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Practical uses of halophytic plants as sources of food and fodder.

© CAB International 2019. Halophytes and Climate Change: Adaptive Mechanisms and
324 Potential Uses (eds M. Hasanuzzaman, S. Shabala and M. Fujita)
20.1 Introduction
20.1.1 Halophytes and the environment
Food production needs to increase by up to 70%
by 2050 to match the food needs of a growing
population that is expected to reach 9 million by
2050 (Panta et al., 2014). This increase is a
challenge because of two main problems we are
facing today: land and water scarcity. Worldwide
urbanization and land degradation are the main
causes of land scarcity for agricultural produc-
tion. Soil salinization is a major factor contribut-
ing to soil degradation. High levels of salts in soil
cause salinity, and in a global context NaCl is the
major cause (Munns and Tester, 2008; Rozema
and Flowers, 2008). Only about 6% of the world’s
land surface is composed of naturally saline
soils that have never been cultivated because of
their high salinity. Irrigated agriculture, which
produces 70% of crops, is the cause of soil
degradation, called secondary salinization. It is
estimated that about 20% of ir rigated land (230
million ha) is seriously affected by salts (Boscaiu
and Vicente, 2013). Irrigation water carries
salts (dissolved ions) that accumulate in the soil
with cycles of ir rigation and cause the loss of
more than 10 million ha of arable land every year
20 Practical Uses of Halophytic Plants
as Sources of Food and Fodder
Tiziana Centofanti1,* and Gary Bañuelos2
1Central European University, Budapest, Hungary; 2San Joaquin Valley
Agricultural Sciences Center, Parlier, USA
* Corresponding author e-mail:
Halophytes are plants that are adapted to saline soils in their natural habitats because they are salt tolerant.
They are found in a range of environments with varied salinity and climatic conditions. These plant species can
be irrigated with saline water and cultivated on saline soils that are unsuitable for commercial crops. Halophytes
are rich in nutrients, such as antioxidants, fatty acids and amino acids, and many species have been used trad-
itionally as herbs and vegetables, feed and fodder. Therefore, halophytes are considered one of the alternative
solutions to problems related to food security, fresh water scarcity, salinization and diversication of diets for
healthier nutrition. However, despite the promising future for multiple uses of halophytes, many problems re-
lated to halophyte cultivation for human and animal consumption and their commercialization have still not
been tackled. This chapter is intended to provide an overview of the development of halophytes as vegetable,
feed and fodder, and to highlight the importance of creating a demand in the marketplace for halophyte con-
Keywords: Alternative foods; Climate change; Marine ecosystem; Medicinal plants; Salinity
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Practical uses of halophytic plants as sources of food and fodder 325
(Owens, 2001). The problem of soil salinization
has occurred since ancient times. Yield losses
and crop failures were documented in old writ-
ten records of early civilizations such as those in
the Indus and Nile valleys. It is thought that the
collapse of the Mesopotamian civilization was
partly due to the similar phenomenon of soil
salinization due to irrigation (Aronson, 1985;
Boscaiu and Vicente, 2013). Civilizations have
historically been shaped by soil degradation (in-
cluding salinity) that restricted population num-
bers, limited where domestication could occur
and forced relocation (Hasegawa, 2013). The
Aral Sea Basin in central Asia represents one of
the worst cases of salinization, where salinity
and waterlogging affect up to 50% of the irri-
gated area (Kijne, 2005; Qadir etal., 2009). In
addition to soil salinity, freshwater aquifers are
becoming increasingly salty because of sea-level
rise and the increase in droughts. In human his-
tory, agriculture has expanded into new land at
the loss of natural ecosystems. Today, we are
aware of the consequences of this process (forest
conversion), which lead to signicant losses of
ecosystem services, such as habitat necessary to
maintain biodiversity, storage of carbon, ood
mitigation, and soil and watershed protection.
Hence, use of degraded lands has often been
suggested as an alternative to the issue of land
scarcity. Denitive data on the annual world-
wide loss of farmland due to soil degradation
and related causes are lacking. Gibbs and Sal-
mon (2015) used four approaches (expert opin-
ion, satellite observations, biophysical models
and inventories of abandoned agricultural land)
to assess degraded land at the global scale. They
concluded that there is a wide disagreement in
the spatial distribution of these degraded lands,
but estimated that the world’s total degraded
area ranges from 1 billion ha to 6 billion ha. The
disagreement stems from the uncertainty in cat-
egorizing an area as degraded, as this classica-
tion depends on a number of characteristics that
are subjective and site- and time-specic.
To meet the future food demand, we need
not only to develop alternative and more ef-
cient technologies to increase yield, but also to
enlarge the diversity of crops that are grown and
consumed. Today, among the thousands of crops
deemed edible, humans consume only about 20
crops on a large scale, of which the major four
(corn, rice, wheat and soybean) represent staple
foods worldwide. A diversication of our diets
with inclusion of traditional or neglected crops
that are rich in micronutrients, require less in-
puts (fertilizer s, water, pesticides) for growth and
can adapt to degraded environments (i.e. saline
soils) could be a win–win situation to tackle the
problem of food security and resources scarcity.
In line with other authors (Panta et al., 2014;
Ventura etal., 2014), we think that halophytes
(plants adapted to saline soil, see section 20.1.2)
may be a commercial, alternative plant group
for the use of saline land and to ease pressure on
the requirement for good-quality land and water
(Panta et al., 2014). Others have shown that
halophytes can be grown to produce oilseeds,
grains, forage, fuel, food, medicine, chemicals,
timber and bre (Kahn and Qaiser, 2006; Glenn
etal., 2013) or be used for soil or water conser-
vation or remediation (Ravindran et al., 2007;
Zhao etal., 2002). In addition, halophytes have
been tested for the treatment of saline aquacul-
ture efuent (Brown et al., 1999) and used as
landscaping ornamentals (Zia etal., 2008).
However, despite the promising future for
multiple uses of halophytes, many problems re-
lated to halophyte cultivation for human and
animal consumption and their commercializa-
tion have still not been tackled. This chapter
considers the development of halophytes as
vegetable, feed and fodder and highlights the
importance of creating market demand for halo-
phyte consumption. We also analyse the envir-
onmental impacts and benets of halophytes in
bio-saline agriculture.
20.1.2 Characteristics
of halophytic plants
Halophytes are plants that are adapted to saline
soils in their natural habitats because they are
salt tolerant. Halophytes are found in a range of
environments with varied salinity and climatic
conditions. They are trees, grasses and saltbushes,
growing in salt marshes, saline deserts and
coastal areas around the world (Epstein et al.,
1980). They represent only a small fraction of
the total number of owering plants (2600 out
of 400,000) and only a small percentage of
halophytes is domesticated and used as food and
fodder. There is no broadly accepted denition of
0004229901.INDD 325 11/2/2018 4:55:22 PM
326 T. Centofanti and G. Bañuelos
halophyte but the most commonly used is: ‘a
halophyte is any plant that can complete its life
cycle and reproduce itself under conditions of
soil-water salinity of 8–10 dS m−1 electrical con-
ductivity (EC) (approximately 20% seawater) or
more’ (Aronson, 1985, 1989).
For a long time researchers have tried to
identify the mechanisms of salt tolerance, but
these have not yet been fully claried. Indeed,
the effect of salinity on growth varies among
halophytes and many differences do exist among
species in the balance of Na+ and Cl in shoot tis-
sues (Flowers and Colmer, 2008).
For example, it has not yet been claried
whether mechanisms for salt tolerance are simi-
lar between all halophytes and whether mech-
anisms for salt tolerance in halophytes are linked
with other environmental variables such as arid-
ity, ooding, etc. (Flowers and Colmer, 2008).
There is clearly still much to learn about halo-
phytes and the diversity of mechanisms that
they employ to cope with salinity.
Salt tolerance comprises morphological,
physiological and biochemical adaptations at
the whole plant, tissue and cellular levels (Koyro
etal., 2011). These adaptations consist of com-
plex polygenic traits controlled by a number of
genes or groups of genes (Türkan and Demiral,
2009). Transferring salinity tolerance traits to
glycophytes (less salt-tolerant crops) would be
useful to obtain conventional crops that are
adapted to saline environments, but is currently
impractical due to complex polygenic traits.
According to Rozema and Schats (2013), glyco-
phytes would not accommodate the appropriate
allelic variants at all the loci involved in halo-
phyte salt tolerance.
