Colchicum Autumnale L. - an Ancient Medicinal Plant and Its Hysteranthous Geophytic Life Strategy

Frankova, L. 2006: Colchicum autumnale L. - An ancient medicinal plant and its hysteranthous geophytic life strategy. Server at URL: www.fyziologia.sav.sk/geophyte-colchicum

INTRODUCTION

Autumn crocus (Colchicum autumnale L.), alias naked lady, meadow saffron, wonder bulb, naked boy mysteria etc., has been used as an approved medical plant for more than 3000 years. Its scientific name comes from the Greek Colchis, an ancient country on the easternmost coast bordering on the Black Sea, where colchicums were found for the first time.

During the last centuries corms and seeds of C. autumnale were used in the treatment of asthma, rheumatism, dysentery and acute gout [1]. The modern medicine uses Colchicum as a source of therapeutically active alkaloids called colchicinoids. One of the most abundant alkaloid - colchicine, is known to have cancerostatic, antirheumatic, antiinflammatory, antimitotic, cathartic and emetic effects [2,3]. Colchicine and its analogues have been used clinically for the treatment of certain forms of leukaemia, familiar Mediterranean fever, cirrhosis, acute attacks of gout, Behcet's disease and solid tumours [2-5]. It is also applied in plant breeding to induce polyploids. The biogenesis of phenylethylisoquinoline skeleton of colchicinoids is well described, but its enzymatic aspects have not been investigated until now [6,7].

Along with the ability to produce colchicinoides, autumn crocus has certain other specific features. This plant has adopted a special life strategy being manifested through the unique life cycle [1,8]. In addition, it is a representative of ornamental clonal, monocotyledonous plants with hysteranthous geophytic life style [8,9].

Generally, geophytes are plants that survive by using a subterranean storage organ with renewal buds [10]. They divide into two groups - synanthous and hysteranthous [11]. The leaves of synanthous geophytes coexist with flowers in the same stage of the life cycle. In the case of hysteranthous plants flowers develop first and leaves later, at the another developmental stage. C. autumnale L. is one of the most common species of Colchicum genus in Central Europe. Because of its frugality and ability to flower during autumn, its hybrids and cultivars are successfully planted in gardens and used in plant horticulture as unpretentious ornamental herbs.

Even though Colchicum has been successfully used in medical treatment for more than 3000 years, only a few papers were devoted to its specific features and unusual life cycle [for

1 details see 1,8,9,15]. Taxonomical aspects of Colchicum autumnale L. are covered in detail by Persson [12] and morphological aspects were partly described by Jaehn et al. [13,14]. The analyses performed at the physiological, structural, biochemical and molecular level are still incomplete.

It is of note, that there is limited information about:

- aspects of clonality of cormous, bulbous and tuberous plants, - features of hysteranthous life forms - and finally about biochemical and molecular analyses of underground storage organs of perennial plants.

The first papers aimed to describe physiological, structural, biochemical characteristics of hysteranthous life form were published by Frankova et al. using C. autumnale as an exemplar plant [1,8,9,15-17].

Characterisation of the life strategy of the autumn crocus might contribute not only to the deciphering of biology of this species, but also to the understanding of hysteranthous and perennial life forms.

REFERENCES

1. KOMJATHYOVÁ, H. - FRANKOVÁ, L. - BÓKA, K. - PŠENÁK, M.: Botanical and developmental aspects of Colchicum autumnale L. (autumn crocus). (in Slovak). Acta Fac. Rerum Nat. Univ. Comenianae, Bot. 40, 2000, p. 67-80.

2. BOYÉ, O. - BROSSI, A.: Tropolonic Colchicum alkaloids and allo congeners. In: The Alkaloids. Vol. 41 (BROSSI, A. - CORDELL, G.A. - Eds.). New York, Academic Press 1992, p. 125-174.

3. KATZUNG, B.G. (Ed.): Basic and clinical pharmacology (9th Ed.). New York, Lange Medical Books, McGraw-Hill 2004, 527 pp.

4. ALALI, F. - TAWAHA, K. - QASAYMEH, R.M.: Determination of colchicines in Colchcium steveni and C. hierosolymitanum (Colchicaceae): Comparison between two analytical methods. Phytochem. Anal. 15, 2004, p. 27-29.

