Ettore Pacini

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.

Nectaries and nectar.

Contributing Authors. Preface. 1. Introduction E. Pacini, S.W. Nicolson. 1.1 Evolutionary origins. 1.2 Secretions analogous to nectar. 1.3 Floral and extrafloral nectarines. 1.4 Nectar components. 1.5 Organization of this volume. 2. A Systematic Survey of Floral Nectaries G. Bernardello. 2.1 Introduction. 2.2 Nectaries in gymnosperms. 2.3 Nectaries in angiosperms. 2.3.1 Diversity. 2.3.1.1 Nectar presentation. 2.3.1.2 Structure. 2.3.1.3 Fate. 2.3.1.4 Symmetry. 2.3.1.5 Number. 2.3.1.6 Colour. 2.3.2 Factors influencing nectary diversity. 2.3.3 Basic types of floral nectarines. 2.3.4 Nectariferous spurs. 2.3.5 Patterns of variability in nectarines. 2.3.5.1 Asteraceae. 2.3.5.2 Brassicaceae. 2.3.5.3 Cucurbitaceae. 2.3.5.4 Euphorbiaceae. 2.3.5.5 Ranunculaceae. 2.3.5.6 Solanaceae. 2.3.6 Nectaries and deceit pollination. 2.3.6.1 Apocynaceae. 2.3.6.2 Bignoniaceae. 2.3.6.3 Orchidaceae. 2.3.7 Relictual nectarines in anemophilous species. 2.3.8 Distribution of nectary types. 2.3.8.1 Early-branching lineages. 2.3.8.2 Magnoliids. 2.3.8.3 Early-branching monocots. 2.3.8.4 Monocots. 2.3.8.5 Commelinids. 2.3.8.6 Ceratophyllales. 2.3.8.7 Eudicots. 2.3.8.8 Core Eudicots. 2.3.8.9 Rosids. 2.3.8.10 Eurosids I. 2.3.8.11 Eurosids II. 2.3.8.12 Asterids. 2.3.8.13 Euasterids I. 2.3.8.14 Euasterids II. 2.3.9 Evolutionary trends. 3. Nectary Structure and Ultrastructure M. Nepi. 3.1 Introduction. 3.2 Nectary structure and ultrastructure. 3.2.1 Epidermis. 3.2.1.1 Secretory trichomes. 3.2.1.2 Nectary-modified stomata. 3.2.2 Nectary parenchyma. 3.2.2.1 Patterns of plastid development in nectary parenchyma cells. 3.2.3 Subnectary parenchyma. 3.2.4 Nectary vasculature. 3.3 Gynopleural (septal) nectarines. 3.4 Extrafloral nectarines. 3.5 Nectary histochemistry. 4. Nectar Production and Presentation E. Pacini, M. Nepi. 4.1 Introduction. 4.2 Nectar secretion mechanism and models of nectary function. 4.3 Dynamics of nectar components. 4.3.1 Nectar reabsortion: resource recovery and homeostasis. 4.3.2 Nectar standing crop. 4.4 The source of nectar components. 4.5 Ecophysiological significance of parenchyma plastids. 4.6 Nectar presentation. 4.6.1 Floral nectarines. 4.6.2 Extrafloral nectarines. 4.7 Fate of nectar and nectarines. 4.8 Variability of nectar characteristics. 4.8.1 Environmental variables. 4.8.2 Intraspecies variability. 4.8.3 Interpopulation differences. 4.8.4 Variability and experimental design. 5. Nectar Chemistry S.W. Nicolson, R.W. Thornburg. 5.1 Introduction. 5.2 Water. 5.2.1 Nectar concentration. 5.2.2 Chemical and microclimatic influences on nectar concentration. 5.2.3 Viscosity and feeding rates. 5.3 Sugars. 5.3.1 Constancy of sugar composition within species. 5.3.2 The use of sugar ratios can be misleading. 5.3.3 Is sugar composition determined by floral visitors or common ancestry? 5.4 Inorganic ions. 5.5 Amino acids. 5.5.1 Non-protein amino acids. 5.5.2 Nectar amino acids are under the control of environmental factors. 5.5.3 Contribution of amino acids to the taste of nectar. 5.6 Proteins. 5.6.1 Proteins in leek nectar. 5.6.2 Nectar redox cycle. 5.7 Other nectar constituents. 5.7.1 Lipids. 5.7.2 Organic acids. 5.7.3 Phenolics. 5.7.4 Alkaloids. 5.7.5 Terpenoids. 5.8 Conclusion. 6. Molecular Biology of the Nicotiana Floral Nectary R.W. Thornburg. 6.1 Introduction. 6.2 The ornamental tobacco nectary. 6.3 Developmental processes. 6.3.1 Origin of the floral nectary. 6.3.2 Conversion of chloroplasts into chromoplasts. 6.3.3 Filling of the nectary. 6.4 Protection of the gynoecium. 6.5 Gene expression. 6.5.1 Macroarray analysis indentifies defence genes. 6.5.1.1 Role of hydrogen peroxide in plant stress and defence. 6.5.1.2 Role of ascorbate in plant stress and defence. 6.5.2 EST analysis. 6.5.3 Nectary-specific gene expression. 6.6 Nectary molecular biology in other species. 6.6.1 Other n

