Chromosome numbers of goldenrods,Euthamia and Solidago (Compositae: Astereae). II. Additional counts with comments on cytogeography

Chromosome number determinations were made from 407 wild or transplanted individuals and seedlings representing 65 taxa and hybrids inEuthamia andSolidago. The following are first reports:Euthamia remota, 2n=9II;Solidago leavenworthii, 2n=54;S. mollis, 2n=36;S. mollis var.angustata, 2n=36;S. rigida var.glabrata, 2n=9II;S. sempervirens var.azorica, 2n=9II; andS. sparsiflora, 2n=54. Most species have been sampled only a few times or are consistently of one cytotype. Sufficient counts have been made to indicate some general patterns of cytotype distribution in the following species complexes:S. gigantea, S. canadensis, S. flexicaulis, S. rugosa, andS. uliginosa.     

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.     

Growth,phenology, and biomass allocation of alien Solidago species in central Europe

Plant invasion is a major threat to the integrity of an ecosystem. Exceptionally successful invaders in Europe are the American species of Solidago genus. In this study we examined growth, reproduction, and phenology of Solidago species, of American origin, growing in central Europe (S. altissima, S. canadensis, S. gigantea, S. graminifolia). These taxa were compared with two native species: Solidago virgaurea and Tanacetum vulgare. We observed high differentiation in height, number of shoots, and biomass production between individuals within taxa. Generally, the invasive species produced substantially (two to five times) more biomass than the native ones, being statistically significant in the comparison of alien Solidago graminifolia and S. gigantea versus native S. virgaurea and T. vulgare. The ratio of biomass of reproductive parts to overall biomass varied considerably among years, but generally the lowest one was for Solidago altissima, and the highest for S. graminifolia. It shows a lack of a clear pattern of differentiation between alien and native species in terms of biomass investment in reproduction. We observed a general tendency of allocation of a major part of biomass in rhizomes by phalanx species (S. graminifolia and S. gigantea), while species with guerrilla strategy (S. altissima and S. canadensis) invested more biomass in stems and leaves. However, because of the high variability there was no clear, stable pattern of statistically significant differences between these two groups. The results suggest that S. graminifolia reveals a strong potential of invasion, in spite of its, so far limited, distribution in Europe.     

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.     

How many heteropteran species can live on alien goldenrods Solidago canadensis and S. gigantea in Europe?

During the vegetation periods of 2001–2003 Heteroptera associated with the invasive alien tall goldenrods Solidago canadensis and S. gigantea were studied in seven model habitats in the north-eastern part of the Czech Republic. Heteropterans associated with adjacent growths were also studied in 2002–2003. A set of 3,042 specimens of 127 samples was analyzed with the aim of estimating average species richness, abundance and trophic structure of the heteropteran assemblages of the studied plant stand. On alien Solidago, 68 heteropteran species were recorded and 71 species were collected in the stands adjacent to the tall goldenrods with 48 shared species. Despite the nearly indentical species richness and similar abundances in Solidago and adjacent stands, there are differences in the trophic structure. The majority of the shared species and species found on Solidago canadensis only are polyphagous contrary to the majority of stenophagous species found on Solidago free stands only. Only a small proportion of heteropteran species that were recorded on alien Solidago stands are specialized to Asteraceae and their abundance was mostly low. Only the lygaeid Nysius senecionis, an Asteraceae specialist, occured in masses on S. canadensis in sunny and warm habitats. Similarly, predatory Orius minutus and O. niger reached high abundance values in Solidago stands compared to adjacent stands.     

Is phenotypic plasticity an explanation for the invasiveness of goldenrods (Solidago and Euthamia) in Europe?

High phenotypic plasticity contributes to invasiveness of alien species. Goldenrods of American origin (Solidago canadensis, S. gigantea and Euthamia graminifolia) have successfully invaded Europe, and this success can be related to their high phenotypic plasticity. The aim of our study was to test the hypothesis of higher phenotypic plasticity of invasive goldenrods by comparing them with native taxa: closely related Solidago virgaurea and similar ecologically, invasive in other regions Tanacetum vulgare. The species studied were grown in a common garden on three different substrates with different fertilizers. After 3 years, the height and number of ramets, biomass production and allocation, phenology and nitrate reductase activity (NRA) were measured. The highest level of phenotypic plasticity was exhibited by the competitively weak native species S. virgaurea. The invasive species produced relatively high biomass and exhibited a moderate level of phenotypic plasticity. Variability in the studied traits did not always correlate with differences in substrates. The invasion success of non-native goldenrods is not caused by their having a higher phenotypic plasticity than native species. Rather, the non-natives should be considered as “Jack of all traits, and master of some”.     

