Plant, Cell and Environment (2010) 33, 1474–1485
Scaling of heat production by thermogenic flowers:
limits to floral size and maximum rate of respiration
ROGER S. SEYMOUR
Ecology and Evolutionary Biology, University of Adelaide, Adelaide, SA 5005, Australia
ABSTRACT
Effect of size of inflorescences, flowers and cones on
maximum rate of heat production is analysed allometrically
in 23 species of thermogenic plants having diverse struc-
tures and ranging between 1.8 and 600 g. Total respiration

rate ( MCO -1
2 , mmol s ) varies with spadix mass (M, g)
according to MCO  2 = 0.482 M 0.58 in 15 species of Araceae.
Thermal conductance (C, mW °C-1) for spadices scales
according to C = 18.5M0.73. Mass does not significantly affect
the difference between floral and air temperature. Aroids
with exposed appendices with high surface area have high
thermal conductance, consistent with the need to vaporize
attractive scents. True flowers have significantly lower heat
production and thermal conductance, because closed petals
retain heat that benefits resident insects. The florets on
aroid spadices, either within a floral chamber or spathe,
have intermediate thermal conductance, consistent with
mixed roles. Mass-specific rates of respiration are variable
between species, but reach 900 nmol s-1 g-1 in aroid male
florets, exceeding rates of all other plants and even most
animals. Maximum mass-specific respiration appears to be
limited by oxygen delivery through individual cells. Reduc-
ing mass-specific respiration may be one selective influence
on the evolution of large size of thermogenic flowers.
Key-words: allometry; cone; flower; inflorescence;
temperature.
Abbreviations: F, rate of heat production; C, thermal con-

ductance; MCO 2 , rate of CO2 production; Ta, ambient
temperature; Tf floral temperature.
INTRODUCTION
Size has an overwhelming influence on the biology of
organisms. Differences in size affect nearly all aspects of
life, including not only morphology and physiology, but also
ecology, behaviour and life history (Calder 1996; Brown &
West 2000). This study concerns the fact that the reproduc-
tive organs of many ancient seed plants can produce
enough heat to raise their temperatures significantly, a char-
acteristic known as thermogenesis. It analyses the effect of
Correspondence: R. S. Seymour. Fax: +61 8 83034364; e-mail:
roger.seymour@adelaide.edu.au
1474
doi: 10.1111/j.1365-3040.2010.02190.
pce_2190 1474..1485
size on heat production, heat loss and temperature eleva-
tion among a broad range of thermogenic plants, and relates
the significant differences to the roles of thermogenesis in
pollination biology.
Depending on taxonomic preference, there are about 14
families of thermogenic plants that have a diverse range of
reproductive structures, including true single flowers (i.e.
Annonaceae, Aristolochiaceae, Hydnoraceae, Illiceaceae,
Magnoliaceae, Nelumbonaceae, Nymphaeaceae, Rafflesi-
aceae and Schisandraceae), inflorescences composed of
many florets (i.e. Araceae, Arecaceae, Cyclanthaceae) and
male and female cones (i.e. Cycadaceae, Zamiaceae) (Tang,
Sternberg & Price 1987; Seymour & Schultze-Motel 1997;
Seymour 2001a; Terry et al. 2004; Thien et al. 2009; Seymour,
Maass & Bolin 2009b). This study involves measurements
from 23 species of thermogenic plant with all three floral
types, including Araceae (15 species), Nelumbonaceae (one
species), Nymphaeaceae (two species), Magnoliaceae
(one species), Hydnoraceae (two species), Cycadaceae (one
species) and Zamiaceae (one species). For simplicity, this
paper will often identify these reproductive structures col-
lectively as ‘flowers’, for want of a better encompassing
word.
The size of flowers ranges between 1 mm to 1 m in diam-
eter, and natural selection on floral size is clearly associated
with pollination biology (Davis, Endress & Baum 2008). It
is significant that the largest floral structures in the world
are thermogenic. The largest single flower is Rafflesia arnol-
dii (Rafflesiaceae) weighing up to 7 kg (Davis et al. 2007),
the largest unbranched inflorescence is Amorphophallus
titanum (Araceae) that can stand over 2 m high (Bown
2000), the largest branched inflorescence is Corypha
umbraculifera (Arecaceae), reaching 6–8 m long (Seifriz
1924), and the largest cycad cone is Dioon spinulosum
(Cycadaceae) at 15 kg (Chamberlain 1909). The genera
Rafflesia (Patiño, Grace & Bänziger 2000), Amorphophallus
(Barthlott et al. 2009) and Dioon (Poisson 1878) are defi-
nitely thermogenic, and members of the Arecaceae
(Búrquez, Sarukhán & Pedroza 1987; Küchmeister et al.
1998) are thermogenic, although Corypha has not been
measured.
All thermogenic flowers are not large, however. Among
the aroids (Araceae), the spadices range in size from 1.5 g in
Arum maculatum (Lance 1972) to nearly 1 kg in Amor-
phophallus titanum (Baumann, Knoche & Noga 1998).
Thermogenic tissue is also not uniformly distributed, so the
© 2010 Blackwell Publishing Ltd

