Annals of Botany 82 (Supplement A): 3-15, 1998
Article No. bo980764
Q
Nuclear DNA Amounts in Gymnosperms
BRIAN G. MURRAY
School of Biological Sciences, The University of Auckland, Private Bag 92019, Auckland, New Zealand
Received: 10 September 1997
Returned for revision: 26 March 1998 Accepted: 8 May 1998
A survey of genome size variation in 117 gymnosperms has found that the systematic coverage of the measurements
is uneven, some families have been well sampled but others have not. There is a 144-fold variation in genome size
within the group; the largest genome being that of Pinus lambertianawith 63'5 pg DNA/2C nucleus and the smallest,
1296 pg DNA/2C nucleus in Metasequoia glyptostroboides. The different methods that have been used to measure
genome size in gymnosperms, the choice of standards, and problems associated with the different methods are
reviewed. The conflicting reports of intraspecific variation in genome size in gymnosperm species are problematic.
Some appear real whereas others can be attributed to problems associated with methodology. Some studies of species
show considerable uniformity of genome size over a wide geographical range whereas others find high levels of
variation which may, in some cases, be correlated with geographical parameters. Possible correlations between
genome size and adaptive features of gymnosperms are examined and a number of correlations are reported between
genome size and growth-related parameters.
© 1998 Annals of Botany Company
Key words: Gymnosperms, genome size, intraspecific and interspecific variation in genome size, flow cytometry,
Feulgen microdensitometry, reassociation kinetics.
INTRODUCTION
The gymnosperms comprise a diverse group of woody
plants, many of which are of considerable economic
importance as sources of wood and wood-derived products.
Despite this diversity, they are frequently grouped together
and are usually divided into two subdivisions, the Coniferophytina, which includes the conifers, and the Cycadophytina, which includes the cycads (Page, 1990). However,
there is some debate as to whether the gymnosperms are a
monophyletic group (Page, 1990); some studies suggest that
this grouping is an artificial one (Doyle and Donoghue,
1992; Chase et al., 1993) whereas others have produced
evidence in support of a monophyletic origin (Goremykin et
al., 1986; Chaw et al., 1997). When taken together, the
gymnosperms form a fairly small group of plants arranged
into 17 families with 83 genera and a total of approximately
730 species. The distribution of species in genera and genera
in families is very uneven, some families like the Pinaceae
and Cupressaceae have many genera and more than 100
species, whereas others like the Sciadopityaceae and
Stangeriaceae contain just a single species (Page, 1990).
Thus, compared to the angiosperms with 250000 species
(Heywood, 1993), the gymnosperms form a fairly small
assemblage of taxa.
Analysis of the chromosomes of gymnosperms reveals a
number of unusual and characteristic features. In general
the chromosomes are large. For example, in a study of 36
Pinus L. species, the smallest chromosome of any complement was approximately 6 m long and the longest was
approximately 16 m long (Hizume, 1988). The chromosomes are relatively few in number, 2n = 24 in the Pinaceae
and 2n = 22 in the Cupressaceae. Polyploids are exceedingly
0305-7364/98/0A0003 + 13 $30.00/0
rare and comprise less than 1% of gymnosperm species
(Khoshoo, 1959). In addition, with a few exceptions such as
the Podocarpaceae and Taxaceae, the families are characterized by a constant, basic chromosome number, for
example, x = 12 in the Pinaceae and x = 11 in the
Cupressaceae. Karyotypes of species within genera can also
show a remarkably uniform structure (Pederick, 1970;
Saylor, 1972; Hizume, 1988) but this may mask an
underlying organizational complexity revealed when differential staining techniques and in situ hybridization are
applied to these chromosomes (MacPherson and Filion,
1981; Doudrick et al., 1995; Lubaretz et al., 1996).
SYSTEMATIC COVERAGE OF THE
MEASUREMENTS
Measurements of genome size are available for 12 of the
families but the distribution of these measurements is very
uneven (Fig. 1). The majority of the estimates are for species
of Pinaceae (127) followed by the Cupressaceae (20) and
Podocarpaceae (18). For the remaining families there are
five or fewer estimates. Several features emerge from this
compilation of measurements. There is a significant range in
genome size, 14-1-fold if the measurement of 63-5 pg for
Pinus lambertianaDougl. by Wakamiya et al. (1993) is used
as the largest value, 19-5-fold if the larger measurement of
Rake et al. (1980) for the same species is used. The more
conservative range is probably the most realistic as the 2C
value of 87'7 pg obtained by Rake et al. (1980) is at least
24 pg higher than the next highest gymnosperm value. This
range is clearly significantly smaller than the 600-fold
variation in the angiosperms (Bennett and Leitch, 1995).
© 1998 Annals of Botany Company
4
Murray-Nuclear DNA Amounts in Gymnosperms
FIG. 1. The distribution of 2C DNA amounts amongst 12 families of gymnosperms. All values given in the appendix are included with the
exception of those of Seo et al. (1979). These were obtained from reassociation kinetic studies and are one to two orders of magnitude lower than
other published values.
The widest range of variation in a family, four-fold, is in the
Pinaceae (Fig. 1) which also contains the gymnosperm with
the largest genome size; Pinus lambertiana. The Cupressaceae and Podocarpaceae have a very similar spread of
variation, 29-fold and 27-fold, respectively. Whether
measurements on more species in these families will increase
their range remains to be seen.
It is interesting that genome size in the Gnetaceae, all be
it based on a single measurement, is significantly smaller
than any of the measurements in all the other families. On
the other hand, the single value for Ephedraceae is well
within the range of other families like the Podocarpaceae,
Taxodiaceae and Cupressaceae. Ehrendorfer (1976) suggested that the karyological features of the Gnetales are in
keeping with their status as a distinct group within the
gymnosperms; a view supported by Ohri and Khoshoo
(1986). Recent phylogenetic treatments (Loconte and
Stevenson, 1990; Doyle and Donoghue, 1992) agree on the
grouping of the Ephedraceae, Gnetaceae and Welwitchiaceae as clades distinct from the rest of the gymnosperms.
However, Chaw et al. (1997) consider, on the basis of
information from nuclear 18S rRNA gene sequences, that
the gymnosperms are monophyletic and comprise three
monophyletic clades: Cycadales-Ginkgoales, Gnetales and
Coniferales. Consequently, the distinct karyological features
of these different families, in particular the differences in
genome size, do not appear to have a direct bearing on their
phylogenetic arrangement. The Gnetales would appear to
be a group where more measurements of genome size are
needed if any meaningful phylogenetic relationship in
genome size is to be revealed.
METHODS FOR MEASURING DNA
AMOUNTS IN GYMNOSPERMS
The measurement of genome size in gymnosperms has been
dogged by controversy centred around wide discrepancies
between measurements made by different laboratories. Four
direct methods have been used: Feulgen microdensitometry
(Fe), chemical analysis, reassociation kinetics (RK) and
flow cytometry (FC), but indirect estimates using the
relationship between nuclear volume and genome size have
also been used. The first of these methods, Fe, has been the
most widely used and provides 63% of the values in the
Appendix. Flow cytometry provides the next largest number
of measurements, 31%, with reassociation kinetics providing the remaining 6 % of measurements. None of the
estimates from chemical methods have been included as
Murray-NuclearDNA Amounts in Gymnosperms
they are all on a per cell basis and consequently cannot
reliably be classified as 2C or 4C.
