J.

Agronomy & Crop Science (2009) ISSN 0931-2250

DROUGHT STRESS

Re-Introduction of Quı́noa into Arid Chile: Cultivation of

Two Lowland Races under Extremely Low Irrigation
E. A. Martı́nez1, E. Veas1, C. Jorquera2, R. San Martı́n3 & P. Jara4
1 Centro de Estudios Avanzados en Zonas Áridas, Universidad de La Serena, La Serena, Chile
2 Departamento de Agronomı́a, Facultad de Ciencias, Universidad de La Serena, La Serena, Chile
3 Facultad de Ingenierı́a, Pontificia Universidad Católica de Chile, Santiago, Chile
4 Álvaro Casanova 1215, Peñalolén, Santiago, Chile

Keywords
Chenopodiaceae; Chenopodium quinoa;
climate; desertification; drought tolerance;
quinoa; water use efficiency; yield
Correspondence
E. A. Martı́nez
Centro de Estudios Avanzados en Zonas
Áridas, Universidad de La Serena, La Serena,
Chile
Tel.: +56 51 204378
Fax: +56 51 334741
Email: enrique.martinez@ceaza.cl
Accepted July 23, 2008
doi:10.1111/j.1439-037X.2008.00332.x
Abstract
Annual rainfall in Chile at 30S decreased from 170 to 70 mm in the last cen-
tury, forcing a search for new low-rain adapted crops. Chenopodium quinoa
Willd. was cultivated by pre-Hispanic cultures, but it disappeared in this region
since the Spanish conquest. Two quı́noa landraces (Don Javi and Palmilla)
were re-introduced from lowlands of central Chile (34S) evaluating seed sapo-
nin content and grain yields under low irrigation. Replicated assays were con-
ducted in two sites with distinct microclimates after august (end of the rains in
2004 and 2005). Treatments included low (40–75 mm) and high (150–
250 mm) irrigation and were distributed along the five cultivation months.
Fertilization, with the humus of the worms, was carried out in the second sea-
son, as soils are poor in organic matter. Results showed significantly higher
saponin content in the seeds of Don Javi landrace (1.2 %) with respect to
Palmilla seeds (0.3 %). However, grain yields were not different between land-
races under the same treatments. Yields were instead affected by microclimate,
irrigation and fertilization. Although higher yields corresponded with higher
irrigation, 2.6 tons ha)1 was obtained under high irrigation, but surprisingly,
also under low irrigation in the more humid site. Yields of 2006 harvesting sea-
son (ca. 7 tons ha)1) were higher than that of the previous season (ca. 5.5 tons
ha)1), mainly because of the addition of organic matter. We suggest that
re-introduction of Quinoa in arid Chile is feasible even under the prevailing
conditions of low rainfall and deficient soils, but better yields will need some
irrigation and addition of organic matter.

market opportunities. Consequently, change of land use in

Introduction
Average rainfall at La Serena (30 latitude S), centre of the
Coquimbo Region in Chile has decreased from 170 to
70 mm year)1 in the last century (Fig. 1, Novoa and López
2001). Data from different studies (Caldentey 1987, Novoa
and Villaseca 1989) indicate that this zone must be consid-
ered as being under an arid climate rather than semiarid
because the ratio of precipitation to potential evapotranspi-
ration (ETo) is between 0.05 and 0.20 (Le Houerou 1996).
Agricultural practices have thus evolved in the last 30 years
towards increased water use efficiency in this region so that
cereal cultivation has been almost abandoned in favour of
high-value fruit cultivation, a trend also driven by export
the past, in a time interval of 20 years, shows a trend of
more than 300 % increase in fruit cultivation surfaces such
as vineyards and other fruit trees. This contrasts with a
strong reduction of 80–100 % of less cereal cultivation and
forested areas (Jorquera 2001). This trend implies that
watering practices will become much more expensive
because of sophisticated technologies and likely unafford-
able for small-scale landowners. The identification of alter-
native annual grain species with high tolerance to drought
stress needs to be investigated in arid Chile. One such alter-
native annual crop is quı́noa (Chenopodium quinoa) a
drought stress tolerant species that produces a high quality
grain for human consumption.