male florets on the spadix of Arum concinnatum can weigh
as little as 0.4 g (Seymour, Gibernau & Pirintsos 2009a).
Thus the mass ranges over three orders of magnitude and
provides ample scope for analysis.
The analysis of how biological characteristics are related
to size is called ‘scaling’ (Schmidt-Nielsen 1984; Calder
1996; Brown & West 2000). Characteristics that vary in
direct proportion to body mass are said to scale ‘isometri-
cally’, whereas those that do not scale ‘allometrically’, and
most relationships are allometric. Curves of selected vari-
ables plotted on body mass are generally fit with a power
equation of the form: Y = aMb, where Y is the variable of
interest, a is the ‘scaling factor’ (which defines the elevation
or height of the curve), M is mass (in defined units, often
grams as in this presentation) and b is the ‘power’ or ‘scaling
exponent’ of the equation (which defines the shape and
direction of the curve). Sometimes data are expressed in
mass-specific units, in a mistaken attempt to remove the
effect of body mass, for example, metabolic rate per unit of
body mass (Packard & Boardman 1999). In these cases, the
a value is unchanged, but the mass-specific b value (bm-s) is
the complement of the whole-body one (bm-s = b - 1). This
shows that taking mass-specific values does not remove the
allometric effect of body mass, except in the unique and rare
situation of isometric scaling (b = 1).
The primary aim of this study is to analyse rates of
respiration, because this is the only valid measurement of
thermogenesis, although temperature elevation is often
assumed to be a proxy. According to Newton’s Law of
Cooling, however, the rate of heat production from a body
(F) equals a value known as the thermal conductance (C)
multiplied by the difference between floral temperature
(Tf) and ambient temperature (Ta): F = C ¥ (Tf – Ta).
Thermal conductance depends on complex relationships
involving surface area and the mechanisms of heat loss by
conduction, convection, radiation and evaporation (Nobel
1999), so it is quite variable and not predictable from mass
alone. Therefore temperature elevation is an inaccurate
measure of thermogenesis. A better measure is either direct
calorimetry (measuring heat loss in a calorimeter) or indi-
rect calorimetry (measuring respiration rate and converting
to heat production with caloric equivalents). When this is
coupled with temperature elevation, it is possible to obtain
a complete picture of thermal balance.
The results of allometric analyses have several uses. Pri-
marily, they can be used to quantify the effect of mass on
respiration so that different thermogenic species can be
validly compared and, if necessary, estimates of respiration
rate from floral thermogenic tissues can be based on mass
alone. Comparisons can be made with other organisms as
well. In particular, there has been considerable recent con-
troversy over the scaling exponent for metabolic rate in
plants and animals – its theoretical basis and empirical
support (White, Blackburn & Seymour 2009). Secondly,
scaling can be used to distinguish different thermogenic
tissues within individual species and relate them to func-
tion. Thirdly, the analysis of thermal conductance can
lead to insights about the functions of thermogenesis in
© 2010 Blackwell Publishing Ltd, Plant, Cell and Environment, 33, 1474–1485
Scaling of thermogenic flowers 1475
pollination biology. Because conductance is the inverse of
insulation, flowers with low conductance tend to retain heat
within the structure, whereas those with high conductance
tend to liberate heat. Floral thermogenesis has been pro-
posed to volatilize fragrance (Fægri & van der Pijl 1979;
Meeuse & Raskin 1988), provide an energy reward to insect
pollinators (Seymour, Bartholomew & Barnhart 1983;
Seymour, White & Gibernau 2003b), facilitate arrival and
departure of insects (Búrquez et al. 1987), assist fertilization
(Li & Huang 2009) and prevent freezing (Knutson 1974).
If scent production is the major role, then thermal con-
ductance should be high to favour volatilization from the
surface, but if a reward to insects is important, then it should
be low to retain heat within the flower.
Previous physiological studies involving the size of ther-
mogenic flowers have been fragmentary. Knutson (1974)
measured the rate of heat loss from inflorescences of skunk
cabbage (Symplocarpus foetidus) and reported that the rate
depended on spadix mass (2.5–9.5 g) with an exponent of
0.73. Seymour et al. (1983) compared respiration in S. foeti-
dus (4.5 g) with Philodendron selloum (123.5 g) and found
an exponent of 0.71. Because surface-specific conductance
was similar in the two species, both appeared equally able to
release heat to the environment through their surfaces. Gib-
ernau et al. (2005) found that the temperature elevation by
inflorescences of 18 species of Neotropical Araceae tended
to increase in larger species. There are also a few other
studies dealing with the biophysics of thermogenesis, but
none has considered floral size (Seymour et al. 1983;
Breidenbach et al. 1997; Lamprecht, Seymour & Schultze-
Motel 1998; Seymour 2001a; Roemer et al. 2005).
MATERIALS AND METHODS
Selection of data
Thermogenesis of flowers is rarely constant, so this analysis
considered only the maximum respiration rates and the
simultaneous floral and ambient temperatures. Data were
selected from the literature and unpublished studies of the
author, and no available data were deliberately excluded.
Where multiple studies on the same species were involved,
the means were taken. The sample sizes varied between
studies. If possible therefore the values were means from
several plants, or calculated from regression equations
derived from several replicates, sometimes from original
data files of the author.
Floral mass
Mass data for each species were fresh weights. The ther-
mogenic part of the flower depended on the species. For
inflorescences of aroids consisting of a thermogenic spadix
surrounded by a spathe, the spathe was never included in
the mass. For aroid species in which the bisexual, male, or
sterile male florets on the spadix are thermogenic (Symplo-
carpus, Philodendron), the mass of the whole spadix was
taken, including the stalk. For aroids with spadices having
1476 R. S. Seymour
two distinct thermogenic tissues, an appendix outside of the
spathe and male florets inside the floral chamber (Amor-
phophallus, Arum, Dracunculus, Helicodiceros, Sauroma-
tum), the appendix and male florets were treated separately.
For species with true flowers (Nelumbo, Magnolia, Victoria)
the mass of the entire flower was recorded. For cycads with
cones (Cycas, Macrozamia) the entire male cone was mea-
sured. For Hydnora, a holoparasitic genus with large, fleshy
flowers that are only partly emergent from the soil, only the
mass of the thermogenic osmophores was taken.
Respiration and heat-production data
Measurements were usually taken from intact flowers,
cones and inflorescences, attached to the plant, generally in
the field. In a few cases, data were included from excised
tissues measured immediately after cutting. In recent
studies, heat production rate was measured indirectly, as
rate of CO2 production, but where O2 consumption was
measured, it was converted to CO2 production according to
the known respiratory quotient (RQ) for the group. In most
cases, the RQ was assumed to be 1, as carbohydrates are
generally metabolized (Seymour & Schultze-Motel 1998;
Seymour & Blaylock 1999), however for Philodendron
species that metabolize a proportion of lipid, the RQ was
assumed to be 0.83 (Seymour, Barnhart & Bartholomew
1984). Mass-specific respiration was obtained by dividing by
fresh mass. Heat production was obtained from rate of
CO2 production according to the calorific equivalent:
1 mmol s-1 = 0.47 W (Wieser 1986). Temperature correction
of respiration rate to 30 °C assumed a Q10 = 2. Heat produc-
tion rates of Amorphophallus titanum and Sauromatum
guttatum were derived from calorimetric analyses and con-
verted into respiration units accordingly.
Temperatures and thermal conductance
Data for temperatures were taken in association with the
respiration rates at the same time. Temperatures were
therefore measured in the respiring tissue and adjacent air,
usually inside of a respirometry hood.Thermal conductance
was calculated according to Newton’s Law of Cooling and
encompasses all four mechanisms of heat transfer: conduc-
tion, convection, radiation and evaporation.
Statistics
Regressions are standard least square linear regressions
done on log-transformed data where appropriate. Mass is
always in grams of fresh tissue. Comparison of regressions
from different groups was done with analysis of covariance
(Zar 1998), with Bonferroni adjustments, using StatistiXL
(v. 1.8; http://www.statistixl.com). For these analyses, n is the
number of species, SEb is the standard error of the slope of
regression line, CIb is the 95% confidence interval of the
slope, and R2 is the coefficient of determination. 95% con-
fidence belts for the regression means were calculated
with JMPin (v. 4; http://www.JMPdiscovery.com).
Phylogenetically-informed analysis was not possible
© 2010 Blackwell Publishing Ltd, Plant, Cell and Environment, 33, 1474–1485
because sufficient species and their cladistic relationships
were unavailable.
RESULTS
Species diversity
The thermogenic species of this study represent five catego-
ries of flowering structures (Table 1). Aroid inflorescences
are the most represented, with ten species having spadices
and appendices wholly or partially enclosed by the spathe
(e.g. Fig. 1a, c, d). Five aroid species have spadices with
appendices that are completely free of the spathe (e.g.
Fig. 1b). Four species produce true flowers (e.g. Fig. 1e, f).
Two species of cycad produce cones (e.g. Fig. 1g). Two
species of parasitic Hydnora have large, fleshy flowers with
specific, scent-producing osmophores (e.g. Fig. 1h). Unfor-
tunately, the study could not include branched inflores-
cences of palms.
Spadices
Among thermogenic species, the Araceae are the most
studied, yielding data over a broad range of size. Therefore,
maximum respiration rate, temperature elevation and
thermal conductance of spadices and appendices of 15 aroid
species are analysed allometrically first, and then other
species are compared with them. Total respiratory rate
increases in larger spadices (Fig. 2). The allometric
equation for respiration (mmol s-1) from spadices is

