ArticlePDF Available

Effects of Shading on the Growth, Development, and Anthocyanin Content of Echeveria agavoides and E. marcus

Authors:

Abstract and Figures

Shading is a key element in the control of light intensity and is usually used during the summer when sunlight is at its most intense. Succulents are ornamental crops that have be extremely popular because of their leaf structure and other characteristics. A study was conducted to determine the effects of shading on the growth and development as well as the anthocyanin content of two Echeveria species: Echeveria agavoides and E. marcus. Three levels of shading were used, namely no shading/full light treatment, partially shaded conditions, and well-shaded conditions, which were achieved by placement inside controlled growing chambers for 4 weeks. The results revealed the bolting of growth in both height and diameter for both species when grown under shaded and partially shaded conditions, which were abnormal compared with the typical development. CIELAB color results also showed that a* was significantly affected by shading levels, exhibiting a higher positive value for succulents under full light or no shading. These results were consistent with the anthocyanin analysis, where the highest contents were identified in the species grown under full light. The image analysis also confirmed a higher percentage area based on the smart segmentation for red pigments compared with that for green pigments.
Content may be subject to copyright.
www.ijfs.org
Abstract Shading is a key element in the control of light
intensity and is usually used during the summer when
sunlight is at its most intense. Succulents are ornamental
crops that have beextremely popular because of their leaf
structure and other characteristics. A study was conducted
to determine the effects of shading on the growth and
development as well as the anthocyanin content of two
Echeveria
species:
Echeveria agavoides
and
E. marcus.
Three levels of shading were used, namely no shading/full
light treatment, partially shaded conditions, and well-shaded
conditions, which were achieved by placement inside controlled
growing chambers for 4 weeks. The results revealed the
bolting of growth in both height and diameter for both
species when grown under shaded and partially shaded
conditions, which were abnormal compared withthe typical
development. CIELAB color results also showed thata* was
significantly affected by shading levels, exhibiting a higher
positive value for succulents under full light or no shading.
These results were consistent with the anthocyanin analysis,
where the highest contents were identifiedinthe species
grown under full light. The image analysis also confirmed a
higher percentage area based on the smart segmentation
for red pigments compared withthat for green pigments.
Additional key words: anthocyanins, CIELB, image analysis,
pigment, shade, succulents
Introduction
The use of succulents as indoor and landscape plants is
mainly due to their ability to survive in harsh environments
with minimal watering and care compared to other ornamental
plants (Fischer and Schaufler 1981; Nyffeler et al. 2008;
Oldfield 1997). Crassulacae family is considered as the third
largest among succulent groups. This consists of plants that
have a wide range of habitat adaptation and temperature
tolerance. These succulents are popularly known as the
‘stonecrop’ and ‘houseleek’ family with a broadcast appeal for
growers, hobbyists and collectors (Rowley 1978; Sevilla et al.
2012). Within this huge family, several subfamilies are
prevalently known including Echeveria, Sedeveria, Sedum,
Crassula, and Graptopetalum. They come in variety of
morphological structures and their leaves evidently create
unique growth patterns.
Most of the researches for succulents that enable
consumers and propagators to improve production and
quality of produce are more inclined to propagation
through in-vitro and leaf cuttings and the discovery of new
species (Raju and Mann 1971; Ruiz et al. 2016; Ruiz and
Costea 2014).
Echeveria is considered as one of the popular genus
among the succulent plants. This genus is comprised of 140
species of which 95% of the total is endemic in Mexico
(Meyran and Lopez-Chavez 2003; Vazquez et al. 2013).
Succulents under this genus is known for the development
of gradient colors on the margins of the lower or mature
leaves of the plant. The colors range from light pink to red
and even deep red hues that are already close to brown or
black. This change in color may be due to the presence of
the anthocyanin pigments.
*Corresponding author: Sang Yong Nam
Tel: +82-2-3399-1732
E-mail: namsy@syu.ac.kr
ORCID: htt
p
s://orchid.or
g
/0000-0003-0863-4721
Flower Res. J. (2017) 25(4) : 270-277
DOI htt
p
s://doi.or
g
/10.11623/fr
j
.2017.25.4.12
ISSN 1225-5009(Print)
ISSN 2287-772X
(
Online
)
O R IG IN AL AR TIC L E
Effects of Shading on the Growth, Development, and
A
nthocyanin
Content of Echeveria agavoides and E. marcus
Raisa Aone M. Cabahug
1,2
, Son Yil Soh
1,2
, and Sang Yong Nam
1,2,*
1Department of Environmental Horticulture, Sahmyook University, Seoul 01795, Korea
2Natural Science Research Institute, Sahmyook University, Seoul 01795, Korea
Received 5 September 2017; Revised 25 October 2017; Accepted 1 November 2017
Copyright © 2017 by The Korean Society for Floricultural Science
www.ijfs.org
Flower Res. J. (2017) 25(4) : 270-277 271
One of the most noticeable classes of flavonoids is
anthocyanin. They are known to be responsible for important
plant pigments like red, pink, purple and blue colors in plants
(Grotewold 2006). One of the widely-accepted function of
anthocyanin is that they protect leaves in plants which are
facing biotic or abiotic stressors and has been discussed as
putative roles as anti-oxidants and sunscreens (Hughes and
Yadun 2015; Landi et al. 2015; Pringsheim 1879). Environmental
conditions of other ornamental flowers and foliage have been
manipulated to produce attractive colors that may enhance
plant attractiveness to potential consumers (Zhao and Tao
2015; Harpaz and Padowicz 2007). With these researches,
determination of pigment content is deemed necessary. They
keep track of actual visible colors using high-performance
liquid chromatography (HPLC), however modern color
instrumentation has made use of CIELAB which is more
practical and easy to use (Wrolstad et al. 2005).
