Academic literature on the topic 'Floral nectar spur'

Floral nectar spurs are hypothesized to have had a major role in the evolution of floral diversity and plant-pollinator coadaptation. We examined variation in the degree of nectar spur curvature in two species of jewelweed (Impatiens capensis and Impatiens pallida) pollinated by different sets of pollinators. To distinguish between adaptive and nonadaptive explanations for between-species differences in curvature, we determined the relationship between spur curvature and pollen removal, which is one estimate of male reproductive success. Spur curvature exhibited considerable variation both within and among three populations, with spur angles ranging from 0° to 297°. A greenhouse experiment determined that spur curvature of I. capensis flowers has a broad-sense heritability of 0.636. Laboratory experiments indicated that flowers having recurved spurs deposit significantly more pollen grains on hummingbird visitors than flowers having perpendicular spurs, apparently as a result of greater contact between the androecium of curve-spurred flowers and the upper bill of hummingbirds. We also found a significant relationship between spur curvature and flower length, suggesting a developmental link between the two traits. We discuss the degree of spur curvature in bird-pollinated I. capensis as a function of both adaptive evolution and developmental constraint.Key words: pollination, nectar spurs, hummingbirds, Impatiens capensis, Impatiens pallida, pollen export, floral morphology, floral polymorphisms.

The evolution of novel features, such as eyes or wings, that allow organisms to exploit their environment in new ways can lead to increased diversification rates. Therefore, understanding the genetic and developmental mechanisms involved in the origin of these key innovations has long been of interest to evolutionary biologists. In flowering plants, floral nectar spurs are a prime example of a key innovation, with the independent evolution of spurs associated with increased diversification rates in multiple angiosperm lineages due to their ability to promote reproductive isolation via pollinator specialization. As none of the traditional plant model taxa have nectar spurs, little is known about the genetic and developmental basis of this trait. Nectar spurs are a defining feature of the columbine genusAquilegia(Ranunculaceae), a lineage that has experienced a relatively recent and rapid radiation. We use a combination of genetic mapping, gene expression analyses, and functional assays to identify a gene crucial for nectar spur development,POPOVICH(POP), which encodes a C2H2 zinc-finger transcription factor.POPplays a central role in regulating cell proliferation in theAquilegiapetal during the early phase (phase I) of spur development and also appears to be necessary for the subsequent development of nectaries. The identification ofPOPopens up numerous avenues for continued scientific exploration, including further elucidating of the genetic pathway of which it is a part, determining its role in the initial evolution of theAquilegianectar spur, and examining its potential role in the subsequent evolution of diverse spur morphologies across the genus.

To date, the structure of the nectary spur of <i>Dendrobium finisterrae</i> has not been studied in detail, and the present paper compares the structural organization of the floral nectary in this species with the spurs of other taxa. The nectary spur of <i>D. finisterrae</i> was examined by means of light microscopy (LM), scanning electron microscopy (SEM), and transmission electron microscopy (TEM). It is composed of a single layer of secretory epidermis and several layers of small and compactly arranged subepidermal secretory cells. The secretory cells have thick cellulosic cell walls with primary pits. The secretory tissue is supplied by vascular bundles that run beneath in ground parenchyma and are additionally surrounded by strands of sclerenchymatous fibers. The flowers of the investigated species displayed morphological features characteristic of bee-pollinated taxa, as they are zygomorphic, creamy-green coloured with evident nectar guides. They also emit a weak but nice scent. However, they possess some characters attributed to bird-pollinated flowers such as a short, massive nectary spur and collenchymatous secretory tissue that closely resembles the one found in the nectaries of certain species that are thought to be bird-pollinated. This similarity in anatomical organization of the nectary, regardless of geographical distribution and phylogeny, strongly indicates convergence and appears to be related to pollinator-driven selection.

