Original
Research Research on Crop Ecophysiology Vol. 9/1, Issue 1 (2014), Pages: 1 -11
Saltgrass, a True
Halophytic Plant Species for Sustainable Agriculture in Desert Regions Mohammad
Pessarakli Professor. School of Plant
Sciences, the University of Arizona Tucson, AZ 85721, USA * Corresponding
author E-mail:pessarak@ag.arizona.edu Received: 4 April
2013 Accepted: 12
November 2013 Abstract Continuous desertification of arable lands due to
urbanization, global warming, and shortage of water mandates use of low
quality/saline water for irrigation, especially in the regions experiencing
water shortage. Using low quality/saline water for irrigation imposes more
stress on plants which are already under stress in these regions characterized
with saline soils and shortage of water. Thus, there is an urgent need for
finding salt/drought tolerant plant
species to survive/sustain under such stressful conditions. Since the native
plants are already growing under such conditions and are adapted to these
stresses, they are the best and the most suitable candidates to be manipulated
for use under these stressful conditions. If stress tolerant species/genotypes
of these native plants are successfully identified, there would be a
substantial savings in cultural practices and inputs in using them by the growers
and will result in substantial savings in the currencies of the countries. My
investigations at the University of Arizona on saltgrass (Distichlis spicata L.), a euhalophytic plant species, have
indicated that this plant has an excellent drought and salinity tolerance with
a great potential to be used under harsh and stressful environmental
conditions. This grass has multi usages, including animal feed, soil
conservation, saline soils reclamation, and combating desertification processes. The objectives of this study were to find the most
salt tolerant of various saltgrass genotypes for use in arid and semi-arid
regions for sustainable agriculture and biologically reclaiming saline soils.
Twelve saltgrass clones were
studied in a greenhouse, using the hydroponics technique to evaluate their
growth responses in terms of shoot and root lengths and DM weights, and general
grass quality under salt stress conditions. Grasses were grown vegetatively in Hoagland solution for 90 days prior to
exposure to salt stress. Then, 4 treatments [EC of 6 (control), 20, 34,
and 48 dSm-1 salinity stress] were replicated 3 times in a RCB design experiment. Grasses were grown
under these conditions for 10 weeks. During this period, shoots were clipped
bi-weekly, clippings were oven dried at 65o C and DM weights were
recorded, and shoot and root lengths were also measured. At the last
harvest, roots were also harvested, oven dried, and DM weights were determined.
General grass quality was weekly evaluated and recorded. Although, all the
grasses showed a high level of salinity tolerance, there was a linear reduction in their growth responses
as salinity level increased. However, there was a wide range of variations
observed in salt tolerance of these saltgrass clones. The superior stress tolerant genotypes were
identified which could be recommended for sustainable production under arid
regions and combating desertification. This grass proved to not only have a
satisfactory growth under the harsh desert conditions, but also to substantially
reduce salinity level of the rhizosphere, which indicates that saltgrass can
effectively be used for biological salinity control or reclamation of desert
saline soils and combating desertification processes. Keywords: Salt stress, Arid regions, Saltgrass, Sustainable agriculture, Saline
soil reclamation, Combating desertification processes Introduction Saltgrass (Distichlis
spicata (L.) Greene var. stricta (Gray) Beetle) (Gould, 1993), indigenous to the Southwest, a potential animal
feed plant, saline soil reclamation, soil establishment/erosion control, and
use as a turfgrass species for
lawns/recreation areas, grows in very
poor to fair condition soils, in both salt-affected soils and soils under poor
fertility as well as drought and harsh
environmental conditions (Gould, 1993 O’Leary and Glenn, 1994). Its dominant and most common habitats are
arid and semi-arid regions (Marcum et al., 2005 Pessarakli and Kopec,
2010 Pessarakli and Kopec, 2011 Pessarakli et al., 2011a, 2011b Pessarakli
et al., 2012). The plant is
abundantly found in areas of the western parts of the United States as well as
on the sea-shores of several Middle-Eastern countries, Africa, South and
Central American countries (Pessarakli et al., 2005 Pessarakli, 2007
Pessarakli and Kopec, 2010 Pessarakli et al., 2011a, 2011b Pessarakli et
al., 2012). The species can be
manipulated to modify its performance and increase its yield and productivity. This plant has multi-purpose usages. It can be
substituted for animal feeds like alfalfa, used for biological reclamation of
saline soils, soil conservation and erosion control for covering road sides and
soil surfaces in lands with high risks of erosion, and use as a turfgrass
species. Recently, the United States Golf Association (USGA)
and the US Bureau of Land Management (BLM) have shown a great deal of interest
in financing research work on this plant to use it as a turfgrass or for soil
erosion control and saline soil reclamation. Most of these research works have
been conducted at the University of Arizona and Colorado State University.