Estimates of genome size by Fe have been the most
controversial for two reasons: firstly because different
authors have obtained very different genome size estimates
using the same species; and secondly, and possibly directly
related to the first reason, because vastly different degrees of
intraspecific variation have been reported in some species.
Greilhuber (1986, 1988) highlighted this problem with
examples of a 2 1-fold variation in Picea glauca (Moench)
Voss between the measurements of Teoh and Rees (1976)
and Rake et al. (1980), and a 25-fold variation in Pinus
lambertiana between the measurements of Dhillon (1980)
and Rake et al. (1980). The basis for the reported differences
has, in part, been attributed to methodological problems
associated with the preparation of material for Feulgen
staining. Greilhuber (1986, 1988) has attributed some of
these differences to the type of fixation used prior to Feulgen
staining in plants that are rich in tannins. He made detailed
studies to compare the effects of two different groups of
fixatives, additive ones like buffered formaldehyde and nonadditive ones such as Carnoy's, on staining. The results of
these studies clearly showed that in species with large
amounts of tannins in the cells, such as gymnosperms, there
was a very significant reduction of Feulgen staining in the
material fixed with non-additive fixatives. Formaldehyde
appears to immobilize tannins, thus preventing them from
inhibiting the Feulgen reaction; a phenomenon called 'selftanning' by Greilhuber (1986). In contrast to the variable
results obtained with non-additive fixatives, in most studies
using formaldehyde-based fixation and Fe the results for the
same species from different laboratories are similar (e.g.
Pinus mugo Turra and Gingko biloba L.: Greilhuber, 1986,
1988; Ohri and Khoshoo, 1986). Greilhuber (1988) concluded that the results of microdensitometric analyses of
gymnosperm nuclei that had been fixed with non-additive
fixatives were likely to be highly suspect. For this reason it
is difficult to decide how significant the observed discrepancies between investigators really are, but examples are
discussed further below.
More recently flow cytometry has been used to measure
genome size in gymnosperms (Dhillon, 1987; Marie and
Brown, 1993; Wakamiya et al., 1993; Valkonen et al., 1994;
Davies, 1996; O'Brien et al., 1996; Davies, O'Brien and
Murray, 1997). In most studies, with the exception of
Dhillon (1987), intercalating dyes, either propidium iodide
or ethidium bromide, have been used as the DNA-specific
fluorochrome. In the most extensive study, where species
were measured by flow cytometry and Feulgen microdensitometry, there was a significant correlation between measurements obtained with the two techniques (Wakamiya et
al., 1993). Comparison of measurements of the same species
from different laboratories, for example Pinus taeda L.,
were very similar, being 43.4 and 44.2 pg/2C nucleus by
Wakamiya et al. (1993) and O'Brien et al. (1996), respectively. An unexpected feature of the Wakamiya et al.
(1993) study was the deviation of measurements in Pinus
eldarica Medw. of haploid, megagametophyte and diploid,
embryo nuclei from the expected 1:2 ratio. Using both Fe
and FC they found a ratio of 1: 172 for Fe and 1: 174 for
5
FC. They suggest that this may be due to amplification or
loss of specific DNA sequences in the different tissues. In a
FC study on Larix x eurolepis Henry, Wyman et al. (1993)
also compared megagametophyte and shoot tissue but their
results, considering the standard deviations of their measurements, were within the expected 1:2 ratio. O'Brien et al.
(1996) also report variation in staining intensity in Pinus
nuclei but they attributed this to differences in the amount
of chromatin condensation affecting the binding of propidium iodide. Nuclei from actively growing tissue showed
a higher fluorescence intensity than those from less actively
growing tissue. When the latter nuclei were treated with
dilute hydrochloric acid there was a significant increase in
fluorescence intensity without nuclear degradation. In,
addition these nuclei showed less sensitivity to DNase
digestion that those from more active tissue. These results
suggest that differential chromatin condensation may be a
contributing factor to variation in fluorescence intensity
when intercalating dyes are used for flow cytometric
measurements of DNA amounts.
The use and choice of standards of known DNA amount
has also been widely discussed (Dhillon, Berlyn and
Miksche, 1977; Berlyn, Berlyn and Beck, 1986; Greilhuber,
1988; Wakamiya et al., 1993; Wyman et al., 1993). Dhillon
et al. (1977) and Berlyn et al. (1986) both recommended
chicken erythrocyte as an internal standard. They suggested
that these were better than plant standards since they
contain nuclei that are known to be at the 2C level; also
their plant standard measurements showed large and
therefore unacceptable levels of variation. Greilhuber (1988)
has criticized this suggestion citing the lack of availability of
chicken erythrocytes to most plant laboratories and the
apparent inability of the authors to correctly identify late
telophase (2C) and early prophase (4C) nuclei. Others have
criticized the use of plant standards on the basis of reported
differences in genome size within species (Wyman et al.,
1993) but appear unaware of the repeated suggestions of
Bennett and co-workers on the importance of using plant
standards with constant DNA amounts (Bennett and Smith,
1976, 1991; Bennett, Smith and Heslop-Harrison, 1982).
These standards are freely available (Bennett and Leitch,
1995). Wakamiya et al. (1993) compared DNA values for
Pinus taeda obtained from a number of studies that had
used different standards including chicken erythrocytes and
a variety of plants such as Allium cepa L., Zea mays L. and
Hordeum vulgare L. The values from the chicken erythrocyte
standard were significantly lower than those obtained with
the plant standards, and consequently, they recommend that
chicken erythrocytes not be used as the calibration standard
for Pinus as its C-value is too low. As an alternative to chick
or trout erythrocytes, Wyman et al. (1993) have suggested
the use of mouse hepatocytes as a suitable standard for
plant flow cytometry. The suggested advantage of the
mouse standard is that it contains nuclei at the 2C, 4C and
8C levels with a corresponding range of DNA values of
75-30 pg which are much closer to the range of DNA values
observed in gymnosperms.
Reassociation kinetics have been used in several studies of
genome size in gymnosperms. Rake et al. (1980) obtained
measurements from Piceaglauca and three species of Pinus
6
Murray-Nuclear DNA Amounts in Gymnosperms
used neutral formaldehyde as the fixative so that the
bleaching effects of the cellular tannins could be discounted.
In P. laricio, the differences in genome size are correlated
with striking differences in plant habit and morphology; the
smaller genomes were found in small, stunted plants raised
from seed from stunted branches on normal plants. The
measurements for Picea glauca and Pinus banksiana also
show highly significant correlations with values obtained by
direct chemical measurements of DNA amounts (Miksche,
1968). Dhir and Miksche (1974) also reported differences
between accessions of Pinus resinosa Ait. using the diphenylamine reaction to chemically measure DNA amount. It is
also relevant that the recent study of Wakamiya et al. (1993)
which combined flow cytometry and Feulgen microdensitometry with non-additive fixation in a study of 19 Pinus
species found a highly significant correlation between values
obtained with the two methods. Other examples of
intraspecific variation with Feulgen microdensitometry, but
with non-additive fixation, include Pinus caribaeaMorelet
(Berlyn et al., 1987), Pinus resinosa (Dhir and Miksche,
1974) and Picea sitchensis (Miksche, 1971).