MCO 2 = 0.482 M
0.58
(n = 15; SEb = 0.09; CIb = 0.19; R2 =
0.76) at the measured tissue temperature. In terms of heat
production (F, Watt), the equation is: F = 0.226M0.58. These
respiration rates are derived from tissues at different
temperatures. When adjusted to a common temperature of
30 °C, assuming a temperature coefficient (Q10 = 2.0), the
equation for spadices is MCO  2 = 0.738 M
0.47
(n = 15;
2
SEb = 0.11; CIb = 0.24; R = 0.58) (Fig. 3). This equation
fits the data less satisfactorily than the unadjusted data
set. When Q10 values of 2.5, 3.0 and 3.5 were tried, the R2
value progressively decreased, indicating yet poorer fits.
The allometry of respiration rate on a mass-specific
basis (nmol s-1 g-1), for spadices yields the equation,
 −0.42
MCO 2 m-s = 482 M (n = 15; SEb = 0.09; CIb = 0.19; R2 =
0.62) (Fig. 4). In terms of mass-specific heat production
(W g-1), the equation is: Fm-s = 0.227 M-0.42. Note that the
exponent for mass-specific respiration (-0.42) equals
the exponent for whole inflorescence respiration (0.58)
minus 1.
Thermal conductance (C, mW °C-1) for spadices is
related to mass according to C = 18.5M0.73 (n = 15;
SEb = 0.13; CIb = 0.29; R2 = 0.70) (Fig. 5). The exponent of
this relationship is not significantly different from 0.67, the
value expected if conductance were proportional to surface
area. Among the spadices, there are five species that have
large appendices that protrude outside of the spathe,
namely, Amorphophallus titanum, A. konjac, Dracunculus
vulgaris, Helicodiceros muscivorus and Sauromatum
 Scaling of thermogenic flowers 1477

Table 1. Maximum rates of respiration and heat production (F) of intact thermogenic flowers, inflorescences and cones, in relation to
mass of the floral part and difference in temperature between the flowers (Tf) and ambient air (Ta). Thermal conductance is calculated
from heat production rate and temperature difference

