Morphology and anatomy of inflorescence and inflorescence axis in Paepalanthus sect. Diphyomene Ruhland (Eriocaulaceae,Poales) and its taxonomic implications

Paepalanthus sect. Diphyomene has inflorescences arranged in umbels. The underlying bauplan seems however to be more complex and composed of several distinct subunits. Despite appearing superficially very similar, the morphology and anatomy of the inflorescences can supply useful information for the understanding of the phylogeny and taxonomy of the group. Inflorescences of Paepalanthus erectifolius, Paepalanthus flaccidus, Paepalanthus giganteus, and Paepalanthus polycladus were analyzed in regard to branching pattern and anatomy. In P. erectifolius, P. giganteus and P. polycladus the structure is a tribotryum, with terminal dibotryum, and with pherophylls bearing lateral dibotrya. In P. flaccidus, the inflorescence is a pleiobotryum, with terminal subunit, and without pherophylls. Secondary inflorescences may occur in all species without regular pattern. Especially when grown in sites without a pronounced seasonality, the distinction between enrichment zone (part of the same inflorescence) and new inflorescences may be obscured. The main anatomical features supplying diagnostic and phylogenetic information are as follows: (a) in the elongated axis, the thickness of the epidermal cell walls and the cortex size; (b) in the bracts, the quantity of parenchyma cells (c) in the scapes, the shape and the presence of a pith tissue. Therefore, P. sect. Diphyomene can be divided in two groups; group A is represented by P. erectifolius, P. giganteus and P. polycladus, and group B is represented by P. flaccidus. The differentiation is based in both, inflorescence structure and anatomy. Group A presents a life cycle and anatomical features similar to species of Actinocephalus. Molecular trees also point that these two groups are closely related. However, inflorescence morphology and blooming sequence are different. Species of group B present an inflorescence structure and anatomical features shared with many genera and species in Eriocaulaceae. The available molecular and morphology based phylogenies still do not allow a precise allocation of the group in the bulk of basal species of Paepalanthus collocated in P. sect. Variabiles. The characters described and used here supply however important information towards this goal.     

Anatomy of Brazilian Eriocaulaceae: correlation with taxonomy and habitat using multivariate analyses

In this study we present a survey on vegetative anatomy in species of Actinocephalus, Blastocaulon, Eriocaulon, Leiothrix, Paepalanthus, Philodice, Syngonanthus, and Tonina (Eriocaulaceae). Multivariate analyses were used to correlate anatomical characters to taxa and the habitats where the species occur. Root and stem anatomical characters seem to be more affected by environmental factors where these species occur, and seem of little value for delimiting major taxonomic groups within the family. Other characters in the leaves, such as epidermis with thickened wall cells, compartmented substomatal chambers, mesophyll with hypodermis, compact chlorenchyma, collenchymatous bundle sheath extensions, and numerous vascular bundles, were shown to be important for defining species clusters in Leiothrix, Syngonanthus, and Paepalanthus subg. Platycaulon. Similarly, loosely aggregated chlorenchyma caused Blastocaulon, Eriocaulon, Philodice, Syngonanthus sect. Carpocephalus, S. sect. Syngonanthus, and Tonina, genera from humid environments, to cluster. Scape characters appear to be more informative in discriminating groups. This situation probably reflects lower selection pressures determining anatomical characters of this organ.     

Distribution of Photosynthetic Enzymes between Mesophyll, Specialized Parenchyma and Bundle Sheath Cells of Arundinella hirta

Arundinella hirta L. is a C₄ plant having an unusual C₄ leaf anatomy. Besides mesophyll and bundle sheath cells, A. hirta leaves have specialized parenchyma cells which look morphologically like bundle sheath cells but which lack vascular connections and are located between veins, running parallel to them. Activities of phosphoenolpyruvate and ribulose-1,5-bisphosphate carboxylases and phosphoenolpyruvate carboxykinase, NADP-and NAD-malic enzymes were determined for whole leaf extracts and isolated mesophyll protoplasts, specialized parenchyma cells, and bundle sheath cells. The data indicate that A. hirta is a NADP-malic enzyme type C₄ species. In addition, specialized parenchyma cells and bundle sheath cells are enzymatically alike. Compartmentation of enzymes followed the C₄ pattern with phosphoenolpyruvate carboxylase being restricted to mesophyll cells while ribulose-1,5-bisphosphate carboxylase and decarboxylating enzymes were restricted to bundle sheath and specialized parenchyma cells.     

