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Vascular Anatomy of the Flower of Some Species of the Pontederiaceae*

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Vascular Anatomy of the Flower of Some Species of the Pontederiaceae* VASCULAR ANATOMY OF THE FLOWER OF SOME SPECIES OF THE PONTEDERIACEAE* BY V. SINGH (School of Plant Morphology, Meerut) Received June 11, 1962 (Communicatedby Prof. V. Puti, t~.A.sc.) INTRODUCTION TIlE 'pickerel-weed' family Pontederiaceae is a small one of aquatic and marsh plants, comprising 6 genera with 21 species (Willis, 1951), and dis- tributed in warmer parts of the world. The systematic position of the family has long remained one of some controversy. Engler (1886) treated it to be a member of Farinosae because of its endosperm and embryo cha- racters. Schwartz (1930) treated it as being closely related to the Liliaceae, but also considered the dorsiventral form of the flower, the variability and reduction in the androecium and the mealy endosperm characters as of sufficient importance to justify its being placed close to the Commelinaceae and Philydraceae. Hutchinson (1934) placed the plants in the Liliales stating "The Pontederiaceae are a difficult family to place. They appear to me to be aquatic Liliaceae, tending towards the Aroid type, the spiciform inflorescence having a sheath-like reduced leaf (leaf-sheath)". Schwartz (1930) divided the family into three tribes, Eichhornieae, Heterantherae and Pontederieae, the first two with a trilocular ovary and the third with a unilocular ovary. A perusal of the literature reveals that nothing is known about the floral anatomy of this family. Therefore, at the suggestion of Professor V. Purl an investigation of the vascular anatomy of the flower of the Pontederi- aceae has been taken up. The present study deals with the Indian repre- sentatives of the family, viz., Monochoria hastaefolia Presl, Monochoria vaginal& Presl and Eichhornia crassipes Solms. The first two species belong to the tribe Heterentherae and the last to Eichhornieae. Attempts are, however, being made to procure the material of other species. * Research contribution No. 45 from the School of Plant Morphology, Meetut College, Meerut, 339 340 V. S~GH MATERIAL AND METHODS The material of all the three species Monochoria hastaefolia Presl, Monochoria vaginalis Presl and Eichhornia crassipes Solms was collected from Hastinapur--a place situated 23 miles north-east of Meerut (Uttar Pradesh)--in the months of April and September 1960. Flowers and flower- buds of different stages of development were preserved in formalin-acetic- alcohol. The usual customary methods of dehydration and embedding were followed. Serial transverse and longitudinal sections 10-14/, thick weIe cut and stained in crystal violet and erythrosin, which gave satisfactory results. Some floral parts cleared by sodium hydroxide, lactic acid and methyl salicylate and stained with basic fuchsin were also studied. OBSERVATIONS Monochoria External Morphology.--The two species studied are M. hastaefolia and M. vaginalis. They are distributed throughout India and Ceylon extend- ing to Malaya, China, Japan and tropical Africa. The plants are aquatic herbs occurring in marshy places or slow-running streams, and remain attached to the soil by means of a subterranean root stock. The root stock is spongy, elongate and creeping (M. hastaefolia) or short and suberect (M. vaginalis) clothed with brown or purple remains of the old leaf-sheaths, emitting tufts of filiform roots clothed with root hairs. The leaves are radical and solitary at the top of the emerged stems or branches, long petioled, cordate-ovate, sagittate or lanceolate, blade upraised (M. hastaefolia) or floating (M. vaginalis). Inflorescence which is always borne above water is centrifugal (M. hastaefolia) or centripetal (M. vaginalis) and reflex after flowering. The flowers are bisexual, slightly irregular, in racemes or subumbellate, arising from the sheaths of the uppermost leaves and encased by an irregular sheathing bract. The perianth is petaloid, brilliant, purplish blue with red spots, campanulate and subequally 6-partite. The segments are nearly free, the three inner ones obovate and wider than the three outer ones, all with three strong parallel median nerves and reticulately veined between them and towards the margins (Fig. 3). The stamens are six, inserted at the base of the perianth, one is large blue and five are smaller yellow. The filament of the long stamen is spurred. In M. vaginalis spurr is present on one side of the filament but in M. hastaefolia it is either present on one side (Fig. 4) or both the sides of the filament (Fig. 5). The anthers are basifixed, dehiscing introrsely by a terminal slit at length elongating. The ovary is ovoid, sessile, Vascular Anatomy of Flower of Species of Pontederiaceae 34i 3 ~' 2 mm f ' 1 i I ! I ~6 O, mm ~ '~ 3 mm -- 3 T'orn FIos. 1-6. Figs. 1-5. Monoehoriahaataefolia. Fig. 1. Semi-diagrammaticreprescntaticn of the longitudinalsection of the flower showing vascular supply to various organs. Fig. 2. A part of the transverse section of the pedicel magnified to show air-spaces and raphides. Fig. 3. A porianth lobe cleared to show reticulate venation. Figs. 4 and 5. Stamens ,,~owing aFFcndagcs. Fig. 6. Eichhornia crasslpes. Semi;diagrammatic representation of the longitudir.al s¢ctic,n ~f the flower showing vascular supply to various organs. 