Maternal salinity influences anatomical parameters, pectin content, biochemical and genetic modifications of two Salicornia europaea populations under salt stress.

Salicornia europaea is among the most salt-tolerant of plants, and is widely distributed in non-tropical regions. Here, we investigated whether maternal habitats can influence different responses in physiology and anatomy depending on environmental conditions. We studied the influence of maternal habitat on S. europaea cell anatomy, pectin content, biochemical and enzymatic modifications under six different salinity treatments of a natural-high-saline habitat (~ 1000 mM) (Ciechocinek [Cie]) and an anthropogenic-lower-saline habitat (~ 550 mM) (Inowroclaw [Inw]). The Inw population showed the highest cell area and roundness of stem water storing cells at high salinity and had the maximum proline, carotenoid, protein, catalase activity within salt treatments, and a maximum high and low methyl esterified homogalacturonan content. The Cie population had the highest hydrogen peroxide and peroxidase activity along with the salinity gradient. Gene expression analysis of SeSOS1 and SeNHX1 evidenced the differences between the studied populations and suggested the important role of Na+ sequestration into the vacuoles. Our results suggest that the higher salt tolerance of Inw may be derived from a less stressed maternal salinity that provides a better adaptive plasticity of S. europaea. Thus, the influence of the maternal environment may provide physiological and anatomical modifications of local populations.

Insights into genomic structure and evolutionary processes of coastal Suaeda species in East Asia using cpDNA, nDNA, and genome-wide SNPs.

Species in the genus Suaeda have few diagnostic characters and substantial morphological plasticity. Hence, regional floras do not provide clear taxonomic information for Suaeda spp. in East Asia. In order to assess the taxonomy of four species in the genus Suaeda (S. australis, S. maritima, S. japonica, and S. heteroptera), cpDNA (rpl32-trnL and trnH-psbA), nDNA (ITS), and MIG-seq analyses were carried out. Genome-wide SNP results indicated three lineages: (1) S. australis in Korea and S. maritima in Japan, (2) S. maritima in Korea and S. heteroptera in China, and (3) S. japionica. In phylogenetic trees and genotype analyses, cpDNA and nDNA results showed discrepancies, while S. japonica and S. maritima in Korea, and S. heteroptera in China shared the same haplotype and ribotype. We suggest that the shared haplotype may be due to chloroplast capture. Based on our results, we assume that S. japonica was formed by homoploid hybrid speciation between the two lineages.

Phenotypic plasticity in diaspore production of a amphi-basicarpic cold desert annual that produces polymorphic diaspores.

Phenotypic plasticity has been studied in diaspore-dimorphic species, but no such study has been done on a diaspore-polymorphic species. Our aim was to determine the effects of abiotic and biotic factors on phenotypic plasticity of the diaspore-polymorphic cold desert annual Ceratocarpus arenarius. Plants produced from dispersal units near the soil surface (a, basicarps) and at the middle (c) and upper (f) parts of the plant canopy were subjected to different levels of soil moisture, nutrient supply and intramorph and intermorph densities. Different levels of these biotic and abiotic factors resulted in significant variation in total plant mass, diaspore mass, mass allocation to stem and reproductive organs and total number and proportion of morphs a, c and f on an individual. The effect of stress on number and mass of a dispersal unit morph varied by treatment, with dispersal unit f having the highest CV and dispersal unit a the lowest. The success of this diaspore polymorphic species in its rainfall-unpredictable environment likely is enhanced by plasticity in production of the different types of diaspores.

Morphological description and DNA barcoding study of sand rice (Agriophyllum squarrosum, Chenopodiaceae) collected in Kazakhstan.