Most of the succulents are being placed inside offices,
schools, homes and other indoor places where the lighting
conditions are shaded. Recent studies have been conducted
for few succulent species on the effect of light intensity and
the type of light that would intensify color change (Nam et
al. 2016; Park et al. 2016; Soh et al. 2015). However, the
color quality of these ornamental plants varies, especially, if
they are used as potted plants inside establishments creating
shaded environments with low source of light. This paper
aimed to identify how shade in Echeveria species impacts
their growth, development and quality.
Materials and Methods
Planting materials
Echeveria agavoides
and E.cheveria marcus species were
procured from Kim Succulent Nursery, Anseong, South
Korea. These species have a common trait which under
unknown conditions, their leaf margins will turn pinkish to
red in color. Young, healthy and disease-free succulents
were chosen as experimental plants which were around 60
days old and were placed inside the greenhouse of
Sahmyook University, Seoul, South Korea.
Experimental design, treatments and growth
conditions
The experiment was laid out in a completely randomized
design with four replications with six plants per replication.
There was a total of seventy-two plants per species. Three
shading levels served as the factor which were no shading/full
light, partially shaded and well-shaded conditions. To achieve
shading levels, the use of two metal frames were placed inside
each chamber and was lined with polyethylene nets. Each
metal frame held a different light intensity because of the density
of the polyethylene nets (Fig. 1). Shading levels were also
secured to be consistent throughout the metal frame by
determining the lux value using the Digital Lux Meter (DX-200,
Centenary Materials Co., Taiwan). The control or no shading/
full light treatment had 10,000 lux followed by the
partially-shaded with 5,000 lux and the well-shaded with 2,500
lux. There was a 14 hours light period and 10 hours dark
period.
All experimental plants were placed inside three plant
growth chambers (KGC-175 VH, Koenic Ltd., South Korea).
Each chamber was equipped with four LED tube lights
(Philips F48T12/CW/VHO 110 Watt, USA) and the relative
humidity was set at 65% and the temperature at 20°C.
Hunter’s CIELAB
Color change was determined using the Hunter’s CIELAB
(Konica Minolta Spectrophotometer CM2600d) which makes
use of the L*a*b* color space to indicate lightness, hue and
saturation of colors. One leaf for each plant was tagged to
trace the color changes. The color value was measured
through two areas within the tagged leaf; margin of the top
and underside of the leaf.
Anthocyanin analysis
Modified quantitative method for anthocyanin (Fuleki and
Francis 1968) was used in this study by gathering an inch
from the tip of the tagged leaves of succulent plants. One
gram fresh-cut leaf samples were macerated using a mortar
and pestle. The macerated sample was added with 1mL of
95% ethanol and 1.5 N HCl (85 : 15) which served as the
www.ijfs.org
272 Flower Res. J. (2017) 25(4) : 270-277
extracting solvent. The mixed solution was transferred to a
separate container. Samples were then centrifuged at 13,000
rpm at 4°C using the Micro Refrigerated Centrifuge Smart R17
(Hanil Science Co. Ltd., Seoul, South Korea). This was then
stored and refrigerated overnight. Samples were placed in a
microplate which was then analyzed for a full-spectrum UV/Vis
absorbance at 535 nm using the Fluostar Optima Microplate
Reader (BMG Labtech, Ortenberg, Germany). Results from this
analysis are expressed using µg/ g of fresh weight (FW).
Image analysis
Photos were taken using a Canon 750D (Canon, Japan)
with the same aperture, brightness and contrast at the same
distance. Individual images were cropped to show the
succulents alone without the pots and were processed
using the Image Pro Premier ver. 9.3 (Media Cybernetics,
USA). Smart segmentation was applied to individual
representative images to determine the ratio of the colors
green and red pigments. Colored overlays of identified
colors was presented as bases for color identification.
Statistical analysis
Data gathering was done every two weeks for a month.
Aside from the Hunter’s CIELAB and anthocyanin analysis,
growth and development parameters were also collected:
plant height and diameter. Statistical analyses were conducted
using Statistical Product and Service Solutions for Windows,
version 16.0 (SPSS Inc., Japan). The data were analyzed
using analysis of variance (ANOVA), and the differences
between the means were tested using Duncan’s multiple
range test (P < 0.05).
Results and Discussion
Plant height and diameter
Results revealed that plant height has significantly affected
plant height for both Echeveria species (Table 1). Succulents
that were grown under shaded conditions was significantly
taller compared to those that were exposed to higher light
exposure which had 44.77 mm (E. agavoides ) and 46.95 mm
(E. marcus ). These were followed by those in partial shading
with 42.67 mm (E. agavoides ) and 42.06 mm (E. marcus ).