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Journal compilation ª 2008 Blackwell Verlag, 195 (2009) 1–10 1
Martı́nez et al.
Fig. 1 Running means for 30 years of rainfall data at La Serena city
(30S, 71W) between 1869 and 2003 (updated from Novoa and
López 2001).
The seeds of quinoa are high in protein (15 % for
soluble proteins in 10 Chilean landraces, data not shown)
and contain an excellent balance of the amino acids com-
prising all the essentials to the human diet (Repo-
Carrasco et al. 2001). Hence, it has been chosen by NASA
(USA) as the plant to be cultivated in artificial extra-
planetary spatial stations (Schlick and Bubenheim 1996)
and with high potential to be introduced in India as a
basic food source (Bhargava et al. 2006). This crop spe-
cies has high frost, salinity and drought resistance as well
as pest resistance (Jacobsen et al. 1998, Mujica et al.
2001). Prior to the European discovery of the Americas,
the Araucanian and Andean civilizations from southern
Chile to northern Colombia extensively cultivated quinoa.
Its cultivation dates back to 5000 years bc (Nuñez 1989),
and in Chile, quinoa has been cultivated for at least the
past 3000 years (National Research Council [NRC] 1989,
Tagle and Planella 2002). The traditional cultivation prac-
tices by these ancient cultures were changed significantly
by the introduction of new European crops at the time of
the Spanish conquest. Then, wheat and other cereal crops
replaced quinoa cultivation, particularly in lowlands of
Chile (Tagle and Planella 2002) and also in the rest of
similar ecosystems of South America, because of various
reasons including ignorance of its nutritional facts, its bit-
ter saponin content and probably an urgent need for
well-known staple food (NRC 1989). Ancient and long-
lasting agricultural practices often resulted in the produc-
tion of stress tolerant ecotypes defined as landraces (after
Zeven 1998). Genetic characterizations separate the
quinoa landraces into two broad ecotypes or categories:
Altiplano and Lowland (Mason et al. 2005). In the
high Andeans regions of Bolivia and Peru, ecotypes are
2 Journal compilation ª 2008 Blackwell Verlag, 195 (2009) 1–10
cultivated by farmers organized in cooperatives for com-
mercial matters (usually involving many villages) for
export to North America, Europe and Japan (Laguna
et al. 2006). In the Chilean Altiplano, quinoa is cultivated
above 3500 m.a.s.l. at 18–22 S. No artificial irrigation is
used as it rains slightly more than 150 mm year)1. The
seeds are buried 15–20 cm deep in the soils where
humidity for germination is higher, and freezing tempera-
tures are less harmful and competition is avoided by a
long sowing distance between plants (ca. 50 cm between
plants) (Lanino 2006). On the other side, lowland land-
races are still cultivated on small family farms in the
southern regions of Chile from elevations of 1000 m.a.s.l.
to near the sea level, practices inherited from Pehuenche’s
ancient cultures at 34–36S and from the Mapuches at
40 S (Martı́nez et al. 2007a). In the lowlands, it rains
from 400 to 2000 mm year)1. In arid Chile (30S),
quinoa cultivation and the Diaguita culture disappeared
during the Spanish colonization (500 years ago), a situa-
tion that affected other native crops in the whole Andean
region of South America (NRC 1989).
To test the potential of quinoa as an alternative crop
species for arid regions in Chile, seeds of two lowland land-
races were tested for yield responses under different irriga-
tion treatments at two arid localities with different
microclimatic conditions (sub-Andean dry vs. coastal
humid). The two landraces did not differ in appearance
but the two farmers that cultivated them had never
exchanged seeds, although they lived close to each other.
This fact probably influenced the genetic distance found
between them when using microsatellite molecular markers
(Fuentes et al. 2008). Thus, we tested possible differences
between them, for example their saponin contents. More-
over, seed coat saponins are considered attractive and not
really undesirable because of their potential use in pest
control for good agricultural practices (San Martı́n et al.
2007). In the second year (2006/2007), the effects of the
addition of organic matter on yield were also evaluated.
Materials and Methods
Study area
Two localities of arid Chile were used in this study: the
experimental field station of Instituto de Investigaciones
Agropecuarias (INIA) at Coquimbo (3003¢S, 7113¢W,
123 m a.s.l.) and the experimental field station of La Serena
University at Ovalle (3034¢S, 7111¢W, 600 m.a.s.l.).
Coquimbo is located close to the coast (123 m. a.s.l.) and
is often under cloud cover called ‘Camanchaca’ that is
used in high hills to collect fog-water (Cereceda and
Schemenauer 1991). These clouds provoke less irradiance
and higher humidity than at Ovalle location. Rainfall
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 Response of Quinoa to Drought Stress in Arid Chile

