Growth and yield of chia (Salvia hispanica L.) in the Mediterranean and desert climates of Chile

Abstract

Chia (Salvia hispanica L.) is a species with seeds that have
high essential fatty acid content, which has encouraged
increased crop production worldwide. However, the
expansion of chia is limited because it is a photoperiodsensitive
plant adapted to areas without cold. The objective
of the present study was to determine the effect of different
climatic conditions on the growth, grain yield and oil
production of chia under irrigation in three geographic
areas of Chile: Valle de Azapa (18°30’ S lat) with a
coastal desert climate, normal desert climate in Canchones
(20°26’ S lat), and Las Cruces (33°30’ S lat) with dry
Mediterranean climate with marine influence, and two chia
phenotypes: white and dark. Results indicated that desert
conditions in the Valle de Azapa (VA) and Canchones (CH)
provided better conditions for plant growth; the highest
yield (> 2900 kg ha-1) and oil production (> 550 L ha-1).
In Las Cruces (LC), at higher latitude, low temperatures
present beginning in April coincided with the reproductive
stage, affecting yield which was no more than 129 kg ha-1;
thus this zone is not recommendable for chia cultivation.
This study also determined an 11.8 h day length threshold
for the beginning of flowering; when plants are exposed to
shorter days flower initiation is more precocious, but when
day length is not adequate plants only begin to flower when
they have accumulated 600-700 ºC d.

Full text

255
CHILEAN JOURNAL OF AGRICULTURAL RESEARCH 76(1) JANUARY-MARCH 2016
CHILEAN JOURNAL OF AGRICULTURAL RESEARCH 76(3) JULY-SEPTEMBER 2016
Chia (Salvia hispanica L.) is a species with seeds that have
high essential fatty acid content, which has encouraged
increased crop production worldwide. However, the
expansion of chia is limited because it is a photoperiod-
sensitive plant adapted to areas without cold. The objective
of the present study was to determine the effect of different
climatic conditions on the growth, grain yield and oil
production of chia under irrigation in three geographic
areas of Chile: Valle de Azapa (18°30’ S lat) with a
coastal desert climate, normal desert climate in Canchones
(20°26’ S lat), and Las Cruces (33°30’ S lat) with dry
Mediterranean climate with marine inuence, and two chia
phenotypes: white and dark. Results indicated that desert
conditions in the Valle de Azapa (VA) and Canchones (CH)
provided better conditions for plant growth; the highest
yield (> 2900 kg ha-1) and oil production (> 550 L ha-1).
In Las Cruces (LC), at higher latitude, low temperatures
present beginning in April coincided with the reproductive
stage, affecting yield which was no more than 129 kg ha-1;
thus this zone is not recommendable for chia cultivation.
This study also determined an 11.8 h day length threshold
for the beginning of owering; when plants are exposed to
shorter days ower initiation is more precocious, but when
day length is not adequate plants only begin to ower when
they have accumulated 600-700 ºC d.
Key words: Chia, date sowing, grain yield, photoperiod.
ABSTRACT
Growth and yield of chia (
Salvia hispanica
L.)
in the Mediterranean and desert climates
of Chile
Cecilia Baginsky1, Jorge Arenas2, Hugo Escobar3, Marco Garrido1, Natalia Valero1, Diego
Tello1, Leslie Pizarro3, Alfonso Valenzuela4, Luís Morales1, and Herman Silva1*
RESEARCH
CHILEAN JOURNAL OF AGRICULTURAL RESEARCH 76(3) JULY-SEPTEMBER 2016
1Universidad de Chile, Facultad de Ciencias Agronómicas, Santa
Rosa 11315, La Pintana, Santiago, Chile.
*Corresponding author (hsilva@uchile.cl).
2Universidad Arturo Prat, Facultad de Recursos Naturales Renovables,
Av. Arturo Prat 2120, Iquique, Chile.
3Universidad de Tarapacá, Facultad de Ciencias Agronómicas, 18 de
Septiembre 2221, Arica, Chile.
4Universidad de Chile, Instituto de Nutrición y Tecnología de los
Alimentos, El Líbano 5524, Santiago, Chile.
Received: 20 April 2016.
Accepted: 20 June 2016.
doi:10.4067/S0718-58392016000300001
RESEARCH
INTRODUCTION
Chia (Salvia hispanica L.) belongs to the Lamiaceae family and
its center of origin is in mountain areas of Mexico and Guatemala
(Cahill, 2004). It was traditionally one of the four basic elements
in the diet of Central American civilizations in the pre-Columbian
epoch. Today chia is being re-introduced into western diets
because of its numerous positive nutritional characteristics. These
include 1) a high concentration of essential fatty acids (29%-32%
extractable), 2) polyunsaturated fatty acids omega 3 and omega 6
(Peiretti and Gai, 2009), and 3) mucilaginous ber content (27%
of the grain) that expands when hydrated and has a low cation
exchange capacity that increases availability of several minerals,
proteins, and vitamins required by humans (Reyes-Caudillo et al.,
2008; Ayerza, 2013). These characteristics are helping to rapidly
increase its production worldwide.
Chia is currently cultivated in Australia, Bolivia, Colombia,
Guatemala, Mexico, Peru, and Argentina (Busilacchi et al., 2013).
The largest production center is located in Mexico; it currently
exports seeds to Japan, USA, and Europe (Alenbrant et al., 2014).