Those grown in partial-shaded condition did not differ from
those under full-light or no shade with 41.97 mm and 41.63
mm, respectively.
Consequent growth of the succulent’s height was also
A
BC
Fig. 1. Experimental set-up inside the growth chamber to achieve light conditions: A, metal frames with polyethylene (PE) nets with
black cloth barriers on the side; B, PE net for shaded condition; and C, PE net for partially-shaded.
www.ijfs.org
Flower Res. J. (2017) 25(4) : 270-277 273
evident in the diameter with similar results. Significant
differences were observed in diameter for E. agavoides
having those grown in shaded conditions had the largest
plants with 90.25 mm followed by partial shading and no
shade with 88.42 mm and 83.61 which were comparable
with each other.
E marcus species had similar results wherein the
treatment of succulents under shaded conditions (79.82 mm)
gave the largest plants followed by those in partial-shading
(77.39 mm) and full-light (75.88 mm). However, these were
not significantly different from each other.
These results are commonly seen in experiment as those
done in germinating bean seeds in dark and light
conditions. Aside from the bean family, studies of Rylski
and Spigelman (1986) have found out that when sweet
peppers were grown in shaded conditions had increased
plant height, number of nodes and leaf size as the light
intensity decreased.
Various explanations are said about profuse growth in
plants in shaded conditions such as its relation to circadian
cycles as well the shade avoidance syndrome. The changes
in plant body and function are the responses of shade
tolerance. Studies of Casal (2012) showed that the different
responses of Arabidopsis thaliana during its different stages
of life cycle. In this study, it was observed that shading
increases the adaptive benefits of elasticity which enables
the plant to elongate its stem, spread its leaf surface area.
It was also considered that the lack of light increases the
production of auxin which allows more movement in the
tips of the growing plants.
Despite of these pants recording these growths are found
to produce less healthy looking plants with lightness of the
green pigments as well as portraying a lanky structure.
Studies of Zhao et al. (2012) on herbaceous peony
concluded that plants grown under shaded condition
reduced its photosynthetic capacity, light saturation and
compensation point because of the declined stomatal
conduction. Because of those decline, it resulted to the
decrease of soluble sugar, protein and malondialdehyde
which in turn produced plants with delayed flowering,
reduced ornamental quality and faded plant color.
Hunter’s CIELAB
Results of the hunter’s CIELAB revealed that E. agavoides
were significantly affected by shading conditions (Table 2).
The top portion or exposed leaf area showed that L*, a*
and b* values were significantly different from each
condition. L* or the brightness of the color shows that there
was a lighter quality of color with 43.00 value on the scale
which was significantly comparable from those of partial
shading (40.67). The darkest color quality was observed
from those grown under no shade or full light condition
with 40.93.
A positive a* was observed in pants that were grown
under full light with the highest value of 4.98 signifying
that it had a more reddish hue compared to those nearing
to the 0 value or negative which was seen in succulents
under partially-shaded (0.39) and shaded conditions (-2.39)
signifying towards the greener hue. Saturation of colors or
b* value was also seen as more intense in full-light
conditions with 14.42 compared to those of partial-shading
(13.51) and shaded (11.50) conditions.
T
able 1. Average plant height and diameter (mm) of Echeveria species in response to shading levels.
Shading conditions Echeveria agavoides Echeveria marcus
Height Diameter Height Diameter
No shade (full light) 41.97 b
z
83.61 b 41.64 b 75.88
Partial-shading 42.67 b 88.42 ab 42.06 b 77.39
Shaded 44.77 a 90.25 a 46.95 a 79.92
F-test
y
** * ns
z
Mean separation within columns by Duncan’s multiple range test at P = 0.05.
y
NS, *, **, Nonsignificant, significant, highly significant at P= 0.05 and P= 0.01, respectively.
www.ijfs.org
274 Flower Res. J. (2017) 25(4) : 270-277
For the bottom area of the leaf not exposed to the light,
there was only a significant difference for the a* value
which showed a positive value for no shading treatment
(7.84) these were significantly different for those with
negative values for succulents grown under partial shading
(-0.23) and shaded (-2.09) conditions. However, the *L and
b* values were not significantly affected by the treatments.
For E. marcus, some of the CIELAB values were
significantly affected by the treatments (Table 3). These had
similar results for those of E. agavoides except for the b*
values. E. marus has a differently natural light blue green.
In the CIELAB sphere of colors, lower values for b* signify
a bluer saturation tone. Thus, in both b* values for the top
and bottom portions of the leaves have lower values for the
no shade treatment compared to shaded conditions.
Anthocyanin analysis
The anthocyanin content was significantly affected by
shading treatments for both species (Table 4). Results
revealed that there is more presence of anthocyanins for
Echeveria succulents that were grown under full light or
non-shaded conditions.
Based on the results, the highest anthocyanin content
was found in plants in full light or no shading with 1.32
µg/g FW for E. agavoides and 0.82 µg/g FW for E. marcus.
These were followed by partial-shading with 0.66 µg/g FW
for E. agavoides and 0.44 µg/g FW for E. marcus. The
lowest anthocyanin contents were taken for shaded levels
for both species.
