Wild Halophytes: Tools for Understanding Salt Tolerance Mechanisms of Plants and for Adapting Agriculture to Climate Change II

A special issue of Plants (ISSN 2223-7747). This special issue belongs to the section "Plant Response to Abiotic Stress and Climate Change".

Deadline for manuscript submissions: closed (30 June 2024) | Viewed by 13483

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Guest Editor
Faculty of Medicine and Biological Sciences, Stefan cel Mare University of Suceava, 13 University Street, 720229 Suceava, Romania
Interests: halophytes; halophytes anatomy and ecology; ecophysiology of halophytes; plant abiotic stress; conceptual and historical approach of halophytes
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Guest Editor
Institute for the Conservation and Improvement of Valencian Agrodiversity (COMAV), Universitat Politècnica de València, Camino de Vera 14, 46022 Valencia, Spain
Interests: androgenesis, microspores; plant reproduction; salinity; genome editing; plant biotechnology; stress tolerance

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Guest Editor
Institute for the Conservation and Improvement of Valencian Agrodiversity (COMAV), Universitat Politècnica de València, Camino de Vera 14, 46022 Valencia, Spain
Interests: climate change; plant biotechnology; plant reproduction; abiotic stress; plant stress physiology; halophytes; drought; salinity; stress tolerance; biostimulants
Special Issues, Collections and Topics in MDPI journals

Special Issue Information

Dear Colleagues,

All our major crops and most plant wild species are glycophytes, sensitive to relatively low salt levels in the soil. On the contrary, a relatively small group of plants—the halophytes—are adapted to natural saline environments and can survive and complete their life cycle in habitats with soil salinity equivalent to 200 mM NaCl, although some can withstand salinities even higher than that of seawater. These saline habitats are fascinating from an ecological perspective, but also very much threatened by human activities and extremely sensitive to climate change effects. The general increase in average temperatures; the more extended, frequent and intense drought periods; floods; changes in normal weather patterns; and the increase in salinity levels in different saline environments brought about by climate change are altering the distribution of wild plants in Nature, significantly affecting halophytes. The study of their response mechanisms to these abiotic stress factors constitutes a relevant topic in current plant biology research.

Climate change also represents a major challenge for agriculture and food security, now and in the foreseeable future. Soil salinity is, together with drought, one of the leading causes of the reduction of crop yields worldwide, and climate change is contributing to the increasing loss of irrigated cropland due to secondary salinization, especially in arid and semiarid regions. The most promising strategy to address this problem should be based on the genetic improvement of crop salt tolerance. This approach, in turn, requires a deep understanding of salt tolerance mechanisms. All plants, tolerant or not, share a series of basic, conserved responses to salt stress (control of ion transport, osmolyte biosynthesis, activation of antioxidant systems, synthesis of ‘protective’ proteins), although the specific mechanisms of tolerance can vary widely in different species. Therefore, halophytes are ideal subjects for fundamental studies of salt-tolerance mechanisms in plants, in general, at the physiological, biochemical, and molecular levels. These studies complement and extend those most frequently carried out using non-tolerant model species, such as Arabidopsis thaliana. Besides a purely scientific interest in halophytes as a source of basic knowledge, halophytes can also provide biotechnological tools (i.e., salt-tolerance genes and salt-induced promoters) for the genetic improvement of the salt tolerance of conventional crops. Furthermore, some halophytes could represent the basis of a sustainable ‘saline agriculture’, being commercially grown for food, feed or the production of metabolites of medical, nutraceutical or industrial interest in salinized land and irrigated with brackish or saline water, not competing with our conventional crops for these limited resources, fertile land and good-quality water for irrigation.

The basic and applied aspects of halophytes research mentioned above have been addressed in a Plants Special Issue, “Wild Halophytes: Tools for Understanding Salt Tolerance Mechanisms of Plants and for Adapting Agriculture to Climate Change”, recently published as an eBook (https://doi.org/10.3390/books978-3-0365-6572-9). Most articles included in the Special Issue dealt with the elucidation of salt (and/or drought) stress tolerance mechanisms under controlled greenhouse conditions, in several cases performing comparative analyses of the stress responses of taxonomically related taxa, using physiological and biochemical approaches. Some other papers refer to field studies, phytochemical analyses, or biotechnological applications of halophytes in phytoremediation or as a source of metabolites of medical/nutritional interest.

