Differential functional traits underlying the contrasting salt tolerance in Lepidium species

Plant and Soil - To explore the mechanisms responsible for salt tolerance in Brassicaceae species, a multifactorial approach was used to clarify the functional traits underlying the differential...     

Does Bienertia cycloptera with the single-cell system of C(4) photosynthesis exhibit a seasonal pattern of delta (13)C values in nature similar to co-existing C (4) Chenopodiaceae having the dual-cell (Kranz) system?

Family Chenopodiaceae is an intriguing lineage, having the largest number of C₄ species among dicots, including a number of anatomical variants of Kranz anatomy and three single-cell C₄ functioning species. In some previous studies, during the culture of Bienertia cycloptera Bunge ex Boiss., carbon isotope values (δ¹³C values) of leaves deviated from C₄ to C₃−C₄ intermediate type, raising questions as to its mode of photosynthesis during growth in natural environments. This species usually co-occurs with several Kranz type C₄ annuals. The development of B. cycloptera morphologically and δ¹³C values derived from plant samples (cotyledons, leaves, bracts, shoots) were analyzed over a complete growing season in a salt flat in north central Iran, along with eight Kranz type C₄ species and one C₃ species. For a number of species, plants were greenhouse-grown from seeds collected from the site, in order to examine leaf anatomy and C₄ biochemical subtype. Among the nine C₄ species, the cotyledons of B. cycloptera, and of the Suaeda spp. have the same respective forms of C₄ anatomy occurring in leaves, while cotyledons of members of tribe Caroxyloneae lack Kranz anatomy, which is reflected in the δ¹³C values found in plants grown in the natural habitat. The nine C₄ species had average seasonal δ¹³C values of −13.9‰ (with a range between species from −11.3 to −15.9‰). The measurements of δ¹³C values over a complete growing season show that B. cycloptera performs C₄ photosynthesis during its life cycle in nature, similar to Kranz type species, with a seasonal average δ¹³C value of −15.2‰. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Mandragora turcomanica (Solanaceae) in Iran: A new distribution record for an endangered species 1

Mandragora turcomanica Mizg. is reported from gardens of Dahaneh village c.7 km north of Kalaleh in Golestan Province (NE Iran). The plants were transplanted from wild populations by the local people many years ago, because of their highly aromatic leaves and fruits. Current information suggests the species is extinct or extremely rare in natural habitats in Iran and protection of extant stands in this village is urgently recommended. It is already known to be endangered in Turkmenistan. The detailed morphological characteristics are discussed and compared with the Mediterranean M. officinarum L., and photographs of the plant are provided.     

Biodiversity and phytogeography of the alpine flora of Iran

Iran is a mountainous country. Zagros and Alborz mountains reach altitudes of more than 4,000 m. Alpine regions are above timber-line, which is not easy to recognize, since aridity is prominent in most regions. The alpine zone in Alborz lies between 3,000 and 4,000 m, the nival zone is above 4,000 m, locally varying by some hundred meters. A first evaluation of vascular flora shows that 682 species belonging to 193 genera and 39 families are known from the alpine zone of Iran. The alpine zone is commonly characterized by many species of hemicryptophytes and thorny cushions. Species numbers decline very strongly with increasing altitude. In this paper biogeographical patterns of the alpine flora of Iran have been discussed and distribution maps of 44 species are illustrated. New data indicate a transitional situation of the Iranian mountains between Anatolia/Caucasus and the Hindu Kush, but with a strong own element with high endemism and remarkable relict species. Ca. 58% of the alpine flora of Iran are endemic and subendemic. The Zagros Mountains harbor high endemism which justify considering this area as a separate floristic province. Based on the evaluation of published data from 682 known alpine species ca. 160 species have been known only by one record, 110 species by 2–3 records and 87 endemic species have been known only based on the type location. These plants need a strong conservation and protection management since the fragile ecosystems are often very restricted, small and very isolated, nonetheless grazing and overgrazing are still common threats.     

Differentiation of cellular and biochemical features of the single-cell C4 syndrome during leaf development in Bienertia cycloptera (Chenopodiaceae)

