First Nuclear DNA Amounts in more than 300 Angiosperms

Author(s): B. J. M. ZONNEVELD, I. J. LEITCH and M. D. BENNETT

Source: Annals of Botany , August 2005, Vol. 96, No. 2 (August 2005), pp. 229-244
Published by: Oxford University Press

Stable URL: https://www.jstor.org/stable/42795994

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Annals of Botany 96: 229-244, 2005
doi : 1 0. 1 093/aob/mci 1 70, available online at www.aob.oupjournals.org

First Nuclear DNA Amounts in more than 300 Angiosperms

B. J. M. ZONNEVELD1, 1. J. LEITCH2andM. D. BENNETT2*
1 Institute of Molecular Plant Sciences, Leiden University, Leiden, The Netherlands and 2Jodrell Laboratory,
Royal Botanic Gardens , Kexv, Richmond , Surrey TW9 3AB , UK

Received: 3 February 2005 Returned for revision: 30 March 2005 Accepted: 10 April 2005 Published electronically: 19 May 2005

• Background and Aims Genome size (DNA C-value) data are key biodiversity characters of fundamental signi-
ficance used in a wide variety of biological fields. Since 1976, Bennett and colleagues have made scattered published
and unpublished genome size data more widely accessible by assembling them into user-friendly compilations.
Initially these were published as hard copy lists, but since 1997 they have also been made available electronically
(see the Plant DNA C-values database www.kew.org/cval/homepage.html). Nevertheless, at the Second Plant
Genome Size Meeting in 2003, Bennett noted that as many as 1000 DNA C-value estimates were still unpublished
and hence unavailable. Scientists were strongly encouraged to communicate such unpublished data. The present
work combines the databasing experience of the Kew-based authors with the unpublished C-values produced by
Zonneveld to make a large body of valuable genome size data available to the scientific community.
• Methods C-values for angiosperm species, selected primarily for their horticultural interest, were estimated by flow
cytometry using the fluorochrome propidium iodide. The data were compiled into a table whose form is similar to
previously published lists of DNA amounts by Bennett and colleagues.
• Key Results and Conclusions The present work contains C-values for 41 1 taxa including first values for 308 species
not listed previously by Bennett and colleagues. Based on a recent estimate of the global published output of
angiosperm DNA C-value data (i.e. 200 first C-value estimates per annum) the present work equals 1-5 years of
average global published output; and constitutes over 12 % of the latest 5-year global target set by the Second Plant
Genome Size Workshop (see www.kew.org/cval/workshopreport.html). Hopefully, the present example will
encourage others to unveil further valuable data which otherwise may lie forever unpublished and unavailable
for comparative analyses.

Key words: Genome size, C-value, plant DNA C-values database, Genome Size Initiative (GSI), Plant Genome Size
workshop and Discussion Meeting, angiosperm, monocots.

