bcd bcd
0.15 cd
de de de
0.1 e
0.05
0

Plant extract (mg L-1)

Fig. 8. Effects of medicinal plant extracts on tomato callus weight in vitro. The control was MS medium without any
plant extract.

Total phenol total phenol content of tomato callus in vitro (Fig.

Plant extracts caused no significant difference in 9).

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90
Total phenol (mg gallic acid
ab ab a a
80 abc
70 abc abc abc
bc bc abc
60 c
c
50
g-1 FW)

40
30
20
10
0

Plant extract (mg L-1)

Fig. 9. Effects of medicinal plant extracts on tomato phenol content in vitro. The control was MS medium without any
plant extract.

Total flavonoids (6.58 ± 0.284 mg g-1) (Fig. 10). The present study
A significant difference (p≤0.01) was observed showed that the total flavonoids increased,
from the effects of plant extracts on total parallel to the increase in extract concentration
flavonoids (Table 1). The highest amount of total from 50 to 100 mg L-1 in all treatments except the
flavonoids (12.16 ± 0.284 mg g-1) was observed Conocarpus erectus treatment. In some
in response to 50 mg L-1 Conocarpus erectus treatments, no statistically significant differences
treatment, whereas the lowest was observed in were observed. Total flavonoids decreased,
response to 50 mg L-1 Rheum ribes treatment parallel to the increase in Conocarpus erectus
extract concentration from 50 to 100 mg L-1.
Total flavonoid (mg quercetin g-1 FW)

14
ab a
12
bc c bc bc
10 c c
c cd
cd cd
8 d
6
4
2
0

Plant extract (mg L-1)

Fig. 10. Effects of medicinal plant extracts on total flavonoid content of tomato callus in vitro. The control was MS
medium without any plant extract.

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Antioxidant activity (DPPH) amount of antioxidant activity (63.97 ± 1.283%)

A significant difference (p≤0.01) was observed in was observed in the control group, while the
antioxidant activity (DPPH) among the treatment application of plant extracts improved the
groups and the control (Table 1). The lowest antioxidant activity to various extents (Fig. 11).
Antioxidant activity (DPPH (%))

90 a a a a a a a a
a a a
80
70 b b
60
50
40
30
20
10
0

Plant extract (mg L-1)

Fig. 11. Effects of medicinal plant extracts on antioxidant activity of tomato callus in vitro. Control was MS medium
without any plant extract.

Discussion cell division and enlargement. These processes

Effect of pure plant extract on stem length depend strongly on internal regulators in plants,
Typically, chemicals have a stimulatory effect on including auxin and cytokinin (Taiz et al., 2015).
plants at low concentrations and a suppressive According to Table 2, plant extracts contain
effect at higher concentrations (Qasem and Foy, numerous compounds such as phenols, phenolic
2001). However, the current findings about stem acids, terpenes, and auxin. As previously
length suggest that plant extracts can exhibit mentioned, plants have various compounds that
various effects at different concentrations. can transform into other compounds through
Biological stimulants are substances of natural biological pathways (Vermerris and Nicholson,
origin that have beneficial effects on the growth 2007). These substances can become precursors
and development of plants, their resistance to of plant regulators during intracellular processes,
stress, their yield, and product quality. The following the biosynthesis pathway of growth
physiological effects of these stimulants depend regulators (Negi et al., 2005; Peer and Murphy,
on their composition, which includes various 2007; Nagata et al., 1992). Using medicinal plants
organic and inorganic compounds that plants can to evaluate their impact on the growth of other
use as metabolites and growth regulators plants has been limited to in vitro culture
(Paradikovic et al., 2019). Previous studies on experiments. Previous studies have mainly
crops have shown that seaweed extract can focused on investigating the effects of different
enhance growth vigor in bean seeds (Carvalho et plant compounds, such as adding allelopathic
al., 2014). Bio-stimulants can include a variety of plant powder to agar (ECAM (equal compartment
compounds, such as amino acids, saponins, agar medium)) (Mushtaq et al., 2020). In another
vitamins, PGR, and plant compounds, as study, researchers compared plant compounds
demonstrated in previous research (Paradikovic and growth regulators that improved the micro-
et al., 2019). However, caffeic acid reportedly propagation process in acacia plants (Robinia
decreased stem length at high concentrations pseudoacacia L.) through tissue culture. Plant
(Batish et al., 2008). As mentioned earlier, plant samples were supplemented with 0, 1000, 2000,
height is influenced by growth and determined by and 3000 mg L-1 Ascophyllum nodosum brown

