Supergene Evolution Via Stepwise Duplications and Neofunctionalization of a Floral-Organ Identity Gene

Total Page:16

File Type:pdf, Size:1020Kb

Supergene Evolution Via Stepwise Duplications and Neofunctionalization of a Floral-Organ Identity Gene Supergene evolution via stepwise duplications and neofunctionalization of a floral-organ identity gene Cuong Nguyen Huua, Barbara Kellerb, Elena Contib, Christian Kappela, and Michael Lenharda,1 aInstitute for Biochemistry and Biology, University of Potsdam, D-14476 Potsdam-Golm, Germany; and bDepartment of Systematic and Evolutionary Botany, University of Zurich, CH-8008 Zurich, Switzerland Edited by June B. Nasrallah, Cornell University, Ithaca, NY, and approved August 5, 2020 (received for review April 7, 2020) Heterostyly represents a fascinating adaptation to promote out- addition, self-incompatibility is present in many distylous species breeding in plants that evolved multiple times independently. (9). In all examined cases, distyly is under simple Mendelian While L-morph individuals form flowers with long styles, short genetic control, with a dominant and a recessive haplotype at the anthers, and small pollen grains, S-morph individuals have flowers single S locus determining the morphs (10). With a few excep- with short styles, long anthers, and large pollen grains. The differ- tions, the dominant S haplotype causes S-morph and the reces- ence between the morphs is controlled by an S-locus “supergene” sive s haplotype L-morph flowers. Rather than a single gene, the consisting of several distinct genes that determine different traits S locus represents a supergene, that is, a chromosomal region of the syndrome and are held together, because recombination with at least two tightly linked genes that determine the different between them is suppressed. In Primula, the S locus is a roughly aspects of a coadapted set of phenotypes (10–12). Similar su- 300-kb hemizygous region containing five predicted genes. How- pergenes underlie many fascinating coadapted phenotypes in ever, with one exception, their roles remain unclear, as does the plants and animals (13–16), and their evolutionary buildup is evolutionary buildup of the S locus. Here we demonstrate that the often only incompletely understood. MADS-box GLOBOSA2 (GLO2) gene at the S locus determines an- The primrose genus Primula is the most studied model for ther position. In Primula forbesii S-morph plants, GLO2 promotes distyly (17). Primula flowers show reciprocal herkogamy, pollen growth by cell expansion in the fused tube of petals and stamen polymorphism, and self-incompatibility, with the dominant S and filaments beneath the anther insertion point; by contrast, neither recessive s haplotypes producing S- and L-morph flowers, re- GLO2 pollen size nor male incompatibility is affected by activity. spectively (3, 18, 19). Based on classical genetic studies, the S GLO1 GLO2 The paralogue , from which arose by duplication, has locus supergene has been proposed to contain at least three maintained the ancestral B-class function in specifying petal and separable individual loci: G, controlling style length; A, deter- GLO2 stamen identity, indicating that underwent neofunctionali- mining anther position; and P, controlling pollen size and zation, likely at the level of the encoded protein. Genetic mapping number (10). Style length and female-compatibility type have and phylogenetic analysis indicate that the duplications giving rise CYP734A50 GLO2 never been genetically separated (20, 21); by contrast, male to the style-length-determining gene and to oc- compatibility is likely controlled by a locus distinct from A and P curred sequentially, with the CYP734A50 duplication likely the (21, 22). At the molecular level, the dominant S-locus haplotype first. Together these results provide the most detailed insight into represents a roughly 280-kb-long genomic segment that is the assembly of a plant supergene yet and have important impli- hemizygous; specifically, it is present only on the chromosome cations for the evolution of heterostyly. with the S haplotype but missing from the chromosome with the recessive s haplotype (23–25). Five genes have been predicted to heterostyly | Primula | supergene | gene duplication | neofunctionalization Significance lowering plants have evolved many different adaptations to Fpromote efficient cross-pollination (1, 2). Some of the most fascinating of these are found in the group of stylar polymor- Heterostyly is an adaptation to promote outbreeding in plants. phisms that often combine