The comparative efficiency of a brown

algal-derived biostimulant extract

(AMPEP), with and without supplemented
PGRs: the induction of direct, axis
shoots as applied to the propagation of
vegetative seedlings for the successful mass
cultivation of three commercial strains of
Kappaphycus in Sabah, Malaysia
Majid Majahar Ali, Mohd Zaidey Bin
Sani, Kim Keon Hi, Suhaimi Md. Yasir,
Journal of Applied Phycology

et al.
ISSN 0921-8971
Volume 30
Number 3

J Appl Phycol (2018) 30:1913-1919

DOI 10.1007/s10811-017-1366-1

1 23
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1 23
 Author's personal copy
Journal of Applied Phycology (2018) 30:1913–1919
https://doi.org/10.1007/s10811-017-1366-1

The comparative efficiency of a brown algal-derived biostimulant

extract (AMPEP), with and without supplemented PGRs:
the induction of direct, axis shoots as applied to the propagation
of vegetative seedlings for the successful mass cultivation of three
commercial strains of Kappaphycus in Sabah, Malaysia
Majid Majahar Ali 1,2 & Mohd Zaidey Bin Sani 2 & Kim Keon Hi 2 & Suhaimi Md. Yasir 2 & Alan T. Critchley 3 &
Anicia Q. Hurtado 4

Received: 31 August 2017 / Revised and accepted: 6 December 2017 / Published online: 16 December 2017
# Springer Science+Business Media B.V., part of Springer Nature 2017

Abstract
Three strains of Kappaphycus spp. (viz. K. alvarezii tambalang brown and green and K. striatus sacol green) were used in the
present study to optimize the use of Ascophyllum (Acadian) marine plant extract powder (AMPEP) as a culture medium
ingredient acting as a biostimulant, applied with, and without, the addition of terrestrial plant growth regulators (PGRs). This
was undertaken in order to develop management tools and best practice recommendations for the mass production of new
plantlets (seedlings) for industrial, nursery, and out-planting purposes in eastern Malaysia, Sabah, and Peninsular Malaysia
(i.e., Langkawi, Kedah and Batu Maung, Penang). After 45 days of laboratory incubation, the three strains tested demonstrated
their best performances at 3 mg L−1 of AMPEP, supplemented with PGR. This evaluation was based on the longest direct axes
formed, the shortest time to their appearance, and also their highest percentage emergence. Kappaphycus alvarezii (tambalang
green) had the longest direct axes (7.0 ± 0.23 mm), followed by K. alvarezii (tambalang brown) at (6.4 ± 0.48 mm) and finally
K. striatus (sacol green). In terms of the highest percentage of direct axes formed, K. alvarezii (tambalang brown), K. alvarezii
(tambalang green), and K. striatus (sacol green) were recorded as follows: 100 ± 0.00, 99 ± 1.34, and 98 ± 2.66, respectively. The
shortest duration taken for the emergence of direct axes was observed in K. alvarezii (tambalang green) followed by tambalang
brown and K. striatus (sacol green) on days 9, 10, and 15, respectively. The use of a brown seaweed-derived extract acting as a
biostimulant and as the main ingredient of the culture medium for the micropropagation of three strains of Kappaphycus was
highly encouraging and one which may be promoted as a protocol for the economic and commercial mass production of new
plantlets (asexual seedlings) which are an urgent requirement for Malaysian seaweed farming to meet its full potential.