5. KAPLAN, M.M. - SCHMID, C. - PROVENZALE, D. - SHARMA, A. - DICKSTEIN, G. - MCKUSIK, A.: A prospective trial of colchicines and methotrexate in the treatment of primary biliary cirrhosis. Gastroenterology 117, 1999, p. 1173-1180.

6. BILECOVÁ, D. - ŠIMÁNEK, V. - JELOKOVÁ, J. - PŠENÁK, M.: Polyphenoloxidase activity and isoforms during ontogenesis of corms of Colchicum autumnale. Biologia 51, 1996, p.323-330.

7. BILECOVÁ A. - PŠENÁK, M.: Biochemical aspects of colchicinoids in Colchicum autumnale L. (in Slovak). Chem. Listy 89, 1995, p.100-110.

2 8. FRANKOVÁ, L. - KOMJÁTHYOVÁ, H. - BÓKA, K. - GAŠPARÍKOVÁ, O. - PŠENÁK, M.: Biochemical and physiological aspects of developmental cycle of Colchicum autumnale L. Biol. Plant. 47, 2003/2004, p. 509-516.

9. FRANKOVÁ, L. - CIBIROVÁ, K. - BÓKA, K. - GAŠPARÍKOVÁ, O. - PŠENÁK, M.: The role of the roots in the life strategy of Colchicum autumnale L. Biologia 59 (Suppl. 13), 2004, p. 87-93.

10. RAUNKIAER, C.: The life forms of plants and statistical geography. Oxford University Press, Oxford UK. 1934, 632 pp.

11. DAFNI, A. - COHEN, D. - NOY-MEIR, I.: Life-cycle variation in geophytes. Ann. Missouri Bot. Gard. 68, 1981, p. 652-660.

12. PERSSON, K.: New and revised species of Colchicum (Colchicaceae) from the Balkan Penninsula. Plant Syst. Evol. 217, 1999, p. 55-80.

13. JAEHN, F. - PFIRSCH, E. - ROUX, J.: Zur Architectur des Jahressprosses der Herbstzeitlose (Colchicum autumnale L.). Beitr. Biol. Pflanzen 60, 1985, p. 303-311.

14. JEAHN, F. - ROUX, J.: Architecture et cycle annuel du colchique adulte (Colchicum autumnale L.). Bull. Soc. bot. Fr. 133, Lettres bot., 1986, p.225-233.

15. FRANKOVÁ, L. - BÓKA, K. - GAŠPARÍKOVÁ, O. - PŠENÁK, M: Metabolic processes behind the autumnal developmental stage of Colchicum autumnale L. Acta Physiol. Plant. 26, 2004, p. 34-35.

16. FRANKOVÁ, L. - BÓKA, K. - PŠENÁK, M: The life strategy of Colchicum autumnale L. during the second developmental stage. In: Book of Abstracts of Plant Physiology Conference of Ph.D. Students and Young Scientists, Brno, Czech Republic, June 24-25, 2003, p. 56.

17. FRANKOVÁ, L.: Biochemical characteristics of developmental cycle of autumn crocus (Colchicum autumnale L.) [PhD. Thesis]. Bratislava, Institute of Botany, Slovak Academy of Sciences 2004.

BOTANICAL CHARACTERISTICS

The plant kingdom: Plantae Subkingdom: Tracheobionta Division: Magnoliophyta Subdivision: Magnoliophytina Class: Liliopsida Subclass: Liliidae Family: Colchcicaceae Subfamily: Colchicoideae Tribus: Colchicaceae Genus: Colchicum Species: Colchicum autumnale L.

3 Other names:

English: Autumn crocus naked lady, meadow saffron, wonder bulb, naked boy fall crocus mysteria son before the father bastard saffron

Slovak: Jesienka obyčajná šafranica bujačina naháč ocún

Czech: Ocún podzimní Ocoun, naháč, naháček, jeseňka, jesenka, luční matečník, oučastník, popovo moudí, sirotka, vocoun, vokoun, zimovít

Polish: Zimowit jesienny

Hungarian: Öszi kikerics homoki kikerics

Bulgarian: Мpазoвец (Mrazovec), Мразовец есенен (Mrazovec jesenen) Кърпи кожух (Krpi kožuch)