Chia grows naturally in tropical and subtropical environments; it
is optimally established from 400 to 2500 m a.s.l., but conditions
below 200 m elevation are not adequate for its cultivation (Orozco
et al., 2014). It is intolerant to freezing in all development stages
(Lobo et al., 2011; Bochicchio et al., 2015). Its minimum and
maximum growth temperatures are 11° and 36 °C, respectively,
with an optimum range of 16-26 °C (Ayerza and Coates, 2009).
It is considered to be a short-day plant with a threshold of 12-13
h (Jamboonsri et al., 2012; Busilacchi et al., 2013), and as such,
its period of growth and fruiting depend on the latitude where
it grows. Jamboonsri et al. (2012) indicated that domesticated
chia germplasm has a owering induction photoperiod of
approximately 12:12 h. Thus in the Northern Hemisphere chia
begins to ower in October and in the Southern Hemisphere in
April. Hildebrand et al. (2013) indicated that with existing plant
germplasm, distribution of chia for grain production is restricted
to 22°55’ N-25°05’ S. At higher latitudes, the probability of
the crop reaching maturity is low, since plants die due to early
frosts (Ayerza and Coates, 2005). Efforts to induce chia to ower
with day lengths greater than 12 h have failed (Cahill, 2004),
with the idea of widening cultivated area to temperate zones
and regions such as the Mediterranean basin. Jamboonsri et al.
(2012) demonstrated the existence of new chia varieties with early
owering, which were able to ower with a photoperiod 15:9 h
in greenhouse and in the eld with a photoperiod of 14 h 41 min.

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There is a new photoperiod-insensitive variety (‘Sahi Alba
914’; Sorondo, 2014), however it is not yet a commercial
variety. The sowing date is an extremely relevant variable,
since it determines the duration of the development period of
the crop due to variations in environmental temperature and
day length to which it is exposed (Lobo et al., 2011). These
conditions are mostly responsible for the potential yield and
seed quality (Ayerza, 2010; 2011).
Grain oil yield ranges from 29.4% to 33.5% depending
on the area of origin of the chia, climatic conditions and the
technique used for its extraction (Ayerza and Coates, 2009).
Studies have shown that seed oil content tends to increase
with altitude in which seeds are grown; however, the
environment also modies oil composition. The temperature
affects the type of fatty acids present in the oil; increase of
temperature during grain development produces a reduction
in the production of polyunsaturated fatty acids. A relation
has been found between altitude, fatty acid composition,
and oil saturation. Ayerza and Coates (2009) found that in
different inter-Andean valleys of Ecuador, the lower altitude
environment (1600 m a.s.l.) showed a higher percentage of
α-linolenic acid, but lower values for linoleic and oleic acids,
indicating that oil saturation decreased at higher altitude
(Ayerza, 2011). This would be related to the negative relation
between altitude and temperature, which produces changes
in the metabolic processes as has been demonstrated in other
oil crops such as jojoba (Simmondsia chinensis (Link) C.K.
Schneid.; Ayerza, 2001). Colder temperatures generally
increase the level of unsaturation of fatty acids (Thomas et
al., 2003).
The objective of this study was to determine the effect
of climatic conditions in regions of coastal desert, normal
desert, and dry Mediterranean with marine inuence climates
of Chile on the growth, yield and oil composition of chia
grown with irrigation. This study also proposes to determine
the most promising sowing date in terms of yield and oil
quality in chia in the different study areas.
MATERIALS AND METHODS
The study was performed from January to June 2013 in
Valle de Azapa in the Arica y Parinacota Region, Canchones
in Alto Hospicio, Tarapacá Region, and Las Cruces in
Valparaiso Region, Chile. In the Valle de Azapa (18°30’ S
lat) prevailing conditions are a coastal desert climate with
abundant cloudiness, no frost, high humidity and high
solar radiation throughout the year (González et al., 2013).
Canchones Experiment Station has a desert climate, hot with
absence of rainfall and strong thermal oscillation and low
relative humidity. It is located at an altitude of 990 m a.s.l.
and therefore exposed to frost that begins in winter (Lanino,
2005). Las Cruces has a mild Mediterranean climate with
very low incidence of frost, located at an altitude of 12 m
a.s.l. Table 1 summarizes climate and soil characteristics of
the experimental sites, and Table 2 shows thermal sum and
number of freezes recorded during January-June 2013 in
each experimental site.
Conguration of the experiment and
agricultural management
We used phenotypes of white and dark (black spotted) chia
from Santa Cruz de la Sierra (Bolivia) provided by the
company Benexia S.A. Five sowing dates were used for each
locality (Table 2), constituting 10 treatments per locality (5
sowing dates × 2 phenotypes). The experimental design
was randomized complete blocks with six replicates. The
experimental unit was dened as a plot 5.0 m long and 2.4 m
wide, with 0.6 m between rows.