More recently, Davies et al. (1997) and Marie and Brown
(1993) have reported intraspecific differences in genome size
following the FC analysis of plants of Manoao colensoi
(Hook.) Molloy and Ginkgo biloba, respectively. Both these
species are strictly dioecious and Davies et al. (1997) found
that males and females had small differences in DNA
amount superimposed on differences between plants from
the North and South Islands of New Zealand. They also
consider, in Manoao, that the North and South Island
plants may possibly be distinct taxa. Marie and Brown
(1993) make no comment on the sex of their plants or on
the differences that they found, but Ginkgo, along with
several other gymnosperms, is reported to have sex chromosomes (Lee, 1954; Pollock, 1957; Hizume, Shiraishi and
Tanaka, 1988). Davies et al. (1997) found no obvious
differences in the morphology or staining pattern of the
chromosome complements of male and female Manoao
colensoi.
In contrast to these reports of intraspecific variation,
Teoh and Rees (1976) found no variation between 26
provenances of Picea glauca, three of Picea engelmanii
Parry ex Engelmann and nine of Pinus contorta Dougl. A
number of more recent studies have also found no evidence
of intraspecific variation, even when measurements have
been made using different methods in different laboratories.
IS THERE INTRASPECIFIC VARIATION IN
One such example is Pinus taeda. Ohri and Khoshoo
GENOME SIZE IN GYMNOSPERMS?
(1986) investigated plants from Alabama west to Texas, and
As mentioned above in the discussion of methods of DNA Wakamiya et al. (1993) plants from Texas and North
measurement in gymnosperms, many of the claims of Carolina; in neither case was any intraspecific variation
intraspecific variation in genome size obtained by Fe have observed. O'Brien et al. (1996) measured genome size in a P.
been discounted. Much of this initial work followed from taeda plant growing in New Zealand and obtained a very
the observation of significant variation in nuclear volume similar estimate to that of Wakamiya et al. (1993). Ohri and
between different provenances of Pinus sylvestris L., P. Khoshoo (1986) also found no variation between 20
banksiana Lamb., Picea glauca and P. sitchensis (Bong.) provenances of Pinus roxburgii Sarg. or between two
Carr by Mergen and Thielges (1967). However, it is provenances of P. wallichiana A. B. Jacks from vastly
important to point out that examples of intraspecific different ecological areas. Clearly, more detailed studies
variation in Picea glauca, Pinus banksiana and P. laricio with both Fe and FC of a wide range of species are needed
Poir. have still been reported when formaldehyde fixation to obtain a clearer idea of the extent of intraspecific
has been used (Miksche, 1968; Lin et al., 1988). Both studies variation in gymnosperms.
by both RK and Fe but found significant differences
between the two groups of measurements. In all cases the
estimates of number of base pairs calculated from the
reassociation kinetics are much lower that those from
recalculations of their and other reports of genome size
using other methods of estimation. In Cycas revoluta
Thunb., Kurdi Haidar et al. (1983) report a DNA value
of 20-2x109bp which is approx. 20% less that the
calculated value of 250 x 109 bp from the 2C value in pg
provided by Ohri and Khoshoo (1986). Even greater
discrepancies are seen in the measurements of Seo, Lee and
Kim (1979). Where comparisons can be made with other
measurements, their estimates from RK are one or two
orders of magnitude smaller than those obtained by other
techniques. A large discrepancy was also found by Kreibel
(1985) for Pinus strobus L. Both Rake et al. (1980) and
Kreibel (1985) comment that the large gymnosperm genome
is not well described by partition into three, four or five
major kinetic components and consequently estimates of
genome size will be significantly distorted. These studies
report that the unique sequences make up 20-30 % of the
genome, which appears unrealistic given the large genome
sizes of gymnosperms, but Kriebel (1985) does stress that
these may be ancient partially diverged repeats and that
probably only 0%
of the Pinus strobus genome is expressed as mRNAs. The DNA values derived from RK
have been included in the Appendix but have not been
used to show the range of genome sizes by families
in Fig. 1.
The initial demonstration of variation between species
and genera (Miksche, 1967) used a combination of two
methods, chemical extraction and Feulgen cytometry, and
found a clear correlation between values obtained by the
two techniques. Miksche (1967) also demonstrated a
significant relationship between nuclear volume and DNA
amount. Based on observations such as these, Price, Sparrow
and Nauman (1973) used nuclear volumes to estimate
genome size in 236 gymnosperms, reporting a 15-fold
range. However, big discrepancies between values obtained
from nuclear volume measurements and other techniques
suggest that these estimates are only approximate at
best and should be used with caution (Hesemann, 1980;
Dhillon, 1987).
Murray-NuclearDNA Amounts in Gymnosperms
THE ADAPTIVE SIGNIFICANCE OF
GENOME SIZE VARIATION IN
GYMNOSPERMS
Only a few studies on genome size in gymnosperms have
commented on the possible adaptive significance of the
variation in this key parameter. Unlike angiosperms,
gymnosperms do not show a wide range of life forms and
are characteristically trees with long minimum generation
times. Also, in the studies on genome size, few measurements
have been made of characters that may show nucleotypic
variation. However, within some families such as the
Podocarpaceae in New Zealand, there are several prostrate
shrubs such as Podocarpusnivalis Hook. and Lepidothamnus
laxifolius (Hook. f.) Quinn. A comparison of the genome
sizes of the 20 endemic New Zealand gymnosperms shows
that mean genome size (2C) is greatest in trees (26 1+ 9.7),
smallest in shrubs (163+3±3) and intermediate in small
trees (210 + 6.7), but that there is a large range of sizes in
each category. Thus, it appears that in this sample with
overlapping genome sizes between categories, the growth
form of the species is not necessarily related to genome size
and the Prumnopitys Philippi species, which are forest trees
reaching 30 m in height, have smaller DNA amounts than
the shrubby, prostate Podocarpusnivalis.
It is equally difficult to find significant correlations
between gymnosperm genome size and cellular parameters.
Independent measurements of pollen volumes (Ueno, 1957,
1958, 1959, 1960) for 13 gymnosperms where DNA
measurements also exist show no correlation between these
two characters. This is perhaps not surprising since
gymnosperm pollen, unlike that of angiosperms, is much
more variable in structure and may contain as many as 40
prothallial cells in genera such as AraucariaJussieu (Sporne,
1965). There is also no clear correlation between genome
size and tracheid volume (Patel, 1967a, b) for a sample,
which includes both forest trees and prostrate shrubs, of 14
New Zealand gymnosperms. These do, however, belong to
different families, which may confound the problem. It is
interesting that Wakamiya et al. (1996) could also find no
significant correlation between genome size and conductive
cell radius in 21-month-old seedlings of six Pinus species,
yet significant correlations were observed in 17-month-old
ones (Wakamiya et al., 1993). Wakamiya et al. (1996) also
found that Pinus species with large genomes had a smaller
lumen radius and thicker cell wall in their conducting tissue
than those with smaller genomes. They proposed that species
with large genomes have less mechanical stress on the walls
of conducting cells, which is caused by increased internal
pressure, when plants are suffering water stress. Wakamiya
et al. (1993) found a number of correlations between DNA
content, growth related parameters and climatic factors in
North American Pinus species. For example, they observed
a positive correlation between minimum seed-bearing age
and genome size, but negative correlations between aridity
of the environment and DNA amount. Ohri and Khoshoo
(1986) also observed these sorts of relationships in an
equally large sample of Pinus species that included both
European and Asian species. The differences in tracheid
wall width associated with larger genome size (Wakamiya et
7
al., 1996) may partly explain the possible adaptation of
pines with large genomes to more arid conditions.