Belorussian: Безвременник осенний (Bezvremennik osennij), Oсенник (Osennik)

Russian: Безвременник осенний (Bezvremennik osennij) Безвременник, подснежник (Bezvremennik podsnežnik) Осенник (Osennik), Дивий-, моровой-шафран (Divij morovoj šafran)

Ukrainian: Пізньоцвіт осінній (Pizňocvit osinnij), Брандушка осiння (Branduška osinnaja), Зимовик (Zimovik), Зимовник (Zimovnik), Зимоцвiт (Zimocvit), Мороз (Moroz), Осiнник (Osinnik), Син без батька (Syn bez baťka), Шафран дикий (Šafran dikij)

German: Herbstreitlose Zeitlose, Butterwecken, Henne, Hennegift, Giftblume, Herbstblume, Herbstlilie, Hundsblume, Hundshode, Hundsknofel, Kuckucksweck, Kühe, Kuheuter, Läuseblume, Leichenblume, Michelwurz, Mönchskappen, Nackte Hure, Nackte Jungfer, Ochsen, Ochsenpinsel, Spindelblume, Spinnblume, Teufelsbrot, Teufelswurz, Wiesenlilie, Wiesensafran, Wildsafran, Winterhaube, Winterhauch

4 French: Colchique d'automne Tue-chien; Veilleuse; Safran des prés, Paternostar

Spanish: Colquico, Colchico Azafrán Bastardo, Cólchico, Flor de Otoño, Mataperros, Narciso de Otoño, Quitamersendas

Portugal: cólquico

Italian: Colchico d'autumno, colchico autumnale desmonia, zafferano bastorda, zafferano salvatico, zafferano bastardo

Greek: κολχικό

Estonian: Harilik sügislill

Finnish: Syysmyrkkylilja, Alastonimpi, Syksyn myrkkylilja.

Swedish: Tidlösa, Hösttidlösa, Nakna jungfrun, Vanlig tidlösa.

Norwegian: Tidlaus, Tidløyse, Naken jomfru, Tidløs

Danish: Høst-Tidløs, Høst-Tidløs, Nøgne jomfruer, Nøgen jomfru, Tidløs

Netherlandish: Herfsttijloos

LIFE FORM: Geophyte LEAVES COEXISTENCE: flowers without leaves - hysteranthous UNDERGROUND STORAGE ORGAN: a corm RELEVANCE: medicinal plant, ornamental plant

Corm: underground basal stem, ovoid, sometimes flattish, a little bit wrinkled, 2-5 cm wide, 3-6 cm high, with well developed protuberance (a hollow diverticulum - an organ helping the new plant to descend into the soil - 10,11). Surface: covered in cinnamon-brown scale leaves - tunic Externally: Yellowish to brownish, wrinkled Internally: White, solid and full of starch. Bitter taste and somewhat acrid. The plant has usually two corms: old mother corm and continual developing new daughter corm so called regular one (10,11) - situated in the wrinkled inner furrow on mother corm.

Flowers: showy pink, purple, lavender, pale-pink to white flowers (6 parted) in groups of 1 to 6 opened up on the leafless stalks sprouting from the corm;. Each petal is about 3 to 4.5 cm long and is fused below into a pale stalk-like tube 5-25 cm long. Each flower has 6 stamens with dark yellow anthers, 3 independent white styles and a superior ovary (1-9). Flowers appear at the beginning of autumn, when there are no leaves above the soil. Within the autumnal developmental stage the flowers appear first and the leaves start to emerge from the soil five months later. Therefore hysteranthous geophytes have special requirements on the accessibility of storage reserves (10,11,13).

5 Leaves: 10-40 cm tall, 1-4 cm wide, lanceolate, slender, dark green, shiny, elongate up to 60 cm when the stem rises. Veins run from the base parallel to the leaf axis (1-9). They form a rose sprout and appear in the spring (hysteranthous), lasting only for 3-4 months (10,12,13)

Fruit: oblong to ovoid green (in mature brown) fruit (three-cased, three-valved dehiscent capsule). Young non mature capsules are hidden under the soil surface after the fertilization. Capsules ripen in the late spring. The seeds are rounded in shape, very hard and tough, when unripe whitish and polished, fat and dark brown when mature (1-9).

