Chap. 3. Classification and System in Flowering Plants: Historical Background
HISTORY OF PLANT SYSTEMATICS 역사시대이전의원시시대의분류 Folk taxonomy 먹을수있나?, 해로운가?, 이용가능하나?
Theophrastus Greek philosopher (BC371-BC287) Platon과 Aristoteles의제자 Father of botany "Historia Plantarum" 480 taxa 4 개의식물 group으로인식 : trees, shrubs, subshrubs, and herbs 주요형질들을기록. corolla type( 화관형태 ), ovary position( 자방위치 ), inflorescence( 화서 ). flowering plant( 현화식물 ) 과 nonflowering plant( 은화식물 ) 을최초로구분 그의저서 De Historia Plantarum 에실린이름몇가지는 Linnaeus 가채택 Asparagas, Crataegus, Daucus, Narcisus
Corolla type Passiflora sp.
Ovary position
Amborella trichopoda Baill. Inflorescence
Dioscorides Roman military surgeon (A.D. 1C) Deoscorea : 마의속명 De Materia Medica 600 medicinal taxa 를기록 이책에기록된것만이진짜약초로인식 배열등에대한분류학적 concept 은없었음. Dioscorides가썼던많은이름들이오늘날 속 명으로서쓰인다. Anemone, Dipsacus, Aloe, Aristolochia
Deoscorea 마
De Materia Medica Emperor Flavius Olybrius 가딸인 Juliana 공주에게선물한화보집그림이유명
Albet Magnus Middle age (12 C) De vegetabilis : medicinal plants Provided excellent descriptions ( 기재 ) 줄기의구조를바탕으로 monocots ( 단자엽식물 ) 와 dicots ( 쌍자엽식물 ) 를최초로인식.
Monocots vs. Dicots
HERBALISTS Herbals Renaissance 시대에약용식물을찾기위해만들어진 botanical book Gart der Gesundheit or Hortus sanitatis Herbalist 약용식물채취하여치료하는사람 이시대에 botany = herbalism
Culpeper's Complete Herbal 1812 Edition
Culpeper's Complete Herbal 1812 Edition
Luca Ghini (Italy, 15 C) Sometimes credited with the invention of the herbarium ( 표본관 ) Herbarium: roles the rapid accumulation of knowledge on classification
16C Germany: center of botanical activity German Herbalist: Brunfels, Bock, Cordus, Funchs 분류 와식물의 group 에대한생각들이시작됨 르네상스시대를맞아인쇄술의발달로많은출판물이나왔고, 이로인해본초학이발달.
Transition of the 16 C 신대륙으로뻗어나가많은식물들을새로발견함 Fuchs 500 sp. Bauhin 6,000 sp. Ray 18,000 sp.
Badianus Manuscript An Aztec Herbal, 1552
Badianus Manuscript An Aztec Herbal, 1552
본초강목 ( 本草綱目 ) - 중국명나라의이시진이지은본초학의연구서 - 흙, 옥, 돌, 초목, 금수, 충어등 1892 종을 7 항목에걸쳐해설
Andrea Cesalpino Italian botanist 아리스토텔레스의학풍과이론을이어받은식물학자 De plantis libri : 1,500 sp. trees ( 관목 ), shrubs ( 아관목 ), herbs ( 초본 ) 으로분류하고다시한개의형질에의한이하그룹을인식 downward classification by logical division 32 개의식물 groups 을인식 후세의 Tournefort, Linnaeus, 등에큰영향을줌
John Bauhin and Caspar Bauhin Swiss botanist Pinax theatri botanici: ca. 6,000 plants Synonym ( 동종이명 ) 들을포함. He used names consisting of only two words (before Linnaeus!) 종이모여속이된다는개념을최초로도입.
