Schisandra

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Schisandra

Description

Woody vines, monoecious. Leaves alternate on long shoots or clustered on short shoots, exstipulate; lamina papyraceous to coriaceous, elliptic to ovate, apex acute to acuminate, base generally acute to decurrent, margins denticulate to entire, pinnate venation; petioles with groove on adaxial surface. Flowers generally solitary and axillary, occasionally in clusters of up to 8, sometimes ramiflorous. Fruit an aggregate of berries attached to elongated receptacle; berries ellipsoid to obovoid, ripening red; pedicel not enlarged in fruit. Seeds (l-)2(-3), smooth to rugulose, hilum lateral.

Distribution

Asia-Tropical:, Jawa (Jawapresent); Lesser Sunda Is. (Balipresent) Northern America: from southern Sakhalin in the north-east to the Himalayas in the west: present
There are c. 25 species in the genus, with an extensive distribution from from southern Sakhalin in the north-east to the Himalayas in the west; the centre of diversity lies in Assam and southern China. The genus shows two major disjunctions: two species, S. propinqua (Wall.) Baill. and S. elongata (Blume) Baill., occur in Java and Bali; and S. glabra (Brickell) Rehder is endemic to the south-east United States. In Malesia: Java and Bali (2 species).

Morphology

The female flowers of Schisandra are characterised by carpels with subulate stigmatic crests, and cylindrical or conical receptacles that are markedly longer than wide and become highly elongated in the fruit. The male flowers show great variation in androecial structure: the most primitive androecia consist of a large number of essentially free stamens that are only fused at the base of the filaments. According to Smith (1947), three distinct evolutionary trends are evident in the genus: 1) reduction of the number of stamens, ultimately giving rise to an extreme of only 4-5 in S. chinensis (Turcz.) Baill.; 2) complete fusion of the filaments to form a carnose mass with the anthers embedded in cavities; and 3) fusion of the filaments to form a 'staminal shield' of five stamens, with the thecae borne on the lower (morphologically dorsal) margins of the connectives. The androecial variation was used by Smith (1947) as the basis for his sectional classification of the genus (discussed below). Floral ontogeny is described and illustrated in detail by Tucker & Bourland (1994).
A. Smith, A.C. in Sargentia 7. 1947, B. Tucker, S.C. & J. A. Bourland - in PI. Syst. Evol. 1994

Anatomy

In addition to the references given in the discussion of the morphology and anatomy of the family, the following aspects of the vegetative anatomy of Schisandra have received particular attention: stomatal structure and development (Rao 1939; Jalan 1962); nodal anatomy (Jalan 1968a); crystalliferous sclereids (Jalan 1968b); ethereal oil cells (Jalan 1965); and mucilage cells (Jalan 1975).
C. Jalan, S. in Beitr. Biol. Pflanzen 44, D. Jalan, S. in Bot. Jahrb. 88, E. Jalan, S. in Curr. Sci. 18. 1965, F. Jalan, S. in J. Ind. Bot. Soc 54. 1975, G. Jalan, S. in Phytomorphology 12. 1962, H. Rao, H.S. in Proc. Ind. Acad. Sci. B 9. 1939

Taxonomy

The most comprehensive taxonomic revision of Schisandra is by A.C. Smith (1947), who accepted 25 species. Smith recognised four different androecial types in the genus, which he used as the primary criterion for the delimitation of sections, as follows:
  • Sect. Pleiostema A.C. Sm., with few to numerous essentially free stamens, arranged in
    irregular whorls on an elongated column or torus;
  • Sect. Maximowiczia (Rupr.) Nakai, with the filaments fused into a short column, bearing few sessile anthers at its apex;
  • Sect. Schisandra ('Euschisandra'), with the androecium composed of a flattened pentagonal shield with five radiating anthers bearing thecae on the dorsal margin;
  • Sect. Sphaerostema (Blume) Nakai, with the androecium composed of a subglobose or ellipsoid carnose mass, derived by the fusion of filaments, with anthers borne in cavities on the surface.

Smith hypothesised that the most primitive section was Pleiostema, and that the other sections have been derived by processes of filament fusion and the reduction in stamen number. Although the delimitation of the sections was primarily based on androecial structure, there is also an imperfect correlation with seed coat structure.

Y.-W. Law (1996) proposed a revised supraspecific classification for the genus, involving both the raising of the sections to subgeneric level, and the recognition of two new subgenera, viz.: 'Sinoschisandra' for those species (previously classified in sect. Pleiostema) which possess a sterile 'shield' over the apex of the torus; and 'Plenischis- andra' solely for S. plena A.C. Sm., which differs from the other species previously classified in sect. Sphaerostema in possessing sessile anthers.

The two Malesian species belong to different supraspecific taxa: S. elongata (Blume) Baill. belongs to sect. Pleiostema sensu Smith (= subg. Pleiostema sensu Law), whereas S. propinqua subsp. axillaris (Blume) R.M.K. Saunders belongs to sect. Sphaerostema sensu Smith (= subg. Sphaerostema sensu Law).
I. Law, Y.-W. in Fl. Reipubl. Pop. Sin 30, 1. 1996, J. Smith, A.C. in Sargentia 7. 1947

Cytology

Six species of Schisandra have been examined cytologically, although none of these species occur in Malesia. Most reports are of n = 14 (Whitaker 1933; Ehrendorfer et al. 1968; Stone 1968; Ratter & Milne 1976; Sandhu et al. 1989) and 2n = 28 (Janaki-Ammal, unpubl., cited in Darlington & Wylie 1955; Stone 1965, 1968; Okada 1975; Singhal et al. 1980; Gill et al. 1982; Chen et al. 1993; Wu & Huang 1995), although Bostick (1965) reports n = 13 for S. glabra (syn. S. coccinea). Other studies have revealed a diploid cytotype of n = 7 for S. grandiflora (Wall.) Hook. f. & Thorns. (Malla et al. 1977; Sandhu et al. 1989). The base number for the genus is therefore regarded as x = 7, 13.
K. Bostick, P.E. in Sida Contrib. Bot. 2. 1965, L. Chen, R.-Y., et al. in Chromosome Atlas of Chinese Principal Economic Plants 1. 1993, M. Darlington, C.D. & A.P. Wylie - in Chromosome Atlas of Flowering Plants. 1955, N. Ehrendorfer, F., et al. in Taxon 17. 1968, O. Gill, B.S., et al. - in The Vegetational Wealth of the Himalayas. 1982, P. Malla, S.B., et al. - in Taxon. 1977, Q. Okada, H. in J. Sci. Hiroshima Univ. B (Bot.) 15. 1975, R. Ratter, J. A. & C. Milne in Notes Roy. Bot. Gard. Edinb. 35. 1976, S. Sandhu, P.S., et al. in Curr. Sci. 58. 1989, T. Singhal, V.K., et al. - in Taxon. 1980, U. Stone, D.E. in J. Elisha Mitchell Sci. Soc. 84. 1968, V. Stone, D.E. in Madroño 18. 1965, W. Whitaker, T.W. in J. Arnold Arbor. 14. 1933, X. Wu, Z. & C. Huang in Guihaia 15. 1995

Uses

Schisandra is not very important economically, although the North Asian species S. chinensis and S. propinqua are used for various medicinal purposes

Citation

DC. - in Syst. 1818: 544
Backer & Bakh.f. - in Fl. Java. 1964: 99
A.C. Sm. - in Sargentia. 1947: 86