Table of Contents
Vollmann, J., A. Damboeck, A. Eckl, H. Schrems, and P.
Ruckenbauer. 1996. Improvement of Camelina sativa, an underexploited
oilseed. p. 357-362. In: J. Janick (ed.), Progress in new crops. ASHS Press,
Improvement of Camelina sativa, an Underexploited Oilseed
Johann Vollmann, Astrid Damboeck, Anna Eckl, Heinrich Schrems, and Peter
- MATERIALS AND METHODS
- Genetic Entries
- Performance Trials
- Selection for Large Seed Size
- Environmental Effects
- Genotype Performance
- Selection for Seed Size
- Table 1
- Table 2
- Fig. 1
- Fig. 2
- Fig. 3
- Fig. 4
- Fig. 5
- Fig. 6
Camelina [Camelina sativa (L.) Crtz., Brassicaceae], known as
false flax or gold-of-pleasure is a spring-planted crop species. Although
camelina has been cultivated in Europe since the Bronze Age (Schultze-Motel
1979), it is an underexploited oilseed crop at present. Recent interest in the
species is mainly due to the demand for alternative low-input oilseed crops
with the potential for a non-food utilization of the seed oil (Seehuber 1984;
Putnam et al. 1993). Camelina oil has a unique fatty acid pattern and is
characterized by a linolenic acid (C18:3) content of 30% to 40% and an
eicosenic acid (C20:1) content of around 15%, with less than 4% erucic acid
(Seehuber 1984; Marquard and Kuhlmann 1986; Budin et al. 1995), which suggests
a utilization of the seed oil as a drying oil with environmentally safe
painting and coating applications similarly to linseed oil (Luehs and Friedt,
1993). Moreover, a rather low content of glucosinolates has been found in
camelina as compared to other brassicaceous species (Lange et al. 1995), which
makes the utilization of meals easier.
Agronomic investigations of camelina have been carried out both in Europe
(Zimmermann and Kuechler 1961; Marquard and Kuhlmann 1986) and North America
(Plessers et al. 1962; Robinson 1987). Unique agronomic features such as the
compatibility with reduced tillage and cover crops, competitiveness with weeds
or winter surface seeding have been emphasized by Putnam et al. (1993) with
regard to the suitability of camelina for sustainable agriculture systems.
Furthermore, the species has a potential as a low-cost crop for green-manuring.
Breeding research and genetic improvement of camelina have been initiated in
Germany during the 1980s; apart from the collection and evaluation of germplasm
(Seehuber 1984), agronomically advanced lines have been developed (Seehuber et
al. 1987), which are available as base population for further improvement
(Seehuber and Dambroth 1987).
In the present investigation, results from an agronomic evaluation of new lines
of camelina are reported. The genotypes tested were derived from a
recombination program, which mainly focused on the improvement of agronomic
performance of camelina as an alternative and low-input oilseed crop for
Camelina populations CA13X and CA2X were derived from crosses BGRC 51558 x BGRC
51572 and Voronezskij 339 x BGRC 51558, respectively. BGRC genotypes were
obtained from the genetic resources collection of the FAL Institute of Agronomy
(Braunschweig, Germany), Voronezskij 339 is an accession obtained from the N.I.
Vavilov Institute (St. Petersburg, Russia). Segregating populations were
advanced to the F4-generation by a bulk breeding method. In the F4, single
plants were harvested separately and F4-derived lines were selected visually
from F5-single row plots for further testing.
Homozygous breeding lines from the populations described above were evaluated
in 1993 (F4-6) and 1994 (F4-7) at the two Austrian locations Gross Enzersdorf
(10 km east of Vienna, 48°12' N, 16°32' E) and Reichersberg (60 km west
of Linz, 48°20' N, 13°20' E) in replicated field trials. Cv. Iwan
(Saatbau Linz, Reichersberg, Austria) was used as a check cultivar. Plot size
was 5 x 1.25 m in Gross Enzersdorf and 8 x 1.25 m in Reichersberg.
Agronomic parameters were set as described by Seehuber et al. (1987). All
experiments were arranged as lattice designs in two replications. As
particular genotypes were discarded after the 1993 experiments due to the
necessity of selection, a complete set of data was available for 10 genotypes
from both locations and for both years for a combined analysis of variance.
