J. Phytol. Res. 19(l) i7782,2006 NODAL ANATOMY OF SOME TIITHERTO UNSTUDIED

H.A.THAKUR and D.A.PATIL Department of Botany, H.P.T" / R.Y.K. College, Nashik-422005 (M.S.), lndia. *Post Graduate Department of Botany, S.S.V.P Sanstha's L.K.Dr P.R.Ghogrey Science College, Dhule - 424005 (M.S.), India.

Nodal anatomy of twenty-six species belonging to twenty-one genera of the family Euphorbiaceae havqbeen studied. The present study revealed trilacunar, three traced; one lacunar, one traced and pentilacunar, five traced condition in the family. Howeveq the latter two conditions are noted only in two species. Obviously, the trilacunar, three traced condition is more prevalent and hence thought basio for the family Euphorbiaceae. The association of stipules, reduction or amplification of nodal vasculature are pertinently discussed.

Keyfuords : Euphorbiaceae; Nodal organization.

Introduction the stem-axis (Fig.3). The two lateral traces on either side Nodal anatomical studf have been carried out in the family emerge first (Fig.4). The median trace departs higher up. It Euphorbiaceae in pastr'6. The present authors also is prominent (Fig.s). The lateral traces, however, split in extended observatiohs on similar line on eleven their upward course (Fig.6). All the traces extend into the euphorbiaceous speciesT. However, some more taxa have petiole. The nodes are trilacunar, three trace. been investigated whi6h forms the subject maffer of this 3) Aporosa lindleyona(Wight) Baillon : Leaves are altemate- communication. and stipulate. The stem-axis contains a continuous ring of Materialsand Methods vascular tissue (Fig.7). The lateral as well as median trace The collected from various places, like Nakane Dam departs simultaneously (Fig.8). The laterals do not bear (Dhule District, Maharashtra), Peint, Harsul Forest (Nasik vascular tissue for the stipules. The stipules are thus District, Maharashtra), Radhanagari, Dajipur Forest evascular. The laterals in their upward course divide (Fig.9). (Kolhapur District, Mahamshfa), Ootakamund Government All the traces, the median and the laterals extend into the Botanical Garden (Ootakamund, Tamil Nadu), were petiole. The nodes are trilacunar three trace. preserved n 70 % alcohol. Healthy herbarium materials 4) Baliospermum uillqre (WighQ Baillon: Leaves are from S INU Botanical Herbarium Singapore, Rancho Santa altemate and stipulate. Stipules are described as biglandular Ana Botanic Garden Claremont (U. S. A.) and preserved at base in taxonomic accounts. A continuous vascular material from Auckland War Memorial Museum cylinder extend in the stem-axis (Fig. l0). It is the median Auckland (New Zealand) were obtained. trace that emerged first from it (Fig.I I ). The lateral traces Herbarium materials whenever used were first on either side departs a little higher up (Fig. l2). The lateral boiled in water for 5 - l0 minutes. Few drops of acetic acid traces in their upward course however do not send vascular were added to soften and to help recovery of tissue to supply towards the biglandular stipules (Fig.l3). The natural state, with a gap of few minutes after boiling, the laterals and median trace extend into the petiole. The node materials were washed in water and kept ready for next is hilacunar, three trace. stage ofoperation. 5) Bridelia retusq Spreng: Leaves are alternate and Results and Discussion stipulate. A continuous ring of vascular tissue extend in l) Acalypha indica Linn.; Leaves are alternate and the stem-axis (Fig.la). The median trace departs first (Fig. stipulate. The stem-axis contain a ring of vascular tissue l5). The lateral do so a little fartherup (Fig.16). The stipules (Fig.l). A median vascular trace departs a little earlier, do not receive vascular supply from the laterals. They are however, a lateral traces do so a little latter (Fig.2). The evascular. The laterals as well as the median trace extend stipules do not receive vascular supply from the laterals. into the petiole. The petiole is slightly three-pronged They are minute and non-vascularised. Thus, the nodes (Fig.l7).Thenodesarethustrilacunar,threehaced. are tilacunar, three tace. 6) Bridelia stipularis Blume: Leaves are alternate and 2) Agrostistachys indica Dalz: Leaves are alternate and stipulate. The stem - axis contains a continuous vascular exstipulate. A continuous ring of vascular tissue extend in cylinder (Fig.l8.). The median and lateral traces emerge

