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Plant systematics and evolution - Comparative embryology of three Ge... (1997)
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40 , M. Akhalkatsi & J. Wagner:

al pecularities, the life histories, and the dynamics of seed formation are pre¬
sented.

Material and methods

Plant material. Gentiana pyrenaica (syn. Gentiana djimilensis C. Koch) is a perennial
mountain plant distributed in the Pyrenees, the Carpathians, in SW Bulgaria, Anatolia and
Iran, and in the Caucasus (Gagnidze 1985, Tutin & al. 1972). In the Central Caucasus this
species can be found between 2000 and 3500 m s.m. Depending on the altitude the flow¬
ering period is June to July, with a possible second flowering in August to October. The
plant material for the present investigation was collected in July at 3100 m s.m. in the sub-
nival zone on Mt. Sabertse in the Central Caucasus (Kazbegi region, Georgia).

Gentianella caucasea (syn. Gentiana caucasica Bieb.) is an annual plant, distributed in
Anatolia, Iran and in the Caucasus (Gagnidze 1985). Various morphological types with
different flowering times from summer to late autumn occur in widely differing habitats
from the subalpine to the subnival zone (Akhalkatsi & Wagner 1996). Plant material was
collected in July at 1950 m s.m. in the subalpine zone on the lower slopes of Mt. Kuro in
the Central Caucasus (Kazbegi region, Georgia).

Gentianella germanica (syn. Gentiana germanica Willd.) is an annual or biennial spe¬
cies, occurring from the colline up to the alpine zone in West and Central Europe and in
the mountain regions of the Balkans (Meusel & al. 1978). Like G. caucasea, this species
occurs in ecotypic variants, showing differences in their morphological characteristics and
time of flowering (Wettstein 1895, Zopfi 1991). The plant material was sampled in Octo¬
ber from a biennial population at 2000 m s.m. in the subalpine, zone of the Mt. Patscher-
kofel in the Central Alps (Tyrol, Austria). Herbarium vouchers of G. pyrenaica and G.
caucasea are deposited in the herbarium of the Institute of Botany of the Academy of Sci¬
ences, Tbilisi (TBI). Voucher specimens of G. germanica are deposited in the herbarium
of the University Institute of Botany, Innsbruck (IB).

Methods. Buds, flowers and fruits at different stages of development were fixed in
FPA50 (formalin, propionic acid, 50% ethanol, 5:5: 90) and stored in 70% ethanol.
Ovules and seeds were dissected out of the ovaries, mounted on microscope slides in
clearing solution according to Herr (1971) and examined with a microscope equipped
with differential interference contrast optics. Mature seeds were cut into slices with a
razor blade before mounting. Photographs were taken with an OLYMPUS Photomicro¬
graphic System, using KODAK Technical Pan 135 film at ISO 25 developed in AGFA
Rodinal. For each of the species the lengths and widths of 50 seeds and the lengths of 20
embryos were measured at 100X magnification with an ocular micrometer.

The gynoecia of the investigated species are superior, unilocular, bicarpellate
(very rarely tricarpellate) and paracarpous; they are terminated by a short style and
a 2-lobed stigma. In Gentiana pyrenaica the gynophore measures 7-8 mm, in
Gentianella caucasea and G. germanica 3-4 mm and 3-5 mm, respectively.
Numerous ovules develop on the parietal placentae along the fused margins of the
carpels.

Gentiana pyrenaica. The ovule is anatropous, tenuinucellar and unitegmic
(Fig. 2 A). The single integument consists of 8-10 layers of cells and forms a long
micropyle. At the micropylar end the nucellus consists of only one cell layer,
which degenerates during the extension of the embryo sac (Fig. 2 B). As is com¬
mon in Gentianaceae (Bouman & Schier 1979, Shamrov 1987), no vascular bun-