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algae extract in MS medium. The results showed accurately capture the effectiveness of these
that the Ascophyllum nodosum extract (1000 mg extracts on plant growth.
L-1) led to the highest number of roots among Our study provided valuable insights into the
other treatments (Kaviani et al., 2016), which potential use of herbal plant extracts in
confirms the stimulatory effects observed in our promoting plant growth and highlighted the need
findings. It should be noted that during the for further research in this field. Our findings are
process of starch decomposition and consistent with previous studies that have
germination, seeds produce free radicals within demonstrated a decrease in sesame (Sesamum
themselves due to biochemical and physiological indicum L.) fresh weight in response to an
interactions (Taiz et al., 2015). These free radicals aqueous extract of billygoat weed (Ageratum
can affect growth through their impact on cell conyzoides L.) (Natarajan et al., 2014).
organelles. However, medicinal plants are rich in Additionally, a decrease occurred in mustard
compounds that counteract these radicals, which plant seed germination, root length, branch
means that seedlings exposed to these specialized length, chlorophyll content, fresh weight (FW),
compounds absorb fewer radicals during seed dry weight (DW), and relative water content
germination and growth, leading to safer and (RWC) in response to aqueous extracts of senna
faster growth. Similar to the current research, tora roots, stems, and leaves (Cassia tora L.)
previous findings showed that (Sarkar et al., 2012). The differences in growth in
cylindrospermopsin (CYN) had various effects at our study can be attributed to variations in the
different concentrations, thus being consistent biochemical and phytochemical profiles of the
with our results (Garda et al., 2015). Other studies extracts (Table 2). These differences relate to
on plant extracts affecting other plants have phenolic, antioxidant, and hormonal substances.
shown that aqueous extracts of Calotropis Research has also shown that using aqueous
procera L. can increase germination and seedling seaweed extract can increase plant biomass in
growth in corn (Naz and Bano, 2014). various plants (De Saeger et al., 2020). These
Interestingly, the same extract had an inhibitory findings are consistent with our results and
effect on wild cabbage (Brassica oleracea L.) and suggest that the application of different plant
botrytis variety (Gulzar and Siddiqui, 2017). This extracts can have varying effects on plant growth
observation is crucial because it reveals that and development. Therefore, it is crucial to
herbal compositions can affect plants differently, carefully select and evaluate appropriate plant
which was also evident in our study. For instance, extracts as they affect specific plants and as
the Juniperus sabina extract (at both researchers aim to achieve optimal growth.
concentrations) stimulated growth, Overall, our study contributes to the current
while Taraxacum officinale at 100 mg L-1 knowledge on potential plant extract usage as a
decreased stem growth and length. Furthermore, natural and sustainable alternative to traditional
another study showed that aqueous extracts chemical fertilizers in promoting plant growth
of Celosia argentea obtained from its leaf, stem, and development.
and flower increased α-amylase activity in lentils
(Lens culinaris Medic) (Kengar and Patil, 2018). Effects of pure plant extract on callus growth
It can be concluded that plant extracts have index
different effects due to variations in chemical During cell division and growth, plant cells
substances and their various levels of PGRs. produce free radicals that can target intracellular
organelles, thus reducing their efficiency. This
Effect of pure plant extract on seedling fresh effect is particularly evident when preparing
weight explants for tissue culture (Taiz et al., 2015).
The changes in SFW further confirmed that However, high levels of phenolic and antioxidant
different extracts have varying effects on plant compounds in the plant extracts (Table 2) can
growth and development. It is important to note neutralize the effects of these free radicals and
that plant growth and development are complex promote better cellular activity and growth.
processes and can change in response to multiple Additionally, compounds in the extracts serve as
factors, including the genetic base of a plant, biological stimulators for plant growth as they
environmental conditions, and the presence of contain abundant precursors of plant growth
various signaling molecules and growth regulators, leading to increased callus formation.
regulators. Therefore, research on the effects of A correct balance between auxin and cytokinin
medicinal plant extracts on tomato plant growth hormone levels is crucial for callus formation
and development needs to consider these factors (Taiz et al., 2015).
and carefully design experiments that can