John Ray English biologist Methodus plantarum nova Historia plantarum New concept of species: define species in the terms of morphology and reproduction Ray and Bauhin: NATURAL CLASSIFICATION ( 자연분류 ) 의시작 the grouping together of those plants that resemble one another
Joseph Pitton de Tournefort French botanist (17 C) Institutiones rei herbariae: 9000 sp. arranged into 800 genera But it was a ARTIFICAL CLASSIFICATION! Only for identification Father of the genus concept Began to develop aggregate system
Carl Linnaeus
Swedish botanist (18 C), professor at the University of Uppsala Father of taxonomy Binominal system ( 이명법 ) of nomenclature referable(~ 에귀착할수있는 ) system referable means that the name of the species immediately indicates the genus into which it is classified Linnaeus was NOT the first to use two names for each species; Bauhin and others had done so, but not consistently. Species plantarum, 1753 Divided plants into 24 classes based on the number, union, and length of stamen ( 수술 ). ARTIFICIAL SYSTEM! simple and easy 그러나독특한수술을갖는몇개의자연적인분류군을이로서인식하게됨
A B C D E F G H I K L M N O P Q R S T U V X Monandria Diandria Triandria Tetrandria Pentandria Hexandria Heptandria Octandria Polyandria Didynamia (2강웅예; 꿀풀과 ) Tetradynamia (4강웅예; 십자화과 ) Monandelphia ( 단체웅예 ; 아욱과 ) Diadelphia ( 이체웅예 ; 콩과 ) Y Z
A Natural System of classification implies that plants presumed to be related are cataloged together. In its original context, the natural system was designed to reflect God s plan of creation and not one of lineages - Artificial classification - Natural classification - Phylogenetic classification
Michael Adanson French naturalist (18 C) Use as many characters as possible 형질의분류학적중요성이라는것은분류후에있는것이지분류전에있는것이아님! 모든형질을동등하게취급 현대 numerical taxonomy의발전에공헌
de Jussieu Family BROTHERS Antoine Jussieu: director of the Jardin des Plantes in Paris Joseph Jussieu: explorer and collector in South America Bernard Jussieu: founder of the Royal Botanical Garden at Versailles THEIR NEPHEW Antoine-Laurent Jussieu: founder of the Musee d Histoire Naturelle de Paris Genera plantarum: 100 group 으로나눔 현재까지도인정받는 group 들이대부분임. Order 라고불렀지만, 현대의 family 와같음. Acotyledons ( 은화식물 ) Monocotyledones ( 단자엽식물 ) Dicotyledones ( 쌍자엽식물 ; 현재의나자식물도포함 )
de Candolle Family Swiss-French family (18-19 C) Taxonomy ( 분류학 ) 이라는용어를최초로만듬 Pyramus de Candolle - Theorie elementaire de la botanique - 분류체계는 Jussieu system 과유사하나 major group 으로 Celluares (nonvascular) Vasculares (vascular plants) 를인식 Alphonse de Candolle (son)
George Bentham and Joseph Hooker (19 C) Genera plantarum 200 families, 7569 genera (1) Polypetale (free petals) 이판화 (2) Gamopetalae (fused petals) 합판화 (3) Monochlamydeae (apetalous) 무판화 - Based on pre-darwinian concept - Widely accepted (Descriptive botany 의표준이죔 ). - British herbaria 인 K, BM 등에서여전히사용되고있음 - 후대에 Bessey 에게큰영향을줌
Charles Darwin The rules of classification by C. Darwin. 1) Organisms have lineages 2) Characters constant over large groups have value in developing classifications 3) Taxa should be based upon correlated complexes of characters 4) Characters should be weighted based on a posteriori information
Phylogenetic system: 계통을바탕으로한분류체계 Eichler - Syllabus der Vorlesungen u"ber Specielle und Medicinisch - Pharmaceutische Botanik (3rd ed.) Engler system 의기초가됨 nonseed plants (Cryptogamae) 비종자식물 algae 조류 fungi 균류 bryophytes 선태류 seedless vascular plants 비종자관속식물 seed plants (Phanerogamae) 종자식물 gymnosperms 나자식물 angiosperms 피자식물 Monocotyledons 단자엽식물 Dicotyledons 쌍자엽식물 Choripetalae (plants with flowers having free petals) 이판화 Sympetalae (those with fused petals) 합판화
