Current Understanding of Role of Vesicular Transport in Salt Secretion by Salt Glands in Recretohalophytes.

Soil salinization is a serious and growing problem around the world. Some plants, recognized as the recretohalophytes, can normally grow on saline-alkali soil without adverse effects by secreting excessive salt out of the body. The elucidation of the salt secretion process is of great significance for understanding the salt tolerance mechanism adopted by the recretohalophytes. Between the 1950s and the 1970s, three hypotheses, including the osmotic potential hypothesis, the transfer system similar to liquid flow in animals, and vesicle-mediated exocytosis, were proposed to explain the salt secretion process of plant salt glands. More recently, increasing evidence has indicated that vesicular transport plays vital roles in salt secretion of recretohalophytes. Here, we summarize recent findings, especially regarding the molecular evidence on the functional roles of vesicular trafficking in the salt secretion process of plant salt glands. A model of salt secretion in salt gland is also proposed.

Morphological and physiological responses of two willow species from different habitats to salt stress.

Plant salt tolerance is a complex mechanism, and different plant species have different strategies for surviving salt stress. In the present study, we analyzed and compared the morphological and physiological responses of two willow species (Salix linearistipularis and Salix matsudana) from different habitats to salt stress. S. linearistipularis exhibited higher seed germination rates and seedling root Na+ efflux than S. matsudana under salt stress. After salt treatment, S. linearistipularis leaves exhibited less Na+ accumulation, loss of water and chlorophyll, reduction in photosynthetic capacity, and damage to leaf cell structure than leaves of S. matsudana. Scanning electron microscopy combined with gas chromatography mass spectrometry showed that S. linearistipularis leaves had higher cuticular wax loads than S. matsudana leaves. Overall, our results showed that S. linearistipularis had higher salt tolerance than S. matsudana, which was associated with different morphological and physiological responses to salt stress. Furthermore, our study suggested that S. linearistipularis could be a promising tree species for saline-alkali land greening and improvement.

Secretory structures in plants: Lessons from the Plumbaginaceae on their origin, evolution and roles in stress tolerance.

The Plumbaginaceae (non-core Caryophyllales) is a family well known for species adapted to a wide range of arid and saline habitats. Of its salt-tolerant species, at least 45 are in the genus Limonium; two in each of Aegialitis, Limoniastrum and Myriolimon, and one each in Psylliostachys, Armeria, Ceratostigma, Goniolimon and Plumbago. All the halophytic members of the family have salt glands and salt glands are also common in the closely related Tamaricaceae and Frankeniaceae. The halophytic species of the three families can secrete a range of ions (Na+ , K+ , Ca2+ , Mg2+ , Cl- , HCO3- , SO42- ) and other elements (As, Cd, Cr, Cu, Fe, Mn, Ni, Pb and Zn). Salt glands are, however, absent in salt-tolerant members of the sister family Polygonaceae. We describe the structure of the salt glands in the three families and consider whether glands might have arisen as a means to avoid the toxicity of Na+ and/or Cl- or to regulate Ca2+ concentrations with the leaves. We conclude that the establishment of lineages with salt glands took place after the split between the Polygonaceae and its sister group the Plumbaginaceae.

Studies on genome size estimation, chromosome number, gametophyte development and plant morphology of salt-tolerant halophyte Suaeda salsa.

BACKGROUND: Soil salinization and alkalization are among the major agricultural threats that affect crop productivity worldwide, which are increasing day by day with an alarming rate. In recent years, several halophytes have been investigated for their utilization in soil remediation and to decipher the mechanism of salt-tolerance in these high salt tolerant genetic repositories. Suaeda salsa is an annual halophytic herb in the family Amaranthaceae, displaying high salt and alkali-resistance and having nutritive value. However, the fundamental biological characteristics of this valuable plant remain to be elucidated until today. RESULTS: In this study, we observed the morphology and development of Suaeda salsa, including seed morphology, seed germination, plant morphology, and flower development. Using microscopy, we observed the male and female gametophyte developments of Suaeda salsa. Also, chromosome behaviour during the meiosis of male gametophyte was studied. Eventually, the genome size of Suaeda salsa was estimated through flow cytometry using Arabidopsis as reference. CONCLUSIONS: Our findings suggest that the male and female gametophyte developments of Suaeda salsa are similar to those of the model plant Arabidopsis, and the diploid Suaeda salsa contains nine pairs of chromosomes. The findings also indicate that the haploid genome of Suaeda salsa is approximately 437.5 MB. The observations and results discussed in this study will provide an insight into future research on Suaeda salsa.

