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Effects of Auxin and Cytokinin Application on Leaf Cutting Propagation in

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Flower Res. J. (2016) 24(4) : 264-273 ISSN 1225-5009(Print)
DOI http://dx.doi.org/10.11623/frj.2016.24.4.04 ISSN 2287-772X(Online)

O R IG IN A L A R T IC L E

Effects of Auxin and Cytokinin Application on Leaf Cutting

Propagation in Echeveria Species
Raisa Aone Cabahug1,2, Soon-Yil Soh1,2 and Sang Yong Nam1,2,*
1
Department of Environmental Horticulture, Sahmyook University, Seoul 01795, Korea
2
Natural Science Research Institute, Sahmyook University, Seoul 01795, Korea

Received 6 December 2016; Revised 13 December 2016; Accepted 20 December 2016

Copyright © 2016 by The Korean Society for Floricultural Science

Abstract A study was conducted to evaluate the effects of
auxin, cytokinin, and their combined application to hasten
their propagation using leaf cuttings. Different hormone levels
were evaluated on two Echeveria species (E. subsessilis and
E. runyonii). Three levels of auxin as represented by the use
of indole-3-butyric acid (IBA) (0, 100, and 500 ppm), three
cytokinin levels as represented by Kinetin (Ki) (0, 100, and
200 ppm) and their combination were applied and observed
for 8 weeks. The use of 100 ppm IBA resulted in the highest
shoot height, diameter, and rooting and shooting rate. Using
500 ppm IBA led the roots to develop the earliest but both
species also had the highest mortality. The application of
100 ppm Ki significantly affected the majority of leaf cuttings
of E. runyonnii which exhibited the highest and thickest
shoots, and the number of leaves. The application of 100
ppm IBA and 100 ppm Ki is recommended to obtain
increased shoot growth and development for leaf cuttings.
In the case of single hormone use, the application of 100
ppm IBA may be preferable. The combination of auxin and
cytokinin significantly stimulated the hastening of production
of succulents using leaf cuttings.
(Benková et al. 2003). Meristems enables plant growth and
development in these organs which are affected by
numerous external and internal factors (Dello Loio et al.
2008). These meristems coordinate the initiation of new
organs, continuously provide development of new cells for
plant and are all regulated by a dynamic system (Heisler
and Jonsson 2007).
However, previous studies suggest that succulents have a
more peculiar way of developing new plants through its
leaves. This is due to the presence of intercalary meristems
attached to the leaf tissues which common for other plants.
Due to abundant moisture in leaves of succulents, leaf
cuttings are able to survive for longer periods until new
organs appear (Donnelly et al. 1999; Gorelick 2015; von
Willert et al. 1990).
Succulents are growing in popularity and demand due
to their minimum maintenance and drought-tolerant

Additional key words: exogenous, hormones, IBA,
kinetin, leaf cuttings, succulents
Introduction
Plants have remarkable plasticity and adaptability which
makes it possible for them to form new organs such as
lateral roots, shoots and flowers including meristems
*Corresponding author: Sang Yong Nam
Tel: +82-2-3399-1732
E-mail: namsyzip@naver.com
ORCID: http://orcid.org/0000-0002-4351-447X
characteristic as an indoor and landscape plant. Aside from
their minimal care, succulents have attractive leaf formation
with geometric shapes which are unique among ornamental
plants (Nyffeler et al. 2008). Nowadays, these succulents
are commercially produced and have been increasing in
popularity for plant collectors, landscapers and in
households (Altman 2001).
The use of Echeveria group among research experiments
is popular for propagation studies due to its large group
encompassing greater commonality for morphological
structures amongst succulent species. Despite its demand,
there is slow production of new plants thus, plenty of