Another set of 32 genotypes was analyzed across 3 environments (Gross
Enzersdorf 1993, 1994; Reichersberg 1994). In general, adjusted entry means
from the lattice analysis were used for the combined analyses of variance; in
the experiment at Gross Enzersdorf 1993, seed yield data were adjusted by a
neighbor analysis procedure (Vollmann et al. 1996), which was most efficient in
controlling the spatial variation present in the particular trial field. In
order to identify genotypes superior to the check cultivar, LSI (= least
significant increase, one-tailed t-statistic) was used as a statistical testing
procedure (Petersen 1994). Oil content of seed was determined
non-destructively by near-infrared reflectance spectroscopy (NIRS) and is
reported in g kg-1 on a dry weight basis.
For an improvement of seed size (mass), sub-populations of crosses CA2X and
CA13X were established by sieving out small fractions (< 0.1 %) of largest
seeds from the F2- (CA2X-1S, CA13X-1S) or from both the F2- and F3-bulks
(CA2X-2S, CA13X-2S). F4-derived lines of these sub-populations were evaluated
for 1000-seed weight in the F5-generation using single row plots with two
replications (Gross Enzersdorf 1992) and a selected number of large-seeded
genotypes was tested for agronomic performance (Gross Enzersdorf 1993) as
In the set of 10 genotypes of camelina evaluated in two locations for two
years, both seed yield and oil content were highly influenced by year and
location effects as well as by a highly significant year x location
interaction. Genotype effects were also significant, but genotype x year and
genotype x location interactions were of minor importance for seed yield and
not significant for oil content indicating a similar response of the different
genotypes to environmental changes. Seed yields were in the range of 1050 to
1700 kg ha-1 in 1993 and from 1450 up to 3250 kg ha-1 in
1994. In both locations, oil contents were high in 1993 (400 to 455 g
kg-1) and clearly lower in 1994 (385 to 425 g kg-1), as
shown in Fig. 1.
Genetic variation in different agronomic characters was highly significant
(F-test) in the set of 32 genotypes tested across 3 environments (Gross
Enzersdorf 1993, 1994; Reichersberg 1994). The check cultivar was
significantly outyielded in both seed and oil yield by several of the new
genotypes evaluated (Table 1), an improved oil content of seed was however
found in only one genotype. In the populations investigated, seed yield and
oil content were positively correlated (Fig. 2), whereas a tightly negative
correlation was found between 1000-seed weight and oil content (Fig. 3); a
negative correlation was also detected between 1000-seed weight and yield. For
the genotypes listed in Table 1, estimates of heritability were rather high in
characters such as oil content or 1000-seed weight and lower for plant height
In particular performance trials (Gross Enzersdorf 1993), spatial variation in
the trial field affected oil content (apart from the influence on seed yield)
in the range of -8 to +7 g kg-1 seed oil. A clear trend in
variation of oil content was found using a neighbor analysis covariate (Fig. 4), which was also helpful in order to adjust individual plot values for field
heterogeneity during analysis of variance (results not shown).
In population CA2X, seed size of F4-5-lines was clearly improved by sieving of
F2- or F3-bulks and lines with a 1000-seed weight of up to 2 g were identified
within sub-population CA2X-2S (Fig. 5), whereas 1000-seed weights of the
respective parent genotypes were only 0.9 and 1.2 g. In population CA13X,
selection response was less pronounced with respect to seed weight. In a
preliminary assessment of agronomic performance of large-seeded genotypes (96
F4-6-lines, Gross Enzersdorf 1993), seed yield was positively affected by
1000-seed weights of up to 1.5 g, whereas seed yield was reduced in genotypes
with larger seed size (Fig. 6). Moreover, seed size and oil content were
negatively correlated, and large-seeded genotypes had a reduced number of seeds
per pod, malformation of pods as well as higher lodging scores (data not
A considerable agronomic potential has been detected in newly developed lines
of camelina during the present study. Furthermore, strong year x location
interactions for seed yield and oil content indicated that specific weather
conditions within a location could modify performance considerably. In the
1993 experiments, a severe drought during the flowering phase limited plant
development and yield potential at both locations, whereas sufficient rainfall
during the seed filling period resulted in the subsequent expression of high
oil contents (Fig. 1). In the 1994 experiments, water conditions were not
limiting plant development throughout the growing season, which allowed seed
yields of up to 3250 kg ha-1, whereas oil contents were lower as a
response to the high yield potential. Similar ranges of seed yield and oil
content have also been reported from experiments in Germany (Seehuber 1984;
Seehuber et al. 1987), whereas both seed yield and oil content were lower in
North American studies (Plessers et al. 1962; Robinson 1987; Putnam et al.
