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Parasitic Plants (Holoparasites and Holoparasite Hemiparasites) “ Scrophulariaceae” S.L

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Parasitic Plants (Holoparasites and Holoparasite Hemiparasites) “ Scrophulariaceae” S.L Phylogeny of Asterids Asterids Lamiids Ericales Campanulids Cornales Cornales Ericales Lamiids: Campanulids: Garryales Aquifoliales Gentianales Apiales Lamiales Dipsacales Solanales Asterales Lamiales contains about 20 family-level clades and ca. 22, 000 species. After APG, 2003; Judd and Olmstead, 2004, and Soltis et al., 2005 “ Scrophulariaceae” s.l. A. Mimulus lewisii C. Mimulus cardinalis B. M. lewisii x cardinalis (F1) D-L F2 offsprings Schemske and Bradshaw, 1999. Proc Natl Acad Sci USA 96(21): 11910-11915. “ Scrophulariaceae” s.l. A. Mimulus lewisii C. Mimulus cardinalis A single gene could produce a pollinator shift, and subsequently cause speciation. Note: the story on the evolutionary genetics of monkey flowers is purely for fun, won’t be on the exam Bradshaw and Schemske, 2003. Nature 426: 176-178 “ Scrophulariaceae” s.l. Photo: Yaowu Yuan Photo: Yaowu Yuan Castilleja sp. Mimulus alsinoides Photo: Yaowu Yuan Photo: Yaowu Yuan Collinsia sp. Penstemon sp. “ Scrophulariaceae” s.l. Textbook DVD KRR Penstemon digitalis Note the bilabiate corolla (usually 2 upper lobes and 3 lower lobes); and the didynamous Textbook DVD DLN Stamens –– with two pairs of stamens of Digitalis purpurea unqual length. “ Scrophulariaceae” s.l. Textbook DVD KRR Textbook DVD KRR & DLN Antirrhinum najus; snapdragon Again, note the bilabiate corolla and didynamous stamens Textbook DVD KRR Penstemon pallidus “ Scrophulariaceae” s.l. Photo: Yaowu Yuan Textbook DVD KRR Verbascum sp.; corolla Veronica arvensis; 4 corolla nearly anctinomorphic; 5 lobes, 2 stamens. stamens. A few genera do not follow the “bilabiate (zygomorphic) corolla, didynamous stamens” rule “ Scrophulariaceae” s.l. Photo: Yaowu Yuan Castilleja sp.; hemiparasite Textbook DVD JDS “Scrophulariaceae” s.l. includse a group Orobanche uniflora; of parasitic plants (holoparasites and holoparasite hemiparasites) “ Scrophulariaceae” s.l. Textbook DVD KRR Textbook DVD MHS Antirrhinum majus “Scrophulariaceae” s.l. fruit a many-seeded capsule Linaria macroccana “Scrophulariaceae” s.l. is polyphyletic This family always has been difficult to characterize by any explicit characters. It seems to have the generalized reproductive morphology of the large order Lamiales, which has ca. 20 families. Recently, Olmstead et al. has determined the family is polyphyletic and it is now being proposed that several families should be recognized. Scrophulariaceae s.s. Orobanchaceae Veronicaceae (or Plantaginaceae) Calceolariaceae Stilbaceae Phrymaceae Olmstead e et al., 2001. Am. J. Bot. 88(2): 348-361. “Scrophulariaceae” s.l. “Scrophulariaceae” s.l. - 269 genera/5100 species, but---- Herbs, shrubs, or small trees. Stem usually round, but sometimes square (e.g., Scrophularia). Leaves alternate or opposite. Flowers zygomorphic (or more rarely actinomorphic). Corolla often bilabiate. 4 stamens, didynamous (occasionally reduced to 2). Carpels 2, connate, ovary superior. Fruit a many-seeded capsule. “ Lamiaceae” (or Labiatae) Textbook DVD KRR Many members of this family have aromatic oils. These include many culinary herbs and ingredients in perfumes, including sage, oregano, marjoram, rosemary, thyme, basil, lavender. Lavendula stoechas “ Lamiaceae” (or Labiatae) Textbook DVD KRR & DLN Photo: Yaowu Yuan Dracocephalum moldavica Note the bilabiate corolla and Lamium purpureum didynamous stamens “ Lamiaceae” (or Labiatae) Textbook DVD KRR Textbook DVD KRR Teucrium canadense Phlomis russeliana Note the opposite (decussate –– square stems) leaf arrangement “ Lamiaceae” (or Labiatae) Photo: Yaowu Yuan Stachys cooleyae Note the 4 nutlets, similar to Boraginaceae Textbook DVD JRA Rosmarinus offcinalis “ Lamiaceae” (or Labiatae) Textbook DVD KRR & DLN Physostegia virginiana Note gynobasic style, similar to Boraginaceae Textbook DVD KRR “ Lamiaceae” (or Labiatae) “Lamiaceae” - 258 genera/6970 species Herbs, shrubs, or small trees. Often aromatic. Stem usually square. Leaves opposite and decussate. Flowers zygomorphic. Corolla bilabiate. 