DOCSLIB.ORG

Female Gamete Competition in an Ancient Angiosperm Lineage

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
Female Gamete Competition in an Ancient Angiosperm Lineage Female gamete competition in an ancient angiosperm lineage Julien B. Bacheliera,b,c and William E. Friedmana,b,c,1 aDepartment of Ecology and Evolutionary Biology, University of Colorado, Boulder, CO 80309; bDepartment of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA 02138; and cArnold Arboretum, Harvard University, Boston, MA 02131 Edited* by Peter H. Raven, Missouri Botanical Garden, St. Louis, MO, and approved May 19, 2011 (received for review March 23, 2011) In Trimenia moorei, an extant member of the ancient angiosperm ly diversification of floral morphology and anatomy are widely clade Austrobaileyales, we found a remarkable pattern of female viewed to have led to increased levels of pollen reception and gametophyte (egg-producing structure) development that strik- hence, male–male competition (4). Collectively, the evolution of ingly resembles that of pollen tubes and their intrasexual compe- the carpel, extragynoecial compitum, transmitting tissue, callose tition within the maternal pollen tube transmitting tissues of most plugs, and insect pollination seems to have resulted in signifi- flowers. In contrast with most other flowering plants, in Trimenia, cantly enhanced levels of prefertilization male competition and multiple female gametophytes are initiated at the base (chalazal maternal choice, and thus may have played a major role in the end) of each ovule. Female gametophytes grow from their tips and early diversification of angiosperms (4–9). compete over hundreds of micrometers to reach the apex of the Despite all of the attention paid to prefertilization mecha- nucellus and the site of fertilization. Here, the successful female nisms of male competition and female choice, there has never gametophyte will mate with a pollen tube to produce an embryo been any discussion of the possibility that female gametophytes and an endosperm. Moreover, the central tissue within the ovules (haploid egg-producing structures within ovules) might engage in of Trimenia, through which the embryo sacs grow, contains starch a process of competition. Interestingly, recent embryological and other carbohydrates similar to the pollen tube transmitting studies of several members of the Nymphaeales and Austro- tissues in the styles of most flowers. The pattern of female game- baileyales have revealed that, in contrast with most angiosperms, tophyte development found in Trimenia is rare but by no means more than one female gametophyte is occasionally or frequently EVOLUTION unique in angiosperms. Importantly, it seems that multiple female initiated in each ovule (10–14). For us, this suggests the potential gametophytes are occasionally or frequently initiated in members for competition between female gametophyte genomes during fi of other ancient angiosperm lineages. The intensi cation of pollen the developmental period leading up to the time of fertilization. tube (male gametophyte) competition and enhanced maternal se- To gain insight into whether female gametophytes, in addition lection among competing pollen tubes are considered to have to male gametophytes, also engage in intrasexual competition been major contributors to the rise of angiosperms. Based on fl Trimenia before fertilization among early lineages of owering plants, we insights from , we posit that prefertilization female ga- examined ovule and gametophyte development in Trimenia (Tri- metophyte (egg) competition within individual ovules in addition meniaceae). Previous studies of Trimenia reported that as many as to male gametophyte (sperm) competition and maternal mate 20 megasporocytes differentiate in each ovule and that several choice may have been key features of the earliest angiosperms. female gametophytes will proceed to develop (11, 15). These brief reports provide circumstantial evidence for potentially high levels evolution of flowering plants | reproduction | Darwin of intragametophytic competition. However, none of these earlier studies examined the development of female gametophytes from ver since Charles Darwin concluded that the early diversi- their initiation to the time of fertilization. As will be seen, because fi “ ” E cation of angiosperms was an abominable mystery, numer- Trimenia occupies a relatively basal position within the Austro- ous hypotheses have been advanced to explain their extraor- baileyales, it has the potential to help resolve the