COMPARATIVE MORPHO-ANATOM1CAL STUDIES OF CYPSELAS OF SIX SPECIES OF THE TRIBE ANTHEM1DEAE...

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BIODIVERSITY

Interrelation between Flora, Fauna and Human

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Biodiversity: Interrelationship between Flora, Fauna and Human

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Proceedings on Biodiversity: Interrelationship between Flora, Fauna and Human Pp. 30-38 (ISBN: 978-81-929410-0-4)

COMPARATIVE MORPHO-ANATOM1CAL STUDIES OF CYPSELAS OF SIX SPECIES OF THE TRIBE ANTHEM1DEAE (COMPOSITAE)

Bidyut Kumar Jana and Sobhan Kr. Mukher jee Department of Botany, University of Kaiyani, Kaiyani-741235, West Bengal, India

Email: [email protected], [email protected]

Abstract

This paper deals with the detail morpho-anatomical structures of cypselas of six spec ies (An themis mar i t ima, A r temis ia vu lgar is , Tanace tum par then ium, Tan ac e tu m vu lgare , T r i p l eu rospe rmum i nodo rum and Tr ip leurospermum mar i t imum ) of the tribe An themideae . Except the cypselas of Tanacetum vulgare and Tr ip leurospermum inodorum, the remain ing 4 studied cypselas are heteromorphic . In the cypsela of An themis mar i t ima, bristelate pappus is present, whereas in remain ing 5 studied cypselas pappus are absent. Sty lopodium is very prominent , en larged and dome shaped in the cypsela of An themis mar i t ima, whereas in remain ing studied cypselas sty lopodia are inconspicuous ly deve loped. In the cypsela of Anthemis mari t ima, Tanace tum vulgare, Ar temis ia vulgar is and Tr ip leurospermum inodorum true carpopod ia are present i.e. histological ly carpopodia l cells are clearly dist inct f rom the remaining part of cypsela. Remain ing 2 studied cypselas, carpopodia l cells are not clearly distinct f rom the remain ing part of cypsela i.e. pseudocarpopod ia . Anatomical ly all the studied cypselas are ell iptical to t r igonos to te t ragonus in cross sect ional conf igurat ion. In the cypse la of An themis mar i t ima and Tr ip leurospermum inodorum, mesocarp ic region is heterogeneous ly deve loped, whereas in remaining 4 s tud ied cypselas, mesocarpic region is homogenously developed. Vellicular cavity is present in the mesocarp ic region of the cypsela of An themis mar i t ima and Tr ip leurospermum mar i t imum. Cypselas of Ar temis ia vulgar is, vel l icular cavity is present at the lower region of testa, near the lateral lobe region of cypsela. Secretary ducts are present in the mesocarp ic region of the cypselas of Tanacetum parthenium. Testal layer is uniser iately d e v e l o p e d in case of all the s tud ied cypse las . In the cypse la of Tr ip leurospermum inodorum and Anthemis mar i t ima endospe rm layers are uniser iately deve lop but in 4 studied cypselas, endospe rm layer is biseriately deve loped. The morpho-anatomica l features of the above studied cypselas have been invest igated to establ ish their potent ial usefu lness in taxonomy.

Keywords : An themideae; Cypselar morpho-anatomy; Compos i tae

Proceedings • 30

In t roduct ion

The fami ly As te raceae is one of the most advanced , h ighly evo l ved fami ly a m o n g the dicotyledons, contain ing 43 tribes, 1 6 0 0 - 1 7 0 0 gene ra and 2 4 0 0 0 spec ies by F u n k e t a l . 1 . The A n t h e m i d e a e is one of the o ldest t r ibe of the As te raceae, w h e r e capsu lar fea tures we re used for better eva luat ion and unders tand ing of tax. Acco rd ing to B remer et al .2, Cypse la prov ides much in fo rmat ion in th is tr ibe, B r e m e r & H u m p h r i e s 3 h a v e m a d e a cladist ic analys is of the gene ra of this tr ibe. T h e u s e of c a p s u l a r f e a t u r e s in t h e c l a s s i f i c a t i o n of A s t e r a c e a e h a s b e e n e m p l o y e d s i n c e t h e w o r k of S c h u l t z -B i p o n t i n u s 4 . K y n c l o v a 5 h a s p r o v i d e d a deta i led invest igat ion of cypse lar s t ructure in s o m e gene ra of the t r ibe An them ideae . The most st r ick ing fea tu res of cypse las of this tr ibe are

a. Ap ica l part of cypse la bears sca les or co rona l ike pappus .

b. Pappus br is t les are usual ly absent

wh i ch are p resen t in o ther major t r ibes of th is fami ly .