Epipogium aphyllumis a European-Asian obligatory mycoheterotrophic orchid containing no chlorophyll. Flowers are not resupinate with a sack-shape spur and cordate lip, which is divided into two parts: the basal (hypochile) and distal one (epichile). The floral analysis provides strong evidence to conclude that nectar is secreted on the upper surface of pink-coloured papillate ridges and epidermal (adaxial) cells at different place in spur, especially at the apex. The exudation on papillae has been observed through the entire anthesis and it has been stained on polysaccharides, proteins, and lipids. The dense cytoplasm of papillae contains profuse endoplasmic reticulum, plentiful vesicles (bigger ones with tannin-like materials), numerous mitochondria, sometimes dictyosomes, starch grains, and plastids with tubular structures. The large electron-dense bodies in cell walls are structurally the same as tannin-like materials from vesicles that are in contact with plasmalemma. The rupture of thin layer of swelled cuticle is caused by pressure of gathered substances exuded due to granulocrine secretion. The idioblasts with raphides occur mainly in tepals tissue. The dynamic changes of the nectar exudation, released through endocrine secretion, have been noticeable during the anthesis: both on the lip and inside the spur. The nectar secretion is not dependent on the colour form ofE. aphyllumblooming shoots. The floral biology and ultrastructure differ from mycoheterotrophic plants known up to date.

An outstanding feature of the orchid family is that approximately 30–40% of the species have non-rewarding flowers and deploy various modes of deception to attract pollinators, whereas the remaining species engage in pollination mutualisms based on provision of floral rewards. Here, we explore the direction, frequency and reversibility of transitions between deceptive and rewarding pollination systems in the radiation of the large African genus Disa , and test whether these transitions had consequences for diversification. By optimizing nectar production data for 111 species on a well-resolved phylogeny, we confirmed that floral deception was the ancestral condition and that nectar production evolved at least nine times and was lost at least once. Transitions to nectar production first occurred ca 17 million years ago but did not significantly affect either speciation or extinction rates. Nectar evolved independently of a spur, which was lost and gained multiple times. These results show that nectar production can be a highly labile trait and highlight the need for further studies of the genetic architecture of nectar production and the selective factors underlying transitions between deception and mutualism.

The anatomy and ultrastructure of floral nectary of <em>Platanthera bifolia</em> were studied. The epidermis inside the nectary spur showed characteristic features of secretory tissue. Many cells of this epidermis were protruded forming unicellular hairs. The protoplasts of secretory cells were characterized by few small vacuoles, a lot of mitochondria and leucoplasts, which stored starch before secretion. Numerous vesicles budded off from the endoplasmic reticulum and the Golgi apparatus were accumulated near plasmalemma and fused with it. This fact probably indicates that these structures are involved in secretory processes. Nectar was released onto the surface through the pores in a ruptured cuticle, which covered the walls of secretory hairs.

The structure of the floral nectaries of <i>Lonicera kamtschatica</i> was examined using light microscopy, scanning electron microscopy and transmission electron microscopy. Nectariferous tissues are located in the lower portion of the corolla tube. It was found that the secretory tissue of the nectary was composed of two layers of epidermal formations: short papillae and about 3x longer unicellular trichomes. They cover the adaxial surface of a small spur. Nectar secretion takes place through the apical portion of the trichomes and papillae. The cell wall of the upper part of the trichome has protuberances participating in nectar transfer to the subcuticular space which reaches large dimensions. The lateral walls of the trichomes are saturated with cutin. The papillae have much thicker walls than the trichomes. In the papillae, there are no wall protuberances. Less secretion accumulates in the subcuticular cavities of the papillae than in the trichomes.

Some of the most spectacular examples of coevolution between flowers and their pollinators are reflected in their morphologies. Tongue length of insects and bill lengths of nectar-feeding birds are some of the most significant characters in pollination studies (Kearns & Inouye 1993). As Darwin noted as early as 1862, the evolution of deep floral tubes or spurs and long tongues of flower visitors can be explained by runaway coevolution (Nilsson 1988). However, such a process has only been shown to be likely in a few cases, e.g. between Malagasy orchids and hawkmoths (Nilsson et al. 1985). The pollinator of the Malagasy orchid Angraecum sesquipedale, with a 30 cm nectar spur, is a sphingid moth with a tongue of a similar size (Darwin 1862, Nilsson 1998). However, most studies of such extreme pollination specialization also report that the interaction is asymmetrical, i.e. the pollinators interact with a guild of plants, whereas the plant often depends on only a few pollinators (Johnson & Steiner 1995).