1993), which might partly be due to a lack of environmental adaptation. The
strong influence of environmental conditions on seed yield has also been
reported for other spring-sown oilseed crops (Diepenbrock et al. 1995).
Sufficient genetic variation was present within the germplasm available, which
allowed the selection of lines with improved agronomic features (Table 1 and
2). A broad genetic variation and moderately high heritabilities of important
agronomic characters have also been identified by Seehuber et al. (1987)
investigating similar genetic materials. The positive correlation between seed
yield and oil content indicates that a simultaneous improvement of both
characters is possible. An improvement of seed oil content would be of
particular interest to make camelina more competitive to other oilseed crops
exhibiting higher oil contents (Plessers et al. 1962; Putnam et al. 1993).
Screening of large numbers of entries for oil content can be accomplished
easily by non-destructive techniques such as near-infrared reflectance
spectroscopy or nuclear magnetic resonance (Thies and McGregor 1989); the
presence of spatial field variations affecting oil content (Fig. 4) can however
reduce the efficiency of selection if not controlled properly (Ball et al.
1993; Vollmann et al. 1996).
In camelina, improvement of seed size would be of interest for a rapid field
emergence and crop establishment under less favorable growing conditions;
moreover, solvent extraction of oil is generally more efficient with
large-seeded cultivars. A considerable increase of 1000-seed weight of up to 2
g has been achieved during the present selection experiment, as compared to the
unselected control population (Fig. 5) as well as to variation in seed size
reported previously (Seehuber 1984; Seehuber et al. 1987; Putnam et al. 1993).
However, the obtained improvement of 1000-seed weight seems to be of low
immediate value due to the drastic reduction of both oil content and seed
yield, which makes further cycles of recombination necessary. Within smaller
ranges of seed size, a significant correlation betweeen seed size and oil
content was not found in camelina (Seehuber 1984), whereas positive
correlations between 1000-seed weight and oil content have been reported for
particular crosses in oilseed rape (Engqvist and Becker 1993).
The results obtained from the present study suggest that a considerable
agronomic potential is present in newly developed lines of camelina. However,
high oil content and a specific fatty acid profile would be the key
requirements to re-establish camelina as an industrial oilseed crop. These
quality characters should therefore deserve most attention in future breeding
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of camelina (Camelina sativa L. Crantz) seeds and oils. J. Am. Oil Chem.
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seed quality of linseed dodder (Camelina sativa Crtz.).
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promising low-input oilseed. p. 314-322. In: J. Janick and J.E. Simon (eds.),
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sativa (L.) Crantz. Archaeo-Physika 8:267-281.
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single-seed-descent method in false flax to increase the yield level.
Landbauforsch. Voelkenrode (Germany) 37:132-136.
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content and fatty acid composition. p. 132-164. In: G. Roebbelen, R.K. Downey,
and A. Ashri (eds.), Oil crops of the world, their breeding and utilization.
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Table 1. Agronomic performance of 32 genotypes of camelina (mean values
across 3 environments).
