沙漠研究 22-1, 147 -151 (2012 )"ICAL 1 / DT X"

Journal of Arid Land Studies－Refereed Paper－

Salt Tolerance of Oilseed Crops during Establishment
Seiichi MIYAMOTO*1), Mike FOSTER2), Calvin TROSTLE3) and Edward GLENN4)
Abstract: Bioenergy production in arid and semi-arid regions is viewed as being limited due to water resource constraints andpotential competition with food production. However, there are crop rotation niches as well as opportunities for utilizing saline soils
and water which are not conducive to high value crop production. An exploratory study was thus conducted in a greenhouse forassessing salt tolerance of various oilseed crops during establishment, which is often the critical stage for successful production.
Canola (Brassica napus), and safflower (Carthamus tinctorius) were salt-tolerant during germination, but emergence was curtailed
owing to soil crusting. Camelina (C. sativa) germinated well, yet emergence was poor, probably due to weak hypocotyl. Field
pennycress (Thlaspi arvense) and Lesquerella (L. fendleri) were salt sensitive, and could not germinate in NaCl solutions higher than
50 mM. Salicornia (S. bigelovii), a halophyte, is extremely salt-tolerant once established, but not during seedling emergence.
Seedling emergence which had little correlation with salt tolerance of established plants, was constrained not only by salt tolerance atgermination and the pattern of salt accumulation at the soil surface, but also by seed size and soil crust development. At the present
state of field management capability, safflower and canola are among the most promising oilseed crops for saline areas, and canpotentially be grown as a winter rotation crop with a comparatively low water requirement. For the species with small seed, crop
improvements towards greater seedling vigor as well as the effective establishment methods have to be developed.
Key Words: Bioenergy, Oilseed, Saline water, Salt tolerance

1. Introduction
Increasing concern over energy supply, especially oftransportation fuel, the carbon balance, as well as theincreasing need to better manage saline soils and water has
generated an interest in salt tolerant species which can be usedas feedstock or fuelwood. Oilseed crops, especially canolaand safflower are among the crops which can readily produce
transportation fuel. In the American West, these crops fit in therotation niches as winter crops, and their water requirement iscomparatively low. These crops are also known to besalt-tolerant once established (Table 1). Soil salinity which
may cause a 15% yield reduction of canola is 12 dS m-1 in thesaturation extract (Francois, 1994), and that for safflower, 9.3
dS m-1 (Francois and Bernstein, 1964). These values can becompared against that for cotton (11 dS m-1), which is among
the most salt-tolerant crops commercially grown. Camelina
with a low fertility requirement and a short growing season
may have potential for biodiesel production (e.g., Gugel andFalk, 2006). Field pennycress has high contents of erucic andlinoleic acids, and grows wild in Northern states and Canada(Moser, 2009). Lesquerella (L. fendleri) is a potential crop to
produce engine lubricants (Adam et al., 2007), and unlike
castor (Ricinus communis), it does not contain toxic substances.
Salt tolerance of established Lesquerella is rated “moderately

tolerant,” 6.9 dS m-1 (Grieve et al., 1997). Salt tolerance ofCamelina and pennycress is unknown. Salicornia is a
halophyte native to estuaries, and was grown with seawater at a
30% leaching fraction in lysimeter (Glenn et al., 1997). It isamong a few halophytes which can be grown with highly
saline or sea water (Glenn et al., 1997; Grattan, et al., 2008).
The seed of this plant is very small (<0.05 g/100 seed), and its
salt tolerance during seedling establishment is unknown.
Difficulties of establishing field crops in saline soils werepointed out years ago (e.g., Bernstein and Fireman, 1957).
The primary cause of poor emergence was attributed to salt
accumulation in crop beds where seed is usually placed.
Difficulties of establishing certain vegetable crops with waterof elevated salinity were shown to be hypocotyl mortality by
the salts deposited at the soil surface (Miyamoto et al., 1985) or
seedling kills by salts splashed during passing showers, but notdue to poor germination (Miyamoto et al., 1986). Field
experience in far West Texas has shown difficulties withestablishing oilseed crops using water having the electricalconductivity (EC) of 4 to 5 dS m-1 (personal communication).
The primary objective of this study was to examine salt
tolerance of promising oilseed crops during germination andemergence. Another objective was to explore strategies for
improving seedling establishment.

