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Nuclear DNA Content of Some Important Plant

Species

Article in Plant Molecular Biology Reporter November 1991

DOI: 10.1007/BF02672069

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Plant Molecular Biology Reporter pa ges 208-218
Volum e 9(3) 1991

Genetic Resources

Nuclear DNA Content of Some Important Plant

Species
K. Arumuganathan and E. D . Earle*
Department o f Plant Breeding a nd Biom etry, Co rnell Univer sity,

Ith aca , NY 14853-1902, USA .

Key words: Geno me size, flow cytometry, pr opidium iod id e, Arabidops is, rice,
toma to .

Ab stract: Nuclear DNA contents of m or e than 100 important plant sp ecies were
measured by flo w cytometry of isol ated nuclei stained with propidium iodide.
A rabidopsis exhibits d ev elopmentally regulated multiploidy an d has a 2C nuclear
DNA content of 0 .30 pg (145 Mbp/1C), twice th e value u su ally cited . Th e 2C
value for ric e is on ly abo ut three times that of A rabidopsis . Tomat o has a 2C value
of ab out 2.0 p g, larger than commonly cited . Th is survey identified sever al
horticultural crop s in a variet y of families with gen omes o nly t wo or three tim es
as large as Arabidopsis; the se include several fruit tr ees (a prirot, ch erry, mango,
o ra nge , papaya, and pe ach). The small gen ome sizes of ri ce a nd th e h orti cultural
plants sho uld facilitate m ol ecular stud ies of th ese cr op s.

uclear DNA con tents of over 100 important plant species, were

N es tima ted by flow cytometry (Arumuganathan and Earle, 1991),

and are presented in Table 1. This table include s the major cro p
plants, some of which are widely used in m olecul ar studies (e.g., rice and
tom ato), as well as Ambidopsis thaliana. For some, one or m ore nuclear
DNA values (usually fr om Feul gen micrcdensitornetry) have been pub
lished; for many o the rs no such value s have been reported . The inforrna
tion list ed may be helpful to plant biologists interested in gen om e
analysis or in the relationship of nu clear DNA content to plant physiol
ogy a nd ecology (Benne tt, 1985).

"To w ho m corresp ondence sho u ld be ad d ressed .

Abbreviation s: C, DNA co nte n t of the unreplic at ed haplo id ch ro moso me

co m ple m en t; CRI3C, chicken red bloo d cells; CE, chemical extractio n; FC, flow

cyt om et ry; FMD, Feulgen m icrod en sito me try; RK, reassoci ati on kin etics.

208
Nuclear DNA Content 209

Material and Methods

For flow cytometric analysis, suspensions of intact nuclei prepared by
chopping pieces of young leaf tissues in MgS0 4 buffer were stained with
propidium iodide and treated with DNAase-free RNAase (for details,
see Arumuganathan and Earle, 1991). The DNA content of 2C nuclei
(those in the G/G] phase of the cell cycle) was calculated by using nuclei
from chicken red blood cells (CRBC) as internal standards. For the
absolute DNA content of CRBC nuclei, we used the value of 2.33 pg/2c,
as estimated chemically by Galbraith et aL (1983)after extraction of total
DNA from a sample of cells.