INTRODUCTION (e.g. Knight et al ., 2005; Leitch et al, , 2005; Price e

2005) and are the main source of information on
The DNA amount in the unreplicated gametic nuclear chro-
genome sizes cited in the literature. Published lists of
mosome complement (1 C-value) is highly characteristic
sperm DNA amounts have been cited over 1500 t
for taxa and varies over 1000-fold between plant
whilst species.
the electronic database has received over 60 000 hits
Species' DNA amounts [C-value and genome
since 2001.size (lCx)]
are key diversity characters of fundamental significance
Bennett et al. (2000) and Bennett and Leitch (2005a)
with many important uses (Bennett and Leitch, 2005a).
noted that C-values for probably a surprisingly large number
However, it is often difficult to know if a genome size
of plant species had been estimated but, for various reasons,
measurement exists for a taxon, or where to find it, as such
not published and hence were unavailable. Indeed, this was
values are widely scattered in the literature or unpublished.
suspected to apply to hundreds of species overall whose
Consequently, since 1976, lists of DNA amounts in angio-
genome sizes together constitute an untapped pool of
sperm species, compiled for reference purposes, have been
information of considerable potential value. Scientists
published (e.g. Bennett and Smith, 1976), and the data have
have therefore been encouraged to communicate them for
been pooled and released in electronic form since 1997 (e.g.
inclusion in the Plant DNA C-values database, thus making
the Angiosperm DNA C-values database). To extend the
them accessible for comparative studies. The present work
coverage to include other plant groups, databases for gym-
is proof of this hypothesis and provides a model example of
nosperms, pteridophytes and bryophytes were added in 2001
collaboration to transfer a large body of previously invisible
to create the Plant DNA C-values database (release 1 0) with
plant genome size data firmly into the public domain.
C-values for 3864 land plants. In December 2004 the Plant
Two of the present authors have worked together for the
DNA C-values database was further updated (release 3 0)
past decade to produce pooled lists of plant DNA C-values
and included, as a novel feature, C-values for 253 algal
for reference purposes in both hard copy (Bennett and
species. Release 3 0 has DNA amounts for almost 5000
Leitch, 1995, 1997, 2005a; Bennett et al ., 2000) and elec-
species from over 500 original sources, including first val-
tronic versions (http://www.kew.org/genomesize/homepage).
ues for 628 angiosperms not previously included (http://
Meanwhile the present first author has made genome size
www.kew.org/genomesize/homepage). These lists and data-
estimates for angiosperm taxa in several genera, including
bases have been widely used for comparative studies
Helleborus (Zonneveld, 2001), Hosta (Zonneveld and
* For correspondence. E-mail m.bennett@kew.orgVan Iren, 2000, 2001), Galanthus (Zonneveld et al ., 2003)
© The Author 2005. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved.
ror Permissions, please email: j ournals .permi ssions (2> oupj ournals . org

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230 Zonneveld et al. - Nuclear DNA Amounts in Angiosperms

and Agapanťhus (Zonneveld and Duncan, 2003), some

Table 1. Species used as calibration standards to estimate
of which have subsequently been included in a recent com- nuclear DNA C-values
pilation by Bennett and Leitch (2005a). While correspond-
Assumed Abbreviation used
ing with Bennett and Leitch about these data and planned
2C DNA in column 13 of
future publications, Zonneveld noted that he had also made
Species amount (pg) the Appendix
C-value estimates for taxa in various genera that seemed
most unlikely to be published. It emerged that this sample strida 7-8 A. stricta
Agave
was substantial and included first estimates for hundreds of Hordeum vulgare 'Sultan'* 10-0 Hordeum
miscellaneous species, measured mostly out of curiosity. Agave americana 15-9 A. americana
Given their potential value, it was decided to combine the Agave sisalana 20-2 A. sisalana
Clivia miniata 39-0 Clivia
taxonomie and databasing experience of the Kew-based
authors with information amassed in The Netherlands to
*The 2C value of 10 pg for Hordeum
produce a further reference list in a form similar to other
Agave americana (2C = 1 5 -9 pg) as the c
recent plant C-value reference lists (e.g. Bennett and Leitch,
1 6 1 q in the Appendix). This value is sli
and
2005ö; Kapraun, 2005). The present work is the product Smith (1991) of 2C = 1 112 pg. In t
calibration standard for only one speci
of that collaboration. Analysis of the completed Appendix
see entry number 82b in Appendix). If t
table shows that it contains C- values for 411 angiosperm
by Bennett and Smith (1991) is used t
taxa, including first values for 308 species not listed inbeany
l-6pg.
previous compilation by Bennett and colleagues.
flow cytometer available for use, as the peaks tended to be
lost in the debris fluorescence.
MATERIALS AND METHODS