reciprocal herkogamy with self- In heterostylous primroses, plants form flowers either with incompatibility (1). Individuals of species with stylar polymor- long styles and low anthers or with short styles and high an- phism fall into two or three classes, or morphs, that differ in their thers. This difference is due to a chromosomal segment con- taining five predicted genes, yet their roles and the evolution arrangement of male and female reproductive organs. In par- of this segment remain unclear. Here we identify the gene ticular, male and female organs are physically separated within a responsible for raising the anthers in short-styled flowers. This flower, and this herkogamy is reciprocal between the morphs in gene arose by duplication from a classical floral-organ identity most species, with anthers in one morph located in the analogous gene and gained a novel function. Surprisingly, the responsible position to the style in the other, and vice versa. As a result, pollen chromosomal segment appears to have evolved by stepwise from the different morphs is deposited in a spatially segregated gene duplications rather than duplication of an entire chro- manner on the bodies of pollinators (3), resulting in preferential mosomal block. These findings thus provide detailed insight cross-pollination between the morphs and preventing pollen into the evolution of complex polymorphisms involving dif- wastage and the formation of less fit inbred progeny. Often, re- ferent individual traits. ciprocal herkogamy is complemented by self-incompatibility to prevent self-fertilization; with only two or three compatibility types Author contributions: C.N.H., B.K., E.C., C.K., and M.L. designed research; C.N.H., B.K., and corresponding to the different morphs this manifests as an intra- C.K. performed research; C.N.H., B.K., C.K., and M.L. analyzed data; and M.L. wrote morph incompatibility (4, 5). the paper. An intensively studied stylar polymorphism is heterostyly The authors declare no competing interest. (6–8). In the simplest case, distyly, individuals either form This article is a PNAS Direct Submission. flowers with short styles and high anthers (S-morph) or flowers Published under the PNAS license. with long styles and low anthers (L-morph). Additional poly- 1To whom correspondence may be addressed. Email: [email protected]. morphisms can be associated with this reciprocal herkogamy; for This article contains supporting information online at https://www.pnas.org/lookup/suppl/ example, in many species S-morph flowers produce larger, but doi:10.1073/pnas.2006296117/-/DCSupplemental. fewer, pollen grains compared to L-morph flowers (9). In First published August 31, 2020. 23148–23157 | PNAS | September 15, 2020 | vol. 117 | no. 37 www.pnas.org/cgi/doi/10.1073/pnas.2006296117 Downloaded by guest on September 27, 2021 reside in this 280-kb segment based on its sequence in Primula brassinosteroids, a class of phytohormones promoting cell elon- vulgaris and the closely related Primula veris (25). This archi- gation, and thus suppresses style elongation in S-morph flowers. tecture of the S locus with a large hemizygous region appears to Despite this important progress, a number of key questions be conserved in five species belonging to different clades of the remain unanswered. One concerns the identity of the other two genus Primula (25), yet whether all five predicted genes are genetically defined loci, A and P, which determine anther posi- present at the S locus throughout the different Primula species is tion and pollen size, respectively. The GLO2 gene represents a unknown. The five genes predicted in P. vulgaris and P. veris are plausible candidate for the A locus; in fact, a transposon inser- tion in GLO2 of P. vulgaris has been reported to result in a short- CYP734A50, encoding a cytochrome P450; GLOBOSA2 (also homostyle phenotype (23). However, no description of the mutant known as GLOT), coding for a B-class homoeotic MADS- phenotype has been provided, nor has this finding been confirmed domain transcription factor; a KELCH F-BOX PROTEIN by an independent allele. Thus, the identity of the GLO2 gene (KFBT) gene; a gene encoding a PUMILIO RNA-binding pro- T with the A locus remains to be firmly established. As for P,no tein (PUM ); and a gene encoding a CONSERVED C-TER- information is currently available about its molecular nature. T MINAL DOMAIN (CCM ) protein. [Naming of the genes The second major unresolved question concerns the evolu- follows priority regarding their first publications (23, 26, 27).] Of tionary