Adolf Engler and Karl Prantle German botanists (19 20 C) "Die Natrlichen Pflanzenfamilien" - Genera (by Prantle) "Syllabus der Pflanzenfamilien" -Family "Das Pflanzenreich" - Species plantarum 시도 ( 완성X) Englar System: 현재까지광범위하게받아들여지는분류체계 PHYLOGENETIC CLASSIFICATION SYSTEM ( 진화를반영한분류체계 ) 이판화에서합판화가기원된것으로다룸. 간단한것이원시적이다! 단순한단성화가원시적임 : ex) Salix, Populus (Amentiferae; 유이화서군을원시적으로봄 ) 그러나이것을지지할 fossil data 를발견못함
Charles E. Bessey American botanist (U. of Nebraska) 20C Bessey의 classification system Bentham and Hooker의 system과비슷하나 Ranalian complex (Ranales; Ranunculaceae, Magnoliaceae를포함한그룹 ) 를가장원시적으로취급 진화의방향성에대한 Bessey s dicta를발표
Caltha sp. (Ranunculaceae; 미나리아재비과 )
Bessey 의선인장 tree 합판화 다판화 방사상칭 좌우대칭 자방상생 = 자방하위 자방하생 = 자방상위 자방주생 = 자방중위 이생심피 ( 참조 : 합생심피 syncarpy)
Polypetaly ( 이판화 ) Sympetaly ( 합판화 )
Actinomorphy ( 방사상칭 ) Zygomorphy ( 좌우대칭 )
Ovary position ( 자방의위치 ) 자방상위 superior ovary 자방하생 hypogynous 자방중위 half-inferior ovary 자방주생 perigynous 자방하위 inferior ovary 자방상생 epigynous
Possible evolutionary trends in angiosperms: modified Bessey s dicta 1. - Woody ( 목본 ) herbs, vines, and climbers ( 초본, 덩굴식물, 만목 ) - Perannial ( 다년생 ) biennial, annual ( 이년생, 일년생 ) - Terrestrial seed plant ( 육상종자식물 ) aquatic, epiphytes, saporophytes, parasites ( 수생, 착생, 부생, 기생식물 ) 2. - Collateral vascular bundle ( 병립유관속 )(dicots) scattered vascular bundle ( 산재유관속 )(monocots) - Scalariform endplate, scalariform side wall, appotracheal wood parenchyma를갖는 unstoried wood가 primitive 사다리꼴끝판과측벽, 도관에서떨어져있는유조직을갖는층지지않은목부가원시적 3. - Alternate leaves ( 호생 ) opposite or whorled ( 대생또는윤생 ) - secretory cell, stipule ( 분비세포, 탁엽 ): present absent - simple ( 단엽 ), pinnately veined ( 우상맥 ), evergreen ( 상록성 ) compound leaves ( 복엽 ) 4. - Bisexual flower ( 양성화 ) 는전통적으로 primitive 그러나 unisexual flower ( 단성화 ) 가 primitive 라는견해도많음 ( 최근분자데이타에의한결과는 Amborella가가장원시적임. Amborella는단성화를갖음 ) - leaflike bract ( 엽형의포 ) 의엽액에서나오는꽃은 primitive 5. - 꽃 : many parted ( 부분의수가많음 ), few parts ( 각부분의수가적음 ) - spirally imbricate flower ( 나선복와상 ) whorled valvate ( 윤생, 섭합상 ) - Actinomorphic ( 방사상칭 ) irregular zygomorphic flower ( 불규칙좌우대칭 )
Possible evolutionary trends in angiosperms: modified Bessey s dicta 6. - Perianth ( 화관 ) part 들이분리 ( 갈래꽃 ) fused perianth ( 화관부분이붙은 ; 통꽃 ) - tepal ( 화피편 ) 상태의꽃잎 꽃잎꽃받침의구분된상태 apetalous ( 꽃잎없음 ) 7. - Many separate stamens ( 분리된많은수술 ) few or united stamens ( 적은수의합쳐진수술 ) 8. Hypogeny ( 자방하생 =superior ovary 자방상위 ) perigyny ( 자방주생 =half inferior ovary 자방중위 ), epigyny ( 자방상생 =inferior ovary 자방하위 ) 9. - Numerous separate carpels ( 많은수의분리된심피 ) fewer fused carpels ( 적은수의합쳐진심피 ) 10. - One pore pollen grains ( 발아구가하나인화분 ) three or more ( 셋또는그이상 ) 11. - Arillate ( 가종피 ) 를갖음 : primitive - small embryo( 배가작고 ), abundant endosperm ( 배유가많고 ) nonendospermic ( 무배유성 ) - crassinucellate ovary ( 후주심성자방 : 주심nucellus이비후됨 ) tenuinucellate ovary ( 박주심성자방 : 주심이얇음 ) 참고 : nucellus ( 주심 ) 는 ovule ( 배주 ) 내의 megasporangium wall ( 대포자낭벽 ) 을말함. 12. - Axile placentation ( 중축태좌 ) free central placentation ( 독립중앙태좌 ) 13. - Single fruits ( 단화과 ) multiple fruits ( 다화과 ) - capsule ( 삭과 ) drupe or berry ( 핵과, 장과 ) 14. - leuco-anthocyanins, ellagitamis와같은화합물을갖는식물 : primitive
soc1 and ful : annual to perrenial Siegbert et al. Nnature Genet. 2008 Hutchinson 의분류에서사용된가장큰 group 의구분인목본 / 초본의구분은현재 evodevo 연구에의해서단지두개의유전자변화로바뀔수있는형질임이밝혀짐.
