M a s h u a
Promoting the conservation and use of underutilized and neglected crops. 25.
Alfredo Grau
Ramiro Ortega
Dueñas
Carlos Nieto
Cabrera
Michael Hermann
Tropaeolum tuberosum Ruíz & Pav.
M a s h u a
Clayton G.
Campbell
Alfredo Grau
University Tucumán,
Argentina
Ramiro Ortega Dueñas
University Cusco,
Peru
Carlos Nieto Cabrera
Instituto Nacional de Investigaciones Agropecuarias,
Quito, Ecuador
Michael Hermann
International Potato Center,
Lima, Peru
Volume Editor: Jan M. M. Engels
Promoting the conservation and use of underutilized and neglected crops. 25.
Tropaeolum tuberosum Ruíz & Pav.
The International Plant Genetic Resources Institute (IPGRI) is an independent inter-
national scientific organization that seeks to advance the conservation and use of plant
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Citation:
Grau Alfredo, Ramiro Ortega Dueñas, Carlos Nieto Cabrera and Michael Hermann. 2003.
Mashua (Tropaeolum tuberosum Ruíz & Pav.). Promoting the conservation and use of
underutilized and neglected crops. 25. International Potato Center, Lima,
Peru/International Plant Genetic Resources Institute, Rome, Italy.
ISBN 92-9043-581-X
IPGRI
Via dei Tre Denari 472/a
00057 Maccarese 
Rome, Italy
© International Plant Genetic Resources Institute, 2003
Acknowledgements
We thank the Swiss Development Corporation and the Japanese government for
their long-standing commitment to research and development of Andean roots and
tubers and their financial support, which made this publication possible. Dr Carlos
Arbizu contributed to the manuscript by sharing his field experience and pointing
out critical aspects of the crop. Mr Coen Bussink of CIP kindly prepared Figure 11.
We thank Dr Thomas Bernet, Dr Jan Engels, Dr Miguel Holle and an anonymous
reviewer for their critical and helpful comments on the final manuscript.

Contents
Foreword vii
1 Introduction 1
2 Crop history and dispersal 3
3 Production and distribution 5
4 Vernacular names 7
5 Crop biology 10
5.1 Crop cycle and development 10
5.2 Plant architecture and morphology 10
5.3 Reproductive biology 13
6 Agronomy 14
6.1 Plant propagation 14
6.2 Cropping systems and crop husbandry 14
6.3 Fertilization 16
6.4 Pests and diseases 16
6.5 Harvest and yields 18
6.6 Post-harvest 20
6.7 Crop ecology 21
7 Composition and uses 23
7.1 Chemical composition 23
7.2 Food uses 23
7.3 Feed uses 25
7.4 Flour and starch 25
7.5 Medicinal uses 26
8 Taxonomy and biosystematics 28
8.1 Taxonomic history and synonyms 28
8.2 Wild species closely related to cultigen 29
9 Genetic resources 32
9.1 Chromosomal variation 32
9.2 Molecular variation 33
9.3 Genetic erosion, germplasm collection and genebank holdings 33
9.4 Maintenance of the collections and conservation strategies 36
10 Crop limitations 38
11 Breeding 39
12 Prospects and research needs 40
12.1 Advantageous features of mashua 40
12.2 Development objectives and research needs 40
References 43
Research contacts 50
Institutions maintaining germplasm collections 54
Foreword
Humanity relies on a diverse range of cultivated species; at least 6000 such species are
used for a variety of purposes. It is often stated that only a few staple crops produce
the majority of the food supply. This might be correct but the important contribution
of many minor species should not be underestimated. Agricultural research has
traditionally focused on these staples, while relatively little attention has been given
to minor (or underutilized or neglected) crops, particularly by scientists in developed
countries. Such crops have, therefore, generally failed to attract significant research
funding. Unlike most staples, many of these neglected species are adapted to various
marginal growing conditions such as those of the Andean and Himalayan highlands,
arid areas, salt-affected soils, etc. Furthermore, many crops considered neglected at a
global level are staples at a national or regional level (e.g. tef, fonio, Andean roots and
tubers, etc.), contribute considerably to food supply in certain periods (e.g. indigenous
fruit trees) or are important for a nutritionally well-balanced diet (e.g. indigenous
vegetables). The limited information available on many important and frequently basic
aspects of neglected and underutilized crops hinders their development and their
sustainable conservation. One major factor hampering this development is that the
information available on germplasm is scattered and not readily accessible, i.e. only
found in ‘grey literature’ or written in little-known languages. Moreover, existing
knowledge on the genetic potential of neglected crops is limited. This has resulted,
frequently, in uncoordinated research efforts for most neglected crops, as well as in
inefficient approaches to the conservation of these genetic resources.
This series of monographs intends to draw attention to a number of species which
have been neglected in a varying degree by researchers or have been underutilized
economically. It is hoped that the information compiled will contribute to: (1)
identifying constraints in and possible solutions to the use of the crops, (2) identifying
possible untapped genetic diversity for breeding and crop improvement
programmes and (3) detecting existing gaps in available conservation and use
approaches. This series intends to contribute to improvement of the potential value
of these crops through increased use of the available genetic diversity. In addition,
it is hoped that the monographs in the series will form a valuable reference source
for all those scientists involved in conservation, research, improvement and
promotion of these crops.
This series was born out of a joint project between the International Plant Genetic
Resources Institute (IPGRI) and the Institute of Plant Genetics and Crop Plant
Research (IPK) with financial support provided by the Federal Ministry of Economic
Cooperation and Development (BMZ) of Germany.
Series editor:
Dr Jan Engels, International Plant Genetic Resources Institute (IPGRI)

1 Introduction
Mashua (Tropaeolum tuberosum R. & P.), along with several potato species (Solanum
spp.), ulluco (Ullucus tuberosus Caldas) and oca (Oxalis tuberosa Molina), pertains to
the group of edible tuber crops indigenous to the cool-temperate Andes. Mashua is
economically much less important than the other Andean tuber crops, but, as this
monograph is intended to show, the crop’s potential has been largely overlooked
and opportunities for wider use within its native range and beyond have remained
unexplored (Fano and Benavides 1992; Hermann and Heller 1997). Mashua
contributes to tuber diversity, resulting in greater production stability in the
heterogeneous Andean environments. So far this has resulted in the conservation of
a wide range of native cultivars. However, the future role of mashua as a crop in the
region seems to be uncertain. Apparently a range of positive attributes such as
rusticity, pest tolerance and high productivity under low levels of inputs cannot
counterbalance the present lack of ‘market interest’.
Mashua possesses a range of phytochemicals, which protect not only the crop
itself, but possibly also associated crops, against pests. Also, except in soils of poor
drainage, mashua has very few diseases. Mashua’s aggressive growth makes it a
very good ground cover, adequate for soil protection on the steep slopes of the Andes.
Low input requirements, pest and disease tolerance, high productivity and
multipurpose uses, are features that make mashua a very attractive plant for organic
agriculture.
Mashua’s particular and strong taste has often been mentioned as a main cause
for its neglect and abandonment. However, similarly pungent phytochemicals occur
in several horticultural crops, such as in the Brassicaceae family (radishes, mustards)
and Allium species (onions, garlic), which are well liked in many cultures and have
a variety of uses in cooking. Rather than treating mashua as a starchy staple, as in
Andean food systems, it might have potential as a spicy resource in raw dishes,
pickles or fermented specialties. For example, thinly sliced raw mashua adds a
pleasant pungency to salads, like Brassica species, which also contain glucosinolates.
Mashua’s reputed antiaphrodisiac action, however, imposes constraints on the
marketing of the crop. Its demonstrated hormonal effect in rats indicates that there
may be a physiological basis for the popular belief that it reduces libido. The possible
hormonal activity in humans, and the effect of different processing methods on that
activity, will have to be investigated in detail.
If a wider market could be identified and marketing resources made available,
mashua would be an excellent candidate for crop improvement. This is because of
the considerable diversity available in mashua germplasm collections, the ease of
hybridization of domesticated with (conspecific) wild mashua and with other
Tropaeolum species, as well as mashua’s prolific seed set, thus making conventional
breeding possible.
Visually distinguishing the tubers of mashua from those of oca takes a little
practice, especially in the case of the yellow forms, which can look confusingly similar
to the untrained eye (Fig. 1). Melchiorre (1985) provides a list of morphological
2 Mashua (Tropaeolum tuberosum Ruíz & Pav.)
features that identify the two species, but a bite into the raw tuber provides an
immediate answer: oca has a tart, acidulous taste owing to its high content in organic
acids, notably oxalate, whereas the taste of mashua is reminiscent of the pungency
of a radish.
Fig. 1. Andean tuber crops: oca (Oxalis tuberosa Molina), ulluco (Ullucus tuberosus Caldas)
and mashua (Tropaeolum tuberosum R. & P.) (Photo M. Hermann).
2 Crop history and dispersal
Wild and cultivated T. tuberosum is found in the high Andes from Colombia to
northwest Argentina. Cárdenas (1989) and Hawkes (1989) assume that
domestication occurred in the region encompassing Ecuador to Bolivia. This is
supported by the large diversity present in this wide area. However, in the absence
of a comprehensive study on the diversity of wild and cultivated forms of mashua,
it is difficult to pinpoint a smaller area as the likely centre of origin of the crop.
Similarly, little is known about mashua crop history and dispersal. Mashua is
more widespread than the minor Andean tubers maca (Lepidium meyenii Walpers)
or mauka (Mirabilis expansa (Ruíz & Pav.) Standl.), species known only from isolated
and ‘insular’ distributions, but is less common than the cool-temperate oca and
ulluco. Excavations at Guitarrero cave in Peru suggest (Pearsall 1992; Smith 1980),
that oca, and probably also ulluco, may have been consumed as early as
8000–7500 years before the present. The first archaeological evidence of mashua dates
from 650–1350 AD (Pearsall 1992) in the Huachumachay cave sediments, located in
the Jauja valley, Peru. The reduced economic scope of mashua in the colonial past
and today, as well as its relatively late appearance in the archaeological record,
suggests that its domestication may have taken place relatively late.
On the basis of current medicinal uses and beliefs surrounding mashua, as well
as the plant’s use as (a rare) ornamental in northwest Argentina (Fernández 1973,
Hermann 1992, personal observation), Johns et al. (1982) suggest that mashua had a
significant place in the distant past for reasons other than food, and that its
domestication may relate to its importance as medicine (see Section 7.5). In the same
vein, Fernández (1973) argues for it to be a relict of a primordial agricultural complex.
Beautiful, unambiguous depictions of mashua are present in the Nazca pottery
at Pacheco, dated 1000 AD (Yacovleff and Herrera 1935; Towle 1961) (Fig. 2),
Fig. 2. Nasca pottery designs
representing Andean tubers.
Clockwise from upper left:
potato, ulluco, oca and
mashua. (Reproduced from
Yacovleff and Herrera, 1935,
p. 308, with permission from
Museo Nacional de la Cultura
Peruana, Lima, Peru.)