Generally, in both glycophytes and halo-
phytes, salt-tolerance mechanisms involve con-
trolled uptake of salt ions in the root system,
compartmentalization of toxic ions in the vacu-
ole and synthesis of organic solutes in the
cytoplasm to metabolically balance the osmotic
potential of the Na+ and Cl accumulated in the
vacuole (Zhu, 2001; Flowers and Colmer, 2008;
Shabala and Mackay, 2011). Sugars (e.g. sucros e),
sugar alcohols (e.g. sorbitol), amino acids (e.g.
proline), methylated proline-related compounds
(e.g. methyl-proline), betaines (e.g. glycinebe-
taine) and methylated sulfonio compounds (e.g.
dimethylsulfonioproprionate, DMSP) have been
proposed to fulll this function in halophytes
(Flowers and Colmer, 2008). In addition, to
counteract the oxidative damage caused by re-
active oxygen species (ROS), halophytes produce
antioxidative enzymes and non-enzymatic
molecules with antioxidant properties – such as
ascorbate, glutathione, alpha tocopherol, avon-
oids, anthocyanins, carotenoids and polyphenolic
compounds (Türkan and Demiral, 2009; Zhu,
2001). These antioxidants and organic solutes
confer nutritional value to the halophytes and
are ultimately responsible for the quality of the
vegetable product (Maggio etal., 2011). The en-
richment of the edible plant part with antioxi-
dant compounds, such as ascorbic acid (vitamin
C), β-carotene and polyphenols, is highly desir-
able for the human diet. Halophytes have gained
considerable attention and have been the subject
of many research studies in the past 30 years be-
cause they are considered one of the alternative
solutions to problems related to food security,
freshwater scarcity, salinization and diversica-
tion of diets for healthier nutrition. Their poten-
tial as alternative crops is multi-faceted: (1) they
can be irrigated with seawater; (2) they can
grow in arid saline environments, e.g. deserts,
marginal and degraded soils; and (3) they can
contain nutraceutical products that may be
benecial to human health. In section 20.2 we
discuss the role of halophytes as alternative
crops for sustainable use of natural resources.
20.2 Halophytes as Food
20.2.1 Traditional wild halophytes
Many halophytic plant species have been used
traditionally as herbs and vegetables, and a re-
view of the history of their uses can be found in
ethnobotanic reviews and literature (Davy etal.,
2001; Simopoulos, 2004; Guarrera etal., 2006;
Tardío etal., 2006).
Sugar beet (Beta vulgaris) is the only major
food crop derived from the halophyte wild pro-
genitor B. vulgaris subsp. maritima (Aronson,
1985). Similarly, the date palm (Phoenix dactylif-
era) cultivated today seems to have derived from
the wild halophytic population that grew in the
Middle East. Other traditional food staples derived
from halophytes that grow in ocean water are
eelgrass (Zostera maritima) and Palmer’s saltgrass
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Practical uses of halophytic plants as sources of food and fodder 327
(Distichlis palmeri) from the Gulf of California.
Eelgrass plant seeds have been used by the Seri
Indians as gruel after being toasted and ground.
The gruel was then mixed with fats or other fatty
seeds oils (i.e. columnar cactus) to prepare a
meal very rich in protein, fats and carbohydrates
(Felger et al., 1980; Aronson, 1985). Distichlis
palmeri (Glenn etal., 2013) is a highly salt-tolerant
grass endemic to the delta of the Colorado River
and in the northern Gulf of California, USA.
A perennial species, its grains are similar to rice
and wheat in size and nutritional value. The
plant can grow in ooded conditions – like paddy
rice – in seawater, and the grains were used by
the Cocopa Indians to make bread (Felger, 1979).
Another halophyte species that has been used
since ancient times is the mangrove. Mangroves
represent an abundant source of timber, re-
wood and charcoal, as well as tannins. The bark
of mangrove trees from genera like Excaecaria
agallocha, Kandelia, Rhizophora, Ceriops, Sonnertia
acida and Carapa are cut and boiled in large ves-
sels of water to extract the tannins that are then
used for shing and in the plywood industry (Ar-
onson, 1985). Many species of mangroves have
palatable foliage that can be used as animal feed.
In particular, the foliage of Avicennia marina,
Sonneratia alba and Rhizophora mangle has served
as camel and cattle feed (Chapman, 1976). In
Colombia, mangrove species including Avicenna
marina and A. germinans are used as food (Lieth
etal., 2000). In different regions of India, young
shoots and leaves of Chenopodium album and Am-
aranthus spp. are used as salads and vegetables,
and raw fruits of Capparis decidua are used for
pickles (Dagar, 2005; Rameshkumar and Es-
waran, 2013). People in the Maldives regularly
eat Sonneratia caseolaris, also known as mangrove
apple or crabapple mangrove fruit, which are eaten
raw and also used to make vinegar.
Sea spinach (Tetragonia sp.) was introduced
to the culinary world in the 18th century. Its ed-
ible leaves are the halophyte version of spinach
and they have been used pickled and cooked to
ght scurvy. Sea kale (Crambe maritima) is an-
other traditional halophyte vegetable precursor
of a widely consumed commercial cabbage. Its
shoots are served steamed or sauced like aspara-
gus (Panta etal., 2014).
Many of the plants described above have
been collected from wild populations or culti-
vated in private backyard and kitchen gardens
and sometimes sold at local farmers’ markets
(Wilson et al., 2000). These halophytic species
have great potential to be transformed into vege-
table crops for saline agriculture. Growing inter-
est over the past decade in cultivating crops
under saline conditions has led to rediscovery of
the potential of several promising halophytic
plant species to be farmed as gourmet vegetables.
Most are eaten as raw vegetables or in fresh
salads, and some are also cooked or pickled.
Apart from their palatability, these plants are –
in general – rich in protein, antioxidant com-
pounds and/or essential nutrients (minerals,
vitamins, amino acids and/or fatty acids). How-
ever, halophytes are wild species and they can
hardly compete with standard crops, which
were domesticated thousands of years ago and
improved over the centuries (Boscaiu and Vi-
cente, 2013). Traits that are adaptive for wild
plants, such as uneven germination, lack of seed
retention (seed shattering) and toxic substances
in tissues (saponins, tannins) are undesirable for
use of halophytes as cultivated plants (Brown
etal., 2014). For example, the agronomic char-
acteristics of Distichlis palmeri and its crop poten-
tial as grain still need to be identied to develop
efcient breeding programmes (Boscaiu and Vi-
cente, 2013). The history of traditional uses of
halophytes in communities is important in over-
coming the problem of domestication. Brown
etal. (2014) discuss the problems related to do-
mestication of halophytic plants for commercial
production. One important aspect to consider is
that the success of halophyte growth in saline
agriculture is related to the environmental con-
ditions. These species can be grown better in en-
vironments similar to those in which the plant
occurs in nature.
To date, most halophytic plants have been
tested for their yield and adaptability to saline
agriculture. A summary of saline agriculture
using domesticated halophytes is provided in
section 20.2.2.
20.2.2 Saline agriculture
Saline agriculture is the cultivation of crops us-
ing seawater (approx. 40 dS m−1) or brackish water.
Seawater agriculture, as proposed by Glenn etal.
(1998), is an extreme case of halophyte farming
0004229901.INDD 327 11/2/2018 4:55:22 PM
328 T. Centofanti and G. Bañuelos
and allows the cultivation of only the most
salt-tolerant halophytes. Instead, the use of
brackish water represents a more viable option
for commercial saline agriculture, because a
higher number of halophyte crops can be grown
in much lower salinity levels than seawater
(Rozema and Flowers, 2008; Ventura and Sagi,
2013). Saline agriculture can be carried out on
marginal land, deserts and degraded soils close
to the sea. About 43% of the Earth’s total land
surface is arid or semi-arid, and seawater in the
oceans makes up 97% of water on earth. Hence,
saline agriculture could be an alternative, sus-
tainable way of increasing food production
without competing with conventional crops for
increasingly scarce resources such as fertile land
and good-quality irrigation water (Khan and
Duke, 2001; Boscaiu and Vicente, 2013).
Despite these positive environmental factors of
saline agriculture, successful agronomic tech-
niques must be developed for growing saline
water-irrigated crops in a sustainable manner.
These methods must also not contribute to fur-
ther damage of natural environments (Khan
and Duke, 2001). One primary consideration for
sustainable saline agriculture is to choose spe-
cies from climate regimes similar to those where
the potential crop is being planted, as the candi-
date species will be better adapted to local envir-
onmental conditions (Zerai etal., 2010; Brown
etal., 2014).