Temperatures

Mass Maximum respiration rate F Tf Ta Tf-Ta Conductance

Species Part g nmol s-1 g-1 mmol s-1 Watts °C °C °C mW °C-1 Reference

Symplocarpus foetidus Spadix 3.76 286 1.07 0.50 18.2 -7.4 25.6 19.4 1
Symplocarpus renifolius Spadix 3.61 287 1.04 0.49 15.0 5.0 10.0 48.7 2
Arum italicum Spadix 1.84 733 1.35 0.63 31.4 20.6 10.9 58.4 3
Philodendron melinonii Spadix 25.6 200 1.88 0.88 39.5 27.2 12.3 71.8 4
Philodendron imbe Spadix 25.9 72.1 1.87 0.88 21.9 12.9 9.0 97.5 5
Philodendron appendiculatum Spadix 9.64 93.9 0.91 0.43 27.1 22.9 4.3 103 6
Philodendron solimoesense Spadix 95.1 52.8 5.02 2.36 41.1 27.5 13.6 174 3
Philodendron sellouma Spadix 137 78.2 10.46 4.91 40.3 18.5 21.8 232 7
Philodendron grandifolium Spadix 97.2 49.0 4.76 2.24 22.6 16.3 6.4 350 5
Amorphophallus konjac Appendix 213 25.4 5.41 2.54 28.5 26.9 1.6 1570 8
Helicodiceros muscivorus Appendix 3.20 450 1.40 0.66 29.8 15.2 14.6 45.1 9
Arum concinnatum Appendix 14.5 169 2.29 1.08 33.2 22.1 11.1 97.1 10
Sauromatum guttatum Appendix 10.2 213 2.17 1.02 28.0 26.0 2.0 510 11
Dracunculus vulgaris Appendix 47.2 72.0 3.62 1.70 26.6 24.1 2.5 681 12
Amorphophallus titanum Appendix 488 122 73.46 34.53 31.0 24.3 6.5 5168 13
Dracunculus vulgaris Male florets 1.89 113 0.21 0.10 18.6 10.3 8.3 12.1 12
Helicodiceros muscivorus Male florets 0.65 820 0.55 0.26 24.3 12.8 11.5 22.5 9
Arum concinnatum Male florets 0.40 846 0.33 0.16 26.2 21.4 4.8 32.7 10
Nelumbo nucifera Flower 42.2 49.7 2.10 0.99 30.0 10.0 20.0 49.4 14
Magnolia ovata Flower 55.6 30.4 1.69 0.79 29.7 24.3 5.4 147 6
Victoria cruziana ¥ amazonicab Flower 282 8.5 2.39 1.12 30.7 25.1 5.6 201 15
Victoria amazonicac Flower 271 9.7 2.63 1.24 32.6 26.5 6.1 203 16
Macrozamia machinii Male cone 167 46.0 7.70 3.62 36.5 30.1 6.4 565 17
Cycas revoluta Male cone 600 23.1 13.86 6.51 34.4 29.0 5.4 1206 15
Hydnora africana Osmophore 8.45 8.3 0.07 0.03 25.0 24.5 0.5 65.4 18
Hydnora abyssinica Osmophore 11.4 27.5 0.31 0.14 31.9 30.8 1.1 131 18

Sources of the data are: 1Knutson 1974, Seymour & Blaylock 1999, Seymour 2004; 2Seymour & Ito unpublished; 3Seymour & Gibernau
unpublished; 4Seymour & Gibernau 2008; 5Seymour & Schultze-Motel unpublished; 6Seymour, Silberbauer-Gottsberger & Gottsberger
2010b; 7Nagy et al. 1972, Seymour et al. 1983, Seymour 1999; 8Lamprecht & Seymour in press; 9Seymour et al. 2003a; 10Seymour et al. 2009a;
11
Lamprecht et al. 1991; 12Seymour & Schultze-Motel 1999; 13Lamprecht & Seymour 2010; Baumann et al. 1998; 14Seymour & Schultze-Motel
1998; 15Seymour unpublished; 16Seymour & Matthews 2006; 17Seymour et al. 2004; 18Seymour et al. 2009.
Notes: aPhilodendron selloum is called P. bipinnatifidum by Mayo (1991), but differs in many characteristics (Gottsberger & Amaral 1984).
b
Hybrid from the Adelaide Botanic Garden. cThe original publication erroneously listed mass-specific respiration rate in pmol s-1 g-1; it should
have been nmol s-1 g-1.

guttatum. Analysis of covariance (ancova) shows that these
species do not have significantly higher respiration rate, but
do have significantly higher thermal conductance (F = 17.4;
P = 0.001) than in ten other spadices with more compact
structures partly or wholly within the spathe.
There is no significant effect of mass on the temperature
elevation measured at the maximum respiration rate of
inflorescences of Araceae (Fig. 6). Because F = C (Tf - Ta),
the scaling exponents of these three factors should balance.
From the allometric equations, therefore F ⬀ M0.58 (Fig. 2),
C ⬀ M0.73 (Fig. 5) and (Tf - Ta) ⬀ M-0.15 (Fig. 6).
Comparisons with spadices
Although the data from other floral types have fewer
samples, it is nevertheless possible to compare them statis-
tically with the data from spadices and arrive at insightful
© 2010 Blackwell Publishing Ltd, Plant, Cell and Environment, 33, 1474–1485
relationships. The four true flowers of the study (Victoria,
Nelumbo, Magnolia) are all significantly different from 15
spadices and appendices when compared with ancova. In
all cases, the exponents of the allometric equations are not
significantly different, but the scaling factors (elevations)
are significantly different. Flowers are significantly lower
in total respiration rate (F = 12.6; P = 0.002) (Fig. 2), Q10-
corrected respiration rate (F = 8.2; P = 0.011) (Fig. 3), mass-
specific respiration rate (F = 12.5; P = 0.003) (Fig. 4), and
thermal conductance (F = 9.2; P = 0.008) (Fig. 5).
When individual points are considered for floral types
with less than n = 3 species, it is instructive to see if any
groups fall outside of the 95% confidence bands of the
regression mean for spadices. Thus, the cones of cycads
(Cycas, Macrozamia) and the male florets of aroids (Arum
concinnatum, Dracunculus vulgaris, Helicodiceros mus-
civorus) are not significantly different from spadices for any
1478 R. S. Seymour