Keywords Kappaphycus alvarezii . K. striatus . AMPEP . Biostimulant . Direct axis shoots . Micropropagation

Introduction
* Anicia Q. Hurtado
anicia.hurtado@gmail.com Kappaphycus is a genus of economically important, red sea-
1
weeds distributed and cultivated in the tropics and sub-tropics,
School of Mathematical Sciences, Universiti Sains Malaysia,
primarily as a source of raw materials for the production of the
11800 Gelugor, Penang, Malaysia
2
industrial gel carrageenan (Bixler and Porse 2011; Porse and
Seaweed Research Unit, Faculty of Science and Natural Resources,
Rudolph 2017). The availability of Kappaphycus as raw ma-
Universiti Malaysia Sabah, Jalan UMS, 88400 Kota
Kinabalu, Sabah, Malaysia terial for processing is possible because of extensive, open-
3 water farming which has been developed and practiced for
Verschuren Centre for Sustainability in Energy and the Environment,
Cape Breton University, Sydney, Nova Scotia B1P 6L2, Canada more than four decades. There are several successful stories
4 of its farming in the Philippines and elsewhere. However, it is
Integrated Services for the Development of Aquaculture and
Fisheries, (ISDA) Inc., McArthur Highway, Tabuc Suba Jaro, not spared technical difficulties such as low productivity and
5000 Iloilo City, Philippines lost production capacity simply due to the lack of availability
 Author's personal copy
1914
and access to high-quality propagules for out-planting. Such
shortages are increasing in frequency due to a high incidence
of disease and pest infestations, at least in part, due to poor
farm management practices (Cottier-Cook et al. 2016). The
increasingly high incidence of disease and pest infestations
in Kappaphycus is likely to be also attributed to its low genetic
variability as well as the fact it is almost exclusively propa-
gated by asexual multiplication and as a monocrop
(Zuccarello et al. 2006; Tan et al. 2012; Lim et al. 2014).
Commercial-scale Kappaphycus farming was introduced
to Sabah, Malaysia, in the late 1970s (Sade et al. 2006).
Farming was introduced in Semporna, east coast of Sabah,
by the Bajau and the Suluk people from Tawi-Tawi,
Philippines (Eranza et al. 2015) and operated commercially
by the American company Aquatic Resources Limited. The
north-east coast of Sabah is endowed with literally thousands
of areas suitable for open-water, extensive cultivation (e.g.,
approximately 38,500 ha, Ali Pers. comm.). The Malaysian
government has worked on several economic plans; each of
which was intended to help local coastal economies and assist
in the alleviation of poverty. However, the annual production
of eucheumatoids in Malaysia has not fulfilled its anticipated
potential, as compared to neighboring countries such as the
Philippines and Indonesia. There appear to be three major
issues requiring resolution in order to enable seaweed farming
to become an economically sustainable activity (Eranza et al.
2015), namely, (1) seaweed farming challenges between
Malaysian islanders and immigrant seaweed farmers, (2) sea-
weed farmers and the middlemen who purchase their cultivat-
ed biomass, and (3) seaweed quality, and the price paid to the
farmer and use of the biomass by companies producing semi-
refined carrageenan (SRC). These issues were confirmed by
the latest report of Nor et al. (2017) which highlighted the
following: (1) participation of indigenous people in seaweed
cultivation must be promoted, (2) there should be legalization
of activities by existing migrant seaweed farmers, (3) there is a
need to strengthen local seaweed cooperative organizations,
(4) there needs to be provision for entrepreneurship skills
training to farmers, and (5) the full integration of all stake-
holders into a seaweed farming decision-making protocol is a
requirement.
Aside from socio-economic issues and concerns, technical
problems related to the productivity and production of
Kappaphycus raw materials in Malaysia are important chal-
lenges to be overcome. As with other geographic jurisdictions,
Malaysia shares common problems of disease and pest infes-
tations in those species used as seaweed crops (Vairappan
2006; Vairappan et al. 2008; Ali et al. Pers. comm.). Since
all cultivation is dependent on the production of asexual
