Production of halophytes as the oilseed provides another dimension of the use of the salinity‐degraded land and gain alternative agriculture practices in the salt‐stress conditions. Several species from the genus Salicornia, Suaeda, and other halophytes showed no accumulation of the salt in the seed, no matter what saline conditions they were grown (Panta et al. 2014). The Salicornia bigelovii showed the higher yield potential of the seeds, even at the soil salinity equivalent to 70 g/l NaCl concentration (Glenn et al. 1999). When the S. bigelovii growing area was irrigated with the seawater having salt concentration equivalent to 40 g/l, it was able to produce seed at a rate of 2 t/ha (Glenn et al. 1999). This yield potential is equivalent to the oilseed production from the conventional crop in the nonsaline conditions (Panta et al. 2014). It must be noted that S. bigelovii not only has the higher yield potential in the salt‐affected areas but its seed is also rich in the oil (30%) with similar properties to the oil derived from the safflower (Glenn et al. 1991; Zerai et al. 2010) and protein (35%) content (Panta et al. 2014). These findings provide hope to the farmers across the globe whose agricultural land is degraded due to the salinity and can help us minimize demand load on the glycophytic crop to increase their productivity to feed the growing population. The higher oil yield potential of the halophyte seed opens the opportunity to use the oil produced from the halophytes for biodiesel production without competing with oilseed crops used for food. In a trial at the Mexico coast for oilseed productivity, the S. bigelovii showed higher seed productivity in seawater irrigated fields than the other oil seeds, such as the mustard, safflower, and sunflower grown under nonsaline conditions (Glenn et al. 2009; Panta et al. 2014). The perennial halophytic Tamarix species can grow and survive in drought and flooding conditions and is salinity tolerant (Panta et al. 2014). The plant has very high yield potential ranging from 19.5 to 52 t/ha/year and heat value relative to standard coal at the ratio of 1/0.7 (Panta et al. 2014). Higher heat value and biomass yield potential make it an important halophytic candidate for the thermal electricity generation and lignocellulosic second generation biofuel production. The cooperation of Boeing, Etihad Airways, and UOP Honeywell with the Masdar Institute of Science and Technology to grow Salicornia species at the Abu Dhabi for production of aviation industry‐grade biofuel can exemplify the importance of biofuel production using the halophytes (Panta et al. 2014).
The unique ability of halophytes to accumulate salt ions in the root and shoot vacuole with very low leakage provides them the unique ability to extract salts from the soil. It thus may act in the desalinization of degraded land. Some halophytes such as S. brachiata (40% of dry weight) and Atriplex sp. (39% of the dry weight) serve as the salt accumulator (Barrett‐Lennard 2002) and therefore can be extensively used for the phytoremediation. Panta et al. (2014) have provided a table of the halophytes from the previously published report, which describes different species from the genus Suaeda, Atriplex, Tectocornia, Sesuvium, Anthrocnemum, Excoecarcia, Ipomoea, Batis, and Salicornia, which were used in past or have a high potential in coming future for the desalinization and removal of the heavy metals from the degraded soil.
2.7 Conclusion and Future Perspectives
Salt stress affects plants’ survival and productivity. Research on understanding the mechanism of salinity stress gave us significant insight into the plants’ response to salt stress and how halophytes respond differently than the glycophytes. However, the complete mechanistic understanding of the salt tolerance mechanism in the halophytes at the developmental and intracellular molecular levels is lacking. To meet the food, feed, fiber, and fuel demand of the growing human population and improve the abiotic stress resilience of the crop, there are two ways to pursue in parallel; one, adopting the halophyte for food, forage, and fuel generation by growing them in the salt‐affected land, and, second, understanding the salt‐stress tolerance mechanism in the halophytes and then engineering the genome of glycophytic crop plants for better salt‐stress tolerance. Halophytes grown by seawater irrigation have shown their significance by producing a higher yield of oil and protein‐rich seeds than a conventional oilseed crop, which suggested that the salt‐degraded land can be used for growing halophytes for edible oil or biofuel. Halophytes were also used to restore the wasteland by desalinization and phytoremediation of the heavy metals, used as forage for the cattle and vegetable salads in different parts of the world. The halophyte C. quinoa seeds are rich in minerals and vitamins and gained popularity as mainstream food because of its gluten‐free nature. For improving the salt‐stress tolerance in the crop plants, the current requirement is to understand salt‐stress sensory system at the root, root to shoot signaling, and intracellular sensory and responses at the cell organelles such as chloroplasts, mitochondria, or peroxisomes in halophytes. In halophytes, yet very little is known about the development and functioning mechanism of the EBC or different types of the salt glands. Understanding these mechanisms and engineering, the crop using the genome‐editing approach may improve the salt‐stress tolerance of the crops within a shorter time frame.
