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Vol.35, No.3, pp.289-299, 2023
Journal of Agricultural, Life and Environmental Sciences
RESEARCH ARTICLE
pISSN 2233-8322, eISSN 2508-870X
https://doi.org/10.22698/jales.20230023
Assessment of the Growth and Ornamental Quality of Senecio
haworthii (Sweet) Sch.Bip. Grown under Different Day/Night
Temperatures
Jae Hwan Lee1,2, Sang Yong Nam3,4*
1Integrated Ph.D. Student, Department of Environmental Horticulture, Sahmyook University, Seoul 01795, Korea
2Researcher, Natural Science Research Institute, Sahmyook University, Seoul 01795, Korea
3Professor, Department of Environmental Horticulture, Sahmyook University, Seoul 01795, Korea
4Director, Natural Science Research Institute, Sahmyook University, Seoul 01795, Korea
*Corresponding author: Sang Yong Nam (E-mail: namsy@syu.ac.kr)
A B S T R A C T
Received: 31 July 2023
Revised: 7 September 2023
Accepted: 8 September 2023
Senecio haworthii is a succulent plant belonging to the Asteraceae family. Optimal growth
conditions for this species have not been well-established. In this study, we investigated the
optimal day/night temperature range for enhancing the growth and ornamental quality of S.
haworthii. Four day/night temperature treatments were designed as follows: 20/15, 24/19, 28/23,
and 32/27°C. Based on the plant sizes and biomass analysis, the best growth performance was
determined to be achieved under a relatively low-temperature treatment of 20/15°C. Conversely,
the 32/27°C treatment resulted in the lowest growth rates, likely attributed to decreased leaf
function caused by prolonged exposure to high-temperature conditions. Additionally, leaf color
analysis using the CIELAB color space revealed that CIELAB L*, representing lightness, and
CIELAB a*, indicating green-red opponent colors, proportionally increased with the rise in
day/night temperature levels. Previous studies have reported an inverse relationship between L*
and plant growth parameters. Consistent with previous research, the treatment at 32/27°C, which
exhibited the lowest growth, showed higher L* values. Conversely, the CIELAB b* value,
representing blue-yellow opponent colors, was highest under the 20/15°C treatment, suggesting
leaf-yellowing tendencies in relatively lower temperature conditions. Based on the color
difference analysis using CIE76 color difference (ΔE*ab), temperatures of 20/15 and 24/19°C
were used as reference points; under these conditions, the temperatures of 28/23 and 32/27°C were
evaluated as having a ‘small color difference’. Therefore, even with significant differences in
day/night temperatures, no noticeable differences were concluded to be found in leaf color. Based
on these findings, cultivating S. haworthii in relatively low-temperature conditions of around
20/15°C is concluded to be advantageous for enhancing both plant growth and ornamental quality
compared to higher temperature conditions such as 32/27°C.
Keywords: Asteraceae, Biomass, Caputia tomentosa, CIELAB, Color difference
Ⓒ Journal of Agricultural, Life and Environmental Sciences. This is an Open Access article distributed under the terms of the Creative Commons
Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/4.0) which permits unrestricted non-commercial use, distribution,
and reproduction in any medium, provided the original work is properly cited.
290 ∙Journal of Agricultural, Life and Environmental Sciences Vol. 35, No. 3, 2023
Introduction
Senecio haworthii, also known as Caputia tomentosa, is a succulent plant belonging to the Asteraceae family. S.
