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Fruits 73(6), 342–348 | ISSN 0248-1294 print, 1625-967X online | https://doi.org/10.17660/th2018/73.6.

4 | © ISHS 2018

Review article – Thematic Issue

Traditional enset [Ensete ventricosum (Welw.) Cheesman]


sucker propagation methods and opportunities for crop
improvement
Z. Yemataw1, K. Tawle1, G. Blomme2,a and K. Jacobsen3
1 The Southern Agricultural Research Institute (SARI-Areka), Areka Agricultural Research Center, P.O. Box 79, Areka, Ethiopia
2 Bioversity International, c/o ILRI, P.O. Box 5689, Addis Ababa, Ethiopia
3 Royal Museum for Central Africa, Leuvensesteenweg 13, 3080 Tervuren, Belgium

Summary Significance of this study


Introduction  –  This review focuses on the enset What is already known on this subject?
seed systems in Ethiopia and explores opportunities • Traditional macro-propagation methods, using entire
to improve the system. Cultivated enset is predomi- rhizomes or rhizome pieces, currently suffice to pro-
nantly vegetatively propagated by farmers. Repro- vide the needed enset suckers at farm, village or land-
duction of an enset plant from seed is seldom prac- scape level.
ticed by farmers and has been reported only from
the highlands of Gardula. Seedlings arising from seed What are the new findings?
are reported to be less vigorous than the suckers • When larger quantities of suckers are needed, e.g.,
obtained through vegetative propagation. Rhizomes when introducing a new enset cultivar or coping with
from immature plants, between 2 and 6 years old, severe disease or pest impacts, improved/novel mi-
are preferred for the production of suckers. The aver- cro- and macro-propagation techniques, as listed in
age number of suckers produced per rhizome ranges this review paper, could offer solutions.
from 40 to 200, depending on soil conditions, culti-
var type, size and age of the parent plant, amount of What is the expected impact on horticulture?
rainfall, land preparation and time of planting. Tra- • Clean enset planting materials as part of an integrated
ditional macro-propagation of enset suckers involves field and landscape management approach will con-
both men and women farmers. Sucker propagation tribute to healthier and more resilient systems.
and transplanting activities often contribute to the
dissemination or maintenance of pests (e.g., the en-
set root mealy bug or nematodes) and diseases (e.g.,
enset bacterial wilt). Conclusion  –  Macro- and mi-
issues de semences seraient moins vigoureuses
cro-propagation are useful technologies to improve
que les rejets obtenus par multiplication végétative.
the efficiency of sucker production and to provide
Les rhizomes de plantes immatures, âgées de 2 à 6
clean replacement plants in locations where diseases
ans, sont préférés pour la production de rejets. Le
have affected plantations or to locally multiply newly
nombre moyen de rejets produits par rhizome varie
introduced cultivars for distribution.
de 40 à 200, en fonction du type de sol et de cultivar,
de la taille et de l’âge de la plante mère, de la pluvio-
Keywords
métrie, de la préparation du sol et de la saison de
enset, Ethiopia, macro-propagation, micro-propagation,
plantation. La macro-propagation traditionnelle des
vegetative propagation
rejets d’ensète mobilise les agriculteurs, hommes et
femmes. Les activités de propagation et de repiquage
Résumé des rejets contribuent souvent à la dissémination ou
Méthodes de propagation des rejets d’en- au maintien d’organismes nuisibles (par exemple, la
sète traditionnel [Ensete ventricosum (Welw.) punaise cornéenne ou les nématodes) et de maladies
(par exemple, le flétrissement bactérien de l’ensète).
Cheesman] et opportunités pour l’amélioration Conclusion  –  La macro- et la micro-propagation sont
culturale. des technologies utiles pour améliorer l’efficacité de
Introduction  –  Cette revue se concentre sur les la production de rejets, et pour fournir des plants
systèmes semenciers de l’ensète en Ethiopie et ex- sains de remplacement aux endroits où les maladies
plore les opportunités pour améliorer le système. ont affecté les plantations ou pour multiplier locale-
L’ensète cultivée est principalement multipliée par ment des cultivars nouvellement introduits.
voie végétative par les agriculteurs. La reproduction
d’un plant d’ensète à partir d’une graine est rarement Mots-clés
pratiquée par les producteurs et n’a été signalée que ensète, Ensete ventricosum, Ethiopie, macro-propagation,
sur les hauts plateaux de Gardula. Les plantules micro-propagation, multiplication végétative

a Corresponding author: g.blomme@cgiar.org.