77 78 Thakur& Patil (Fig.25). The lateral tracegemerges, however, little farther more or less simultaneously fiom it (Fig. 19.). All these tacqs i.. extend upwards and enter into the petiole (Fie.20 '2).tr'...' up (Fig.26). The mediar.,{1d lateral traces extend into the The stipules do not receive vascular supply from the lateral periole (Fig.27). The stipulds do not receive vascular supply taces. Thus they are non-vascular. The nodes are thus from the lateral traces. Thus, they are non-vascular. The three mce. nodes are three lacunar, three trace. trilacunar, 'g) l)Cicca acida Merr.: Leaves are altemate and exstipulate' Drypetes Wnusto (Wight) Pax. & Hoffm: Leaves a ring of vascular tissue are alternate and stipulate. The vascular supply of The stem-axis contains continuous ' (Fig.22). The laterals as well as the median trace emerge stem-axis contains a continuous vascular cylinder from simultaneously from it (Fig'23)' They extend upwards into (Fig.28). It is the median trace that departs first a farther up the petiole (Fig.23). The nodes are three lacunar, three trace' it (Fig.29). The lateral traces depart little the 8) lawianas Hook: Leaves are altemate (Fig.30). They do not bear vascular supply to three and stipulate. The stem- axis contains a continuous stipules. Stipules are evascular. The nodes are vascular cylinder (Fig.zq. The median trace departs first lacunar, three trace.

3-Lacunar,3-traced 3-lacunar,3-traced stipules absent stpules present and e.g. AgrostistachYs indica wrthout vascular suPPlY Blachia denudata e.g. Acalypha indica Cjcca acida Aporosa lindleyana Euphorbid helioscoPia Baliospermum axillare Excaecaria bicolor Bridelia retusa Hura crepitans Bridelia stipularis Jonnesia princiPes Dimorphocalyx lawianus Sapium insigne Drypetes venusta Securinega virosa geniculata Trewia polycarPa Euphorbia heterophylla Euphorbia leucocephala Glochidion hohenckeri Glochidion neilgherrence polyandrus Homalanthus populifolius 1 - Lacunar;"tr: taced atd Homonoia riparia stipules absent Jatropha panduraefolia e.g. Simmondsia chinensis Kirganelia reticulata Mallotus phillippensis Mallotus stenanthes Manihot esculenta 1-Lacunar,1-traced Neoscortechinia kingii and stiPules without Tragia involucrata vascular suPPlY e.g. Ac{ephila excelsa

5-Lacunar,5-traced I - Lacunar, I -traced and stipules without and stiPules without vascular supply vascular suPPIY e.g. Macaranga peltata e.g. Breynia nivosa

3 - Lacunar, 3 - traced and stipule with vascula r suPPlY. e.g. Ghrozophora rottleri