Bcuttings^ or Bseedlings,^ the availability of good-quality
propagules for cultivation is a major bottleneck, facing the
entire industry of warm-water, carrageenophyte farming.
J Appl Phycol (2018) 30:1913–1919
Due to its importance as a critical production stage, the
micropropagation of Kappaphycus and Eucheuma spp. has
been reported by earlier authors as a reliable source of new
and improved propagules for commercial cultivation (Dawes
and Koch 1991; Dawes et al. 1993, 1994; Hurtado and
Cheney 2003; Reddy et al. 2003; Hurtado and Biter 2007;
Hayashi et al. 2008; Hurtado et al. 2009; Yunque et al. 2011;
Sulistiani et al. 2012; Yong et al. 2014; Neves et al. 2015;
Tibubos et al. 2017) using multiple varieties of culture media.
This study was conducted using a commercial extract of
the cold-water, brown seaweed Ascophyllum nodosum, named
AMPEP for these studies (Ascophyllum marine plant extract
powder; Hurtado et al. 2009; Borlongan et al. 2011; Yunque
et al. 2011; Hurtado et al. 2012; Tibubos et al. 2017) and
applied for its biostimulatory properties, taking a lead from
several reviews of its benefits when applied to terrestrial plants
(Craigie 2011; Khan et al. 2009). The studies required the
optimization of applied concentrations for the induction of
direct, axis shoots of three Kappaphycus spp. (viz.
K. alvarezii—tambalang brown and green morphotypes and
K. striatus—sacol green) in order to develop management
protocols to be used in their mass production for sourcing
propagules to supply to open-water field cultivation for the
Malaysian seaweed industry.
Materials and methods
Kappaphycus alvarezii (tambalang brown and green) and
Kappaphycus striatus (sacol green) plants were taken from
the Universiti Malaysia Sabah (UMS) mini-estate farm in
Semporna, Sabah. While on site, the seaweeds were cleaned
of any adhering particles and organisms such as sand, shells,
and obvious ecto-epiphytes, neatly packed and finally covered
with seawater-moistened cheesecloth, and transported in an
ice chest to the Seaweed Unit Laboratory of UMS.
Following the procedures described by Hurtado and Cheney
(2003), the seaweeds were acclimated in a fiberglass tank (5 ×
1.5 m) for 5–7 days using UV-treated seawater provided with
mild aeration. A daily change of UV-treated seawater was
made.
Only clean, healthy, and epiphyte-free apical segments (3–
5 cm long) were cut and used as explants. These apical seg-
ments were cleaned and sterilized using Povidone iodine (1%)
by brushing the surface with a soft artist’s brush, and then
rinsed three times with autoclaved seawater in order to provide
clean, healthy, and diatom-free explants. Apical segments
were cut into 2–3-mm-thick segments, washed, and rinsed
three times with autoclaved seawater. Five sections (1 section,
10 mL) were placed inside a 45-mL T2 flask containing a
range of concentrations of Ascophyllum (aka Acadian) marine
plant extract powder (AMPEP) + PGR and used to induce
formation of direct axis shoots.
 Author's personal copy
J Appl Phycol (2018) 30:1913–1919
Plant growth hormones (or plant growth regulators—
PGRs, i.e., indole acetic acid (IAA) and kinetin) were added
also to each treatment at 1 mg L−1 following the procedure of
Hurtado and Cheney (2003). The following concentrations
were used for solutions of AMPEP alone: 0.1, 0.5, 1.0, 3.0,
5.0, and 10 mg L−1; these were also prepared in combination
with PGR at 1 mg L−1. The treatment concentrations for
AMPEP and the PGR were taken after Tibubos et al. (2017).
The present study was designed to determine the conditions
necessary to optimize Kappaphycus cultivation in the
Malaysian seaweed industry. A total of 90 T2 flasks (i.e., 3
strains × 6 concentrations of AMPEP, with and without, PGR
× 5 replicates) were used. All treatments were placed on a
shaker (Laboratory Shaker Benchtop-HY-5 Shaker) at 4–
8 rpm and incubated for 45 days, in a walk-in culture room
at 23–24 °C, 13L:11D photoperiod, and an irradiance of 30–
40 μmol photons m−2 s−1. Changes of media were made every
7 days. The following parameters were observed and record-
ed: (1) the first emergence of direct axes and (2) the length
(mm), number, and percentage of new direct axes formed. The