342 International Journal of Tropical and Subtropical Horticulture


Yemataw et al. | Traditional enset sucker propagation methods and opportunities for crop improvement

Introduction ular food scarcity, family or social obligations and annual


The arable Ethiopian highlands, ranging from 1,200 ceremonies, and an often limited number of enset plants
to 3,100 m a.s.l., and located between 35°17’23.56” and per farm, households most often harvest enset before the
40°23’24.35”E and 5°18’22.38” and 14°09’47.70”N, are plant reaches full maturity and a bunch/fruits with seeds
considered as one of the foremost centres of origin for the is developed (Pankhurst, 1996). In addition, enset plants
domestication of a wide variety of plants, including wheat should preferably be harvested at or immediately after flow-
(Triticum), barley (Hordeum vulgare), coffee (Coffea), vari- er emergence to produce optimum quantities and quality of
ous pulses, tef (Eragrostis tef) and enset (Ensete ventricosum) kocho (i.e., fermented starch obtained from grated rhizome,
(Vavilov, 1926; Brandt, 1996; Haile et al., 1996). The domes- real stem and pseudostem leaf sheath tissue) (Tsegaye and
tication of enset is estimated to have arisen in Ethiopia as Struik, 2002; Yemataw et al., 2014). At the farm-level, propa-
early as 10,000 years ago (Brandt et al., 1997). As a genus gation of enset cultivars is primarily vegetative, through the
in the Musaceae family, enset shares many botanical char- production of suckers.
acteristics with its close relative, the banana. The wild and In a typical enset farm, the main activities include prepar-
cultivated Ensete ventricosum is a large, tree-like monocot, ing and planting rhizomes for sucker production, production
with stiff, semi-erect leaves and a flower that resembles that of young suckers (Figure 1), subsequent one or more steps of
of banana, but which produces no edible fruit. transplanting of the suckers (i.e., plants are uprooted, most of
As with wild enset, cultivated enset often produces seed. the roots and leaves are removed and plants are replanted at a
However, cultivated enset is predominantly propagated veg- wider spacing), final transplanting, management of the mature
etatively. Traditional vegetative macro-propagation methods enset plot, harvesting plants, processing of pseudostem leaf
can rapidly multiply large quantities of planting materials sheaths and rhizomes, and fermentation and storage of kocho/
and are sustainable in the absence of pests (e.g., the enset starch. This review focuses on the enset sucker propagation
root mealy bug) or diseases (e.g., Xanthomonas wilt of en- methods/system (i.e., all steps before the first transplanting)
set). However, when pest/disease-affected plants are pres- and opportunities to improve the system. This review is based
ent in a field, it becomes more difficult to produce healthy on publications, grey literature and reports which were com-
planting material using traditional propagation methods. piled by the Southern Agricultural Research Institute (SARI)
This makes enset farming systems vulnerable, particularly and Bioversity International. Various aspects of enset prop-
in the context of biotic factors and climate change/higher agation have been researched on over the past decades, and
temperatures that could worsen the situation (Brandt et al., this review provides a structured overview of results and rec-
1997; McKnight, 2013). ommendations, and discusses research knowledge gaps in the
Propagation by seed is possible, but for reasons of reg- light of possible future research endeavours.

Figure 1.  Traditional macro-propagation of enset seedlings in a backyard in southern Ethiopia (Source: Guy Blomme).