Optimization of irrigation quantity at each
salinity level (brackish or seawater) is another
important aspect for sustainable saline agricul-
ture, to allow salt leaching below the root zone,
and consequently avoiding soil salinization
(O’Leary, 1988; Lieth, 2000). Multi-year long-
term experiments with halophytes have been
conducted in various parts of the world; and no
decrease in yield, or negative impact on soil
structure has been observed because of saline
water irrigation. Sandy soils in coastal areas or
inland sand dunes may be readily available for
large-scale halophyte production without the
risk of salt contamination occurring on fertile
soils through leaching Ca2+/Na+ exchange and
subsequent clay dispersion. Likewise, under-
ground freshwater contamination should be
avoided by the existence of sufciently deep
water tables or adequate drainage. In addition,
saline agriculture must justify the expense of
pumping seawater for irrigation by producing
useful crops at high yields (Glenn etal., 1998).
However, seawater agriculture should be cost-
effective in desert regions – even though the yields
may be smaller than in traditional agriculture –
because it is cheaper to pump seawater at sea
level than to pump freshwater from wells (Glenn
etal., 1998). Typical agricultural wells lift water
from 20 m to as deep as 100 m, whereas the lift
of typical coastal seawater wells is only 3–10 m.
Furthermore, in some locations tides can be
used to irrigate crops without the need for pump-
ing (Glenn etal., 1994; Brown etal., 2014). In
addition, coastal desert farms on sandy soils
generally have unimpeded drainage back to the
sea, thus avoiding groundwater salt contamin-
ation. Furthermore, coastal and inland salt des-
ert aquifers often already have elevated concen-
trations of salts and so should not be damaged
by seawater irrigation (Glenn et al., 1998).
Glenn et al. (1998) have shown that normal
farm irrigation equipment can be modied to
protect it from seawater damage. Successful
long-term eld trials have been carried out with
the halophyte Salicornia bigelovii in saline agri-
culture for fodder production, as discussed in the
section 20.4 below. Some halophytes have also
been cultivated in saline agriculture as gourmet
vegetables. The long history of halophyte vege-
table cultivation and consumption has devel-
oped consumers’ appreciation for halophytes as
gourmet vegetables because of their salty taste
and high nutritional value (Mudie etal., 2005;
Lu etal., 2010). In section 20.2.3 we describe
some of the most cultivated halophyte gourmet
20.2.3 Halophytes as gourmet
vegetables Salicornia and Sarcocornia
Perhaps the most cultivated and consumed halo-
phyte vegetables are Salicornia and Sarcocornia
spp., which have been studied for their poten-
tial as gourmet food, animal feed and oils for
biodiesel. Salicornia and Sarcocornia species are
distinguished by their growth habit and ower
morphology (Kadereit etal., 2006, 2007).
In coastal communities, both Salicornia and
Sarcocornia have traditionally been used as
vegetables in fresh salads for self-consumption
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Practical uses of halophytic plants as sources of food and fodder 329
or for sale in local markets. Different species of
the Salicornia genus are suitable for vegetable
production. These halophytes are rich in min-
erals, fatty acids and antioxidant compounds,
such as polyphenols (Boscaiu and Vicente,
2013; Ventura and Sagi, 2013). In Salicornia
and Sarcocornia, total polyphenols are high, rep-
resenting 1.2 and 2.0 mg GAE g−1 fresh weight
(FW), respectively (Ventura et al., 2011). The
lower limit of other, non-halophytic leafy veget-
ables rated as rich in phenolic compounds is >
0.5 mg GAE g−1 FW (Isabelle etal., 2010). Leaves
of S. bigelovii are a source of omega-3 polyunsat-
urated fatty acids, and antioxidant β-carotene,
with quantities ranging from 4.7 (Ventura etal.,
2011) to 15.9 mg 100 g−1 FW (Lu etal., 2010).
These values are similar to those found in seaweeds
(4.0 mg/100 g FW) and spinach (5.1 mg/100 g
FW) (Isabelle et al., 2010). The highly saline-
adapted shoots of S. bigelovii contain ascorbic
acid content of 6 mg 100 g−1 FW (Lu et al.,
2010). These values are in a similar range to
those for non-halophyte leafy vegetables, such
as spinach (7 mg 100 g−1) and lettuce (< 2 mg
100 g−1) (Proteggente etal., 2002).
S. europaea has been grown in several trials
and also in small-scale commercial cultivation
using seawater in Ensenada, Mexico (Ventura
and Sagi, 2013). S. bigelovii is also grown in sub-
tropical climates, particularly in Mexico, where
growers can supply European markets with Sali-
cornia shoots in the off-season from September to
June (OASE Foundation, 2009). Indeed, most of
the halophyte vegetable production is sold as
gourmet vegetables in Europe and the USA
(Böer, 2006; Zerai et al., 2010; Ventura et al.,
2011). The young shoots are sold in the market
as ‘samphire’ or ‘sea asparagus’. They are succu-
lent and have a salty taste, which makes them
suitable as vegetables (Ventura et al., 2011).
High-yield production of Salicornia under saline
conditions is a necessary requirement for its
cultivation to be commercially viable (O’Leary,
1988). In addition to irrigation with seawater,
Salicornia has been tested in small lysimeter ex-
periments for irrigation with hypersaline drain-
age water (Grattan etal., 2008) or saline aqua-
culture efuent (Brown etal., 1999). Glenn etal.
(1998) proposed to combine shrimp farms with
halophyte production. Seawater is used to grow
the shrimps and the effluent water is then
used to grow halophytes (Buhmann etal., 2013;
Turcios and Papenbrock, 2014). This water is
also a source of nutrients. Salicornia and Sarco-
cornia spp. have proved to be efcient biolters
for the removal of nutrients from aquaculture
efuents (Shpigel et al., 2013). It is estimated
that a production level of 20–30 kg m−2 year−1
of fresh Salicornia biomass in the constructed
wetlands can remove 1–3 g m−2 day−1 of nitro-
gen (Shpigel etal., 2013).
Some of the problems encountered in halo-
phyte vegetable production are related to har-
vest and germination. Only the young shoots are
used in fresh market vegetable production, and
these must be harvested manually, thus creating
a labour-intensive element critical to halophyte
crop production (Panta etal., 2014). Often these
halophytes have repeated harvests during the
growing cycle. However, the disadvantage of
labour-intensive manual harvest can be offset by
the advantage of accumulating higher nal yiel ds
per area than crops harvested once per season.
In addition, high crop quality is obtained by har-
vesting only the young shoot tips. Ventura etal.
(2011) obtained the highest yield (16 kg m−2)
during a 6-month harvest period, using the
3-week interval for Salicornia; this is when the
harvest is done every three weeks for a total dur-
ation of 6 months and only the shoots of the Sal-
icornia plant are harvested, hence the harvest
can be repeated multiple times.These types of
studies are crucial to assess the commercial
feasibility of halophytes as gourmet vegetables.
Various factors affect germination rate: for
example, salinity levels of irrigation water applied
to the seeds, and temperature, both inuence
germination potential. Salicornia and Sarcocornia
spp. can be germinated with irrigation water
comprising concentrations of up to 75% sea-
water. Germination limits for Salicornia and Sar-
cocornia spp. occur only at extreme hypersaline
conditions of double the seawater concentra-
tion. Temperature has a strong impact on the
germination rate of Salicornia spp. Germination
usually happens during the winter and spring
months when the sediment salt concentration is
lowest (Davy etal., 2001, 2006). Crithmum maritimum
Humans have consumed C. maritimum (marine
fennel) for centuries, and to date it is still often
gathered from the coastal salt marshes and inland
0004229901.INDD 329 11/2/2018 4:55:22 PM
330 T. Centofanti and G. Bañuelos
saltpans of Europe (Franke, 1982; Wagenvoort
et al., 1989; Davy et al., 2001; Simopoulos,
2004; Tardío etal., 2006).
C. maritimum is a facultative halophyte and
has been largely used for nutritional and medi-
cinal purposes. The plant is rich in several bio-
active substances, minerals, vitamin C, essential
oils and other biomolecules (Atia et al., 2009;
Meot-Duros et al., 2010). It grows in rocky
coastal environments in the Mediterranean re-
gion, where it is often subjected to sea spray
(Ben-Hamed et al., 2004). Traditionally, C. mar-
itimum has been collected and eaten for its anti-
scorbutic properties, owing to the vitamin C
content. This halophyte has the potential to be-
come a multipurpose cash crop (Ventura et al.,
2014). Oils extracted from the plant have shown
the presence of high concentrations of fatty
acids of the omega-3 and omega-6 series, which
have benecial effects against coronary heart
diseases (Guil-Guerrero and Rodriguez-Garcia,
1999). The succulent leaves are consumed fresh
or pickled as salty vegetables and its inores-
cence, in the shape of an umbrella, can be used
for ornamental purposes. As discussed for Sali-
cornia, the germination potential of C. mariti-
mum can hinder its cultivation for large-scale
commercial purposes. Atia etal. (2006) reported
that germination was signicantly inhibited
when NaCl concentrations exceeded 50 mM
(approx. 5 dS m−1). Salsola soda
Salsola soda, more commonly known as ‘agretti’,
is a halophyte native to the Mediterranean basin.