(2004–2006), temperature and humidity (September 2004

to March 2005) were recorded at both sites and the same
data were also recorded in Coquimbo between December
2005 and April 2006. ETo (August 2005 to August 2006)
was estimated by using Penman-Monteith equation (Allen
1986). Afterwards, maximum, minimum and average tem-
peratures and humidity were compared between sites and
seasons using anova (systat Software Inc., San Jose, CA,
USA). Soil fertility and salinity were evaluated by samples
analysed by standard methods (at AGROLAB A.S.).
Landraces origin
Quinoa seeds were obtained from farmers of two locali-
ties in central Chile (3434¢S, 7148¢W), both located
close to the sea level near the estuary of the Nilahue
River. The first landrace, named ‘Don Javi’ was collected
in a farmer field at 3431¢40.2¢S, 7156¢46.1¢W. The sec-
ond landrace, named ‘Palmilla’ was collected from a
farmers field at approximately 25 km from the first land-
race (3432¢08.7¢S, 7157¢24.7¢W). Seeds from both land-
races were collected in the summer of 2004 and they are
stored as part of the National Seed Bank collection at
Vicuña, Chile (INIA-Intihuasi). These two landraces did
not differ in morphology, but they were chosen because
the available information indicated that they were histori-
cally cultivated without seed exchange between farmers
(strong isolation) and, in good agreement with that, they
were genetically distinct as revealed by the genetic dis-
tance evaluated using 20 microsatellite loci (see Palmilla
as Palmix and Don Javi as Javi accessions in Fuentes et al.
2008). The saponin content was ignored when the choice
was made.
Saponin content
Field studies
Because of logistic limitations, the essays of Coquimbo
and Ovalle are not identical in irrigation conditions.
Besides, the essays of 2004/2005 were repeated only at
Coquimbo the next season (2005/2006). Some modifica-
tions were made during the second season to evaluate a
broader spectrum of irrigation conditions.
Experiments of 2004/2005 season
Six 9 m2 interspersed parcels were established randomly
along three lines for each landrace and irrigation
condition in October 2004 at Coquimbo and Ovalle. Seed
density was equivalent to 3 kg ha)1 and they were hand-
sown in four lines per parcel at 1–2 cm depth. Soils at
both locations are described in Table 1. They were soft-
ened by using a harrow. No additional fertilization was
carried out during the season 2004/2005.
Watering conditions
Irrigation was applied at Ovalle by drip lines releasing
4 l m)1 h)1. There were four drip lines of 3 m in length
each, totalizing 12 linear metres in each parcel. This gives
48 l h)1 per parcel. Watering time during the irrigation
days lasted between 2.5 and 4 h day)1. At Coquimbo,
furrows with a flow of 2 l s)1 irrigated parcels of the
same size. This is equivalent to 0.12 m3 min)1 and not
more than 15 min of watering was given to each parcel of
60 m2. Figure 2 depicts the cumulated irrigation data.
Table 1 Soil characterization (salinity, fertility and texture) of the
experimental plots at Ovalle and Coquimbo

Saponins were extracted from the original seeds of both Coquimbo Ovalle
landraces and from those harvested in Coquimbo in 2006 Salinity (units)
(one sample per landrace and harvest season was used pH 7.5 8.1
because seeds were not separated from individual plants). Electric conductivity (dS m)1) 3.9 1.1
Seeds for extraction (30 g) were thoroughly grounded
Fertility (units)
and then extracted with water at 60 C for 3 h. The ratio pH suspension in H2O (1 : 2.5) 7.3 8.0
of water to seeds was 15 to 1 (by weight). The extract Organic matter (%) 1.3 3.2
was centrifuged and the supernatant filtered (pore size Total nitrogen (%) 0.09 0.17
0.45 lm) and then analysed by reverse phase-high C/N ratio 8 11
pressure liquid chromatography (RP-HPLC) following the Available nitrogen (ppm) 115 80
Available phosphorus (ppm) 28 21
protocol described by San Martı́n and Briones (2000).
Available potassium (ppm) 172 395
The standard (saponin from Quinoa Real) contained
approximately 80 % w/w saponins, and it was prepared Texture (units)
as described by San Martı́n et al. (2007). Saponin yields Clay (<0.002 mm) (%) 22 49
Silt (0.002–0.05 mm) (%) 25 22
were reported as % w/w of dry weight of seeds and their
Sand (0.05–2.00 mm) (%) 53 29
contents were compared using Mann–Whitney U-test for
Texture class Sandy-clay-loam Clay
small samples.

