Hostname: page-component-8448b6f56d-tj2md Total loading time: 0 Render date: 2024-04-25T03:35:57.968Z Has data issue: false hasContentIssue false

Seed afterripening and germination photoinhibition in the genus Crocus (Iridaceae)

Published online by Cambridge University Press:  26 August 2015

Evangelia Skourti
Affiliation:
Department of Botany, Faculty of Biology, National and Kapodistrian University of Athens, Panepistimiopolis, Athens15784, Greece
Costas A. Thanos*
Affiliation:
Department of Botany, Faculty of Biology, National and Kapodistrian University of Athens, Panepistimiopolis, Athens15784, Greece
*
*Correspondence E-mail: cthanos@biol.uoa.gr

Abstract

Mediterranean characteristics are attributed to the genus Crocus, which is inadequately studied in terms of seed germination. An afterripening requirement is very common in environments with warm and dry periods, and photoinhibition has been detected in many angiosperms inhabiting dry and open areas. The effects of afterripening and light on seed germination were investigated for the first time in 23 native Greek Crocus taxa, collected from various localities with either a Mediterranean or a temperate climate. Germination experiments were conducted in continuous darkness and in light at the optimal temperature for each taxon, with both freshly collected and afterripened seeds; warm stratification (20°C, darkness) was also examined in 22 taxa. A number of selected taxa were additionally investigated with respect to afterripening outdoors, afterripening and warm stratification at higher temperatures (35 and 25°C, respectively), stratification at 20/10°C, dry storage at low temperatures, response to gibberellic acid and phenology of embryo growth. It was postulated that an afterripening requirement is a characteristic of the genus Crocus, and we found that it can be fulfilled in nature during the Mediterranean dry summer. Also, for the vast majority of the taxa, warm stratification and stratification at 20/10°C can both meet the afterripening requirement. Embryos of the taxa studied are underdeveloped and have to grow prior to germination. Intrageneric differences of seed germination were observed only towards light, with photoinhibition being predominant in taxa from drier environments.

Type
Research Papers
Copyright
Copyright © Cambridge University Press 2015 

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Allen, P.S. and Meyer, S.E. (1998) Ecological aspects of seed dormancy loss. Seed Science Research 8, 183191.Google Scholar
Antonidaki-Giatromanolaki, A., Dragassaki, M., Papadimitriou, M. and Vlahos, I. (2008) Effects of stratification, temperature and light on seed germination of Colchicum macrophyllum B.L. Burtt. Propagation of Ornamental Plants 8, 105107.Google Scholar
Ascough, G.D., Erwin, J.E. and Van Staden, J. (2007) Temperature-dependent seed germination in Watsonia species related to geographic distribution. South African Journal of Botany 73, 650653.CrossRefGoogle Scholar
Bair, N.B., Meyer, S.E. and Allen, P.S. (2006) A hydrothermal after-ripening time model for seed dormancy loss in Bromus tectorum L. Seed Science Research 16, 1728.CrossRefGoogle Scholar
Barbour, M.G. (1968) Germination requirements of the desert shrub Larrea divaricata . Ecology 49, 915923.Google Scholar
Baskin, C.C. and Baskin, J.M. (1985) The annual dormancy cycle in buried weed seeds: a continuum. BioScience 35, 492498.Google Scholar
Baskin, C.C. and Baskin, J.M. (1998) Ecology of seed dormancy and germination in grasses. pp. 3083 in Cheplick, G.P. (Ed.) Population biology of grasses. New York, Cambridge University Press.CrossRefGoogle Scholar
Baskin, C.C. and Baskin, J.M. (2007) A revision of Martin's seed classification system, with particular reference to his dwarf-seed type. Seed Science Research 17, 1120.Google Scholar