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Effects of pure plant extract on auxin content confirmed that these compounds effectively
High levels of auxin and cytokinin can direct cell induce some hormonal changes in plants. For
division in specific ways. However, the ratio instance, some flavonoid aglycones can disturb
between these two regulators is the final the polar transport of auxin (Brunn et al., 1992).
determining factor in growth (Taiz et al., 2015). In another study, benzoic acid and its derivatives
Cell division and cell enlargement are two (such as 3, 4-dihydroxybenzoic acid (DHB))
important mechanisms that influence growth, stimulate callus formation at low concentrations
and they are influenced by growth regulators but have a suppressive effect on root growth at
such as auxin, cytokinin, gibberellin, ethylene, high concentrations in tobacco plants
abscisic acid, jasmonic acid, brassinosteroids, and (Mucciarelli et al., 2000). It is important to note
salicylic acid. Research over many years has that higher auxin concentrations stimulate
demonstrated that two regulators, auxin and ethylene production (Taiz et al., 2015). Therefore,
cytokinin, play a vital role in cell division. if we attribute the role of auxin analog to 3, 4-
Therefore, regarding the developmental process, DHB, the suppressive effect is proven at higher
these two regulators can be relative indicators for concentrations of this compound. This fact may
physiological studies (Taiz et al., 2015). Auxin, a explain the concentration-dependent effect of
regulator of cell growth, is a basic factor in the cell allelochemicals that mimic or affect auxin
cycle. A study demonstrated that when tobacco synthesis. In contrast, other studies have reported
plant cells were cultured without an auxin source that some phenolic compounds can prevent auxin
in the culture medium, the cells performed DNA degradation, leading to auxin accumulation (Mato
centration but could not complete the cell cycle et al., 1994; Cvikrova et al., 1996). However, gene
and stopped in the G1 phase. Later, auxin was studies currently under investigation in other
added to the culture medium, and the expression studies may provide stronger confirmations.
of genes was checked, revealing that arcA gene Growth is not solely under the control of a plant
expression was induced (Ishida et al., 1993). growth regulator, and the ratio of auxin to
Auxin acts as a growth stimulant in low cytokinin is an important factor for growth
concentrations, a growth inhibitor, and a regulation (Taiz et al., 2015). Some compounds
herbicide (2-4-D) in high concentrations (Taiz et can affect the biosynthesis pathway and
al., 2015). Medicinal plants contain many production of a compound, such as lovastatin,
compounds (Vermerris and Nicholson, 2007) that which inhibits the production of mevalonic acid
can affect auxin content in various ways. Some and is a significant anti-cytokinin that prevents
compounds can act as inhibitors of auxin normal plant growth (Laureys et al., 1998). It is
biosynthesis, while others can stimulate auxin valuable because it highlights that compounds in
production. For example, flavonoids, a type of plant extracts can affect the biosynthesis pathway
plant pigment, can increase auxin levels in some of plant growth regulators in specific
plants by up regulating the expression of auxin concentrations, as shown in the results of the
biosynthesis genes (Peer and Murphy, 2007). In present study. For example, coumaric acid is one
contrast, some alkaloid compounds, such as of the dominant and important compounds in
cinchonine and quinine, can inhibit auxin dandelion (Piccolella et al., 2023), and it can
biosynthesis in plants (Nagata et al., 1992). The transform into salicylic acid in its biochemical
relationship between plant compounds and auxin pathway (Taiz et al., 2015). Salicylic acid can act
content in plants is complex and varies depending as a stimulant at low concentrations and a growth
on the specific compound and plant species inhibitor at high concentrations (Horvath et al.,
involved. Plant extracts can change auxin content 2007). Various studies have shown that plant
depending on the type of plant or concentration compounds (plant extracts) can suppress or
used (Fig. 7). For instance, past studies with stimulate enzymes involved in the biosynthesis of
elevated carbon dioxide have demonstrated that plant compounds. For instance, the production of
polyphenols increase growth by intensifying the IAA is suppressed and stimulated by different
effects of auxin by preventing IAA concentrations of different plant extracts
decarboxylation (Tomaszewski and Thimann, (ALHaithloul et al., 2022; Abou El-Ghit et al.,
1996). Sinapic and ferulic acids can increase 2016), or the production of ethylene and ABA
growth regulators by preventing IAA increases with an increase in allelopathy
decarboxylation, while monophenols can do the (Bogatek et al., 2005). In another study, Artemisia
opposite and increase IAA decarboxylation. argyi water extract caused auxin accumulation in
Phenolic compounds can change growth roots (Li et al., 2022). Considering that growth
regulator contents (Tomaszewski and Thimann, conditions (light, temperature, culture medium,
1996). Studies on different plant compounds samples) were identical, the differences in
internal auxin content were attributable to plant