(a) (b) (c) (d)

(e) (f) (g) (h)

Figure 1. Representative diversity of inflorescences, flowers and cones of thermogenic plants included in this study. All structures have
been cut vertically to reveal the internal parts, and 2 cm scale bars indicate relative sizes. Thermogenic parts are identified below, but it is
important to realize that thermogenesis in multiple organs of a species are rarely synchronous. (a) Inflorescence of Philodendron selloum
showing sectioned spathe at the back and the spadix consisting of thermogenic fertile male florets at the top, thermogenic sterile male
florets in the middle and non-thermogenic female florets at the bottom. (b) Inflorescence of Dracunculus vulgaris with broad purple
spathe subtended by the floral chamber. The thermogenic appendix of the spadix protrudes over the spathe, and thermogenic male florets,
exserting pollen strings, are above non-thermogenic female florets in the floral chamber. (c) Inflorescence of Arum italicum showing the
club-shaped, thermogenic appendix within the cut spathe above a band of non-thermogenic occlusion hairs, thermogenic male florets and
non-thermogenic female florets. (d) Inflorescence of Symplocarpus foetidus with thermogenic spadix consisting of bisexual florets inside of
the cowl-like spathe. (e) True flower of Victoria amazonica showing the white exterior petals, pink interior petals above pink thermogenic
stamens and staminodes, yellow thermogenic stylar processes (food bodies) and a capacious floral chamber. (f) True flower of Nelumbo
nucifera showing the nearly closed petals surrounding the conical, thermogenic receptacle to form the floral chamber. Thermogenic
staminal appendages block access to the stamens during the main thermogenic period. (G) Male cone of the cycad Macrozamia machinii
showing thermogenic sporophylls surrounding the thermogenic rachis. (H) Flower of the holoparasitic Hydnora africana showing two
pink tepals with orange, thermogenic osmophores on the inner faces. Two floral chambers are divided by the light-colored,
non-thermogenic antheral ring, and a light-colored, mildly thermogenic gynoecium is at the bottom.

100
Spadices
Male florets
Flowers
10
Cones
Heat production rate (Watts)
Respiration rate (mmol s–1)

10 Osmophores

0.1
Figure 2. Maximum rate of respiration,
0.1 as rate of CO2 – production, by whole
reproductive structures of thermogenic
plants. Five types of thermogenic organs
0.01
are indicated. The data are from Table 1.
0.01 Tissue temperatures were variable. The
0.1 1 10 100 1000 regression equation for whole spadices
Mass (g) 
only is MCO 2 = 0.482 M
0.58
.

© 2010 Blackwell Publishing Ltd, Plant, Cell and Environment, 33, 1474–1485
 Scaling of thermogenic flowers 1479

100
Spadices
Male florets
Q10-corrected respiration rate (mmol s–1)

Flowers
Cones
10 Osmophores

0.1
Figure 3. Maximum respiration rate
of the reproductive structures of
thermogenic plants, given as CO2 –
production rate, as in Fig. 2, but corrected
0.01 to a common temperature 30 °C assuming
0.1 1 10 100 1000 that Q10 = 2.0. The regression equation for
Mass (g) 
spadices only is MCO 2 = 0.738 M
0.47
.

variable. However, the osmophores of two species of
Hydnora have significantly lower respiration rates (by all
three measures), but similar thermal conductance values,
compared with spadices.
There is no significant effect of floral mass on the differ-
ence between floral temperature (Tf) and ambient tempera-
ture (Ta) at the point of maximum thermogenesis (Fig. 6).
There is also no significant correlation between tempera-
ture elevation (Tf – Ta) and either total respiration rate
[analysis of variance (anova): F = 0.06; P = 0.80) or mass-
specific respiration rate (anova: F = 0.86; P = 0.36). Except
for the osmophores of Hydnora species that are weakly
thermogenic, there is no apparent effect of floral type on
temperature elevation.
Mass-specific respiration of floral parts
Data on mass-specific respiration rates of floral tissues dem-
onstrate the diversity of metabolic intensity of floral tissues
(Fig. 7). Sources of the data are the same as listed in Table 1
for the same species, except where noted. The values repre-
sent the maxima recorded, from either intact flowers or
parts of them removed from the flowers and measured
immediately, but under variable temperature conditions.
The data therefore may not be the maximum possible, but
they are useful to compare species and tissues broadly,
because measured and maximal temperatures are unlikely
to be more than 10 °C apart, which would not change the
values by more than a factor of approximately twofold. In
contrast, respiration rate varies nearly 200-fold from the
lowest in the osmophores of Hydnora to the highest in the
male florets of Arum.
There are important limitations of these mass-specific
data that need to be understood. In species that were
exposed to a range in ambient temperatures, the data reli-
ably represent the maxima possible for the species. This is
the case in physiologically thermoregulatory species that
were exposed to a broad range of ambient temperatures

1000
Mass-specific respiration rate (nmol s–1 g–1)

100

10
Spadices
Male florets
Flowers
Cones Figure 4. Mass-specific respiration rate
Osmophores of reproductive structures of thermogenic
1 plants, not corrected to a common
0.1 1 10 100 1000 temperature. The regression equation for
Mass (g) 
spadices only is MCO 2 = 482 M
−0.42
.
© 2010 Blackwell Publishing Ltd, Plant, Cell and Environment, 33, 1474–1485
1480 R. S. Seymour

10000
Spadices
Male florets
Flowers
Thermal conductance (mW °C–1)

Cones
Osmophores
1000

100

Figure 5. Thermal conductance of

10 reproductive structures among
0.1 1 10 100 1000 thermogenic plants. The equation for
Mass (g) spadices is C = 18.5M0.73.

