Clonal propagation of cashew (Anacardium occidentale L.) by stem cuttings and in vitro adventitious shoots and roots formation
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Kouakou et al., 2021 Journal of Animal & Plant Sciences (J.Anim.Plant Sci. ISSN 2071-7024)
Vol.49 (2): 8845-8855 https://doi.org/10.35759/JAnmPlSci.v49-2.2
Clonal propagation of cashew (Anacardium
occidentale L.) by stem cuttings and in vitro
adventitious shoots and roots formation
Kouakou Camille1, *Kouakou Kouakou Laurent1, Beugré Manehonon Martine2, Zoro BI Irié
Arsène1
1 Université Nangui Abrogoua, UFR Sciences de la Nature, Laboratoire de Biologie et Amélioration des Productions Végétales,
02 BP 801 Abidjan, Côte d’Ivoire
2 Université Jean Lorougnon Guédé, UFR Agroforesterie, Laboratoire d’Amélioration de la Production Végétale, BP 150 Daloa,
Côte d’Ivoire
*Author corresponding: laukouakou@gmail.com
Key words: Anacardium occidentale, micropropagation, cuttings propagation, non-mist polypropagator, genetic
resources
Date of Acceptance 21/05/2021, Publication date 31/08/2021, http://m.elewa.org/Journals/about-japs/
1 ABSTRACT
The cashew main production is derived from orchards obtained with unselected trees.
Consequently, in Côte d’Ivoire, Cashew production is characterized by poor yield and a high
susceptibility to pathogens. Conservation of elite trees in germplasm collection in order to
create genotype with high yield is a priority for research programs on cashew. This work aims
to contribute to the improvement of the cashew’s productivity. Thus, a successful cuttings
propagation method and in vitro adventitious shoots and roots formation protocol was
developed. Propagation by cuttings was performed using softwood and semi-hardwood
cuttings collected from young seedlings. Cuttings were treated with Indole-butyric-acid
(IBA) and set in non-mist poly-polypropagator. Concerning micropropagation, vitroplantlets
were cut into three parts (proximal, median, and distal) and cultured on Murashige and Skoog
medium supplemented with various concentrations of Benzyl-aminopurine (BAP) and IBA.
The time required for buds and leaf emissions was estimated for both technics of propagation.
Number of roots and emission times were also registered. Results revealed that the highest
percentages of sprouting (95%) were obtained with semi-hardwood cuttings set in non-mist
poly-propagator. The best average emergence times of buds (7 days) and leaves (13 days)
were obtained with the same cutting type treated with IBA against, 10 and 16 days respectively
for the control. Moreover, proximal and median parts induced 3 to 6 shoots per explant on
MS medium supplemented with BAP. Eighty four percent of proximal explants produce
shoots against 63% and 50% for median and apical parts, respectively. In addition, proximal
part produces more than 10 roots per explants when MS medium is supplemented with 22.5
mg l-1 IBA.
2 INTRODUCTION
Considered a forest species to combat erosion role, the planting of the cashew tree has changed
and desertification, the cashew tree (Anacardium to a socio-economic role since the 1990s, due to
occidentale L.) is now recognized as presenting the increasing demand for cashew on the
opportunities for income diversification for the international market. Production of cashews has
rural populations of central and northern Côte increased remarkably with production of raw
d'Ivoire. Thus, from an essentially ecological nuts which has increased from 19 000 tons in
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Kouakou et al., 2021 Journal of Animal & Plant Sciences (J.Anim.Plant Sci. ISSN 2071-7024)
Vol.49 (2): 8845-8855 https://doi.org/10.35759/JAnmPlSci.v49-2.2
1990 to more than 800 000 tons in 2019 (Cashew cashew is highly recalcitrant to in vitro culture and
Info, 2020). Côte d’Ivoire has become the cutting propagation. Micropropagation is
world’s largest producer of cashew nuts, affected by excessive contamination, phenolic
producing 700,000 tons, since 2015. This recent exudation, slow growth, difficulty in elongation
increase in cashew production is due in large part and microshoots rooting. It has been establish
to the expansion of land under cultivation, that the limited potential to form adventitious
which replaced areas previously devoted to other roots may in part result from inappropriate time
crops (Kouakou et al., 2020). The land area under of propagation or the physiological state of
cashew tree cultivation was about 234,000 ha in tissues (Ermel et al., 2000; Cameron et al., 2003).