FVIGURE
olum 1. eTraditional
7 3 | I s smacro-propagation
u e 6 | N o v e mof
b enset
e r - Dseedlings
e c e m b in
e ra backyard
2018 in southern Ethiopia. 343
Yemataw et al. | Traditional enset sucker propagation methods and opportunities for crop improvement

Propagation by seed ulate sucker production. When the apical meristem is left
Wild enset reproduces through seed, while cultivated intact, only one shoot will arise (Diro et al., 1996).
landraces are vegetatively multiplied, which over centuries Rhizomes from immature vegetative stage plants,
of cultivation might have altered seed morphology (Zerihun between 2 and 4 years old, are preferred for the production
Yemataw, 2017, pers. commun.). Enset seeds from cultivated of suckers (Bezuneh and Feleke, 1966; Negash, 2001;
types are characterized by low germination rates (Negash, Yemataw et al., 2014). The apical meristem is slightly raised/
2001). This is attributed partly to the structure of the seed grows upward within the pseudostem (3–12 cm for 2–3
coat, which is hard, impermeable and contains certain chem- year old plants) and therefore a portion of the pseudostem
ical inhibitors which affect germination efficiency, and partly (10–20 cm) must remain intact when harvesting a rhizome
due to the considerable number of so-called empty seeds, for sucker production. This is in order not to remove large
which are seeds containing no embryo or endosperm (Ne- sections of tissues situated around the apical meristem,
gash, 2001). In contrast, seeds from wild enset types often tissues that can give rise to numerous suckers (Diro et al.,
have a far higher germination rate, as has been observed in 1996). After uprooting the rhizome (sometimes it is left
Ethiopia and along the rift valley highlands in eastern Dem- in the ground), the apical meristem and ideally as little as
ocratic Republic of Congo (Zerihun Yemataw, 2016, pers. possible of the surrounding tissues is removed, making the
commun.; Guy Blomme, 2014, pers. commun.). Karlsson et center of the pseudostem hollow (Negash, 2001). The hollow
al. (2013) found that germination of mature enset seeds ob- area is refilled with manure, soil, humus and perhaps soft
tained from five wild and six cultivated enset plants ranged crushed stones (Simmonds, 1958; Bezuneh and Feleke, 1966;
from 5–55% depending on genotype. In this study, the av- Olmstead, 1974) or the rhizome is split into two or four equal
erage germination rate did not differ significantly between parts. The rhizome may be left in the sun or shade for 2–5
seed lots of wild and cultivated origin, which might be due days before planting to allow the wounds to heal (Tsegaye
to the used landraces and wild genotypes. Overall, time to and Struik, 2002; Bezuneh and Feleke, 1966). Rhizomes or
50% of final germination was 8.5 weeks and no germination rhizome pieces are planted (20–30 cm deep) in loosened
occurred after 28 weeks of incubation. Dalbato et al. (2014) soil, often mixed with manure (Diro and Tsegaye, 2012).
did not find a positive effect of chemicals, mechanical scar- Field trials demonstrate that the practice of filling the
ification, or warm water pre-treatments on germination. In hollow space with clod or manure has no significant effect on
contrast, Diro and Tsegaye (2012) reported an improved ger- sucker production; nor does the practice of leaving a rhizome
mination rate after scarification, which may be linked to the in the sun for a few days before re-planting (Diro et al., 1996;
assessed enset landraces. Makiso, 1996). Replicated field trials have also shown that
Reproduction of the enset plant from seed is seldom prac- suckers from half and quarter split-rhizomes emerged earlier