buildup of the S locus, both regarding its chromosomal these, CYP734A50 has been shown to correspond to the G locus assembly and the sequence of trait changes. Clear paralogues of (26). The encoded cytochrome P450 monooxygenase inactivates CYP734A50, GLO2, and CCMT are found outside of the S locus, A B C ns ns GLO1 GLO2 ns *** PLANT BIOLOGY Relative expression of Relative expression of GLO2 GLO2 GLO2 GLO2 L-morph L-morph S-morph S-morph S-morph S-morph L-morph L-morph VIGS-GLO2 VIGS-GLO2 L-morph VIGS- L-morph VIGS- S-morph VIGS- S-morph VIGS- DFEGH ns *** 6.0 2.5 40 * 4.0 * *** *** 2.0 30 *** *** 30 3.0 4.0 *** 1.5 *** *** 20 2.0 1.0 20 2.0 Anther height (mm) 1.0 0.5 10 Length of corolla tube (mm) 10 Cell length (below anther) (µm) Cell length (above anther) (µm) Diameter of corolla mouth mm) 0.0 0.0 0.0 GLO2 GLO2 GLO2 GLO2 GLO2 L-morph L-morph L-morph S-morph S-morph L-morph S-morph S-morph L-morph S-morph S-morph VIGS- S-morph VIGS- S-morph VIGS- S-morph VIGS- S-morph VIGS- Fig.
Recommended publications
  • Outline of Angiosperm Phylogeny
    Outline of angiosperm phylogeny: orders, families, and representative genera with emphasis on Oregon native plants Priscilla Spears December 2013 The following listing gives an introduction to the phylogenetic classification of the flowering plants that has emerged in recent decades, and which is based on nucleic acid sequences as well as morphological and developmental data. This listing emphasizes temperate families of the Northern Hemisphere and is meant as an overview with examples of Oregon native plants. It includes many exotic genera that are grown in Oregon as ornamentals plus other plants of interest worldwide. The genera that are Oregon natives are printed in a blue font. Genera that are exotics are shown in black, however genera in blue may also contain non-native species. Names separated by a slash are alternatives or else the nomenclature is in flux. When several genera have the same common name, the names are separated by commas. The order of the family names is from the linear listing of families in the APG III report. For further information, see the references on the last page. Basal Angiosperms (ANITA grade) Amborellales Amborellaceae, sole family, the earliest branch of flowering plants, a shrub native to New Caledonia – Amborella Nymphaeales Hydatellaceae – aquatics from Australasia, previously classified as a grass Cabombaceae (water shield – Brasenia, fanwort – Cabomba) Nymphaeaceae (water lilies – Nymphaea; pond lilies – Nuphar) Austrobaileyales Schisandraceae (wild sarsaparilla, star vine – Schisandra; Japanese
    [Show full text]
  • An Additional Nomenclatural Transfer in the Pantropical Genus Myrsine (Primulaceae: Myrsinoideae) John J
    Nova Southeastern University NSUWorks Marine & Environmental Sciences Faculty Articles Department of Marine and Environmental Sciences 9-13-2018 An Additional Nomenclatural Transfer in the Pantropical Genus Myrsine (Primulaceae: Myrsinoideae) John J. Pipoly III Broward County Parks & Recreation Division; Nova Southeastern University, [email protected] Jon M. Ricketson Missouri Botanical Garden Find out more information about Nova Southeastern University and the Halmos College of Natural Sciences and Oceanography. Follow this and additional works at: https://nsuworks.nova.edu/occ_facarticles Part of the Marine Biology Commons, and the Oceanography and Atmospheric Sciences and Meteorology Commons NSUWorks Citation John J. Pipoly III and Jon M. Ricketson. 2018. An Additional Nomenclatural Transfer in the Pantropical Genus Myrsine (Primulaceae: Myrsinoideae) .Novon , (3) : 287 -287. https://nsuworks.nova.edu/occ_facarticles/942. This Article is brought to you for free and open access by the Department of Marine and Environmental Sciences at NSUWorks. It has been accepted for inclusion in Marine & Environmental Sciences Faculty Articles by an authorized administrator of NSUWorks. For more information, please contact [email protected]. An Additional Nomenclatural Transfer in the Pantropical Genus Myrsine (Primulaceae: Myrsinoideae) John J. Pipoly III Broward County Parks & Recreation Division, 950 NW 38th St., Oakland Park, Florida 33309, U.S.A.; Nova Southeastern University, 8000 N Ocean Dr., Dania Beach, Florida 33004, U.S.A. [email protected]; [email protected]; [email protected] Jon M. Ricketson Missouri Botanical Garden, 4344 Shaw Blvd., St. Louis, Missouri 63110, U.S.A. [email protected] ABSTRACT. Rapanea pellucidostriata Gilg & Schellenb. TYPE: Democratic Republic of the Congo. Ruwen- is transferred to Myrsine L.