The terrestrial plant Bienertia cycloptera has been shown to accomplish C(4) photosynthesis within individual chlorenchyma cells by spatially separating the phases of carbon assimilation into distinct peripheral and central compartments. In this study, anatomical, physiological, and biochemical techniques were used to determine how this unique compartmentation develops. Western blots show ribulose-1,5-bisphosphate carboxylase (Rubisco) (chloroplastic) is present in the youngest leaves and increases during development, while levels of C(4) enzymes-pyruvate,Pi dikinase (chloroplastic), phosphoenolpyruvate carboxylase (PEPC) (cytosol), and NAD-malic enzyme (mitochondrial)-increase later in development. Immunolocalization confirmed this for Rubisco and PEPC. The youngest chlorenchyma cells have a central nucleus surrounded by monomorphic granal chloroplasts containing Rubisco. Later stages show progressive development of a central cytoplasmic compartment enriched with chloroplasts and mitochondria and of a peripheral cytoplasm with chloroplasts. A complex reticulum of connections between the compartments also developed and was characterized. δ(13)C isotope analyses show mature leaves have distinct C(4)-type isotope composition, while the composition in younger leaves is "C(4)-like." Based on the results, this form of single-cell C(4) photosynthesis develops from a common pool of organelles through partitioning to separate compartments, and the development of biochemically and ultrastructurally dimorphic chloroplasts.     

Photosynthetic pathways inChenopodiaceae from Africa, Asia and Europe with their ecological, phytogeographical and taxonomical importance

Photosynthetic pathways are reported for 305 species ofChenopodiaceae from Africa, Asia and Europe. Ecological characteristics, phytogeography and life forms of all species are given, and their correlation with relevant CO₂-fixation types are discussed. 205 species (67.2%) exhibit the C₄-pathway and 100 species (32.8%) the C₃-pathway of CO₂-fixation. Most of the C₄ species are of Irano-Turanian origin. The diversity of C₄ species of the Irano-Turanian phytochoria, with very harsh winters, are interpreted by the active period of Chenopods in summer. There is a close relationship between some special morphotypes and respective photosynthetic type. Halophytes and xerophytes with articulated stems and stem succulents ofAnabasis-type are exclusively C₄. Leaf succulent halophytes and xerohalophytes are predominantly C₄. Hygrohalophytes with leaf or stem succulence are often C₃. Probably many C₄ Salsoleae have been evolved in the Afroasiatic arid zone after the climatic changes of the Miocene. Among them there is a high proportion of annuals that have a younger origin. There is some evidence that the present Chenopods of the subfamily Salsoloideae may have their ancient stock in NW Africa. We found close taxonomic relationships between photosynthetic pathways and infrageneric classification in genera with both types of C₃ and C₄, likeAtriplex andSuaeda. Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer, Wien, on the occasion of his 70th birthday     

Salt tolerance mechanisms in three Irano-Turanian Brassicaceae halophytes relatives of <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis>

Salt tolerance mechanisms were studied in three Irano-Turanian halophytic species from the Brassicaceae ??(Lepidium latifolium, L. perfoliatum and Schrenkiella parvula) and compared with the glycophyte Arabidopsis thaliana. According to seed germination under salt stress, L. perfoliatum was the most tolerant species, while L. latifolium and S. parvula were rather susceptible. Contrastingly, based on biomass production L. perfoliatum was more salt sensitive than the other two species. In S. parvula biomass was increased up to 2.8-fold by 100 mM NaCl; no significant growth reduction was observed even when exposed to 400 mM NaCl. Stable activities of antioxidative defense enzymes, nil or negligible accumulation of superoxide anion and hydrogen peroxide, as well as stable membrane integrity in the three halophytes revealed that no oxidative stress occurred in these tolerant species under salt stress. Proline levels increased in response to salt treatment. However, it contributed only by 0.3?2.0% to the total osmolyte concentration in the three halophytes (at 400 mM NaCl) and even less (0.04%) in the glycophyte, A. thaliana (at 100 mM NaCl). Soluble sugars in all three halophytes and free amino acids pool in S. parvula decreased under salt treatment in contrast to the glycophyte, A. thaliana. The contribution of organic osmolytes to the total osmolyte pool increased by salt treatment in the roots, while decreased in halophyte and glycophyte, A. thaliana leaves. Interestingly, this reduction was compensated by a higher relative contribution of K in the leaves of the halophytes, but of Na in A. thaliana. Taken together, biomass data and biochemical indicators show that S. parvula is more salt tolerant than the two Lepidium species. Our data indicate that L. latifolium, as a perennial halophyte with a large biomass, is highly suitable for both restoration of saline habitats and saline agriculture.     

