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Atriplex

Atriplex is a of approximately 250 of and herbs and shrubs belonging to the family , commonly known as saltbushes or oraches, and renowned for their adaptation to saline and arid environments worldwide. These exhibit diverse growth habits, ranging from low-sprawling, deep-rooted subshrubs to erect s, often featuring leaves that are alternate, simple, and entire to dentate, with many displaying a distinctive scaly or mealy covering formed by collapsed bladder-like hairs. Flowers are inconspicuous and typically arranged in spherical clusters, spikes, or panicles, with staminate flowers bearing 3–5 sepals and stamens, while pistillate flowers lack a and produce fruits enclosed by two enlarged, persistent bracts that aid in dispersal. Seeds are generally erect and flattened, contributing to the genus's resilience in harsh conditions. Distributed primarily in temperate and subtropical regions across all continents except , Atriplex species thrive in alkaline, saline, or disturbed soils, with many acting as halophytes that accumulate salts and even in some cases. Ecologically significant for stabilizing soils and providing in arid ecosystems, certain like A. hortensis are cultivated as edible greens similar to , while others serve as drought-tolerant feed or in revegetation efforts. The 's monoecious or dioecious and morphological variability, including dimorphic fruits in some taxa, underscore its evolutionary adaptations to extreme habitats.

Description

Morphological Features

Atriplex species exhibit a diverse range of growth habits, encompassing or and that can be prostrate, spreading, or erect, with some reaching heights of up to 3 meters, as seen in A. nummularia, a sprawling or erect . These are typically monoecious or dioecious, often featuring bladderlike hairs on their surfaces that collapse upon drying to produce a characteristic silvery, scurfy, or mealy vestiture. Leaves in the genus are generally alternate, though sometimes or subopposite, and , with blades that are entire, serrate, or lobed, measuring 1–10 cm in length and displaying shapes from rhombic and triangular to ovate or lanceolate. They are sessile or petiolate, and while many species, such as A. canescens, bear grayish-green leaves covered in dense trichomes that impart a silvery appearance, others like the cultivated A. hortensis have green, glabrous to sparsely scaly, ovate-lanceolate to cordate blades. Stems are branched, ranging from herbaceous in annuals to woody at the base in perennials, often bearing the same scurfy or farinose coating from collapsed hairs, with rigid, brittle forms in species like A. canescens. Flowers are small and unisexual, arranged in axillary or terminal inflorescences that form spikes, panicles, or clusters; staminate flowers possess a 3–5-parted and 3–5 stamens without bracts, while pistillate flowers typically lack a or have a reduced one and are enclosed by 2 foliaceous bracteoles with two stigmas. Fruits are indehiscent utricles tightly enclosed within persistent, enlarged bracteoles that vary in shape—often compressed, free to fused, and appendaged with wings, tubercles, or spines to facilitate wind dispersal—as exemplified by the four-winged bracts in A. canescens.

Physiological Traits

Most species of Atriplex employ the photosynthetic pathway, characterized by , which features distinct bundle sheath and mesophyll cells that facilitate CO2 concentration around to enhance efficiency under high light and temperature conditions typical of arid environments. This adaptation likely originated in the genus during the , approximately 15 to 5 million years ago, coinciding with increasing aridity and declining atmospheric CO2 levels that favored carbon-concentrating mechanisms. The syndrome provides a in water-limited habitats by reducing and improving water-use efficiency compared to the ancestral C3 pathway. Water and uptake in Atriplex is supported by extensive deep systems, such as the in A. halimus that can extend up to 10 meters, enabling access to subsurface moisture during prolonged droughts and enhancing overall . Some species, like A. halimus, also exhibit and succulence, where enlarged cells store water to maintain turgor and sustain physiological processes under stress. These traits collectively allow efficient foraging in saline or nutrient-poor soils, with facilitating selective uptake to avoid toxicity. Reproductive strategies in Atriplex vary between monoecious and dioecious forms, with many species producing separate male and female flowers on the same or different plants to promote . is primarily anemophilous, relying on dispersal of lightweight , which suits the open, sparse vegetation of arid regions. Seeds often exhibit dormancy mechanisms, such as physical barriers from persistent bracteoles or physiological inhibition, enabling persistence in soil seed banks until favorable moisture conditions break dormancy in unpredictable arid environments. Growth patterns differ markedly across life forms: annual and ephemeral species, such as A. sagittata, display rapid vegetative growth and reproduction during brief wet periods to capitalize on ephemeral resources. In contrast, perennial shrubs like A. confertifolia exhibit slower development, with gradual accumulation over years to establish resilient structures in stable but harsh conditions. This dichotomy optimizes survival in variable arid ecosystems. A distinctive biochemical feature of Atriplex is the synthesis of compatible solutes like glycine betaine and , which accumulate in response to osmotic stress from or to lower cellular and protect proteins and membranes. In species such as A. canescens and A. halimus, betaine contributes significantly to osmotic adjustment in leaves under high , while plays a complementary role in roots, enhancing tolerance without disrupting metabolism. These osmoprotectants enable sustained growth and in hypertonic environments.