4 Mashua (Tropaeolum tuberosum Ruíz & Pav.)
suggesting that, in spite of being a highland crop, it was also well known in coastal
Peru. The first historical reference to the plant appears in a chronicle cited by Yacovleff
and Herrera (1935) as: ‘Various, Bello Gayoso, on Cuenca and its Province, 1582’.
Under its alternative common names ‘añus’ and ‘masuas’ (plural), mashua is later
described by Garcilazo de la Vega (1966 [first published in 1609]) and Guamán Poma
de Ayala (1936 [first published 1615]). The Jesuit Bernabe Cobo (1956 [1639]), gives
a detailed account of mashua, emphasizing its similarity with oca in size and shape.
This author also mentions the antiaphrodisiac effects attributed to mashua, and how
these were exploited by the Inca military to keep their soldiers quiet. Much later,
Ruíz and Pavón (1802) made the first and still valid botanical description,
accompanied by a detailed drawing, which is reproduced in Fig. 3.
Although less common in the past and present than oca and ulluco, mashua was
important enough to deserve significant selection efforts by the Andean peasants.
This resulted in a large number of cultivars. In the last decades the demand for
mashua seems to have decreased, to judge from its diminished presence in city
markets. However, it is still available in small amounts in rural markets and from
farmers who use it largely for home consumption.
Fig. 3. Mashua plant (upper right )
and reproductive parts (lower right)
depicted in Ruíz and Pavón’s ‘Flora
Peruviana et Chilensis’ (1802).
3 Production and distribution
According to agricultural statistical data provided for Peru by Deza (1977), total
mashua production in that country reached 19 500 t in the mid 1970s, with a total
area of 5290 ha and an average yield of 3.7 t/ha. More recent data (Anonymous 1999)
suggest an increase in yields and cropped area to 32859 t and 7244 ha, respectively,
with an average yield of 5.2 t/ha. However, these figures must be treated with
caution, as the small and fragmented plots in remote mountain areas are difficult to
estimate, and agricultural census data in the Andean countries are notoriously
unreliable. Peru is generally believed to be the largest Andean producer.
One of the few rural women seen in 2001 offering a small amount of mashua in
the market of Huánuco, a mid-sized town in the Peruvian highlands, is pictured in
Fig. 4. This photograph illustrates and typifies the limited scale of mashua production
and commercialization in Peru. The woman also displays yacon (Smallanthus
sonchifolius (Poepp.) H. Rob.), another neglected Andean root crop, as well as roasted
calabaza (Cucurbita ficifolia Bouché). Her attire, and the choice of these ‘orphan
products’—rarely present on urban markets—clearly identifies her as a specialist
supplier of indigenous crops.
Fig. 4. Mashua, yacon roots and
roasted calabaza fruits (Cucurbita
ficifolia Bouché) in a market display in
Huánuco town, Peru (Photo M.
Hermann, September 2001).
6 Mashua (Tropaeolum tuberosum Ruíz & Pav.)
Data for the other Andean countries are extremely scarce. Tapia (1994) estimates
the cultivated mashua area in Bolivia at no more than 100 ha, with yields of 2–3 t/ha.
However, both figures seem too low since production characteristics similar to those
in Peru can be expected in Bolivia. There are no statistics for Ecuador, where we
estimate the cultivated area to be around 50 ha, all located in isolated areas of the
Sierra region. Nieto (1993) provides an overview of the use of mashua in Ecuador
and the status of germplasm conservation. The area dedicated to mashua in Colombia
is concentrated in Boyacá.
In Argentina, mashua is restricted to the northwestern provinces, where one of
us has seen cultivated plants in Colanzulí, Salta, and Chalguamayoc, Jujuy (Hermann
1992, personal observation). However, mashua, or ‘sisaño’ as it is known locally, is
hardly more than a botanical curiosity, not seen on local markets and known to only
a few farmers, who occasionally grow it for food or as an ornamental.
Latcham (1936) lists mashua in his book on pre-Colombian plants of Chile, but
neither the crop nor the product was found on a collecting mission in the high-
altitude oases of the Atacama desert in 1993, an area once belonging to Peru, where
other Andean tubers such as native potatoes, oca and ulluco are still fairly common
(Hermann 1993, personal observation).
4 Vernacular names
Mashua (or maswa), a Quechua word for which no meaning is known to us, is the
most widely used common name for the plant and the tuber, especially in southern
Colombia, Ecuador and Central Peru, and is often understood even where other
names prevail. Masua (or maswa) is the form used in Quechua from the Ayacucho
region. A set of other terms, such as maxua, majua, mazuco, maswallo, mascho, are
probably post-Columbian variations under the influence of the Spanish language.
Quechua speakers also refer to mashua as ‘añu’, a word derived from the Aymara
term isañu or isaño. Añu is used in the Cusco region, while isaño is common around
Lake Titicaca and further south in Bolivia, including the Cochabamba region. In
Colombia, mashua is also referred to as ‘cubio’, a term whose origin is unknown to
us but could be from a pre-Columbian language, thus perhaps indicating the
antiquity of mashua cultivation in this country. ‘Navo’, ‘navios’ or ‘nabos’ are also
terms for mashua restricted to Colombia that appear to be modifications of ‘nabo’,
the common Spanish name for Brassica napus. A list including less common names
is presented in Table 1. The presence of at least three completely different name
groups—mashua, añu and cubio—suggests that the crop was widely established
before the homogenizing effect of the Inca and Spanish conquests. This phenomenon
supports the notion that either the crop was domesticated and dispersed very early,
or it was domesticated independently in different areas in the Andes.
Mashua tubers show a notably wide range of colours and qualifiers in varietal
names often refer to them. A list of the more frequent names is presented in Table 2.
Some varietal names apparently refer to attributes such as shape or taste
(Hermann and Cruz 1991): ‘Huaka hasta’ or ‘Huagra hasta’ (cow horns) refers to
elongated, curved mashuas. ‘K’eya-añu’ indicates foetid smell. ‘Take-añu’ or ‘Taqui
añu’ refers to tuber stores common in Cusco, and might indicate that this type
holds well in storage. ‘Kita-añu’ and ‘Añu-añu’ are terms used for wild mashua
(Herrera 1921).
8 Mashua (Tropaeolum tuberosum Ruíz & Pav.)
Table 1.  Vernacular names of mashua
Name Language Region/country Reference
Allausu Peru Mejía 1931
Añu Quechua Peru Cárdenas 1989
Apilla Quechua Bolivia Cárdenas 1989
Apiñamama Quechua Peru Herrera 1941
Capuchinha tuberosa Portuguese Portugal Sánchez-Monge 1981
Capucine tubéreuse French France Sánchez-Monge 1981
Cubios Spanish (?) Colombia Pérez Arbelaez 1947
Gallu Gallu Sparre and 
Andersson 1991
Isaño Aymara Bolivia/Titicaca Cárdenas 1989
Majua Spanish Ecuador Lescano 1994, 
Espinosa et al. 1997, 
Patiño 1964
Mashua Quechua Ecuador Tapia et al. 1996, 
Espinosa et al. 1997, 
Estrella 1986
Mashua Peru Herrera 1941
Maxua Spanish Ecuador Lescano 1994, Patiño 
1964
Mishua Peru Mejía 1931
Navios Spanish Colombia Pérez Arbelaez 1947
Navos Spanish Colombia Pérez Arbelaez 1947
Ocaquisaño Quechua Bolivia Cárdenas 1989
Pane Guambiano Colombia Patiño 1964
Peruanische German Germany Sánchez-Monge 1981
Knollenkresse
Puel Paéz Southern National Research 
Colombia Council 1989, Patiño 1964
Sisaño Aymara (?) Argentina, Jujuy Hermann 1992, 
personal observation
Tropeolo del Peru Italian Italy Sánchez-Monge 1981
Tuber nasturtium English United Kingdom Sánchez-Monge 1981
Promoting the conservation and use of underutilized and neglected crops. 25. 9
Table 2.  Mashua varietal names
Name Tuber attributes Source
Occe-añu Leaden Herrera 1941
Yana-añu Black Herrera 1941
Checche-añu Grey Herrera 1941
Ckello-añu, K’ello-añu Yellow Herrera 1941
Muru-añu Purple Herrera 1941
Phutilla añu Red Hermann and Cruz 1991
Puca-añu Red Herrera 1941
Yana-añu Black Hermann and Cruz 1991
Yurac-añu White Herrera 1941
Zanahoria-añu like a carrot Hermann and Cruz 1991
Zapallo-añu like a pumpkin Herrera 1941
Quillu-mashua Yellow Espinosa et al. 1997
Putsu-mashua Yellow background  Espinosa et al. 1997
covered in red stripes
Sucsu-mashua Yellow background Espinosa et al. 1997
covered in pink stripes
Mashua yana-saco Black Espinosa et al. 1997
Mashua-chaucha Early-maturing Tapia et al. 1996
Mashua-shira Yellow with purple dots Tapia et al. 1996
Mashua-zapallo Yellow and red Tapia et al. 1996
Mashua-zapallo Yellow Tapia et al. 1996
Sangre de Cristo Yellow background Tapia et al. 1996, 
(‘Christ’s blood’) covered in red stripes Espinosa et al. 1997
Yawar waqac (‘weeping  Yellow background Hermann and Cruz 1991
tears of blood’) covered in red stripes
Huaka hasta or Huagra hasta Long curved Hermann and Cruz 1991
K’ella añu Foetid smell Hermann and Cruz 1991
Take-añu or taqui-añu Good for storage Hermann and Cruz 1991
Kita-añu Wild mashua Herrera 1921
Añu-añu Wild mashua Herrera 1921
5 Crop biology
5.1 Crop cycle and development
Mashua has often been mistakenly described as an annual, because of its yearly
production cycle (e.g. Arbizu and Tapia 1992; Zela et al. 1997). However, the term
‘annual’ is reserved for plants in which only sexually formed seeds give rise to a new
generation, whereas both cultivated and wild mashua persist through tubers,
although aerial organs may die back at the end of the growing season. Mashua is
therefore clearly a perennial species, with some genotypes presumably being very
old clones. Small tubers are frequently left in the field and sprout easily, especially
under the wetter conditions in the northern Andes. Under such circumstances,
mashua can become a weed problem, thus fully confirming the species’ perennial
nature.
Unlike ulluco, mashua flowers profusely, starting 3–5 months after planting and
lasting for 1–2 months. The plants also set abundant seed. Tubers are ready for harvest
6–9 months after planting. A schematic representation of mashua’s development is
shown in Fig. 5.