Nectar biodiversity: a short review

Abstract. Nectaries differ in many aspects but a common feature is some kind of advantage for the plant conferred by foraging of consumers which may defend the plant from predators in the case of extrafloral nectaries, or be agents of pollination in the case of floral nectaries. This minireview is concerned mainly with floral nectaries and examines the following characteristics: position in flower; nectary structure; origin of carbohydrates, aminoacids and proteins; manner of exposure of nectar; site of nectar presentation; volume and production of nectar in time; sexual expression of flower and nectary morphology; nectar composition and floral sexual expression; variability of nectar composition; fate of nectar; energy cost of nectar production. The species of certain large families, such as Brassicaceae, Lamiaceae and Asteraceae, resemble each other in nectary organisation; other families, such as Cucurbitaceae and Ranunculaceae, have various types of organisation. A scheme is presented to illustrate factors influencing nectary and nectar biodiversity.

Sexual Reproduction in Higher Plants

OF POSTER PRESENTATIONS 459 AUTHOR INDEX ••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••• 493 SUBJECT INDEX •••••••••••••••••••••••••••.••••••••••••••••••••••••••••••••••• 497 Gene-Expression and Transcription DifTerential Gene-Expression During Microsporogenesis with Nicotiana tabacum J.A.M. Schrauwen, M.W.M. Derks, P.F.M. de Groot, M.M.A. van Herpen & G.J. Wullems Department of Experimental Botany Research Group Molecular Plant Physiology University of Nijmegen Toernooiveld NL-6525 ED Nijmegen The Netherlands INTRODUCTION W.H. Reijnen, 3 The changes in gene-expression during the development of the plant result in the formation of new transcription and translation products and in formation of other cell types. Pollen, which plays a key roll in the fertilization process, is a typical product of differential geneexpression occuring in a mature plant. The formation of pollen from vegetative tissue is concomitant with a series of processes by which type specific gene products, like mRNAs, are formed. These transcription products can be distinguished by specificity and function in relation to moment of activity and

Types and meaning of pollen carbohydrate reserves

During pollen development, soluble carbohydrates of sporophytic origin may be consumed immediately, polymerized to form starch reserves or intine, or transformed into other molecules. Disregarding intine, in mature pollen there are three different types of carbohydrates: (1) polysaccharides such as starch in amyloplasts or polysaccharides in cytoplasmic vesicles, (2) disaccharides such as sucrose and (3) monosaccharides such as glucose and fructose. At dispersal, pollen may be partly or slightly dehydrated, or not dehydrated at all. Partly dehydrated pollen has the capacity to lose or acquire water within limits without detriment to its viability. Slightly and non-dehydrated pollen is vulnerable to water loss and quickly becomes inviable. In partly dehydrated of pollen the carbohydrates consist of cytoplasmic polysacharides and sucrose; in slightly and non-dehydrated pollen these are absent or in low concentrations but there may be reserves of cytoplasmic callose. Starch, glucose and fructose are found in both types. It is postulated that cytoplasmic carbohydrates and sucrose are involved in protecting pollen viability during exposure and dispersal.

Lichens and higher plants on stone: a review

This review article deals with the site and colonization mode of plants and lichens on the wall of historical buildings, freezes and statues. Higher plants and lichen colonization is essentially conditioned by the adaptability of the species and the efficiency of their mode of reproduction. A list of the more common plants and lichens living on historical building is given as well as the type of damage. Nine niche types are distinguished for plants. The chemical and mechanical effects of plant colonization of walls, and the methods of conserving the aesthetic and functional integrity of the walls, are also presented and discussed.