Taxonomy

Classification History

The genus Atriplex derives its name from the Latin atriplex, which originates from the ἀτράφαξυς (atráphaxus), a term used for orache-like plants. established the genus in his in 1753, designating A. hortensis ( orache) as the based on its polymorphic leaves and inflorescences. This initial description encompassed a broad range of annual and perennial herbs and shrubs characterized by scurfy vestiture and fruits, reflecting the genus's early recognition as a diverse group adapted to saline environments. Historically, Atriplex was classified within the family Chenopodiaceae from its inception until molecular phylogenetic studies prompted its merger into the expanded sensu lato under the APG III system in 2009. Early taxonomists subdivided the genus into sections such as Atriplex (for monoecious species with samara-like bracteoles) and Obione (for dioecious taxa with obcompressed, non-samaroid bracteoles), a distinction first formalized by Friedrich Adalbert Maximilian von Meyer in 1833 and elaborated by Alfred Moquin-Tandon. These sections highlighted morphological differences in reproductive structures and habit, aiding in the organization of the genus's global diversity across arid and coastal habitats. Key taxonomic revisions in the 19th and 20th centuries addressed the genus's expanding known diversity; Moquin-Tandon's 1840 monograph Chenopodearum Monographica Enumeratio provided a comprehensive of approximately 100 , incorporating new descriptions from explorations in the and . By 1978, A. J. Scott's revisions within Chenopodiaceae recognized around 250 worldwide, emphasizing anatomical and floral traits to refine infrageneric groupings. Recent molecular analyses have led to synonymy reductions, such as the consolidation of certain North American variants, by revealing cryptic relationships obscured by . The genus's high morphological variability—manifest in leaf shape, indumentum, and bracteole form—has historically prompted over-splitting, as seen in A. semibaccata (creeping saltbush), where synonyms like A. flagellaris arose from misinterpretations of prostrate habit and fruit dimorphism before their merger based on consistent traits.

Phylogenetic Relationships

Atriplex belongs to the subfamily within the family (formerly Chenopodiaceae), a placement supported by chloroplast analyses that confirm its monophyletic grouping with related genera. Phylogenetic studies indicate that Atriplex shares its closest relatives with , forming sister clades that diverged approximately 61 million years ago, while , in the sister subfamily , represents a broader alliance within . A comprehensive biogeographic and phylogenetic analysis in 2022 reconstructed the evolutionary history of Atriplex using multi-locus molecular , revealing its origin in continental during the around 30-25 million years ago. The genus diversified into approximately 5-7 major , with subsequent global spread driven by long-distance dispersal events, including transoceanic jumps to , the , and in the Miocene and later periods. Molecular phylogenetic investigations have employed markers such as the nuclear ribosomal (ITS) region, along with chloroplast genes matK and ndhF, to resolve relationships within Atriplex. These studies demonstrate in several traditional sections, such as Obione and Pterochiton, necessitating taxonomic revisions to reflect monophyletic groupings based on genetic evidence. Regarding photosynthetic pathways, C4 evolution occurred multiple times within , with at least 10 independent origins across the subfamily, though a single origin characterizes the core C4 of Atriplex itself during the Middle to . Recent phylogenetic reassessments have been prompted by field discoveries in 2025, including the rediscovery of Atriplex acutiloba in , which challenges its prior extinct status and highlights the need for updated listings based on genetic confirmation of viability. Additionally, the invasive spread of Atriplex semilunaris to new regions in the , documented in 2025, underscores ongoing evolutionary dynamics and potential hybridization risks within Atriplex clades.