In addi t ion to this, apica l part of cypse la p o s s e s s e s p e r s i s t e n t s t y l a r b a s e in assoc ia t ion wi th g landu la r nectary, k n o w n as s ty lopodium. Di f ferences in size, shape, colour of cypse la , or ientat ion of cypse la in relat ion to recep tac le , na tu re of ap ica l ma rg i n of the corona, number and d is t r ibut ion of vascu la r t races and ribs and dist r ibut ion of carpopod ia l cel ls differ f rom taxa to taxa. The present s tudy is a imed to supp lemen t the p rev ious w o r k s ( Bhar and M u k h e r j e e 6, Ka l le rs jo7 etc.), for bet ter unders tand ing of taxa, wh i ch may he lp in r e s h a p i n g t h e e x i s t i n g s y s t e m of c lass i f icat ions. The p resent s tudy dea ls w i th the deta i led s t ructure of cypse la in 6 spec ies b e l o n g i n g t o 4 g e n e r a o f t h e t r i b e An them ideae .

Mater ia ls and M e t h o d s

T h e p r e s e n t w o r k is b a s e d o n the He rba r i um s p e c i m e n s ob ta ined f rom 3 herbar ia of the wo r l d wh i ch w e r e g iven in the t a b l e - 1 .

SI. No. NAME OF TAXA SOURCES

1. An them is mar i t ima L. Botan ischer Gar ten der Univers i tat Zur ich. X X O B R I S S - 2 0 0 5 3 2 6 1

2. A r temis ia vu lgar is L. Botanic G a r d e n Of T h e Universi ty, Jyeangont ie 2, SF- 00550 Helsinki , Finland.

3. Tanace tum pa r then ium Sch. Bip. Botanischer Gar ten der Universi tat Zur ich. X X O Z 19965019

4. Tanace tum vu lga re L. Humbo ld t - Univers i ta t zu Berl in. Ins t i tu t fu r Bio logie. Spezie l le Botan ik u. A rbo re tum. Berl in, Ge rmany .

5. T r i p l eu r spe rmum inodo rum (L.) Sch. Bip.

B o t a n i c G a r d e n of T h e U n i v e r s i t y , Jy rangon t ie 2, SF- 0 0 5 5 0 Helsinki , F in land

6. T r i p l eu rspe rmum mar i t imum (L.) W. D. J. Koch

Humbo ld t - Univers i ta t zu Berl in. Institut fur Bio logie. Spezie l le Botan ik u. A rbo re tum. Berl in, Germany .

Table -1 : S h o w i n g the n a m e of s tud ied taxa of the t r ibe A n t h e m i d e a e a n d their s o u r c e s

Proceedings • 31

For morpho log ica l observa t ion , mature cypse las we re so f tened by d ipp ing in 4 % N a O H solut ion and after that they we re s ta ined in 0 .2 -0 .5% a q u e o u s saf ran in - l ight g reen comb ina t ion to obse rve the di f ferent morpho log ica l parts under s imp le d issect ing m ic roscope and s tereo d issec t ing b inocular m ic roscope. For ana tomica l study, c ross sect ions we re done f rom the midd le port ion

of cypse las , w i th the he lp of sha rpe b lade to observe the d i f ferent ana tomica l reg ions wi th the help of c o m p o u n d light microscope.

Resul ts and D i s c u s s i o n

C o m p a r a t i v e m o r p h o l o g i c a l a n d ana tomica l fea tu res of six s tud ied spec ies w e r e g iven i n . the tab le - 2 and tab le - 3 respect ively.

Table - 2 : S h o w i n g the morpho log ica l fea tures of s t u d i e d cypse las

Anthemis maritima (Fig 1 A-C)

Artemisia vulgaris (Fig 1 D-F)

Tanacetum parthenium (Fig 1 G-H)

Tanacetum vulgare (Fig 11-J)

Tripleurspermum inodorum (Fig 1K)

Tripleurspermum maritimum

(Fig 1L-M)

Cypsela heteromorphic. Ray cypsela 2mm x 0.5 mm; disk cypsela 1.5mm x 0.5mm.

