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1

Travers, Steven E., Ethan J. Temeles, and Irvin Pan. "The relationship between nectar spur curvature in jewelweed (Impatiens capensis) and pollen removal by hummingbird pollinators." Canadian Journal of Botany 81, no. 2 (February 1, 2003): 164–70. http://dx.doi.org/10.1139/b03-014.

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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.
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2

Ballerini, Evangeline S., Ya Min, Molly B. Edwards, Elena M. Kramer, and Scott A. Hodges. "POPOVICH, encoding a C2H2 zinc-finger transcription factor, plays a central role in the development of a key innovation, floral nectar spurs, inAquilegia." Proceedings of the National Academy of Sciences 117, no. 36 (August 26, 2020): 22552–60. http://dx.doi.org/10.1073/pnas.2006912117.

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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.
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3

Kamińska, Magdalena, and Małgorzata Stpiczyńska. "The structure of the spur nectary in Dendrobium finisterrae Schltr. (Dendrobiinae, Orchidaceae)." Acta Agrobotanica 64, no. 1 (2012): 19–26. http://dx.doi.org/10.5586/aa.2011.003.

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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.
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4

Święczkowska, Emilia, and Agnieszka K. Kowalkowska. "Floral Nectary Anatomy and Ultrastructure in Mycoheterotrophic Plant,Epipogium aphyllumSw. (Orchidaceae)." Scientific World Journal 2015 (2015): 1–11. http://dx.doi.org/10.1155/2015/201702.

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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.
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5

Johnson, Steven D., Nina Hobbhahn, and Benny Bytebier. "Ancestral deceit and labile evolution of nectar production in the African orchid genus Disa." Biology Letters 9, no. 5 (October 23, 2013): 20130500. http://dx.doi.org/10.1098/rsbl.2013.0500.

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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.
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6

Stpiczyńska, Małgorzata. "The structure of nectary of Platanthera bifolia L. Orchidaceae." Acta Societatis Botanicorum Poloniae 66, no. 1 (2014): 5–11. http://dx.doi.org/10.5586/asbp.1997.001.

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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.
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7

Weryszko-Chmielewska, Elżbieta, and Małgorzata Bożek. "Structure of trichomatous nectaries in flowers of Lonicera kamtschatica (Sevast.) Pojark." Acta Agrobotanica 61, no. 1 (2012): 13–26. http://dx.doi.org/10.5586/aa.2008.002.

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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.
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8

Antoń, Sebastian, and Magdalena Kamińska. "Comparative floral spur anatomy and nectar secretion in four representatives of Ranunculaceae." Protoplasma 252, no. 6 (March 15, 2015): 1587–601. http://dx.doi.org/10.1007/s00709-015-0794-5.

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9

LINDBERG, ANNIKA BÜCHERT, and JENS MOGENS OLESEN. "The fragility of extreme specialization: Passiflora mixta and its pollinating hummingbird Ensifera ensifera." Journal of Tropical Ecology 17, no. 2 (March 2001): 323–29. http://dx.doi.org/10.1017/s0266467401001213.

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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).
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10

Martínez‐Salazar, Sebastián, Favio González, Juan F. Alzate, and Natalia Pabón‐Mora. "Molecular framework underlying floral bilateral symmetry and nectar spur development in Tropaeolum , an atypical member of the Brassicales." American Journal of Botany 108, no. 8 (August 2021): 1315–30. http://dx.doi.org/10.1002/ajb2.1719.

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11

Singh, Shweta, Vacha Bhatt, Virender Kumar, Surbhi Kumawat, Praveen Khatri, Pankaj Singla, S. M. Shivaraj, et al. "Evolutionary Understanding of Aquaporin Transport System in the Basal Eudicot Model Species Aquilegia coerulea." Plants 9, no. 6 (June 26, 2020): 799. http://dx.doi.org/10.3390/plants9060799.

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Aquaporins (AQPs) play a pivotal role in the cellular transport of water and many other small solutes, influencing many physiological and developmental processes in plants. In the present study, extensive bioinformatics analysis of AQPs was performed in Aquilegia coerulea L., a model species belonging to basal eudicots, with a particular focus on understanding the AQPs role in the developing petal nectar spur. A total of 29 AQPs were identified in Aquilegia, and their phylogenetic analysis performed with previously reported AQPs from rice, poplar and Arabidopsis depicted five distinct subfamilies of AQPs. Interestingly, comparative analysis revealed the loss of an uncharacterized intrinsic protein II (XIP-II) group in Aquilegia. The absence of the entire XIP subfamily has been reported in several previous studies, however, the loss of a single clade within the XIP family has not been characterized. Furthermore, protein structure analysis of AQPs was performed to understand pore diversity, which is helpful for the prediction of solute specificity. Similarly, an AQP AqcNIP2-1 was identified in Aquilegia, predicted as a silicon influx transporter based on the presence of features such as the G-S-G-R aromatic arginine selectivity filter, the spacing between asparagine-proline-alanine (NPA) motifs and pore morphology. RNA-seq analysis showed a high expression of tonoplast intrinsic proteins (TIPs) and plasma membrane intrinsic proteins (PIPs) in the developing petal spur. The results presented here will be helpful in understanding the AQP evolution in Aquilegia and their expression regulation, particularly during floral development.
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12

Hodges, Scott A. "Floral Nectar Spurs and Diversification." International Journal of Plant Sciences 158, S6 (November 1997): S81—S88. http://dx.doi.org/10.1086/297508.