A possible optical role of the bundle sheath extensions of the heterobaric leaves of Vitis vinifera and Quercus coccifera

Heterobaric leaves are characterized by transparent regions in their lamina, due to the occurrence of bundle sheath extensions. Fused silica fibre‐optic microprobes were used to monitor light gradients and part of the spectral regime along the bundle sheath extensions, as well as along the mesophyll in the heterobaric leaves of two representative plants, one mesomorphic (Vitis vinifera L.) and one xeromorphic (Quercus coccifera L.). It was found that the attenuation of collimated visible light by the bundle sheath extensions of both plants was weaker than the attenuation by the photosynthetic parenchyma layers. However, only a small portion of the amount of light that strikes the leaf surface is transmitted through these structures. The adaxial epidermis covering the bundle sheath extensions, as well as the mesophyll, afforded similar effective protection against UV radiation in both tissues. The relative amount of the forward‐scattered visible light inside the bundle sheath extensions approached that detected by the microprobe at the adaxial illuminated leaf surface. Moreover, light transmitted through the bundle sheath extensions was enriched mainly in the blue and red regions, compared to light transmitted through the photosynthetic tissue. The time course of photosynthetic starch formation in the leaves of V. vinifera detected by iodine staining showed that the accumulation of starch during the first minutes of illumination was high within photosynthetic parenchyma cells adjacent to the bundle sheath extensions. The data showed that bundle sheath extensions act as transparent ‘windows’ which enrich the neighbouring mesophyll areas with high levels of photosynthetically active radiation (400–700 nm). The phenomenon was more pronounced in the thick and compact sclerophyllous leaves of Q. coccifera by virtue of the greater abundance of bundle sheath extensions as compared to that in V. vinifera. The enhancement of the light micro‐environment within the deep internal layers of the mesophyll may affect the photosynthetic performance of such leaves, giving adaptive advantages.     

Morphology, anatomy and vasculature of leaves in Pinus (Pinaceae) and its evolutionary meaning

The morphology, anatomy and vasculature of Pinus leaves was studied. The results indicate that the Pinus leaf is always supplied with only one single vascular bundle, which is surrounded by a prominent bundle sheath. In several taxa, especially of subgenus Pinus, the vascular bundle may however be subdivided by longitudinal parenchymatic bands in the middle part of the leaf. As a result, the single bundle gets the appearance of two individual bundles, which are surrounded by a common bundle sheath. The general rule that a bundle sheath does sheath only one single bundle, as in other gymno- and angiospermous seed plants, applies therefore also for Pinus. The morphological and anatomical similarities between cladodes of Sciadopitys (Sciadopityaceae) and Pinus leaves are based on a completely different bauplan. The idea of “hidden cladodes” in some Pinaceae is therefore obsolete.     

COMPARATIVE LEAF ANATOMY OF SOLIDAGO AND RELATED ASTERACEAE

Leaf anatomy of 63 taxa is investigated to elucidate generic relationships among Brachychaeta, Brintonia, Chrysoma, Euthamia, Gundlachia, Oligoneuron, Oreochrysum, Petradoria, and Solidago. All these genera have been included at one time or another within Solidago. Aster ptarmicoides is also studied because it hybridizes with some species of Solidago (sens. str.). Qualitative and quantitative differences in mesophyll, storage parenchyma, secretory apparatus, bundle sheath extensions, and midvein structure allow rather precise grouping of the taxa. Brachychaeta, Brintonia, Oligoneuron, Oreochrysum, and Aster ptarmicoides should be considered as constituents of Solidago. They all have bundle sheath extensions and little or no water storage parenchyma. In Solidago secretory cavities, when present, are shaped and positioned differently from those in Euthamia. The absence of bundle sheath extensions and various combinations of other anatomical features suggest that Chrysoma, Euthamia, Gundlachia, and Petradoria are generically distinct from one another and from Solidago.     