References
1 Abbasi, F.M. and Komatsu, S. (2004). A proteomic approach to analyze salt‐responsive proteins in rice leaf sheath. Proteomics 4: 2072–2081.
2 Babu, G.A. and Reddy, S.M. (2011). Diversity of arbuscular mycorrhizal fungi associated with plants growing in fly ash pond and their potential role in ecological restoration. Curr. Microbiol. 63: 273–280.
3 Bao, H., Chen, X., Lv, S. et al. (2015). Virus‐induced gene silencing reveals control of reactive oxygen species accumulation and salt tolerance in tomato by γ‐aminobutyric acid metabolic pathway. Plant Cell Environ. 38: 600–613.
4 Barrett‐Lennard, E.G. (2002). Restoration of saline land through revegetation. Agric. Water Manag. 53: 213–226.
5 Bose, J., Munns, R., Shabala, S. et al. (2017). Chloroplast function and ion regulation in plants growing on saline soils: lessons from halophytes. J. Exp. Bot. 68: 3129–3143.
6 Busch, K., Piehler, J., and Fromm, H. (2000). Plant succinic semialdehyde dehydrogenase: dissection of nucleotide binding by surface plasmon resonance and fluorescence spectroscopy. Biochemistry 39: 10110–10117.
7 Chattopadhyay, A., Subba, P., Pandey, A. et al. (2011). Analysis of the grasspea proteome and identification of stress‐responsive proteins upon exposure to high salinity, low temperature, and abscisic acid treatment. Phytochemistry 72: 1293–1307.
8 Chaudhary, D.R., Rathore, A.P., and Jha, B. (2018). Aboveground, belowground biomass and nutrients pool in Salicornia brachiata at coastal area of India: interactive effects of soil characteristics. Ecol. Res. 33: 1207–1218.
9 Che‐Othman, M.H., Millar, A.H., and Taylor, N.L. (2017). Connecting salt stress signalling pathways with salinity‐induced changes in mitochondrial metabolic processes in C3 plants. Plant Cell Environ. 40: 2875–2905.
10 Chiang, C.P., Yim, W.C., Sun, Y.H. et al. (2016). Identification of ice plant (Mesembryanthemum crystallinum L.) microRNAs using RNA‐seq and their putative roles in high salinity responses in seedlings. Front. Plant Sci. 7: 1143.
11 Choi, W.G., Toyota, M., Kim, S.H. et al. (2014). Salt stress‐induced Ca2+ waves are associated with rapid, long‐distance root‐to‐shoot signaling in plants. Proc. Natl. Acad. Sci. U. S. A. 111: 6497–6502.
12 Christmann, A., Grill, E., and Huang, J. (2013). Hydraulic signals in long‐distance signaling. Curr. Opin. Plant Biol. 16: 203–300.
13 Dassanayake, M. and Larkin, J.C. (2017). Making plants break a sweat: the structure, function, and evolution of plant salt glands. Front. Plant Sci. 8: 1–20.
14 Dumont, S. and Rivoal, J. (2019). Consequences of oxidative stress on plant glycolytic and respiratory metabolism. Front. Plant Sci. 10: 1–16.
15 Fahy, D., Sanad, M.N.M.E., Duscha, K. et al. (2017). Impact of salt stress, cell death, and autophagy on peroxisomes: Quantitative and morphological analyses using small fluorescent probe N‐BODIPY. Sci. Rep. 7: 1–17.
16 FAO