Cypsela heterom6rphic. Ray cypsela 1.5 mm x 0.5 mm; disk cypsela 1 mm X 0.5 mm.

Cypsela heteromorphic. Ray cypsela 1 mm x 0.05 mm; disk cypsela 1.05 mm x 0.05 mm.

Cypsela homomorphic, 2 mm x 0.5 mm.

Cypsela homomor-phic, 2 mm x 0.05 mm.

Cypsela hetero-morphig. ray cypsela 2 mm x 1 mm; disk cypsela 2.05 mm x 0.5 mm.

Ray cypsela yellow brown, oblong, slightly curved; disk cypsela yellow brown, straight, obovate.

Ray cypsela pale brown, wide oblong, slightly curved; disk cypsela pale yellow, ovate, slightly curved.

Ray cypsela pale yellow, straight, obovate; disk cypsela pale yellow, linear-ovate, straight.

Pale yellow, linear, straight. Upper part truncate, lower part rounded.

Black, fistulose, straight, upper part truncate, lower part tapered

Ray cypsela pale yellow, ovate, upper part truncate, lower part tapered

Surface glabrous, containing 9 ribs.

Surface rough and glabrous, containing 4-5 ribs.

Surface rough and glabrous, containing 10 ribs.

Surface rough and glabrous, containing 5 ribs.

Surface glabr-ous, containing 4 ribs

Surface rough and glabrous, containing 3 ribs.

Pappus bristelate type.

Pappus absent. Pappus absent. Pappus absent.

Pappus absent. Pappus absent.

Stylopodium prominent

Stylopodium inconspicuous.

Stylopodium inconspicuous.

Stylopodium inconspicuous.

Stylopodium inconspicuous.

Stylopodium inconspicuous.

Proceedings • 32

Anthemis Artemisia Tanacetum Tanacetum Tripleurspermum Tripleurspermum maritima vulgaris parthenium vulgare inodorum maritimum (Fig 1A-C) (Fig 1 D-F) (Fig 1 G-H) (Fig 11-J) (Fig 1 K) (Fig 1 L-M)

Carpopodium Carpopodium Carpopodium Carpopodium Carpopodium Pseudocarpopodium symmetric. symmetric, symmetric, symmetric, symmetric, bi- present. Carpopodial ring like. carpopodial 4lobed. convex. Carp-cells arranged Carpopodial cells not clearly Carpopodial opodial cells in single row. cells arranged distinct from the cells arranged arranged in 3-4

in single row remaining part in single row. rows. of cypsela i.e. pseudo-carpopodium.

L M

~ 0.C5 mm. C, F, J

1mm. A, B, D, E, G, H, I, K, L, M

Fig.1 Morphology of studied cypselas

Proceedings • 33

A-C-Anthemis maritima: A-Ray cypsela, B- Disk cypsela, C- Carpopodial cells; D-F- Artemi-sia vulgaris: D-Disk cypsela, E- Ray cypsela, F- Carpopodial cells; G-H Tanacetum parthenium: G- Ray cypsela, H- Disk cypsela; l-J- Tanacetum vulgare: l-Cypsela, J- Carpopodial cells; K-Cypsela of Tripleurspermum inodorum; L-M- Tripleurspermum maritimum: L-Ray cypsela, M-Disk cypsela.

Table -3 : S h o w i n g the ana tomica l fea tures of s tud ied cypse las

Anthemis Artemisia Tanacetum . Tanacetum Tripleurspermum Tripleurspermum maritima vulgaris parthenium vulgare inodorum maritimum (Fig. 2A) (Fig. 2B,C) (Fig. 2D) (Fig. 2E) (Fig. 2F) (Fig. 2G)

Cypsela elliptic Cypsela elliptic Cypsela elliptic Cypsela elliptic in Cypsela Cypsela in cross in cross section. in cross section. cross section. tetragonous in tetragonous in section. cross section cross section Cypselar wall Cypselar wall Cypselar wall Cypselar wall Cypselar wall Cypselar wall 0.04 0.03 mm x 0.02 0.04 mm x 0.02 0.03 mm x 0.01 0.04 mm x 0.02 0.04 mm x 0.03 mm x 0,02 mm wide at mm wide at mm wide at ribs mm wide at ribs mm wide at ribs mm wide at ribs ribs and furrow ribs and furrow and furrow and furrow and furrow region and furrow region region region respec- region respec- region respec- respectively respectively. respectively. tively. tively tively

