Table Of ContentArnaldoa 22 495 506, 2015 ISSN: 1815-8242 (edición impresa)
(2): -
ISSN: 2413-3299
(online edition)
The chromosomes
of and endemic genus
the rare
Famatinanthus
(Famatinanthoideae, Asteraceae)
cromosomas
Los endémico género
del raro
y
Famatinanthus
(Famatinanthoideae, Asteraceae)
ARNALDOA
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I
The chromosomes Famatinanthus
Chiarini eta!:. of
F.
Classical staining and fluorescent hybridization with probes for the 18-5.8-26S and 5S
in situ
know
genes were performed in Famatinathus decussatus (endemic from Argentina), in order to
main chromosomal and compare them The
characteristics to to related genera. species a
its is
=
paleopolyploid with 2n likely originated from x=9, and karyotype features are conservative:
54, its
one pair of 5S signáis and two of 18-5.8-26S per complement were found. All the data are congruent
with the basal position of this species in the Asteraceae phylogenies.
Keywords: Chromosomes, endemism, yDNA.
Famatinanthus polyploidy,
decussatus,
&
99%
Famatinanthus Ariza E. Freire of Asteraceae. These characteristics
S. is
made
a recently described genus of Asteraceae Famatinanthus to diserve a subfamily
endemic to a restricted area in the province only for itself Considering this background
.
know
La chromosomal
of Rioja, Argentina (Freire et al, 2014). Its it is important to the
single species, decussatus (Rieron.) Ariza characteristics of the species.
F.
&
was
formerly placed within
S. E. Freire,
From
the cytological point of view,
Wedd. However,
Aphyllocladus analyses of
chromosome
counts provide a key tool
floral characters corollas, anthers, style,
(i.e.
phylogeny and
in studies of systematics,
achenes), stem anatomy, trichomes and
and
evolution, they are especially useful for
pollen (Freire et al, 2014), indicate that the
and
understanding speciation hybridization
species has a combination of features that
and
(Stebbins, 1971). Structural quantitative
do not correspond with the circumscription
karyotypes have
characteristics of
of Aphyllocladus or any other genus within
been and
evolutionary
significant in
the Mutisioideae Consequently, a
tribe.
many
taxonomic angiosperm
studies in
new
genus was proposed accommodate
to
groups being
(Stebbins, decisive in
1971),
The f
not only
singularities of decussatus
it.
. groups and
establishing linkage natural
morpho-anatomical
involve features; in
fact,
Weiss-Schneeweiss
classifications.
(e.g.
molecular phylogenetic study using 14
in a
The
et al, 2003; Pellicer et al, 2010).
DNA
Panero
chloroplast al
loci. et (2014)
FISH homologous
procedure allows
named
found genus has
that the recently
chromosomes complement
be
in a to
two
the chloroplast inversions present in
differentiated and permits the comparison
Asteraceae except the nine genera of the
all
among
related species (Heslop-Harrison,
subfamily Barnadesioideae and concluded
2000; Garcia et al, 2007; Leitch et al, 2008).
that is sister to the Mutisioideae-
it
The procedure provides information
also
more
Asteroideae clade that represents than
ARNALDOA
496 22(2): Julio -Diciembre, 2015
I
The chromosomes Famatinanthus
Chiarini eta/:.
F.
on genome organization and allows stored at -20 °C. Some of these slides were
chromosomal evolutionary questions to used for classical staining with Giemsa.