1980~1990 년대의식물분류학계는 Takhtajan 과 Cronquist system 이가장널리받아들여짐
1990 년대말피자식물의분류는분자계통학적연구결과들의축적으로급격한 변화를가져왔으며이러한결과들을바탕으로전세계의많은식물분류학자들 이공동으로참여하여 Angiosperm Phylogeny Group (APG) system 을만들었음. APG system 의특징 - 분자계통학적연구를주축으로한현재까지의모든연구결과들을집약하여만듬 - 한두명의학자들이아닌다수의학자들이참여함 - 앞으로부분적으로는수정되어야할부분이있겠지만 APG system 이보여주는 전체적인피자식물유연관계의윤곽은변하지않고지속되리라예상됨. - 현재우리가배우는교과서 (Judd et al., 2009) 에서도 APG system 에의한배열 을채택함. Angiosperm Phylogeny Group (APG) system http://en.wikipedia.org/wiki/apg_system Angiosperm Phylogeny Website http://www.mobot.org/mobot/research/apweb/
APG system 의기초가되는분자계통학적연구결과들을종합한계통수 Asterales Dipsacales Apiales Aquifoliales Garryales Gentianales Laminales Solanales Ericales Cornales Sapindales Malvales Brassicales Fagales Cucurbitales Rosales Fabales Zygophyllales Celestrales Oxalidales Malpighiales Geraniales Myrtales Saxifragales 범의귀, 돌나물 Caryphyllales 카네이션, 선인장 Santalales 단향, 겨우살이 Beberidopsidales Gunnerales Buxaceae 회양목 Trochodendraceae Proteales 연꽃, 버즘나무 Sabiaceae 나도밤나무 Euptelea Ceratophyllales 국화, 초롱꽃인동, 산토끼꽃당귀, 인삼감탕나무 두충용담, 꼭두서니꿀풀, 금어초토마토, 메꽃진달래, 앵초층층나무, 수국 귤, 단풍나무무궁화애기장대, 무우자작나무, 상수리박, 베고니아장미, 느릅나무콩, 자귀나무남가새노박덩굴굉이밥버드나무, 제비꽃쥐손이풀바늘꽃 미나리아재비양귀비 붕어마름 Campanulids (Euasterids II) Lamiids (Euasterids I) Malvids (Eurosids II) Fabids (Eurosid I) Ranunculales Asterids Rosids Coreeudicots 핵심진정쌍자엽식물 Basal eudicots EUDICOTS진정쌍자엽식물 Acorus Canellales Piperales Magnoliales Laurales Chloranthus Austrobailales Nymphaeaceae Hydatellaceae Amborella 벼, 백합옥수수 창포 후추, 족도리풀목련, 튜립나무녹나무, 아보카도 홀아비꽃대 붓순나무, 오미자수련 MONOCOTS 단자엽식물 Magnoliids Basal Angiosperms 원시 ( 기부 ) 피자식물 EXTENT GYMNOSPERMS 현생나자식물
Amborella trichopoda Baill., the basal-most angiosperm
END