It is a relatively small plant that grows to about
0.7 m on average in soils located within coastal
regions that are at times saturated with salt
water. Throughout history, the plant was a very
important source of soda ash, as people would
extract the ashes from S. soda. The plant is no
longer grown for the use of its soda ash, but ra-
ther is farmed as a vegetable in Mediterranean
countries such as Italy. Various Salsola species
are traditionally used in folk medicine for the
treatment of hypertension, constipation and in-
ammation (Tundis etal., 2009). In this regard,
alkaloid extracts from Salsola species have been
evaluated for the treatment of Alzheimer’s dis-
ease (Tundis et al. 2009). A preliminary study
conducted by Centofanti and Bañuelos (2015)
in central California, USA, showed that Salsola
soda can grow in saline (EC 2 >10 dS m−1) and
B-laden soils (10 mg L−1) of the San Joaquin Val-
ley, CA, USA, and easily tolerate irrigation with
saline and B-rich water (EC 3 dS m−1 and 4 mg B
L−1). In this poor-quality soil, under these grow-
ing conditions, the plant accumulated and re-
moved high amounts of Na (80 g Na kg−1 dry
weight (DW), B (100 mg B kg−1 DW) and Se (3–4
mg Se kg−1 DW) without showing any toxicity
symptoms (Figs 20.1 and 20.2). Hence, S. soda
showed promising potential as a plant species
that can be irrigated and grown in saline and
B-laden conditions and accumulate and harvest
unwanted ions from the soil. We have no direct
evidence of its ability to grow under wetland
growing conditions, although the plant is com-
monly grown under salty marshy regions in
coastal areas of Mediterranean Europe. The plant
can, apparently, survive under continuous wet
and saline growing conditions.
Currently, the authors of this chapter are
investigating the potential of agretti as a vege-
table crop in large-scale eld trials (Fig. 20.3) at
Red Rock Ranch, Five Points, central California,
USA. The goal of the study is to understand how
it can be grown and used on a sustained basis for
phyto-harvest of Na, B and Se eld soil drip-
irrigated with saline drainage water, as well as in
wetland conditions with simulated drainage water
representative of the salinity, B and Se levels
found on the west side of the San Joaquin Valley
in central California. In addition, consumers’
preferences and the potential market demand for
alternative vegetable crops such as this will be
performed in these current studies through eco-
nomic analysis and consumer surveys carried out
at farmers’ markets and retail centres. Currently,
agretti produced in the eld trials at Red Rock
Ranch is sold at farmers’ markets and served in
local restaurants in Fresno, California (Fig. 20.4). Minor gourmet vegetables
Inula crithmoides is consumed in salads or pickled
in vinegar and is a dietary source of iodine. It
also contains a high total lipid content with a
signicant portion of omega-3 fatty acids. The
attractive yellow owers of I. crithmoides make it
a candidate for owering pots and landscaping
in arid saline areas such as Lebanon (Zurayk and
Baalbaki, 1996).
0004229901.INDD 330 11/2/2018 4:55:23 PM
Practical uses of halophytic plants as sources of food and fodder 331
Various minor halophytes used in fresh
salads or pickled are sold as gourmet vegetables
in local markets. Among these are Aster tripoli-
um, Batis maritima and Plantago coronopus,
which are rich in minerals and polyphenols, es-
sential amino acids and tocopherol antioxidants
(vitamin E); there are also high levels of vita-
mins A, C and K, and minerals. Perennial wall
rocket (Diplotaxis tenuifolia) is another halophyte
leafy vegetable commonly cultivated in many
parts of the worlds (de Vos etal., 2013).
Three halophytic plant species, Atriplex
hortensis, A. triangularis and Tetragonia tetrago-
niodes (New Zealand spinach), are alternatives to
spinach (Wilson etal., 2000; Słupski etal., 2010)
owing to their similar chemical composition
Fig. 20.2. Salsola soda (agretti) grown on non-saline soil (Hanford sandy loam) with EC <1 dS m−1, water
soluble B concentration of 0.12 mg L−1, water soluble Se concentration of 0.002 mg L−1 and water soluble
Na concentration of 97 mg L−1. Plants were grown on the soil for 3 weeks and were 37 days old when the
picture was taken.
Fig. 20.1. Salsola soda (agretti) grown on saline soil (Oxalis silty clay loam) with EC >10 dS m−1, water
soluble B concentration of 10 mg L−1, water soluble Se concentration of 0.2 mg L−1 and water soluble Na
concentration of 1470 mg L−1. Plants were grown on the soil for 3 weeks and were 37 days old when the
photograph was taken.
0004229901.INDD 331 11/2/2018 4:55:25 PM
332 T. Centofanti and G. Bañuelos
(Carlsson and Clark, 1983) and their edible
leaves. These halophytes are used for human
consumption in the Netherlands, Belgium and
Portugal (Lieth, 2000; Panta etal., 2014). Yield
of A. triangularis was 21.2 t ha−1 on a FW basis
when grown under seawater irrigation (30 g
NaCl L−1) (Gallagher, 1985). Nutritional aspects of gourmet
In section we have listed the types of
secondary metabolites and nutraceutical com-
pounds that are naturally present in the edible
parts of halophyte vegetables. However, it is
possible to enhance the content of secondary
metabolites by specic agronomic practices (Ven-
tura etal., 2011), including amount and salinity
of irrigation water, plant fertilization, harvest
time and cycle, and harvested plant material
(young or old leaves) (Ventura etal., 2010, 2011,
2013). Little is known about how to implement
these agronomic techniques in large- scale halo-
phyte production, because these studies have
often been conducted in small eld trials for re-
search purposes, rather than for improvement
of commercialization of halophytes.
Microelement deciency may result from
soil alkalization as a side effect of soil saliniza-
tion (Grattan and Grieve, 1999). Ventura et al.
(2013) obtained an improvement of the quality
of leafy vegetable A. tripolium by iron fertilization
because, during cultivation, a leaf chlorosis was
Fig. 20.4. Salsola soda (agretti) grown in a
farmer’s field at Red Rock Ranch, central
California, and served in a restaurant in Fresno,
California, USA.
Fig. 20.3. Salsola soda (agretti) grown in a farmer’s field at Red Rock Ranch, central California, USA.
0004229901.INDD 332 11/2/2018 4:55:29 PM
Practical uses of halophytic plants as sources of food and fodder 333
indirectly induced by the high soil pH (> 8) that
affected the product quality. This species has
been tested for its potential as a halophytic cash
crop in pilot projects in The Netherlands, Bel-
gium and Portugal (Lieth and Mochtchenko,
2002; Geissler etal., 2009).
It is often mentioned that halophyte veget-
ables have potential negative health impacts
because of the high Na and Cl levels in the edible
shoot. However, they may only have a minor
impact on the plant’s nutritional value, because
a gourmet product is consumed in small quan-
tities. In addition, adding little or no salt to the
halophyte dishes during food preparation can
counterbalance the high NaCl content (Ventura
and Sagi, 2013). Many of those aspects related
to the taste of halophytes have not been tested
for consumers’ preferences and attitudes, be-
cause yield has received much more attention
than the effects of salinity on the quality param-
eters of the marketable product (Maggio et al.,
2011). Quinoa
We discuss here the South American seed crop
Chenopodium quinoa because some of the 2500
accessions can tolerate salinity levels present in
seawater (approx. 40 dS m−1). Quinoa can also
tolerate other abiotic stresses such as drought,
frost and water stress (Boscaiu and Vicente,
2013), and it also withstands temperatures
from −4°C to almost 40°C. Interest in this seed
crop is increasing worldwide, owing to its excep-
tional nutritional quality (e.g. Vega-Gálvez etal.,
2010). C. quinoa is considered to contribute to
global food security because of the exceptional
nutritional quality of the seeds, which are rich
in vitamins, minerals, essential amino acids and
fatty acids (Adolf etal., 2013). C. quinoa is con-
sidered a ‘pseudocereal’ as its seeds contain
starch, proteins, essential amino acids (i.e. lysine)
in the nucleus of the grain, unlike wheat and
rice. In addition, quinoa seeds are gluten free
(James, 2009). C. quinoa has a long history of
cultivation, being grown in the Andean region
for thousands of years. Because of the high nu-
tritional quality of its seeds, and its adaptability
to a wide array of climates and environmental
conditions, it has been successfully grown – at a
small scale – in the USA and Canada, several EU
countries, Africa (Kenya) and Asia (the Himalaya
and the plains of northern India) (Boscaiu and
Vicente, 2013). Panta et al. (2014) reported
that C. quinoa had a grain yield potential of up to
5.2 t ha−1 when grown in temperate environme nts
in Argentina. Despite the high environmental
adaptability and high nutritional value of C. qui-
noa, its market is still relatively small, and it is
mainly sold in specialized shops at high prices. Commercialization
of halophytes as food: is there a market
for halophytes as vegetables?