* Corresponding Author: s-miyamoto@tamu.edu
1380 A&M Cir., El Paso, TX 79927, USA
1) Texas A&M University AgriLife Research Center at El Paso

3) Texas AgriLife Extension at Lubbock, Texas A&M University

2) Texas A&M University Agricultural Research Station at Pecos

4) Environmental Research Laboratory, University of Arizona

Table 1. The oilseedp crops tested.
1┘

CommonScientific ECe
Name
Name
15%
11.9
Canola (Brassica napus )Safflower (Carthamus
9.3
tinctorious)
Camelina (C. sativa )?
Pennycress (T. arvense )?
Lesquerella (L. fendleri )6.9
Salicornia (S. bigelovii )>30.0

Cultivars tested
for this study 2┘
DKW 45-10, 47-15
PI 485984, 406002
CW99OL
BSX-WG1, WG3
Patton, USDA W12
USDA, A3219, 01L0
U of A selection

1┘

Salinity of the soil saturation extract which causes
a 15% reduction in growth or yields.
2┘
DKW, Dekalab; PI, USDA lines; CW, Cal West,
BSK, Bluesun;

2. Material and Methods
2.1. Germination experiment
Seed germination of cultivars shown in Table 1 wasobserved in petri-dishes containing various saline solutions.
The temperature of the incubator was 23 ± 2°C, except for
Salicornia which required a higher temperature of 8 °C (Rivers
and Weber, 1971). The saline solutions were prepared byadding NaCl to tap water (0.8 dS m-1) at three rates, 100, 200,
and 300 me L-1. In addition, 100 me L-1 of CaCl2 andNa2SO4 were added for assessing salt type effects. For
Lesquerella and pennycress only, 50 me L-1 saline solutions
were added. Sterilized glass petri-dishes with a sheet of afilter paper contained 5 ml of each solution which was changed
every other day. Seed germination was observed in triplicatesusing 50 seeds per dish. The seed was considered germinatedwhen the length of radicle exceeded that of the seed. The
length of hypocotyl was also measured using five randomly
selected samples, after the incubation period of 5 to 7 days.
2.2. Seedling emergence and mortality experiments
A preliminary experiment was conducted in a greenhouse
which was regulated at 15°C at night and 30°C during day
hours. It was for determining appropriate seeding depths
under two watering methods; one subirrigated by placing
potted soils in a shallow pan of water, and another irrigatedfrom the top. Three soil types were used; Bluepoint loamy
sand, Harkey silt loam, and Glendale silty clay (Typic
Torrifluvents).
The electrical conductivity of the soil
saturation extract (ECe) of these soil samples were 0.8, 2.2 and1.2 dS m-1, respectively. Soil samples (<4 mm) were placedin greenhouse pots (15 cm in diameter and depth). Seed (20each per pot) was placed on a smoothed soil surface, and wascovered with the soil which provides a thickness of 0.25, 0.5,
1.0, and 2.0 cm. The entire greenhouse was covered with ashade cloth which provides 70% light transmission. The pan
evaporation in the greenhouse averaged 7 mm/day or the
potential evaporation rate of 5 mm/day. Irrigation water wasapplied once a week at 4.5 cm per application. Seedlings thatemerged were counted daily or bi-daily.
Subsequently, two experiments were conducted. The first
one utilized Harkey silt loam with three levels of soil salinity(1.2, 3.8 and 6.8 dS m-1), and tap water for irrigation. The
second experiment used three saline solutions with the EC of
4.1, 8.2 and 12.1 dS m-1. The saline water was prepared byadding NaCl, CaCl2 and Na2SO4 to tap water at the chemical
equivalent ratio of 2:1:1 to yield the total salt concentration of40, 80 and 120 me L-1. All other procedures used were thesame as those used for the preliminary experiment, except for
irrigation. The irrigation frequency was increased to twice a
week, but using the same amount of water per week, except for
Salicornia which was watered daily. Seedlings which haveemerged as well as experienced mortality were recorded.
One week to 10 days after emergence, seedlings were
sprayed with NaCl solutions having concentrations of 100, 200,
300 and 400 mM, until completely wet. The sprayedseedlings were placed in humidity and temperature controlled
glass chamber (85% humidity at 22°C). Seedling mortality
was then observed for a week, and this process was repeated
for another week.
The analysis of variance (ANOVA) was performed for
germination and seedling emergence data using a split plot
design (Steele and Torrier, 1960). The mean separation was
performed at a 5% level.
2.3 Soil Salinity and Strength Measurements
Additional potted soils were prepared for measuring soil
salinity and strength. These pots were irrigated with thesaline solutions mentioned above, one set under subirrigation,
and another set top-irrigated. Soil strength was measured infour replicates on 6th day into the experiment with a soil
penetrometer. The potted soils were then sectioned to 0-0.5,
0.5-1, 1-3, and 3-5 cm. The samples in duplicate were analyzed
for salinity of the saturation extract (Rhoades and Miyamoto,1990).