六种芦荟叶的解剖结构及其与芦荟素含量的相关性

应用半薄切片法、高效液相色谱法(HPLC)和荧光显微镜研究了6种芦荟叶的结构、芦荟素的含量和储藏芦荟素的组织.结果表明,6种芦荟叶均由表皮、光合组织、储水组织和维管束组成,都表现出明显的旱生植物肉质叶的结构特征,表皮由一层扁平的细胞组成,其外壁加厚,并覆盖着厚的角质膜,气孔器凹陷,储水组织发达.6 种芦荟叶的结构存在显著差异.木立芦荟(Aloe arborescens Mill.)和易变芦荟(A. mutabilis Pillans)的光合组织细胞呈长柱状,类似栅栏薄壁组织.中华芦荟(A. vera L. var. chinensis Berg.)、库拉索芦荟(A. vera L.)、皂叶芦荟(A. saponaria Hawer)和绿芦荟(A. greenii Bak.)则为等直径薄壁细胞.木立芦荟、中华芦荟、易变芦荟和库拉索芦荟的维管束中有大型薄壁细胞,皂叶芦荟和绿芦荟的维管束中无大型薄壁细胞.木立芦荟、易变芦荟和库拉索芦荟在光合组织和储水组织之间有一层不含叶绿体的小型薄壁细胞,包围着储水薄壁组织,称之为储水组织鞘.中华芦荟、皂叶芦荟和绿芦荟则没有储水组织鞘.HPLC测量结果表明,木立芦荟叶芦荟素含量最高,库拉索芦荟和易变芦荟次之,中华芦荟、皂叶芦荟和绿芦荟含量最低.荧光显微镜观察结果表明,在紫外光和蓝光下,黄色和黄绿色小球体仅存在于维管束的大型薄壁细胞、维管束鞘和储水组织鞘中,而光合组织和储水组织中没有黄色和黄绿色小球体.因此,维管束中大型薄壁细胞、维管束鞘和储水组织鞘是芦荟素等蒽醌类物质的储藏场所.综上所述,芦荟素含量与维管束的大型薄壁细胞、维管束鞘和储水组织鞘的情况密切相关.     

六种芦荟叶的解剖结构及其与芦荟素含量的相关性

应用半薄切片法、高效液相色谱法(HPLC)和荧光显微镜研究了6种芦荟叶的结构、芦荟素的含量和储藏芦荟素的组织。结果表明,6种芦荟叶均由表皮、光合组织、储水组织和维管束组成,都表现出明显的旱生植物肉质叶的结构特征,表皮由一层扁平的细胞组成,其外壁加厚,并覆盖着厚的角质膜,气孔器凹陷,储水组织发达。6种芦荟叶的结构存在显著差异。木立芦荟(Aloe arborescens Mill．)和易变芦荟(A．mutabilis Pillans)的光合组织细胞呈长柱状,类似栅栏薄壁组织。中华芦荟(A．vera L.var．chinensis Berg．)、库拉索芦荟(A．vera L．)、皂叶芦荟(A．saponaria Hawer)和绿芦荟(A．greenii Bak．)则为等直径薄壁细胞。木立芦荟、中华芦荟、易变芦荟和库拉索芦荟的维管束中有大型薄壁细胞,皂叶芦荟和绿芦荟的维管束中无大型薄壁细胞。木立芦荟、易变芦荟和库拉索芦荟在光合组织和储水组织之间有一层不含叶绿体的小型薄壁细胞,包围着储水薄壁组织,称之为储水组织鞘。中华芦荟、皂叶芦荟和绿芦荟则没有储水组织鞘。HPLC测量结果表明,木立芦荟叶芦荟素含量最高,库拉索芦荟和易变芦荟次之,中华芦荟、皂叶芦荟和绿芦荟含量最低。荧光显微镜观察结果表明,在紫外光和蓝光下,黄色和黄绿色小球体仅存在于维管束的大型薄壁细胞、维管束鞘和储水组织鞘中,而光合组织和储水组织中没有黄色和黄绿色小球体。因此,维管束中大型薄壁细胞、维管束鞘和储水组织鞘是芦荟素等蒽醌类物质的储藏场所。综上所述,芦荟素含量与维管束的大型薄壁细胞、维管束鞘和储水组织鞘的情况密切相关。     

Calcium,Calcium Oxalate Crystals,and Leaf Differentiation in the Common Bean (Phaseolus vulgaris L.)*

Young plants of Phaseolus vulgaris were grown in nutrient solutions at different levels of calcium concentration. When the calcium concentration was low more palisade parenchyma and less extended bundle sheath was formed at the adaxial side of minor veins of the leaves as compared to leaves of plants grown with higher calcium supply. The number of calcium oxalate crystals in the bundle sheath extensions was positively correlated to the amount of calcium fed to the plants. The ion induces additional cell divisions in the bundle sheath extensions. A high supply of calcium leads to the formation of a second type of crystal in the bundle sheath.     