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Journal compilation ª 2008 Blackwell Verlag, 195 (2009) 1–10 3
Martı́nez et al.
(a)
(b)
Fig. 2 Cumulated watering applied for each treatment, high (closed
circles) and low (open circles). Irrigation in season 2004/05 (a) at
Coquimbo (solid and dotted lines) and Ovalle (dashed line and dot-
dashed lines). Irrigation at Coquimbo in 2005/06 (b) shows only high
and low treatments.
Experiments of 2005/2006 season
During the 2005/2006 season, the study was repeated only
in Coquimbo with five repetitions for each landrace and
watering condition. Plants were sown in November, and
the seed density was increased to 4 kg ha)1. The ran-
domly interspersed parcels of 5 m2 each were fertilized on
the seeding lines only with humus of Eisenia phoetida
worms, rich in organic matter (nutritional description in
Table 2), at a dose of 20 tons ha)1. Weed control was
managed by hand removal in both the seasons.
Watering conditions
Irrigation was applied in the second season, in
Coquimbo, also by drips (1.5–4 h of watering/day releas-
ing 4 l linear-m)1 h)1). However, as the parcels were
smaller, each parcel had only four linear metres of drips,
totalling 16 l h)1 per parcel. Watering time during the
4 Journal compilation ª 2008 Blackwell Verlag, 195 (2009) 1–10
Table 2 Humus characterization from Eisenia phoetida worms used
in this study
Nitrogen (N) % 0.75
Phosphorus (P2O5) % 0.53
Potassium (K2O) % 0.30
pH suspension 1:5 8.70
Conductivity mho cm)1 2.60
Organic matter % 18.70
Organic Carbon % 10.90
Ratio C/N 14.50
Humidity % 27.50
Total humic extract % 3.6
Percentages based on dry matter.
Food source for worms: Opuntia ficus indica branches.
irrigation days lasted between 2.5 and 4 h day)1, and the
cumulated data are depicted in Fig. 2. The first watering
period was always aimed to obtain soil with 3/4th of field
capacity, which is equivalent to a rainfall of 10 mm. A
tensiometer was placed with a bulb at 50 cm depth. The
water table does not reach the surface levels in both the
localities, being always below 5 m at least (after well level
observations by Dr F. Squeo, personal communication).
The described two extreme watering conditions, applied
in each site and season, simulated extreme precipitation
regimes, one of normal irrigation mimicking an accumu-
lated precipitation of above 200 mm/cultivation period
(high irrigation) and the other of more stressing condi-
tion of <60 mm/period (low irrigation). Exact amounts of
applied water were estimated by flow and watering time
measurements in each locality and season, trying to adjust
the applied water to the two mentioned extreme condi-
tions. Such conditions were relaxed during the second
season by slightly increasing the low irrigation to 75 mm/
period, but on the other side they were also hardened by
reducing the high irrigation condition to 150 mm/period.
This 150 mm is the lowest annual rainfall reported for
not irrigated quinoa cultivation lands in Chile (Altiplano)
(Lanino 2006). The experiments were started after the
rainy seasons ended (Fig. 3a) so that all the available water
in the soil was provided by the irrigation treatments.
During the driest season of 2004/2005, an extra irrigation
period had to be applied because of evident signs of stress
(leaf bending) at Ovalle. This was in December 2004
(Fig. 2a) and at this date tensiometers with a bulb located
at 50 cm depth, lost negative pressure at )40 kPa because
of the disconnection of bulb–soil interphase.
Harvesting and yield evaluations
Seeds were hand-harvested at maturity (14 % humidity
obtained in late February 2005 and late March in 2006) and
quinoa grain yields were expressed as tons ha)1. Significant
ª 2008 The Authors