    [Show full text]
  • Doctorat De L'université De Toulouse
    En vue de l’obt ention du DOCTORAT DE L’UNIVERSITÉ DE TOULOUSE Délivré par : Université Toulouse 3 Paul Sabatier (UT3 Paul Sabatier) Discipline ou spécialité : Ecologie, Biodiversité et Evolution Présentée et soutenue par : Joeri STRIJK le : 12 / 02 / 2010 Titre : Species diversification and differentiation in the Madagascar and Indian Ocean Islands Biodiversity Hotspot JURY Jérôme CHAVE, Directeur de Recherches CNRS Toulouse Emmanuel DOUZERY, Professeur à l'Université de Montpellier II Porter LOWRY II, Curator Missouri Botanical Garden Frédéric MEDAIL, Professeur à l'Université Paul Cezanne Aix-Marseille Christophe THEBAUD, Professeur à l'Université Paul Sabatier Ecole doctorale : Sciences Ecologiques, Vétérinaires, Agronomiques et Bioingénieries (SEVAB) Unité de recherche : UMR 5174 CNRS-UPS Evolution & Diversité Biologique Directeur(s) de Thèse : Christophe THEBAUD Rapporteurs : Emmanuel DOUZERY, Professeur à l'Université de Montpellier II Porter LOWRY II, Curator Missouri Botanical Garden Contents. CONTENTS CHAPTER 1. General Introduction 2 PART I: ASTERACEAE CHAPTER 2. Multiple evolutionary radiations and phenotypic convergence in polyphyletic Indian Ocean Daisy Trees (Psiadia, Asteraceae) (in preparation for BMC Evolutionary Biology) 14 CHAPTER 3. Taxonomic rearrangements within Indian Ocean Daisy Trees (Psiadia, Asteraceae) and the resurrection of Frappieria (in preparation for Taxon) 34 PART II: MYRSINACEAE CHAPTER 4. Phylogenetics of the Mascarene endemic genus Badula relative to its Madagascan ally Oncostemum (Myrsinaceae) (accepted in Botanical Journal of the Linnean Society) 43 CHAPTER 5. Timing and tempo of evolutionary diversification in Myrsinaceae: Badula and Oncostemum in the Indian Ocean Island Biodiversity Hotspot (in preparation for BMC Evolutionary Biology) 54 PART III: MONIMIACEAE CHAPTER 6. Biogeography of the Monimiaceae (Laurales): a role for East Gondwana and long distance dispersal, but not West Gondwana (accepted in Journal of Biogeography) 72 CHAPTER 7 General Discussion 86 REFERENCES 91 i Contents.