Seeds were sown by hand at 1 cm depth with 5 kg
seeds ha-1, producing plant densities between 120 and
160 plants m-2. Once the rst pair of leaves had completely
unfolded, plants were thinned to a density of 80-90 plants m-2
(50 to 60 plants m-1). Plants were fertilized by fertirrigation
Characteristics
Elevation (m a.s.l.) 230 996 12
Temperature, °C Max. 24.1 33.8 22.5
Min. 13.3 3.7 1.8
Mean 18.6 19.2 12.2
Precipitation, mm Mean 0.0 0.0 166.0
VPD, kPa 1.1 1.2 0.9
Soil
Order Aridisol Aridisol Mollisol
pH 7.8 8.1 7.8
Salinity, dS m-1 7.9 28.4 0.8
Nutrient, mg kg-1 N 123.0 117.0 10.0
P 37.0 20.0 92.0
K 914.0 1060.0 264.0
Organic matter, % 1.4 1.5 2.3
Irrigation water
pH 7.8 8.2 7.5
Salinity, dSm-1 1.1 8.2 7.8
Carbonate, mmol L-1 HCO2-3 2.1 1.4 5.3
Sulfate, mg L-1 SO-2 239.0 286.0 262.0
Chloride, mg L-1 Cl- 146.0 113.0 258.0
Boron, mg L-1 B 0.7 nd nd
Sodium, mmol L-1 Na+ 4.2 7.7 7.5
Table 1. Location climatic characteristics, and soil and water
quality from sowing to harvest in the sites where chia was sown.
*Dry Mediterranean with maritime inuence (Csb).
VPD: Vapor pressure decit.
nd: not detected.
Location
Valle de Azapa Canchones Las Cruces
General
Climate Coastal Desert Normal Desert Mediterranean*
Latitude 18°30’ S 20°26’ S 33°30’ S
Longitude 70°00’ W 69°32’ W 71°36’ W
Month
January 332.3 0 363.2 0 194.2 0
February 350.2 0 346.0 0 181.6 0
May 310.2 0 337.7 0 113.9 0
April 196.0 0 223.1 0 54.3 0
May 183.5 0 207.8 4 44.3 0
June 176.9 0 161.0 9 22.0 5
Total 1549.1 0 1638.8 13 610.3 5
Table 2. Accumulated growing degree-days (GDDi) and frost
number.
GDDi: Accumulated growing degree days (> 10 ºC).
Locality
GDDi GDDi GDDi
Valle de Azapa
Frost
number Frost
number Frost
number
Canchones Las Cruces

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to maintain 45, 60, and 100 units N, P2O5, and K2O,
respectively, in the soil. We used drip irrigation controlled
by a Rain Bird programmer (Rain Bird Corporation, Tucson,
Arizona, USA). Water requirements were programmed
to replenish 70% of reference evapotranspiration demand
(ET0), calculated daily using the Penman Monteith method.
Weeds were controlled by hand when they appeared; no
pests or diseases were observed.
Sampling and analysis
Plants were evaluated every 2-3 d to determine their
phenological stages of emergence (visible cotyledons and
completely expanded over the soil), initiation of branching
(aerial branches with a node), ower initiation (inorescence
of the central stalk with rst ower open), physiological
maturity (inorescence of the central stalk with brown
color), and harvest maturity (grains with 14% humidity).
The development stage was dened when 50% plants of the
experimental unit reached that stage.
At the beginning of owering the accumulation of aerial
biomass was measured by sampling 0.5 m of plants per
experimental unit. These were dried in an oven at 70 °C. This
measurement was repeated at harvest, when we also evaluated
plant height and length, and number of inorescences. Yield
was measured along 3 m per experimental unit by separating
inorescences from the rest of the plant. The inorescences
were dried in a forced air oven at 40 °C until constant
weight. Dry inorescences were threshed, separating the
grains. Yield per plant was estimated and 500 grains sampled
randomly were weighed. We calculated the harvest index
(HI) as the quotient of dry weight to total plant weight.
Growing degree days (GDDi) were calculated with the
following function:
[1]
where GDDi is the accumulated growing degree days,
Ti is the mean air temperature on day i and Tb is the base
development temperature, in this case 10 °C, given the area
of origin of the species.
The day length for each locality was estimated as:
[2]
where DL is day length, LAT is the latitude of the locality,
and d is the corresponding calendar day.
The meteorological variables were obtained from weather
stations closest to the study sites.
To quantify oil content and lipid prole after harvest we
sampled 500 g seeds per treatment; we used the modied cold
method of Folch et al. (1957) to extract oil and determine
lipid prole of each oil sample. To each 100 g chia seeds
ground in a processor we added 500 mL 2:1 (v:v) mixture of
chloroform:methanol and mixed for 5 min. Mixtures were
then ltered twice, obtained ltrate was placed in a decantation
funnel and 40-50 mL distilled water was added, agitated
vigorously for 2 min and then left to stand until phases were
completely separated (1.5-3 h). Two phases were obtained,
the lower phase is chloroform, which was rescued by rotary
evaporation at 60 °C in a vacuum. This separated the solvent,
remaining chia oil was 25%-39% total seed weight. The oil
samples of each treatment were homogenized and then total
fatty compounds were extracted in methanol:chloroform 2:1
with 0.5 N Mg2Cl. Total fatty compounds and chloroform were
recovered; the latter was evaporated using N2, thus obtaining
the total lipid fraction available in seeds. To determine fatty
acid prole of fatty fraction extracted it was rst methylated
with boron triuoride (BF3) in 12% methanol and then with
NaOH in methanol to obtain free methylated esters of fatty
acids. These were suspended in hexane in order to be injected
into a gas chromatograph.
