0012-4966/03/1112- © 2003 MAIK “Nauka / Interperiodica” 0553
Doklady Biological Sciences, Vol. 393, 2003, pp. 553–556. Translated from Doklady Akademii Nauk, Vol. 393, No. 4, 2003, pp. 570–573.
Original Russian Text Copyright © 2003 by Nikolaev, Berney, Fahrni, Mylnikov, Petrov, Pawlowski.
Several years ago, the taxonomy of protists (unicel-
lular eukaryotes) was still based exclusively on mor-
phological and ultrastructural characters. For example,
the taxonomic group Heliozoa was distinguished on the
basis of the body shape and the presence of radial
axopodia emerging from the protist body and rein-
forced from inside by microtubules. For a long time,
this group was considered to be monophyletic. Studies
on ultrastructural organization demonstrated that the
mitochondrial cristae in centrohelid and desmothoracid
Heliozoa were lamellar and tubular, respectively.
Because it is generally believed that the shape of the
mitochondrial cristae is one of the most conserved
characters related to the basal radiation of eukaryotes
[1], this finding cast doubt on the suggestion that cen-
trohelid and desmothoracid Heliozoa are closely
related [2]. On the other hand, it was suggested on the
basis of these data that centrohelid Heliozoa were
closely related to gymnophryid amoebas (with lamellar
mitochondrial cristae), whereas desmothoracid Helio-
zoa (with tubular mitochondrial cristae) were classified
as a taxon related to Cercomonadea [3].
Although the majority of protistan taxa identified on
the basis of ultrastructural characters and the shape of
mitochondrial cristae have retained their status, the use
of molecular data for phylogenetic reconstruction pro-
vides many examples of uniting protists with different
shapes of mitochondrial cristae into monophyletic
groups. For example, the use of molecular data based
on the marker 18S rRNA suggested that there was a
morphologically heterogeneous group combining the
following taxa: Cercomonadea, Chlorarachnea, shell
and naked filose amoebas, and Phytomyxea [4, 5].
Later, a representative of gymnophryid amoebas (Gym-
nophrys cometa) which has lamellar mitochondrial
cristae was added to this group of protists with tubular
mitochondrial cristae. The life cycle of most represen-
tatives of this new group of Cercozoa is complicated; it
includes the double-flagellar amoeboid and the cyst
stages. The results of molecular analysis suggest that
this life cycle is original. Therefore, the life cycles of
both amoeboid organisms and Flagellata have under-
gone secondary reduction, with of one of stages being
lost [6]. The complicated life cycle and the presence of
extrusomes of the kinetocyst type that give grounds to
suggest that desmothoracid Heliozoa and Cercozoa are
closely related.
Because recent molecular phylogenetic studies
failed to demonstrate that centrohelid Heliozoa was a
closely related to any other group of protists [7], the
goal of this work was to test the hypothesis that des-
mothoracid Heliozoa were closely related to the main
group of Cercozoa. To test this hypothesis, we
sequenced the 18S rRNA gene in two representatives of
desmothoracid Heliozoa, Clathrulina elegans
(AY305009) and Hedriocystis sp. (AY305010). The
resulting sequences were aligned with the sequences of
the 18S rRNA gene in representatives of different
groups of protists and subjected to phylogenetic analy-
sis. The results of the study provide unambiguous evi-
dence that desmothoracid Heliozoa and gymnophryid
amoebas are closely related subgroups of Cercozoa.
The 18S rRNA gene of Clathrulina elegans was
obtained by synthesis of cDNA on an RNA template
using a universal eukaryotic primer sB [8]. The result-
ing cDNA was used for amplification of the 18S rRNA
gene using universal primers sA and sB [8], and the
obtained product was reamplified using two specific
primers, s14nBr (5'-AGCCGC A(AG)(AG)CTC CAC
TCC TGG-3') and s13Cl (5'-CCG ACT AGA(AT)AT
GGG TAT T(AC)G-3') along with universal primers sA
and sB. The complete sequence of the 18S rRNA gene
of Hedriocystis sp. was obtained by amplification of
three overlapping gene segments limited by the follow-
GENERAL BIOLOGY
Genetic Relationships between Desmothoracid Heliozoa
and Gymnophryid Amoebas as Evidenced by Comparison
of the Nucleotide Sequences of 18S rRNA Genes
S. I. Nikolaev*, S. Berney**, J. Fahrni**, A. P. Mylnikov***, N. B. Petrov*, and J. Pawlowski**
Presented by Academician A.A. Bogdanov June 30, 2003
Received July 8, 2003
*Belozersky Institute of Physico-Chemical Biology,
Moscow State University, Vorob’evy gory, Moscow,
119992 Russia
**Department of Zoology and Animal Biology,
University of Geneva, Switzerland
***Papanin Institute for Biology of Inland Waters,
Russian Academy of Sciences, Borok,
Yaroslavl oblast, 152742 Russia