3. Results
3.1. Germination
The final germination counts as well as the days required toobtain 80% of the final counts are shown in Table 2. Becauseof the space limitation, only the cultivar which provided thebest germination are shown. Both canola and safflower
germinated in 2 to 4 days when incubated in tap water and in
100 mM solutions. In 200 mM solution, seed germination
slowed (4 to 5 days), and the final germination percentage was
reduced to 40 to 50%. No germination occurred at 300 mM.
Camelina also germinated in 2 days, but the germination inTable 2. The final germination percentage, and the days required toobtain 80% of the final germination.

Table 3. Seed weight, hypocotyl length and appropriate seedling
depth when subirrigated with low salt water.

Crops Treatments Tap
NaCl
CaCl2 Na2SO4
Conc. (me L-1) 8
50 100 200 100
100-------------------------------------------------------------------------------Canola (DKW 45-10)
2
Germ. (%)
70a - 57a ┘ 47b 60a
63a
Days1┘
2.7
2.8 3.9 3.2
2.8
Safflower (485984)
Germ. (%)
82a - 63b 39c 65b
62b
Days
4
4.1 4.9 5.1
4.5
Camelina (BSX-WG1)
Germ. (%)
88a - 81a 23c 75b
73b
Days
1.9
2 5.8 2.4
2Pennycress (Patton)
Germ. (%)
100a 68b 5e
0
40c
30d
Days
3
4.5
8
9
(50) 3┘ (50) 3┘
Lesquerella (01L0)
Germ. (%)
58a 30c 11d 0
46b
32c
Days
4
4
5
7
7
NaCl (me L-1)
Salicornia (U of A)
300 400 500
600
Germ. (%)
90a - 80b 70c 40d
28e
Days
5
6.4 6.6 7.2
7.8
1┘Days required for attaining 80% of the final germination
counts2┘The numbers in columns followed by the same letter are
not significantly different
3┘The numbers in parenthesis are concentrations of CaCl2
and Na2SO4

Seed Hypocotyl length1┘ Optimum
weight Tap NaCl (100) Depths
g/ 100 ------------------ mm -----------------Canola (DKW 45-10)
0.39
28
8
2.5 - 10
Safflower (PI485984)
2.79
32
22
2.5 - 10
Camelina (BSX-WG1) 0.10
226
2.5 - 5
Pennycress (Patton)
0.09
23
2
2.5 - 5Lesquerella (01L0)
0.07
11
4
2.5 - 5
Salicornia (U of A)
0.03
22
0.0 - 2.5
1┘The length of hypocotyl measured after 5 to 7 daysof incubation

200 mM solution was reduced to 23%. Pennycress
germinated in 50 mM NaCl solution, but little in 100 mM
solutions. It germinated better in CaCl2 or Na2SO4 solutions
than in NaCl solutions at 100 me L-1. Lesquerella had thegermination pattern similar to pennycress. Salicornia seedgerminated in 5 to 6 days in tap water as well as in 300 mM
NaCl solutions. However, germination was reduced andslowed with increasing salinity (Table 2). The germination ofcultivars not listed in Table 2 varied, but the difference among
the tested cultivars became mostly insignificant whennormalized by the germination counts from tap water.
3.2. Seedling Emergence and Mortality
The preliminary test conducted has shown that seedlingemergence when subirrigated with tap water was largely
independent of the seeding depths examined as long as it waswithin the range specified in Table 3. This table also includes
the length of hypocotyl measured after 5 to 7 days of
incubation in tap water as well as in a 100 me L-1 NaClsolution during the germination study. The optimum seeding
depth for emergence appeared to be less than the length of
hypocotyl, and usually less than 1 cm, but not as deep as 2.0
cm. Camelina, pennycress and Lesquerella have to be seeded
shallow, 0.25 to 0.5 cm, and even shallower for Salicornia.
The seed size of these species is too small to yield long
hypocotyl.
Seedling emergence of canola and safflower from salinizedcultivars

Table 4. Seedling emergence from saline soils with seeds placed atthree depth, and irrigated with tap water.
g
p
Crops
Emergence: Seeded at
2┘
Irrigation Soil Days 0.25 cm 0.5 cm 1.0 cm
method ECe1┘------------------%-----------------Canola (DKW 45-10)60a2┘ 55ab
Sub
1.2
4-5
55b
irrig.
3.8
5-6
25d
60a
70a
6.4
8-7
15e
35c
40c
Top
1.2
4-5
45c
55ab
50b
3.8
4-5
irrig.
50bc
50ab
10d
6.4
4-5
60a
60a
15d
Safflower (PI 485984)
Sub
1.2
4-5
60ab
80a
80a
irrig.
3.8
5-6
65a
75b
60c
6.4
6-7
55b
60c
40d
Top
1.2
4-5
55b
60c
75ab
irrig.
3.8
5-6
55b
65c
60c
6.4
6-7
65a
60c
65c
Camelina (BSX WG1)
Sub
1.2
5-6
20c
30ab
0
irrig.
3.8
6-7
20cd
30ab
0
6.4
7-9
10d
30ab
0
Top
1.2
5-6
35b
25b
0
irrig.
3.8
6-7
25c
30ab
0
6.4
7-9
55a
40a
0
1┘