ticed by farmers (as plants are most often harvested before/ than those from the whole rhizome (Tabogie and Diro, 1992).
at flower emergence in order to obtain maximum amounts of On average 22, 76 and 102 suckers emerged from whole, half
starch). Harvesting at flower emergence prevents relocation and quarter rhizomes, respectively (Tabogie and Diro, 1992).
of nutrients from rhizome and stem to the growing inflores- These observations might indicate that apical dominance
cence. Reproduction via seed has only been reported from of the central meristem is not the only factor involved in
the highlands of Gardula, which might indicate an interest suppressing suckers/regulating sucker development on
from the farmers to compare vegetative propagated land- enset rhizomes. Early emergence is associated with more
races with seedlings obtained from seed (which represent vigorous suckers, which promotes success of establishment,
new genotypes). However, seedlings obtained from seed are higher and earlier yield. Moreover, failure of suckers to fully
less vigorous in terms of speed of growth than the suckers develop is higher in the case of whole rhizome (25%) and
obtained from vegetative propagations (Alemu and Sand- lowest for the half rhizome (8%) (Tabogie and Diro, 1992).
ford, 1991). Farmers in the Gurage and Wolaita areas are known to
The difficulty propagating enset from seeds has been a sort undersized/underdeveloped suckers (i.e., suckers that
limiting factor for enset breeding efforts (Morpurgo et al., are not growing vigorously enough) and replant them in
1996; Zerihun Yemataw, 2017, pers. commun.). The Ethiopi- the nursery for further growth, and eventual field planting
an national research organization (SARI) at its Areka research (Spring, 1996).
station, initiated enset breeding efforts in 1999 through the Variations in rhizome preparation are reported between
collection of landraces and wild genotypes as parent materi- different ethnic groups (Diro et al., 1996). The ethnic
als, with the aim to breed improved enset genotypes for vari- affiliation is a stronger determinant for decision-making
ous use values (e.g., kocho, bulla, fiber and amicho). than the wealth category to which the farming household
belongs (Tsegaye and Struik, 2002). Differences are seen in
Traditional vegetative propagation whether or not farmers uproot the rhizome, and whether or
Vegetative propagation is the primary method used by not it is split after uprooting (Diro et al., 1996). For example,
farmers to produce enset suckers. Enset has a large under- farmers of the Wolaita, the Gamo and the Gofa use a split
ground stem (i.e., rhizome, also called a corm), with nodes rhizome to produce suckers. Farmers from other areas prefer
and internodes. Suckers arise on the rhizome surface. The the whole rhizome. Where whole corms are used, farmers
growing buds are not located on the outer surface of the rhi- either destroy the apical meristem, but leave the rhizome
zome as ‘eyes’ but are found in concentric circles on the up- in situ (Jemjem, Gedeo and some areas of Gofa awrajas), or
per sections of the rhizome, and around the apical meristem, they first uproot the rhizome, destroy the apical meristem
and can be revealed though removal of leaf sheaths layer by and then replant the rhizome (Kambata-Hadiya and Chebo-
layer from the rhizome (Diro et al., 1996). In contrast to ba- Gurage areas). Farmers from the Sidama area and the Ari
nana, naturally occurring sucker production of field-grown either use split rhizome or whole rhizome preparation
enset plants is rare due to the dominance of the apical mer- methods to produce suckers (Diro et al., 1996; Yntiso, 1996;
istem, which must be destroyed/removed in order to stim- Tsegaye and Struik, 2002).