5.2 Plant architecture and morphology
Mashua is a stout climber with thick or slender, often reddish, glabrous stems. Stems
arising from sprouting tubers are erect but the plant soon adopts a semiprostrate or
prostrate habit, producing a very dense soil cover. Stems and petioles are twining
and able to climb on any available support, reaching 2 m or more in height. The
tubers are usually 5–15 cm long and 3–6 cm broad in their distal part. Most tuber
Fig. 5. Mashua development: 1, Emergence; 2, Stolon formation; 3, Tuberization; 4–5, Flowering;
6, Fruiting; 7, Tuber maturity (Source: Lescano 1994, p. 74).
Promoting the conservation and use of underutilized and neglected crops. 25. 11
growth is due to pith proliferation. The tuber is covered by an epidermis with thick
exterior walls, which gives a waxy aspect to the surface. Tuber colour and shape vary
considerably, as seen in the cultivar samples from Peru and Colombia in Fig. 6 and
Fig. 7 respectively. Some cultivars are of spectacular beauty, such as the Peruvian
Yawar waqac (Fig. 8), which has a skin pattern reminding the name-giving Quechua
Indians of ‘tears of blood’ (yawar = blood, waqac = to cry).
Petioles are reddish, 4–20 cm long, with thin, reddish, inconspicuous and
caducous stipules. Leaf blades are peltate, suborbicular, 4–6 cm long and 5–7 cm
Fig. 6. Sample of
mashua variation in
Peru (Photo G. Chang).
Fig. 7. Mashua
cultivars from the
market in Tunja,
Boyacá, Colombia,
locally called ‘cubios’
or ‘nabos’ (Photo M.
Hermann, November
1986; coin is 2 cm in
diameter).
12 Mashua (Tropaeolum tuberosum Ruíz & Pav.)
broad, at base rounded to subtuncate, with normally 5 lobes (sometimes 3–4),
rotundate, with an obtuse to truncate apex, mucronate, upwards and slightly
sideward directed. The upper surface is dark green while the lower surface is pale
green with marked purple palmate nervation. The hypogynous flowers are solitary,
zygomorphic, placed on long (15–25 cm) peduncles (Fig. 9). Calyx with 5 sepals, 5-
lobed, mostly red or reddish, but sometimes yellow; inferior lobes lanceolate,
12–14 mm long, 4–5 mm broad at the base; sepals fused at the base forming a nectar
containing spur, often referred to as calcar, of the same colour, tubular with a very
broad, straight or curved appendix, 18–22 mm long, 5–6 mm in diameter at the
base. Petals, 5, free, commonly yellow or orange with darker veins, sometimes light
lilac or reddish; posterior petals are unguiculate with a rotundate blade, 6–9 mm
long, 5–8 mm broad; anterior petals are elliptical, unguiculate, 10–15 mm long and
4–6 mm broad. Stamens 8; the syncarpic ovary is superior from 3 carpels, 3 locules
and a simple style with 3 lobbed stigmata; each locule contains one ovule in axilar
position.
The described floral morphology is typical for Tropaeolum. A number of
aberrations have been described by Pocco (1976) and Vallenas (1977) in clones of the
Fig. 9. Flowering mashua plants, near
Quito, altitude 3050 m (Photo M. Hermann,
May 1990).
Fig. 8. Mashua cultivar ‘Yawar waqac’ from
Cuzco, Peru. The Quechua name of this
cultivar means ‘tears of blood’ (Photo M.
Hermann, 1991).
Promoting the conservation and use of underutilized and neglected crops. 25. 13
Puno area. They include an increased number of sepals and petals to 6 or 7 per flower,
flowers with 2 spurs, increased number of stamens (up to 13), increased number of
carpels to 4 or 5 and a corresponding number of stigmata.
At maturity the fruit, a schizocarp, separates into 3 mericarps, each one with 3
pronounced ribs and a rugged surface and containing 1 seed. The pericarp is thin
but epicarp, mesocarp and endocarp can be easily distinguished. Some morphotypes
from Ecuador present 2–5 mericarps per fruit at maturity. This may be an
environmentally modified trait. The bulk of the seed is formed by the cotyledons
rather than endosperm, with cells containing aleuron grains and thick cell walls
formed by galactoxyloglucans that are mobilized following seed germination
(Bewley and Black 1994).
The mashua tuber is morphologically a thickened stem—it has ‘eyes’ from which
it produces aerial stems and adventitious roots, which are slender and filiform.
Lateral roots are less developed than the main ones (Chacón 1960).
5.3 Reproductive biology
Unlike oca and ulluco, mashua flowers profusely and sets seed easily. Unfortunately,
little is known about mashua’s reproductive biology. To date the most detailed
research on the subject was carried out by Pocco (1976) and Vallenas (1977).
According to Vallenas (1977) mashua begins to flower 3–4 months after plant
emergence. Flowering lasts for 1–1.5 months. Each flower remains open for
8–15 days. Stigmata are receptive at the beginning of anthesis, and remain receptive
for 40 hours. Pollen is viable 24 hours before anthesis and pollen viability reaches a
peak 24 hours after anthesis. The abundant flowering attracts a considerable number
of insects and birds. Pollen viability at anthesis was estimated at 50–53% by Pocco,
but Johns and Towers (1981) found much higher values (95% viability) in T. tuberosum
ssp. tuberosum and T. tuberosum ssp. silvestre. Information on pollination biology of
Tropeolaceae is still limited. The best-known species so far is T. pentaphyllum Lam.
(Fabbri and Valla 1998), a species of Southern Brazil, Uruguay and Northeast
Argentina, which is pollinated by hummingbirds and several Hymenoptera.
In our experience mashua seeds can be desiccated to low moisture levels at the
ambient conditions of the Andean highlands and will germinate after several months
of storage, but formal experiments confirming the orthodox behaviour and
germinability of the seeds have still to be conducted. The fact that mashua fields
consisting of only one clonal cultivar form ample seeds suggests that mashua is self
fertile. Plants grown from botanical seeds, apart from their distinct tuber
morphology, resemble mother plants in terms of crop duration, phenology and
capacity to reproduce asexually via the tubers. To our awareness, botanical seeds are
currently not used to conserve germplasm.
6 Agronomy
6.1 Plant propagation
Mashua is propagated for production purposes in traditional Andean agriculture
exclusively via the tubers, which need to be set aside from the harvest for that
purpose. Rooted young tubers or aerial stems can also be used for accelerated bulking
of particular clones, although this practice is limited to research purposes.
As in the case of other Andean tubers (potatoes, oca, ulluco) botanical seeds of
mashua are not used for propagation purposes in current agricultural practice. The
reasons are probably the same as for the other tubers: failure to breed true to type
by sexual propagation, extended crop duration of seed-propagated plants and
predominance of vegetative agriculture in traditional Andean systems. The
abundant seed set of mashua and its apparent good germinability, however, afford
plant breeders and researchers with opportunities to produce and select new
genotypes with relative ease.
6.2 Cropping systems and crop husbandry
Mashua is part of the traditional, subsistence-oriented cropping systems of the
Andean highlands. Like most of the agricultural production systems in the region
these are polycropping arrangements, very intensive in land use, and labour intensive
(Camino 1997). Mashua is interplanted with other tubers such as potato, oca or ulluco
or with other crops such as faba bean (Vicia faba L.), quinoa (Chenopodium quinoa Willd.)
or Andean lupin (Lupinus L.) (Fig. 10). In other cases, mashua together with oca and
ulluco is part of crop rotation systems; they are planted in small monocrop plots before
or after barley, broad bean or lupine. At lower altitudes, as in the inter-Andean valleys
of Ecuador (altitude less than 3000 m), mashua is often intercropped with maize (Zea
mays L.) and squashes (Cucurbita spp.). In any case, mashua is seldom monocropped
Fig. 10. Mashua
intercropped in the
foreground with oca and
ulluco, next to faba beans
and barley, Cotopaxi
Province, Ecuador, 3700 m
altitude (Photo M. Hermann,
July 1988).
Promoting the conservation and use of underutilized and neglected crops. 25. 15
in plots exceeding 2000 m2. In Colombia, mashua is often used in border rows
surrounding potato fields. This practice is believed to repel potato pests. Johns et al.
(1982) refer to a similar practice in Peru. This is supported by Rea’s (1984) mention
of ornamental Tropaeolum serving as pest barriers in Ecuador.
In Ecuador and in large parts of the Colombian Andes mashua has been reduced
to the status of a botanical curiosity or that of a medicinal plant, often tolerated in
backyard gardens rather than cultivated. Some farmers consider mashua a potential
weed problem in successive crops, as it can persist in the field for two or three crop
seasons. Mashua regenerates easily from tubers left in the field.
Because of agroecological and ontogenetic similarities and its common
intercropping with oca and ulluco, mashua is subjected to similar cultivation
practices, to the point that they are often dealt with together in the literature (e.g.
Arbizu and Tapia 1992).
Small mashua tubers are preferred for planting (or are the only material left over
at planting time), which may in fact be a poor agricultural practice. By using small
tubers for seed, farmers may unconsciously select for less vigorous types, which may
either represent somatic mutations or material affected by pathogens.
The tubers are planted on ridges 70–80 cm apart. Distances within ridges vary
from 30 to 40 cm (Meza et al. 1997), giving total plant populations of 31 000–48 000
plants/ha. Barrionuevo (1975) indicates a wider distance, up to 100 cm between rows
and up to 70 cm within rows. Under favourable growing conditions mashua can
respond efficiently to the wider spacing. In Peru planting is done with the
‘chaquitaclla’, which is used to open a hole, at the bottom of which one or two tubers
are placed. Planting is done from mid-August to mid-October, coinciding with the
onset of the first rains.
When mashua is planted as a monocrop (Fig. 11), planting distance varies from
80 to 100 cm between rows and from 30 to 50 cm within rows, depending on soil
Fig. 11. Mashua (right)
and oca plot (left),
Cotopaxi Province,
Ecuador, 3700 m altitude
(Photo M. Hermann, July
1988).
16 Mashua (Tropaeolum tuberosum Ruíz & Pav.)
quality. The wider distances are applied in soils with high fertility. Weed control is
necessary only during the early stages. Once in the ground, tubers sprout quickly
and mashua develops a very dense canopy that effectively starves competing plants
of light. Quispe et al. (1997) found 160 days after planting a leaf area index (LAI) of
3.6. This was less than the LAI in potato (4.5) but more than the LAI in oca (2.5)
grown under the same conditions. Similar results were obtained by Valladolid et al.
(1982). Fertilized mashua can reach much higher LAI values of 4.7–5 (Valdivia et al.
1999).
Plants are hilled twice. Hilling is believed to increase yields by stimulating stolon
and tuber formation, but no formal experiments have been conducted to confirm
this. The first hilling is normally done after the plants have emerged and the soil has
been moistened by the first rains. A second hilling is done at the beginning of
flowering.
According to Valdivia et al. (1999), mashua harvest index varies between 0.52
and 0.69.