Species Diversity

The genus Atriplex comprises approximately 250–300 species of herbs, subshrubs, and shrubs, making it one of the most species-rich genera in the family. Diversity is highest in , with about 69 species (of which around 62 are native) recorded, the majority of which are endemic to the continent. hosts around 62 species, while has approximately 45 species, reflecting centers of diversification in arid and semi-arid regions across these continents. Species of Atriplex exhibit a range of life forms, predominantly annual herbs but including perennial herbs, subshrubs, and shrubs; plants are typically monoecious, though some are dioecious. Notable examples include A. hortensis, an annual herb cultivated as orache for its edible leaves in temperate regions; A. canescens, a shrub known as fourwing saltbush and valued as widespread in North American arid zones; A. nummularia, a shrub called old man saltbush and native to inland ; and A. halimus, an Mediterranean shrub adapted to saline coastal habitats. Endemism is pronounced in Australian hotspots of arid ecosystems, where many are restricted to specific saline or alkaline soils, contributing to regional . Some taxa face threats from alteration, such as A. yeelirrie, a rare tetraploid shrub described in 2015 and confined to two genetically distinct populations in Western Australia's palaeodrainage channels. Certain also exhibit invasive potential outside their native ranges, including A. suberecta (peregrine saltbush), which has naturalized and spread in disturbed sites across parts of and the Mediterranean.

Distribution and Habitat

Global Range

The genus Atriplex has a cosmopolitan native distribution, occurring across temperate and subtropical regions worldwide, from subarctic areas like and to subtropical zones in and , encompassing all continents except . This broad range reflects its origin in continental during the epoch around 30 million years ago, followed by global dispersal primarily through long-distance seed transport, likely aided by birds, which facilitated colonization of distant landmasses such as in the approximately 15–20 million years ago, as well as the , , and . Centers of and are concentrated in several arid and semi-arid hotspots, including Australia's arid interior where extensive radiations produced high during the and ; the in western , featuring numerous shrubby species adapted to saline deserts; the Andean highlands of , with temperate lineages showing significant ; the , a key invasion corridor for multiple dispersals from Asia; and , particularly the Aralo-Caspian and Pontic regions, which served as ancestral hubs for onward migrations. Beyond native ranges, Atriplex species have been widely introduced for , , and ornamental purposes, becoming naturalized in non-native regions. Notable examples include A. nummularia, originally from , which is established in arid parts of ; A. micrantha, native to , naturalized across central and including and ; and various species like A. semibaccata in Pacific islands such as . A recent example of range expansion involves the Australian A. semilunaris, previously recorded only on and in the , which was documented as an on and in 2025, potentially signaling further spread in Macaronesian archipelagos.

Habitat Preferences

Atriplex species predominantly inhabit saline, alkaline, and gypsum-rich soils, demonstrating tolerance to a wide of 6 to 9 and electrical (EC) levels up to 40 dS/m, which enables them to colonize areas with high sodium and accumulation. They also thrive in heavy clay loams, sandy loams, and silty soils, particularly in disturbed or reclaimed sites where supports their root systems. These plants favor arid to semi-arid climates characterized by annual rainfall between 100 and 500 mm, often with bimodal patterns including winter rains and summer thunderstorms that facilitate establishment. They are well-adapted to regions with extreme temperatures, ranging from -20°C to 50°C, allowing persistence in hot deserts and cold steppes. Preferred microhabitats include inland , sinks in the , Mediterranean shrublands, coastal dunes, and salt flats, as well as disturbed areas like overgrazed rangelands and eroded slopes. Some species occur in freshwater wetlands adjacent to saline zones, though most are restricted to hypersaline environments. High in open, unshaded settings further defines their niches, promoting vigorous growth in full sun exposure.