|No. ||Genotype name ||Seed yield (kg ha-1) ||Oil content (g kg-1) ||Oil yield (kg ha-1) ||1000-seed weight (g) ||Plant height (cm)|
|1 ||CA13X-11 ||2091 ||413.3 ||858 ||1.26 ||72.5|
|2 ||CA13X-17 ||2381 ||424.0 ||1002 ||1.21 ||70.0|
|3 ||CA13X-20 ||2032 ||425.7 ||861 ||1.16 ||64.2|
|4 ||CA13X-6 ||2143 ||427.4 ||911 ||1.16 ||71.7|
|5 ||CA13X-1S-9 ||2392 ||424.9 ||1011 ||1.13 ||79.2|
|6 ||CA13X-2S-23 ||1947 ||427.7 ||828 ||1.21 ||69.2|
|7 ||CA13X-2S-44 ||2210 ||423.1 ||927 ||1.19 ||75.0|
|8 ||CA13X-2S-96 ||2370 ||400.7 ||940 ||1.34 ||75.8|
|9 ||CA2X-2S-29 ||2040 ||405.1 ||820 ||1.28 ||70.8|
|10 ||Iwan (check) ||2029 ||422.9 ||853 ||1.29 ||71.7|
|11 ||CA13X-13 ||1965 ||434.9 ||847 ||1.20 ||72.5|
|12 ||CA13X-1S-19 ||1951 ||411.6 ||797 ||1.28 ||78.3|
|13 ||CA13X-2S-17 ||1799 ||410.9 ||735 ||1.35 ||70.8|
|14 ||CA13X-2S-20 ||1818 ||404.0 ||731 ||1.43 ||76.7|
|15 ||CA13X-2S-22 ||1746 ||393.5 ||677 ||1.41 ||70.8|
|16 ||CA13X-2S-26 ||1904 ||415.0 ||783 ||1.24 ||74.2|
|17 ||CA13X-2S-29 ||2041 ||408.4 ||824 ||1.32 ||75.0|
|18 ||CA13X-2S-4 ||1692 ||414.3 ||698 ||1.31 ||70.0|
|19 ||CA13X-2S-53 ||1945 ||424.2 ||822 ||1.26 ||70.8|
|20 ||CA13X-2S-56 ||1746 ||423.1 ||733 ||1.22 ||67.5|
|21 ||CA13X-2S-6 ||1775 ||405.4 ||715 ||1.42 ||75.0|
|22 ||CA13X-2S-63 ||1898 ||415.2 ||786 ||1.40 ||71.7|
|23 ||CA13X-2S-69 ||1953 ||401.7 ||782 ||1.47 ||73.3|
|24 ||CA13X-2S-7 ||2025 ||411.7 ||828 ||1.34 ||73.3|
|25 ||CA13X-2S-75 ||2009 ||410.3 ||819 ||1.31 ||74.2|
|26 ||CA13X-2S-85 ||1821 ||410.4 ||747 ||1.33 ||70.8|
|27 ||CA13X-2S-95 ||1797 ||399.8 ||714 ||1.36 ||70.8|
|28 ||CA2X-1S-1 ||1622 ||383.5 ||617 ||1.68 ||72.5|
|29 ||CA2X-1S-14 ||1888 ||400.3 ||769 ||1.47 ||67.4|
|30 ||CA2X-2S-17 ||1663 ||381.8 ||633 ||1.62 ||72.5|
|31 ||CA2X-2S-20 ||1605 ||378.0 ||604 ||1.66 ||70.2|
|32 ||CA2X-2S-22 ||1733 ||388.1 ||661 ||1.57 ||72.5|
| ||Total mean ||1939 ||410.0 ||792 ||1.34 ||72.0|
| ||LSD 0.05 ||215 ||8.8 ||88 ||0.06 ||5.0|
| ||LSI 0.05 ||183 ||7.5 ||75 ||0.05 ||4.5|
Table 2. Estimates of heritability calculated from components of
variance for different agronomic characters of camelina (32 genotypes tested in
|Character ||Heritability (%)|
|Seed yield ||86.5|
|Oil content ||95.6|
|Oil yield ||90.5|
|1000-seed weight ||97.6|
|Plant height ||62.5|
||Fig. 1. Oil content of a set of 10 genotypes of camelina as affected by environmental conditions (locations: GE = Gross Enzersdorf, RE = Reichersberg; years: 1993, 1994).
||Fig. 2. Relationship between seed yield and oil content in 32 genotypes
of camelina (genotype mean values across 3 environments).
||Fig. 3. Relationship between 1000-seed weight and oil content in 32
genotypes of camelina (genotype mean values across 3 environments).
||Fig. 4. Trends of oil content due to spatial variation in the trial
field (Gross Enzersdorf 1993) as visualized by a neighbour analysis covariate
||Fig. 5. Distributions of seed size (1000-seed weight) in F4-5 lines of
population CA2X. a: unselected control population (n = 25 lines); b: F2-bulk
sieved for large-seededness (n = 25); c: both F2- and F3-bulks sieved (n =
||Fig. 6. Relationship between 1000-seed weight and seed yield in a set
of 96 genotypes selected for large seed size (environment: Gross Enzersdorf
1993; regression line and coefficient of correlation according to 2nd order
Last update August 21, 1997