percentage of direct axes formed was computed as follows:
The percentage of direct axes formed (%) = (number of
shoots formed at day 45 total number of explants incubated
at day 1) × 100 (after Tibubos et al. (2017)).
The results obtained were tested statistically using a one-
way analysis of variance (ANOVA, P < 0.01) and pairs of
treatment means were compared by Duncan’s multiple range
test (DMRT) at P < 0.01 and P < 0.1 using the SPSS Software
Program 21.0.
Results
Length of direct axis shoots
Results showed that the average length of those direct axes
formed after 45 days of incubation varied according to both
the strain selected and the applied concentration of AMPEP in
combination with the PGR (IAA and kinetin).
The direct axis shoots formed in K. alvarezii (tambalang
brown) ranged from 1.0 ± 0.07 to 6.4 ± 0.48 mm with
0.5 mg L−1 AMPEP (control) and 3 mg L−1 AMPEP + PGR
treatments, respectively; for K. alvarezii (tambalang green)
from 0.8 ± 0.06 to 7.0 ± 0.23 mm with 0.1 mg L−1 AMPEP
alone (control) and 3 mg L−1 AMPEP + PGR, respectively;
and K. striatus (sacol green) from 0.6 ± 0.06 to 3.3 ± 0.08 mm
at 5 mg L−1 AMPEP (control) and 3 mg L−1 AMPEP + PGR,
respectively. Amongst the three strains tested, K. alvarezii
(tambalang green) produced the longest direct axes (6.4 ±
0.48 mm), while K. striatus (sacol green) had the shortest
(0.6 ± 0.06 mm).
There was a significant (P < 0.01) interaction of strain with
various concentrations of AMPEP + PGR. The effect in length
1915
of the axes formed was dependent on the strain and concen-
tration of AMPEP in the presence of PGRs. Amongst the
interactions, the concentration of AMPEP applied and the
presence of PGR were the major factor that contributed to
significance of the combination of factors. As shown in
Fig. 1a–c, the mean length of the shoots was highest in all
strains at the 3.0 mg L−1 AMPEP, with or without PGR.
However, the response of axis length was the best with the
presence of PGR for the K. alvarezii brown and tambalang
green morphotypes. There were no significant differences in
the length of the direct axes formed between tambalang green
and brown morphotypes at 3 mg L−1 AMPEP + PGR. In all
strains used, the length of those direct axes formed had their
best response at 3 mg L−1 AMPEP singly, and in combination
with PGR.
Percentage of direct axes shoots formed
The average percentage of direct axes formed after 45 days of
incubation ranged from 55 ± 5.73 to 100 ± 0.00 in K. alvarezii
(tambalang brown) at 0.5 mg L −1 AMPEP K +(control)
and 3 mg L−1 AMPEP + PGR, respectively; 65 ± 6.47 to 99
± 1.34 in K. alvarezii (tambalang green) at 10 mg L−1 AMPEP
(control) and 3 mg L−1 AMPEP + PGR, respectively; and 52
± 5.73 to 98 ± 2.66 in K. striatus (sacol green) at 0.1 mg L−1
AMPEP (control) and 3 mg L−1 AMPEP + PGR, respectively
(Table 1).
In considering the percentage of direct axes, formed after
45 days, there was a significant (P < 0.01) interaction between
strain and AMPEP, and between strain and PGR, but not in
combination with AMPEP and PGR (Figs. 2a–c). The inter-
actions of strain, levels of AMPEP, and supplemental presence
of PGR were significant (P < 0.01). The effects on shoot
length were dependent on the combination of strain and levels
of AMPEP and the presence of PGR. Amongst the interac-
tions, the various levels of AMPEP and supplemental pres-
ence of PGR were the major factors contributing to signifi-
cance (partial eta squared = 0.475 or 47.5%) of the factorial
combinations. The mean lengths of direct shoots were longest
in all strains at 3.0 mg L−1 AMPEP with or without PGR.
However, the greatest response in terms of length was to the
supplementation of AMPEP with addition by PGRs treatment
in K. alvarezii, both tambalang brown and green strains.
Emergence of direct axes
The shortest treatment time taken for the first emergence of
direct axes (i.e., day 9) was observed in K. alvarezii
(tambalang green), both at 3 mg L−1 AMPEP + PGR and
the corresponding AMPEP only (control), followed by
K. alvarezii (tambalang brown) on day 11, with 3 mg L−1
AMPEP + PGR, and lastly by K. striatus (sacol green) on
day 13 with 3 mg L−1 AMPEP + PGR (Table 1). On the other
 Author's personal copy
1916 J Appl Phycol (2018) 30:1913–1919