be addressed (Chacón ei al, 2012; Pellicer The reamining stored slides were used
The most and number
determining
et al, 2010; Chiarini et al, 2014). for the location
rDNA
common One
molecular-cytogenetic markers are of by FISH. of the probes
sites
ribosomal genes (5S and 18-5.8-26S rDNA), was the pTa71 containing the 18-5.8-26S
rDNA &
which are abundant and highly conservad in (Gerlach Bedbrook, labeled
1979)
&
higher plant species (Schmidt Heslop- with biotin-14-dATP (BioNick; Invitrogen,
all
rDNA,
Harrison, 1998). Variations in the number, Carlsbad, USA). For the 5S a
rDNA
signal intensity and position of loci probe was obtained from the genome of
common PCR
seem to be in several plant groups Prionopsis ciliata by (Moreno et al,
&
(e.g. Datson Murray, 2006; Urdampilleta 2012) labeled with digoxigenin-ll-dUTP
The FISH
et al, 2013), suggesting their mobility. (Roche Diagnostics). procedure
Among
number and was accordance with Schwarzacher
relatad species, the in
rDNA &
may
location of be conservad Heslop-Harrison with minor
loci
(2000),
among
or vary considerably populations modifications. The preparations were
with different ploidy levels Adachi incubated in 100 Ig/ mi RNase, post-fixed in
(e.g. et
Lan Considering 4 % (w/v) paraformaldehyde, dehydrated
al, 1997; et al, 2011). this
background, the aim of our work was to in a 70-100% graded ethanol series, and air-
On
describe the basic cytogenetic features of dried. each slide 15 pl of hybridization
Famatinathus, together with the patterns mixture was added ng/pl of probe, 100%
(3
of 18-5.8-26S and 5S genes, in order to formamide, 50% dextran sulfate, 20 x SSC
compare them to relatad genera. and 10% SDS), previously denatured at 70°C
Chromosome
10 min. denaturation/
for
and Methods
Materials
hybridization was done 90°C 10 min,
at for
Seeds were bulked from plants in 48°C 10 min, and 38°C 5 min using
for for
several natural populations in La Rioja thermal Eppendorf,
a cycler (Mastercycler,
Famatina
province, Dpt. (S28°52'00.9” Hamburg, Germany), and were
placed
slides
W67°41'28.1", G. Barboza et al 4268; in a humid chamber 37°C overnight. The
at
S28“50'28" W67°40'56.3", Barboza
G. et probes were detected with avidin-FITC
al 4270; S28°39'58.2" W67°42'05.5", G. and
conjúgate antidigoxigenin-rodamine
Voucher specimens
Barboza et al 4271). are conjúgate and counterstained and mounted
Herbarium
depositad the of the National
at with 25 pl antifade (Vectashield Vector
Córdoba (CORD).
University of Burlingame, USA), containing 2ng/pl
Lab.,
DAPI.
of
chromosomes were examinad
Mitotic in
root tips obtained from seeds germinated in
Results
Root were
Petri dishes. tips pre-treated in
most
Seeds resulted inviable in collected
saturated p-dichlorobenzene in water for 2
Only showed
capitula. accession G.B. 4271
h room
at temperatura, fixed in 3:1 ethanol/
good
a germination thus the analyzed
rate,
washed
acetic acid, in distilled water,
were from sample and
they
cells all this
PECTINEX ® min
digested with 37
(45 at
= Chromosomes
2n
presentad
54
(Eig.
1).
and squashed drop 45%
°C), in a of acetic
compared
are small to relatad genera of
removal
acid. After coverslip in liquid
Asteraceae with 2n = being the average
54,
were and
nitrogen, the slides air dried
pm
=
length c 1.75 ± 0.21 and the total
ARNALDOA
497
22 Julio - Dicie
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I
The chromosomes Famatinanthus
Chiarini eta!:. of
F.
pm
LT =
haploid lenght of 94.05 ± 2.95 (Table in several review articles Soltis
(e.g.
&
The chromosomes homogeneous Hegarty
are in et al, 2003; Hiscock, 2008;
1).
&
Van
size, with a ratio between the largest and the Leitch Leitch, 2008; de Peer et
common
complement Notable Autopolyploidy
shortest of the ca. 1.87. al, 2009). is a
secondary constrictions were detected phenomenon and regarded as frequent
it is
&
two chromosome most angiosperms (Ramsey Schemske,
in pairs in of the in
metaphases lA). Centromeres were Within Asteraceae, Barker al
(Fig. 1998). et
difficult to visualize, and the high number (2008) revealed at least three ancient whole-
of chromosomes hindered identifying the genome duplications: a one (shared by
first
homologue
Mutisioideae, Carduoideae, Cichorioideae
pairs.
and
Helianthoideae) placed near the
FISH
technique with 18-5.8-26S probe
origin of the family just prior to the rapid
evidenced four loci in terminal position,
and two independent
radiation of
tribes,
its
coinciding with the secondary constrictions
genome
duplications near the base of the
visualized with the conventional staining,
We
and
Mutisieae Heliantheae. can
tribes
showed two
while the 5S probe terminal
genome
speculate that the duplication
signáis per cell (Fig. IB), located in asynteny
near the base of Mutisieae also affected
chromosomes
respect the bearing 18-5.8-
to
Famatinanthus since they are sister clades.