6.3 Fertilization
In the Andean rotation system, the first crop after fallow is always the potato. It is
the preferred crop because of its importance in subsistence and for income
generation. The benefits of restored soil fertility after fallow and what little manure
or fertilizer may be available are targeted to potato productivity. Mashua frequently
follows the potato. In Peru, such fields are called ‘kqalpares de papa’, a term
indicating the condition of residual soil fertility. Mashua may also be used to conclude
the rotation before fallow, which indicates the crop’s comparatively low nutrient
demands.
Knowledge on mashua response to the application of mineral fertilizers is still
limited. Valdivia et al. (1999) found a significant increase response to fertilization
(from 28.8 t/ha in the unfertilized control to 36.1 t/ha using 80 kg N, 160 kg P and
80 kg K per hectare). However, mashua was found to be significantly less responsive
than oca to fertilization. This result was obtained in the relatively poor soils of the
Toralapa research station (Cochabamba, Bolivia). According to these authors an even
weaker response is to be expected in the soils normally used for mashua, which
usually contain more NPK.
6.4 Pests and diseases
Rusticity is one of the attractive attributes of mashua. Information on the severity of
mashua pests and diseases is still very limited. Some reports, such as Morales and
Rea (1982), indicate no relevant damage through pests or diseases. Of the 131 mashua
accessions observed by these authors in Belén (3820 m altitude) and Italaque (2800 m),
La Paz Department, Bolivia, few were affected by biotic agents. Nevertheless some
pests and diseases are reported in the literature and are discussed in the following
paragraphs. Rarely, though, do they cause significant damage. No pests or diseases
are currently known that constitute serious constraints to mashua cultivation.
Promoting the conservation and use of underutilized and neglected crops. 25. 17
6.4.1 Viruses
The first virus reported in mashua was a mosaic, which can be easily transmitted to
a range of hosts including Amaranthaceae, Chenopodiaceae, Asteraceae, Fabaceae,
Solanaceae and other Tropaeolaceae (Delhey and Monasterios 1977). Brunt et al.
(1996) have isolated and named two mosaic virus species: Tropaeolum 1 potyvirus
and Tropaeolum 2 potyvirus. They frequently occur together and are transmitted by
mechanical inoculation. The aphid Myzus persicae can act as a vector. However, these
viruses are not transmitted by seed. A similar potyvirus has been reported by Soria
et al. (1998). The authors named the virus Tropaeolum mosaic potyvirus (TropMV).
This virus can also be transmitted mechanically and by Myzus persicae aphids. A
survey of Ecuadorian germplasm from INIAP’s Santa Catalina Experimental Station
revealed that 34 out of 46 accessions were infected by TropMV; all eight fields
surveyed contained infected plants. Observations at Santa Catalina also suggest that
virus infection increases when mashua is cultivated in the same field for more than
one cropping season.
There are still important aspects of mashua viral diseases that are unknown, such
as the actual impact of these viruses under field conditions, how easily virus-free
material can be obtained and how fast reinfection would occur in the field (Lizarraga
1993).
A wide spectrum of viruses has been reported to infect the garden nasturtium
Tropaeolum majus L. (Soria et al. 1998), which suggests that other viruses might be
found in the closely related mashua. Viruses have not, however, been found to affect
mashua cultivation significantly.
6.4.2 Fungi
A comprehensive treatment of mashua diseases by Ames de Icochea (1997) describes
several fungal pathogens, some of them causing considerable damage, particularly
those that affect the harvested tubers. Acroconidiella leaf spot is always present in
mashua crops, and its symptoms increase at the end of the cropping season, when
all plants are affected to some extent. The causal agent, Acroconidiella tropaeoli, affects
other wild and cultivated Tropaeolaceae. Sclerotinia sclerotiorum includes mashua
within its wide host range. Sclerotinia usually affects localized sectors of the crop at
the middle of the cropping season. It can cause considerable damage but this appears
to be restricted to exceptionally wet conditions, as is the case with an as yet
unidentified species of Phytophthora which can affect mashua.
Ascochyta pinodes affects mashua under field conditions. However, this so-called
black tip disease is only significant as a storage problem. Species of the fungal genera
Mucor, Rhizopus, Cylindrosporium, Penicillium and also bacteria such as Erwinia and
Pseudomonas have been isolated from mashua; however, they represent opportunistic
pathogens affecting the crop in the wake of frost or desiccation damage (Ames de
Icochea 1997).
López (1986) reported Rhizoctonia spp. and Phytium affecting tubers and roots;
Fusarium spp. that produces wilting plants and rotting roots; Verticillium spp. and
18 Mashua (Tropaeolum tuberosum Ruíz & Pav.)
Acrocylindrium spp., causing rotting in low stems and roots. Falconi and Morales
(1983) reported mashua as host for several fungal species: Acrocylindrium,
Cylindrophora, Phytium, Arthrobrotis, Fusarium, Verticillium and Mucor, producing
some rotting in tuber roots; Graphium, Epiccocum, affecting the aerial stems; Pyronema,
Alternaria, Ovularia, Heterosporium, Stemphylium, Eriocercospora, Mycosphaerella and
Ulocladium, producing leaf spots. Nevertheless, none of these fungal diseases appear
to significantly decrease yields. Mashua is still considered a highly tolerant crop and
the application of fungicides is not recommended, even in the presence of the fungal
pathogens listed above.
6.4.3 Nematodes
In a screening for nematodes in Andean tubers Montero (1993) found species of
several genera affecting mashua: Criconemoides, Paraphelenchus, Discocriconemella,
Drylaimus, Aphelenchus, Neocriconema and Heiciclophora, a larger number than those
affecting oca and ulluco. Members of the Criconematidae (Criconemoides,
Discocriconemella and Neocroconema) were present only on mashua, suggesting that
mashua has a particular interaction with members of this family.
Mashua has been mentioned as a host of the oca nematode, Thecavermiculatus
andinus (Franco and Mosquera 1993). However, its sensitivity is low, similar to wild
herbs like Salvia spp. and Senecio vulgaris, and well below potato, ulluco and oca.
Similarly, mashua can be infected by the potato nematode Nacobbus aberrans, but
most genotypes showed no symptoms. Only one from eight genotypes developed
nematode nodules and two were not infected at all (Balderrama and Franco 1994).
Similar results were reported in Toralapa, Bolivia (Anonymous 1995b), where three
accessions tested did not show any infection with N. aberrans. These results strongly
suggest that mashua has a high level of tolerance to nematodes.
6.4.4 Insects
INIAP (1988) indicates that the mashua germplasm in the INIAP collection was
affected by Copitarsia turbata (‘gusano cortador’) and the leaf miner (Philionorycter
sp.). A later report (INIAP 1993) found that most of the accessions held at Santa
Catalina, Ecuador, had been affected by Copitarsia spp. However, 33% of the
accessions suffered no or only minor damage (0–2% of the tubers damaged).
Interestingly, mashua was more sensitive than oca to the Andean weevil,
Premnotrypes vorax, and was less effective when used as a control barrier (Calvache
1991). Ruales and Moscoso (1983) reported the following insects in mashua at farm
level: Copitarsia turbata, attacking aerial stems; Phyllonoryeter spp., mining leaves,
and Thysanoptera spp., also affecting leaves.
6.5 Harvest and yields
Information on mashua yields still reflects predominantly small trial plot results,
rather than actual farm yields (Table 3). Experimental plot results indicate that
mashua is very productive, particularly on a fresh weight basis. Yields frequently
Promoting the conservation and use of underutilized and neglected crops. 25. 19
Ta
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32
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20 Mashua (Tropaeolum tuberosum Ruíz & Pav.)
are well over 30 t/ha . Single plant yields under favourable conditions can exceed
2 kg of tubers (Fig. 12). Nevertheless, mashua yields at commercial level reach only
5–10 t/ha (Vimos 1987; Romero et al. 1989). Harvest time is 6–9 months after planting.
The crop is ready to harvest when plants become yellowish or brown, when they
lose their lower leaves or, in some cases, when they produce the tubers on the soil
surface (Cabrera 1986; Vimos 1987).
Owing to the small size of cultivated areas and predominant intercropping,
mashua is harvested manually. The soil is opened using a hoe and tubers are collected
in baskets or sacs. Farmers prefer to harvest in dry soil, which falls off the tubers
completely and leaves them clean. Mature tubers can be left in the soil for some time
before harvest. They will not sprout immediately after the aerial parts have senesced.
Once tuber dormancy is over, sprouts emerge from the tuber eyes. Sprouting tubers
lose weight and their quality deteriorates rapidly, because of water loss and nutrient
translocation to the new tissues.
6.6 Post-harvest
Since mashua is a secondary crop, very little research has been done on post-harvest
practices. Standard traditional practices include cleaning the tubers to eliminate soil
(if harvested in wet, clayey soil), tuber classification according to size and colour, and
the elimination of injured or bruised material. Like other tubers, mashua are often
stored in farmhouses. South of central Peru the tubers are commonly held outdoors
Fig. 12. Two freshly harvested mashua
plants with senescent aerial parts shown by
Ing. Pedro Cruz, Fierroccata, Cusco, Peru,
3500 m altitude (Photo M. Hermann, May
1991).
Promoting the conservation and use of underutilized and neglected crops. 25. 21
in heaps, called ‘phinas’ (Fig. 13), that are covered with a layer of dry ichu grass (Stipa
ichu (R. & P.) Kunth). This layer is a few centimetres thick, enough to maintain the
tubers in darkness and insulate them against night frosts (which are common after
the harvest period) and high day temperatures. The temperature variation underneath
the cover is small and oscillates minimally around the average ambient temperature,
which in the high-altitude location shown in Fig. 13 is probably below 8°C.
Before consumption mashua tubers are exposed to direct sunlight for a few days
to improve sweetness and palatability. This practice increases sugar content,
probably by inducing starch degradation; it may also reduce isothiocyanates.
Mashua tubers have relatively high hourly respiration rates: 75 mg CO2/kg of
tuber fresh weight 3 hours after harvest and 54 mg CO2/kg 3 days after harvest, at
20ºC. This is approximately the same as oca and twice as much as potato and yacon
stored under the same conditions at 17°C in the shade (Grau 1993).
6.7 Crop ecology
As with most other Andean root and tuber crops, the information on mashua ecology
is scarce, and the crop’s ecological requirements must be derived from its
distributional record. The latitudinal range of mashua cultivation in the Andes is
from 8°N to 24°S, and the altitude ranges from 2400 to 4300 m, with the most frequent
occurrence between 3000 and 3700 m, where annual mean temperatures are in the
range of 8–11°C.
Fig. 13. Farmhouses with ‘phina’ tuber
stores at Accha Alta, Calca Province,
Cusco, Peru, at 3900 m altitude (Photo M.
Hermann, September 1990).
22 Mashua (Tropaeolum tuberosum Ruíz & Pav.)
Morales and Rea (1982) indicate that mashua aerial organs behave similar to
those of oca and ulluco in relation to frost, tolerating –1°C, but they are completely
destroyed at –5°C. Reports from Romero et al. (1989) indicate that at early crop stages
mashua is able to recover completely after exposure to –2 to –4°C. Recovery may not
be complete at later stages. For instance, Valdivia et al. (1999) indicate that a frost
event of –3.5°C at Toralapa, Cochabamba, Bolivia, killed aerial plant parts. Above
freezing temperatures, mashua tolerates long chilling periods. It is able to tolerate
temperatures of 30°C or slightly higher, at least for short periods.
