BULL. BOX SURV. INDIA Vol. 12, Nw. 1-4: pp. 92-96, 1870

EPIDERMAL AND VENATION STUDIES IN -I11

Natirmol Botanic Gardens, Lucknow

ABSTRACT Epidermal characters and venation pattern of the leaves of ten specie of Apocynaceae, viz., Agano~macquophyIlata (Roxb. ex Sirns.) C. Don, Beamontia grand&!wa Wight, Catharanthus rosnrs (Linn.) en,.RawrfIb sar&ntina ,%th. ex Kun, R. tr~llaLim., Roq%llicr grata Wall. a Hook., Strophanthur dwarrtah Wall., S. walldur A. DC., Vallari.~solanuma 0. Ktze. and Vwcanga granch~5liaRolfe have been described.

INTRODUCTION ikficro-characters: Epidermaz cells usually penta- The present paper, in continua~ionof the authors' or hexa-gonal, lower almost straight walled or walls studies on epidermal characters and venation slightly curved, upper straight walled. Stomatn pattern in Apocynaceae (Chandra et al. 1969, confined to lower surface, 19.4-21.7-23.4 x 13.4- Kapoor et al. 1969), deals with ten more species. rg.7-23.4p, frequently as long as broad, rubiaceous, &z., Aganosma caryophyllata (Roxb. ex Sims.) stomatal index 14-19-20. Trichomes unicellular or G. Don, Beaumontia grandiflora Wight, Catha- septate (when septate 2- to 4-celled ; Fig. 12b), uni- ranthus roseus (Linn,) Don, Rauvolfia serpentinn seriate, nonglandular, unbranched. Vein islets Benth. ex Kurz, R. tetraphylla Linn., Rozrpelli~z 5-8-10 per sq. mm. Vein endings 14-20-23 per sq. grata Wall. ex Hook., divaricatus mm. Palisade fatio 5-7.8-12. Wall., S. wallichii A. DC., solanacea 0. Ktze. and Voucanga grandifolia Rolfe. Bemumontip ~dilimWight MATERIAL AND METHODS Macro-charrcclers : Leaves opposite, simple, peti- Normal and mature fresh leaves were collected date ; petioles 1-4 cm long, minutely villo-puberu- locally in the month of August 1970. The same lus ; lamina elliptic-oblong, ovate-oblong, obovate- method, as described earlier (Chandra et al,, loc. oblong or oblong, abruptly narrowed at the apex, cit.), was followed. shortly caudate, obtuse or acute, base cuneate or sub- cuneate, often oblique, slightly decurrent down to OBSERVATIONS some length, 12-30 cm xg-18 cm, may or may not AIplloama cpryopbyllPto (Roxb. ex Sims,) G. - Don be undulate, membranous, upper surface glabrous, (A. dichotoma K, Schum.) very dark shining green, midrib and lateral nerves Macro-characters : Leaves opposite, rarely sub- impressed, lower surface villo-puberulus or sparsely opposite, simple, petiolate ; petioles 0.3-2.6 cm long, tomento-villous particularly on the nerves, lighter stout, copper-red, adpressedly puberulus or vilio- green, veins raised, lateral nerves 14-32, distant, puberulus; lamina ovate or elliptic, shortly cau- subparallel, arcuate, looping towards the margin, date, obtuse, acute or acuminate, base subcordate. nervation conspicuous. Young leaves villo-pube- rounded or cuneate, 3.5- 16 cm x 2.5- I I .g cm, margin rulus on the nerves above and tomento-villous on thickened, entire, rather coriaceaus, upper surface the nerves beneath, indumentum rufous-brown. dark shining green, adpressedly puberulus or Very young leaves villo-tomentose on both the sparsely villo-pubescent, midrib, laterals and ter- surfaces. tiaries copper-red, impressed, lower surface paler, Micro-characters : E@dermal cells--lower slightly very sparsely villo-puberulus or glabrescent, mid- sinuous walled, upper straight walled, usually penta- rib, laterals and some of the tertiaries copper-red, or hexa-gonal. Stomata restricted to lower surface, raised, main lateral nerves 6-10, arcuate, upper ones 13.4-18-19.4 x 13.4-16.7-19.4 p, frequently as long as very oblique, nervation conspicuous forming a broad, rubiaceous, stomatal index 9-12-16. Tri- beautiful reticulate pattern. Young leaves copper chomes multicellular (2- to ~g-celled), uniseriate, ting-ed. aon-glandular, unbranched, with or without one or 19701 SHARMA et al. : EPIDERMAL AND VEN more cells collapsed. Veirr islets 4-5-7 per sq. mm. islets 5.7-10 per sq. nun. Vein endings. 7.8-1o per sq. Vein endkgs 19-23-27 per sq. mm. Palisade ratia mm, Palisade ratio 1-3.2-4. 5-8.5-14.