Results and Discussion

In Table 1 plant species are presented in alphabetical order together
with the DNA amount in 2C nuclei; a calculation of the number of base
pairs per lC (the unreplicated haploid genome of the species) is also
provided . For plants that are not diploid in the cultivated form, the
ploidy level of the species is included to permit estimation of actual
genome size . For some species, several varieties or cultivars were
assayed; nuclear DNA content often varied somewhat among the differ
ent cultivars, as has been previously reported (Bennett, 1985). Table II
presents data for the same species in increasing order of genome size.
Arabidopsis has been reported to have the smallest genome known
among flowering plants (Leutwileret al., 1984). In our survey Arabidopsis
also had the lowest value; its 2C nuclei contained 0.30 0.012 pg DNA
(mean standard deviation, n = 36). This is, however, about twice the
widely cited genome size (70 Mbp or -0.0725 pg DNA per lC genome),
deduced from DNA reassociation kinetics (Leutwiler et al., 1984).
Arabidopsis showed multiploidy in many of its tissues. Nuclei from
leaves produced four or five distinct peaks of fluorescence (Fig. I A),
representing the 2C, 4C, 8C, 16C, and 32C complements of the genome.
Fewer than 30% ofthenuclei were in the 2C peak. The fourth (16C) peak
coincides with or falls very close to the peak for nuclei of CRBC, used as
an internal DNA standard (Fig.1B). Although nucleifrom rosette leaves
and roots showed up to six peaks, nuclei from leaves on the flowering
stem showed three or four; those from the flowering stem itself had only
three. Nuclei from flower buds fall into only two peaks of fluorescence
(Fig. 10. Similar results were seen withcv. Columbia and cv. Landsberg
(erecta mutant) and with plants grown either in sterile nutrient agar
medium or in soil. These results suggest tha t multiploidy in Arabidopsis
210 Arumuganathan and Earle

800
A 8C 800 B CRBC,
4C
:8 600 :8 600
Debris
~ 400 ~ 400 4C
<J
;:l
Z 200 ~ 200
16C

O+-.--r-..........,,..........."'T"'"'"'......opI...~.......,.-.-J,
32 64 96 128 iso 192 224 256 o 32 64 96 128 160 192 224 256
Chan nel number Chann el number

i ~llC-. . . ._.
. . . .l ;
o 31 64 96 128 160 192
{""CI
224 256
i ~
0

O-o~--""''''''''''''''''''''''''''''''''1
32 64 96 l2 8 160 192 224 256 111
Cha nnel number Channel number
300 T-::,----- - - - - - - - - - ..., 500 "r."F,----------::-;:-.,----==-::-:::-1
E I
CRDC
Tomato Gl ~4()()
\ ~
Q
300
' Q;
U 200
::l
Z IOO
0 -1--0...,......,....
o 32 64 96 128 160 192 224 256
o+-__........... ~~ .... ~~

o 32 64 96 128 160 192 224 256

Cha nnel number Channel number
Fig. 1. H istograms of numbers of nuclei p er channel as a function of relative
fluore scence intensity. Signals from nucl ei w er e g ate d to elimina te much of the
d ebris from analysis (exce p t in N . Histo grams acqu ired on EPICSPROFILE flow
cyto meter ar e presented in Macintosh for ma t (Ca meron, 1990). Th e cha nnel
number is proportional to the log of the fluor escence int en sity (except in E), such
that a differ ence o f 26 channe ls represent s a doubling of flu orescen ce intensity.
Nuclei fr om chicken red blood cells (CRBC) wer e included (except in A) to serve
as an internal sta ndard for the d eterminations of plant nuclear DNA contents .
Nuclear DNA content s were calculated by com pari ng mean peak posit ions of th e
plant nucl ei to the me an peak positions of th e nucl ei of CRBe. ( A) Five
flu or escen ce p eaks representing five ploid y classes (2C, 4C, SC, 16 C and 32C) for
nuclei isolated fr om leaftissues of A. ihalian a. (B) The fourth nucl ear pe ak from
mature leaf of A. thaliana coincides with the nuclear p eak of CRI3C. (C) Only two
peak s of flu or escenc e are present fr om nuclei (2C and 4C) from flower buds of A.
ihaliana. (0) The th ird nud ear pe ak from mature lea f of a tran sg eni c A rabidopsis
plant co incides wi th th e CRBC peak, ind icating that the plant is a tetraploid with
at least four ploidy lev els, as in normal plants. (E) The relative peak positions for
the nudei from leaves o f rice and tom ato, a nd CRBe. (F) Relativ e flu orescence
peak positions of nucl ei from Arabidopsis, rice, tomato and CRBe.
Table I. Nucl ear DNA co n te n t of a number of important plant s peci es as determined by fl ow cyto m e try. '2:
l'::
('>
~
Nuclea r DNA con te n t l::>