haworthii exhibits high drought tolerance (WOS, 2023) and possesses a remarkable white hairy covering over the
entire plant, presumed to serve as a mechanism for reflecting intense direct sunlight in arid and high-temperature
environments, resulting in its striking and fascinating appearance, which makes it highly valuable for horticultural
interest. Plants within the Senecio genus are known to contain pyrrolizidine alkaloids, which exhibit toxic
properties to animals and humans, categorizing them as poisonous plants (Cheeke, 1988; McLean, 1970). Various
cases of Senecio poisoning have been reported for about a hundred years (Basson, 1987; Gava and Barros, 1997;
Giles, 1983; Selzer and Parker, 1951; Stillman et al., 1977; Van Schalkwyk et al., 2021; Willmot and Robertson,
1920). On the other hand, from a pharmacological perspective, Senecio species have shown effects in anti-
microbial and anti-bacterial (Pérez et al., 1999), anti-fungal (Loizzo et al., 2004), anti-inflammatory (Pérez González
et al., 2013; Zahoor et al., 2023), anti-tumor (Dou et al., 2017), anti-diabetic (Ayoola et al., 2019; Mahmoud et al.,
2011), and anti-oxidant (Faraone et al., 2018; Lounis et al., 2018), suggesting their potential for future medicinal
purposes. Therefore, under appropriate management, Senecio species can be considered suitable for use as
ornamental succulents or medicinal crops. Despite these advantages, little is known about the optimal growth
conditions for Senecio species, necessitating research to identify the most suitable environment for cultivation.
Growth temperature is one of the critical factors influencing plant growth and quality (Morison and Lawlor,
1999; Oh et al., 2022). Each plant species has specific temperature preferences and study on the day/night
temperature range is challenging due to its complex nature, requiring various experimental approaches. Succulent
plants have developed unique physiological abilities to adapt to extreme habitats and dry environments (Nam et al.,
2016). Succulent plants have a unique carbon dioxide assimilation pathway called Crassulacean acid metabolism
(CAM) (Grace, 2019; Ting, 1989; Winter and Holtum, 2014), and they are known to be particularly sensitive to
night temperature increases during cultivation (Sutton, 1975; Yamori et al., 2014). Particularly, high night
temperatures have been reported to significantly inhibit CO2 nocturnal uptake and accumulation of titratable acidity
in succulent plants (Neales, 1973). Therefore, determining the suitable growth temperatures for each succulent
species is necessary. Previous studies have investigated the effect of temperature on flowering in S. bellum (Heinze,
1971), and the impact of temperatures on the growth and anthocyanin content in Echeveria species (Cabahug et al.,
2019). However, studies on the optimal day/night temperature setting for cultivating Senecio species are virtually
nonexistent, necessitating research to address this knowledge gap.
For ornamental plants, leaf color is an important factor that significantly influences their ornamental value in
addition to plant growth. Leaf color serves as an element enabling consumers to intuitively evaluate the external
quality of plants, it is an essential consideration (Lee and Nam, 2023b). CIELAB is a color space created by the
International Commission on Illumination (CIE) in 1976 (Lee et al., 2022c). Developed to overcome the limitations
of Hunter Lab, which was introduced in 1958, CIELAB is characterized by the inclusion of asterisks in its notation,
represented as L*, a*, and b* as opposed to L, a, and b in Hunter Lab (Lee and Nam, 2022a). CIELAB has been
Jae Hwan Lee and Sang Yong Nam / Assessment of the Growth and Ornamental Quality of Senecio haworthii (Sweet) Sch.Bip. ~ ∙291
widely applied in various horticultural studies, such as external quality evaluation of fruits and vegetables (Cho et
al., 2021; Kim et al., 2022; Lee et al., 2022a; 2022e), ornamental flower crops (Jang et al., 2023; Park et al., 2023;
Shin et al., 2022), and succulent plants (Lee, 2023; Lee and Nam, 2022b; Nam et al., 2022), and serves as an
excellent parameter for describing the qualitative quality of plants. Therefore, CIELAB is expected to be useful for
evaluating leaf color in succulent plants.
Accordingly, we selected S. haworthii, one of the most widely distributed Senecio species in the South Korean
ornamental succulents market, as our experimental plant, and investigated and analyzed the effects of day/night
temperatures on the growth and leaf color of S. haworthii.
Materials and Methods
Plant Material
Senecio haworthii cultivated within the experimental greenhouse of the Department of Environmental
Horticulture at Sahmyook University, were used for this study. The plant sizes, measured as shoot height and shoot
width, were about 5 and 3 cm, respectively.