Catlmranthm ~CMHBU (Ikm.) Don (Vinca rosen Linn.) Macro-characters : Leaves opposite, simple, peri- olate ; petioles 0.3-1~2cm long, glabrous or softly pubescent ; lamina elliptic, obovate or oblong- elliptic, obtuse or retuse, mucronulate, base cune- ate or subcuneate, often oblique, slightly decurrent, 2-6.5 cm x 1-3.5 cm, margin entire, may or may not be hairy, inembranous or thinly coriaceous, upper surface puberulus or glabrescent, dark shining green. lower surface pubescent or nearly glabrous, lighter green, main lateral nerves upto 26, rather close, arcuate, nervation not conspicuous. Micro-characters: Epidermal cells-lower sinu- ous walled, upper slightly sinuous walled or walls slightly curved. Stomata on lower surface, 23'4- 25.1-30. 1 x 19.4-20.&23.4~,ranunculaceous. stomata! index 20-22-24, on upper surface 23.+28.1-30.1 x I 8.4-20.8-~3.~p , ranunculaceous and crucifer~us~ shrivelled stomata occasional, stornatal index 6 I 2- I 4, non-stomatiferous areas observed. ~richomes unicellular of septate (when septate a-+-celled ; Fig. 12 C), uniseriate, nonglandular, unbranched. Vein islets ro-11-14 per sq. mm. Vein endings 1-3-7 per sq. Fig. 1-10: Vmation@ttcm: 1. Beatmrontbgrand;flara. 2. Agmosnur mm, areas witti nil vein endings also observed. pali- ~cny&dIa&. 3. Rawolfia serpmfrira. 4. R. i%lra@yUa. 5. Sh+- thus waliichii. 6. S. diuaricotus. 7. Rq19~Uiagrata. 8. CatharanthuF sade ratio 2-2.8-4. rosnu. 9. Valhsolanucea. 10, Vaacang4gr&golia. Fig. 11. Faliar &dermis (lower surface) :Catharanthus rosm. Flg. 12 a-c: TricW: a-Ramvia tatra@ylla (lower surface). b-Agananna carvophrllata (upper surface). c-Catkaranthus rosm (margin). RNIVO~epdiaa Benth, ex Kurz Mauro-characters: Leaves opposite or verticil- he, generally 3- to 4-nately whorled, simple, peti- R. tetr8pbyb Linn. date ; petioles 0.2-2 cm long, glabrous ; larnina lan- Macro-characters: Leaves verticillate, 3- to 4- ceolate, oblanceolate, oblong-lanceolate or elliptic- nately whorled, usually with two unequal pairs in lanceolate, oblong or even obovate, acute or acumi- the whorl, simple, shortly petiolate ; petioles 0.1-0.9 nate, base cuneate, tapering gradually, decurrent, cm long, tomento-villous ; lamina elliptic, elliptic- 6.5-18 cm x 1.~4.~cm, entire, rather membranous oblong, ovate or obovateelliptic, acute, base round- or thinly coriaceous, upper surface glabrous, shining ed or subcuneate, often oblique, 1-5-1 I cm x I .2-5.5 green, lower surface glabrous, pale green, midrib cm, entire, may or may not be undulate, rather prominently raised, main lateral nerves 16-24, slen- coriaceous, upper surface softly pubescent, green, der, arcuate, oblique; lamina green when dry, not shining, lower surface villo-pubescent or nervation not conspicuous, to~nento-villous, paler, midrib prominently raised, Micro-characters: Epidermal cells-lower sinu- main lateral nerves upto 25, very fine, spreading, ous walled, upper almost straightwalled or walls remotely arcuate, nervation more or less conspicu- slightly curved, usually penta- or hexa-gonal. ous. Stomata restrict4 to lower surface, 23.4-27.733.4 x Microcharacters : Epidermal cells--lower slightly 16.,[email protected] p, rubiace0~8 and ranunculaceo~s, sinuous walled, upper straight walled ; usually stomata1 index 12-13-15. Ttichomes absent. Vein penta- or hexa-gonal. Stomata confined to lower 94 BULLXTIN OF THE BOTANICAI, SURVEY OF INDIA p'01. 12 surface, 23.4-3 1-40. I x 19.4-20.7-26.7p rubiaceous and absent. Vein islets 679 per sq. mm. Vein endings ranunculaceous, shrivelled stomata occasional, 23-28-33 per sq. mm. Large rosettes of calcium oxa- stomata1 index 5-14-17. Trichomes multicellular late crystals present in mesophyll. Palisade ratio (3- to 15-celled ; Fig. I za), uniseriate, non-glandular, 8-13-15. unbranched, with or without one or more cells col- S. wpuichii A. DC. (S. dichotoma auct. non DC.) lapsed. Vein islets 6-8-1 I per sq. mm. Vein end- Macrocharacters : Leaves opposite, occasionally ings 12-16-21 per sq, mm. Palisade ratio 3-98 verticillate, 3-whorled, rarely hubopposite, simple, Ro-a grata Wall. ex Hook. (Strophanthrrs petiolate ; petioles 0.3-1 .P cm long, glabrous ; gratus Franch.) lamina elliptic, elliptic-oblong, oblong or obovate, Macrocharacters : Leaves opposite, simple, peti- abruptly narrowed at the apex, shortly caudate, olate ; petioles 0.5-1.5 cm long, nearly glabrous, acuminate, mucronate or obtuse, base rounded, purple tinged ; lamina oblong, elliptic or elliptic- cuneate or subcuneate, slightly decurrent, 2-14.0 crn oblong, abruptly narrowed at the apex, shortly x 1.5-5.5 cm, entire undulate, coriaceous, upper sur- caudate, acute or acuminate, base rounded or sub- face glabrous, dark shining green, midrib impressed, acute, slightly decurrent, 8-18 cmx4.5-7.5 cm, lower surface glabrous, paler, main lateral nerves margin thickened, entire, coriaceous, upper surface 16, arcuate, anastomosing into an intramarginal glabrous, dark shining green, lower surface glab- nerve ; nervation more or less conspicuous. rous, light green, flashed with purple, midrib Micro-characters : EppdermaE cdls on lower sur- raised, purple tinged, main lateral nerves up to 22, face sinuous walled, on upper surface slightly sinu- almost straight or slightly arched uniti.ng in an ous walled. Stomata confined to lower surface, 30.1- intramarginal nerve, nervation not conspicuous. 