Succulents are considered to mature within the four
weeks treatment. According to Kliewer (1970), anthocyanin
T
able 2. Average Hunter’s CILEAB values of E. agavoides in response to shading levels.
Shading conditions Top Bottom
L* a* b* L* a* b*
No shade (full light) 40.93 b
z
4.98 a 14.42 a 41.00 7.84 a 10.61
Partial-shading 40.67 ab 0.39 b 13.51 b 40.39 -0.23 b 12.14
Shaded 43.00 a
-2.39 c 11.50 c 37.46 -2.09 c 11.96
F-test
y
****ns * ns
z
Mean separation within columns by Duncan’s multiple range test at P = 0.05.
y
NS, *, **, Nonsignificant, significant, highly significant at P = 0.05 and P = 0.01, respectively.
T
able 3. Average Hunter’s CILEAB values of E. marcus in response to shading levels.
Shading conditions Top Bottom
L* a* b* L* a* b*
No shade (full light) 39.31 a
z
3.16 a 10.05 41.00 3.80 a 9.53 b
Partial-shading 39.85 a -2.27 b 11.73 40.39 -2.58 b 12.82 a
Shaded 37.64 b -3.83 b 12.67 37.46 -4.44 b 11.99 a
F-test
y
***nsns * *
z
Mean separation within columns by Duncan’s multiple range test at P = 0.05.
y
NS, *, **, Nonsignificant, significant, highly significant at P = 0.05 and P = 0.01, respectively.
T
able 4. Average anthocyanin (µg/g FW) content of Echeveria species in response to shading levels.
Shading conditions Echeveria agavoides Echeveria marcus
No shade (full light) 1.32 a 0.82 a
Partial-shading 0.66 b 0.44 b
Shaded 0.37 c 0.34 c
F-test ** **
z
Mean separation within columns by Duncan’s multiple range test at P = 0.05.
y
NS, *, **, Nonsignificant, significant, highly significant at P= 0.05 and P= 0.01, respectively.
www.ijfs.org
Flower Res. J. (2017) 25(4) : 270-277 275
pigment which are predominantly red are supposed to
accumulate and be enhanced by low light intensities.
However, studies of Fukuoka et al. (2014) with Gynura
bicolor suggested that there was a limited increase of
anthocyanin content for shaded treatments despite
expansion of leaves during maturity.
These contrasting results have been controversial in
determining anthocyanin tendencies, on the other hand,
studies of Jeong et al. (2004) revealed that anthocyanins can
also be affected by expression of anthocyanin biosynthetic
genes and the application of shading treatments suppressed
the accumulation.
Image analysis
Smart segmentation of images coupled with the original
images for three levels of shading and their corresponding
histogram results is presented on Fig. 2a for E. agavoides
and Fig. 2b for E. marcus.
Based on the histogram of each level, it was evident that
red pigments are more evident in succulents under full light
and has slowly decline with shading. Results of the smart
segmentation suggested that there is a high percentage of
red color compared to green color for both species for full
light or no shading. Thus, there is a negative correlation
between red pigments and shading while a positive
correlation between green pigments.
Shading level Segmented image Histogram
Full light or no shading
Partially shaded
Shaded
Fig. 2. Original image, processed segmentation image and histogram determining area ratio of green and red pigments using numbe
r
of pixels of E. agavoides in response to shading levels.
www.ijfs.org
276 Flower Res. J. (2017) 25(4) : 270-277
Shading level Segmented image Histogram
Full light or no shading
Partially shaded
Shaded
Fig. 3. Original image, processed segmentation image and histogram determining area ratio of green and red pigments using numbe
r
of pixels of E. marcus in response to shading levels.
T
able 5. Red and green pigment ratio of using smart segmentation in response to shading levels of Echeveria species.
Shading level Green pixels Red pixels Total % Green % Red
E. agavoides
Full light 1028.50 (± 3.15)
z
701.88 (± 1.02) 1730.38 59.44 b
y
40.56 a
Partial shading 1518.81 (± 3.50) 343.00 (± 5.75) 1861.82 81.58 a 18.42 b
Shading 4053.09 (± 7.96) 589.26 (± 3.99) 4642.35 87.31 a 12.69 b
E. marcus
Full light 811.72 (± 1.66) 2603.82 (± 2.74) 3415.54 23.77 c 76.23 a
Partial shading 501.68 (± 1.82) 365.56 (± 7.88) 867.24 57.85 b 42.15 b
Shading 1253.08 (± 3.67) 316.07 (± 1.99) 1569.15 79.86 a 20.14 c
z
Mean ± standard deviation.
y
Mean separation within columns by Duncan’s multiple range test at P= 0.05.
www.ijfs.org
Flower Res. J. (2017) 25(4) : 270-277 277
Acknowledgement
Succulents Export Innovation Model Development towards
Chinese Market (514006-03-1-HD040)’, Ministry of Agriculture,
Food and Rural Affair and Sahmyook University Research
Fund.