Although mashua is normally grown at high altitude, it is the cool conditions
and not altitude per se which are required by the plant. Mashua is able to tuberize
and produce good crops outdoors at sea level at 46°S in Christchurch, New Zealand
(Martin et al. 1996) and at 49°N in Vancouver, Canada (Johns and Towers 1981).
In general mashua is not irrigated. However, it has high water requirements and
receives between 700 and 1600 mm on most cultivation sites. It tolerates the misty
and cloudy weather typical of the eastern Andean ranges, and indeed thrives there,
but it also seems capable of tolerating dry spells.
Mashua is a short-day plant, requiring less than 12 hours for tuberization.
Razumov (1931) obtained no tubers with 18 hours, while plants exposed to 10-hour
or 12-hour days produced tubers. Bukasov (1930) indicates that 9 hours is the
optimum day length for tuberization. Under long-day conditions (14–20 hours, in
Finland) mashua is unable to tuberize (Kalliola et al. 1990). Day length also affects
bud number, stem number, leaf size and pigments in stem and leaves. Tuberization
in New Zealand at 46ºS begins when day length becomes less than 12 hours. In South
Island, New Zealand and Vancouver Island, Canada, tuberization and yield
formation are still possible because autumn temperature minima are moderated by
maritime influence. Under colder continental conditions crop development is
truncated by frost before tuberization occurs. This may explain why there is no report
for the successful production of mashua tubers in continental Europe, although the
species may have been introduced several times by botanists and hobby gardeners
(Bukasov 1930, Kalliola et al. 1990).
Apparently, mashua has a wide tolerance to soil pH, growing from 5.3 to 7.5
(National Research Council 1989). However, there has been little systematic research
in this area.
7 Composition and uses
7.1 Chemical composition
The water content of mashua tubers is comparatively high, ranging from 79 to 94%
in the fresh or edible matter (Table 4). As seen in Table 4, the main nutritional
contribution of mashua is its high carbohydrate content, particularly of starch but
also of sugars. The protein content of the mashua tuber in the fresh matter approaches
potato values, although potatoes are much higher in dry matter content. Amino acid
analyses of mashua tuber protein show a satisfactory nutritional composition
compared with the WHO recommendations (Shah et al. 1993; Stegemann and Shah
1993). These authors also indicate that there is considerable variation between the
different clones analysed. The high content of ascorbic acid or vitamin C as determined
by Collazos et al. (1996) (77.5 mg per 100g fresh matter) is nutritionally also important.
Like other Tropaeolaceae, mashua contains isothiocyanates present as
glucosinolates (Kjær et al. 1978), compounds similar to those found in other crucifers
and also known as mustard oils. Isothiocyanates are well known for their antibiotic,
insecticidal, nematocidal and diuretic properties, which substantiates mashua’s
extensive use in Andean folk medicine (Johns et al. 1982). The characteristic piquant
flavour of mashua is produced by p-methoxybenzyl isothiocyanate. This compound
appears to be specific to subspecies T. tuberosum ssp. tuberosum, which also contains
small amounts of 2-propyl isothiocyanate. This compound and 2-butyl isothiocyanate
are the main isothiocyanates in subspecies T. tuberosum ssp. silvestre (Johns and Towers
1981). Thiocyanates liberate cyanide by hydrolysis and could lead to poisoning.
Thiocyanate values higher than 20 mg/100 g are common in uncooked mashua.
Dolores and Espín (1997) found values of 23–33 mg/100 g in mashua accessions of
the Ecuadorian collection at INIAP.
7.2 Food uses
As explained above, most mashua tuber types contain high levels of isothiocyanates,
which give them a pungent taste, and make them unsuitable for raw consumption
except in small quantities. Boiling causes the isothiocyanates to hydrolyse,
eliminating cyanide and improving taste. Mashua tubers are usually boiled to form
a stew with other tubers and meat (Cortés 1981). Mashua can be used as a component
in many dishes, ranging from soups and stews to desserts.
The widespread Andean practice of exposing tubers and roots to direct sunlight
is also used for mashua, as mentioned above, in order to increase sweetness and
reduce cyanide levels before cooking (Dolores and Espín 1997). Another common
procedure is baking mashua in the huatia, an earthen field oven, which is typically
prepared at harvest time from sods or stones. Baked mashua has an aromatic flavour
and a mushy texture like that in moist sweet potatoes.
Cárdenas (1989) indicates that in the recent past it was a common practice for
the middle class of La Paz, Bolivia, to eat cooked and frozen mashua soaked in
sugarcane syrup (‘thayacha’) during the winter. This is a rare practice today.
24 Mashua (Tropaeolum tuberosum Ruíz & Pav.)
Ta
bl
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4.
  C
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)
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19
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Tr
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to
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Cy
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4–
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*
 R
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s i
nd
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 m
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 a
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 va
lue
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 b
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s, 
typ
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 w
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wi
th
 d
iffe
re
nt
 cu
ltiv
ar
s.
Promoting the conservation and use of underutilized and neglected crops. 25. 25
The per capita consumption of mashua tubers is difficult to assess and its
derivation from national production and population figures makes little sense
because consumption varies hugely from place to place. In rural areas, where native
Amerindian people predominate, consumption can be considerable, especially
during and after harvest, when mashua is a frequent side dish or ingredient in main
dishes, and weekly per capita consumption may be well over 1 kg. However,
consumption is limited by the seasonal availability of tubers.
Mashua flowers can be consumed fresh in salads, and flower buds can be
consumed pickled in vinegar in a similar way to capers (King 1987).
7.3 Feed uses
Peasants in Ecuador (Tapia et al. 1996) and in the Candelaria area, Cochabamba,
Bolivia (Terrazas and Valdivia 1998), feed mashua tubers to pigs. Few studies have
been published on the use of mashua tubers as feed, but this use is presumably much
more widespread than the reports suggest. The first research trial was apparently
conducted by Bateman (1961) in Costa Rica. According to this author raw mashua
plus skimmed milk and grains performed better than grains alone, and acceptability
of mashua was considerably improved by cooking. Similar results were obtained by
Ramos et al. (1976) in a more comprehensive experiment comparing a standard ration
(barley, oats (Avena sativa L.), maize, cotton paste, hay and milk) and a ration in which
the cereals and cotton had been replaced by cooked mashua. They concluded that
the increase of weight in calves fed with mashua did not differ significantly from
that of the control, and that including mashua in the diet was a cheaper alternative.
They suggested that cooked mashua can be replaced by raw mashua, to feed animals
over 12 months of age
However, there is evidence indicating that raw mashua can be toxic. One of us
observed a donkey in San Juan, Chimborazo, Ecuador, dying immediately after
ingesting a portion of fresh mashua tubers (Nieto 1988, personal observation). Lethal
effects of mashua intake on donkeys or mules have also been reported during
ethnobotanical field work in Pisac, Cusco (King 1988), and in Colanzulí, Salta,
northern Argentina (Hermann 1992, personal observation). Presumably, cultivars
producing such effects contain high levels of isothiocyanide.
There are no reports of aerial parts being used as forage, which is somewhat
surprising, because mashua produces much above-ground biomass. This may be
partly explained by the presence of the same antinutritional substances that
accumulate in the tubers. Salcedo (1986) indicates that ‘stems and leaves are as
poisonous as tubers’. There is no information on hay production or silage, treatments
that may reduce or eliminate isothiocyanates.
7.4 Flour and starch
Information on mashua processing into flour and on starch properties is limited. The
first published report on mashua flour production indicates disappointing results.
Processing yielded a piquant and strongly smelling product with a yellow greyish
26 Mashua (Tropaeolum tuberosum Ruíz & Pav.)
colour, not suitable for human consumption (Deza 1977; Cortés et al. 1982). Recent
studies (Ramallo 1999) suggest that mashua flour could be an interesting alternative
as pig feed. Mashua flour has an acceptable protein (8%) and high energy content
(416 kcal/100 g). Protein levels are similar to those of maize but energy content is
30% higher (Ramallo 1999). The high water content of mashua, however, implies
relatively high drying costs.
Although production of flour for human consumption still has technical
problems to be solved, mashua starch has interesting characteristics. Compared to
potato, ulluco, oca, achira (Canna indica L.), maize, rice and manioc (Manihot esculenta
Crantz), mashua and arracacha (Arracacia xanthorrhiza Bancroft) starch gels are the
most resistant to freezing (minimal syneresis), after 4 weeks at –20ºC (Dufour et al.
1997). However, these results are partially contradicted by Villacrés and Espín (1996),
who found high syneresis values after a freeze–thaw cycle. These authors also found
that gelatinization temperature of mashua starch (60°C) is similar to that of oca starch
(60°C) and ulluco starch (63°C), and higher than that of potato starch (55°C). A factor
making mashua starch use economically unfeasible would be its very low extraction
yield—2.3 %, according to Villacrés and Espín (1996). However, starch recovery rates
reported by these authors for a number of tuber species appear to be unreasonably
low. Mashua starch granules are small, with an average size of 8.8 µm (Cortella and
Pochettino 1995) to 15.6 µm (Villacrés and Espín 1996).
7.5 Medicinal uses
‘. . . llamada añus. Dizen los indios que comida es contraria a la potencia
generativa; para que no les hiziesse daño, los que se preciavan de galanes
tomavan en la una mano una varilla o un palillo mientras la comían, y comida
assí dezían que perdía su virtud y no dañava. Yo les oí la razón y algunas vezes
vi el hecho, aunque davan a entender que lo hazían más por vía de donaire que
no por dar crédito a la burlería de sus mayores’1
Garcilaso de la Vega, writing in the sixteenth century in his
‘Comentarios Reales’, in reference to folk beliefs of
antiaphrodisiac properties of mashua in what is today Peru
(as quoted in Patiño 1964).
Mashua diets are reputed to have beneficial effects on liver and kidneys (Hodge
1946; Oblitas 1969) and to alleviate prostate and urinary disorders (Salcedo 1986;
__________________________
1
‘It is called añus; and the Indians say that it reduces the procreative powers, but in order to
prevent it from harming them, those who prided themselves on their gallantry used to hold a
little rod or stick in one hand while they ate it, saying that in this way it lost its peculiar property
and did them no harm. I heard them give this explanation and often saw them do this, though
they implied that it was intended rather as a joke than as a serious acceptance of the foolish
tradition of their elders.’ (Translation taken from Garcilaso de la Vega (1966)).
Promoting the conservation and use of underutilized and neglected crops. 25. 27
Brack 1999). In Bolivia mashua is still used in treating prostate ailments and is sold
in small quantities in urban areas (Terrazas and Valdivia 1998). Bateman (1961)
indicates that mashua fed to pigs seemed to have a diuretic effect. This is consistent
with similar effects on humans as recognized by Andean folk medicine (Cárdenas
1948, 1958; Brack 1999).