4 stamens, didynamous (occasionally reduced to 2). Carpels 2, connate, each with 2 ovules and a false septum between ovules of each carpel; with a gynobasic style arising from between 4 mericarps Fruit a schizocarp forming 4 nutlets. Summary of stamen arrangement Monadelphous Filaments fused into a tube - Malvaceae Diadelphous 9 fused at the filaments, 1 free - Fabaceae Tetradynamous 6 stamens, 4 long, 2 short - Brassicaceae Didynamous 4 stamens, 2 long, 2 short - Lamiaceae & Scrophulariaceae Connivent anthers anthers held together but not actually fused, surrounding the gynoecium - Violaceae & Solanaceae (Solanum) Parasitic Plants Textbook DVD WSJ Monotropa uniflora Autotrophic: capable of feeding oneself; For plants, ones that are photosynthetic. Heterotrophic: Referring to oraganisms that do not produce their own food (nonphotosynthetic) and which must then obtain organic compounds from exogenous sources (such as another plant). Many forms Textbook DVD KRR of heterotrophism exist such as saprophytism, Monotropa hypopithys myco-heterotropism, and parasitism. Parasitic Plants Note: All materials on parasitic plants are from the website www.parasiticplants.siu.edu (presented by Dan Nickrent), unless otherwise indicated. Parasitic Plants Haustorium: The morphologically modified root which physically connects the parasite to the host. Parasitic Plants http://www.cas.vanderbilt.edu/bioimages/p/wphle14fr30464.jpg http://www.cas.vanderbilt.edu/bioimages/image /p/phle14wpin-fruit-and-flower56193.htm Phoradendron flavescens; Christmas mistletoe Hemiparasite: A parasite that is photosynthetic (during at least one stage of its life cycle) that obtains water and nutrient from the host xylem. Holoparasite: A nonphotosynthetic parasite that obtains water and nutrients as well as photosynthates from the host. Parasitic Plants Obligate parasite: A plant that must attach to a host to complete its life cycle. All holoparasites are obigate whereas only some hemiparasites are obligate. Facultative parasite: one that does not require a host to complete its life cycle. Note, however, that in nature, parasitism is nearly always observed. Parasitic Plants Parasitic Reduction Syndrome (becoming holoparasites): Textbook DVD WSJ Loss of leaves - leaves reduced to scales; Small overall size of the plant - no need for large plants to hold leaves; Loss of chlorophyll; Loss of roots - reduced to short, stumpy projections with haustoria; Loss of genes needed for photosynthesis; Rapid DNA divergence in genes that are not lost; Parasitic Plants Wolfe et al., 1992. Proc Natl Acad Sci USA 89(22): 10648-10652 Wakasugi e et al., 1998. Pl. Mol. Biol. Reporter. 16: 231-241 Note the size of the parasitic plant chloroplast genome is much smaller than tobacco’s –– most genes needed for photosynthesis are lost from the parasite. Parasitic Plants Note the rapid divergence of chloroplast genes of parasites compared with their non- parasitic relatives Stefanovic e et al., 2002. Am. J. Bot. 89(9): 1510-1522. Parasitic Plants Parasitism have evolved perhaps 20 times among flowering plants. In each case the plants have become parasitic on either other plants or fungi. These include 4 families we know: Broginaceae, Scrophulariaceae, Ericaceae, and Orchidaceae..
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