Baskin, C.C. and Baskin, J.M. (2014) Seeds: ecology, biogeography and evolution of dormancy and germination (2nd edition). San Diego, Academic Press.Google Scholar
Bazin, J., Batlla, D., Dussert, S., El-Maarouf-Bouteau, H. and Bailly, C. (2010) Role of relative humidity, temperature, and water status in dormancy alleviation of sunflower seeds during dry after-ripening. Journal of Experimental Botany 62, 627640.Google Scholar
Bell, D.T., Plummer, J.A. and Taylor, S.K. (1993) Seed germination ecology in southwestern Western Australia. The Botanical Review 59, 2473.CrossRefGoogle Scholar
Bell, D.T., Rockich, D.P., McChesney, C.J. and Plummer, J.A. (1995) Effects of temperature, light and gibberellic acid on the germination of seeds of 43 species native to Western Australia. Journal of Vegetation Science 6, 797806.Google Scholar
Bell, D.T., King, L.A. and Plummer, J.A. (1999) Ecophysiological effects of light quality and nitrate on seed germination in species from Western Australia. Australian Journal of Ecology 24, 210.Google Scholar
Bewley, J.D., Bradford, K.J., Hilhorst, H.W.M. and Nonogaki, H. (2013) Seeds: Physiology of development, germination and dormancy (3rd edition). New York, Springer.Google Scholar
Buide, M.L., Sánchez, J.M. and Guitián, J. (1998) Ecological characteristics of the flora of the Northwest Iberian Peninsula. Plant Ecology 135, 18.Google Scholar
Carasso, V., Hay, F.R., Probert, R.J. and Mucciarelli, M. (2011) Temperature control of seed germination in Fritillaria tubiformis subsp. moggridgei (Liliaceae) a rare endemic of the South-west Alps. Seed Science Research 21, 3338.Google Scholar
Carpenter, W.J., Wilfret, G.J. and Cornell, J.A. (1991) Temperature and relative humidity govern germination and storage of Gladiolus seed. HortScience 26, 10541057.Google Scholar
Carta, A., Bedini, G., Müller, J.V. and Probert, R.J. (2013) Comparative seed dormancy and germination of eight annual species of ephemeral wetland vegetation in a Mediterranean climate. Plant Ecology 214, 339349.Google Scholar
Carta, A., Probert, R., Moretti, M., Peruzzi, L. and Bedini, G. (2014) Seed dormancy and germination in three Crocus ser. Verni species (Iridaceae): implications for evolution of dormancy within the genus. Plant Biology 16, 10651074.Google ScholarPubMed
Côme, D., Corbineau, F., Footitt, S. and Holdworth, M. (2006) Dormancy breaking – temperature. pp. 211213 in Black, M.; Bewley, J.D.; Halmer, P. (Eds) The encyclopedia of seeds: Science, technology and uses. Wallingford, CAB International.Google Scholar
Conversa, G., Lazzizera, C. and Elia, A. (2010) Effects of after-ripening, stratification and GA3 on dormancy release and on germination of wild asparagus (Asparagus acutifolius L.) seeds. Scientia Horticulturae 125, 196202.CrossRefGoogle Scholar
Copete, E., Herranz, J.M., Ferrandis, P., Baskin, C.C. and Baskin, J.M. (2011) Physiology, morphology and phenology of seed dormancy break and germination in the endemic Iberian species Narcissus hispanicus (Amaryllidaceae). Annals of Botany 107, 10031016.Google Scholar
Copete, E., Herranz, J.M., Copete, M.A. and Ferrandis, P. (2014) Interpopulation variability on embryo growth, seed dormancy break, and germination in the endangered Iberian daffodil Narcissus eugeniae (Amaryllidaceae). Plant Species Biology 29, E72E84.Google Scholar
Corbineau, F., Neveur, N. and Côme, D. (1989) Characteristics of Cyclamen persicum Mill. seed germination. Acta Horticulturae 261, 337346.Google Scholar
Council of Europe (1979) Convention on the Conservation of European Wildlife and Natural Habitats. European Treaty Series, No. 104. Bern, Council of Europe. Google Scholar
Davies, T.J., Savolainen, V., Chase, M.W., Goldblatt, P. and Barraclough, T.G. (2005) Environment, area, and diversification in the species-rich flowering plant family Iridaceae. The American Naturalist 166, 418425.Google Scholar