A correlation between seed weight and genome size was
also observed by Wakamiya et al. (1993) in North American
Pinusspecies. Donoghue and Scheiner (1992), in a discussion
on the evolution of endosperm, suggest that double
fertilization followed by the evolution of endosperm is a
major difference between the Gnetales and angiosperms on
one hand and the remainder of the gymnosperms on the
other. They suggest that without endosperm it is more
efficient for large seeds, which are characteristic of most
gymnosperms, to have large cells as these are more efficient
as storage tissues. Since larger genome sizes equate with
larger cells this may provide a selective force for maintaining
large genome sizes in the gymnosperms; in the angiosperms
a triploid endosperm results in a tissue-specific increase in
genome size which might allow the evolution of smaller
genomes. Gymnosperms are characteristically forest plants
growing in communities where a premium can exist for the
provision of large food reserves to the developing seedling.
DNA measurements on more gnetophytes and a comparative study of genome and seed size in the cycads would
be interesting, since members of this latter group have both
a very wide range of seed and genome sizes.
Where intraspecific variation has been reported, this may
or may not be correlated with the latitudinal distribution of
the seed provenances. For example, in Picea glauca, P.
sitchensis, Pinus banksiana and P. sylvestris Mergen and
Thielges (1967) found that nuclear volumes were correlated
with latitude; larger volumes were found in the more
northern latitude. However, Miksche (1968) did not find
any correlation between latitude and the DNA variation
observed in Picea glaucaand Pinus banksiana. On the other
hand there did appear to be a correlation between these
characters in Picea sitchensis with the northern populations
having larger DNA amounts than the southern ones
(Miksche, 1971). In P. laricio, Lin et al. (1988) found two
different genome sizes that were correlated with different
growth patterns. Dwarf plants grown from seed of compact,
mutant branches had very significantly smaller genomes
than normal control plants. Thus, the gymnosperms offer
few insights into the possible nucleotypic effects of genome
size due to the very limited number of investigations to date.
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Murray-NuclearDNA Amounts in Gymnosperms
9
APPENDIX
Nuclear DNA amount, chromosome number, family and ploidy level in gymnosperm species
Species
Entry
number'
Voucher
Family
2n Ploidy
IC
DNA (D)
2C
4C
2C bp
xlO
Oriinal Present Standard Method'
rer
amount species'
1
Abies alba Mill.
no
Pinacece
24
2
16.6
33.1
66.2
32.4
18
O
E
Fe
2
Abies balsamea (L.) Miller
no
Pinaceae
24 2
13.2
26.3
52.6 25.8
11
O
F
Fe
3
Abies sibiricaLedeb.
no
Pinaceae
24
2
15.8
31.6
63.2 31.0
15
O
B*
Fe
4
Agathis australisSalisb.
yes
Armucaiaceae
26
2
15.8
31.6
63.2
31.0
3
O
A&C
FC:PI
5
Araucariacooki R. Brown ex.
no
Araucariaceae
26
2
9.6
19.1
38.2
18.7
15
O
B*
Fe
Lind.
6
Araucwria cmninghamii D. Don
no
Araucariaceae
26 2
10.9
21.8
43.6
21.4
15
O
B*
Fe
7
Araucmria robusta(C. Moore)
no
Araucariaceae
26 2
10.8
21.6
43.2
21.2
15
O
B*
Fe
no
Cupressaceae
22 2
11.4
22.8
45.6
22.3
15
O
B*
Fe
B*
Fe
F.M. Bailey
8
Biotaorientalis Endl.
9
CaUitrisglaucaR. Br.
no
Cupressaceae
22
2
8.3
16.5
33.0
16.2
15
O
10
CallitrisrhomboideaR. Br. ex
no
Cupressaceae
22
2
11.2
22.3
44.6 21.9
15
O
B*
Fe
no
Cupressaceae
22 2
10.4
20.8
41.6 20.4
15
O
B*
Fe
no
Cupressaceae
22 2
15.1
30.1
60.2
29.5
15
O
B*
Fe
Chamaecyparisobtusa (Sieb. & no
Cupressaceae
22
2
13.7
27.4
54.8 26.9
15
O
B*
Fe
Cupressaceae
22
2
11.1
22.1
44.2 21.7
15
O
B*
Fe
no
Taxodiaceae
22
2
13.6
27.1
54.2 26.6
15
O
B*
Fe
L.C. Rich.
11
CalitrisverrucosaA. Cunn. ex
Endl. Muller
12
Chamaecyparislawsonicma (A.
Muffrr.) Parl.
13
Zucc.) Endl.
14
Chamaecyparispisifera (Sieb. & no
Zucc.) Endl.
15
Cuwwnghamialanceolata
(Lamb.) Hook. f.
16
CupressusarizonicaGreene
no
Cupressaceae
22 2
11.8
23.6
47.2 23.1
15
O
B*
Fe
17
Cupressusglabravar. conica
no
Cupressaceae
22 2
11.5
23.0
46.0
22.5
15
O
B*
Fe
Sudw.
18
Cupressus guadalupensisSarg.
no
Cupressaceae
22
2
11.9
23.7
47.4 23.2
15
O
B*
Fe
19
CupressusmacrocarpaHartw.
no
Cupressaceae
22
2
14.2
28.4
56.8 27.8
15
O
B*
Fe
20
Cupressussempervirens L.
no
Cupressaceae
22
2
11.4
22.7
45.4 22.2
15
O
B*
Fe
21
Cupressus sempervirens var.
no
Cupressaceae
22
2
11.6
23.1
46.2 22.6
15
O
B*
Fe
strictaAiton
22
Cycas circinalisL.
no
Cycadaceae
22 2
14.8
29.5
59.0 28.9
15
O
B*
Fe
23a
Cycas revoluta Thunb.
no
Cycadaceae
22
2
12.8
25.5
51.0
25.0
15
O
B*
Fe
23b
Cycas revolutaThunb.
no
Cycadaceae
22
2
10.3
20.6
41.2 20.2
10
O
none
RK
24
Dacrycarpus dacryioides(A.
yes
Podocarpaceae
20 2
18.1
36.2
72.4
35.5
3
O
A&C
FC:PI
Dacrydiumcupressinum Lambert yes
Podocarpaceae
20
18.0
36.0
72.0 35.3
3
O
A &C
FC:PI
Rich.) Laubenf. d
25
2
10
Murray-Nuclear DNA Amounts in Gymnosperms
Entry
number'
Species
Voucher
Family
2n Ploidy
DNA (p)
2n o 2
4C
2C b
xlO'
Oriinal Present Standard Method'
rer
amount species'
26
Encephalatos villosus Lem.
no
Zamiaceae
18 2
21.1
42.2
84.4
41.4
15
O
B*
Fe
27
Ephedra tweediana C.A. Mey
no
Ephedraceae
14
2
8.9
17.8
35.6
17.4
15
O
B*
Fe
28a
Ginkgo biloba L.d
no
Ginkgoaceae
24
2
10.0
19.9
39.8
19.5
15
O
B*
Fe
28b
Ginkgo bilobaL.d
yes
Ginkgoaceae
24
2
9.9
19.8
39.6
19.4
9
O
B*
Fe
28c
Ginkgo blloba L.d
no
Ginkgoaceae
24
2
2.3
4.6
9.2
4.5
19
O
none
RK
28d
Ginkgo biloba L.
no
Ginkgoaceae
24
2
9.8
19.5
39.0
19.1
12
O
D
FC:PI,EB
29
Gnetum ula Brongn
no
Gnetaceae
22
2
2.3
4.5
9.0
4.4
15
O
B*
Fe
30
Halocarpusbidwillii (Kirk)
yes
Podocarpaceae
18
2
7.9
15.7
31.4
15.4
3
O
A&C
FC:PI
yes
Podocarpaceae
24
2
11.7
23.4
46.8
22.9
3
O
A&C
FC:PI
Quinn
31
Halocarpusbiformis (Hook.)