Statistical analyses
The results obtained for each locality were submitted to
ANOVA using the InfostatGenprogram (Balzarini et al.,
2011); when signicant differences were found among
treatments (P < 0.05), Duncan post hoc test was used to
identify signicantly different means. We also performed a
combined ANOVA to evaluate the effect of locality on sowing
date and chia accession, and a principal components analysis
to associate growth parameters (biomass at owering and
harvest, plant height, number and length of inorescences
and yield) with the combination of locality and sowing date.
RESULTS
Climatic condition and crop development
The day lengths to which the crops were exposed were similar
at VA and CH, which are at similar latitudes (18º and 20º S
lat, respectively). At 33°30’ S lat, LC had longer days before
the autumn equinox and shorter days afterwards. Canchones
had the greatest temperature oscillation and LC the least.
Mean temperatures were similar in VA and CH, and lower
in LC (Figure 1). Since there were nonsignicant differences
between white and dark chia phenotypes in their growing
cycle length, values of each type were averaged. Sowing
date did not modify cycle length in VA or CH. However,
in VA owering and maturity were more rapid when plants
were sown later (Figure 2). In CH, frosts affected plants and
did not allow them to reach maturity on the latest sowing
dates (Figures 1 and 2). The low thermal accumulation in
LC did not allow plants to reach and complete adequately
their phenological development (Figures 1, 2, and 3). In VA,
the fth sowing date (F5) reached physiological maturity
with a thermal sum of 728.7 °C d, while in LC the earliest
sowing (F1) only accumulated 585.1 °C d at harvest. Plants
did not accumulate sufcient temperature in LC, due to
which they could not adequately complete their phenological
development. This is shown by the lack of yield, since there
was not an adequate phase for grain lling. Chia thermal
requirements were affected by day length. Photoperiods of
less than 11.8 h dene the lower thermal requirement of
plants to start owering, which corresponded to about 500
°C d, while longer day length increases the thermal time
necessary to initiate owering (Figures 1 and 3).
GDDi = Σti (Ti _ Tb) if GDDi < 0 then GDDi = 0
tf
DL = ACOS _ TAN(LAT)TAN 23.45
24
π
( )
(
[
SIN( (d + 284)
360
365
)
]

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CHILEAN JOURNAL OF AGRICULTURAL RESEARCH 76(3) JULY-SEPTEMBER 2016
Figure 1. Variation of photoperiod (A), minimum temperature (B), maximum temperature (C), and accumulated evapotranspiration
(ET0) (D) from January to June 2013 in each experimental site.
Figure 2. Days after sowing to ower initiation (A) and physiological maturity (B) of chia plants with different sowing dates (F1-F5)
in three localities. The missing data of physiological maturity for Canchones and Las Cruces resulted from plant death due to the
effect of frost and/or low temperature.

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Growth parameters and crop yield
The behavior of biomass production, yield, and harvest index
did not present the same tendency in the different localities
(Table 4). Only VA had a signicant interaction between
sowing date and phenotype (P < 0.05), thus in this locality the
effect of the individual factors (sowing date and accession)
are not clear (Table 3); however, in all experimental sites
the effect of sowing date was signicant (P < 0.01), thus
the tendency in VA was a decrease in biomass production
with later sowing and increase in yield and harvest index.
This is explained by the fact that in the latest sowing dates
there were lower temperatures, which did not reach the level
of plant damage, which prolong the period of grain lling
and thus increase yield and harvest index (Tables 3 and 4).
Biomass production increased in CH with later sowing,
but yield and harvest index decreased (Table 3). In this
case there was less thermal oscillation, which implies high
temperature accumulation, however at the end of summer
there were periods of cold during the day and freezes in
autumn (Figure 1B, 1C). Due to these conditions, length
of grain ll period was relatively constant between sowing
dates, but the later sowings had less metabolic activity during
grain lling due to cold stress and later damage from freezes.
This resulted in lower yield in the later sowing dates (Table
4). In LC, the three parameters mentioned above decreased
with later sowing dates. The difference between types of
chia was signicant for biomass and harvest index variables
only in LC; dark phenotype produced greater values. The
lowest yields were observed in LC (70 to 129 kg ha-1). The
presumed reason was that low temperatures experienced
during plant reproductive stage, which affected grain lling
during sowing dates F1, F2, and F3, resulted in most plants
not reaching necessary physiological maturity, some did not
even start owering in period F5. Similarly, CH plants were
damaged during owering by low temperature for the last
two sowing dates (F4 and F5).
The combined ANOVA indicated a signicant interaction
between locality and sowing date. The greatest yields were
achieved during periods F5 and F1 in VA and CH, with yields
of 2500 and 1900 kg ha-1, respectively, while the lowest
yields were obtained in LC for all sowing dates, with a mean
of 105 kg ha-1 (Figure 4). Valle de Azapa produced high
and stable yields independent of sowing date, in contrast to
CH where F3 produced a signicant reduction in yield with
delayed planting date (49% reduction between F1 and F3).
Components of yield and association
among variables
Delaying sowing date tended to generate larger inorescences
in all localities (from 13.6 to 34.1 cm). However, in CH there
was a signicant interaction between sowing date and accession
(P ≤ 0.05). Here the white phenotype, planted at the earliest
date generated relatively small inorescences (Table 5). The
number of inorescences increased with later sowing dates in
all localities except CH, where the incidence of frost altered
this tendency without signicant differences (P > 0.05) between
Figure 3. Relation between accumulated growing degree days
according to Equation [1] and base temperature of 10 °C and
day length according to Equation [2], and owering initiation
of chia in different localities and sowing dates.