BACKGROUND: Sand rice (Agriophyllum squarrosum (L.) Moq.) is an annual shrub-like plant adapted to the mobile sand dunes in desert and semi-desert regions of Asia. It has a balanced nutrient composition with relatively high concentration of lipids and proteins, which results in its nutrition being similar to legumes. Sand rice's proteins contain the full range of essential amino acids. However, calories content is more similar to wheat. These features together with desert stress resistance make sand rice a potential food crop resilient to ongoing climate change. It is also an important fodder crop (on young stages of growth) for cattle in arid regions of Kazakhstan. In our work, sand rice samples were collected from two distant regions of Kazakhstan as a part of the nation-wide project to determine genetic variation of the native flora. RESULTS: Samples were collected in western and southeastern parts of Kazakhstan separated by distances of up to 1300 km. Sequences of the nuclear ribosomal DNA ITS1-5.8S-ITS2 region and the chloroplast matK gene confirmed the identity of species defined by morphological traits. Comparison with GenBank sequences revealed polymorphic sequence positions among Kazakh populations and GenBank references, and suggested a distinction among local populations of sand rice. The phylogenetic analysis of nucleotide sequences showed a clear partition of A. squarrosum (L.) Moq. from Agriophyllum minus Fisch. & C.A. Mey, which grows in the same sand dunes environment. CONCLUSIONS: DNA barcoding analyses of ITS and matK sequences showed a segregation of A. squarrosum from A. minus into separate clades in Maximum-Likelhood dendrograms. ITS analysis can be successfully used to characterize A. squarrosum populations growing quite distant from each other. The data obtained in this work provide the basis for further investigations on A. squarrosum population structure and may play a role in the screening of sand rice plants growing in desert and semi-desert environments of Central Asia and China.

The unique structural and biochemical development of single cell C4 photosynthesis along longitudinal leaf gradients in Bienertia sinuspersici and Suaeda aralocaspica (Chenopodiaceae).

Temporal and spatial patterns of photosynthetic enzyme expression and structural maturation of chlorenchyma cells along longitudinal developmental gradients were characterized in young leaves of two single cell C4 species, Bienertia sinuspersici and Suaeda aralocaspica Both species partition photosynthetic functions between distinct intracellular domains. In the C4-C domain, C4 acids are formed in the C4 cycle during capture of atmospheric CO2 by phosphoenolpyruvate carboxylase. In the C4-D domain, CO2 released in the C4 cycle via mitochondrial NAD-malic enzyme is refixed by Rubisco. Despite striking differences in origin and intracellular positioning of domains, these species show strong convergence in C4 developmental patterns. Both progress through a gradual developmental transition towards full C4 photosynthesis, with an associated increase in levels of photosynthetic enzymes. Analysis of longitudinal sections showed undeveloped domains at the leaf base, with Rubisco rbcL mRNA and protein contained within all chloroplasts. The two domains were first distinguishable in chlorenchyma cells at the leaf mid-regions, but still contained structurally similar chloroplasts with equivalent amounts of rbcL mRNA and protein; while mitochondria had become confined to just one domain (proto-C4-D). The C4 state was fully formed towards the leaf tips, Rubisco transcripts and protein were compartmentalized specifically to structurally distinct chloroplasts in the C4-D domains indicating selective regulation of Rubisco expression may occur by control of transcription or stability of rbcL mRNA. Determination of CO2 compensation points showed young leaves were not functionally C4, consistent with cytological observations of the developmental progression from C3 default to intermediate to C4 photosynthesis.

When do different C4 leaf anatomies indicate independent C4 origins? Parallel evolution of C4 leaf types in Camphorosmeae (Chenopodiaceae).

