nuclear dnacontent varies with cell size across human cell
TRANSCRIPT
Nuclear DNA Content Varies with Cell Sizeacross Human Cell Types
James F. Gillooly, Andrew Hein, and Rachel Damiani
Department of Biology, University of Florida, Gainesville, Florida 32611
Correspondence: [email protected]
Variation in the size of cells, and the DNA they contain, is a basic feature of multicellularorganisms that affects countless aspects of their structure and function. Within humans, cellsize is known tovary by several orders of magnitude, and differences in nuclear DNA contentamong cells have been frequently observed. Using published data, here we describe how thequantity of nuclear DNA across 19 different human cell types increases with cell volume.This observed increase is similar to intraspecific relationships between DNA content and cellvolume in other species, and interspecific relationships between diploid genome size andcell volume. Thus, we speculate that the quantity of nuclear DNA content in somatic cells ofhumans is perhaps best viewed as a distribution of values that reflects cell size distributions,rather than as a single, immutable quantity.
Our understanding of the complexity ofmulticellular organisms, and the diverse
cells of which they are comprised, has dramat-ically increased over the past several decades.Yet, we still lack an understanding of some ofthe most basic features of the cells that consti-tute multicellular organisms. For example, thenumber of different cell types in an organism,or the rate at which different cells grow, divide,and die, remain poorly understood (see Niklas2015). But perhaps most important, we lack anunderstanding of the size and abundance ofcells that constitute an organism (see Amodeoand Skotheim 2015). Cell size, in particular, af-fects virtually all structural and functional attri-butes of multicellular organisms, from the mo-lecular level to the whole organism level.
One key feature of organisms that may varywith cell size is the amount of nuclear DNA.
Across species, genome size has long beenknown to correlate positively with cell and nu-clear volume (Price et al. 1973; Szarski 1976;Olmo 1983). But within species, too, the nucle-ar DNA content of somatic cells has been shownin a few instances to increase with cell size inspecies such as Daphnia (Beaton and Hebert1989) and Arabidopsis (Jovtchev et al. 2006).Such increases in nuclear DNA content canhave important consequences for cell function,in general, and gene expression, in particular(Hancock et al. 2008; Lee et al. 2009; De Veylderet al. 2011; Marguerat and Bahler 2012).
In the case of humans, substantial differenc-es in DNA content have been observed in manyhuman cell types. Indeed, since Watson andCrick described the structure of DNA, studiesof healthy human tissues have reported thepresence of polyploid cells (Winkelmann et al.
Editors: Rebecca Heald, Iswar K. Hariharan, and David B. Wake
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1987; Biesterfeld et al. 1994). The cell types inwhich this has been observed appear to havelittle in common, except that they are generallystable, fully differentiated cells (Winkelmann etal. 1987). Still, these observations have done lit-tle to change the traditional view that all healthysomatic cells in the human body hold the samecharacteristic quantity of DNA (�7 billion basepairs) based on the long-standing principle ofDNA constancy (Mirsky and Ris 1949). Devia-tions from the diploid quantity of DNA in hu-mans, like other animals, are still often viewed asexceptional, tissue-specific, or indicative of pa-thology. A more synthetic view of differences innuclear DNA content across human cell typesmay provide some clarity on these and otherissues.
In this review, we compile and analyze pub-lished data to examine the extent to which nu-clear DNA content varies across diverse humancell types, and whether such variation is corre-lated with cell size. We then compare these re-sults with previously reported relationships be-tween nuclear DNA content and cell size withinfour other species. Finally, we compare theseresults with the relationships between diploidgenome size and cell size observed across speciesin several broad taxonomic groups. These anal-yses suggest that systematic variation in nuclearDNA content is a more ubiquitous phenome-non in human cells than was previously appre-ciated. However, as we later discuss, the mecha-nisms underlying these patterns remain inquestion.
THE RELATIONSHIP OF NUCLEAR DNACONTENT TO CELL SIZE IN HUMANS
Methodology
Our analysis for this work used published datafrom healthy human cell populations represent-ing 19 different cell types, as designated in theoriginal studies (data provided in Table 1). Inthe original studies, DNA content was estimatedusing the Feulgen staining method, and the sizeof cells or cell nuclei were directly measured.Feulgen staining (Feulgen and Rosenbeck1942) has been the most widely used method
for estimating DNA content for several decades,and is still generally considered a reliable meth-od for making quantitative measurements ofDNA content (Chieco and Derenzini 1999; Bies-terfeld et al. 2011). Measurement errors are typ-ically ,5% using this method (Gregory 2005),which for our analyses is negligible given theorders of magnitude variation in DNA contentamong the cells we consider. The method worksby staining DNA owing to the reaction of Schiffor pseudo-Schiff reagents with aldehydes, whichare converted from deoxyribose in DNA afterHCl hydrolysis (Chieco and Derenzini 1999).The light absorbance of the stained genetic ma-terial is then measured to quantify the relativeDNA content of cells.
In cases in which the relative content of DNAwas originally expressed in “arbitrary units” inthe original studies, we converted these mea-sures to pg DNA given that the Feulgen stainingmethod results in a linear relationship betweenlight absorbance and DNA content (Biesterfeldet al. 2011). In each case, a specific conversionfactor was used for each cell type based on theobserved relationship between light absorbanceand DNA content in the particular study (seeTable 1). This was done to avoid any bias as-sociated with differences in DNA compactionlevel or uptake of stain across cell types. Theconversions used for each cell type are given inTable 1.
Finally, in many studies considered here,only the population means of cell size andDNA content were reported. Thus, to remainconsistent, for those studies that reported indi-vidual cell measurements, we calculated meanvalues and used these in the analyses. So, allpoints shown in the figures for humans repre-sent means of DNA content and cell size fromcell populations. In Table 1, we give all values forcell size and DNA content reported in the orig-inal sources, both population mean values andindividual cell values. In the majority of cases,cell volumes were not directly reported, so weestimated these volumes from measures of nu-clear size based on a published relationship be-tween nuclear volume and cell volume acrosshuman cell types (see Table 1 for further detailsof methodology and data) (Swanson et al. 1991).
J.F. Gillooly et al.
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We evaluated the relationship between meannuclear DNA content (pg) and cell volume (m3)using a linear mixed model of the form log10
(DNA) ¼ a þ b log10 (cell volume) þ C þ Dþ E, where a and b are regression coefficients, Cand D represent random effects of cell type (ran-dom slope and intercept, respectively), and Erepresents residual errors (Pinheiro and Bates2000). This statistical model accounts for with-in-group autocorrelation, allowing us to includecells from multiple cell types in a single analysis.To determine whether DNA content was corre-lated with cell volume within cell types, we usedordinary least-squares regression. We verifiednormality of residuals using quantile–quantileplots.
Results
Our analyses of these data indicate that theamount of DNA in human cells varies system-atically with cell size among cell types, such thatlarger cells contain substantially more DNA(Fig. 1A) (mean log10 [DNA] ¼ 21.4 þ 0.77log10 [cell volume], r2 ¼ 0.82, F1, 175 ¼ 36.6,P , 1 � 1029). Figure 1A shows that the meanDNA content forcell populations varies from 1.8to 68.6 pg across cell types ranging from haploidsperm to polyploid megakaryocytes. For 17 ofthe 19 cell types shown in this relationship,cell volume was calculated based on some mea-sure of nucleus size. Without converting to cellvolume, we still observed a significant, positive
1
10
100
Human cell volume (µ3)
Hum
an n
ucle
ar D
NA
con
tent
(pg
)
HumanRattusPseudacrisDaphniaArabidopsis
1
10
100
0.1
Nuc
lear
DN
A c
onte
nt (
pg)
10 102 103 104 105
Cell volume (µ3)
102 103 104
A B
Figure 1. Mean DNA content for cell populations. (A) Mean nuclear DNA content versus cell volume for healthyhuman cell populations from 19 cell types (log10 [DNA]) ¼ 21.3 þ 0.74 log10 [cell volume], r2 ¼ 0.81; data inTable 1). The characteristic diploid human cell contains 7 pg DNA. (B) Relationship between DNA content andcell size in humans (blue) in comparison to previously reported relationships in the rat (Rattus norvegicus, log10
[DNA] ¼ 21.3 þ 0.77 log10 [cell volume] [Heizer 1955]), frog (Pseudacris obscura, log10 [DNA] ¼ 20.6 þ0.57 log10 [cell volume] [Bachmann et al. 1966]), crustacean (Daphnia pulex, log10 [DNA] ¼ 22.3 þ 0.87 log10
[cell volume] [Beaton and Hebert 1989]), and plant (Arabidopsis thalania, log10 [DNA] ¼ 23.1 þ 1.1 log10
[cell volume] [Jovtchev et al. 2006]). Lines were fitted to data for nonhumans using ordinary least-squaresregression. The human data shown here are the same as those shown in A.
DNA—Cell Size Relationship in Humans
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relationship between nuclear volume and DNAcontent among these cell types (log10 [DNA,pg] ¼ 20.25 þ 0.54 log10 [nuclear volume,m3], P ¼ 3.8 � 10210). For the two remainingcell types in which we had direct measures of celldiameter (cardiomyocytes and megakaryocytes),we also observed a significant positive relation-ship between DNA content and cell volume(log10 [DNA] ¼ 20.71 þ 0.52 log10 [cell vol-ume], P ¼ 1.0 � 10210).