32.4-33.4 x 19.4-20.443.4 p, rubiaceous, stornat~~lin- Yoling leaves purple beneath. dex 8-12-13. Trichomes absent. Vein islets. 6-8-10 hiicro-characters: Epidermal cells on lower sur- per sq. mm. Vein endilzgs 10-13-13 per sq. mm. face sinuous walled, on upper surface slightly sinu- Large rosettes of calcium oxalate crystals present in ous walled. Stomata confined to lower surface, mesophyI1. Palisade ratio I 6-1 925. 26.7-29.1-33.4 x 19.4-21.4-23.4 p, rubiaceous, stomata2 V- solanaees 0.Ktze. (V. heynei Spreng.) 1-12, Vein I index 9-1 Trichomes absent. islets 57-1 Macro-characters: Leaves opposite, rarely sub per sq. mm.. Vein endings 15-20-25 per sq, mm. opposite, simple, petiolare ; petioles 0.5-2.0 cm long, Large rosettes of calcium oxalate crystals present in glabrous ; lamina lanceolate, oblong~lanceolate, mesophyll. Palisade ratio 4-8-10. elliptic-lanceolate. oblong, elliptic.oblong, or elliptic, StrophWb divaariatug Wall. may or may not be shortly caudate, acuminate or Macro-characters: Leaves opposite or verticillate, mucronate, base rounded or cuneate, slightly de- in latter case 3- to 4-nately whorled, simple, shortly current, 3.5-14 cm x I .56 cm, entire, undulate, rather petiolate ; petioles, 0.3-0.7 cm long, glabrous ; membranous, upper surface glabrous, pelluciddot- lamina oblong or elliptic-oblong, obl~ng-oblanceo- ted, shining green, midrib. prominently and other late, occasionally shortly caudate, qspidate-acumb nerves slightly raised, lower surface nearly glabrous, nate, base cuneate, rarely oblique, slightly decurrent, midrib prominently raised, main lateral nerves 10- 5-12 cm x 2.5-4.5 cm, entire, undulate, coriaceous, 18, arcuate, these and tertiaries fine but distinct, upper surface glabrous darker green than S. nervation more or less conspicuous. New leaves wallich.ii, midrib impressed, lower surface glabrous, light bright green. paler, midrib prominently raised, main lateral Micro-characters : Epidermul cells on lower sur- nerves upto IS, nearly straight or shortly arched, face slightly sinuous walled, on upper surface al- looping near the margin, these and tertiaries also most straight walled or walls slightly curved, usually slightly raised, nervation more or less conspicuous. penta- or hexa-gonal, occasionally tetragonal. Micro-characters: Epidermal cells usually penta- Stomata confined to lower surface, 23.4-24-26.7 x or hexa-gonal, lower almost straight walled of walls 19.4-23.4-26.7 p , frequently as long as broad, rubia- slightly curved or slightly sinuous walled, upper ceous, stomata1 index 14-21-23. Trichomes absent. gtraight walled. Stomata confined to lower surface, Vein islets 79-13 per sq. mm. Vkfz endings 16-22-28 23.4-31-33.4 x 16.7-18.4-23.4 p ,, rubiaceous, rarely ran- per sq. mm. Large rosettes of calcium oxalate crys- ribculaceous, stornatal index 4919.. Trichomes tals present in rnesophyll. Palisade ratio g-13-17:. 19701 SHARMA et al.: EPIDERMAL AND VENATION STUDIES IN A'POCYNACEAE-1x1 95 Voacanga granltolis Rolk. slightly sinuous walled or walls 'slightly curved, Macro-characters : Leaves opposite, in mostly un- upper straight walled, small rosettes of calcium oxa- equal pairs, simple, shortly petiolate ; petioles 0.1-0.5 late crystals present in epidermal cells on both the cm long, glabrous ; lamina elongate-obovate or obo- surfaces. Stomata confined to lower surface, 19.4-21.7- vate-elliptic, oblong-oblanceolate or broadly oblan- 23.4~10-15.4-19.4~, rubiaceous and ranuncula- ceolate, abruptly narrowed at apex, caudate, acute or ceous, shrivelled stomata occasional, stornatal index acuminate, base rounded, often oblique, slightly de- 5-9-17. Trichomes absent. Vein islets 7-10-14 per current, 8.0-20.5 cm x 3.0-8.5 cm, entire, undulate, sq. mm. Vein endings 4-7-12 pr sq. mm. Large membranous, upper surface glabrous, dark shining rosettes of calcium oxalate present in rnesophyll. green, lower surface glabrous, paler, midrib and Palisade ratio 2-3.5-6. lateral nerves prominently raised, main lateral nerves upto 30, nearly straight or arcuate, nervation more or less conspicuous. DISCUSSION Microcharacters : Epidermal cells usually pentii- The important data regarding the present or hexa-gonal, lower almost straight walled or study are sumniarised in Table I.

