R.P.Barykina T.A.Gulanjan, T.V.Klytchkova

Ontomorphogenesis of some herbaceous representatives of the genus PAEONIA L.

2. P. lactiflora Pall. and P. obovata MAXIM.

Vestnik Moskovskovo Universiteta Ser.6 Biol. No.3 p.39-47 (1976) Translation from the Russian of Dr. Carsten Burkhardt, Cottbus

Summary

R.P.Barykina T.A.Gulanjan, T.V.Klytchkova

ONTOMORPHOGENESIS OF SOME HERBACEOUS PAEONIA L. SPECIES.

II.. P.LACTIFLORA PALL. AND P.OBOVATA MAXIM.

These Paeonia species are tuberose herbaceous polycarpics, P.lactiflora - tapracemoseradical, P.obovata - racemoseradical. Tubers are formed as a result of adventitiosus roots thickening, storage tissue is parenchymatous secondary xylem. Monopodial growth of main shoot is changing by sympodial growth in virginal period. Monocarpic shoots are monocyclic, long. Short rhizome has gypogeogenous origin. Axial organs vessel members have scalariform perforation plates. Leaves are mesomorphous shadowy.

For the observation of the development of seedlings of the plants, a material collection was accomplished for the anatomical investigation in mixed forests in the Kavalerovskij Rayon of the Primorskij Kraj and in the BG of the Moscow State University on the Lenin - mountains ; there were explored seedlings, juvenile and mature plants.

Paeonia lactiflora Pall. is a plant of the vacancies and stone slopes , the riverbanks, more rarely the deciduous forests and bushes. It is native in Japan, China, on the peninsula Corea, in Mongolia (Tibet); in the USSR in the Far East and in the Eastern Siberia (Schipczinski, 1937, Ohwi, 1965).

The construction of seedling is underground (Fig. 1, à), which proceeds further ontomorphogenesis similarly like them from P.tenuifolia L. (Barykina et al., 1976). The buds remain in the year of the germination resting or form 1 - 2 longstemned leaves with biternate plastines (Fig. 1, á). The epicotyl is little developed. In the leaf axils of the seedling side buds line up. The hypocotyl is short. The mean root reaches on a depth of 8 cm, forming few thin side roots. According to the supply with nutrients the above ground drive of the hypocotyl and the basis of the mean root thickens . The anatomical construction of the plastines (Tab. 1) and the stems of the first leaf is similarly with that of P. tenuifolia. The knots are 3-lacunar. di-, rarer triarchic. The mean root in the first year begins the secondary thickening and coverage with periderm (Fig. 2, à). In the following 2 - 3 years is formed the terminal bud on the mean stem, the annual increase is unimportant, and is shown on 1 - 2 joined assimilating leaves and 2 - 3 scale-shaped at the basis (Fig. 1, â, ã). In the 4 - year-old age the monopodial sprout-shoot early becomes taken off through a sympodial (Fig. 1. ä). The buds of the renewal are found in the essential in the leaf axis - buds of the lower scale-shaped leaf of the growth of the preceding year. As well as at P. tenuifolia,passes over the further growth the mean root and the hypocotyls in the thickness to the formation spindle-shaped bundle. There develop in his hypocotyl - part side roots, which can thicken also measurably (Fig. 1, ã), the ends of the roots remain relatively thin.

As result of the submersion in the bud at the basis of the above ground stems short vertical small roots line up. On his knot the side roots appears, latter is thickened tuberous. With the time become the small roots more - pointed (Abb.3) and increase clearly in the thickness (Vereshtchagina, 1971). At mature individuals open themselves every year 2 - 5 buds, which are found on a depth of 2 - 3 cm. The other buds of the rootsystem remain resting. On the lower page their lower root-scale subepidermal, displaces with phellogene, forms the layer cork. Along with the stem - side roots, they reach after N.P.Nikolajenko (1958), a length to 100 cm and present themselves as extended spindle-shaped bulge; in the course of the further time the system develops the mean root. As well as at P. tenuifolia, is in the fleshy roots a massive developed greatly parenchymatisised secondary xyleme, but the tracheal elements differentiate themselves further.