question of dinary evolutionary radiation (1, 2). These hypotheses suggest that whether female gametophyte competition before fertilization was fl anumberofkey owering plant synapomorphies were intimately a critical feature of the earliest phases of angiosperm evolution. associated with their early evolutionary history and subsequent diversification (3). Foremost among the defining features of an- Results giosperms is the carpel, which encloses the ovule/seed and increases Initiation of Multiple Female Gametophytes Occurs Far from the Site the distance of pollen tube growth between the site of pollen re- of Fertilization. In T. moorei, numerous diploid megaspore mother ception and point of fertilization. This key innovation fundamen- cells (megasporocytes) within a single ovule initiate meiosis to tally altered the basic interactions between the male and female create haploid functional megaspores (a megaspore is the founding components of the reproductive process, and it resulted in higher cell of a female gametophyte). Female gametophytes are initiated levels of male gametophyte competition and greater opportunities deep within the ovule (proximate to the chalaza) in a position re- for maternal selection among potential sperm donors (4). mote from the future site of fertilization (Fig. 1). This phenomenon Recent character state reconstructions show that the common stands in marked contrast to most flowering plants, where the fe- ancestor of angiosperms likely had an extragynoecial compitum, male gametophyte is initiated and develops in a region of the ovule a construction of the carpels that enhances competition among that is close to the location where the pollen tube will arrive to pollen tubes and their selection by the maternal plant before deposit its sperm (Discussion)(16). fertilization (5, 6). In addition, the differentiation of a trans- mitting tissue within the carpel that supports pollen tube growth to the ovule has long been viewed as promoting male competi- Author contributions: W.E.F. designed research; J.B.B. performed research; J.B.B. and tion and increasing maternal choice of sperm donors (7, 8). The W.E.F. analyzed data; and J.B.B. and W.E.F. wrote the paper. formation of callose plugs within pollen tubes recently has been The authors declare no conflict of interest. hypothesized to have enabled male gametophytes to grow at *This Direct Submission article had a prearranged editor. higher rates and compete over greater distances than was the 1To whom correspondence should be addressed. E-mail: [email protected]. case for the immediate ancestors of angiosperms (8, 9). Finally, This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10. the evolution of insect pollination in conjunction with the ear- 1073/pnas.1104697108/-/DCSupplemental. www.pnas.org/cgi/doi/10.1073/pnas.1104697108 PNAS Early Edition | 1of6 Downloaded by guest on September 26, 2021 A B * A B ii ii B fun ii ii * C nuc C fg nuc D D zm zm mmc ch ch Fig. 2. Young ovule with multiple female gametophytes. (A)3Dre- Fig. 1. Young ovules showing the zone of megasporogenesis at the cha- construction (from transverse serial sections) of ovule nucellus (gray) con- lazal pole of the nucellus. (A) Very young ovule with one of the megaspore taining tube-like female gametophytes (each a different color) initiated at mother cells (mmc) in the zone of megasporogenesis (zm) at the chalazal the base of the nucellus and growing to the site of fertilization at the apex of pole (ch) of the nucellus (nuc). The asterisk marks the future site of fertil- the nucellus (micropylar pole of ovule at the top). One female gametophyte ization at the micropylar end of the nucellus. (B) Ovule showing several (green) has become dominant. (Scale bar: 200 μm.) (B–D) Transverse sections degenerated megaspores within the zone of megasporogenesis. The arrow from the series used for reconstruction in A and stained with periodic acid points to the tip of a female gametophyte (fg) that has grown out of the Schiff’s (PAS) reagent to reveal starch and other carbohydrates in pink. Each zone of megasporogenesis and is beginning to ascend through the nucellus star marks a female gametophyte of the corresponding color in the computer central tissue (indicated by arrowheads) to the future site of fertilization reconstruction; dashed lines in A indicate position of each transverse section. (asterisk). ii, inner integument; fun, funiculus. (Scale bars: 100 μm.) (B) Central tissue of the nucellus filled with starch grains (black arrows). (C) Central tissue of the nucellus with fewer and smaller starch grains and one dominant two-nucleate female gametophyte (green in A; one of two nuclei Female Gametophytes Undergo Tip Growth and Appear to Compete. shown by the green arrowhead). (D) Base of nucellus, where additional fe- Although some megaspores abort, as many as 10 megaspores male gametophytes (black stars) were initiated. (Scale bars: 25 μm.) within an individual ovule initiate female gametophytes that seem to crush and displace the diploid maternal tissue (nucellus) and Movie S3). The two free nuclei of a dominant female ga- situated