TABLE 1

STO&TA EPIDERMAI,CELL WALL VENATION S1. Name of species Pali- No. SIZEIN p (Average) ~~TAL INDEX Lower Upper Vein Vein sade Tw* islets endii ratio Lower Upper Lower Upper per sq. .per sq. (Aver- mm mm age) (Aver- (Aver- aec) we)

1. Aganosmcr caryo#hyllata Ru 21.7 x 19.7 Absent 14-19-20 - Almost Straight 8 20 7.8 (Roxb. ex Sims.) G.Don straight ' 2. ~emontiagrandvlora Ru 18 X 16.7 Absent 9-12-16 - Slightly Straight 5 23 8.5 Wight sinuous 3. Cdlaranthus mseus(Linn.) Ra, 25.1 x 20.8 28.1 x20.8 20-22-24 6-12-14 Sinuous Slightly 11 3 2.8 Don Cr sinuous or almost streight 4. Ratmfia stlpantrna Ru, 27.7 x20.4 Abaeni 12-13-15 - Sinuous Almost 7 8 3.2 .' Benth. ex Kun Ra straight 5. R. ktrapI&a Linn. Ru, 31 x 20.7 Absent 5-14-17 - Slightly Straight 8 16 5 Ra sinuous 6. ~cllirrgrata Wall. q Ru 29.1 x21.4 Abaent $11-12 Sinuous Slightly 7 20 8 Hook. - ainuous 7. sh.opkntiuu diwritatus Ru, 31 x 18.4 Absent 49-19 - Almost Straight 7 28 13 Wall. Ra straipht (Rare) qc dqntly smuous 8. S. zvullichii RU 32.4X20.4 Absent 812-13 - Sinuous S.&htly 8 13 19 A. DC. sinuous 9. VaUaris solanuma RU 24~23.4 Absent 1421-23 - Slightly Almost 9 22 13 0. Ktze. sinuous straight 10. Voaeanga grandiYdiu 21.7 x 15.4 Absent 59-17 - Almost Sh:ai&t 10 7 3.5 Rolfe 2 straight or slightly rinuous g6 BULLETIN OF THE BOTANICAL SURVEY OF INDIA {VO~.XZ