Microhair on the adaxial leaf surface of salt secreting halophytic Oryza coarctata Roxb. show distinct morphotypes: Isolation for molecular and functional analysis.

Halophytic Oryza coarctata is a good model system to examine mechanisms of salinity tolerance in rice. O. coarctata leaves show the presence of microhairs in adaxial leaf surface furrows that secrete salt under salinity. However, detailed molecular and physiological studies of O. coarctata microhairs are limited due to their relative inaccessibility. This work presents a detailed characterization of O. coarctata leaf features. O. coarctata has two types of microhairs on the adaxial leaf surface: longer microhairs (three morphotypes) lining epidermal furrow walls and shorter microhairs (reported first time) arising from bulliform cells. Microhair morphotypes include (i) finger-like, tubular structures, (ii) tubular hairs with bilobed and flattened heads and (iii) bi-or trifurcated hairs. The unicellular nature of microhairs was confirmed by propidium iodide (PI) staining. An efficient method for the isolation and enrichment of O. coarctata microhairs is presented (yield averaging ˜2 × 105/g leaf tissue). The robustness of the microhair isolation procedure was confirmed by subsequent viability staining (PI), total RNA isolation and RT-PCR amplification of O. coarctata trichome-specific WUSCHEL-related homeobox 3B (OcWox3B) and transporter gene-specific cDNA sequences. The present microhair isolation work from O. coarctata paves the way for examining genes involved in ion secretion in this halophytic wild rice model.

Palyno-morphological investigations of halophytic taxa of Amaranthaceae through SEM from Salt range of Northern Punjab, Pakistan.

The pollen morphology of 11 salt tolerant plant species of family Amaranthaceae from the salt range of Northern Punjab, Pakistan has been studied. The palyno-morphological characters were examined using light and scanning electron microscope. The examined all salt tolerant species have a slight difference in size but have similarity in shape, pore ornamentation, and polarity. The observed morphological characters of pollen grains were pollen symmetry, size, shape, pore ornamentation, pore size, number of pores, exine thickness, polar and equatorial diameter and, P/E ratio. Apolar type of pollens has been observed in all species. Shape of pollens was spheroidal. Exine sculpturing of pollen grains was scabrate (six spp), microechinate (four spp), and microechinate-scabrate (one spp). Different pori numbers were observed in different species. The pantoporate aperturate and sunken pore ornamentation have been reported in all species. A pollen taxonomic key was developed using examined morphological characters for the accurate identification of halophytic taxa. The high fertility and low sterility of pollens confirmed that the selected halophytes are well-established in the salt region. The findings highlight the taxonomic significance of pollen morphology in correct identification and differentiation of salt tolerant plant species.

Leaf anatomical traits determine the 18 O enrichment of leaf water in coastal halophytes.

Anatomical adaptations to high-salinity environments in mangrove leaves may be recorded in leaf water isotopes. Recent studies observed lower 18 O enrichment (ΔL ) of leaf water with respect to source water in three mangrove species relative to adjacent freshwater trees, but the factors that govern this phenomenon remain unclear. To resolve this issue, we investigated leaf traits and ΔL in 15 species of true mangrove plants, 14 species of adjacent freshwater trees, and 4 species of semi-mangrove plants at five study sites along south-eastern coast of China. Our results confirm that ΔL was generally 3-4‰ lower for mangrove species than for adjacent freshwater or semi-mangrove species. We hypothesized that higher leaf water content (LWC) and lower leaf stomatal density (LS) both played important roles in reducing ΔL in mangroves relative to nearby freshwater plants. Both differences acted to elongate effective leaf mixing length (L) in mangroves by about 200% and lower stomatal conductance by about 30%. Péclet models based on both LWC and LS could accurately predict ΔL . Our findings highlight the potential species-specific anatomical determinants of ΔL (or L), which has important implications for the interpretation of environmental information from metabolites produced by leaf water isotopes in palaeoclimate research.

Polyamine and tyramine involvement in NaCl-induced improvement of Cd resistance in the halophyte Inula chrithmoides L.