A key question in biology is how changes in gene function or regulation produce new morphologies during evolution. The nectar spur is an evolutionarily labile structure known to influence speciation in a broad range of angiosperm taxa. Here, the genetic basis of nectar spur development, and the evolution of differences in nectar spur morphology, is investigated in Linaria vulgaris and two closely related species of orchid, the primitively longer-spurred Dactylorhiza fuchsii, and more derived short-spurred D. viridis (Orchidinae, Orchidaceae). Despite considerable morphological and phylogenetic differences, nectar spur ontogeny is fundamentally similar in each of the study species, proceeding from an abaxial bulge formed on the ventral petal relatively late in petal morphogenesis. However, spur development is progenetically curtailed in the short-spurred orchid D. viridis. In each case spur development involves class 1 KNOTTED1-like homeobox (KNOX) proteins. KNOX gene expression is not restricted to the spur-bearing petal, indicating that additional components are required to define nectar spur position, e.g. canonical ABC genes, determinants of floral zygomorphy, and additional (currently unknown) factors. However, constitutive expression of class 1 KNOX proteins in transgenic tobacco produces flowers with ectopic outgrowths on the petals, indicating that KNOX proteins alone are, to some degree, capable of inducing structures similar to nectar spurs in a heterologous host. Interestingly, KNOX gene expression is high in the ovary of all study taxa, suggesting that KNOX proteins may also have been involved in the evolution of this key angiosperm feature. Although principally involved in maintaining indeterminacy in the shoot apical meristem (SAM), members of the KNOX gene family have been co-opted in the evolution and development of compound leaves where they suppress differentiation and extend the morphogenetic potential of the leaf. A similar model is presented here to explain the role of KNOX proteins in nectar spur development. Co-option of KNOX gene expression to the maturing perianth delays cellular differentiation, facilitating the development of the nectar spur but requiring additional, unknown factors, to determine nectar spur fate. As facilitators of nectar spur development, changes in the spatio-temporal patterns of KNOX gene expression may alter the potential for nectar spur development and explain the critical length differences observed between the orchids D. fuchsii and D. viridis (and among other angiosperm taxa). Taken together, the available data indicate that KNOX genes confer a meristematic state upon plant tissues in a variety of morphogenetic contexts, making the gene family a potentially versatile tool to mediate a wide variety of evolutionary transformations.

The main objective of this research project was to study the growth and development of the floral nectar spur of Centranthus ruber (L.) DC. Nectar spurs are tubular floral outgrowths, generally derived from the perianth organs, which typically contain secreted floral nectar. The morphological characteristics of the spur, particularly the length, determine which floral visitors will be able to access the nectar reward pooled at the spur tip. Therefore, nectar spurs are ecologically important for the development of specialised pollinator interactions and have been demonstrated to act as key innovations in the evolution of some taxa. Morphological and anatomical characteristics of the spur and floral nectary were investigated using light and scanning electron microscopy. Ultrastructural features of the nectar spur, particularly the floral nectary within, were assessed using transmission electron microscopy. Nectar in C. ruber is produced by a trichomatous nectary which runs along the entire, inner abaxial surface of the spur. The nectary is aligned with the single vascular bundle which runs along the abaxial side of the spur, through the sub-nectary parenchyma, and back up the adaxial side. The secretory trichomes are unicellular and, in late development, they develop a thick layer of secondary wall ingrowths which vastly increases the surface area of the plasma membrane for nectar secretion. Elongate, non-secretory trichomes occupy the entire remaining circumference of the spur’s inner epidermis, but their density is reduced compared to the secretory trichomes. The cellular basis for spur growth is poorly characterized in the literature. Until recently, it was assumed that all nectar spurs grow by the constant production of new cells via up to three potential meristematic regions (the meristem hypothesis, Tepfer 1953). The cellular basis for spur growth in C. ruber was investigated by cell file counts and cell length and width measurements along the lateral side of nectar spurs in each of the developmental stages. DAPI stained spurs were also examined with Confocal/Apotome microscopy to determine the timing and position of cell division activity throughout spur development. It was determined that elongation of the spur epidermal cells contributes much more to spur growth than cell division. In early development, division is the primary driver of spur growth and the cells are isotropic. However, as development progresses, cell division activity slows down and the spur cells become increasingly anisotropic until anthesis. The patterns of nectar secretion were determined by assessing the volume, solute concentration and carbohydrate composition of the nectar throughout flowering phenology in two C. ruber plants. Nectar volumes and solute amounts rose initially, followed by an eventual decline in both as phenology progressed towards senescence. Because this study was conducted on greenhouse grown plants, it can be assumed that nectar was not removed by insects, suggesting that it is likely reabsorbed following secretion. High performance liquid chromatography (HPLC) analysis determined that C. ruber's nectar is sucrose dominant and that nectar composition remains stable following anthesis throughout floral phenology.