When DNA content and cell volume wereaveraged over all cells for each cell type, we ob-served a similar relationship between meanDNA content and cell volume (Fig. 2) (meanlog10 [DNA] ¼ 21.3 þ 0.72 log10 [cell vol-ume], r2 ¼ 0.82, F1,17 ¼ 86.8, P ¼ 4.3 �1028). Note, however, that the range in DNAcontent of individual cells from these popula-tions was even greater, as one would expect. Atone extreme, the largest megakaryocytes con-tained 448 pg of DNA (see Royere et al. 1988in Table 1). This indicates that these cells havelikely undergone six genome duplication eventsfrom their diploid state.
Moreover, within cell types, the same corre-lations between nuclear DNA content and cellvolume were observed in five of the six cases inwhich mean cell volume varied by a factor of5 or more (megakaryocytes, hepatocytes, lym-phocytes, hepatic parenchymal cells, and amni-on epithelial cells). In these five cases, DNAcontent was positively correlated with cell vol-ume (ordinary least-squares [OLS] regression,all P � 1.6 � 1029). In the case of cardiomyo-cytes, however, we did not observe a statisticallysignificant correlation (P ¼ 0.1).
RELATIONSHIP OF NUCLEAR DNACONTENT TO CELL SIZE: HUMANSCOMPARED WITH OTHER SPECIES
Methodology
To determine whether the relationships betweencell size and DNA content were similar in hu-mans and in other species, we used data fromthe few studies available on intraspecific var-iation in cell size and nuclear DNA content
Amnion epithelial cellCardiomyocyteCytotrophoblastEndocrine pancreatic cellEndothelial cellExocrine pancreatic cellHepatic parenchymal cellHepatocyteLymphocyteLymphocyte polymorphMegakaryocyte
Human cell volume (µ3)
102 103 104
MonocytePrimary spermatocyteSecondary spermatocyteSpermatidSpermatongiumSpermatozoonSyncytiotrophoblastVillous stroma
1
10
100
Hum
an n
ucle
ar D
NA
con
tent
(pg
)
Figure 2. Relationship between mean nuclear DNA content and mean cell size in diploid and polyploid humancells by cell type for data shown in Figure 1. Each point represents mean cell size and DNA content from all cellpopulations of a given cell type.
J.F. Gillooly et al.
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(Heizer 1955; Bachmann et al. 1966; Beaton andHebert 1989; Jovtchev et al. 2006). Studies ofnonhuman cells typically reported nuclear sizeinstead of cell size. To convert nuclear volume tocell volume, we used the formula log10 (cell vol-ume) ¼ 0.88 þ log10 (nuclear volume), whichwe obtained by fitting a linear function to log10-transformed cell and nuclear size measurementsfrom vertebrate (Olmo 1983) and plant (Priceet al. 1973) cells from a diverse assortment ofspecies using ordinary least squares.
Results
The relationship between nuclear DNA contentand cell volume in humans is similar to thatpreviously reported for other species. DNA con-tent is similarly correlated with cell volumein the rat Rattus norvegicus, the frog Pseudacrisobscura, the crustacean Daphnia pulex, andthe plant Arabidopsis thalania (Fig. 1B). Theseresults indicate that concomitant changes in
cell volume and nuclear DNA content mayoccur across very different cell types and spe-cies, although this has only rarely been inves-tigated.
The relationship between nuclear DNA con-tent and cell volume in humans also appearssimilar to the relationships observed betweendiploid genome size and cell volume in plants(Price et al. 1973), vertebrate animals (Olmo1983), and unicellular eukaryotes (Fig. 3A)(Shuter et al. 1983). Interestingly, the range ofnuclear DNA content in human cells is compa-rable to the range of diploid genome sizes acrossall vertebrates (Fig. 3B). However, one must becautious in interpreting similarities and differ-ences among these interspecific relationshipsgiven possible differences in methodology, pos-sible differences in the relationship betweennuclear volume and cell volume among taxaand, in some cases, a fairly limited samplingof species from these groups (e.g., unicellulareukaryotes).
100 200 300 400 5000
Nuclear DNA content (pg)
Human 2–448
Fishes 1–266
Amphibians 2–260
Reptiles 2–11
Mammals 3–17
Birds 2–4
Nuclear DNA content in humans relativeto diploid genome size in vertebrates
Human
Vertebrates
Plants
Unicellular eukaryotes
Cell volume (µ3)
Nuc
lear
DN
A c
onte
nt (
pg)
10–1
1010
3
103 105104102
A B
Figure 3. Relationship between nuclear DNA content and cell volume in humans. (A) Relationship be-tween nuclear DNA content and cell size in diploid and polyploid human cells (blue) in comparison topreviously reported relationships for diploid cells of vertebrates (log10 [DNA] ¼ 21.91 þ 1.07 log10 [cellvolume]), unicellular eukaryotes (log10 [DNA] ¼ 22.81 þ 0.945 log10 [cell volume]), and angiosperms(log10 [DNA] ¼ 21.75 þ 0.912 log10 [cell volume]). Lines were fitted to previously reported relationshipsusing ordinary least-squares regression. The range of unicellular eukaryote cell and genome sizes is truncated toclearly show data for other groups. Note that the human data shown in this figure are the same as those shown inFigure 1A. (B) Range of nuclear DNA content in individual human cells in comparison to ranges of diploidgenome sizes within vertebrate groups. DNA content was rounded to the nearest whole number.
DNA—Cell Size Relationship in Humans
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CONCLUSIONS
Our analysis of previously published data raisesquestions regarding the prevalence of endo-replication in human cells and its relationshipto cell size. It remains to be seen whether therelationship presented here extends across thehundreds of different cell types in the humanbody. Our common assumption of DNA con-stancy across somatic cells in humans and otherspecies has not been adequately investigated.The number of animal species for which wehave measured the amount of DNA in differentcell types of different sizes is restricted to only ahandful of model organisms. Endoreplicationmay be a common and/or functional trait inthe somatic cells of humans and other animals(Lee et al. 2009; Ullah et al. 2009; Anatskaya andVinogradov 2011; De Veylder et al. 2011), as ithas long been viewed in plants. Some have ar-gued that perhaps this process serves to provideextra gene copies as an insurance policy againstDNA damage, or that larger cells simply requiregreater levels of protein synthesis for mainte-nance and production (Lee et al. 2009; DeVeylder et al. 2011). Certainly, in some humancell types (e.g., megakaryocytes), polyploid nu-clei are considered normal and functional. Andin other cell types not considered here (e.g.,muscle cells), multinucleated cells are commonand functional. The relationship between cellsize, nuclear DNA content, and gene expressionwithin and across cell types is only beginning tobe explored (Marguerat and Bahler 2012), butclearly genomes are much more dynamic thanwas previously thought (Parfrey et al. 2008).
We have moved beyond the view that ge-nomes are immutable blueprints. Most of thecells shown in Figure 1 are presumably postmi-totic, but many show quantities of nuclear DNAwell beyond the expected value of about 7 pg.
The notion that human cells may have quitedifferent genetic constitutions depending ontheir size would have profound implicationsfor our understanding of cell structure andfunction.
The data shown here suggest that perhapsthe quantity of nuclear DNA content in humancells is best viewed as a distribution of valuesthat reflects cell size distributions, rather than
as a single value. That is, most cells may be of asimilar size and contain something close to thecharacteristic amount of nuclear DNA, but arelatively small number of cells may be largewith high amounts of DNA.
Changes in the DNA content of somaticcells may occur by endoreplication of individualgenes or the entire genome (i.e., polyploidiza-tion). Often, it is assumed that genome dupli-cation via endoreplication is responsible for anyincreases in nuclear DNA content beyond thetypical diploid quantity. Of the cell types shownhere, it is simply not possible to generalize theextent to which this may be the case becausemost data are averages of cell populations. How-ever, for the two cell types for which we havedata on the individual cells of a cell population(amnion epithelial cells and hepatocytes), acloser look at the data provides some clues(see Table 1). In these data, the increases inDNA content with cell size appear to clusteraround values expected from endopolyploid-ization (e.g., 7, 14, 28, etc.), but there is consid-erably more variation than one would expectsimply based on error. Thus, the data suggestDNA content has increased as a result of somecombination of partial and complete genomeduplication in these two cell types.
However, the results shown here do not ad-dress the question of whether genome size con-trols cell size, or cell size controls genome size.This has long been debated in the literature, andhas yet to be resolved. It is our hope that ourcomparisons of DNA–cell size relationshipswithin species to genome size–cell size relation-ships across species may help point to a moregeneral understanding. We were surprised bythe qualitatively similar relationships betweenthe two. They highlight the fact that we havemuch to learn about the various mechanismsthat lead to increases in cellular DNA content,and how such increases affect gene expression(Therman et al. 1983; Marguerat and Bahler2012). Again, here, broadly comparative ap-proaches may be insightful. In particular, wesuggest that it may be useful to relate relevantstructural features (genome size and cell size)and functional features (gene expression andpolyploidization) to the life span, replicationrate, and metabolic rate of cells.
J.F. Gillooly et al.
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Tabl
e1.
Hum
annucl
ear
DN
Aco
nte
ntan
dce
llvo
lum
edat
a
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Am
nio
nep
ith
elia
lce
ll(s
)3.
040
.7c,
a10
72.1
5.9
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
641
.6c,
a95
8.0
6.0
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
941
.7c,
a10
48.5
6.0
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)4.
042
.4c,
a14
37.0
6.1
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
442
.7c,
a12
27.5
6.2
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
543
.0c,
a89
5.3
6.2
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
543
.0c,
a91
5.8
6.2
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
744
.0c,
a96
8.9
6.4
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
844
.6c,
a10
02.2
6.5
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
944
.6c,
a10
36.5
6.5
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
244
.7c,
a11
47.2
6.5
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)4.