Broad-scale phylogenetic studies give first insights in numbers, relationships, and ages of C4 lineages. They are, however, generally limited to a model that treats the evolution of the complex C4 syndrome in different lineages as a directly comparable process. Here, we use a resolved and well-sampled phylogenetic tree of Camphorosmeae, based on three chloroplast and one nuclear marker and on leaf anatomical traits to infer a more detailed picture of C4 leaf-type evolution in this lineage. Our ancestral character state reconstructions allowed two scenarios: (i) Sedobassia is a derived C3/C4 intermediate, implying two independent gains of C4 in Bassia and Camphorosma; or (ii) Sedobassia is a plesiomorphic C3/C4 intermediate, representing a syndrome ancestral to the Bassia/Camphorosma/Sedobassia lineage. In Bassia, a kochioid leaf type (Bassia muricata and/or Bassia prostrata type) is ancestral. At least three independent losses of water-storage tissue occurred, resulting in parallel shifts towards an atriplicoid leaf type. These changes in leaf anatomy are adaptations to different survival strategies in steppic or semi-desert habitats with seasonal rainfall. In contrast, Camphorosma shows a fixed C4 anatomy differing from Bassia types in its continuous Kranz layer, which indeed points to an independent origin of the full C4 syndrome in Camphorosma, either from an independent C3 or from a common C3/C4 intermediate ancestor, perhaps similar to its C3/C4 intermediate sister genus Sedobassia. The enlarged bundle sheath cells of Sedobassia might represent an important early step in C4 evolution in Camphorosmeae.

Fruit and seed anatomy of Chenopodium and related genera (Chenopodioideae, Chenopodiaceae/Amaranthaceae): implications for evolution and taxonomy.

A comparative carpological study of 96 species of all clades formerly considered as the tribe Chenopodieae has been conducted for the first time. The results show important differences in the anatomical structure of the pericarp and seed coat between representatives of terminal clades including Chenopodium s.str.+Chenopodiastrum and the recently recognized genera Blitum, Oxybasis and Dysphania. Within Chenopodium the most significant changes in fruit and seed structure are found in members of C. sect. Skottsbergia. The genera Rhagodia and Einadia differ insignificantly from Chenopodium. The evolution of heterospermy in Chenopodium is discussed. Almost all representatives of the tribe Dysphanieae are clearly separated from other Chenopodioideae on the basis of a diverse set of characteristics, including the small dimensions of the fruits (especially in Australian taxa), their subglobose shape (excl. Teloxys and Suckleya), and peculiarities of the pericarp indumentum. The set of fruit and seed characters evolved within the subfamily Chenopodioideae is described. A recent phylogenetic hypothesis is employed to examine the evolution of three (out of a total of 21) characters, namely seed color, testa-cell protoplast characteristics and embryo orientation.

Nitrogen cycle of a typical Suaeda salsa marsh ecosystem in the Yellow River estuary.

The nitrogen (N) biological cycle of the Suaeda salsa marsh ecosystem in the Yellow River estuary was studied during 2008 to 2009. Results showed that soil N had significant seasonal fluctuations and vertical distribution. The N/P ratio (15.73 +/- 1.77) of S. salsa was less than 16, indicating that plant growth was limited by both N and P. The N absorption coefficient of S. salsa was very low (0.007), while the N utilization and cycle coefficients were high (0.824 and 0.331, respectively). The N turnover among compartments of S. salsa marsh showed that N uptake from aboveground parts and roots were 2.539 and 0.622 g/m2, respectively. The N translocation from aboveground parts to roots and from roots to soil were 2.042 and 0.076 g/m2, respectively. The N translocation from aboveground living bodies to litter was 0.497 g/m2, the annual N return from litter to soil was far less than 0.368 g/m2, and the net N mineralization in topsoil during the growing season was 0.033 g/m2. N was an important limiting factor in S. salsa marsh, and the ecosystem was classified as unstable and vulnerable. S. salsa was seemingly well adapted to the low-nutrient status and vulnerable habitat, and the nutrient enrichment due to N import from the Yellow River estuary would be a potential threat to the S. salsa marsh. Excessive nutrient loading might favor invasive species and induce severe long-term degradation of the ecosystem if human intervention measures were not taken. The N quantitative relationships determined in our study might provide a scientific basis for the establishment of effective measures.

The need to re-investigate the nature of homoplastic characters: an ontogenetic case study of the 'bracteoles' in Atripliceae (Chenopodiaceae).