Plant materials
RESULTS AND DISCUSSION
The present work examined angiosperm taxa available as
Comparison of C-value data in the Appendix table wit
seed or plants growing in The Netherlands selected prim-
previously
arily for their horticultural interest. The Netherlands has a reported
long tradition of growing and breeding many angiosperm
To assess the quality of C-value data in the Append
taxa for horticulture, consequently such plant materials are
comparisons were made.
readily available for research purposes. (1) C-values for 103 species in the Appendix were
pared with 'prime' (see note (a) in 'Notes on com
Estimation of DNA C-values the Appendix') C-values for the same species listed
Plant DNA C-values database (Bennett and Leitch
DNA 2C-values were estimated using flow cytometry
with the fluorochrome propidium iodide (PI) as described Figure 1 A shows that generally the two data sets were
by Zonneveld and Van Iren (2001). Briefly, somatic nuclei agreement (R2 = 0-9068). However, there were 18 s
were isolated from leaves by co-chopping them with one where of previously published 'prime' C-values differe
the internal calibration standards (Table 1) in nuclei isola->30% from those given in the Appendix. These
tion buffer. After adding 2 mL PI solution (50 mg PI/1 are in represented by open circles in Fig. 1A. The
half-strength isolation buffer) the suspension was filtered. of the discrepancies was not clear but possibilities
Nuclei fluorescence was measured 30 min after the addition taxonomie errors or genuine intraspecific variation.
of PI using a Partee CA-II flow cytometer. In most cases, of the species the different C-values reported were m
several samples, each with at least 5000 nuclei, were meas- of previous estimates, suggesting that polyploidy m
ured. The 2C- value of the sample was calculated as follows: responsible. Nevertheless, as no chromosome counts
(mean of sample peak -r mean of standard peak) x 2C DNAmade for any species in the Appendix (except Hydr
amount (pg) of the standard species (see Table 1). Othermacrophylla ), the role of polyploidy is unconfirme
peaks corresponding to higher C-values (i.e. 4C, 8C and (2) A comparison was made between the C-valu
16C, etc.) were sometimes observed but their DNA amounts seven species in the Appendix and the C-values for
were not calculated. calibration standards given in Bennett and Smith (1
These two datasets showed very close agreement (F
Internal calibration standards (R2 = 0-9965). Overall, it was concluded that the Ap
C-value data are of high quality.
Five calibration standards, differing in DNA amount,
were used (Table 1). In most cases Agave americana was
Representation of C-value data in Appendix
chosen, as previous work showed it to be a reliable and
reproducible standard, giving clean peaks with low CVs The present Appendix gives C-values for 411 s
(Zonneveld and Van Iren, 2001). If nuclear DNA content
of which 308 (75 %) are for species not included
of a test species overlapped with that of A. americana ,
previous compilation by Bennett and colleagues (Be
another calibration standard was used whose C-value fell
and Smith, 1976, 1991; Bennett et al ., 1982, 2000; B
and Leitch, 1995, 1997, 2005a) or listed in the Plant
close to that of the unknown. Species with 2C- values below
C-values database (release 3 0, December 2004). B
2 0 pg could usually not be measured with the Partee CA-II

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 Zonneveld et al. - Nuclear DNA Amounts in Angiosperms 231

60p A y = 0-8582*: + 0-8914 Mr y = 0957* + 0-5 1 72

A R2 = 0-9008 'S Rl = 09965
££ 3 % Triticum aestivum
^ # -S Allium cepa
g
0 •A('
# /_#/• #
co •Ē co
15 -/
- /
& A(' _ "g y Vicia faba
1 I io-
«u « y PO Y Secale cereale
Oh <ü
Oh 20 20
- * o « */ /yo^ o-Z y/
3 Jr ° ao
13 ^ < -_
^ ^ Hordeum
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-Z Hordeum
I . I
^ 0 o > y/ Pisum sativum
Sjw ^ • Senecio vulgaris
0®-

0 20 40 60 0 5 10 15 20

lC-value in Appendix (pg) lC-value in Appendix (pg)

Fig. 1. Comparisons between C-values for species listed in the Appendix and 'prime' C-values for the same species listed in
database (release 3 0, December 2004). (A) Comparison for all 103 species. Open circles correspond to the 1 8 species whose C-valu
by >30 % from the 'prime' C-values in the database. (B) Comparison between C-values for seven species listed in the Appendix and
these calibration standards given in Bennett and Smith (1991).

