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

Ontomorphogenesis of some herbaceous representatives of the species PAEONIA L.

1. P. tenuifolia L.

Vestnik Moskovskovo Universiteta Ser.6 Biol. 2 p.32-39 (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.

1. P. TENUIF0LIA L.

The results of anatomo-morphological study of Paeonia tenuifolia in the ontogenesis are given. The structure of the seedlings, the juvenile and adult plants, shoots, peculiarities of underground organs formations and age-related changes were discribed. P. tenuifolia is rachisracemoseradical tuberose herbaceous polycarpic. Tubers are forming as result of local thickening of roots, storage tissue is parenchymatous secondary xylem. Monocarpic shoots are monocyclic, long, with some shotened internodes which form rhizome. The leaves are mesomorphous whith some light structure features, nodes are three-, fivelacunar. The secondary xylem tracheal elements of underground organs are more specialised then in shoots; the vessel members have simple perforation.

The species Paeonia L. contains 40 - 50 species (Stern, 1946), spread in Europe, the mediterranean region, in the eastern Asia and the western states of north - America. These several years of plants with thickened fleshy roots form often large shrubs.

The species Paeonia is a relic the old mesophilischen arctic -tertiary flora. Usually are the primitive species in the mountain -forests of the subtropical zones the eastern Asia, on an hight of 23604250 m spreads . In the USSR finds one 15 species (European part, Siberia, long-Far East, Crimea ,Caucasus).

The botanists dedicated their attention since long time the unique Charakteritiska of the construction of the peonies and arranged itas independent group. Before the species Paeonia in the family was arranged the Ranunculaceae (order Ranales) as monotypic stem or monotypic subfamily. Some systematics organize the Paeonia to the Berberidaceae, other to the Magnoliaceae. There were submitted also suggestions, to examine the peonies as independent family. In latter time became the independence of the family Paeoniaceae not only from the systematic persons but also by whom Embryologen, Cytologen, anatomists, Morphologen and biochemists underlined. Some authors count the family Paeoniaceae among the Magnoliaceae, other among the Helleboraceae or Dilleniaceae. A.L.Takhtakshjan (1966) cites the possibility the Einordnung in the order proposed by Nakai (Nakai, 1949) Paeoniales.

The monograph of the species Paeonia of F.C. Stern (Stern, 1946) subdivides the species in 3 sections: Moutan DC., in which the shrubby species are arranged, Paeonia, including the herbaceous species, at which the petals are large, and Onaepia Lindl., which is formed by the herbaceous North American species with unique features the petals and petals . L.M.Kemularia-Nathadse(1961) divided the species Paeonia in 5 sections. She organized the yellow flowerd species in the section Flavonia Kem.-Nath., which raised subsection Dissectifolia Stern of the section Paeonia it in the rank of a section Sternia Kem.-Nath. Many examiners consider the shrubby species as the more primitive the species. So noticed A. Ims (1964) , that the representatives of the section Moutan are charkterisised through a larger amount of carpels and more primitive tracheal elements and tomentum, than in herbaceous peonies. A.L.Takhtakshjan (1948) noticed besides the primate of the woody drive - type at the peonies, ascertained however a lower specialization of the vessels the herbaceous species. Evident are these contradictions over the judgment of the level of the specialization of the elements the xyleme ties with insufficient knowledges of the comparing anatomy both the herbaceous as well as the shrubby peonies altogether, as well as the individual species of the section Moutan.

In the following we explored with morphologic - anatomical methods the vegetative organs of Paeonia tenuifolia L. (Sect.. Paeonia) [The results of the investigation of the other species of this section are disclosed in a further publication.] from the initial stages up to the mature individual. The material became collected in the Lugansker Oblast (Stavropoler steppe); besides, plants were raised in the years 1973 - 1975 by us in the BG of the Moscow university (Lenin - mountains).

P. tenuifolia is a plant of the steppe, the open steppe areas and the brushwood. It occurs in means - Europe, on the Balkan and in Asia, in the USSR in the European part and in the Caucasus. It is a typical geophyt. It increases essentially through seeds (Dudar. 1966). The seedling is composed of above ground and underground parts. First the main -root and the hypocotyl (Fig. 1, à) appear, the seedling fulfills to this time gaustorial functions and serves as place for the meanwhile deposit of nutrients, absorbed in the endosperm. At the Stem base of the seedling firmly joined, the short pipe develops the Keimsprosses. In the further arise the plastines of the seedling over the earth's surface as result of interkalären fiber - pipes and free segments of the stem. The plastines of the seedling are oval - lanzettlich (Fig. 1, á), fleshy. Their epidermalen cells are outlined imperfectly and contain few small Chloroplasten (Fig. 2, î, â). The mouths are anaomozytisch, sometimes two -, more often more - zählig in the upper epidermis (103 facing 33 on 1 mm ²). The mesophyllum is 9 - 10-layerd; them 2 - 3 uppermost layers appear as little expanded perpendikuläre surfaces - organs of the cells (Fig. 2, á). The cells of the following layers are round. The accompanying bundles are closed small, collateral,. The expansion of the vessels totals 270 mm on 1 cm ².