2002, but is estimated at about 1,567,000 ha in According to Stankova and Panetsos (1997) and
2016. All of this Ivorian production is provided Voyiatzi et al. (2002), some species may be slow
by an orchard composed of unselected trees to form adventitious roots and the cutting may
from various origins with heterogeneous fail prior to the formation of functional roots. In
agronomic and technological characteristics, slow-to-root species, reasons for cutting failure
making them highly vulnerable to insect pests are sometime associated with tissue dehydration,
and diseases. In addition, the failure to follow loss or an inability to photosynthesis new
appropriate technical itineraries by producers is carbohydrates (Grange and Loach, 1983; Del
also one of the main causes of these low yields Rio et al., 1991; Aminah, 2003) and a limited
(350 to 500 kg / hectare) compared to the response to exogenous auxins. Many
minimum of 1000 kg/ha harvested from investigations revealed that leafy ‘softwood’
orchards in India and Brazil. The cashew cuttings need to form adventitious roots quickly
farming has recently begun research for (Howard and Harrison-Murray, 1995; Aiello and
improvement, with selection of elite trees and Graves, 1998) Only limited successes have been
more productive (Kouakou et al., 2020). Since achieved (Leva and Falcone, 1990;
2015, the Ivorian authorities have implemented Ananthakrishnan et al., 1999; Thimmappaiah et
a research program to improve cashew al., 2002) and African cashew, particularly,
production. One of the main activities is the Ivorian orchards are less documented. This
conservation in Cashew Development Centers study therefore tried to develop a cutting
(CDC) or in germplasms collection of propagation method and in vitro regeneration for
Potentially High-Producing Trees (APHP) Ivorian elite cultivars. To circumvent these
identified in the three major agroecological difficulties, non-mist polypropagators, which
zones of the country. The grafting technique ensure a greater uniformity of moisture content,
(Djaha et al., 2019; Kambou et al., 2019) usually is used for cuttings macropropagation (Leakey,
used in the production of improved plants does 1990; Newton and Jones, 1993; Kouakou et al.,
not allow a fairly faithful conservation of plant 2016), on the one hand and micropropagation
genetic resources, because of the rootstock using vitroplantlets treated with different
which always comes from another tree. It is concentrations of auxin and cytokinin for shoots
therefore necessary to seek an alternative for and roots production, on the other hand was
cloning these APHPs in order to make them achieved. The aim of this work is to develop
available to the improver. Tissue culture and efficient methods of clonal propagation of
cutting regeneration offer prospects for faster Anacardium occidentale for a germplasm collection.
multiplication of elite trees. Unfortunately,
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Kouakou et al., 2021 Journal of Animal & Plant Sciences (J.Anim.Plant Sci. ISSN 2071-7024)
Vol.49 (2): 8845-8855 https://doi.org/10.35759/JAnmPlSci.v49-2.2
3 MATERIALS AND METHODS
The experiments were carried out in Abidjan, at formation, stem cuttings were treated with a
University Nangui Abrogoua (former University commercial ‘rooting’ hormone; indole-3-butyric
Abobo-Adjame) research station (05°23´N, acid (IBA, Sigma-Aldrich Co, St Louis, USA) or
04°00´W, forest zone). Stem and seeds were left as an untreated (Control). Indeed, the basal
collected in field at Yamoussoukro (district of end of each treated cutting was sprinkled with
Yamoussoukro) situated 250 km north of IBA powder before planting. Cuttings were then
Abidjan. This site is located in the cashew planted out vertically, with the apical end up
agroecological zone. For the both sites, the (Kouakou et al., 2016). The cuttings were
rainfall pattern is bimodal with two dry seasons watered one times per week. Experiment was
(from December to March and from July to laid out in a randomized block design with two
August), and two rainy seasons (from April to treatments: the cutting type (softwood/semi-
June and from September to November). Mean hardwood), and IBA application (sprinkled) with
annual rainfall for Abidjan varies between 1800– 60 cuttings arranged in three replicates (20
2000 mm, while those of Yamoussoukro vary cuttings per replicate block) were planted out
between 1100-1200 mm. Mean monthly directly on the substrate. Five quantitative traits,
temperature varies between 27 and 30 °C, generally studied in vegetative propagation were
whereas the mean relative humidity ranges from investigated: (1) The viability estimated 60 days
70 to 84% for the both sites. after planting (cuttings remained green and alive,
3.1 Plant material: Two types of plant or dead); (2) and (3) the percentage of sprouting
materials were used (stems and vitro seedlings). and rooting, (4) the mean numbers of sprouts
(i) Stem cuttings collected from elite trees and. produced per cutting, (5) the mean emergence
(ii) Vitro plants of 30 to 45 days old (10 to 20 cm time of first leaves. The cutting was said to be
in height) obtained from seed vitrogermination. sprouted when they had one bud exceeding 0.5
3.2 Methods cm in length. Sprouting was assessed every two
3.2.1 Propagation by stem cuttings: Stems days and final percentages were calculated 60
were excised from elite plants with a disinfected days after planting.