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Yemataw et al. | Traditional enset sucker propagation methods and opportunities for crop improvement

Karlsson et al. (2014) assessed the effect of rhizome ± 89 surviving plants are transplanted for a second time to
splitting, application of cow manure just next to buried one of two 200 m2 fields (at 1.5 × 1.5 m spacing). Two years
rhizomes and regular watering on sucker emergence and later (year 4), the ± 80 surviving plants are transplanted for
growth. The first suckers started emerging at 50 days after the third time to one of four 500 m2 fields (at 2.5 × 2.5 m
rhizome burial. Time to sucker emergence was longer for spacing). The plants are now left to mature for 4 years (year
entire than for split rhizomes, while a higher number of 8) until harvest, and every year, one of these four final fields,
suckers were obtained per rhizome when it was split. Less consisting of 80 plants, can be harvested. This system takes
than 60 suckers were recorded for entire rhizomes, while 8 years to become productive, after which harvesting can
between 60 and 140 suckers were most often recorded per be carried out annually. The number of plants in Makiso’s
rhizome for the rhizomes that were split in two or four pieces. (1976) example is based on the observation that 30–100
Watering decreased average time to sucker appearance and enset plants harvested annually supplies the enset needs for
resulted in more even sucker emergence and growth. For a 5–6 person household. The example also assumes a 10%
example, time to sucker emergence ranged from 60 to 65 loss of plants at each transplanting phase.
days for watered rhizomes of the ‘Zerita’ enset cultivar, while
it ranged from 60 to 85 days for non-watered rhizomes. This Enset cultivar diversity and sucker
is contradictory to local belief which often says that it is production
impossible to water newly buried rhizomes, as this practice Throughout the enset-growing regions, the maintenance
will cause rhizome rotting. In areas where extended droughts of enset cultivar diversity contributes significantly to food
are common and watering is impossible, it may be preferable and livelihood security, whereby wealthier households tend
to bury entire rhizomes, to, as much as possible, utilize the to have more land, grow more enset and maintain more
rhizome’s own water. Blomme et al. (2008) reported that diversified cultivars (Negash, 2001). Farmers base their
enset rhizomes contain 83% water, while rhizomes used for choice of cultivar on multiple criteria including ecological
propagation (sourced from 2 to 4 years old plants) weigh adaptation, tolerance to various diseases, rate of growth and
on average around 2–4 kg. The application of cow manure maturity, fibre quality and quantity, quality and quantity of
adjacent to rhizomes, to give the emerging suckers quick food yield, ease of work during decortication, size and taste
access to nutrients, resulted in a high sucker emergence of rhizome after cooking, food and fodder quality, medicinal
rate and subsequent vigorous growth. Karlsson et al. (2014) attributes (Makiso, 1996; Negash, 2001; Tsegaye and Struik,
concluded that vegetative propagation using buried disease- 2002; Yemataw et al., 2014).
free rhizomes in the field results in a large amount of healthy A loss of cultivar diversity has been reported due to
and vigorous suckers, ideal for re-planting, 9 months after disease encroachment (Negash, 2001). Propagation and
burial. transplanting activities may aggravate the dissemination and
Tsegaye and Struik (2002) reported that suckers appear maintenance of certain pests and diseases, such as bacterial
2–3 months after the rhizome or rhizome pieces have been wilt caused by Xanthomonas campestris pv. musacearum, the
buried and remain undisturbed for at least a year before the enset root mealy bug (Cataenococcus ensete), the root lesion
first transplant is carried out. Diro et al. (1996) observed nematode Pratylenchus goodeyi, the root-knot nematode
that new roots initiate from growing points on the suckers’ Meloidogyne sp., mosaic and chlorotic streak viruses (Quimio
developing rhizome after transplanting. From 2 to 6 new and Tessera, 1996; Morpurgo et al., 1996; Negash, 2001).
roots of 1.5 to 19 cm length can be expected 15 days after
transplanting (DAT) and about 50 cm at 35 DAT. Leaves Effects of gender in enset sucker
desiccate following transplanting, except for the central production
cigar leaf, which continues to grow up to 15 DAT. For ease The production of suckers consists of a series of
of handling, farmers may remove roots and leaves upon consecutive tasks, whereby some are more gender-specific
transplanting since their absence does not affect the growth than others. As a backyard crop, enset is primarily considered
of the transplants (Diro et al., 1996). a women’s crop (Negash, 2001; Pankhurst, 1996). However,
The number of suckers produced per rhizome ranges decision-making regarding cultivar diversity is shared
between 40 and 200, depending on soil conditions, cultivar between husband and wife, as is the time of planting
type, size and age of mother plant rhizome, amount of (Negash, 2001). Women are more involved in carrying
rainfall, land preparation and time of planting (Shambulo suckers for transplanting and manuring the transplants,
et al., 2012). An average enset farming household cultivates and also contribute with plant harvesting, leaf sheath and
between 200 and 400 plants and consumes about 10 to 20 rhizome processing, kocho storage and marketing. Whereas
plants annually. So, 1 to 4 parent rhizomes can fulfil the men are more involved in propagation, transplanting,
annual requirement for enset planting materials (Bezuneh digging the pits (for fermentation of the harvested enset),
and Feleke, 1966; Negash, 2001). and weeding. There is some flexibility in gender-division,
Overall, traditional propagation of enset suckers is with males helping during harvest and females responsible
resource intensive, in terms of space and labour requirement. for transporting manure from the homestead to the field and
For example, Makiso (1976, cited in Hiebsch, 1996) noted for weeding (Olmstead, 1974).
that in a typical entire 8-year enset production system, with
4 transplanting steps, with each step taking respectively 1, 1, Sucker movement and markets
2 and 4 years, an overall surface area of 2,455 m2 is needed There are very few reports describing sucker markets
to harvest 80 nearly mature enset plants from a space of and movement of enset suckers. In Kaffa Shaka, enset suckers
500 m2 annually. This production system starts with a 5 m2 of various cultivars are traditionally exchanged together
nursery with 5 rhizomes (at 1 × 1 m spacing), planted at with indigenous knowledge about the crop/cultivars as a
year 0. After one year (year 1), the ± 100 suckers produced means of increasing cultivar diversity on farm. Exchanges
in this nursery are transplanted for the first time to a 50 m2 are made between neighbours, relatives or kin and within
field (at 1.0 × 0.5 m spacing). After another year (year 2), the communities, between neighbouring villages and sometimes

