Floral Morphology
꽃 (Flower): - 피자식물의특징 characteristic of angiosperms. - 특수화된부속지를갖는변형된잎으로생각됨 A flower is a highly modified shoot bearing specialized ppendages that are referred to as modified leaves. 왜식물은꽃을갖는가? Why plants have flowers? 꽃은피자식물의생식기관 정자와난자의결합을매개함. Flowers are the reproductive structures of plants--they mediate the union of a sperm with an egg 꽃은과실로성장 Flowers become fruits 과실은종자를갖음 Fruits contain seeds 종자는산포되어다음세대가됨. Seeds grow to produce the next generation and are the primary means by which individuals of a species are dispersed across the landscape
Goethe 와 Schiller 가활동한독일의 Weimar 궤테 : 은행나무를관찰하여은행나무의생식기관이잎의변형체임을최초로주장.
Does Size Matter? 세계에서가장큰꽃 Rafflesia arnoldii. Amorphophallus titanum 이가장크다고생각할수도있지만이것은꽃이아니라화서임! (1m) The world's largest flower is Rafflesia arnoldii, which can be a meter in diameter (over 3 feet) and weigh 11 kg (24 pounds). - [NOTE: Some web sources list the titan arum, Amorphophallus titanum, as the world's largest flower, but the structure is an inflorescence, not an individual flower.] 가장작은꽃 : Wolffia globosa ( 개구리밥종류 ) (1mm) The world's smallest flower is probably Wolffia globosa, and the entire plant is less than 1 mm long.
앉은부채 (Symplocarpus renifolius Schott ex Miq.) 천남성과 (Araceae) 애기앉은부채 (Symplocarpus nipponicus Makino) 천남성과 (Araceae)
앉은부채는
Floral Terminology Parts Receptacle 화탁 the more or less enlarged or elongated stem axis on which the floral parts are attached Peduncle 화병 the stalk of a solitary, terminal flower (or the stalk of an inflorescence) A Flower Typically has Four Parts 1. Sepals 꽃받침열편 (calyx 꽃받침전체 ) First series; the outermost whorl or spiral of a typical flower (asexual/sterile) 2. Petals 꽃잎열편 (corolla 꽃잎전체 ) Second series; the second whorl or spiral of a typical flower (asexual/sterile); petals alternate with sepals Perianth 화관 collective term for calyx + corolla
3. Stamen(s) 수술 (androecium 웅예군 ) Third series; the third whorl or spiral; the male reproductive unit; stamens alternate with petals and are opposite the sepals Filament 화사 stalk of a stamen Anther 약 pollen producing portion of a stamen (meiosis occurs within the pollen sacs) Pollen 화분 the male gametophytes Connective 약들의연결부위 the sterile tissue connecting the two locules of an anther
4. Carpel(s) 암술, 심피 (gynoecium 자예군 ) Fourth series; the terminal or centermost component; the female reproductive unit (the site of pollination and fertilization); 1 to many separate or fused carpels comprise a gynoecium Stigma 주두 the pollen receptive portion of the gynoecium Style 화주 the elongated portion between stigma and ovary specialized for pollen tube growth Ovary 자방 the basal portion that surrounds and protects the ovules (meiosis occurs within the ovule) ovule 배주 ; embryo 배
1 st whorl 2 nd whorl 3 rd whorl 4 th whorl
Tepal ( 화피편 ): 꽃잎과꽃받침의구분이없는상태
Completeness (refers to flower) Complete 완전화, 갖춘꽃 all four floral series present (Ca, Co, A, & G) Incomplete 불완전화, 안갖춘꽃 one or more floral series absent The four series of a complete flower Perfect (bisexual) 양성화 flower with both functional androecium and gynoecium Imperfect (unisexual) 단성화 flower lacking either a functional androecium or a functional gynoecium: Staminate 수꽃 a male flower; one that has a functional androecium but lacks a functional gynoecium Carpellate 암꽃 a female flower; one that has a functional gynoecium but lacks a functional androecium
A quiz question: Can an incomplete flower be perfect?