differences among landraces, treatments and localities were
tested using Tukey a posteriori test after one-way anova
analysis (systat software). When yields between the two
landraces were found to be not significantly different, they
were pooled to have higher statistical power, particularly to
test differences between localities or microclimate.
Results
Microclimatic conditions in both localities
The two chosen localities (Coquimbo and Ovalle) showed
similar rainfall patterns (Fig. 3a) but, over the year, Ovalle
showed reference ETo that was 330 mm higher than at
Coquimbo, particularly because of the higher ETo during
the driest months between December and March 2004/
2005 (Fig. 3b). Ovalle also showed significantly higher
maximum temperatures (F8,1806 = 732.165, P < 0.001) and
(a)
(b)
Fig. 3 Monthly rainfall between 2004 and 2006 (a) and reference
evapotranspiration (ETo) between August 2005 and August 2006 (b)
in Coquimbo (black bars) and Ovalle (shaded bars).
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Journal compilation ª 2008 Blackwell Verlag, 195 (2009) 1–10
Response of Quinoa to Drought Stress in Arid Chile
lower daily relative humidity during the experimental culti-
vation period of 2004/2005 (F2,602 = 14.884, P < 0.001).
Coquimbo was then significantly colder (i.e. lower daily
maximum temperature and daily means) and it was also
significantly more humid than Ovalle (Fig. 4a and b).
Coquimbo did not show differences between both
seasons, except for a slightly higher maximum temperature
in 2005–2006 (Tukey a posteriori test, P < 0.05), but
relative humidity was not significantly different (Tukey
a posteriori test, P < 0.05) between both seasons (Fig. 4b).
Saponin content
Saponin content in seeds of different harvests from the
‘Don Javi’ landrace varied between 1.20 % and 1.56 %
w/w, while in the seeds of ‘Palmilla’ landrace saponin
contents were four times lower varying between 0.20 % and
0.46 % (Table 3). The observed contents were significantly
(a)
(b)
Fig. 4 Mean annual air temperature (a) and humidity (b) at Ovalle
and Coquimbo. Vertical lines correspond to 95 % confidence limits.
Significant differences are shown by different letters (P < 0.05, Tukey
a posteriori test).
5
Martı́nez et al.
different (Mann–Whitney U-test = 9.0, P = 0.05) in spite
of the small sample size. Lack of between plant replication
prevents between-harvests or between-treatments compar-
isons within landraces.
Yield responses
When comparing yields of both landraces, although ‘Don
Javi’ seemed to have yielded lower grain weight than
‘Palmilla’ in the locality of Ovalle (harvest 2005). No signif-
icant differences were found between landraces when they
were cultivated under the same conditions in either
growing season (Figs 5 and 6). Significant differences in
yields were only found when comparing different watering
treatments and different environments (F7,40 = 47.559,
P < 0.001). In the 2005 harvest, yields ranged from
4.8 tons ha)1 (the more humid condition at Coquimbo,
with higher watering) to 1.3 tons ha)1 at the drier Ovalle
condition with lower irrigation (Fig. 5). Interestingly,
yields were not significantly different (P > 0.05) when
compared between the more humid Coquimbo site with
low irrigation (40 mm in the 5-month period) and the
yields obtained at the drier Ovalle site supplemented with
an extensive irrigation (205 mm/period; Fig. 5).
In the 2005/2006 harvest, yields averaged between
4.0 tons ha)1 (Palmilla-low irrigation) and 7.7 tons ha)1
(Don Javi-high irrigation), but these yields were not sig-
Table 3 Seed saponin contents of the initial material (harvest 2004)
of Don Javi and Palmilla landraces (% of dry weight) and those in the
harvest of Coquimbo (2006) with high and low watering conditions
Harvest season and watering treatment Don Javi Palmilla
2004 1.20 0.46
2006 – high irrigation 1.10 0.34
2006 – low irrigation 1.50 0.20
6 Journal compilation ª 2008 Blackwell Verlag, 195 (2009) 1–10
nificantly different (F3,16 = 2.871, Tukey a posteriori test,
P > 0.05, Fig. 6).
When comparing yields between seasons (2006 vs.
2007) within the same locality (Coquimbo) and for the
same treatment (high irrigation), the highest yield
(7.7 tons ha)1) was obtained for the ‘Don Javi’ landrace,
fertilized with humus with an irrigation of 150 mm/per-
iod. The lowest yield was also obtained for ‘Don Javi’
landrace (4.9 tons ha)1) even if irrigation was higher
(250 mm/period). To see more clearly the locality and
irrigation effects, yield values of both landraces were
pooled. They were not significantly different within the
same treatments. Pooled data showed more clearly that
higher watering does not always coincide with higher
yields in quinoa lowland landraces. Although comparisons
between different years are not so prone to be directly
interpreted, it is surprising that at Coquimbo, a combina-
tion of lower watering (150 mm/period) with addition of
worm humus (rich in organic matter) produced signifi-
cantly higher yields in 2006 (F5,62 = 23.702, Tukey a pos-
teriori test P < 0.001) than at higher irrigation practice
(250 mm/period) as tried in the previous year (2005)
with no addition of organic matter (Fig. 7).
Discussion
This study shows for the first time in arid Chile that low-
land landraces of Quinoa can grow and produce interesting
experimental yields, even when mimicking the dominant
conditions of low local rainfall through extremely reduced
irrigation (<75 mm year)1). Our irrigation treatments
were performed after the rainy seasons and the amount of
applied water at high irrigation was even lower than those
reported in other field experiments, at the Altiplano, where
the applied water (irrigation plus rain) reached over
300 mm during the cultivation period (Garcı́a et al. 2003).
Fig. 5 Mean grain yields obtained in 2005
with two quinoa landraces (J: Don Javi,
P: Palmilla) for high (205/250 mm) and low
(50/40 mm) watering treatments at Ovalle
and Coquimbo. Vertical lines correspond to
95 % confidence limits. Significant differences
are shown by different letters (P < 0.05,
Tukey a posteriori test).
ª 2008 The Authors