pair of secateurs. The average length of semi- 3.2.2 Micropropagation: Mature seeds of
hardwood and softwood cuttings was 10.2 ± 2.2 cashew were surface sterilized in 70% ethanol
cm and 9.7 ± 1.8 cm, respectively. The average for 2 min followed by 30% sodium hypochlorite
diameters of semi hardwood and softwood for 30 min and rinsed three times in sterile
stems were 1.5 ± 0.2 cm and 1.20 ± 0.5 cm, distilled water. The seeds were then decocted
respectively. The cuttings propagation method and cotyledons were trimmed to half. The basal
was a non-mist poly-propagator set in a nursery, part containing embryos was cultured on
measuring 200 × 120 ×150 cm modified from Murashige and Skoog (1962) basal medium,
the design of Leakey (2014) and Kouakou et al. supplemented with 30 g l-1 sucrose and 0.7% agar
(2016). This propagator was a wooden frame and incubated in growth chamber for
enclosed in clear polythene. Covered, at the base, germination. Vitro seedlings of 30 to 45 days old
with a thin layer of fine river sand to prevent the (Figure 1) obtained were used for
polythene from being perforated by the stones micropropagation. Indeed, these vitroplants
and successive layers of small stones (Kouakou were divided in three parts: proximal, Median
et al., 2016). According to Leakey et al. (1990; and apical explant (Figure 2). Explants were
2014) and Kouakou et al. (2016), this culture cultured on MS basal medium supplemented
condition reduces the post-severance with various concentrations of BAP (22.5 and 50
physiological stress resulting from water loss. µg l-1) and IBA (50, 75, 100, 125, 150, 150, 175
Relative humidity and air temperature registered µg l-1). The pH of the media was adjusted to 5.7
under this nursery were 95 % and 29 ± 2 °C, before autoclaving at 121 °C for 30 min and 1 x
respectively. In order to stimulate root 10 Pa (1.1 kg cm-2). Three replicates were
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Kouakou et al., 2021 Journal of Animal & Plant Sciences (J.Anim.Plant Sci. ISSN 2071-7024)
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performed by the all type of explant whatever After 12 weeks of culture, (1) the percentage of
the media, each consisting of one rack with 15 explants inducing shoots with 0.3 cm height and
explants. Cultures were incubated in growth (2) the mean number of shoots produced per
chamber at 24 ± 2 °C and illuminated with cool- explants, (3) the mean number of shoots
white fluorescent lights (PPFD: 25 µmol m-2s-1) produced roots, (4) the mean time of leaves
under 14 h photoperiod and 60 – 75% humidity. emergence, was recorded.
Figure 1: Plantlets of Anacardium occidentale L. (30 days old) developed from seeds cultivated on MS
basal medium.
B C
A
Figure 2: Explants used for shoot production
A: Proximal part of explant, B: Median part of explant, C: Apical of explant
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3.3 Statistical analyses: Data were different media on the traits. When the null
analysed using Statistical package version 9.1 hypothesis of ANOVA1 was rejected, multiple
(SAS, 2004) statistical software. After checking comparisons using the Least Significant
the normal distribution of data by using Difference (LSD) test were carried out to test for
Kolmogorov-Smirnov test, One-way analysis of significant pairwise differences between cuttings
variance (ANOVA1) was performed for and media. All LSD tests were carried out at α =
assessing the influence of cutting types and 0.05 significance level.