Load more

Recommended publications
  • Antioxidant Effects of Schisandra Chinensis Fruits and Their Active Constituents
    antioxidants Review Antioxidant Effects of Schisandra chinensis Fruits and Their Active Constituents Dalia M. Kopustinskiene 1 and Jurga Bernatoniene 1,2,* 1 Institute of Pharmaceutical Technologies, Faculty of Pharmacy, Medical Academy, Lithuanian University of Health Sciences, Sukileliu pr. 13, LT-50161 Kaunas, Lithuania; [email protected] 2 Department of Drug Technology and Social Pharmacy, Faculty of Pharmacy, Medical Academy, Lithuanian University of Health Sciences, Sukileliu pr. 13, LT-50161 Kaunas, Lithuania * Correspondence: [email protected] Abstract: Schisandra chinensis Turcz. (Baill.) fruits, their extracts, and bioactive compounds are used in alternative medicine as adaptogens and ergogens protecting against numerous neurological, cardiovascular, gastrointestinal, liver, and skin disorders. S. chinensis fruit extracts and their active compounds are potent antioxidants and mitoprotectors exerting anti-inflammatory, antiviral, anti- cancer, and anti-aging effects. S. chinensis polyphenolic compounds—flavonoids, phenolic acids and the major constituents dibenzocyclooctadiene lignans are responsible for the S. chinensis antioxidant activities. This review will focus on the direct and indirect antioxidant effects of S. chinensis fruit extract and its bioactive compounds in the cells during normal and pathological conditions. Keywords: Schisandra chinensis; lignan; schisandrin B; antioxidant; pro-oxidant; mitochondria Citation: Kopustinskiene, D.M.; 1. Introduction Bernatoniene, J. Antioxidant Effects Schisandra chinensis Turcz. (Baill.) belongs to the Schisandraceae family. The plants of Schisandra chinensis Fruits and are native to northeastern China, Japan, Korea, Manchuria, and the Far East part of Russia. Their Active Constituents. Their purple-red berries are called five-flavor fruits because of the sweet, bitter, pungent, Antioxidants 2021, 10, 620. https:// salty, and sour taste [1–5]. S.
    [Show full text]
  • Analysis of Genetic Diversity of Piper Spp. in Hainan Island (China) Using Inter-Simple Sequence Repeat ISSR Markers
    African Journal of Biotechnology Vol. 10(66), pp. 14731-14737, 26 October, 2011 Available online at http://www.academicjournals.org/AJB DOI: 10.5897/AJB11.2342 ISSN 1684–5315 © 2011 Academic Journals Full Length Research Paper Analysis of genetic diversity of Piper spp. in Hainan Island (China) using inter-simple sequence repeat ISSR markers Yan Jiang 1,2 and Jin-Ping Liu 1,2 * 1Key Laboratory of Protection and Development Utilization of Tropical Crop Germplasm Resources (Hainan University), Ministry of Education, Haikou, Hainan Province, 570228, China. 2College of Agronomy, Hainan University, Danzhou, Hainan Province, 571737, China. Accepted 28 September, 2011 Inter-simple sequence repeat (ISSR) analysis was used to evaluate the genetic variation of Piper spp. from Hainan, China. 247 polymorphic bands out of a total of 248 (99.60%) were generated from 74 individual plants of Piper spp. The overall level of genetic diversity among Piper spp. in Hainan was high, with the mean Shannon information index (I) of 0.2843 and the mean Nei’s genetic diversity (H) of 0.1904. The genetic similarity (GS) coefficient ranged from 0.548 to 0.976 within 74 individual plants of Piper spp., and the within-species genetic distance ranged from 0.104 to 0.28. Unweighted pair group method with arithmetic mean (UPGMA) dendrogram showed that P. kadsura is the most divergent and the most distant of the 11 species, and that P. hainanense and P. bonii , are closely related as well as P. sarmentosum and P. betle . The diversity analysis unambiguously distinguished all Piper spp. The high levels of genetic diversity in Jianfengling and Diaoluoshan demonstrate that conservation of wild resources of Piper in these two localities is more effective than that in Limushan, Wuzhishan, Xinglong Tropical Botanical Garden and Danzhou.