Fig. 6 Mean grain yields obtained in 2006 with two quinoa landraces
(J: Don Javi, P: Palmilla) for high (150 mm) and low (75 mm) watering
treatments at Coquimbo. Vertical lines correspond to 95 % confi-
dence limits.
Fig. 7 Mean yields of grain production (tons ha)1) for the harvest of
2005 and 2006 obtained by pooling the non-significantly different
data of Fig. 5, i.e. every pair of the two tested Quinoa landraces (Don
Javi and Palmilla) for each watering condition. All significant differ-
ences correspond to the differences between high and low watering
treatments, and between localities. Vertical lines correspond to 95 %
confidence limits. Different letters indicate significantly different yields
(a posteriori Tukey test, P < 0.05).
Besides, our conditions were also extreme regarding ETo
(20 times higher than precipitation) and poor soil fertility
(low nitrogen and organic matter contents). We do not
fully understand how quinoa plants can reach such high
yields under these extreme environmental conditions. Esti-
mations of low available soil water and high ETo rates do
not seem to allow plant growth nor the high observed
yields. However, three physiological/anatomical mecha-
nisms can be proposed as possible alternative or comple-
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Journal compilation ª 2008 Blackwell Verlag, 195 (2009) 1–10
Response of Quinoa to Drought Stress in Arid Chile
mentary explanations. First, evidences from other plants
indicate that mucopolysaccharides located in the plant
xylem not only help to decrease ETo without affecting pho-
tosynthesis but also provoke a phenomenon called reverse
transpiration, providing water to the plant xylem through
the leaves (Zimmermann et al. 2007). Although, the evi-
dences are described for trees, something similar could be
occurring in quinoa plants too. Secondly, it has been
observed that quinoa stomata do not seem to respond to
abscisic acid (ABA), except under extremely high drought
and it has been observed that quinoa plants can perform
photosynthesis for a long period under extreme low irriga-
tion, even for 3 days after stomata are closed (Jacobsen
et al. 2007). This phenomenon can be hypothetically
explained by a mechanism where carbon dioxide is taken
by open stomata and stored as oxalic acid. This molecule is
present in succulent and in non-succulent plants such as
quinoa (Bown 1995). Then, when stomata are closed,
oxalic acid is reconverted to carbon dioxide for photo-
synthesis, allowing strong water use efficiency, a phenome-
non occurring in many plant species (Sen et al. 1971).
Furthermore, a third alternative and/or complementary
explanation is postulated derived from an interesting recent
field observation on quinoa root morphology. At the end
of spring season of 2007, a local farmer located close to the
zone influenced by coastal fog (Quilimarı́ valley), cultivated
a lowland quı́noa landrace of the same zone rather than
‘Don Javi’ and ‘Palmilla’. His parcel had no irrigation at all
and the sowing was performed after the last rain of 2007
and the harvesting was carried out in February 2008. Plants
grew of very small sizes (<50 cm height) and roots seemed
to modify their normal growth pattern. That is, under these
extreme drought, quinoa pivotal roots grew to a very short
distance in depth, but one lateral root grew a much longer
distance, shallow and parallel to the surface (Fig. 8). At this
depth of 2–5 cm soils have probably more water availability
coming from sources like morning dew or from dense fog
during foggy days. In fact, at Coquimbo, the cloudy days
produced by the fog called ‘Camanchaca’ (Cereceda and
Schemenauer 1991) are particularly common during the
drier summer months, reaching more than 40 % of the
summer days (Martı́nez et al. 2007b). This common coastal
fog comes inland from the Pacific Ocean because of the
strong heat of the summer. Under conditions of more uni-
form soil humidity roots seem to develop in a more normal
growth pattern, with a deeper root axis (Fig. 9). Thus, the