    [Show full text]
  • Bioactive Phenolic Compounds from Primula Veris L.: Influence of the Extraction Conditions and Purification
    molecules Article Bioactive Phenolic Compounds from Primula veris L.: Influence of the Extraction Conditions and Purification Maria Tarapatskyy 1,* , Aleksandra Gumienna 1, Patrycja Sowa 1 , Ireneusz Kapusta 2 and Czesław Puchalski 1 1 Department of Bioenergetics, Food Analysis and Microbiology, Institute of Food Technology and Nutrition, College of Natural Sciences, University of Rzeszów, 35-601 Rzeszów, Poland; [email protected] (A.G.); [email protected] (P.S.); [email protected] (C.P.) 2 Department of Food Technology and Human Nutrition, Institute of Food Technology and Nutrition, College of Natural Sciences, University of Rzeszów, 35-601 Rzeszów, Poland; [email protected] * Correspondence: [email protected]; Tel.: +48-17-7854834 Abstract: Our experiments may help to answer the question of whether cowslip (Primula veris L.) is a rich source of bioactive substances that can be obtained by efficient extraction with potential use as a food additive. A hypothesis assumed that the type of solvent used for plant extraction and the individual morphological parts of Primula veris L. used for the preparation of herbal extracts will have key impacts on the efficiency of the extraction of bioactive compounds, and thus, the health- promoting quality of plant concentrates produced. Most analysis of such polyphenolic compound contents in extracts from Primula veris L. has been performed by using chromatography methods such as ultra-performance reverse-phase liquid chromatography (UPLC−PDA−MS/MS). Experiments ◦ demonstrated that the most effective extraction agent for fresh study material was water at 100 C, whereas for dried material it was 70% ethanol. The richest sources of polyphenolic compounds Citation: Tarapatskyy, M.; were found in cowslip primrose flowers and leaves.
    [Show full text]
  • Assessment Report on Primula Veris L. And/Or Primula Elatior (L.) Hill, Radix
    19 September 2012 EMA/HMPC/113577/2012 Committee on Herbal Medicinal Products (HMPC) Assessment report on Primula veris L. and/or Primula elatior (L.) Hill, radix Based on Article 16d(1), Article 16f and Article 16h of Directive 2001/83/EC as amended (traditional use) Final Herbal substance(s) (binomial scientific Primula veris L., Primula elatior (L.) Hill, radix name of the plant, including plant part) Herbal preparation(s) A) Dry extract (DER 3-9:1), extraction solvent ethanol 40-50% v/v B) Liquid extract (DER 1:1), extraction solvent ethanol 70% v/v C) Liquid extract (DER 1:2.0-2.5), extraction solvent ethanol 70% v/v D) Tincture (ratio of herbal substance to extraction solvent 1:5), extraction solvent ethanol 70% v/v E) Soft extract (DER 5-10:1), extraction solvent water F) Soft extract (DER 1-4:1), extraction solvent ethanol 20-55% v/v G) Soft extract (DER 6-10:1), extraction solvent methanol, water, ammonia solution 10% (50.0:49.5:0.5) H) Soft extract (DER 6-10:1), extraction solvent methanol 50% I) Comminuted herbal substance Pharmaceutical form(s) Comminuted herbal substance as herbal tea for oral use. Other herbal preparations in liquid and solid dosage forms for oral use. Rapporteur R. Länger Assessor(s) R. Länger 7 Westferry Circus ● Canary Wharf ● London E14 4HB ● United Kingdom Telephone +44 (0)20 7418 8400 Facsimile +44 (0)20 7523 7051 E -mail [email protected] Website www.ema.europa.eu An agency of the European Union © European Medicines Agency, 2013.
    [Show full text]
  • Evolutionary Consequences of Dioecy in Angiosperms: the Effects of Breeding System on Speciation and Extinction Rates
    EVOLUTIONARY CONSEQUENCES OF DIOECY IN ANGIOSPERMS: THE EFFECTS OF BREEDING SYSTEM ON SPECIATION AND EXTINCTION RATES by JANA C. HEILBUTH B.Sc, Simon Fraser University, 1996 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in THE FACULTY OF GRADUATE STUDIES (Department of Zoology) We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA July 2001 © Jana Heilbuth, 2001 Wednesday, April 25, 2001 UBC Special Collections - Thesis Authorisation Form Page: 1 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. The University of British Columbia Vancouver, Canada http://www.library.ubc.ca/spcoll/thesauth.html ABSTRACT Dioecy, the breeding system with male and female function on separate individuals, may affect the ability of a lineage to avoid extinction or speciate. Dioecy is a rare breeding system among the angiosperms (approximately 6% of all flowering plants) while hermaphroditism (having male and female function present within each flower) is predominant. Dioecious angiosperms may be rare because the transitions to dioecy have been recent or because dioecious angiosperms experience decreased diversification rates (speciation minus extinction) compared to plants with other breeding systems.