The stem is in the cross-section densely - sattelshaped (Fig. 2, ã). The basale thin-walled parenchyme has large luftgefüllte cavities. 3 the accompanying bundles the Versorgungsparenchyms are overcast, arched ordered, which are double medianen. The pipe the Keimsprosses besitz a length of 3 - 4 mm and contains 6 - 8 bundles. The knot the Keimsprosses is 1 - lakunär - ++, 2-bundleig.

The hypocotyl remains in the earth, his length totals 8 - 10 mm. His bud forms the sucker with 1 - 2 stems of the triply @+cut leaves (Fig. 1, á). If the first leaf arises over the surface, the seedling is resorbed in the essential. In his leaf-axis the Nebenknospen develop. In the course of the year the Basalteil thickens unimportantly to the little branched main - root and the hypocotyl.

diarchisch is The mean root. Their Exoderm is composed of the 1. row of thin-walled cells. The primary root is broadly (5 - 7 layers cells). The Endoderm has zones of Êàñïàðè. The pericyclum is on the largest expansion 1-layerd. The hypocotyl is set up similarly like the root (Exarchie of the primary xylemes, Endoderm with zones of Êañïàðè): the passage to the endarchischem xyleme ends closely the basis of the Keimspross - pipe. In the central cylinder of the top parts the hypocotyls of the seedling in the stage by 2 developed leaves owns it 10 bundles: 4 by them are in pairs joined bundle of the seedling, which are 6 others leaf - attempts. In the Hypokotyl is in the difference to the root well developed parenchymatisches marks. The secondary bulge in the underground organs begins early. The bundle - cambium replaces intensely the elements the secondary xylemes. leitershaped (with 1 - 2 crossbeam) are the perforated plastines of the vessel - segments; the rare perforation are simple. Small groups of bast-fibers contains his primary phloeme. As result of the work of the cells the pericyclum the tangentially - detached outer connective tissue develops a broad zone of stärkehaltigem parenchyme. Later the derivative pericyclum develops in the peripheral layers ring - phellogene, set up from cork of mehrlayerdem phelloderme. The primary root dies out and dissolves . The secondarily thickened root and the hypocotyl are composed of closed thick periderm.

Up to the fall the terminale lines up on the tip of the main axis bud with 3 - 4 leaf - Primordien, covers through 2 Knospenschuppen. In the second year is the increase of the main axis slight: The amount of the above ground part totals approx. 5 cm. After the scale-shaped develop 1 - 2 joined leaves with doubly - triply tailored plastines. The buds the basalen knots the mean stem remain resting. The bulge the hypocotyls and the basis the mean root leads to the formation an oval bundles of Stielstrukturen. On it side roots develop few Beside - and. In the largest expansion remains the mean root thin, his Sprossung and branching continues. Before the formation the bundle breitet itself secondary xyleme greatly from (Fig. 3, à); it owns parenchyme, but rather little tracheale elements. The vessels are individually (Fig. 3, á) or in small groups. Broad sectors produces The Zwischenbundle-cambium radiating parenchymees, at their place (Fig. 3, à) little solid accompanying elements, especially phloemic (nichtpolare bundle). In the following years decreases the activity the cambiums greatly. In the thin-walled parenchymebundles (xylemeic, phloemic, beam -, phellodermic) reserve -amyloid stores surely .

At 3 - 4 - year-old plants is the development the mean stem furthermore in the type rosettig monopodial. At the boundary of the slight annual increase the scale-shaped leaves takes off ita langstielige lower, at which gradually with the age the power of the division begins the plastines (Fig. 1, â). The leaf the 4 - year-old seedling is triply - or triply - feathery - cut with linealisch-fadenshaped segments, as well as at mature plants. The basalen areas the mean stem serve the buds -renewal, eingesenkt in the bud from the Kontraktilität going off from their root, line up short epigeogenische vertical small roots. It continues the growth also the reason - bundle, but at 4 - year-old plants appear as result of a strong bulge of the basis the hypocotyls and the stem the side roots still 1-2 bundle. The construction of this bulb is similarly the construction the basalen, but the number the beam the primary xyleme is large at them (3 - 4).