We are proud to launch this Special Issue’s second edition, which will again cover all biological and biotechnological aspects of halophytes research mentioned above, reflected in original research papers, reviews, minireviews or opinion papers. Those topics or experimental strategies not addressed or underrepresented in the first edition will be especially welcome: halophyte ecophysiology, investigation of stress-tolerance mechanisms using molecular biology or ‘omics’ approaches, and agronomic assessments of halophytes as ‘new’ crops for saline agriculture.

Dr. Marius-Nicusor Grigore
Dr. Ricardo Mir
Prof. Dr. Oscar Vicente
Guest Editors

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Keywords

  • abiotic stress
  • antioxidant systems
  • climate change
  • ecophysiology
  • drought
  • ion transport
  • osmolyte accumulation
  • plant breeding
  • saline agriculture
  • salinity
  • salt stress responses
  • salt tolerance

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Related Special Issue

Published Papers (8 papers)

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Research

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19 pages, 3368 KiB  
Article
Thriving under Salinity: Growth, Ecophysiology and Proteomic Insights into the Tolerance Mechanisms of Obligate Halophyte Suaeda fruticosa
by Bilquees Gul, Abdul Hameed, Muhammad Zaheer Ahmed, Tabassum Hussain, Sarwat Ghulam Rasool and Brent L. Nielsen
Plants 2024, 13(11), 1529; https://doi.org/10.3390/plants13111529 - 31 May 2024
Cited by 1 | Viewed by 1001
Abstract
Studies on obligate halophytes combining eco-physiological techniques and proteomic analysis are crucial for understanding salinity tolerance mechanisms but are limited. We thus examined growth, water relations, ion homeostasis, photosynthesis, oxidative stress mitigation and proteomic responses of an obligate halophyte Suaeda fruticosa to increasing [...] Read more.
Studies on obligate halophytes combining eco-physiological techniques and proteomic analysis are crucial for understanding salinity tolerance mechanisms but are limited. We thus examined growth, water relations, ion homeostasis, photosynthesis, oxidative stress mitigation and proteomic responses of an obligate halophyte Suaeda fruticosa to increasing salinity under semi-hydroponic culture. Most biomass parameters increased under moderate (300 mmol L−1 of NaCl) salinity, while high (900 mmol L−1 of NaCl) salinity caused some reduction in biomass parameters. Under moderate salinity, plants showed effective osmotic adjustment with concomitant accumulation of Na+ in both roots and leaves. Accumulation of Na+ did not accompany nutrient deficiency, damage to photosynthetic machinery and oxidative damage in plants treated with 300 mmol L−1 of NaCl. Under high salinity, plants showed further decline in sap osmotic potential with higher Na+ accumulation that did not coincide with a decline in relative water content, Fv/Fm, and oxidative damage markers (H2O2 and MDA). There were 22, 54 and 7 proteins in optimal salinity and 29, 46 and 8 proteins in high salinity treatment that were up-regulated, down-regulated or exhibited no change, respectively, as compared to control plants. These data indicate that biomass reduction in S. fruticosa at high salinity might result primarily from increased energetic cost rather than ionic toxicity. Full article
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25 pages, 3360 KiB  
Article
Temporal Changes in Biochemical Responses to Salt Stress in Three Salicornia Species
by Hengameh Homayouni, Hooman Razi, Mahmoud Izadi, Abbas Alemzadeh, Seyed Abdolreza Kazemeini, Ali Niazi and Oscar Vicente
Plants 2024, 13(7), 979; https://doi.org/10.3390/plants13070979 - 29 Mar 2024
Cited by 3 | Viewed by 1264
Abstract
Halophytes adapt to salinity using different biochemical response mechanisms. Temporal measurements of biochemical parameters over a period of exposure to salinity may clarify the patterns and kinetics of stress responses in halophytes. This study aimed to evaluate short-term temporal changes in shoot biomass [...] Read more.
Halophytes adapt to salinity using different biochemical response mechanisms. Temporal measurements of biochemical parameters over a period of exposure to salinity may clarify the patterns and kinetics of stress responses in halophytes. This study aimed to evaluate short-term temporal changes in shoot biomass and several biochemical variables, including the contents of photosynthetic pigments, ions (Na+, K+, Ca2+, and Mg2+), osmolytes (proline and glycine betaine), oxidative stress markers (H2O2 and malondialdehyde), and antioxidant enzymes (superoxide dismutase, peroxidase, catalase, and ascorbate peroxidase) activities of three halophytic Salicornia species (S. persica, S. europaea, and S. bigelovii) in response to non-saline, moderate (300 mM NaCl), and high (500 mM NaCl) salinity treatments at three sampling times. Salicornia plants showed maximum shoot biomass under moderate salinity conditions. The results indicated that high Na+ accumulation in the shoots, coupled with the relative retention of K+ and Ca2+ under salt stress conditions, contributed significantly to ionic and osmotic balance and salinity tolerance in the tested Salicornia species. Glycine betaine accumulation, both constitutive and salt-induced, also seems to play a crucial role in osmotic adjustment in Salicornia plants subjected to salinity treatments. Salicornia species possess an efficient antioxidant enzyme system that largely relies on the ascorbate peroxidase and peroxidase activities to partly counteract salt-induced oxidative stress. The results also revealed that S. persica exhibited higher salinity tolerance than S. europaea and S. bigelovii, as shown by better plant growth under moderate and high salinity. This higher tolerance was associated with higher peroxidase activities and increased glycine betaine and proline accumulation in S. persica. Taking all the data together, this study allowed the identification of the biochemical mechanisms contributing significantly to salinity tolerance of Salicornia through the maintenance of ion and osmotic homeostasis and protection against oxidative stress. Full article
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25 pages, 14466 KiB  
Article
Arthrocnemum Moq.: Unlocking Opportunities for Biosaline Agriculture and Improved Human Nutrition
by Esteban Ramírez, Nuria Rodríguez and Vicenta de la Fuente
Plants 2024, 13(4), 496; https://doi.org/10.3390/plants13040496 - 9 Feb 2024