References
Casal JJ (2012) Shade avoidance. The arabidopsis book
Fischer CC, Schaufler EF (1981) Artificial lighting for decorative
plants. http://www.gardening.cornell.edu/houseplants/pdf/
artificiallighting/pdf
Fuleki T, Francis FJ (1968) Quantitative methods for
anthocyanins. 1. extraction and determination of total
anthocyanin in cranberries. J Food Sci 33:72-77
Fukuoka N, Suzuki T, Minamide K, Hamada T (2014) Effect
of shading on anthocyanin and non-flavonoid polyphenol
biosynthesis of Gynura bicolor leaves in midsummer. Hort
Sci 49:1148-1153
Grotewold E (2006) The genetics and biochemistry of floral
pigments. Annu Rev Plant Biol 57:761-780
Harpaz S, Padowicz D (2007) Color enhancement in the
ornamental dwarf cichlid microgeophagus ramirezi by
addition of plant carotenoids to the fish diet. The Israeli
J Aquaculture - Barnidgeh 59:195-200
Hughes NM, Yadun SL (2015) Red/purple leaf margin
coloration: potential ecological and physiological functions.
Environ Expt Bot 119:27-39
Jeong ST, Yamamoto NG, Kobayashi S, Esaka M (2004)
Effects of plant hormones and shading on the
accumulation of anthocyanins and the expression of
anthocyanin biosynthetic genes in grape berry skins. Plant
Science 167:247-252
Kliewer WM (1970) Effect of day temperature and light
intensity on coloration of Vitis vinifera L. grapes. J Amer
Soc Hort Soc 95:693-697
Landi M, Tattini M, Gould KS (2015) Multiple functional roles
of anthocyanins in plant-environment interactions. Environ
Expt Bot 119:4-17 DOI:10.1016/j.envexpbot.2015.05.012.
Meyran J, Lopez L (2003) Las Crasuláceas de México.
Sociedad Mexicana de Cactología, Arca Contenental,
Mexico, p 234
Nam SY, Lee HS, Soh SY, Cabahug RA (2016) Effects of
supplementary lighting intensity and duration on
hydroponically grown Crassulaceae species. Flower Res
J 24:1-9
Nyffeler R, Eggli U, Ogburn M, Edwards E (2008) Variations
on a theme: repeated evaluation of succulent life forms
in the Portulacineae (Caryophyllales). Haseltonia J
14:26-36
Oldfield S (comp) (1997) Cactus and succulent plants-status
and conservation action plan. International Union for
Conservation of Nature and Natural Resources/ SSC
Cactus and Succulent Specialist Group. IUCN, Gland,
Switserland and Cambridge, UK, p 212
Park SY, Nam YW, Ko EK (2016) Effects of different
supplemental lighting lamps on growth and quality of
succulent plants. J Natural Science 20:30-44
Pringsheim N (1879) Ueber Lichtwirkung und Chlorophyll
Function in der Pflanze. Jahrbücher für Wissenschaftliche
Botanik
Raju MV, Mann, HE (1971) Regenerative studies on the
detached leaves of Echevevia elegans. patterns of
regeneration of leaves in sterile culture. J Canadian Bot
49:2015-2021
Rowley G (1978) The Illustrated Encyclopedia of Succulents.
Crown Publishers Inc. New York, USA, pp 116-135
Ruiz GI, Costea M (2014) Echeveria marianae (Crassulaceae),
a new species from Jalisco, México. Phytotaxa 170:
35-40
Ruiz GI, Martinez DV, Reyes PC, Costea M (2016) Taxonomic
and floristic novelties for Echeveria (Crassulaceae) in
Central Michoacan, Mexico. Phtokeys 75:1-12
Rylski I, Spigelman M (1986) Effect of shading on plant
development, yield and fruit quality of sweet pepper grown
under conditions of high temperature and radiation.
Scientia Horticulturae 29:31-35
Sevilla H, Reyes P, Calix E, Basanez M (2012) Additions to
the Crassulaceae of the state of Veracruz, Mexico.
Haseltonia 18:140-152
Soh SY, Park SY, Cabahug RA, Nam SY (2015) Analysis of
supplemental lighting lamps and color on growth of
succulent plants. J Natural Science 19:47-58
Vazquez-Garcia JA, Jimeno SD, Cuevas GR, Chazaro BM,
Muñiz-Castro MA (2013) Echeveria yalmanantlanensis
(Crassulaceae): A new species from Cerro Grande, Sierra
de Manantlán, western Mexico. Brittonia 65:273-279
Wrolstad RE, Durst RW, Lee JM (2005) Tracking color and
pigment changes in anthocyanin products. Trends in Food
Sci Technol 16:423-428
Zhao D, Tao J, Han C, Ge J (2012) Flower color diversity
revealed by differential expression of flavonoid biosynthetic
genes and flavonoid accumulation in herbaceous peony
(Paeonia lactiflora Pall.) Mol Biol Reports 39:11263-11275
... Já no estudo de Kim;Kim (2015), mudas de Sedum middedorffianum enraizaram independente do sombreamento, porém o desenvolvimento das raízes foi maior na condição sem sombreamento. Avaliando duas espécies de Echeveria, Cabahug et al. (2017) observaram que ambas apresentaram maior crescimento em altura sob sombreamento, entretanto o desenvolvimento foi considerado anormal em relação ao típico das espécies. Algumas espécies crassuláceas ainda respondem melhor ao aumento da intensidade luminosa e tempo de exposição à luz, como a Sedeveria 'Letizia', Echeveria 'Momotaro' e Graptoveria opalina, enquanto para a Sedum 'Sun Red' não há efeito . ...