045
.4c,
a14
37.0
6.6
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
645
.5c,
a93
6.7
6.6
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
945
.6c,
a10
36.5
6.6
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
646
.8c,
a12
98.5
6.8
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
047
.2c,
a10
72.1
6.8
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
447
.6c,
a86
5.5
6.9
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
847
.7c,
a10
02.2
6.9
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
847
.7c,
a10
13.5
6.9
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
948
.0c,
a14
05.1
7.0
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)4.
748
.7c,
a16
82.4
7.1
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
748
.8c,
a96
8.9
7.1
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
948
.8c,
a10
36.5
7.1
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
148
.9c,
a11
09.2
7.1
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
849
.1c,
a13
73.6
7.1
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
649
.3c,
a93
6.7
7.1
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
849
.4c,
a10
13.5
7.2
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
749
.9c,
a96
8.9
7.2
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)4.
250
.3c,
a15
03.2
7.3
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
950
.5c,
a10
36.5
7.3
Kli
nge
ran
dSc
hw
arza
cher
1960
Con
tin
ued
DNA—Cell Size Relationship in Humans
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Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Am
nio
nep
ith
elia
lce
ll(s
)3.
150
.6c,
a11
34.2
7.3
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
450
.7c,
a12
13.8
7.3
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
551
.0c,
a90
5.6
7.4
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)4.
052
.0c,
a14
37.0
7.5
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)2.
752
.2c,
a96
8.9
7.6
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
252
.4c,
a11
73.3
7.6
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
553
.0c,
a12
55.3
7.7
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)1.
853
.0c,
a66
0.8
7.7
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)4.
453
.2c,
a15
72.4
7.7
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
053
.5c,
a10
72.1
7.8
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
953
.8c,
a14
05.1
7.8
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
954
.4c,
a13
89.4
7.9
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
554
.9c,
a12
55.3
8.0
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
257
.4c,
a11
73.3
8.3
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
958
.2c,
a14
05.1
8.4
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
662
.2c,
a13
13.2
9.0
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
864
.4c,
a13
73.6
9.3
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
971
.4c,
a13
89.4
10.3
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
880
.8c,
a13
58.1
11.7
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
682
.6c,
a13
13.2
12.0
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)4.
385
.7c,
a15
37.5
12.4
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)3.
887
.5c,
a13
58.1
12.7
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)5.
689
.1c,
a19
92.2
12.9
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)4.
089
.6c,
a14
20.9
13.0
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)5.
390
.1c,
a19
04.3
13.1
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)4.
591
.9c,
a16
08.4
13.3
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)4.
092
.7c,
a14
20.9
13.4
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)4.
393
.9c,
a15
37.5
13.6
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)4.
694
.0c,
a16
63.6
13.6
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)4.
297
.1c,
a15
20.3
14.1
Kli
nge
ran
dSc
hw
arza
cher
1960
Con
tin
ued
J.F. Gillooly et al.
8 Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Am
nio
nep
ith
elia
lce
ll(s
)5.
897
.7c,
a20
60.6
14.2
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)4.
998
.5c,
a17
60.2
14.3
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)5.
598
.7c,
a19
70.0
14.3
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)4.
310
0.5
c,a
1537
.514
.6K
lin
ger
and
Sch
war
zach
er19
60A
mn
ion
epit
hel
ial
cell
(s)
5.8
101.
0c,
a20
60.6
14.6
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)5.
110
2.0
c,a
1820
.414
.8K
lin
ger
and
Sch
war
zach
er19
60A
mn
ion
epit
hel
ial
cell
(s)
6.0
102.
2c,
a21
31.4
14.8
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)4.
610
2.9
c,a
1645
.114
.9K
lin
ger
and
Sch
war
zach
er19
60A
mn
ion
epit
hel
ial
cell
(s)
5.7
104.
5c,
a20
14.8
15.1
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)5.
110
6.7
c,a
1820
.415
.5K
lin
ger
and
Sch
war
zach
er19
60A
mn
ion
epit
hel
ial
cell
(s)
5.3
106.
8c,
a18
83.0
15.5
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)5.
010
7.9
c,a
1780
.015
.6K
lin
ger
and
Sch
war
zach
er19
60A
mn
ion
epit
hel
ial
cell
(s)
4.9
109.
1c,
a17
40.3
15.8
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)6.
911
4.8
c,a
2439
.816
.6K
lin
ger
and
Sch
war
zach
er19
60A
mn
ion
epit
hel
ial
cell
(s)
5.5
115.
7c,
a19
70.0
16.8
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)6.
711
8.7
c,a
2358
.917
.2K
lin
ger
and
Sch
war
zach
er19
60A
mn
ion
epit
hel
ial
cell
(s)
7.2
120.
2c,
a25
52.4
17.4
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)5.
912
4.0
c,a
2107
.418
.0K
lin
ger
and
Sch
war
zach
er19
60A
mn
ion
epit
hel
ial
cell
(s)
5.7
126.
7c,
a20
14.8
18.4
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)5.
913
2.7
c,a
2107
.419
.2K
lin
ger
and
Sch
war
zach
er19
60A
mn
ion
epit
hel
ial
cell
(s)
7.0
142.
5c,
a24
67.4
20.7
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)7.
814
9.4
c,a
2761
.621
.7K
lin
ger
and
Sch
war
zach
er19
60A
mn
ion
epit
hel
ial
cell
(s)
7.6
156.
3c,
a27
00.3
22.7
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)8.
717
1.5
c,a
3056
.524
.9K
lin
ger
and
Sch
war
zach
er19
60A
mn
ion
epit
hel
ial
cell
(s)
7.8
185.
4c,
a27
61.6
26.9
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)8.
318
5.5
c,a
2921
.726
.9K
lin
ger
and
Sch
war
zach
er19
60A
mn
ion
epit
hel
ial
cell
(s)
8.9
187.
9c,
a31
26.2
27.2
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)8.
319
2.0
c,a
2921
.727
.8K
lin
ger
and
Sch
war
zach
er19
60A
mn
ion
epit
hel
ial
cell
(s)
7.3
209.
8c,
a25
81.4
30.4
Kli
nge
ran
dSc
hw
arza
cher
1960
Am
nio
nep
ith
elia
lce
ll(s
)6.
822
4.3
c,a
2412
.532
.5K
lin
ger
and
Sch
war
zach
er19
60
Con
tin
ued
DNA—Cell Size Relationship in Humans
Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091 9
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Car
dio
myo
cyte
(m)
1740
1.0
20.0
v,p
g56
46.7
20.0
Vli
egen
etal
.19
90C
ard
iom
yocy
te(m
)20
794.
021
.2v,
pg
6339
.921
.2V
lieg
enet
al.
1990
Car
dio
myo
cyte
(m)
43.8
3.6
cc,
p14
63.2
12.6
Wo
hls
chla
eger
etal
.20
10C
ard
iom
yocy
te(m
)60
.74.
1cc
,p
2013
.514
.4W
oh
lsch
laeg
eret
al.
2010
Car
dio
myo
cyte
(m)
55.5
4.5
cc,
p18
44.6
15.8
Wo
hls
chla
eger
etal
.20
10C
ard
iom
yocy
te(m
)13
5.5
4.6
cc,
p44
03.8
16.1
Wo
hls
chla
eger
etal
.20
10C
ard
iom
yocy
te(m
)15
6.4
5.1
cc,
p50
64.6
17.9
Wo
hls
chla
eger
etal
.20
10C
ard
iom
yocy
te(m
)12
7.0
5.8
cc,
p41
32.1
20.2
Wo
hls
chla
eger
etal
.20
10C
ard
iom
yocy
te(m
)65
.15.
9cc
,p
2154
.920
.7W
oh
lsch
laeg
eret
al.
2010
Car
dio
myo
cyte
(m)
68.1
5.9
cc,
p22
52.3
20.7
Wo
hls
chla
eger
etal
.20
10C
ard
iom
yocy
te(m
)77
.35.
9cc
,p
2546
.120
.7W
oh
lsch
laeg
eret
al.
2010
Car
dio
myo
cyte
(m)
71.1
6.0
cc,
p23
46.7
21.0
Wo
hls
chla
eger
etal
.20
10C
ard
iom
yocy
te(m
)11
9.2
6.6
cc,
p38
85.0
23.1
Wo
hls
chla
eger
etal
.20
10C
ard
iom
yocy
te(m
)60
.36.
8cc
,p
2000
.223
.8W
oh
lsch
laeg
eret
al.
2010
Car
dio
myo
cyte
(m)
84.1
7.2
cc,
p27
65.2
25.2
Wo
hls
chla
eger
etal
.20
10C
ard
iom
yocy
te(m
)10
3.7
7.9
cc,
p33
92.5
27.6
Wo
hls
chla
eger
etal
.20
10C
ard
iom
yocy
te(m
)10
4.0
7.9
cc,
p34
02.4
27.7
Wo
hls
chla
eger
etal
.20
10C
ard
iom
yocy
te(m
)11
2.1
8.4
cc,
p36
58.9
29.4
Wo
hls
chla
eger
etal
.20
10C
ard
iom
yocy
te(m
)99
.88.
5cc
,p
3266
.229
.8W
oh
lsch
laeg
eret
al.
2010
Car
dio
myo
cyte
(m)
161.
58.
6cc
,p
5224
.030
.1W
oh
lsch
laeg
eret
al.
2010
Car
dio
myo
cyte
(m)
152.
58.
8cc
,p
4939
.630
.8W
oh
lsch
laeg
eret
al.
2010
Car
dio
myo
cyte
(m)
105.