26S.
would an
Thus, Famatinanthus be ancient
Discussion
polyploid.
Chromosome As
number. most
well as Chromosome
There
a
size. is
basal Asteraceae, Famatinanthus resulted phenomenon genome
to downsizing
called
be a high polyploid with a hypothetical basic whereby do have genome
polyploids not a
number = chromosome
x Polyploids with
9. size exactly equal to the múltiple of their
numbers derived from x = 9 have been which somewhat
diploid progenitors but
is
reported for Barnadesioideae (Wulff, 1990), removal
smaller, as a result of a of
DNA
Waisman &
Mutisioideae (Ward,
1983; et al, redundant (Leitch Bennett, 2004).
and
Grau, (Gibbs
1984; 1987) Stifftioideae Famatinanthus has small chromosomes
&
Ingram, 1982) (Table In fact, this compared hexaploid members
1). to of
number
has been pointed out as basic for and
Barnadesiodeae Mutisioideae (Table
which congruent with
the entire family, Only Doniophyton have
is species of
c
1).
the basal position of Famatinanthus within and LT valúes minor than
decussatus.
F.
Semple
Asteraceae (Panero Genome
et al, 2014). size related to the size of single
is
&
Watanabe
hypothesized
that chromosomes
(2009) Garnatje
(e.g. et al, 2004),
downward
from
múltiple dysploid events
and
therefore the relatively small size of the
on =
polyploids based x 9 account for nearly chromosomes would be due
of decussatus
F.
numbers
all the base reported for Mutisieae, to the longer time that the species have had
numbers = would
the x 8 to 32 be the result from
polyploidization rearrange
to
its its
=
from
of a long dysploid series x2 27. In genome and redundant
eliminate elements.
Famatinanthus remarkable by
this sense,
is
rDNA
Regarding Asteraceae
sites.
conserving the paleopolyploid number.
FISH
species related to Famatinanthus,
Recurrent polyploidy. Polyploidy
is experiments with 18-5.8-26S probe were
many
important for aspects described conducted only in the Mutisioideae species
ARNALDOA
498 22(2): Julio -Diciembre, 2015
I
The chromosomes Famatinanthus
Chiarini eta/:.
F.
Fig. Metaphasic chromosomes of Famatinanthus decussatus with (A) Giemsa staining and (B)
1.
FISH Arrowheads NORs, and
technique. point to arrows point to 18-5.8-26S loci (green signáis)
pm.
asterisks indícate 5S loci (red signáis). Bars represent
3
2n = The amount and
Chaptalia nutans, 100 (Fregonezi other. great variability in
50,
rDNA
with two and four pairs of position of in related species
et al, 2004) loci is
known
signáis per complement, respectively, a fact already (Hasterok et al, 2006;
&
and two
Heslop-Harrison Schwarzacher,
also in species of Chaetanthera 2011;
(Baeza where and Chacón Morales For
et al, 2005a), 1 3 pairs et al, 2012; et al, 2012).
of signáis in diploids with 2n = 22 were these rapid changos of copy number and
rDNA,
found. Regarding the 5S, data in species chromosomal location of different
more
or less related to Famatinanthus are types of transposable elements have been
known
only for Chaetanthera (Baeza et al, postulated as responsible (Raskina et al,
2005a) in which 3 and 4 pairs of 5S signal 2004a, b; Belyayev et al, 2005; Altinkut et al,
&
in diploid species with 2n = 22 have 2006; Datson Murray, 2006).
