The tapetum inSchizaea pectinata (Schizaeaceae) and a comparison with the tapetum inPsilotum nudum (Psilotaceae)

A combination tapetum consisting of a cellular, parietal component and a plasmodial component occurs inSchizaea pectinata. A single, tapetal initial layer divides to form an outer parietal layer which maintains its cellular integrity until late in spore wall development. The inner tapetal layer differentiates into a plasmodium which disappears after the outer exospore has developed. In the final stages of spore wall development, granular material occurs in large masses and is dispersed as small granules throughout the sporangial loculus. No tapetal membrane develops. Comparisons are drawn with the combination tapetum found inPsilotum nudum.     

Taxonomy ofCarminatia (Asteraceae,Eupatorieae)

A taxonomic study of the largely Mexican genusCarminatia is rendered. It is comprised of three closely related species:C. tenuiflora, C. recondita andC. anomala spec. nova. Illustrations, dot maps, keys to species and complete synonymy are presented.     

Taxonomy and phylogeny of the tribePlucheeae (Asteraceae)

The tribePlucheeae (Benth.)A. Anderb., has been analysed cladistically by means of a computerized parsimony program (Hennig 86), using theArctotideae as outgroup. The results of the analysis are presented in a consensus tree and one cladogram. Four major monophyletic subgroups can be recognized: TheColeocoma group (3 genera), thePterocaulon group (3 genera), theLaggera group (6 genera), and thePluchea group (12 genera). All recognized genera are described and most genera are supplied with taxonomical notes including comments on their taxonomic status. Genera such asBlumea, Pluchea, andEpaltes are demonstrated to be unnatural assemblages.Monarrhenus andTessaria are both closely related to thePluchea complex. The old generic nameLitogyne Harv. has been taken up for one species ofEpaltes, the genusRhodogeron is reduced to a synonym ofSachsia, and the following new combinations are made;Litogyne gariepina (DC.)A. Anderb., andSachsia coronopifolia (Griseb.)A. Anderb.     

Systematic study of the genusDelilia (Asteraceae,Heliantheae)

A taxonomic study of the genusDelilia is presented. Two species are recognized:D. biflora, a widespread weed of tropical and subtropical regions of the New World, extending from Mexico to northern Argentina and recently introduced in the Cape Verde Islands; andD. repens, endemic to the Galapagos islands. A key to the species, illustrations, distribution maps, and complete synonymy are presented.     

Synopsis ofVerbesina sect.Ochractinia (Asteraceae)

A taxonomic revision ofVerbesina sect.Ochractinia is presented. 40 species plus 7 additional infraspecific taxa are recognized. A table of the distribution of the species, key to the species and synonymy are included. Relationships to other sections ofVerbesina and species relationships within sect.Ochractinia are discussed.     

The tapetum: Its form,function, and possible phylogeny inEmbryophyta

It appears that the tapetum is universally present in land plants, even though it is sometimes difficult to recognize, because it serves mostly as a tissue for meiocyte/spore nutrition. In addition to this main function, the tapetum has other functions, namely the production of the locular fluid, the production and release of callase, the conveying of P.A.S. positive material towards the loculus, the formation of exine precursors, viscin threads and orbicules (= Ubisch bodies), the production of sporophytic proteins and enzymes, and of pollenkitt/tryphine. Not all these functions are present in all land plants:Embryophyta. Two main tapetal types are usually distinguished in theSpermatophyta: the secretory or parietal type and the amoeboid or periplasmodial type; in lower groups, however, other types may be recognized, with greater or lesser differences. A hypothetical phylogenesis of the tapetum is proposed on the basis of its morphological appearance and of the nutritional relations with meiocytes/spores. The evolutionary trends of the tapeta tend towards a more and more intimate and increasingly greater contact with the spores/pollen grains. Three evolutionary trends can be recognized: 1) an intrusion of the tapetal cells between the spores, 2) a loss of tapetal cell walls, and 3) increasing nutrition through direct contact in narrow anthers.     

Biology of afroalpineDendrosenecio (Asteraceae)

The genusDendrosenecio (giant groundsels), encompassing three species and 12 subspecies, is endemic to the high mountains of East and Central Africa where it constitutes the most conspicuous components of the afroalpine vegetation. Two lifeforms, the arborescent and the prostrate rhizomatous, are regarded as the results of evolution from forest-living woody or herbaceous ancestors. Due to the uninterrupted growth period in the tropics, there are no anatomical or morphological features which allow conventional age determination. However, stem elongation rates have been determined (3–5.5 cm per year) and indicate an age of about 250 years for the tallest arborescent Dendrosenecios which may reach a height of 10 m. 30 to 120 large leaves are clustered in an enormous terminal rosette, justifying the term giant rosette plants. A leaf bud, consisting of about as many developing leaves as the rosette contains, is found in the center. During the nocturnal frost period the adult rosette leaves form a so-called night-bud by nyctinastic upwards bending and thus protect the leaf bud from freezing by insulation. The stem is surrounded by a mantle of persistent dead leaves; this ameliorates the microclimate of the pith-cells which greatly contribute to water transport into the leaves. Below the leaf rosette a zone of putrefaction is found, from where the decay of the dead leaves apparently provides nutrients directly to the growing stem. The population dynamics of the arborescentD. keniodendron is characterized by a simultaneous inflorescence development at irregular intervals of up to more than twenty years. Due to sporadic flowering and a seedling survival rate of less than 1%, oscillations of the population size are to be expected.     