19.
0cc
,p
3436
.831
.5W
oh
lsch
laeg
eret
al.
2010
Car
dio
myo
cyte
(m)
175.
49.
5cc
,p
5660
.733
.3W
oh
lsch
laeg
eret
al.
2010
Car
dio
myo
cyte
(m)
97.6
11.9
cc,
p31
98.5
41.7
Wo
hls
chla
eger
etal
.20
10C
ard
iom
yocy
te(m
)12
9.9
13.6
cc,
p42
25.6
47.6
Wo
hls
chla
eger
etal
.20
10C
yto
tro
ph
ob
last
(m)
133.
01.
9cc
,p
1061
.56.
7G
alto
n19
62C
yto
tro
ph
ob
last
(m)
109.
02.
1cc
,p
932.
77.
4G
alto
n19
62C
yto
tro
ph
ob
last
(m)
125.
02.
4cc
,p
1019
.68.
3G
alto
n19
62C
yto
tro
ph
ob
last
(m)
128.
02.
4cc
,p
1035
.48.
5G
alto
n19
62E
nd
ocr
ine
pan
crea
tic
cell
(m)
98.4
2.0
cc,
p87
2.7
7.0
Eh
rie
and
Swar
tz19
74
Con
tin
ued
J.F. Gillooly et al.
10 Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
En
do
crin
ep
ancr
eati
cce
ll(m
)11
6.5
2.0
cc,
p97
3.9
7.0
Eh
rie
and
Swar
tz19
74E
nd
ocr
ine
pan
crea
tic
cell
(m)
119.
02.
0cc
,p
987.
57.
0E
hri
ean
dSw
artz
1974
En
do
crin
ep
ancr
eati
cce
ll(m
)20
5.4
4.0
cc,
p14
08.0
14.0
Eh
rie
and
Swar
tz19
74E
nd
ocr
ine
pan
crea
tic
cell
(m)
217.
84.
0cc
,p
1462
.714
.0E
hri
ean
dSw
artz
1974
En
do
crin
ep
ancr
eati
cce
ll(m
)22
1.8
4.0
cc,
p14
80.1
14.0
Eh
rie
and
Swar
tz19
74E
nd
ocr
ine
pan
crea
tic
cell
(m)
435.
58.
0cc
,p
2294
.928
.0E
hri
ean
dSw
artz
1974
En
do
crin
ep
ancr
eati
cce
ll(m
)44
7.3
8.0
cc,
p23
35.1
28.0
Eh
rie
and
Swar
tz19
74E
nd
ocr
ine
pan
crea
tic
cell
(m)
453.
18.
0cc
,p
2354
.828
.0E
hri
ean
dSw
artz
1974
En
do
thel
ial
cell
(m)
36.5
4.2
cc,
pg
1225
.44.
2D
orm
anet
al.
1990
En
do
thel
ial
cell
(m)
42.4
4.2
cc,
pg
1418
.24.
2D
orm
anet
al.
1990
En
do
thel
ial
cell
(m)
41.1
4.6
cc,
pg
1375
.84.
6D
orm
anet
al.
1990
En
do
thel
ial
cell
(m)
63.5
5.3
cc,
pg
2102
.65.
3D
orm
anet
al.
1990
Exo
crin
ep
ancr
eati
cce
ll(m
)10
0.1
2.0
cc,
p88
2.5
7.0
Eh
rie
and
Swar
tz19
74E
xocr
ine
pan
crea
tic
cell
(m)
107.
72.
0cc
,p
925.
57.
0E
hri
ean
dSw
artz
1974
Exo
crin
ep
ancr
eati
cce
ll(m
)11
2.8
2.0
cc,
p95
3.7
7.0
Eh
rie
and
Swar
tz19
74H
epat
icp
aren
chym
alce
ll(m
)34
.71.
8cc
,a
1124
.64.
8Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
34.4
2.0
cc,
a11
18.2
5.4
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)45
.12.
1cc
,a
1333
.55.
6Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
34.8
2.3
cc,
a11
26.7
6.1
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)26
.12.
3cc
,a
934.
66.
2Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
38.9
2.3
cc,
a12
11.3
6.2
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)37
.92.
3cc
,a
1191
.06.
3Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
36.6
2.4
cc,
a11
64.2
6.5
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)33
.92.
5cc
,a
1107
.76.
6Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
33.2
2.5
cc,
a10
92.8
6.6
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)32
.92.
5cc
,a
1086
.36.
7Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
41.7
2.5
cc,
a12
67.3
6.7
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)27
.62.
5cc
,a
969.
16.
8Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
51.2
2.5
cc,
a14
48.1
6.8
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)35
.22.
6cc
,a
1135
.16.
8Sw
artz
1956
Con
tin
ued
DNA—Cell Size Relationship in Humans
Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091 11
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Hep
atic
par
ench
ymal
cell
(m)
41.8
2.6
cc,
a12
69.2
6.9
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)29
.52.
6cc
,a
1012
.07.
0Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
29.4
2.6
cc,
a10
09.7
7.1
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)40
.42.
6cc
,a
1241
.57.
1Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
47.4
2.7
cc,
a13
77.3
7.2
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)39
.92.
7cc
,a
1231
.47.
2Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
32.5
2.7
cc,
a10
77.7
7.2
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)55
.32.
7cc
,a
1522
.57.
2Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
30.6
2.7
cc,
a10
36.3
7.4
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)36
.92.
8cc
,a
1170
.57.
4Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
22.3
2.8
cc,
a84
3.7
7.5
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)36
.42.
8cc
,a
1160
.17.
5Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
28.1
2.8
cc,
a98
0.5
7.5
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)33
.92.
8cc
,a
1107
.77.
5Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
54.5
2.8
cc,
a15
08.1
7.5
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)38
.62.
9cc
,a
1205
.27.
7Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
49.3
2.9
cc,
a14
13.0
7.7
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)36
.12.
9cc
,a
1153
.97.
7Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
44.6
2.9
cc,
a13
23.9
7.8
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)33
.32.
9cc
,a
1094
.97.
9Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
31.4
3.1
cc,
a10
53.9
8.4
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)36
.03.
2cc
,a
1151
.88.
5Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
71.4
3.4
cc,
a17
97.6
9.1
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)98
.73.
8cc
,a
2218
.610
.1Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
78.1
4.4
cc,
a19
05.5
11.8
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)44
.54.
8cc
,a
1321
.912
.8Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
65.8
4.8
cc,
a17
04.6
12.9
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)71
.64.
9cc
,a
1800
.813
.1Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
65.0
5.0
cc,
a16
91.1
13.3
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)69
.05.
1cc
,a
1758
.113
.7Sw
artz
1956
Con
tin
ued
J.F. Gillooly et al.
12 Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Hep
atic
par
ench
ymal
cell
(m)
64.1
5.2
cc,
a16
75.9
13.9
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)72
.95.
3cc
,a
1822
.014
.1Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
70.7
5.3
cc,
a17
86.1
14.1
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)91
.75.
3cc
,a
2115
.014
.2Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
87.0
5.4
cc,
a20
43.9
14.5
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)78
.45.
4cc
,a
1910
.214
.5Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
147.
25.
4cc
,a
2876
.914
.5Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
70.3
5.5
cc,
a17
79.5
14.7
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)81
.65.
6cc
,a
1960
.615
.0Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
72.9
5.7
cc,
a18
22.0
15.2
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)73
.66.
1cc
,a
1833
.416
.4Sw
artz
1956
Hep
atic
par
ench
ymal
cell
(m)
65.3
6.4
cc,
a16
96.2
17.2
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)17
7.5
7.4
cc,
a32
49.1
19.9
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)10
6.7
9.3
cc,
a23
33.9
24.9
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)12
9.4
9.8
cc,
a26
45.7
26.2
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)14
1.9
9.8
cc,
a28
09.1
26.4
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)13
6.8
9.9
cc,
a27
43.1
26.5
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)14
2.1
11.6
cc,
a28
11.7
31.1
Swar
tz19
56H
epat
icp
aren
chym
alce
ll(m
)14
4.7
13.6
cc,
a28
45.0
36.6
Swar
tz19
56H
epat
ocy
te(s
)35
.718
.3cc
,a
1200
.98.
7M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
46.1
20.7
cc,
a15
37.3
9.9
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)41
.921
.0cc
,a
1400
.810
.0M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
59.3
22.4
cc,
a19
65.6
10.7
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)39
.022
.6cc
,a
1306
.210
.8M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
42.2
23.6
cc,
a14
11.4
11.3
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)73
.423
.8cc
,a
2422
.611
.4M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
64.7
24.4
cc,
a21
42.5
11.6
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)46
.725
.6cc
,a
1558
.212
.2M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
68.0
25.8
cc,
a22
46.3
12.3
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)64
.126
.7cc
,a
2121
.712
.7M
eek
and
Har
bis
on
1967 Con
tin
ued
DNA—Cell Size Relationship in Humans
Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091 13
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Hep
ato
cyte
(s)
89.5
29.5
cc,
a29
39.3
14.1
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)54
.130
.6cc
,a
1798
.814
.6M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
73.1
32.1
cc,
a24
12.3
15.3
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)89
.532
.5cc
,a
2939
.315
.5M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
64.1
33.3
cc,
a21
21.7
15.9
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)73
.134
.4cc
,a
2412
.316
.4M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
96.3
35.8
cc,
a31
55.6
17.1
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)53
.835
.9cc
,a
1788
.317
.1M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
68.0
38.3
cc,
a22
46.3
18.2
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)84
.139
.1cc
,a
2763
.918
.6M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
78.9
39.7
cc,
a25
98.6
18.9
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)63
.839
.9cc
,a
2111
.319
.0M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
68.3
40.3
cc,
a22
56.7
19.2
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)72
.840
.9cc
,a
2401
.919
.5M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
113.