Cited by 1 | Viewed by 1414
Abstract
(1) Background: This study provides novel insights into the elemental content and biomineralization processes of two halophytic species of the genus Arthrocnemum Moq. (A. macrostachyum and A. meridionale). (2) Methods: Elemental content was analyzed using ICP-MS, while biominerals were detected through [...] Read more.
(1) Background: This study provides novel insights into the elemental content and biomineralization processes of two halophytic species of the genus Arthrocnemum Moq. (A. macrostachyum and A. meridionale). (2) Methods: Elemental content was analyzed using ICP-MS, while biominerals were detected through electron microscopy (SEM and TEM) and X-ray diffraction. (3) Results: The elemental content showed significant concentrations of macronutrients (sodium, potassium, magnesium, and calcium) and micronutrients, especially iron. Iron was consistently found as ferritin in A. macrostachyum chloroplasts. Notably, A. macrostachyum populations from the Center of the Iberian Peninsula exhibited exceptionally high magnesium content, with values that exceeded 40,000 mg/kg d.w. Succulent stems showed elemental content consistent with the minerals identified through X-ray diffraction analysis (halite, sylvite, natroxalate, and glushinskite). Seed analysis revealed elevated levels of macro- and micronutrients and the absence of heavy metals. Additionally, the presence of reduced sodium chloride crystals in the seed edges suggested a mechanism to mitigate potential sodium toxicity. (4) Conclusions: These findings highlight the potential of Arthrocnemum species as emerging edible halophytes with nutritional properties, particularly in Western European Mediterranean territories and North Africa. They offer promising prospects for biosaline agriculture and biotechnology applications. Full article
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20 pages, 3116 KiB  
Article
Salt Tolerance of Sea Flax (Linum maritimum L.), a Rare Species with Conservation Interest in Eastern Spain
by Diana M. Mircea, P. Pablo Ferrer-Gallego, Inmaculada Ferrando-Pardo, Oscar Vicente, Ricardo Mir and Monica Boscaiu
Plants 2024, 13(2), 305; https://doi.org/10.3390/plants13020305 - 19 Jan 2024
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Abstract
Seldom found in saltmarshes, Linum maritimum is a halophyte of great conservation interest in the eastern Iberian Peninsula. Although the species has been reported in different plant communities, there is no information on its range of salinity tolerance or mechanisms of response to [...] Read more.
Seldom found in saltmarshes, Linum maritimum is a halophyte of great conservation interest in the eastern Iberian Peninsula. Although the species has been reported in different plant communities, there is no information on its range of salinity tolerance or mechanisms of response to environmental stress factors. In this study, L. maritimum plants were subjected to increasing salt concentrations in controlled conditions in a greenhouse. After six months of watering with salt solutions, only plants from the control, 50 mM and 100 mM NaCl treatment groups survived, but seeds were produced only in the first two. Significant differences were found between the plants from the various treatment groups in terms of their growth parameters, such as plant height, fresh weight, and the quantity of flowers and fruits. The main mechanism of salt tolerance is probably related to the species’ ability to activate K+ uptake and transport to shoots to partly counteract the accumulation of toxic Na+ ions. A biochemical analysis showed significant increases in glycine betaine, flavonoids and total phenolic compounds, highlighting the importance of osmotic regulation and antioxidant compounds in the salt tolerance of Linum maritimum. These findings have implications for the conservation of the species, especially under changing climatic conditions that may lead to increased soil salinity in its Mediterranean distribution area. Full article
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19 pages, 2205 KiB  
Article
Salicylic Acid- and Potassium-Enhanced Resilience of Quinoa (Chenopodium quinoa Willd.) against Salinity and Cadmium Stress through Mitigating Ionic and Oxidative Stress
by Sameera A. Alghamdi, Hesham F. Alharby, Ghulam Abbas, Habeeb M. Al-Solami, Afshan Younas, Majed Aldehri, Nadiyah M. Alabdallah and Yinglong Chen
Plants 2023, 12(19), 3450; https://doi.org/10.3390/plants12193450 - 30 Sep 2023
Cited by 5 | Viewed by 1542
Abstract
Salinity and cadmium (Cd) contamination of soil are serious environmental issues threatening food security. This study investigated the role of salicylic acid (SA) and potassium (K) in enhancing the resilience of quinoa against the combined stress of salinity and Cd. Quinoa plants were [...] Read more.
Salinity and cadmium (Cd) contamination of soil are serious environmental issues threatening food security. This study investigated the role of salicylic acid (SA) and potassium (K) in enhancing the resilience of quinoa against the combined stress of salinity and Cd. Quinoa plants were grown under NaCl (0, 200 mM) and Cd (0, 100 µM) stress, with the addition of 0.1 mM SA and 10 mM K, separately or in combination. The joint stress of Cd and NaCl caused >50% decrease in plant growth, chlorophyll contents, and stomatal conductance compared to the control plants. The higher accumulation of Na and Cd reduced the uptake of K in quinoa tissues. The joint stress of salinity and Cd caused an 11-fold increase in hydrogen peroxide and 13-fold increase in thiobarbituric acid reactive substances contents, and caused a 61% decrease in membrane stability. An external supply of 0.1 mM SA and 10 mM K helped plants to better adapt to salinity and Cd stress with less of a reduction in plant biomass (shoot 19% and root 24%) and less accumulation of Na and Cd in plant tissues. The activities of superoxide dismutase (SOD), catalase (CAT), peroxidase (POD), and ascorbate peroxidase (APX) were enhanced by 11-fold, 10-fold, 7.7-fold, and 7-fold, respectively, when SA and K were applied together to the plants subjected to the joint stress of Cd and salinity. Based on the values of the bioconcentration factor (>1), the translocation factor (<1), and the higher tolerance index, it was clear that Cd-contaminated, salty soils could be stabilized with quinoa under the combined supply of SA and K. Full article
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Review