and Leitch (2005a) analysed the global output

Table 2. of DNA
Representation of C-value data in the Appendix at
C-values for angiosperms during the past half century. various taxonomie levels
They concluded that the 1997 Angiosperm Genome Size
No. listed in No. of first
Workshop had stimulated a large increase from an average
Appendix table C-values
of about 100 per annum in the early 1990s to almost 200 per
annum for 1998-2002. The total number of first C-values
Species 411 308
for species in the present work therefore equals 1 -5 yearsBasal
of angiosperm 8 5
average global published output at the record levels
Monocots 253 196
Eudicots 1 50 1 07
achieved in recent years. By any measure, it is a significant
Genera 187 85
contribution to knowledge of this important character.
Basal angiosperm 6 2
Indeed the present data sample (all from one source)
Monocots 117 56
increases the total sample of angiosperm species with
Eudicots 64 27
genome size estimates by about 7-4%. Moreover, it con-
Family 67 1
Basal
stitutes over 12% of the global target set by the Second angiosperm 5 0
Monocots 2 1 0
Plant Genome Size Workshop (namely to estimate first
Eudicots 4 1 1
DNA C-values for about 2500 angiosperm species within
a quinquennium; see http://www.rbgkew.org.uk/cval/
workshopreport.html). Clearly, one worker can still make
a significant contribution to the global pool of informationconcluded th
on genome sizes. Failing to make such information visible representation
could seriously impair the scope of the available database careful target
and limit the scope of comparative studies. of untargeted
Table 2 shows how many of the C-values in the Appendix the Appendi
only
are first values at the species, generic and family level for one of t
the three major angiosperm groups (i.e. basal angiosperms, Tamaricaceae)
monocots and eudicots). Although monocots constitute only Also as expec
approx. 20% of all angiosperm species, most of the new the present sa
C-values in the Appendix are for monocots (i.e. 64 and 65 filling
% gaps fo
of first C-values are for monocot species and genera,
under-repres
respectively). bog, fen, tun
Nevertheless, while the present data make a significant halophytic, in
contribution to knowledge of C-values at the species level, phytic species
there has been no significant improvement in representation the present sam
at the family level. The 1997 Angiosperm Genome Size such interest
Workshop recommended a goal of complete familial rep- hederae and
resentation by 2002, as a first C-value was available for with 1C valu
only about 30% of families. Bennett et al. (2000) noted table also incl
that in a sixth supplementary list of C-values for 69 1 species (1C = 2-24 pg
only 12 were also first estimates for families. Later they in the Cactaceae, and a number of succulents from the

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232 Zonneveld et al. - Nuclear DNA Amounts in Angiosperms

Table 3. The minimum , maximum , mean , median ,Future

mode, and
plans
range of IC DNA values for 411 species in the Appendix com-
Participants at the 2003 Plant Genome Size Workshop
pared with the corresponding statistics for 4119 species in thetheir international collaboration as the Genome
formalized
Plant DNA C-values database ( release 3-0, December 2004)
Size Initiative (GSI; Bennett and Leitch, 2005ft), whose aims
Plant DNA C-values are to improve the availability, quality and understanding of
database (release 3 0, genome size data, and to provide a focus for monitoring
Appendix December 2004) progress and facilitating discussions in a holistic genomic
context. Further, in a keynote address opening the 2003 Plant
Minimum 1C- value (pg) 0-6 01 Genome Size Discussion Meeting, Bennett reiterated the
Maximum 1C- value (pg) 95 0 127-4
potential value of unpublished genome size data, and the
Mean lC-value (pg) 1 1*7 61
Median 1C- value (pg) 6-6 2-7 need to make it visible as follows: 'There are probably over
Mode 1C- value (pg) 5 0 0-6 1000 prime C-value estimates for plant species, equal to
Range (max/min) 94-4 1274 0 4 years of known global output, unlocated in publications
or unpublished. Such genome size data are almost valueless
if potential users cannot easily access and assess them. It is
impossible to over-emphasise the importance of publishing
families Asparagaceae (Agave, Ledebouria), Bromeliaceae
C-value data or linking them to the database by informing
( Tillandsia ), Crassulaceae ( Aeonium , Graptopetalum ,
the co-ordinators of its existence, so that others can assess its
Sedum ), Apocynaceae (Ceropegia, Hoodia , Hoya,
quality and use it'. The present work supports this sentiment
Catharanthus , Stapelia ), Xanthorrhoeaceae (Aloe) and
and the aims of GSI. Hopefully, it will encourage others to
Asteraceae (Senecio) in which the C-values ranged from
emulate this example and unveil further useful data which
Ml to 16-85 pg. otherwise may lie forever unpublished.