Ecology, Biogeography and Pollen Morphology of Bienertia cycloptera Bunge ex Boiss. (Chenopodiaceae), an Enigmatic C4 Plant without Kranz Anatomy

Abstract: Bienertia cycloptera is a most interesting terrestrial plant which applies C₄ photosynthesis without Kranz cells, but with dimorphic chloroplasts in one cell. In order to study this special quality and provide more information about the natural habitat of the species and probable evolutionary trends, we have examined the plant communities associated with B. cycloptera, the climatic conditions in which this species occurs, soil characteristics and pollen morphology in comparison with several other possible relatives. B. cycloptera is distributed around Central Iranian deserts and subdeserts with northern and southern radiation in the Persian Gulf countries and Central Asia, from 24° to 49° N latitude and from 43° to 67° E longitude. It grows in areas with mean daily temperatures between 20 and 30 °C during the hot months of the year, and annual precipitation of 400 to 76 mm (but mostly of 200 to 100 mm). B. cycloptera grows on high salty and temporarily moist clay soils with a salinity ranging from 21.7 to 89.6 ds/m, both in Na⁺ and SO₄^2‐ rich salty soils. In most plots, the percentage of C₄ species is over 50 % and in some cases 100 %. Climacoptera turcomanica is the most frequently associated species with B. cycloptera in most habitats. In order to look for further comparative data to find the affinity and evolutionary trends in Suaedeae and Salicornioideae, the pollen morphology of B. cycloptera, Borszczowia aralocaspica, Halopeplis pygmaea, Salicornia persica and six species of Suaeda, each representing one section, are compared by light and SEM microscopy. The pollen grains of B. cycloptera have the lowest pore number, with a mean of 42 pores per grain. The pore numbers of B. cycloptera are very close to Halopeplis pygmaea, a member of the tribe Halopeplideae of subfamily Salicornioideae. Among the studied Suaedeae, the pollen of B. cycloptera shows similarities with Suaeda physophora with regard to pore number and pore diameter. The affinity of B. cycloptera is a matter of discrepancy. Three different possible origins and relationships in Suaeda sect. Immersa, sect. Physophora, sect. Schanginia and tribe Halopeplidae are discussed. We conclude that the evolution, diversification and adaptation of Central and SW Asian halophytes is the result of intricate environmental and geological history of the area and needs intensive multidisciplinary research.     

Species Diversity and Life-Form Patterns in Steppe Vegetation along a 3000 m Altitudinal Gradient in the Alborz Mountains, Iran

Biodiversity pattern and life-form spectra were studied along a 3,000 m altitudinal gradient from a semi-desert area to the alpine peak of Tochal Mountain. The gradient is located on the southern slopes of Central Alborz with a Mediterranean continental climate. DCA ordination was applied to 1,069 relevés and 7 quantitative variables to discover the relation of diversity and altitude. A biodiversity pattern was obtained by relating values for species richness and Shannon-Wiener’s index to 100-m altitudinal sections. Altitude was determined as the major ecological gradient. Both diversity indices are negatively correlated with altitude and show a decreasing trend beyond a peak in species richness at 1,800–1,900 m a.s.l. towards a very low diversity in the high alpine zone. The biodiversity peak does not match with the potential tree line in the area (2,500–3,000 m a.s.l.). The high diversity in foothills can be related to habitat heterogeneity, longer suitable climatic conditions, and diverse disturbance factors, while unfavorable conditions at high-altitude alpine and low-altitude desert areas reduce the number of species at both extremes. Life-form patterns clearly change along altitudinal gradient. Annuals with decreasing trend, and hemicryptophytes and chamaephytes with increasing trend along the altitudinal gradient are notable patterns of life form in the area. Temperature, soil moisture and nutrients are the main factors that explain the ecological influence of altitude on species diversity and life-form patterns in the semi-arid steppe vegetation of the area.     

Vegetation Patterns of the Irano-Turanian Steppe along a 3,000 m Altitudinal Gradient in the Alborz Mountains of Northern Iran

The Irano-Turanian floristic region is a major center of endemism in the Holarctic of Eurasia. The Alborz Mountains of northern Iran are a complex and heterogeneous environmental system with rich water resources and great habitat diversity. We have investigated steppe plant communities along an altitudinal gradient ranging from approximately 1,000 m a.s.l. in the semi-desert steppes near Tehran to a height of 3,966 m a.s.l. at the summit of Mount Tochal. Our two-way indicator species analysis of 1,069 vegetation samples resulted in classification of five major vegetation zones: (1) a semi-desert Artemisia steppe near Tehran, (2) a Stipa grassland in the alluvial undulating hills north and west of Tehran, (3) a submontane and steppe zone, (4) a subalpine cushion formation zone and (5) an alpine meadow and subnival zone of Mount Tochal. Annuals and ephemerals in the semi-desert vegetation decline as altitude increases and almost disappear in the alpine zone. Past human impacts of ancient Persian civilization and a traditional pastoral economy have affected the physiognomy of plant communities; thorny dwarf shrubs now dominate the treeless vegetation of the region. Lower competition for space, different phenology and the presence of edaphic and hydrological layers associated with anthropogenic impacts are major reasons for entanglement of different plant communities in the arid- and semi-arid steppe. The phytogeography of the region changes from omni-Irano-Turanian and Saharo-Sindian transgressive species at lower altitudes to a more limited range of western Irano-Turanian species and local endemics at higher altitudes.