8 (a) K. alvarezii (tambalang brown)

a hand, the longest duration for new shoot formation was days
7
30 and 27, observed in K. alvarezii (brown) at 0.5 mg L−1
6 AMPEP alone (control) and K. striatus (sacol green) with
5 c 0.1 mg L−1 AMPEP + PGR and at 0.5 mg L−1 AMPEP alone
4 de
(control), and K. alvarezii (tambalang green) at 0.1 mg L−1
de de
3 AMPEP control, respectively. Table 2 shows a summary of
fgh the highest/longest and lowest/shortest shoots in relation to the
2 de
Length (mm) of direct axes after 45 days of incubation

ef length of direct axes (mm), percentage of direct axes formed,

1
fghi fghi fghi and first day on which direct axes emerged.
0 hi
Amongst the interactions (strain × AMPEP, strain × PGR,
AMPEP × PGR, and strain × AMPEP × PGR), strain ×
8 (b) K. alvarezii (tambalang green)
a AMPEP had the highest contribution to the response in num-
7
ber of days of emergence. However, strain, AMPEP, and PGR
6 b
c were dependent variables on each other on the impact of treat-
5 ment days taken before the first emergence of shoots.
cd
4 de Likewise, significant differences were found between those
3 concentrations of AMPEP used with and without PGR which
fghi cd
2 varied by strain used (P < 0.01).
efg
1 fgh
fghi fghi
0 hi
Discussion
8
7 (c) K. striatus (sacol green)
Several reports on the micropropagation of Kappaphycus and
6
Eucheuma from callus have been published (Dawes and Koch
5 AMPEP with PGR
AMPEP Without PGR
1991; Dawes et al. 1993; Hurtado and Cheney 2003; Reddy
4 cd et al. 2003; Hurtado and Biter 2007; Hayashi et al. 2008;
3 ef Sulistiani et al. 2012). However, few recent micropropagation
fghi
2
hi ghi fghi studies have been reported from Sabah, Malaysia, which
1 hi ranged from direct regeneration (Yong et al. 2014), evaluation
hi fghi hi
0 i hi of growth rate and semi-refined carrageenan (Yong et al.
0.1 0.5 1 3 5 10
2014), acclimation in a nursery system (Yong et al. 2015),
Concentration of AMPEP (mg L-1) with PGR at 1mg L-1
chemical composition (Yong et al. 2015) to the presence of
Fig. 1 Average (x ± SEM) length of direct axes (mm) in a K. alvarezii heavy metals in the tissues of the cultivated materials (Yong
(tambalang brown), b K. alvarezii (tambalang green), and c K. striatus
(sacol green) (n = 5)
et al. 2017). Fundamentally, the earlier studies used inorganic
sources of nutrients in the preparation of their culture media.
Interestingly, only the reports of Hurtado et al. (2009), Yunque