Day/Night Temperature Treatments
This study was conducted for 10 weeks, inside growth chambers (KGC-175VH, KOENCON, South Korea)
located in the experimental greenhouse of the Department of Environmental Horticulture at Sahmyook University
in South Korea. Four different temperature treatments were designed, with day/night temperatures set at 20/15,
24/19, 28/23, and 32/27°C, respectively. The photoperiod was set at 14/10 hours (day/night). We utilized
light-emitting diodes (LEDs) with a ratio of 4:2:8:2 in red:blue:white:far-red as the internal light sources for the
growth chambers. The light intensity inside the growth chamber was adjusted to a photosynthetic photon flux
density (PPFD) of 500 µmol m-2 s
-1 using a portable spectroradiometer (SpectraPen mini, Photon Systems
Instruments, Czech Republic). The plants were planted in a succulent media, consisting of decomposed granite,
river sand, and fertilized horticultural substrate (Hanareumsangto, Shinsung Mineral, South Korea) mixed in a ratio
of 6:3:1 (v/v/v). Rectangular pots measuring 48.5 × 33 × 8 cm (length × width × height) were used. The plants were
watered once a week with 1.5 L of purified water per pot.
Parameters and Measurement Methods
The parameters assessed in this study included survival rate, shoot height, shoot width, stem diameter, root
length, number of leaves, leaf length, leaf width, shoot and root fresh weight, shoot and root dry weight, shoot and
root moisture content, CIELAB values (L*, a*, and b*), and CIE76 color difference (ΔE*
ab). Shoot height was
measured as the distance from the soil surface to the highest point of the plant. Shoot width was measured as the
widest part of the plant when observed from above. Root length was measured as the length of the longest root
292 ∙Journal of Agricultural, Life and Environmental Sciences Vol. 35, No. 3, 2023
among the plant’s roots. Fresh weight was measured after the soil attached to the plant was washed off and then
naturally air-dried in a sealed space for 24 hours. Dry weight was measured after heat drying the plants using a heat
drying oven (HK-DO135F, HANKUK S&I, South Korea) at 85°C for 24 hours. Moisture content was calculated
based on the comparison between fresh weight and dry weight, and it was determined using the following equation
(Eq. 1).
⋅
(1)
(
is plant moisture content,
is fresh weight, and
is dry weight)
For CIELAB measurements, we referred to the leaf color measurement method described by Lee et al. (2022b)
and obtained CIELAB L*, a*, and b* values with a spectrophotometer (CM-2600d, Konica Minolta, Japan) set to
CIELAB D65/10° with the specular component included (SCI). To convert the CIELAB L*, a*, and b* values to
visually assessable colors, we used the Converting Colors tool operated by Zettl (2023) for color conversion. To
compare the effect of temperature levels on the leaf color of S. haworthii in the day/night temperature treatments,
we calculated color differences (ΔE*
ab) using CIE76 color difference formula, setting each day/night temperature
level as a reference. The formula (Eq. 2) used for ΔE*
ab in this study was as follows (CIE, 2004).
(2)
(In this study, ΔE*
ab ≤ 1.5 is regarded as ‘no color difference’ or ‘subtle color difference’, 1.6-3.0 as ‘very small
color difference’, 3.1-6.0 as ‘small color difference’, 6.1-9.0 as ‘color difference’, 9.1-12.0 as ‘big color difference’,
and ≥ 12.1 as ‘very big color difference’ or ‘completely different color’)
Data Analysis
The results were analyzed using SAS 9.4 (SAS Institute, USA) with analysis of variance (ANOVA). Duncan’s
multiple range test at the p < 0.05 significance level was performed for mean comparisons. The study was in a
completely randomized design, with five plants assigned to each treatment in three replications.
