Re-Introduction of Quinoa Into Arid Chile

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D R O U G H T S T R E S S

Re-Introduction of Quı noa into Arid Chile: Cultivation ofTwo Lowland Races under Extremely Low IrrigationE. A. Martınez1, E. Veas1, C. Jorquera2, R. San Martın3 & P. Jara4

1 Centro de Estudios Avanzados en Zonas A ridas, Universidad de La Serena, La Serena, Chile

2 Departamento de Agronomı a, Facultad de Ciencias, Universidad de La Serena, La Serena, Chile

3 Facultad de Ingenierı a, Pontificia Universidad Cato lica de Chile, Santiago, Chile

4 A lvaro Casanova 1215, Pen ˜ alole n, Santiago, Chile

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 Lopez

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

market opportunities. Consequently, change of land use in

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 quınoa (Chenopodium quinoa) a

drought stress tolerant species that produces a high quality

grain for human consumption.

Keywords

Chenopodiaceae; Chenopodium quinoa;

climate; desertification; drought tolerance;

quinoa; water use efficiency; yield

CorrespondenceE. A. Martı nez

Centro de Estudios Avanzados en Zonas

A ridas, Universidad de La Serena, La Serena,

Chile

Tel.: +56 51 204378

Fax: +56 51 334741

Email: [email protected]

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 quınoa 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.

J. Agronomy & Crop Science (2009) ISSN 0931-2250

ª 2008 The Authors

Journal compilation ª 2008 Blackwell Verlag, 195 (2009) 1–10 1



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 (Nunez 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

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 wherehumidity 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 (Martınez 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 differencesbetween 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 Martın 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

Fig. 1 Running means for 30 years of rainfall data at La Serena city

(30S, 71W) between 1869 and 2003 (updated from Novoa and

Lo pez 2001).

Martı nez et al.

ª 2008 The Authors

2 Journal compilation ª 2008 Blackwell Verlag, 195 (2009) 1–10



(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 andseasons 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

Vicuna, 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 choicewas made.

Saponin content

Saponins were extracted from the original seeds of both

landraces and from those harvested in Coquimbo in 2006

(one sample per landrace and harvest season was used

because seeds were not separated from individual plants).

Seeds for extraction (30 g) were thoroughly grounded

and then extracted with water at 60 C for 3 h. The ratio

of water to seeds was 15 to 1 (by weight). The extract

was centrifuged and the supernatant filtered (pore size

0.45 lm) and then analysed by reverse phase-high

pressure liquid chromatography (RP-HPLC) following the

protocol described by San Martın and Briones (2000).

The standard (saponin from Quinoa Real) contained

approximately 80 % w/w saponins, and it was prepared

as described by San Martın et al. (2007). Saponin yields

were reported as % w/w of dry weight of seeds and their

contents were compared using Mann–Whitney U -test for

small samples.

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 abroader 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

Coquimbo Ovalle

Salinity (units)

pH 7.5 8.1

Electric conductivity (dS m)1) 3.9 1.1

Fertility (units)

pH suspension in H2O (1 : 2.5) 7.3 8.0

Organic matter (%) 1.3 3.2

Total nitrogen (%) 0.09 0.17

C/N ratio 8 11

Available nitrogen (ppm) 115 80

Available phosphorus (ppm) 28 21

Available potassium (ppm) 172 395

Texture (units)

Clay (<0.002 mm) (%) 22 49

Silt (0.002–0.05 mm) (%) 25 22

Sand (0.05–2.00 mm) (%) 53 29

Texture class Sandy-clay-loam Clay

Response of Quinoa to Drought Stress in Arid Chile

ª 2008 The Authors




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

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

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.

(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 highand low treatments.

Martı nez et al.

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

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 temperaturein 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. 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).

(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).


ª 2008 The Authors




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-

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 (Garcıa 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).

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

Martı nez et al.

ª 2008 The Authors




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-

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 beoccurring 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 (Quilimarı valley), cultivated

a lowland quınoa 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 theseextreme 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 (Martınez 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

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).

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.


ª 2008 The Authors




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

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 thoseobtained experimentally in Chillan (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

Fig. 9 Normal root development is shown where the root grows

longer downwards than sidewards.

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 Quilimarı valley (Region of Coquimbo).

Martı nez et al.

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 onecological 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). Addingcomposted 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

kindly updated by Dr F. Squeo. Help during hand-sowing

and harvesting was provided by Olga and Manuel Martı-

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 Jose Garcıa a small-scale farmer of the Quilimarı

Valley. Aid in figures art was kindly provided by Manuel

Martınez. Two anonymous reviewers and Dr Andres

Zurita made very valuable comments and suggestions to

the text. Many thanks to all of them.

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Re-Introduction of Quinoa Into Arid Chile

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