The forest tree species generally display very high level of
genetic diversity (Müller-Starck et al. 1992) and therefore
an accurate analysis of the entity and distribution of their
genetic resources requires the screening of many popula-
tions and many individuals within populations. Hence the
choice of appropriate genetic markers assumes great
importance in order to reduce costs and to save time. In
particular genetic markers should be highly polymorphic
and inherited in a Mendelian codominant manner.
Microsatellites, or simple sequence repeats (SSR), that are
tandem repeats of six base pairs or less (Litt & Luty 1989;
Weber & May 1989), have these characteristics and there-
fore seem to be suitable genetic markers for population
genetic studies also in forest tree species. The length vari-
ation of the tandem repeats can easily be scored following
amplification of the SSR-containing fragments by the poly-
merase chain reaction using unique flanking primers. In
plants the possibility to screen highly polymorphic
microsatellite regions also in the chloroplast genome,
which shows lower mutation rates than the nuclear
genome and is usually uniparentally inherited, might
increase the knowledge about the biology of these organ-
isms. In particular chloroplast microsatellites may be very
useful for studying cytoplasmic diversity, cytonuclear
interactions and monitoring gene flow.
Powell et al. (1995a,b) recently demonstrated the utility
of the analysis of the length variation of chloroplast
microsatellites. In particular they analysed (Powell et al.
1995b) a mononucleotide microsatellite located in the
intergenic region between the trnK and psbA genes for
assaying genetic variation in pine species. It was shown
that the primers worked in all the species tested and that
the amplified fragment showed high levels of polymor-
phisms due to size variations between and within species.
Sequencing of the fragments confirmed that the size dif-
ferences were due to variation in the repeat units forming
the microsatellite region. The level of within and between
population diversity detected in Pinus leucodermis Ant., a
tertiary relic and highly endangered species, were much
higher than those previously reported for the same popu-
lations with isozyme markers (Boscherini et al. 1994).
PCR–RFLP analysis (Boscherini et al. 1994) had previously
failed to detect any polymorphism in the cpDNA molecule
in this species.
These observations suggest that the identification of
other tandem repeats localized in the chloroplast genome
may be very useful. The search for cp-microsatellites is
greatly facilitated by the availability of the full sequence of
the cpDNA for an increasing number of species. Due to the
high degree of sequence conservation of the chloroplast
genome (Clegg et al. 1992), especially in its coding
sequences, it should be possible to design primers that
could work across a wide range of species. The persistence
of homologous A/T stretches in the chloroplast DNA of
distantly related species such as Oenothera and Nicotiana
has been observed beyond theoretical expectations
(Tachida & Iizuka 1992) and provides further support to
the use of primers targeting such regions across different
species.
Recently Wakasugi et al. (1994) sequenced the complete
chloroplast genome of the black pine (Pinus thunbergii
Parl.). As reported by Powell et al. (1995a,b) 19 A/T and
one G/C mononucleotide SSRs (flanking another A/T
stretch) of 10 or more repeat units are present in it and no
dinucleotide SSRs of similar length are found. 20 primer
pairs flanking the mononucleotide stretches in the P. thun-
bergii sequence were designed (Table 1) using the comput-
er program PRIMER version 0.5 provided by S. Lincoln, M.
Daly and E. Lander (Whitehead Institute, Cambridge, MA,
USA). One region with a complex stretch of Ts and Cs
(Pt15169) was added to the 19 above mentioned ones, even
though neither of the repeated regions in it was 10 repeats
long. Primers were designed in order to allow multiplex-
ing by size and thus save considerable time in amplifica-
tion and/or running gels. The primers were then used to
amplify DNA samples extracted from P. leucodermis. DNA
was extracted from germinating embryos according to the
CTAB procedure outlined by Doyle & Doyle (1990). PCR
PRIMER NOTE
A set of primers for the amplification of 20 chloroplast
microsatellites in Pinaceae
G. G. VENDRAMIN, L. LELLI, P. ROSSI andM. MORGANTE*
Istituto Miglioramento Genetico Piante Forestali, CNR, Via Atto Vannucci 13, 50134 Firenze, Italy and *Dipartimento di
Produzione Vegetale e Tecnologie Agrarie, Università di Udine, Viale delle Scienze 208, 33100 Udine, Italy
Molecular Ecology 1996, 5, 595–598
Keywords: chloroplast genome, microsatellites, Pinaceae, simple
sequence repeats
Received 25 November 1995; revision accepted 6 February 1996
Correspondence: G. G. Vendramin, Istituto Miglioramento
Genetico Piante Forestali, CNR, Via Atto Vannucci 13, 50134
Firenze, Italy. Tel.: number: + 39-55-461071, Fax: + 39-55-486604,
E-mail: vendramin@imgpf.fi.cnr.it
© 1996 Blackwell Science Ltd