Article
Full-text available
O cultivo de plantas suculentas se popularizou no Brasil, aumentando a necessidade da compreensão do comportamento das espécies sob condições diferentes de produção. Assim, objetiva-se com este trabalho avaliar a influência de fertilizante enraizador, temperatura e luminosidade na emergência de brotações e raízes para espécies suculentas. Três experimentos foram conduzidos de novembro 2020 a fevereiro 2021, em casa de vegetação, com delineamento inteiramente casualizado, com quatro repetições, cada estudo com uma espécie: Echeveria elegans, Graptosedum Francesco Baldi e Sedum adolphii. Para cada espécie, os tratamentos foram arranjados em esquema fatorial (3x2), o fator A comparou fertilizante enraizador comercial e duas testemunhas (água e controle); e o fator B, as estacas foliares das espécies foram submetidas a dois ambientes de temperatura e luminosidade (24,8 oC e 449,8 µmol/m2s; 5 oC e 0 µmol/m2s). Avaliou-se aos 30 dias de instalação dos estudos: índice de velocidade de brotação; porcentagem de brotação; comprimento médio das raízes; comprimento médio do broto e massa seca total. O fertilizante beneficia a emissão de brotações e raízes na Echeveria elegans, porém tais efeitos não são verificados na ausência de luminosidade e baixa temperatura (5 oC). Esse ambiente também prejudicou as demais espécies avaliadas. Palavras-chave: Echeveria elegans; Graptosedum Francesco Baldi; Propagação; Sedum adolphii. Emergency of sprouts and roots of suculent species due to the application of rooting fertilizer, in contrasting environments ABSTRACT: The cultivation of succulent plants became popular in Brazil, increasing the need to understand the behavior of species under different conditions. Thus, the aim of this work is to evaluate the influence of plant rooting fertilizer, temperature, and light intensity on the emergence of new shoots and roots for succulent species. Three experiments were conducted from November 2020 to February 2021, in greenhouse, with completely randomized design, with four replications, each study with one species: Echeveria elegans, Graptosedum Francesco Baldi and Sedum adolphii. The treatments for each species were arranged in factorial scheme (3x2), factor A compared commercial plant fertilizer and two checks (water and control); and factor B, the leaf cuttings of the species were submitted to two environments in relation to temperature and light (24.8 oC and 449.8 µmol/m2s; 5 oC and 0 µmol/m2s). It was evaluated at 30 days of installation: sprouting speed index; sprouting percentage; average root length; average sprout length and total dry mass. The fertilizer benefits the emission of shoots and roots in Echeveria elegans, but such effects are not observed for absence of light and low temperature (5 oC). This environment also harmed the other species evaluated. Keywords: Echeveria elegans; Graptosedum Francesco Baldi; Propagation; Sedum adolphii.
... In domesticating and cultivating these plants, germplasm facilities of ornamental plants are generally indoors and plant species are grown under greenhouse conditions. During the summer season, shading is considered a key element in controlling light intensities and has been found to be an important factor in growing succulent plants like that of S. zokuriense (Cabahug et al. 2017). In regulating light intensities, it is essential to consider how this factor affects plant morphology, productivity, and photosynthetic activities (Kim et al. 2011). ...
Article
Full-text available
An endemic plant to South Korea, Sedum zokuriense Nakai, has medical and floricultural potential and is of ecological importance. Today, many species under the genus Sedum are used as green-roofing systems, sold as ornamental plants, and studied for breeding programs. As such, optimization studies should be conducted to identify key environmental and cultivation factors that would affect their survival and vegetative growth. In this study, shading levels (50%, 65%, 80%, 95%, and 98%), potting mixes (decomposed granite, fertilizer-amended media, perlite, river sand, burnt husk, and vermiculite), and fertilization rates [(control, 0 ppm), 500, 750, 1,000, and 2,000 ppm] were investigated and the responses of S. zokuriense in terms of their survival rate, plant growth and development, CIELAB color reading, and chlorophyll content under greenhouse conditions. Results showed that these stonecrop species are shade-loving and thrive in low-light conditions. Although the fertilizer use had minimal impacts, growing plants at 65% shading, planted with RS:VL:PL (6:2:2, v/v/v) potting media have substantially produced a high survival rate in propagation using stem cuttings. Furthermore, this allowed plants to be established while supporting high vegetative growth, green and healthy plants with high chlorophyll content.