Quinn
32
Halocarpuskirkii(Parl.) Quinn
yes
Podocarpaceae
22
2
7.9
15.7
31.4
15.4
3
O
A&C
FC:PI
33
Juniperus virginianaL.
no
Cupressaceae
22
2
9.7
19.3
38.6
18.9
7
O
F
Fe
34
Lagarostrobosfranklinii(Hook. yes
Podocarpaceae
30
2
15.2
30.4
60.8 29.8
4
O
A&C
FC:PI
f.) Quinn
35a
Larix decidua Miller
yes
Pinaceae
24
2
11.5
22.9
45.8 22.4
8
O
B*
Fe
35b
Larix decidua Miller
no
Pinaceae
24
2
9.9
19.7
39.4
7
O
F
Fe
36
Larix gmelini (Rupr.) Kuz.
no
Pinaceae
24
2
14.0
28.0
56.0 27.4
15
O
B*
Fe
37
Larix aricina(Duroi) K. Koch
no
Pinaceae
24
2
9.5
19.0
38.0
18.6
7
O
F
Fe
38
Larixsibirica Ledeb.
no
Pinaceae
24 2
12.3
24.6
49.2
24.1
15
O
B*
Fe
39a
Larixx eurolepisHenry
no
Pinaceae
24 2
15.4
30.8
61.6 30.2
15
O
B*
Fe
39b
Larixx eurolepis Henry
no
Pinaceae
24 2
16.1
32.2
64.4 31.6
23
O
G
FC:PI
39c
Larixx eurolepis Henry
no
Pinaceae
24 2
17.5
35.0
70.0 34.3
23
O
H
FC:PI
40
Lepidothamnus intermedius
yes
Podocarpaceae
30 2
6.6
13.1
26.2
12.8
3
O
A&C
FC:PI
Lepidothamnus laxifolius (Hook. yes
Podocarpaceae
30
6.7
13.4
26.8
13.1
3
O
A&C
FC:PI
19.3
(Kirk) Quinn
41
2
f.) Quinn
42
Libocedrus bidwillii Hook. f.
yes
Cupressaceae
22 2
19.5
39.0
78.0 38.2
3
O
A &C
FC:PI
43
Libocedrus plumosa (Don)
yes
Cupressaceae
22 2
20.0
40.0
80.0 39.2
3
O
A&C
FC:PI
yes
Podocarpaceae
20 2
13.8
27.6
55.2 27.0
3
O
A&C
FC:PI
no
Taxodiaceae
22 2
6.5
13.0
26.0
12.7
15
O
B*
Fe
Sargent
44
Manoao colensoi (Hook.)
Molloy
45
d
Metasequoiaglyptostroboides
Hu &W.C. Cheng
46
PhyllocladusalpinusHook. f.
yes
Phyllocladacese
18
2
11.1
22.1
44.2
21.7
3
O
A&C
FC:PI
47
Phyllocadusglaucus Carr.
yes
Phyllocladaceae
18
2
11.4
22.8
45.6 22.3
3
O
A&C
FC:PI
48
Phyllocladus trichomanoides
yes
Phyllocladaceae
18
2
10.0
19.9
39.8 19.5
3
O
A&C
FC:PI
Don
Murray-Nuclear DNA Amounts in Gymnosperms
Entry
number'
Species
Voucher
Family
2n Plddy
IC
DNA (P)
2C
4C
2C
11
Ori'nd! Present Standard Method'
xl~wr
amount species
49a
Picea abies(L) Karsten
no
Pinaceae
24 2
14.8
29.6
59.2
29.0
7
O
F
Fe
49b
Piceaabies (L.) Karsten
no
Pinaceae
24 2
14.8
29.6
59.2 29.0
7
O
?F
FC:M
50
Picea engebnmi'i Parry ex
no
Pinaceae
24
2
19.5
38.9
77.8 38.1
20
O
B*
Fe
2
20.2
40.4
80.8 39.6
15
O
B*
Fe
7
O
F
Fe
Engelmann
51a
Piceaglauca(Moench) Voss
no
Pinaceae
24
51b
Piceagaua (Moench) Voss
no
Pinaceae
24 2
8.5
17.0
34.0
51c
Piceaglauca(Moench) Voss
no
Pinaceae
24 2
19.5
38.9
77.8 38.1
20
O
B*
Fe
51d
Picea gauca(Moench) Voss
no
Pinaceae
24 2
4.5
9.0
18.0 8.8
16
O
none
RK
51e
Piceaglauca(Moench) Voss
no
Pinaceae
24 2
9.7
19.3
38.6
18.9
16
O
F
Fe
52a
Picmea
mia
(Mill.) Britt.
no
Pinaceac
24 2
15.8
31.6
63.2 31.0
15
O
B*
Fe
52b
Picea maiaa (Mill.) Britt
no
Pinaceae
24 2
11.1
22.2
44.4 21.8
7
O
F
Fe
53
PiceaorimalsL.
no
Pinaceae
24 2
18.6
37.2
74.4 36.5
15
O
B*
Fe
54
PiceapungenEgelm.
no
Pinaceae
24 2
20.0
40.0
80.0
15
O
B*
Fe
55
Piceapungens Englm.f. glauca
yes
Pinaceae
24
2
18.2
36.3
72.6 35.6
9
O
B*
Fe
56a
Pinus aenuata Lemmon
no
Pinaceae
24 2
25.1
50.1
100.2 49.1
22
O
C
Fe
56b
Pinus atenuataLemmon
no
Pinaceae
24 2
22.1
44.2
88.4 43.3
22
O
C
FC:PI
57
Pinusaurescens (Sic.)
no
Pinaceae
24 2
0.6
1.1
2.2
19
O
none
RK
58a
Pinus bankuiana Lamb d
no
Pinaceae
24
2
17.2
34.4
68.8 33.7
15
O
B*
Fe
58b
Pinus banksianaLamb d
no
Pinaceae
24 2
14.9
29.8
59.6 29.2
16
O
F
Fe
58c
Pinus banksi
Lamb d
no
Pinacre
24
2
9.0
18.0
36.0
17.6
16
O
none
RK
58d
Pfnus bankianaLamb d
no
Pnaceae
24
2
14.1
28.1
56.2 27.5
24
O
I
FC:PI
59a
Pinus carbaea Morelet
no
Pinaceae
24
2
19.6
39.1
78.2 38.3
15
O
B*
Fe
59b
PinuscaribaeaMorelet
no
Pinacee
24
2
22.8
45.6
91.2 44.7
12
O
D
FC:PI,EB
60
Pinus caribaeavar. bahamens
no
Pinace
24
2
12.6
25.2
50.4 24.7
1
O
F
Fe
no
Pinaceae
24
2
5.8
11.5
23.0
11.3
1
O
F
Fe
no
Pinaceae
24
2
10.6
21.2
42.3 20.7
1
O
F
Fe
Pinaceae
24
2
24.2
483
96.6 47.3
8
O
B*
Fe
24
2
22.1
44.2
88.4 43.3
22
O
C
Fe
24
2
20.0
39.9
79.8 39.1
22
O
C
FC:PI
16.7
39.2
1.1
Barr. &Golf.