Sequential flowering of neighboring goldenrods and the movements of the flower predator Epicauta pennsylvanica

Summary As neighboring plants flower sequentially, do flower feeders preferentially remain in the area, rather than move to another area with flowering plants? I examined the movements of the meloid beetle Epicauta pennsylvanica, a flower predator specializing on Solidago, in four types of replicated experimental plots — monocultures of Solidago altissima, or S. altissima interplanted with members of the same genus, same family, or different taxonomic orders. I released marked beetles only in the genus plots, which contained four species of Solidago, two that bloom before S. altissima. The number of beetles in the genus plots declined steadily as S. altissima came into flower in all the plots; the total number of beetles in all the plots remained fairly constant. I found no evidence that plant neighborhoods affected beetle distribution. Beetles foraging on the early blooming Solidago species did not remain in the genus plots as S. altissima came into flower. In addition, beetles that left the genus plots did not differentially accumulate in any of the other plot types, even though one type of plot was a monoculture with four times the density of S. altissima than the other plots.     

Correlations between genet architecture and some life history features in three species of Solidago

Summary Members of the genus Solidago are among the most widely studied model systems in plant population biology. A comparative study of Solidago canadensis, S. altissima, and S. gigantea in an experimental garden showed that the three species had different patterns of shoot growth and development, leaf morphology and physiology, and biomass allocation at harvest. These differences were also found in the field. Contrary to some current taxonomic usage, our results show that S. canadensis should ecologically be treated as a separate taxon distinct from S. altissima, and that the latter may be grouped together with S. gigantea. Many of the biological differences between S. canadensis and the other two taxa, such as differential investment into sexual reproduction versus clonal growth, may be explained by differences in genet architecture. These architectures concern high compared to lower within-genet shoot density resulting from differences in rhizome lengths among the taxa (shorter in S. canadensis than in S. altissima and S. gigantea).     

Minor vein structure and sugar transport in Arabidopsis thaliana

Leaf and minor vein structure were studied in Arabidopsis thaliana (L.) Heynh. to gain insight into the mechanism(s) of phloem loading. Vein density (length of veins per unit leaf area) is extremely low. Almost all veins are intimately associated with the mesophyll and are probably involved in loading. In transverse sections of veins there are, on average, two companion cells for each sieve element. Phloem parenchyma cells appear to be specialized for delivery of photoassimilate from the bundle sheath to sieve element-companion cell complexes: they make numerous contacts with the bundle sheath and with companion cells and they have transfer cell wall ingrowths where they are in contact with sieve elements. Plasmodesmatal frequencies are high at interfaces involving phloem parenchyma cells. The plasmodesmata between phloem parenchyma cells and companion cells are structurally distinct in that there are several branches on the phloem parenchyma cell side of the wall and only one branch on the companion cell side. Most of the translocated sugar in A. thaliana is sucrose, but raffinose is also transported. Based on structural evidence, the most likely route of sucrose transport is from bundle sheath to phloem parenchyma cells through plasmodesmata, followed by efflux into the apoplasm across wall ingrowths and carrier-mediated uptake into the sieve element-companion cell complex. Received: 5 October 1999 / Accepted: 20 November 1999     

Light reduction predicts widespread patterns of dominance between asters and goldenrods

Here we investigate the long-cited pattern that throughout the eastern United States, Solidago species (goldenrods), and in particular S. canadensis displace Aster species and dominate old-field communities. Theory predicts that such a ubiquitous pattern of repeated dominance should be linked to competitive ability for a limiting resource. However, no one has investigated this possibility in old-fields, representing a potentially significant gap in our understanding of a common human-altered environment. We tested the hypothesis that S. canadensis is the superior competitor for light compared to other common co-occurring goldenrod species, and that the goldenrods in general are the superior competitors for light compared to coexisting aster species, which are typically less abundant. We tested this hypothesis by comparing the light attenuation abilities of four goldenrod species, S. canadensis, S. rugosa, S. gigantea, and Euthamia graminifolia, and three aster species, Aster novae-angliae, A. pilosus, and A. prenanthoides. Consistent with our hypothesis, S. canadensis had a greater ability to attenuate light than any of the other goldenrods at higher densities, and the goldenrods overall had a greater ability to attenuate light than the asters. By conducting a census in our study area, we verified that S. canadensis is locally the most abundant goldenrod and that goldenrods are more locally abundant than asters. Furthermore, by conducting a literature survey we found evidence that S. canadensis replaces A. pilosus through time. Thus we found a close correspondence between relative abundance in the field and light attenuation ability in field experiments. These results are consistent with theory predicting that competition for limiting resources, in this case light, explains patterns of dominance and relative abundance in old-field plant communities.     

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.     

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.     