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12 pages, 1057 KiB  
Review
ROS Homeostasis and Antioxidants in the Halophytic Plants and Seeds
by Hadi Pirasteh-Anosheh, Maryam Samadi, Seyed Abdolreza Kazemeini, Munir Ozturk, Agnieszka Ludwiczak and Agnieszka Piernik
Plants 2023, 12(17), 3023; https://doi.org/10.3390/plants12173023 - 22 Aug 2023
Cited by 8 | Viewed by 1835
Abstract
Reactive oxygen species (ROS) are excited or partially reduced forms of atmospheric oxygen, which are continuously produced during aerobic metabolism like many physiochemical processes operating throughout seed life. Previously, it was believed that ROS are merely cytotoxic molecules, however, now it has been [...] Read more.
Reactive oxygen species (ROS) are excited or partially reduced forms of atmospheric oxygen, which are continuously produced during aerobic metabolism like many physiochemical processes operating throughout seed life. Previously, it was believed that ROS are merely cytotoxic molecules, however, now it has been established that they perform numerous beneficial functions in plants including many critical roles in seed physiology. ROS facilitate seed germination via cell wall loosening, endosperm weakening, signaling, and decreasing abscisic acid (ABA) levels. Most of the existing knowledge about ROS homeostasis and functions is based on the seeds of common plants or model ones. There is little information about the role of ROS in the germination process of halophyte seeds. There are several definitions for halophytic plants, however, we believed “halophytes are plants that can grow in very saline environment and complete their life cycle by adopting various phenological, morphological and physiological mechanisms at canopy, plant, organelle and molecular scales”. Furthermore, mechanisms underlying ROS functions such as downstream targets, cross-talk with other molecules, and alternative routes are still obscure. The primary objective of this review is to decipher the mechanisms of ROS homeostasis in halophytes and dry seeds, as well as ROS flux in germinating seeds of halophytes. Full article
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18 pages, 335 KiB  
Review
Economic Uses of Salt-Tolerant Plants
by Pedro Garcia-Caparros, Mohammed J. Al-Azzawi and Timothy J. Flowers
Plants 2023, 12(14), 2669; https://doi.org/10.3390/plants12142669 - 17 Jul 2023
Cited by 8 | Viewed by 2453
Abstract
Climate change is likely to affect the ability of world agricultural systems to provide food, fibre, and fuel for the growing world population, especially since the area of salinised land will increase. However, as few species of plants (less than 1% of all [...] Read more.
Climate change is likely to affect the ability of world agricultural systems to provide food, fibre, and fuel for the growing world population, especially since the area of salinised land will increase. However, as few species of plants (less than 1% of all plant species) can tolerate saline soils, we believe it is important to evaluate their potential as crops for salinised soils. We have analysed the economic and potential economic uses of plants that are listed in the database eHALOPH, including the most tolerant species, halophytes. For nine main categories of economic value, we found a total of 1365 uses amongst all species listed in eHALOPH as of July 2022; this number reduced to 918 amongst halophytes. We did not find any obvious differences in rankings between the more tolerant halophytes and the whole group of salt-tolerant plants, where the order of use was medical, followed by forage, traditional medicine, food and drink, fuel, fuelwood, and bioenergy. While many species are potentially important as crops, the effects of salt concentration on their uses are much less well documented. Increasing salt concentration can increase, decrease, or have no effect on the concentration of antioxidants found in different species, but there is little evidence on the effect of salinity on potential yield (the product of concentration and biomass). The effect of salinity on forage quality again varies with species, often being reduced, but the overall consequences for livestock production have rarely been evaluated. Salt-tolerant plants have potential uses in the bioremediation of degraded land (including revegetation, phytoremediation, and extraction of NaCl) as well as sources of biofuels, although any use of saline water for the sustainable irrigation of salt-tolerant crops must be viewed with extreme caution. Full article