61
Pnuscaribaeava. carbaea
Barr. &Golf.
62
Pinus caribaea var. hondurensis
Barr. &Golf.
63
Pinus cembra L.
no
64a
Pins clausa (Chapm.) Vasey
no
64b
Pinus clausa (Chapm.) Vasey
no
65a
Pinuscontorta Dougl.
no
24
2
18.9
37.8
75.6 37.0
14
O
A&C
FC:PI
65b
Pinus contortaDougl.
no
24 2
20.2
40.3
80.6 39.5
20
O
B*
Fe
65c
Pinus contorta Dougl.
no
24
2
12.2
24.4
48.8 23.9
7
O
F
Fe
66
Pinus contortavar. atifoliaS.
no
24 2
17.8
35.6
71.2 34.9
15
O
B*
Fe
24
28.7
57.4
114.8 56.3
22
O
C
Fe
Pinaceae
Watson
67
Aruts coultenD.Don
no
Pinaceae
2
Murray-NuclearDNA Amounts in Gymnosperms
12
Entry
number'
Species
Voucher
Family
2n Ploidy
DNA (gi)_
-. 4C
2 I ~2C
2C bo
x
Oriinal Present Standard Method'
r10
e
amount species
67b
Pinus coulter D. Don
no
Pinaceae
24 2
28.4
56.7
113.4 55.6
22
0
C
FC:PI
67c
Pinus coulter D. Don
no
Pinaceae
24 2
8.5
17.0
34.0
16.7
2
O
F
Fe
68
Pinus densiflora Sieb. &Zucc.
no
Pinaceae
24 2
21.5
43.0
86.0
42.1
15
O
B*
Fe
69
Plnusdivaricata(Ait.) Dum.-
no
Pinaceae
24
2
17.3
34.5
69.0
33.8
15
O
B*
Fe
Cours.
70a
Pinus echinataMill.
no
Pinaceae
24
2
22.8
45.5
91.0
44.6
22
C
Fe
70b
Pinus echinataMill.
no
Pinaceae
24
2
21.8
43.5
87.0 42.6
22
C
FC:PI
71
Pinus excelsa Hook.
no
Pinaceae
24
2
24.1
48.2
96.4 47.2
15
B*
Fe
72a
Pinus edaricaMedw.
no
Pinaceae
24
2
31.4
62.7
125.4 61.4
22
C
Fe
72b
Pinus eldaricaMedw.
no
Pinaceae
24
2
27.8
55.5
111.0 54.4
22
C
FC:PI
73a
Pinus elliottiiEngelm.
no
Pinaceae
24 2
23.3
46.6
93.2 45.7
22
C
Fe
73b
Pinus elliotiiEngelm.
no
Pinaceae
24 2
22.4
44.7
89.4 43.8
22
C
FC:PI
73c
Pinus elliottii Engelm.
no
Pinaceae
24
2
17.7
35.3
70.6 34.6
15
B*
Fe
74a
Pinus flexdis James
no
Pinaceae
24 2
29.2
58.4
116.8 57.2
22
C
Fe
74b
Pinusfiexilis James
no
Pinaceae
24 2
29.6
59.2
118.4 58.0
22
C
FC:PI
75
Pinus gerardianaWall.
no
Pinaceae
24 2
28.7
57.4
114.8 56.3
15
B*
Fe
76a
Pinusjeffreyi Grev. &Balf.
no
Pinaceae
24 2
27.7
55.4
110.8 54.3
22
C
Fe
76b
Pinusjeffreyi Grev. &Balf.
no
Pinaceae
24 2
24.9
49.8
99.6
48.8
22
C
FC:PI
77
Pinuskoraiensis Sieb &Zucc.
no
Pinaceae
24
0.1
0.2
0.4
0.2
19
none
RK
78a
Pinus lamberianaDougl.
no
Pinaceae
24 2
29.6
59.1
118.2 57.9
22
C
Fe
FC:PI
2
78b
Pinus ambertianaDougl.
no
Pinaceae
24 2
31.8
63.5
127.0 62.2
22
C
78c
Pinus lambertianaDougl.
no
Pinaceae
24 2
17.4
34.7
69.4
6
F
Fe
78d
Pinus lambertanaDougl.
no
Pinaceae
24 2
43.9
87.7
175.4 85.9
16
F
Fe
78e
Pinus amberihna
Dougl.
no
Pinaceae
24 2
10.6
21.2
42.4 20.8
16
none
RK
78f
Pinus lambertnaDougL
no
Pinaceae
24 2
17.6
35.2
70.4
7
F
Fe
79a
Pinus laricioPoir.
no
Pinaceae
24
2
25.4
50.7
101.4 49.7
11
B*
Fe
79b
Pinus laicioPoir. '
no
Pinaceae
24
2
15.0
30.0
60.0 29.4
11
B*
Fe
80
Pinus maritina Lamb.
no
Pinaceae
24
2
24.1
48.1
96.2 47.1
15
B*
Fe
81a
Pinus monophylla Torrey
no
Pinaceae
24
2
30.2
60.4
120.8 59.2
22
C
Fe
81b
Pinus monophylla Torrey
no
Pinaceae
24
2
27.4
54.7
109.4 53.6
22
C
FC:PI
A&C
FC:PI
34.0
34.5
82
Pinus montezwae Lamb.
no
Pinaceae
24 2
26.7
53.4
106.8 52.3
14
83a
Pinus monticola Dougl.
no
Pinacese
24 2
27.2
54.4
108.8 53.3
22
C
Fe
83b
Pinus monticola Dougl.
no
Pinaceae
24 2
29.3
58.6
117.2 57.4
22
C
FC:PI
84a
Pinus mugo Turra
no
Pinaceae
24 2
20.2
40.3
80.6 39.5
8
B*
Fe
84b
Pinus mugo Turra
no
Pinaceae
24 2
20.1
40.2
80.4 39.4
15
B*
Fe
85a
PinuspalustrisMill.
no
Pinaceae
24 2
24.1
48.1
96.2 47.1
22
C
Fe
85b
PinuspalustrisMill.
no
Pinaceae
24 2
23.1
46.1
92.2
45.2
22
C
FC:PI
86
PitwpaulaSchlecht. &Char.
no
Pinaceae
24 2
18.5
36.9
73.8
36.2
15
B*
Fe
O
Murray-Nuclear DNA Amounts in Gymnosperms
Entry
number'
Species
Voucher
Family
2n Ploidy
DNA ()_
IC0 2C 4C
2C b
xl0'
13
Original Present Standardb Method'
ret
amount species
87
PinuspinasterAit.
no
Pinaceae
24
2
24.4
48.7
97.4 47.7
15
O
B*
Fe
88a
Pinusponderosa Dougl. ex
no
Pinaceae
24
2
24.2
48.4
96.8 47.4
14
O
A&C
FC:PI
no
Pinaceae
24
2
19.9
39.7
79.4 38.9
15
0
B*
Fe
Laws.