An example of incomplete but perfect flower Assarum 족도리풀 Aristolochiaceae
Plant condition (refers to ENTIRE individual plant) Monoecious 단성화를갖는자웅동주 both staminate and carpellate flowers occur on the same plant - Synoecious 양성화를갖는자웅동주 a sexual condition in which the flowers on a plant are all perfect Dioecious 자웅이주 staminate and carpellate flowers occur on different plants
Another quiz question! What plant condition is exemplified by this member of the birch family? Note that the male and female inflorescences are separate but on the same plant
오리나무 Alnus japonica (Thunb.) Steud. Betulaceae
Fusion Distinct similar parts (of same embryonic origin) not fused Connate 동합 similar parts fused Adnate 이합 parts of different origin that are fused Free unlike parts that are not fused There are various other specialized terms that are used for various types of connation and adnation: apocarpous 이생심피, epipetalous 화피상생, sympetalous 합판화, syncarpous 합생심피, monadelphous 단체웅예, diadelphous 이체웅예 Perianth 화관 Apetalous 무판화 lacking petals Sympetalous 합판화 having the petals united (connate) at least at the base Floral tube 화관 (cylindrical, at base), throat 화통 (gradually opening), and lobes 열
여러가지화관의종류
3-merous 3수성, 4-merous 4수성, etc. indicates number of parts in a specific floral series (e.g., a 4-merous flower might have four sepals, four petals, eight stamens, and four carpels) Symmetry: Actinomorphic 방사상칭 (radial, regular) divisible into equal halves by two or more planes Zygomorphic 좌우대칭 (bilateral, irregular) divisible into equal halves in one plane only A few flowers have no plane of symmetry and are called asymmetrical 비대칭
Another quiz question! What is the symmetry of this flower? It can be divided into two equal halves by only one plane.
이축좌우대칭방사대칭 / 방사상칭좌우대칭 / 양면대칭 이종화피 동종화피 일렬오수성이열사수성이열삼수성
Androecium Staminodes 가수술 ; 기능이없는수술 sterile stamens (they do not produce pollen); variable in form and size, may be petaloid or secrete nectar Epipetalous 화피상생 stamens adnate to corolla Alternating with petals or corolla lobes Opposite petals or corolla lobes
2013. Freeman Co.
Amborella trichopoda female flower Amborella 의암꽃의수술은형태적으로는완전한수술로보이지만화분을만들어내지않는다. 그러므로 staminode!
수술 Didynamous 이강웅예 ex, 꿀풀과, 현삼과 Tetradynamous 사강웅예 ex, 십자화과 Diadelphous 양체웅예 ex, 콩과 Monadelphous 단체웅예 ex, 아욱과, 원지과 Syngenesious 취약웅예 ex, 국화과 Epipetalous 화판상생 - 수술의열개종개횡개공개 ex, 진달래속판개 ex, 녹나무속, 생강나무속 - 수술의붙는위치저착측착 T 자착
수술의유합 이생웅예화판상생양체웅예단체웅예취약웅예 약의유형과구성 약의부착 약격 반약 이반약성단반약성저착측착아저착 정자착
약열개의유형 종개공개횡개판개 수술이어떻게진화했을까에대해서는여러가지견해가있지만 Cronquist 는엽형의수술에서약과수술대로분화하는진화의방향성을설명하였다. 이는 Bessey 의진화의방향성에대한 dicta 와도일치한다.
Hypanthium 악통 (floral cup) 꽃잎, 꽃받침과수술이기부에서융합하여통상구조를이룬것이다. 대표적으로장미과식물들이악통을갖는다. Perigynous insertion in black cherry, Prunus serotina
심피의발생 자방 자방 자방실배주주병 태좌 P381 이생심피성합생심피성단심피성 암술군의유합
심피유합의진화과정에대한설명 c.s.: cross section
Gynoecium 자예군 One carpel per a flower (monocarpous gynoecium 단심피성자예 ) Carpels more than 1 per flower carpels distinct (apocarpous gynoecium 이생심피성자예 ) carpels connate (syncarpous gynoecium 합생심피성자예 )
3 개의자방 (ovary), 화주, 주두 1 개의자방, 3 개의화주, 3 개의주두 1 개의자방, 1 개의화주 ( 끝부분만가지침 ), 3 개의주두 1 개의자방, 1 개의화주, 3 개의주두 3 개의심피 (carpel) 가여러가지로 connate 된경우들
Another quiz question! Is the gynoecium of this Hepatica flower monocarpous, apocarpous, or syncarpous? Note the many distinct carpels in the center of the flower.
노루귀 Hepatica asiatica Nakai Ranunculaceae
Insertion ( 삽입 / 부착법 ): 하나의구조가다른부분에붙는방법 SUPERIOR OVARY 상위자방 자방이화관과수술에완전히독립적이면서이들의위에위치함 ( 수술은꽃잎에이합할수도있다 ) - HYPOGYNOUS FLOWER 자방하생 화관과수술이자방 ( 암술 ) 의밑에서나오는것. - PERIGYNOUS FLOWER 자방주생 ( 악통인경우자방이이합하지않는다 ) 화관과수술이자방과이합하지않는악통에서나온다 자방은여전히상위! INFERIOR OVARY 하위자방 자방은화관과수술이붙은자리아래쪽에위치함. 즉화관과수술이자방에이합함. 악통은있을수도있지않은경우도있다. - EPIGYNOUS FLOWER 자방상생 ( 악통인경우자방에이합한다 ) 화관과수술이 ( 화탁이아닌 ) 자방의위에서나온다.