Table 3. Planting date, days after planting and accumulated growing degree days in chia plants in different development stages for
each locality.
DAS: Days after sowing; GDDi: accumulated growing degree days (> 10 ºC).
Locality
Valle de Azapa
F1 4 Jan 56 1 March 649.1 93 7 April 1001.4 136 20 May 1.271.6
F2 18 Jan 47 6 March 558.0 84 12 April 1067.2 136 4 June 1.186.1
F3 4 Feb 49 25 March 551.4 87 2 May 811.6 134 18 June 1.084.7
F4 18 Feb 49 8 April 489.0 96 25 May 782.9 123 21 July 1.115.3
F5 6 March 48 23 April 392.6 105 19 June 728.7 148 1 Aug 927.7
Canchones
F1 4 Jan 58 3 March 709.3 106 20 April 1165.3 139 23 May 1.393.0
F2 18 Jan 53 11 March 643.5 110 8 May 1110.4 137 5 June 1.282.8
F3 1 Feb 51 24 April 606.9 109 21 May 1048.4 130 11 June 1.159.4
F4 18 Feb 54 13 April 568.2 - - - - - -
F5 1 March 60 30 April 547.6 - - - - - -
Las Cruces
F1 4 Jan 111 25 April 519.1 - - - 170 23 June 585.1
F2 18 Jan 97 25 April 436.5 - - - 158 26 June 502.6
F3 2 Feb 84 27 April 333.4 - - - 144 26 June 396.8
F4 18 Feb 81 10 May 238.1 - - - - - -
F5 4 March - - - - - - - - -
DateDAS GDDi
Flowering beginning
DateDAS GDDi
Physiological maturity
DateDAS GDDi
Harvest maturity
Planting
date

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sowing dates (Table 5). There was interaction between sowing
date and phenotype only in LC; white seeds sown earlier (F1)
produced a mean of two inorescences less than the rest of
treatments. It should be noted that there were nonsignicant
differences between the phenotypes for any of these variables.
The results of the principal components analysis are
represented in Figure 5. There was a signicant association
among growth variables with sowing dates in VA and CH.
Las Cruces was excluded from this analysis due to the lack
of thermal accumulation (Tables 2 and 3 and Figure 3) that
generated different situations. Components 1 and 2 explained
76% of the observed variance; seed yield was positively and
signicantly associated with inorescence length (r = 0.48,
P ≤ 0.05), harvest index (r = 0.85, P ≤ 0.001), and with the
number of inorescences per plant (r = 0.85, P ≤ 0.001) and
thus with important components in determining yield. Yield
and length of the main inorescence were negatively and
signicantly associated with day length at the beginning of
owering. The lowest yields observed in CH and sowing
dates F1 and F2 in VA were associated with the longest
day lengths. Photoperiod was positively and signicantly
associated with plant height and biomass at owering.
Table 5. The length of central axis inorescence and number
of inorescences per plant of chia in different localities and
different sowing dates (SD).
*, **Signicant at the 0.05 and 0.01 probability levels, respectively.
VA: Valle de Azapa; CH: Canchones; LC: Las Cruces; ns: nonsignicant.
Factor
SD
F1 13.6d 14.6c 15.1c 11.2b 8.6a 5.4b
F2 14.0d 18.2b 16.1b 10.3b 11.3a 6.8a
F3 17.3c 20.4a 18.6a 9.8b 8.2a 6.5a
F4 19.6b 10.1b
F5 34.1a 13.8 a
** ** ** ** ns **
Accession
White 19.3a 17.7a 16.5a 10.8a 10.0a 6.1a
Dark 20.1a 17.8a 16.7a 11.2a 8.7a 6.3a
ns ns ns ns ns ns
SD × Accession
F1×White 13.3a 13.5d 15.4a 11.5a 8.0a 4.5b
F1×Dark 14.0a 15.8c 14.9a 10.8a 9.2a 6.2a
F2×White 13.4a 19.0ab 16.1a 9.3a 13.6a 6.9a
F2×Dark 14.5a 17.5bc 16.2a 11.2a 9.0a 6.6a
F3×White 17.5a 20.5a 18.1a 9.7a 8.3a 7.0a
F3×Dark 17.0a 20.2a 19.1a 9.8a 8.1a 6.1a
F4×White 17.0a 9.8a
F4×Dark 22.2a 10.3a
F5×White 35.3a 13.7a
F5×Dark 32.9a 14.0a
ns * ns ns ns **
CHVA LC
Central axis
inorescence length
CHVA LC
cm
Nr inorescences
per plant
Table 4. Biomass, yield and harvest index for chia grown in different localities and with different sowing dates (SD).
*, **Signicant at the 0.05 and 0.01 probability levels, respectively.
HI: Harvest index; VA: Valle de Azapa; CH: Canchones; LC: Las Cruces; ns: nonsignicant.