88b
PinusponderosaDougl. ex
Laws.
89
L indl.
Pinuspseudostrobus
no
Pinaceae
24
2
21.4
42.7
85.4 41.8
14
O
A&C
FC:PI
90a
Pinusradita D. Don
no
Pinaceae
24 2
22.0
44.0
88.0 43.1
14
O
A&C
FC:PI
90b
PinusradiataD. Don
no
Pinaceae
24 2
24.3
48.6
97.2
47.6
22
O
C
Fe
90c
Pinusradi'aD. Don
no
Pinaceae
24 2
23.1
46.2
92.4 45.3
22
O
C
FC:PI
90d
Pimnus radiataD. Don
no
Pinaceae
24 2
11.0
22.0
44.0 21.6
6
O
F
Fe
90e
Pnu radiata D. Don
no
Pinaceae
24 2
11.2
22.3
44.6
21.9
7
O
F
Fe
90f
PinusradiataD. Don
no
Pinaceae
25 2n+1
23.1
46.1
92.2
45.2
14
O
A&C
FC:PI
91a
Pinusresinosa Ait.
no
Pinaceae
24 2
23.4
46.7
93.4 45.8
15
O
B*
Fe
91b
Pinus resinosa Ait.
no
Pinaceae
24 2
21.6
43.1
86.2
42.2
16
O
F
Fe
91c
Pinusresinosa Ait.
no
Pinaceae
24 2
4.2
8.4
16.8
8.2
16
O
none
RK
92a
PinusrigidaMill.
no
Pinaceae
24 2
20.8
41.6
83.2 40.8
15
O
B*
Fe
92b
Pinus rigidaMill.
no
Pinaceae
242
7.8
15.5
31.0
15.2
5
O
F
Fe
92c
PinusrigidaMill.
no
Pinaceae
24
8.8
17.6
35.2
17.2
6
O
F
Fe
92d
PinusrigidaMill.
no
Pinaceae
242
8.9
17.8
35.6
17.4
7
O
F
Fe
92e
Pinus rigidaMill.
no
Pinaceae
242
0.1
0.2
0.4
0.2
19
O
none
RK
93
Pinusroxburghii Sarg.
no
Pinaceae
24 2
19.4
38.8
77.6 38.0
15
O
B*
Fe
94a
Pinus sabinlanaDoug.
no
Pinaceae
24 2
29.6
59.2
118.4 58.0
22
O
C
Fe
94b
Pinus sabinianaDougl.
no
Pinaceae
24 2
28.4
56.7
113.4 55.6
22
O
C
FC:PI
95a
Pinusserotina Mihx.
no
Pinaceae
24 2
21.5
43.0
86.0
42.1
22
O
C
Fe
95b
Pinus serotina Michx.
no
Pinaceae
24
2
21.0
42.0
84.0 41.2
22
O
C
FC:PI
96a
Pinus strobus L.
no
Pinaceae
24 2
25.7
51.3
102.6 50.3
14
O
A&C
FC:PI
96b
Pinus strobus L.
no
Pinaceae
24
26.7
53.3
106.6 52.2
22
O
C
Fe
96c
Pinus strobus L.
no
Pinaceae
242
29.1
58.1
116.2 56.9
22
O
C
FC:PI
19.4
38.8
77.6 38.0
7
O
F
Fe
2
2
96d
Pinus srobus L.
no
Pinaceae
24 2
96e
Pinus strobus L.
no
Pinaceae
242
19.4
38.8
77.6 38.0
7
O
?F
FC:M
97a
Pinus sylvestris L.
no
Pinaceae
242
27.8
55.6
111.2 54.5
21
O
C
FC:PI
97b
Pinus sylvestris L.
no
Pinaceae
24
2
13.8
27.6
55.2 27.0
7
O
F
Fe
98a
Pinus taedaL.
no
Pinaceae
24
2
22.1
44.2
88.4 43.3
14
O
A&C
FC:PI
98b
Pinustaeda L.
no
Pinaceae
242
23.2
46.3
92.6 45.4
22
O
C
Fe
98c
Pinus taedaL.
no
Pinaceae
24 2
21.7
43.4
86.8 42.5
22
O
C
FC:PI
98d
Pinus taedaL.
no
Pinaceae
24 2
19.0
37.9
75.8
15
O
B*
Fe
98e
Pinstaeda L.
no
Pinaceae
242
11.0
21.9
43.8 21.5
7
O
F
Fe
98f
Pinus taedaL.
no
Pinaceae
242
13.0
26.0
52.0 25.5
17
O
F
Fe
37.1
Murray-Nuclear DNA Amounts in Gymnosperms
14
Entry
number
Species
Voucher
Family
2n Ploidy
IDNA ()
2C
_C
4C
C
2C bxl Ori inal Present Standard Method'
ref
amount species'
xlO'
19
0
none
RK
88.0 43.1
15
O
B*
Fe
58.1
116.2 56.9
22
0
C
Fe
26.4
52.7
105.4 51.6
22
0
C
FC:PI
24 2
16.6
33.2
66.4 32.5
15
O
B*
Fe
Pinaceae
24 2
21.2
42.3
84.6 41.5
22
0
C
Fe
no
Pinaceae
24 2
20.4
40.7
81.4 39.9
22
0
C
FC:PI
Pinus wallichianaA.B. Jacks
no
Pinaceae
24 2
24.6
49.2
98.4 48.2
15
0
B*
Fe
103
Podocarpusacutifolius Kirk
yes
Podocarpaceae
34
8.2
16.4
32.8
3
0
A&C
FC:PI
104
Podocarpusgraci/orPilger
no
Podocarpaceae
242
11.4
22.8
45.6 22.3
15
0
B*
Fe
105
Podocarpushallii Kirk
yes
Podocarpaceae
34
8.7
17.4
34.8
17.1
3
0
A&C
FC:PI
106
Podocarpusnivalis Hook.
yes
Podocarpaceae
38 2
10.0
19.9
39.8
19.5
3
O
A&C
FC:PI
107
Podocarpus otaraG. Benn. ex
yes
Podocarpaceae
342
113
22.5
45.0 22.1
3
0
A&C
FC:PI
Prwnopitysferruginea(D. Don) yes
Podocarpaceae
36 2
7.9
15.8
31.6
15.5
3
0
A&C
FC:PI
Podocarpaceae
38
2
8.0
15.9
31.8
15.6
3
0
A&C
FC:PI
Pinaceae
24
2
19.1
38.1
76.2 37.3
14
O
A&C
FC:PI
98g
Pinustaeda L.
no
Pinaceae
24 2
0.7
1.3
2.6
99
Pinus thunbergtiParl.
no
Pinaceae
242
22.0
44.0
100a
Pinus torreyanaParry
no
Pinaceae
24 2
29.1
100b
Pius torreyanaParry
no
Pinaceae
24 2
101a
Pinus virgianaMill.
no
Pinaceae
101b
Pinus virginianaMill.
no
-101c
Pinus virgnianaMill.