Results and Discussion
Plant Sizes Analysis
Senecio haworthii, subjected to different day/night temperatures, exhibited diverse growth responses (Fig. 1 and
Table 1). The survival rate of S. haworthii was 100% within the day/night temperature range of 20/15-28/23°C;
however, at 32/27°C, the survival rate declined to 86.6%. The results of the plant sizes analysis revealed that shoot
Jae Hwan Lee and Sang Yong Nam / Assessment of the Growth and Ornamental Quality of Senecio haworthii (Sweet) Sch.Bip. ~ ∙293
height and shoot width showed no significant differences within the day/night temperature range of 20/15-28/23°C,
based on Duncan’s multiple range test. Consequently, it is estimated that the shoot sizes of S. haworthii plants are
minimally affected unless exposed to high temperatures, such as 32/27°C. The stem diameter was found to be the
thickest at 0.56 cm under 20/15°C conditions. Meanwhile, the root length reached its maximum at 9.29 cm under
28/23°C conditions, while under the relatively higher temperature of 32/27°C, it sharply decreased to 6.35 cm. The
orchid cultivar Phalaenopsis cv. KS Little Gem, known for its Crassulacean acid metabolism (CAM) plant, was
reported to exhibit the lowest root length at the highest temperature treatment of 30°C within the temperature range
of 20-30°C (Vo et al., 2021). The specific root length, a parameter used for fine roots, was reported to exhibit a
negative response as the growth temperature increased (Ostonen et al., 2007). Consistent with this previous study
report, it was observed that at excessively high temperatures beyond the plant’s optimal growth range, the root
length could sharply decrease. The number of leaves reached its maximum at 26.2 under 20/15°C conditions, while
conversely, at 32/27°C treatment, it exhibited the lowest number of leaves at 5.7, similar to other parameters
indicating plant size. The Peperomia caperata, within the temperature range of 15-21°C, an increase in temperature
was reported to lead to an increase in leaf length, leaf width, and leaf number (Brøndum and Friis, 1990). For
succulent plants, it has been reported that with an increase in night-time temperature, the stomatal conductance
Fig. 1. Shape of Senecio haworthii plants as affected by day/night temperatures for 10 weeks.
Table 1. Survival rate and plant growth parameters of Senecio haworthii as affected by day/night temperatures for 10
weeks
Temperatures
(day/night)
Survival
rate
(%)
Plant sizes (cm)
No. of leaves
Leaf sizes (cm)
Shoot
height
Shoot
width
Stem
diameter
Root
length Length Width
20/15°C 100.0 az8.22 a 4.34 a 0.56 a 8.32 ab 26.2 a 3.43 a 0.77 a
24/19°C 100.0 a 8.19 a 4.76 a 0.49 b 8.27 ab 24.3 ab 3.38 a 0.75 a
28/23°C 100.0 a 8.14 a 4.17 a 0.46 b 9.29 a 20.8 b 3.62 a 0.68 b
32/27°C 86.6 b 7.59 b 1.13 b 0.48 b 6.35 b 5.7 c 2.37 b 0.45 c
Significancey*** ** *** ** * *** *** ***
zMeans separations within columns were performed using Duncan’s multiple range test, with a significance level set at p < 0.05.
y*, **, and ***: significant at p < 0.05, 0.01, or 0.001, respectively.
294 ∙Journal of Agricultural, Life and Environmental Sciences Vol. 35, No. 3, 2023
pattern is altered, resulting in reduced photosynthetic efficiency (Neales, 1973). Succulent species with thicker leaf
tissues, like S. haworthii, are considered to experience decreased leaf function under excessively high-temperature
conditions compared to their optimal growth temperature (Medina et al., 2021; Schoffl et al., 1998; Schroda et al.,
2015), emphasizing the need for caution when setting prolonged exposure to high-temperature conditions. The leaf
length showed comparable significant levels within the temperature range of 20/15-28/23°C, consistent with the
results of shoot height and shoot width. However, leaf width, unlike the leaf length, appeared broader at 0.77 and
0.75 cm under 20/15-24/19°C treatments, respectively. A previous study on Chrysanthemum cv. Orange Egg
subjected to shade with temperature treatments showed a similar result to this study, as leaf length increased with a
decrease in relative growth temperature (Park and Kim, 2021). According to Vo et al. (2021), the P. cv. KS Little
Gem exhibited a decrease in both leaf length and width as the growth temperature increased within the temperature
range of 20-30°C, which aligns with the findings of this study.
In conclusion, when considering the overall analysis of plant sizes, a growth temperature of 20/15°C was deemed
most favorable for significant enlargement of the shoot part of the plants. However, for root length, the results were
contrary to those of the plant shoot sizes, as a growth temperature of 28/23°C was considered most advantageous
for significant root elongation.