Leaf anatomical studies of Chamaecrista subsect. Baseophyllum (Leguminosae, Caesalpinioideae): new evidence for the up-ranking of the varieties to the species level

This study aims to determine whether morphoanatomical characters of the leaves of the species of Chamaecrista sect. Absus subsection Baseophyllum could be used to support an up-ranking of C. cytisoides varieties to the species level as previously defined by molecular studies. The significance of anatomical adaptive strategies to arid environments is also discussed. Standard light microscopy techniques and histochemical tests were used for both morphoanatomical and histochemical characterization of the leaves. All the species studied share a single-layered epidermis, actinodromous–camptodromous–brochidodromous type of venation, vascular bundles surrounded by layers of fibers bounded externally by a sheath of cells containing solitary crystals, enlarged tracheids at the vein endings, vascular petiole/rachis tissue with a parenchymatous pith, accessory vascular bundles, mucilage idioblasts and hypodermis in the mesophyll, colleters, and non-secretory trichomes. The position and type of stomata, type of mesophyll, number of palisade and spongy parenchyma layers, position of mucilage idioblasts in the mesophyll, and the number of accessory vascular bundles of petiole/rachis provided useful characters for discriminating the eight species of the subsect. Baseophyllum. Histochemical tests reveal the presence of total polysaccharides, pectins, mucilage and phenolic compounds in the idioblasts. Other xeromorphic characters such as thickened outer periclinal cell walls, compact mesophyll, mucilage idioblasts and hypodermis in the mesophyll may help enable the studied species to survive in exposed sunny areas.     

Leaf structure of Bromelia and its significance for the evolution of Bromelioideae (Bromeliaceae)

This study investigated the leaf structure of Bromelia and its importance for understanding the evolution of Bromelioideae (Bromeliaceae). Because of the scarcity of informative molecular characters in Bromeliaceae, this study evaluates the relevance of anatomical characters for the taxonomy and phylogeny of Bromelia and the subfamily Bromelioideae. Anatomical studies in monocots have shown that the combined application of anatomical and external morphological characters from leaf structure can improve the taxonomic delimitation of species, genera and subgenera, and is very informative for inferring phylogenies. The current study analyzed the leaves of 27 species of Bromelia and found that the most important characters for the systematics of this group are the occurrence of a water storage hypodermis, the number of stalk cells of peltate scales, the presence of a ribbed abaxial surface, the occurrence of palisade parenchyma on the adaxial side, the shape of the cells that surround the air lacunae, the presence of raphides and secretory channels, and the occurrence of fibrous extensions on the bundle sheath on minor veins. Combining our results with those described for the family, we made a list of the anatomical characters that can be used in phylogenetic studies of Bromelioideae.     

Correlations among anatomical,morphological, chemical and agronomic characteristics of leaf blades in Panicum maximum genotypes

Tropical forage grasses present high growth rates and biomass yields, partly due to its C₄ photosynthetic pathway. Considering this, the anatomy of the grasses related to morphology and chemical composition of leaf blades may influence consumption and digestibility, thus interfering in the forage quality. Agronomic, morphological, anatomical and chemical characteristics of leaves of nine Panicum maximum genotypes were evaluated in Brazil, to verify if these characteristics and the associations among them may influence the quality of the leaf blades. A randomized complete blocks design was used with nine treatments and three replications. Three evaluation harvests were done in the rainy season and one in the dry season to evaluate forage yields and quality. Quality was determined through NIRS – Near Infrared Spectrometer. One day prior to each harvest, four leaf blades per plot were harvested for morphological and anatomical evaluations. Results were subjected to analysis of variance and mean comparison by Tukey test, and to simple linear and canonical correlations by SAS. Leaf width was positively correlated with mesophyll. The parenchyma bundle sheath was associated with leaf area and specific leaf area. Neutral detergent fibre was positively correlated with the parenchyma bundle sheath area and specific leaf area. The specific leaf area was negatively correlated with in vitro organic matter digestibility. Morphological differences among P. maximum genotypes did not interfere in biomass accumulation. Considering this, leaf width may be a supplementary tool, that may be used in the early phases of the process of genotype selection, for discriminating qualitatively promising high yielding materials.     