The plant blooms in that 5. - 6. Year. Monocyclic are the generative stems, extended, with 4 - 5 lower leaves at younger generative stems and 7 - 10 at more mature. The stems of the future years along with the terminal blossom develop already to fall - beginning . The central cylinder of the stems of the above ground stems are larger, than at P. tenuifolia, their amount is 3 - 6-rows at the primary and 1 - 2-rows at the secondary beam. Relatively short are the segments of the vessels in the xyleme (Tab. 2) with ladder-shaped plastines perforated, including by 2 to 10 (mostly 3 - 5) crossbeams and reticular perforated plastines with 5 - 8 openings. In the primary root there is no well developed angle - collenchyme of the cortical bundles (Fig. 2, á). In the root are basal the knot of the monocarpic stems (annual increase of the rootsystem), above ground ordered and do not unfold a strong expansion of a series of years, the cells of the primary root are thin-walled, collenchyme and bast-fibers are absent. In the fall on the boundary of the central cylinder and the primary root, displaced with phellogene, 4 - 5 layers cells cork with little suberinisised and partially lignifisised thin coverage produced. The primary root dies out after that. The central cylinder contains 8 - 10 accompanying bundles of divided broad mark - beam. In the secondary xyleme of the several years roots a the clear annual increase is visible (Fig. 2, â). The capacity at marks is slight in comparison with the above ground extended internodes.

The plastines of the mature leaves of P. lactiflora are mesoform (Fig. 4, à - â) and thicker, than at the other by us explored species of Paeonia (Tab.. 1). The mesophyllum 8 - 9 - layerd (Fig. 4, á). The rows of the parenchyme put themselves as densely unites there, expanded perpendicular to the surface of the leaf is composed of äëàíå-shaped cells, the sponge is narrow - shovel-shaped. The medium expansion of the vessels and the frequency of openings is slight (Tab.. 1). The accompanying bundles are chlorophyllised. In the lower epidermis against the large vessels short simple small hairs develop. The stems are round with a groove (Fig. 4, ã). The bundles (in the number 6 - 10) are open, the cambium - zone consists in May of 4 - 5 cell layers. Them 2 - 3 lateral bundles, nearest to dorsal, unite themselves with it in form of broad hard connective tissue; Well developed is subepidermal the layer of the collenchyme. The knots are 5-lakunary.

Paeonia obovata Maxim. occurs in small groups in river valleys, in conifer - and deciduous forests in the Far East, in Corea and Manchuria (Komarov, 1950).

The leaf (Fig. 4, å - ç) is mesoform (Tab. 2). The mesophyllum is large-cubic. The row - parenchyme is presented from short long - schaufelshaped cells, which own 1 - 3 furrows, sometimes meet H-shaped cells (Fig. 4, æ). The stems in the centerpiece are round - flügelshaped with 5 large and 4 small bundles, ordered in the wings (Fig. 4, ä); The root zone is narrow. The knots are 5-lacunar.

The particularities the ontomorphogenesis of P.obovata stand more closely to P.lactiflora, than to P.tenuifolia. The construction the seedling is underground. Embryos and young seedlings own a morphological structure (Fig. 5, à, á), which is similar with the structure of P.lactiflora, but the root system is more shallow. Clearly thicken not only them Beside -but also which dies out side roots (Fig. 5, á, â). The system the mean root frequently in early stadiums the Ontogenese. The mature plant owns small number long, thickened clearly expanded side roots, which spread more or less horziontal, essentially in 15 - 20 - cm depth (Fig. 5. ã). The sympodiale renewal the mean stems begins with that 3. - 4. Year , if the rosette does not become form his rung - rosettig. At mature individuals 3 extended flower holding develop every year not more than 2 - stems. In his central cylinder larger compact areas of secondary accompanying web line up, as at P.tenuifolia., and P.lactiflora, (Fig. 2, ã) divided in narrow Markstrahlen; the spring - tracheal elements are broad (Tab. 2), which perforated plastines have mostly with 4 - 5 crossbeams (Fig. 2. ä). In the annual increase the secondary xyleme develop thick fiber - tracheides.