Phenotypic plasticity inCrepis tectorum (Asteraceae)

Two experiments were carried out using two different approaches to compare populations ofCrepis tectorum (Asteraceae). One was based on a comparison of means of various vegetative and reproductive characters and another was based on a comparison of response patterns of the same characters in a series of environments. Population divergence within two earlier recognized form series, one from weed habitats and one from alvar habitats on Baltic islands, resulted in a partially overlapping pattern in cluster analyses based on character means. However, the pattern revealed by a comparison of the direction and amount of plastic response suggested that populations within the two form series had more similar response patterns than other combinations of populations. It was concluded that patterns of plasticity may provide useful additional information on the overall similarity among taxa. An hypothesis that plants in weed populations should exhibit a greater phenotypic response to the environments than plants in alvar populations was rejected.     

Karyological notes onCentaurea sect.Acrocentron (Asteraceae)

The karyology ofCentaurea sect.Acrocentron is surveyed. 19 chromosome counts on 8 species are reported; those onC. acaulis, C. crocata, C. galianoi, C. pubescens, andC. malinvaldiana are new. The basic chromosome numbers of the section are x = 11 and x = 10. Karyological arguments have been used to show that evolution was from x = 11 to x = 10. This is supported by biogeographical data. Two main centres of diversification of sect.Acrocentron were studied from that point of view: the East and the Southwest Mediterranean region.     

The common occurrence of incompletely developed pollen of Eupatorium (Compositae: Eupatorieae)

Mature anthers of Eupatorium serotinum Michx., E. coelestinum L. and E. purpureum L. (Compositae: Eupatorieae) contain both loose mature pollen grains and acetolysis-resistant extratapetal sacs enclosing clusters of immature pollen. Many sacs contain incompletely developed pollen grains that differ in size diameter, with spines of varying lengths and degrees of acuity and with colpal areas having narrow to markedly protracted margins. We presume that all nutrients for pollen development within sacs come from plasmodial tapetum included therein. Thus, pollen in portions of the sac or in isolated sacs with inadequate plasmodial tapetum may be incompletely developed. Such partial development may result from stress, e.g. insufficient nutrients and water. While our data are from three species of Eupatorium, we have long noted the syndrome in other Eupatorieae and other tribes in the Compositae. Curtailing supplies for development of some portions of an anther could be an evolutionary strategy for survival of the species through periods of stress.     

New views of tapetum ultrastructure and pollen exine development in Arabidopsis thaliana

Background and Aims

The Arabidopsis thaliana pollen cell wall is a complex structure consisting of an outer sporopollenin framework and lipid-rich coat, as well as an inner cellulosic wall. Although mutant analysis has been a useful tool to study pollen cell walls, the ultrastructure of the arabidopsis anther has proved to be challenging to preserve for electron microscopy.

Methods

In this work, high-pressure freezing/freeze substitution and transmission electron microscopy were used to examine the sequence of developmental events in the anther that lead to sporopollenin deposition to form the exine and the dramatic differentiation and death of the tapetum, which produces the pollen coat.

Key Results

Cryo-fixation revealed a new view of the interplay between sporophytic anther tissues and gametophytic microspores over the course of pollen development, especially with respect to the intact microspore/pollen wall and the continuous tapetum epithelium. These data reveal the ultrastructure of tapetosomes and elaioplasts, highly specialized tapetum organelles that accumulate pollen coat components. The tapetum and middle layer of the anther also remain intact into the tricellular pollen and late uninucleate microspore stages, respectively.

Conclusions

This high-quality structural information, interpreted in the context of recent functional studies, provides the groundwork for future mutant studies where tapetum and microspore ultrastructure is assessed.     

The origin ofSenecio vulgaris (Asteraceae)

Senecio vulgaris L. (2n = 40) is suggested to be of autotetraploid origin fromS. vernalis Waldst. & Kit. (2n = 20). This conclusion is based on results obtained from experimental hybridisations and cytological observations, and the consideration of morphological affinities, patterns of geographical distribution and hybrid formation under natural conditions. The morphological differences between the two species are related to a difference in the breeding system. WhilstS. vernalis is self-incompatible,S. vulgaris is self-compatible and strongly self-pollinating. Equally, other self-pollinating taxa traditionally associated withS. vulgaris are shown to have evolved independently from outbreeding relatives. Within a narrow frame of relationship, annual weeds have evolved in different cytotaxonomic circumstances.     