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Flower Res. J. (2016) 24(4) : 264-273 265

studies are conducted using the use of tissue culture selected Echeveria species using leaf cuttings.
demands sophisticated techniques and higher finances (Lee
and Park 2013; Lee et al. 2012). Materials and Methods
The use of leaf cuttings is a cheaper option for plant
Planting Materials and Growth Conditions
growers which require simple methods but may take longer
The experiment was conducted in the greenhouse of the
time to develop individual plants (Heisler and Jonsson
Department of Environmental Horticulture, Sahmyook
2007; Zinkan 2010).
University for duration of two months. Two Echeveria
Leaves and stem cuttings or any method of vegetative
species (E. subsessilis and E. runyonii) were utilized as
propagation in general have naturally occurring substances
experimental units of the study.
that are able to stimulate and produce necessary hormones
Whole leaves located at the base of the mother plant
for root and shoot formation, however, most cuttings
were carefully removed close to the petiole of the stem.
require additional stimulation (Altman and Waisel 1997;
Leaves were reselected to obtain a uniform size, weight (2
Carlson 1950). Bio-stimulators were identified in 1930’s as
- 3 g) and stage of development. These mother plants
natural forming substances which are now called ‘plant
completed 80 - 85% of their apical and marginal expansion
growth regulators’ (PGRs). These PGRs are molecules which
and reached their optimal maximum thickness of leaves.
affects numerous plant growth responses even in extremely
These leaves were procured a week before planting to
low concentrations. These plant functions include seed
provide enough time for drying (Raju and Mann 1971).
growth, flowering, fruit development and ripening, plant
These leaves were placed in a dry and well-aerated room
longevity and, vegetative development (Brown 2006; Davis
in the conditions of an average temperature of 20°C and
and Haissig 1990).
average relative humidity (RH) of 70% for 5 - 10 days in
The use of auxin, such as indole-3-butyric acid (IBA),
order to induce callusing before any application or contact
has been studied as rooting hormones for stem cuttings
with aqueous substances which will prevent rotting of
while cytokinin use is reported to help promote
leaves.
adventitious bud and shoot formation in leaf and root
After application of treatments, the leaf succulents were
cuttings (Abdullateef and Osman 2012; Kassahun and
planted in their respective growing trays and were grown
Mekonnen 2011). Both plant hormones have synergistic or
inside the greenhouse with an average temperature of 24°C
antagonistic effect on several important developmental
and 75% RH and an average daylight intensity of 15
processes in growth in plants which includes the formation
µmol·m-2·s-1.
and maintenance of meristems for plant propagation
(Davies 2002; Su et al. 2011). Experimental Design and Treatments
Studies of Paterson and Rost (1978) reported that the use The study was done in a 3 x 3 factorial arrangement in
of auxin alone had only minimal stimulating effect. Their completely randomized design (CRD) with three replications
study also concluded that the use of regenerating new having 10 leaf cuttings per replication with a total of 30
plants using leaf cuttings took longer periods. However, leaves per treatment for each species. Three levels of auxin
they recommended that the use of these exogenous as represented by the use of IBA (0, 100, 500 ppm),
substances, auxin in combination with cytokinin, to quicken cytokinin represented by Kinetin (0, 100, 200 ppm) and
the shoot and root initiation should be explored for study. their combination served as treatments.
Thus, this study aims to determine the effects of auxin,
cytokinin and their combination on the propagation of