This study P r eviou s w o rk

...
tJ
C ommon pgl2C '2:
~
Scientific name nam e Fa m i ly pg/ 2ca (N) -MbpbllC (meth od u sed, re n c
n
a
:::s
....
Aegilops sqUtm osa Gram inea e 8.34 4024 7.2-11.6 (FMO, 1)
;:::
""
Allium ampeloprasum Leek Am aryllidaceae 50.27 24255 NO ....

Allium cepa Onion Ama r yllida ceae 31.69,32.74 (2) 15290, 15797 3.55 (FMO, 1)

Aromas bracteatus Red pineapple Brorneliaceae 0.92 444 NO

Aromas comosus Pineapple Brome liaceae 1.09 526 N O

Arabidopsisthaliana Arabido psi s Cruciferae 0.30 (2) 145 0.5 (FMO, 1);

0.15 (RK, 2)
Arachis hypogam (2n=4X) Pean u t/ groundnut Leguminosae 5.83 2813 3.5 (FMO, 5)
Asparagus officinalis Asparagus Liliaceae 2.71 1308 NO
Avena sa/iva Oa ts Gra mineae 23.45 11315 26.5-27.5 (FMO, 1)
Beta vulgaris ssp. esculenia Beet / beetroot Ch enopodiaceae 1.48 714 2.5 (FMO, 1)

Beta VUlgaris ssp. sacchariiera Suga r beet Che nopodi aceae 1.57 758 2.6 (FMO, 1)

Brassicacampestris ssp. chinensis Pak choi Crucifera e 1.05 507 1.6 (FMO, 1)

Brassicacampestris ssp. oteiiera Turnip rap e Cruciferae 0.97-1.07 (6) 468-516 1.6 (FMO, 1)

Brassicacampesiris ssp. rapifera Turnip Crucifera e 1.06 511 1.6 (FMO, 1)

Brassica hirta (=Sil1l1pisalba) White mu st ard Cruci ferae 1.02 492 1.0 (FMO,3)

Brassicajuncea Brown mustar d Cr ucifera e 2.29 1105 3.1 (FMO, 1)

Brassica napus Rapeseed Cruci fera e 2.34-2.56 (4) 1129-1235 3.2 (FMO, 1)

Brassica nigra Black must ar d Cruciferae 0.97 468 1.6 (FMO, 1)

Brassica oleracea ssp. botrytis Ca uliflower Crucifera e 1.30-1.37 (4) 628-662 1.8 (FMD, 1)

Brassica oleracea ssp. capitata Cabbage Cruciferae 1.25 603 1.8 (FMO, 1)

Brassica oleracea ssp.gemmifera Bruss els sprouts Cruciferae 1.30 628 1.8 (FMO, 1)

Brassicaoleracea ssp. italica Broccoli Cruci ferae 1.24, 1.28 (2) 599,618 1.8 (FMO, 1)