Other

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12 pages, 646 KiB  
Opinion
The Sustainable Use of Halophytes in Salt-Affected Land: State-of-the-Art and Next Steps in a Saltier World
by Nadia Bazihizina, Jutta Papenbrock, Henrik Aronsson, Karim Ben Hamed, Özkan Elmaz, Zenepe Dafku, Luísa Custódio, Maria João Rodrigues, Giulia Atzori and Katarzyna Negacz
Plants 2024, 13(16), 2322; https://doi.org/10.3390/plants13162322 - 20 Aug 2024
Viewed by 1562
Abstract
Salinization is a major cause of soil degradation that affects several million hectares of agricultural land, threatening food security and the sustainability of agricultural systems worldwide. Nevertheless, despite the negative impact of salinity, salt-affected land also provides several important ecosystem services, from providing [...] Read more.
Salinization is a major cause of soil degradation that affects several million hectares of agricultural land, threatening food security and the sustainability of agricultural systems worldwide. Nevertheless, despite the negative impact of salinity, salt-affected land also provides several important ecosystem services, from providing habitats and nurseries for numerous species to sustainable food production. This opinion paper, written in the framework of the EU COST Action CA22144 SUSTAIN on the sustainable use of salt-affected land, therefore, focuses on the potential of halophytes and saline agriculture to transform and restore key functions of these salt-affected and marginal lands. As the current knowledge on sustainable saline agriculture upscaling is fragmented, we highlight (i) the research gaps in halophyte and salinity research and (ii) the main barriers and potentials of saline agriculture for addressing food security and environmental sustainability in terms of population growth and climate change. Full article
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