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266
Preparation and Application of Hormones
Powder formulations were prepared by mixing the
growth hormones in required quantities to be adjusted to
their respective concentrations following the procedure
of Hartmann and Kester (1983). Different levels of
concentration were made by weighting 0.5 g (500 ppm)
and 0.1 g (100 ppm) of the hormone and were added with
1 - 2 drops of ether for IBA and NaCl for Kinetin to freely
dissolve substances. Hormone combinations were diluted to
1 L distilled water and were stirred together inside an
Erlenmeyer flask. Treatment combinations composed of two
hormones were added together and also separately
prepared.
The nodal part of the leaves was dipped into their
respective solutions for 10 - 15 minutes based on the
studies of Carpenter and Cornell (1992) on the application
of auxin or other hormones for rooting cuttings. In higher
concentrations (e.g. 8,000 - 1,000 ppm), cuttings or
planting materials should be dipped in a shorter time
(between 2 - 6 min) and at lower concentrations (lower
than 1,000 ppm) should have at least 10 - 15 min. Within
the allotted 15 min of treatment, optimum absorption of
hormones was considered to be most effective.
Planting of Leaf Cutting
Leaf cuttings were laid carefully in 60 x 30 cm growing
trays using commercial growing media (Nursery Bed Soil,
Seoul Bio Inc., South Korea) without any addition of
fertilizers. Growing trays were equipped with tiny holes on
its base to facilitate drainage of excess water. These were
laid carefully in an upright position with a planting distance
of 1 cm away from each other. Each treatment had been
separately placed in one growing tray. These were also
grouped and properly labelled based on their
corresponding treatments, treatment combinations and
replications.
Watering and Management
Watering was carefully done by drenching the growing
medium and ensuring that no water droplets stay contained
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Flower Res. J. (2016) 24(4) : 264-273
in the ridge of the leaves which would promote rotting and
attract fungi. Growing conditions were recorded including
temperature and relative humidity by using a data logger
(Center 342 Temperature Humidity Recorder, Center
Technology Corp, Taiwan). Rotten leaf cuttings that did not
develop any roots and/or buds were removed from the
growing trays to avoid fungal growth and spread towards
other growing leaf cuttings.
Data Collection
Shoot height and diameter, the number of leaves and
shoots, shooting and rooting rate, the number of days to
rooting, and percent mortality were measured as parameters
to evaluate the response of leaf cuttings to treatments.
Statistical Analysis
The data were analyzed using SPSS ver. 22. Duncan
Multiple Range Test (DMRT) was conducted to test the
significant differences between groups.
Results and Discussion
Echeveria subsessilis
Also known as ‘Morning Beauty’, margins of this species
have touches of pink in the fully rounded by gray-blue
rosettes which is native to Mexico (Zinkan 2010). E.
subsessilis has been found to be very distinct aside from
those of E. peacockii which are grown with short stem or
none at all and accessorized with numerous and crowded
leaf formation as slightly dusty and crumbly. This species is
known to quickly leach away water from its roots and
could not tolerate submergence in water (Walter 1972).
Results of the application of auxin and cytokinin as IBA
and Kinetin for this species is shown on Table 1. Plant
height of this species was affected by auxin levels alone.
Application of 500 ppm IBA significantly gave the tallest
shoots (11.27 mm) which was followed by descending
levels of IBA, 100 ppm and 0 ppm, respectively. This was
also similar for the shoot diameter which had 14.97 mm
(500 ppm IBA) which significantly differed from those of
Flower Res. J. (2016) 24(4) : 264-273 267

Table 1. Response of E. subsessilis leaf cuttings to levels of auxin and cytokinin application.
Shoot Shoot Shooting Root No. of Rooting Percent
No. of No. of
Height Diameter Rate Length Days to Rate Mortality
Leaves Shoots
(mm) (mm) (%) (mm) Rooting (%) (%)
Auxin Level (IBA)
0 ppm 8.64 bz 11.35 b 17.29 2.46 70.56 a 25.83 a 13.81 b 65.00 b 48.89 c
100 ppm 9.54 ab 12.07 b 18.90 2.42 67.22 ab 21.88 a 11.66 c 73.33 a 56.67 b
500 ppm 11.27 a 14.97 a 14.31 1.95 62.78 b 16.26 b 20.23 a 56.67 c 73.33 a
F-test * ** NS NS * ** ** ** **
Cytokinin Level (Ki)
0 ppm 10.95 13.23 15.04 2.11 68.33 24.27 a 14.89 b 68.33 55.56 b
100 ppm 8.84 11.41 18.57 2.42 67.22 23.30 a 13.84 b 61.67 67.78 a
500 ppm 9.65 13.74 16.90 2.29 65.00 16.00 b 16.97 a 65.00 55.56 b
F-test NS NS NS NS NS ** * NS **
Interaction (IBA + Ki)
100 ppm IBA + 100 ppm Ki 9.29 10.87 26.94 a 3.11 68.33 a 21.92 10.75 c 70.00 a 60.00 b
100 ppm IBA + 500 ppm Ki 8.38 11.89 14.56 b 2.25 56.67 b 18.13 15.31 b 70.00 a 66.67 a
500 ppm IBA + 100 ppm Ki 8.19 11.70 14.33 b 1.89 65.00 a 15.86 19.80 b 50.00 b 66.67 a
500 ppm IBA + 500 ppm Ki 12.69 18.43 15.17 ab 1.83 56.67 b 13.40 22.95 a 53.33 b 63.33 ab
F-test NS NS * NS ** NS ** * **
z
Means separation within columns by Duncan’s multiple range test at P ≤ 0.05 (n = 30).
NS, *, **
mean not significant or significant at P = 0.05 or 0.01, respectively.

