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Ontogenetical Shifting Indigenous BLUMEA 23 (1976) 189—201 The vascular pattern in the flower of some Mesembryanthemaceae: Aptenia cordifolia and Dorotheanthusbellidiformis. The effect ofan ontogenetical shifting on the vascular pattern and vascular conservatism C. van der Meulen-Bruijns Afdeling Plantensystematiek, Biologisch Centrum Harcn, Rijksunivcrsiteit Groningen, Netherlands. Contents I. Introduction 189 2. Materials and Methods 190 2.1. Complete flowers 190 2.2. Sections offlowers 191 3. Observations 191 3.1. Aptenia cordifolia 191 3.2. Dorotheanthus bellidiformis 192 4. Discussion 193 4.1. Comparison between the vascular patterns of Aptenia cordifolia and Dorotheanthus bellidi- formis 193 4.2. Comparison with observations of Ihlenfeldt 198 4.3. Comparison with other Centrospermae 199 4.4. The interpretation of the inferior ovary 199 4.5. Conclusions 200 5. References 201 Summary The vascular in the flower various of of and Dorotheanthus 1. pattern at stages maturity Aptenia cordifolia bellidiformis is examined. The vascular ofDorotheanthus has been with that of Dorotheanthus 2. pattern compared Aptenia: typologically, is derived from Aptenia. 3. The vascular pattern of Apteniahas been compared with the observations ofIhlenfeldt: the differences are due of different methods. probably to the use 4. The vascular pattern in the ‘ovary wall’ ofAptenia has been compared with that in the outer floral whorls of Melandrium rubrum the decision which ofinferior (Caryophyllaceae):: itis necessary to postpone as to type in ovary is present Aptenia and Dorotheanthus untill more evidence is available. 1. Introduction The Mesembryanthemaceae (order Centrospermae) are indigenous to South Africa and show more or less succulence. The flowers are characterized by (Ihlenfeldt, Schwantes & —6 Straka, 1962): 1) a calyx with 4 sepals, mostly of unequal size; 2) several whorls of staminodes and semi-inferior petaloid stamens; 3) a plurilocular, to inferior, syncarpous of attached to the ovary, compounded 3—23 carpels; 4) mostly many ovules, placenta by and a very long funicle; 5) characteristic loculicide, hygrochastic capsules. VOL. 190 BLUMEA 23, No. I, 1976 The sub-families of the Mesembryanthemaceae used to be classified in the family Aizoaceae (=Ficoidaceae) (e.g. Eckardt, 1964). However, according to Ihlenfeldt & Straka (1961), they have to be treated as a separate family. The Mesembryanthemaceae can be dividedinto two groups according to the placentation, and into four subfamilies, as follows: group A: axile placentation. 1. Mesembryanthemoideae sensu Ihl., Schw. et Str. (syn. with Aptenioideae Schwant.). 2. Hymenogynoideae Schwant. 3. Caryotophoroideae Ihl., Schw. et Str. basal to placentation. group B: pseudo-parietal 4. Ruschioideae Schwant. The members of to be than those of group A are supposed more primitive group B. both are identical, and have axile (Ihlenfeldt, Very young stages of types placentation the of the Ruschioideae the shifts from 1961). In genera placentation ontogenetically This of axile to basal or parietal (already noted by Huber, 1924). shifting is the result in in changes in relative growth rates the ovary. These changes relative growth rates have known for and Buxbaum illustrated them in the Mesem- been a very long time, (1951) This the basis of the bryanthemaceae (his fig. 27). means that, on carpel concept, it is in- describe Ruschioideae. should be correct to use the term parietal to placentation in the It but still called interpreted as being axile, is pseudo-parietal. the In a study on Mesembryanthemaceae, Ihlenfeldt (1961) made a comparison between the vascular patterns in ovaries with diverse types ofplacentation. Although in his study vascular eluci- some very complicated patterns (e.g. Dorotheanthus) were not completely Ihlenfeldt concluded that the vascular be reduced a dated, various patterns can to common groundplan; and that, with regard to the vascular pattern, the subfamily Ruschioideae derived should be considered to be more than the subfamily Mesemhryanthemoideae. On account of the foregoing it was decided to examine the effect of ontogenetical vascularisation. that the vascular in a flower with shifting on For purpose pattern pseudo- will be with the vascular a flower with axile parietal placentation compared pattern in placentation. As an example of the latter the flower of Aptenia cordifolia (L./) Schwant. (subfamily Mesembryanthemoideae) will be examined. Preparatory studies demonstrated, of Dorotheanthus than from that the vascular pattern is much more complicated appears of the study of Ihlenfeldt (1961). So as an example the group with pseudo-parietal placen- tation the flower of Dorotheanthus bellidiformis (Burm./) N. E. Br. (subfamily Ruschioideae) will be examined. 