Plasmodesmatal frequency in relation to short-distance transport and phloem loading in leaves of barley (Hordeum vulgare). Phloem is not loaded directly from the symplast

We investigated the phloem loading pathway in barley, by determining plasmodesmatal frequencies at the electron microscope level for both intermediate and small blade bundles of mature barley leaves. Lucifer yellow was injected intercellularly into bundle sheath, vascular parenchyma, and thin-walled sieve tubes. Passage of this symplastically transported dye was monitored with an epifluorescence microscope under blue light. Low plasmodesmatal frequencies endarch to the bundle sheath cells are relatively low for most interfaces terminating at the thin- and thick-walled sieve tubes within this C₃ species. Lack of connections between vascular parenchyma and sieve tubes, and low frequencies (0.5% plasmodesmata per μm cell wall interface) of connections between vascular parenchyma and companion cells, as well as the very low frequency of pore-plasmodesmatal connections between companion cells and sieve tubes in small bundles (0.2% plasmodesmata per μm cell wall interface), suggest that the companion cell-sieve tube complex is symplastically isolated from other vascular parenchyma cells in small bundles. The degree of cellular connectivity and the potential isolation of the companion cell-sieve tube complex was determined electrophysiologically, using an electrometer coupled to microcapillary electrodes. The less negative cell potential (average –52 mV) from mesophyll to the vascular parenchyma cells contrasted sharply with the more negative potential (–122.5 mV) recorded for the companion cell-thin-walled sieve tube complex. Although intercellular injection of lucifer yellow clearly demonstrated rapid (0.75 μm s^-1) longitudinal and radial transport in the bundle sheath-vascular parenchyma complex, as well as from the bundle sheath through transverse veins to adjacent longitudinal veins, we were neither able to detect nor present unequivocal evidence in support of the symplastic connectivity of the sieve tubes to the vascular parenchyma. Injection of the companion cell-sieve tube complex, did not demonstrate backward connectivity to the bundle sheath. We conclude that the low plasmodesmatal frequencies, coupled with a two-domain electropotential zonation configuration, and the negative transport experiments using lucifer yellow, precludes symplastic phloem loading in barley leaves.     

The relation of carbon dioxide compensation and chlorenchymatous vascular bundle sheaths in leaves of dicots

Low CO₂ compensation points have been found to be associated with several unusual characteristics related to photosynthesis. One such characteristic is a prominent, chlorenchymatous vascular bundle sheath in the leaves. It has been suggested that the presence of this sheath in dicotyledons can serve as a means of detecting low CO₂-compensating species. We collected 88 dicotyledon species from 22 families reported to have chlorenchymatous sheaths. Of the 88, only three, Tribulus terrestris, L., Boerhaavia paniculata, L. C. Rich, and Trianthema portulacastrum L., had low CO₂ compensation points. Cross sections of the leaves of the other species revealed that they did have chlorenchymatous vascular bundle sheaths. However, these sheath cells contained chloroplasts which were not specialized for starch formation as were the bundle sheath chloroplasts of the low CO₂-compensating species.     

大车前与平车前营养器官的比较解剖学研究

利用光学显微镜和石蜡切片技术对生长在盐碱地的大车前和平车前的营养器官进行了比较解剖学研究。结果表明二者的解剖结构存在差异:平车前根的木栓层较大车前的狭窄,栓内层细胞层数较少;平车前叶表皮的细胞角质膜较薄,孔下室较小,构成维管束鞘的薄壁细胞较大车前小,束鞘细胞两层。二者都具有适应盐渍环境的结构特征:营养器官中通气组织发达;根结构中薄壁组织和木栓发达;叶表皮角质层发达。     

On the morphological position of Paepalanthus subgenus Psilandra (Eriocaulaceae)