The substitute the monopodialen Sprosshauptachse through a sympodiale occurs in the essential at the passage of the plants to the generative phase of the development, more rarely in the virgile period. The first blossom appears in that 5. - 6. Year. monocyclic are the generative stems, terminal, leafy (up to the blossom develop 9 - 10 lower leaves), extends, with some joined knots in the basis, at an elevation of 10 - 50 cm. The lower buds -development of the above ground stems begins in June and ends in October - November. The stem the generative of drive is gerippt (Fig. 3, â). The epidermalen cells have a thickened outer wall and a smooth cuticula. Under the epidermis a broad layer of angle - collenchyme differentiates itself, intermediate to the large -cubic parenchyme. In the zone the primary root each internodess are the Blattansätze of the both uppermost leaves visibly (4 - 6 accompanying bundles). The medianen bundles the Blattansätze are collateral in the essential, isolates on very short expansion, quickly deviating to the center of the stem and at the boundary the 1. internodess unites with his accompanying system, but the lateral bundles are more amphicribalic, which are ordered in the Perifere of the stems the primary root, which draw themselves to some internodes up to the fusion with it equal bundles of the leaf - attempts, them go out from the lower leaf. The Vorhandensein of cortical bundles in the stems of the peonies was observed by Eames (Eames, 1953). you limit ripped surface organs. The central cylinder of the stems owns a ring of densely ordered accompanying bundles. The primary Kernstrahl is 3 - 4-rows, mehrzählig, the secondary ray 1 - 2-rows. The primary xyleme cuts measurably in the mark. Slight groups of protophloemicn contains His phloeme fibers. The limbs the siebshaped pipes own complicated siebshaped plastines. The vessels the secondary xyleme have a slight diameter (approx. 30 mm) with woody and equal type (Fig. 4, à). The limbs of the vessels are long (medium length 770 mm); plastines perforated the ladder-shaped mostly with 3 - 4 crossbeams, and meet the reticular perforated plastines with 5 - 7 openings. Besides the vessels are at the construction the xyleme tracheides, fibers (medium length 940 mm. Diameters 17 mm) with spaltenshaped framed pores , and hard parenchyme involves (Fig. 4, à).

The typical anatomical structure of the leaf develops in the Ontogenese early. The leaf is mesoform with several marks of blossoms - organization. The cells the epidermis have wound a thickened convex outer wall and are antiklinal (Fig. 2. ä, æ). asset developed is the cuticula. The openings (90 on 1 mm2) on the lower leaf side are wenigzählig . The upper layer the mesophyllums becomes presented from expanded perpendicularn surfaces - cells, which own 1 - 3 furrows ( Fig. 2, å). The sponge the 5 - 6-layerd parenchyme is composed of shovel - and star - cells. The leaf plastines at mature plants are thicker than at juvenilen (230 mm against 190 mm), for which the size of their cells is the reason. The medium vein stands out a little on the lower page, their phloeme is equips from collenchyme. The expansion vessels totals 570 mm on 1 cm ². The rachis in the centerpiece is round - sattelshaped (Fig. 2, à). 3 - 4 subepidermale cellsrows own the character of collenchyme. 7 accompanying bundles are ordered broadly arched. The phloeme has soft primary bast-fibers. stems first and following order own a slighter amount of bundles, than which in the rachis, (Fig. 2, è, ê).Die knots are 3 - ++ or 5-lakunär.

The above ground part the flower holdingn stems dies out in the fall, at the life remain the lower rootinternodes, which have developed from the year - increase the rootsystemes. One from their leaf-axis - buds takes over the function of the renewal, which remain resting others. The bud enlarges itself by year to year, branches, on the extended segments of the stem develops with the time of cork. Some of the resting buds open first at mature generative individuals. Before the submersion in the bud time increases them long delimited living basis the monocarpicn stems the small roots, at arise copies can reach ita length of 6 - 12 cm at a diameter of 4 - 8 mm (Fig. 5). The several years of small roots own closed cork from 10 - 14 layers of thin-walled cells. Under them the Brachysklerid develops. The central cylinder in 1 - ++ year-old small roots is compact, with ring - cambium and narrow Kernstrahlen. With beginning the 2. - 3. year are produced at the places the bundlecambiums broad (from 6 - 15 row cells) sectors of parenchyme and arranged connective tissue like on the particular bundles (Fig. 3, ã). In the secondary xyleme is the annual increase clearly distinctive. The tracheal elements in the small roots are more specialized, than which in the above ground stems (Fig. 4, á). The number of the crossbeams is slight in the woody perforated plastines (1 - 3), the broad segments are describes through simple perforations. The Porenhäufigkeit the Seitenwand is considerable. Besides the vessels in it the xyleme from detailed tracheides (leitershaped and porous), hard and beam - parenchyme put together (Fig. 4, á), In each hard on 2 - 3 cells. The secondary Kernstrahl is narrow. In the thin-walled cells are the nuclei broad, as well as in the beam -, raffia - and xylemeischen parenchymecells, there are found several amyloid grains. In the mark Drusen of Calcium-Oxalat. are found

With the time the rootsystemes from einzenen lines up with the annual increase side roots new lumps. The basale bulb and the thickened hypocotyle root stop gradually, to be the Hauptvorrats - organs of the plant. The rootverdickung at mature copies can total 7 - 12 and more (Fig. 1, ã). The side roots take over the functions the mean root . After V.N.Golubejew (1962), arrive some roots on a depth of 100 cm and reach in the width up to 40 - 50 cm.