Roots: white, wrinkled, smooth, contractile (11-13). The root system is formed after flowering (11,13,14).

Appearance: C. autumnale is a native plant of south, west and central Europe, extending to the eastern banks of the Black Sea, in Georgia (14,15). North Africa, middle Asia, and also south Scandinavia are grater extent of the autumn crocus. The plant is cultivated throughout much of the world, primarily as an outdoor ornamental plant. For example in the U.S. Kentucky, Maryland, Massachusetts, New Hampshire, New York, North Carolina, Oregon, Utah can be mentioned as the new cultivable areas of the autumn crocus (15).

Colchicum autumnale L. native to Slovakia prefers wet meadows, pastures, moorlands and flood-plain forests. It needs nutritive, loamy, humid, permeable and neutral or slightly acid soils respectively. It can be found around the area of Levice, Kozárovce, Nový Tekov (12) Malacky-Kostolište, Krupina, Devínska Nová Ves – flood plain of the Morava and Čadca- Milošová.

REFERENCES

1. HUXLEY, A.H. ed-in-chief (1992). The new Royal Horticultural Society dictionary of Gardening, vol 1. London, MacMillan.

2. HEGNAUER, R., 1963: Chemotaxonomie der Pflanzen II. Birkhänser Verlag, Basel und Stuttgart, 301-304.

3. KRESÁNEK, J., KREJČA, J., 1982: Atlas of medical plants and forest fruits. Osveta, Martin, 127-128. [in Slovak]

4. LIST, P. H., HORHAMMER, L., 1972: Hagers Handbuch der Pharmazeutischen praxis II. Springer - Verlag Berlin, Heidelberg, New York, 235-240. [in German]

5. TREASE, G. E., EVANS, W. C., 1978: Pharmacognosy. Bailliere Tindall - London, 584- 588.

6. HEGI, G., 1909: Illustrierte Flora von Mittel Europa II. A. Picher's Wittwe & Sohn, Wien, 195-199.

6 7. DOSTÁL, J., 1989: Nová květena ČSSR 2. Academia, Praha, 1196-1197. [in Czech]

8. SÃVULENSEN, T., NYÁRÁDY, E., 1966: Flora Republich Socialiste Romania XI. Editura Academici Republicii Socialiste Romania, 115-128.

9. SOÓ, R., 1973: A magyar flóra és vegetáció rendszetrani - növényföldrajzi kézikönyve V., Akadémiai Kiadó, Budapest, 56-58. [in Hungarian]

10. FRANKOVÁ, L. - KOMJÁTHYOVÁ, H. - BÓKA, K. - GAŠPARÍKOVÁ, O. - PŠENÁK, M.: Biochemical and physiological aspects of developmental cycle of Colchicum autumnale L. Biol. Plant. 47, 2003/2004, p. 509-516.

11. FRANKOVÁ, L. - CIBIROVÁ, K. - BÓKA, K. - GAŠPARÍKOVÁ, O. - PŠENÁK, M.: The role of the roots in the life strategy of Colchicum autumnale L. Biologia 59 (Suppl. 13), 2004, p. 87-93.

12. KOMJATHYOVÁ, H. - FRANKOVÁ, L. - BÓKA, K. - PŠENÁK, M.: Botanical and developmental aspects of Colchicum autumnale L. (autumn crocus). (in Slovak). Acta Fac. Rerum Nat. Univ. Comenianae, Bot. 40, 2000, p. 67-80.

13. FRANKOVÁ, L.: Biochemical characteristics of developmental cycle of autumn crocus (Colchicum autumnale L.) [PhD. Thesis]. Bratislava, Institute of Botany, Slovak Academy of Sciences 2004.

14. FRANKOVÁ, L. - CIBIROVÁ, K., - BILKA, F. - BILKOVÁ, A. - BALÁŽOVÁ, A. - PŠENÁK, M.: Nitrate reductase from the roots of Colchicum autumnale L., Acta Facult. Pharm. Univ. Comenianae 52: 1-10, 2005 - in print.

