Industrial Crops and Products 7 (1998) 169 – 174

Developmental stability of quinoa under European conditions

Sven-Erik Jacobsen
Department of Agricultural Sciences, The Royal Veterinary and Agricultural Uni6ersity, Thor6aldsens6ej 40,
DK-1871 Frederiksberg C, Denmark

Received 6 August 1996; accepted 28 October 1996

Abstract

Five quinoa lines, from different maturity groups, were investigated for their adaptability to European conditions.
Results from experiments conducted over three growing seasons indicate that seed types originating from Chile are
adapted for growth under northern European conditions. Ranking of the lines for earliness was probably determined
by the beginning of July and was consistent over years, indicating that selection for this character could take place
at an early stage in plant development. Discussion of the plant breeding implications of these and other results
suggested that a model genotype for seed production for northern European agriculture would be early, uniformly
maturing, non-branching and short, with a high seed yield and a low saponin content. © 1998 Elsevier Science B.V.

Keywords: Chenopodium quinoa; Development; Model plant; Selection criterion

1. Introduction The prospects of introducing quinoa into Eu-

Quinoa (Chenopodium quinoa Willd.) has been
cultivated in the Andean region for thousands of
years. Recently, however, increased interest has
been shown in this crop by agronomists and
consumers alike because of its potential as a
palatable and nutritious food. Possible uses are in
either gluten-free breads, in brown breads mixed
with wheat flour, in soups, salads and infant
foods and as an alternative to rice. Furthermore,
its protein quality is such as to make it of value as
an animal feedstuff.
ropean agriculture have been discussed in a num-
ber of papers (Risi and Galwey, 1984, 1989a,
1991; Galwey, 1989; Fleming and Galwey, 1995;
Jacobsen, 1993; Jacobsen and Stølen, 1993; Jacob-
sen et al., 1994, 1996). In 1994 an EU-project,
embracing 11 organisations in six member states,
was initiated to examine the industrial uses of
quinoa (Galwey, 1993).
As a potential new crop in European agricul-
ture, problems relating to the breeding and selec-
tion of quinoa may arise. Risi and Galwey (1991)
showed that testing, even in a limited range of
carefully chosen environments, could give infor-

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PII S 0 9 2 6 - 6 6 9 0 ( 9 7 ) 0 0 0 4 5 - 9
170 S.-E. Jacobsen / Industrial Crops and Products 7 (1998) 169–174
Table 1
Origin and morphology of quinoa lines
Group/origin line Basic colour
1/205 Baer×Faro (Chilean cultivars) Green
2/210 Kcancolla×Amarilla de Green
Marangani (Peruvian cultivars)
3/224 Germplasm collection Chilean Green
origin
4/233,240 Field selection from Chilean Green
material
5/Olav Standard variety Chilean origin Green
a
See Table 2.
mation into those factors which must be taken
into account when adapting a crop to a new
region. Probably the most important single fea-
ture which will determine whether quinoa can be
successfully introduced into Europe will be its
developmental pattern and particularly its earli-
ness, in these new conditions. Results from a
study by Risi and Galwey (1989b), describing 294
quinoa accessions, indicated that significant dif-
ferences existed for the duration of all growth
phases, with the period from two true leaves to
bud formation ranging from 41 to 89 days; from
bud formation to anthesis 7 – 53 days and the
period from anthesis to maturity lasting between
65 and 137 days.
Duration of developmental stages and stability
of developmental pattern across years, were inves-
tigated under Danish conditions, to determine the
most appropriate time in a breeding programme
to perform selection and to assess which lines
could serve as potential parents in such a pro-
gramme (Jacobsen et al., 1996). Various measures
of stability were employed to analyse the data,
including those proposed by Francis and Kannen-
berg (1978) and Lin and Binns (1988), modified
for the purposes of the investigation. Selection for
developmental stage, height and inflorescence size
could be performed satisfactorily at an early stage
of the breeding programme. For saponin content,
however, the measuring techniques then available
were too insensitive to enable a recommendation
to be made (Jacobsen et al., 1996).
Colour of top Developmental stage for top Leaf size
colour expressiona
Red 16 – 17 Large
Light red 15 Large
White 11 Small
Orange ( “red) 14 – 15 (/16 – 17) Medium
Yellow 15 Large
The aim of this investigation was to define the
range of quinoa genotypes with the potential to
adapt to European conditions, for which purpose
lines from five different maturity classes were
studied over a number of growing seasons.
2. Materials and methods
The experiment was carried out in 1991, 1992
and 1995, on sandy loam soil at the research
station of the Royal Veterinary and Agricultural
University in Tåstrup, Denmark. Quinoa lines,
including the standard variety Olav, were grown
at a target density of 200 plants/m2. Seed rate was
adjusted on the basis of a laboratory germination
test and an estimated field germination of 60%,
though actual plant density differed due to vary-
ing field germination.
The lines, whose origins are shown in Table 1,
can be regarded as representative of the groups
described by Jacobsen and Stølen (1993), which
were known to differ markedly in length of grow-
ing period. Groups 1 and 4 were early, group 3
medium-early, group 5 medium-late and group 2
late maturing. All lines were uniform.
Sowing took place when the soil temperature
was about 8°C. In 1991, the experimental design
was a split-plot, with quinoa lines as sub-plots.
Five lines were grown at two row spacings, 25 and
50 cm, with and without hoeing. Each combina-
tion was represented by duplicate plots, giving 40
sub-plots in all. Sowing took place on 24 April. In
 S.-E. Jacobsen / Industrial Crops and Products 7 (1998) 169–174 171