Plant Biomass Analysis
In the results of biomass analysis, the shoot fresh weight was heaviest in the 20/15°C treatment, measuring 16.15
g (Fig. 2). Conversely, in the 32/27°C treatment, it was considerably lower at 0.97 g, representing a reduction of
approximately 94.0% compared to the 20/15°C treatment. On the other hand, the root fresh weight did not show
significant differences between the 20/15-28/23°C treatments, but it was the lowest in the 32/27°C treatment,
measuring 0.63 g. This finding aligns with a previous study on the P. cv. KS Little Gem, where the lowest root
biomass was observed in the treatment with the highest growth temperature of 30°C (Vo et al., 2021), which is
consistent with the results of our study. Similarly to the shoot fresh weight, the shoot dry weight was highest in the
20/15°C treatment at 1.12 g. Conversely, in the 32/27°C treatment, it was the lowest at 0.31 g, indicating a
significant reduction of approximately 72.4% compared to the 20/15°C treatment. This considerable decrease in
shoot biomass allocation under high-temperature conditions indicates likely heat damage to the shoot part,
suggesting that the growth conditions exceed the optimal growth temperature range. The root dry weight was
relatively high in the 20/15 and 24/19°C treatments, measuring 0.35 and 0.34 g, respectively. Despite the relatively
lesser decrease in root biomass compared to shoot biomass, the negative effects on root development under
high-temperature conditions appeared to be similar. Both shoot and root moisture content did not show significant
differences within the temperature range of 20/15-28/23°C. However, in the 32/27°C treatment, both shoot and root
moisture contents were the lowest, measuring 68.1% and 54.9%, respectively. In a previous study by Way and Oren
(2010), trees grown in high-temperature environments were reported to allocate less biomass to the roots.
According to the results of the biomass analysis, S. haworthii exhibited higher biomass in both shoot and root
Jae Hwan Lee and Sang Yong Nam / Assessment of the Growth and Ornamental Quality of Senecio haworthii (Sweet) Sch.Bip. ~ ∙295
under the 20/15°C treatment. Therefore, it appears that S. haworthii performs more carbon dioxide assimilation
under lower temperature conditions. Based on the results of this study, we suggest cultivating S. haworthii at a
relatively low temperature, approximately around 20/15°C, to encourage the growth of sturdy plants. However, it is
important to note that further research may be necessary to explore the potential for even better growth at lower
temperatures. Additionally, it is imperative to take into account that uniform irrigation conditions were consistently
applied throughout this study, regardless of growth temperatures.
Leaf Color Analysis
In the leaf color analysis using CIELAB values, the lightness parameter of CIELAB L* was observed to increase
slightly with the rise in S. haworthii’s growth temperature, measuring 78.01 and 77.29 for the 28/23 and 32/27°C
treatments, respectively (Table 2). Previous studies have reported an inverse relationship between L* and plant
growth parameters (Lee and Nam, 2023a; Lee et al., 2022d), and our study also showed similar results. The
green-red opponent color parameter of CIELAB a* was highest at -0.61 under the 32/27°C treatment, suggesting
that under high-temperature conditions, the leaf color tends to shift towards the red. On the other hand, the
blue-yellow opponent color parameter of CIELAB b* was highest at 7.80 in the 20/15°C treatment, indicating a
subtle yellowing tendency of the leaves under lower temperature conditions. Furthermore, the CIE76 color
Fig. 2. Plant biomass and moisture content of S. haworthii as affected by day/night temperatures for 10 weeks: A) shoot
fresh weight; B) shoot dry weight; C) shoot moisture content; D) root fresh weight; E) root dry weight; and F) root
moisture content. Vertical bars indicate standard error and asterisks (*, **, and ***) indicate significance at p < 0.05,
0.01, or 0.001, respectively. Different lowercase letters indicate significant differences at p < 0.05 based on Duncan's
multiple range test.
296 ∙Journal of Agricultural, Life and Environmental Sciences Vol. 35, No. 3, 2023
difference (ΔE*
ab) analysis revealed various leaf color differences in S. haworthii according to growth temperature.
