Molecular Ecology Notes (2001) 1, 103–105
© 2001 Blackwell Science Ltd
Blackwell Science Ltd
TECHNICAL NOTE
Technique for analysis of bryozoan mitochondrial DNA
J. S. PORTER, D. O. F. SKIBINSKI, J. LEAMON and P. J. HAYWARD
School of Biological Sciences, University of Wales, Swansea, Singleton Park, Swansea, SA2 8PP, UK
Abstract
The use of non-specific mitochondrial DNA (mtDNA) primers can frequently result in the
erroneous amplification of DNA from contaminating organisms or from mtDNA nuclear
inserts. A reliable method for deriving species-specific mtDNA polymerase chain reaction
(PCR) primers using genomic and mtDNA enriched preparations from the same individual
is described.
Keywords: Alcyonidium diaphanum, Bryozoa, mitochondria, mtDNA, PCR
Received 14 September 2000; revision accepted 16 November 2000
Polymerase chain reaction (PCR) analysis of mitochondrial
DNA (mtDNA) from a relatively little studied group such
as bryozoa is hampered by lack of prior sequence data and
by the small amount of tissue available in many species.
This hinders design of species-specific PCR primers. The
task is made more difficult by the presence of bacteria
and phytoplankton on colony surfaces and in the gut.
Use of nonspecific mtDNA primers with genomic DNA
preparations might result in amplification of foreign DNA
or bryozoan nuclear DNA inserts.
To date, universal mtDNA primers have been used in
bryozoa. Dick et al. (2000) amplified, cloned and sequenced
a 16S rRNA gene segment and constructed phylogenetic
trees of 23 Gymnolaemate bryozoan species; Schwaninger
(1999) sequenced cytochrome oxidase I (COI) PCR products
to study population differentiation in the marine bryozoan
Membranipora. Our testing of universal primers for COI
and for the 12S and 16S rRNA genes gave multibanded
PCR profiles with genomic DNA from the bryozoan
Alcyonidium diaphanum (Hudson 1762), and raised concern
about primer specificity.
Nuclear copies of mtDNA sequences occur in many spe-
cies and, although interesting, may confound phylogenetic
analyses (see Zhang & Hewitt 1996). Nuclear insertions can
occur repeatedly during evolution and be highly or little
diverged from normal mtDNA (e.g. Bensasson et al. 2000).
Thus, phylogenetic affinity cannot be relied upon to dis-
tinguish mtDNA from nuclear pseudogenes. Nuclear
contamination can frustrate interpretation in several
ways. In monkeys, mtDNA primers prepared from related
species gave PCR bands containing multiple nuclear DNA
as well as mtDNA sequences even with mtDNA purified
by caesium chloride centrifugation (Collura & Stewart
1995). Differences in the relative amounts of nuclear DNA
and mtDNA between elephant blood and hair caused
differential amplification of the two genomes (Greenwood
& Paabo 1999). Apparent mtDNA heteroplasmy in humans
was subsequently attributed to coamplification of nuclear
mtDNA pseudogenes (Wallace et al. 1997). Although appar-
ent frameshift or nonsense mutations are often present in
mtDNA like nuclear sequences, protein-coding potential
has also been observed (Collura et al. 1996).
The use of mtDNA enriched preparations for PCR ana-
lysis has been advocated previously (e.g. Burgener &
Hubner 1998). However, this cannot solve all the above
problems when nonspecific primers alone are available.
Because it is easy to extract genomic DNA and apply such
primers, pitfalls can occur easily. We describe here a tech-
nique exploiting the high concentration of mtDNA in
oocytes. A mtDNA extraction procedure is applied to
oocyte rich tissue: a conventional genomic DNA extrac-
tion procedure to somatic tissue. The former procedure
enriches for mtDNA generally compared with the latter,
whether this mtDNA be bryozoan or from a contaminating
organism. In addition, the use of oocyte rich tissue enriches
for mtDNA that is specifically bryozoan, compared with
the use of somatic tissue. PCR with universal primers gives
distinctively different banding patterns for the two pre-
parations. Bands specific to the mtDNA preparation are
likely to derive from mtDNA rather than nuclear DNA.
mtDNA was extracted from the bryozoan A. diaphanum
using the method of Fisher & Skibinski (1990). Tissue
rich in oocytes was homogenized in an equal volume of
Correspondence: D. O. F. Skibinski. Fax: 01792 295447; E-mail:
d.o.f.skibinski@swansea.ac.uk