The aim of the present work was to analyze the impact of salinity on the plant response to Cd toxicity in the Mediterranean halophyte species Inula crithmoides. For this purpose, cuttings were cultivated hydroponically during 21d in the presence of 0, 25 or 50µM CdCl2 combined or not with 0, 100, 200 and 400mM NaCl. The obtained data demonstrated that, in the absence of Cd, NaCl strongly increased plant growth (the maximal dry weight being observed at 100mM) and enhanced the Na+/K+ ratio in the shoot. Cd alone strongly affected plant growth in this halophyte. However, in Cd-treated plants, NaCl protected Inula crithmoides from Cd toxicity and contributed to reduce Cd absorption and translocation. Small aliphatic polyamine (putrescine, spermidine, spermine) increased in response to both NaCl and CdCl2, the highest concentration in plants being observed when both agents are present in the medium. The recorded increase preferentially concerned the polyamine bound fraction, which might be related to their involvement in the protection of endogenous cellular structures. The aromatic monoamine tyramine also strongly increased in response to Cd toxicity and its putative role is discussed in relation to conjugation processes. Salinity and Cd increased ammonium/nitrate ratio in leaves and roots and the involvement of stress-induced modification of N nutrition on polyamine oversynthesis is also discussed.

Cambios anatómicos en raíces e hipocótilos de plántulas de Prosopis ruscifolia (Fabaceae) sometidas a estrés salino / Anatomical changes in roots and hypocotyls of Prosopis ruscifolia (Fabaceae) seedlings exposed to saline stress

ResumenProsopis ruscifolia es una especie arbórea pionera en áreas inundadas o salinas. El objetivo de este trabajo fue determinar cambios anatómicos en raíces e hipocótilos de plántulas de P. ruscifolia sometidas a estrés salino, bajo condiciones controladas. Las semillas se recolectaron en bosques nativos de la Región Chaqueña Occidental de Argentina. Las semillas se sembraron sobre toallas de papel humedecidas con soluciones salinas de 100, 200 y 300 mM de NaCl y un control humedecido con agua destilada. Se sembraron cuatro repeticiones de 50 semillas cada una, correspondientes a cada tratamiento, se ubicaron en cajas plásticas herméticas dentro de cámara de siembra a 27 ºC y con fotoperíodo de 12 horas. Doce días después de la siembra, se extrajeron plántulas para estudios anatómicos. Se estudiaron 35 plántulas correspondientes a cada tratamiento. Se midieron en raíces e hipocótilos las siguientes variables anatómicas: diámetro de la raíz principal e hipocótilo (µm), espesor de la corteza (µm), número de estratos celulares en la corteza, diámetro del cilindro central (µm), diámetro de la médula (µm), número de estratos celulares en el periciclo y diámetro tangencial de los vasos (µm). Se realizó ANOVA con diámetro de la raíz o hipocótilo como variable dependiente y espesor de la corteza, número de estratos celulares en la corteza, diámetro del cilindro central, diámetro de la médula, número de estratos celulares en el periciclo, diámetro tangencial de los vasos y concentración salina como variables independientes. El diámetro de la raíz disminuyó significativamente con el aumento de la concentración salina (P < 0.0001). El espesor de la corteza redujo su espesor a 100 mM (P < 0.0001) e incrementó el número de estratos celulares que la componen (P < 0.0002). El diámetro del cilindro central se redujo a la concentración salina de 100 mM (P < 0.0001) y el diámetro de la médula y el número de estratos celulares del periciclo (P < 0.0003) disminuyó progresivamente hasta 300 mM. El diámetro tangencial de los vasos (P < 0.0001) se redujo recién a 300 mM de NaCl. Estos cambios anatómicos podrían estar relacionados con la alteración de la expansión y división celular causada por la salinidad y comprometer la formación de raíces laterales y el almacenamiento de reservas. Los hipocótilos no mostraron cambios anatómicos significativos en respuesta al incremento en la salinidad, con excepción de la variación en la posición de estomas y un incremento en el espesor de la hipodermis. Estos cambios parecen indicar el estrés hídrico impuesto por el bajo potencial osmótico causado por las sales. Las plántulas de P. ruscifolia experimentaron cambios anatómicos en respuesta a las concentraciones salinas analizadas, en rasgos vinculados al almacenamiento de reservas, a la absorción y la conducción de agua y la formación de raíces laterales.