Rea (1984) reported the use of mashua by people with diabetes, using both the
cooked tubers and water in which they were boiled. In their catalogue of Ecuadorian
mashua germplasm Tapia et al. (1996) list several accessions with collector
information on medicinal uses for the treatment of tonsillitis, dengue and malaria
fever, and postpartum conditions. Pérez Arbelaez (1947) indicates that mashua is
useful for the treatment of skin ailments, such as eczema and ‘skin spots’. The same
dermatological properties are attributed to kita-añu (T. tuberosum ssp. silvestre)
(Herrera 1921). Without specifying used plant parts, Oblitas (1969) indicates that
mashua is used as vermifuge and ‘to induce menstruation’.
Among its numerous reputed medicinal effects, mashua is best known in the
Andes for its alleged capacity to suppress sexual appetite and decrease reproductive
potential and erectile function in men (Herrera 1933; Hodge 1951; Leon 1967; Oblitas
1969; Brack 1999). According to traditions recorded by the sixteenth century
chroniclers, the Inca fed mashua to their troops ‘so that they would forget their
women’ while on military operations (Patiño 1964, citing Padre Bernabé Cobo). With
similar intentions, rural women in Cusco today prepare concoctions from mashua
and add these surreptitiously to their men’s food, in the hope of preventing them
from becoming unfaithful (Hermann 1992). Johns et al. (1982), who review such folk
beliefs, showed experimentally that they are perhaps not unsubstantiated. Male rats
fed with mashua tubers had significantly reduced testosterone and
dihydrotestosterone levels (45%). However, test animals maintained their capability
to impregnate females.
Hormonal effects may also be responsible for the reputed effects on menstruation.
Johns et al. (1982) failed to find any oestrogenic activity in guinea-pigs. However,
their results indicate that N,N-di-(methoxy-4-benzil)thiourea present in mashua can
competitively inhibit radioactively marked estradiol using a preparation of estrogen
receptor in calf uterine cytosol, suggesting that mashua may still have estrogenic
activity.
Considering the hormonal effects of mashua in rats, and its range of active
substances, it would not be surprising if these attributes were found to be the basis
for some of the reputed medicinal properties. However the experimental background
is still limited to the work of Johns and co-workers (1981, 1982).
8 Taxonomy and biosystematics
8.1 Taxonomic history and synonyms
The Tropaeolaceae is a small and quite homogeneous family of herbaceous, mostly
climbing species. The family includes three genera, with two of them, Trophaeastrum
Sparre and Magallana Cav., restricted to Patagonia. The largest genus is Tropaeolum,
which contains 86 species, distributed from southern Mexico throughout South
America (Sparre and Andersson 1991).
The tuber forming capacity is present in all genera of the family. Species that
have been positively identified as tuber forming are listed in Table 5. The list may
not be complete, as in many cases Sparre and Andersson (1991) report ‘subterranean
parts not seen, probably tuberous’. Trophaeastrum patagonicum (Speg.) Sparre yields
white edible tubers that can reach a length of 10 cm. These tubers were used by
several Patagonian tribes, baked or boiled with milk (Martínez Crovetto 1982). There
Table 5.  Tuber-forming species in the Tropaeolaceae (after Sparre and
Andersson 1991)
Species Distribution
Magallana porifolia Patagonia
M. trialata Northern Patagonia
Tropaeolum rhomboideum * Chile
T. azureum * Chile
T beuthii * Chile
T. brachyceras * Chile
T. ciliatum * Chile
T. hookerianum * Chile
T. leptophyllum ssp. Chile, Argentina
Leptophyllum = T. edule *
T. pentaphyllum † Central Argentina, Southern Brazil, Uruguay
T. polyphyllum * Chile, Argentina
T. sessilifolium * Chile 
T. tricolor * Chile
T. umbellatum ‡ Southern Ecuador
T. patagonicum = T. patagonicum Patagonia
* Section Chilensia; † Section Chymocarpus; ‡ Section Umbellata.
Promoting the conservation and use of underutilized and neglected crops. 25. 29
are also reports of T. leptophyllum G. Don being eaten by Chilean indigenous people
in times of scarcity (Bridges 1842).
Tropaeolum L. was referred to originally as Cardamindum Adans. Linnaeus (1735)
introduced the name Trophaeum ( = trophy), because of the flowers resembling a
warrior’s helmet and leaves resembling shields; he later changed the name to
Tropaeolum, using the original Greek word ‘trópaion’, which has the same meaning.
Tropaeolum tuberosum was described by Ruíz and Pavón (1802) in their magnificent
work ‘Flora Peruviana et Chilensis’, in which for the first time they gave an original,
detailed and illustrated (Plates CCCXIII, CCCXIV) account of the species (Fig. 3).
Synonyms of T. tuberosum are T. mucronatum Meyen, T. suberosum Walpers (1857)
and T. denticulatum (Kuntze 1891). Heynhold (1840) proposed to transfer the species
to a different genus as Chymocarpus tuberosus (Ruíz & Pav.) Heynh.
Bukasov (1930) proposed the name T. cubio for the mashua forms found in
Colombia, which he considered taxonomically distinct from those cultivated in
Ecuador, Peru and Bolivia. Sparre and Andersson (1991) dismissed this proposal,
indicating that the features pointed out by Bukasov are of little taxonomic value.
The only other widely cultivated Tropaeolum species is T. majus L., the garden
nasturtium, which is a popular ornamental in temperate areas.
8.2 Wild species closely related to cultigen
T. tuberosum has been placed within the section Mucronata by Sparre (1973). This
section includes five well-defined species, of which T. longiflorum Killip, T.
crenatiflorum Hook. and T. purpureum are endemic to Peru; T. cochabambae Buchenav.
occurs in Peru and Bolivia. All species within the section, except T. tuberosum, have
relatively reduced geographic ranges.
Sparre (1973) and Sparre and Andersson (1991) recognize two subspecies of T.
tuberosum; the cultivated T. tuberosum ssp. tuberosum and the wild T. tuberosum ssp.
silvestre, with the latter being smaller and more slender in all parts (Fig. 14).
Fig.14. A–C: Wild mashua (Tropaeolum
tuberosum ssp. silvestre). D–E:
Cultivated mashua (Tropaeolum
tuberosum ssp. tuberosum). C, E: Petals:
i, inferior petal; s, superior petal. Scale A,
B, D, 3 cm; C, E, 1 cm (Reproduced from
Sparre and Andersson 1991, p. 53, with
permission from the Nordic Journal of
Botany/Opera Botanica).
30 Mashua (Tropaeolum tuberosum Ruíz & Pav.)
According to Sparre and Andersson (1991), the only consistent character to
differentiate both subspecies is the lack of tubers in T. tuberosum ssp. silvestre. This
appears to be a questionable distinction, for the authors state that the presence or
absence of tubers ‘can usually not be ascertained in herbarium material’ of T.
tuberosum. Moreover, the authors’ classification does not accommodate clearly wild
but tuberous T. tuberosum, such as the ‘kipa isaño’ used by Johns and Towers (1981)
and material known to the authors of this monograph from Paruro Province, Cusco,
Peru. This material has elongated and twisted tubers, a feature retained in
cultivation and setting this species apart from the domesticate (Fig. 15). Ortega
(2000) in his thesis on ‘noncultivated’ mashua pictures over a dozen accessions with
similar tuber shapes, predominantly collected in Cusco’s Calca and Paucartambo
provinces. Notably, the same author reports the frequent occurrence of escaped
mashua cultivars that could be mistaken by the collecting botanist for wild
Tropaeolum species.
Nevertheless, phytochemical analyses of isothiocyanates by Kjær et al. (1978) and
Johns and Towers (1981) support the assessment of two subspecies in T. tuberosum:
cultivated mashua and a tuber-bearing wild species, referred to by these authors as
T. tuberosum ssp. silvestre. Mashua produces p-methoxybenzyl isothiocyanate,
whereas the wild material is characterized by benzyl-2-propyl isothiocyanate and
2-butyl isothiocyanate.
Fig. 15. Tubers of wild mashua from Paruro Province, Cusco, Peru. (Material shown is
from cultivation in a field genebank near Cusco, 3500 m alt; planted October 1990,
harvested May 1991, Photo M. Hermann).
Promoting the conservation and use of underutilized and neglected crops. 25. 31
Both wild and cultivated T. tuberosum extend from Venezuela to northwest
Argentina along an impressive latitudinal gradient (Fig. 16). Sparre and Andersson
(1991) show two areas of high concentration of ssp. silvestre, one in northern Peru
and a larger one in Ecuador. However, Hermann (field observations) found wild
mashua with sizeable tubers in Peru, Bolivia and Argentina, but not in Ecuador or
Colombia. To accommodate the different observations within a model it seems
necessary to consider the presence of wild T. tuberosum and plants escaped from
culture overlapping at least partially along a wide range.
Fig. 16. Distribution of (A) cultivated mashua (Tropaeolum tuberosum ssp. tuberosum) and (B)
wild mashua (T. tuberosum ssp. silvestre) (Reproduced from Sparre and Andersson 1991, p. 54,
with permission from the Nordic Journal of Botany/Opera Botanica).
Table 6.  Chromosome numbers of Tropaeolum species
Species 2n Cultivated/ Source
Wild
T. canariense 28 n.a. Heitz 1926*
T. cochabambae 26 n.a. Huynh 1967
T. majus 28 n.a. Sugiura 1925*
T. minus 28 n.a. Heitz 1926*
T. peltophorum 28 n.a. Sugiura 1925*
T. peregrinum 24, 26–30 n.a. Warburg 1938*
T. pubescens 28 n.a. Huynh 1967
T. tricolor 52 n.a. Huynh 1967
T. tuberosum 42 n.a. Sugiura 1925*
T. tuberosum 42 n.a. Acosta Solís 1980
T. tuberosum 48 Cultivated? Marks 1976†
T. tuberosum ssp. tuberosum 52, 64, 51, Cultivated Gibbs et al. 1978
48, 43
T. tuberosum ssp. silvestre 42 Wild Johns and Towers 1981
T. tuberosum ssp. tuberosum 52 Cultivated Johns and Towers 1981
T. tuberosum ssp. tuberosum 18, 27, 36 Cultivated Román and García 1997
* Cited by Darlington and Ammal 1945; † cited by Gibbs et al. 1978.
9 Genetic resources
9.1 Chromosomal variation
Chromosome studies of mashua have shown contradictory results (Table 6). The
presence of two subspecies, high ploidy levels, hybridization, and sexual and
asexual reproduction, are the likely factors that contribute to a blurred picture.