102
2
2
1.3
16.1
Don
108
Laubenf.
109
Prwnnopitys taxifolia (D. Don)
yes
Laubenf.
110
Pseudotsugamenziesii (Mirabel) no
Franco
111
Taxodium mucronatwn Ten.
no
Taxodiaceae
22
2
8.8
17.5
35.0 17.2
15
O
B*
Fe
yes
Taxaceae
24
2
11.1
22.1
44.2 21.7
9
0
B*
Fe
47.2 23.1
112
Taxus baccataL.
113
Tetraclinisarticulaa(Vahl) Mast. no
Cupressaceae
22
2
11.8
23.6
15
0
B*
Fe
114a
Thuja occidentalisL.
no
Cupressaceae
22
2
11.7
23.3
46.6 22.8
15
0
B*
Fe
114b
Thuja occidentais L.
no
Cupressaceae
22
2
6.3
12.6
25.2
7
0
F
Fe
115
Thuja picataD. Don
no
Cupressaceae
22
2
12.3
24.6
49.2 24.1
15
0
B*
Fe
116
Tsuga caradensis(L.) Can.
no
Pinaceae
24
2
13.5
27.0
54.0 26.5
7
0
F
Fe
117
Zamnia angustifoliaJacquin
no
Zamiaceae
16 2
12.1
24.1
48.2 23.6
15
O
B*
Fe
Key Standard species
A
B
C
D
E
F
G
H
I
Hordeum vulgare 'Sultan'
Allium cepa 'Ailsa Craig'
Triticum aestivum 'Chinese Spring'
Petunia hybrida 'PxPc6'
Pisum sativum 'Kleine Rheinlinderin'
Gallus domestica
Mus musculus 'Balb/C' or 'B6/AF1'
Salmo trutta
Rattus rattus
DNA
amount (pg)
22'24
67-00
69-27
570
17-68
4-66
1504
550
11-40
Notes to the appendix
(a) The original references for the DNA values are listed
below by their corresponding number.
12.3
(b) Five plant and four animal standard species have been
used to determine absolute DNA amounts. Where named or
specified cultivars have not been used an asterisk follows the
appropriate letter in the table. The key gives the 4C DNA
amounts of the standard species.
(c) Where several estimates are available, the 'a' value is
the preferred estimate for that species.
(d) These species have been reported to show intraspecific
variation and the reader is advised to check the original
reference for detailed information.
(e) In Pinus laricio the intraspecific variation is discontinuous and is reported to occur in distinct types of
plant.
(f) In the method column the following abbreviations are
used: Fe, Feulgen microdensitometry; FC, flow cytometry
(with PI, propidium iodide; EB, ethidium bromide; M,
mithramycin); RK, reassociation kinetics.
Murray-Nuclear DNA Amounts in Gymnosperms
Originalreferencesfor DNA values
1. Berlyn GP, Anoruo AO, Beck RC, Cheng J. 1987. DNA content
polymorphism and tissue culture regeneration in Caribbean pine.
CanadianJournal of Botany 65: 954-961.
2. Berlyn GP, Berlyn MKB, Beck RC. 1986. A comparison of internal
standards for plant cytophotometry. Stain Technology 61: 297302.
3. Davies BJ, O'Brien IEW, Murray BG. 1997. Karyotypes, chromosome bands and genome size variation in New Zealand endemic
gymnosperms. Plant Systematics and Evolution 208: 169-185.
4. Davies BJ. 1996. Chromosome evolution and cytogenetics in the
New Zealandconifers. MSc. Thesis. University of Auckland, New
Zealand.
5. Dhillon SS, Berlyn GP, Miksche JP. 1978. Nuclear DNA content
in populations of Pinus rigida. American Journal of Botany 65,
192-196.
6. Dhillon SS. 1980. Nuclear volume, chromosome size and DNA
content relationships in three species of Pinus. Cytologia 45:
555-560.
7. Dhillon SS. 1987. DNA in tree species In: Bouga JM, Deuzan DJ,
eds. Cell and tissue culture in forestry. Dordrecht: Martinus Nijhoff,
298-313.
8. Greilhuber J. 1986. Severely distorted Feulgen-DNA amounts in
Pinus (Coniferophytina) after non-additive fixation as a result of
meristematic self-tanning with vacuole contents. CanadianJournal
of Genetics and Cytology 28: 409-415.
9. Greilhuber J. 1988. Critical reassessment of DNA content in plants
In: Brandham PE, ed. Kew Chromosome Conference III. London:
HMSO, 39-50.
10. Kurdi Haidar B, Shalhoub V, Dib Haj S, Deeb S. 1983. DNA
sequence organization in the genome of Cycas revoluta. Chromosoma 88: 319-327.
11. Lin YQ, Bitonti MB, Ciolli M, Innocenti AM. 1988. Somatic mutagenesis in Pinus laricio. A cytophotometric analysis of DNA and
histone content in 2C meristematic nuclei. Caryologia 41: 137-142.
12. Marie D, Brown SC. 1993. A cytometric exercise in plant DNA
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
15
histograms, with 2C values for 70 species. Biology of the Cell 78:
41-51.
Mellerowicz EJ, Riding RT, Little CHA. 1989. Genomic variability
in the vascular cambium of Abies balsamea. Canadian Journal of
Botany 67: 990-996.
O'Brien IEW, Smith DR, Gardner RC, Murray BG. 1996. Flow
cytometric determination of genome size in Pinus. Plant Science
115: 91-99.
Ohri D, Khoshoo TN. 1986. Genome size in gymnosperms. Plant
Systematics and Evolution 153: 119-132.
Rake AV, Miksche JP, Hall RB, Hansen KM. 1980. DNA reassociation kinetics for four conifers. CanadianJournal of Genetics and
Cytology 22: 69-79.
Renfroe MH, Berlyn GP. 1984. Stability of nuclear DNA content
during adventitious shoot formation in Pinus taeda L tissue
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Roth R, Ebert I, Schmidt J. 1997. Trisomy associated with loss of
maturation capacity in a long-term embryogenic culture of Abies
alba. Theoretical and Applied Genetic 95: 353-358.
Seo JS, Lee KY, Kim TU. 1979. Genome analysis of several
gymnosperm pollens. Seoul Journal of Medicine 20: 70-76.
Teoh SB, Rees H. 1976. Nuclear DNA amounts in populations of
Picea and Pinus species. Heredity 36: 123-137.
Valkonen JPT, Nygren M, Yl6nen A, Mannonen Y. 1994. Nuclear
DNA content of Pinus sylvestris (L.) as determined by laser flow
cytometry. Genetica 92: 203-207.
Wakamiya I, Newton RJ, Johnston JS, Price HJ. 1993. Genome
size and environmental factors in the genus Pinus. American Journal
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Wyman J, Guertin F, Mansour S, Fournier M, Laliberte S. 1993. Use
of mouse hepatocytes for the flow cytometric determination of
DNA levels of nuclei extracted from fresh tissue of hybrid larch
Larix x eurolepis Henry. Cytometry 14: 217-222.
Wyman J, Laliberte S, Tremblay M-T. 1997. Nuclear DNA content
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banksiana, Pinaceae). American Journalof Botany 84: 1351-1361.