The range of C-values in the Appendix compared

LITERATURE CITED
with existing data
APG II. 2003. An update of the Angiosperm Phylogeny Group cl
Table 3 compares the minimum, maximum, mean, for the orders and families of flowering plants. Botanical Jour
median, mode and range of DNA amounts for all 41 1 spe- Linnean Society 141: 399-436.
Bennett MD, Leitch IJ. 1995. Nuclear DNA amounts in an
cies listed in the Appendix, with the 4119 species listed in
the Plant DNA C-values database (release 3 0, December Annals of Botany 76: 113-176.
Bennett MD, Leitch IJ. 1997. Nuclear DNA amounts in angios
2004). new estimates. Annals of Botany 80: 169-196.
The range of C-values in the present sample (1C = 0-6- Bennett MD, Leitch IJ. 2004. Plant DNA C-values database (r
95-5 pg) falls within that reported for the Plant DNA December 2004). http://www.kew.org/cval/homepage.html
C-values database (1C = 0- 1-127-4 pg), although it is some- Bennett MD, Leitch U. 2005a. Nuclear DNA amounts in an
progress, problems and prospects. Annals of Botany 95: 45-
what enriched in species with larger genomes, leading to
Bennett MD, Leitch IJ. 2005¿>. Plant genome size research: a fie
higher mean and modal values compared with the larger Annals of Botany 95: 1-6.
sample in the database (Table 3); thus the minimum C-value Bennett MD, Smith JB. 1976. Nuclear DNA amounts in angiosperms.
in the Appendix is the same as the modal value for species Philosophical Transactions of the Royal Society of London . Series
B. Biological Sciences 274: 227-274.
in the database. This probably reflects the greater proportion
Bennett MD, Smith JB. 1991. Nuclear DNA amounts in angiosperms.
of monocots in the present sample with large genomes. Philosophical Transactions of the Royal Society of London. Series
Indeed the 43 species (approx. 10 %) with the largest gen- B. Biological Sciences 334: 309-345.
omes in the Appendix are all monocots from the orders Bennett MD, Smith JB, Heslop-Harrison JS. 1982. Nuclear DNA amounts
Asparagales, Commelinales or Liliales; these groups have in angiosperms. Proceedings of the Royal Society of London. Series B.
Biological Sciences 216: 179-199.
previously been shown to contain species with the largest
Bennett MD, Bhandol P, Leitch U. 2000. Nuclear DNA amounts in angio-
recorded angiosperm genomes (Leitch et al ., 1998; Soltis sperms and their modern uses: 807 new estimates. Annals of Botany 86:
et al ., 2003). (NB The presence of only eight species with 859-909.

2C- values <2-0 pg is, in large part, due to technical diffi- Brandham PE, West JP. 1993. Correlation between nuclear DNA values
and differing optimal ploidy levels in Narcissus, Hyacinthus and Tulipa
culties encountered in estimating C-values in species with cultivars. Genetica 90: 1-8.
smaller genomes, as noted in Materials and methods.) Cavalier-Smith T. 1985. The evolution of genome size. Chichester: John
Analysis at the family level showed that in half of the Wiley & Sons.
66 families already represented in the Plant DNA C-values Dabrowska J. 1992. Chromosome number and DNA content in taxa of
database, the new data did not increase the range of Achillea L. in relation to the distribution of the genus. Prace Botaniczne
49: 1-84.
C-values previously reported for a family. In only eight
Kapraun DF. 2005. Nuclear DNA content estimates in multicellular
families was the minimum C-value decreased as a result
eukaryotic green, red and brown algae: phylogenetic considerations.
of new data, whereas in 25 families the maximum C-valueAnnals of Botany 95: 7-44.
Knight CA, Molinari N, Petrov DA. 2005. The large genome constraint
increased, sometimes greatly. For example, in Garryaceae
and Escalloniaceae, which both previously had a C-valuehypothesis:
177-190.
evolution, ecology and phenotype. Annals of Botany 95:

for just one species, the addition of an estimate for oneLeitch

new IJ, Chase MW, Bennett MD. 1998. Phylogenetic analysis of
species increased the range of C-values for each family overDNA C-values provides evidence for a small ancestral genome size
8- and 15-fold, respectively. in flowering plants. Annals of Botany 82: 85-94.