    [Show full text]
  • New Dibenzocylooctadiene Lignans from Kadsura Heteroclita
    Chinese Journal of Natural Chinese Journal of Natural Medicines 2014, 12(9): 06890692 Medicines Heteroclitins R-S: new dibenzocylooctadiene lignans from Kadsura heteroclita CHEN Min1*, LUO You-Ping1, 2, ZOU Yan-Lin1, LANG Ling-Hu1, CHEN Dao-Feng3 1 Key Laboratory on Luminescence and Real-Time Analysis (Ministry of Education), College of Pharmaceutical Sciences, Southwest University, Chongqing 400715, China; 2 College of Chemistry and Chemical Engineering, HaiNan Normal University, Hai Kou 571158, China; 3 Department of Pharmacognosy, School of Pharmacy, Fudan University, Shanghai 201203, China Available online September 2014 [ABSTRACT] AIM: To study the dibenzocylooctadiene lignans from the stems of Kadsura heteroclita. METHOD: Chromatographic separations of silica gel and semi-preparative HPLC were used. All of the structures were elucidated on the basis of spectroscopic analysis ,including 2D-NMR and HR-MS techniques. RESULTS: Four dibenzocylooctadiene lignans were isolated from K. heteroclita. Their structures were identified as heteroclitin R (1), heteroclitin S (2), gonisin O (3), and schisanlignone A (4). CONCLUSION: Heteroclitin R (1) and heteroclitin S (2) are new natural lignans. [KEY WORDS] Dibenzocylooctadiene Lignan; Kadsura heteroclita; Schisandraceae; Heteroclitins R-S [CLC Number] R284.1 [Document code] A [Article ID] 2095-6975(2014)09-0689-04 ern area of China. Repeated column chromatography of the Introduction Et2O extract of the stems of K. heteroclita led to the isolation The stems of Kadsura heteroclita (Roxb.) Craib (Schi- and identification of a new dibenzocylooctadiene lignan named sandraceae) are commonly used in Chinese traditional medi- heteroclitin R (1), a new natural product named heteroclitin S [9] cine to promote vital energy and blood circulation, to expel (2), and two known compounds, gonisin O (3) and schi- [10] wind-evil, and to remove wetness-evil [1].
    [Show full text]
  • Read Book Schisandra Chinensis: an Herb of North Eastern China Origin
    SCHISANDRA CHINENSIS: AN HERB OF NORTH EASTERN CHINA ORIGIN PDF, EPUB, EBOOK Kam Ming Ko | 252 pages | 28 Feb 2015 | World Scientific Publishing Co Pte Ltd | 9789814651226 | English | Singapore, Singapore Schisandra Chinensis: An Herb Of North Eastern China Origin PDF Book Due to its ability to positively affect the immune system and fight inflammation , schisandra seems to help stall the development of atherosclerosis hardening of the arteries , balances blood sugar, prevents diabetes and bring the body into an optimal acid-base balance. Safety: High, but may cause over-stimulation if overused. The average increase in the mean concentration of Tac in the blood was percent for the group receiving higher doses of SchE and percent for the group receiving lower a dose. Repeated expansions and fragmentations linked to Pleistocene climate changes shaped the genetic structure of a woody climber, Actinidia arguta Actinidiaceae. Structure-activity relationships of lignans from Schisandra chinensis as platelet activating antagonists. All rights reserved. Schisandra berry demonstrates significant adaptogenic activity. To assure optimal quality, they directly purchase from China berries that have been graded as premium quality only. But some berries are deep refrigerated, and eventually used to make health juices, primarily for the Korean market. Schisandra supports the immune system and protects against microbes. Schisandra chinensis can also function as a convalescent tonic herb when the kidney system is involved. Evolution 38, — Antao, T. Share: facebook twitter pinterest. We recommend that you consult with a qualified healthcare practitioner before using herbal products, particularly if you are pregnant, nursing, or on any medications. When it comes to cancer prevention, active lignans have been isolated from schisandra especially one called schisandrin A that have chemo-protective abilities.