    [Show full text]
  • Allopatric Speciation with Little Niche Divergence Is Common Among
    Journal of Biogeography (J. Biogeogr.) (2016) 43, 591–602 ORIGINAL Allopatric speciation with little niche ARTICLE divergence is common among alpine Primulaceae Florian C. Boucher1*, Niklaus E. Zimmermann2,3 and Elena Conti1 1Institute of Systematic Botany, University of ABSTRACT Zurich,€ 8008 Zurich,€ Switzerland, 2Dynamic Aim Despite the accumulation of cases describing fast radiations of alpine Macroecology, Swiss Federal Research plants, we still have limited understanding of the drivers of speciation in alpine Institute WSL, 8903 Birmensdorf, Switzerland, 3Department of Environmental floras and of the precise the timing of their diversification. Here, we investi- Systems Science, Swiss Federal Institute of gated spatial and temporal patterns of speciation in three groups of alpine Technology ETH, CH-8092 Zurich,€ Primulaceae. Switzerland Location Mountains of the European Alpine System. Methods We built a new phylogeny of Primulaceae including all species in three focal groups: Androsace sect. Aretia, Primula sect. Auricula and Soldanella. Combining phylogenetic information with a detailed climatic data set, we investigated patterns of range and ecological overlap between sister-species using an approach that takes phylogenetic uncertainty into account. Finally, we investigated temporal trajectories of diversification in the three focal groups. Results We found that a large majority of sister-species pairs in the three groups are strictly allopatric and show little differences in substrate and cli- matic preferences, a result that was robust to phylogenetic uncertainty. While rates of diversification have remained constant in Soldanella, both Androsace sect. Aretia and Primula sect. Auricula showed decreased diversification rates in the Pleistocene compared to previous geological epochs. Main conclusions Allopatric speciation with little niche divergence appears to have been by far the most common mode of speciation across the three groups studied.
    [Show full text]
  • From New Guinea
    Phytotaxa 442 (3): 133–137 ISSN 1179-3155 (print edition) https://www.mapress.com/j/pt/ PHYTOTAXA Copyright © 2020 Magnolia Press Article ISSN 1179-3163 (online edition) https://doi.org/10.11646/phytotaxa.442.3.1 A new species of Myrsine (Primulaceae-Myrsinoideae) from New Guinea TIMOTHY M.A. UTTERIDGE1,3 & BRENDAN J. LEPSCHI2,4 1 Identification & Naming Dept, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, UK. 2 Australian National Herbarium, Centre for Australian National Biodiversity Research, GPO Box 1700, Canberra, ACT, 2601, Australia. 3 [email protected]; https://orcid.org/0000-0003-2823-0337 4 [email protected]; https://orcid.org/0000-0002-3281-2973 Abstract Myrsine exquisitorum Utteridge & Lepschi (Primulaceae-Myrsinoideae) is described and illustrated as a new species endemic to the Western Highlands Province from Papua New Guinea. The new species is unique in the relatively large, almost orbicular leaves with entire margins, and the tetramerous flowers arranged in axillary fascicles without forming short shoots. Keywords: Papuasia, Malesia, Myrsinaceae, Rapanea, taxonomy Introduction Myrsine Linnaeus (1753: 196) is a pantropical genus of c. 280 species, and now includes all taxa previously placed in the paleotropical genus Rapanea Aublet (1775: 121) and the New World genus Suttonia A.Richard (1832: 349). Traditionally, Rapanea was considered distinct from Myrsine based on the absence of well-defined filaments and style, but these characters have been shown to intergrade between species of the different genera. Pipoly (2007) and Takeuchi & Pipoly (2009) further discuss character variation in the genera in New Guinea, and formally completed the transfer of the eastern Malesian names of Rapanea to Myrsine.