and the maximum could be clearly identified (P. selloum, N. DISCUSSION

nucifera, D. vulgaris, S. foetidus, S. renifolius). In other cases
where ambient temperature was not controlled, it is
possible that under different conditions, rates might be
different. For instance, the potentially thermoregulatory
Philodendron solimoesense, which is a member of the sub-
genus Meconostigma along with P. selloum, and possibly
Victoria amazonica, which shows evidence of thermoregu-
lation (Seymour & Matthews 2006), would probably show
higher values at lower ambient temperature. In apparently
non-thermoregulatory species, the maxima are those mea-
sured in the field. A few measurements of mass-specific
respiration come from in vitro measurements of isolated
tissues, including sporophylls of Macrozamia moorei (Tang
et al. 1987) and appendix of Arum maculatum (Lance 1972).
It is not clear whether these are maxima or whether they
were influenced by excision from the plant.
Scaling of heat production in
thermogenic flowers
This study demonstrates that the maximum, total rate of
floral heat production increases allometrically in ther-
mogenic aroid inflorescences (Fig. 2). The exponent of the
relationship (0.58) is not significantly different from 0.67,
which is the exponent expected for surface area in relation
to mass, if shape does not change. Because the exponent is
significantly less than 1.0, mass-specific respiration scales
with spadix mass to the -0.42 power, so it is apparent that
heat generation depends on floral size and is not a constant
characteristic of cells in isolation. Is there an underlying
cause of this relationship? It might be argued that allomet-
ric scaling of metabolic rate often results in an exponent

100
Spadices
Male florets
Flowers
Cones
Osmophores
Temperature excess (°C)

10

Figure 6. The difference between tissue

temperature and ambient temperature in
0.1 relation to mass of the reproductive
0.1 1 10 100 1000
structures of thermogenic plants. The
Mass (g) equation for spadices is Tf – Ta = 12.2M-0.15.
© 2010 Blackwell Publishing Ltd, Plant, Cell and Environment, 33, 1474–1485
 Scaling of thermogenic flowers 1481

Arum concinnatum male florets (night)

Arum maculatum appendix
Helicodiceros muscivorus male florets
Dracunculus vulgaris male florets
Arum concinnatum male florets (day)
Symplocarpus foetidus bisexual florets
Philodendron selloum sterile male florets
Symplocarpus renifolius bisexual florets
Philodendron melinonii spadix
Nelumbo nucifera receptacle
Arum concinnatum appendix
Victoria amazonica stylar processes
Philodendron solimoesense male and sterile florets
Magnolia ovata anthers
Figure 7. Mass-specific respiration rates
Arum idaeum male florets
Dracunculus vulgaris appendix from tissues of thermogenic flowers and
Macrozamia machinii male cone cones. Rates are the maximal measured
Arum creticum male florets values (but see text). Respiration is CO2
Magnolia ovata petals production or converted from O2
Macrozamia moorei male cone consumption assuming an appropriate
Hydnora africana osmophore RQ. Sources of data for each species are
Hydnora africana anthers given in Table 1, except for Arum
0 100 200 300 400 500 600 700 800 900 1000
maculatum (Lance 1972) and Macrozamia
Respiration rate (nmol s–1 g–1) moorei (Tang et al. 1987).