Huynh (1967), Gibbs et al. (1978) and Johns and Towers (1981) have reported a
sporophytic number of 2n = 52 for T. tuberosum ssp. tuberosum. However, Gibbs et
al. (1978) also propose 2n: 43, 48, 51 and 64 as less frequent values emerging from
abnormal meiotic configurations. Johns and Towers indicate 2n = 42 for subspecies
silvestre. Huynh (1967) and Gibbs et al. (1978) propose that mashua is of
autopolyploid origin, on the basis of the frequent formation of multivalent and
secondary associations during meiosis. However, Gibbs and co-workers do not rule
out segmental allopolyploidy with a closely related species, such as T. cochabambae.
Promoting the conservation and use of underutilized and neglected crops. 25. 33
Hybridization between species has been observed in several other cases in the
Tropaeolaceae (Sparre and Andersson 1991). Combining phytochemical and
chromosomal information, Johns and Towers (1981) reinforce this hypothesis. They
maintain that T. tuberosum ssp. tuberosum is of allopolyploid origin and resulted
from the hybridization of T. tuberosum ssp. silvestre with another Tropaeolum species
of 2n = 26, probably T. cochabambae. Nevertheless these authors analysed a relatively
narrow sample of T. tuberosum germplasm. Recent findings by Román and García
(1997) working with 20 Peruvian accessions of cultivated mashua indicate different
ploidy levels, 2n = 18, 27 and 36, with a basic number of x = 9, and thus have added
further complexity to the issue. To shed more light on this complex picture a more
comprehensive study is warranted, including a range of accessions of both
subspecies along the whole latitudinal gradient.
9.2 Molecular variation
Some variation has been detected at the molecular level in different mashua clones.
Shah et al. (1993) studied the protein patterns in mashua tubers using polyacrylamide
gel electrophoresis (PAGE). They found that porosity gradient PAGE was the most
suitable method for discriminating varieties, and SDS disk PAGE was the least
suitable. The main protein bands in mashua are located in the 43–110 kD range.
Monteros et al. (1997) found variation for phosphoglucomutase and malate
dehydrogenase and were able to classify the INIAP mashua collection in 10 groups
using this isoenzymatic information.
9.3 Genetic erosion, germplasm collection and genebank holdings
There is general agreement that mashua has suffered a progressive reduction in its
production and consumption, particularly during the last decades of the twentieth
century. However, there has been no systematic evaluation of that process and there
is no quantitative information on its present status.
Efforts to safeguard genetic resources of mashua and other Andean roots and
tubers began relatively early in the Andean countries. The first record of mashua
accessions being maintained date from 1958 (Holle 1986). In Peru, the first
collecting activities are documented in the 1970s. However, mashua ex situ
conservation gained momentum during the 1980s. At present Peru maintains the
largest number of holdings and the largest number of accessions (Table 7). In
Ecuador, collecting and conservation efforts have been spearheaded by INIAP, at
the Santa Catalina research station, which also holds a few accessions from
Colombia (Tapia et al. 1996).
The initiative of the Collaborative Program of Andean Root and Tuber Crop
Diversity funded by the Swiss Development Cooperation and implemented by CIP
and numerous national programmes ensured that exploration and collecting
activities continued in Bolivia, Ecuador and Peru during the period 1993–1997.
In conclusion, large areas of the Andean countries have been covered by
collecting missions. As seen in the plot of mashua genebank accessions reported
34 Mashua (Tropaeolum tuberosum Ruíz & Pav.)
Table 7.  Mashua holdings reported for Bolivia, Ecuador and Peru
Organization Number of Report Source
accessions year
Bolivia
IBTA 110 1980 Holle 1986
IBTA 131 1983 Holle 1986
IBTA-PROINPA 71 1995 Ugarte 1995
Ecuador
Santa Catalina, INIAP-DENAREF 43 1985 Holle 1986
Santa Catalina, INIAP-DENAREF 36 1986 Holle 1986
Santa Catalina, INIAP-DENAREF 89 1996 Tapia et al. 1996
Peru
Camacani, Puno 19 1981 Holle 1986
Camacani, Puno 28 1986 Holle 1986
University of Puno 47 1995 Velazco 1995
Ayacucho University 78 1979 Holle 1986
Ayacucho University 94 1983 Holle 1986
Ayacucho University 146 1986 Holle 1986
Ayacucho University 14 1995 Anonymous 1995a
Ayacucho University 10 (in vitro) 1995 Anonymous 1995a
INIA Ayacucho 68 1995 Anonymous 1995a
University of Cajamarca 7 1995 Velazco 1995
INIA Cajamarca 103 1995 Velazco 1995
INIA Huanuco 22 1995 Velazco 1995
University of Huanuco 50 1995 Velazco 1995
INIA Huancayo 125 1995 Velazco 1995
Kayra Research Station 101 1986 Holle 1986
INIA Cusco 14 1995 Velazco 1995
University of Cusco 302 1995 Velazco 1995
University of Pasco 35 1995 Velazco 1995
University of San Marcos 48 (in vitro) 1986 Holle 1986
University of San Marcos 112 (in vitro) 1995 Velazco 1995
Experimental Station La Molina 151 1986 Holle 1986
University of La Molina 65 1995 Velazco 1995
CIP 119* 2002 CIP corporate 
databases
* 55 of these accessions are FAO designated.
Promoting the conservation and use of underutilized and neglected crops. 25. 35
from 1986 to 1998 (Fig. 17), there are several distributional discontinuities within the
species range from Colombia to northern Argentina. Mashua appears to be absent
from large areas of the Colombian highlands, except for Boyacá where mashua
consumption and market presence is atypically high, with no obvious explanation
for this phenomenon. Mashua also seems to be absent in Peru north of Cajamarca
and in Loja, Ecuador, perhaps because of the low altitude of the Andes there and the
lack of contiguous paramo environments suitable for mashua cultivation. Finally,
there are no genebank records of mashua accessions collected in the inhospitably
dry southern Altiplano in Bolivia. It is important to note that not all reported
accessions, perhaps only a fraction of the total ever collected, are actually available
from genebanks. This is mainly due to the fact that large numbers of accessions were
lost from field genebanks in the 1980s and early 1990s.
Fig. 17. Distribution
of cultivated
mashua (dots
represent collecting
sites of genebank
accessions reported
from 1986 to 1998,
source: CIP
databases; shaded
area is over 2000 m
above sea level).
36 Mashua (Tropaeolum tuberosum Ruíz & Pav.)
In situ conservation has been proposed as a strategy to complement genebanks
and ex situ conservation (for a review see Maxted et al. 1997, Ortega 1997). The concept
of in situ conservation has become fashionable and often a political issue. In the case
of mashua in situ and ex situ conservation should complement each other. However,
at present a systematic framework for in situ conservation is largely lacking. As in
the case of most other crops, on-farm maintenance of mashua germplasm by farmers
just happens as a normal part of farming practice (Fig. 18).
9.4 Maintenance of the collections and conservation strategies
Conservation of genebank holdings has been considered difficult to some extent.
This is because mashua needs to be replanted and harvested in field collections every
cropping season. This (clonal) conservation method is not only labour-intensive, but
can also be unsafe, because of the associated risks posed by pathogens, weather
conditions or poor management practices.
Micropropagation of mashua as plantlets grown in vitro using modified
Murashige and Skoog media (MS 4.6 g, AG3 0.25 ppm, sucrose 3%, agar 0.8%) as
first reported by Castillo (1991) is now widely practised to maintain clonal collections
under sterile conditions. A comprehensive study including protocols for callus and
Fig.18. Germplasm collectors examining
mashua on-farm diversity at Ajchahuata,
Calca Province, Peru, 3700 m altitude (Photo
M. Hermann, September 1990).
Promoting the conservation and use of underutilized and neglected crops. 25. 37
meristem culture, micropropagation by multiple shoot formation and nodal cuttings
has been published by Torres et al. (1992).
Conservation in genebanks as seed could be a less expensive and safer alternative
for germplasm conservation, as mashua produces sexual seeds that are easy to collect
and to store. However, more research is needed, in relation to the segregation of seed
progeny, seed physiology, storage conditions, longevity and germination conditions
(see Section 5.3). As far as we are aware, sexual mashua seed is currently not used
for mashua conservation, which entirely relies on clonal maintenance in field
collections and, to a lesser extent, on tissue culture.
No protocols are currently available for pollen storage and cryopreservation of
mashua.
10 Crop limitations
Pests and diseases do not seem to pose significant constraints to mashua culture.
The main limitation appears to be the low acceptance outside family or local
consumption. Urban demand for mashua is far below that for ulluco and oca, as
evidenced by the marginal presence of mashua in markets. The flavour of some
mashua lines is so strong that it deters potential consumers, even those used to spicy
foods and eager to try ‘novel’ foods. The shelf life of mashua is short as the tubers
lose water rapidly under the prevailing (rustic) storage conditions and deteriorate
within weeks after harvest, yielding a product with little or no market value. Some
cultivars do not tolerate storage at all: almost immediately after harvest, the tubers
of such cultivars sprout or rapidly rot, especially if they have been damaged during
harvest or transport. In summary, the main limitations for expanded use of mashua
at the present relate essentially to demand, marketing and post-harvest aspects, not
to production.
11 Breeding
Several potentially high-yielding clones have been identified in Ecuador (Vimos
1987, 1989; Romero et al. 1989; Monteros 1996), and these could be used to increase
the fresh tuber yields. Similarly, clones already identified for high protein content
or low isothiocyanate content should be used in breeding programmes.
Rea (1984) mentions the selection of high-protein mashua clones from plants
reproduced sexually. Some clones already identified by high protein content are:
• Asuti from Huancavelica, Peru, with 17% protein (on a dry matter basis)
• IBTA 3132, also from Huancavelica, Peru, with 16% protein
• Amarillo, from Italaque, Bolivia, with 15% protein
• the ornamental clone Pajarillo, from Puno, Peru, with 11% protein.
However, Rea suggests that the ‘bitterness’ trait (high levels of isothiocyanate)
in mashua may be associated with higher protein content. Thus, selecting for high
protein may reduce palatability but simultaneously increase tolerance to pests and
diseases.
Some potentially useful correlations have already been established in mashua.
Yield is correlated with size of tubers, number of tubers per plant, plant height and
crop duration. It seems that clones with black tubers are higher yielding than clones
with yellow or white tubers (Vimos 1987; Romero et al. 1989). Yield is not associated
with stem width, stem number and tuber length (Delgado 1978a, 1978b). In any case,
because mashua produces sexual seeds, crosses between different clones are possible.
Although seed germination in mashua is low, breeding programs by hybridization
should be an easy way to increase variation and to select clones with desired
characters.
12 Prospects and research needs
12.1 Advantageous features of mashua 
Mashua has many attractive features, actual or potential:
• high productivity, in terms of tuber fresh and dry weight
• high pest and disease tolerance
• good ground cover and prevention of erosion
• feed potential for tubers and aerial parts
• adaptation to chilling temperatures and poor soils
• reasonably good post-harvest life, if managed properly
• medicinal properties
• potential as bioinsecticide
• potential as an ornamental.