441
.9cc
,a
3699
.819
.9M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
126.
243
.3cc
,a
4109
.220
.6M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
90.2
43.7
cc,
a29
59.9
20.8
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)10
6.9
44.1
cc,
a34
94.7
21.0
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)10
1.1
45.7
cc,
a33
09.8
21.8
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)96
.047
.0cc
,a
3145
.322
.4M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
90.2
47.6
cc,
a29
59.9
22.7
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)14
0.1
48.0
cc,
a45
48.1
22.9
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)95
.348
.7cc
,a
3124
.723
.2M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
107.
249
.1cc
,a
3504
.923
.4M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
120.
849
.8cc
,a
3935
.323
.7M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
90.2
50.3
cc,
a29
59.9
23.9
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)14
0.1
50.9
cc,
a45
48.1
24.3
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)10
1.4
52.7
cc,
a33
20.1
25.1
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)11
3.7
53.4
cc,
a37
10.1
25.4
Mee
kan
dH
arb
iso
n19
67H
epat
ocy
te(s
)15
6.2
56.6
cc,
a50
57.1
27.0
Mee
kan
dH
arb
iso
n19
67 Con
tin
ued
J.F. Gillooly et al.
14 Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Hep
ato
cyte
(s)
113.
058
.7cc
,a
3689
.527
.9M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
120.
863
.0cc
,a
3935
.330
.0M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
140.
779
.6cc
,a
4568
.437
.9M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
170.
491
.6cc
,a
5503
.943
.6M
eek
and
Har
bis
on
1967
Hep
ato
cyte
(s)
177.
597
.9cc
,a
5726
.946
.6M
eek
and
Har
bis
on
1967
Lym
ph
ocy
te(m
)17
.94.
0cc
,p
g61
1.8
4.0
Do
rman
etal
.19
90Ly
mp
ho
cyte
(m)
19.8
4.0
cc,
pg
675.
04.
0D
orm
anet
al.
1990
Lym
ph
ocy
te(m
)18
.54.
2cc
,p
g63
1.8
4.2
Do
rman
etal
.19
90Ly
mp
ho
cyte
(m)
37.6
5.8
cc,
pg
1261
.45.
8D
orm
anet
al.
1990
Lym
ph
ocy
te(s
)18
.117
.2cc
,a
618.
76.
3B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
te(s
)19
.917
.4cc
,a
679.
46.
3B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
te(s
)18
.617
.7cc
,a
633.
96.
4B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
te(s
)23
.518
.5cc
,a
796.
56.
7B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
te(s
)21
.218
.6cc
,a
721.
06.
7B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
te(s
)22
.818
.8cc
,a
773.
86.
8B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
te(s
)21
.418
.9cc
,a
728.
56.
9B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
te(s
)22
.918
.9cc
,a
777.
66.
9B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
te(s
)24
.919
.0cc
,a
845.
46.
9B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
te(s
)22
.419
.0cc
,a
762.
56.
9B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
te(s
)24
.319
.0cc
,a
822.
96.
9B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
te(s
)23
.019
.1cc
,a
781.
46.
9B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
te(s
)22
.819
.3cc
,a
773.
87.
0B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
te(s
)23
.819
.7cc
,a
807.
87.
1B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
te(s
)28
.019
.9cc
,a
946.
97.
2B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
te(s
)7.
42.
7n
,a
1437
.34.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)5.
53.
0n
,a
793.
24.
5P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
03.
0n
,a
1293
.64.
6P
etra
kis
1953
Lym
ph
ocy
te(s
)5.
23.
0n
,a
712.
04.
6P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
73.
0n
,a
1562
.64.
6P
etra
kis
1953
Con
tin
ued
DNA—Cell Size Relationship in Humans
Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091 15
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Lym
ph
ocy
te(s
)7.
53.
1n
,a
1462
.04.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
03.
1n
,a
1278
.24.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
13.
1n
,a
1340
.84.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)5.
63.
2n
,a
824.
14.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
03.
3n
,a
1293
.65.
0P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
03.
3n
,a
1666
.45.
0P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
53.
3n
,a
1462
.05.
0P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
23.
3n
,a
1746
.75.
0P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
83.
4n
,a
1597
.15.
2P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
53.
4n
,a
1118
.15.
2P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
93.
5n
,a
1239
.85.
3P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
73.
5n
,a
1562
.65.
3P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
03.
5n
,a
1666
.45.
3P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
53.
5n
,a
1495
.05.
3P
etra
kis
1953
Lym
ph
ocy
te(s
)5.
03.
5n
,a
680.
85.
4P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
03.
5n
,a
953.
55.
4P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
43.
5n
,a
1453
.65.
4P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
53.
6n
,a
1118
.15.
4P
etra
kis
1953
Lym
ph
ocy
te(s
)5.
03.
6n
,a
671.
65.
5P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
23.
6n
,a
1737
.65.
5P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
03.
6n
,a
950.
45.
5P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
03.
6n
,a
1293
.65.
5P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
53.
7n
,a
1462
.05.
6P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
83.
7n
,a
1579
.65.
6P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
03.
7n
,a
1286
.05.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)5.
03.
7n
,a
671.
65.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
23.
8n
,a
1028
.45.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
03.
8n
,a
956.
95.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
03.
8n
,a
953.
55.
8P
etra
kis
1953
Con
tin
ued
J.F. Gillooly et al.
16 Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Lym
ph
ocy
te(s
)6.
63.
8n
,a
1142
.75.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
13.
8n
,a
1317
.15.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
53.
8n
,a
1113
.75.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
03.
9n
,a
953.
55.
9P
etra
kis
1953
Lym
ph
ocy
te(s
)10
.13.
9n
,a
2627
.95.
9P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
83.
9n
,a
1217
.05.
9P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
13.
9n
,a
980.
35.
9P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
33.
9n
,a
1035
.56.
0P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
23.
9n
,a
1364
.76.
0P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
53.
9n
,a
1495
.06.
0P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
03.
9n
,a
1299
.06.
0P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
24.
0n
,a
1370
.26.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
04.
0n
,a
1657
.96.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
14.
0n
,a
1711
.16.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)10
.04.
0n
,a
2594
.96.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
54.
0n
,a
1113
.76.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
84.
0n
,a
1209
.76.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)5.
64.
0n
,a
836.
66.
2P
etra
kis
1953
Lym
ph
ocy
te(s
)9.
04.
1n
,a
2115
.16.
2P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
04.
1n
,a
1278
.26.
2P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
84.
1n
,a
1217
.06.
3P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
04.
2n
,a
950.
46.
4P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
54.
2n
,a
1113
.76.
4P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
04.
2n
,a
1286
.06.
4P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
64.
2n
,a
1512
.16.
4P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
64.
2n
,a
1142
.76.
4P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
04.
3n
,a
1299
.06.
5P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
64.
3n
,a
1503
.66.
5P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
04.
3n
,a
950.
46.
5P
etra
kis
1953
Con
tin
ued
DNA—Cell Size Relationship in Humans
Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091 17
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Lym
ph
ocy
te(s
)6.
94.
3n
,a
1252
.56.
5P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
34.
3n
,a
1818
.26.
6P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
04.
3n
,a
966.
96.
6P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
64.
3n
,a
1150
.16.
6P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
04.
3n
,a
1286
.06.
6P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
14.
3n
,a
1325
.16.
6P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
24.
3n
,a
1356
.66.
6P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
54.
4n
,a
1485
.06.
6P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
54.
4n
,a
1106
.46.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
64.
4n
,a
1503
.66.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
44.
4n
,a
1435
.46.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
34.
4n
,a
1056
.66.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
64.
4n
,a
1135
.66.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
04.
4n
,a
966.
96.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
04.
4n
,a
1299
.06.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
04.
5n
,a
1666
.46.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)9.
04.
5n
,a
2095
.06.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)5.
54.
5n
,a
811.
76.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
24.
5n
,a
1014
.66.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
84.
5n
,a
1221
.96.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
64.
5n
,a
1503
.66.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)5.
84.
5n
,a
897.
36.
9P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
04.
6n
,a
1666
.46.
9P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
04.
6n
,a
1299
.06.
9P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
44.
6n
,a
1453
.67.
0P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
84.
6n
,a
1588
.37.
0P
etra
kis
1953
Lym
ph
ocy
te(s
)9.
04.
6n
,a
2105
.07.
0P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
94.
6n
,a
2075
.57.
0P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
14.
6n
,a
1000
.97.
0P
etra
kis
1953
Con
tin
ued
J.F. Gillooly et al.
18 Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Lym
ph
ocy
te(s
)7.
04.
6n
,a
1286
.07.
0P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
34.
6n
,a
1035
.57.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
64.
6n
,a
1150
.17.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
64.
6n
,a
1503
.67.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
94.
7n
,a
1239
.87.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
84.
7n
,a
1605
.57.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)9.
04.
7n
,a
2085
.57.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
84.
7n
,a
1595
.57.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
24.
7n
,a
1007
.97.
2P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
54.
7n
,a
1478
.57.
2P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
04.
7n
,a
1666
.47.
2P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
54.
7n
,a
1132
.67.
2P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
84.
7n
,a
1209
.77.
2P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
04.
7n
,a
1286
.07.
2P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
44.
7n
,a
1437
.37.
2P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
04.
7n
,a
1307
.07.
2P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
44.
8n
,a
1429
.37.
3P
etra
kis
1953
Lym
ph
ocy
te(s
)5.