On the basis of the study the Ontomorphogenesis and anatomy of the vegetative organs can be described Paeonia tenuifolia as kern-traubenrootige knollenbildende staudige polycarpic plant. stielrootiger is the basale bulb nature; the following bulb develops in the result of the growth the basalen parts the side roots, emerged from a short rootsystem. The reserve - web - bundle develops as greatly parenchymatisised secondary being xyleme.

The construction the seedling is above ground, which in case of an underground germination is rare. For the mean stem is in the course the first 3 - 5 years the rosette form the rung characteristic. monocyclic are form the generative stems, extends, leafy, with few root - internodes in the basis, which, eingesenkt in the bud, vertical or slanting small roots. The renewal of the bud results hidden in the bud.

The limbs - vessels the secondary xyleme are stems of the above ground stems with woody perforated plastines; the tracheal elements in the underground organ are specialized more, some from them own simple perforations. The leaf is mesoform, with marks of light - organization. The knots are 3 - ++, rare 5-lakunär.

Literature

Golubejev V.N. 1962. Bases the Biomorphologie the herbaceous plants the central Waldsteppe. «Beiträge of the company of the central Schwarzmeer - protectorates nam. Alexin No.7

Dudar Ju.A. 1966. The year - cycle the Morphogenese the netzblättrigen Paeonie (Paeonia tenuifolia L.) and the possibility of their increase. «Beiträge the Stavropoler NII Vol.2, 191197.

Ims A. 1964. Morphology of the flowering plants. «Mir».

Kemularia-Nathadse L.M.. 1961. The Caucasian representatives of the species Paeonia L. «Trud. Tiflis. Offered. Inst. AT GrusSSR», 21, 351.

Takhtadshjan A.L. 1948. The morphological evolution the Bedecktsamer. ., MOIP

Takhtadshjan A.L. 1966. System and phylogeny of the flowering plants. M.L., «Nauka».

Eames, A 1953. Floral anatomy as at aid in generic litation. «Chron. Bot.», 14,126132.

N a k a i T. 1949. Classes, ordines, families, familiae, subfamiliae, tribus, genera nova quae attinent ad plantas Koreanas. «J. Jap. Bot.», 24, 814.

Stern F.C.. 1946. A study of the genus Paeonia. London.

Assumed from the editing 22.4 1975 , Department of morphology and systematic persons higher plants

Captions:

Fig. 1:. P. tenuifolia (June): à - embryo á - 1 - year-old, â - 3 - year-old, ã - several years of plant; ãï - hypocotyl, ãê - mean root. ñ - seedling. ÷ë - scale-shaped leaf, íïá - above ground drive. êë - bulb

Fig. 2. P. tenuifolia. - Anatomical construction of the seedling (à - ã) and the leaves of mature plants (ä - ê): a, ä - upper, â, æ - lower epidermis with surface, á,å - cross-section the plastines; Cross-section of the stem of the seedling (ã), rachis (ç), stems 1. (and) and 2. (ê) order

Fig. 3. P. tenuifolia. diagram in the cross-section: bundle (à), stems the flowering stem (â) and 10 - year-old rootsystem-à (ã); ïðä periderm. Ôë phloeme. ê - cambium. ããê - Border the Jahresringes, ñêð - sclereides. ïñë - primary heart-shaped beam. ÷ïï - nichtpolare accompanying bundle, ýï - epidermis, êï - rootbundle, êëí collenchyme-à, ëb - bast-fibers, âñë - secondary heart-shaped beam ??- Cut offed the secondary xyleme the bundle - ++, ñ -vessels

Fig. 4. Histological composition the xyleme: à - stems 1 - ++ year-old stems, á - rootsystem-à; ÷ñ - Cut offed the vessels, òð -tracheides. ïð - hard woody parenchyme

Fig. 5. P. tenuifolia (May, June): à - 6 - year-old (from culture), á - 8 - year-old (Stavropoler steppe) plant; ë - stems the assimilation leaf, ÷ë - scale-shaped leaf, ï÷ - bud - è, ïê - side roots, êë - bulb , íïá - above ground stems, îïá - dying out stems, êðí - small roots