Consequently, the USGA and the BLM funds for the investigations on this grass
species have been allocated to these institutions. Positive and promising
results have already been obtained from these studies (Gessler and Pessarakli,
2009 Kopec et al., 2000, 2001a, 2001b, 2006 Marcum et al.,
2001, 2005 Pessarakli, 2005a, 2005b, 2007, 2008 Pessarakli and Kopec, 2005,
2006, 2008a, 2008b, 2010 Pessarakli and Marcum, 2000 Pessarakli et al.,
2001a, 2001b, 2001c, 2003, 2005, 2008 2011a, 2011b, 2012). Most of the published reports on saltgrass, including
those of Sigua and Hudnall (1991), Sowa and Towill (1991), Enberg and Wu
(1995), Miyamoto et al. (1996), Rossi et al. (1996), and Miller et
al. (1998) are concern only with the growth of this species, usually
concentrated only on one grass genotype or the species of a specific location. The objectives
of this study were to find the most salinity tolerant of various saltgrass
genotypes and to recommend them as the potential species for use under arid,
semi-arid, and areas with saline soils and limited water supplies for
sustainable agriculture and combating desertification. Materials and
Methods Plant Materials Twelve
inland saltgrass (Distichlis spicata L.) clones (A37, A49, A50, A60, 72,
A86, A107, A126, A136, A138, 239, and 240), collected from different locations
in several western states of the United States (Arizona, California, Nevada,
and Colorado) were used in a greenhouse experiment to evaluate their growth
responses in terms of shoot and root lengths as well as shoot and root dry
weights, and visual grass quality under different levels of salinity stress
conditions, using a hydroponics technique. Plant Establishment The
plants were grown as vegetative propagules in cups, 9 cm diameter and cut to 7
cm height. Silica sand was used as the plant anchor medium. The cups were
fitted in plywood lid holes and the lids were placed on 42 cm X 34 cm X 12 cm
Carb-X polyethelene tubs containing half strength Hoagland nutrient solution
(Hoagland and Arnon, 1950). Three replications of each treatment were used in a
randomized complete block (RCB) design in this investigation. The plants were
allowed to grow in this nutrient solution for 8 weeks. During this period, the
plant shoots were harvested weekly in order to reach full maturity and develop
uniform and equal size plants. The harvested plant materials were discarded.