Halophytes have been studied for decades but
the main focus of past research efforts has been
understanding salt tolerance mechanisms. Large-
scale commercial production of halophytes is
still lacking. C. quinoa is one halophyte crop with
a larger market. Commercial cultivation of halo-
phytes requires the selection of superior geno-
types as gourmet vegetables, and denition of
the plant’s reproducible growing conditions
(Ventura and Sagi, 2013). Halophytes should
also be produced at a cost that is competitive
with conventional crops (Brown etal., 2014). In
addition, Brown etal. (2014) point out that the
halophytic crop should not adversely affect water
supplies or land that could support conventional
agriculture. Some of the common objectives of
traditional plant breeding and agricultural prac-
tices should be applied to the domestication of
halophytes for large-scale commercialization.
Some of the most important objectives are: se-
lection of the best genotypes for particular
agro-ecological conditions, reduction in toxic
compounds or anti-nutrient content, increased
yields, improvement of marketing characteris-
tics (uniformity of the product with respect to
taste, size, colour, etc.) and tailoring standard
agricultural practices to particular species (Bos-
caiu and Vicente, 2013).
Besides the development of plant breeding
strategies and agronomic practices for cultiva-
tion of halophytes, another crucial factor for
successful commercialization is consumers’ pref-
erences toward halophyte products and their
willingness to pay for them. There is very little
knowledge about these important aspects, which
are fundamental to the commercialization of
halophytes because farmers will start investing
in the crop only if there is marketing potential.
Consumers’ food choices depend on many factors
0004229901.INDD 333 11/2/2018 4:55:29 PM
334 T. Centofanti and G. Bañuelos
(Clark, 1998) but taste, avour and freshness
are considered the most important attributes in
fresh food quality (Migliore etal., 2015). Migliore
et al. (2015) showed that ‘healthiness’ has an
important impact on consumers’ choice, espe-
cially with regard to prickly pear fruit. Some of
the ‘credence attributes’ (i.e. environmental
conservation and nutritional aspects) can repre-
sent added value in products obtained from
halophytes, and should be studied in depth to
understand their role in consumers’ preferences
towards halophytes. This knowledge could cre-
ate a bigger market demand and trigger farmers’
investment in saline agriculture. The company
OceanDesertFood in The Netherlands represents
one example of halophyte commercialization.
Farmers in Mexico produce Salicornia and then
sell it to OceanDesertFood. Jeannette Hoek, the
company president, is creating a market demand
for halophyte food, fodder and products (OASE
Foundation, 2009). The company produces Sali-
cornia crackers and seaweed chips, which are
among its most popular products.
In general, the gourmet vegetable and herb
market requires the marketed products to be
fresh and visually appealing in respect of colour
and packaging. The product should also have a
particular taste and nutritional value cherished
by its consumers (van der Voort etal., 2007). In
particular, some halophytes can meet the stand-
ards of being ‘functional foods’ for health enthu-
siasts, owing to the high levels of nutritionally
valuable metabolites such as omega-3 fatty acids,
phenolic compounds, antioxidants or minerals
(Ares and Gámbaro, 2007; Del Giudice and Pas-
cucci, 2010). Other factors (credence attributes)
such as environmental conservation claims,
support to local farming, organic or origin certi-
cation, and knowledge and trust of the science,
are more difcult to evaluate and need further
analysis (Moser et al., 2011; Dentoni et al.,
20.3 Oilseed Production for Human
Consumption and Biodiesel
Halophytes are grown commercially for the bio-
fuels generated from the seeds and biomass.
Growing halophytes for biofuels has an import-
ant environmental advantage because these
plants can be gown on marginal saline land and
irrigated with seawater, thus reducing the com-
petition for land with agriculture. Salicornia bige-
lovii is probably the most widely grown halophytes
for oilseed and biodiesel production. Global Sea-
water Inc., a Mexican company, supports com-
mercial biodiesel production of S. bigelovii in
farms in Mexico. Christiansen (2008) reported
that 1 ha of Salicornia grown on coastal land can
produce between 890 and 950 L of biodiesel.
The Masdar Institute of Science and Technology
started a seawater-irrigated cultivation of Sali-
cornia in Abu Dhabi to produce biofuel for the
aviation industry, with the support of Boeing,
Ethiad Airways and UOP Honeywell (ICBA,
2011; Panta et al., 2014). Tamarix spp. and
Euphorbia tirucalii (a desert succulent from East
Africa) are saline- and drought-tolerant plants
that have been grown for biomass production to
generate biofuel. T. jordanis is most suited for
ethanol fermentation as it contains preferred
characteristics such as high cellulose, and low
hemicellulose and phenol content (Eshel et al.,
2010, 2011; Santi etal., 2014). T. aphylla irri-
gated with reclaimed sewage (EC approx. 3 dS
m–1) or brine (EC approx. 7–10 dS m–1) produced
52 and 26 t ha−1, respectively (Ventura et al.,
2014). Further studies on the yield potential of
halophytes for the production of biomass or oil-
seed for biofuels are needed.
In addition to biofuel, seeds from halo-
phytes can also be used to produce edible oils.
Panta etal. (2014) reported that among the 50
seed-bearing halophytic species that are poten-
tial sources of edible oil and proteins the most
studied are: S. bigelovii (Glenn et al., 1991),
Suaeda moquinii (Weber etal., 2001), Kosteletzk-
ya virginica (Gallagher, 1985; He et al., 2003),
S.aralocaspica (Wang etal., 2012), Salvadora per-
sica (Rao etal., 2004; Reddy etal., 2008), Batis
maritima (Marcone, 2003), Crithmum maritimum
and Zygophyllum album (Zarrouk et al., 2003),
Nitraria sibiria, S. salsa, Chenopodium glaucum
and Descurainaia sophia (Yajun et al., 2003).
There have been many trials of S. bigelovii as an
oilseed plant in various parts of the world (ICBA,
2007). Commercial Salicornia production trials
have been carried out in the USA, Middle East
(Jaradat, 2005; Abdal, 2009), India (Ramesh-
kumar and Eswaran, 2013), Mexico (Grattan
et al., 2008) and Africa (Zerai et al., 2010).
S.bigelovii has shown high seed yield (2 t ha−1)
0004229901.INDD 334 11/2/2018 4:55:29 PM
Practical uses of halophytic plants as sources of food and fodder 335
when cultivated under seawater irrigation (40g
NaCl L−1) or at root zone salinities greater than
70 g L−1 of total dissolved salts (Glenn et al.,
1997, 1999). These yield values are equivalent
or higher than freshwater-irrigated oilseed crops
such as sunower and soybean. With respect to
the nutritional value of S. bigelovii oil, the pro-
tein content is about 35% and its properties are
similar to those of safower oil (Glenn et al.,
1991; Zerai etal., 2010). The seeds of Salicornia
are very small (only about 1 mg in weight); this
makes retention at harvest difcult, with seed re-
coveries of only about 75% compared to 90% for
most crops. However, Salicornia seeds have been
successfully harvested using ordinary mechan-
ical equipment (Panta etal., 2014). A 25% lar-
ger seed size was reported in Salicornia plants
that had been cultivated for several crop cycles
and harvested with mechanical equipment, sug-
gesting that at least this species is subject to
improvement through mass selection (Zerai etal.,
2010). Other problems affect the success of
commercial production of Salicornia as oilseed.
At maturity, the plant tends to lie at in the eld
(lodge) and the seeds may shatter before harvest.