    [Show full text]
  • Two New Species of Primula (Primulaceae) from China
    Two New Species of Primula (Primulaceae) from China Hu Chi Ming South China Institute of Botany, Academia Sinica, Guangzhou 510650, People's Republic of China. [email protected] Geng Yu Ying West China Subalpine Botanical Garden, Institute of Botany, Academia Sinica, Dujiangyan 611830, China ABSTRACT. Two new species of Primula from Species af®nis P. hylobiae et P. klaverianae, a quibus western China are described, illustrated, and com- foliis scapisque glabrissimis, calyce tantum ad tertiam partem in lobos ovatos ®sso differt; ab altera etiam ¯oribus pared with putatively related species. Both new heterogeneis differt. species belong to section Petiolares subsect. Davi- dii (Balfour f.) Craib, which is characterized by the Perennial herbs, efarinose, with a short compar- capsule crumbling irregularly at maturity, the basal atively stout rhizome, basally with brown palea- buds always having a covering of paleaceous scales, ceous bud-scales at ¯owering time. Leaves forming and the leaves being more or less coriaceous, per- a spreading rosette, to 30 cm diam.; leaf blade ob- sisting into the following spring, with veins often long-elliptic to elliptic-obovate, 12±20 3 6±9 cm, prominently raised and alveolate abaxially and im- apex rounded, tapering from the middle into a very pressed adaxially. In this subsection there are about short or almost obsolete winged petiole, irregularly 18 species all distributed in Sichuan or Yunnan, crenate-dentate, papery when dry, 6 bullate on the with only one species extending to Guizhou, Hubei, margin and covered with minute glandular hairs Hunan, and Jiangxi. when just unfolded from the resting buds, soon gla- Key words: China, Primula, Primulaceae.
    [Show full text]
  • Phylogenetic Relationships in the Order Ericales S.L.: Analyses of Molecular Data from Five Genes from the Plastid and Mitochondrial Genomes1
    American Journal of Botany 89(4): 677±687. 2002. PHYLOGENETIC RELATIONSHIPS IN THE ORDER ERICALES S.L.: ANALYSES OF MOLECULAR DATA FROM FIVE GENES FROM THE PLASTID AND MITOCHONDRIAL GENOMES1 ARNE A. ANDERBERG,2,5 CATARINA RYDIN,3 AND MARI KAÈ LLERSJOÈ 4 2Department of Phanerogamic Botany, Swedish Museum of Natural History, P.O. Box 50007, SE-104 05 Stockholm, Sweden; 3Department of Systematic Botany, University of Stockholm, SE-106 91 Stockholm, Sweden; and 4Laboratory for Molecular Systematics, Swedish Museum of Natural History, P.O. Box 50007, SE-104 05 Stockholm, Sweden Phylogenetic interrelationships in the enlarged order Ericales were investigated by jackknife analysis of a combination of DNA sequences from the plastid genes rbcL, ndhF, atpB, and the mitochondrial genes atp1 and matR. Several well-supported groups were identi®ed, but neither a combination of all gene sequences nor any one alone fully resolved the relationships between all major clades in Ericales. All investigated families except Theaceae were found to be monophyletic. Four families, Marcgraviaceae, Balsaminaceae, Pellicieraceae, and Tetrameristaceae form a monophyletic group that is the sister of the remaining families. On the next higher level, Fouquieriaceae and Polemoniaceae form a clade that is sister to the majority of families that form a group with eight supported clades between which the interrelationships are unresolved: Theaceae-Ternstroemioideae with Ficalhoa, Sladenia, and Pentaphylacaceae; Theaceae-Theoideae; Ebenaceae and Lissocarpaceae; Symplocaceae; Maesaceae, Theophrastaceae, Primulaceae, and Myrsinaceae; Styr- acaceae and Diapensiaceae; Lecythidaceae and Sapotaceae; Actinidiaceae, Roridulaceae, Sarraceniaceae, Clethraceae, Cyrillaceae, and Ericaceae. Key words: atpB; atp1; cladistics; DNA; Ericales; jackknife; matR; ndhF; phylogeny; rbcL. Understanding of phylogenetic relationships among angio- was available for them at the time, viz.