Article
Full-text available
Plants under the genus Orostachys have been known as medicinal plants. This study deems to determine the growth and leaf color of Orostachys japonica and O. boehmeri when subjected to various LED light sources. A total of seven LED light treatments were used, i.e. red (630 nm), green (520 nm), blue (450 nm), purple (650 and 450 nm), 3000 K white (455, 600 nm), 4100 K white (455, 590 nm), and 6500 K white (450, 545 nm) LEDs. Results showed that O. japonica plants showed favorable growth under 4100 K white LED, while O. boehmeri plants had a positive growth response under white light LEDs (3000, 4100, and 6500 K). In leaf color analysis, the use of green LED showed the greatest change in CIELAB L * and b * values which were relatively higher compared to other treatments indicating that leaves turned yellowish. Further statistical analysis using Pearson’s correlation also suggested that there is a small negative association between dry weight and b * values of O. japonica, and a negative moderate association between plant weights (fresh and dry weight) and leaf color (L * and b * ) and positive association between said plant weights and a * color values of O. boehmeri. Therefore, it is recommended to cultivate O. japonica under 4100 K white LED and O. boehmeri under 3000, 4100, 6500 K white LEDs.
Article
Full-text available
Delosperma cooperi is a perennial herbaceous succulent that grows wild in Lesotho and the Republic of South Africa. As a ground cover plant and as an indoor ornamental plant, it has a high horticultural value. It is essential to fully understand the conditions and other factors that play a key role in higher propagation success to reduce labor costs and efficiently propagate. Optimal cultivation methods for the vegetative propagation of D . cooperi has not yet been studied. In this study, the effects of shading levels (50%, 65%, 80%, 95%, and 98%), various soil mixes (decomposed granite, perlite, river sand, fertilizer-amended media, and vermiculite), and fertilization rates (Control, 500, 750, 1,000, and 2,000 ppm) on the propagation success, growth, and development of D. cooperi were investigated. According to the results, the best growth condition for propagating this succulent was subjecting them to 50% shading conditions using vermiculite (VL):fertilizer-amended media (FM):perlite (PL) (3:2:5, v/v/v) as soil media with the application of 1000 ppm fertilization rates to enhance plant growth. The leaf color of D. cooperi was dark green (RHS N137A, 147A) at a shading level of 65% or less, and the leaf color changed to yellow (RHS 146A, 147B, 148A) at the shading level of 80% or more. The correlations between CIELAB L* and b* values with plant growth parameters were analyzed in the shading levels study, and they showed a negative correlation with each other. However, there was no correlation between growth parameters and leaf color in the fertilization rates study.
Article
Full-text available
This study was conducted to determine the effects of supplementary lighting intensity and duration on selected Cassulaceae species grown in a hydroponic system. Five subfamilies in Crassulaceae with corresponding species were chosen as experimental units namely Sedeveria ‘Letizia ’, Sedum ‘Sun Red’, Crassula rupestris, Echeveria ‘Momotaro’, and Graptoveria opalina. Light duration (3 and 6 hours) and intensity (4,000 lux or 60 μmol • m−2 • s−1 and 8,000 lux or 120 μmol • m−2 • s−1), and their combinations served as factors which were replicated twice. Results revealed that the use of supplementary lighting using LED fixtures had influenced selected species under Crassulaceae. The use of three hours supplementary lighting under low light intensity had statistically similar results with those of the control S. letizia, C. rupestris and G. opalina in particular parameters. Meanwhile, succulents under six-hour with high intensity condition grew well, compared to species S. letizia, C. rupestris and E. ‘Momotaro,’ demonstrating that the data was significantly different. Interestingly, there were no statistical significant differences between species C. rupestris and the control regardless of change of variables (duration and intensity) in all parameters. S. letizia, acquired from an Italian collection (Sedum cuspidatum × Echeveria setosa var. ciliate), belongs to subfamily Sedoidae. Plants under this group are well-known for being in ‘splits’ or having another included within them (Fig. 1). They are drought tolerant species and leaf color may change depending on the shading or lighting (Lee and Kim 2008; Rowley 1978). Results of the analysis revealed that supplemental lighting significantly affected S. letizia height, diameter, and *b among parameters as shown on Table 1.
Article
Full-text available
Leaf margins of many plant species belonging to the floras of several continents feature a conspicuous band of red/purple color around their periphery. Despite the widespread distribution of this leaf trait, very few studies have proposed or tested hypotheses to explain its significance (if any). Common explanations for leaf coloration such as photoprotection, plant camouflage, attraction of seed dispersers, or undermining herbivorous insect camouflage do not seem, at first glance, to be applicable to this color pattern. Could pigments localized at the leaf margin still function in these traditional ecological or physiological roles? Or should new hypotheses be devised that are more specific to coloration at the leaf margin? The purpose of this paper is to review and explore potential ecological and physiological functions of pigmented leaf margins, in hopes of inspiring further inquiry into this topic.
Article
Full-text available
The effects of high (86°F) and low (68°F) day temperature, and of high (2,500 to 5,000 ft-c) and low (500 to 1,200 ft-c) light intensity, on the coloration of ‘Cardinal’ and ‘Pinot noir’ grapes grown in sunlit, temperature-controlled rooms during the ripening period were investigated. Night temperature (7 PM to 7 AM) was 59°F in all treatments. Low day temperature significantly increased the level of anthocyanin pigments in the skins of both cultivars at both high and low light intensity. Anthocyanin synthesis was almost completely inhibited in the skins of ‘Cardinal’ berries that had average daytime temperatures between 91 and 95°F. Low light intensity greatly reduced coloration of ‘Pinot noir’ grapes at both low and high day temperatures but decreased the level of pigments in grapes grown at 86°F. It either increased or had little effect on fruit coloration of ‘Cardinal’ grapes grown at 68°F.