Brassica tourneiorthii Cruciferae 1.64 791 NO tv

.....
.....
Capsicum annuum Pep per / chil i Solanaceae 5.6-7.51 (7) 2702-3420 8.0- 10.8 (FMD, 5);
5.52 (FC, 7)
Carica paptrya Papa ya Caricaceae 0.77 372 NO tv
......
Cicer arieiinum Chi ck pea Leguminosae 1.53 738 1.9 (FMD, 1) tv
Citrullus vulgaris (= lanalus) Watermelon Cucurbitaceae 0.88, 0.90 (2) 425,434 NO
Citrus sinensis Ora nge Rutaceae 0.76, 0.82 (2) 367,396 NO
Crepis capillaris Crepis Composil ae 3.87 1867 4.1 (FMD, 1); 3.55
(FC,8)
Cucumis melo Ca n talo u pe Cucurbitaceae 0.94, 1.04 (2) 454, 502 1.9 (FMD, 3)
Cucumis saiious Cucumber Cucur bitaceae 0.76 367 2.1 (FMD, 1)
Cucurbila pepo Zucc hini Cucur bitaceae 1.04, 1.08 (2) 502,521 5.6 (FMD, 3)
Datura stramoniun [im son w eed Solanaceae 4.11 1983 NO
Daucus carota Carrot Umb elliEer ae 0.98 473 2.0 (FMD, 1)
Dioscorea 0.10.10. Yam Dioscor eac eae 1.15 555 NO
Diplotaxis erucoides Cruciferae 1.31 632 NO
Eruca satit/a Cru ciferae 1.16 560 NO
Glycine max (2n=4X) Soybean Leguminosae 2.31 1115 1.9 (FMD,3)
Goss ypium hirsulum (2n=4X) Co tto n Malv aceae 4.39,4.92 (2) 2118, 2374 6.1-6.5, 6.06 (FMD, 1,
6); 5.6 (FC, 6)
Helianthus annuus Sunflower Composit ae 5.95-6.61 (3) 2871-3189 4.9-9.9,7.22 (FMD, 3,
6); 3.67, 6.96
(FC, 7, 6)
Hordeum vulgare Barley Graminea e 10.10 4873 10.7-11.1 (FMD, 1)
Iporrvea balalas (2n=6X) Sw eet potato Conv olvul aceae 3.31 1597 NO
Lactucasativa Lettuce Com posita e 5.47 2639 5.32 (FC, 6);6.06 ...;;::~
(FMD,6) ::i
;;::
Lensculinaris (e esculenta} Lentil Leguminosae 8.42 4063 9.2 (FMD, 1)
Lyco persicon chsesemanii Solan aceae 1.83 883 NO ~
Lycopersicon esculenium Solan aceae 1.88-2.07 (6) 907-1000 1.48, 1.9 (Fe, 7, 6); ~
Toma to S.
2.0-5.1, 2.05 (FMD, 1, 6) ~
;:l
Lycopersicon pennellii Solan aceae 2.47-2.77 (3) 1192-1337 NO ~
;:l
Lycopersiconperut/ianum Solanaceae 2.27 1095 NO 1:l..
Malus x domestica (2n =2X) Ap p le Rosac eae 11.54-1.65 (3) 743-796 NO tTl
~
Mangifera indica Mango Ana cardia ccae 0.91 439 NO -e
~
Manihotesculen ta (=utilissima) Cassava / manioc Euph orbiaceae 1..43-1.72 (17) 690-830 NO ~
l':
....,
Medicago so. tiva (2n=4X) Alfalfa / lucern e Legu minosae 3.13 1510 3.5 (FMO, 1) nl
Medicago truncalula Leguminosae 0.94- 1.09 (6) 454-526 NO l':l
....
Me/ilotusofficina/is Sweet clover Legu minosae 2.25 1086 NO tJ
Musa sp. Banan a Musaceae 1.81 873 NO
Nicoiiana plumbaginifolia Solanaceae 4.74 2287 NO ~
Nicotiana tabacum (2n ::4X) Tobacco Solanacea e 8.75-9.63 (4) 4221-4646 7.8 (FMO, 1);9 .67 (FC, 7) n
a
1.5, 1.25 (FMO, 3, 4) ~
Oryza Iongistaminata African rice Grami neae 0.78 376
~
Oryza sativassp.Indica Rice Gramineae 0.87-0.96 (49) 419-463 1.2, 1.67, 2.1 ;:!
...,.
(FMO, 1, 4, 3)
Oryza sativassp. [aponica Rice Gram ineae 0.86-0.91 (20) 415-439 1.2, 1.55, 1.9
(FMO, 1, 4,3)
Oryza sativa ssp.[atanica Rice Gramineae 0.88 (3) 424 NO
Passiflora menspermiiolia Pas sion fruit Passifloraceae 4.54 2191 ND