4 RESULTS
4.1 Propagation by stem cuttings semi-hardwood cuttings, versus 9±4 and 15±2.5
4.1.1 Effect of cutting type and IBA days for bud and leaf emergence in control
application on sprouting ability of stem cuttings in a non-mist propagator. The mean
cuttings and subsequent shoot growth: numbers of sprouts (2.8±0.4) per cutting were
Cutting types affected significantly sprouting, higher from semi-hardwood (Figure 3) cuttings
sprouts and leaves emergence. The shortest time treated or not against 1.6±0.7 buds for
to sprouting was 7±2.3 days and for leaf softwood.
emergence was 13±3.5 days from IBA-treated
Figure 3: Multiple shoots of Anacardium occidentale L. induced from semi-hardwood cuttings
4.1.2 Effect of cutting type and IBA significant effect on shoots (Figure 4) and roots
application on rooting ability of stem production (P < 0.01). Whatever BAP
cuttings: Three weeks after planting, the basal concentration, proximal part gave the highest
part of all the cuttings was covered with a whitish percentages (84 to 92%) of explants bearing
powder characterizing fungus attacks. None of shoot and mean number of shoot per explant (7
the stem cuttings cultured (IBA treated and to 8 shoots). The lowest values were obtained
control) has regenerated root. Leaves have with apical and median parts. Concerning roots
dropped and cuttings, whatever the diameter production, only 11% of proximal part induced
died 65 days after planting. roots. Median and apical parts, whatever BAP
4.2 Micropropagation content, were unable to produce root (table 1).
4.2.1 Effect of BAP on shoots and roots
production: The type of explant has a
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Kouakou et al., 2021 Journal of Animal & Plant Sciences (J.Anim.Plant Sci. ISSN 2071-7024)
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A B
Figure 4: Multiple shoots of Anacardium occidentale L. induced from explants
A: proximal part of plantlets, B: median and apical part of plantlets.
4.2.3 Effect of IBA on shoots and roots explant as well as the number of explants bearing
production: According to table 2, sprouting is shoots were obtained with the proximal parts.
lowest in presence of IBA contrary to cuttings IBA high concentrations inhibited the
treated of BAP (Table 1). None of apical and production of shoots and roots. Cytokinine
median part have induced root. All the proximal (BAP) and Auxine (IBA) don’t affect
parts, whatever IBA concentration promoted significantly leaves emergence, regardless of the
significantly roots induction (Figure 5). type of explant. Therefore, leaves were emerged
Therefore, the highest percentage of roots from shoots produced for basal and median part.
production was obtained with proximal parts The mean time for leaf emergence, after shoot
cultured on 100-µg l-1 IBA. In addition, the sprouting, was 29.5±7.0 days for these both
highest values of the mean number of shoots per types of explants.
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Figure 5: Rooted proximal part of plantlets treated with indole-3-butyric acid (56 days after
incubation).
Table 1: Combined effect of BAP and explant type on sprouting and roots induction
Hormones Concentrations Type of Mean Mean Mean
(µg l-1) explant percentage of number of percentage of
explant bearing shoots per explants
shoots explant bearing roots
Apical 50±12c 2±0.1c -
22.5 Median 62.96±23b 4±1.0b -
Proximal 84±17a 7±3.2a 11.22±5.3d
BAP Apical 47.11±15c 3±0.9bc -
50 Median 61.33±13b 4±1.1b -
Proximal 92±18a 8±2.2a 11.09±8.4d
Table 2: Combined effect of IBA and explants type on sprouting and roots induction
Hormones Concentrations Type of Mean percentage Mean Mean
-1
(µg l ) explant of explant number of percentage of
bearing shoots shoots per explants bearing
explant roots
50 Apical 22.22±13de 1.9±1.1c
Median 30.66±11td 2.4±1.2c
Proximal 66.66±23b 5.7±3.2ab 68.34±23.1ab
75 Apical 20.03±18de 2.3±1.9c
Median 28.57±14d 4.2±1.1b
Proximal 47.12±22c 6.8±3.2a 66.65±17.5b
100 Apical 23.33±13de 1.1±1.9cd -
Median 22.24±21de 2.1±1.6c -
IBA Proximal 61.56±15b 4.7±3.9b 78.81±27.4a
125 Apical 19.20±17de 0.9±0.5cd -
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Median 31.01±23d 2.2±1.1c -
Proximal 57.07±24b 2.8±1.2c 65.23±18.4b
150 Apical 33.56±18d 1.3±0.6cd
Median 45.12±22c 1.9±1.6cd
Proximal 49.61±10c 2.7±3.9c 33.76±21.8c
175 Apical 07.56±01f 0.5±0.5d -
Median 18.33±11e 1.2±0.9cd -
Proximal 22.14±09de 1.8±1.2cd 19.53±5.2d
5 DISCUSSION
5.1 Stem cuttings regeneration: The cuttings would lead to a depletion of water and
chronological age differences in cutting carbohydrate reserves present in these stems.