Article
Full-text available
A new species, Echeveria marianae (Crassulaceae) is described from Sierra del Tigre, Valle de Juárez, State of Jalisco, Mexico. The species belongs to ser. Gibbiflorae due to of its acaulescent or short caulescent rosette habit, paniculiform inflorescence, conical-urceolate corolla, and tricolpate pollen grains. Within ser. Gibbiflorae it shares morphological affinities with E. novogaliciana and E. dactylifera from which it differs in the shape, color and margin of leaves, corolla size and color, stamen length, nectaries morphology, and its geographical distribution.
Article
Full-text available
During the ongoing studies of the Crassulaceae family for the Flora of Veracruz (Mexico), we found two new species of Crassulaceae (Echeveria uxorium and Sedum jarocho), and eight new records for the state: Crassula connata var. connata, Echeveria bifida, E. coccinea, E. halbingeri, Sedum corynephyllum, S. ebracteatum, S. guatemalense, and Villadia patula. Data on the distribution and habitat of each species are given.
Article
Full-text available
The present research examined the effects of adding carotenoids from oleoresin paprika to fish feeds for ornamental dwarf cichlid, Microgeophagus ramirezi. The growth rate, survival, carotenoid accumulation level, and color intensity were evaluated. Post larvae and near-adult (three months old) fish were tested to determine when carotenoids are better assimilated. The addition of carotenoids had no effect on the growth rate or survival in either life stage, however, they had a clear effect on color enhancement. After 45 days, near-adult fish that consumed carotenoid-supplemented diets at 60, 120, or 240 mg/kg had significantly higher levels of carotenoids (72.19±4.55, 84.81±5.29, and 86.55±4.50 µg/g dry matter, respectively) than con- trol fish (33.69±1.06 µg/g), with no significant differences between treatments. After 75 days, post larvae that consumed 240 mg/kg carotenoids accumulated significantly more carotenoids in their body (59.34±3.93 µg/g dry matter) than fish that consumed only 60 mg/kg carotenoids (40.53±2.37 µg/g dry matter) or no supplemental carotenoids (29.18 µg/g dry matter). Visual examination revealed a strong correlation between level of pigment accumulation and color appearance in the fish. Results indicate that addition of 60 mg oleoresin paprika per kg diet is sufficient to obtain good coloration in M. ramirezi.
Article
The effect of shading in midsummer on anthocyanin and non-flavonoid polyphenol biosynthesis of Gynura bicolor DC leaves was examined using a control (full solar radiation) and a shade treatment (50% shading of full solar radiation). Leaf temperature in the shade plot remained ∼40 °C in the daytime, ∼6 °C lower than in the control. Plants in the shade plot grew better than the control. The content of chlorogenic acid (CGA) in leaves decreased with leaf maturation for both treatments, and a larger amount of CGA was detected in leaves from the control than the shade treatment. The profiles of reactive oxygen species (ROS) scavenging activity exhibited an identical pattern to the content of CGA. Although there was an abrupt increase in the content of anthocyanin in the early stage of leaf expansion, the content decreased rapidly as the leaves matured. The increase in anthocyanin early during leaf expansion was much more limited in control leaves than shaded leaves. There were no correlation between the profiles of anthocyanin and gene expression such as GbPAL, GbC4H, Gb4CL, GbCHS, GbCHI, GbF3H, and GbUFGT. However, the profiles of expression of genes such as GbMYB2, GbF3#H, GbDFR, and GbANS were similar to the anthocyanin profiles. These results suggest that artificial shading in midsummer is an effective method to promote anthocyanin accumulation but reduces ROS scavenging capacity as a result of lowered CGA production.
Article
SUMMARY— The method developed consists of extracting the anthocyanins with ethanol-1.5N hydrochloric acid (85:15) and measuring the O.D. of the extract, diluted with the extracting solvent, at 535 nm. The total anthocyanin content was calculated in absolute quantities with the aid of the extinction coefficients established for the four major cranberry anthocyanins dissolved in the alcoholic solvent system.
Article
The effect of different levels of shading of sweet pepper under high solar radiation (> 600 cal cm−2day−1) at 2 different spacings during the summer months in the northwestern Negev desert of Israel (31°N) was investigated. When light intensity was reduced, plant height, number of nodes and leaf size increased. However, shading inhibited the development of lateral shoots on the main stem of the plant below the first terminal flower. The changes in plant development due to shading affected fruit set, number of fruits per plant, fruit location on the plant, fruit development and yield.The lateral shoots which developed under high light intensity provided 25% of the total yield, whereas only a few fruits were picked from the lateral shoots of plants under low light intensity. The lowest number of fruits per plant was obtained under 47% shading at 5 plants m−2 density, under 47 and 26% shading at 6.7 plants m−2 density. Under shading, individual fruits were larger and had a thicker pericarp. Shading reduced sun-scald damage of the fruits from 36% in full sunlight to 3–4% under 26 and 47% shading. The highest yield of high-quality fruits was obtained with 12–26% shade.