P382 화탁통있음 화탁통없음 화탁통없음 상위자방하위자방반하위자방 자방하생 자방주생 자방상주생자방상생자방중생
자방의삽입 / 부착의위치 (insertion) 유형 자방이화탁또는화축에부착되는위치에따라 Hypogenous flower 자방하생 Perigenous flower 자방주생 Epigenous flower 자방상생 자방의다른부분들 ( 화관, 수술 ) 과의상대적위치에따라 Superior ovary 상위자방 Inferior ovary 하위자방 자방하생상위자방 자방주생상위자방 자방상생하위자방 자방상생하위자방
Quiz questions! Insertion type?
Placentation ( 태좌 / 태자위 ) - 배주는 1 개또는여러개의자방실 (locule) 들로이루어져있다. - 자방실들은격벽 (septum) 으로나뉨 -Placenta ( 태좌 ): 배주가자방벽에붙는자리. Placentation: Placentation 태좌 the arrangement of ovules within the ovary Ovule 배주 the structure containing the female gametophyte; it differentiates into the seed after fertilization Locule 방 the chamber within the ovary; there may be one or more locules Placentae 배주들이붙는위치 the region or line along which the ovules are attached Septum (pl. septa) 격막 an interior wall which separates the locules in those instances where two or more chambers occur (walls=septa)
PLACENTATION TYPES: MARGINAL 변연태좌 only found in a monocarpous or an apocarpous gynoecium; the ovules are attached to the folded margins of the carpel. Monocarpous gynoecium with marginal placentation Apocarpous gynoecium with marginal placentation
AXILE 중축태좌 only found in a syncarpous gynoecium; the placental area of the ovary is attached to an axis derived from the connate margins of the component carpels such an ovary is divided into two or more locules by septa. The ovules are borne along the central axis. PARIETAL 측막태좌 only found in a syncarpous gynoecium; the placental areas are attached to the side walls of the ovary (or extrusions of the wall) such an ovary usually has one locule (therefore no septa). Syncarpous Gynoecia
APICAL 정단태좌 attachment of ovules to the top of the ovary (one locule, no septa) BASAL 기저태좌 attachment of ovules to the botton of the ovary (one locule, no septa) FREE-CENTRAL 독립중앙태좌 attachment of ovules to a free-standing central column in a syncarpous unilocular ovary (one locule, no septa) Basal and apical placentation in syncarpous gynoecia Free-central placentation in a syncarpous gynoecium
Placentation Types (Judd et al. 2009) 측막태좌 측막태좌 Marginal 변연태좌 독립중앙태좌 중축태좌정단태좌기저태좌
Walters and Keil, 1988, Vascular Plant Taxonomy, 3rd ed.
고란초 Crypsinus hastatus (Thunb.) Copel.
Quiz questions! Placentation type?
Evolutionary Development of the Carpel What is a carpel( 심피 )? Basic unit of gynoecium 웅성기 ( 암술 ) 의기본단위 The foliar, ovule bearing unit of a flower that forms either all (monocarpous 단심피 or apocarpous 이생심피 ) or part (syncarpous 합생심피 ) of the gynoecium. 배주를갖고있는잎형태의단위. Leaf-like megasporophyll with infolded or inrolled connate margins bearing one or more ovules on the inside. 접혀진대포자엽으로주변부가서로붙어하나이상의배주를감싼다. Walters and Keil, 1988, Vascular Plant Taxonomy, 3rd ed.
How can the number of carpels comprising a gynoecium be determined? 암술에서심피의수는어떻게알수있나? Count the stigmas or stigmatic lobes Count the styles Count the lobes of the ovary Count the locules in the ovary Count rows of placentae NOTE: Not all are necessarily present or equally useful in a given flower. Extreme connation may cause difficulties.
Quiz questions! Number of carpels?
