Nodal Organization in Some Species of the Ranunculaceae^
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S a r in , M N. 1960. Physiological studies on S in g h , M., P. L al, a n d K.S. Sinoh. 1971. ■alt tolerance of crop plants XV. Influence Preliminary studies on the salt tolerance of of sodium sulphate on chemical composition moong (Phaseohs aureus Roxb. var. R_s-5) of Cicer arletinum seedlings. Uoydia. 23: seeds at germination stage. Ann. Arid Zone 65-71. 10. 255-260. J. Indian bot. See. 58: 14S-15.1. 1979. NODAL ORGANIZATION IN SOME SPECIES OF THE RANUNCULACEAE^ By M eena K ashyap* Botany Department, Meerut University, Meerut* A bstract The nodal structure of sixteen species of Raounculaccae reveals four types of nodal organization (I) Trilacunar three traced (2) Bilacunar two traced (3) Unilacunar one traced (4) Multilacunar multitraced. Twelve of the species {Ranunculus taelus, R. arvensis, R. muricatus, R. scteralus, R. aqualilis. R. diffusus. Delphinium, ajacix. Clematis triloha, C. grata, C. puberula, C. paniculate!. and Thalictrum sanictilaej'orme) show a trilacunar three traced condition, Bllacunar condition has been iiotcj in Ranunculus auriconus and in one node of Clematis pentculatn. Helleborus faetidus aloiK presents a unilacunar condition. The multilacunar condition is present in Aquilegia Bulgaria with seven traces and in Ranunculus hirtellus with four traces. Based on these obiervatioas a probable couise of evolution in ihc nodal vasculature has been suggested, Trilacunar three traced condition in which all the tlireo traces show branching as in Thalictrum saniculaeforme is thought to be the least advanced. From this bisjc type, unilacunar and multilacunar nodes have been derived through intermediate stages of trilacunar, bilacunar and tetralacunar condition. Clematis with opposite leaves is supposed to be forming a separate line in which reduction of laterals has progrcsssd. The absence of laterals or their extension only into the cortex of the node is taken as an indication how progression has taken place from a trilacunar to bi or unilacunar condition. I ntroduction the node in some Anemoneae to be multilacunar. Most of the Helle- Sianott (1914) while investigating the boreae including sometimes Aquilegia nodal pattern in dicotyledons described were described as trilacunar. Ezelerab and Dormer (1963) made a detailed study of the vascular patterns in Ranunculaceae 1. Accepted for publication on October 12, 1978. and concluded that the family shows such 'Present address: Botany School, Cam diversified characters that it is possible to bridge University, Cambridge, U.K. derive many lines of progress of evolution. 1 am tluinkful to Professor Y.S. Murty for his He found, in almost all the plants he constant intertst in my work and for going studied, a multilacunar or trilacunar tbroitgh the manuscript so critically and making UMfbl suWB*tions and also to Dr Vishnu Kuraai condition at the base and a gradual reduc for help in various ways- tion in the number of traces and gaps higher up reaching unilacunar condition (2) Multilacunar multitraced. in the inflorescence node. With a view (3) Bilacunar two traced. to study the nodal vasculature in the (4) Unilacunar one traced. family in greater detail this work was (1) Trilacunar three-traced.—This is undertaken and the vascular supply of the the most common condition noted in node of sixteen species belonging to six Ranunculus lactus, R. arvensis. R. muricatus, genera has been studied. R. scleratus, R. aquatilis, R. diffusus. Del phinium ajacis. Clematis triloba, C. grata, M aterial a n d M etho d C. puberula, C. paniculata and Thalictrum saniculaeformc. The nodal anatomy of All the material for the present study R. lactus is described in detail and varia was collected or obtained from various tions in other species are recorded where- places. Except for HcUehours foctidus and ever neccssary. Ranunculus auriconus which were collected A transverse section of the intcrnodc by Dr N.P. Saxena in Switzerland, shows a peripheral ring of conjoint coll all other species {Ranunculus lactus, R. ateral endarch bundles of ditrcrent sizes arvensis, R. aquatiHs, R. muricatus, R. (Fig. 1). The leaves arc alternate as such diffusus, R. hirteUus, R. sderatus, R. traces for one leaf leave the vascular scleratus, Clematis, triloba, C. grata, C. cylinder at a particular level. The bundles paniculata, C. puberula, AquHegia vulgaris. present on the side of the appearing leaf Delphinium ajacis, Thalictrum saniculae- show some anastomosing (I'ig. 2). Three farme), are Indian. All the material was traces consisting of one large prominent fixed in F.A.A. After washing with 70% bundle occupying the median position and alcohol it was passed for dehydration in two smaller ones from the lateral position tertiary butyl alcohol and embedded in diverage out from the central va.scularcylin- paraffin wax. Sections were cut 