84.
8n
,a
894.
07.
3P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
04.
8n
,a
1675
.47.
3P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
64.
8n
,a
1520
.37.
3P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
04.
8n
,a
1275
.97.
4P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
04.
9n
,a
1291
.57.
4P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
24.
9n
,a
1346
.37.
4P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
44.
9n
,a
1437
.37.
4P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
54.
9n
,a
1470
.47.
4P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
74.
9n
,a
1194
.47.
5P
etra
kis
1953
Lym
ph
ocy
te(s
)9.
04.
9n
,a
2095
.07.
5P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
04.
9n
,a
953.
57.
5P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
04.
9n
,a
1666
.47.
5P
etra
kis
1953
Con
tin
ued
DNA—Cell Size Relationship in Humans
Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091 19
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Lym
ph
ocy
te(s
)7.
15.
0n
,a
1314
.97.
6P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
15.
0n
,a
1719
.87.
6P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
05.
0n
,a
1666
.47.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
95.
1n
,a
1614
.37.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
05.
1n
,a
966.
97.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
05.
1n
,a
1283
.67.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
05.
1n
,a
1293
.67.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
45.
1n
,a
1846
.47.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
75.
1n
,a
1571
.37.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
65.
1n
,a
1150
.17.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
05.
1n
,a
1657
.97.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
15.
1n
,a
1711
.17.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
85.
2n
,a
1224
.77.
9P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
55.
2n
,a
1495
.07.
9P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
05.
2n
,a
1684
.47.
9P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
05.
2n
,a
970.
67.
9P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
05.
3n
,a
1691
.88.
0P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
05.
3n
,a
940.
28.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
05.
3n
,a
1684
.48.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
15.
3n
,a
1719
.88.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)5.
65.
4n
,a
843.
08.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
05.
4n
,a
1307
.08.
2P
etra
kis
1953
Lym
ph
ocy
te(s
)9.
85.
4n
,a
2486
.48.
2P
etra
kis
1953
Lym
ph
ocy
te(s
)9.
05.
4n
,a
2085
.58.
2P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
05.
4n
,a
1674
.18.
2P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
05.
5n
,a
1666
.48.
3P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
15.
5n
,a
1314
.98.
4P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
05.
5n
,a
1691
.88.
4P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
35.
5n
,a
1388
.88.
4P
etra
kis
1953
Con
tin
ued
J.F. Gillooly et al.
20 Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Lym
ph
ocy
te(s
)8.
65.
6n
,a
1912
.58.
5P
etra
kis
1953
Lym
ph
ocy
te(s
)9.
05.
6n
,a
2095
.08.
5P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
55.
6n
,a
1487
.08.
5P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
25.
6n
,a
1356
.68.
6P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
15.
7n
,a
977.
28.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
85.
7n
,a
1229
.68.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
65.
7n
,a
1512
.18.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
45.
7n
,a
1856
.28.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)5.
65.
8n
,a
824.
18.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
15.
8n
,a
1309
.18.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)9.
25.
9n
,a
2215
.09.
0P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
36.
0n
,a
1049
.49.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
86.
0n
,a
1597
.19.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
06.
0n
,a
1675
.49.
1P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
06.
1n
,a
1684
.49.
3P
etra
kis
1953
Lym
ph
ocy
te(s
)9.
06.
1n
,a
2095
.09.
4P
etra
kis
1953
Lym
ph
ocy
te(s
)7.
06.
2n
,a
1307
.09.
5P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
96.
3n
,a
1267
.89.
5P
etra
kis
1953
Lym
ph
ocy
te(s
)9.
26.
3n
,a
2205
.29.
6P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
06.
4n
,a
1684
.49.
7P
etra
kis
1953
Lym
ph
ocy
te(s
)9.
26.
4n
,a
2205
.29.
8P
etra
kis
1953
Lym
ph
ocy
te(s
)9.
26.
5n
,a
2215
.09.
9P
etra
kis
1953
Lym
ph
ocy
te(s
)9.
16.
6n
,a
2134
.810
.0P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
06.
6n
,a
1666
.410
.1P
etra
kis
1953
Lym
ph
ocy
te(s
)8.
06.
7n
,a
1666
.410
.2P
etra
kis
1953
Lym
ph
ocy
te(s
)6.
66.
8n
,a
1162
.110
.4P
etra
kis
1953
Lym
ph
ocy
te(s
)9.
06.
9n
,a
2124
.710
.5P
etra
kis
1953
Lym
ph
ocy
te(s
)9.
16.
9n
,a
2164
.810
.5P
etra
kis
1953
Lym
ph
ocy
te(s
)9.
17.
0n
,a
2144
.510
.7P
etra
kis
1953
Con
tin
ued
DNA—Cell Size Relationship in Humans
Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091 21
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Lym
ph
ocy
te(s
)11
.17.
4n
,a
3158
.411
.3P
etra
kis
1953
Lym
ph
ocy
te(s
)12
.18.
5n
,a
3762
.413
.0P
etra
kis
1953
Lym
ph
ocy
tep
oly
mo
rph
(m)
25.1
17.9
cc,
a84
9.2
6.5
Bed
ian
dG
old
stei
n19
76Ly
mp
ho
cyte
po
lym
orp
h(m
)22
.818
.0cc
,a
773.
86.
5B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
tep
oly
mo
rph
(m)
21.4
18.0
cc,
a72
8.5
6.5
Bed
ian
dG
old
stei
n19
76Ly
mp
ho
cyte
po
lym
orp
h(m
)20
.018
.1cc
,a
683.
16.
6B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
tep
oly
mo
rph
(m)
24.4
18.2
cc,
a82
6.6
6.6
Bed
ian
dG
old
stei
n19
76Ly
mp
ho
cyte
po
lym
orp
h(m
)21
.318
.2cc
,a
724.
86.
6B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
tep
oly
mo
rph
(m)
27.1
18.2
cc,
a91
6.9
6.6
Bed
ian
dG
old
stei
n19
76Ly
mp
ho
cyte
po
lym
orp
h(m
)22
.918
.5cc
,a
777.
66.
7B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
tep
oly
mo
rph
(m)
25.7
18.9
cc,
a87
1.8
6.9
Bed
ian
dG
old
stei
n19
76Ly
mp
ho
cyte
po
lym
orp
h(m
)28
.019
.0cc
,a
946.
96.
9B
edi
and
Go
ldst
ein
1976
Lym
ph
ocy
tep
oly
mo
rph
(m)
26.2
19.0
cc,
a88
6.8
6.9
Bed
ian
dG
old
stei
n19
76M
egak
aryo
cyte
(m)
15.5
3.9
d,
p19
49.8
13.7
Ish
ibas
hi
etal
.19
86M
egak
aryo
cyte
(m)
15.3
4.4
d,
p18
75.3
15.4
Ish
ibas
hi
etal
.19
86M
egak
aryo
cyte
(m)
16.1
4.4
d,
p23
52.1
15.4
Ish
ibas
hi
etal
.19
86M
egak
aryo
cyte
(m)
16.5
4.4
d,
p21
85.1
15.4
Ish
ibas
hi
etal
.19
86M
egak
aryo
cyte
(m)
16.3
4.6
d,
p22
67.6
16.1
Ish
ibas
hi
etal
.19
86M
egak
aryo
cyte
(m)
16.8
5.3
d,
p24
82.7
18.6
Ish
ibas
hi
etal
.19
86M
egak
aryo
cyte
(m)
21.3
7.0
d,
p50
59.8
24.5
Ish
ibas
hi
etal
.19
86M
egak
aryo
cyte
(m)
20.8
7.3
d,
p47
11.8
25.6
Ish
ibas
hi
etal
.19
86M
egak
aryo
cyte
(m)
22.4
8.6
d,
p58
84.9
30.1
Ish
ibas
hi
etal
.19
86M
egak
aryo
cyte
(m)
27.5
10.4
d,
p10
889.
236
.4Is
hib
ash
iet
al.
1986
Meg
akar
yocy
te(m
)27
.610
.6d
,p
1100
8.4
37.1
Ish
ibas
hi
etal
.19
86M
egak
aryo
cyte
(m)
30.2
10.9
d,
p14
421.
838
.2Is
hib
ash
iet
al.
1986
Meg
akar
yocy
te(m
)30
.612
.6d
,p
1500
2.5
44.1
Ish
ibas
hi
etal
.19
86M
egak
aryo
cyte
(m)
32.3
12.8
d,
p17
644.
444
.8Is
hib
ash
iet
al.
1986
Meg
akar
yocy
te(m
)33
.314
.1d
,p
1933
4.4
49.4
Ish
ibas
hi
etal
.19
86M
egak
aryo
cyte
(m)
33.8
14.5
d,
p20
218.
550
.8Is
hib
ash
iet
al.
1986
Meg
akar
yocy
te(m
)33
.816
.1d
,p
2021
8.5
56.4
Ish
ibas
hi
etal
.19
86
Con
tin
ued
J.F. Gillooly et al.
22 Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Meg
akar
yocy
te(m
)34
.216
.8d
,p
2094
4.8
58.8
Ish
ibas
hi
etal
.19
86M
egak
aryo
cyte
(m)
34.1
16.9
d,
p20
761.
659
.2Is
hib
ash
iet
al.
1986
Meg
akar
yocy
te(m
)34
.417
.2d
,p
2131
4.4
60.2
Ish
ibas
hi
etal
.19
86M
egak
aryo
cyte
(m)
34.4
17.2
d,
p21
314.
460
.2Is
hib
ash
iet
al.