Persea americana Avocado Lauraceae 1.83 883 NO
Petroselinum crispum Parsley Umbelliferae 3.96 1911 ND
Petuniahybrida Pe tunia Solanaceae 2.64 1274 3.1,3.9 (FMO, 1, 3)
Petunia parodii Solanaceae 2.53 1221 ND
Phaseolus acutiiolius Tepary bean Legurnlnosae 1.34 647 NO
Phaseolus coccineus Scarle t runn er bea n Legu minosae 1.47 709 3.5, 1.9 (FMO, 1,3)
Phaseolus lunalus Lima bean Leguminosae 1.29 622 2.5 (FMO, 1)
Phaseolus vulgaris Com mo n bean Leguminosae 1.32 637 3.7, 2.7 (FMO, 1,3)
Pisum satiuum Garden pea Leguminosae 8.18,9.11 (2) 3947,4397 9.8-10.5,9.75(FMO, 1,6);
7.72,8.55 (Fe, 7,6)
Prunus armenaica Aprico t Rosaceae 0.61 294 ND
Prunus avium Sweet Cherry Rosaceae 0.70 338 NO
Prunus avium x cera sus (2n =4X) Che rry Rosaceae 1.42 685 NO
Prunus ceras us (2n=4X) Sour cherry ' Rosaceae 1.24 599 NO
Prunus domesiica (2n=6X) Prune Rosacea e 1.83 883 NO
Prunus persica Peach Rosaceae 0.54, 0.55 (2) 262,265 NO
Pyrus communis Pear Rosaceae 1.03, 1.11 (2) 496,536 ND N
Raphanus satiuus Radish Cruciferae 1.09 526 0.9 (FMO, 3) >-'
w
 N
Ricinus communis Castor bean Euphorbiaceae 0.67 323 NO >--'
Rubus idaeus ~
Raspberry Rosaceae 0.58 280 NO
S. barbed X S. spontaneum Sugarcane Gram ineae 6.12 2953 NO
Saccharumbarberi Suga rcane Crarnineae 6.54,8.54 (2) 3156,4121 NO
Saccharum afficinarum Suga rcane Gramineae 5.28-7.47 (3) 2547-3605 8.1- 8.7 (FMO, 1)
Saccharum rabustum Suga rcane Gram ineae 6.53 3151 7.7-8.6 (FMO, 1)
Saccharum sinense Suga rcane Gra mineae 8.67 4183 NO
Sesbania rostrata Leguminosae 2.46 1187 NO
Sinapis aroensis Cruciferae 0.76 367 NO
Solan um benhaultii Solanaceae 1.74 840 NO
Solanum melongena Eggpl ant Solanaceae 2.28,2.48 (2) 1100, 1197 2.33 (FC, 7)
Sola num tuberosum (2n =4X) Pota to Solana ceae 3.31-3.86 (3) 1597-1862 4.2 (FMO, 1)
Sorghum bicolor Sorghum Grami neae 1.55, 1.60 (2) 748,772 1.56-1.74 (FMO, 5);1.74
(FC,6)
Spinaciaoleracea Spi nach Chenopodiaceae 2.05 989 1.9 (FMO, 3)
Trifolium pratense Red dover Leguminosae 0.97 468 NO
Trifolium repens White clover Leguminosae 2.07 999 NO
TripsaCLi m dactvloides Gama grass Gramineae 7.73 3730 NO
Trit icum aesiioum ( 2n~X) Wheat Grami neae 33.09 15966 34.6 (FMO, 1, 6); 36.11
(Fe,6)
Triticum monococcu m Grami neae 11.92 5751 12.4-13.4 (FMO, 1)
Tulipa sp, Garden tulip Liliaceae 51.2,63 .6 (2) 24704,30687 72.0 per cell (CE, 1) :.
Vanilla planifalia Vanilla Or chidaceae 15.90 7672 NO ....
>::
Vigna mungo Black gram I urud Leguminosae I.l9 574 NO ~
>::
Vigna radiata Mung bean Leguminosae 1.20 (2) 579 I.l (FMO, 3)
Vigna unguiculala ( = sinensis) Legum inosae 1.27
~
Cowpea 613 NO
Vitis oini[era Grape Vitaceae 1.00 483 NO E
.....
~
X Ciiroio nunella mitis Ca lamondin orange Rutaceae 0.80 386 NO i::l
;:1
Zea diplope rennis Gra mineae 3.57 1723 5.28 (FMO, 5) i::l
lea mays Com Gra mineae 4.75-5.63 (6) 2292-2716 4.42-6.75 (FMO, 5);5 .99 ;:1
l':>.
(Fe,7) t'"rl
....l::l
<i>
Nuclear DNA Content 215