Contributions to the Wood Anatomy of the Rubioideae (Rubiaceae)

Damnacanthus , Lasianthus, Saldinia, and Trichostachys are also included. Wood anatomical characters are compared with recent phylogenetic insights into the study group on the basis of molecular data. The observations demonstrate that the delimitation and separation of several taxa from the former Coussareeae/Morindeae/Prismatomerideae/Psychotrieae aggregate is supported by wood anatomical data. The Coussareeae can be distinguished from the other Rubioideae by their scanty parenchyma, septate libriform fibres, and the combination of uniseriate and very high multiseriate rays with sheath cells. Axial parenchyma bands and fibre-tracheids characterise Gynochtodes and some species of Morinda (Morindeae s.str.), but the latter genus is variable with respect to several features (e.g. vessel groupings and axial parenchyma distribution). Wood data support separation of Rennellia and Prismatomeris from Morindeae s.str.; vessels in both genera are exclusively solitary and axial parenchyma is always diffuse to diffuse-in-aggregates. Damnacanthus differs from the Morindeae alliance by the occurrence of septate fibres, absence of axial parenchyma, and the occasional presence of fibre wall thickenings. There are interesting similarities between members of the Lasianthus clade and the Pauridiantheae/Urophyleae group such as the sporadic occurrence of spiral thickenings in axial parenchyma cells. Received 26 January 2001/ Accepted in revised form 6 June 2001     

Microsatellite loci for studies of population differentiation and range expansion in Solidago sempervirens L. (Asteraceae)

For studies of population differentiation and range expansion in the seaside goldenrod, Solidago sempervirens, we identified hypervariable molecular markers by screening genomic libraries enriched for microsatellite motifs. We designed primers that reliably amplified nine polymorphic loci. High polymorphism in a population from Delaware Bay, USA suggests that the loci will be useful in population studies. The success of cross‐amplifications in 11 species of Asteraceae varied among loci and did not appear to reflect phylogenetic relationships within Solidago.     

Ultrastructural changes during early infection of Vigna unguiculata and Phaseolus vulgaris leaves by Xanthomonas axonopodis pv. phaseoli and an unexpected association between chloroplast and mitochondrion

Common bacterial blight, caused by Xanthomonas axonopodis pv. phaseoli, is an important disease affecting beans causing considerable yield losses in many producing areas. Our aim was to investigate Xap–beans interactions in the first 48 h after infection analyzing structurally relevant changes by electron microscopy. Scanning electron microscopy analysis demonstrated morphological changes on leaf surfaces of both species. Transmission electron microscopy showed different types of chloroplast structural disorganization in both parenchyma and bundle sheath cells of Vigna unguiculata which were only observed in parenchyma cells of Phaseolus vulgaris. Additionally, we report an unexpected physical association between the chloroplasts and mitochondria observed only to P. vulgaris-infected plants.     

Idioblasts and other unusual internal foliar secretory structures in Scrophulariaceae

 Leaf samples of mostly herbarium specimens (237 species of 172 genera) were cleared. Internal secretory structures of large size or unusual shape were detected and observed. Selected samples were processed into resin and sectioned for light microscopy or prepared for scanning electron microscopy. Adding results from two earlier publications, our survey includes 365 species from 174 genera. Five types of internal secretory structure, mostly unknown previously in the traditional Scrophulariaceae, occur in only nine genera: 1) single-celled subepidermal idioblasts, empty at maturity, in Scrophularia and Verbascum (Lersten and Curtis 1997), 2) 2–16 (or more)-celled nodules, with fibrous contents, in mid-mesophyll strata of Graderia and Radamaea, 3) epithelium-lined oil cavities in Leucophyllum (Lersten and Beaman 1998) and Capraria, 4) Kranz-type enlarged bundle sheath in Anticharis, and 5) paraveinal mesophyll (PVM) in Picria and Bonnaya. Received April 24, 2000 Accepted October 27, 2000     

Anatomy, chloroplast structure and compartmentation of enzymes relative to photosynthetic mechanisms in leaves and cotyledons of species in the tribe Salsoleae (Chenopodiaceae)