    [Show full text]
  • Characterization of the Omija (Schisandra Chinensis) Extract and Its Effects on the Bovine Sperm Vitality and Oxidative Profile During in Vitro Storage
    Hindawi Evidence-Based Complementary and Alternative Medicine Volume 2020, Article ID 7123780, 15 pages https://doi.org/10.1155/2020/7123780 Research Article Characterization of the Omija (Schisandra chinensis) Extract and Its Effects on the Bovine Sperm Vitality and Oxidative Profile during In Vitro Storage Eva Tvrda´ ,1 Jaroslav Michalko,2,3 Ju´ lius A´ rvay,4 Nenad L. Vukovic,5 Eva Ivanisˇova´,6 Michal Dˇ uracˇka,1 Ildiko´ Matusˇı´kova´,7 and Miroslava Kacˇa´niova´ 8,9 1Department of Animal Physiology, Faculty of Biotechnology and Food Sciences, Slovak University of Agriculture in Nitra, Tr. A. Hlinku 2, 949 76 Nitra, Slovakia 2BioFood Center, Faculty of Biotechnology and Food Sciences, Slovak University of Agriculture in Nitra, Tr. A. Hlinku 2, 949 76 Nitra, Slovakia 3Detached Branch of the Institute of Forest Ecology-Arbore´tum Mlynˇany, Slovak Academy of Sciences, Vieska Nad Zˇitavou 178, 951 52 Slepˇcany, Slovakia 4Department of Chemistry, Faculty of Biotechnology and Food Sciences, Slovak University of Agriculture in Nitra, Tr. A. Hlinku 2, 949 76 Nitra, Slovakia 5Department of Chemistry, Faculty of Science, University of Kragujevac, 34000 Kragujevac, Serbia 6Department of Technology and Quality of Plant Products, Faculty of Biotechnology and Food Sciences, Slovak University of Agriculture, Tr. A. Hlinku 2, 94976 Nitra, Slovakia 7Department of Ecochemistry and Radioecology, Faculty of Natural Sciences, University of SS. Cyril and Methodius in Trnava, Na´m. J. Herdu 2, 917 01 Trnava, Slovakia 8Department of Fruit Sciences, Viticulture and Enology, Faculty of Horticulture and Landscape Engineering, Slovak University of Agriculture, Tr. A. Hlinku 2, 94976 Nitra, Slovakia 9Department of Bioenergy, Food Technology and Microbiology, Institute of Food Technology and Nutrition, University of Rzeszow, Zelwerowicza St.