15. BRUNETON, J.: Pharmacognosy, phytochemistry, medicinal plants. Paris, Lavoisier, 1995.

16. TUTIN TG., - HEYWOOD VH., - BURGES NA., - MOORE DM., - VALENTINE DH., - WALTERS, SM., - WEB DA. eds. Flora Europaea, vol 5. UK, Cambridge, Cambridge University Press, 1980

17. Server at URL: http://plants.usda.gov

PLANT COMPOUNDS AND STORAGE MATERIAL

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EFFECT OF COLCHICINOID ALKALOIDS

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7 THE HYSTERANTHOUS GEOPHYTIC LIFE FORMS

Colchicum autumnale L. is the Perennial (also clonal) Cormous Hysteranthous Geophyte

Hysteranthous - definition:

• Having the leaves expand after the flowers have opened. (http://dict.die.net/hysteranthous/) (http://en.thinkexist.com/dictionary/meaning/hysteranthous/) (http://www.dictionaryhead.com/definition/hysteranthous-definition.html) (http://www.websters-online-dictionary.org/definition/english/Hy/Hysteranthous.html) (http://onlinedictionary.datasegment.com/word/hysteranthous) (http://www.webster-dictionary.net/d.aspx?w=hysteranthous)

8 • a plant with leaves appearing after the flowers. (http://www.zimbabweflora.co.zw/glossary.php)

Synanthous - definition:

• Having flowers and leaves which appear at the same time; -- said of certain plants. (http://dict.die.net/synanthous/) (http://www.dolally.com/dictionary/definition.asp?Word=Synanthous) (http://www.webster-dictionary.net/definition/Synanthous) (http://www.websters-online-dictionary.org/definition/english/Sy/Synanthous.html) (http://onlinedictionary.datasegment.com/word/synanthous) (http://en.thinkexist.com/dictionary/meaning/synanthous/)

Geophyte - definition:

• a plant that stores food underground and has subterranean buds which form aerial growth. (http://www.succulents.co.za/botanical_terms.shtml)

• (1) Perennial herb with its perennating bud (s) located well below the soil surface. • (2) Perennial plant with an annual shoot and perennial underground parts. (http://www.geobotany.uaf.edu/toolikgeobot/definitions.html)

• a perennial plant that propagates by underground bulbs or tubers or corms (http://wordnet.princeton.edu/perl/webwn)

• a storage organ is a part of a plant specifically modified for storage of energy (generally in the form of starch) or water. Storage organs often grow underground, where they are better protected from attack. Underground storage organs are called geophytes. In common parlance, underground storage organs may be generically called roots, tubers, or bulbs, but to the botanist these are specific, technical terms, which apply only more narrowly. (http://en.wikipedia.org/wiki/Geophyte)

• A perennial plant, such as a crocus or tulip, propagated by buds on underground bulbs, tubers, or corms. (http://www.thefreedictionary.com/geophyte) (http://davesgarden.com/terms/go/2108/)

• a perennial plant that propagates by underground bulbs or tubers or corms (http://dict.die.net/geophyte/) (http://www.wordwebonline.com/en/GEOPHYTE) (http://www.beetfoundation.com/words/g/alt.geophyte.html) (http://www.wordreference.com/definition/geophyte.htm) (http://define.ansme.com/words/g/geophyte.html)

9 Perennial - definition:

• Lasting or active through the year or through many years. 1. Botany - A perennial plant. 2. Something that recurs or seems to recur on a yearly or continual basis (http://www.thefreedictionary.com/perennial)

• a plant or plant tissue which lives more than one growing season (https://www.uwsp.edu/natres/nres743/Definitions/Perennial.htm)

• (1) A plant or flower that usually takes more than one growing season to mature and usually lives for more than one year. • (2) Perennials are plants that survive winter outdoors to produce new growth each spring. Replanting is not necessary. • (3) A perennial plant is a plant that usually grows for three or more years. It usually does not flower in it's first year of growth. A perennial is usually a flower. (http://davesgarden.com/terms/go/659/)