Abstract:Prosopis ruscifolia is a pioneer tree species in flooding or saline areas. The aim of this work was to assess anatomical changes in roots and hypocotyls of P. ruscifolia seedlings induced to saline stress under controlled conditions. Seeds, collected in natural forests of Western Chaco region in Argentina, were sown on paper towels moisturized with saline solutions of 100, 200 and 300 mM of NaCl, and a control group with distilled water. Four repetitions of 50 seeds per treatment were sown, located in hermetic polystyrene boxes, and included in a seeding chamber, at 27 ºC and 12 hours photoperiod. Were studied 35 seedlings from each saline concentration; these seedlings were processed 12 days after sown to obtain microscopic samples. The anatomical variables measured in roots and hypocotyls were the following: main root diameter (µm), bark thickness (µm), number of cell strata in bark, central cylinder diameter (µm), pith diameter (µm), number of cell strata in the pericycle and the tangential diameter of vessels (µm). ANOVA analysis were performed with hypocotyl and root diameters as the dependent variable, and bark thickness (µm), number of cell strata in the bark, the central cylinder diameter (µm), the pith diameter (µm), number of cell strata in the pericycle, the tangential diameter of vessels and the saline concentration as independent variables. Results showed that the root diameter decreased with increasing saline concentrations (P < 0.0001). The bark thickness decreased at 100 mM (P < 0.0001) and the number of cell strata of bark increased to 300 mM (P < 0.0002). The central cylinder diameter decreased at 100 mM saline concentration (P < 0.0001) and the number of cell strata of the pericycle and the pith diameter reduced progressively until 300 mM. The tangential diameter of vessels decreased at 300 mM. These anatomical changes suggested alterations in the expansion and cell division caused by the salinity, and could limit lateral roots formation and reserves storage. Hypocotyls did not show significant anatomical changes in response to increasing salinity, with exception of stomata position and an increase of the hypodermis thickness. These changes indicated that the water stress imposed by low osmotic potential is caused by increasing saline concentration. The seedlings of P. ruscifolia experienced anatomical changes in response to tested saline concentrations in traits related to reserve storage, the absorption and conduction of water, and lateral roots formation. Rev. Biol. Trop. 64 (3): 1007-1017. Epub 2016 September 01.

[Anatomical changes in roots and hypocotyls of Prosopis ruscifolia (Fabaceae) seedlings exposed to saline stress].

Prosopis ruscifolia is a pioneer tree species in flooding or saline areas. The aim of this work was to assess anatomical changes in roots and hypocotyls of P. ruscifolia seedlings induced to saline stress under controlled conditions. Seeds, collected in natural forests of Western Chaco region in Argentina, were sown on paper towels moisturized with saline solutions of 100, 200 and 300 mM of NaCl, and a control group with distilled water. Four repetitions of 50 seeds per treatment were sown, located in hermetic polystyrene boxes, and included in a seeding chamber, at 27 ºC and 12 hours photoperiod. Were studied 35 seedlings from each saline concentration; these seedlings were processed 12 days after sown to obtain microscopic samples. The anatomical variables measured in roots and hypocotyls were the following: main root diameter (µm), bark thickness (µm), number of cell strata in bark, central cylinder diameter (µm), pith diameter (µm), number of cell strata in the pericycle and the tangential diameter of vessels (µm). ANOVA analysis were performed with hypocotyl and root diameters as the dependent variable, and bark thickness (µm), number of cell strata in the bark, the central cylinder diameter (µm), the pith diameter (µm), number of cell strata in the pericycle, the tangential diameter of vessels and the saline concentration as independent variables. Results showed that the root diameter decreased with increasing saline concentrations (P < 0.0001). The bark thickness decreased at 100 mM (P < 0.0001) and the number of cell strata of bark increased to 300 mM (P < 0.0002). The central cylinder diameter decreased at 100 mM saline concentration (P < 0.0001) and the number of cell strata of the pericycle and the pith diameter reduced progressively until 300 mM. The tangential diameter of vessels decreased at 300 mM. These anatomical changes suggested alterations in the expansion and cell division caused by the salinity, and could limit lateral roots formation and reserves storage. Hypocotyls did not show significant anatomical changes in response to increasing salinity, with exception of stomata position and an increase of the hypodermis thickness. These changes indicated that the water stress imposed by low osmotic potential is caused by increasing saline concentration. The seedlings of P. ruscifolia experienced anatomical changes in response to tested saline concentrations in traits related to reserve storage, the absorption and conduction of water, and lateral roots formation.

Growth responses of the mangrove Avicennia marina to salinity: development and function of shoot hydraulic systems require saline conditions.