Salicornia must grow for 100 days at cool tem-
peratures before owering, to obtain high seed
yields (Glenn et al., 1998). Hence, commercial
production of Salicornia is strictly bound to cli-
matic regions of the subtropics that have cool
winters and hot summers. This region is where
large areas of coastal desert are found, thus fa-
cilitating the use of marginal land for halophyte
Other minor halophytes have been trialled
to obtain data on oilseed yield and quality. The
seeds of Kosteletzkya virginica, the perennial
seashore mallow, contain 32% protein and 22%
lipids (Gallagher, 1985). Suaeda persica can be
cultivated in soils with EC values ranging from
25 to 65 dS m−1, but the maximum seed yield is
obtained at 25–35 dS m−1 because, at higher EC
values (55–65 dS m−1), the seed yield can decline
by 40%–47% (Rao etal., 2004). The seeds con-
tain 40%–45% of oil rich in lauric (C12) and
myristic (C14) acid, widely used in the cosmetic
and pharmaceutical industries (Reddy et al.,
2008). Weber etal. (2007) reported that seeds of
S. fruticosa could be used as a source of edible
oil for human consumption owing to their 74%
unsaturated fatty acid content.
20.4 Halophytes as Forage
In principle, animals could be fed all halophytes
that are used as food for humans, either as vege-
table crops (e.g. Aster tripolium) or grains (e.g.
C. quinoa) (Boscaiu and Vicente, 2013). How-
ever, several factors such as biomass production,
the voluntary feed intake by animals and the nu-
tritional value of the product are still poorly
known, and should be further analysed to make
forage saline agriculture efcient (Norman etal.,
One of the most challenging agricultural
problems in dry lands that have been degraded
through overgrazing is to nd enough forage for
cattle, sheep and goat herds. Livestock such as
sheep, camels and cattle thrive well on certain
halophyte feeds (Swingler etal., 1996; Khan and
Ansari, 2008). Pigs and poultry are the most
sensitive to feed composition and the least sa-
line-tolerant farm animals (Masters etal., 2007).
Some of the most productive halophytes used for
forage are shrubby species of Salicornia, Suaeda,
and Atriplex from the family Chenopodiaceae.
Average yield potential of these halophytes
when irrigated with saline water is 10-20 t ha−1;
this is comparable to the productivity of conven-
tional forage species under non-saline condi-
tions (O’Leary et al., 1985; Glenn et al., 1999;
Masters et al., 2007). Other highly productive
species are the salt grass Distichlis and the succu-
lent viney ground covers such as Batis (Glenn
et al., 1998). D. spicata was examined and
showed considerable promise for selection as a
fodder crop for ruminants (Bustan etal., 2005).
In 2011, the International Centre of Biosa-
line Agriculture (ICBA) started large-scale halo-
phyte forage model farms to study the adaptability
and yield potential of salt-tolerant shrubs, trees
and grasses in the western regions of Abu Dhabi
Emirates (ICBA, 2012). Other studies are cur-
rently ongoing in cooperation with ICBA and
other research organizations in Saudi Arabia,
Pakistan and Bangladesh for the development of
biosaline fodder and forage production using
Sporobolus virginicus, D. spicata and Atriplex spp.
(Panta etal., 2014). The most well-studied halo-
phyte for forage is Atriplex spp. Many farmers
around the world feed Atriplex saltbush to sheep,
which creates a leaner, more hydrated lamb
meat, with higher levels of vitamin E. Of all the
0004229901.INDD 335 11/2/2018 4:55:29 PM
336 T. Centofanti and G. Bañuelos
Atriplex species, A. lentiformis is an important
forage halophyte because its leaves have a nutri-
tional quality (rich N source up to 17% crude
protein under 100% seawater) similar to that of
alfalfa (Pasternak etal., 1985; Pasternak, 1990).
A. lentiformis is a perennial, deep-rooted desert
shrub, which can be grown with seawater irri-
gation, but its seeds need to be germinated in
nurseries because they do not germinate under
high-salt conditions (Boscaiu and Vicente, 2013).
In addition to its salt tolerance, A. lentiformis is
also highly tolerant to drought and extremely ef-
cient in water use. However, the energy content
of the seeds is lower (approximately 15 Mcal kg
DW) than other conventional forages, and the
seeds contain some antinutrients. For example,
the seeds of S. bigelovii contain saponins, which
can have deleterious effects on animal growth
(Glenn et al., 1991, 2013. As a consequence of
these negative attributes the animal intake of
Atriplex spp. was low (0.3–0.4 kg dry matter d−1)
(Pasternak et al., 1985; Pasternak, 1990). The
plant is a perennial but it is advisable to cultivate
it as an annual because it tends to become woody
with age (Glenn etal., 2013).
Forage halophytes are often used as a com-
plementary feed source because of their high
salt content, to avoid meat containing levels of
Na that are too high and unhealthy. The use of
halophytes as feed or fodder for livestock will de-
pend on the specic animals to be fed, some ani-
mals being more sensitive to feed composition
than others. For example, the seed meal remain-
ing after extraction of oil from S. bigelovii seeds
can be used as a protein supplement in sh and
ruminant diets, but it cannot be used as a pro-
tein source in poultry feeds because of its high
saponin content (Glenn et al., 2013). In dry
lands of different climatic zones in Faisalabad,
Peshawar, Bhawalpur and Karachi in Pakistan,
forage halophytes (i.e. Atriplex and Maireana
spp.) represent a viable additional source of for-
age (Hollington etal., 2001). In arid coastal re-
gions where freshwater for crop irrigation is
limited, the straw or seed meal of S. bigelovii fed
to lambs turned out to be an acceptable feed
substitute (Swingle et al., 1996). For example,
S.bigelovii grown in seawater-irrigated agricul-
ture in the eastern coastal region of the Arabian
Peninsula can replace 25% of alfalfa in animal
diets (e.g. sheep) (Abdal, 2009). Other halophytes
are also successful as complementary feed for
animals that tolerate a halophyte-based diet,
such as goat and sheep. Feeding trial with goats
and sheep using 70% Sporobolus virginicus and
Distichlis spicata and 30% conventional feed
(Rhodes grass, Chloris gayana), showed that ani-
mals performed better in meat–fat–bone ratio
and body composition than animals fed on
100% conventional feed (ICBA, 2007). Simi-
larly, 1-year-old cow calves fed only on halophyt-
ic grass (Panicum turgidum) in Pakistan were
leaner than animals fed on maize (Khan and An-
sari, 2008). Other halophytic grasses, such as
Leptochloa fusca and Spartina patens, are grown in
arid lands for forage production. Mangrove and
other coastal halophytic species (i.e. Terminalia
catappa, Aeluropus lagopoides, Cynodon dactylon
and Brachiaria mutica) are fodder for cattle,
camel and goats (Dagar, 2005).
To increase the palatability and voluntary
feed intake in animals it is important to select for
low-salt-accumulating varieties of halophytes
(Ventura etal., 2014).
Animals that feed on a salty diet due to the
presence of halophytes in the diet may increase
their water consumption, which may in turn af-
fect animal weight. In addition to higher animal
requirement for freshwater when the diet is hal-
ophyte-based, all salt ingested by grazing animals
needs to be processed through the kidneys and
this may increase the energy requirement to me-
tabolize the feed (Panta et al., 2014). Further-
more, levels of metabolizable energy come from
the organic matter, which is usually lower in
halophytes (O’Connell etal., 2006). Finally, the
high salinity in the feed of halophyte-based diet
limits the amount an animal can eat and dilutes
the nutritional value, because salt has no calor-
ies but takes up space (Glenn et al., 1998). In
open grazing situations, animals turn to halo-
phytes only when the more palatable plants are
20.5 Conclusion
In this chapter, we have provided an overview of
the advantages and disadvantages of halophytes
as alternative plants for production of vegetables,
feed and fodder. One of the most important
0004229901.INDD 336 11/2/2018 4:55:29 PM
Practical uses of halophytic plants as sources of food and fodder 337
environmental benets is that halophytes can
grow on land that would be otherwise unused
for agricultural production, thus reducing land
competition for food production. In addition,
halophytes can grow on extremely saline soils in
dry lands and can be irrigated with seawater or
brackish water. However, there are many prob-
lems associated with halophyte cultivation. First
and most important, many halophytes need to
be domesticated to reach yields, taste and agri-
cultural practices comparable to conventional
crops. Second, but no less important, is the cre-
ation of a market for successful commercializa-
tion of halophytes as vegetables and animal feed.
Although there are many success stories of
halophyte production and commercialization
(Panta et al., 2014), large-scale and long-term
eld studies are needed to attain protable and
consumer-acceptable products. Studies on con-
sumers’ acceptance of alternative crops are
needed for halophytes to become a recognized
crop that starts to generate economic prots
(Ventura and Sagi, 2013). As in the case of qui-
noa, halophytes’ high nutritional value can be a
leverage to attract consumers’ attention and de-
velop large-base consumption, other than just
being gourmet vegetables in high-end restaur-
ants. Educational programmes and marketing
campaigns can be targeted toward the con-
sumption of halophytes, not only as functional
foods, but also as environment-friendly and less
resource-intense food.