10 to 12(x der causing three gaps (Figs. 3). The two thick and stained with crystal violet large bundles on either side of the median erytJirosin combination. trace show .splitting and form a number of bundles constituting the supply of the O bservations axillary branch (Fig. 4|. Ihe remaining vascular bundles of the node show organi The sixteen species chosen for the study zation forming a complete ring of vascular show variation in their habit i.e., annual bundles that continue into the next inter herbs {Delphinium, Ranunculus) {perennial node. The leaf base containing three herbs {Helleborus), climbing shrubs bundles separates from the node. These {Clematis) aquatic herbs (ii. aquatiHs) bundles remain unbranched within the leaf mostly alternate (Ranunculus, Thalicrum, base. The vascular bundles of the main Aquilegia etc.) or opposite leaves axis and the axillary branch reorganize {Clematis). The genus Thalicitrum also into a ring. Following the separation of shows conspicuous sheathing leaf base as the leaf base the axillary bud also separates well as stipules. , from the main axis (Fig. 5). This type The nodal vasculature of these species of nodal vasculature is exhibited by all can be distinguished into four categories species of Ranunculus listed above and viz. Delphinium ajacis (Fig. 6), (1) Trilacunar three traced. However, in Clematis the leaves are opposite and as such after supplying traces the departure of the leaf traces the bundles for both the leaves the central vascular near the median gap show some branching tissue divides into three parts - the central and organize into a ring of vascular tissue one representing the main axis while the which constitutes the vascular supply of two lateral ones supplying the two axillary of the axillary branch (Fig. 15). The branches of the two opposite leaves. In remaining vascular tissue in the node Clematis triloba the median trace, as soon reorganizes into a ring of bundles and as it leaves the central vascular cylinder, continues in the main axis. divides into three while each lateral divides In Ranunculus hirtellus four traces having into two (Fig. 7). forming in all seven independent gaps leave the vascular traces that enter each leaf base (Fig. 8). cylinder for the leaf (Fig. 16) and all of In one of the nodes of Clematis pani~ them enter the leaf base. The axillary culaia, the median bundle gave oif two branch is produced in the same manner short branches (Figs. 11-13) which did not as in other species of Ranunculus described reach the leaf base. The laterals of the for the trilacunar type. One of the laterals opposite leaves arose on one side con divides into two, as such there are five jointly leaving a single gap, but they bundles at the base of the leaf. The separated immediately outside the gap axillary branch appears bigger than the (Fig. 12). Laterals of the other side main axis (Fig. 17). however, arose independently from (3) Bilacunar two-traced condition.— separate gaps. This type of nodal anatomy is exhibited Thallctrum saniculaeforme that has a by Ranunculus awicanus. In the intcr- sheathing leaf base and slipules, shows yet node there is a ring of many bundles. At another deviation in the trilacunar condi the nodal level a prominent median and tion. The median trace divides into three a lateral trace pass out of the ring of (Tig. 9). The laterals divide and one of vascular bundles and each trace leaves a the two branches supplies the stipule on gap behind (Figs. 18, 19). Although the its side (Fig. 10). node is bilacunar and only two traces are (2) Multilacunar multi-traced condi given out the median branches immediately tion.—Tina condillon exists in Aquilegia and hence the leaf base shows three vulgaris and Ranunculus hirtellus. In the bundles. All these three bundles branch internodc Aquilegia there is a ring of profusely in the leaf base. The production conjoint, collateral, endarch bundles of and supply of the axillary branch is as in different sizes. At the base of the node other species of Ranunculus. the vascular bundles show some reorgani One of the nodes of Clematis paniculata zation. Seven large bundles and one or differed from other nodes which are two very small bundles diverge from the trilacunar and three traced. This node central vascular tissue at the nodal level belongs to the bilacunar type. The leaves (Fig. 14). These bundles enter the leaf being opposite, the median traces of the base as such and do not show any bran two opposite leaves arise on two opposite ching (Fig. 15). The smaller bundles sides of the nq|le. The median, as soon coatinue only for a short distance during as, it differentiates, branches into three their upward course. The leaves being (Fig. 20). The laterals for both the leaves alternate, only one leaf and one axillary do not arise at all on one side while on branch are produced at a node. After the other side they appear very incons Flos. 1-3.