Factor
SD
F1 7.300a 7.594b 5.152a 1.479b 1.912a 127a 0.22d 0.27a 0.04a
F2 4.151c 9.291b 3.066b 1.375b 1.622b 114a 0.34c 0.21a 0.04a
F3 3.529c 12.330a 2.051c 1.337b 941c 75b 0.38c 0.09b 0.03b
F4 3.710c 2.285a 0.56a
F5 5.611b 2.468a 0.44b
** ** ** ** ** ** ** ** **
Accession
White 4.856a 9.817a 2.931b 1.669b 1.460a 104a 0.38a 0.19a 0.03b
Dark 4.864a 9.660a 3.915a 1.908a 1.524a 107a 0.40a 0.19a 0.04a
ns ns ** ** ns ns ns ns **
SD × Accession
F1×White 8.971a 6.504a 4.281a 1.324d 1.823a 129a 0.15e 0.28a 0.04a
F1×Dark 5.629bc 8.684a 6.024a 1.634d 2.002a 125a 0.29d 0.25a 0.04a
F2×White 3.635d 10.703a 2.660a 1.222d 1.808a 113a 0.34cd 0.20a 0.03a
F2×Dark 4.667cd 7.879a 3.473a 1.527d 1.437a 115a 0.33cd 0.22a 0.04a
F3×White 3.498d 12.245a 1.854a 1.280d 941c 70a 0.37bcd 0.08a 0.02a
F3×Dark 3.559d 12.415a 2.248a 1.393d 941c 81a 0.39bc 0.10a 0.03a
F4×White 3.559d 2.484b 0.58a
F4×Dark 3.860d 2.085c 0.54a
F5×White 4.618cd 2.032c 0.44b
F5×Dark 6.604bc 2.903a 0.44b
** ns ns ** ns ns * ns ns
CHVA LC
Biomass
CHVA LC
Yield
CHVA LC
Harvest index
kg ha-1
Different letters on bars indicate signicant differences between sowing
dates and localities, according to Duncan test (P < 0.05). n = 12.
Figure 4. Effect of ve sowing dates on chia grain yield established
in three locations (Valle de Azapa, Canchones, and Las Cruces).

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CHILEAN JOURNAL OF AGRICULTURAL RESEARCH 76(3) JULY-SEPTEMBER 2016
Total oil, -linolenic (omega 3) and linoleic
acid content in seeds
The total seed oil content and the content of α-linolenic and
linoleic acids obtained in VA and CH are shown in Table
6. These parameters were not determined for LC due to the
scarce seed production and the notorious damage effect
observed in them as a result of the low temperatures during
grain lling. The results show that there were nonsignicant
differences between phenotypes for total oil content;
however, α-linolenic and linoleic acid content tended to
be higher in VA. There was signicant interaction between
sowing date and phenotype in all localities except CH,
where there was no interaction with total oil content and
no difference between sowing dates (Table 6). The highest
values of total oil content, α-linolenic and linoleic acid
content were found in the latest sowing dates; F4 (February
18) and F5 (March 6) showed signicantly greater values
in white and dark phenotypes, respectively (Table 6). There
was no clear tendency in CH with respect to sowing date;
phenotype was more relevant in this case, as indicated above.
DISCUSSION
Flower induction by short days is common in tropical
crops such as chia and this helps the plant to synchronize
development with the rainy season, which occurs in the
warmest time of year (Jamboonsri et al., 2012; Busilacchi
et al., 2013). The geographic location (between 18° and 33°
S lat) and the climatic conditions of the sites (VA - coastal
desert, CH - normal desert, and LC - Mediterranean) caused
ower induction to occur at the beginning of autumn, when
temperatures were decreasing (Figure 1). The result of this
was that in CH and LC, for the later sowings, owering
occurred in the presence of low temperatures and even frosts.
The short-day character of chia was reected by the timing
of owering, which was stable in VA and CH independent
Figure 5. Biplot of principal components for yield and growth variables: Biomass at owering and harvest, length of central axis
inorescence, number of inorescences per plant, plant height, and harvest index.
HI: Harvest index; Photoperiod: day length at the beginning of owering; BF: biomass at the beginning of owering. Pentagons represent different sowing
dates in localities of Valle de Azapa (VA) and Canchones (CH). B: dark chia accession; W: white chia phenotype; 1, 2, 3, 4 and 5: sowing dates F1, F2, F3, F4,
and F5, respectively.
Table 6. Total oil, alpha-linolenic (omega 3) and linoleic acid
content in seeds in two localities (Valle de Azapa and Canchones)
established in different sowing dates (SD).
*, **Signicant at the 0.05 and 0.01 probability levels, respectively.
VA: Valle de Azapa; CH: Canchones; LC: Las Cruces; ns: nonsignicant.
