Karyological and cytogenetic studies of conifers from West Siberia and Far East

Karyological and Cytogenetic Studies of Conifers

from West Siberia and Far East

E. N. Muratovaa, T. S. Sedel’nikova

a, T. V. Karpyuk

b, O. S. Vladimirova

a,

A. V. Pimenova, N. A. Mikheeva

a, E. V. Bazhina

a, and O. V. Kvitko

a

a

Sukachev Institute of Forest, Siberian Branch of the Russian Academy of Sciences,

Akademgorodok 55/28, Krasnoyarsk, 660036 Russiab

Krasnoyarsk State Agrarian University, pr. Mira 88, Krasnoyarsk, 660049 Russia

Abstract—The results summarize many years of karyological and cytogenetic studies of conifers from the

boreal zone, done at the Sukachev Forest Institute. The species under study belong to four genera of the family

Pinaceae: Larix (larch), Pinus (pine), Picea (spruce), and Abies (fir). Overall, the research covers more than 100

populations and habitats of the species from these genera. The study concerns species growing under both

optimal and marginal conditions. The studies of the family Cupressaceae have been initiated. A special attention

is given to conifers growing on bogs and in planted forests.

DOI: 10.1134/S1995425508020148

Karyological and cytogenetic studies contribute sig-

nificantly to the studies of biodiversity of forest trees

and their polymorphism, problems of taxonomy and

evolutional genetics, structure and genetic composition

of natural populations, and practical aspects of selec-

tion. In the Sukachev Institute of Forest, karyological

studies of tree species were initiated by L. F. Pravdin in

the 1960s. His book [1] presented data on the chromo-

some number and morphology in Scots pine, at the time

one of the first attempts of karyological study of trees in

USSR. In the late 1960s — early 1970s the karyological

and cytogenetic studies were significantly advanced by

M. V. Kruklis who investigated karyotypes of Siberian

and Dahurian larch and their hybrids, meiosis in these

species, and the karyotype of Siberian spruce and dis-

covered B chromosomes in conifers [2–5].

In the 1970s, karyological studies of the main for-

est-forming conifer species of Siberia and Far East

were continued in the Institute of Forest. Several spe-

cific features distinguish conifers from other plants.

They are mostly diploid throughout the life cycle, with

polyploids being exceedingly rare [6–8]. Plants of this

group have large amounts of DNA and large genomes

[9–11]. The chromosomes in the karyotype are numer-

ous; they are big, stain well, and are well differentiated

morphologically but similar in length. In addition, spe-

cies of one genus have invariable number and morpho-

logical type of chromosomes [12]. This makes conifers

a special and complicated object of karyological stud-

ies.

The species under study belong to four genera of

the family Pinaceae: Pinus L. (pine), Picea A. Dietr.

(spruce), Larix Ledeb. (larch), and Abies Mill. (fir). At

the moment, the study includes six species of larch,

seven species of pine, eight species of spruce, and four

species of fir. Overall, about 150 populations and habi-

tats have been investigated. The studies involved both

optimal and marginal growth conditions and ecosys-

tems disturbed by human activity. The karyotypic di-

versity and wide spectrum of chromosomal abnormali-

ties in conifers have been revealed, chromosomes from

the species of Larix, Pinus, Picea, and Abies have been

exhaustively studied. Studies of species from the family

Cupressaceae (Juniperus L., Thuja L.) have been initi-

ated.

The species of the genus Pinus L. represent two sub-

genera, Strobus and Pinus. Of the former, three species

of section Strobi, and subsection Cembrae have been

investigated in detail: P. sibirica Du Tour, P. koraiensis

Siebold et Zucc., and P. pumilla (Pall.) Regel, and P.

cembra was studied for comparison. Five of the species

under study represent the subgenus Pinus (section Pi-

nus, subsection Sylvestres). Scots pine (P. sylvestris L.)

was investigated in most detail, and supplemented with

studies of P. densiflora Siebold et Zucc., P. thunbergii

Parl., P. funebris Kom., and P. eldarica Medw.

Karyotypes of pines consist of 24 chromosomes

(2n = 24). Cembrae possess 11 pairs of symmetric

(metacentric) and one pair of asymmetric (submeta-

centric or similar to that) chromosomes [13–18]. Pines

of the Sylvestris group possess 10 pairs of symmetric

and two pairs of asymmetric chromosomes [19–26].

The pine species differ in the number and localization

of secondary constrictions. Figures 1–3 show chromo-

some sets and karyograms of several species from the

genus Pinus.