Daws, M.I., Crabtree, L.M., Dalling, J.W., Mullins, C.E. and Burslem, D.F.R.P. (2008) Germination responses to water potential in neotropical pioneers suggest large-seeded species take more risks. Annals of Botany 102, 945951.CrossRefGoogle ScholarPubMed
Delipetrou, P. (1996) Ecophysiology of seed germination in maritime plants with emphasis on the action of light. PhD thesis, University of Athens, Greece (in Greek). Google Scholar
Del Vecchio, S., Mattana, E., Acosta, A.T. and Bacchetta, G. (2012) Seed germination responses to varying environmental conditions and provenances in Crucianella maritima L., a threatened coastal species. Comptes Rendus Biologies 335, 2631.CrossRefGoogle Scholar
De Vitis, M., Seal, C.E., Ulian, T., Pritchard, H.W., Magrini, S., Fabrini, G. and Mattana, E. (2014) Rapid adaptation of seed germination requirements of the threatened Mediterranean species Malcolmia littorea (Brassicaceae) and implications for its reintroduction. South African Journal of Botany 94, 4650.Google Scholar
Dimopoulos, P., Raus, T.H., Bergmeier, E., Constantinidis, T.H., Iatrou, G., Kokkini, S., Strid, A. and Tzanoudakis, D. (2013) Vascular plants of Greece: An annotated checklist. Berlin, Botanic Garden and Botanical Museum Berlin-Dahlem; Athens, Hellenic Botanical Society.Google Scholar
Doussi, M.A. and Thanos, C.A. (2002) Ecophysiology of seed germination in Mediterranean geophytes. 1: Muscari spp. Seed Science Research 12, 193201.Google Scholar
Esler, K.J., Rundel, P.W. and Voster, P. (1999) Biogeography of prostrate-leaved geophytes in semi-arid South Africa: hypotheses on functionality. Plant Ecology 142, 105120.Google Scholar
Fenner, M. and Thompson, K. (2005) The ecology of seeds. Cambridge, Cambridge University Press.Google Scholar
Finch-Savage, W.E. and Leubner-Metzger, G. (2006) Seed dormancy and the control of germination. New Phytologist 171, 501523.Google Scholar
Fournaraki, C. (2010) Conservation of threatened plants of Crete: seed ecology, operation and management of a gene bank. PhD thesis, University of Athens, Greece (in Greek). Google Scholar
Fu, Z., Tan, D., Baskin, C.C. and Baskin, J.M. (2013) Seed dormancy and germination of the subalpine geophyte Crocus alatavicus (Iridaceae). Australian Journal of Botany 61, 376382.Google Scholar
Geneve, L.R. (2005) Some common misconceptions about seed dormancy. Combined Proceedings International Plant Propagators' Society 55, 912.Google Scholar
Grime, J.P., Mason, G., Curtis, A.A., Rodman, J., Band, S.R., Mowforth, M.A.G., Neal, A.M. and Shaw, S. (1981) A comparative study of germination characteristics in a local flora. Journal of Ecology 69, 10171059.Google Scholar
Gutterman, Y. (2006) Deserts and arid lands. pp. 122127 in Black, M.; Bewley, J.D.; Halmer, P. (Eds) The encyclopedia of seeds: Science, technology and uses. Wallingford, CAB International.Google Scholar
Harpke, D., Meng, S., Rutten, T., Kerndorff, H. and Blattner, F.R. (2013) Phylogeny of Crocus (Iridaceae) based on one chloroplast and two nuclear loci: ancient hybridization and chromosome number evolution. Molecular Phylogenetics and Evolution 66, 617627.Google Scholar
Harpke, D., Peruzzi, L., Kerndorff, H., Karamplianis, T., Constantinidis, T., Randelović, V., Randelović, N., Jušković, M., Pasche, E. and Blattner, F.R. (2014) Phylogeny, geographic distribution, and new taxonomic circumscription of the Crocus reticulatus species group (Iridaceae). Turkish Journal of Botany 38, 11821198.Google Scholar
Hidayati, S.N., Walck, J.L., Merritt, D.J., Turner, S.R., Turner, D.W. and Dixon, K.W. (2012) Sympatric species of Hibbertia (Dilleniaceae) vary in dormancy break and germination requirements: implications for classifying morphophysiological dormancy in Mediterranean biomes. Annals of Botany 109, 11111123.Google Scholar