Funds for part of the research reported in this
chapter were provided by the California De-
partment of Water Resources, Proposition 204
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... They are used as gourmet foods, animal feed, and a source of oil for biodiesel production. They have traditionally been used in fresh salad for consumption and for sale at local markets as a vegetable (Centofanti and Bañuelos, 2019). For example, Salicornia europaea L. is one species used as a livestock feed. ...
... Likewise, date palm (Phoenix dactylifera L.) has also been derived from wild halophytes, while the seeds of eelgrass (Zostera marina L.) (Glenn et al. 2013) and Palmer's saltgrass (Distichlis palmeri (Vasey) Fassett ex I.M. Johnston) are edible halophytes that can grow in ocean water and are both sources of food. The grains of D. palmeri are used for the production of bread by Cocopah Indians (Centofanti and Bañuelos 2019). The grains of perennial species are similar to rice (O. ...
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Salinity is one of the major causes of abiotic stress that leads to a reduction in crop yield. One strategy to alleviate and improve crop yield is to use halophytes. These types of plants naturally produce bioactive secondary metabolites, proteins, carbohydrates, and biopolymers that are involved in specialized physiological adaptation mechanisms to alleviate soil salinity. These traits could be leveraged and in turn be the focus of future breeding programs aimed to improve salinity resistance in traditional crops. Recently, the field of nanotechnology has gained the attention of researchers involved in agricultural science and associated disciplines. However, information on salinity tolerance mechanisms of halophytes, based on nanoparticles in agricultural crop plants is limited. Recently, the use of selected halophytic-based nanoparticles has shown to improve crop performance by enhancing the plants ion-flux, improving water efficiency, root hydraulic movement in the favor of plant photosynthesis, the production of proteins involved in oxidation-reduction reactions, reactive oxygen species (ROS) detoxification and hormonal signaling pathways under stress. Therefore, the aim of this review is to highlight the application of halophytic nanoparticles in alleviating salt stress in plants by understanding the mechanisms of plant growth, water relation, ion flux, photosynthesis and the antioxidant defense system. This review also addresses uncertainties, ecotoxicological concerns and associated drawbacks of nanoparticles on the environment. Future research perspectives with respect to the sustainable usage of nanoparticles in saline agriculture has also been presented.
... Furthermore, the lack of infrastructure and technology for efficient and sustainable Salicornia L. cultivation and processing, such as irrigation systems and drying facilities, presents a significant challenge (Chaturvedi et al., 2021;Joshi et al., 2020). Limited human capacity and technical expertise in Salicornia L. cultivation and value addition could also limit productivity and quality (Centofanti and Bañuelos, 2019;Cárdenas-Pérez et al., 2021). Other challenges include limited availability and accessibility of high-quality seeds, fertilizers, and other inputs (Gelfand et al., 2013), environmental factors such as soil salinity, water scarcity, and extreme temperatures that may affect crop productivity (Parida, 2005), competition with other crops and land-use practices that may be more based or profitable (Blanc, 2012), ...
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Halophyte plant Salicornia has potential uses in farming and environmental management. Salicornia is one of the most important families of halophytes and known for its exceptional salt tolerance. It thrives well in saline habitats near coastal areas. A comprehensive review paper provides an overview of Salicornia, including details on the impact of temperature and salinity on the germination of different ecotypes, as well as the influence of day length and salinity on seedling establishment. Salicornia L. presents a promising opportunity for sustainable agriculture and economic development as it may improve the lives and livelihoods of underprivileged groups while also benefiting the environment through carbon sequestration, soil preservation, and biodiversity preservation.
... In order to cope with the high salinity and drought conditions during the summer, A. halimus may use osmotic adjustment (Nedjim 2012; Eissa and Selmy 2015). The tissues of halophyte plants contain more sodium than potassium, so the plants use sodium instead of potassium in physiological processes (Centofanti and Bañuelos 2019;Oi et al. 2022). The same phenomenon was discovered for A. lentiformis, A. undulata, and A. amnicola by Irshad et al. (2000) and by Eissa and Selmy (2015). ...
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Saline agriculture offers promising opportunities for the production of halophytic forage crops under arid and semi-arid climatic conditions. The Mediterranean saltbush (Atriplex halimus L.), a halophytic forage plant that can withstand a variety of harsh climatic conditions, is affected by seasonal climate changes in terms of growth and productivity. The objective of the current study is to comprehend how climatic changes affect the Mediterranean saltbush's growth and productivity in semi-arid environments. Plant samples were collected every 3 months from the Mediterranean saltbush cultivated on saline sandy loam soil (15 dS m −1). Biomass yields, nutritional value, and biochemical components were recorded. The studied plant produced 8.55 Mg ha −1 of dried stems and leaves yearly, with the leaves contributing 31% of the plant's overall dry yield. The dried stems and leaves yield of the Mediterranean saltbush declined by 50 and 45% in the summer season compared to the spring. The summer leaves of the Mediterranean saltbush had higher concentrations of Na, K, and Cl than the winter leaves. In contrast to chlorophyll concentrations, which increased in the spring and decreased throughout the other growing seasons, proline concentrations in the leaf tissues changed throughout the year in the opposite direction. The Mediterranean saltbush reduced the chlorophyll and relative water content in the summer months and increased the leaf Na, K, Cl, and proline. The Mediterranean saltbush plants regulate the proline levels as well as some ions like Na, K, and Cl to achieve an osmotic adjustment in the leaf's tissues. The Mediterranean saltbush plants can produce fodder that is high in protein, nutrients, and nutritional value under conditions of extremely salty soil and irrigation water. These results provide a good opportunity to exploit water and saline lands in the production of animal feed, which helps in implementing sustainable development plans in semi-arid regions.
... Molecular studies have recently shed light on some mechanisms involved in plant salt tolerance, and this may translate into more rapid selection of salt-tolerant genotypes and development of suitable transgenic cultivars (Arzani and Ashraf, 2016;Schroeder et al., 2013;Shah et al., 2018;Sun et al., 2019). Some halophytes have been explored for their potential to be used as crop plants, for example in the production of oil, animal fodder, and even grains (Centofanti and Banuelos, 2019;Esfahan et al., 2010;Glenn et al., 2013). Halophytes as well as salt-tolerant species may also be used in phytoremediation of saline soils (Hayat et al., 2020;Shao et al., 2019;Yue et al., 2020). ...
... The available studies on the quality of animal feed mostly targeted the chemical composition of selected plant species which are used as animal feed, such as those of genera Atriplex, Kochia, Juncus, Suaeda, Salsola, etc. In addition, individual halophytic species from coastal saline areas were screened for their nutritional quality (El-Shaer and Attia-Ismail, 2002), as well as some of terrestrial halophytes as a viable supplementary food source (Centofanti and Banuelos, 2019). It is known that species of the genus Atriplex are often used as an alternative animal feed, especially in conditions of deficiency of adder fodders, such as in early spring, during hot and dry summer months and late autumn (Osman et al., 2006). ...
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Continental halophytic grasslands are known for performing of range of ecosystem services especially remarkable in the regions where they are much distributed – in arid and semi-arid areas. Continental halophytic grasslands of the Central and South-East Europe are not considered as favourable for arable farming, however, traditional animal husbandry plays a crucial role in maintaining biodiversity and preserving these natural habitats. The particular interest of this study is the assessment of the chemical composition of biomass and the quality of insufficiently studied halophytic grassland communities of the central Balkans. In addition, the differences in pastoral value were monitored along the geographical gradient, i.e., between grasslands situated in the Pannonian plain (grasslands of Alliance Puccinellion limosae) and those distributed on the south Serbia (Alliance Festucion pseudovinae). The study was carried out at 18 representative sites, focusing on the saline grasslands of the two distinct regions – on the north and on the south of Serbia. The obtained results show that the halophytic grasslands have satisfying quality characteristics (average values: dry matter - 93.67%, crude protein - 8.66%, cellulose - 30.36%, crude fat - 2.27%) compared with other studied grasslands of saline habitats. There were no significant differences in quality and chemical composition of grasslands of the two regions, despite differences in floristic composition, indicating that salinity is the key determinant for pastoral value of the halophytic vegetation. The certain variations were attributed to the specific floristic composition related to ecological conditions and halophytic community characteristics.
... From all the above referred it is easy to deduce that halophyte plants may find niches in the demanding market for novelties, as for example as herbs, vegetables, fresh gourmet products, and animal feed. One challenge that limits halophyte cultivation is the lack of knowledge about consumers' acceptance of halophyte products, which are fundamental to the commercialization of halophytes because farmers will start investing in the crop only if there is marketing potential (Centofanti and Banuelos, 2019). Halophyte's species will take a little to make people believe that they are good food for them, although some are used by certain communities for this purpose (Menzel and Lieth, 1999). ...