1986
Meg
akar
yocy
te(m
)35
.017
.7d
,p
2244
9.3
62.0
Ish
ibas
hi
etal
.19
86M
egak
aryo
cyte
(m)
36.0
17.7
d,
p24
429.
062
.0Is
hib
ash
iet
al.
1986
Meg
akar
yocy
te(m
)35
.418
.6d
,p
2322
7.8
65.1
Ish
ibas
hi
etal
.19
86M
egak
aryo
cyte
(m)
35.4
19.3
d,
p23
227.
867
.6Is
hib
ash
iet
al.
1986
Meg
akar
yocy
te(m
)36
.419
.6d
,p
2525
2.4
68.6
Ish
ibas
hi
etal
.19
86M
on
ocy
te(s
)21
.419
.3cc
,a
728.
57.
0B
edi
and
Go
ldst
ein
1976
Mo
no
cyte
(s)
32.1
19.9
cc,
a10
81.8
7.2
Bed
ian
dG
old
stei
n19
76M
on
ocy
te(s
)26
.320
.0cc
,a
890.
67.
3B
edi
and
Go
ldst
ein
1976
Mo
no
cyte
(s)
30.8
20.1
cc,
a10
36.9
7.3
Bed
ian
dG
old
stei
n19
76M
on
ocy
te(s
)30
.620
.2cc
,a
1033
.17.
3B
edi
and
Go
ldst
ein
1976
Mo
no
cyte
(s)
30.9
20.3
cc,
a10
40.7
7.4
Bed
ian
dG
old
stei
n19
76M
on
ocy
te(s
)46
.221
.5cc
,a
1543
.37.
8B
edi
and
Go
ldst
ein
1976
Pri
mar
ysp
erm
ato
cyte
(m)
37.6
2.6
cc,
a11
84.8
7.6
Leu
chte
nb
erge
ret
al.
1956
Pri
mar
ysp
erm
ato
cyte
(m)
28.1
3.3
cc,
a98
0.5
9.7
Leu
chte
nb
erge
ret
al.
1956
Pri
mar
ysp
erm
ato
cyte
(m)
59.3
3.3
cc,
a15
93.2
9.7
Leu
chte
nb
erge
ret
al.
1956
Pri
mar
ysp
erm
ato
cyte
(m)
42.1
3.9
cc,
a12
75.2
11.3
Leu
chte
nb
erge
ret
al.
1956
Pri
mar
ysp
erm
ato
cyte
(m)
69.4
4.6
cc,
a17
64.7
13.3
Leu
chte
nb
erge
ret
al.
1956
Pri
mar
ysp
erm
ato
cyte
(m)
45.1
4.6
cc,
a13
33.5
13.4
Leu
chte
nb
erge
ret
al.
1956
Pri
mar
ysp
erm
ato
cyte
(m)
48.6
5.1
cc,
a13
99.9
14.7
Leu
chte
nb
erge
ret
al.
1956
Pri
mar
ysp
erm
ato
cyte
(m)
48.2
5.4
cc,
a13
92.4
15.8
Leu
chte
nb
erge
ret
al.
1956
Seco
nd
ary
sper
mat
ocy
te(m
)14
.21.
5cc
,a
629.
24.
2L
euch
ten
ber
ger
etal
.19
56Se
con
dar
ysp
erm
ato
cyte
(m)
13.1
1.9
cc,
a59
7.0
5.4
Leu
chte
nb
erge
ret
al.
1956
Seco
nd
ary
sper
mat
ocy
te(m
)17
.61.
9cc
,a
723.
45.
5L
euch
ten
ber
ger
etal
.19
56Se
con
dar
ysp
erm
ato
cyte
(m)
19.9
2.3
cc,
a78
3.5
6.8
Leu
chte
nb
erge
ret
al.
1956
Seco
nd
ary
sper
mat
ocy
te(m
)33
.42.
6cc
,a
1097
.07.
7L
euch
ten
ber
ger
etal
.19
56
Con
tin
ued
DNA—Cell Size Relationship in Humans
Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091 23
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Seco
nd
ary
sper
mat
ocy
te(m
)25
.92.
7cc
,a
929.
98.
0L
euch
ten
ber
ger
etal
.19
56Se
con
dar
ysp
erm
ato
cyte
(m)
20.6
2.9
cc,
a80
1.3
8.4
Leu
chte
nb
erge
ret
al.
1956
Seco
nd
ary
sper
mat
ocy
te(m
)26
.23.
0cc
,a
936.
98.
7L
euch
ten
ber
ger
etal
.19
56Sp
erm
atid
(m)
5.2
0.6
cc,
a32
7.5
1.8
Leu
chte
nb
erge
ret
al.
1956
Sper
mat
id(m
)8.
71.
0cc
,a
457.
62.
9L
euch
ten
ber
ger
etal
.19
56Sp
erm
atid
(m)
5.3
1.0
cc,
a33
1.6
3.0
Leu
chte
nb
erge
ret
al.
1956
Sper
mat
id(m
)8.
01.
1cc
,a
433.
33.
2L
euch
ten
ber
ger
etal
.19
56Sp
erm
atid
(m)
8.7
1.4
cc,
a45
7.6
4.0
Leu
chte
nb
erge
ret
al.
1956
Sper
mat
id(m
)9.
01.
5cc
,a
467.
84.
3L
euch
ten
ber
ger
etal
.19
56Sp
erm
atid
(m)
9.5
1.5
cc,
a48
4.5
4.4
Leu
chte
nb
erge
ret
al.
1956
Sper
mat
id(m
)6.
51.
5cc
,a
378.
64.
5L
euch
ten
ber
ger
etal
.19
56Sp
erm
ato
gon
ium
(m)
39.3
1.9
cc,
a12
19.4
5.6
Leu
chte
nb
erge
ret
al.
1956
Sper
mat
ogo
niu
m(m
)23
.42.
3cc
,a
870.
56.
7L
euch
ten
ber
ger
etal
.19
56Sp
erm
ato
gon
ium
(m)
19.7
2.6
cc,
a77
8.4
7.4
Leu
chte
nb
erge
ret
al.
1956
Sper
mat
ogo
niu
m(m
)38
.62.
8cc
,a
1205
.28.
1L
euch
ten
ber
ger
etal
.19
56Sp
erm
ato
gon
ium
(m)
51.1
3.4
cc,
a14
46.3
10.0
Leu
chte
nb
erge
ret
al.
1956
Sper
mat
ogo
niu
m(m
)26
.73.
6cc
,a
948.
410
.5L
euch
ten
ber
ger
etal
.19
56Sp
erm
ato
gon
ium
(m)
35.9
3.6
cc,
a11
49.7
10.5
Leu
chte
nb
erge
ret
al.
1956
Sper
mat
ogo
niu
m(m
)51
.93.
8cc
,a
1461
.011
.0L
euch
ten
ber
ger
etal
.19
56Sp
erm
ato
zoo
n(m
)11
.76.
4c,
a40
4.5
3.1
Ro
yere
etal
.19
88Sp
erm
ato
zoo
n(m
)11
.96.
7cc
,a
410.
93.
3R
oye
reet
al.
1988
Sper
mat
ozo
on
(m)
11.1
6.7
cc,
a38
4.9
3.3
Ro
yere
etal
.19
88Sp
erm
ato
zoo
n(m
)11
.07.
0cc
,a
380.
53.
4R
oye
reet
al.
1988
Sper
mat
ozo
on
(m)
11.6
7.0
cc,
a40
0.8
3.5
Ro
yere
etal
.19
88Sp
erm
ato
zoo
n(m
)12
.07.
3cc
,a
414.
23.
6R
oye
reet
al.
1988
Sper
mat
ozo
on
(m)
11.5
7.4
cc,
a39
8.4
3.6
Ro
yere
etal
.19
88Sp
erm
ato
zoo
n(m
)11
.47.
5cc
,a
394.
03.
7R
oye
reet
al.
1988
Sper
mat
ozo
on
(m)
13.0
7.7
cc,
a44
7.9
3.8
Ro
yere
etal
.19
88Sp
erm
ato
zoo
n(m
)11
.77.
9cc
,a
405.
13.
9R
oye
reet
al.
1988
Syn
cyti
otr
op
ho
bla
st(m
)81
.01.
8cc
,p
769.
06.
4G
alto
n19
62
Con
tin
ued
J.F. Gillooly et al.
24 Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
Tabl
e1.
Continued
Cel
lty
pe
Cel
l/nucl
ear
size
(ori
ginal
valu
es)
DN
Aco
nte
nt
(ori
ginal
valu
es)
Size
,co
nte
nt
(ori
ginal
units)
Cel
lvo
lum
e
(m3)
DN
Aco
nte
nt
(pg)
Sourc
es
Syn
cyti
otr
op
ho
bla
st(m
)72
.01.
9cc
,p
712.
36.
7G
alto
n19
62Sy
ncy
tio
tro
ph
ob
last
(m)
79.0
2.2
cc,
p75
6.6
7.7
Gal
ton
1962
Syn
cyti
otr
op
ho
bla
st(m
)77
.02.
2cc
,p
744.
17.
8G
alto
n19
62V
illo
us
stro
ma
(m)
82.0
2.1
cc,
p77
5.2
7.4
Gal
ton
1962
Vil
lou
sst
rom
a(m
)10
8.0
2.1
cc,
p92
7.1
7.5
Gal
ton
1962
Vil
lou
sst
rom
a(m
)96
.02.
4cc
,p
858.
88.
5G
alto
n19
62V
illo
us
stro
ma
(m)
102.
02.
5cc
,p
893.
38.