Nine representatives from the genus Picea A. Dietr.

have been studied: P. abies (L.) aest., P. fennica Regel

(Kom.), P. obovata Ledeb., P. schrenkiana Fisch. et

Mey, P. koraiensis Nakai, P. glehnii (Fr. Schmidt)

ISSN 1995-4255, Contemporary Problems of Ecology, 2008, Vol. 1, No. 2, pp. 263–271. © Pleiades Publishing, Ltd., 2008.

Original Russian Text © E.N. Muratova, T.S. Sedel’nikova, T.V. Karpyuk, O.S. Vladimirova, A.V. Pimenov, N.A. Mikheeva, E.V. Bazhina, O.V. Kvitko, 2005, published in

Sibirskii Ekologicheskii Zhurnal, 2005, Vol. 12, No. 4, pp. 573–583.

263

Mast., P. meyeri Rehd. from the Picea section; P. ajan-

ensis (Lindl. at Gord.) Fisch. ex Carr. and P. jezoensis

(Siebold et Zucc.) Carr. from the Casicta section. The

karyotypes of these spruce species contain eight pairs

of long metacentric, two pairs of short metacentric, and

two pairs of short submetacentric chromosomes [3,

27–40]. The karyotypes of the studied species may con-

tain one or several B chromosomes: P. schrenkiana 2n =

24 + 1B, P. ajanensis 2n = 24 + 1–3B, P. meyeri 2n = 24

+ 1–3B, P. obovata 2n = 24 + 1–4B, P. glehnii 2n = 24 +

1–5B [3, 4, 28, 29, 31, 37–39, 41–51]. Figures 4–6

show chromosome sets and karyograms of several spe-

cies from the genus Picea.

The larch species included in the analysis represent

two related groups of species. One group includes

Larix sibirica Ledeb. and L. sukaczewii Dylis, while the

second group consists of L. gmelinii (Rupr.) Rupr.,

L. cajanderi Mayr, L. ochotensis Kolesn., and L. amur-

ensis Kolesn. The diploid set of larches consists of

24 chromosomes (2n = 24). The karyotype includes six

pairs of symmetric (metacentric) and six pairs of asym-

metric (submetacentric or intercentric) chromosomes

[4, 52–61]. Different larch species vary in the number

and localization of secondary constrictions. An addi-

tional chromosome, 2 24 1n � � B, was found in L. gme-

linii [60, 62]. The ordered karyotype of the Okhotsk

larch is shown in Fig. 7, and a chromosome set of the

Dahurian larch with a B chromosome, in Fig. 8.

The study involved the following species of fir:

A. sibirica Ledeb., A. lasiocarpa (Hook.) Nutt., A. ho-

lophylla Maxim. from the Piceaster section; A. sa-

chaliensis Fr. Schmidt from the Elatae section.

Karyotypes of Abies species contain seven pairs of

metacentric and four pairs of submetacentric and sub-

acrocentric chromosomes [63]. Different fir species

also vary in the number and localization of nucleolar re-

gions. The karyogram of Siberian fir is presented in

Fig. 9.

Our studies show that the main difference between

karyotypes of conifers is the number and localization of

secondary constrictions. The secondary constrictions

are functionally very important chromosomal loci: In

most eukaryotes, they contain ribosomal RNA genes

and mark places of ribosome assembly [64–66]. The ac-

tivity of these genes becomes morphologically appar-

ent in the telophase of mitosis with the formation of

nucleoli.

The nucleolar organizers are the best studied loci

with respect to their structure and function. Direct

rRNA and DNA hybridization shows that the nucleolar

organizers of conifers contain actively transcribed 18S

and 25–26S rRNAgenes [67, 68]. The existing methods

of staining with silver nitrate allow one to visualize

functional rRNA genes in chromosomes. This tech-

nique have been developed by us and used to stain chro-

mosomes in conifers.

This approach reveals that chromosomes of the

Larix and Picea species are stained not only in second-

ary constrictions, but also in telomeres of some chro-

mosomes [69]. Figure 10 shows chromosomes of

Siberian larch with silver-stained nucleolar organizers,

and the karyogram of this species with marked nucleo-

lar regions. It can be suggested that telomeric regions of

chromosomes in conifers contain additional nucleolar

organizers, which are turned on under extreme condi-

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 1 No. 2 2008

264 MURATOVA et al.

Fig. 1. Ordered karyotype of Scots pine (Pinus sylvestris L.).

2n = 24. I–IX, X, XI, XII are chromosome numbers.

Fig. 2. Chromosome set of Siberian pine (Pinus sibirica Du Tour).

2n = 24.