BACKGROUND AND AIMS: Halophytic eudicots are characterized by enhanced growth under saline conditions. This study combines physiological and anatomical analyses to identify processes underlying growth responses of the mangrove Avicennia marina to salinities ranging from fresh- to seawater conditions. METHODS: Following pre-exhaustion of cotyledonary reserves under optimal conditions (i.e. 50% seawater), seedlings of A. marina were grown hydroponically in dilutions of seawater amended with nutrients. Whole-plant growth characteristics were analysed in relation to dry mass accumulation and its allocation to different plant parts. Gas exchange characteristics and stable carbon isotopic composition of leaves were measured to evaluate water use in relation to carbon gain. Stem and leaf hydraulic anatomy were measured in relation to plant water use and growth. KEY RESULTS: Avicennia marina seedlings failed to grow in 0-5% seawater, whereas maximal growth occurred in 50-75% seawater. Relative growth rates were affected by changes in leaf area ratio (LAR) and net assimilation rate (NAR) along the salinity gradient, with NAR generally being more important. Gas exchange characteristics followed the same trends as plant growth, with assimilation rates and stomatal conductance being greatest in leaves grown in 50-75% seawater. However, water use efficiency was maintained nearly constant across all salinities, consistent with carbon isotopic signatures. Anatomical studies revealed variation in rates of development and composition of hydraulic tissues that were consistent with salinity-dependent patterns in water use and growth, including a structural explanation for low stomatal conductance and growth under low salinity. CONCLUSIONS: The results identified stem and leaf transport systems as central to understanding the integrated growth responses to variation in salinity from fresh- to seawater conditions. Avicennia marina was revealed as an obligate halophyte, requiring saline conditions for development of the transport systems needed to sustain water use and carbon gain.

Taxonomic complexity in the halophyte Limonium vulgare and related taxa (Plumbaginaceae): insights from analysis of morphological, reproductive and karyological data.

BACKGROUND AND AIMS: Limonium is a well-known example of a group of plants that is taxonomically complex due to certain biological characteristics that hamper species' delineation. The closely related polyploid species Limonium vulgare Mill., L. humile Mill. and L. narbonense Mill. are defined species and can be used for studying patterns of morphological and reproductive variation. The first two taxa are usually found in Atlantic Europe and the third in the Mediterranean region, but a number of intermediate morphological forms may be present alongside typical examples of these species. This study attempts to elucidate morphological, floral and karyological diversity representative of these taxa in the Iberian Peninsula. METHODS: The extent of morphological differentiation was tested through comparison of 197 specimens from both Portugal and Spain using 17 descriptive morphological characters and 19 diagnostic morphometric characters. Analyses of floral morphisms (heterostyly and pollen-stigma dimorphism) and karyological determinations were also conducted. KEY RESULTS AND CONCLUSIONS: Discriminant analysis using morphometric variables reliably assigned individuals in natural populations to their respective groups. In addition, the results provide the first direct evidence that L. narbonense and a new species, LIMONIUM MARITIMUM: Caperta, Cortinhas, Paes, Guara, Espírito-Santo and Erben, SP NOV: , related to L. vulgare are present on Portuguese coasts. Most of these species are found together in mixed populations, especially L. vulgare and L. narbonense. It is hypothesized that taxonomic biodiversity found in sites where distinct species co-occur facilitates the evolutionary processes of hybridization, introgression and apomixis. This study therefore contributes to the elucidation of the taxonomic diversity in L. vulgare-related species and may also help in implementing future conservation programmes to maintain the evolutionary processes generating biodiversity.

Salt bladders: do they matter?

Soil salinity is claiming about three hectares of arable land from conventional crop farming every minute. At the same time, the challenge of feeding 9.3 billion people by 2050 is forcing agricultural production into marginal areas, and providing sufficient food for this growing population cannot be achieved without a major breakthrough in crop breeding for salinity tolerance. In this Opinion article, we argue that the current trend of targeting Na(+) exclusion mechanisms in breeding programmes for salinity tolerance in crops needs revising. We propose that progress in this area will be achieved by learning from halophytes, naturally salt-loving plants capable of surviving in harsh saline environments, by targeting the mechanisms conferring Na(+) sequestration in external storage organs.

Seagrass proliferation precedes mortality during hypo-salinity events: a stress-induced morphometric response.