When examining a flower, ask yourself: What parts are present? How many of each part? What is the shape of each part? Are any parts connate? If so, which? Are any parts adnate? If so, what to what? What are the relationships of parts to each other? FLORAL FOMULAR: See Chap 4, Box 4A
다른교과서 (Plant Systematics, Simpson) 에서의화식 화식은책마다약간씩틀리지만기본적인형식과원리는동일하다. 꽃받침 (K), 꽃잎 (C), 화피편 (P), 수술 (A), 암술 (G) 자방상위 / 하위는따로기재 동합의경우숫자를괄호안에표시 구성요소가여러그룹으로나뉠때는 + 로표시 대활호 [ ] 는드문경우를의미 12개이상다수는 로표시 이합의경우를따로표시안함 예 : K(5)[(4)] C5[4] A5+5[4+4] G5[4], 상위
Pollination Biology ( 수분생물학 ) - Wind pollination : 풍매화 : 가볍고작은화분, 화관이없거나발달안함. 많은양의화분방출 - Animal pollination : 비교적큰화분. 점성, 특수구조등으로곤충및척추동물을통한수분. - Self-incompetability ( 자가불화합성 ): 자신의화분이자신의주두를통해수정되지못하게하는기작시간적, 공간적, 화학적기작 - protogyny ( 자예선숙 ): 수술이먼저성숙 - protandry ( 웅예선숙 ): 암술이먼저성숙 - distyly ( 이화주성 ): 장주화 (pin; 암술이수술보다길다 ) 와단주화 (thrum; 암술이수술보다짧다 ) 를가져단주화의화분은장주화의암술에장주화의화분은단주화의암술에만수정됨 Distyly Tristyly
Inflorescence 화서 : 꽃의축에대한꽃들의배열상태 Parts of an Inflorescence Peduncle ( 화경 ): 화서를지지하고있는줄기 ( 정단화서의경우에도씀 ). Pedicel ( 소화경 ): 화서내에서한꽃을지지하고있는줄기. Rachis ( 화축 ): 신장된화서의주된축. Bracts ( 포 ): 화서나하나의꽃을보호하고있는변형된잎. Bracteoles ( 소포 ) or bractlets: 작은포들또는 2차적포. Involucre ( 총포 ): 꽃이나과실을지지하는포들이환상을이루는것.
Involucre
Sequence of Flowering Indeterminate 무한화서 : 바깥부위부터꽃이성숙하며, 중앙축이신장하며정단에꽃을계속만듬. Determinate 유한화서 : 중앙부위부터꽃이성숙하고축의정단에꽃이달림 ; 축이계속성장하지못함.
Indeterminate Determinate 냉이 (Capsella) Kalanchoe marmorata Crassulaceae ( 돌나물과 )
Indeterminate inflorescence Spike ( 수상화서 ): 화경이없이축에붙어꽃이달림 보리 (Hodeum )
Indeterminate inflorescence Raceme ( 총상화서 ): 화경이있음. 냉이 (Capsella)
Indeterminate inflorescence Panicle ( 원추화서 ): 총상화서와비슷하나가지를침.
Indeterminate inflorescence Corymb ( 산방화서 ): 총상화서와비슷하나꽃이평면에배열하여마치산형화서처럼보임. 마타리 (Patrinia)
Determinate inflorescence Umbel ( 산형화서 ): 여러가지가한지점에서방사형으로뻗어우산의형태로보임. 산형과의특징 Simple umbel Compound umbel 복산형화서 신감채 (Angelica)
Determinate or indeterminate inflorescence Head (or Capitulum 두상화서 ): 화경이퇴화하여줄기정점에많이꽃이밀생함. 국화과의특징 감국 (Chrysanthemum)
Determinate inflorescence Simple (or basic) cyme ( 취산화서 ): 세개의꽃이한곳으로부터나오는유한화서 Dichasial cyme ( 복취산화서 ): 가지를치는취산화서. Silene pratensis Caryophyllaceae ( 석죽과 ) Hypericum cistifolium Hypericaceae ( 물레나물과 )
Inflorescence ( 화서 ) - determinate/indeterminate inflorescence ( 유한 / 무한화서 )
취산화서 원추상취산화서 총상취산화서 신장형취산화서 권산상취산화서안목상취산화서두상화서산형화서
총상화서 원추화서 밀추화서 산형화서 수상화서 두상화서 : 화경이퇴화 불염포 산방화서 속생 육수화서
어떤화서? 육수화서! 어떤식물? 토란 Colocasia esculenta L.
Ament=catkin 유이화서 : 화축이연하여늘어지며, 단성화로구성됨. Ex, 버드나무과 Spadix 육수화서 : 수상화서의변형으로육질의중축에소화경이없는작은꽃들이밀생. 불염포 (spathe) 로둘러싸여있음. Ex, 천남성과 )
P387 수상화서총상화서단산방화서복산방화서원추화서 밀집화서 단집화서 단산형화서 ( 무한화서 ) 단산형화서 ( 유한화서 )
P388 내화영 외화영 윤산화서 배상화서 포영 미상 / 유이화서 육수화서 은두화서 소수화서 두상화서