Epicarp Epicarp Epicarp Epicarp Epicarp Epicarp uniseriate, uniseriate, uniseriate, uniseriate, made uniseriate, made uniseriate, made up of thin made up of thin made up of thin up of thin walled, up of thick- made up of thin walled, rectangular, walled, walled, rectangular, walled, rectangu- walled, rectangu- compactely rectangular, rectangular, compactely lar - oval, lar, arranged paren-compactely compactely arranged compactely compactely chyma cells, arranged, arranged parenchyma arranged, arranged provided with parenchyma parenchyma cells, provided parenchy-ma parenchyma cuticle cells. cells,provided with cuticle cells, provided cells, provided

with cuticle. with cuticle. with cuticle

Mesocarp Mesocarp Mesocarp Mesocarp Mesocarp Mesocarp heterogenous, homogenous, homogenos, homogenous, heterogenous, homogenous, containing both made up of only made up of sclerenchyma- containing parenchymatous, parenchyma sclerenchyma parenchyma tous. parenchyma and containing vellicular and scleren- cells. cells containing sclerenchyma cavity. chyma cells. secretary duct. cells. Mesocarpic region containing vellicular cavity

Proceedings • 34

Anthemis maritima (Fig. 2A)

Artemisia vulgaris

(Fig. 2B,C)

Tanacetum parthenium

(Fig. 2D)

Tanacetum vulgare

(Fig. 2E)

Tripleurspermum inodorum

(Fig. 2F)

Tripleurspermum maritimum

(Fig. 2G)

Testa 0.01 mm thick, uniseriate,. parenchyma-tous.

Testa 0.01 mm thick,uniseriate, parenchyma-tous

Testa 0.01 mm thick, uniseriate, parenchymatous.

Testa 0.01 mm thick, uniseriate, parenchymatous

Testa 0.02 mm thick,uniseriate, parenchymatous.

Testa 0.01mm thick, uniseriate, parenchymatous.

Endosperm uniseriate, parenchyma-tous.

Endosperm biseriate, parenchyma-tous

Endosperm biseriate, parenchymatous.

Endosperm biseriate, parenchymatous.

Endosperm uniseriate, parenchymatous.

Endosperm biseriate, paren-chymatous.

Cotyledons conta-ining 6 resin ducts ( 3 ducts in each cotyledon).

Cotyledons containing 6 resin ducts (3 ducts in each cotyledon).

Cotyledons containing 6 resin ducts ( 3 ducts in each cotyledon).

Cotyledons containing 6 resin ducts ( 3 ducts in each cotyledon).

Cotyledons containing 6 resin ducts (3 ducts in each cotyledon).

Cotyledons containing 6 resin ducts ( 3 ducts in each cotyledon).

Fig.2 Anatomy of studied cypselas 0.05 mm A, B, C, D, E, F, G

Proceedings • 35

A- Anthemis maritime, B- Artemisia vulgaris (Through ribs), C- Artemisia vulgaris (Through lateral lobe), D- Tanacetum parthenium, E- Tanacetum vulgare, F- Tripleurspermum inodorum, G- Tripleurspermum maritimum

Detailed morpho-anatomy of cypselas of 6 species of the tribe Anthemideae have been studied. Regarding the cypselas of this tribe, Bremer et al 8 have indicated a valu-able statement "The fruits (cypselas) provide much information in the Anthemideae". Except the cypse la of An them is mar i t ima and Tr ip leurspermum inodorum, the remaining 4 studied cypselas are homomorphic. Colour quality of cypselas are also variable, though it is not a reliable dist inguishing character because it changes with the degree of matu-rity of cypselas. Among the studied cypselas, in c a s e of A n t h e m i s m a r i t i m a a n d Tr ip leurspermum mar i t imum, yel low brown; Tanacetum vulgare , Tanacetum parthenium and Ar temis ia vu lgar is , pale ye l low and Tr ip leurspermum inodorum, brown black in colour. Shape of studied cypselas are vari-able from oblong to linear to obovate. Though size, shape, colour are not important taxo-nomic character, for the separat ion of taxa. Cypselar surface in the Anthemideae is char-acterized by the presence of myxogenous tri-chomes by Kallersjo9. Though, among the studied cypselas, these structures are not observed. Structure of sty lopodium is clearly visible in young cypsela. Therefore, this struc-ture may not be valuable in mature cypsela. In the cypsela of Anthemis marit ima, sty-l opod ium is e l o n g a t e d , d o m e shape , whereas in remaining 5 studied cypselas, stylopodia are inconspicuously developed. M u k h e r j e e l O , has studied the stylopodial characters of some members of Asteraceae. Actually, stylopodia are the persistent style base with associate nectar. At the basal region of cypselas, carpopodium is present. It is a meristematic zone with variable