Fig. 1. Echeveria species with no application of any hormones A: E. subsessilis; B: E. runyonii.

100 ppm (12.07 mm) and 0 ppm (11.35 mm). These Zhao (2010) concluded that auxins, exogenous or
individual hormone responses are evident in species biosynthesis, are also responsible for seedling growth and
response shown in Fig. 2 and Fig. 1 for the control. vascular patterning which are mainly involved in
Despite its reputation to inhibit shoots, the positive developmental processes. Thus, this level of concentration
response of this species did exhibit unorthodox response. or the use of higher auxin levels, specifically IBA, has been
Auxin substances in different forms at high concentrations used for treating apple, camellia and rose cuttings (Sun and
are already classified as toxic and often retards protoplasmic Bassuk 1991; Susaj et al. 2012; Wazir 2014).
streaming which may also inhibit the morphological Interaction of auxin and cytokinin significantly affected
growing parts of the plant where it was applied (Bonner the number of leaves for this species. More number of
and Bandurski 1952; Thimann 1939). However, findings of leaves were found in plants that were pretreated with 100

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Fig. 2. E. subsessilis representative plants treated with auxin and cytokinin alone at different levels of concentration. Succulents treated
with A: 100 ppm Ki; B: 500 ppm Ki; C: 100 ppm IBA; D: 500 ppm IBA.
ppm IBA and 100 ppm Ki (26.94 leaves) which significantly
differed from higher levels: 500 ppm IBA + 500 ppm Ki
(15.17), 100 ppm IBA + 500 ppm Ki (14.56) and 500 ppm
IBA + 100 ppm Ki (14.33) that were statistically did not
differ from each other (Fig. 3).
Adjusted according to species, the formation of leaves is
flexible and changes based on certain environmental
circumstances and developmental circumstances as well as
the initiation of shoot apical meristem with variable sizes
(Bar and Ori 2014). These leaf developments are regulated
by internal and external cues which includes certain
hormones like auxins and their interaction with other plant
hormones, transcription regulators that affects its mechanical
properties of certain leaf tissues (Kaplan 2001; Zimmerman
1952). Cleland (2001) concluded that in localizing the
application of the auxin hormone would result to the
wall-loosening protein expansion which is sufficient to
induce the primordium and provide more mature leaves.
Treatments did not give a significant difference on the
number of shoots of E. subsessilis leaf cuttings. However,
the root length had a significant difference between
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Flower Res. J. (2016) 24(4) : 264-273
application of auxin and cytokinin as separate factors.
Among treatments the control (Fig. 1) gave the highest root
length with 25.83 mm for auxin levels which had similar
effects with those of 100 ppm (21.88 mm) while application
of 500 ppm gave the shortest roots with 16.26 mm. This
case was also similar to the IBA levels having the control
(24.27 mm) as the plants with highest roots which were
also similar to those of 100 ppm IBA (23.30 mm) and 500
ppm IBA (16.26 mm) having the shortest roots as well.
Su and Zhang (2011) stated that shoot meristems would
give rise to above-ground parts of the plant while the
root meristems make below-ground parts wherein
auxin-cytokinin interactions are responsible for both
meristem developments and in the excess of its levels, it
may avert the effects of its supposedly its intended
purpose. This seems to be the case for E. subsessilis.
It was observed that the plants applied with 500 ppm
gave tallest plants and largest shoot diameter, however, it
caused a decline on the root growth (16.26 mm) and also
a larger mortality (36.67%) of leaf cuttings. It was observed
that the use of hormones increased the mortality rate
Flower Res. J. (2016) 24(4) : 264-273 269

Fig. 3. E. subsessilis representative plants treated with auxin and cytokinin combinations. Succulents treated with A: 100 ppm Ki +
100 IBA; B: 500 ppm Ki + 500 ppm IBA; C: 500 ppm Ki + 100 ppm IBA; D: 100 ppm Ki + 500 ppm IBA.