Table 1 Average first day of

emergence of direct axes after Concentration K. alvarezii (brown) K. alvarezii (green) K. striatus (green)
incubation in K. alvarezii (mg L−1)
(tambalang brown), K. alvarezii
(tambalang green), and K. striatus AMPEP with PGR 0.1 19 22 31
(sacol green) (n = 5) 0.5 15 15 26
1 15 12 23
3 10 9 13
5 13 11 18
10 15 14 25
AMPEP only (control) 0.1 18 27 18
0.5 30 15 30
1 23 20 23
3 23 9 23
5 23 18 23
10 23 19 23
 Author's personal copy
J Appl Phycol (2018) 30:1913–1919 1917

ab a
abc
100 abcd
abcd
abcd The present study reports results for optimization of the
90 micropropagation of commercially important eucheumatoids,
80
70 abcde
viz. two K. alvarezii (tambalang brown and green) and one
60 def abcdef cdef cdef K. striatus (sacol green) commercially grown in Semporna,
50
ef Sabah, Malaysia, using AMPEP at different concentrations,
40
singly or in combination with PGRs. The formation of direct
Percentage (%) of direct axes formed after 45 days of incubation

30
20 (a) K. alvarezii (tambalang brown) axes in the three strains of Kappaphycus used in the present
10 study required 3 mg L−1 of AMPEP + PGR (at 1 mg L−1 IAA
0
a and kinetin) which is comparable with the results of Hurtado
100 abcd
abcd et al. (2009) using the same concentration of AMPEP and IAA
and zeatin as the PGR stimulant at 1 mg L−1. However, the
90
80 cdef def def
70 abcdef report of Yunque et al. (2011) required much lower concen-
abcdef abcdef
60 cdef def ef
trations of AMPEP + PGR for successful regeneration in
50
K. alvarezii (brown, 0.1 mg L −1 ), 0.001 mg L −1 in
K. alvarezii (green), and 1.0 mg L−1 in K. striatus (green),
40
30
20 (b) K. alvarezii (tambalang green) viz. the present study which required 3 mg L−1 AMPEP with
10 additional PGR. The differences might be attributed to the
0
ab type of PGR used which induced both cell division and elon-
abcd
100
abcde abcdef
gation, as well as the influence of the geographical location,
90 ab
80 def provenance of the material used to produce seedlings, and the
70 ef
abcde
abcdef
season when the Kappaphycus strains were taken. However,
60
def def in terms of the length (mm) of the direct axes formed, the
50
40
ef present study is comparable to the results of Yunque et al.
30 AMPEP with PGR (2011). It is rather difficult to compare the results of the pres-
20
AMPEP Without PGR (c) K. striatus (sacol green) ent study to those of Tibubos et al. (2017) which reported on
10
0
the use of AMPEP K+, though the same sources of PGRs were
0.1 0.5 1 3 5 10 used (i.e., IAA and kinetin).
Concentration of AMPEP (mg L-1) with PGR at 1 mg L-1
Amongst the three strains tested in the present study, it was
Fig. 2 Average (x ± SEM) percentage of direct axes formed in: a only K. alvarezii (brown) at 3 mg L−1 AMPEP with PGR
K. alvarezii (tambalang brown), b K. alvarezii (tambalang green), and c
which produced 100% direct axes after 45 days, as compared
K. striatus (sacol green) (n = 5)
to Tibubos et al. (2017) (for K. alvarezii brown strain) which
reported 100% direct axes at 0.5, 1, 5, and 10 mg L−1 with the
et al. (2011), and Tibubos et al. (2017) reported the successful exception of 0.1 mg L−1 AMPEP K++ PGRs. The differences
micropropagation of various strains of Kappaphycus and de- in responses of the brown strain might be attributed to the
scribed the used of organic sources of nutrients and stress and health status of the explants used and also
biostimulant activity when using a commercial seaweed ex- probably due to the geographical location where the explants
tract from the brown alga Ascophyllum nodosum (i.e., in both were gathered. The explants of Tibubos et al. (2017) were
AMPEP and K+ forms—a potassium fortified AMPEP; see collected in a commercial farm in central Vietnam.
references above for details). Kappaphycus alvarezii, originally indigenous to the