Table 2
Developmental stages in quinoa (Jacobsen and Stølen, 1993)

Stage Description Stage Description Stage Description

0 Vegetative phase 8 Anthesis 14 Seed set

Onset of flowering
1 Bud formation 33% Seed set
Bud covered 9 50% Flowering 15 50% Seed set
2 Bud visible 10 100% Flowering 16 67% Seed set
3 Bud distinct 11 Floral dehiscence 17 100% Seed set
4 Bud approximately 0.5 cm long Onset 18 Maturity
5 Bud approximately 1.0 cm long 12 Majority of flowers dehisced Plant colour:
6 Onset of pyramid shape Green\yellow
19 Yellow\green
7 Distinct pyramid shape 13 Only wilted anthers present 20 Mature
21 Wilted

1992 and 1995, the experimental design was a 3. Results

randomized block design with three replicates of
five lines, giving 15 plots in all. Row spacing was
25 cm and plots were hoed to combat weeds.
Sowing took place on 6 and 3 May in 1992 and
1995, respectively. In all years nitrogen was ap-
plied as a top dressing at a rate of 120 kg/ha 2
weeks after emergence. Plot size was 15 m2
throughout.
In order to have a reliable estimate of the
number of plants, two samples were taken at
random from each plot, each sample comprising
plants harvested from a 25 cm length of row.
Within each sample the number of plants was
recorded. Developmental stage, according to the
scale shown in Table 2, was measured on five
occasions (time 1–5) at fortnightly intervals from
1 July.
For the analyses of variance a successive test
was used, in which all factors in the model were
tested sequentially. Since it had already been
demonstrated that the number of plants had an
effect on inflorescence size and height (Jacobsen,
1993), it was included as a covariate in the analy-
sis in order to reduce the error variance. Differ-
ences between lines were tested using a
Ryan–Einot–Gabriel – Welsch multiple range
test.
In 1991, the spring was wet and cold, especially
in April and June. In 1992, drought was experienced
in May and June. In 1995, the spring was very cold,
whereas July and August were dry and warm.
In 1991 and 1992 seed and fodder types developed
differently over the growing season, whereas in 1995
the effect of plant type was generally less pro-
nounced. The overall results from the three seasons
(Table 3) show that the vegetative stage was longest
in those years with cold springs, 1991 and 1995.
Anthesis, floral dehiscence and seed set periods were
longest in 1995, when the spring was cold and the
summer dry. All growth periods were short in 1992,
resulting in a total average growth period of 121
days, compared to 146 days in 1991 and 153 days
in 1995.
The developmental patterns of the lines, shown
in Fig. 1, change from year to year. The shape of
the curve for 1992 is exponential, which reflects a
rapid emergence and early growth before the ap-
pearance of drought in May–June. In 1991 and
1995 growth began later than in 1992. In the late
summer of 1995 developmental rate decreased
temporarily between 110 and 135 days after emer-
gence, due to a rainy period which kept the plants
green. Ranking of lines, however, was consistent
over the growth season.
172 S.-E. Jacobsen / Industrial Crops and Products 7 (1998) 169–174