The root (Fig. 2, å) is triarchisch or tetrarchisch. The groups the tracheal elements the secondary xylemes are usually compact, but the mark - beam are more narrow, as at P.tenuifolia and P.lactiflora. The parenchymecells own the derivative pericyclum thickened cellulose - coverages. Along with amyloid grains one meets in the roots small drops of fat and mehrzählige Drusen of Calcium-Oxalat.

It allowed the morphologic - anatomical investigation, P.lactiflora to the tapracemos-radicalen, to associate P.obovata to the racemos - ++ radicalen knollenbildenden herbaceous polycarpicn plants. The bulb lines up as result of the growth the side roots, his basal Vorratsgewebe owns greatly developed parenchyme the secondary xyleme. The form of the bulb varies by spindle - shaped - extends (P.lactiflora) to zylindisch - extends (P.obovata). The construction the underground mean stems is at the juvenilen form in the course of the first 2-3 years rosettig monopodial. monocyclic is extended the flower holding drive, leafy,, with few rootigen underground internodes, closed periderm. The small roots are short and hypogeogen. Woody perforated plastines have The Gliedergefässe the secondary xyleme in the stems, rootsystem and roots. His leaf is mesoform shady?. The knots are 5-lacunig, what correlates with the most his leaf - plastines. The differences in the anatomical structure of both species are basically quantitative nature.

Literature

Barykina R.P.. Gulanjan T.A., Klytchkova T.V. 1976. Ontomorphogenesis some herbaceous representatives of the species Paeonia L. 1. P. tenuifolia L. Vest. Mosk. Univ.», biology .Vs 2. 37 - 44.

Vereshtchagina I.V. 1971. Over the morphology of the underground parts Paeonia. «Bull. GBS AT USSR UdSSR», 78, 7072.

Komarov V.L. 1950. Selected works, vol. 4. M.L., Izd. Akad.der Wiss. the USSR.

Nikolajenko N.P. 1958. Over the underground parts of Paeonia. «Zvetovodstvo»,No.6. 2425.

Schipczinsky N.V.. 1937. The species Paeonia. «Flora USSR UdSSR», vol. 7. M.L..

Ohwi J. 1965. Flora of Japan. Washington.

Assumed from the editing 22.4 1975. Biological faculty Department of higher plants

Fig. 1. P.lactiflora (July, August) : à. á - embryo, â - 2 - year-old and ã,ä - 3 - year-old seedlings, å - underground part one 7 - year-old plant; ý - Endosperm, ãê - main - and ïê - side roots, ï÷ - leaf-axis and òï÷ - terminale bud, ë - assimilation leaf, ÷ë - scale-shaped and îë - dying out leaf, îïá - dying out above ground stems

Fig. 2. Diagram of the cross-section the secondarily thickened mean root (a), stems (á), rootsystem (â) P.lactiflora; stems (ã) and bundle (å) P.obovata: ä - Histological composition the xyleme of the stems 1 - ++ year-old drive P. obovata; ýï - epidermis, ïðä - periderm, ëâ - primary bast-fibers, ôë - phloeme, ê -cambium, âêñ - secondary xyleme, ïêñ - primary xyleme, ïåë - primary and âcë - sekundärå Markstrahlen, íïï - nichtpolare accompanying bundle, ÷ñ - segments of the vessels, òð - tracheides, ïð - hard woody parenchyme, êëí - collenchyme

Fig. 3. P.lactiflora. underground part of a mature plant: íïá - above ground and îïá - dying out stems, ï÷ - bud, êðí - small roots, ïê - side roots

Fig. 4. Leaf of mature plants P.lactiflora (à - ã) and P.obovata (ä - ç). à, å - upper epidermis, â,ç - lower epidermis with surface, á,æ - cutting the leaf plastines, ã, ä cross-section of the stems (medium part)

Fig. 5. P.obovata (July): à - 2 - year-old, á - 5 - year-old, â - 9 - year-old seedlings, ã - underground part a several years of plant; ë - assimilation -leaf, îë - dying out leaf, ï÷ - bud, ãê - mean root, áê - pages - and ïê - side roots, òï÷ - terminale bud, íïá - above ground drive, îïá - dying out stems