The culture solutions were changed biweekly to ensure adequate amount of plant
essential nutrient elements for normal growth and development. At the last
harvest, 10th week, the roots were also cut to 2.5 cm length to have
plants with uniform roots and shoots for the stress phase of the experiment. Salt Treatments The
salt treatments were initiated by gradually raising the EC (electrical
conductivity) of the culture medium to 6, 20, 34, and 48 dS m-1 by
adding Instant Ocean salt to the nutrient solutions, followed procedures used
by Pessarakli and Kopec (2005, 2006). The EC of the culture solutions were
raised by increments of 6 (first day) and 7 every other day until the desired
EC levels were reached. Four treatments were used, including control (EC = 6 dS
m-1, several of my salinity stress experiments showed that saltgrass
at relatively low level of salinity for this high salinity tolerant halophytic
grass performs better than growing in normal condition, therefore, for the
control, usually, I use EC = 6 dS m-1), 20, 34, and 48 dS dS m-1
(EC = 48 dS dS m-1 is a good representative of the EC of sea water
which is normally between 30 and 60 dS dS m-1). The culture solution
levels in the tubs were marked at the 10 liter
volume, and the solution conductivities were monitored/adjusted to maintain the
prescribed treatment salinity levels. After the final salinity levels were
reached, the shoots and the roots were harvested and the harvested plant
materials were discarded prior to the beginning of the data collection of the
salinity stress phase of the experiment. Then, plant shoots were harvested bi-weekly for 10
weeks for the evaluation of the dry matter (DM) production. At each harvest,
both shoot and root lengths were measured and recorded. The harvested plant
materials were oven dried at 65o C and DM weights were measured and
recorded. The recorded data were considered the bi-weekly plant DM production.
At the termination of the experiment, the last harvest, plant roots were also
harvested, oven dried at 65o C, and dry weights were determined and
recorded. Weekly visual evaluation of the grass quality was also performed and
recorded. The data were subjected
to Analysis of Variance, using SAS statistical package (SAS Institute, Inc. 1991). The means were separated, using Duncan Multiple
Range test. Results and Discussion Shoot Dry Matter (DM) Weight The
shoot dry matter (DM) weights of all the saltgrass clones decreased with
increased salinity stress level. A marked reduction in shoot dry weights
occurred at the higher salinity levels (EC 34 and EC 48 dS m-1)
across all the clones (Table 1). For the dry weights of the shoots, the gap
between the means of the stressed plants and the control (EC = 6 dS m-1)
was wider as the exposure time to salinity stress progressed. Root Dry Matter (DM) Weight The
effect of salinity on root dry weight was less severe compared to that of shoot
dry mass (Table 2). Similar results were reported on different genotypes/ accessions/clones
of this grass in other studies by this author and his co-workers Table 1. Saltgrass shoot dry weight (DM) under salt
stress Grass ID Shoot 6 DM (g)* 20 at EC 34 dS m-1 48 A37 1.10cde** 0.57bcde 0.27cde 0.15c A49 1.26bcd 0.77ab 0.32bcde 0.13c A50 1.65ab 0.60bcd 0.21de 0.17bc A60 1.03cde 0.38e 0.17e 0.13c 72 1.38bc 0.82a 0.38abc 0.19bc A86 1.66ab 0.86a 0.26cde 0.14c A107 0.95de 0.52cde 0.30bcde 0.20bc A126 0.83e 0.41de 0.18e 0.15c A128 1.37bc 0.73abc 0.52a 0.30a A138 1.09cde 0.46de 0.36abcd 0.25ab 239 1.67ab 0.88a 0.44ab 0.15c 240 1.94a 0.91a 0.49a 0.24ab *The values are the means of 3
replications of each treatment. **The values
followed by the same letters in each column are not statistically significant
at the 0.05 probability level. Table 2. Saltgrass root dry weight (DM) (cum. values)
under salt stress Grass ID Root 6 DM (g)* 20 at EC 34 dS m-1 48 A37 0.74cde** 0.99def 1.10cdef 0.78cd A49 1.61b 1.11cdef 1.56bcd 1.03bcd A50 1.83b 1.65a 1.94abc 0.74cd A60 1.46bc 1.71a 1.31bcde 0.84bcd 72 0.77cde 0.93def 0.72def 0.50d A86 1.06bcde 1.18bcde 0.76def 0.81bcd A107 0.68de 0.84ef 0.53ef 0.68cd A126 0.50e 0.68f 0.26f 0.48d A128 3.46a 1.50abc 2.05ab 1.18bc A138 1.17bcde 0.88def 0.43ef 2.28a 239 1.31bcd 1.30abcd 2.82a 1.21bc 240 3.36a 1.63ab 1.25bcde 1.42b *The values are the means of 3 replications of each