The loss of agro-biodiversity, climate changes and food insecurity are major challenges in the Mediterranean countries with potentially multidimensional consequences. With respect to salinity, approximately 18 million ha, corresponding to 25 % of total irrigated land in the Mediterranean area, are salt affected. Intensive cropping and the excessive use of expensive inputs such as water and fertilizers aggravate this situation. Understanding how we could improve crop productivity in salinized environments is therefore critical to face these challenges. Our comprehension of fundamental physiological mechanisms in plant salt stress adaptation has greatly advanced over the last decades. However, many of these mechanisms have been linked to salt tolerance in simplified experimental systems whereas they have been rarely functionally proven in real agricultural contexts. The sustainability of farming systems in salt affected Mediterranean soils can be effectively achieved by the use of salt-tolerant halophyte plants even more effective through the use of intercropping, crop rotation and aquaponics.. Moreover, if these halophyte plants are removed from the soil to grow other species, pressure on generating salt-tolerant crop plants would be reduced and much healthier crop plants would be cultivated in less stressed saline soils. This paper will focus on the sustainable practices based on the cultivation of halophytes in saline soils by highlighting some experimental activities carried out at laboratory and field levels in the last few years.
Aquaculture is an established technique for producing marine species. However, it creates large amounts of nutrient‐rich waste. Therefore, it is worth examining the methods available to use these nutrients and close the nutrient cycle. This review covers research activities of different saline water methods for food production. These are Integrated Multi‐Trophic Aquaculture (IMTA), Biofloc Technology (BFT) and marine aquaponics. IMTA combines on‐ and offshore aquaculture with macroalgae and filter feeder production. Commercial implementations have been achieved but are not widely used. Especially shrimp are suitable for cultivation in saline BFT systems. Few commercial deployments exist. There is a well‐established market for the products of IMTA and BFT systems. Marine aquaponics is increasingly being tested as part of an expansion of Recirculating Aquaculture Systems (RAS) and BFT systems, but commercial implementations are not known. Moreover, there is no established market for the resulting salt‐tolerant plants (halophytes). Market prices and popularity were evaluated to assess the economic potential of the different plant and aquaculture species farmed with IMTA, BFT and marine aquaponics. There are established markets for well‐known products, but therefore the competition is greater. Niche products are offered occasionally, but often achieve top prices. Closing nutrient cycles and increasing saline water usage for food production as well as high product diversity are necessary to mitigate the increasing pressure on freshwater resources and to be able to feed the growing world population. Therefore, IMTA, BFT and marine aquaponics are promising techniques for reducing pressure on freshwater resources and improving food supply.
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Salinity is one of the oldest and most serious environmental problems in the world. The increasingly widespread salinization of soils and water resources represents a growing threat to agriculture around the world. A strategy to cope with this problem is to cultivate salt-tolerant crops and, therefore, it is necessary to identify plant species that are naturally adapted to high-salinity conditions. In this review, we focus our attention on some plant species that can be considered among the most representative halophytes of the Mediterranean region; they can be potential resources, such as new or relatively new vegetable crops, to produce raw or minimally processed (or ready-to-eat) products, considering their nutritional properties and nutraceuticals. The main biological and agronomic characteristics of these species and the potential health risks due to mycotoxigenic fungi have been analyzed and summarized in a dedicated section. The objective of this review is to illustrate the main biological and agronomical characteristics of the most common halophytic species in the Mediterranean area, which could expand the range of leafy vegetables on the market.
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Sesuvium sesuvioides(Fenzl) Verdc. (Aizoaceae) is commonly known as BarriUlwaiti and used in folklore remedies; i.e. arthritis, gout, epistaxis, hemorrhage, smallpox, chickenpox, cold and flu by the local practitioners in the Cholistan desert. In the current study, fresh and dried plant material was examined macroscopically and microscopically. Transverse sections of plant parts such as leaf, stem, root and flower were also examined. Physico-chemical and fluorescence analysis according to WHO recommendations for standardization of plant material were performed. Phytochemical screening maybe helpful in determining the secondary metabolites responsible for their biological activities. Mineral analysis (Na + , K + , Li + , Ca 2+ , Mg 2+ , Cl ˗ , Zn 2+ , Cu 2+ and Fe 2+), total fat and crude proteins were estimated to evaluate the nutritional value of the plant. In in-vitro cytotoxic activity, n-hexane fraction (50µg) showed significant results against Human T-lymphoblastic Leukemia CCRF-CEM cell lines followed by methanol and chloroform fractions. This study will be worthwhile for the correct identification and for observing any type of adulteration. This observation will be helpful for differentiating this species from closely related species of the same genus or family.
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Not less than about 400 million ha and perhaps as much as 950 million ha of land in arid and semi-arid regions may be salt-affected from natural and anthropogenic causes (Massoud 1974, Ponnamperuma 1977). Definitive data on the annual worldwide loss of farmland due to salinization and related causes is lacking. However salinity is unquestionably the most important problem of irrigated agriculture (Dregne 1977), and one-fifth of the irrigated lands of the world, approximately 47 million ha, is salt-affected today (Maas & Hoffman 1977).
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Some Tunisian salty areas have been explored for halophytes presenting an economical interest. Plants were mainly chosen which could be considered as oilseed species. Three interesting species have been identified as oil producers, called Zygophyllum album, Zygophyllaceae, Cakile maritima, Cruciferous and Crithmum maritimum, Umbelliferous. Oils extractions were carried out on ripe seeds collected from their natural biotope. Some physiological and biochemical aspects have been studied such as dry weight of 100 seeds, oil content, lipid and fatty acid composition. Considering seeds size criteria, it appeared that seeds of investigated species have a suitable size for harvesting. In fact, the dry weight of 100 seeds varied from 132.6 mg in Zygophyllum to 773.9 mg in Cakile. The content of dry matter in seeds of the latter halophyte is nearly 3 times higher when compared to rape (Brassica napus) seeds, a conventional oleaginous plant. In contrast, seeds of the two other halophytes contain a lot less dry matter than Cakile, representing approximately 50 % of that of rape ones. Concerning oil content and fatty acid composition, analyses showed that seeds of Cakile and Crithmum present high levels of oil forming respectively 42 and 30 % on dry weight basis. However, oil content in Zygophyllum seeds is very low, no more than 6 %. Like for olive, fatty acid composition of Crithmum seeds is characterized by a high level of oleic acid (81 %) whereas that of Zygophyllum, characterized by a high percentage of linoleic acid (64 %), looks like sunflower oil composition. Therefore, the two latter species contain oils of good quality which can be used without any further modification. In opposite, oil of Cakile, rich in erucic acid (25 %) may be used for industrial applications.
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Many aquaculture systems generate high amounts of wastewater containing compounds such as suspended solids, total nitrogen and total phosphorus. Today, aquaculture is imperative because fish demand is increasing. However, the load of waste is directly proportional to the fish production. Therefore, it is necessary to develop more intensive fish culture with efficient systems for wastewater treatment. A number of physical, chemical and biological methods used in conventional wastewater treatment have been applied in aquaculture systems. Constructed wetlands technology is becoming more and more important in recirculating aquaculture systems (RAS) because wetlands have proven to be well-established and a cost-effective method for treating wastewater. This review gives an overview about possibilities to avoid the pollution of water resources; it focuses initially on the use of systems combining aquaculture and plants with a historical review of aquaculture and the treatment of its effluents. It discusses the present state, taking into account the load of pollutants in wastewater such as nitrates and phosphates, and finishes with recommendations to prevent or at least reduce the pollution of water resources in the future.
We discuss the process of domesticating wild halophytes to serve as crop plants using seawater irrigation. First steps in this domestication involve determining whether halophyte species exist that may produce significant amounts of a usable product under seawater irrigation and that this is a sustainable agronomic practice. This is followed by development of strategies to improve crop productivity via selecting appropriate species for domestication and then affecting agronomic traits through plant breeding. We demonstrate that halophytes may be productive under seawater irrigation, that this management system may be sustainable, and there are demonstrated pathways toward domestication.
In China, the saline soil area, as an important soil resource, is about 99,130,000 ha, in which the modern saline soil area is 36,930,000 ha, and the potential saline soil area is about 17,330,000 ha (Wangzunqin, 1993). The saline soil is mainly distributed in low and coastal regions, such as Hebei, Shandong, Liaoning, Jiangsu provinces. On the other hand, in inland, Inner Mongolia and Xinjiang, there is a larger area of saline soil too in arid regions.