fog that seems to be an effective cause of the observed
reduced ETo may also be a source of available water, for
the reverse transpiration of the leaves and also for modified
superficial roots. The reverse metabolism of oxalic acid and
carbon dioxide may in turn allow highly efficient mecha-
nisms for the carbon and water budget of quinoa plants,
this hypothesis needs to be further tested. In fact, root
7
Martı́nez et al.
Fig. 8 Root morphology of small plants with longer horizontal than
deep roots, grown with no irrigation, sown after the last rain of 2007
at Quilimarı́ valley (Region of Coquimbo).
Fig. 9 Normal root development is shown where the root grows
longer downwards than sidewards.
morphology is crucial for water uptake as has been previ-
ously suggested, particularly under local salty soils (Schleiff
2008). At Ovalle, the environmental conditions were much
drier. However, water retention could have been slightly
increased because of the three times higher content of
8 Journal compilation ª 2008 Blackwell Verlag, 195 (2009) 1–10
organic matter and a less sandy soil. Also the water use
efficiency of quinoa must have been facilitated at Ovalle by
the observed three times lower electric conductivity and
lower sodium content in these soils.
The observed saponin content was also different between
landraces and stable within the same order of magnitude
between the harvests. However, we cannot provide conclu-
sions about the effects of the irrigation conditions on
saponin content. The observed agronomic yields between
both landraces were not different when submitted to
similar treatments. This behaviour may allow genetic
improvement, for instance, with respect to saponin
content, with no apparent decrease on the expected yields.
According to our results, quinoa cultivation without
artificial irrigation, as it is normally cultivated in the rest
of the country, is theoretically possible in this arid region
of Chile even for the low annual rainfall records observed
during the last 50 years (50–70 mm). However, early sow-
ing is recommended during winter to receive a maximum
of the annual rainfall. Additional tests of frost resistance
will be needed in those zones where freezing conditions
are common. However, rains in the coastal region are not
only scarce but also highly unpredictable. Then, it is
highly recommended to ensure water availability for all
critical phenological stages (panicle formation, flowering
and grain filling); therefore, it will be needed to invest in
artificial irrigation, a decision that will inevitably increase
farmer’s production costs.
The observed experimental yields are similar to those
displayed by other landraces of central Chile, previously
tested in experimental conditions in more rainy sites of
southern Chile. For instance, our yields are similar to those
obtained experimentally in Chillán (36–37S), where the
highest yield reached 3.8 tons ha)1 (Berti et al. 1998,
2000). With respect to other previously reported yields, our
maximum values were similar to those obtained in north-
ern Chile, i.e. 4–9 tons ha)1 (Delatorre et al. 1995, 2001).
However, when comparing with real farmer’s yields, our
experimental results, which were obtained in small parcels
(5–10 m2) often seem to be much higher. The obtained
yields by small-scale farmers in central Chile reached
2–3 tons ha)1 at the most, where lowland landraces are also
cultivated (R. Valdebenito, personal communication).
Thus, direct extrapolations of our experimental yields
should not be expected for commercial larger farms.
Besides, our parcels did not have borderlines with shading
plants to imitate more real large-scale farming conditions.
However, one farmer in our region obtained, in his 2007
harvest, a yield of 5 tons ha)1 in one-third of a hectare after
high irrigation and nitrogen fertilization. When water is
provided by artificial waterways, a normal practice in arid
and semiarid regions of Chile, a strong invasion of weedy
species is observed. This invasion is avoided when water
ª 2008 The Authors