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Abblwations M - Median Trace. L -LateralTrace. Legends for Figures 10-13' 1 -2.Acalyphaindica, Fig. 3-6.Agrostistachysindica, Fig. 7'9'Aporosalindleyona'Fig' Fig. 22'23.ciccaacida, Baliospermumuillare,rig.'r+ -il.grid"liir"tusa,Fig. 18-2l.aiidrliostipularis,Fig- Drypetes venusta, Fig' 3l -33' Euphorbia geniculata' Fig. 24 -27, DimopnocZryx hwianus, Fig. 28 - 30, Fig. 39 '42. Homalanthus polyandrus' Fig. 34 -36. Euphorbia helioscopia, rig. si-:s. Euphorb'ia heteropltylla, Fig.50- 52. Huracrepitans, Fig' 53-55' Fig. 43 -46. HomalannirpopitifitiuiRig.4T -49.-Homonoiariparia, 63. Macarangapeltata, Fig' 64 '67' Mallotus Jatrophapanduraefolia, Fig. 56 -Ss.Jonnesiaprincipes, Fig. 59- esculenta, Fig. 75 ' 77 Neoscortechinia phillippensis, Fig. 6g - TllMoilotu, stenanthei, rig. lz - li . tvtanihot ' Fig' 83 85' Trewia polycarpa' kingii,Ftg.78 - 80. Sapium insigne,Fig. 8l -82. Si**ondsia chinensrs, - J. Phytol. Res. 19(l) :77-82,2006 8l ll) Euphorbia geniculataLinn. : Leaves are alternate and extend upwards in the petiole. The lateral traces do not exstipulate. The vascular tissue ofthe stem - axis contains provide vascular supply to the stipules (Fig. 54 -55). The a continuous ring of vascular tissue (Fig.3 I ). The laterals stipules are evascular. The nodes are thus trilacunar, three and the median trace depart simultaneously (Fig. 32). They trace. extend upwards into the petiole (Fig. 33). The nodes are 19) Jonnesia principes Vella.: Leaves are altemate and trilacunar, three race. exstipulate. A continuous ring ofvascular tissue is present 12) Euphorbia helioscopia Linn. : Leaves are alternate in the stem - axis (Fig. 56). The three traces - two laterals and exstipulate. The stem axis contains a continuous ring and a median one emerge simultaneously from it. (Fig. 57). of vascular tissue (Fig.34). All the three traces - a median The laterals as well as the median one continue in the petiole and two laterals, departs simultaneously from it (Fig'35). in their upward course (Fig. 58). The nodes are thus They extend upwards into the petiole (Fig.36). The nodes trilacunar, three trace. are trilacunar, three trace. 20) Macaranga peltataMuell. - Arg.: Leaves are alternate 13) Euphorbia heterophylla Linn.: Leaves are altemate and stipulate. The stem axis contains a wavy ring of and exstipulate. A continuous ring of vascular tissue continuous vasculartissue (Fig. 59). Five traces are resolved extend in the stem - axis (Fig.37). The laterals as well as upwards but below the nodal region. They ultimately median trace emerge simultaneously (Fig.38). They extend represent as - the median trace and two others on each upwards into the petiole. The nodes are trilacunar three side representing lateral traces - L I , L2 (Fig.60). The median trace. trace and the adjacent lateral trace (Ll) departs slightly 14) Horualanthus polyandrozs Cheesem. : Leaves are early and extends upwards (Fig. 6l). Afterwards the alternase a.nd stipulate. The stem - axis contains a remaining lateral traces (L2) emerge from the vascular ring csntirruaous rfurg of vascular tissue (Fig. 39). The median and then extend upwards (Fig. 62). Thus all the five traces ftace departs first from it (Fig.40). The lateral traces departs - the median one and total four lateral traces continue in little higherup (Fig. al). They do not bear vascular supply the petiole (Fig. 63). The stipules do not receive vascular to the stipules. The stipules are non - vascular. However, supply. The nodes are pentalacunar, five trace. they spli,t in their upward course towards the petiole (Figs. 2l) Mallotus phillippensis (Lamk.) Muell.