Salinity of the soil saturation extract
The number in columns followed by the same letterare not significantly different

2┘silt loam subirrigated with tap water was significantly lower
when seeded at 0.25 cm as compared to seeding at 0.5 or 1.0
cm. When the salinized soils were top-irrigated, however,canola planted at 0.25 cm emerged better than subirrigated case.
Seedling emergence of safflower, when subirrigated, was not
reduced significantly when initial soil salinity was 3.8 dS m-1,
but did so at 6.4 dS m-1 (Table 4). Seedling emergence of
canola from salinized silt loam decreased significantly at 3.8
dS m-1 when seeded at 0.25 cm, then subirrigated. Seedlingemergence of Camelina was poor (<30%), regardless of theirrigation methods tested.
Seedling emergence of canola decreased significantly
when salinity of irrigation water used increased beyond either
4.1 or 8.2 dS m-1, depending on the irrigation methods used
(Table 5), (Seedling emergence data of canola and safflower
at4.1 dS m-1 are omitted as they were similar to that from tap
water). Top-irrigation suppressed emergence in canola,Table 5. Seedling emergence from potted soils irrigated with various
saline solutions under two irrigation methods at a seeding
depth of 0.5 cm.

Water Days1┘

Crops

ECw
(dS m-1)
0.8
8.2
12.1
0.8
8.2
12.1
0.8
4.1
8.2
0.8
4.1
0.8
4.1
0.8
4.1

CanolaSafflower

Camelina

Pennycress
Lesquerella
Salicornia

Water
EC
dS m-1

Emergence

silt loam
clay loam
Sub Top
Top
Sub------------------- % -----------------85a 40a
45a
65a
4-5
55b 10b
10c
5-6
65a
50b
0b
0c
30b
6-7
75a 50a45a
4-5
85a
75a 10b
75a
20b
6-7
0b
40b
0c
7-8
50b
50a 10a
30a
20a
5-6
0b
40a
15a
30a
6-7
0b
20b
0b
20b
7-9
75a 80a70a10-12 50a
0b
0b
0b
0b
20a 10a10a
14-20 20a
0b
0b
0b
0b
0
10a
30a
0
18-23
30a
0
10a
0
18-23

1┘

Days required for attainting 80% of the final emergence
under subirrigation
2┘
Numbers in column followed by the same letter are notsignificantly different at a 5% levelsafflower and Camelina. Seedling emergence from Glendale
silty clay was better than from the silt loam, except for
Lesquerella and Salicornia (Table 5).
Seedlings under top-irrigation emerged almost exclusively
through soil cracks.
None of the seedlings emerged have died, except for a few
seedlings of safflower. Likewise, no seedling mortality was
noted when seedlings were sprayed with the saline solutions up
to 400 mM.
3.3 Salinity Distribution and Soil Strength
Soil salinity measured 6 days into the experiment are
shown in Figure 1. The data are from Harkey silt loam (ECeof 2.4 dS m-1), and those for the silty clay loam were similar,
and are not shown. Salt accumulation occurred mostly in the
top 0.5 cm.
The soil strength measured on the 6th day averaged 1 and 5kg/cm2 in sub and top-irrigated silty clay respectively, and 1
and 7 kg/cm2 in silt loam. There was no significant effect ofwater salinity on soil strength (Table 6).

A

Table 6. Soil strength of Harkey salt loam and Glendale silty clay
sud or top-irrigated with saline solutions.

B

Harkey silt loam

Fig. 1. Salinity of the saturation extract (ECe) measured 3 days after
previous top-irrigation (A) and 6 days after subirrigation (B).

0.8
4.1
8.2

Soil strength
Silt loam
Clay loam
sub
top
sub
top
-------------------------- kg/cm2 ------------------------1.0a
4.5b
1.0a
6.5b
1.0a
5.0b
7.0b
1.0a
1.0a
7.0b
1.0a
4.5b

┘The numbers in row followed by the same letter are notsignificantly different.