Hilton, J.R. (1984) The influence of temperature and moisture status on the photoinhibition of seed germination in Bromus sterilis L. by the far-red absorbing form of phytochrome. New Phytologist 97, 369374.Google Scholar
Hoffman, A.J., Liberona, F. and Hoffman, A.E. (1998) Distribution and ecology of geophytes in Chile. Conservation threats to geophytes in Mediterranean-type regions. pp. 231253 in Rundel, P.W.; Montenegro, G.; Jaksic, F.M. (Eds) Landscape disturbance and biodiversity in Mediterranean-type ecosystems. Berlin, Springer.Google Scholar
Holloway, P.S. (1987) Seed germination of Alaska Iris, Iris setosa ssp. interior. HortScience 22, 898899.Google Scholar
Hoyle, G.L., Daws, M.I., Steadman, K.J. and Adkins, S.W. (2008) Mimicking a semi-arid tropical environment achieves dormancy alleviation for seeds of Australian native Goodeniaceae and Asteraceae. Annals of Botany 101, 701708.Google Scholar
Iglesias-Fernández, R., Rodríguez-Gacio, M.C. and Matilla, A.J. (2011) Progress in research on dry afterripening. Seed Science Research 21, 6980.Google Scholar
ISTA (2013) International rules for seed testing (edition 2013). Bassersdorf, Switzerland, International Seed Testing Association. Google Scholar
Jankowska-Blaszczuk, M. and Daws, M.I. (2007) Impact of red:far red ratios on germination of temperate forest herbs in relation to shade tolerance, seed mass and persistence in the soil. Functional Ecology 21, 10551062.Google Scholar
Karlsson, L.M. and Milberg, P. (2008) Variation within species and inter-species comparison of seed dormancy and germination of four annual Lamium species. Flora 203, 409420.Google Scholar
Karlsson, L.M., Hidayati, S.N., Walck, J.L. and Milberg, P. (2005) Complex combination of seed dormancy and seedling development determine emergence of Viburnum tinus (Caprifoliaceae). Annals of Botany 95, 323330.Google Scholar
Kigel, J. (1995) Seed germination in arid and semiarid regions. pp. 645700 in Kigel, J.; Galili, G. (Eds) Seed development and germination. New York, Marcel Dekker.Google Scholar
Koller, D. (1957) Germination-regulating mechanisms in some desert seeds, IV. Atriplex dimorphostegia Kar. et Kir. Ecology 38, 213.Google Scholar
Koller, D., Sachs, M. and Negbi, M. (1964) Spectral sensitivity of seed germination in Artemisia monosperma . Plant and Cell Physiology 5, 7984.Google Scholar
Kondo, T., Miura, T., Okubo, N., Shimada, M., Baskin, C.C. and Baskin, J.M. (2004) Ecophysiology of deep simple epicotyl morphophysiological dormancy in seeds of Gagea lutea (Liliaceae). Seed Science Research 14, 371378.Google Scholar
Koutsovoulou, K., Daws, M.I. and Thanos, C.A. (2014) Campanulaceae: a family with small seeds that require light for germination. Annals of Botany 113, 135143.Google Scholar
Leishman, M.R., Wright, I.J., Moles, A.T. and Westoby, M. (2000) The evolutionary ecology of seed size. pp. 3157 in Fenner, M. (Ed.) Seeds: the ecology of regeneration in plant communities. Wallingford, CAB International.Google Scholar
Leopold, A.C., Glenister, R. and Cohn, M.A. (1988) Relationship between water content and afterripening in red rice. Physiologia Plantarum 74, 659662.Google Scholar
Leubner-Metzger, G. (2005) β-1,3-Glucanase gene expression in low-hydrated seeds as a mechanism for dormancy release during tobacco after-ripening. The Plant Journal 41, 133145.Google Scholar
Mamut, J., Tan, D.Y., Baskin, C.C. and Baskin, J.M. (2014) Intermediate complex morphophysiological dormancy in seeds of the cold desert sand dune geophyte Eremurus anisopterus (Xanthorrhoeaceae; Liliaceae s.l.). Annals of Botany 114, 991999.CrossRefGoogle ScholarPubMed
Marques, I. and Draper, D. (2012) Seed germination and longevity of autumn-flowering and autumn-seed producing Mediterranean geophytes. Seed Science Research 22, 299309.Google Scholar
Martin, A.C. (1946) The comparative internal morphology of seeds. American Midland Naturalist 36, 513660.Google Scholar
Mathew, B. (1982) The crocus: A revision of the genus Crocus (Iridaceae). Portland, Timber Press.Google Scholar