Table 3
Ranking analysis of duration of growth stages, combined over years

Vegetative phase/bud forma- Anthesis Floral dehiscence Seed set Total

tion

0–8 8–11 11 – 14 14 – 18 0 – 18

Type 2 65.90a 2 12.58a 2 19.85a 2 51.72a 2 150.05

1 59.06b 1 8.49b 1 16.40b 1 46.00a 1 129.95
Line 210 67.86a 210 12.91a 210 20.81a 210 55.12a 210 156.70
Olav 63.94b Olav 12.25a 224 19.58a,b Olav 48.32b Olav 143.40
224 61.80c 233/240 9.66b Olav 18.89a,b 224 47.42b 224 136.00
233/240 58.44d 205 8.62b 205 15.93b,c 205 46.35b 205 127.85
205 56.95d 224 7.20b 233/240 13.68c 233/240 44.24b 233/240 126.02
Year 1991 73.26a 1995 12.33a 1995 19.47a 1995 54.87a 95 152.80
1995 66.13b 1992 10.60a 1991 18.26a 1991 46.70b 91 145.51
1992 49.95c 1991 7.29b 1992 16.15a 1992 44.55b 92 121.25

Type 1, seed producing; type 2, fodder producing. Letters a– d indicate significant differences between lines within time (PB0.05).

4. Discussion render harvesting difficult, increases in the au-

The lines tested, representing genotypes able to
mature and yield satisfactorily under North Eu-
ropean conditions, had an average total growing
period of between 126 and 157 days over the 3
years. This compares with a total growth period
of 131–200 days in Peru (Flores, 1977), while the
usual growth period in South America ranges
from 110 to 190 days (Jacobsen and Stølen, 1993).
The reason for the longer growth period in Peru
compared to Denmark is due mainly to a longer
vegetative phase, though anthesis and seed set are
also longer in Peru. Risi and Galwey (1989b)
suggest that the bud formation period is first to
respond to differences in day length. Thus, results
from Britain and Denmark indicate that variation
for earliness leads to corresponding variability in
the length of the vegetative phase. Flowering also
extends over a longer period in South America
than was observed in Denmark (Rea, 1948; Gan-
darillas, 1979).
Usually plant breeding seeks to improve well-
adapted varieties. In this case, however, when a
crop is being introduced to a new region, substan-
tial modification may be required. First of all, in
quinoa, the ideal variety for seed production
would be uniformly early maturing. This is partic-
ularly important for North European conditions,
where the risk of cold, humid weather, which will
tumn. A growing period longer than 150 days
would therefore normally be regarded as too
risky. Nevertheless, in the three years under inves-
tigation it was possible to harvest the crop even
with a growing period up to 1 month longer.
Besides earliness, quinoa should have a consis-
tently high seed yield and a low saponin content
(Jacobsen et al., 1996). It should also be short and
non-branching, to facilitate mechanical harvest-
ing. Size, shape and compactness of the inflores-
cence may be important for the rate of
maturation. A large, open inflorescence will dry
quicker after rain and morning dew than a small,
compact one, but it may also be more prone to
seed scattering. Fodder types, on the other hand,
should be tall, leafy and late-maturing, with a
high dry-matter yield and preferably a low sa-
ponin content. Genotypes for this purpose will
probably not be grown in northern Europe be-
cause of their late maturity.
As quinoa is predominantly an inbreeding spe-
cies, breeding and selection programmes such as
those commonly used in cereals could be adopted.
Moreover, previous experiments have shown that
considerable variation exists between lines for
many of the desired characters. When considering
the level of adaptation, many analytical tech-
niques are available, one of which, suggested by
Lin and Binns (1988) and modified by Jacobsen et
 S.-E. Jacobsen / Industrial Crops and Products 7 (1998) 169–174 173

Fig. 1. Developmental stage, 1991 – 1995; dae, days after emergence.

al. (1996), requires the calculation of a distance
mean square across environments between the
candidate genotype and an optimum value. Those
genotypes whose overall mean for a certain char-
acter does not depart significantly from the opti-
mum and which do not have significant
heterogeneity across environments, can be recom-
mended as being generally adapted for a whole
region. Genotypes which exhibit specific adapta-
tion are identified by a low distance mean square
in specific environments. In quinoa, one of the
lines tested in Denmark exhibited general adapta-
tion for earliness and other characters, while other
lines exhibited general adaptation for individual
characters (Jacobsen et al., 1996).
Potential parents for any future crossing pro-
grammes have been identified. After making the
crosses, the best procedure would probably be to
develop recombinant inbred lines by single seed
descent and defer selection to the F6 or even later
generations, as suggested by Jinks and Pooni
(1981). By this stage the effects of dominance can
safely be ignored and much of the genetic varia-
tion between the recombinant lines will be due to
additive effects of the genes concerned. Thus by a
combination of appropriate breeding and selec-
tion techniques it will hopefully be possible to
combine many of the desired characteristics in a
single genotype, which in turn could establish
quinoa as a novel crop for European agriculture.
174 S.-E. Jacobsen / Industrial Crops and Products 7 (1998) 169–174
Acknowledgements
The author wish to acknowledge Dr John Hill
for commenting on the script.
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