Germination and early tube development in vitro of Lycopersicum peruvianum pollen: Ultrastructural features

Morphologic changes occurring during pollen grain activation and ultrastructural features of Lycopersicum peruvianum Mill. pollen tube during the first stages of growth in vitro have been studied. The more evident morphologic changes during activation, in comparison to those already described for mature inactive pollen, concern dictyosomes, rough endoplasmic reticulum (RER), and ribosomes. The dictyosomes are very abundant and produce “large” and “small” vesicles. Near the germinative pores both types of vesicles are present, while all along the remaining cell wall only the large type is observed. These latter react weakly to Thiérys test and probably contain a callose precursor necessary for the deposition of a callosic layer lining at first only the inner side of the functioning pore and occasionally the other two pores, and subsequently the entire inner surface of the cell wall. The small vesicles, highly positive to Thiérys test, are present only near the pores and could be involved in the formation of the pectocellulosic layer of the tube wall. The setting free of RER cisterns, which in the mature inactive pollen were aggregated in stacks, coinciding with polysome formation and resumption of protein synthesis, is in accord with the hypothesized role of RER cistern stacks as a reserve of synthesizing machinery. The pollen tube reaches a definitive spatial arrangement soon after the generative cell and vegetative nucleus have moved into it. At this stage four different zones that reflect a functional specialization are present. In the apical and subapical zone two types of dictysosome-originated vesicles, similar to those found in the activated pollen grain, are present. Their role in the formation of the callosic and pectocellulosic wall layers seems to be the same as in the activated pollen grain.

Pollen carbohydrates and water content during development, presentation, and dispersal: a short review.

Summary.Pollen accumulates starch reserves during development and the final stage of ripening. Before the anther opens, starch is totally or partially converted to pectins, glucose, fructose, sucrose, and to some unknown polysaccharides. Pollen is exposed to dispersing agents in an arrested developmental state which differs according to pollen water content. Pollen is classified as partially dehydrated or partially hydrated. The final water content may be reached before or after anther opening. Especially during exposure and dispersal, partially dehydrated pollen may interconvert soluble and insoluble reserves, modifying internal turgor pressure and hindering water loss or gain. Partially hydrated pollen is commonly devoid of mechanisms to conserve viability in time but has the advantage of quickly emitting pollen tubes on reaching the stigma.

Occurrence of mono- or disaccharides and polysaccharide reserves in mature pollen grains

Abstract Pollen from 13 species of gymnosperms and angiosperms was studied for soluble and insoluble carbohydrates at dispersal. Starch reserves stored during pollen development give rise to carbohydrates at maturity. Combinations of different types of carbohydrates in mature pollen may depend on the extent of starch hydrolysis. An inverse relationship was found between the extent of starch hydrolysis and sucrose content. If the starch was scarcely de-polymerized, the cytoplasm had very low levels of soluble sugars and none of the periodic acid-Schiff (PAS)-positive material as found in pollen not subject to high dehydration (Cucurbita pepo L., Zea mays L.). After total or partial starch hydrolysis, insoluble PAS-positive oligo/polysaccharides were found in the cytoplasm associated with much soluble sugar, and the pollen grains were dehydrated at dispersal as in Typha latifolia L., Chamaerops humilis L., Trachycarpus excelsa Wendl., and other specimens. Intermediate levels of starch and soluble sugars, together with cytoplasmic PAS-positive material, characterized species with dehydrated pollen such as Pinus halepensis Miller. Carbohydrates may be related to pollen longevity, which largely depends on the abundance of sucrose, which is known to protect membrane integrity. The relationship between PAS-positive material and pollen viability is unclear at present.

From anther and pollen ripening to pollen presentation

The events and processes occurring between pollen maturation, opening of the anther and presentation of pollen to dispersing agents are described. In the final phases of pollen development, starch is always stored; this occurs before the anther opens. Depending on the species, this starch may be totally or partially transformed into: (a) other types of polysaccharides (fructans and rarely callose); (b) disaccharides (sucrose); (c) monosaccharides (glucose and fructose, all situated in the cytoplasm. While awaiting dispersing agents and during dispersal, polysaccharides, especially fructans, and sucrose may be interconverted to control osmotic pressure and prevent loss and uptake of water. Opening of the anther is preceded by disappearance of the locular fluid and in many cases by partial dehydration of the pollen. Pollen generally has a water content between 5 and 50%. Pollen with a high water content may or may not be able to control water retention during pollen exposure and dispersal. Pollen may be dispersed in monads or grouped in pollen dispersing units by the following mechanisms: (i). tangling of filamentous pollen; (ii). adhesion by viscous substances (pollenkitt, tryphine, elastoviscin) derived from the tapetum; (iii). common walls. When the anther opens, the pollen may be dispersed immediately, remain until dispersed (primary presentation), or be presented to pollinators in another part of the flower (secondary presentation).