Milberg, P., Andersson, L. and Thompson, K. (2000) Large-seeded species are less dependent on light for germination than small-seeded ones. Seed Science Research 10, 99104.Google Scholar
Mira, S., González-Benito, M.E., Ibars, A.M. and Estrelles, E. (2011) Dormancy release and seed ageing in the endangered species Silene diclinis . Biodiversity and Conservation 20, 345358.Google Scholar
Mondoni, A., Probert, R., Rossi, G. and Hay, F. (2009) Habitat-related germination behaviour and emergence phenology in the woodland geophyte Anemone ranunculoides L. (Ranunculaceae) from northern Italy. Seed Science Research 19, 137144.Google Scholar
Mondoni, A., Orsenigo, S. and Rossi, G. (2013) Ecophysiology of embryo development and seed germination of the European woodland herbaceous perennial Corydalis cava (L.) Schweigg & Körte subsp. cava (Fumariaceae). Plant Species Biology 28, 215223.Google Scholar
Murdoch, A.J. and Ellis, R.H. (2000) Dormancy, viability and longevity. pp. 183214 in Fenner, M. (Ed.) Seeds: The ecology of regeneration in plant communities. Wallingford, CAB International.Google Scholar
Narbona, E., Delgado, A., Encina, F., Miguez, M. and Buide, M.L. (2013) Seed germination and seedling establishment of the rare Carex helodes Link depend on the proximity to water. Aquatic Botany 110, 5560.Google Scholar
Pérez-Fernández, M.A., Lamont, B.B., Marwick, A.L. and Lamont, W.G. (2000) Germination of seven exotic weeds and seven native species in south-western Australia under steady and fluctuating water supply. Acta Oecologica 21, 323336.Google Scholar
Petersen, G., Seberg, O., Thorsøe, S., Jørgensen, T. and Mathew, B. (2008) A phylogeny of the genus Crocus (Iridaceae) based on sequence data from five plastid regions. Taxon 57, 487499.Google Scholar
Phitos, D., Strid, A., Snogerup, S. and Greuter, W. (1995) The Red Data Book of rare and threatened plants of Greece. Athens, World Wide Fund for Nature.Google Scholar
Phitos, D., Constantinidis, T. and Kamari, G. (2009) The Red Data Book of rare and threatened plants of Greece. Patra, Hellenic Botanical Society.Google Scholar
Pons, T.L. (2000) Seed responses to light. pp. 237260 in Fenner, M. (Ed.) Seeds: The ecology of regeneration in plant communities. Wallingford, CAB International.Google Scholar
Probert, R.J. (2000) The role of temperature in the regulation of seed dormancy and germination. pp. 261292 in Fenner, M. (Ed.) Seeds: The ecology of regeneration in plant communities. Wallingford, CAB International.Google Scholar
Probert, R.J., Smith, R.D. and Birch, P. (1985) Germination responses to light and alternating temperatures in European populations of Dactylis glomerata L. New Pytologist 99, 305316.Google Scholar
Procheş, Ş., Cowling, R.M., Goldblatt, P., Manning, J.C. and Snijman, D.A. (2006) An overview of the Cape geophytes. Biological Journal of the Linnean Society 87, 2743.Google Scholar
Rukšāns, J. (2010) Crocuses: A complete guide to the genus. Portland, Timberpress.Google Scholar
Schütz, W. and Rave, G. (1999) The effect of cold stratification and light on the seed germination of temperate sedges (Carex) from various habitats and implications for regenerative strategies. Plant Ecology 144, 215230.Google Scholar
Schütz, W., Milberg, P. and Lamont, B.B. (2002) Seed dormancy, after-ripening and light requirements of four annual Asteraceae in south-western Australia. Annals of Botany 90, 707714.CrossRefGoogle ScholarPubMed
Skordilis, A. and Thanos, C.A. (1995) Seed stratification and germination strategy in the Mediterranean pines Pinus brutia and P . halepensis. Seed Science Research 5, 151160.Google Scholar
Skourti, E. and Thanos, C.A. (2013) Seed germination requirements in the genus Crocus . p. 93 in Zhimin, L.; Bo, L.; Jinlei, Z.; Qiaoling, Y. (Eds) Seed Ecology IV Conference Proceedings. Beijing, China Meteorological Press.Google Scholar