7G
alto
n19
62
Dat
afr
om
19ce
llty
pes
use
din
anal
yses
show
nin
Fig
ure
s1
and
2.T
he
ori
gin
al(i
.e.,
rep
ort
ed)
valu
esfo
rce
ll/n
ucl
ear
size
and
DN
Aco
nte
nta
reli
sted
inco
lum
ns
2an
d3,
and
the
ori
gin
al
un
its
inw
hic
hth
ese
valu
esw
ere
exp
ress
edar
eli
sted
inco
lum
n4.
For
anal
yses
,all
of
the
mea
sure
so
fce
ll/n
ucl
ear
size
wer
eco
nve
rted
tou
nit
so
fce
llvo
lum
e(m
3)
show
nin
colu
mn
5,an
d
mea
sure
so
fD
NA
con
ten
tw
ere
con
vert
edto
pic
ogr
ams
of
DN
Ash
own
inco
lum
n6.
Let
ters
inp
aren
thes
esfo
llow
ing
the
list
ing
of
cell
typ
ein
colu
mn
1d
eno
tew
het
her
the
valu
eis
a
mea
sure
men
to
fasi
ngl
ece
ll(s
)o
rth
em
ean
ofa
po
pu
lati
on
ofc
ells
(m).
Th
efi
rst
no
tati
on
use
din
colu
mn
4in
dic
ates
wh
eth
erce
ll/n
ucl
ear
size
mea
sure
men
tsw
ere
ori
gin
ally
rep
ort
edas
cell
dia
met
er(d
),n
ucl
ear
cro
ss-s
ecti
on
alar
eain
cali
bra
ted
arb
itra
ryu
nit
s(c
),n
ucl
ear
cro
ss-s
ecti
on
alar
eainm
2(c
c),n
ucl
ear
dia
met
erinm
(n),
orc
ellv
olu
me
inm
3(v
).T
he
seco
nd
no
tati
on
inco
lum
n4
ind
icat
esw
het
her
DN
Aco
nte
nt
mea
sure
men
tsw
ere
rep
ort
edin
un
its
of
plo
idy
leve
l(p
),ar
bit
rary
abso
rban
ceu
nit
s(a
),o
rp
ico
gram
so
fD
NA
(pg)
.Cel
ltyp
esar
ere
po
rted
bas
edo
nce
ll-t
ype
des
ign
atio
ns
give
nin
ori
gin
alst
ud
ies.
Co
mp
uta
tio
no
fm
ean
cell
volu
mes
:St
ud
ies
rep
ort
edce
llo
rn
ucl
ear
size
usi
ng
on
eo
ffo
ur
mea
sure
men
ts:
cell
volu
me,
cell
dia
met
er/
rad
ius,
nu
clea
rcr
oss
-sec
tio
nal
area
,o
rn
ucl
ear
dia
met
er/r
adiu
s.M
easu
rem
ents
of
cell
dia
met
er/r
adiu
sw
ere
con
vert
edto
cell
volu
me
assu
min
gce
lls
wer
esp
her
ical
.Mea
sure
men
tso
fn
ucl
ear
cro
ss-s
ecti
on
alar
eaan
dd
iam
eter
/rad
ius
wer
eco
nve
rted
ton
ucl
ear
volu
me
assu
min
gn
ucl
eiw
ere
sph
eric
alan
dth
atn
ucl
ear
cro
ss-s
ecti
on
alar
eas
wer
eci
rcu
lar.
All
esti
mat
eso
fn
ucl
ear
volu
me
wer
eth
enco
nve
rted
toce
llvo
lum
e
usi
ng
ap
revi
ou
sly
rep
ort
edre
lati
on
ship
bet
wee
nce
llan
dn
ucl
ear
volu
me
ofh
um
ance
lls
(Sw
anso
net
al.1
991)
:cel
lvo
lum
e¼
44.2
(nu
clea
rvo
lum
e)0
.65
asin
dic
ated
inth
ete
xt.I
no
ne
case
(Kli
nge
ran
dSc
hw
arza
cher
1960
),m
easu
rem
ents
ofn
ucl
ear
area
wer
egi
ven
inca
lib
rate
dar
bit
rary
un
its.
Inth
isca
se,w
eco
nve
rted
bac
kfr
om
arb
itra
ryu
nit
sto
un
its
ofs
qu
ared
mic
rom
eter
s
bef
ore
com
pu
tin
gn
ucl
ear
rad
ius
(in
mic
rom
eter
s).
Co
mp
uta
tio
no
fmea
nD
NA
con
ten
t:D
NA
con
ten
tm
easu
rem
ents
wer
eo
rigi
nal
lyre
po
rted
ino
ne
oft
hre
ew
ays:
asp
ico
gram
so
fDN
A,a
sp
loid
yle
vel,
or
asar
bit
rary
abso
rban
ceu
nit
so
f
DN
A.T
oco
nve
rtp
loid
yle
velt
op
gD
NA
,we
mu
ltip
lied
by
the
typ
ical
hap
loid
DN
Aco
nte
nt
ofh
um
ance
lls
(3.5
pg
[Gre
gory
2012
]).T
oco
nve
rtar
bit
rary
un
its
top
gD
NA
,we
det
erm
ined
the
mea
nar
bit
rary
un
itva
lue
of
cell
sth
atth
eo
rigi
nal
inve
stig
ato
rso
fea
chst
ud
yd
esig
nat
edas
dip
loid
,an
dth
enas
sum
edth
isco
rres
po
nd
edto
2�
3.5¼
7p
go
fD
NA
toco
nve
rtva
lues
fro
mar
bit
rary
un
its
top
gD
NA
.Th
us,
each
oft
he
seve
nst
ud
ies
inw
hic
hth
isco
nve
rsio
nw
asp
erfo
rmed
had
au
niq
ue
con
vers
ion
fact
or:
Bed
ian
dG
old
stei
n19
76(2
.75
arb
itra
ryu
nit
s/p
g
DN
A),
Kli
nge
ran
dSc
hw
arza
cher
1960
(6.9
arb
itra
ryu
nit
s/p
gD
NA
),L
euch
ten
ber
ger
etal
.195
6(0
.34
arb
itra
ryu
nit
s/p
gD
NA
),M
eek
and
Har
bis
on
1967
(2.1
arb
itra
ryu
nit
s/p
gD
NA
),
Pet
raki
s19
53(0
.66
arb
itra
ryu
nit
s/p
gD
NA
),R
oye
reet
al.1
988
(2.0
3ar
bit
rary
un
its/
pg
DN
A),
and
Swar
tz19
56(0
.37
pg)
.Th
isco
nve
rsio
nis
po
ssib
leb
ecau
seth
eFe
ulg
enst
ain
ing
met
ho
d
resu
lts
ina
lin
ear
rela
tio
nsh
ipb
etw
een
ligh
tab
sorb
ance
and
DN
Aco
nte
nt
asd
iscu
ssed
inth
ete
xt.
Ave
ragi
ng
ind
ivid
alce
llm
easu
rem
ents
:Fo
rca
ses
inw
hic
hin
div
idu
alce
llm
easu
rem
ents
wer
ere
po
rted
,w
eav
erag
edal
lce
llvo
lum
ean
dD
NA
con
ten
tm
easu
rem
ents
toyi
eld
asi
ngl
e
po
pu
lati
on
mea
nfo
rea
chce
llty
pe
wit
hin
each
stu
dy.
Mea
nce
llvo
lum
eso
fth
ese
cell
typ
esw
ere
1574
m3
(am
nio
nep
ith
elia
lcel
ls[K
lin
ger
and
Sch
war
zach
er19
60])
,293
6m
3(h
epat
ocy
tes
[Mee
kan
dH
arb
iso
n19
67])
,144
1m
3(l
ymp
ho
cyte
s[P
etra
kis
1953
]),7
64m
3(l
ymp
ho
cyte
s[B
edia
nd
Go
ldst
ein
1976
]),8
17m
3(l
ymp
ho
cyte
po
lym
orp
hs
[Bed
ian
dG
old
stei
n19
76])
,an
d
1051
m3
(mo
no
cyte
s[B
edia
nd
Go
ldst
ein
1976
]).C
orr
esp
on
din
gm
ean
DN
Aco
nte
nts
wer
e12
,19.
9,7.
1,6.
8,6.
7,an
d7.
3p
g,re
spec
tive
ly.T
hes
em
ean
sw
ere
use
din
the
anal
yses
pre
sen
ted
in
the
text
.
DNA—Cell Size Relationship in Humans
Cite this article as Cold Spring Harb Perspect Biol 2015;7:a019091 25
on July 1, 2015 - Published by Cold Spring Harbor Laboratory Press http://cshperspectives.cshlp.org/Downloaded from
ACKNOWLEDGMENTS
We thank the population biology group at Uni-versity of Florida for feedback.
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DNA—Cell Size Relationship in Humans
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2015; doi: 10.1101/cshperspect.a019091Cold Spring Harb Perspect Biol James F. Gillooly, Andrew Hein and Rachel Damiani Nuclear DNA Content Varies with Cell Size across Human Cell Types
Subject Collection Size Control in Biology: From Organelles to Organisms
Small but Mighty: Cell Size and BacteriaPetra Anne Levin and Esther R. Angert Human Cell Types
Nuclear DNA Content Varies with Cell Size across
DamianiJames F. Gillooly, Andrew Hein and Rachel
Subcellular SizeWallace F. Marshall DrosophilaSignaling in
Control of Organ Growth by Patterning and Hippo
Kenneth D. Irvine and Kieran F. HarveyA Phyletic Perspective on Cell Growth
Karl J. Niklas
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