Heliantheae sensu lato (Asteraceae), clades and classification

The interrelationships within theHeliantheae s. lato and the closely relatedEupatorieae are analyzed and discussed. The basis to this discussion is a cladistic analysis of 141 morphological characters (172 apomorphic states) scored for 97 genera. TheHeleniae s. lato, a subgroup of theHeliantheae s. lato, are paraphyletic, and a monophyletic group corresponding largely to theHeliantheae s. str. is recognized. TheEcliptinae sensuRobinson are polyphyletic. TheCoreopsidinae form an ingroup in theHeliantheae s. str. TheTageteae (Pectidinae) and theMadieae (Madiinae) are two separate branches within the helenioid assemblage of taxa.     

Genetic components of heterocarpy inMicroseris hybrid B87 (Asteraceae,Lactuceae)

Microseris B87 is derived from a single hybrid specimen betweenM. pygmaea with few, weakly hairy peripheral achenes and aM. bigelovii with many, strongly hairy peripheral achenes. Offspring through the F4 and F5 generations obtained by spontaneous selfing were analyzed for the segregation of quantitative and qualitative characters relating to achene dimorphism. The phenotypic effects of two previously identified unlinked genes determining the relative number of outer achenes are characterized in partially and completely homozygous sublines. We show that two morphological markers genetically linked to one of these genes are themselves regulated by the system inducing heterocarpy. Not more than two more unlinked genes are involved in the genetic basis of the heterocarpic response. The interaction of these genes in determining the heterocarpy phenotypes is discussed in the framework of a model postulating genes for a morphogen gradient across the capitulum and genes responding to this gradient.     

Ontogeny of intruding non-periplasmodial tapetum in the wild chicory,Cichorium intybus (Compositae)

The tapetal development ofCichorium intybus L. is investigated using LM and TEM and discussed in relation to the development in other species. During the second meiotic division the tapetal cells become binucleate and lose their cell walls. They intrude the loculus at the time of microspore release from the meiotic callose walls, which means that a locular cavity is never present in this species. During pollen development they tightly junct the exine, especially near the tips of the spines. During the two-celled pollen grain stage they degenerate and most of their content turns into pollenkitt. Until anther dehiscence they keep their individuality, which means that these intruding tapetal cells never fuse to form a periplasmodium. The ultrastructural cytoplasmatic changes during this development are discussed in relation to possible functions.     

Biochemistry and species problems inAmbrosia (Asteraceae-Ambrosieae)

The shrubby to herbaceous genusAmbrosia is centred in the arid regions of N. America. Its remarkable variability appears to be related to occurrence as pioneers and weeds in unstable habitats, extensive migrations and hybridization, seed-longevity, inbreeding, polyploidy and dysploidy. Volatile oils underline that the genus is a natural assemblage. Sesquiterpene lactones profiles parallel migration routes and characterize geographical races in several groups, but patterns of chemical and morphological differentiation sometimes are not congruent. Seed proteins are ± individual-specific and useful as parameters of genetic polymorphism. In connection with other approaches analyses of chemical constituents can greatly help to clarify problems of species, speciation and evolution.Presented at the symposium Speciation and the Species Concept during the XIIth International Botanical Congress, Leningrad, July 8, 1975.     

Geographical variation and genetic analysis of leaf shape inCrepis tectorum (Asteraceae)

Canonical variate analysis of plants raised in a uniform environment was used to study the pattern of geographical variation in leaf shape ofCrepis tectorum (Asteraceae). The diversity in leaf shape was much greater among populations confined to areas with exposed bedrock in the Baltic region than among weed populations scattered throughout Europe and Canada. A Ward's clustering linked outcrop populations from the archipelago of SW. Finland and the islands of Öland (Sweden) and Saaremaa (Estonia) due to the deeply lobed leaves characterizing these populations, while outcrop populations along the coast of E. Sweden were grouped due to their weakly lobed, narrow and dentate leaves. Most of the weed populations were grouped together but there was no tendency for the variation in this group to be related to habitat or geographical location. A mosaic of variation reflected in sharp (random) differentiation among local populations was superimposed on the large-scale ecogeographical pattern.—Crossing data indicated that most of the variation in degree of leaf dissection is governed by one major gene with deeply lobed leaves dominant over weakly lobed leaves. I suggest that the simple pattern of inheritance may have favoured rapid evolutionary changes in leaf shape, particularly in the Baltic area which emerged relatively late from the sea. Genetic correlations may have constrained the pattern of variation at higher taxonomic levels, since some of the trait associations detected in a segregating F₂ generation were also found at the among-population level.     

A cladistic study of North AmericanCoreopsis (Asteraceae: Heliantheae)

A cladistic study of all 44 species of North AmericanCoreopsis was performed using 35 characters. The resulting cladogram indicated that all 11 sections are monophyletic. At the intersectional level, two lineages were revealed, one consisting of six sections occurring almost exclusively in Mexico and California, and another comprising five sections restricted largely to the eastern and southeastern United States. The cladogram is similar to phylogenies produced by less explicit methods but it differs in two major respects: the monotypic sect.Silphidium is placed with other sections from the southeastern United States rather than with Mexican sections, and sect.Anathysana from Mexico is more closely allied with the three California sections than with sect.Electra from Mexico.