BACKGROUND AND AIMS: Within Chenopodioideae, Atripliceae have been distinguished by two bracteoles enveloping the female flowers/fruits, whereas in other tribes flowers are described as ebracteolate with persistent perianth. Molecular phylogenetic hypotheses suggest 'bracteoles' to be homoplastic. The origin of the bracteoles was explained by successive inflorescence reductions. Flower reduction was used to explain sex determination. Therefore, floral ontogeny was studied to evaluate the nature of the bracteoles and sex determination in Atripliceae. METHODS: Inflorescences of species of Atriplex, Chenopodium, Dysphania and Spinacia oleracea were investigated using light microscopy and scanning electron microscopy. KEY RESULTS: The main axis of the inflorescence is indeterminate with elementary dichasia as lateral units. Flowers develop centripetally, with first the formation of a perianth primordium either from a ring primordium or from five individual tepal primordia fusing post-genitally. Subsequently, five stamen primordia originate, followed by the formation of an annular ovary primordium surrounding a central single ovule. Flowers are either initially hermaphroditic remaining bisexual and/or becoming functionally unisexual at later stages, or initially unisexual. In the studied species of Atriplex, female flowers are strictly female, except in A. hortensis. In Spinacia, female and male flowers are unisexual at all developmental stages. Female flowers of Atriplex and Spinacia are protected by two accrescent fused tepal lobes, whereas the other perianth members are absent. CONCLUSIONS: In Atriplex and Spinacia modified structures around female flowers are not bracteoles, but two opposite accrescent tepal lobes, parts of a perianth persistent on the fruit. Flowers can achieve sexuality through many different combinations; they are initially hermaphroditic, subsequently developing into bisexual or functionally unisexual flowers, with the exception of Spinacia and strictly female flowers in Atriplex, which are unisexual from the earliest developmental stages. There may be a relationship between the formation of an annular perianth primordium and flexibility in floral sex determination.

Soil determinants for distribution of Halocnemum strobilaceum Bieb. (Chenopodiaceae) around Lake Tuz, Turkey.

In this study we aimed to reveal the ranges and effectiveness of soil parameters on the distribution of H. strobilaceum. Halocnemum strobilaceum Bieb. is a widespread species in saline habitats and the distribution pattern of this halophytic species around Lake Tuz in Central Anatolia was examined according to the soil characteristics. pH, electrical conductivity, soil humidity, salt percentage, soluble Ca2+, Mg2+, Na+, K+, CO3(2-), HCO3(-), Cl(-) and SO4(2-) values, total cations, cation exchange capacity, sodium adsorption ratio, exchangeable Na+, K+, Ca2+ and Mg2+ values were the examined soil properties. The most effective soil parameters for flowering period were found as Na, SO4, total cations, SAR and EC and for seed bearing period as EC, Mg, total cations, Cl, Na, SO4 and salt (%) content of the soil.

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.

Morphological study of Salicornieae (Chenopodiaceae) native to Iran.

The tribe Salicornieae Dum. belonging to the subfamily Salicornioideae Kostel (Chenopodiaceae Vent.) includes halophyte plants. These 5 genera and 6 species are distributed in different habitats of Iran. Members of this tribe have reduced vegetative parts, scale like leaves and articulated stems. In this study quantitative and qualitative morphological characters for 46 accessions of Salicornieae were evaluated. Vegetative characters are not sufficient to distinguish these taxa. Our results indicated that vegetative form, globular buds, plant color, stem base disarticulation and presence of node at the base of inflorescence are diagnostic character in this tribe. Besides floral arrangement in inflorescences, bracts shape, color and shape of seeds are important key features in members of Salicornieae in Iran. Seed coat ornamentations and its hairs and color are found to be helpful in distinguishing these taxa. Quantitative characters show variation too, but as they were not significant, they can not help to resolve the taxonomic problems of this tribe in Iran. Based on studied morphological characters, an identification key for members of this tribe in Iran is provided.