Table 2 Summary of highest/

longest and lowest/shortest shoots Highest/longest
in relation to length of direct axes
(mm), percentage of direct axes Length of direct axes (mm) 7 ± 0.23
formed, and first day on which 3 mg L−1 AMPEP + PGR
direct axes emerged K. alvarezii (tambalang green)
Percentage of direct axes formed 100 ± 0.0
3 mg L−1 AMPEP + PGR
K. alvarezii (tambalang brown)
First day on which direct axes emerged 31 ± 0.98
0.1 mg L−1 AMPEP only
K. striatus (sacol green)
Lowest/shortest
0.7 ± 0.11
5 mg L−1 AMPEP only
K. striatus (sacol green)
52 ± 5.73
0.1 mg L−1 AMPEP only
K. striatus (sacol green)
9 ± 0.37
3 mg L−1 AMPEP + PGR
K. alvarezii (tambalang green)
 Author's personal copy
1918
Philippines, was first introduced from cultures held in Japan
(Ohno et al. 1995, 1996) to the Institute of Applied Material
Technology in Nha Trang and the Fisheries Extension
Department in Ninh Thuan, in 1993; after which, its commer-
cial cultivation started 3 years later in the central and southern
regions (Nang and Dinh 1998).
The advantages of using a brown seaweed extract as the
primary source of ingredient for the micropropagation and
field cultivation of the rhodophyte Kappaphycus strains lies
in its source as a natural source of biostimulant and bioeffector
properties (van Oosten et al. 2017). These come in the form of
efficacy derived from natural, phytohormone-like com-
pounds, physiological primers, and bio-signalling molecules
(as yet unidentified), as reported by earlier studies in their
applications to terrestrial, agricultural crops (Crouch and van
Staden 1992; Leclerc et al. 2006; Rayirath et al. 2008; Fan
et al. 2011; Spann and Little 2011; van Oosten et al. 2017).
Indeed, as marine agronomy becomes increasingly important
in the food supply required for the earth’s burgeoning popu-
lations, much more research will be required in keeping the
pests and diseases of cultivated seaweed crops at bay, no dif-
ferent from the intervention methods developed for terrestrial
agronomy (see Cottier-Cook et al. (2016) and Buschmann
et al. (2017)). The positive benefits specifically derived from
Ascophyllum extracts are covered under the descriptions of
biostimulant/bioeffector properties, as reviewed by van
Oosten et al. (2017), namely, as an enhancer or inducer for
the formation of direct axis formation and growth (i.e., abiotic
responses; Hurtado et al. 2009; Yunque et al. 2011; Hurtado
et al. 2012; Tibubos et al. 2017) and as a priming agent or
mitigator to biotic responses to damaging endophytes
(Borlongan et al. 2011; Hurtado et al. 2012; Loureiro et al.
2012), epiphytes, and epibionts (Loureiro et al. 2010; Marroig
et al. 2016) and as stimulant in in-tank cultivation (Pedra et al.
2017). However, an in-depth economic analysis for the use of
a variety of seaweed extracts, as the main ingredient for the
culture medium of commercially important red seaweeds of
the Kappaphycus genus, as compared to the use of inorganic
sources of ingredients, is something worthy in future studies.
Hurtado et al. (2017) provided an extensive update to the
challenges of warm-water carrageenophyte farming on a glob-
al basis; future prospects for Malaysia and sustainable seed-
ling production are major issues covered in the collated book.
The present study provides evidence for the positive benefits
of using a given seaweed extract in the micropropagation of
Kappaphycus strains in Sabah, Malaysia, and, as such, pro-
vides an important management tool which could be used in
commercial, university, or state-sponsored seaweed nurseries
for the mass production of new plantlets to become an abun-
dant and healthy source of propagules for the future of im-
proved, sustainable commercial cultivation practices. Taken
together, these interventions will hopefully lead to achieving
the full potential of carrageenophyte farming and the
J Appl Phycol (2018) 30:1913–1919
following socio-economic benefits for coastal communities
in Malaysia.
Acknowledgements We thank the two reviewers of this article for useful
comments leading to an improvement of the manuscript. The authors are
thankful to Acadian Seaplants Limited, Nova Scotia, Canada, for their
donation of the AMPEP sample used for this study.
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