around 3 - 5% compared to the control. However, without
exogenous application of hormones, there were evident
growth and quality differences of shoots produced.
After 500 ppm having the highest percent mortality, this
was then followed by plants treated with 100 ppm IBA
(28.36%) and the control (24.45%) which significantly
differed within the IBA or auxin application. The application
of 100 ppm Ki gave the highest percent mortality with
33.89% which was followed by those of 500 ppm Ki
(27.78% which were similar with those of the control.
Based on the hormone interactions, application of 500 ppm
IBA + 500 ppm Ki gave the lowest mortality rate of 21.67
% followed by other combinations which did not
statistically differ from each other.
Echeveria runyonii
Native to Mexico, numerous cultivars have been
described and cultivated with this species and also belongs,
like that of E. subsessilis, to the Crassulaceae family
(Zinkan 2010). These species are described to have
glaucous pinkish-white color of leaves that are spatulate to
oblong in shape and has similar leaves with those of E.
peacokii in terms of its glaucous feature. It is distinctively
different due to its wedged shape tips (Rose 2011).
Results on the application of auxin and cytokinin
hormones, and their combinations are shown on Table 2.
Upon the analysis of these results, the application of auxin
using IBA significantly affected the shoot height, diameter
and percentage mortality of leaf cuttings. Leaf cuttings
treated with 500 ppm IBA gave the tallest shoots with 9.22
mm which significantly differed from those of the 100 ppm
IBA (11.96 mm) and control (11.07 mm), however, this
also resulted in minimal growth of its shoots (diameter)
with the smallest shoots with 14.40 mm compared to 100
ppm IBA and the control. The application of this IBA level
also gave the highest percent morality of 42.22%. The
application of 100 ppm IBA significantly gave largest shoots
(diameter) with 16.77 mm and the lowest percent mortality
of 31.11%. Although not statistically significant, it may be
noted that the application of 100 ppm IBA also gave the
highest number of leaves (23.27) and shoots (3.05) and the
longest roots (28.47 mm).
Mahonen et al. (2014) reported that the application of
500 ppm IBA or higher concentrations of auxins rapidly

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Table 2. Response of E. runyonii leaf cuttings to levels of auxin and cytokinin application.
Shoot Shoot No. of No. of Shooting Root No. of Rooting Percent
Height Diameter Leaves Shoots Rate Length Days to Rate Mortality
(mm) (mm) (%) (mm) Rooting (%) (%)
Auxin Level (IBA)
0 ppm 11.07 az 16.03 a 21.29 2.92 85.00 a 25.77 14.94 c 62.22 32.22 b
100 ppm 11.96 a 16.77 a 23.27 3.05 81.67 ab 28.47 17.85 b 62.78 31.11 b
500 ppm 9.22 b 14.40 b 21.38 2.61 80.56 b 24.38 19.96 a 57.22 42.22 a
F-test ** ** NS NS NS NS ** NS **
Cytokinin Level (Ki)
0 ppm 11.48 a 16.08 a 21.73 b 2.69 83.33 a 26.53 17.73 ab 72.22 a 44.44 a
100 ppm 11.90 a 17.09 a 24.62 a 3.19 77.22 b 27.52 16.04 b 50.56 b 36.67 b
500 ppm 9.59 b 14.03 b 19.60 b 2.70 86.67 a 24.57 18.99 a 59.44 b 24.44 c
F-test ** ** ** NS ** NS * ** **
Interaction (IBA + Ki)
100 ppm IBA + 100 ppm Ki 10.90 a 16.23 a 20.31 2.84 81.67 b 27.97 15.40 c 65.00 a 20.00 b
100 ppm IBA + 500 ppm Ki 8.69 c 13.23 b 17.52 2.72 73.33 c 23.16 23.16 a 41.67 b 43.33 a
500 ppm IBA + 100 ppm Ki 10.06 b 15.29 b 20.82 2.45 71.67 c 23.76 19.96 b 43.33 b 56.67 a
500 ppm IBA + 500 ppm Ki 8.71 c 12.96 c 19.89 2.41 90.00 a 20.06 22.84 a 61.67 a 33.33 b
F-test ** ** NS NS ** NS ** ** **
z
Means separation within columns by Duncan’s multiple range test at P ≤ 0.05 (n = 30).
NS, *, **
mean not significant or significant at P = 0.05 or 0.01, respectively.