3-ceUed, each cell with many anatropous ovules arranged in a double series. The style is slender, filiform; the stigma is terminal and minutely three-lobed. The fruit is a membranous loculicidal many-seeded 3-valved capsule enclosed in a persistent twisted perianth. 342 V. S~GI~ Vascular Anatomy of the Inflorescence.--The flowers of M. hastaefolia are long pediceUed arranged spirally on a short, stout, and spongy peduncle. The inflorescence axis has a large number of air spaces and numerous scattered vascular bundles. The vascular bundles are collateral with poor lignification, the xylem is very much reduced and is represented by a few tracheids. The peripheral bundles anastomose and divide to give off a large number of traces for the sheathing bract which encloses the inflorescence. On entering the bract they divide repeatedly formnig a more or less continuous band of vascular bundles. Each bundle is collateral and its xylem faces towards the inner side. There are many air chambers which alternate with the vascular bundles. Many traces for each flower pass away in a spiral manner. Vascular Anatomy of the Flower.--In a transverse section the pedicel shows a single layer of epidermis of closely fitted cells, followed by a few layers of hypodermis consisting of large thin-walled loosely packed ceils. There is a considerable amount of aerenchyma in the cortex. The air chambers are separated by one cell thick partitions (Fig. 2). In the cells of the epidermis, hypodermis and cortex occur frequently the sheaf-like bundles of long acicular calcium crystals (raphides). The vascular supply of the pedicel consists of many collateral vascular bundles with much reduced xylem as typical of hydrophytes. Apparently they may be differentiated into three whorls--outer with fourteen, middle with eight and the innermost with five bundles (Fig. 7). At the base of the receptacle the vascular bundles of the pedicel anasto- mose to form a more or less complete unbroken vascular cylinder in which the xylem is distinct at about six places. The saccate bases of the perianth are first to appear in cross-sections (Fig. 8). The perianth is many layered in the middle, but gradually thins out towards the margin. The epidermis is formed of thin-walled cells with irregularly lobed outline. Stomata occur on both upper and lower surface but they seem to be vestigial. The mesophyll is composed of loosely arranged parenchymatous ceils. There are also many air chambers alternating with the vascular bundles. From the receptacular stele six traces pass out simultaneously (Fig. 9). At a slightly higher level within the cortex of the receptacle each of these traces branches into three, two perianth laterals and one median conjoint perianth stamen trace that immediately branches into the perianth midrib bundle and the stamen bundle (Fig. 10). Thus, each perianth lobe receives three traces. On entcring the perianth the laterals divide to form a large number of bundles which are arranged in a single file in a transverse section. Vascular Anatomy of Flower of Species of Pontedertaceae 345 9 9o s t mm r OC Imm ' 19 17 18 Ftos. 7-20. Figs. 7-18. Monochoria hastaefolia. Figs. 7-16. Diagrams of serial trans- verse sections of the flower from base upwards. In Fig. 14 note the appendages of the stamen, In Fig. 15 except ovary other parts are omitted. Figs. 17 and 18. Transverse sections of the style and stigma respectively of the same flower. Figs. 19 and 20. Monochoria vaginalts. Trans- verse sections of the ovary and the style respectively. In Fig. 19 note the globular masses of cells in the air-c~vitics formed by the stretching of the ovary wall. 3.44 V, SINGH However, the median bundle does not divide and remains prominent. All the bundles are endarch and collateral. Almost at the same level where carpellary dorsals depart from the central cylinder, three lacunae begin to appear outer to the three dorsals (Figs. 10, 11). These lacunae merge with one another, thus separating the corolla stamen tube from the central gynoecium part (Fig. 12). Simultaneously, spaces also appear inner to the perianth bundles sepa- rating the stamen tube from the perianth tube. The stamen tube splits up into six bits which represent the somewhat flattened bases of the filaments of the six stamens (Fig. 13). Each bit contains a single bundle which is con- centric. The filament of the larger stamen has flattened appendages or outgrowths, which continue almost up to the level of the style (Fig. 14). These outgrowths are composed of homogeneous parenchymatous cells and are without any vascular supply. The single stamen bundle runs through the filament into the connective unbranched.
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