Certain members of the family Chenopodiaceae are the dominant species of the deserts of Central Asia; many of them are succulent halophytes which exhibit C4-type CO2 fixation of the NAD- or NADP-ME (malic enzyme) subgroup. In four C4 species of the tribe Salsoleae, the Salsoloid-type Kranz anatomy in leaves or stems was studied in relation to the diversity in anatomy which was found in cotyledons. Halocharis gossypina, has C4 NAD-ME Salsoloid-type photosynthesis in leaves and C3 photosynthesis in dorsoventral non-Kranz cotyledons; Salsola laricina has C4 NAD-ME Salsoloid-type leaves and C4 NAD-ME Atriplicoid-type cotyledons; Haloxylon persicum, has C4 NADP-ME Salsoloid-type green stems and C3 isopalisade non-Kranz cotyledons; and S. richteri has C4 NADP-ME Salsoloid-type leaves and cotyledons. Immunolocalization studies on Rubisco showed strong labelling in bundle sheath cells of leaves and cotyledons of organs having Kranz anatomy. The C4 pathway enzyme phosphoenolpyruvate carboxylase was localized in mesophyll cells, while the malic enzymes were localized in bundle sheath cells of Kranz-type tissue. Immunolocalization by electron microscopy showed NAD-ME is in mitochondria while NADP-ME is in chloroplasts of bundle sheath cells in the respective C4 types. In some C4 organs, it was apparent that subepidermal cells and water storage cells also contain some chloroplasts which have Rubisco, store starch, and thus perform C3 photosynthesis. In non-Kranz cotyledons of Halocharis gossypina and Haloxylon persicum, Rubisco was found in chloroplasts of both palisade and spongy mesophyll cells with the heaviest labelling in the layers of palisade cells, whereas C4 pathway proteins were low or undetectable. The pattern of starch accumulation correlated with the localization of Rubisco, being highest in the bundle sheath cells and lowest in the mesophyll cells of organs having Kranz anatomy. In NAD-ME-type Kranz organs (leaves and cotyledons of S. laricina and leaves of H. gossypina the granal index (length of appressed membranes as a percentage of total length of all membranes) of bundle sheath chloroplasts is 1.5 to 2.5 times higher than that of mesophyll chloroplasts. In contrast, in the NADP-ME-type Kranz organs (S. richteri leaves and cotyledons and H. persicum stems) the granal index of mesophyll chloroplasts is 1.5 to 2.2 times that of the bundle sheath chloroplasts. The mechanism of photosynthesis in these species is discussed in relation to structural differences.     

Composite bundles,the host/parasite interface in the holoparasitic angiosperms Langsdorffia and Balanophora (Balanophoraceae)

Composite bundles are not simply a type of vascular bundles, but an integrated host/parasite interface. We investigated their structure in tubers of Langsdorffia and Balanophora. Composite bundles in both genera have similar components: 1) a central mass of host vascular tissues among which are located large parasite transfer cells; 2) a sheath of parasite parenchyma surrounding the central host vascular tissues; 3) specialized conducting tissues in the sheath; and 4) apical meristems composed of both host and parasite meristematic cells. Sheath parenchyma is recognizable from parasite tuber matrix by having thinner cell walls, and, especially in Langsdorffia, by the presence of collapsed matrix cells between the bundle sheath and tuber matrix. Sheath-conducting tissues consist of densely cytoplasmic transfer cells and small sieve tube members; in Langsdorffia, tracheary elements are also present. These sheath bundles connect with vascular bundles of the tuber matrix. Direct host/parasite contact only occurs by means of parasite transfer cells in the composite bundles. There is no xylem-xylem contact at the host/parasite interface. Abundance of parasite transfer cells suggests that they play an important role in nutrient absorption and translocation.     

Effects of fusicoccin and abscisic acid on glucose uptake into isolated beetroot protoplasts

Uptake of glucose, 3-O-methylglucose and sucrose into beetroot protoplasts is considerably stimulated by 10^–6M fusicoccin. This effect is decreased in the presence of 10mM Na⁺ or K⁺, 2 mM Mg²⁺ or Ca²⁺. Whereas fusicoccin causes no change in the pH-optimum of the sugar uptake (pH 5.0), the apparent K_m of this uptake which obeys a biphasic kinetics is decreased by the action of fusicoccin. In the protoplast suspension, fusicoccin induces an acidification which is suppressed by uncoupling agents. Correspondingly, uncouplers as well as vanadate and diethylstilbestrol markedly inhibit the effect of fusicoccin on sugar uptake. The present data support the view that glucose uptake into beetroot protoplasts depend on the proton-pumping activity of the plasmalemma-ATPase. cis–Abscisic acid diminishes significantly the fusicoccin-enhanced glucose uptake. By using a radioimmunoassay, the internal abscisic acid content of the protoplast was estimated to be in the range of 10^–6 M. Protoplasts isolated from bundle tissue contain twice as much abscisic acid as those derived from storage parenchyma. Because protoplasts from the bundle tissue were shown to take up sugars much faster than those from the storage cells, the observed effect of abscisic acid might reflect an involvement of this hormone in the regulation of carbohydrate partitioning in the beet.Abbreviations ABA cis–abscisic acid - bundle protoplast protoplasts isolated from the conducting tissue of beetroots - DES diethylstilbestrol - FC fusicoccin - 3-OMG 3-O-methylglucopyranose - PCMBS p–chloromercuribenzenesulfonic acid - storage protoplasts protoplasts isolated from storage parenchyma