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extracts. Further enzymatic and molecular unguiculata L., which is consistent with a recent
investigations can gain a deeper understanding of study by Bingol et al. (2022). It showed that
this process. extracts of Xanthoparmelia somloensis had
varying effects on the growth and physiology of
Effect of pure plant extract on callus weight tomato plants. A study by Laureys et al. (1998)
Several studies have reported that plant showed that lovastatin, a compound against
compounds can affect plant growth and mevalonic acid, prevents cytokinin production. It
development by regulating various physiological highlighted the importance of studying the effects
processes, including cell growth and division. For of plant compounds on other plants, as observed
example, Alsharekh et al. (2022) suggested that in the present study. A study by Hassanzadeh
different plant compounds, such as secondary (2014) suggested that the stimulation of the plant
metabolites, can act as growth regulators by immune system (SAR) may be the main factor
influencing cell division and differentiation. responsible for the observed effects in the present
Similarly, a study by Bashar et al. (2023) study, which is consistent with a recent study by
demonstrated that secondary metabolites can Li et al. (2021) that the application of plant
serve as allelochemicals and affect plant growth extracts can activate the SAR pathway and
and development by regulating gene expression enhance the resistance of plants to various
and metabolic pathways. These results suggest stresses.
that plant compounds in the extracts can affect However, further studies are needed to elucidate
cell growth and division, leading to changes in specific mechanisms underlying the observed
callus weight. The present study is in line with effects in the present study. Overall, the present
previous research on the effects of plant study and recent research provide insights into
compounds on growth regulation. The results the mechanisms underlying the effects of plant
suggest that plant extracts containing specific extracts on neighboring plants.
compounds can be used as growth regulators to
enhance plant growth and development. Further Effect of pure plant extract on total
research is needed to explore the mechanisms of flavonoids
action of these compounds and their potential Our findings are consistent with a recent study by
applications in agriculture and medicinal Mutale-Joan et al. (2020), which showed that the
applications. concentration of microalgae extracts can
significantly affect the metabolite profile in
Effect of pure plant extract on total phenol tomatoes. Also, this finding is in agreement with a
The concept of allelopathy, which refers to the recent study by El-Shora et al. (2022), which
ability of plants to influence the growth and demonstrated that the application of Rumex
development of neighboring plants through the dentatus L. plant extracts can increase the
release of chemical compounds, has been well- production of flavonoids in Portulaca oleraceae.
established in the literature (Mushtaq et al., An ex vitro study by Salvi et al. (2019) also
2020). A study by Singh et al. (2006) showed that foliar application of seaweed extract
demonstrated that alpha-pinene, a major increased the amount of anthocyanin and
compound in Juniperus sabina, increased the flavonols in grape plants, further supporting the
activity of antioxidant enzymes. Singh et al. findings of the present study. However, as
(2006) showed that alpha-pinene effectively mentioned earlier, this may be due to the
enhanced the activity of antioxidant enzymes in stimulation and suppression of the plant's
rice plants under drought stress. immune system, and further enzyme studies are
Furthermore, a study by Khatun et al. (2023) needed to provide stronger evidence. A study by
demonstrated that extracts from Trewia Kengar and Patil (2018) demonstrated that the
nudiflora Linn. had a significant effect on the aqueous extracts of the leaf, stem, and flower
growth and physiology of lettuce (Lactuca of Celosia argentea L. increased the α-amylase
sativa L.) and foxtail fescue (Vulpia myuros L.). activity of lentils, which supports the argument
The bell-shaped response observed in the present that plant extracts can have varying effects on the
study, where a low concentration of compounds is growth and development of neighboring plants,
stimulatory while a high concentration inhibits as observed in the present study.
growth, is common in plant biochemistry (Batish Finally, the results of the present study suggest
et al., 2008). A study by Desoky et al. (2020) that allelopathic substances can have varying
demonstrated the differential effects of different effects on different plant species, depending on
plant extracts on the enzymatic and non- the type of plant and its concentration. This
enzymatic defense systems in Vigna finding is consistent with previous studies