Paepalanthus subgen. Psilandra comprises two species known only from the type collections. The aim of this study is to evaluate their unique floral morphology within a given phylogenetic context and its implications for taxonomy. Flowers of both species were analyzed, described, and photographed using a digital microscope. Taxonomically relevant characters were mapped on previous molecular phylogenies. The flowers of both species fit the Paepalanthoideae pattern and are quite similar to each other. The petals of the pistillate flowers, although free, show evidence of previous fusion in the middle. The free condition is interpreted to be secondary, probably because of the handling of the specimens or to the current developmental stage. Character mapping indicates that both species share floral features with Syngonanthus but not with Paepalanthus. Therefore, based on the median fusion of the pistillate flower petals, absence of floral bracts, petals of the pistillate flower being shorter than the sepals, simple stigmas in the commissural position, the pedicellate pistillate flower, and the glabrous petals and sepals, we transfer P. subgen. Psilandra to synonymy of Syngonanthus sect. Syngonanthus and combine both species in this genus.     

Leaf anatomical studies of Eugenia L. (Myrtaceae) species from the Malay Peninsula

KHATIJAH, H. H., CUTLER, D. F. & MOORE, D. M., 1992. Leaf anatomical studies of Eugenia L. species from the Malay Peninsula. The leaf and petiole anatomy of 25 Malaysian species of Eugenia L. sensu lato was investigated to determine the value of vegetative anatomy for the identification of non-flowering material. Twelve characters proved useful for the identification of species: type of stomata; shape of leaves in T.S.; shape of midrib bundle; sclerenchyma sheath; cutinization of outer epidermal wall; presence or absence of sclereids, idioblasts, hypodermis, columnar epidermal cells, solitary crystals; number of palisade layers and the shape of vascular strand, sclerenchyma sheath and sclereids in the parenchyma of the petiole. Using these characters an anatomical key was constructed to assist identification of the species. Only the closely related species E. stapfiana and E. nilidula could not be separated on anatomical characters.
The anatomical data support the treatment of Henderson (1949) of section Acmena but not of sections Cleistocalyx and Fissicalyx , which fit better in section Syzygium.     

费菜和长药八宝叶形态结构及其与耐旱性关系的研究

利用光学显微镜观察比较了费菜（Sedum aizoon L.）和长药八宝（S. spectabilis Boreau）的叶表皮形态及解剖结构,并对其进行不同程度的水分胁迫,研究干旱逆境条件对其叶片显微结构的影响。结果表明：两种景天的叶形态、解剖结构与耐旱性之间存在较密切的相关性。两种植物成熟叶均具较薄的角质层和发达的储水组织,叶肉组织中没有栅栏组织和海绵组织的明显分化。二者的维管组织较发达,且具较发达的孔下室,维管束周边发现一些吸水能力较强的异细胞,这些特征是植物耐旱的关键。     

9种芦荟属植物叶的结构和芦荟素含量的比较研究

9种芦荟属植物叶的比较解剖研究结果表明,它们都具有明显的旱生叶的结构特征,其维管束的韧皮部内都有大型薄壁细胞,但其表皮角质膜的厚度,表面纹饰,气孔上,下腔的形状和大小,同化组织 导 ,细胞分化情况,维管束的大小,分布密度和其大型薄壁细胞占维管束的比例,中央贮水组织占叶横切面的比例等特征,在各种间存在差异,且性状稳定,可以作为该属内种间分类的解剖学指标,植物化学分析结果表明,9种植物叶内蒽醌类物质的主要种类和含量不同,其含量高,低与叶内维管束密度,大型薄壁细胞占维管束的比例以及同化组织的厚度密度切相关,从而为芦荟属植物选育商业用良种提供了植物解剖学依据。     

Leaf morphology and anatomy of Manilkara Adans. (Sapotaceae) from northeastern Brazil

Aiming to complement the group’s taxonomy, the objective of this work is to describe and characterise, searching for morpho-anatomical patterns, the leaf anatomy of the Manilkara species occurring in the northeast region of Brazil. The leaves of all Manilkara species were analysed, using herbaria specimens and collections made in the field by the authors. The characterisation of the leaf blades was performed according to the usual protocols in vegetal anatomy. As diagnosed by anatomical characters, the Manilkara Adans. genus presents petioles with a thick cuticle, a uniseriate epidermis, a sclerenchymatous sheath involving the vascular bundle, and laticifers on the cortical and medullar regions. The anatomical data increase the M. elata and M. huberi synonymisation and show the importance of the anatomy as a complementary tool to the taxonomy based on external morphology.