been found. This situation evidenced the
In higher eukaryotes, the 18-5.8-26S
rDNA
pathways
different that the loci
rDNA rDNA
and
5S are transcribed
loci
two from
took since the lineages each
split
RNA
by polymerases and
different
ARNALDOA
22 Julio - Dicie
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The chromosomes Famatinanthus
Chiarini eta!:. of
F.
more
usually located in different positions of hexaploid, than a pair of 5S signáis
&
chromosomes complement
(Srivastava Schlessinger, per could be expected. Instead,
1991). Colocalization of 18-5.8-26S and 5S only a pair was found, which would be
rDNA
commonly
has been reported an indication of an ancient polyploidy.
loci
It
number
in animáis (Dobigny et al, 2003) but has been found that the of loci of
it
less frequent in plants Garcia 5S subject to dynamic, rapid, changes
is (e.g. et is
&
Abd
2007, 2009b; El-Twab Kondo, Loss of gene copies might be
al, sup.).
(cfr.
A
Chang
2006; et al, 2009). dominant taking place at these loci, as demonstrated
rDNA
was members
linked genotype found within in of other several plant families
three large groups of Asteraceae (García (i.e. Kotseruba et al. 2003; Renny-Byfield et
A
was
Anthemideae, Gnaphalieae observed
similar situation
et al, 2010): al, 2012).
and
in the "Heliantheae alliance" in polyploid species in other Asteraceae
The remaining genera such Brachyscome
(Asteroideae). five tribes of as (Asteroideae,
non Adachi
the Asteroideae displayed canonical Artemisia (Asteroideae,
et al, 1997),
rDNA,
linked arrangement of and also the Garcia et al, 2009a; Pellicer et al, 2013),
Xeranthemum
remaining 12 subfamilies, with sepárate (Carduoideae, Garnatje
et al,
Tragopogón Buggs
organisation. Famatinanthus perfectly (Cichorioideae,
2004),
fits
into this scheme, with the 5S and 18-5.8- et al, 2009; Chester et al, 2012), in which
26S loci in sepárate chromosomes. The the number of 18-5.8-26S sites seemed to
results of García al indícate that evolve faster than the 5S as they do
et (2010) sites,
may
25% genome
nearly of Asteraceae species have not increase with the successive
By genome
developed unusual arrangement
linked of additions. constrast, sizes of
rRNA
genes, and the 5S gene integration some polyploid taxa present additivity,
genome
within the 35S unit might have repeatedly although data pointing upsizing
to
and
occurred during plant evolution, in older polyploids are limited (Garcia
et al,
probably once
in Asteraceae. 2009; Leitch et al, 2008).
rDNA
Genome phenomenon FISH would
downsizing, with probes
a studies
rDNA
more
referred before, also affects genes. be desirable in genera closely related to
During Famatinanthus
the polyploidization process, Barnadesia, Duseniella,
(e.g.
may
gene reordering and gene silencing Dasyphyllum) in order to achieve a more
chromosomal
occur (Stebbins, 1985; Soltis et al, 2003; complete picture of evolution
&
Leitch Bennett, 2004; Pires et al, 2004). in basal Asteraceae.
In Solanaceae, for example, an analysis of
Acknowledgements
rDNA
the 18-5.8-26S of Nicotiana indicated
The authors thank Consejo Nacional
were maintained
that parental loci initially
in newly formed polyploids, although de Investigaciones Cientifíficas y Técnicas
(CONICET, SECyT-UNC, and
Argentina),
the sequences within a locus might be
SECyT-UNRC
Thanks
and for financial support.
subject to concerted evolution, over
tendered
periods greater than myr, individual are also to
1 loci
would
disappear (Kovafík In
et al, 2008).
D. Olivera for his unvaluable assistance
was
found
Asteraceae, a similar situation in
work
in Famatina's field (La Rioja).
Tragopogón (Buggs Chester
et al, 2009; et
al, 2012).
Given Famatinanthus
that a
is
ARNALDOA
500 22(2): Julio -Diciembre, 2015
I
The chromosomes Famatinanthus
Chiarini eta/:.
F.
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