    [Show full text]
  • Schisandra Chinensis Sam Schmerler
    Plainly Unique: Schisandra chinensis Sam Schmerler he plants of the Arnold Arboretum dis- into elongated fruits with numerous bright red, play incredible floral diversity. Magnolia berrylike fruitlets. Winter will reveal exfoliating Tmacrophylla’s huge waxy blooms open bark resembling that of climbing hydrangea. twice, partly closing in between for an over- Evolutionary biologists (including Arboretum night sex change. Helwingia japonica sprouts director Ned Friedman) have discovered that tiny green umbels in the center of otherwise Schisandra and the other Austrobaileyales can unremarkable leaves. Davidia involucrata for- offer insight into many key events in the his- goes petals entirely, but shelters its reproductive tory of flowering plants. Aspects of Schisandra’s organs with massive white bracts. Even wild vascular system may represent an early step in Viola sororia, flagging down bees with its iconic the development of vessels, the structures that violets, surreptitiously sends out discrete, self- allow most flowering plants to rapidly trans- pollinating flowers underground. port water and ecologically dominate hot and With all this bizarre and beautiful reproduc- dry habitats. Schisandra also retains a relatively tion going on, most of us overlook the most simple anatomy during its haploid stage, with evolutionarily distinctive flowering plant in only four nuclei and one developmental module the collection: Schisandra chinensis. An unas- in each female gametophyte (almost all flower- ing plants have eight nuclei and two modules). suming woody vine, it represents a unique and The endosperm of Schisandra seeds conse- ancient lineage that parted ways with most other quently contains only one complement of genes flowering plants at least as far back as the early from each of its parents, while most flowering Cretaceous, before even “living fossils” like plants acquire an additional copy of their moms’ Magnolia.
    [Show full text]
  • Evaluation of Neuroactive Effects of Ethanol Extract of Schisandra
    Chinese Journal of Natural Chinese Journal of Natural Medicines 2018, 16(12): 09160925 Medicines Evaluation of neuroactive effects of ethanol extract of Schisandra chinensis, Schisandrin, and Schisandrin B and determination of underlying mechanisms by zebrafish behavioral profiling WANG Jia-Wei1Δ, LIANG Feng-Yin2Δ, OUYANG Xiang-Shuo1, LI Pei-Bo1, PEI Zhong2*, SU Wei-Wei 1* 1 Guangdong Engineering and Technology Research Centre for Quality and Efficacy Re-evaluation of Post-marketed TCM, Guangdong Key Laboratory of Plant Resources, School of Life Sciences, Sun Yat-sen University, Guangzhou 510275, China; 2 Department of Neurology, National Key Clinical Department and Key Discipline of Neurology, Guangdong Provincial Key Laboratory for Diagnosis and Treatment of Major Neurological Diseases, The First Affiliated Hospital, Sun Yat-sen University, Guangzhou 510080, China Available online 20 Dec., 2018 [ABSTRACT] Schisandra chinensis, a traditional Chinese medicine (TCM), has been used to treat sleep disorders. Zebrafish sleep/wake behavioral profiling provides a high-throughput platform to screen chemicals, but has never been used to study extracts and components from TCM. In the present study, the ethanol extract of Schisandra chinensis and its two main lignin components, schisan- drin and schisandrin B, were studied in zebrafish. We found that the ethanol extract had bidirectional improvement in rest and activity in zebrafish. Schisandrin and schisandrin B were both sedative and active components. We predicted that schisandrin was related to serotonin pathway and the enthanol extract of Schisandra chinensis was related to seoronin and domapine pathways using a database of zebrafish behaviors. These predictions were confirmed in experiments using Caenorhabditis elegans. In conclusion, zebrafish behavior profiling could be used as a high-throughput platform to screen neuroactive effects and predict molecular pathways of extracts and components from TCM.