treatment. **The values
followed by the same letters in each column are not statistically significant
at the 0.05 probability level. (Marcum
et al., 2005 Pessarakli, 2007, 2008 Pessarakli and Kopec, 2005, 2006,
2010 Pessarakli and Marcum, 2000 Pessarakli et al., 2001c, 2005, 2008,
2011a, 2012). Sagi et al. (1997) and Pessarakli and Tucker (1985, 1988)
also found the adverse effect of salinity stress was more pronounced on plant
shoots than the roots. This is a common phenomenon in halophytic plant species
that usually under salinity stress conditions, their shoots are more severely
affected than their roots. Clone 240 had excellent root growth at EC 6 dS m-1
and the second highest root production at EC 48 dS m-1 (Table 2),
but had poor quality under high salinity level. The same was true for clone
239. Clone A138 had twice the root mass of most other clones at EC 48 dS m-1,
but essentially had no green foliage at EC 48 dS m-1 at the close of
the test. At EC 6 dS m-1, clone A128 produced twice
the test mean average for roots (3.46 g) with fairly good absolute root
production afterwards, but showing a significant change in root production as
EC levels increased (Table 2). Although the root dry weight was enhanced at the lower
level of salinity for most of the clones, there was not statistically
significant difference detected between the means of the different treatments
(Table 2). Grass
Visual Quality Any level of salinity stress had a significant adverse
effect on the grass visual quality (Table 3). Quality scores for various clones
ranged from 9.7 to 2.6 at different salinity stress levels. At EC 20 dS m-1,
quality scores ranged from 5.1 to 9.7 (Table 3) throughout the entire test. As
shown in Table 3, all clonal entries had good quality and full maintenance of
green tissue retention at EC 6 dS m-1 at the end of the trial. After 10 weeks growth at EC 34 dS m-1 (salinity
level equal to that of sea level salinity), entries 239 and 240 were the only
clones to have quality ratings of 6 (acceptable quality, on the scale of 1 -
10) or greater (Table 3). These two clones represented the best quality clones
at EC 34 dS m-1 at the end of the test. At EC 48 dS m-1, no entries produced an
acceptable plant quality (scores of 6 or higher). Most clones decreased in
(final) quality as EC increased from EC 6 to EC 48 dS m-1, but the
entries 239 and 240 showed a more of typical halophytic response, having an
increase in quality at EC 20 dS m-1 over that observed at EC 6 dS m-1
(Table 3). Table 3. Saltgrass visual quality under salinity stress Grass ID General 6 quality*
20 at 34 EC 48 A37 8.0cde** 5.1f 3.3g 2.6e A49 7.7def 6.4d 4.3ef 2.8e A50 8.6abc 7.2bc 5.0cd 4.0bc A60 8.2bcd 5.5ef 3.9fg 3.5cd 72 9.0a 7.4bc 5.9b 4.8a A86 8.5abc 6.7cd 5.7b 3.9bc A107 7.5def 5.9def 5.4bc 4.4ab A126 6.7g 5.3f 4.6de 3.9bc A128 7.1fg 6.2de 5.0cd 3.0de A138 8.6abc 7.9b 5.4bc 4.2ab 239 8.9ab 9.3a 6.6a 4.2ab 240 9.2a 9.7a 7.1a 2.8e *The quality values are the means of 3 replications
of each treatment and 10 weekly evaluations. **The values followed by the same letters in each column
are not statistically significant at the 0.05 probability level. Salt
Tolerance Ranking of the Various Clones of Saltgrass Salinity tolerance ranking of the various saltgrass
clones used in this study based on shoot DM weight, root DM weight, grass
visual quality, or overall ranking considering all the study parameters
together, are presented in Table 4. Although there are some
minor differences in salt tolerance ranking of the clones when compared based
on shoot DM weight, root DM weight, or grass visual quality, the overall
ranking is the best representation of the salinity tolerance of the various
tested clones. Considering all the study
parameters together, there was a wide range of salinity tolerance found among
the 12 saltgrass clones. The 240 and 239 clones were the most salt tolerant
clones (especially, up to EC of 34 dS m-1) followed by A128, 72, A138. These were
closely followed by A50, A86, and A49 in salinity tolerance. A49 clone laid