Smith, H. (2000) Phytochromes and light signal perception by plants – an emerging synthesis. Nature 407, 585591.Google Scholar
Stefani, A., Meletti, P., Sbrana, V. and Onnis, A. (2000) Low temperature storage of caryopses of Triticum durum: viability and longevity. Annals of Botany 85, 403406.Google Scholar
Swart, P.A., Kulkarni, M.G., Finnie, J.F. and Van Staden, J. (2011) Seed physiology of four African Romulea species. Seed Science and Technology 39, 354363.CrossRefGoogle Scholar
Thanos, C.A. (1993) Germination and the high irradiance reaction. pp. 187190 in Hendry, G.A.F.; Grime, J.P. (Eds) Methods in comparative plant ecology. A laboratory manual. London, Chapman & Hall.Google Scholar
Thanos, C.A., Georghiou, K. and Skarou, F. (1989) Glaucium flavum seed germination – an ecophysiological approach. Annals of Botany 63, 121130.Google Scholar
Thanos, C.A., Georghiou, K., Douma, D.J. and Marangaki, C.J. (1991) Photoinhibition of seed germination in Mediterranean maritime plants. Annals of Botany 68, 469475.Google Scholar
Thanos, C.A., Georghiou, K., Kadis, C. and Pantazi, C. (1992) Cistaceae: a plant family with hard seeds. Israel Journal of Botany 41, 251263.Google Scholar
Thanos, C.A., Georghiou, K. and Delipetrou, P. (1994) Photoinhibition of seed germination in the maritime plant Matthiola tricuspidata . Annals of Botany 73, 639644.Google Scholar
Thanos, C.A., Fournaraki, C. and Makri, N. (2005) Photoinhibition of seed germination ecology, phylogeny and presumptive mechanisms. p. 44 in Book of abstracts from the 8th International Workshop on Seeds, May 2005, Brisbane, Australia .Google Scholar
Tobe, K., Zhang, L. and Omasa, K. (2005) Seed germination and seedling emergence of three annuals growing on desert sand dunes in China. Annals of Botany 95, 649659.Google Scholar
Tuckett, R.E., Merritt, D.J., Hay, F.R., Hopper, S.D. and Dixon, K.W. (2010) Dormancy, germination and seed bank storage: a study in support of ex situ conservation of macrophytes of southwest Australian temporary pools. Freshwater Biology 55, 11181129.Google Scholar
Turner, S.R., Merritt, D.J., Ridley, E.C., Commander, L.E., Baskin, J.M., Baskin, C.C. and Dixon, K.W. (2006) Ecophysiology of seed dormancy in the Australian endemic species Acanthocarpus preissii (Dasypogonaceae). Annals of Botany 98, 11371144.Google Scholar
Turner, S.R., Merritt, D.J., Renton, M.S. and Dixon, K.W. (2009) Seed moisture content affects afterripening and smoke responsiveness in three sympatric Australian native species from fire-prone environments. Austral Ecology 34, 866877.Google Scholar
Vandelook, F. and Van Assche, J.A. (2008) Temperature requirements for seed germination and seedling development determine timing of seedling emergence of three monocotyledonous temperate forest spring geophytes. Annals of Botany 102, 865875.Google Scholar
Vandelook, F., Van de Moer, D. and Van Assche, J.A. (2008) Environmental signals for seed germination reflect habitat adaptations in four temperate Caryophyllaceae. Functional Ecology 22, 470478.Google Scholar
Vandelook, F., Verdú, M. and Honnay, O. (2012) The role of seed traits in determining the phylogenetic structure of temperate plant communities. Annals of Botany 110, 629636.Google Scholar
van der Pirl, L. (1969) Principles of dispersal in higher plants. Berlin, Springer-Verlag.Google Scholar
Veasey, E.A., Karasawa, M.G., Santos, P.P., Rosa, M.S., Mamani, E. and Oliveira, G.C.X. (2004) Variation in the loss of seed dormancy during after-ripening of wild and cultivated rice species. Annals of Botany 94, 875882.Google Scholar
Wang, J.H., Baskin, C.C., Chen, W. and Du, G.Z. (2010) Variation in seed germination between populations of five sub-alpine woody species from eastern Qinghai-Tibet Plateau following dry storage at low temperatures. Ecological Research 25, 195203.Google Scholar