• (1) A perennial plant or perennial (Latin per, "through", annus, "year") is a plant that produces flowers and seeds more than one time in its lifespan, and therefore lives for more than one year. This term is usually applied to herbaceous plants or small shrubs rather than large shrubs or trees, but used strictly it also applies to all plants which flowers and produces seeds more than once. • (2) Plants that flowers and produces seeds only once in its lifetime are called 'monocarp' as annual plants, which flowers in its first living year, then dies, or biennial plants, which only live for two seasons. Some monocarp plants can live for many years before flowering (and dying) as bamboo and agave. • (3) Herbaceous perennials are plants that do not form permanent woody tissue. In warmer and more clement climates they may grow continuously. In seasonal climates, their growth pattern is adapted to the growing season. In cooler temperate regions they generally grow and bloom during the warm part of the year, and the foliage dies back every winter. Regrowth is from their existing tissue or root-stock rather than from seed, as with annuals and biennials. In some cases, these perennials may retain their foliage all year round, even in seasonal climates. Herbaceous perennials that retain their foliage all year round may be called evergreen perennials. Others are called deciduous. Examples of evergreen perennials; Begonia; banana Examples of deciduous perennials; Goldenrod; mint Woody perennials (ie. trees and shrubs) retain their woody structure permanently, but may lose their foliage in seasonal climates. Perennial plants dominate most natural ecosystems. For example, grasses and most forbs on the prairie are perennial. Wild perennial plants are usually better competitors than annual plants, especially under resource-poor conditions. This is due to larger root systems which can access water and soil nutrients deeper in the soil and to earlier emergence.

(http://encyclopedia.laborlawtalk.com/Perennial)

• (1) Noun: A plant lasting for three seasons or more • (2) Adjective: Lasting three seasons or more "the common buttercup is a popular perennial plant"

10 (http://www.wordwebonline.com/en/PERENNIAL) (http://www.wordreference.com/definition/perennial)

Annual - definition:

• (1) Adjective: Recurring, done, or performed every year; yearly: an annual medical examination. Of, relating to, or determined by a year: an annual income. Botany: Living or growing for only one year or season. • (2) Noun: A periodical published yearly; a yearbook. Botany: A plant that completes its entire life cycle in a single growing season. (http://www.answers.com/topic/annual)

• (1) Adjective: (botany) completing its life cycle within a year; "a border of annual flowering plants" [syn: one-year] [ant: biennial, perennial] • (2) Noun: a plant that completes its entire life cycle within the space of a year (http://dict.die.net/annual/) (http://www.wordwebonline.com/search.pl?w=annual)

• Annual, from the Latin annuum, or year means pertaining to a year or happening every year. • An annual plant has a lifespan of a year or less. These are generally plants adapted to life as weeds or in difficult habitats where quick flowering and seeding are necessary. (http://encyclopedia.laborlawtalk.com/Annual) (http://en.wikipedia.org/wiki/Annual)

Clonal - definition:

• Of or relating to a clone. • See also clone: A group of genetically identical cells or organisms derived from a single cell or individual by some kind of asexual reproduction (http://www.wordwebonline.com/search.pl?w=clonal) (http://www.wordreference.com/definition/clonal)

• Pertaining to a clone. (http://www.biology-online.org/dictionary/clonal)

• Any of a group of genetically identical cells or organisms derived from a single parent cell or organism by asexual reproduction. (http://www.allwords.com/word-clonal.html) (http://www.thefreedictionary.com/clone)

• Clonal propagation: Asexual reproduction of plants that are considered to be genetically uniform and originated from a single individual or explant. • Clone: A clone is not necessarily homogeneous and, therefore, the terms clone and cloned do not indicate homogeneity in a cell population, genetic or otherwise. In plant culture terminology, the term may refer to a culture derived as above or it

11 may refer to a group of plants propagated only by vegetative and asexual means, all members of which have been derived by repeated propagation from a single individual. (http://www.sivb.org/edu_terminology.asp)

• Clonal plants spread vegetatively within their habitats by forming rooted ramets on stolons or rhizomes. Each of these ramets is capable of an independent existence after establishment. Nevertheless, ramets remain physically connected by stolon or rhizome internodes for variable periods of time, thereby allowing for resource movement and signal transduction within clones. Interconnected ramets of clonal plants, though potentially independent and totipotent, can specialize functionally in the performance of limited numbers of tasks such as the uptake of resources from above- vs below-ground sources, carbohydrate storage, vegetative spread and sexual reproduction. Such specialization and cooperation is comparable to a division of labour in economic systems or in colonies of social animals. The ecological significance of division of labour in clonal plants may be found in the increased efficiency of entire clones in exploiting their environments. (Stuefer J.F.1998. Two types of division of labour in clonal plants: benefits, costs and constraints. Perspectives in Plant Ecology, Evolution and Systematics: 1: 47-60)

• The ultimate elaboration of modularity and totipotency is clonal growth: one plant gives rise to another gives rise to another gives rise to another with no sex in between. A large proportion of plants possesses the capacity for clonal growth and, as a group, plants clone themselves in an intriguing variety of ways. (Eckert CG. 1999. Clonal plant research: proliferation, integration, but not much evolution 86:1649-1654.)