Fig. 3. Karyogram of Siberian dwarf pine (Pinus pumila Regel).

Bar, 5 �m. 2n = 24. I–XII are chromosome pair numbers.

tions or if the activity of the main nucleolar organizer

decreases.

Until the late 1970s, the karyotypes of conifers were

regarded as stable. However, this was only due to insuf-

ficient knowledge of karyology and cytogenetics of this

group at that time. Our studies of many populations re-

vealed wide karyotypic diversity and a number of chro-

mosomal mutations. Under optimal growth conditions,

chromosomal abnormalities in conifers are exceedingly

rare. However, deviations from the chromosome num-

ber and morphology are observed at the margins of the

natural species habitat and in ecosystems disturbed by

human activity. Genomic and chromosomal mutations

appear, the number of nucleolar regions in the chromo-

somes increases, and chromosomal mutations occur

more frequently.

Scots pine is the species for which the most karyo-

logical and cytogenetic data is available. These meth-

ods reveal a wide spectrum of chromosomal mutations

observed at the northern and souther borders of the nat-

ural species habitat [70–79]. These include ring and

polycentric chromosomes, deletions, fragmentation

and other anomalies (Fig. 11). The same trees show ab-

normal mitosis and meiosis, suggesting the distur-

bance in the development of their reproductive system

[75–77, 80]. The abnormalities in mitosis and meiosis

in Scots pine are shown in Figs. 12 and 13.

Chromosomal mutations have been observed in Si-

berian larch from Kazakhstan and Mongolia, Sukachev

larch from the South Urals, southern populations of

Dahurian larch, northern populations of East Siberian

larch, Siberian dwarf pine, and Siberian spruce, Sibe-

rian fir from the Khamar-Daban Mountains, Jezo

spruce from Primorye and Yakutia, and in Japanese red

and black pines introduced to new habitats [23, 29, 34,

56, 58–61, 63, 74, 76, 78, 79]. The studied species of

pine, larch, spruce, and fir also showed genomic muta-

tions, such as aneuploidy and mixoploidy, with a few

occurrences of polyploidy. Chromosomal abnormali-

ties of this kind are presented in Fig. 14.

An example of homozygous pericentric inversion,

simultaneously affecting two pairs of chromosomes,

has been found in Dahurian larch from Eastern Trans-

baikalia [60]. The affected chromosomes, normally

submetacentric, turned into metacentric. This type of

chromosomal mutations has not been described in coni-

fers before, but was later found in the Siberian fir [81].

Our studies showed that pericentric inversion is often

observed in various spruce species and could possibly

promote speciation in this genus [36, 37, 39, 44, 45].


KARYOLOGICAL AND CYTOGENETIC STUDIES OF CONIFERS 265

Fig. 4. Chromosome set of Jezo spruce (Picea ajanensis (Lindl. et

Gord.) Fisch. ex Carr.). 2n = 24.

Fig. 6. Karyogram of Siberian spruce (Picea obovata Ledeb.) with

an additional chromosome, 2n = 24 + 1B. Bar, 5 �m. I–XII are chro-

mosome pair numbers.

Fig. 5. Chromosome sets of spruce species with different numbers of additional chromosomes: (a) Jezo spruce, one additional chromosome

(2n = 24 + 1B); (b) Siberian spruce (P. obovata Ledeb.) with two B chromosomes (2n = 24 + 2B); (c) Meyer’s spruce (P. meyeri Rehd.) with

three B chromosomes (2n = 24 + 3B).

Analysis of meiosis in the Siberian fir has shown

that it generally follows the scheme common for all co-

nifers. In most cases, the meiosis was regular and the

chromosomes normally segregated to the poles. How-

ever, some abnormalities in various phases of meiosis

have been observed. A comparison of meiosis in the

trees growing naturally and in an arboretum has shown

that introduction to the unnatural habitat increases the

frequency of abnormalities and broadens their spec-

trum in comparison with the natural stand [82, 83]. Ex-

amples of meiotic abnormalities in the Siberian fir are

given in Fig. 15.

A special attention was given to studies of popula-

tions of various conifer species in sandy-loam dry val-

leys and bogs of various types in the southern taiga

subzone of the West Siberian Plain (Tomsk oblast). The

number and types of chromosomal abnormalities in

metaphase and anatelophase were used to assess the

conditions of stands of Scots pine, Siberian pine, Sibe-


266 MURATOVA et al.

Fig. 7. Ordered karyotype of Okhotsk larch (Larix ochotensis

Kolesn.). 2n = 24. I–VI, VII–XII are chromosome numbers. Bar,

10 �m.

