Anatomía de los órganos vegetativos de dos especies de Atriplex (Chenopodiaceae) de Venezuela / Anatomy of the vegetative organs of two species of Atriplex (Chenopodiaceae) from Venezuela

In Venezuela, Atriplex is represented by A. cristata and A. oestophora, the latter being endemic; they inhabit coastal areas with high temperatures, high solar radiation and sandy soils with high salt content. This work aimed to provide information to facilitate and clarify these species taxonomic delimitation, throughout the study of the anatomy of their vegetative organs; this may also clarify our understanding of their adaptability to soil and climatic conditions prevailing in areas they inhabit. The plant material was collected from at least three individu- als of each species in Punta Taima Taima and Capatárida, Falcon. Segments of roots, located near the neck and towards the apex, apical, middle and basal internodes of stems, were taken; and of leaves, located in the middle portion of plants. This material was fixed in FAA (formaldehyde, acetic acid, 70% ethanol) until processing. Semipermanent and permanent microscope slides were prepared with transverse or longitudinal sections, made using a razor (free-hand) or a rotation microtome, in this latter case, after paraffin embedding; besides, additional plates were mounted with portions of leaf epidermis, obtained by the maceration technique. The sections were stained with aqueous toluidine blue (1%) or safranin-fast-green, and mounted in water-glycerin or in Canada balsam. In order to calculate the vulnerability index, the vessel diameter in the vascular rings of roots, as well as their density, were quantified. Our results revealed structural features in the different organs, that resulted of taxonomic value and allowed the distinction of the species: in the leaf, the presence of aquifer tissue, the number of vascular bundles and their organization in the midrib, and the collenchyma differentiation in this part of the leaf; in the roots, the xylem and phloem arrangement in the growth rings, the nature of conjunctive tissue, and the presence of included phloem in one species. In addition, the species showed typical anatomical features of halophytes and xerophytes, such as: high density of trichomes on leaves and young stems which act as salt secreting glands, abundant sclerenchyma in stems and roots, water storage tissue and Kranz anatomy in leaves, narrow cortical region in young roots, presence of cambial variants in stems and roots, as well as short and narrow xylem vessels. Vulnerability index calculations indicated that both species tend to assure conduction but not the efficiency of the system. Atriplex species have anatomical characters which facilitate their adaptation to the special conditions prevailing in their habitats and that may be used for taxonomic delimitation.

En Venezuela, Atriplex está representado por A. cristata y A. oestophora, siendo esta última endémica, las mismas habitan zonas costeras con altas temperaturas, alta radiación solar y suelos arenosos con alto contenido de sales. Se caracterizaron anatómicamente sus órganos vegetativos con el fin de aportar rasgos para delimitarlas taxonómicamente y precisar caracteres que contribuyan a su adaptabilidad a las condiciones edafoclimáticas imperantes en su hábitat. El material vegetal fue recolectado en tres individuos de cada especie en Punta Taima Taima y Capatárida (Falcón). Se recolectaron segmentos de raíces próximos al cuello y al ápice; entrenudos basales, medios y apicales, y hojas ubicadas en la porción media de las plan- tas. Este material fue fijado en FAA (formaldehido, ácido acético y etanol 70%) hasta su procesamiento. Se prepara- ron láminas semipermanentes y permanentes con secciones transversales y longitudinales hechas a mano alzada o con micrótomo, en este último caso posterior a la inclusión en parafina. Adicionalmente, se realizaron macerados con el fin de obtener las epidermis foliares. Las secciones fueron teñidas con azul de toluidina acuosa (1%) o con safranina- fastgreen, montándolas en agua/glicerina o en bálsamo de Canadá. Se cuantificó el número de vasos y su densidad en los anillos vasculares de las raíces, para calcular el índice de vulnerabilidad. Se encontraron rasgos estructurales de valor taxonómico: la presencia de tejido acuífero en la lámina foliar, el número de haces vasculares y su organización en el nervio medio, así como la diferenciación de colénquima en el mismo; además, el arreglo del xilema/ floema en los anillos de crecimiento, la naturaleza del tejido conjuntivo, así como la presencia de floema incluso, en las raíces. Se detectaron caracteres anatómicos típicos de halófitas y xerófitas, como son: alta densidad de tricomas en hojas y tallos jóvenes, que actúan como glándulas secretoras de sal, abundancia de esclerénquima en tallos y raíces, tejidos reservantes de agua y anatomía Kranz en hojas, región cortical estrecha en raíces jóvenes, presencia de variantes cambiales en tallos y raíces, así como vasos cortos y estrechos en el xilema. El índice de vulnerabilidad indica que ambas especies tienden a asegurar la conducción, pero no la eficiencia del sistema. Las especies de Atriplex que crecen en Venezuela tienen caracteres que permiten su delimitación taxonómica y que facilitan su adaptación a las condiciones particulares de su hábitat.