    [Show full text]
  • New Cadinane Sesquiterpenes from the Stems of Kadsura Heteroclita
    molecules Article New Cadinane Sesquiterpenes from the Stems of Kadsura heteroclita 1, 1,2, 1,2 1 1 Liang Cao y, Nuzhat Shehla y, Shumaila Tasneem , Mengru Cao , Wenbing Sheng , Yuqing Jian 1, Bin Li 1, Caiyun Peng 1, M. Iqbal Choudhary 1,2,*, Atta-ur-Rahman 2, Duan-fang Liao 1 and Wei Wang 1,2,* 1 TCM and Ethnomedicine Innovation & Development International Laboratory, Innovative Materia Medica Research Institute, School of Pharmacy, Hunan University of Chinese Medicine, Changsha 410208, China; [email protected] (L.C.); [email protected] (N.S.); [email protected] (S.T.); [email protected] (M.C.); [email protected] (W.S.); [email protected] (Y.J.); [email protected] (B.L.); [email protected] (C.P.); dfl[email protected] (D.-f.L.) 2 H.E.J. Research Institute of Chemistry, International Center for Chemical and Biological Sciences, University of Karachi, Karachi 75270, Pakistan; [email protected] * Correspondence: [email protected] (M.I.C.); [email protected] (W.W.); Tel.: +86-136-5743-8606 (M.I.C.); +92-21-4824924 or +92-21-4819010 (W.W.) They contributed equally to this work. y Received: 10 March 2019; Accepted: 26 April 2019; Published: 28 April 2019 Abstract: As part of our continual efforts to exploit ‘Tujia Ethnomedicine’ for their pharmacophoric functionalities, we herein investigated Kadsura heteroclita collected from a deep Wulin mountain area in northern Hunan province. The current study resulted in the isolation of three new sesquiterpenes: 6α,9α,15-trihydroxycadinan-4-en-3-one (1), (+)-3,11,12-trihydroxycalamenene (2), (–)-3,10,11,12-tetrahydroxy-calamenene (3), along with four known sesquiterpenes (4–7), and a cytochalasin H (8).
    [Show full text]
  • Rhodiola Rosea L., Rhizoma Et Radix
    12 July 2011 EMA/HMPC/232100/2011 Committee on Herbal Medicinal Products (HMPC) Assessment report on Rhodiola rosea L., rhizoma et radix Based on Article 16d(1), Article 16f and Article 16h of Directive 2001/83/EC as amended (traditional use) Draft Herbal substance(s) (binomial scientific name of Rhodiola rosea L., rhizoma et radix the plant, including plant part) Herbal preparation(s) Dry extract (DER 1.5-5:1), extraction solvent ethanol 67-70% v/v Pharmaceutical forms Herbal preparations in solid dosage forms for oral use. Note: This Assessment Report is published to support the release for public consultation of the draft Community herbal monograph on Rhodiola rosea L., rhizoma et radix. It should be noted that this document is a working document, not yet fully edited, and which shall be further developed after the release for consultation of the monograph. Interested parties are welcome to submit comments to the HMPC secretariat, which the Rapporteur and the MLWP will take into consideration but no ‘overview of comments received during the public consultation’ will be prepared in relation to the comments that will be received on this assessment report. The publication of this draft assessment report has been agreed to facilitate the understanding by Interested Parties of the assessment that has been carried out so far and led to the preparation of the draft monograph. 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, 2011.
    [Show full text]
  • Thailand’S Sixth National Report on the Implementation of the Convention on Biological Diversity
    Thailand’s Sixth National Report on the Implementation of the Convention on Biological Diversity Thailand’s Sixth National Report on the Implementation of the Convention on Biological Diversity June 2019 Forward As a Party of the Convention on Biological Diversity (CBD), Thailand is responsible for addressing the challenges of safeguarding biodiversity and ecosystem services, which are national priorities. CBD’s parties are required to submit the National Report in accordance with the Convention’s objectives (Article 26 of the CBD). Thailand’s Sixth National Report covers the period of January 2015 to December 2018 and aligns with the agreed reporting guidelines of the CBD. We hope this report can contribute to the development of the Global Biodiversity Outlook and the Post 2020 Global Biodiversity Framework. Thailand will continue to take concrete steps to protect its biodiversity, the unique natural heritage, and precious ecosystems, for the next generations. All Thais – including all levels of government, the public, private, local stakeholders and non-government organizations are responsible for biodiversity conservation. 1 Preface Thailand’s Sixth National Report (6NR) is one of the obligations under the Convention on Biological Diversity (CBD). As a party to the CBD, Thailand has been implementing prevalent policies and measures to fulfill the country’s commitments to the CBD and its Protocols. In addition, Thailand has been actively participating in various international cooperation and regional biodiversity initiatives. This 6NR
