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Studies in Geraniale: 1. the Nodal Organization*

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Studies in Geraniale: 1. the Nodal Organization* Prec. Indian Acad. Sci., Vol. 86 B, No. 2, August 1977, pp. 99-106, © Printed in India. Studies in Geraniale: 1. The nodal organization* ASHOK KUMAR School of plant morphology, Meerut College, Mearut 250 001 MS received 21 February 1977 Abstract. A study of the nodal organization of 29 members of this order revealed four nodal types: I. Trilacunar three-traced, II. Bilacunar three-traced, III. Bilacunar two-traced and IV. Unilacunar one-traced. Trilacunar three-traced condition has been observed in 25 out of 29 species investigated. Whereas bilacunar two-traced condition is represnted by some half a dozen species, bilacunar three-traced condition and unilacunarone-traced conditions have been observed only in Geranium robertianum and Linum usitatissimum respectively. It is significant to note that Geranium robertia- hum in addition shows a trilacunar three-traced and bilacunar two-traced conditions. Trilacunar three-traced condition is considered to be basic for this group and it is believed that the unilacunar one-traced condition is derived by approximation and coalescence of the laterals with the median, followed by the obliteration of their gaps. Such a tendency has been observed in some members of this group and a reduction series has been traced. The mode of branching in Geraniaeeae is referred to as sympodial. This has been confirmed by the behaviour of the vascular cylinder at the node. Keywords. Geranium robertianum; Linum usitatissimum. 1. ~u~on The nodal organization of Geraniales has not received much attention in the past. Sinnott (1914) pointed out that this order agrees with Rosales and the great majority of Archichlamydae in possessing, for the most part, a trilacunar nodal structure. Singh (1972) described the nodal vasculature of 18 species of the non-cyatheous Euphorbiaceae. The present study deals with the nodal organization of some 29 species belonging to the families Oxalidaceae, Geraniaoeae, Tropaeolaceae, Linaceae and Zygophyllaceae. 2. Materials and methods The material was collected and procured from different places and the following is the list of the species available for work. The arrangement of families and genera is after Engler and Praatl (1931). Oxalidaceae: Oxalis corniculata L.; Biophytum sensitivum DC., Averrhoa caram- *Research contribution No. 121 99 100 Ashok Kumar bola L.; Geraniaceae: Geranium endressi J. Grey, G. erianthum DC., G. lucidum L.; G. neglectum Carolin, G. nepalense Sweet, G. nodosum L; G. ocellatum Facquem., G. Polyanthes Edgew and Hook. F., G. pusillum Brum. f., G. robertianum L.; G. rotundi- folium L.; G. sanguineum L.; G. sylvaticum L.; G. versicolor L. and G. wallichianum D. Don., Erodium cicutarium L'Herit ex Air., E. moschatum L'Herit ex Ait., E. stephanianum Willd., Monsonia angustifolia E. Mey., Pelargonium australe Facq., P. hortorum Bailey; Tropaeolaceae: Tropaeolum majus L.; Liaaceae; Reinwardtia indica Dum., Linum usitatissimum L.; Zygophyllaceae: Peganum harmala L. and Tribulus terrestris L. All the materials were preserved in 70 ~o alcohol after fixing them in F.A.A. They were dehydrated by passing through alcohol-xylol as well as tertiary butyl alcohol grades and embedded in paraffin wax by the traditional technique (Johansea 1940). Nodes of Geranium endressi, G. neglectum and Biophytum sensitivum were given a prior treatment of 5 % KOH for 12-24 hours. Serial microtome sections were cut from 12-15 microns thick and stained in crystal violet and erythrosia combination. 3. Observations 3.1. Anatomy of the node The nodal anatomy in this group may be classified into four categories: I. Trilacunar three-traced, II. Bilaeunar three-traced, III. Bilacunar two-traced and IV. Unilacuaar one-traced. I. Trilacunar three-traced condition: This is the most prevailing condition observed in some 25 species viz. Oxalis corniculata, Biophytum sensitivum, Averrhoa caram- bola, Geranium erianthum, G. endressi, G. nepalense, G. nodosum, G. ocellatum, G. polyanthes, G. pusillum, G. robertianum, G. sanguineum, G. sylvaticum, G. versicolor, G. wallichianum, Erodium cicutarium, E. moschatum, E. stephanianum, Momonia angustifolia, Pelargonium australe, P. hortorum, Tropaeolum majus, Reinwardtia indica, Peganum harmala and Tribulus terrestris. A cross section of the internode shows either one ring or two flags of vascular bundles (figure 2). However, in Biophytum sensitivum where the iaternodes are very much compressed, a complete vascular cylinder is present (figure 7). In each of these forms three traces are given off in the nodal region from three distinct gaps. Usually the median trace enters as such into the leaf base, however, in certain members as Geranium erianthum (figure 3), G. wallichianum, Tropaeolum, etc. it divides into three branches at the very base and the side branches anastomose with the laterals of their own side. In most of the Geraaiaceae where the