Halophytes, such as seagrasses, predominantly form habitats in coastal and estuarine areas. These habitats can be seasonally exposed to hypo-salinity events during watershed runoff exposing them to dramatic salinity shifts and osmotic shock. The manifestation of this osmotic shock on seagrass morphology and phenology was tested in three Indo-Pacific seagrass species, Halophila ovalis, Halodule uninervis and Zostera muelleri, to hypo-salinity ranging from 3 to 36 PSU at 3 PSU increments for 10 weeks. All three species had broad salinity tolerance but demonstrated a moderate hypo-salinity stress response--analogous to a stress induced morphometric response (SIMR). Shoot proliferation occurred at salinities <30 PSU, with the largest increases, up to 400% increase in shoot density, occurring at the sub-lethal salinities <15 PSU, with the specific salinity associated with peak shoot density being variable among species. Resources were not diverted away from leaf growth or shoot development to support the new shoot production. However, at sub-lethal salinities where shoots proliferated, flowering was severely reduced for H. ovalis, the only species to flower during this experiment, demonstrating a diversion of resources away from sexual reproduction to support the investment in new shoots. This SIMR response preceded mortality, which occurred at 3 PSU for H. ovalis and 6 PSU for H. uninervis, while complete mortality was not reached for Z. muelleri. This is the first study to identify a SIMR in seagrasses, being detectable due to the fine resolution of salinity treatments tested. The detection of SIMR demonstrates the need for caution in interpreting in-situ changes in shoot density as shoot proliferation could be interpreted as a healthy or positive plant response to environmental conditions, when in fact it could signal pre-mortality stress.

Sodium instead of potassium and chloride is an important macronutrient to improve leaf succulence and shoot development for halophyte Sesuvium portulacastrum.

Soil salinity is contributed largely by NaCl but some halophytes such as Sesuvium portulacastrum have evolved to adapt salinity environment and demonstrate optimal development under moderate salinity. To elucidate the detail mechanisms of the great salt tolerance and determine the respective contributions of Na(+), K(+) and Cl(-) on the development of S. portulacastrum, morphological and physiological analysis were performed using plants supplied with 200 mM of different ions including cations (Na(+), K(+), Li(+)) and anions (Cl(-), NO(3)(-), Ac(-)) respectively. The results revealed that the salt-treated plants accumulated large amounts of sodium in both leaf and stem. There was a greater shoot growth in presence of external Na(+) compared to K(+) and Cl(-). Na(+) was found more effective than K(+) and Cl(-) in cell expansion, leaf succulence, and shoot development. Flame emission and X-Ray microanalysis revealed the relative Na(+) content was much higher than K(+) and Cl(-) in both leaf and stem of well developed S. portulacastrum, leading to a higher Na(+)/K(+) ratio. The effects of different ions on the development of S. portulacastrum were listed as the following: Na(+) > NO(3)(-) > CK > Cl(-) > K(+) > Ac(-) > Li(+). These results demonstrated NaCl toxicity is attributable largely to the effect of Cl(-) but rarely to Na(+), and thus sodium is concluded as a more important macronutrient than potassium and chloride for improving leaf succulence and shoot development of halophyte S. portulacastrum.

Germination of dimorphic seeds of the desert annual halophyte Suaeda aralocaspica (Chenopodiaceae), a C4 plant without Kranz anatomy.

BACKGROUND AND AIMS: Suaeda aralocaspica is a C4 summer annual halophyte without Kranz anatomy that is restricted to the deserts of central Asia. It produces two distinct types of seeds that differ in colour, shape and size. The primary aims of the present study were to compare the dormancy and germination characteristics of dimorphic seeds of S. aralocaspica and to develop a conceptual model of their dynamics. METHODS: Temperatures simulating those in the natural habitat of S. aralocaspica were used to test for primary dormancy and germination behaviour of fresh brown and black seeds. The effects of cold stratification, gibberellic acid, seed coat scarification, seed coat removal and dry storage on dormancy breaking were tested in black seeds. Germination percentage and recovery responses of brown seeds, non-treated black seeds and 8-week cold-stratified black seeds to salt stress were tested. KEY RESULTS: Brown seeds were non-dormant, whereas black seeds had non-deep Type 2 physiological dormancy (PD). Germination percentage and rate of germination of brown seeds and of variously pretreated black seeds were significantly higher than those of non-pretreated black seeds. Exposure of seeds to various salinities had significant effects on germination, germination recovery and induction into secondary dormancy. A conceptual model is presented that ties these results together and puts them into an ecological context. CONCLUSIONS: The two seed morphs of S. aralocaspica exhibit distinct differences in dormancy and germination characteristics. Suaeda aralocaspica is the first cold desert halophyte for which non-deep Type 2 PD has been documented.