Ecology

Environmental Adaptations

Atriplex species exhibit remarkable halophytism, enabling survival in high-salinity environments through multiple integrated mechanisms. Salt exclusion occurs primarily via specialized glandular structures, including trichomes and epidermal bladder cells that secrete and accumulate excess sodium s on leaf surfaces, thereby preventing toxic buildup in photosynthetic tissues. These bladders, characteristic of the family, sequester NaCl and other s, reducing their transport to metabolically active cells. Additionally, Atriplex compartmentalizes salts within vacuoles, maintaining cytosolic ; species such as A. portulacoides tolerate external NaCl concentrations up to 200 mM without significant toxicity, as ions are isolated via transporters like H+-ATPases. Drought resistance in Atriplex is facilitated by efficient photosynthesis, which concentrates CO2 at the site of , allowing higher carbon fixation with reduced stomatal opening and thus minimizing transpirational water loss. Stomatal regulation, mediated by accumulation, further enhances water use efficiency by closing stomata under water stress, while osmotic adjustment lowers tissue to as low as -4.20 . Complementing these traits, deep systems in like A. halimus extend up to 5 m, accessing in arid soils and sustaining growth during prolonged dry periods. Atriplex also demonstrates adaptations to other abiotic stresses, including . Flavonoids, such as those upregulated in A. canescens under stress, scavenge to protect cellular integrity. In species like A. canescens, tolerance arises from resprouting capability; while not highly fire-resistant, can regenerate from roots if fire intensity is moderate. Recent research underscores Atriplex's , particularly in saline land . A 2025 study on A. canescens found that with (electrical conductivity >16 dS/m) enhances accumulation and amelioration under combined and , provided is managed, offering sustainable strategies for rehabilitating salt-affected lands.

Biotic Interactions

Atriplex are primarily wind-pollinated, a common to the Chenopodiaceae family, facilitating reproduction in arid and saline environments where insect pollinators may be scarce. Seed dispersal often involves a combination of abiotic and agents, with fruits of like Atriplex semibaccata attracting , mammals, reptiles, and , which aid in spreading seeds over short to moderate distances. In some cases, such as are particularly drawn to the nutrient-rich elaiosomes on fruits, promoting and enhancing in disturbed . Herbivory on Atriplex encompasses a range of consumers, from large mammals to small . Species such as Atriplex nummularia serve as important for domesticated like sheep and camels, providing drought-resistant that supports in saline rangelands. In native ecosystems, Atriplex has historically been a dietary staple for herbivores including , contributing to its evolutionary adaptations against browsing pressure. Insect herbivores include larvae of , notably Coleophora atriplicis, which feed on seeds and foliage of coastal Atriplex species like A. littoralis and A. portulacoides, often creating silken cases for protection while consuming plant tissues. Additionally, foliage of Atriplex shrubs hosts diverse arthropods, including spiders and other that utilize the dense leaves for and predation on smaller pests. Symbiotic relationships in Atriplex involve potential associations with arbuscular mycorrhizal fungi (AMF), which enhance nutrient uptake, particularly , in saline or nutrient-poor soils. with AMF like Glomus mosseae has been shown to improve growth of Atriplex nummularia under varying levels, suggesting a beneficial role despite the genus's general non-mycorrhizal reputation. These fungi also influence broader soil microbial communities, promoting shifts in bacterial composition that support plant establishment and in revegetation efforts. Several Atriplex species exhibit invasiveness in introduced ranges, where they compete aggressively with native . For instance, Atriplex prostrata forms dense stands in coastal habitats, displacing local salt-tolerant plants through rapid colonization of disturbed saline areas. In the , the Australian species Atriplex semilunaris has shown rapid invasive spread, with new populations documented in and as of 2025, outcompeting endemic coastal vegetation and altering local biodiversity.