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(Mushtaq et al., 2020) and highlights the compounds found in various plant species
importance of investigating specific compounds increases the chance of achieving desirable
responsible for the observed effects in future results by different applications. However, as the
studies. results of this study demonstrated, the effects of a
plant extract on plant growth and development
Effect of pure plant extract on antioxidant vary significantly from plant to plant, as well as on
activity (DPPH) the type and concentration of the extract used.
Reactive oxygen species (ROS) are compounds The results showed that the plant extracts
with high energy levels that can have destructive significantly influenced the growth of seedlings
effects on cells, causing disruptions in growth from seeds, sample regeneration, and tomato
processes (Taiz et al., 2015). Antioxidant activity callus formation efficiency. This relationship is
can reduce the harmful effects of these important to mention that the main difference is
compounds and create a low-stress environment between the control group and other treatments,
for cell growth. The biochemical profile of herbal although in some treatments there was no
treatments shown in Table 2 also indicates that significant difference. Therefore, achieving the
the plant extracts are rich in phenolic compounds desired results from plant extracts requires
and antioxidants, which may contribute to stress- extensive and targeted research, taking into
free growth for the plant. The observed increase account the type and composition of the extract as
in antioxidant activity is consistent with the well as the plant species. The findings of this
findings of several previous studies. For example, study highlighted the potential of natural plant
Desoky et al. (2020) reported that foliar extracts as effective and environmentally friendly
application of the extracts of two medicinal biological compounds. However, to fully realize
plants, fennel (Foeniculum vulgare L.) and their potential, further research is needed to
toothpick (Ammi visnaga L.), at a concentration of understand their mechanisms of action and
2000 mg L-1, increased the antioxidant potential optimize their use in different plant species and
compared to the control plant. Similarly, a study applications. Generally, this study underscores
by Singh et al. (2015) showed that the internal the importance of conducting further research to
chemical compounds of tobacco leaves (N. uncover the full potential of natural plant extracts
plumbaginifolia) stimulated some enzymes (CAT as sustainable alternatives to synthetic chemicals
and SOD) in sunflowers. These findings suggest and to promote the development of
that plant extracts can cause changes in environmentally friendly approaches to
specialized compounds in plants, leading to an agriculture and crop production. Finally, these
increase in antioxidant activity. Further enzyme plant extracts in a tissue culture medium are
studies can be conducted in future stages of the advisable for inducing an increase in callus
current research to investigate the potential formation.
mechanisms underlying the observed effects on
antioxidant activity. For example, the activities of Author contributions
antioxidant enzymes such as catalase, peroxidase, Author contributions as mentioned below: OS:
and superoxide dismutase can be measured to Methodology, Formal analysis, Investigation, Data
provide insights into the mechanisms of action of curation, writing original draft. AGH: Project
the plant extracts. In conclusion, the present advisor in sabbatical leave, Validation, Writing-
study showed that plant extracts can increase review and editing. AH: Project administration,
antioxidant activity in plants. The findings are Validation, Writing-review and editing. HS:
consistent with previous studies highlighting Consulting in data analysis. VEM: Consulting in
plant extracts as a natural and sustainable means data analysis.
of enhancing plant growth and development.
Acknowledgements
Conclusion The authors gratefully acknowledge the support
In conclusion, natural plant extracts are a of the Ministry of Science and Research. The
promising source for various applications due to authors acknowledge Mrs. Fatemeh Taheri for her
their compatibility with the environment and help.
potential to replace synthetic chemicals. The
indiscriminate use of chemical-based fertilizers Funding
and pesticides can have environmental Partial financial support was received from Iran
implications, emphasizing the importance of Ministry of Science and Research.
research that focuses on introducing herbal
compounds. The high diversity of natural Conflict of Interest
The authors indicate no conflict of interest in this

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work. allelopathy stress. In Fourth World Congress on

Allelopathy.
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