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13

Martins, D. J., and S. D. Johnson. "Hawkmoth pollination of aerangoid orchids in Kenya, with special reference to nectar sugar concentration gradients in the floral spurs." American Journal of Botany 94, no. 4 (April 1, 2007): 650–59. http://dx.doi.org/10.3732/ajb.94.4.650.

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14

Zhang, Wenliu, and Jiangyun Gao. "High fruit sets in a rewardless orchid: a case study of obligate agamospermy in Habenaria." Australian Journal of Botany 66, no. 2 (2018): 144. http://dx.doi.org/10.1071/bt17182.

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Low fruit set and pollination limitation are common characteristics of non-autogamous orchids, especially in rewardless species. The flowers of many Habenaria species are often characterised by long spurs and are mostly pollinated by long-tongued hawkmoths or butterflies. Unlike the flowers of other Habenaria species, the flowers of Habenaria malintana (Blanco) Merr. have very short spurs with no nectar or scent; however, this species is able to maintain high fecundity in south-west China. Breeding system experiments suggested that H. malintana is an obligate agamospermous orchid. Seed set did not need to be triggered by pollen grain deposition on stigmas, and ~100% fruit set was found in different populations and years. In pollen germination experiments, hand-deposited pollen failed to germinate on stigmas. The flowers of H. malintana failed to attract any pollinators, as we did not observe any floral visitors, and no pollinia removal or deposition occurred in both 2013 and 2014 at two study sites. These results strongly suggested that H. malintana has completely abandoned sexual reproduction and has adopted obligate agamospermy to achieve high reproductive output. We suggest that this strategy may have evolved to provide reproductive assurance and reduce the cost of flowers in response to unreliable pollinator service.
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15

Stpiczyńska, Małgorzata, Magdalena Kamińska, Kevin L. Davies, and Emerson R. Pansarin. "Nectar-Secreting and Nectarless Epidendrum: Structure of the Inner Floral Spur." Frontiers in Plant Science 9 (June 20, 2018). http://dx.doi.org/10.3389/fpls.2018.00840.

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16

Conway, Stephanie J., Cristina L. Walcher-Chevillet, Kate Salome Barbour, and Elena M. Kramer. "Brassinosteroids regulate petal spur length in Aquilegia by controlling cell elongation." Annals of Botany, September 11, 2021. http://dx.doi.org/10.1093/aob/mcab116.

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Abstract Background and Aims Aquilegia produce elongated, three-dimensional petal spurs that fill with nectar to attract pollinators. Previous studies have shown that the diversity of spur length across the Aquilegia genus is a key innovation that is tightly linked with its recent and rapid diversification into new ranges, and that evolution of increased spur lengths are achieved via anisotropic cell elongation. Previous work identified a brassinosteroid response transcription factor as being enriched in the early developing spur cup. Brassinosteroids (BRs) are known to be important for cell elongation, suggesting that brassinosteroid-mediated response may be an important regulator of spur elongation and potentially a driver of spur length diversity in Aquilegia. In this study, we investigated the role of brassinosteroids in the development of the Aquilegia coerulea petal spur. Methods We exogenously applied the biologically active BR brassinolide to developing petals spurs to investigate spur growth under high hormone conditions. We used virus induced gene silencing and gene expression experiments to understand the function of brassinosteroid-related transcription factors in Aquilegia coerulea petal spurs. Key Results We identified a total of three Aquilegia homologs of the BES1/BZR1 protein family and found that these genes are ubiquitously expressed in all floral tissues during development, yet consistent with the previous RNAseq study, we found that two of these paralogs are enriched in early developing petals. Exogenously applied brassinosteroid increased petal spur length due to increased anisotropic cell elongation as well as cell division. We found that targeting of the AqBEH genes with VIGS resulted in shortened petals, a phenotype caused in part by a loss of cell anisotropy. Conclusions Collectively, our results support a role for brassinosteroids in anisotropic cell expansion in Aquilegia petal spurs and highlight the BR pathway as a potential player in the diversification of petal spur length in Aquilegia.
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17

"Spurring plant diversification: are floral nectar spurs a key innovation?" Proceedings of the Royal Society of London. Series B: Biological Sciences 262, no. 1365 (December 22, 1995): 343–48. http://dx.doi.org/10.1098/rspb.1995.0215.

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