Notes to Table I.
'Value for ea ch cu ltivar was determined by tw o or more me a sur ement s of at lea st
2000 nuclei. N, number o f cu ltivars or v arietie s examined .
bl p icogra m (pg ) = 965 million base p airs (Mbp ) (1).
<FMD, Feulgen m icrod en sit om etry; FC, flow cytometry; ND, not d et ermined ;
RK, reassociat ion kin et ics; CE, chemical ex tractio n; Ref., referen ce : 1, Bennett
a nd Smi th (1 976); 2, Leurwiler et a L (1984); 3, Bennett e t al. (1982); 4, Iyengar
an d Sen (1 978); 5, Laurie and Smith (1982); 6, Micha el son et a l. (1991). 7,
Gal br aith et aL (1983). 8, Galbraith (1990).

is under d evelopmental control, wi th younger tissues showing less or no

multiploidy, as in the succu len t speci es with small geno mes examined
by De Roche r et al. (1990).
Anal ysis of nuclei from lea ves of a transgenic Arabidopsis pl ant sus
pect ed of bein g tetr apl oid su ppor ted our es timation of the nuclear DN A
value in this species. These nuclei pro duced four distinct peak s of
fluor escence as in normal pl ant s, but the third, rather than th e fourth,
peak coincid ed with the CRBC peak (Fig . 1D). This indicat es that the
plant is a tetr apl oid wi th four lev els of ploidy. The nucl ei w ith the low est
fluorescen ce intensity in the tetraploid plant h ad a DN A con ten t of 0.60
0.02 pg. These res ults are co nfirm ation that th e 2C value for d iploid
plan ts is indeed 0.30 p g; i.e., diploids ha ve no smaller peak not visible on
our flow histogr ams.
It is not abl e that several horticultural cro ps in a varie ty of families ha ve
ge no mes on ly two or th ree tim es as lar ge as that o f A mbidopsis (Table Il).
Among these are fru it trees (e.g. peach , aprico t, swee t ch erry ) in the
family Rosaceae (Ta ble 1). Thus lo w nucl ear DNA con ten t is not always
as sociated with sm all size or sho r t life cycles. No multiploidy was
observed in an y of these pl ants, so that phenomenon is also not a
univer sal feature of small genome size.
Th e n uclear geno me sizes of rice and tomato ar e of particular interest
becau se these plants are being used extensively for mapping, isolation,
and transfer of genes and are the most lik ely candid ates for possibl e
complet e geno me seque ncing. Rice has a nucl ear geno me onl y abo u t
three tim es as lar ge as that of A rabidopsis . The 2C geno me size of Oryza
sativa ssp. indica (cv. IR 36) is 0.90 0.03 pg (mean standard deviation,
n= 31). The 2C value for tomato (cv.VFNT cherryhs2.02 0.13 pg (m ean
s tandard dev iation, n=43), about 2.2 times as larg e as the rice geno me
(Fig. 1E). Th ese val ues ha ve been confirmed in many separa te trial s in
our laboratory using other cultiva rsofrice(ra ngeof O.86-0.96 p g/2C) and
tomato (ran ge of 1.88-2.07 pg /2C). The rela tive fluorescen ce of a mixture
216 Arumuganaihan and Earle