Volume 73 | Issue 6 | November-December 2018 345


Yemataw et al. | Traditional enset sucker propagation methods and opportunities for crop improvement

beyond (Negash, 2001). Rahmato (1996) reports that poor Conclusion


farmers may occasionally sell their young plants to others Traditional macro-propagation methods, using entire
who need them for propagation purposes. Woldetensaye rhizomes or rhizome pieces, currently suffice to provide
(1997) reported that farmers from Dale Yirgalem district the needed enset suckers at farm, village or landscape level.
(located higher than 2,100 m in the Sidama zone of the However, when larger quantities of suckers are needed, e.g.,
Southern Nations, Nationalities, and Peoples’ Region of when introducing a new enset cultivar or coping with severe
Ethiopia) produce suckers as a main source of cash income disease or pest impacts, improved/novel micro- and macro-
and sucker markets are widely practiced. However, selling propagation techniques, as listed in this review paper, could
and buying of suckers in Sodo and Butajera districts hardly offer solutions. Current research in Ethiopia on fine-tuning
occurs, as planting material is mainly distributed as gifts. In in vitro multiplication technologies and ongoing breeding
addition, households in these districts are most often self- efforts for improved enset genotypes for various use values
sufficient for sucker production. will contribute to a high-quality clean planting material
supply system.
Improved/novel rapid propagation
technologies Acknowledgments
Several novel methods for rapid propagation of enset We would like to thank the Directorate General for
are found in the literature. Alternative pathways for the Development, Belgium, for funding this research through the
rapid propagation of enset are discussed extensively by Diro Consortium for Improving Agriculture-based Livelihoods in
and Van Staden (2004), including zygotic embryo culture Central Africa (CIALCA). We would also like to acknowledge
(Bezuneh, 1980; Negash et al., 2000, 2001; Diro and Van the financial contribution of the CGIAR Research Program
Staden, 2003), shoot tip culture (George and Sherrington, on Roots, Tubers and Bananas and the CGIAR Fund Donors.
1984; Nehra and Kartha, 1994; Afza et al., 1996; Morpurgo A special appreciation is due to the Southern Agricultural
et al., 1996; Zeweldu, 1997; Negash et al., 2000), and callus Research Institute (SARI), the Ethiopian Institute of
culture and somatic embryogenesis (George and Sherrington, Agricultural Research (EIAR) and the Areka Agricultural
1984; Zeweldu, 1997; Afza et al., 1996; Morpurgo et al., 1996; Research Centers for providing all necessary facilities/
Mathew et al., 2000; Mathew and Philip, 2003). support to carry out this review.
Macro-propagation and micro-propagation (syn. tissue
culture or in vitro) are useful technologies to provide large References
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Received: Jan. 30, 2018


Accepted: Oct. 19, 2018

348 International Journal of Tropical and Subtropical Horticulture

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