12.2 Development objectives and research needs
Subsistence Andean farmers need to maintain mashua’s good characteristics, such
as rusticity, high tolerance to pests and growth habit, while improving palatability
and post-harvest life. However, improving the palatability of mashua for both
humans and domestic animals would mean reducing the isothiocyanate content,
and this may reduce its capacity to resist pests and diseases. Developing mashua as
a commodity for city consumption will require focusing on external aspect, colour
and shape, palatability and post-harvest life, and may require a trade-off with
maintaining pest tolerance and rusticity.
Mashua has attracted some research attention in recent years. Nevertheless, it is
the least well known of the Andean tubers. Several research areas and specific
activities can be suggested:
• Mashua’s reputed medicinal properties deserve more attention. In the same way
that ‘aphrodisiac’ properties have been a key element of the recent success of
maca (Lepidium meyenii Walpers), reputed libido-reducing properties may be
responsible for the failure of mashua. No research has been conducted in this
field since the pioneering work of Johns et al. (1982). It is essential to quantify the
effect of mashua diets in humans, especially in the long term. Similar attention
needs to be paid to all the reputed medicinal properties of mashua.
• If substantiated, mashua’s libido-reducing property may be an attribute of
economic interest for the formulation of animal feed, in order to reduce sexual
drive and accelerate growth and weight gain.
• Some research has been conducted on the use of mashua tubers for feed, but
more detailed information is needed to promote its use, and to assess safe intake
levels. The chemical composition and nutritional properties of the aerial parts of
the plant is poorly known, as is the potential for silage production that may
reduce or eliminate isothiocyanates.
• Sound dietary information of the tubers may be one of the elements required to
overcome lack of market demand in the cities, particularly by social groups with
Promoting the conservation and use of underutilized and neglected crops. 25. 41
higher purchasing power. The development of city markets would also require
the development of post-harvest technologies and associated information on
tuber physiology. Mashua starch may have some interesting properties, but more
information is required to evaluate if it is sufficiently attractive to succeed in the
competitive starch market. Processing technologies focusing on pickling and
canning, pigment and texture preservation need experimentation.
• Although mashua appears to be highly tolerant to a range of pests and diseases,
recent studies have shown that there is a range of potentially harmful pathogens.
It is necessary to evaluate whether mashua’s apparent tolerance will remain intact
if it is cultivated on a larger scale, and if it could still be produced as an essentially
‘organic’ crop as is currently the case.
• The extent to which mashua is affected by virus disease should be quantified
under field conditions, and the advantages of using virus-free material need to
be assessed. Fungal diseases also require consideration, particularly those
affecting mashua after harvest.
• Although little is known about mashua’s floral biology and its crossability with
wild species and other Tropaeolum species, the crossability observed in other
Tropaeolum species provides pointers for the potential ease of mashua breeding.
Studies on pollen viability and longevity, seed viability, germination, dormancy
and longevity need to be conducted.
• The present germplasm collections probably contain a representative sample of
the variability available. Field collecting should continue, especially in Colombia
and southern Bolivia, which are the least explored areas. Wild T. tuberosum and
related wild species have received only limited attention so far. All species of
section Mucronata of Tropaeolum tuberosum and the closely related T. umbellatum
should be the first targets. T. umbellatum, with tubers reputedly reaching
3–4 pounds per plant (Sparre and Andersson 1991), was last collected in its native
Ecuador in the nineteenth century, and may be on the brink of extinction, if not
extinct already. Section Chilensia of Tropaeolum, with most members producing
tubers, also deserves attention.
• In spite of several chromosomal studies, mashua cytology still requires
clarification. A comprehensive chromosomal study is necessary, including a
range of accessions of both subspecies along the whole latitudinal gradient.
Chromosomal studies should be combined with molecular analysis of the
different mashua morphotypes in order to understand their relationships. This
analysis should be extended to other relevant wild species.
• The taxonomic revision of the genus by Sparre and Andersson (1991) using
conventional tools is sound and comprehensive. However, numerical and
molecular analysis would certainly add a quantitative dimension to the
relationships between the different species and help to identify ‘wild’ mashua
types as members of subspecies silvestre or as escapes from culture.
• A separate aspect would be the development of mashua as an ornamental. It is
an exceptionally beautiful species. Nasturtium (T. majus) and Tropaeolum hybrids
42 Mashua (Tropaeolum tuberosum Ruíz & Pav.)
already have a place as ornamentals worldwide. Mashua would add the
advantages of asexual propagation, as begonias and lilies do, to nasturtium
beauty. Beckett (1979) describes its use in Britain. A place in the home garden, as
an ornamental edible plant may be a way to introduce mashua to a wider public.
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Research contacts
Argentina
Dr Alfredo Grau Taxonomy, utilization, 
Universidad Nacional de Tucumán ethnobotany
CC 34
4107 Yerba Buena,
Tucumán, Argentina
Tel 54 81 304957
Fax 54 81 254468
Email graua@tucbbs.com.ar
www.unt.edu.ar
Bolivia
Mr Augusto Guidi Feed uses of mashua
Promoción e Investigación de Productos 
Andinos (PROINPA)
Av. Blanco Galindo, Km 12.5
Calle C. Prado s/n (Quillacollo)
Cochabamba, Bolivia
Tel 591 4 4360800
Fax 591 4 4360802
Email aguidi@proinpa.org www.proinpa.org
Ms María L. Ugarte Ex situ conservation, 
Promoción e Investigación de Productos morphological 
Andinos (PROINPA) characterization
Av. Blanco Galindo, Km 12.5
Calle C. Prado s/n (Quillacollo)
Cochabamba, Bolivia
Tel 591 4 4360800
Fax 591 4 4360802
Email mugarte@proinpa.org
www.proinpa.org
Promoting the conservation and use of underutilized and neglected crops. 25. 51
Colombia
Dr Mario Lobo Ex situ conservation
Corporación Colombiana de Investigación 
Agropecuaria (CORPOICA)
Km. 7 Vía Las Palmas
Río Negro Antioquia, Vereda Llano Grande, 
Colombia
Tel 57 94 5371133
Fax 57 94 5370146
Email mlobo@corpoica.org.co, pnrgv@epm.net.co
www.corpoica.org.co
Ecuador
Dr Carlos Nieto Agronomy, cropping 
Proyecto de Desarrollo de los Pueblos systems, utilization
Indígenas y Negros del Ecuador (PRODEPINE)
Bosmediano E 11–50 y
Carlos Guerrero
Quito, Ecuador
Tel 593 2 2464900
Fax 593 2 2922453
Email: carlosn@prodepine.org
www.prodepine.org
Mr César Tapia Germplasm conservation,
Instituto Nacional de Investigaciones tissue culture, molecular 
Agropecuarias (INIAP) characterization
Panamericana Sur, Km 14
Estación Experimental Santa Catalina
Quito, Ecuador
Tel 593 22 693359
Fax 593 22 693359
Email: denaref@ecnet.ec
www.denaref.org
52 Mashua (Tropaeolum tuberosum Ruíz & Pav.)
Peru
Dr Teresa Ames de Icochea Pests and diseases
Centro Internacional de la Papa
Av. La Molina 1895,
Lima 12, Perú
Tel 51 1 3496017
Fax 51 1 3175326
Email: t.ames@cgiar.org
www.cipotato.org
Dr Carlos Arbizu Ethnobotany, germplasm 
Centro Internacional de la Papa characterization
Av. La Molina 1895,
Lima 12, Perú
Tel 51 1 3496017
Fax 51 1 3175326
Email: c.arbizu@cgiar.org
www.cipotato.org
Prof Rolando Estrada Tissue culture
Instituto Nacional de Investigación 
Agraria (INIA)
Av. La Molina 1981,
Lima 12, Perú
Tel 51 1 3496131
Fax 51 1 3496131
Email restradaj@unmsm.edu.pe, 
restrada@fenix.inia.gob.pe
www.inia.gob.pe
Dr Michael Hermann Agronomy, ethnobotany, 
Centro Internacional de la Papa post-harvest utilization
Av. La Molina 1895,
Lima 12, Perú
Tel 51 1 3496017
Fax 51 1 3175326
Email: m.hermann@cgiar.org
www.cipotato.org
Promoting the conservation and use of underutilized and neglected crops. 25. 53
Prof Ramiro Ortega Ethnobotany, on-farm  
Centro Regional de Investigación en germplasm management,
Biodiversidad Andina (CRIBA) ex situ germplasm 
Centro Agronómico K’ayra-UNSAAC conservation, wild
Distrito San Jerónimo s/n Tropaeolum species
Cusco, Perú
Tel 51 84 277323
Fax 51 84 221632
Email criba@terra.com.pe
Prof Juan Seminario Ethnobotany, post-harvest 
Universidad Nacional de Cajamarca utilization
Carretera Baños del Inca, Km 3.5
Oficina 2C-111
Cajamarca, Perú
Tel 51 44 824938
Fax 51 44 823263
Email seminari@terra.com.pe
www.unc.edu.pe
Mr Mauro Vallenas Agronomy, reproductive 
Acción Agraria biology
Av. Raymondi 161-C,
Sandia, Juliaca,
Puno, Perú
Tel 51 54 320327
Email sandia@hotmail.com
Institutions maintaining germplasm collections
Bolivia
Promoción e Investigación de Productos Andinos (PROINPA)
Av. Blanco Galindo, Km 12.5
Calle C. Prado s/n (Quillacollo)
Cochabamba, Bolivia
Tel 591 4 4360800
Fax 591 4 4360802
Email xcadima@proinpa.org
www.proinpa.org
Curator: Ms Ximena Cadima
Ecuador
Instituto Nacional de Investigaciones Agropecuarias (INIAP)
Panamericana Sur, Km14
Estación Experimental Santa Catalina
Quito, Ecuador
Tel 593 22 693359
Fax 593 22 693359
Email: denaref@ecnet.ec
www.denaref.org
Curator: Mr César Tapia
Peru
Instituto Nacional de Investigación Agraria (INIA)
Av. La Molina 1981
Lima 12, Perú
Tel 51 1 3482703
Fax 51 1 3495646
Email pnirgb@fenix.inia.gob.pe
www.inia.gob.pe
Curator: Mr Santiago Pastor
Centro Regional de Investigación en Biodiversidad Andina (CRIBA)
Centro Agronómico K’ayra-UNSAAC
Distrito San Jerónimo s/n
Cusco, Perú
Tel 51 84 277323
Fax 51 84 221632
Email criba@terra.com.pe
Curator: Prof Ramiro Ortega
Promoting the conservation and use of underutilized and neglected crops. 25. 55
Centro Internacional de la Papa (CIP)
Av. La Molina 1895,
Lima 12, Perú
Tel 51 1 3496017
Fax 51 1 3175326
Email c.arbizu@cgiar.org
www.cipotato.org.pe
Curator: Dr. Carlos Arbizu
ISBN 92-9043-581-X
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