morphology between softwood and semi- This has favoured the rapid splitting of leaves
hardwood significantly affected propagation in emitted by these cuttings. The significant
terms of rooting ability, numbers of roots, emission of leaves for semi-hardwood cuttings
sprouting and leaf emergence (Kouakou et al., could be explain by the fact that this type of
2016). Vegetative regeneration is generally best cuttings contains more carbohydrate reserve and
with leafy, softwood cuttings from young water which ensure the first metabolisms of
seedlings. This emphasises the importance of the cutting before their deterioration.
physiological state of the stockplant to 5.2 Micropropagation: Reports on cashew
regenerate (Leakey, 2014). The failure of stem indicate low rates of regeneration and abnormal
cuttings to regeneration, is always attributed to morphogenetic responses. Several factors such
several factors like the age-related physiology as tissue source and age, and composition of the
and morphology of donor plants and their culture media influence regeneration frequency
phenology (Snedden et al., 2010; Teklehaimanot (Ananthakrishnan et al., 2002). Application of
et al., 2012); including factors such as plant cytokinin at relatively high doses has been
carbohydrate and nutrient contents and the recommended for in vitro cashew bud
content of growth regulators (Dick and Leakey, development in microshoots and cotyledonary
2006 ; Maile and Nieuwenhuis 2010). According nodes (Lievens et al., 1989; Das et al., 1996). BAP
to several authors, physiological stress caused for is the most widely used cytokinin for multiple
the time between the collections of cuttings in shoots induction of several plant species such
field and planting in nursery could induce the rattan (Goh et al., 2001; Kouakou et al., 2009a; b),
loss of water and carbohydrate reserves from the Hibiscus (Samanthi et al., 2004). According to
tissues due to transpiration and respiration Samanthi et al. (2004), a single exposure of
during this period (Sanoussi et al., 2012; Leakey, Hibiscus cannabinus L. explants to a low
2014). In this study, cuttings have been cultured concentration of BAP improve the development
the same days of their harvest. The stem failure of the shoots. In this present study, BAP was
could probably due to fungus attacks. Indeed, effective in inducing shoots bud from proximal
although substrates have been treated with a explant (basal part of seedling bearing
fungicide (mancozeb Callivoire-Abidjan 880 W cotyledon). The root initiation is visible only
in water at 15 g l–1), stem cuttings were not when the adventitious buds differentiated into
treated. The fungal spores present on stem well-developed shoots. The interaction between
cuttings would be responsible for infections auxin and carbohydrates has been recognised as
observed at the base of cuttings and which vital for root formation (Sorin et al. 2005).
would have caused the obstruction of Indeed, root formation is stimulated by IBA
conducting vessels of stems and their which hydrolyses the polysaccharides, thus
deterioration. Also, non-emission of roots by the increasing the metabolic activities required to
8852Kouakou et al., 2021 Journal of Animal & Plant Sciences (J.Anim.Plant Sci. ISSN 2071-7024)
Vol.49 (2): 8845-8855 https://doi.org/10.35759/JAnmPlSci.v49-2.2
provide vital energy for the formation and seedlings. As auxin is the specialized hormone in
elongation of meristematic tissues responsible root production, the concentrations of AIB used
for root production and sprouting (Georges et or the nature of this auxin, used in this study,
al., 2007; Husen et al., 2007). The proximal would be inadequate to the neoformation of
explant is the part of the seedling that ends with roots on these both parts. The production of
the radicle. The proximal part contains cells roots would also be promoted by the high
already differentiated and predisposed to roots number of shoots produced on this part.
production unlike median and apical parts of the
6 CONCLUSION
This preliminary work on West African needed to determine the optimal conditions for
Anacardium micropropagation gives quite robust and larger scale propagation of
promising results. In fact, this plant has always Anacardium occidentale. Efficient conditions of
been regarded as recalcitrant to culture such as temperature, relative humidity
micropropogation, because of the difficulties in and a further improvement of culture medium
rooting. The significant production of shoots composition and substrate is required to increase
and roots of the proximal part of the seedlings is the time of rooting as well as in nursery and vitro
a significant step towards the exponential culture. Studies on the survival and growth of
cultivation of cultivars with intersecting plantlets under nursery and field conditions are
agronomic characteristics. Further research is necessary.
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