between this and the last group in regards to salinity tolerance. A60, A107,
A37, and A126 were among the lowest salinity tolerant grasses which the A126
was the least tolerant clone. Table 4. Salt tolerance ranking of Saltgrass based on shoot
weight, root weight, or grass visual quality Tolerance Salt Shoot wt. tolerance Root wt. based Quality on Overall High 240a* A128a 240a 240a
A128ab 240ab 239a 239a 239ab 239ab 72ab A128ab 72ab A50ab A138ab 72ab A86ab A60abc A50abc A138ab A138abc A138abc A86bc A50b A50bc A49bc A60bcd A86b A49bc A86bc A49cde A49bc A107cd A37cd A128de A60cd A37cd 72cd A107de A107cd A126d A107cd A37de A37cd Low A60d A126d A126e A126d *The clones followed by the same letters in each column are
not statistically significant at the 0.05 probability level. Overall, the results of the shoot and the root dry
mass and the visual grass quality showed that the maintenance of green foliage
and tolerance under saline hydroponic conditions are under physiological
conditions/adjustments that are not totally related to dry matter (DM)
production in shoots and roots. This was corroborated by the results that
clones which maintained the highest quality under EC 34 dS m-1
exhibited either a large increase in root mass (i.e., clone 239), or only a
small increase of the root mass (i.e., clone 240) produced at EC 6 dS m-1.
Likewise, clone A138 produced a large increase of its EC 6 dS m-1
root mass at the highest EC level of 48 dS m-1. However, it could
not maintain green foliage at 10 weeks of exposure to this high EC. The same
was true for shoot DM production that occurred in a more narrow range of values
than did root DM production. In short, saltgrass shoot DM
weight decreased linearly with increased salinity levels for all clones. For
most clones, there was no difference among the root DM of the grass at
different salinity levels. Visual quality of the grass followed the same
pattern as the shoot DM weight. It decreased linearly with increased salinity
levels for all clones. Clones differed
greatly in their maintenance of green color retention (quality) as EC levels
(salinity) increased. Two clones which produced acceptable quality at the EC
level of 34 dS m-1 were clones 239 and 240. No clones could maintain
adequate quality color at EC level of 48 dS m-1 after 10 weeks of
exposure at this EC level. The difference in salinity tolerance level among the
clones was significant. The grasses were separated in several groups with
different degrees of salt tolerance. Considering
all the study parameters together, there was a wide range of salinity tolerance
found among the 12 saltgrass clones. The 240 and 239 clones were the most salt
tolerant clones (especially, up to EC of 34 dS m-1) followed by A128, 72, and A138. These were
closely followed by A50, A86, and A49 in salinity tolerance. A49 clone laid
between this and the last group in regards to salinity tolerance. A60, A107,
A37, and A126 were among the lowest salinity tolerant grasses which the A126
was the least tolerant clone. Conclusions In terms of salinity tolerance (quality), green
foliage retention was empirically the best assessment of the clonal response to
increased salinity. For large scale screening of saltgrass germplasm, the
maintenance of green tissue at a specific EC level would seem to be adequate as
a simple selection method for salinity tolerance. Shoot and root lengths and dry weights decreased with
increased salinity stress. However, shoots were more severely affected by
salinity stress than the roots. Grass visual quality was significantly affected
(lower quality) as the salinity levels of the culture solutions increased. Overall, the results of this investigation indicate
that saltgrass is a very high salinity tolerant species, and the results further
suggest that this grass growing even under poor soil conditions (salt-affected
desert soils) can be a suitable and beneficial plant species for growth and
production in arid regions, and still show a favorable growth. Acknowledgments This
study was financially supported by a grant from the United States Golf
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