1

4. Discussion
Traditionally, salt tolerance of agricultural crops has been
evaluated after plants were established with water of low
salinity. This protocol is not always consistent with fieldpractices. In the present study, seedling emergence of canola
was reduced at ECe of 3.8 dS m-1 (Table 4), while establishedcanola plants were shown to tolerate soil salinity of as high as
11.9 dS m-1 (Table 1). Likewise, Lesquerella failed toestablish in soils with ECe of as low as 1.2 dS m-1 (Table 4),although salt tolerance of established plants was shown to be
6.9 dS m-1 (Table 1). This conjuncture also applies to
Salicornia. These findings indicate that salt tolerance of
established plants is not a good indicator of seedlingestablishment in saline areas.
Seedlings of certain plants are known to have salt tolerance
which is different from established plants. Salicornia seems
to fall into this category. The development of succulent tissue
did not take place until seedlings had emerged. The slow rateof hypocotyl growth of Salicornia can also compound the
problem as it allows for a high degree of salt accumulation
when subirrigated.
The species tested did not suffer seedling kills, even whensprayed with a 400 mM of NaCl solution. Salt-sensitive
species can undergo mortality when sprayed once with saline
solutions at the concentration as low as 100 mM (Miyamoto, etal., 1986). Brassica sp as well as safflower seedlings have theleaves which repel water, a trait desirable for establishment insaline areas.
The salt accumulated at the soil surface might haveaffected seed germination, especially in pennycress andLesquerella. The seed of these species, especially that ofLesquerella, germinated in 50 mM of NaCl, but not at 100 mM
with the corresponding EC of 5.5 and 10.5 dS m-1, respectively.
When the seed was placed at 0.25 or 0.5 cm, soil salinity might
have been at least 2 to 13 dS m-1 in ECe when top-irrigatedwith the saline solution of 4.1 dS m-1, and 5 to 24 dS m-1 undersubirrigation (Fig. 1). Salinity of soil solutions to which theseed is exposed is 2 to 3 times higher than ECe (Rhoades andMiyamoto, 1990). Poor seedling emergence observed with

pennycress and Lesquerella (Table 5) can be attributed togermination failure.
If the germination failure was the only reason for poor
emergence, Camelina should have provided better seedlingemergence than the observed (Table 4), because it issalt-tolerant at germination (Table 2). Likewise, canola
should have provided seedling emergence comparable tosafflower, as their salt tolerance at germination is comparable(Table 2). The salts accumulated near the soil surface might
have reduced hypocotyl growth of canola as well as ofCamelina and Lesquerella (Table 3). This may have lead to
reduced emergence.
Seedling emergence of the tested species, except for
pennycress and Salicornia did not improve throughtop-irrigation, in spite of the fact that top-irrigation provided
lower salinity (Fig. 1). Since seedling emergence under
top-irrigated conditions occurred nearly exclusively throughsoil cracks, it would be realistic to assume that soil crusting
was a factor under top-irrigated conditions. In fact, soil
strength increased markedly with top-irrigation, especially inHarkey silt loam (Table 6). Glendale silty clay containedsome organic matter, and registered lower soil strength and in
most cases, better emergence than in the silt loam (Tables 5 and
6). The reduction in emergence from top-irrigated conditions
was pronounced when saline water was used as opposed tosaline soils (Tables 4 and 5). We suspect that salts in theirrigation water had accumulated at the wall of soil cracks, thus
limiting emergence through the cracks.
This study indicates that seedling emergence is influenced
by many factors, such as seed size which may affect hypocotyl
and seedling vigor; irrigation methods which affect vertical
distribution of salts as well as soil strength, besides salt
tolerance. In oilseed crops having small seed and low salt
tolerance at germination, such as pennycress and Lesquerella,crop breeding efforts towards improving these traits seem to be
prerequisite. In the case of Salicornia, a detailed study isneeded to clarify the cause(s) of establishment difficulties.
Safflower and canola can be established with water of elevatedsalinity (4 dS m-1or somewhat higher), using furrow or
corrugation irrigation as long as soil crusting is minimized andthe field is leached prior to seeding or with the use of reshaped
bed after pre-irrigation. The use of disk-type seeders whichplant seeds in V-shaped shallow indentation on the reshaped
bed tops may help improve seedling establishment.
Camelina, which has spindly and thin hypocotyl, has to beseeded shallow, and may be planted using corrugated beds with
the V-shaped notches. Under the existing field management
capability, safflower and canola appear to be among thepromising oilseed crops in saline areas.

Acknowledgement
This project was supported in part by the Texas State
Bioenergy Research Grant, and the Rio Grande Basin Initiative,
USDA Grant No. 2005-34461-15661
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