Fig. 8. Chromosome set of Dahurian larch (L. gmelinii Rupr.) with a

B chromosome. 2n = 24.

Fig. 9. Karyogram of Siberian fir (Abies sibirica Ledeb.). 2n = 24.

I–XII are chromosome pair numbers. Bar, 5 �m.

Fig. 10. Nucleolar regions of Siberian larch stained with silver ni-

trate: (a) chromosomal nucleolar organizers (black dots); (b) karyo-

gram, I–XII are chromosome pair numbers.

Fig. 11. Chromosomal abnormalities (arrows) in Scots pine:

(a) ring chromosome; (b) dicentric chromosome.

rian spruce, Siberian larch, and Siberian fir [25, 26, 40,

79, 81, 84–90].

Mixoploidy (2n = 24, 36, 48) was found in all stud-

ied species from both dry valleys and bogs, with Sibe-

rian larch also showing aneuploidy (2n = 24, 25, 26).

The spectrum of genomic mutations and chromosomal

rearrangements in metaphase and anatelophase in vari-

ous species was broader in the specimens from bogs

than in those from dry valleys. The Siberian fir was an

exception, showing more chromosomal aberrations and

pathologic mitoses in dry valleys [81]. The aberrations

were represented by ring and polycentric chromo-

somes, fragmentation, and abnormal chromosome spi-

ralization (Fig. 16). The aberrations in anatelophase

included multipolar mitoses, lagging chromosomes,

single and paired chromosomal bridges, fragmentation,

chromosome scattering, and chromosome agglutina-

tion.

In a specimen of the Scots pine with witch’s broom,

unusually shaped and residual nucleoli were observed,

together with mitotic abnormalities and C-mitosis [84,

85, 89]. Mixoploidy (2n = 24, 36, 48) and high fre-

quency of pathologic mitoses were common in artifi-

cially planted Scots pine on a drained eutrophic bog

[90]. The main types of aberrations are multipolar

mitoses, chromosome scattering, lagging and preco-

cious chromosomes, single and paired chromosomal

bridges, fragmentation, C-mitoses, and residual nucle-

oli.

A karyological study of common juniper (Juniperus

communis L.) from bogs and dry valleys showed that

the karyotype of this species contains 22 chromosomes

(2n = 22) (Fig. 18). All chromosomes are metacentric,

with one pair tending to the submetacentric type. No

significant difference in the karyotypes of the bog and

dry valley populations were found, possibly because



Fig. 13. Meiotic aberrations in Scots pine: (a) split spindle in anaphase I; (b) irregular chromosome segregation in anaphase I; (c) triple nu-

clei in telophase II; (d) bridges in anaphase I; (e) possible phenotypic manifestation of the mutant ameiotic gene in metaphase I.

Fig. 12. Mitotic aberrations in Scots pine: (a) lagging chromosome

in anaphase; (b) lagging chromosome fragment in anaphase; (c) lag-

ging pair or fragments in anaphase; (d) broken chromosomal bridge.

the soil hydrological regime of eutrophic bogs is favor-

able for this species [91, 92]. The karyotype of another

representative of the family Cupressaceae (Thuja orien-

talis L.) introduced into an unnatural habitat also con-

tains 22 chromosomes [93].

Karyotypes of plants and animals usually consist of

A chromosomes — the invariable chromosomes of the

set. Some species contain additional B chromosomes.

Over 20 conifer species containing B chromosomes

have been described by now. More than half of them are

different spruce species [50, 51]; recently, B chromo-

somes have been found in larches. Their role in the pop-

ulations and the influence on viability are not clear, and

are subject of active current investigation.

This work was supported by grants from RFBR

(03-04-49719), Krasnoyarsk Territorial Science Foun-

dation (12F0006c and 11F023c), SB RAS Integrative

Program (53, 145), RAS program (12.1), and the Basic

Research Program of the RAS Presidium “Dynamics of


268 MURATOVA et al.

Fig. 15. Meiotic aberrations in Siberian fir: (a) chromosome frag-

mentation in the first division; (b) asynchronous division of the mei-

otic cell with the late anaphase at one pole and the early anaphase

with unequal segregation on another pole.

Fig. 14. Chromosome abnormalities (arrows) in various species of Larix, Picea, and Abies: (a) ring chromosome in Sukachev larch;

(b) chromosome fragments in Sukachev larch; (c) two ring chromosomes in Siberian spruce; (d) dicentric chromosome in Siberian spruce;

(e) dicentric chromosome.

gene pools of plants, animals, and humans” (state con-

tract 10002-251/P-24/154-319/190-504-294).

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