-Arg.: Leaves 4142).The nodes are trilacunar, three trace. are alternate and stipulate. A continuous ring ofvascular 15) Homalanthus populifolizs Graham. : Leaves are tissue present in the stem - axis (Fig. 64). The lateral on alternate and stipulate. The stem - axis contains a either side emerge first from it (Fig. 65). The rngdian trace continuous ring of vascular tissue (Fig. a3)' The median resolves little upward (Fig.67). The median as well as lateral tace departs first from it (Fig.44). The lateral traces departs traces continue in the petiole (Fig. 66, 67). The stipule do little higher up (Fig. a5). They do not bear vascular supply not receive vascular supply from laterals (Fig . 66,67).They to the stipules. The stipules are nonvascular. However, are non- vascular. The nodes are frilacunar, three fface. they splits ire their upward course towards the petiole (Figs. 22) Mallotus stenqnthes Muell. - Arg. : Leaves are alternate 45 46). The nodes are trilacunar, three trace. and stipulate. A continuous ring ofvascular tissue present 16) Hamonoia ripa ia Lour : Leaves are alternate and in the stem -axis (Fig. 68). The laterals on either side emerge stipulate. The stem-axis contains a continuous ring of frst from it(Fig.69). The medianfiaceresolves little upward vascular tissue (Fig. 47). The median trace departs first (Fig 70). The median as well as lateral traces continue in the from it (Fig. 48). The Lateral traces emerge little higher up petiole (Fig. 70 -71). The stipule do not receive vascular (Fig.a8). All the three traces extend in the petiole. The supply from laterals (Fig.70 -71). They are non-vascular. stipules however do not receive vascular supply from the The nodes are trilacunar, three trace. lateral traces (Fig. a9) and thus are evascular. The nodes 23) Manihot esculenta Crantz.: Leaves are alternate and are frilacunar, three tace. stipulate. The stem - axis contains a continuous vascular 17) Hura crepitans Linn. : Leaves are alternate and cylinder (Fig.72). The two lateral traces and median one exstipulate. A continuous ring ofvasculartissue is present depans simultaneously (Fig. 73). They continue upward in the stem - axis (Fig.50). The lateral as well as median course in the petiole (Fig. 73-:74). The lateral traces do not trace departs more or less simultaneously from it (Fig'S I ). provide vascular supply to the stipules, thus stipules are All the tkee fiaces - two laterals and median one, continue evascular. The nodes are trilacunar, three faced. in the petiole in their upward course (Fig. 52). The nodes 24) Neoscortechinia kingii Hook. f.: Leaves are altemate are fiilacunar, three tace. and stipulate. The stem - axis contains a continuous ring 18) Jatropho panduraefolia Andr, : Leaves are altemate of vascular tissue (Fig. 75). The lateral as well as median and stipulate. The stem-axis contains aring of continuous trace departs more or less simultaneously (Fig. 76). The vascular tissue (Fig. 53). The lateral traces and the median lateral traces do not bear vascular supply to the stipules departs simultaneously from it (Fig. 5 )' All the tlree traces (Fig.76 A7). The stipules are evascular. All the three trace n Thakur& Patil continue in the petiole upwards (Fie. 7'l). The nodes are appear evolved in three different ways. First, onty the thus trilacunar, three trace. stipules are lost and their corresponding laterals are not 25') Sapium insigne Benth.: Leaves are alternate and suppressed. Second, both the stipules and their exstipulate. The stem - axis contains a continuous ring of corresponding laterals are suppressed totally culminating vasculartissue (Fig. 78). It is the median trace that departs into unilacunar nodes with exstipulate condition. Third, first from it (Fig.79\. The lateral traces departs a little higher sometimes only the corresponding laterals are lost, and up (Fig. 80). All these continues in their upward course the stipules are still present. This is also noted in some into the petiole. The nodes are trilacunar, three trace. Rubiaceaee and celastralean taxar0. Obviously, the 26') Simmondsia chinensis (Link.) C. K. Schneid.: Leaves unilacunar condition appears a later attainment. The are alternate and exstipulate. The stem - axis contains a conclusions deducted from the earlier