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 Zonneveld et al. - Nuclear DNA Amounts in Angiosperms 233
Leitch IJ, Soltis DE, Soltis PS, Bennett MD. 2005. Evolution S.
ofpendulus
DNA and S. radie ans), however, the different C-values
amounts across land plants (Embryophyta). Annals of Botany 95:
so strongly suggested the presence of polyploidy that they
207-217.
were given separate entry numbers in accordance with pre-
Price HJ, Dillon SL, Hodnett G, Rooney WL, Ross L, Johnston JS. 2005.
Genome evolution in the genus Sorghum (Poaceae). Annals ofvious practice
Botany (e.g. Bennett and Leitch, 2005a).
95: 219-228. For 103 species, where one or more C-value had previously
Soltis DE, Soltis PS, Bennett MD, Leitch IJ. 2003. Evolution of been listed by Bennett and colleagues, the estimate in the
genome size in the angiosperms. American Journal of Botany 90:Appendix was given a number and the next available letter in
1596-1603.
United Nations Environment Programme. 1992. Convention on bio-
the alphabet. For example, three previous C-values have been
logical diversity. Nairobi, Kenya: United Nations Environment reported for Allium ampeloprasum , thus the entry for this
Programme. species in the Appendix was assigned the letter *d'
Zonneveld BJM. 2001. Nuclear DNA contents of all species of Helleborus
(b) Taxonomie authorities . Taxonomie authorities of
(Ranunculaceae) discriminate between species and sectional divisions.
Plant Systematics and Evolution 229: 125-130. the species studied were taken from the International
Zonneveld BJM, Duncan GD. 2003. Taxonomie implications of genome Plant Names Index (IPNI) (http://www.ipni.org/index.html).
size and pollen colour and vitality for species of Agapanthus L'Heritier
A superscript 'b' following a species name indicates that the
(Agapanthaceae). Plant Systematics and Evolution 241: 1 15-123.
Zonneveld BJM, Van Iren F. 2000. Flow cytometric analysis of DNA taxonomie authority was unknown or unclear (i.e. more than
one entry in IPNI).
content in Hosta reveals ploidy chimeras. Euphytica 111: 105-110.
Zonneveld BJM, Van Iren F. 2001. Genome size and pollen viability as
(c) Family and higher order group . The family, order and
taxonomie criteria: Application to the genus Hosta. Plant Biology 3:
176-185. higher group given for each species in Appendix columns 3,
Zonneveld BJM, Grimshaw JM, Davis AP. 2003. The systematic value4ofand 5, respectively, follows the Angiosperm Phylogeny
nuclear DNA content in Galanthus. Plant Systematics and Evolution
Group (APG) system (APG II, 2003). The following abbre-
241: 89-102.
viations were used: BA = basal angiosperm; M = monocot;
E = eudicot.
APPENDIX
(d) Chromosome number, ploidy and voucher informa-
Notes on compiling the Appendix tion . The sentiment of the Convention on Biological Divers-
C- value data were compiled into the Appendix ity (United
tableNations
whose Environment Programme, 1992), which
form is similar to that of previous lists of noted
DNA theamounts
need to make biodiversity data available despite
by Bennett and colleagues (i.e. Bennett and imperfections,
Smith, merits
1976,support. Thus, while the species listed
1991; Bennett et al., 1982; Bennett and Leitch, 1995, in the Appendix generally lack a voucher and information
1997; Bennett et al ., 2000). Additional information on on chromosome number and ploidy level, they can make a
compiling the Appendix is given below. valuable contribution to knowledge of plant C-values.
Chromosome numbers and/or ploidy information were
(a) Assignment of entry numbers and * prime ' entriesentered
. for 90 taxa where previous C-value reports also
The numbering of the Appendix table generally follows that
included this information and the C-values agreed closely
for previous lists of DNA amounts (see above). Taxa arewith those reported here. For example, a C-value of 2-9 pg
arranged alphabetically by genus and species with a new for Achillea filipendulina was previously estimated by
number usually being assigned for each new species. Dabrowska (1992), who also reported that the accession
For 42 species in the Appendix more than one estimate was studied had 2n = 2x = 18. As the 1C- value of 2-88 pg for
made for the same species (e.g. for different cultivars, vari-
the same species in the Appendix was very similar to that of
eties, etc.). In most cases, the C-values were very similar and
Dabrowska (1992), the chromosome number and ploidy data
the different accessions were assigned the same entry number were also entered.
but were distinguished by letters, sorted here by increasing
DNA amounts. For example, C-values are reported for two (e) Life cycle types. Information on the type of life cycle
cultivars of Crocus chrysanthus , a species not included pre-for species in the Appendix was either taken from the liter-
viously in any of the lists mentioned above. The smaller of ature or internet. The following abbreviations were used: A =
annual; B = biennial; AP = annual-perennial; P = perennial.
the two values, for 'Dorothy', is listed as entry number 1 13a,
while the larger, for 'Blue Pearl', is 113b. Where several A superscript 'e' indicates that life cycle information was
estimates are available for the same species, the 'a' value either unknown or unclear to the present authors.
is automatically chosen in any arithmetical or statistical (f) DNA amounts and conversion factor. 1C and 4C values
calculations. In this context, a single estimate for a species
listed in the Appendix were determined by appropriate cal-
and 'a' values are referred to as 'prime entries'. culation of the 2C- value. For 1 C-values in megabase pairs
For 18 species the C-values between accessions differed(Mbp) the conversion factor of 1 pg = 980 Mb (Cavalier-
by >10 %. Taxonomie heterogeneity, polyploidy and/or Smith, 1985; Bennett et al., 2000) was used.
intraspecific variation are the most likely explanations
(g) Standard species. The abbreviation used to indicate
(especially where a range of chromosome numbers has pre-
which of the five calibration standards was used to estimate
viously been reported for the species), yet until chromosome
counts are made, the cause(s) of the differences remainsthe 2C- value for each species in the Appendix is given in
Table 1.
unknown and the different accessions of a species were usu-
ally assigned the same entry number. In six Senecio species (h) Chromosome numbers in Hyacinthus. Brandham
(S. herreianus , S. kleiniiformis , S. mweroensis , S. nyikensisand
, West (1993) reported chromosome numbers in