layers of cellular arrangement. In the cypsela of Tr ip leurspermum inodorum, carpopodial cells are arranged in 3-4 layers whereas in the cypsela of Tanacetum parthenium and T r i p l e u r s p e r m u m m a r i t i m u m , pseudoca rpopod ia are present . In the cypsela of Tanacetum vulgare, Anthemis marit ima and Artemisia vulgaris carpopodial cells are arranged in single row. Actually carpopodium is the basal, abscission zone of cypsela by Mukher jee and Nordenstam 11 and it helps in the detachment of cypselas from the thalamus.

Anatomical ly all the studied cypselas are elliptical to t r igonous or tetragonus in cross sectional configurat ion. Mesocarpic , region exhibits cellular variat ions. In the cypsela of Anthemis mar i t ima and Tr ip leurspermum inodorum, mesocarpic regions are heterog-enous. In remaining 4 studied cypselas, this regions are homogenous ly developed. Heterogenously developed mesocarpic region is also found in some spec ies of the tr ibe L a c t u c e a e by J a n a a n d Mukher jee l 2.Vellicular cavity is present in the mesocarp ic region of cypse la of Anthemis mar i t ima and Tr ip leurspermum mar i t imum whereas in remaining 4 studied cypselas, this cavity is absent. Vellicular cavity is also present in some species (Aster alpinus, Felicia heterophylla) of the tr ibe Astereae by Jana, Bar and Mukher jee l 3. Another important char-acter is the p resence of resin duct in the mesocarpic region of cypsela of Tanacetum parthenium, whereas in remaining 5 studied cypselas, this cavity is absent. Secretary duct is also present in the mesocarp ic region of the cypsela of Elephantopus carol inianus of the t r i b e M o q u i n i e a e by J a n a , Bar a n d

Proceedings • 36

Mukher jeel 4. Presence of vellicular cavity and resin ducts are most important characters on the basis of taxonomic view point. In all the studied cypselas, testal layers are uniseriately develop. Except the cypse las of An themis marit ima and Tr ip leurspermum inodorum, the remaining studied cypselas endosperm lay-ers are biseriately developed.

Key to the studied species

1a. Cypsela homomorph ic ; pale yellow-back (2)

2a. Cypsela pale yel low; surface contain-ing 5 ribs; carpopodial cells arranged in s ing le row; tes ta l layer 0.01 m m in t h i c k Tanacetum vulgare

2b. Cypsela black; surface containing 4 ribs; carpopodial cells arranged in 3-4 rows; t es ta l l a ye r 0 . 0 2 m m in t h i c k n e s s

Tr ipleurspermum inodorum

1b. C y p s e l a h e t e r o m o r p h i c ; ye l l ow brown-pale brown-pale yel low

3a.Vellicular cavity present; secretary duct absent (4)

4a. Surface containing 9 ribs

Anthemis marit ime

4b . S u r f a c e c o n t a i n i n g 3 r ibs

Tr ip leurspermum mar i t imum

3b. Vellicular cavity absent; secretary duct may or may not be present (5)

5a. Surface containing 4-5 ribs; secre-tary duct absent Artemisia vulgaris

5b. Surface containing 10 ribs; secretary d u c t p r e s e n t T a n a c e t u m parthenium

List of abbriviat ions used

Ep- Ep icarp , Me- M e s o c a r p , Pa- Paren-chyma, Scl- Sc lerenchyma, V.C.- Vellicular cavity, S. D.- Secretary duct, T- Testa, E-Endosperm, V. T.- Vascular trace, Pal- Pali-sade parenchyma.

Acknowledgements

We are thankful to Dr. Peter Enz, Curator, Botanischer Garten der Universitat Zurich, Zurich; Curator of Humboldt- Universitat zu Berlin. Institut fur Biologie. Spezielle Botanik u. Arboretum. Berlin, Germany and Aune Koponen, Scientif ic Curator, Botanic Gar-den of The University, Jyrangont ie 2, SF-00550 Helsinki, Finland for sending mature, identif ied seeds for this study.

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Proceedings • 37

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Proceedings • 38