Factor
SD
F1 343.2c 249.1a 188.5bc 194.3a 56.3c 70.1a
F2 324.9c 312.5a 157.3c 147.2b 45.4d 50.9a
F3 321.3c 330.7a 192.2bc 169.2ab 53.5cd 55.5a
F4 446.5b 267.3a - 74.7b -
F5 508.1a 295.6a - 84.8a -
*** ns *** *** *** ***
Phenotype
White 396.4a 291.2a 234.0a 150.7b 65.7a 53.2b
Dark 381.2a 303.7a 206.4a 189.8a 60.2a 64.4a
ns ns ns *** ** ***
SD × Phenotype
F1×White 324.8c 238.4b 181.3cd 185.4b 53.8c 67.8ab
F1×Dark 361.7bc 259.7ab 195.8cd 203.2ab 58.7c 72.4a
F2×White 325.0c 353.1ab 152.6d 167.2b 43.2d 58.6b
F2×Dark 324.7c 272.0ab 162.1cd 127.3c 47.5cd 43.1c
F3×White 321.0c 282.1ab 201.2c 99.6c 54.7cd 33.1c
F3×Dark 321.5c 379.4a 183.2cd 238.8a 52.3cd 77.8a
F4×White 583.9a - 376.5a - 104.1a -
F4×Dark 309.0c - 158.1cd - 45.3cd -
F5×White 427.3bc - 258.6b - 72.5b -
F5×Dark 588.9a - 332.7a - 97.0a -
*** ns *** *** *** ***
CHVA
Total oil Omega 3 Linoleic acid
CHVA
L ha-1
CHVA

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CHILEAN JOURNAL OF AGRICULTURAL RESEARCH 76(3) JULY-SEPTEMBER 2016
of sowing date (Figure 3) and where mean temperature and
day length did not show large variation during the growing
season (Figure 1). By contrast, in LC, where variation in day
length was greater, there was a reduction in days to ower
initiation as the day length decreased. Similar behavior
has been observed in other species that are sensitive to
photoperiod (Christiansen et al., 2010). We found a
photoperiod threshold of 11.8 h for ower initiation, similar
to that indicated by Jamboonsri et al. (2012), who reported
12 h for domesticated chia germplasm, while Busilacchi et
al. (2013) indicate values of 12-13 h. Above this threshold,
thermal requirements of the crop increased linearly, with
a difference of up to 200 °C d between day lengths above
and below the threshold (Figure 3). One result of this was
that chia thermal requirements in LC were lower than in
VA and CH, since in LC the plants always experienced a
day length below this threshold. However, life cycles were
longer in LC (Table 2) due to a slower development rate
as a product of lower thermal accumulation, which did not
allow grain lling. This situation is characteristic of some
short-day species such as sorghum (Craufurd and Qi, 2001),
Miscanthus sacchariorus (Jensen et al., 2013) and soybean
(Jiang et al., 2011) under these conditions.
Together, these results suggest that day length sensitivity
of chia is quantitative; when exposed to decreasing day
lengths oral initiation is accelerated. However, if plants
do not experience a sufciently long day length, owering
will occur when a certain quantity of day degrees is
accumulated. In this study the quantity of day degrees
was between 600 and 700 °C d, with a base development
temperature of 10 °C. These results concur with studies in
other quantitative short-day species from tropical climates
such as Miscanthus, which requires 600 °C d to owering
(Jensen et al., 2013) and sorghum which requires 870 °C d
(Ritchie and Alagarswamy, 1989). The study of Lobo et al.
(2011) in Tucumán, Argentina, showed that January sowing
generated plants with a development cycle of approximately
160 d, which is very similar to that observed in LC, which
is at a similar latitude (Table 3). These authors indicated that
sowing at a later date could be risky because of the possibility
of frost during the reproductive stages. This sensitivity to
frost connes the development of chia to zones with few or
no freezing events or to areas with temperatures that are not
lower than 5 °C during owering (Lukatkin et al., 2012).
It has been observed that many tropical plants suffer
important frost damage when they are exposed to temperatures
slightly below 0 °C and cold damage has been sometimes
been reported at temperatures close to 5 °C (Lukatkin et al.,
2012). Critical periods for ower initiation and for ower
development have also been observed in various species of
genus Salvia. For example, Salvia leucantha Cav. requires 12
h light for oral induction and 10 h light to continue ower
development (Armitage and Laushman, 1989). Further
studies are needed to verify if chia behaves similarly.
Our observed yields were all superior to those that have
been reported in the literature with the exception of LC,
where low temperatures during owering and grain-lling
periods signicantly reduced yields (Figure 4). This is
similar to studies performed at relatively high latitudes,
such as in Choele-Choel (39°11’ S lat), Argentina, and
Tucson (31°14’ S lat; Arizona, USA), where chia plants
died due to freezing before owering. It is not surprising
that low temperatures affect growth and yield of chia,
considering that the species is adapted to temperatures that
uctuate between 11 and 36 °C (Ayerza and Coates, 2009).
Studies performed in Argentina yielded from 606 to
1400 kg ha-1 (Lobo et al., 2011). In Paraguay yields of
1600 kg ha-1 were reported (Bochicchio et al., 2015), while in
the state of Jalisco, Mexico, the main productive zone of chia
in this country, mean yields of 1200 kg ha-1 were obtained.
In our study, the yields of sowing dates F1 and F2 (between
4 January and 4 February) in CH were systematically and
signicantly superior to those of VA (Figure 4). In spite of
being at similar latitudes, these localities have important
climate differences, given that VA has coastal inuence, with
higher minimum and lower maximum temperatures than CH,
and an atmospheric demand (reference evapotranspiration
accumulated to Julian day 100) of 387.7 mm, compared to
495 mm in CH. Although the meteorological conditions were
similar at the beginning of the season, from February on the
minimum temperatures in CH were substantially less than
in VA, which may explain low yield in third sowing (F3) in
the latter locality. The high yields obtained from F4 and F5
in VA were associated with favorable day-length conditions,
lower mean temperatures and the high levels of radiation
that characterize these latitudes. The lower temperatures
in these two sowing dates in VA would have generated
higher levels of unsaturated fatty acids such as α-linolenic
and linoleic (Table 6). This was reported by Ayerza (2011),
who found changes in chia oil composition (higher level of
unsaturation of fatty acids) in higher altitude ecosystems
in which temperature was lower than in localities at lower
altitudes. This would also explain the lack of signicance
in oil contents in the rst three sowing dates in VA and CH,
since temperatures observed in these three dates were similar
in both localities (Figure 1).