    [Show full text]
  • Forest Habitats and Flora in Laos PDR, Cambodia and Vietnam
    See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/259623025 Forest Habitats and Flora in Laos PDR, Cambodia and Vietnam Conference Paper · January 1999 CITATIONS READS 12 517 1 author: Philip W. Rundel University of California, Los Angeles 283 PUBLICATIONS 8,872 CITATIONS SEE PROFILE Available from: Philip W. Rundel Retrieved on: 03 October 2016 Rundel 1999 …Forest Habitats and Flora in Lao PDR, Cambodia, and Vietnam 1 Conservation Priorities In Indochina - WWF Desk Study FOREST HABITATS AND FLORA IN LAO PDR, CAMBODIA, AND VIETNAM Philip W. Rundel, PhD Department of Ecology and Evolutionary Biology University of California Los Angeles, California USA 90095 December 1999 Prepared for World Wide Fund for Nature, Indochina Programme Office, Hanoi Rundel 1999 …Forest Habitats and Flora in Lao PDR, Cambodia, and Vietnam 2 TABLE OF CONTENTS Introduction 1. Geomorphology of Southeast Asia 1.1 Geologic History 1.2 Geomorphic Provinces 1.3 Mekong River System 2. Vegetation Patterns in Southeast Asia 2.1 Regional Forest Formations 2.2 Lowland Forest Habitats 2.3 Montane Forest Habitats 2.4 Freshwater Swamp Forests 2.5 Mangrove Forests Lao People's Democratic Republic 1. Physical Geography 2. Climatic Patterns 3. Vegetation Mapping 4. Forest Habitats 5.1 Lowland Forest habitats 5.2 Montane Forest Habitats 5.3 Subtropical Broadleaf Evergreen Forest 5.4 Azonal Habitats Cambodia 1. Physical Geography 2. Hydrology 3. Climatic Patterns 4. Flora 5. Vegetation Mapping 6. Forest Habitats 5.1 Lowland Forest habitats 5.2 Montane Forest Habitats 5.3 Azonal Habitats Vietnam 1. Physical Geography 2.
    [Show full text]
  • Dynamic Evolution and Phylogenomic Analysis of the Chloroplast Genome
    www.nature.com/scientificreports OPEN Dynamic evolution and phylogenomic analysis of the chloroplast genome in Received: 19 March 2018 Accepted: 31 May 2018 Schisandraceae Published: xx xx xxxx Bin Li1,2,3 & Yongqi Zheng1,2,3 Chloroplast genomes of plants are highly conserved in both gene order and gene content, are maternally inherited, and have a lower rate of evolution. Chloroplast genomes are considered to be good models for testing lineage-specifc molecular evolution. In this study, we use Schisandraceae as an example to generate insights into the overall evolutionary dynamics in chloroplast genomes and to establish the phylogenetic relationship of Schisandraceae based on chloroplast genome data using phylogenomic analysis. By comparing three Schisandraceae chloroplast genomes, we demonstrate that the gene order, gene content, and length of chloroplast genomes in Schisandraceae are highly conserved but experience dynamic evolution among species. The number of repeat variations were detected, and the Schisandraceae chloroplast genome was revealed as unusual in having a 10 kb contraction of the IR due to the genome size variations compared with other angiosperms. Phylogenomic analysis based on 82 protein-coding genes from 66 plant taxa clearly elucidated that Schisandraceae is a sister to a clade that includes magnoliids, monocots, and eudicots within angiosperms. As to genus relationships within Schisandraceae, Kadsura and Schisandra formed a monophyletic clade which was sister to Illicium. Chloroplasts are the photosynthetic organelle that provides energy for plants. Te chloroplast has its own genome. In angiosperms, most chloroplast genomes are composed of circular DNA molecules ranging from 120 to 160 kb in length and have a quadripartite organization consisting of two copies of inverted repeats (IRs) of approximately 20–28 kb in size, which divide the rest of chloroplast genome into an 80–90 kb large single copy (LSC) region and a 16–27 kb small single copy (SSC) region.
    [Show full text]