account (c/ Thakur continuous ring (Fig. 8l ). A soliary prominent ttace departs and PatilT) and the present one are represented in the flow from it(Fig.82)which continues in thepetiole in its upwards diagram - I. course. The nodes are unilacunar, one trace. Acknowledgement 27)TrewiapolycarpaBenth: Leaves are opposite as well Junior author (l{ A T) is thankful to Dr. S.R.Yadav, as subopposite and exstipulate. The stem axis contains a Department ofBotany, Delhi University, Delhi for his help continuous ring of vascular tissue (Fi9.83.). median and in identification and collection of some plant materials. He two lateral mces departs simultaneously (Fig. 84). They is gratefulto Principal Dr. B.M.Patil, L.K.Dr. P.R. Ghogrey continue upwards in the petiole (Fig. 85). The nodes are Science College, Dhule and Principal and colleagues of filacunar, three trace. Botany Dept. of H.P.T./ R.Y.K. College, Nashik, for The earlier study by the authorsT revealed a necessary facilities and encouragement. trilacunar, three traced condition more prevalent, whereas References unilacunar, one mce is noted in few euphorbiaceous taxa. l. Sinnott E W 1914, Investigations on the phylogeny of They also reached to a conclusion that the trilacunar, three the angiosperms I.The anatomy of the node as an aid raced condition is basic for the family. The present attempt in the classification ofthe angiosperms. Amer.J.Bot.l includes nodal anatomical information of mature nodes of 343-322.. unstudied twenty-six species belonging to twenty-one Z Von Veh R 1928, Beitrage Kennfiis der Anisophyllum genera of the Euphorbiaceae. Majority of taxa show Euphorbiaceen und einige vergleichnde und trilacunar, three traced condition. An unilacunar, one trace entwickh.mgs schicht liche Untersuchungen uber die condition is observed nsimmondsia chinensis.It is also Druse des Qathiuns . AnnJ ard Bot. Buit en 38 I 3 1 - I 43. to be noted that in Macaranga peltata the nodes are 3. Reynolds 1942, Development of the nodes in Ricinus pentalacunar, five taced. which appears a derived condition communis. Bot.G u. lO4 I 67 -17 0. in relation to the larger size of the leaves. Likewise, the 4. Sharma R 1972, Trends of specialization in the reduction of vascular supply to the leaves in case of Euphorbiaceae I. The node. In: Biology ofthe Land Simmondsia chinensis can be conceived into relation to Plants 78 - 89(Ed.V.Puri, Y.S.Murty, P.K.Gupta, and the size of leaves. Thakur and PatilT, reached to a similar D.Banerjee) Sarita Prakashan, Meerut, India. conclusion in their earlier communication on some 5. Patil D A 1988, Nodal anatomy of seedlings of some euphorbiaceous taxa. Euphorbiaceae IBC 5 l4l-143. The trilacunar, three traced is usually associated 6. Singh A 1992, Studies in Euphorbiaceae I. Trends of with the stipulate condition in majority of species except specialization in nodal Vasculature.l Indian Bot Soc. Agroslistachys indica, Cicca acida, Euphorbia 51 350-355. helioscopia, Euphorbia heterophylla, Hura crepitans, 7. Thakur and Patil 2002, Nodal Vasculature in some Sapiam iwigne and Trewia polycorpa of the present Euphorbiaceae. Swami Botqnical J. Bot. 19 59 - 62, account. It is also generally thought that the leaves with 8. Sinnot E W and Bailey I W 1914, Investigations on the unilacunar nodes are exstipulate8. The taxon viz., phylogeny of the angiosperms 3:Nodal anatomy and Simmondsia chinensis have unilacunar nodes but the morphology ofthe stipules lrzer J.Bot.l 144-459. associated with stipulate condition of leaves e.g. Actephilo 9. Majumdar G P and Pal P K 1988, The stipules of the ucelsoand Breynia nivosa. The stipules are non-vascular Rubiaceae: A Review. Tran. Bose Res. Inst.2257 -68 in Actephila *elsa and Simmondsia chinensis, whereas 10. Shisode S B and Patil D A 2000, Nodal anatomical they are vascularised n Breynia nivosa. It appears that studies in some Celastrales. J. Swamy Bot. CL22 rilacunar nodes, vascularised stipules and stipulate leaves 57-68