Specifically, the ΔE*
ab values between the 20/15 and 24/19°C treatments were ΔE*
ab = 1.76, and between the
28/23 and 32/27°C treatments, it was ΔE*
ab = 1.43, indicating ‘very small color difference’ and ‘subtle color
difference’, respectively, within each set of treatments. However, the ΔE*
ab values between the 20/15 and 24/19°C
treatments and the 28/23 and 32/27°C treatments were in the range of ΔE*
ab = 4.47-4.95, showing ‘small color
difference’ Therefore, even with significant differences in day/night temperature levels, noticeable differences in
leaf color were not observed.
Based on the comprehensive evaluation of the leaf color analysis results, S. haworthii exhibited an increase in L*
and a* at higher day/night temperature ranges, while b* showed a decreasing trend. Additionally, leaf color
differences were evaluated as ‘small color difference’ when comparing the 28/23 and 32/27°C treatments to the
20/15 and 24/19°C treatments based on ΔE*
ab values.
Conclusion
In this study, we investigated the optimal day/night temperature range for enhancing the growth and ornamental
quality of Senecio haworthii. Four day/night temperature treatments were designed as follows: 20/15, 24/19, 28/23,
and 32/27°C, respectively. Based on the plant sizes and biomass analysis, it is determined that the best growth
performance can be achieved under the relatively lower temperature treatment of 20/15°C. Conversely, the 32/27°C
treatment resulted in the lowest growth rates, likely attributed to decreased leaf function caused by prolonged
exposure to high-temperature conditions. Additionally, leaf color analysis using the CIELAB color space revealed
that CIELAB L*, representing lightness, and CIELAB a*, indicating green-red opponent colors, proportionally
increased with the rise in day/night temperature levels. Previous studies have reported an inverse relationship
between L* and plant growth parameters. Consistent with this finding, the treatment at 32/27°C, which exhibited the
lowest growth, showed higher L* values, aligning with the tendencies observed in previous research. On the other
Table 2. Leaf color reading values of CIELAB, converted color, and CIE76 (
Δ
E*
ab) values of S. haworthii as affected by
day/night temperatures for 10 weeks
Temperatures
(day/night)
CIELAB values (leaf color) Converted
colorz
(color chip)
CIE76 (ΔE*
ab) by day/night temperatures
L*a*b*20/15°C 24/19°C 28/23°C 32/27°C
20/15°C 73.48 by-2.37 c 7.80 a Reference 1.76 4.85 4.55
24/19°C 73.06 b -2.06 bc 6.11 b 1.76 Reference 4.95 4.47
28/23°C 78.01 a -1.85 b 6.14 b 4.85 4.95 Reference 1.43
32/27°C 77.29 a -0.61 a 6.03 b 4.55 4.47 1.43 Reference
Significancex* *** **
zColors converted using CIELAB L*, a*, and b* values.
yMeans separations within columns were performed using Duncan’s multiple range test, with a significance level set at p < 0.05.
x*, **, and ***: significant at p < 0.05, 0.01, or 0.001, respectively.
Jae Hwan Lee and Sang Yong Nam / Assessment of the Growth and Ornamental Quality of Senecio haworthii (Sweet) Sch.Bip. ~ ∙297
hand, the CIELAB b* value, representing blue-yellow opponent colors, was highest under the 20/15°C treatment,
suggesting leaf yellowing tendencies in relatively lower temperature. Based on the color difference analysis using
CIE76 color difference (ΔE*
ab), temperatures of 20/15 and 24/19°C were used as reference points, under these
conditions, the temperatures of 28/23 and 32/27°C were evaluated as having ‘small color differences’. Therefore, it
was concluded that even with significant differences in day/night temperatures, there were no noticeable
differences in leaf color. Taking all these findings into consideration, it can be concluded that cultivating S.
haworthii in relatively lower temperature conditions around 20/15°C would be advantageous for enhancing both
plants growth and ornamental quality compared to higher-temperature conditions like 32/27°C.
Acknowledgement
This paper was supported by the Sahmyook University Research Fund in 2023.
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