close to 0.75, because it is limited by fractal organization
of rate-limiting supply networks (West, Brown & Enquist
1997), however this cannot apply in the present case,
because the energy substrates for respiration already reside
in the cells of most species and the oxygen delivery system
is not fractal in organization (Seymour 2001b). An alterna-
tive explanation relies on the functions of heat production
in thermogenic flowers. It would be expected that heat pro-
duction is related to surface area, because the common
explanation for thermogenesis is volatilization of insect-
attracting fragrances that are released through the surfaces
(Meeuse & Raskin 1988). This idea seems more reasonable,
despite that fact that floral shapes differ greatly between
species.
Since attraction by scent is important to thermo-
genic flowers, it is reasonable to hypothesize selection for
larger size, especially surface area, to be coupled with ther-
mogenesis. Increasing floral size alone can increase scent
production (Valdivia & Niemeyer 2006), but adding heat
production magnifies the effect. Pollination of Arum itali-
cum is more effective in inflorescences with larger appen-
dices and also plants that are closer together (Méndez &
Díaz 2001), so selection should promote both large size and
high population density. However, evolutionary theory pre-
dicts that there should be a trade-off between floral size
and floral number, presumably because limited resources
cannot accommodate both (Worley & Barrett 2000).
Empirical studies demonstrate that there is an inverse rela-
tionship between flower size and number (Sargent et al.
2007). Although not studied quantatively, this relationship
appears to hold for thermogenic flowers. The population
densities of large-blossomed plants such as Amorphophal-
lus (Beath 1996) and Rafflesia (Davis et al. 2008) are quite
low in comparison with smaller species such as Arum
(Méndez & Díaz 2001). Floral scents are powerful attracta-
nts for insects (Raguso 2008), but dispersal may be a
problem in the still air of tropical forests. The number of
© 2010 Blackwell Publishing Ltd, Plant, Cell and Environment, 33, 1474–1485
beetles flying between inflorescences of Amorphophallus
johnsonii decreases as distance increases, and if inflores-
cences are over 40 m apart, they fail to set seed (Beath
1996). Evolution of large inflorescences under such circum-
stances may be favoured, because, not only is the volatiliza-
tion enhanced, but also convective currents generated by
the warmed air can carry the scent to the canopy (Barthlott
et al. 2009).
The present study indicates that temperature excess is
largely independent of floral size. Among the spadices,
ranging from 1.84 g Arum italicum to 137 g Philodendron
selloum, there is no significant effect on temperature excess,
and even no tendency for the regression line to rise (Fig. 6).
At first this seems surprising, because one would expect that
a larger structure would become warmer with a given rate
of heat production. However, the maximum metabolic rate
of spadices increases with M0.58 (Fig. 2), and thermal con-
ductance increases with M0.73 (Fig. 5), neither of which is not
significantly different from M0.67, the exponent for surface
area, so there is no significant change in temperature excess.
These results differ from those of Gibernau et al. (2005)
who measured maximum spadix temperatures of 18 species
of Neotropical Araceae in relation to the volume of the
thermogenic tissue (male and sterile male florets). They
showed that temperature excess of the largest spadices
(53 cm3) was higher than smaller spadices (<1 cm3). The
allometric equation for temperature excess derived from
their data is (Tf – Ta) = 1.97M0.57 (assuming volume is
proportional to mass). According to the equation,
F = C ¥ (Tf - Ta), and assuming that thermal conductance is
proportional to surface area, then total F ⬀ M1.24. This is
clearly unreasonable, because scaling of metabolic rate of
any organism almost never exceeds M1.0. In fact, Gibernau
et al. (2005) assumed a constant mass-specific heat produc-
tion of 29.5 mW g-1 in all species, or F ⬀ M1.0, in several
biophysical models of convective, conductive and radiant
heat loss. Under this assumption, C ⬀ M0.43 in their species.
1482 R. S. Seymour
This seems more reasonable, because the spadices of many
of the smaller species are completely exposed, whereas
those of the larger ones tend to be surrounded by the
spathe. Exactly how thermal conductance scales with size of
neotropical aroids will eventually require knowledge of the
rate of heat production.
Thermal conductance and roles
of thermogenesis
Thermal conductance is a measure of how easily heat is lost
from the flower. Comparisons of conductance between
species and between organs can give insight into the roles of
heating of thermogenic flowers (Fig. 5). On one hand,
aroids with large appendices that project outside of the
spathe (Amorphophallus, Dracunculus, Helicodiceros and
Sauromatum) have significantly higher conductance than
those with appendices or spadices within the spathe. Free
appendices facilitate the dispersal of scent, but also lose
heat easily. Therefore the total heat production of free
appendices can be high, but the temperature elevation rela-
tively low (Table 1). This shows that temperature elevation
is not always a good indication of the level of thermogen-
esis, particularly if evaporative cooling by transpiration is
involved. For instance, heat production in the appendix of
Dracunculus vulgaris peaks at 1.7 W, yet the temperature
excess is only 2.5 °C (Seymour & Schultze-Motel 1999). On
the other hand, thermogenic true flowers (Victoria,
Nelumbo, Magnolia) all have significantly lower thermal
conductance than aroid inflorescences (Fig. 5). In contrast
with Dracunculus, flowers of Nelumbo nucifera produce
less heat (1 W), but develop a 20 °C temperature elevation.
This is interpreted as an adaptation of overlapping and
closed petals to retain heat in the flower as a thermal
reward to insect visitors. It is significant that the petals
of these flowers form a chamber during the female phase
of the protogynous sequence, when thermogenesis is
maximum, and open fully only during the male phase, when
heat production declines. Such ‘Chamber Blossoms’
coevolved with diverse beetles (Bernhardt 2000), and gen-
erally maintain thermogenesis while beetles are in resi-
dence. Thermogenesis by fertile or sterile male florets or
bisexual florets on the aroid spadix is associated with
moderate thermal conductance (Fig. 5). The floral chamber
walls or the spathe offers some protection from heat loss,
but not as effective as petals, because they are essentially a
single layer and more or less open, either on the top of the
floral chamber or one side of the spathe. The male florets
are thermogenic overnight, when resident insects benefit
from the heat as they climb over the florets.
It is important to distinguish thermogenesis from ther-
moregulation. Some thermogenic flowers or their organs
show discrete episodes of heat production that rise to a
peak and then decline (Gibernau, Macquart & Przetak
2004), and there is no clear relationship between heat
production and ambient temperature. Others show longer
periods of thermogenesis during which heat production
responds inversely to changes in ambient temperature such
© 2010 Blackwell Publishing Ltd, Plant, Cell and Environment, 33, 1474–1485
that floral temperature is nearly constant; these are the
physiologically thermoregulating species (Seymour 2001a).
Because insects are resident during the thermoregulatory
period, the phenomenon is thought to benefit them. It is
significant that thermal conductance is relatively low in
thermoregulating species. This is certainly true of Nelumbo
nucifera that thermoregulates very precisely at about
30–36 °C (Seymour, Schultze-Motel & Lamprecht 1998;
Seymour & Schultze-Motel 1998), and Victoria amazonica
that maintains temperatures between 29 and 35 °C in the
floral chamber. Thermoregulation by male florets inside the
floral chamber is evident in Symplocarpus species (Knutson
1974; Seymour & Blaylock 1999; Ito 2003; Seymour,
Lindshau & Ito 2010a), Philodendron selloum (Nagy, Odell
& Seymour 1972; Seymour 1999), Dracunculus vulgaris
(Seymour & Schultze-Motel 1999) and in the ‘pseudother-
moregulatory’ Helicodiceros muscivorus (Seymour, Giber-
nau & Ito 2003a), consistent with moderate thermal
conductance. However, there are exceptions to the general
association between thermoregulation and thermal reward.
Magnolia ovata flowers do not thermoregulate, but have
two thermogenic episodes associated with arrival and
departure of beetles (Seymour, Silberbauer-Gottsberger &
Gottsberger 2010b), and thermoregulation is ineffective in
male florets of Arum concinnatum (Seymour et al. 2009a)
and Philodendron melinonii (Seymour & Gibernau 2008),
although the mechanism may be present. Nevertheless,
there are absolutely no examples of appendices that physi-
ologically thermoregulate, which is consistent with high
thermal conductance, strong scent production and little
contact with insect pollinators.
Thermal conductance values of the cones of cycads are
indistinguishable from the aroid spadices (Fig. 5). The
sporophylls of the cones in the studied species are relatively
tightly packed and retain heat among them. Activity of
weevils exposed to the 6 °C elevation among the sporo-
phylls is enhanced (Seymour, Terry & Roemer 2004). The
osmophores of Hydnora species are recessed on the
inward-facing sides at the tops of three tepals, therefore
occupying a site similar to that of aroid florets and resulting
in similar values of thermal conductance. However, in these
cases, insects attracted to the flower do not touch the osmo-
phores, but fall into a chamber below (Bolin, Maass &
Musselman 2009). Therefore they do not benefit from
osmophore thermogenesis, which in any case is exception-
ally low (Table 1).
Limits to total floral thermogenesis
The large size of Amorphophallus titanum testifies that
there is probably no limit to thermogenesis other than the
ability to support the tissue that generates heat. But ther-
mogenesis can be limited biochemically, by rising tempera-
ture, as clearly shown in thermoregulating species. Here,
respiration gradually increases with rising tissue tempera-
ture to a maximum, called the ‘switching temperature’, and
then declines at higher temperatures within the regulated
range (Seymour et al. 2010a). There is also evidence that the