• Clonal plant is capable of naturally producing independent offspring by means of vegetative growth (http://enri.uaa.alaska.edu/Staff_home_pages/MattCarlson/EVO%20ECO%20Lectures/Lectur e%2017%20-%20Clonal%20growth-reproduction%20&%20Senescence.pdf)

• The word clone comes from the Greek word klon meaning twig or cutting. Comment: Originally clone referred to the process of vegetative propagation, used by gardeners for many centuries to propagate grape vines, apple trees, rose bushes and the like. Vegetative propagation depends on taking cuttings (slips) of shoots and rooting them or grafting them to a rootstock. This produces a new plant (or shoot, if grafted on a rootstock) that has the same genetic characteristics as the cutting from which it was taken. • A modern definition: The process of asexually producing a group of independent organisms or cells, all genetically identical, from a single ancestor. Comment: In current usage a clone may be a bacterial cell line, a rose bush or a culture of mouse cells. They must all derive directly from a common ancestor without an intervening sexual cross or genetic manipulation. (http://www.mobot.org/jwcross/duckweed/cloning-duckweed.htm)

12 ....Colchicum autumnale L. subspecies autumnale is the perennial cormous hysteranthous geophyte....

The other examples of true hysteranthous plant life forms:

Colchicum autumnale 'Alboplenum' Colchicum autumnale 'Album' Colchicum autumnale 'Nacy Lindsay' Colchicum autumnale 'Plenum' Colchicum autumnale var. atropurpureum (Stapf apud Stearn) Colchicum tunicatum L. - annual geophyte Colchcicum speciosum (Steven) - annual geophyte Colchicum umbrosum (Steven) - annual geophyte Colchicum boissieri (Orph.) - annual geophyte Colchicum graecum (K. Persson) - annual geophyte Colchicum hierosolymitanum Colchicum bivonae (Guss) Colchicum feinbruniae (Persson) Colchicum silicicum ((Boiss.) Dammer) Colchicum polyphyllum (Boiss. & Heldr., natural hybrid) Colchicum sfikasianum (Kit Tan & Iatroú., natural hybrid)

Crocus damascenus - annual geophyte Crocus ochroleucus - annual geophyte Crocus pulchellus - leaves absent at anthesis Crocus speciosus - leaves absent at anthesis Crocus banaticus - leaves absent at anthesis Crocus nudiflorus - leaves absent at and for long after anthesis Crocus robertianus - leaves absent at anthesis Crocus medius - leaves absent at anthesis Crocus cancellatus - leaves absent at anthesis, perennial cormous geophyte Scilla hunburyi – perennial geophyte Scilla autumnalis – perennial geophyte Urginea maritima – perennial geophyte Pancratium maritimum – perennial geophyte Pancratium sickenbergeri (C. et Barbey)

Sternbergia colchiciflora - perennial geophyte Haemanthus pubescens L. Scadoxus multiflorus – tropical species Scadoxus puniceus Nerine alta – tropical species Nerine humilis Nerine marincowitzii Eucrosia aurantiaca (Baker) Merendera Montana

The examples of synanth/hysteranthous plant life forms: (leaves rise during or slightly after flowering)

13 Colchicum baytopiorum (C.Brickell) Colchicum tenorii (the leaves are dwarfed) Colchicum feinbruniae Colchicum freynii Crocus longiflorus (just or after blossoming, perennial geophyte) Crocus pallassii (leaves often absent at anthesis) Nerine sarniensis (deciduous-hysteranthous)

Lenka Franková E-mail: [email protected] Institute of Botany, Slovak Academy of Sciences Tel.: ++421 2 59426 144 Dept. of Plant Physiology Fax: ++421 5477 1948 Dubravska cesta 14 SK-845 23, Bratislava Slovak Republic