Reflectance properties and physiological responses of Salicornia virginica to heavy metal and petroleum contamination.

Wetland ecosystems of California are located in highly populated areas and subject to high levels of contamination. Monitoring of wetlands to assess degrees of pollution damage requires periodic retrieval of information over large areas, which can be effectively accomplished by rapidly evolving remote sensing technologies. The biophysical principles of remote sensing of vegetation under stress need to be understood in order to correctly interpret the information obtained at the scale of canopies. To determine the potential to remotely characterize and monitor pollution, plants of Salicornia virginica, a major component of wetland communities in California, were treated with two metals and two crude oil types to study their sensitivity to pollutants and how this impacted their reflectance characteristics. Several growth and physiological parameters, as well as shoot reflectance were measured and correlated with symptoms and contamination levels. Significant differences between treatments were found in at least some of the measured parameters in all pollutants. Reflectance was sensitive to early stress levels only for cadmium and the lightweight petroleum. Pollutants that differ in their way of action also had different plant reflectance signatures. The high degree of correlation between reflectance features and stress indicators highlights the potential of using remote sensing to assess the type and degree of pollution damage.

Morphology, anatomy and histochemistry of Salicornioideae (Chenopodiaceae) fruits and seeds.

BACKGROUND AND AIMS: The subfamily Salicornioideae (Chenopodiaceae) are a taxonomically difficult group largely due to the lack of diagnostic characters available to delineate tribal- and generic-level boundaries; a consequence of their reduced floral and vegetative features. This study examined the variation in fruits and seeds across both tribes of the Salicornioideae to assess if characters support traditional taxonomic sections. METHODS: Light microscopy, environmental scanning electron microscopy and anatomical ultra-thin sectioning were employed to examine variation in fruits and seeds. Sixty-eight representatives across 14 of the 15 genera currently recognized within the tribes Halopeplideae and Salicornieae were examined to determine whether characters support current taxonomic groups. KEY RESULTS: Characters such as seed coat structure, embryo shape, seed orientation, the forms of seed storage proteins and carbohydrates show variation within the Salicornioideae and may be phylogenetically useful. The campylotropous ovule typical of the Chenopodiaceae generally results in a curved embryo; however, many Halosarcia and Sclerostegia species have straight embryos and in Salicornia and Sarcocornia the large peripheral embryo appears bent rather than curved. Seed coat ornamentation of Microcnemum and Arthrocnemum is distinct from other Salicornioideae as the elongated epidermal cells of the exotesta have convex walls. Histochemical stains of anatomical sections of cotyledon cells showed protein bodies were variable in shape, and starch grains were present in some species, namely Salicornia bigelovii, S. europaea and Allenrolfea occidentalis. CONCLUSIONS: While fruits and seeds were found to be variable within the subfamily, no synapomorphic characters support the tribe Halopeplideae as these genera have crustaceous seed coats, curved embryos and abundant perisperm; features characteristic of many of the tribe Salicornieae. The endemic Australian genera are closely related and few seed and fruit characters are diagnostic at the generic level. Nineteen characters identified as being potentially informative will be included in future phylogenetic analyses of the subfamily.

Physico-chemical and technological studies on sugar beet roots.

The morphological and chemical characteristics of ten varieties of sugar beet roots were evaluated during the season of 1996-1997. The best morphological characteristics were found in Top, Ito and Pamela varieties. The highest expected technological yield of sugar was found in Pamela and Top. The chemical constituents of different varieties of sugar-beet roots showed significant differences. The suitability of sugar beet juice for preparing beet syrup as a new product was also evaluated and compared with sugar cane syrup. Purified beet-syrups concentrated under vacuum or under atmospheric pressure showed more stability during storage than sugar cane syrup. The effect of storage conditions (at cold storage and at room temperature) and packaging materials (glass and polyethylene high-density bottles) on properties of beet-syrup and sugar cane syrup were also studied.