The dark phenotype tended to have higher yield, although
this was only signicant in VA (Table 4). Something similar
occurred in the fatty acid prole, where the dark phenotype
showed higher levels of α-linolenic and linoleic acids,
although the differences were only signicant in CH. The
similarity between phenotypes may be due to the fact that
both came from the same area (Santa Cruz de la Sierra in
Bolivia) and that both were growing within their adaptation
limits, as has been observed for other species of arid zones
such as jojoba (Ayerza and Zeaser, 1987). The dark phenotype
only had a greater level in CH (in the three sowing dates in
which seeds were obtained), coinciding with the report of
Ayerza and Coates (2009), who indicated that this phenotype
probably has greater capacity to adapt to the environmental
conditions in which it was growing. However, other studies
of Ayerza (2011; 2013) found nonsignicant difference in
oil content and fatty acids prole between spotted-black
seeds and white seed of chia grown in different ecosystems.

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CHILEAN JOURNAL OF AGRICULTURAL RESEARCH 76(1) JANUARY-MARCH 2016
CHILEAN JOURNAL OF AGRICULTURAL RESEARCH 76(3) JULY-SEPTEMBER 2016
The studies indicate that the larger differences found in
oil content and fatty acid composition are due to location
(because of environmental differences) rather than chia seed
coat color (Ayerza, 2010).
Plant height generally decreased with later sowing and
thus shorter day length; in VA heights did not surpass 60 cm
when plants were sown in the middle of February (F4, data
not shown). Similar results were obtained by Busilacchi et
al. (2013) in chia as well as in other short day plants such
as Amaranthus (Al Hakimi, 2005) and soybean (Jiang et al.,
2011). However, plant height was greater in the last sowing in
VA (beginning of March, F5); plant height reached almost 90
cm (data not shown) due to the large size of the inorescence
of the central axis, which averaged more than 30 cm in length
(Table 5). Larger inorescence found in later sowings would
be associated with better distribution of assimilates, as has
been observed in soybean (Han et al., 2006).
Plant height was not positively associated with yield
in chia, in contrast with species such as wheat, in which
introduction of dwarng genes in varieties produced superior
yields relative to those obtained in taller plants by increasing
the harvest index (Zapata et al., 2004). Greater plant height in
chia was linked to a lower harvest index (Figure 5), consistent
with the results of Berti et al. (2011) in camelina (Camelina
sativa [L.] Crantz). Plants had greater height under long
days (data not shown), along with shorter inorescences
and smaller number of inorescences (Table 5), all of which
indicate less assimilates used for grain production and thus
lower harvest index (Table 4). This concurs with the report
of Han et al. (2006) in short-day soybean plants, in which
shorter days in autumn promoted the partition of dry material
to the seeds.
Inorescence height was negatively associated with the
number of vegetative nodes, and number of vegetative nodes
was positively associated with day length (data not shown).
Thus owering dynamics in chia is almost completely
determined by the interaction of day length and photoperiod
(data not shown), and not by the number of vegetative nodes
as in Chrysanthemum (Mattson and Erwin, 2005), a short
day plant in which owering is also determined by a critical
number of vegetative phytomers (Jensen et al., 2013).
The greatest yields observed in this study were associated
with greater biomass accumulation, which was mainly
generated by producing large inorescences on shorter
plants. These were plants that had better assignation of their
resources and thus had higher harvest indices. Additionally,
the highest yields were also associated with latitudes
with longer day lengths. These results have important
implications for the management of chia, since shorter,
higher yielding plants can also be more efciently harvested
using mechanized harvesting methods.
CONCLUSIONS
The conditions of coastal desert and normal desert climates
in Valle de Azapa (VA) and Canchones (CH), respectively,
provide the best conditions for chia production under the
conditions of this study. However, the coastal desert climate,
with less thermal oscillation, less extreme temperatures and
without freezes during the entire season, make VA even more
appropriate for the production of this species. Thus in the VA
sowing between middle of February and middle of March
may produce yields greater than 2000 kg ha-1, oil content
above 550 L ha-1 and α-linolenic and linoleic acid content
above 350 and 90 L ha-1, respectively.
Chia cultivation is not advisable in LC, due mainly to
the low thermal accumulation in a dry Mediterranean
climate with marine influence, along with temperatures
below 5 °C in April, the period in which the reproductive
stage begins. Although there are low temperatures and
freezes in CH beginning in April, there is high thermal
accumulation that allows plants established in January
to complete their phenological development adequately.
Thus in CH, sowing should ideally be confined to the
beginning of January. Here we determined a day length
threshold of 11.8 h for floral initiation. When plants
are exposed to shorter days flower initiation is more
precocious, but when the day length is not adequate
plants only begin to flower when they have accumulated
600-700 °C d. This suggests that chia has a quantitative
type day length sensitivity; however, further studies are
required to corroborate this hypothesis.
ACKNOWLEDGEMENTS
This work was supported by grant FONDECYT Nº1120202,
CONICYT, Government of Chile.
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