Uses

Agricultural and Forage Applications

Atriplex species, particularly A. nummularia and A. canescens, serve as valuable forage in arid and semi-arid regions due to their high nutritional content, including crude protein levels ranging from 15% to 25% on a dry matter basis and elevated mineral concentrations from ash content often exceeding 20%. These shrubs are widely planted for livestock feed, especially for sheep and goats, where they provide drought- and salt-tolerant browse that sustains grazing in marginal environments with annual rainfall as low as 200-400 mm. Feeding regimes incorporating Atriplex have demonstrated tolerance to moderate to heavy grazing, with regrowth supported by short rotational periods of 4-6 weeks every 6-8 months, enhancing overall rangeland productivity without severe degradation. In production, Atriplex contributes to improved quality; for instance, sheep diets supplemented with saltbush have shown elevated (an ) in intramuscular and subcutaneous fat compared to non-saltbush feeds, potentially enhancing the nutritional profile of and . These plants are particularly suited to arid zones in , the , where they support pastoral systems by providing reliable during dry seasons. Atriplex plays a key role in soil rehabilitation through phytoremediation of saline soils, where species like A. nummularia and A. hortensis accumulate salt ions in their leaves and bladders, reducing over time while maintaining production. Their deep root systems also aid in rangelands, stabilizing degraded desert landscapes in regions such as and the by binding particles and preventing and runoff. Global planting initiatives have established Atriplex stands in saline deserts to restore on marginal lands previously unsuitable for conventional . Cultivation of Atriplex for agricultural purposes involves direct seeding at rates of 4-8 kg/ha for de-winged seeds, achieving establishment densities of 1,000-3,000 plants/ha in rows spaced 4-6 m apart to optimize growth in saline conditions. These shrubs exhibit strong grazing tolerance, allowing integration into rotational systems that promote resilience, and selected varieties, such as improved cultivars of A. nummularia, enhance palatability for better livestock acceptance without compromising salt tolerance. Hybrids between Atriplex species have been developed to further refine traits like forage quality, though field adoption remains focused on pure lines in most arid applications. Economically, Atriplex supports on marginal lands by enabling production where traditional crops fail, thereby bolstering livelihoods in saline-affected areas and contributing to regional . Recent emphasizes its potential in saline , with studies highlighting Atriplex's role in sustainable systems that mitigate impacts and expand for in arid zones.

Culinary and Medicinal Uses

Certain species of Atriplex, particularly A. hortensis (commonly known as garden orache or mountain ), have been utilized in culinary applications as a leafy green vegetable. The tender leaves of A. hortensis serve as a viable substitute for in various dishes, including salads, soups, and steamed preparations, due to their mild flavor and ability to retain crispness when lightly cooked. This usage dates back historically, with the plant employed in European and Asian cuisines for its versatility in fresh and cooked forms. Nutritionally, Atriplex species like A. hortensis offer significant benefits, being rich in vitamins A and C, iron, and , which contribute to their value as a nutrient-dense . The leaves also contain antioxidants such as betalains and , which provide protective effects against . Recent 2025 research highlights Atriplex as a promising climate-smart crop for saline environments, enhancing in arid regions through its nutritional profile and tolerance. In , Atriplex halimus exhibits properties, attributed to its polyphenolic compounds, which have been used to alleviate conditions like intestinal and renal pain. Extracts from A. halimus have shown antidiabetic effects by reducing blood glucose levels and inhibiting enzymes like α-amylase in experimental models. Additionally, traditional remedies involving A. halimus address wounds and injuries, with studies confirming its wound-healing activity through enhanced tissue regeneration in topical applications. The plant's extracts demonstrate activity against various bacterial strains, supporting its ethnobotanical use for infections. For culinary cultivation, A. hortensis is grown as an annual in home gardens and small-scale plots, thriving in well-drained soils with moderate watering. Yields of fresh biomass can reach up to 14-20 tons per hectare under optimal conditions, making it suitable for vegetable production in temperate to subtropical climates.

Other Practical Applications

Atriplex species, particularly A. lentiformis and A. canescens, are valued in landscaping for their drought tolerance and adaptability to arid, saline environments, making them ideal for xeriscaping in regions like the southwestern United States. These shrubs provide aesthetic appeal with their silvery foliage and rounded forms, while reducing water demands in urban and residential designs; for instance, A. lentiformis thrives in low-water landscapes, supporting erosion control and wildlife habitat without supplemental irrigation once established. In industrial applications, Atriplex shows promise as a feedstock for production due to its high yield in marginal lands. Species like A. crassifolia have been pretreated for bioethanol conversion, yielding fermentable sugars through methods such as alkaline and enzymatic processes, with potential for co-production of and biomethane via . Additionally, pigments extracted from A. rubra and A. hortensis var. rubra serve as natural dyes, offering red hues for coloring through solvent-based , as demonstrated in studies on betacyanin . Beyond biofuels and dyes, several Atriplex taxa exhibit potential for in contaminated soils; A. halimus accumulates , lead, , and from , stabilizing pollutants while producing harvestable , and A. hortensis tolerates high levels of , lead, and in polluted environments. For conservation efforts, Atriplex plays a key role in revegetation of saline and mined lands, aiding habitat restoration in arid ecosystems. A. canescens (fourwing saltbush) is widely used to reclaim salt-affected soils, stabilizing surfaces, reducing , and supporting recovery in areas like North American salt deserts, where it forms foundational communities. Similarly, A. nummularia facilitates of sodic and saline sites by improving and , as seen in and projects that promote native resurgence. Historically, utilized Atriplex species, known as saltbush, in cultural practices, including as a source by grinding seeds for and applying leaves as poultices for wounds in arid regions. In modern contexts, saltbush-dominated plains in attract eco-tourism, showcasing resilient ecosystems through guided tours that highlight and in places like national parks.