2. MATERIALS AND METHODS From plants, cultivated in Hortus de Wolf, Haren (Gr.), flowers at various stages of in maturity (buds, flowers, and fruits) were collected, and preserved ethyl-alcohol 70% FPA Examination carried flowers and sections of flowers. or 70%. was out on complete on 2.1. Complete flowers. bleached with and the elements the 1. Preserved specimens were NaOH lignified in vascular bundles stained with basic fuchsin (Fuchs, 1963). 2. Preserved specimens were bleached with lactic acid and the vascular bundles (phloem and xylem) stained with lacmoid (Aloni & Sachs, 1973). C. van der Meulen-Bruijns: The flower of some Mesembryanthemaceae 191 2.2. Sections of flowers. in Preserved specimens were embedded paraplast according to the standard technique, via TBA-dehydration. The thickness of the transverse and sections longitudinal was 7 fx. The sections were stained with astra blue-auramine-safranine (Maasz & Vagas, 1963). 3. OBSERVATIONS is it 3.1. Aptenia cordifolia tetramerous, has sepals of unequal size, two bigger and two smaller ones. The ovary, with four locules and four complete septa, is inferior in the the fruit axile. young flower, in ripened it is semi-inferior. Placentation is Vascular architecture and 3.1.1. (fig. 1 3). In the pedicel four vascular bundles, called primary bundles, occur in one circle (A, ascend in wall 1 in between the Two of them fig. 1-1). They the ovary ) septa. do not branch and supply the central portions of the two bigger sepals. The other two bundles A give off two bundles each (B, fig. 1-2), and ascend towards the smaller sepals. The four bundles B are situated inside thecircle ofprimary bundles, and give off three bundles each the floral base and The bundles and in (Ci C2, fig. 1-3; B', fig. 1-4). Ci C2 form commissure C The bundles B and B' ascend in the wall in a (fig. 1-3). ovary the radii. On the level of the rim the bundles B form commissure septal hypanthium split up, a The with the bundles A, and supply the lateral portions of thesepals (fig. 1-9, 10). bundles B' form which rise the staminodes a commissure, gives to many bundles, supplying and stamens (fig. 1-8, 9, 10). Commissure C gives rise to four bundles which ascend in the radii of the bundles A (Ci,2, fig. 1-4). Higher up the bundles Ci,2 split into three bundles each (fig. 1-8), the into central one (C3) enters finally the style. The styles are situated in between the septa, with one vascular bundle each (C3, fig. 1-10). The other two bundles, branching several form the roofof times, a plexus in the ovarian cavity (Ci,2, fig. 1-9). This plexus gives rise in in lower to some bundles, descending theseptal radii, ending blindly the part of the ovary (C', fig. 1-7). Commissure C gives rise to another eight bundles (D, fig. 1-4). The two bundles D at both sides ofone bundle B fuse and run centripetally in the septal radius into the central column (fig. 1-4,5). In the central columnthese four bundles D ascend, still in the septal off branches in the half of the radii (fig. 1-6), giving some upper ovary (E, fig. 1-7) supplying the placentae. So each bundle D gives off some bundles E into the placentae in the two adjacent locules. (A placenta is a narrow strip of tissue on the central column, the the locule. There of ovules borders projecting into are two rows on it.) Furthermore, the there each in placenta are two conspicuous parallel bundles, that are joined to other branches. The bundles by fusion or are connected by small placentary are connected with the bundles E (fig. 1-7). The vascular supply in the funicles is derived from the placentary in the central column the bundles D unite bundles. High up split and with the placentary bundles. Generally, there is not much variation in the vascular architecture in various flowers. it is in A representative pattern can thereforebeconstructed; represented a three-dimensional model in fig. 3. Since is in wall' and *) there no unanimity the definition of the terms 'ovary 'hypanthium'(see e.g. Eames, Merxmiiller and Puri, in the the 1961; Leins, 1972; Leins, Sattler, 1972; 1951, 1952), present study term 'ovary wall' is used to indicate the total wall outside the ovarian cavity. BLUMEA VOL. No. 192 23, I, 1976 of the vascular bundles. 3-1.2. Orientation of is turned the In the centripetally running part bundle D, xylem downwards, cen- orientated The bundles are halfinverted. the bundles trally ascending part is normally. E In towards each E, innervating one placenta, the xylem strands are turned other. The bundles in the placenta seem to be amphicribral. The remaining bundles A, B, and C have normal orientation. Dorotheanthus has of The inferior 3.2. bellidiformis is pentamerous, it sepals equal size. has five locules and five Placentation ovary complete septa. is pseudo-parietal. Vascular architecture and 3.2.1. (fig. 2 4). vascular called and In the pedicel ten bundles, primary bundles, occur in one circle (A B, A in in fig. 2-1). The bundles ascend the ovary wall between the septa, without branching, and supply the central portions of the sepals. the sometimes In floral base, or at a higher level, the bundles B give off two bundles B in in each (Ci and C2, fig.
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