    [Show full text]
  • Lysimachia Huangsangensis (Primulaceae), a New Species from Hunan, China
    RESEARCH ARTICLE Lysimachia huangsangensis (Primulaceae), a New Species from Hunan, China Jian-Jun Zhou1, Xun-Lin Yu1*, Yun-Fei Deng2*, Hai-Fei Yan2, Zhe-Li Lin2,3 1 School of Forestry, Central South University of Forestry & Technology, 410004, Changsha, People’s Republic of China, 2 Key Laboratory of Plant Resources Conservation and Sustainable Utilization, South China Botanical Garden, the Chinese Academy of Sciences, Guangzhou 510650, People’s Republic of China, 3 University of Chinese Academy of Sciences, Beijing 100049, People’s Republic of China * [email protected] (XLY); [email protected] (YFD) Abstract A new species, Lysimachia huangsangensis (Primulaceae), from Hunan, China is described and illustrated. The new species is closely related to L. carinata because of the crested calyx, but differs in the leaf blades that are ovate to elliptic and (3–)4.5–9×2–3.4 cm, 2–5- OPEN ACCESS flowered racemes, and the calyx lobes that are ovate-lanceolate and 5–6×3–4 mm. The Citation: Zhou J-J, Yu X-L, Deng Y-F, Yan H-F, Lin Z- systematic placement and conservation status are also discussed. L (2015) Lysimachia huangsangensis (Primulaceae), a New Species from Hunan, China. PLoS ONE 10(7): e0132713. doi:10.1371/journal.pone.0132713 Editor: Nico Cellinese, University of Florida, UNITED STATES Introduction Received: February 19, 2015 Lysimachia L. belongs to the tribe Lysimachieae Reich. and consists of 140–200 species with an Accepted: June 15, 2015 almost worldwide distribution but exhibits striking local endemism [1–7]. China is one of the Published: July 22, 2015 centers of diversity for Lysimachia, being home to approximately 140 species [5, 8–20].
    [Show full text]
  • Pharmacognostical and Preliminary Phytochemical Profile of the Leaf
    Journal of Pharmacognosy and Phytochemistry 2019; 8(1): 1861-1864 E-ISSN: 2278-4136 P-ISSN: 2349-8234 Pharmacognostical and preliminary JPP 2019; 8(1): 1861-1864 Received: 08-11-2018 phytochemical profile of the leaf extracts of Accepted: 12-12-2018 Embelia ribes Burm. F. Ananth V Research Scholar, Department of Botany, Bishop Heber College Ananth V, Anand Gideon V and John Britto S Autonomous, Tiruchirappalli, Tamil Nadu, India Abstract Embelia ribes Burm. f. is a valuable medicinal plant used in Indian system of medicine. This article deals Anand Gideon V with gross morphology, anatomy and preliminary phytochemicals presence of alkaloids, flavonoids, Associate professor and Head, amino acids, proteins and carbohydrates. Department of Botany, Bishop Heber College, (Autonomous), Tiruchirapalli, Keywords: Embelia ribes, microscope, preliminary phytochemical Tamil Nadu, India Introduction John Britto S Plants are the invaluable sources of medicines, especially in traditional systems of health care. The Rapinat Herbarium and The recent global resurgence of interest in herbal medicines has led to an increase in the Centre for Molecular Systematics, St Joseph’s College demand for them. The research on the analysis and quality control of herbal medicines now is Autonomous, Tiruchirappalli, in the direction of an integrative and comprehensive approach, in order to better address the Tamil Nadu, India inherent holistic nature of herbal medicines. Embelia ribes Burm. f. of family Primulaceae. (APG IV 2016) is seen in hills of India up to 1500m elevation extending from outer Himalayas to Western Ghats. It is an endangered medicinal plant. Due to its over exploitation it is reported in red list data book as vulnerable [Ravikumar & Ved 2000].
    [Show full text]