INTRODUCTION
Astragalus L. (Fabaceae: tribe Galegeae), with about
3000 species worldwide, is the largest genus of flow-
ering plants. The high variation of morphological
characters has made the taxonomic division of the
genus uncertain and problematic (Liston and
Wheeler, 1994; Sanderson and Liston, 1995;
Sanderson and Wojciechowski, 1996; Wojciechowski
et al., 1999; Zarre, 2000). Several previous systemat-
ic studies attempted to classify this huge genus into
formal taxonomic groups, that is, subgenera and sec-
tions (Bunge, 1868; Podlech, 1982), but only a few
groups found support from molecular systematics
(Wojciechowski et al., 1999; Kazempour Osaloo et al.,
2003, 2005). Detailed cladistic investigations indicat-
ed the homoplastic nature of most gross morphologi-
cal characters in delimitation of subgenera and sec-
tions in Astragalus.
Embryological studies have provided useful
characters in assessing subgeneric classifications in
Papilionoideae (Rembert, 1969; Palser, 1975;
Lersten, 1983; Prakash, 1987; Cameron and
Prakash, 1994; Soverna et al., 2003) as well as in the
genus Astragalus (Akhalkatsi et al., 1988; Gvaladze
and Akhalkatsi, 1996; Riahi et al., 2003). However,
the methods of such investigations in Astragalus are
time-consuming and difficult to carry out due to prob-
lems in cultivating the plants under greenhouse con-
ditions or in gardens (Riahi et al., 2003).
In this work we made a detailed embryological
study of A. cemerinus G. Beck (sect. Microphysa)
and A. ruscifolius Boiss. (sect. Dissitiflori), two
endemic species of Astragalus in Iran, with the aim
of providing valuable characters useful in assessing
relationships within this genus.
MATERIALS AND METHODS
Flowers and young pods of A. cemerinus and A. rus-
cifolius were collected in May 2004 from mountains
(~1580 m a.s.l.) above Mashhad Ardahal in Esfahan
Province, Iran. At least 60 developing seeds and
*
e-mail: zarre@khayam.ut.ac.ir
SEED DEVELOPMENT IN ASTRAGALUS CEMERINUS AND A. RUSCIFOLIUS
(FABACEAE), AND ITS SYSTEMATIC IMPLICATIONS
MEHRSHID RIAHI AND SHAHIN ZARRE
*
Department of Plant Sciences, University of Tehran,
PO Box: 14155–6455, Tehran, Iran
Received February 17, 2009; revision accepted May 15, 2009
This study focuses on seed development in Astragalus cemerinus and A. ruscifolius, two endemic species of
Astragalus in Iran. In both species the ovules are campylotropous, bitegmic and crassinucellate. Two polar
nuclei fuse before fertilization, forming the diploid secondary nucleus. Division of the primary endosperm nucle-
us gives rise to coenocytic endosperm; however, part of it becomes cellular at the late globular stage. The first
division of the zygote is transverse and the embryo proper forms after several divisions of the terminal cell. The
mature suspensor consists of a mass of cells equal in size to the globular embryo proper, with several inflated
cells towards its base. This massive suspensor seems to be plesiomorphic, as compared with the biseriate sus-
pensor known only in section Incani. Abnormalities in the embryo proper as well as in the suspensor are
observed at the globular stage. In both A. cemerinus and A. ruscifolius, fusion of the polar nuclei occurs in the
median regions of the central cell and before fertilization occurs, as is the rule in most of the Papilionoideae, but
in species of section Incani as in a few other species of the family, the polar nuclei approach the egg apparatus
before fertilization and do not fuse until fertilization. The embryological characters of A. cemerinus and A. rus-
cifolius are compared with those of other species of Astragalus, and the taxonomic application of these charac-
ters as well as their phylogenetic significance are discussed.
Key words: Abnormal embryology, Astragalus cemerinus, Astragalus ruscifolius, megagameto-
phyte, embryo proper, suspensor, phylogeny.
ACTA BIOLOGICA CRACOVIENSIA Series Botanica 51/1: 111–117, 2009
PL ISSN 0001-5296 © Polish Academy of Sciences and Jagiellonian University, Cracow 2009