comes from underground sources. Weed control certainly
will increase costs to small farmers in our region. Up to
now, no herbicides have been proven to be effective in weed
control for quinoa as many herbicides are often produced
to kill its close Chenopodiaceae relatives, eliminating
quinoa plants as well. This condition should be considered
as an invitation and a challenge for more research on
ecological weed control in quinoa cultivation such as weed
incorporation to soils before they release seeds; a manage-
ment suggested for the season previous to plant sowing.
The more humid conditions and lower ETo rates found
in coastal areas of arid Chile, like Coquimbo site in this
study, favour the re-introduction of quinoa cultivation in
this region. This plant with its associated agricultural prac-
tices have been locally absent for almost 500 years. For
other drier localities, more drought stress tolerant geno-
types might be needed so that other landraces should be
tested under extreme conditions similar to those shown in
this study. Finally, an interesting result to remark is that
the addition of organic matter (worm humus with 18 % of
organic matter and low nitrogen content) during the sea-
son 2005/2006 increased quinoa yields by 50 % even under
conditions of lower irrigation. Such increase was higher
than expected with respect to the higher seed density used
in that season (sowing density in the second year was only
25 % higher than in the previous season). Adding organic
matter, particularly compost, seems very effective on soils
under water deficit. For instance adding organic matter
seems much more effective in improving the plant growth
than the higher water retention provided by the application
of mulch, as it has been recently demonstrated in wheat
submitted to drought stress (Eneji et al. 2008). Adding
composted organic matter is thus strongly recommended,
particularly for the clayish soils of our arid region. Obtain-
ing such organic matter can be facilitated from urban recy-
cling of organic matter and also from wastes derived from
rural sites, even from other agricultural by-products. The
costs of obtaining organic matter from such sources, seem-
ingly high now-a-days, might be justified in the near future
if both higher yields and higher water use efficiency are
combined. Sustainable soil management based on the
incorporation of organic matter will produce higher yields
and this will allow easier integration of such practices by
local farmers, to see the benefits of adding organic matter
to their historically arid and poor soils.
Acknowledgements
Funding was provided by grants Innova Chile CORFO
04CR9PAD04 and Fondecyt 1060281. Meteorological data
were kindly provided by CEAZA-MET network of meteo-
rological stations (Coquimbo) and by Agronomic high
school Liceo Tadeo Parribarnes (Ovalle). Figure 1 was
ª 2008 The Authors
Journal compilation ª 2008 Blackwell Verlag, 195 (2009) 1–10
Response of Quinoa to Drought Stress in Arid Chile
kindly updated by Dr F. Squeo. Help during hand-sowing
and harvesting was provided by Olga and Manuel Martı́-
nez, Nacho and Lela Quiroz. Worm humus was kindly
provided by Mr Ricardo Walsen (Walmaster). Advices for
sowing and agricultural practices in Ovalle by Pedro
Astorga are also appreciated. Field facilities provided
by the Instituto de Investigaciones Agropecuarias (INIA-
Intihuasi at Coquimbo) and Universidad de La Serena
(at Ovalle), are also deeply appreciated. Plants with modi-
fied roots were obtained, thanks to the tenacity and effort
of José Garcı́a a small-scale farmer of the Quilimarı́
Valley. Aid in figures art was kindly provided by Manuel
Martı́nez. Two anonymous reviewers and Dr Andrés
Zurita made very valuable comments and suggestions to
the text. Many thanks to all of them.
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