Comparative studies of C3 and C4 Atriplex hybrids in the genomics era: physiological assessments.

We crossed the C3 species Atriplex prostrata with the C4 species Atriplex rosea to produce F1 and F2 hybrids. All hybrids exhibited C3-like δ(13)C values, and had reduced rates of net CO2 assimilation compared with A. prostrata. The activities of the major C4 cycle enzymes PEP carboxylase, NAD-malic enzyme, and pyruvate-Pi dikinase in the hybrids were at most 36% of the C4 values. These results demonstrate the C4 metabolic cycle was disrupted in the hybrids. Photosynthetic CO2 compensation points (Г) of the hybrids were generally midway between the C3 and C4 values, and in most hybrids were accompanied by low, C3-like activities in one or more of the major C4 cycle enzymes. This supports the possibility that most hybrids use a photorespiratory glycine shuttle to concentrate CO2 into the bundle sheath cells. One hybrid exhibited a C4-like Г of 4 µmol mol(-1), indicating engagement of a C4 metabolic cycle. Consistently, this hybrid had elevated activities of all measured C4 cycle enzymes relative to the C3 parent; however, C3-like carbon isotope ratios indicate the low Г is mainly due to a photorespiratory glycine shuttle. The anatomy of the hybrids resembled that of C3-C4 intermediate species using a glycine shuttle to concentrate CO2 in the bundle sheath, and is further evidence that this physiology is the predominant, default condition of the F2 hybrids. Progeny of these hybrids should further segregate C3 and C4 traits and in doing so assist in the discovery of C4 genes using high-throughput methods of the genomics era.

[Anatomy of the vegetative organs of two species of Atriplex (Chenopodiaceae) from Venezuela]. / Anatomía de los órganos vegetativos de dos especies de Atriplex (Chenopodiaceae) de Venezuela.

In Venezuela, Atriplex is represented by A. cristata and A. oestophora, the latter being endemic; they inhabit coastal areas with high temperatures, high solar radiation and sandy soils with high salt content. This work aimed to provide information to facilitate and clarify these species taxonomic delimitation, throughout the study of the anatomy of their vegetative organs; this may also clarify our understanding of their adaptability to soil and climatic conditions prevailing in areas they inhabit. The plant material was collected from at least three individuals of each species in Punta Taima Taima and Capatárida, Falcon. Segments of roots, located near the neck and towards the apex, apical, middle and basal internodes of stems, were taken; and of leaves, located in the middle portion of plants. This material was fixed in FAA (formaldehyde, acetic acid, 70% ethanol) until processing. Semipermanent and permanent microscope slides were prepared with transverse or longitudinal sections, made using a razor (free-hand) or a rotation microtome, in this latter case, after paraffin embedding; besides, additional plates were mounted with portions of leaf epidermis, obtained by the maceration technique. The sections were stained with aqueous toluidine blue (1%) or safranin-fast-green, and mounted in water-glycerin or in Canada balsam. In order to calculate the vulnerability index, the vessel diameter in the vascular rings of roots, as well as their density, were quantified. Our results revealed structural features in the different organs, that resulted of taxonomic value and allowed the distinction of the species: in the leaf, the presence of aquifer tissue, the number of vascular bundles and their organization in the midrib, and the collenchyma differentiation in this part of the leaf; in the roots, the xylem and phloem arrangement in the growth rings, the nature of conjunctive tissue, and the presence of included phloem in one species. In addition, the species showed typical anatomical features of halophytes and xerophytes, such as: high density of trichomes on leaves and young stems which act as salt secreting glands, abundant sclerenchyma in stems and roots, water storage tissue and Kranz anatomy in leaves, narrow cortical region in young roots, presence of cambial variants in stems and roots, as well as short and narrow xylem vessels. Vulnerability index calculations indicated that both species tend to assure conduction but not the efficiency of the system. Atriplex species have anatomical characters which facilitate their adaptation to the special conditions prevailing in their habitats and that may be used for taxonomic delimitation.

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.