Table II. DNA con ten t of unreplicated haploid genome of various plants
arranged according to genome size.
DNA DNA
Pl ant species Mbp/l C Plant species Mb p/lC

Arabidopsis thaliana 145

Diploiaxis erucoides 632

Prunus persica 262, 265

Phaseolus vulgaris 637

Rubus idaeus 280

Phaseolus aeutifolius 647

Prunus arme naica 294

Prunus avium x cera sus (2n =4X) 685

Ricinus communis 323

Man ihotesculenta (= util issima ) 690-830

Prun us avium 338

Phaseolus coccineus 709

Cucumis saiiuus 367

Bela vulgaris ssp. esculen /a 714

Sinapis aruensis 367

Cicerarietinum 738

Citrus sinensis 367,396

Malus x domestica (2n =2X) 743-796

Carica papaya 372

Sorghum bieolor 748,772

Orqza iongis tamina ta 376

Beta vulgaris ssp. saccharilera 758

X Ciiroiortunet! mitis 386

Brassiea iourn eiorth ii 791

Oryza saliva ssp. [aponica 415-439

Solanum berthauui i 840

Orylil sativa ssp. Indica 419463

M usa sp. 873

Oryza saliva ssp. Javanica 424

Persea americana 883

Citru llu s vulgaris (= lanatus ) 425,43 4

Prunu5 domes tica (2n=6X ) 883

Mangifera indica 439

Lycopersicon cneesemanii 883

A nanas bracteatus 444

Lyeopersicon esculent um 907-1000

Cueum is melo 454,502

Spinacia oleracea 989

M edieago truncatula 454-526

Trifolium repens 999

Triiolium pretense 468

Melilotus officinalis 1086

Brassica nigra 468

Lycopersicon peruvianum 1095

Brassiea campesi ri s ssp. oieiiero 468516

Solanum melongena 11 00,11 97

Daucus carola 473

Brassica [uncea 11 05

Vilis u in iiera 483

Glycine max (2n=4X ) 111 5

Brassiea h irta (=Sinapis alba) 492

Brassica napus 1129-1 235

Pyrus commu nis 496,536

Sesbania rostraia 11 87

Cucurbila pepo 502,521

Lycopersicon pennellii 11 92-1337

Brassica campestris ssp. chinensis 507

Petun ia parodii 1221

Brassica campestris ssp. rapifera 511

Petunia hybrida 1274

Raphanus salivus 526

Asparagus officinalis 1308

A nanas comosus 526

Medicagosaliva (2n=4X) 1510

Dioscorea alata 555

I ponrUQ batalas (2n =610 1597

Eruca saliva 560

Solanum iuberosum (2n =4X) 1597-1 862

Vigna .mungo 574

Zea diploperennis 1723

V igna radiata 579

Crepis capillaris 1867

Prunus cerasus (2n=4X) 599

Petroselinum erispum 1911

Brassica oieracea ssp. italica 599,618

Datura slramoniun 1983

Brassica oleracea ssp. capitata 603

Gossypium hirsuium (2n=4X) 2118,2374

V igna unguicu tata (= sinensis) 613

Passijlo ra menspermijolia 219/

Phaseolus lunatus 622

Nicotiana plumbaginifolia 2287

Brassica oleracea ssp. gemmifera 628

Zea mays 2292-2716

Brassica oleracea ssp. bOlrytis 628-662

Lactuea saliva 2639

Nuclear DNA Content 217

Table II, continued .