Safety and Toxicity

Health Risks

Atriplex , commonly known as saltbushes, contain high levels of oxalates in their leaves, primarily in the form of crystals such as sphaeraphides, which can irritate the upon consumption, leading to pain, swelling, and excessive salivation in both humans and . These oxalates bind to calcium, potentially causing , muscle tremors, and kidney stone formation in susceptible individuals or animals if ingested in large quantities. Atriplex can accumulate nitrates under environmental stress such as or high inputs, potentially leading to (characterized by respiratory distress and ) in ruminants like and sheep when plants are the primary source and exceed safe thresholds (e.g., >9000 dry matter). Toxicity varies by species; for instance, Atriplex nummularia (old man saltbush) is generally safe for livestock when fed in moderation as part of a mixed diet but can become toxic if it constitutes the sole forage, due to high oxalate levels (typically 6-8% dry weight, with toxicity risks above ~8%) that induce calcium deficiency and related symptoms. In contrast, Atriplex rosea (red orache) poses risks from selenium accumulation in selenium-rich soils, which can cause chronic poisoning in grazing animals, manifesting as hoof deformities and hair loss when consumed excessively; mitigation includes soil testing and avoiding high-selenium areas. Environmental stresses, such as or , elevate toxin levels in Atriplex plants; for example, saline conditions increase accumulation as a response, heightening risks for grazers. Overgrazed areas exacerbate sheep incidents, as animals are forced to consume higher proportions of Atriplex, leading to acute or intoxication cases reported in arid rangelands. Human incidents involving Atriplex are rare but include warnings against raw consumption of leaves due to oxalate variability, which can contribute to urinary tract stones in predisposed individuals. Recent studies from the highlight seasonal and edaphic factors influencing content, with higher concentrations in spring-stressed plants potentially amplifying these risks for foragers or supplemental feeders.

Mitigation Strategies

To mitigate the risks associated with accumulation in Atriplex used as , dietary guidelines recommend gradually introducing the plants to over four days, incrementally increasing time to allow adaptation and reduce . Mixing Atriplex with low- feeds, such as grasses or , dilutes intake and limits potential for calcium binding and kidney damage, with saltbush comprising no more than 30-50% of the diet to minimize risks. For human consumption, cooking or blanching Atriplex leaves effectively reduces soluble oxalates; for 5-10 minutes can lower levels by 30-87%, while blanching achieves 30-50% reduction, with the cooking water discarded to remove leached compounds. Breeding and selection programs focus on developing low-toxin cultivars of Atriplex, such as hybrids of fourwing saltbush () with species like A. polycarpa or A. gardneri, which exhibit naturally lower concentrations compared to A. halimus (averaging 6-7% ). Selecting varieties with reduced accumulation under saline conditions, through phenotypic screening for lower shoot concentrations, supports safer integration into saline systems, where ongoing monitoring of chemistry ensures toxin levels remain below thresholds. Regulatory frameworks from the USDA and FAO provide guidelines for safe forage planting of Atriplex, emphasizing establishment in early spring via transplants or late fall seeding on saline soils, with deferred rotation grazing to prevent overbrowsing and toxin buildup. For edible species, routine testing for nitrates is required (e.g., using the Nitrate QuikTest or laboratory analysis on dry matter samples from lower plant portions), with levels above 9000 ppm prompting ration limits to half the diet or less to avoid methemoglobinemia. Recent advances in 2025 research highlight processing techniques to minimize risks in farming, including optimizing fertilization with ammonium-dominant sources (e.g., 100:0 NH₄⁺:NO₃⁻ ratios) to significantly reduce levels in Atriplex nummularia, combined with blanching for further post-harvest reduction. These methods, integrated with genetic selection for low- traits, enhance the viability of Atriplex in sustainable saline while addressing nutritional concerns.

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