inhibited cell division and expansion but does not include
cell differentiation. This concept is the application of the
development of auxin herbicides such as 2 - 4 D in the
markets which has been an effective weed control in
modern production systems (Grossmann 2007). It was also
concluded besides increasing and stimulating production of
ethylene and gibberellin, auxin herbicides are able to
directly trigger gene activation of NCED.
Terminal buds or growing shoots and their development
may also be inhibited but are variable on the quantity
applied. Application of its appropriate concentrations may
also increase developing shoot growth which includes
lateral bud development and quantity of leaves (Thimann
1939). This is true to the results of leaf cuttings applied
with 100 ppm IBA.
Shoot height, diameter, number of leaves and percent
mortality were significantly affected by the application of
cytokinin and their different levels of concentration.
Application of 100 ppm Ki gave the tallest (11.90 mm),
largest shoots (17.09 mm) and more abundant leaves
(24.62), however the soot height and diameter did not
significantly differ from those of the control or 0 ppm with
11.07mm (shoot height) and 16.08 mm (shoot diameter).
Although it did not significantly affect the treatments, results
revealed that the highest number of leaves (3.19) and
shoots (27.52) were observed from plant cuttings that were
previously treated with 100 ppm Ki. However, the use of
500 ppm Ki did have the lowest percent mortality of
24.44% compared to 100 ppm Ki (36.67%) and the control
which had the highest mortality (44.44%) for E. runyonii
species. Sole application of hormones IBA and Ki on E.
runyonii species are shown on Fig. 3.
Cytokinins are important key players in the cell cycle and
influences numerous developmental programs in plant
development including cell division, shoot initiation and
growth, gametophyte, embryonic development, leaf
senescence, apical dominance, sink/source relationships,
nutrient uptake, phyllotaxis, and vascular and plant
responses to biotic and abiotic factors (Kieber and Schaller
2014; Werner et al. 2001). Thus, its deficiency effects may

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Flower Res. J. (2016) 24(4) : 264-273 271

Fig. 4. E. runyonii representative plants treated with auxin and cytokinin alone at different levels of concentration. Succulents treated
with A: 100 ppm Ki; B: 500 ppm Ki; C: 100 ppm IBA; D: 500 ppm IBA.

Fig. 5. E. runyonii representative plants treated with auxin and cytokinin combinations. Succulents treated with A: 100 ppm Ki + 100
IBA; B: 500 ppm Ki + 500 ppm IBA; C: 500 ppm Ki + 100 ppm IBA; D: 100 ppm Ki + 500 ppm IBA.

include stunted shoots and smaller apical meristems as higher concentrations it may limit the production of other
studied in tobacco plants (Werner et al. 2001). However, in hormones and may be also be detrimental for shoot

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272
development and enlargement (Brzobohaty et al. 1994).
The combination of both hormones significantly affected
shoot height and diameter including percent mortality.
Among interaction treatments, the use of lower combinations
(100 ppm IBA + 100 ppm Ki) with 10.90 mm which was
followed by leaf cuttings applied with 500 ppm IBA + 100
ppm Ki (10.06 mm), 100 ppm IBA + 500 ppm Ki (8.69
mm) and 500 ppm IBA + 500 ppm Ki (8.71 mm),
respectively. Largest shoots (diameter) were also observed
from those of 100 ppm IBA + 100 ppm Ki with 16.23 mm
and had the lowest percent mortality rate of 20.0% which
significantly differed from other treatments. Similar results
were observed with those of the other species. Combination
of hormones auxin and cytokinin on E. runyonii species
are shown on Fig. 4.
Several studies concluded from genetic and biochemical
studies of the development of these plants have very
intricate relationship for hormone and are both required for
differentiation and maintenance of meristems as well as for
continuous growth of plants which are all grounded in the
interaction of auxin and cytokinin as well as their levels of
application (Dettmer and Helariutta 2009; Su et al. 2011;
Zhao 2010).
Acknowledgement
This research was supported by ‘Succulents Export
Innovation Model Development towards Chinese Market
(514006-03-1-HD040)’, Ministry of Agriculture, Food and
Rural Affair and Sahmyook University Research Fund.
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