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 Zonneveld et al. - Nuclear DNA Amounts in Angiosperms 243
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244 Zonneveld et al. - Nuclear DNA Amounts in Angiosperms

13 Hyacinthus orientalis culti vars and noted that are built up from three layers, LI, L2 and L3, which develop
aneuploidy
was common. Two of the cultivars studied ('Pink fromPearl'
the apical meristem. In the variegated S. infasciata
and 'Delft Blue') were the same as those listed in the the yellow leaf edge corresponds to cells derived
'Laurentii'
from LI and the green cells in the leaf centre from L3.
Appendix. Since the cultivars are clonal the chromosome
BetweentoLI and L3 is a single layer of cells which comprise
numbers given in Brandham and West can be assumed
be the same as those studied here. Thus 'Pink Pearl' L2,
is listed
from which the floral meristem and hence the gametes
as a diploid with 2n = 16, while 'Delft Blue' is anwill arise. Measurements of 2C values from leaf tissue taken
aneuploid
tetraploid with 2n = 4x - 2 = 30. Based on similar 'from the yellow edge gave a 2C value of 5-2 pg whereas
C-values
in cultivars 'Pink Pearl' and 'White Pearl' (28-4 and nuclei
27-3isolated
pg, from the green central tissue (L3) were 2C
= 2-6 pg. As
respectively), the latter is also entered as diploid. Further, as it was difficult to be sure of obtaining cells
the DNA amounts for cultivars 'Delft Blue' and 'Anna from L2, the 1C value of the gametes is unclear. Thus
Marie' are similar (43-4 and 42-4 pg, respectively), the this value has been omitted from the Appendix, and just
latter
the 2C and 4C values from the layer with the lowest DNA
is entered as circa tetraploid, although the exact chromosome
number is unknown. The higher C- value cultivar 'L'Inno- amount (i.e. from L3) are entered. In S. infasciata 'Forescate'
cence' (51-60pg) suggests it is pentaploid. cells in the green central tissue had a 2C value of 7-8 pg (L3)
whereas those at the yellow edge (LI) were 2C=5-2pg.
Again,
(i) Sanse veria tricasciata ploidy chimeras . The leaves of the 1C value was omitted and just the 2C and
4C values for the layer with the lowest DNA amount (i.e.
Sanseveria infasciata 'Laurentii' and 'Forescate' are ploidy
chimeras (Zonneveld and Van Iren, 2000). Most plant from leavesLI) were entered.

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