DNA DNA
Plant species Mbp/lC Plant species Mbp/1C
Capsicum annuum 2702-3420 Lensculmans (= esculen ia} 4063
Arachis hypogaea (2n=4X) 2813 N icotiana tabacum (2n=4X) 4221-4646
Helianthu s annuus 2871-3189 Hordeum vulgare 4873
Saccharum officinarum 2547-3605 Triticum monococcum 5751
S. barberi X S. sponian eum 2953 Vanilla planifolia 7672
Saccharum robustum 3151 Avena saliva 11315
Saccharum barberi 3156,4121 Allium cepa 15290,15797
Saccharum sinense 4183 Triticum aeslitrum (2n=6X) 15966
Tripsacum dactvloides 373 0 Allium ampeloprasum 24255
Pisum sa!ivum 3947, 4397 Tulipa sp. 24704,30687
Aegilops squarrosa 4024

of nuclei from Arabidopsis, rice, tomato and CRBC is shown in Fig. 1F; the
peak for rice G, nuclei falls between the 4C and 8C nuclear peaks of
Arabidopsis; the tomato G 1 peak falls between the 8C and CRBC peaks.
Comparisons of our nuclear DNA values to those from other methods
and / or studies are included in Table 1. For most plant species, including
ones with diverse genome sizes (e.g. mung bean, sorghum, com, pea,
barley, oats, onion, wheat), our results from flow cytometry are in good
agr eement with those obtained in th e laboratory of M. D. Bennett by
Feulgen microdensitornetry (although not with some of the values he
compiled from earlier work elsewhere). The Arabidopsis values esti
mated by reassociation kinetics (0.0725 pg/l0 and by Feulgen micro
densitometry (0.2 pg/1C) are substantially different; our results (0.15
pg/10 are intermediate b etween the two but closer to the latter (Table
I). Our values for rice (0.45/10 are somewhat lower than those from
microdensitometry (0.6 pg /1C or more). The widely used valueof 0.74
pg/1C for tomato, obtained by flow cytometry (Galbraith et al., 1983), is
about25 % lower than ours. This may be due to their use of mithramycin,
a fluorescent stain that preferentially binds to G-C nucleotide pairs.
Because the tomato genome has a G-C content of only 37% (Messeguer
et al., 1991), staining with mithrarnycin is likely to underestimate actual
genome size . Variation in nuclear DNA contents of different cultivars
may also have contributed to the differences seen.
The flow cytometry protocol used in our study (Arumuganathanand
Earle, 1991) should permit easy determination of nuclear DNA values for
additional plants not listed in Table lor previously published reports.
We would be glad to learn of such results for compilation of a more
complete table.
218 Arumuganathan and Earle

Acknowledgments: Wethank S.Tanksley, R. Last, R. Wu,M .Hanson.M . Canal,

e. Martin, R. Wing and J. Slattery for reviewing early ve rsions of th e manuscript;
M . Rangel-Lugo for providing chicken blood; L. A. Batts for technical assistance;
R. Last for the tetraploid transgenic Arabidopsis plant; H. Kik uchi and C. Martinez
for some of the rice and cassava values; D. Heather for some of the values for
Brassica and other Cruciferae. This work wa s supported by a grant from the
Cornell I3iotechnology program (w hich is sp onsored by the New York State
Science and Technology Foundation, a consortium of ind us tries, US Army
Research Office and the National Science Foundation), and by the United States
Department of Agriculture.

Note added in p roof: A recent manuscript [Galbraith, D. W, K. R. Harkins and

S. Knapp. 1991. Systemic endopolyploidy in Arabid opsis ihaliana. Plant Physio!.
(in press)] also reports th e pr esence of multiploidy in th e vegetative tiss ues of
Arabidopsis and estimates the 2C nuclear DNA content as 0.32, 0.33 and 0.34 pg
using mithrarnycin , Hoechst 33258 and DAFl, respectively .

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