leaves are opposite or sub-opposite, (except P. hortorum, where they are alternate) of the two laterals of the leaf, one arises conjointly with the lateral of the other leaf and the second independently (figure 3). However, in G. nodosum and G. wallichianum both the lateral traces of a leaf arise conjointly with those of the other leaf and in Monsonia angustifolia both the laterals for the two leaves arise independently. The stipules which are present in all the Geraniaceae (except Monsonia angustifolia) and Zygophyllaceae, derive their vascular supply from the lateral traces (figure 4). They are, however, non-vascular in Erodium cicutarium. The lateral traces either traverse as such into the leaf base or after branching into two each as in some species The nodal organization 101 ~ z ~e +7/Vil+? .,~ J \':I z'V" I0 \V ~'; II k,3 I [(Fignres 1-15. Captions given in p. 106)] 102 Ashok Kumar O ot G',ZO t "lh~ " Figure 16. Diagrammatic representation of the evolution of the nodal types in Gcranialcs. A, trilacunar three-traced cond.; B, bilacunar three,traced, cond.; C, bilacunar two-traced cond.; D, hypothetical stage leading to the formation of unilacunar one. traced cond.; E, hypothetical stage leading to the formation of unilacunar three- traced cond.; F, unilacunar three-traced cond.; G & H, unilacunar one-traced cond.; H, represents the climax with broad arc-shaped trace. The nodal organization 103 of Geranium, Peganum and Erodium. Biophytum sensitivum is exceptional in that, that both the laterals fuse with the median before entering into the leaf (figure 7). II. Bilacunar three-traced condition: In several nodes of Geranium robertianum, of the two leaves present at a node, orte is typically trilacurtar three-traced and the other shows, that one of the laterals arises conjointly with that of the opposite leaf and the second independently but from the same gap which sends off the median trace (figure 8). In few nodes of this species, it has been observed that one of the lateral traces of a leaf arises from the same gap as the median and the other which arises normally disappears before entering into the leaf (figure 9). In another node, the lateral traces maastomose with the median for a short distance (figure 10). III. Bilacunar two-traced condition: This type of nodal structure is observed in half a dozen species of Geranium, namely G. nepalense (Kumar 1976) G. lucidum, G. neglectum, G. polyanthes, G. robertianum and G. rotundifolium. The nodes of these forms exhibit that generally in the lower leaf only one lateral is present and that the median gives off a prominent branch at its very base which takes the position and function of the missing lateral (figure 11). The stipular supply in all the above species is derived from the lateral traces except in G. lucidum where the stipules are non-vascular. In G. rotundifolium, the median trace gives off two marginal branches which fuse with the laterals of their own side. However, in G. nepalense, G.polyanthes and G. robertianum, both leaves at some of the nodes also show a trilacunar three- traced condition. IV. Unilacunarone-traced condition: This type of node has been observed only in Linum usitatissimum. The single trace that arises leaving a gap in the central vascular cylinder divides into three within the cortex (figures 13 and 14). The lateral branches divide once before entering into the leaf. 3.2. Mode of branching In those members, where the leaves display an alternate arrangement, one axillary branch arises in the axil of each leaf. The mode of branching in Geraniaceae, with opposite or sub-opposite leaves, is of more than passing interest. Here the central vascular cylinder, after giving rise to foliar traces, breaks into three units. The central unit forms the vaseulature of the main axis and the lateral ones, those of the axillary branches (figure 5). The main axis either terminates in a flower or inflorescence as in many nodes of G. ocellatum (figure 15) G. lucidum etc. or its growth is arrested (figure 6). One of the axillary branches then grows more vigourously, thus pushing the main axis to one side and occupying its position. Therefore, these members show sympodial branching (figures 1 and 6). There are two axiUary buds in the axil of one of the two leaves at a node in some nodes of G. robertianum (figure 12) and G. ocellatum. In many species of Geranium instances are, however, not uncommon where there is one axillary bud in the axil of one leaf and none in the axil of the other. 4. Discussion and conclusions This study of 29 members of Geraniales brings out marked variations in the nodal structure, not only among the different families but sometimes even within the same 104 Ashok Kumar species. The nodal structure in this group may be classified into four categories; I. Trilacunar three-traced, II. Bilacunar three -traced, III. Bilacunar two-traced and IV. Unilacunar one-traced. The most prevailing condition is trilacunar three-traced as it has been observed in 25 out of 29 species investigated.
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