maximum temperature of non-thermoregulating aroid
inflorescences is somewhat independent of ambient tem-
perature (Seymour et al. 2003a; Wagner et al. 2008; Seymour
et al. 2009a). In both cases, thermogenesis may be limited by
a similar biochemical mechanism that appears to be related
to the function of the cyanide-insensitive pathway, catalysed
by the alternative oxidase (AOX). In Arum maculatum,
there is a steep inhibition of AOX activity at temperatures
exceeding 32 °C (Wagner et al. 2008). The fact that respira-
tion decreases at temperatures above 30 °C in N. nucifera,
15 °C in S. renifolius and 20 °C in D. vulgaris indicates that
the upper limit of temperature varies among thermoregu-
latory species (Seymour et al. 2010a), and probably among
non-thermoregulating species as well. If the mechanism is
widely spread among in thermogenic plants, then it could
account for limitations of thermogenesis and maximum
temperatures.
Limits to mass-specific respiration rate
The highest respiration rates occur in the smallest floral
parts, the male florets of aroids that reach about
900 nmol s-1 g-1 (Figs 4 & 7). The rates exceed those of any
other plant tissue, and therefore thermogenic inflorescences
are often used in the study of plant respiration. In fact,
the rates are equivalent to the highest rates in the
animal kingdom. For example, respiration reaches 900–
1000 nmol s-1 g-1 in flying insects (White, Matthews &
Seymour 2008; Schippers, Dukas & McClelland 2010), and
560 nmol s-1 g-1 in hummingbirds (Norberg 1990).
The diffusion pathway for oxygen may ultimately limit
maximum thermogenesis in flowers. Because flowers have
no convective system for gas transport, they rely on diffu-
sion through the gas phase. The only thermogenic species
to have its diffusion pathway analysed is Philodendron
selloum (Seymour 2001b). The successive barriers to diffu-
sion in the sterile male florets are (1) the interforal space (2)
the stomata (3) the network of intercellular gas spaces, and
(4) the cytoplasm of parenchymal cells. Each of these bar-
riers in the oxygen cascade accounts for about 2–5 kPa
decrease in oxygen partial pressure, amounting to a
decrease from about 20.5 kPa in air to 5 kPa at the lowest
point at the center of the floret. Lowering external oxygen
to about 80% of atmospheric causes decreased oxygen
uptake by the florets (Seymour et al. 1984), showing that the
florets are near the threshold for diffusion limitation. The
anatomy of the diffusion pathway is not known in other
aroids, and there is the possibility that the size and density
of stomata and interstitial gas spaces might differ. However,
one invariant is the intracellular gas diffusion coefficient.
Following a model of Thumfort, Atkins & Layzell (1994),
one calculates that the parenchyma cells of Philodendron
selloum would have a gradient of 4 kPa from the outside
to the center, assuming the measured rate of oxygen
consumption of 310 nmol s-1 g-1 and a cell radius of 16.5 mm
(Seymour 2001b). Assuming the same size of cell, and a
respiration rate of 890 nmol s-1 g-1 for Arum maculatum
appendix tissue (Fig. 7), the model predicts a gradient of
© 2010 Blackwell Publishing Ltd, Plant, Cell and Environment, 33, 1474–1485
Scaling of thermogenic flowers 1483
11.6 kPa. Using a radius of 33 mm as reported in Arum
maculatum (Simon & Chapman 1961), the gradient
becomes 46 kPa, which is clearly impossible because atmo-
spheric oxygen is only 20.5 kPa. Perhaps there is an error in
Simon and Chapman’s data for cell size or the mitochondria
are not uniformly distributed. It is known that the cells of
the Sauromatum guttatum appendix are about 11 mm in
radius (Skubatz & Kunkel 2000), and the mass-specific rate
is about 213 nmol s-1 g-1 (Lamprecht et al. 1991), although
this may not be maximal. The gradient across such a cell
would be 5.2 kPa, and similar to that in P. selloum. These
considerations lead to the hypothesis that the maximum
respiration rate is limited by oxygen delivery to the cells.
Only a morphometric analysis of the diffusion pathway in
highly thermogenic tissue could confirm this. If this hypoth-
esis is true, then it may help explain why thermogenic
species have the largest flowering structures. To increase the
effect of heating without increasing mass-specific respira-
tion, it is necessary to become large.
ACKNOWLEDGMENTS
The original research for many of the studies used for this
paper was funded by the Australian Research Council, the
Alexander von Humboldt Foundation and the University
of Adelaide. I thank many colleagues who were involved
with the studies that provided the data for this analysis. I
acknowledge the advice of Robert Raguso.
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Received 22 April 2010; received in revised form 18 May 2010;
accepted for publication 23 May 2010