In the species studied here, the epidermal cells The venation patterns of all the species have been if straight walled are usually penta- or hexa-gonal illustrated (Figs. 1-10), as had been done in previous like the ones dealt with in the previous communi- com.munications to help identification. The rela- cations of the series (Chandra et al., ioc. cit., Kapoor tively thicker veins in Aganosma caryophyllatn et at., loc. cit.). (Fig. 2) and the narrow and elongated vein islets in The upper epidermal cells are straight or al- Catharanthus roseus (Fig. 8) are worth mentioning. most straight . walled in Aganosm caryophyllata, The trichomes have been observed to be present Beauwtentia grandifiora, Rauvolfia serpentina, R. in four species. In Aganosmn cayophyllata and tetraphylla, Strophanthus divaricatus, Vallaris sola- Catharanthus roseus they are either unicellular or nacea and Voacanga grandifolia, and slightly sinu- few-celled ; and in Beaumontia grandiflora and ous walled in Catharadthus roseus, Roupellia grata Rauzmlfia tetraphylla they are multicellular (few- and Strophanthus wallichiz. However, clearly to many-celled). In the remaining species the tri- sinuous walls have not hen observed in any of the chomes are absent. species studied here ,in case of upper epidermis. The two species of Rauuolfia studied here, viz., The lower :epidermis is dearly sinuous walled in R. serpentina and R. tetraplrylla, can be distin- Catharanthus roseus, Rauvolfia serpentina, Roupel- guished from each other by the number of their lia grata and Strophanthus wallichii. In the re- vein endings per unit area. maining species the cells are almost straight or Similarly the two species of Strophanthus, via., slightly sinuous walled. S. wallichii and S. divaricatus may be distinguished The stomata in all the species except Catharan- on the basis of vein endings per unit area and pali- thus roseus are restricted.to lower surface. a feature sade ratio. also recorded in most of rases reported in previous communications (Chandra et al., loc. cit. and Kapoor ACKNOWLEDGEMENTS et al., loc. cit). The stoma& are generally rubiaceous, but in Rauvolfia serpen%ina, R. tetraphylla, Stro. Authors are thankful to Dr. R. V. Sitholey, Act- phanthus divaricatus and Voacanga grandifolis ing Director, National Botanic Gardens, Lucknow ranunculaceous type have also been observed. A for the facilities, Shri S. N. Vishwakarma for illus- remarkable feature in Catharanthus roseus is the trations and Shri R. S. Ojha for help in collection absence of i-ubiaceous type of stomata and presence of material. of cruciferous alongwith the ranunculaceous ones (Fig. I I). This spdcies possesses stomata on upper REFERENCES surface also. Though the stomatal index on upper &hNDRA, V., S. L. &POOR, P. C. SHMLMAAND L. D. -R. surface varies from 612-14, some non-stomatiferous Epidermal and venation atudiea in Amaceae-I. Bull. bot. areas have also been seen. Shrivelled stomata have Sm. In& 11 :286389, 1969. also been observed occassidnally in this case. Our &AUDHURI, RAI H. N. COmpwative Phacrnwgn~~~ticstuclie on the Ieavts of Cathurdhw WCUJ (Linn.) G. Don and C. data Fable I) on this sp~cies,regarding stomatal @.GIuF (Murr.). G. Don. Indian J. Phmm. 25 (10): 338-341, index, vein endings per sq. mm and palisade ratio, 1963. IDsooa, S. L., P. C. Sy,V. CHANLJRAAND L. D. KAPOOR. however, do not tally with those given by Chaudhuri Epidermal and Venatlon atudies in Apayaacxm-11. Bull. (1963). bot. Sm. India 11 :372-376, 1969.